BULLETIN

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BULLETIN DU MUSÉUM NATIONAL D'HISTOIRE NATURELLE

3 e série, n° 258, septembre-octobre 1974, Zoologie 180

Observations upon some specimens of the genus Periclimenaeus

Borradaile (Decapoda Natantia, Pontoniinae)

originally described by G. Nobili

by A. J. Bruce *

Résumé. — Les spécimens attribués par Nobili à C. rhodope et à C. hecate, et placés main¬

tenant dans le genre Periclimenaeus Borradaile ont été réexaminés. Il est nécessaire d’attribuer

les spécimens de P. rhodope à trois, peut-être quatre espèces de Periclimenaeus : P. rhodope s. str.,

P. arabicus (Caïman), P. djiboutensis Bruce et Periclimenaeus sp.

Les spécimens types de P. djiboutensis déjà décrits sont considérés comme faisant partie du

matériel de Nobili. P. hecate est distinct de P. tridentatus (Miers) et le spécimen identifié par

Nobili comme une « anomalie » de P. hecate est décrit comm" une nouvelle espèce, P. nobilii. Un

lectotype est désigné pour P. rhodope (Nobili) et une clé est fournie pour l’identification provi¬

soire de P. tridentatus (Miers) et des espèces proches.

In 1904, Nobili briefly described some new Pontoniinid shrimps collected from Dji¬

bouti by H. Coutière. They were then placed in the genus Coralliocaris Stimpson, sub-

genus Onycocaris Nobili. In 1906, another specimen Iront the Persian Gulf, collected

by Bonnier and Pérez, was referred to the same genus. Subsequently, also in 1906,

Nobili provided a moie detailed and illustrated report upon these specimens as Corallio¬

caris (O.) rhodope and C. hecate. Later, Holthuis (1952) referred a further single speci¬

men from Indonesia to Nobili’s C. rhodope, but transferred it from Coralliocaris to the

genus Periclimenaeus Borradaile. C. hecate was similarly transferred to Periclimenaeus.

These specimens hâve been preserved in the collections of the Muséum national d’His-

toire naturelle, Paris, and through the kindness of Dr J. Forest, were made available for

the présent examination.

The specimens are unfortunately incompletely preserved and more than half the

complément of second pereiopods is lacking and very few of those preserved are attached

to the bodies. Very few ambulatory pereiopods hâve been preserved.

The collection consists of six tubes, four labelled as Coralliocaris ( Onycocaris) rhodope

types, one as Coralliocaris hecate and one as “ ? anomalie de Coralliocaris hecate ”. The

tube with most of the specimens of C. rhodope was found to consist of several readily sépa¬

rable species, the details of which are provided later, together with some information

upon other related species.

* East African Marine Fisheries Research Or ganization, Zanzibar.

Présent address : 26 St. Peter’s Grove, Canterbury , Kent, Great Britain.

258, 1

1558

A. J. BRUCE

According to Nobili’s 1906 publications there were fourteen specimens of C. (O.)

rhodope and two of C. hecate from the Red Sea, and one specimen of C. (O.) rhodope from

the Persian Gulf. An additional specimen from the Red Sea, identified by Nobili, but

apparently to which no published référencé was made, was also examined.

NOTES ON THE SPECIMENS

1. Periclimenaeus rhodope (Nobili)

(Fig. 1-2, 3 A-R, 7 A-R)

Coralliocaris ( Onycocaris ) rhodope Nobili, 1904 : 233.

Material examined : 2 <$, 2 ovigerous $, Djibouti, coll. H. Coutière.

Description

Four out of the fourteen specimens originally placed by Nobili to tliis taxon are con-

sidered to belong to the species referred to in bis description. Tliey consist of one male,

with both second pereiopods still attached, together with another male and two ovigerous

fe males.

In the males there are 9/2 and 6/0 rostral teeth and in the females 7/0 and what could

be interpreted either as 9/1 or 10/0. In ail specimens tlie fîrst rostral tooth is just at or

behind the posterior orbital margin. In the four specimens the rostrum shows considérable

variation in dentition, and depth of the lamina, but in ail is more or less horizontal. The

supra-orbital spines are small and conical. The orbit is obsolescent.

The antennae are remarkable particularly for the very great size of the antero-lateral

spine of the scaphocerite, which far exceeds the rounded anterior margin of the lamina,

and the acutely produced disto-lateral angle of the proximal segment of the antennular

peduncle, which extends well beyond the anterior margin of the intermediate segment.

The eyes are moderately slender with the cornea oblique and narrower in diameter than

the stalk.

The first pereiopod is slender with the carpus and merus subequal. The chela is also

slender, about 0.6 of the length of the carpus. The palm in the female is 3.8 as long as

wide and 2.5 times the length of the fingers, which are slender, with simple cutting edges

and small pointed hooked tips. In one male both second pereiopods were still attached

and can therefore be defînitely attributed to this species. The chelae are markedly unequal

and dissimilar. In the major chela the palm is broad and flattened, about 0.6 times as

long as wide and tapering distally. The médian and latéral margins bear numerous small

conical tubercles and a few short setae. The dactylus is stout with the tip strongly curved

distally. The molar process is rather small and only very vaguely demarkated, particu¬

larly distally, where it merges with the cutting edge. The dactylus is about 0.4 of the

length of the palm and is at least twice as long as broad. The fixed fînger is curved, with

a single small blunt tooth on the proximal half of the dorsal border. The tips of the dac-

1560

A. J. BRUCE

Fig. 2. — Periclimenaeus rhodope (Nobili). Lectotype, male : A, scaphocerite ; B, posterior telson spines ;

C, first pereiopod ; D, chela of first pereiopod ; E, third pereiopod ; F, propod and dactylus of third

pereiopod ; G, disto-lateral angle of exopod of uropod. — Female : H, disto-lateral angle of exopod

of uropod ; I, posterior telson spines ; J, dorsal telson spines. — Male : K, disto-lateral angles of exo¬

pod of uropod.

tyls are smoothly continuons with the cutting edges and are poorly demarkated. The

minor chela is essentially similar but smaller, the whole chela being the length of the palm

of the larger chela. The fingers are relatively longer and slimmer and distinctly exceed

half the length of the palm. The borders of the palm are spinulate. The cutting edge

of the dactylus is sinuous and of the fixed finger broadly concave, with a small triangular

tooth on the proximal third of the edge. The tips of the fingers are again feebly demar-

ON SOME SPECIMENS OF PERICLIMENAEUS

1561

kated from the cutting edges. The carpus is short and stout, broadly expanded distally

and with numerous small acute tubercles over the dorso-lateral aspect. The merus bears

several similar small acute tubercles (4-6) on the ventral border and a few minute tubercles

are also présent on the ischium.

