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TRANSACTIONS 


OF 


THE ZOOLOGICAL SOCIETY 


OF LONDON. 


e 


Vou. XIV.—Parr 1. 


LONDON: 


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TRANSACTIONS 


OF 


THE ZOOLOGICAL SOCIETY 


OF LONDON. 


VOLUME XIV. 


LONDON: 


PRINTED FOR THE SOCIETY: 
SOLD AT THEIR HOUSE IN HANOVER-SQUARE; 
AND BY MESSRS, LONGMANS, GREEN, AND CO., PATERNOSTER-ROW. 


1898, 


PRINTED BY TAYLOR AND FRANCIS, 


R&D LION COURT, PLEET STREKT. 


CONTENTS. 


On a Nothosaurian Reptile from. the Trias of Lombardy, apparently referable to 
Lariosaurus. by G. A. Boutenaer, /.R.S. (Platel.) . . . . . page | 


TI. On the Lepidosiren of Paraguay, and on the external characters of Lepidosiren and 7 
Protopterus. By KE. Ray Lanxesrer, W.A., F.RS., F.Z.S8., Linacre Professor 
of Comparative Anatomy in the University of Oujord. “(Platerle aero aeace ltl 


Til. On a Collection of Fishes from the Rio eee By G. A. Boutenaer, 
HBS. 4(Plates: IT—VITL) ae 4y.: 4. ahs. eee ints py Peay Ve Aces 


IV. Contributions to the Comparative Anatomy and Histology of the Suprarenal Capsules. 
—The Suprarenal Bodies in Fishes, and their Relation to the so-called Head- 
Kidney. By Swave Vincent, M.B. Lond., Demonstrator of Physiology and 
Assistant Lecturer on Iistology, Mason College, Birmingham. (Plates IX.- 
PROP E \iis stecie Se ast din ho. Mc / exc ad Seah spectre nisi Se | 


V. A Supplementary Report on the Crustacea of the Group Myodocopa obtained during 
the ‘ Challenger’ Expedition, with Notes on other new or imperfectly-known 
Species. By G. Srewarpson Brapy, WD, LL.D, F.RS. (Plates XV. 
PROVMUIE a2) Sos ee el ek eS ee ee 


VI. A Revision of the Oriental Hesperiide. By H. J. Euwes, ARS, VLS., EZS., 
and James Epwarps, #.#.8. (Plates XVIJIL-—XXVII.) ... . . . 101 


VIL. On the Morphology of the Skull in the Paraguayan Lepidosiren and in other 
Dipnoids. By Professor T. W. Bripvex, Se.D., F.ZS., Mason College, 
buminghame (Plates XXXVI & XXIX.) .. © 5 om. | ws 2). 325 


VIII. On the Mammals obtained by Mr. John Whitehead during his recent Expedition 
to the Philippines. By Ouvrieip Tuomas. With Field-notes by the Collector. 
(IIS NO NONE INGNONONED ep oi gee een a ew 


lv CONTENTS. 


IX. On the Lepidosiren of the Amazons ; being Notes on five Specimens obtained between 
1895-97, and Remarks upon an Example living in the Paré Museum. By 
Dr. Emm A. Gorn, C.M.Z.S., Director of the Pard Museum. (Plates 
EN VT he OR VES os > eee ee ie ‘ . a ose ee pape ais 


X. On a Collection of Fishes from the Rio Jurua, Brazil. By G. A. BouLENGEr, 
ERS, FOS. “(Plates ROX RIM) A te ee eee 


XI. On new or imperfectly-known Species of Ostracoda, chiefly from New Zealand. By 
G. Stewarpson Brapy, M.D., LL.D., D.Sc., F.R.S. (Plates XLIII.--XLVII.) 429 


List of the Papers contained in (Vols MV... 5. foe 5 os Sees 
Index. of Species; icici ice Sap ep. LS Wado w Se eh, ye ee 


Tea SA CrlT ON s 


OF 


PEE ZOO Oe CAs  SOOCREWR ¥ 
OT LON DON: 


I. On a Nothosaurian Reptile from the Trias of Lombardy, apparently referable to 
Lariosaurus. By G. A. Boutencer, F.R.S. 


Received October 14th, 1893, read November 7th, 1893. 


[Puate I.] 


THE Directors of the Senckenberg Museum, Frankfort-on-Main, have, at the 
suggestion of my friend Prof. O. Boettger, entrusted to me for description a very 
valuable paleontological specimen, a slab from the Upper Trias of Perledo, near 
Varenna, on the Lake of Como, Lombardy, with the nearly complete skeleton of a 
small Nothosaurian ; the counter-slab is also preserved, showing the impression of the 
skeleton. ‘The fossil was presented to the Senckenberg Museum by the late Dr. Riippell 
in 1850, under the name of Macromerosaurus plinii, Curioni. It appears from Curioni’s 
paper (8, p. 166) that, prior to 1847, Riippell had examined some of the Reptilian 
remains from those beds, then regarded as Liassic ; one of these specimens he acquired, 
together with Fish-remains1, and subsequently placed in the Museum at Frankfort, 
where it has remained ever since, apparently ignored of all who have written on this 
interesting group of early Plesiosaurians. 

Some confusion has arisen with regard to the specimen for which the name Wacro- 
merosaurus plinii was proposed by Curioni. On referring to his first paper (8) 
published in 1847, it appears as if the name (p. 161) was intended for the Perledo 
Saurian, of which he gives a detailed description and figure; whilst the name Lario- 
saurus balsami applies to the much larger specimen previously noticed by Balsamo- 


1 Described by Deecke, Paleontogr. ly. 1889, p. 110. 


2 MR. G. A. BOULENGER ON A NOTHOSAURIAN REPTILE 


Crivelli (7) as Paleosaurus. But in 1863 Curioni explains (12, p. 2661) that Macro- 
merosaurus (originally misspelt Macromirosaurus) was the name intended for another 
Saurian, from Perledo and Viggiti, with longer humerus, to which he merely alludes in 
his first paper, and which is, in fact, the Pachyplewra edwardsii of Cornalia (g). If 
we could accept this interpretation, the name Macromerosaurus might be used in 
preference to that of Pachypleura, which is preoccupied in zoology, as pointed out by 
Lydekker (21); but I do not find any justification for such a transposition, and, 
following tradition, consider Macromerosaurus as a synonym of Lariosaurus, the former 
name applying to the young, the latter to the adult. 

The Reptile lies on its back, the neck and tail twisted to the right, the hind limbs 
spread out nearly at right angles to the body. The head is intact, with the palate and 
mandible completely exposed. The 15 anterior cervical vertebre are in natural juxta- 
position, whilst the remainder are dislocated. The pectoral arch is likewise dislocated, 
and does not show all the elements. The fore limb is represented by the right 
humerus, and the perfect left propodials and manus. The dorsal vertebre are 
partly imbedded in the matrix, partly hidden under the plastron, most of the bones of 
which are very well preserved. The pelvis is crushed, but the tail is complete and 
exposed to its very tip. The Plate (Trans. Zool. Soc. XIV. Pl. I.) represents the fossil 
natural size; the matrix is represented lighter than it really is, in order to render the 
outlines of the bones more distinct. 

The specimen is now in a very different condition from what it was when sent to 
me. The manus was a complete fraud, imaginary phalanges having been sculptured 
out of the matrix in order, evidently, to give the Reptile a more Plesiosaurian appear- 
ance ; and, deceived at first, I considered the fossil as representing a new type. I must 
add that in all probability the specimens figured by Curioni were developed by the 
same “artist”; and that, consequently, some of the conclusions which have been 
derived from an examination of those specimens, especially as regards the digital 
formula, the shape of the “ claws” or “ hoofs,” and the aspect of the palate are, to say 
the least, very doubtful. The specimen figured by Zittel (20) as Zariosaurus balsami, 
and of which a cast is preserved in the British Museum, is imperfect as regards the 
digits, and the exact number of phalanges is difficult to make out. This explains the 
contradiction between text and figure in Zittel’s Manual, the number of phalanges in 


1 This passage of his second paper is here reproduced :—“ Da una nota che il Cornalia aggiunse alla sua 
Memoria pare che egli ritenga che i rettili di Besano e quelli di Perledo, uno dei quali yenne da me descritto 
e figurato col nome di Lariosaurus balsami, Cur. (vedi Memoria citata, Milano, 1847), spettino ad una sola 
specie di saurii, e debbano riferirsi tutti al suo Pachypleura. Nel 1847 io pensaya che due fossero le specie di 
Perledo; una il Lariosauro, affato speciale a Perledo, e un’ altra di cui trovayansi esemplari anche a Viggit, 
meglio conservati di quelli di Perledo, che io credeva non dover riferire alla medesima specie. Applicaia questa 
seconda specie il nome di Macromerosuwro, giudicando che il carattere pil sagliente di questa specie fosse la 
lunghezza degli omeri rettilinei, eccedente di una terza parte quella dei femori: il che non é nel Lariosauro 
(vedi fig. 2, tay. vii.).” The figure refers to the Reptile with straight and longer humerus, the Pachypleura 
of Cornalia. 


FROM THE TRIAS OF LOMBARDY. 3 


the manus being given as 2, 4, 4, 4, 2, which appears wrong @ priori, whilst the figure 
shows 2, 3, 4, 4, 2, which is likewise probably erroneous as regards the outer finger. 
Curioni, on the other hand, describes and figures the number of phalanges as 2, 3, 4, 5, 3, 
in both manus and pes. In the face of such contradictions, I think it safest to refer 
the specimen provisionally to Lariosawrus balsami, and, contenting myself with 
describing and figuring the specimen entrusted to me, leave it to those who have access 
to the typical specimens to decide whether the determination is correct or not. I 
beg to thank Dr. H. Woodward, F.R.S., for kindly allowing the specimen to be further 
developed in the British Museum, and Mr. Barlow for the painstaking and skilful 
manner in which he has acquitted himself of the task entrusted to him. 

The skull, which displays the palatal aspect with the mandible in situ, and the tips 


Fig. 1. 


Lower view of skull and mandible. 


ang. Angular. p. Palatine. s.ang. Supra-angular. 
ar. Articular, pt. Pterygoid. spl. Splenial. 
d. Dentary. q. Quadrate. v. Vomer. 


ept. KEctopterygoid. 


of a few anterior teeth, is elongate-acuminate, its length once and two thirds its 
greatest width at the mandibular articulation. The structure of the palate is better 
shown than in any of the small Nothosaurians with which we are at present acquainted, 
yet some of the sutures are not to be traced as distinctly as might be desired. I trust 


the annexed diagrammatic restoration (fig. 1) will ultimately prove tolerably correct. 
B2 


4 MR. G. A. BOULENGER ON A NOTHOSAURIAN REPTILE 


Among the points on which there can be no doubt may be mentioned the very anterior 
opening of the choanz, separated by a narrow septum formed by the vomers; the 
presence of large oval suborbital vacuities, such as are known to exist in Newstico- 
saurus, and as are shown on Curioni’s figure; and the backward extension and union 
on the median line of the pterygoids, as far back as the basioccipital, which they cover 
over, as well as the basisphenoid, in this respect agreeing with most Nothosaurians. 
These pterygoids are produced and narrowed anteriorly, where they join the vomers, 
whilst posteriorly they form two raised ridges on each side, between which the quadrate 
is wedged in. Before the skull had been thoroughly cleared from the matrix, the 
outer of these ridges conveyed to my mind the remains of ossified hyoid cornua (cerato- 
branchials). But, in the course of development, these ridges were found to bear a 
series of minute subconical teeth. This discovery is a very startling one, for, unless 
the Placodonts belong to the Plesiosauria, no member of this order was yet known 
to have possessed teeth on any of the bones of the palate. It is true that the 
following remark is made by Prof. Seeley (25, p. 591) respecting Mesosaurus tenut- 
dens :—“ Two elevated ridges, nearly parallel to each other and close together, extend 
along its [the palate’s] length, and converge backward. There is a possibility that 
these ridges carried single rows of teeth like the teeth on the ridges on the palate of 
Pareiasaurus, as the impression from the cast shows at regular intervals a few white 
dots along each ridge.” 

The bone between the suborbital and subtemporal fosse I take to be the ecto- 
pterygoid, known to be largely developed in Simosaurus, but absent from the restora- 
tions of Nothosawrus. Hence, the palatines would be small, and relegated to between 
the choane and the suborbital fosse. 

The mandible has a short symphysis, with median suture, and projects posteriorly 
considerably beyond its articulation with the cranium; on this process the angular, 
supra-angular, and articular bones are perfectly distinguishable. 

The neck consists of 21 vertebree. Its length is once and a half that of the skull, 
and nearly equals that of the body. The vertebre are short, the centra being a little 
broader than long; they bear short ribs, which are fork-headed, as is clearly shown on 
the counter-plate which bears the right rib of the second vertebra, and further by a 
detached rib lying on the right side of the eighteenth vertebra (fig. 2). The dislocated 
seventeenth vertebra shows the posterior articular surface moderately cupped. The 
atlas shows the proatlanto-atlantic hypapophysis separating the neuroids, and followed 
by the centrum, behind which there is no hypapophysis. 

Little is to be seen of the dorsal vertebral column and ribs, obscured as they are by 
the overlying pectoral arch and plastron; the number of vertebre may be computed at 
about 20. 

The tail is long and slender, tapering to a fine point; its length is nearly three times 
that of the skull; it comprises 42 or 43 vertebre. 


FROM THE TRIAS OF LOMBARDY, 5 


Immediately behind the pectoral arch begins the series of small bones which form 
the plastron; this does not differ from that of the other Nothosaurians the plastron of 
which is known, 34 transverse series of bones can be counted between the pectoral 
and pelvic arches, each series consisting of a median angulate piece and two straight 
ones on each side (fig. 3), as in Plesiosaurs 1, which reptiles lack, however, the angular 
bend of the median piece. 


Fig. 2. Fig. 3. 


cat 


Posterior cervical rib. Anterior plastral bones. 


It is to be regretted that the dislocation of the pectoral arch renders the recognition 
of its elements somewhat difficult, There is one bone as to the nature of which no 
doubt can be entertained: that is, the right coracoid, lying on the inner side of the 
right humerus, which agrees essentially with the same element in Neusticosaurus. It 
is nearly equally expanded at both extremities, its width somewhat exceeding half its 
length, which nearly equals that of the humerus ; its inner border is shorter and more 
concave than the outer. Two large, flattened, curved bones anterior to the coracoid I 
take to be clavicles, the left of which appears to cover the left coracoid, and is separated 
from the other coracoid by a small bone with concave surface, which is probably the 
proximal (acromial) portion of the scapula, ‘Two displaced anterior dorsal vertebrae 
and a rib are exposed between the coracoids. 

Only the right humerus is completely exposed, having slipped out of the glenoid 
cavity. It is a short, stout, curved bone with strong preaxial crest ; it is provided with 
a small, elliptic, entepicondylar (ulnar) foramen; its length is exactly half that of the 
skull. The distal extremity of the left humerus projects from below the left clavicle, 
showing distinctly the epicondylar foramen. For the other bones of the fore limb we 
must turn to the left side of the specimen, where the whole forearm and manus are 
exposed in natural juxtaposition, the radius overlapping the proximal extremity of the 
ulna, The radius is large and much flattened, with concave inner border, its length 
twice its distal width and half the length of the humerus; the ulna is much more 


slender. The carpal bones are seven in number: a large intermedium, a radiale, and 


1 Mr. C. W. Andrews has recently pointed out (Geol. Mag. 1895, p, 243) that some Plesiosaurs have three 


paired bones to each row. [June, 1895.] 


6 MR. G. A. BOULENGER ON A NOTHOSAURIAN REPTILE 


a small bone to each digit. The digits are five, increasing in length to the third, 
which equals the fourth, the fifth intermediate in length between the first and second. 
The first metacarpal is as long as the fifth but more slender; the second, third, and 
fourth are equal, half as long as the radius; the first finger has two phalanges, the 
second three, the third and fourth have four, the fifth three; the distal phalanx is 
obtusely pointed, flattened, and did not support a claw. The phalangeal bones are 
devoid of condyles, 

The pelvic bones are somewhat crushed and displaced. The left pubis is probably 
hidden under the ischium, but the right is exposed, and does not show a notch in its 
posterior border. The ischia form large plates with radiating strie. 

The femur is straight, subcylindrical, but feebly expanded at the extremities; its 
length exceeds by one fourth that of the humerus. The tibia isa little longer and 
considerably broader than the fibula, and not quite half as long as the femur. In the 
tarsus, a large intermedium and a smaller fibulare are present; only two very small 
bones, at the base of the third and fourth metatarsals, are distinguishable in the second 
row. ‘The metatarsals are more elongate than the metacarpals, and the third and 
fourth the longest and equal. The phalanges of the pes are stronger than those of the 
manus, and their formula is different, viz. 2, 3, 4,5, 4; the toes increase gradually in 
length from the first to the fourth ; the fifth is nearly as long as the third. The pes 
exceeds the manus by the length of the two distal phalanges of the fourth toe. 

It seems probable, from the structure of the extremities, that this Saurian was better 
adapted for locomotion in the water than on land, and the digits must have been 
connected by a broad web. 

On comparing the hand and foot of this Reptile with those of Mesosaurus, one 
cannot fail being struck by the concordance in the number of phalanges, and the great 
similarity in shape of the epipodials and phalangeals. The number of carpal bones is 
the same in both, but in Mesosaurus the tarsus comprises five elements in the distal 
row, and the fifth toe, instead of being shorter than the fourth, is a little longer. This 
agreement, together with other characters, such as the structure of the humerus and 
the form of the ribs, points to special affinity between the Wesosauria and Nothosauria, 
as first recognized by Prof. Seeley. This author, in his recent paper on Vesosaurus 
(25), places Newsticosawrus, a close ally of the Reptile here described, in a division, 
Neusticosauria, of the Suborder Mesosauria, characterized by flat articular surfaces of 
the centrum, coracoid separated from scapula, a notch in the pubis, a long neck and a 
short tail. As regards the latter character, which, even if founded ona well ascertained 
fact, would be at most a generic character, Prof. Seeley overlooks his former account 
of Neusticosaurus, in which he expresses the opinion that the tails of the specimens 
described by him are imperfect, and also the fine photograph, exhibited in the Geolo- 
gical Gallery of the British Museum, of a slab from the Lettenkohl of Hoheneck, 
Wiirtemberg, representing a group of Neusticosawrus pusillus, which have the tail 


FROM THE TRIAS OF LOMBARDY. 7 


nearly as long as the neck and body, thus proportionately quite as long as in Meso- 
saurus or Macromerosaurus. So far as Iam able to make out from the photograph, 
the tail of WV. pusil/us comprises at least 40 vertebrae. The notch in the pubis, to 
which Prof. Seeley alludes, is not distinctly shown on the lithographed figure published 
in 1882, but is well marked both on the woodcut, p. 364 of the description, and on 
the specimen No. R. 53 in the British Museum, and agrees in every respect with the 
figure of the Reptile referred by Deecke (16) to Lariosawrus. Dames (22) has already 
pointed out that Deecke’s Lariosaurus could hardly be the same as the Lariosaurus of 
Crivelli and Curioni, from which it differs in the shape of the clavicle, the feebly curved 
and distally expanded humerus, the longer propodials, and the notched pubis. Apart 
from the supposed absence of an ectepicondylar foramen in the humerus, which may 
after all be due to its having been overlooked, I can see no ground for regarding 
Deecke’s specimen as generically distinct from Neusticosaurus pusillus. Of course, so 
long as the skull remains unknown, no certain conclusion can be arrived at, but I am 
much struck by the agreement of Deecke’s and Fraas’s specimens in the following 
points :— 

1. The cervical vertebre are less massive than in Lariosaurus, 

2. The dorsal ribs are much thicker in their proximal than in their distal portion ; 

the head is expanded and slightly notched. 

3, The number of dorsal vertebre, 23 to 25, which is higher than in Lariosaurus. 
. The distally flattened and strongly expanded humerus. 
5. The slender, proximally expanded radius, which exceeds half the length of the 

humerus. 
6. The shape of the coracoid. 
7. The shape of the pubis. 


With regard to Weusticosaurus pusillus I must observe that there are certainly 21, 
and perhaps 22 cervical vertebre; and that therefore the generic identity of this 
Reptile with Pachypleura, suggested by Lydekker (21) and accepted by Zittel (20), is 
out of question, as well as its identification with Lariosawrus proposed by Bassani (18)1. 

Pachypleura edwardsii, as known from the descriptions and figures of Cornalia (9) 
and Curioni (12), agrees in many respects with Zariosawrus and Neusticosaurus; but 
the neck contains only 16 vertebre, and the humerus is longer than the femur; the 
radius and ulna are slender, and two thirds the length of the humerus; the exact 
number of phalanges is still unknown, 

Dactylosawrus, Giirich (15), from the Muschelkalk of Silesia, agrees in the number 
of cervical vertebra and the shape and proportion of the radius and ulna with Pachy- 
pleura; the phalanges number 2, 3, 3, 4, 3, and the fourth finger is longer than the third. 


aS 


1 The specimen from Viggit, figured on plate ii. of Cornalia’s memoir, must be taken as the type of Pachy- 
pleura edwardsii. The specimen from Besano, figured on plate i., is possibly a Neusticosaurus. 


8 MR. G. A. BOULENGER ON A NOTHOSAURIAN REPTILE 


Assuming the specimen described in this paper to be the young of the larger Lario- 
saurus balsami, the genus Lariosaurus is to be distinguished from Neusticosaurus by 
the larger and more elongate skull, the shorter and broader radius, the shape of the 
humerus, and the absence of notch in the pubis; from Pachypleura and Dactylosaurus 
in the greater number of cervical vertebra, the shorter and broader radius, and the 
shape of the humerus ; and from the latter more particularly in the relative proportions 
of the third and fourth fingers and the number of phalanges in the third finger. These 
four genera may be conveniently grouped, for the present, under one family, Lario- 
sauride, in many respects intermediate between the Mesosauride and the Notho- 
sauride, although nearer the latter, as shown in the following attempt at a classification 
of the Order Plesiosauria :— 


Order PLESIOSAURIA. 


Skull with a single temporal arch. Dentition thecodont. Plastron present, not connected with 
the clavicles. No ossified praecoracoid. Sternum absent. Dorsal ribs single-headed. 

Divided into three Suborders. 

Before passing on to the diagnosis of the minor groups, I must observe that I reckon the so- 
called “pectoral” vertebre as cervicals, and that I accept Seeley’s interpretation of the shoulder- 
girdle. There is no evidence to show that the bar termed “ precoracoid ” in Chelonians constitutes 
an element distinct from the scapula; therefore, I revert to the nomenclature of Cuvier and 
Rathke, and term this bar acromial process of the scapula in both Chelonians and Plesiosaurians. 


I. M&sosauria. 


Clavicles present; scapule fused with coracoids, the axes of which are transverse. Humerus 
with entepicondylar foramen ; propodial bones elongate ; digits with not more than 5 phalanges. 
Vertebr conically cupped, with persistent notochordal canal, without transverse processes ; 
cervical vertebrz 10 or 11; sacral vertebre 4. 

A single family :— 

1. Mesosavripz. 


II. NorHosavria. 


Clavicles strongly developed, forming a bar across the pectoral region, embracing a small inter- 
clavicle; scapulz with rudimentary acromial process; coracoids elongate, forming an angle, in 
contact by their extremities. Humerus with entepicondylar foramen ; propodial bones elongate ; 
digits with not more than 5 phalanges. Vertebre biplane or feebly biconcave ; transverse processes 
short or absent; cervical vertebre 16 to 21; sacral vertebree 3 to 51. 


1. Lartosaurip#.—Palate with infraorbital foramen. Dorsal vertebre without transverse 
processes, 


2. Nornosaurip#£.—Palate without infraorbital foramen. Dorsal vertebree with transverse 
processes. 


* Their number is still unknown in the Nothosauride. However, Anarosaurus, Dames, which I would refer 
to this family, is stated to have three sacrals. 


FROM THE TRIAS OF LOMBARDY. 9 


III. Savropreryera. 


Clavicles small, if present, applied to the dorsal surface of the scapulz ; scapule with much- 
developed acromial process; coracoids enormously developed, with their long axes parallel, their 
inner borders in contact. Limbs paddle-shaped, with the propodials more or less abbreviated, and 
an increased number of phalanges. Vertebre biplane or feebly biconcave; dorsals with long 
transverse processes ; cervical vertebre 20 to 72; sacral vertebre 1 or 2". 

Three families :— 


1, Priosavrma.—No median prolongation of the scapule and coracoids, which are widely sepa- 
rated from each other on the median line, thus enclosing a single large foramen; scapule not 
meeting in the middle. About 20 cervical vertebre, with forked ribs. 


2. PLesiosaurip#.—Scapule separated on the middle line; coracoids with median anterior 
prolongation, meeting the interclavicle and clavicles; the pectoral arch thus enclosing two 
foramina. Cervical vertebre 28 to 40, with forked ribs. 


3. Exasmosauripa.—Scapulz extensively in contact with each other on the middle line, where 
they join corresponding median processes of the coracoids ; the pectoral arch thus enclosing two 
foramina. Cervical vertebree 35 to 72, with single-headed ribs. Propodial bones much modified, 


simulating mesopodials. 


REFERENCES TO THE LITERATURE ON NOTHOSAURIANS. 


1. Meyer, H. von. Conchiosaurus clavatus, em Saurus aus dem Muschelkalke von Bayreuth. 
Mus. Senckenb. i. 1834, pp. 8-14, pl. i. 

2. Minsrur, G. zu. Vorlaufige Nachricht tiber einige neue Reptilien im Muschelkalk von 
Baiern. N. Jahrb. Min. 1834, pp. 521-527. 

3. Batsamo-Crivetut, G. Descrizione di un nuovo rettile fossile, della famiglia dei Palzosauri. 
Il Politecnico (Milano), May No., 1839, p. 421.—Abstract in N. Jahrb. Min. 1848, 
p. 246. 

4. Braun, F. Verzeichniss der in der Kreis-Naturalien-Sammlung zu Bayreuth befindlichen 
Petrefakten. Leipzig, 1840, 4to. 

5. Meyer, H. von. Simosaurus, die Stumpfschnautze, ein Saurier aus dem Muschelkalke von 
Luneville. N. Jahrb. Min. 1842, pp. 184-197. 

6. Meyer, H. von, and Purenincrr, J. Beitrage zur Paleontologie Wiirttembergs, enthaltend 
die fossile Wirbelthierreste aus den Triasgebilden. Stuttgart, 1844, 4to. 

7. Meyer, H. von. Zur Fauna der Vorwelt. Die Saurier des Muschelkalkes. Frankfort/M., 
1847-1855, fol. 

8. Curton1, G. Cenni sopra un nuovo Saurio fossile dei monti di Perledo sul Lario. Giorn. 

; Ist. Lomb. xvi. 1847, pp. 157-170, pl. —. 

g. Cornauia, HE, Notizie zoologiche sul Pachyplewra edwardsii, nuovo sauro acrodonte degli 
strati triasici di Lombardia. Giorn. Ist. Lomb. (2) vi. 1854, pp. 45-58, pls. i. & ii. 

10. Buainvittz, H.M.D. px. Ostéographie. Publication posthume. Reptiles. Paris, 1855, fol. 


1 This character should probably be erased from the diagnosis. Mr. Andrews (Ann. & Mag. Nat. Hist. 6, 
xy. 1895, p. 334) states that there appear to be three or four sacral vertebre in Cryptoclidus. [June, 1895. ] 


VOL. XIv.—PakT I. No. 2.— April, 1896. c 


10 


II, 
12. 


13. 
T4. 


15. 


16. 


17. 


18, 


19. 


20. 


21. 


22. 
De 


24. 


25. 


26, 


27. 


ON A NOTHOSAURIAN REPTILE FROM THE TRIAS OF LOMBARDY. 


Gervais, P. Zoologie et Paléontologie frangaises. 2° édition. Paris, 1859, 4to. 

Curron1, G. Sui giacimenti metalliferi e bituminosi triasici di Besano. Mem. Ist. Lomb. 
ix, 1864, pp. 241-268, pls. v. & vi. 

Fraas, O. Ueber Simosaurus pusillus. Wiirtt. Jahresh. xxxvii. 1881, pp. 319-324, pl. i. 

Szrtry, H.G. On Neusticosaurus pusillus. Q. Journ. Geol. Soc. xxxviii. 1882, pp. 350-366, 
pl. xiii. 

Guricu, G. Ueber einige Saurier des Oberschlesischen Muschelkalks. Zeitschr. Deutsch. 
Geol. Ges. xxxvi. 1884, pp. 125-144, pl. ii. 

Dexzcxe, W. Ueber Lariosaurus und einige andere Saurier der Lombardischen Trias. 
Zeitschr, Deutsch. Geol. Ges. xxxviil. 1886, pp. 170-197, pls. iii. & iv. 

Baur, G. Bemerkungen iiber Sauropterygia und Ichthyopterygia. Zool. Anz. 1886, 
pp. 245-252, & p. 323. 

Bassani, F. Sui fossili e sull’ eta degli schisti bituminosi triasici di Besano in Lombardia. 
Atti Soe. Ital. xxix. 1886, pp. 15-72. 

Kouniscu, H. Ueber eine Saurierplatte aus dem Oberschlesischen Muschelkalke. Zeitschr. 
Deutsch. Geol. Ges. xl, 1888, pp. 671-693, pls. xxix. & xxx. 

Zrrret, K. A. Handbuch der Palzontologie. Palzozoologie. III. Munich & Leipzig, 
1888, 8vo. 

Lyprexxer, R. Catalogue of the Fossil Reptilia and Amphibia in the British Museum. II. 
London, 1889, 8vo. 

Dames, W. Anarosaurus pumilio. Zeitschr. Deutsch. Geol. Ges. xlii. 1890, pp. 74-85, pl. i. 

Koxen, E. Ueber die Bildung des Schidels, der Gehirnhéhle und des Gehérorgans bei der 
Gattung Nothosaurus. Sitzb. Ges. Naturf. Fr. 1890, pp. 108-111. 
Giricu, G. Ueber einen neuen Nothosaurus von Gogolin in Oberschlesien. Zertschr. 
Deutsch. Geol. Ges. xliii. 1891, pp. 967-970. fi 
Srrrzy, H. G. The Mesosauria of South Africa. Q. Journ. Geol. Soc. xlviii. 1892, 
pp. 586-604, pl. xviii. 

SxurHos, T.G. Vorliufige Mittheilung tiber Parthanosaurus zitteli, een neuen Saurier 
aus der Trias. Zool. Anz. 1893, pp. 67-69. 

Koxen, E. Beitrage zur Kenntniss der Gattung Nothosaurus. Zeitschr. Deutsch. Geol. Ges. 
xlv. 1893, pp. 337-877, pls. vii-—xi. ; 


EXPLANATION OF PLATE I. 