The ambulatory pereiopods bear a characteristic dactylus which is distinctly biun-

guiculate with minute denticles along the ventral margin of the unguis and the corpus

proximal to the accessory spine. The dactylus is 2-2 % times longer than the basal width

and that of the third pereiopod is more slender than the fifth. The corpus has 4-6 ventral

teeth and the unguis 5-6. In one male the tip of the unguis appears to be articulated.

The ventral border of the propod is strongly spinulate with 10 well developed spines. The

carpus is unarmed and the ventral margin of the merus bears a sériés of small acute tubercles

only.

The uropods are broad with entire unarmed latéral borders to the exopods but the

disto-Iateral angle is acutely produced and may be assymetrieal, and bears from 1-3 strong

spines.

The telson is narrower in the female than in the male, 2.1 and 1.8 times longer than

wide respectively. The two pairs of dorsal spines are well developed, subequal in the

male, the posterior slightly shorter than the anterior in the female. In the female the

anterior spines are doser to the anterior margin of the telson and the posterior spines are

more posteriorly situated than in the male, in which they are at about the middle of the

telson length. The latéral posterior spines are short and stout and dorsally situated.

The tips of the intermediate and submedian spines ail reach to about the same level poste¬

riorly. They are distinctly more slender in the male than in the female. The distal halves

of the submedian spines are plumose.

Remarks

Periclimenaeus rhodope is known only from Djibouti. The specimen referred to this

species by Holthuis (1952) has been separated as P. holthuisi by Bruce (1969). The

main différence is that the disto-lateral spine of the scaphocerite greatly exceeds the lamina

in P. rhodope (Nobili). Other différences are that the ventral borders of the dactyls of

the ambulatory pereipods are spinulate, whereas in P. holthuisi they are crenulate, inclu-

ding the accessory spine, and there is no post-rostral tooth situated behind the level of

the orbital margin, a feature that is présent in P. holthuisi.

It must be emphasized that only the male and one female can be attributed with

certainty to Nobili’s species as only in these two specimens was one of the diagnostic ambu¬

latory pereiopods still attached. The other two specimens that hâve been here identified

as belonging to this species cannot with complété certainty be distinguished from P. ara-

bicus (Caïman) as no distinctive features could be found in rostrum, carapace, uropods

or telson. The anterior border of the lamina of the scaphocerite is rounded in P. rhodope

but more truncate in P. arabicus and this feature appears to separate the two species.

The male specimen is designated as the lectotype of Periclimenaeus rhodope (Nobili).

The specimen is moderately well preserved, with ail pereiopods except the right fourth

pereiopod.

Fig. 3. — Second pereiopods of Coralliocaris (O.) rhodope « types ». Periclimenaeus rhodope (Nobili) s. str.,

lectotype (J : A, carpus and chela of major second pereiopod ; B, minor second pereiopod. — Pericli¬

menaeus arabicus : C, major second pereiopod ; D, fingers of major chela ; E, major second pereiopod ;

F, minor second pereiopod. — Periclimenaeus djiboutensis Bruce : G, major second pereiopod ; H,

minor second pereiopod.

ON SOME SPECIMENS OF PERICLIMENAEUS

1563

2. Periclimenaeus arabicus (Caïman)

(Fig. 3, G-F ; 4-6 ; 7, C-H)

Periclimen.es ( Periclimenaeus ) arabicus Caïman, 1939 : 210-211, fig. 4.

Matériel examined : 2 (J, 2 ovigerous Ç, Djibouti, 1906, coll. H. Coutière.

Description

Four specimens are referred to this species. In ail specimens one or more of the cha-

racteristic ambulatory pereipods is still attached to the body, thus enabling the specimens

to be separated from the P. rhodope specimens.

None of the four specimens hâve any of the second pereiopods attached. Calman

(1939) lias provided a figure of the major second pereiopod, and Dr R. W. Ingle has kindly

provided further information concerning the holotype, so that it is felt that three of the

second detached pereiopods can be allocated to these specimens with a considérable degree

of confidence. The relatively long and slender setose fingers of the minor second pereio¬

pod of the holotype, particularly show a close resemblance to the présent specimens.

One male and the smaller ovigerous female clearly resemble each other, particularly

in the form of the rostrum, telson and uropods. The second male and larger ovigerous

female show différences in these parts and should possibly be referred to a separate but

closely related species. This step is not felt to be justifîed in view of the absence of any

associated second pereiopods.

Fig. 4. — Periclimenaeus arabicus (Caïman), anterior carapace and rostrum : A, male; B, smaller ovi¬

gerous female ; C, larger ovigerous female.

258 , 3

1564

A. J. BRUCE

Fig. 5. -— Periclimenaeus arabicus (Caïman) : A, antennule ; B, antenna. — Small ovigerous female :

C, first pereiopod ; D, chela of first pereiopod; E, third pereiopod ; F, propod of third pereiopod. —

Larger ovigerous female : G, third pereiopod. — Male : H, third pereiopod.

In the first pair, the male’s rostrum has nine rather small acute teeth dorsally with

a single small distal ventral tooth, and the ventral border is straight. The posterior tooth

is just situated on the carapace. The rostrum reaches to the middle of the distal segment

of the antennular peduncle. The rostrum in the female is basically similar but there are

ON SOME SPECIMENS OF PERICLIMENAEUS

1565

eight dorsal teeth and the ventral border is convex. In the larger female of the second

pair the rostrum is shorter with only six dorsal teeth and without a ventral tooth. The

teeth are slightly larger and more acute, and the fîrst tooth is situated slightly more poste-

riorly. The rostrum of the male is siniilar. In ail specimens a small acute conical supra-

orbital tubercle is présent.

The basal segment of the antennule is broad with a large acute disto-lateral process

and a short, broad stylocerite. A very strong médial ventral spine is présent. The basi-

cerite is unarmed laterally. The scaphocerite is broad, with the lamina almost twice as long

as wide and the anterior margin truncated rather than rounded. The disto-lateral spine

is very well developed and far outreaches the lamina.

The fîrst pereiopod of the small female is slender, with the merus slightly longer than

the carpus, which is 1.75 times the length of the chela. The chela is slender with medially

curved fmgers. The palm is 4.4 times longer than wide, and tapers slightly distally. The

fîngers are 0.43 of the palm length, slender with strong hooked tips and straight cutting

edges.