Lariosaurus balsami, of the natural size. The matrix is represented lighter than it 


really is, in order to render the outlines of the bones more distinct (see p. 2). 


Trans. Zoot. Soc Vb AN FEL 


A.M. Woodward del.et hth 


Mirtern Bros. imp 
LARIOSAURUS BALSAM]. 


at 


Il. On the Lepidosiren of Paraguay, and on the external characters of Lepidosiren and 
Protopterus. by E. Ray Lanxesrer, W.A., F.RS., P.Z.S., Linacre Professor of 
Comparative Anatomy in the University of Oxford. 


Received and read June 19th, 1894. 
[Puate II. | 


THE object of the present communication is to place in the hands of zoologists a 
carefully-executed drawing of a South-American Lepidosiren (Pl. II. fig. 1), which, as 
will be seen below, there is reason to consider as identical with Natterer’s Lepidosiren 
paradoxa (Fitzinger), side by side with a drawing of the African Protopterus annectens, 
Owen, from the Gambia (Pl. II. fig. 2). Measurements are also given of the two 
species, and illustrations of the proportionate size and of the structure of the scales of 
the three species—Ceratodus forsteri, Krefit, Protopterus annectens, Owen, and Lepi- 
dosiren, from Paraguay (Pl. II. figs. 4-9). The remarkable villi of the posterior limbs 
of the male specimens of the Paraguay Lepidosiren are also carefully represented 
(Pl. II. fig. 3). 

From these data zoologists will be able to form a more correct conception of the 
appearance of Lepidosiren than is possible from Natterer’s figure, whilst the differences 
between it and Protopterus become obvious. 

I do not propose to attempt to decide critically whether the specimens of Lepidosiren 
from Paraguay, which I have examined, are the Lepidosiren paradoxa of Fitzinger or 
not’. The specimens described by Natterer, and named by Fitzinger, were obtained 
from ponds in the neighbourhood of Borba (not Bahia, fide Castelnau), on the Madeira 
River, a tributary of the Amazon system. One specimen is recorded by Castelnau as 
captured since Natterer’s discovery—having been taken by himself in the Lake Ucayale, 
which is in Eastern Peru, also connected with a tributary of the Amazons,—and two 
specimens (one only a skeleton) are in the possession of Prof. Giglioli, of Florence, one 
from Manaos, the other from Madeira, both localities in the Amazon basin. 

The Paraguay specimens, which were brought to Europe in the beginning of 1894 by 
a German traveller, Dr. Bohls’, were obtained from the neighbourhood of the Upper 


* See, however, the postscript to this paper, p. 20. 

2 Dr. Bohls has communicated to the Royal Society of Sciences of Gottingen (‘ Nachrichten,’ 1894, p. 80) an 
account of his capture of these specimens and of the various points relating to the natural history of Lepidosiren 
observed by him. The account is so interesting that I shall give here some extracts from it. Dr. Bohls 
states that he discovered these Lepidosiren in the interior of the swamps of the Chaco (on the right bank of the 
Paraguay River). The Lepidosiren does not occur in the river itself, but only in the pools of the swampy 

c2 


12 PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 


Paraguay River, in “the swamps of the Chaco.” Six of these specimens I have had 
the opportunity of examining: two more closely, which I purchased for the Oxford 
University Museum; three purchased by the British Museum, Natural History (by kind 
permission of the Keeper of the Zoological Collections) ; and one in the possession of 
Mr. E. Gerrard, Jun. 

Since these specimens, from one of which (now in the Oxford Museum) the drawing 
in the accompanying Plate has been prepared, were from the more southern river- 
system of the La Plata, and not from the great Amazon basin, there is, primé facie, a 
question possible as to whether we have here to deal with Natterer’s species or a 
distinct and new species. Castelnau did not hesitate to assign his specimen from 
Lake Ucayali, in the Amazonian area, to a new species; but probably, in view of the 


district near it. It appears that Natterer’s and Castelnau’s specimens were also from pools and not from 
rivers. Dr. Bohls believes that, whilst there is a southern limit to the distribution of Lepidosiren, it will be 
be found everywhere in the northern parts of South America in stagnant pools which are of fair depth (5 to 6 
feet). The particular swamp in which Dr. Bohls captured his specimens was four days’ journey west of the 
Paraguay River, and its borders were inhabited by a primitive tribe of Lengua Indians. They make the 
Lepidosiren and other fish captured in the pools their chief article of diet. The Lepidosiren cannot be caught 
with nets (on account of the weeds), nor by hook and line; they are caught with a spear or harpoon of about 
eight feet in length. The Indians plunge into the water in parties, prodding the bottom of the pools with 
these instruments. Dr. Bohls himself went out with a party of ten Indians. The ovaries of the Lepidosiren 
are preferred as an article of food to the salmon-like flesh, and are pressed into a kind of cake. The Lenguas 
Indians call the Lepidosiren “ Loalach.” The Lepidosiren feeds chiefly on a marsh-snail (Ampullaria) which 
grows as large as a man’s fist, and has a dense shell which the powerful teeth of the Lepidosiren are well fitted 
to crush. The bite of the animal is much feared by the Indians. Vegetable matter is found in the alimentary 
canal of the Lepidosiren together with the remains of the snails, but Dr. Bohls thinks it probable that this may 
have been swallowed accidentally, and not as food. Few examples of the fish are taken in a complete condition ; 
one of the limbs may be missing, or the tail injured (as often noted also with Protopterus). The Jacare 
(Alligator sclerops) feeds on the Lepidosiren in these pools, and specimens were found with the whole region 
of the body posterior to the anus in a state of regeneration—the amputation haying been probably due to the 
bite of an Alligator. 

Dr. Bohls is unable to say from actual observation that Lepidosiren can live in the dry mud of the pools, 
but as the swamps do dry up when the weather is hot and little rain falls, they must either die or pass through 
a period of non-aquatic life. Natterer relates that his Zepidostren could give a cry like that of a cat. Those 
observed by Dr. Bohls gaye out a sound when removed from the water, caused by expelling air through the 
narrow aperture of the branchial chamber. 

The alcohol in which the Lepidosiren were preserved acquired a green colour, and this colouring matter was 
given out even after months, when the alcohol had been replaced several times (cf. green colour of the bones of 
Protopterus). When Dr. Bohls left the Chaco the females were quite ripe with distended ovaries, but no eggs 
apparently had been laid. The specimens brought home by him therefore are, many of them, in a state 
approaching that cf the spawning period, but not actually arrived at that condition (see above as to my 
observations on the villi of the pelvic limbs of the males). Dr. Bohls tried to keep specimens alive in tanks, 
but, as all were necessarily injured by the spear in capture, they invariably died on the second or third day 
after they were obtained, 


PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 13 


absence from his description of any distinctive characters, no one will be found to 
agree with him. 

The view taken as to the Paraguayan Lepidosiren being distinct from that of the 
Amazons will no doubt depend, to some extent, on the conclusion which is accepted 
as to the existence of a single species of Protopterus in the African continent. If 
we agree with Dr. Giinther that Peters’s Protopterus amphibius from the Zambesi 
(Quillemane) is not really distinct from the Protopterus annectens, Owen, of the 
western water-shed of Africa (Gambia), then it will seem not unlikely that a single 
species of Lepidosiren may similarly inhabit two distinct river-systems in South 
America’. 

From an examination which I have made of specimens of Protopterus in the British 
Museum and elsewhere, I cannot regard the question of the species of African 
Protopteri as finally settled, and would venture to point out that it is very desirable 
that large and well-preserved specimens should be secured, by those who may have 
the opportunity, from each of the different river-systems of Africa and deposited in 
European collections for study. It seems that we must either admit that a very 
marked range of variation is exhibited (as is @ priori likely enough) in specimens of 
one species of both Protopterus and Lepidosiren, or that more than one species has to 
be recognized in each genus. 

On the whole, I think it will be best to assume for the present that Bobls’s Paraguay 
Lepidosiren? is identical with the L. paradoxa of Fitzinger and Natterer, and I now 


1 Tt appears that Professor Peters refers some of the specimens from the Zambesi and also those from the 
Nile and from the Tanganyika district to the species P. annectens of Owen, and that his P. amphibius is repre- 
sented only by certain specimens brought by him from the district of the delta of the Zambesi (Quillimane), 
and preserved in the Berlin Museum. P. amphibius is stated to possess only thirty pairs of ribs instead of 
thirty-five (P. annectens), and as its specially distinctive character is given not merely that a fin-membrane 
(« Flossensaum’) is present on one border of both pectoral and pelvic limbs (instead of on the pectorals only, as 
in P. annectens), but that the membrane in both limbs is supported by minute cartilaginous rods, which are 
wanting in P. annectens. (See Schneider, ‘ Zoologische Beitriige,’ vol. ii. 1890, p. 97). 

The determination of the presence or absence of the minute fringing cartilages of the fins requires the use of 
the microscope. I have examined specimens from the Zambesi (in the British Museum) with a fin-membrane 
well developed on the hinder limbs, but was not able to apply the microscope. As to how far these characters 
aro really indicative of specific separation is still doubtful, and can be cleared up only by minute examination of 
large series of specimens, which it is to be hoped will soon be forthcoming now that our communication with 
Equatorial Africa is so greatly improved and extended. 

2 Prof, Ehlers, in an interesting communication to the Royal Society of Sciences of Gottingen (‘ Nachrichten,’ 
1894, p. 84), gives reasons for considering that Dr. Bohls’s collection of Lepidosiven from Paraguay consists 
of two species—one which he considers identical with Natterer’s L. paradoxa, and a second which he distin- 
guishes as L. articulata, n. sp. Prof. Ehlers had the opportunity of looking over thirty-two specimens from 
Paraguay, of which he considers that twenty-seven belong to his new species, whilst five are to be referred to 
L. paradoua of Fitzinger. 

The characters adduced by Prof. Ehlers for separating these two species appear to me to be insufficient 


14 PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 


propose to point out what are the chief differences of form and proportion between 
Lepidosiren and Protopterus as represented on the one hand by the Paraguay specimens 
of Lepidosiren, and on the other hand by specimens of Protopterus from both the 
Gambia and the Zambesi. 

Sir Richard Owen, in his original description of Protopterus annectens (Trans. Linn. 
Soc. xviii. 1841, p. 327), pointed out important differences in his new form from Africa 


depending as they do on the general form of the head and snout, which is liable to distortion owing to the 
specimen resting on the head when placed in a jar containing the alcohol in which it is preserved. The males, 
both of the series assigned to the old species, L. paradowa, Fitz., and of the series assigned by Ehlers to his 
new species, presented equally the remarkable villi on the pelvic limbs hitherto unknown in Lepidosiren, In 
the females of both the villi are absent—the specimens brought home formerly by Natterer were females. 

The only character which Ehlers adduces for separating the Paraguay specimens into two distinct species, 
which seems to me likely to be of value, is that upon which he bases the specific name ‘ articulata.’ 

One (or more? Prof. Ehlers does not state) of the specimens belonging to the series of twenty-seven, which 
on account of shape of head and colour he had set apart as a new species distinct from the remaining five, was 
‘found to exhibit a segmentation of the cartilaginous skeletal axis of the limbs, both pectoral and pelvic. Now 
it is well known that according to Bischoff’s description and figure the skeletal axis of the limb (of each girdle). 
of L. paradowa, Fitz., is an unsegmented continuous tapering rod of cartilage. In the African Protopterus the 
corresponding skeletal element consists of successive segments. 

As Prof. Ehlers justly observes, the character which he has thus put forward requires testing on a large 
number of specimens. It does not appear from his statement that he has so tested it. It would not even follow 
that two species of Lepidosiren exist side by side in Paraguay, because some specimens are found to have 
segmented skeletal axes to the limbs, and a certain proportion are found to haye an unsegmented axis. 

The fact is that the specific determination of the Paraguay Lepidosiren has yet to be made by comparison, 
either with the five specimens from the Amazons now in European Museums (one in Paris, two in Vienna, two 
in Florence), or with new specimens to be obtained from the Amazon system. (See postscript, p. 20.) 

Prof. Ehlers’s identification of five specimens, out of thirty-two brought by Dr. Bohls, with L. paradowa, 
Fitz., cannot be regarded as resting on satisfactory grounds, any more than the separation of the other 
twenty-seven specimens as a distinct species. That one of the latter had a jointed axis to the limb cannot be 
held to go very far; we do not really know at present whether such jointing is common, or occasional, or never 
present in Lepidosiren annectens, and until we do it will be best to consider Dr. Bohls’s specimens from 
Paraguay provisionally as identical with the Amazonian L. annectens, Fitz. (See, however, the postscript to 
this paper, p. 20.) 

Bischoff’s figure and description of the cartilaginous axis of the limb of Z. annectens, Fitz., in the ‘ Annales 
des Sciences Naturelles,’ 1840, does not justify a final conclusion as to the absence of segmentation in that 
axis. Protopterus with its corresponding segmented axis was not known at the time, and in an insufficiently 
cleaned preparation it is possible that segmentation may have escaped Bischoff’s notice. He makes no emphatic 
statement on the subject. 

I am unable to gather from Prof. Ehlers’s statement whether he he has actually seen a Lepidosiren with an 
unsegmented cartilaginous axis to the limb, or whether he is merely quoting and assuming the finality of 
Bischoff’s statement. He says: “‘ Bei dem von mir so gedeuteten Thiere (Z. annectens) ist auch das Skelet der 
Gliedmassen ein einfacher ungegliederter Knorpelstab.” Among Dr. Bohls’s specimens there were five “so 
gedeutet ” by Prof. Ehlers. It would be satisfactory to know whether all five, or how many, were ascertained 
to have the simple unsegmented cartilage-axis, or whether any specimen at all was really ascertained by 
Prof. Ehlers to exhibit “ ein einfacher ungegliederter Knorpelstab ” as the fin-skeleton, 


PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 15 


and the description given by Natterer of the South-American Lepidosiren. He especially 
drew attention to the relative distance (and number of vertebrz) intervening in the 
two species between the anterior and posterior limbs. The difference thus indicated 
is very striking, and is expressed in general terms by saying that Lepidosiren is long 
and eel-like, whilst Protopterus is comparatively short and thick-set. 

In order to place the distinction on a definite numerical footing, it is found convenient 
to take the following points of measurement:—A. Total length from the extremity of 
the snout to the extremity of the caudal fin; B. From the extremity of the snout to 
the anterior margin of the pectoral fin where it springs from the body; C. From the 
anterior margin of the pectoral fin to the anterior margin of the pelvic fin; D. From 
the anterior margin of the pelvic fin to the extremity of the tail; E. From the anterior 
margin of the pectoral fin to the vertical drawn from the anterior limit of the dorsal fin. 

In the following lists I give these measurements, A, B, C, D, E, for five specimens 
of the Paraguay Lepidosiren, and for six specimens of Protopterus. It is no doubt true— 
that the ratio of these lengths one to another differs in larger specimens as compared 
with quite small specimens; also that the distortion caused by stuffing may entirely 
obscure the natural proportions. With the exception of two large stuffed specimens 
of Protopterus in the British Museum, all the specimens, the measurements of which 
are here recorded, were preserved in alcohol. 

If we take the length of B (from tip of snout to anterior margin of pectoral fin) as a 
unit, these measurements may be expressed in a form facilitating a direct comparison 
of all the specimens examined. 