The second pereiopods are ail detached. The major second pereiopods attributed

to these specimens are robust. The palm of the chela is subcylindsical, slightly flattened,

tapering slightly distally, and about 1.7 times longer than wide. The surface is covered

with numerous small tubercles in the larger example but these are less marked in the smal-

ler. The fîngers are short and stout, about 0.45 of the length of the palm. The dacty-

lus is moderately compressed and has a strongly convex outer margin, with an acute stron-

gly hooked tip. The molar process is large and distinct from the anterior cutting edge,

and lits into deep fossa on the proximal cutting edge of the fixed fînger. The cutting

Fig. 6. — Periclimenaeus arabicas (Caïman), telson and disto-lateral angle of exopod of uropod : A-B,

male ; C D, smaller ovigerous female ; E-F, larger ovigerous female.

1566

A. J. BRUCE

ëdges of both Angers are short and blunt. The dorsal proximal rnargin of the fixed fînger

bears a long acute process and a short blunt lobe is présent in a similar position on the

ventral aspect. The carpus is short and stout and devoid of spines or tubercles. The

merus and ischium are robust, with numerous small acute tubercles along the ventral

borders.

The minor second pereiopod is densely tuberculate but bears numerous long setae

on its distal dorsal aspect. The palrn is 1.5 times longer than broad and tapers strongly

distally. The Angers are about 0.85 of the length of the palm. The dactylus is compres-

sed and slender and about 4 times longer than the greatest width. The tip is strongly

hooked and the cutting edge long, straight and entire. The Axed Anger is similar but

bears a small tooth proximally, separated by a semi-circular gap from a larger tooth situated

-at the level of the proximal end of the dactylar cutting edge. The carpus, merus and ischium

are similar to those of the major second pereiopod.

The third pereiopods are represented in each specimen and show no essential différences.

The ventral rnargin of the propod is strongly spinulate in ail, with a tendency to develope

a double row of spines proximally. The carpus bears a few small conical tubercles ven-

trally. Many acute tubercles are présent along the ventral border of the merus and a

few are also présent on the ischium. The fourth and Afth pereiopods are nnich less stron¬

gly spinulate. In the fourth, the merus has only Ave ventral spines and in the Afth onlv

two.

Only the male of the Arst pair of specimens has the third to Afth pereiopods preserved

with the dactyls, which are long and slender and al least four times longer than the basal

width. In this pair of specimens the ventral border bears a sériés of small teeth, 3-7 in

numher, situated on the proximal two thirds or one half, and best developed on the third

pereiopod. The distal ventral extremity of the corpus bears a small acute accessory spine.

The unguis is distinctly demarkated from the corpus, curved, with a sériés of 7-9 small

acute teeth along the ventral rnargin. In the larger ovigerous female, the unguis is similar

but the ventral teeth of the corpus are more numerous, 9 in number, acute posteriorly

and blunter distally, extending along the whole ventral border. The distal ventral angle

is bluntly produced but may hâve been damaged.

The uropods are broad. The protopodite is not produced posteriorly and the latéral

rnargin of the exopod is entire, without spines or denticulations. In the Arst pair of spe¬

cimens the disto-lateral angle is acutely produced with a small latéral and a larger médian

spine on both sides in male and female. In the second pair of specimens the disto-lateral

angle is acutely subrectangular with a single large spine on both sides.

In the Arst pair of specimens the telson is moderately broad, about 1.7-1.8 times longer

than wide, and tapering strongly posteriorly. In the second pair of specimens the poste-

rior rnargin has a distinct acute médian process but in the larger female of the second pair

this is very much reduced. Two pairs of large dorsal spines are présent, ail situated on

the anterior half of the telson, with the anterior pair slightly larger than the posterior pair.

The posterior spines are situated at 0.36 of the telson length and are overlapped by the

tips of the anterior spines. In the second pair, the posterior spines are at 0.48-0.53 of

the telson length and are remote from the tips of the anterior spines. The usual three

pairs of posterior telson spines are présent, with the small latéral spines slightly dorsally

situated. The submedian spines are coarsely plumose and relatively stout.

ON SOME SPECIMENS OF PERICLIMENAEUS

1567

Fig. 7. — Periclimenaeus rhodope (Nobili) : A, dactylus of third pereiopod ; B, dactylus of fifth pereio-

pod. — Periclimenaeus arabicas (Caïman), male : C, dactyl of third pereiopod ; D, unguis ; E, dacty¬

lus of fifth pereiopod; F, dactylus of fifth pereiopod. — Female : G, dactylus of third pereiopod ; H,

carpo-propodal joint of male third pereiopod.

Remarks

In view of the mixed nature of Nobili’s Periclimenaeus material, it seems probable

that the two pairs of specimens and the larger ovigerous female are of separate origin,

probably front different host sponges. The fîrst pair closely resemble Calman’s descrip¬

tion of P. arabicus in the telson but the second pair agréés more closely in the fornt of the

1568

A. J. BRUCE

rostrum and the uropods. The dactylus of the third pereiopod shows slight différences

in ail specimens, which may he possibly only due to individual rather than spécifie varia¬

tion. There is a very close similarity to the dactylus of the holotype (fig. 15, A). The

two pairs may représente distinct species but without the associated second pereiopods

no conclusion can be reached.

Periclimenaeus arabicus has not been reported since its original description and is

known only from the South Arabian coast. P. ohshimai Fujino (1967) is very closelv rela-

ted and appears to differ only in that the posterioi dorsal telson spines are about half the

length of the anterior.

3. Periclimenaeus djiboutensis Bruce

(Fig. 8 ; 9 ; 13 A-D)

Periclimenaeus djiboutensis Bruce, 1970 : 307-308.

Material examined : 1 (J, 2 ovigerous Ç, Djibouti, 1906, coll. II. Coutière.

Description

The three specimens are ail incomplète with no ambulatory pereiopods and most of

the mouthparts missing. Noue of the second pereiopods are attached, but two isolated

detached second pereiopods are considered to belong to these specimens. The remaining

mouthpaits hâve not heen examined.

The male and one of the ovigerous females hâve the rostrum intact. In botli, it is

well developed, strongly compressed and ventrally inclined. The ventral border is straight

and without teetli. The dorsal border is gently convex with nine long slender dorsal teeth

in the male and ten in the female. Ail dorsal teeth are situated well in advance of the

posterior orbital margin. The tip of the rostrum is acute and reaches to the middle of

the intermediate segment of the antennular peduncle.

Supra-orbital spines are completely absent. The orbit is obsolescent but the antennal

spine is long and slender, particulaily in the female where it appears to be rather dorsally

orientated. The anterior margin of the branchiostegite is deeply notched and the antero-

ventral angle is produced and rounded.

The proximal segment of the antennular peduncle is very broad basally, about 0.8

of the length of the segment. It tapers strongly distally, where the latéral border is con¬

cave, and the disto-lateral angle is strongly produced to the middle of the intermediate

segment, and is acutely pointed. The stylocerite is remote from the latéral margin of

the segment, slightly exceeding the middle of the length of the segment and is very acutely

pointed. The médial border bears a small ventral spine at half its length. The upper

flagellum consists of five fused segments, with the shorter ïamus consisting of two segments

only in the male specimen.

The antenna has a robust basicerite without any latéral spine. The dorso-lateral

aspect bears a large rounded lamina that lits into the emargination of the anterior border

of the branchiostegite. The carpocerite is elongated and flattened and reaches to the base

ON SOME SPECIMENS OF PERICLIMENAEUS

1569

Fig. 8. — Periclimenaeus djiboutensis Bruce, anterior carapace, rostrum and antennal peduncles : A,

male ; B, female. — Larger ovigerous female : C, telson, latéral view ; D, telson, dorsal view ; E, uro-

pod.

of the disto-lateral spine of the scaphocerite. The lamina of the scaphocerite is about 2.6

times longer than wide in the female, of relatively uniform width with a bluntly rounded

anterior margin that is slightly exceeded hy the well developed acute disto-lateral spine.

A single detached first pereiopod, probably from the smaller ovigerous female is still

1570

A. J. BRUCE

Fig. 9. — Periclimenaeus djiboutensis Bruce, female : A, antennule ; B, antenna. — Male : C, first pereio-

pod ; D, chela of fîrst pereiopod ; E, Angers of chela of first pereiopod.

présent. The palm of the chela is subcylindrical, tapering slightly distally and about 3.7

times longer than the greatest width. The Angers are short, less than one tliird of the length

of the palm, and with numerous groups of coarse setae. The fîngers are subspatulate and

taper distally, ending in a gioup of acute interdigitating teeth. The teeth on the dacty-

lus are shorter and stouter and slightly curved compared to those on the fixed finger, which

are longer straighter and more slender. The margins of the opposing edges of the Angers

are entire. The carpus is moderately slender, nearly seven times longer than wide distally,

and equal to about 0.9 of the length of the merus, and 1.6 times the length of the chela.

Two second pereiopods are attributed to this species and appear to constitute a natural

ON SOME SPECIMENS OF PERICLIMENAEUS

1571

pair of appendages, being very similar in texture and colouration and were found in the

same tube. The major second pereiopod is robust with a subcylindrical, slightly flattened

and distally tapering palm. The palm is about 1.6 times longer than wide and is almost

completely devoid of tubercles and setae. The fîngers are equal to half the length of

the palm. The dactylus is compressed, feebly upcuived, and twice as long as wide. The

tip is broadly hooked and the anterior cutting edge is convex, with a well developed molar

process proximally. The fixed finger is relatively elongated and slender, exceeding the

tip of the dactylus. The tip is acute and not hooked. The cutting edge is entire with

a proximal fossa. The dorsal border bears a large blunt process proximally. The car-

pus is short and unarmed. The merus bears several acute tubercles along the ventral

border and a few small tubercles are also présent on the ischium.

The chela of the minor second pereiopod is much smaller than the major. The palm

is smooth, 0.6 of the length of the palm of the major chela and 1.5 times longer than wide,

tapering distally, with several long setae along tha ventral margin. The dactylus is about

0.4 of the length of the palm, compressed, with an evenly semi-circular outer border and

a strongly hooked bidentate tip. The cutting edge is entire and smoothly convex. The

fixed finger is short and broad, acute distally but without a hooked tip. Proximally a

distinct groove is présent, into which the cutting edge of the closed dactylus fîts. A small

blunt process is also présent proximally on the dorsal border. The carpus, merus and

ischium are similar to the major second pereiopod but less noticeablv tuberculate.

The uropods hâve the disto-lateral angle of the protopodite unarmed. The latéral

border of the exopodite is convex, without spines or teeth and ends in a strong acute distal

process with a large strongly hooked spine medially, which extends beyond the posterior

margin of the lamella. The endopod distinctly exceeds the exopod. The telson is rather

narrow, 2.5 times longer than wide anterioily and strongly concave ventrally. The laté¬

ral margins converge slightly posteriorly to a broadly rounded posterior margin. The

two pairs of dorsal spines are ail situated on the anterior fiftli of the telson length and arise

fronr the anterior part of a well marked longitudinal dorso-lateral groove that extends

along most of the length of the telson. The anterior spines arise more elosely to the midline

than the posterior spines, at about 0.08 of the telson length and the posterior spines at 0.15

of the length. The posterior spines are stout, tapering strongly distally, ecfual to about 0.15

of the telson length. The anterior spines are slightly shorter. Both pairs of spines are

slightly sinuous in latéral view. The latéral posterior spines are short and small and

situated slightly more dorsallv than the other spines. In the larger ovigerous female they

are also separated by a distinct gap hom the intermediate spines, which are stout and about

equal to 0.2 of the telson length. The submedian spines are similar to the intermediate

spines but slightly more slender and are without setules.

Remarks

The original description of Periclimenaeus djiboutensis was based upon three specimens

found in the collections of the Muséum, labelled “ Onycocaris sp., Djibouti ”, and without

any further indication of their origin. In his 1906 description of CoraUiocaris (O.) rhodope,

Nobili mentions that he had fourteen specimens from Djibouti. The présent material

under studv, clearly labelled as Nobii.i’s material, consists of twelve specimens of which

1572

A. J. BRUCE

three are P. djiboutensis. It seems highly probable that the three type speeimens of P. dji-

boutensis are also part of Nobili’s material that had become separated from his labelled

speeimens. However, this gives a total of fifteen speeimens from Djibouti, one more than

Nobili stated that he had.

The speeimens here reported upon, although incomplète, agréé closely with the original

description of the type speeimens, which also lacked several appendages. To augment

the original description, detail is provided of the second pereiopods of this species. The

minor second pereiopod described in the preliminary description is now considered to hâve

belonged to one of the P. arabicus speeimens. Unfortunately, the only ambulatory pereio¬

pod is no longer extant.

In the original description of this species, it was considered most closely related to

Periclimenaeus fimbriatus Borradaile. This relationship is no longer considered correct.