It will facilitate the statement of these results if we call A the total length, B the 
head-length, C the inter-membral length, D the post-pelvic length, and E the cervico- 
dorsal length. If we give the total length in centimeters it will be unnecessary to 
burden the reader with the other measurements in centimeters; it will be sufficient 
to give them in terms of the unit of comparison adopted, viz. the head-length. 


Tastr A.— Measurements of Lepidosiren from Paraguay. 


Total Ratio Head- Ratio of Ratio of Ratio of 
No. Specimen. length in of total length inter-membral | post-pelyic |cervico-dorsal 
centimeters. | length. | (unit). length. length. length, 
1. | Smaller male Lepidosiren from 57 8°77 1 4:7 3:15 3 
| Paraguay (Oxford Museum) . 
2. | Larger male Lepidosiren from 73°3 10°47 1 5°85 3°57 2-92 | 
Paraguay (Oxford Museum) . | 
8. | Large male Lepidosiren from 72 9°27 1 5 3°38 258 | 
Paraguay (British Museum) . 
4, |Small male Lepidosiren from 40°5 10°12 1 5 41 3-06 
Paraguay (British Museum) . 
5. | Female Lepidosiren from Para- 67 10:3 1 5°53 3°77 354 | 
guay (British Museum) .... | 
Averaper Gove eenieak ere 975 | 1 5-21 3°59 | 32 | 


16 PROF, E, RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 


Taste B.—Measurements of specimens of Protopterus annectens, Owen. 


Total Ratio of total| Head- Ratio of Ratio of Ratio of 
No. Specimen. length in length to | length | inter-membral | post-pelvie | cervico-dorsal 
centimeters. | head-length. | (unit). length. length. length. 
1. | Specimen in Oxford Museum, 
killed, and measured three 
hours after death (see Pl. IT., 
and explanation where this 
specimen is figured), from the 
Cems. Sa otgosoneno gdcs 50 6:25 1 25 2°63 1:3 
2. | Quite small specimen in the 
Oxford Museum .......... 23 7°66 1 33 3:3 14 
3. | From the Zambesi (British 
WHEN) SG ngacbeadno oDno 39 7 1 3 3 1:2 
4. |A second from the Zambesi 
(British Museum) ........ 32 8 1 3°25 3°75 15 
5. | Stuffed specimen in British 
Museum, formerly living in 
the Crystal Palace, Sydenham. 79 9:3 i 3°53 417 1:64 
6. | Very large stuffed specimen in 
British Museum (purchased 
of M. Parzudaki): Nile .... 90 vi 1 3°46 2-46 2 


The facts indicated by these two sets of measurements (excluding any close following 
of the figures derived from the stuffed specimens of Protopterus) are, firstly, that the 
proportional value of the measurements differs greatly in different specimens; and 
secondly, that if we take a fair well-grown specimen of Protopterus on the one hand 
and of Lepidosiren on the other, the former from 40-50 centimeters, and the latter 
60-70 centimeters in length, they contrast most strongly in the following points— 
namely, that in Protopterus the whole animal is about 64 times as long as its head, 
whereas in Lepidosiren the whole animal is about 10 times as long as its head. Next, 
that whereas in Protopterus the inter-membral length is about 23 times the length of 
the head, it is about 5 times the length of the head in Lepidosiren; and, lastly, that 
whereas in Protopterus it is about once and a half the length of the head from the 
pectoral to the commencement of the dorsal fin, it takes as much as three head-lengths 
to cover the same line in Lepidosiren. The post-pelvic ratio of length is nearly the 
same in the two genera. 

These external differences are, of course, accompanied by corresponding numerical 
differences in the number of myocommata and vertebre, which have already been noted 
by Owen, and there are corresponding differences in the number of scales between the 
points taken. It is not my purpose on the present occasion to deal with points of 
anatomy, and with regard to the enumeration of scales I will record that it is by no 
means an easy task in Lepidosiren on account of the delicacy of the scales, their small 
size and deep implantation, as well as the abundant coagulated and strongly pigmented 
tissue which covers them in specimens recently preserved in spirit and therefore not. 
macerated. 


PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 17 


The form and ornamentation of the scales have been more or less fully described 
by Natterer and Bischoff in Lepidosiren, by Owen in Protopterus, and by Giinther 
in Ceratodus. From Wiedersheim we have a more minute account of the denticles 
which are set on the polygonal aree marking the scales of Protopterus. In Pl. II. 
figs. 4-9, I have reproduced drawings prepared for me by Mr. HE. S. Goodrich, F.L.S., of 
Merton College, my assistant in the Oxford Museum, showing the relative size of the 
scales in the three genera Ceratodus, Protopterus, and Lepidosiren, and also showing 
the denticulate ornament of the scales, equally magnified in each case for comparison. 
The scale chosen in each case is taken from a point near the middle of the inter- 
membral area, and about halfway between the dorsal and ventral mid-line; the object 
has been to select in each case a similarly-conditioned scale. It will be seen that the 
scale of Ceratodus is enormous compared with those of. the other two genera, and that 
that of Protopterus is larger than that of Lepidosiren. (See further, as to the scales, 
in the note to the Explanation of the Plates.) 

The denticulations of the scales do not vary in size proportionately to the size of the 
scales, but are only somewhat coarser in Ceratodus than in Protopterus and Lepidosiren 
(Pl. II. figs. 7-9). 

The last point, in reference to the Paraguayan Lepidosiren, to which I desire to draw 
attention in the present communication is the remarkable development of the villi on the 
hind limbs. ‘hese were described and figured by me in outline in a letter to ‘ Nature’ 
in April 1894. They have also been described, but not figured, by Prof. Ehlers in a 
communication to the Gottingen ‘ Nachrichten,’ 1894, p. 87, which was not published 
until June, and was unknown to me when my letter in ‘Nature’ was published, 
although some author’s reprints had been distributed before that date. It appears 
that these villi occur only on male specimens. They are present on all the male 
specimens I have examined, and are absent from the females. They occur on the 
postero-medial surface of the elongated pelvic fins, and are seen both in Pl. Il. fig. 1 
and in the enlarged drawing of the limb (PI. IT. fig. 3), which is represented as turned 
forward so as to expose them fully to view. The villi are small, pale-coloured, blunt 
processes, from three to four times as long as broad; they are either simple or joined, 
to the number of two, three, or four, on a common base. I have examined the micro- 
scopic structure of these villi by means of transverse sections. They are covered with 
epidermic cells, which are two or three deep, and show evidence, in the condition of 
their nuclei, of being in course of active multiplication. But I did not observe any 
special characters of interest in these cells; they were block-like, solid, and not 
flattened. Below the epithelial cells is a finely fibrous connective tissue, which in 
transverse section of the villi is seen to occupy the axis, and to take the form of trabecule 
enclosing oblong cavities with rounded contours. Numerous nuclei belonging to the 
connective tissue—some in karyokinetic activity—are apparent in such sections, and 
here and there a minute blood-vessel. ‘The cavernous structure of the connective tissue 

you. Xiv.—Parr 1. No. 3.—April, 1896. D 


18 PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 


and the vascularity of the villi are in favour of the supposition that the cavernous tissue 
may be erectile under certain conditions, probably at the time of sexual congress’. 


External Gills of Protopterus. 


The presence of external gills was given in the Brit. Mus. Cat. Fishes, viii. p. 322 
(1870), by Dr. Giinther, as diagnostic of the genus Protopterus in contrast to Lepidosiren. 
The diagnostic is adopted also by Schneider (loc. cit.). I believe that the distinguished 
zoologist of the British Museum would not at the present moment put forward that 
character as one to be insisted upon. There is no doubt that moderate-sized specimens 
of Protopterus, such as that figured in PI. II. fig. 2, do not possess external gills. 
While small specimens of Protopterus possess remarkable pigmented (yellow and 
black) external gills, it is impossible at present to use this character as distinguishing 
Protopterus from Lepidosiren, since small specimens of Lepidosiren have not been 
examined, and may therefore, for all we know, possess such gills also. 


The Specimen of Protopterus annectens, figured in Plate II. fig. 2. 


The Protopterus figured in the Plate accompanying this paper, for the purpose of 
comparison with Lepidosiren, was living in the tank of the Reptile House in Regent's 
Park three hours before it was sketched. The drawing has a special value, therefore, 
as being made from a perfectly fresh and undistorted specimen. It was kindly placed 
in my hands for the purpose of study by Dr. P. L. Sclater, Secretary to the Zoological 
Society of London. It proved to be a female, and is one of six from the Gambia River 
presented to the Society by Mr. H. H. Lee. 

It is remarkable that both the limbs and the tail of well-grown specimens of Protopterus 
seem to be specially liable to injury during life. I believe this injury arises not only 
from bites inflicted by the Protopteri on one another, but also from the attacks of a 
parasite which is allied to, if not identical with, the Saprolegnia ferox of salmon 
disease. In captivity they are, as a fact, attacked by Saprolegnia, and the consequent 
ulceration leads to loss of parts of the caudal and lateral fins. 

The female specimen which is drawn in PI. II. fig. 2 is remarkable for the complete- 
ness of the caudal fin and for the fact that a minute median filament is present, 
extending beyond the main outline of the fin posteriorly. 

Iam not able at present to give any further detail as to the nature of this process, 
the specimen being retained in a complete condition for exhibition in the University 
Museum, Oxford. 

The lateral fins are of more than average length as compared with specimens of this 
size. In young Protopteri the pectorals and pelvics are very long; in specimens 


* A demonstration that such is the nature of the villi would, however, require specimens in which the 
cond:tion of turgescence was actually retained in preservation, or fresh specimens upon which injection could 
be practised. 


PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS, 19 


5 inches long the pectoral fins may have a length of 52 inches. Apparently the fins 
are continually being shortened by violence or disease, and continually grow again, not 
to the relative length seen in the young, but so as to repair to a great extent the loss. 
Hence it seems that little importance can be attached, in the diagnosis of Protopterus 
and Lepidosiren, to the relative length of the fins, unless a very large series of each 
is studied. 

I am also unwilling to attach much importance to the general form of the head and 
the relative position of the eye in relation to the angle of the mouth as specific or 
generic characters, since we have no drawing or record of freshly killed or living 
specimens of Lepidosiren, while preservation in spirit is liable to be attended by 
considerable distortion of the head. 

But it is fairly apparent, as shown in the Plate, that the head of Protopterus is less 
blunt anteriorly than that of Lepéidosiren, and that the eye of Lepidosiren is placed 
much further forward than that of Protopterus, so as to be in front of the angle of - 
the mouth. 


EXPLANATION OF PLATE II. 


Fig. 1. Lepidosiren from the swamps of the Chaco (Upper Paraguay River), natural 
size: male. Drawn from a spirit-preserved specimen in the Oxford University 
Museum by J. Bayzand *. 

Fig. 2. Protopterus annectens,Owen. Drawn by J. Bayzand* from a specimen recently 
killed and untouched by alcohol. The specimen is a female, was brought 


* Note, Dec. 20, 1895.—It is only fair to the artist, Mr. Bayzand, to point out that a curious inaccuracy has 
crept into the lithographic reproduction of his drawing which is not present in the original as made by him, 
It will be observed, in the two large figures of Lepidosiren and Protopterus respectively, that there is a 
marking on the surface of the body, especially strongly rendered in the drawing of Protopterus, which 
appears to indicate “scales,” and was interpreted as such by the lithographer. As a matter of fact, no scales 
at all or parts of scales are visible on the surface of the body of a fresh or well-preserved specimen of Proto- 
pterus. The scales are entirely overlaid, in both Protopterus and Lepidosiren, by soft vascular connective tissue 
in addition to a well-developed epithelium. The arew which are marked out on the surface of the body and 
have been wrongly rendered in the lithograph as scales are in reality lozenge-shaped arew outlined by the 
greater abundance along their margins of the large branching pigment-cells of the connective tissue which 
overlies as a uniform and continuously flat layer the subjacent scales. The lithographer, thinking he was 
called upon to represent protruding imbricated scales, such as appear on the surface in many common 
Teleostean fishes, has changed the outline of the pigment-areze and represented the posterior border as a portion 
of a circular curve, a form which it does not present either in Protopterus or Lepidosiren, and which was not 
given to it in Mr. Bayzand’s original drawing. The true form of these arew delimited by pigment-cells is 
seen in the woodcut, fig. 1. The arex are seen to have a pointed angular posterior border, and not a curved 
The anterior border, on the contrary, shows a rounding-off of the angle, so that the form of the markings 


one. 
given in the lithograph should be reversed. The arew are less elongated near the head and near the origin of 


the median fins than in other regions. 

Though these are are in no way to be regarded as scales, they yet correspond in position and number with 
the subjacent imbricated scales, Each scale in Protopterus and Lepidosiren has a large sculptured area carrying 
the denticulations. figured in the Plate, and, extending posteriorly beyond this, a softer unsculptured portion 


D2 


20 PROF, E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS, 


from the Gambia district, West Africa, and had been living two years in the 
Society’s Gardens. 
Fig. 3. Left hind limb of the specimen of Lepidosiren drawn in fig. 1, reflected forward 


which is as it were cut off above and below so as to form a strong median angle (Plate II. figs. 5 & 6). The 
line separating the stronger sculptured part of the scale from the smooth softer piece presents a fine curvature 
(aa in figs. 5 & 6 of the Plate), In old spirit-specimens of Protopterus (woodcut, fig. 2) and also in similar 


Fig. 2. 


Fig. 1. 


ani. 


ost. 
B pst. 


Diagram of the lozenge-shaped ares outlined by pigment- 
cells (which are also seen scattered and in some places 
aggregated on the general surface), from the mid-body 
region of Protopterus annectens, as seen in a freshly- 


Diagram of the same are as those drawn in fig. 1, as 
seen in a specimen which has been long preserved in 
weak spirit. The pigment has become altered in colour 
(from blue-black to a warm brown) and has diffused 


preserved specimen (six months in spirit). 


into and stained adjacent tissue. The subepidermic 
connective tissue is wrinkled and semitransparent, and 
permits one to see the outline of the semicircular curve 
(a, @) which separates the sculptured from the unsculp=- 
tured portion of the subjacent scale. 


‘specimens of Lepidosiren, this curved line shows through the softened and wrinkled dermis (woodcut, fig. 2, aa) 
owing to the greater thickness and differentiated texture of the sculptured portion of the buried scale, 


Fig. 3. 


Diagram to show the relation of the scale of Protopterus 
to the superjacent lozenge-shaped area outlined by 
Ppigment-cells. a, the line separating the thicker 
sculptured portion of the scale from the softer and 
smooth, pointed portion, c; 6, the position of the 
pigment-cells disposed in the form of a lozenge. 


The relation of an entire scale to the lozenge-shaped 
pigment-area of the surface-tissues is shown in woodcut 
fig. 3, where the scale is drawn so as to show its sculp- 
tured and unsculptured portions, and the superjacent 
lozenge-area of pigment-cells is dotted in. 

In the figures 5 and 6 of Plate II. the dotted line 
indicates similarly the anterior border of a superjacent 
pigment-lozenge. 

If we now compare the figure of the scale of Cera- 
todus forsteri (Pl. II. fig. 4) with the diagrams and 
drawings of the scales of Protopterus and Lepidosiren, 
it becomes apparent that there is here also a smooth 
unsculptured posterior portion of the scale. In Cera- 
todus, however, this part of the scale is firm and thick, 
and is freely exposed, covered neither by dermis nor 
epidermic epithelium. 


The most probable interpretation of these facts, from the point of view of the ancestral history of the scale 


PROF, E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 21 


so as to expose the anus on the left side of the animal’s body, and showing 
the villi (found only in the male) on the dorso-mediad face of the limb. 
Drawn by J. Bayzand: twice the natural size, linear. 