The type speeimens of P. fimbriatus hâve been reexamined and the latéral margin of the

exopod of the uropod is spinulate (fig. 15, B). P. djiboutensis now appears to occupy a

rather isolated systematic position in the genus Periclimenaeus but on the available infor¬

mation, appears to be most closely related to P. hebedactylus Bruce.

4. Periclimenaeus sp.

(Fig. 10)

M ATE RI AL EXAMINE!) : 1 çj, Djibouti, 1906, Coll. H. CoUTlÈRE.

Remarks

The single specimen is without second pereiopods and ambulatory pereiopods. The

disposition of the dorsal telson spines exclude P. djiboutensis from considération and the

form of the anterior margin of the scaphocerite appears to resemble that of P. rhodope

rather than P. arabicus. The first pereiopod is also more similar to P. rhodope. The ros-

trum shows small différences from the P. rhodope speeimens that prevent this specimen

being referred to that species with any certainty. The rostrum is more depressed and

the first dorsal tooth is situated in advance of the posterior orbital margin and its posterior

border is continuons with the dorsal midline of the carapace. The eight more anterior

teeth are longer and more slender and acute and reach their greatest development at, the

sixth or seventh teeth. The ventral margin is feebly concave and without teeth. The

supraorbital spine is small and acute.

The anterior margin of the lamina of scaphocerite is broadly rounded and the disto-

lateral spine is long and slender with some small subterminal setae laterally, but extending

beyond the lamina less than in P. rhodope speeimens. The first pereiopod clearly resembles

that of P. rhodope with the chela about 0.6 of the length of the carpus, but the chela is

rather stouter and the fingers are equal to half the length of the palm.

The postero-lateral angles of the sixth abdominal segment are acutely produced,

more markedly than in the speeimens of P. rhodope or P. arabicus. The telson is slightly

narrower than that of female P. rhodope or P. arabicus but is generally similar. The pos¬

terior dorsal telson spines are much smaller than the anterior spines, equal to about half

ON SOME SPECIMENS OF PERICLIMENAEUS

1573

Fig. 10. — Periclimena-eus sp., female : A, anterior carapace, rostrum and antennal peduncles ; B, sixth

abdominal segment ; C, telson ; D, scaphocerite ; E, uropod ; F, chela and carpus of first pereiopod ;

G, fingers of chela of major second pereiopod ; H, minor second pereiopod.

their length, and are situated at 0.45 and 0.08 of the telson length respectivelv. The uro-

pods closely resemble those of P. rhodope, with a single large disto-lateral spine and an

acute disto-lateral angle on the exopod.

A pair of second pereiopods are provisionally attributed to this specimen, as they

show slight différences from those of P. rhodope and P. arahicus. Tuberculation of the

chela is lacking but nuinerous long slender setae are présent on the dorso-ventral aspect

of the distal palm and fixed fînger and to a lesser extent on the dactylus. The fingers of

1574

A. J. BRUCE

the minor second pereiopod are mucli shorter than in P. arabicus and hâve only a single

acute tooth in the cutting edge. The proximal process on the dorsal aspect of the cutting

edge of the major chela is also much less well developed. The carpus is glabrous dorsally

in contrast to P. rhodope in which it is spinulate, and the cutting edges of the fîngers of

the minor second pereiopod are almost straight, witli distinctly hooked tips, rather than

concave, with feehly indicated tips.

5. Periclimenaeus hecate (Nobili)

(Fig. 11-12 ; 13, E)

Coralliocaris hecate, Nobili, 1904 : 232 ; 1906 : 58, pl. 3, fig. 2, —- Borradaile, 1917 : 385.

? Periclimenaeus tridentatus, Holthuis, 1952 : 140-146, figs 63-65 ( partim ).

Material examined : 1 1 ov. Ç, Djibouti, coll. H. Coutière.

Description

Nobili (1906) lias provided a description and illustration of the male specimen, which

is preserved in the attitude of his illustration. The second pereiopods are still attached

to the body together with most of the ambulatory pereiopods.

The rostrum of the. female is similar to that of the male but five slender dorsal teeth,

ail well in advance of the carapace are présent. In both, the ventral border is feebly convex

and without teeth. There is no trace of any supra-orhital spine or tubercle.

The first pereiopods présent no spécial features. The chela is slightly compressed, with

ON SOME SPECIMENS OF PERICLIMENAEUS

1575

fingers slightly less than half the length of the palm, and 0.6 of the carpus. The carpus

is very slightly longer than the merus.

The larger second pereiopod is smooth, subcylindrical, tapering slightly distally with

the fingers short and stout, equal to 0.4 of the palm length. The cutting edge of the dac-

Fig. 12. — Periclimenaeus hecate (Nobili), types : A, anterior carapace and rostrum, male ; B, female ;

C, telson, female ; D, male ; E, first pereiopod ; F, chela of first pereiopod ; G, fingers of minor second

pereiopod.

tylus bears a stout molar process opposing into a fossa on the fixed finger. The carpus,

merus and ischium présent no spécial features. The minor second pereiopod is small,

less than half the length of the major chela. The palm is smooth and subcylindrical and

three times the length of the fingers, which are elongated. The dactylus is 2.6 times longer

than wide, with the proximal two thirds of the outer rnargin subparallel to the cutting

edge, which is straight, finely dentate throughout its length with over 40 acute teeth that

diminish graduallv in size proximally. The tip of the dactylus bears a large blunt tooth

distally, discontinuous with the smaller sériés along the cutting edge. The fixed finger has

the cutting edge entire, with a small blunt tooth proximally and a small acute tooth at the

tip. The dactylus only barely exceeds the fixed finger.

The third pereiopod is stout, particularly the propod, which tapers distally and bears

a short simple dactylus with a distinct margin but without proximal or distal accessory

1576

A. .1. BRUCE

Fig. 13. — Periclimenaeus djiboutensis Bruce, sraall ovigerous female : A, terminal telson spines ; B, disto-

lateral angle of exopod of uropod. Large ovigerous lemale : C, terminal telson spines ; D, dorsal telson

spines. — Periclimenaeus hecate , type : E, dactylus of third pereiopod. — Periclimenaeus tiobilii sp.

nov., holotype : F, dactylus of third pereiopod.

spines. The distal end of the propod bears médial and latéral terminal spines and a ventral

preterminal spine. The dactyls are similar on ail pereiopods and in both sexes.

The caudal fan présents no unusual features. The dorsal telson spines are small and

situated submarginally at thirds of the telson length.