Fig. 4. Scale from middle of the side of the body of Ceratodus forsteri, Krefft. 
xX 3diam. 46, dotted line marking off the exposed portion of the scale (to 
the right) from that which is sunk beneath the surface. 

Fig. 5. Scale from a similar position from Protopterus annectens, X 3 diam. 

Fig. 6. Scale from a similar position from the Paraguayan Lepidosiren, X 3 diam. 
For explanation of reference-letters a and 6 see footnote. 

Fig. 7. Denticulation of the scale of Ceratudus.  X 180 diam. 

Fig. 8. Denticulation of the scale of Protopterus. 180 diam. 

Fig. 9. Denticulation of the scale of Lepidosiren. X 180 diam. 


Postscript.—July 8th, 1895. 


Since the foregoing paper was written and the figures of the Plate drawn, I have made 
it my business to examine as far as possible the specimens of Lepidostren from the 
Amazons which exist in European Museums. 

I visited Paris in April, and by the courtesy of the Director of the Jardin des Plantes, 
M. Milne-Edwards, and of Prof. Le Vaillant was permitted to} examine carefully and to 
measure Castelnau’s specimen, as also a fragmentary skin from an old Portuguese 
collection, which is probably referable to the Amazonian Lepidosiren. 

I proceeded to Florence, where my old friend Prof. Giglioli placed at my service— 
for the purpose of determining the justification of Ehlers’s proposal to form the new 
species, Lepidosiren articulata—his two specimens from the Amazons. 

Lastly, I wrote to Dr. Steindachner, of Vienna, to ask him to examine Natterer’s two 
original specimens in order to determine whether the skeletal axis of the limb is 
“articulated,” or a simple continuous rod of cartilage. Dr. Steindachner wrote to 
inform me that the hinder limbs of the specimen anatomized by Bischoff appear to 
be lost: they cannot be found. At the same time he had the great kindness to send to 
me Natterer’s untouched second specimen in order that I might examine it myself. 
I have examined it, and returned it to Dr. Steindachner a month ago. 

Thus I have myself examined every specimen of Brazilian Lepidosiren known to 


and of the Dipnoi as a natural group, appears to be that the scales of Protopterus and Lepidosiren are in a state 
of retrogressive modification or degeneration as compared with those of Ceratodus. It would seem that, 
whereas the ancestral form presents a freely exposed posterior margin to the scale, the whole scale has shrunk 
and withdrawn, so to speak, within and beneath the outermost layers of the dermis in Protopterus and Lepido- 
siren, whilst Ceratodus retains the ancestral character. Of course, behind and beyond that (in historical 
sequence) there is a hypothetical series of more primitive ancestral phases which lead to the multitudinous 


dermic denticles of primitive fishes. : 


22 PROF, E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS, 


exist in European Museums, excepting the one of Natterer’s two which has lost the 
hinder limbs. 

The main point to which my observation was directed was the existence or non- 
existence of a segmented structure in the cartilaginous skeletal axis of the pectoral 
and pelvic fins. 

Tue Paris SpeciMENs.—I did not ask permission to cut into the limb of Castelnau’s. 
specimen in Paris; hence it remains intact, and may be appealed to should any doubt 
remain after what I have to relate of the other specimens. The small second specimen 
in Paris is very incomplete, and of no value for the present enquiry. 

Castelnau’s specimen measures 85 cm. in length. The head-length (as defined in the 
preceding memoir) is 9 cm., the inter-membral length is 48-5 cm., the post-membral 
length is 27:5 cm., and the cervico-dorsal length is 26 cm. The greatest depth of the 
body is 10 cm. The specimen is very soft, and somewhat flattened and collapsed, 
hence deeper than natural. The colour is very pale—greyish brown. ‘The anus is on 
the left side; there are no villi on the pelvic fin. The lower jaw was damaged in 
capture. The eye is far in front of the angle of the mouth (halfway between it and 
the border of the snout). The eye is flat, without raised cornea, as in all examples of 
Lepidosiren I have seen, whether from Brazil or Paraguay. 

Tue Fiorence Sprecimens.—One of these had been Fig. 4. 
macerated and more or less cleaned. The pelvic limbs 
were attached to the girdle, and presented the appearance 
shown in the woodcut (fig. 4.). The segmentation of the 
axis of the limb was most obvious! The fore limbs were 
very small, and still covered by integument. I dissected this 
away and found the axis “segmented” or “articulated” 
as in the hind limbs. This specimen was not a large 
one (about 40 cm.in length). It was sent to Prof. Giglioli 
by Senhor J. Barbosa Rodriguez, of Rio de Janeiro, 
and was procured by him at Antaz, Madeira, Amazons. 
The sex was not determined. ‘The local native name is 
“ Trayraboia.” 

Prof. Giglioli’s second specimen is larger, and is com- 
plete, except that the viscera have been removed (the 
existence of ova had been noted by Prof. Giglioli). It is 
82°25 cm. long. The head-length is 8°25 em., the inter- 
membral length 47 cm., the post-membral length 27 em., 
the cervico-dorsal 30cm, There are no villi on the pelvic fin, and the anus is on the 
left side. 

With Prof. Giglioli’s permission, I made a partial section of the right fore limb and 
of the right hind limb. In both the cartilaginous skeletal axis was seen to be made 


PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 23 


up of successive segments—the separate cartilages being divided by connective tissue. 
The fore limb in this specimen was very small and flattened. 

This specimen was also sent by Senhor Barbosa Rodriguez to Prof. Giglioli, He 
obtained it from Igarape do Aterro, Manaos, Amazons, where its native name is 
“ Piramboia.” 

Senhor Rodriguez recorded the capture of this specimen in the ‘ Jornal do Commercio,’ 
of Rio de Janeiro, 15th November, 1886, and while furnishing the reader with,a rough 
but characteristic figure (which would enable an up-country naturalist to recognize 
specimens should they come to hand), spoke of it as Lepidosiren gigliolana, n. sp. 

This specimen is a female, and is (as compared with Bohls’s Paraguayan specimens) 
singularly light-coloured. It is of a clear brown. I have, however, little doubt that 
the paleness of colour is due to the prolonged action of sunlight and of the spirit used 
as preservative. 

Tae Vienna Specimen (Narrerer’s oRIGINAL).—It was a rare pleasure to be able to 
handle one of the two specimens which fifty years ago were brought to Europe by 
the celebrated traveller Natterer, and were the first indication to zoologists of the 
existence of the group of Dipnoi, since expanded by the discovery of Protopterus and 
Ceratodus and of many Paleozoic allies. 

The specimen kindly sent to me by Dr. Steindachner was in beautiful condition, in 
spite of its fifty years’ sojourn in spirit. Like Castelnau’s and Giglioli’s specimens, it 
is a clear brown and not greenish black, as are Bohls’s specimens from Paraguay. 
Probably this is due to long soaking in spirit (see Bohls’s remarks, cited above, as to 
the pigment of his specimens). The length of this specimen was 58 cm., its head- 
length 55 cm., the inter-membral length 31 cm., the post-membral length 22 cm., and 
the cervico-dorsal length 16 cm. There were no villi on the pelvic fin. I did not 
determine the sex. I opened the integument of both a pectoral and a pelvic fin, and 
found that the skeletal axis was built up of separate articulated segments as in 
Prof. Giglioli’s two specimens and as in the three Paraguayan specimens which I 
have examined for the determination of this point. This specimen came from Madeira, 
Amazons. 

SumMARy oF ResuLts oF EXAMINATION oF THE AMAZONIAN LEPIDOSIREN.—Of the five 
specimens of Amazonian Lepidosiren which exist in Europe (for we may exclude the 
small Lisbon skin in the Jardin des Plantes) I have examined three in regard to the 
articulation of the skeletal axis of the limbs, and have found that that axis is in all 
segmented. Of the two remaining specimens, one (Natterer’s dissected by Bischoff ) 
has now lost the hinder limbs, the other (Castelnau’s) remains for reference; for 
although I examined it and took measurements, I did not cut into the integument of 
the limbs. 

I have not seen a Paraguayan Lepidosiren with non-segmented or non-articulated 
skeletal limb-axis, and it is desirable that Prof. Ehlers should clearly say whether he 


24 PROF. E. RAY LANKESTER ON LEPIDOSIREN AND PROTOPTERUS. 


has or has not. The words which he uses do not amount to a statement to the effect 
that he has. He says that five specimens out of thirty-two submitted to him by 
Dr. Bohls from Paraguay differed in certain features of the head from the other 
twenty-seven ; further, that one (or more?) of the twenty-seven had segmented limb- 
axes, and therefore (it seems) he concludes (but does not state as the result of 
observation) that the remaining five must be referred to Fitzinger’s L. paradowa, and 
must haye an unsegmented limb-axis, since Bischoff figured such an axis in one of the 
origin specimens of LZ. paradoxa. The probability, from my observations, is that 
Bischoff erroneously omitted the segmentation of the limb-axis in his figure, and did 
notelook into the matter carefully. It seems highly probable that all Brazilian 
specimens of Lepidosiren, and all Paraguayan specimens of J.epidosiren, have, like 
the African Protopterus, a segmented limb-axis. The chief ground put forward by 
Ehlers for erecting a new species, “Lepidosiren articulata” from Paraguay, would, if 
my surmise be correct, fall to the ground. 

Whether any specific distinction between Paraguayan and Amazonian specimens of 
Lepidosiren can be based upon other grounds remains to be seen, The characters 
derived from small differences in the shape of the head, relied upon by Ehlers, cannot, 
in my opinion, be considered as of any value in specimens preserved in spirit, and 
pressed down in jars or travelling-cases. The most striking and uniform difference 
which I have observed is in colour: the Paraguayan specimens are black, the Brazilian 
specimens are clear brown. ‘There is, however, little doubt that this is largely, if 
not altogether, due to the longer action of alcohol and sunlight upon the Brazilian 
specimens. 

Lastly, we may compare the measurements of the Brazilian and Paraguayan specimens. 

If we write down, for the three Brazilian specimens, the measurements already given 
in terms of head-length, it appears that Castelnau’s specimen has a total length of 
9°54 units, an inter-membral length of 5°5, a post-membral of 3, and a cervico-dorsal 
of 3; for Giglioli’s large specimen the corresponding figures are—10, 5-7, 3-4, and 
3°63; for the Vienna specimen they are—10-06, 5-64, 4, and 3. If we now take the 
average of these three Brazilian specimens, and compare it with the average of the five 
Paraguayan specimens given on p. 15, we obtain the following striking result :— 


Total length. Inter-membral. Post-membral.  Ceryico-dorsal. 
Brazilian average in head-length units ...... 9:87 5:25 35 3°21 
Paraguayan average in head-length units.... 9°75 521 3°59 32 


The specimens in both series were taken altogether by hazard, yet there is the closest 
agreement. No doubt such a result does not exclude the possibility of a specific 
difference being discovered separating the Paraguayan from the Brazilian Lepidosiren. 
But it renders the existence of such a difference very unlikely, and I have already 
shown that it is extremely probable that the character relied on by Prof. Ehlers in 
order to separate some Paraguayan specimens from the Brazilian species has no existence 
in fact. 


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CONTENTS. 


I. On a Nothosaurian Reptile from the Trias of Lombardy, apparently referable to 
Lariosaurus. By G. A. Boutencer, P.R.S. (PlateL.) . . . . - pagel 
Il. Qu the Lepidosiren of Paraguay, and on the external characters of Lepidosiren and 


Protopterus. By E. Ray Layxester, VA, FBS, F.Z.S., Linacre Professor 


of Comparative Anatomy in the University of Oxford. (PlateIL) . . . 


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IIT. On a Collection of Fishes from the Rio Paraguay. 
By G. A. Bouencer, F.R.S. 


Received May 20th, 1895, read June 18th, 1895. 
[Puares III.—VIII.] 


THE Trustees of the British Museum have lately acquired a large collection of 
Fishes formed by Dr. C. Ternetz at various localities in Matto Grosso and Paraguay. 
So few fishes have been collected in the Paraguay System since the time of Natterer 
that it seemed to me desirable to draw up an account of the collection, and to give a 
full list of all the species represented, several of which are new to science !. 


SCIANIDA. 


1. PLacioscion TERNETZI. (Plate III.) 
Plagioscion ternetzi, Boulenger, P. Z. S. 1895, p. 523. 

Outer upper and inner lower teeth considerably enlarged. Depth of body 3 times 
in total length, length of head 3 to 3% times; snout a little longer than diameter of 
eye, which is 5 to 54 times in length of head; interorbital width equal to iength of 
snout, 4 times in length of head; maxillary extending to below posterior border of 
eye; preopercular border rounded, finely denticulated behind, with wide-apart small 
serre at the angle and below. 15 gill-rakers on lower part of anterior arch, the 
longest as long as or a little shorter than longest gill-filaments. Lower pharyngeal 
teeth villiform. Dorsal X, I 33-35; third and fourth gr fourth and fifth spines 
longest, + to 3 length of head. Pectoral nearly }# length of head. Anal II 6; second 
spine very strong, 4 length of head. Caudal pointed, densely scaled. The distance 
between base of anal and base of caudal 24 depth of caudal peduncle. Scales nearly 
al! ctenoid, 95-100 44; lat. 1. 48-50. Uniform silvery. 

Total length 450 millim. 


Two specimens from Remanso, Rio Grande, Paraguay. 
This species is allied to Sciwna surinamensis, Blkr. 


2. PACHYURUS SCHOMBURGEI, Gthr. 
Paraguay. 
1 Diagnoses of the new species have appeared in P. Z. S, 1895, p. 525. 
VOL, xIv.— Part 11. No. 1.— October, 1896. 


26 MR. G. A. BOULENGER ON FISHES 


CICHLID. 


3. ACARA BIMACULATA, L. 
Descalvados, Matto Grosso; Paraguayan Chaco. 


4, ACARA TETRAMERUS, Heck. 


Descalvados, Matto Grosso; Paraguayan Chaco. 


5. Mmsonauta INsianis, Heck. 


Descalvados, Matto Grosso. 


4. CRENICICHLA JOHANNA, Heck. 


Paraguay. 


7. CRENICICHLA SAXATILIS, L. 


Paraguay. 


8. GEOPHAGUS DUODECIMSPINOSUS. (Plate IV. fig. 1.) 
Geophagus duodecimspinosus, Boulenger, P. Z. 8. 1895, p. 524. 

Depth of body 14 in total length, length of head 3 times. Eye nearer gill-opening 
than end of snout, its diameter 34 times in length of head and a little less than 
interorbital width; preorbital 14 diameter of eye; scales on cheek small, in 7 series. 
Dorsal XII 14; spines increasing in length to the fourth, which is 2 length of head ; 
soft portion scaly at the base, longest rays nearly as long as head. Pectoral one fourth 
longer than head. Ventrals shorter, reaching anal. Anal III 9; third spine longest, 
as long as third dorsal. Caudal truncate. Scales 30 ai lat. 1. = Yellowish, with a 


dark vertical streak below the eye, and a dark round spot on the side below the 
lateral line; vertical fins brownish, with some small, round, white spots on the soft 
dorsal and anal; pectorals white, ventrals blackish. 

Total length 90 millim. 

A single specimen from Paraguay. 


PLEURONECTID. 
9. SOLEA JENyNSU, Gthr. 
Paraguay. 
SILURID. 


10. Sorvsim Lima, Schn. 


Paraguay. 