Remarks

Periclimenaeus hecate was included in the synonymy of P. tridentatus (Miers) hy Holt-

huis (1952). The holotype of P. tridentatus is preserved in the collection of the British

Muséum (Natural History) and although now in a poor state of préservation, the specinten

still has some of the ambulatory pereiopods. These show not only a distinct distal acces-

sory spine on the dactyl but also an acutely pointed proximal ventral spine, which lies

between the inner and outer distal ventral spines of the propod (fîg. 15, C). On account

of the absence of these spines from the dactyls of the ambulatory pereiopods, P. hecate

(Nobili) is considered a distinct and valid species.

The specimens referred to P. tridentatus (Miers) in the material collected by the Siboga

Expédition, appear to consist partly of specimens of P. hecate or a closely related species,

as the characteristic dactyls of the ambulatory pereiopods are illustrated. However, the

ON SOME SPECIMENS OF PERICLIMENAEUS

1577

minor second pereiopods appear to be of two types and it seems possible that two sepa-

rate species are présent.

Also svnonymized with P. tridentatus (Miers) were P. crassipes (Caïman) and P. qua-

dridentatus (Rathbun). In the holotype specimens of these species also, the acute proxi¬

mal tooth on the body of the dactylus of the ambulatory pereiopods is lacking and these

are therefore also considered to be distinct from Miers’ species. P. quadridentatus has

well developed distal accessory spines on the dactyls of the ambulatory pereiopods and is

therefore considered to be distinct from P. crassipes (fig. 15, D).

The type specimens of P. crassipes (Caïman) need to be re-examined to détermine whe-

ther the species is a synonym of P. hecate (Nobili) or not. In almost ail features the two

species show a very close resemblance. The most striking feature of P. crassipes is the

swollen appearence of the propod and carpus of the third pereiopod (fig. 15, D). The

propod is about three times longer than the greatest width, which is close to the proximal

end, and tapers strongly distally where it is about one quarter of the proximal width. In

the minor second pereiopod of P. crassipes the fingers are longer and more slender than

in P. hecate, with large blunt feebly curved tips. The dactylus extends well beyond the

tip of the fixed finger and the cutting edge is straight with a sériés of low blunt teeth. It is

provisionally concluded that P. crassipes (Caïman) is distinct from P. hecate (Nobili).

In P. quadridentatus (Rathbun) the holotype lacks the minor second pereiopod. The

major second pereiopod is remarkably stout with a dactylus bearing a well developed

rnolar process proximally and with the distal half of the cutting edge strongly thickened

and rounded. The carpus has the distal dorsal margin produced into a distinct lobe that

projects forwards over the proximal part of the palm. This feature has not been noticed

in any other species of Periclimenaeus. In comparison with P. crassipes, and also P. hecate,

the propod of the third pereiopod is slender and not swollen proximally. The dactyls

of the ambulatory pereiopods are without an acute proximal spine but possess well deve¬

loped, robust, blunt distal accessory spines. It is concluded that P. quadridentatus (Rath¬

bun) is also a valid and distinct species.

Ralss (1921) referred a single specimen from Cape Jaubert, Western Austraha to ?

Coralliocaris hecate. The dactyls of the ambulatory pereiopods are not described and the

specimen needs to be re-examined in order to confirme the identification.

6. Periclimenaeus nobilii sp. nov.

Material examined : 1 Ç, coll. Dr. Jousseaume, Red Sea, 1897.

Description

Closely related to P. hecate and P. tridentatus. The rostrum slightly exceeds the proxi¬

mal segment of the antennular peduncle and is acute, slender and depressed, with two

acute dorsal teeth and a convex toothless ventral margin. Supra-orbital spines or tubercles

are lacking. The antennal spine is well developed, slender and acute. The orbit is feebly

developed and the inferior orbital angle is obsolescent. The antero-lateral angle of the

carapace is slightly produced. The sixth abdominal segment is about 1.5 times longer

1578

A. J. BRUCE

Fie. 14. — Periclimenaeus nobilii sp. nov., holotype : A, carapace, antennae and first pereiopod ; B,

chela of major second pereiopod ; C, cliela of minor second pereiopod ; D, fîngers of chela of minor

second pereiopod ; E, propod and dactylus of third pereiopod ; F, telson.

than deep. The pleura of the first five segments are rounded. The posterior ventral

angle of the sixth segment is bluntly pointed and the posterior latéral angle is rounded.

The telson is broad anteriorly, tapering, about twice as long as wide, with two pairs of

slender submarginal dorsal spines situated at 0.35 and 0.62 of the telson length. The

posterior dorsal spines are slightly longer than the anterior pair and are subequal to the

ON SOME SPECIMENS OF PERICLIMENAEUS

1579

latéral posterior spines. The intermediate spines are stout, equal to 0.23 of the telson

length. The submedian spines are almost as long and stout as the intermediate spines

but are more slender distally and are non-setose.

The proximal segment of the antennular peduncle is narrow with a very small disto-

lateral process. The stylocerite is short and acute and the latéral border is feebly concave.

The intermediate and distal segments are short and stout The basicerite is without a

dorsal flange and is unarmed. The carpocerite is slender and exceeds the anterior mar-

gin of the scaphocerite, which is broadly rounded and extends well beyond the small disto-

lateral spine. The eyes are short and stout, almost globular, with an obliquely hemispheri-

cal cornea of slightly smaller width than the stalk.

The first pereiopod is moderately robust, and exceeds the carpocerite by the length

of the carpus and chela. The chela is stout with thick Angers that are feebly subspatulate

and slightly longer than the palm. The chela is 0.78 of the length of the carpus, which

is 4.5 times longer than wide, and increases in size distally. The carpus is 0.91 of the length

of the merus.

The second pereiopods are dissimilar and very unequal, sinooth. The chela of the

major second pereiopod is about three times the post-orbital carapace length. The palm

is stout, subcylindrical, slightly flattened and tapering distally, about 2.3 times longer

than wide and with a few short simple setae only. The fingers are moderately dorsally

curved. The dactylus is 0.37 of the length of the palm and is about 2.5 times longer than

wide. A stout hooked tip is présent and the anterior cutting edge is thickened. A well

developed, distinct molar process is présent posteriorly. The fîxed fînger lias a smaller

hooked tip présent distally with a large fossa proximally. The dorsal margin of the fossa

bears a small blunt triangular process and a blunt lobe is présent ventrally. The carpus

is relatively small and feeble, unarmed and without an anterior dorsal lobe. The ischium

is twice as long as wide, tapers gradually distally and is without ventral spines or tubercles.

The ischium is about 0.6 of the meral length and is also ventrally unarmed.