FROM THE RIO PARAGUAY. 


11. PLatysroMaA oRBIGNIANUM, Val. 


Paraguay. 


12. HEMISORUBIM PLATYRHYNCHUS, C. & V. 


Paraguay. 


15. Scraps pictus, Mill. & Trosch. 
Matto Grosso. 


14. Pimenopvus corroipss, Bler. 


Descalvados, Matto Grosso. 


15. PIMELODUS ALBICANS,C. & V. 


Paraguay. 


16. PimELoDUS MacuLATUS, Lacép. 


Paraguay. 


17. PiweLopus LaTEeRisTRiGA, M. & T. 


Descalyados, Matto Grosso. 


18. PimeLopus eracius, C. & VY. 


Descalvados. 


19. Pimeiopus LaBrosus, Litk. 


Descalvados. 


20. PimeLopus PLATANUS, Gthr. 


Paraguay. 


21. Pimenopvus pati, Val. 


Paraguay. 


22. PIMELODUS PIRINAMPUS, Ag. 


Paraguay. 


23. AGENIOSUS BREVIFILIS, C. & V. 


Paraguay. 


E 2 


28 MR. G. A. BOULENGER ON FISHES 


24. EUANEMUS NIGRIPINNIS. (Plate IV. fig. 2. 
Euanemus nigripinnis, Boulenger, P. Z. 8. 1895, p. 524. 

Depth of body equal to length of head, 43 to 5 times in total length; width of head 
equal to length without snout; groove extending from end of snout to occipital bone ; 
eye as long as snout, 33 times in length of head, half interorbital width; maxillary 
barbel extending nearly to end of pectoral spine, mandibular barbels to a little beyond 
base of pectoral spine. Humeral process covered with skin, ¢ length of pectoral 
spine. Dorsal I 6; spine serrated behind, as jong as head. Pectoral spine a little 
shorter than head. Ventral as long as pectoral, reaching a little beyond origin of 
anal. Anal 36-37. Caudal deeply notched. Upper surface of head, back, and a 
lateral stripe dark grey, rest white; pectorals and ventrals deep black, white at the 
base; caudal black at the base. 

Total length 190 millim. 

Four specimens from Paraguay. 


25. AUCHENIPTERUS GALEaTUS, L. 


Matto Grosso. 


26. TRACHELYOPTERUS CoRIAcEUS, C. & V. 


Descalvados, Matto Grosso. 


27. Doras costatus, L. 


Descalvados, Matto Grosso. 


28. Doras macuxatus, Val. 


Paraguay. 


29. DoRAS WEDDELII, Cast. 


Paraguay. 


30. OXYDORAS EIGENMANNI. (Plate IV. fig. 3.) 
Oxydoras eigenmanni, Boulenger, P. Z. 8. 1895, p. 524. 


Both jaws with small teeth. Depth of body equal to length of head, 4 times in 
total length. Posterior nostril nearer the eye than to the anterior nostril. Diameter 
of eye 4 times in length of head, 1} in interorbital width, 13 in length of snout; 
bases of the six barbels united by the fold of the lower jaw; maxillary barbels 
branched, extending a little beyond base of pectoral spine, twice as long as man- 
dibulars. Gill-cleft extending to below posterior border of eye. Humeral process 


FROM THE RIO PARAGUAY. 29 


striated, half as long as pectoral spine. Pectoral spine a little longer than the 
head. Dorsal 16; spine a little shorter than the head, very strongly serrated in 
front, very feebly behind. Adipose fin a little shorter than base of anal. No shields 
between the dorsal fins. Lateral shields moderate, their vertical diameter equal to the 
eye, serrated behind, 26 or 27. Caudal bifurcate. Brown above, whitish beneath ; 
fins uniform or with small blackish spots. 

Total length 80 millim. 

Several specimens from Descalvados, Matto Grosso. 

This species is named after Prof. C. Eigenmann, the author of the excellent 
synopsis of South American Siluroid Fishes. It connects the sections Oxydoras and 
Rhinodoras as defined by that author, agreeing with the former in the serrature of the 
dorsal spine, with the latter in all other respects. 


51. RumoporaAs KNERI, Blkr. 


Paraguay. 


32. CALLICHTHYS ASPER, Q. & G. 


Paraguay. 


33. CALLICHTHYS PECTORALIS. (Plate IV. fig. 4.) 
Callichthys pectoralis, Boulenger, P. Z. 8. 1895, p. 525. 

Depth of body 34 to 34 times in total length. Head depressed, broader than deep, 
34 times in total length, without bristles on the sides; diameter of eye 6 times in 
length of head, 4 times in interorbital width ; suborbital bone narrow; occipital bone 
pointed in front, not reaching frontal fontanelle ; outer barbels half total length. 
Pectoral spine 2 length of head, covered with fine bristles, serrated on the inner side. 
Dorsal I 7 ; spine nearly } length of head. Anal 15. A pair of large pectoral plates, 
in contact anteriorly or narrowly separated, their inner borders diverging behind. 
Shields on body reaching to base of dorsal fins, 25 above arid 22 below lateral line ; 
3 or 4 pairs and 5 to 7 azygos shields between the two dorsal fins. Caudal rounded. 
Dark brown, with small blackish spots. 

Total length 85 millim. 

Several specimens from Monte Sociedad, Paraguayan Chaco. 

This species is most nearly allied to C. thoracatus, C. & V., and C. longifilis, C. & V., 
but differs in the larger pectoral plates, in the wide separation between the frontal 
fontanelle and the occipital bone, and in the smaller number of anal rays. 


534. CALLICHTHYS PALEATUS, Jen, 


Descalvados, Matto Grosso. 


50 MR, G. A. BOULENGER ON FISHES 


35, PLECOSTOMUS TERNETZI. (Plate V.) 
Plecostomus ternetzi, Boulenger, P. Z. S. 1895, p. 525. 

Head as long as broad, 5 times in total length, with three very obtuse keels ; snout 
rounded, entirely granulated ; diameter of eye 7 times in length of head, 4 times in 
length of snout, 24 times in interorbital width; barbel as long as diameter of eye ; 
44 teeth on each side in the upper jaw, and about as many in the lower jaw; inter- 
operculum with very small spines. Sides of throat, thorax, and middle of belly covered 
with small shields. Dorsal I 7; first ray as long as head, reaching adipose fin. 
Pectoral spine as long as head, strong, with small curved spines. Ventral I 4, first 
ray a little shorter than head, Anal I 4. Lower caudal ray very long, a little longer 
than head, twice as long as upper. Depth of caudal peduncle 23 in distance between 
anal and caudal fins. Posthumeral keel very obtuse and short, not extending beyond 
base of ventral, Scutes on body rough and spinulose, but not keeled; lat. 1. 25; 14 
scutes between anal and caudal! fins. Uniform olive above, white beneath. 

Total length 240 millim. 

A single specimen from Paraguay. 

This species appears to be nearest allied to P. francisci, Liitk., which is only known 
to me from the description. 


36. PLECOSTOMUS VERMICULARIS, Higenm. 


Paraguay. 
) 


537. PLeECOSTOMUS COCHLIODON, Kner, 


Paraguay. 


38. Cia&TosTomus GIGas. (Plate VL). 
Chetostumus yigas, Boulenger, P. Z. 8. 1895, p. 526. 

15 upper and 19 lower teeth on each side. Depth of body 43 times in total length, 
length of head 23 times. Head a little longer than broad, entirely rough with small 
spines ; snout broadly rounded; diameter of eye 11 times in length of head, 42 times 
in interorbital width, 6 times in length of snout; anterior border of orbit with enlarged 
spinules ; no postorbital groove; erectile preopercular spines very strong, the longest 
23 diameter of orbit; barbel a little longer than diameter of orbit. Throat and belly 
studded with small rough shields. Dorsal 110; all the rays rough with spinules, the 
first measuring 3 length of head. Pectoral spine as long as head, covered with small 
spines, the longest of which are hooked and nearly equal diameter of orbit. Ventral I 5, 
as long as first dorsal ray. Anal I 5. Caudal obliquely truncate, lower ray longest. 
Shields on body rough with ridges of strong spinules, 25 in a longitudinal series. 
Brown, densely covered all over with round black spots. 

Total length 630 millim. 

A single specimen from Paraguay. 


FROM THE RIO PARAGUAY. 5) 


This fish, the largest known of the genus, is possibly the adult of C. aculeatus, 
Perugia, in which, however, the ventral region is entirely naked. 


39. CH2ZTOSTOMUS cIRRHOSUS, Val. 


Paraguay; Descalvados, Matto Grosso. 


40. CHa&tTostomuUs LEUcosticTus, Gthr. 


Descalvados, Matto Grosso. 


41. HypopTopoMA GUENTHERI. 

Hypoptopoma guentheri, Boulenger, P. Z. S. 1895, p. 526. 

Head not narrowed behind the eyes, its width equal to length to posterior border of 
orbit, its length 2% to 3 times in total; diameter of orbit 5 to 53 times in length of 
head, 22 to 23 times in length of snout, 3 to 34 times in interorbital width; barbel 
very small; head-shields as in H. thoracatum. Dorsal I 6, first ray as long as head to 
upper angle of gill-cleft. Pectoral as long as first dorsal ray, extending as far as 
ventrals, not reaching anal. Caudal deeply notched, middle rays half as long as outer. 
Two pairs of large transverse pectoral shields, preceded by a transverse series of four 
small shields, the outer of which are in contact with the suborbital shields; three or 
four large ventral shields on each side, the anterior of which are usually separated by 
an azygos shield. Shields on body spinulose, but not keeled, 20 to 22 on each side ; 
3 shields between the occipital and the dorsal, 12 between the dorsal and the caudal. 
Olive; dorsal and caudal fins with black spots, which may form two curved bands on 
the lower lobe of the caudal. 

Total length 65 millim. 

Numerous specimens from Descalvados, Matto Grosso. 

The fish described and figured by Steindachner as H. thoracatum, Gthr., belongs to a 
distinct species, for which I have proposed the name /. steindachneri. His H. bilo- 
batum is distinct from Cope’s, which I regard as identical with H. thoracatwm, and 
is the same as that named by me H. guentheri. In fact, 3 species are confounded 
under H. thoracatum in Dr. and Mrs. Eigenmann’s Synopsis; they are distinguishable 


as follows :— 
I. Throat entirely naked in front of the two pairs of pectoral shields; 6 to 8 pairs of ventral 
shields separated by a series of azygos shields ; spine of adipose fin present; caudal deeply 
emarginate, middle rays half as long as outer. H. thoracatum, Gthr. 
TI. A transverse series of four shields in front of the pectoral shields ; 3 to7 pairs of ventral shields 
and a single azygos shield in front ; posterior dorsal spine usually absent. 
Pectoral spine not reaching origin of anal ; caudal deeply emarginate, middle rays half as long 
Aly ONLI Sse oF Ve ghee oc H. guentheri, Bigr. 
Pectoral spine reaching origin of 
than half as long as outer . 


anal; caudal less deeply emarginate, middle rays much more 
H. steindachneri, Blgr. 


32 MR. G. A. BOULENGER ON FISHES 


42. OTOCINCLUS AFFINIS, Stdr. 


Descalvados, Matto Grosso. 


45. LoRICARIA ROSTRATA, Spix. 


Paraguay. 


44. Loricaria parva. (Plate VIII. fig. 1.) 
Loricaria parva, Boulenger, P. Z. S. 1895, p. 527. 


Teeth small, well developed in both jaws. Head 13 as long as broad, 42 to 5 times 
in total length ; snout obtusely pointed, feebly projecting beyond the lip; head-shields 
with longitudinal, spinulose striz, without keels; diameter of orbit 54 to 6 times in 
length of head, 24 to 2% times in length of snout, 14 to 12 in interorbital width; a 
bread postorbital notch ; lower labial fold moderately large, papillose, feebly notched, 
with a fringe of obtuse papille. Dorsal I 7; first ray 1} to 14 as long as head, just 
above base of ventral. Pectoral I 6, as long as head or a little shorter, reaching 
beyond base of ventral. Ventral I 5, as long as pectoral, reaching beyond origin of 
ventral. Anal I 5, Outer caudal rays much produced, filiform, upper longest. 
Lateral scutes 26-28, with two spinose ridges approximating on the 13th or 14th; nuchal 
shields without keels; 16 or 17 scutes between dorsal and caudal, 14 or 15 between 
anal and caudal. Breast and belly shielded; pectoral shields numerous, polygonal, 
irregular; ventrals 7 to 9 transversely enlarged ones on each side and 3 series of small 
ones in the middle. All the shields spinulose and striated. Olive above, with ill- 
defined dark cross-bars; a dark streak on each side of the snout, from the tip to the 
eye; fins with dark spots. 

Total length 110 millim. 

Numerous specimens from Descalvados, Matto Grosso. 

This species is nearest related to L. filamentosa, Stdr. 


45. LORICARIA LABIALIS. (Plate VII. fig. 1.) 
Loricaria labialis, Boulenger, P. Z. S. 1895, p. 527. 


Teeth minute, rudimentary, in both jaws. Head 13 as long as broad, 44 to 43 times 
in total length; snout obtusely pointed, feebly projecting beyond the lip; head-shields 
rough with villose spinules, except on the edge of the end of the snout, without keels ; 
diameter of orbit 6 to 63 times in length of head, 3 times in length of snout, 13 in 
interorbital width; a broad postorbital notch; lower labial fold moderately large in 
females, very large and extending to the pectoral shields in males, without notch and 
without fringe. Dorsal I 7; first ray nearly as long as head, just above base of ventral. 
Pectoral I 6, as long as head to posterior border of orbit, not reaching base of ventral. 
Ventral I 5, as long as pectoral, reaching origin of anal. Anal 15. Upper caudal ray 


FROM THE RIO PARAGUAY. 33 


produced ina short filament. Lateral shields 30, with two spinose ridges approximating 
on the 21st or 22nd; nuchal shields without keels; 18 shields between dorsal and 
caudal, 16 between anal and caudal. Breast and belly shielded; pectoral shields 
numerous, polygonal, irregular; ventrals 4 to 6 transversely enlarged ones on each 
side, and one or two series of smaller ones in the middle. All the shields finely 
granulate and spinulose. Olive above; dorsal and caudal with small dark spots along 
the rays; pectorals and ventrals blackish. 

Total length 220 millim. 

Three specimens from Paraguay. 

Allied to L. nudirostris, Kner, and L. spixi, Stdr. 


46. LoricariA anus, Val. 
Paraguay. 


47. LoRICARIA MACULATA, Bl. 
Paraguay. 


48. LoRICARIA APELTOGASTER. (Plate VII. fig. 2.) 
Loricaria apeltogaster, Boulenger, P. Z. S. 1895, p. 528. 


A few slender teeth in both jaws. Head slightly longer than broad, 5 times in total 
length; snout acutely pointed, feebly projecting beyond the lip; head-shields strongly 
spinulose ; occipital shield with two closely approximated, parallel keels; diameter of 
orbit 74 to 8 times in length of head, 4 times in length of snout, 14 to 14 in interorbital 
width ; no postorbital notch ; lower labial fold rather larger with long fringes; barbel 
long. Dorsal 17; first ray a little longer than the head, just above base of ventral. 
Pectoral I 6; first ray more or less produced, at least as long as the head, reaching 
much beyond base of ventral. Ventral I 5; first ray produced, but shorter than 
pectoral, reaching much beyond origin of anal. AnalI5. Upper caudal ray produced 
in a very long filament. Lateral shields 31 or 2, with two spinose ridges uniting on 
the 17th to 20th; nuchal shields bicarinate; 21 or 22 shields between dorsal and 
caudal, 19 or 20 between anal and caudal. Breast and belly naked, or with small 
stellate shields ; a series of 6 to 8 small transverse shields may be present on each side 
of the belly. All the shields finely granulate and spinulose. Pale brown above, with 
three or four darker cross-bars on the body and two on the head; fins partially blackish. 

Total length 210 millim. 

Four specimens from Paraguay. 

Allied to ZL. nudiventris, C. & V., L. evansi, Blgr., and especially to LZ. macrodon, 
Kner. 


49. Loricaria LATA, Kigenm. 
Paraguay. 
VOL. x1v.—Part 11. No. 2.—October, 1896. FB 


4 Mk. G. A. BOULENGER ON FISHES 


50. LorrcariaA LAMINA, Gthr. 


Paraguay. 
51. ACESTRA OXYRHYNCHUS, Kner. 
Descalvados, Matto Grosso. 


52. BUNOCEPHALUS IHERINGII, Blgr. 
Descalvados, Matto Grosso, and Monte Sociedad, Paraguayan Chaco. 


53. TRICHOMYCTERUS BRASILIENSIS, Liitk. 


Descalvados, Matto Grosso, and North Paraguay. 


CHARACINID&. 
54. PYRRHULINA SEMIFASCIATA, Stdr. 


Descalvados, Matto Grosso, Monte Sociedad, Paraguayan Chaco. 


55. NANOSTOMUS LATERALIS, Blgr. 


Descalvados and Monte Sociedad. 


56. CuRIMATUS ALBURNUS, M & T. 


Descalvados. 


57. CURIMATUS RUTILOIDES, Kner. 


Paraguay. 

58. CURIMATUS LATIOR, Spix. 
Paraguay. 

59. HEMIODUS SEMITANIATUS, Kner. 
San Luis, Matto Grosso. 


60. HEMIoDUS MICROLEPIS, Kner. 
San Luis. 
61. PARoDON AFFINIS, Stdr. 


North Paraguay. 


62. LEPORINUS STRIATUS, Kner. 


Descalvados, Matto Grosso. 


63. Leporinus EQuEs, Stdr. 


Descalvados and San Luis, Matto Grosso. 


FROM THE RIO PARAGUAY. 35 


64. TmTRAGONOPTERUS MULTIRADIATUS, Stdr. 


Descalvados and San Luis. 


65. TETRAGONOPTERUS THRNETZI. (Plate VIII. fig. 2.) 
Tetragonopterus ternetzi, Boulenger, P. Z. S. 1895, p. 528. 
Depth of body 12 to 1# in total length, length of head 35 to 23 times. Diameter 


of eye 24 to 24 times in length of head, twice length of snout, equal to interorbital 


width; maxillary toothless, extending to below anterior border of eye. Dorsal 11, 
originating behind vertical of ventrals, longest rays as long as head. Anal 40-42, 
deepest anteriorly, longest rays a little shorter than head. Caudal bifurcate. Scales 


30-33 i: lateral line complete. Pale brownish, upper surface of head, dorsal, anal, 


and lower part of caudal region of body blackish ; two blackish vertical bands on upper 
half of body, the first behind the head, the second below the origin of the dorsal fin. 


Total length 45 millim. 
Several specimens from Descalvados, Matto Grosso. 


66. ‘TETRAGONOPTERUS ARGENTEUS, Cuv. 
Descalvados, Matto Grosso ; North Paraguay. 


67. TETRAGONOPTERUS LACcuSTRIS, Liitk. 


Descalvados and North Paraguay. 


68. [ETRAGONOPTERUS ABRAMIS, Jen. 


Descalvados. 


69. TETRAGONOPTERUS DICHROURUS, Kner. 


San Luis and Descalvados. 


70. TETRAGONOPTERUS AGASSIZH, Stdr. 
Descalyados and Monte Sociedad. 


71. TETRAGONOPTERUS LINEATUS, Perugia. 


San Luis. 


72. TerraGonoprerus ULREYI. (Plate VIII. fig. 3.) 
Tetragonopterus ulreyi, Boulenger, P. Z. 8. 1895, p. 529. 

Depth of body 23 to 22 in total length, length of head 53 to 35 times. Diameter 
of eye half length of head, twice length of snout, equal to interorbital width ; 


maxillary toothless, extending to below anterior third of eye. Dorsal 10, originating 
F2 


36 MR. G. A. BOULENGER ON FISHES 


behind base of ventrals, longest rays as long as head. Anal 23-25, anterior rays 
elongate, a little shorter than the head. Caudal bifurcate. Scales 32-33 = lateral 
line reduced to 8 or 9 scales. Yellowish, with a small black humeral spot, a black 
line along the middle of the side, and a black line along the base of the anal fin; 
dorsal blackish at the end. 

Total length 35 millim. 

Several specimens from Descalvados, Matto Grosso. 

Named after Prof. A. B. Ulrey, the author of a very useful key to the determination 
of the species of this genus. 


73. BRACHYCHALCINUS RETROSPINA, Blgr. 
Descalvados and San Luis, Matto Grosso. 


74. CHALCINUS NEMATURUS, Kner. 
Asuncion, Paraguay. 


75. CHALCINUS PARANENSIS, Gthr. 


Descalvados and Paraguay. 


76. GASTROPELECUS STELLATUS, Kner. 


Descalvados. 


77. ANACYRTUS MACROLEPIS, Kner. 


Descalvados. 


78. ANACYRTUS MICROLEPIS, Reinh. 


Descalvados and Paraguay. 


79. ANACYRTUS PROGNATHUS. (Plate VIII. fig. 4.) 
Anacyrtus prognathus, Boulenger, P. Z. S. 1895, p. 529. 

Depth of body 24 in total length, length of head 4 times. Snout strongly projecting 
beyond the mouth, squarely truncate, with two strong tooth-like spines pointing 
outwards and forwards, several smaller teeth on each side of the upper jaw and two on 
each side of the lower; diameter of eye + length of head, 14 in length of snout, 14 in 
interorbital width ; maxillary extending to below anterior third of eye; upper profile 
of head very concave. Dorsal 11, equally distant from eye and base of caudal, longest 
rays a little shorter than head. Anal 52. Caudal deeply forked. Yellowish. with a 
silvery lateral stripe; end of snout and a small spot at base of caudal blackish. 

Total length 125 millim. 

A single specimen from Paraguay. 


FROM THE RIO PARAGUAY. 


80. SALMINUS MAXILLOsUS, C. & V. 
Paraguay. 


81. XIPHORHAMPHUS FEROX, Gthr. 
Paraguay. 


82. SERRASALMO SERRULATUS, C. & V. 


Paraguay. 


83. SERRASALMO SPILOPLEURA, Kner. 


Descalvados and Paraguay. 


84. SERRASALMO HUMERALIS, C. & V. 


Descalvados and Paraguay. 


85. My eres asterias, M. & T. 


Descalvados. 


86. MyYLETES BRACHYPOMUS, Cuv. 


Paraguay. 


87. MYLETES DURIVENTRIS, Cuv. 


Paraguay. 


88. MyLEeTes HYPSAUCHEN, M. & T. 


Descalvados. 
SCOMBRESOCIDZ. 

89. BeLoNE Ta@yiaTa, Gthr. 

Descalvados. 

CLUPEIDZ. 
90. EneravLis oLipus, Gthr. 
Paraguay. 
GYMNOTIDz. 


91. STERNARCHUS ALBIFRONS, L. 


Descalvados, Matto Grosso. 


37 


38 MR. G. A. BOULENGER ON FISHES 


92. RHAMPHICHTHYS BLOCHII, Kaup. 


Paraguay. 


93. RHAMPHICHTHYS BREVIROSTRIS, Stdr. 


Descalvados, Matto Grosso. 
94. STERNOPYGUS cARAPUS, L. 


95. STERNOPYGUS VIRESCENS, Val. 


Descalvados. 


SYMBRANCHID. 


96. SYMBRANCHUS MARMORATUS, Bl. 


Descalvados, Matto Grosso. 


TRYGONID&. 


97. 'CaNIURA DUMERILU, Casteln. 


Paraguay. 


EXPLANATION OF THE PLATES. 


PLATE III. 


Plagioscion ternetzi, 4 nat. size, with upper view of head: p. 29. 


PLATE LV; 
Fig. 1. Geophagus duodecimspinosus: p. 26. 
Fig. 2. Euanemus nigripinnis: p. 28. 
2a. Upper view of head. 
Fig. 8. Oxydoras eigenmanni: p. 28. 
Fig. 4. Callichthys pectoralis: p. 29. 
4a. Lower view of head and pectoral region. 


‘PLATE V. 


Plecostomus ternetzi, $ nat. size, with upper and lower views of head: 


FROM THE RIO PARAGUAY. 


PLATE VI. 


Chetostomus gigas, + nat. size, with upper and lower views of head (4): p. 30. 


PLATE VIL. 


Fig. 1. Loricaria labialis: p. 32. 

1a, upper, 1 4, lower view of head and anterior part of body. 
Fig. 2. Loricaria apeltogaster: p. 33. 

2 a. upper, 2 4, lower view of head and anterior part of body. 


PLATE VIII. 
Fig. 1. Loricaria parva, with enlarged upper (a) and lower (0) views of head: p. 32. 
Fig. 2. Tetragonopterus ternetzi: p. 35. 


1 
2 

Fig. 3. Tetragonopterus ulreyi: p. 35. 
4. Anacyrtus prognathus: p. 36. 


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3.TETRAGONOPTERUS ULREY1.4.ANACYRTUS PROGNATHUS. 


J Green del. et Lith. 


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ILI. On a Collection of Fishes from the Rio Paraguay. By G. A. BouLENGER, PRS. 
(Plagee LT SVE ie secre, eg a OY ee agg tie Miter tine ee bee 


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[ 41 ] 


IV. Contributions to the Comparative Anatomy and Histology of the Suprarenal 
Capsules—The Suprarenal Bodies in Fishes, and their Relation to the so-called 


Head-Kidney. 


Assistant Lecturer on Histology, Mason College, Birmingham}. 


Received September 28th, 1895, read November 19th, 1895. 


[Prates 1X.-XIV.] 


ee IMtOAUCLOLYA crore .sicic oc = 0.0\0)= 1015 «estos 
II. Historical 
III. Classified List of Species examined .... 
Vm GMOs A TIAL OMY 5) << et0y010r0/0Vei=ie1) «ysl eieiate 


Ce i ce i id 


1. Elasmobranchii 
2. Ganoidei 
3. Teleostei 
APADUPNOL jc taws oils vte.e Aetbcate ateleie os 
VY. Histology 


1. Elasmobranchit 


ee ee ay 


2. Ganoidei 

3. Teleostei 

4, Dipnoi 

VI. On the Relation of the Suprarenal Bodies 
to the Head-Kidney............ 

1. Historical 


i i oe ee 


ConTENTs. 
41 2, The Nature of the Intertubular 
43 Material in the Kidneys .... 
50 3. Histology of the Head-Kidney 
51 4. Function of the Lymphatic Head- 
51 NGGNGi Mae Rneaeonoe on SoC 
58 5. Relation of the Suprarenals to the 
56 Head-Kidney...........-.. 
63. VII. Summary and Conclusions............ 
64 A) GrosseAnatomy? 2) crt) neater = 
64 IBS Histology;.i-yojete raster severe deus 
69 (Ch Head Kidney svteterere) islets iaieta etetsiere 
7 VAUNIS Linen? SomcanoadocdsounheoSoc 
72 A. On the Suprarenal Bodies ...... 
B. Head-Kidney, &. ...........- 
73 Explanation of the Plates,................. 
73 


I. IytRopvuctory. 


By Swate Vincent, M.B.Lond., Demonstrator of Physiology and 


Page 


I HAVE been led to investigate the suprarenal capsules in Fishes because their 


importance in Man and Mammals generally is no longer a matter of doubt. 


Their 


function is not yet completely known, but we shall, I imagine, not go far wrong if we 
state that in Mammals the suprarenal bodies are secreting-glands, and that they pour 
some material into the blood which is essential for the maintenance of the normal tone 
of the muscles throughout the body. But this function appears to be confined to the 
medullary portion of the gland, and, since the two portions are so strikingly different 


1 Towards the expenses of this research a grant has been made by the British Medical Association, at the 
recommendation of the Scientific Grants Committee of the Association. 


VOL. X1v.—PaRT mI. No. 1.—A pril, 1897. 


G 


42 MR. SWALE VINCENT ON THE 


in structure and origin, and being inclined to the opinion that these bodies are 
homologous throughout Vertebrates, I considered that a careful enquiry into their 
anatomy and histology in the lowest class of Vertebrates could not fail to be of some 
value as a contribution to our knowledge of their exact distribution and relationships. 

Moreover, I felt that there was definite need for such an enquiry and clear 
statements of its results. Although some good work has been done upon the subject, 
it is embodied only in scattered, often antiquated and inaccessible memoirs; the 
text-books scarcely refer to the subject, and various writers who mention the supra- 
renals make serious errors as to points of fact. 

Some authors have attempted to establish the view that (in Teleosts), where 
suprarenals are present, there is no lymphatic head-kidney, and vice versd; in other 
words, that the suprarenals are interchangeable with the head-kidney, that where one 
exists the other does not. I shall be able to show that this idea is quite groundless. 

Again, certain writers appear to think that suprarenals are comparatively rare 
objects among the Teleosts. It will be seen that, so far as my observations have gone, 
they are universally present in this order. I will not enlarge further on these points 
here, as they will be dealt with in detail in the body of the paper. 

As for methods of work, these have been very various. I have examined fifty-five 
species in all, and many individuals of several species. Most of these have been 
perfectly fresh, indeed all but nine, which were preserved specimens. 

In each case I have observed 77 situ the position and relations of the suprarenals 
and the head-kidney. After careful removal, both these have been examined 
microscopically. Numerous sections of various parts of different kidneys have also 
been made and examined. 

Some preparations were made quite fresh, 7. e. sections cut with Swift’s “ Ether 
Freezing Microtome.” Others have been hardened in alcohol or Miiller’s fluid, 
stained in buik, imbedded in paraffin, and cut with the “ Rocking Microtome.” The 
precise method I have used to the greatest extent is the following :— 

The material is hardened in Miiller’s fluid in the usual way for about six weeks. 
[In some cases the process has been hastened by keeping the tissue at a temperature of 
about 36° C.] It is then double-stained in bulk with Ehrlich’s hematoxylin and 
eosin, and imbedded in paraffin. 

In working at the naked-eye anatomy in the Elasmobranchii, I have made use of 
both the chromic-acid method of Semper and the osmic-acid method of Chevrel. 

I have made a special point of obtaining, in all cases where this was possible, 
perfectly fresh specimens, and no results, with one or two exceptions, have been 
considered worth recording, especially in a histological direction, unless the tissues 
were practically living at the time of dissection. 

I was convinced that what was particularly needed was a series of accurate drawings 
of the suprarenal bodies and their relation to the kidney. Consequently I have given 


SUPRARENAL BODIES IN FISHES. 43 


an outline sketch of the kidney from the ventral surface in nearly every species 
examined, putting in the suprarenals in plain lines when on the ventral surface, in 
dotted lines when on the spinal surface. Such a series of drawings I cannot find and 
do not believe to exist. In the Elasmobranchii the only drawing I am acquainted 
with is that in Semper’s paper, and this, I was sure, might easily be improved upon. 