The chela of the minoi second pereiopod is 0.43 of the length of the major second pereio¬

pod. The palm is subcylindrical and about 2.6 times longer than wide. The ventral médian

aspect bears numerous long simple setae. The fingers are about 0.4 of the length of the

palm. The dactylus is compressed, about 2.6 times longer than wide. The cutting edge

is sinuous with about 26 small acute teeth on the anterior two thirds, which increase in

size distally and blend without interruption with the small acute hooked tip. The proxi¬

mal end of the cutting edge also bears a small isolated blunt tootli. The fîxed fînger is

broad, tapering distally to an acute hooked tip. The cutting edge is straight and entire,

with a distinct tooth proximally.

Only two ambulatory pereiopods are preserved. The third pereiopod is stout with a

small dactylus. The dactylus is compressed, broad at the base and stronglv tapering

to a distinct acute unguis. There is no distal accessory spine on the body of the unguis

but a well developed acute spine is présent proximally on the ventral border, where it

lies between the two distal ventral spines of the propod. The propod is robust, three times

longer than wide and tapering distally. The disto-ventral end bears a pair of stout spines

and the rest of the ventral border is devoid of spines. The carpus is 0.7 of the length of

the propod, stout, broadest distally, twice as wide as long. The merus is about 1.1 times

the length of the propod, uniform in width and unarmed. The (?) fifth pereiopod is simi-

1580

A. J. BRUCE

lar, but with more numerous setae along the distal ventral border of the propod. The

proximal accessory spine on the dactylus is smaller and more acute than on the third pereio-

pod.

The uropods présent no spécial features. The protopodite is blunt disto-laterally.

The latéral margin of the exopod is entire and terminâtes in a small acute tooth with a

mobile spine medially.

Type : The single female specimen is designated as the holotype and is deposited in

the collections of the Muséum national d’Histoire naturelle.

Remarks

The specimen was examined by Nobili in 1905 and tentatively identified as “ ? anoma¬

lie. 15. — Periclimenaeus arabicas (Caïman), holotype : A, dactyl of third pereiopod. —■ Periclimenaeus

fimbriatus Borradaile, type : B, exopod of uropod. — Periclimenaeus tridentatus (Miers), holotype :

C, dactyl of third pereiopod. — Periclimenaeus quadridentatus (Rathbun), holotype : D, dactyl of

third pereiopod. — Periclimenaeus crassipes (Caïman), type : E, third pereiopod.

ON SOME SPECIMENS OF PERICLIMENAEUS

1581

lie de Coralliocaris hecate There does not appear to hâve been any published record

of this specimen. As with many of the shrimps collected by M. Jousseaume, the précisé

locality in the Red Sea was not recorded.

This species is most closely related to P. tridentatus (Miers). These two species are

the only two species of the genus that possess a proximal accessory spine on the dactyls

of the ambulatory pereiopods. The two species may be separated by the absence of a

distal accessory spine in P. nobilii and its presence in P. tridentatus.

7. Periclimenaeus arabicus (Caïman)

Coralliocai is ( Onycocaris ) rhodope, Nobifi, 1906 : 49.

Material examined : 1 Ç, Côtes d’Arabie, Stn. XLVII, dragages banc d’huîtres per¬

lières entre 25°10' N., 24°55' N. et 55°10' E., 24°40' E., Mission J. Bonnier & Ch. Pérf.z,

1907.

Remarks

The single specimen is macerated and lacks both second pereiopods. The rostrum

bas six dorsal and one ventral tooth. The supraorbital spines are distinct. The scapho-

cerite has the lamina subrectangular distally. Both first pereiopods are attached and ail

the ambulatory pereiopods are présent on the right side of the body. The dactyls and

the spinulation of the other segments correspond closely to the specimens described above.

The telson and uropods are also in agreernent.

DISCUSSION

It is now known that most of the Indo-West-Pacifîc species of the genus Periclime¬

naeus are associated with sponges and that a few are also found in association with compound

tunicates. In the case of the specimens reported upon above, none of the hosts were recor¬

ded. In order to clarify the relationships of the various species further intact specimens,

preferably from carefully identified hosts, are urgently needed. At présent it appears

that P. rhodope and P. arabicus are sponge associâtes but that P. tridentatus and its relatives

may be associated with tunicates. In the literature there are a number of records of

Periclimenaeus species being found in association with corals. There appear to be no

carefully authenticated instances of such an association and these records hâve most pro-

bably been derived from specimens that hâve been displaced from small sponges that com-

monly encrust the base of coral colonies or occur between their branches. Small tunicate

colonies can also be found in these situations.

Although a wide variety of sponge hosts appear to be involved in the Periclimenaeus

association, as yet few generic or spécifie host identifications are available. The range of

variations exhibited by the dactyls of the ambulatory pereiopods indicates a high degree

1582

A. J. BRUCE

of adaptation to the host and represents characters of considérable taxonomie value in this

genus. In general, these morphological characteristics are best developed on tbe dacty-

lus ol the third pereiopod and shows some réduction of the characteristic features on the

fourth and fîfth pereiopods. Other features that are of partieular importance in assessing

the relationships of the various species that are probably direct adaptations to the hosts,

are the fingers of the minor second pereiopod and the chela of the first pereiopods. Other

characters, such as the rostrum, supraorbital spines or tubercles, the scaphocerite and the

major second pereiopod are probably less concerned in adaptation to the host. As yet,

the range of variations in many of these characters is unknown, as many of the species are

still only known from one or two specimens, which are frequently of different sexes as

Periclimenaeus species are usually found in pairs. The second pereiopods are also readily

autotomized and incomplète specimens often bave to he identifîed mainly on the basis

of the dactyls of the ambulatory pereiopods.

The type specimens of P. djiboutensis were found in the collections of the Muséum

without any indication that they were part of the Djibouti material reported upon by

Nobili. In his report Nobili mentions that he has fourteen specimens of Coralliocaris

( Onycocaris ) rhodope. The material upon which this report is based, included twelve spe¬

cimens under that name. There seems no reason to doubt that the three type specimens

of P. djiboutensis were also part of Nobili’s material. This gives a total of fifteen instead

of fourteen. Although the origin of the discrepancy is uncertain, it is possible that Nobili

had an additional specimen that was too incomplète to report upon satisfactorily. In the

material at présent available the specimen referred to as Periclimenaeus sp. seems the most

likely specimen as it is the only one that is represented hy a single specimen. The présent

identifications of Nobili’s fifteen specimens may be summarized as follows :

P. rhodope (Nobili) 2 <$, 2 ov. $.

P. arabicus (Caïman) 2 2 ov. $.

P. djiboutensis Bruce lçj, 2 ov. Ç (présent report).

P. djiboutensis Bruce 2 1 ov. Ç (Bruce, 1970).

Periclimenaeus sp. 1 <$.

To facilitate the séparation of Periclimenaeus tridentatus (Miers) and related species,

the provisional key below is provided. The information concerning several of the species

is very limited. Four of the species are known from the holotype specimens only and the

other two species are also known only from the two original specimens.

A provisional kf.y to the identification of P. tridentatus (Miers)

AND RELATED SPECIES

1. Dactyls of ambulatory pereiopods without accessory spines. 2

— Dactyls of ambulatory pereiopods with accessory spines. 3

2. Dactvl of minor second pereiopod much longer than fixed finger, with about ten blunt teeth

distally ; carpus and propod of first pereiopod greatly swollen, propod three times broader

proximally than distally. P. crassipes (Caïman)

— Dactyl of minor second pereiopod subequal to fixed finger with about 40 acute teeth along

the whole length of culting edge ; carpus and propod of third pereiopod less stout and not

greatly swollen. P. hecate (Nobili)

ON SOME SPECIMENS OF PERICLIMENAEUS

1583

3. Dactyls of ambulatory pereiopods without an acute accessory spine proximally. 4

— Dactyls of ambulatory pereiopods with an acute accessory spine proximally. 5

4. Rostrum with four acute distal teeth only ; supraorbital tubercles absent, carpus of second

major pereiopod with a lobvdar process d'sto-dorsally. P. quadridentatus (Rathbun)

— Rostrum with digitiform teeth ; supraorbital tubercles présent ; carpus of second major pereio¬

pod without a lobular disto-dorsal process. P. pachydentatus Bruce

5. Body of dactylus of ambulatory pereiopods with a distinct distal accessory spine.

P. tridentatus (Miers)

— Body of dactylus of ambulatory pereiopods without a distal accessory spine.

P. nobilii sp. nov.

Acknowledgements

1 am particularly grateful to Dr J. Forf.st for the opportunity to examine and report upon

these specimens and also to Dr A. L. Rice, Dr R. W. Ingle, Dr C. B. Goodhart, and Dr F. A.

Chace jr, for access to and information upon other Periclimenaeus specimens in tlieir care.

LITERATURE CITED

Balss, H., 1921. — Stomatopoda, Macrura, Paguridea und Galatheidea. Results of Dr. E. Mjô-

bergs Swedish Scientific Expédition to Australia 1910-13. XXIX. K. svenska Vetensk.-

Akad. Handl. , 61 (10) : 1-24, fig. 1-12.

Bruce, A. J., 1970. — Further preliminary descriptions of new species of the genus Periclimenaeus

Borradaile, 1915 (Crustacea, Decapoda Natantia, Pontoniinae). Zoôl Meded., Leiden,

44 (21) : 305-315.

Calman, W. T., 1939. — Crustacea : Caridea. Scient. Rep. John Murray Exped., 6 : 183-224, figs 1-8.

Holthuis, L. B., 1952. —• The Decapoda of the Siboga Expédition. Part XI. The Palaemonidae

collected by the Siboga and Snellius Expéditions with remarks on other species. II. Sub-

familv Pontoniinae. Siboga Exped., 39 a 10 : 1-252, fig. 1-110, tab. 1.

Nobili, G., 1904. — Diagnoses préliminaires de vingt-huit espèces de Stomatopodes et Décapodes

Macroures de la Mer Rouge. Bull. Mus. nain. Hist. nat. Paris, l re sér., 10 : 228-238.

— 1906. — Crustacés Décapodes et Stomatopodes, Mission J. Bonnier et Ch. Pérez (Golfe

Persique, 1901). Bull, scient. Fr. Belg., 40 : 13-159, fig. 1-3, pl. 2-7.

— 1906. — Faune carcinologique de la Mer Rouge. Décapodes et Stomatopodes. Annls Sci.

nat., Zool., 4 : 1-347, fig. 1-12, pl. 1-11.

Rathbun, M. .1., 1906. — The Brachyura and Macrura of the Hawaiian Islands, Bull. U. S. Fish.

Commn, 23 : 827-930, fig. 1-79, pl. 3-24.

Manuscrit déposé le 10 octobre 1973.

Bull. Mus. natn. Hist. nat., Paris, 3 e sér., n° 258, sept.-oct. 1974,

Zoologie 180 : 1557-1583.

Achevé d’imprimer le 30 avril 1975.

IMPRIMERIE NATIONALE

4 564 004 5

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gnés d’un résumé en une ou plusieurs langues. L’adresse du Laboratoire dans lequel le

travail a été effectué figurera sur la première page, en note infrapaginale.

Le texte doit être dactylographié à double interligne, avec une marge suffisante, recto

seulement. Pas de mots en majuscules, pas de soulignages (à l’exception des noms de genres

et d’espèces soulignés d’un trait).

Il convient de numéroter les tableaux et de leur donner un titre ; les tableaux

compliqués devront être préparés de façon à pouvoir être clichés comme une figure.

Les références bibliographiques apparaîtront selon les modèles suivants :

Bauciiot, M.-L., J. Daget, J.-C. Hureau et Th. Monod, 1970. — Le problème des

« auteurs secondaires » en taxionomie. Bull. Mus. Hist. nat., Paris, 2 e sér., 42 (2) : 301-304.

Tinbergen, N., 1952. — The study of instinct. Oxford, Clarendon Press, 228 p.

Les dessins et cartes doivent être faits sur bristol blanc ou calque, à l’encre de chine.

Envoyer les originaux. Les photographies seront le plus nettes possible, sur papier brillant,

et normalement contrastées. L’emplacement des figures sera indiqué dans la marge et les

légendes seront regroupées à la fin du texte, sur un feuillet séparé.

Un auteur ne pourra publier plus de 100 pages imprimées par an dans le Bulletin,

en une ou plusieurs fois.

Une seule épreuve sera envoyée à l’auteur qui devra la retourner dans les quatre jours

au Secrétariat, avec son manuscrit. Les « corrections d’auteurs » (modifications ou addi¬

tions de texte) trop nombreuses, et non justifiées par une information de dernière heure,

pourront être facturées aux auteurs.

Ceux-ci recevront gratuitement 50 exemplaires imprimés de leur travail. Ils pourront

obtenir à leur frais des fascicules supplémentaires en s’adressant à la Bibliothèque cen¬

trale du Muséum : 38, rue Geoffroy-Saint-Hilaire, 75005 Paris.