Many parts of the paper are not nearly so complete as I should have wished. 
Indeed, in many respects I wish the paper to be considered as merely preliminary to 
future work on the same subject. This applies particularly to the histological part. The 
head-kidney I hope to make the subject of an exhaustive research at some future time. 

I have not touched upon the subject in the Cyclostomata, having so far only been 
able to get specimens of the small river-lamprey, but I hope to deal with their blood- 
vascular glands in a separate contribution’. Further, I have not made any investigation 
of the development of the suprarenal bodies. 

I am deeply indebted to Prof. T. W. Bridge, M.A., D.Sc., of Mason College, 
Birmingham, for his kindly and repeated advice on many subjects connected with this 
research, and for the generous manner in which he has placed specimens at my 
disposal; to Prof. F. J. Allen, M.A., M.D., for much important assistance; to 
Mr. Walter E. Collinge, F.Z.S., for help on many points; and to Prof. W. N. Parker 
for specimens of Protopterus and the use of some of his slides. I must also express 
my thanks to Prof. G. B. Howes for many suggestions, bibliographical and otherwise. 

I take this opportunity of expressing my thanks to the Research Committee of the 
British Medical Association for a grant in aid of these investigations. 


II. Hisroricat. 


The literature of this subject is so very scattered and often so inaccessible, and I 
have become acquainted with some papers touching upon it in such accidental ways, 
that I fear my account may not be complete. However, I trust that most of the 
important papers dealing with the general subject have fallen into my hands; at the 
same time it is possible that accounts of the suprarenals and discussions of their 
morphology in monographs may have escaped me. ‘Then, too, some text-books may 
have accounts which I have overlooked; but, from the exceedingly unsatisfactory 
account given by Eberth in Stricker, I did not consider it would be profitable to 
explore these very thoroughly. 


1 Since the above was written I have, in conjunction with Mr. Walter E. Collinge, F.Z.S., investigated the 
subject in the Cyclostomata, with the result that nothing in the way of suprarenals can be made out. (Anat. 
Anz. Bd. xii. Nr. 9 & 10, 1896.) 

[More recently Pettit (Thése, Paris, 1896) has been unable (p. 86) to ascertain definitely whether supra- 
renals are present in the Cyclostomata or not.—S, Y., 10. 1. 97.] 

G2 


44 MR. SWALE VINCENT ON THE 


The earliest account of the suprarenal bodies in fishes appears to be that of Retzius. 
His description, written in 1819, has reference only to cartilaginous fishes. 

Retzius (18) pointed out as suprarenals certain structures in Squalus glaucus, 
S. acanthias, Raja clavata, R. fullonica, and R. batis. From his description it 
is evident that he refers to what is now usually termed the “ interrenal body.” 
He considered this organ to be the suprarenal on account of its resemblance in 
texture to the suprarenals of birds !. 

The suprarenals in Teleostei appear to have been discovered by Stannius (20) in 
1839. 

In 1843 J. Miiller (16) described in Myxinoids a clustered gland without a duct on 
each side of the cardia, which he at first considered to be a suprarenal, but afterwards 
thought to be thymus. In Petromyzon he found instead of this gland “certain white 
plugs with which the trunks of the posterior veins of the body are beset.” These 
structures had been previously described by Rathke 2. 

Three years later, in 1846, Stannius and Kcker threw considerable light on the 
subject. Stannius (21) describes the suprarenals as existing both in the higher 
cartilaginous and in the bony fishes. His account of the suprarenals in Elasmobranchs 
obviously applies only to the interrenal. The segmentally-arranged bodies were not yet 
discovered. He gives also in this first edition a fairly good account of the general 
position of the suprarenals in several Teleosts and in the Sturgeon. In addition this 
observer questions the right of Miiller’s “clustered gland ” to rank as a suprarenal. 

Ecker (6), in addition to verifying the results of the above observer in regard to the 
gross anatomy of the suprarenals, gives an account of their minute anatomy, which was 
for many years the common store of information on this subject. 

In 1851 Hyrtl (10) mentions some facts about the suprarenals in several Teleosts. 
He examined 222 species. He gives first a short and very imperfect account of their 
general position and appearances, and then mentions them very briefly under the 
special heading of the species. But, although he states in his introductory general 
description that suprarenals were found in almost all the fish he examined, yet I find, 
on looking through the paper, that they are only specifically noted in 28 species. In 
some two or three cases he states that he could not find them. In all the rest he never 
mentions them. He definitely found them in :— 


Uranoscopus scaber. Silurus glanis. 


Cottus quadricornis. Schilbe mystus. 


Diagramma punctatum. Pimelodus bayard. 


Saurus lacerta. 


| 
. . | . 
Lophius piscatorius. | Salmo fario. 
Chironectes punctatus. | 


Normyrus oxyrhynchus. Clupea nilotica. 


* See also Nagel, Miiller’s Archiv, 1836. 
* See Collinge and Vincent (Joc. cit.), also Pettit (loc. cit.).—S. V., 10. 1. 97. 


SUPRARENAL BODIES IN FISHES. 45 


Elops salmoneus. Lota vulgaris. 

Gadus barbatus. Phycis blennoides. 
Cobitis fossilis. Platessa passer. 
Acanthopsis tenia. Rhombus nuchus. 

Esox lucius. Echeneis remora. 
Esococtus exiliens. Ammodytes tobianus. 
Belone vulgaris. Symbranchus marmoratus. 
Motella mustela. Diodon novemmaculatum. 


No doubt a large proportion of Hyrtl’s specimens were not fresh but preserved, and 
this would often prevent the discovery of the suprarenal bodies. 

In the same year Leydig (13) described the suprarenals of Chimera monstrosa thus: 
“Each suprarenal forms a narrow streak #’ long, of an ochre-yellow colour. The 
posterior end is somewhat thicker and more rounded. ‘They lie on the inner border of the 
kidney.” This, of course, is the “ interrenal.” In Elasmobranchs we find no mention 
yet of the paired segmental bodies on the branches of the aorta, but only a body “ in 
Form eines schmales ockergelben Streifens an der Riickseite der Nieren” (Stannius). 

After this period we come to researches of much more importance, in reference at 
any rate to the subject in Elasmobranchs, and it will be necessary to review it in some 
detail. 

In 1852 Leydig published his famous ‘Rochen und Haie’ (11). In this work he 
insists that the “so-called axillary hearts and their continuations on the sympathetic 
ganglia are to be considered as the real suprarenals of Cartilaginous Fishes, and not, as 
heretofore reckoned, the yellow stripes and bodies behind the kidneys.” He bases 
this view on the fact that the paired bodies resemble the suprarenals in Mammalia in 
“consisting of closed bladders filled with cells,” also in their abundant vascular and 
nervous supply. On the other hand, he says “ the yellow stripes” consist simply of 
“masses of fat-globules and bright nuclei.’ He emphasizes the close connection 
between the suprarenals and the sympathetic nervous system. “As the pituitary 
body is an integral part of the brain, so the suprarenals are part of the sympathetic.” 
In the same work Leydig expresses his opinion that the organs previously considered 
as suprarenals are “ fat-glands”” analogous to those known in the Amphibia. Referring 
to the Cyclostomes this author definitely states his belief that the “clustered gland” 
of Miiller and the “white plugs” of Rathke and Miiller are analogous to the 
suprarenals. 

About this time too (1852) appeared Frey’s article on the Suprarenals in Todd’s 
‘Cyclopedia’ (7). It was obviously written before the above researches of Leydig, so 
that the only suprarenals mentioned in Elasmobranchs are the bodies now termed 
‘interrenal.” This writer gives a very good account of the anatomy and histology, 
mostly after Ecker, and in the part relating to fishes, at any rate, there seems little that 


may be considered original. 
In the next year (1853) Leydig published his ‘ Fische und Reptilien’ (12). In this 


46 MR. SWALE VINCENT ON THE 


work he gives a description of the suprarenals in the Sturgeon, and finds them to be 
composed of “an aggregation of fat-globules.” He states that they are always placed 
on the walls of the blood-vessels. But he changes his view entirely on the subject of 
the suprarenals in cartilaginous fishes. He now considers that both the paired bodies 
in connection with the sympathetic ganglia and the “ ochre-yellow stripes” behind the 
kidney belong to the same system and constitute the suprarenal capsules in this order 
of fishes. 

Stannius, in a later edition (22), seems to have recognized both kinds of bodies. 
Balfour says his description is ‘‘not quite intelligible.” This I consider to be quite a 
euphemism. I have found the description totally incomprehensible as applied to the 
Elasmobranchs. On the other hand he gives a very excellent account of the supra- 
renals in Teleosts. He states that in the Pike these bodies have been found studded 
in the kidney from the middle to the tail-end, and believes them to be absent in Clupea 
harengus and Ammodytes tobianus. 

The next important step in advance was made by Semper (1g) in 1875, who 
emphasizes the segmental arrangement of the suprarenals, and believes them to be the 
same kind of structure as the interrenal. In fact he appears to believe that there 
exists a direct anatomical continuity between them. “Hier freilich gehen sie bei 
manchen Formen (Rochen, Chimera, Scymnus, Acanthias, Mustelus, etc.), also 
wahrscheinlich wohl bei den meisten Plagiostomen in einen bald weissen, bald 
hell- oder dunkel-gelben Kérper iiber, welcher, zwischen den Enden der beiden Nieren 
liegend, dicht an der einfachen Caudalvene sitzt.” 

F. M. Balfour (1) in 1878 has also dealt with this subject with considerable care in 
his monograph on Elasmobranch Fishes. He gives an account of the history up to 
date, and it is to him we owe the term “interrenal” as applied to the unpaired body 
in Scyllium. He expresses his opinion that there is very probably “a third kind of 
body in connection with the kidney,” and regrets he could not settle the point with 
fresh specimens. He refers to Stannius’s description as possibly indicating a third 
structure; but, so far as I could understand this author, he seems to allude to broken-off 
or scattered portions of the interrenal, which, as we shall see later on, are frequently 
found. The “lymphoid masses” which Balfour mentions in connection with the 
larger vessels of the kidney do not appear to me to be of any importance in connection 
with this subject, as lymphoid tissue is very common in all fishes both in and 
surrounding the renal organs. 

Balfour's researches were both anatomical and developmental. With the latter I 
shall not concern myself, but the former must be dealt with in some detail. He 
describes the general anatomical relations of both “supra-” and “ inter-”renals, and 
then gives an account of their histology, which suffers from the fact that he had only 
been able to obtain specimens preserved in chromic acid. He lays great stress on the 
relations between the paired suprarenals and the sympathetic nervous system, but 
states that there is a “much smaller ganglionic development” in connection with the 


SUPRARENAL BODIES IN FISHES. AT 


posterior bodies than with the anterior. The typical suprarenal structure, he says, is 
best exhibited in a posterior one. “ Externally there is present a fibrous capsule which 
sends in the septa, imperfectly dividing up the body into a series of alveoli or lobes. 
Penetrating and following the septa there is a rich capillary network. ‘The parenchyma 
of the body itself exhibits a well-marked distinction, in the majority of instances, into 
a cortical and medullary substance. The cortical substance is formed of rather irregular 
columnar cells, for the most part one row deep, arranged round the periphery of the 
body. The cells measure on an average ‘03 mm. in their longest diameter. The 
medullary substance is more or less distinctly divided into alveoli, and is formed of 
irregularly polygonal cells... . &c.” 

Balfour mentions also that the protoplasm of both sorts of cells has a yellowish 
tinge, and that the suprarenals are more or less surrounded by lymphoid tissue. He 
gives a drawing of these appearances in pl. xviii. fig. 6. As for the connection between 
the sympathetic ganglia and the suprarenal bodies, he says :—“ In the case of one of 
the posterior bodies, a small ganglion is generally found attached to both ends of the 
body, and invested in the same sheath; in addition to this a certain number of 
ganglion-cells (very conspicuous by their size and other characters) are to be found 
scattered through the body. In the anterior suprarenal bodies the development of 
ganglion-cells is very much greater. If a section is taken through the region where 
the large sympathetic ganglion is attached to the body, one half of the section is 
composed mainly of sympathetic ganglion-cells and nerve-fibres, and the other of supra- 
renal tissue, but the former spread in considerable numbers into the latter.” At one 
point (n in pl. xviii. fig. 7) a nerve is shown entering. He states that the ganglion and 
nerves are so intimately united with the suprarenal body as not to be separable from it. 

Balfour leaves it an open question whether there are cells of an intermediate character 
between the ganglion cells and the cells of the suprarenal body. Then follow the 
developmental researches, the general conclusion of which is that the paired “ supra- 
renal” bodies develop as parts of the sympathetic nervous system. 

Passing on to the interrenal body, this author gives a sketch of its general anatomy, 
and differs from Semper as to the continuity of the two kinds of structure. It will be 
seen that my own observations agree on this point with Balfour. With regard to the 
histology of the interrenal, he gives a drawing (pl. xvii. fig. 8), and describes it thus: 
“Tt is invested by a fairly thick tunica propria, which sends in septa, dividing it into 
rather well-marked lobules or alveoli. These are filled with polygonal cells, which 
form the true parenchyma of the body. These cells are in my hardened specimens not 
conspicuous by the number of oil-globules. . . . They are rather granular in appear- 
ance, and are mainly peculiar from the somewhat large size of the nucleus. . . .” 

Balfour notes as differences between the “ supra-” and “ inter-”renals the distinction 
into cortex and medulla in the former, and the large size of the nuclei in the cells of 
the latter. From the developmental standpoint he concludes that the interrenal does 
not belong to the same system as the suprarenals, and he proceeds to enquire with which 


48 MR. SWALE VINCENT ON THE 


(if with either) of these two bodies the suprarenal bodies of the higher vertebrates are 
homologous. He does not decide the question definitely, but inclines to the view that 
the paired bodies of Scyllium are homologous with the suprarenals of Mammalia !. 

The only recent account of the suprarenals in a Teleost which I have found is that 
of M‘Kenzie (15), written in 1884, who gives a description of the suprarenals of 
Amiurus catus, with drawings of their histological appearances. They are, according 
to M‘Kenzie, sometimes entirely concealed by the kidney-substance. This is note- 
worthy as being decidedly rare in this order. He describes alveoli containing large 
and small granular cells, the longest of them reaching from wall to wall of an alveolus. 
These two kinds of cells bear no constant relation to each other. This author does 
not believe that the bodies have anything to do with the elaboration of the blood, 
and is opposed to Weldon’s view that the suprarenals are metamorphosed parts of 
the kidney just as the head-kidney is. This fish, he states, has a well-developed 
(lymphatic?) head-kidney in addition to the undoubted suprarenals. The interest and 
importance of this will be seen later on. 

Chevrel (3) appears to be the last author who has written upon this subject”. He, 
like Leydig, Semper, and Balfour, has discussed the subject in Elasmobranchs, and 
chiefly from the standpoint of the sympathetic nervous system. He has picked out the 
nerve-fibrils and suprarenal bodies with osmic acid. In this excellent memoir he gives 
a very good historical account, and carefully reviews Balfour’s work on the subject. 
Chevrel states that the interrenal body is situated between the inferior surface of the 
dorsal aorta and the superior surface of what he calls the “interrenal vein.” He 
describes also small detached parts of the interrenal on the superior and posterior 
parts of the kidneys. 

In the case of the paired bodies, Chevrel denies Balfour’s division into cortex and 
medulla, and explains the appearance obtained by the latter observer as due to the 
action of reagents. As will be seen below, this view is entirely corroborated by my 
own researches. Chevrel got no appearances anything like Balfour described. He 
says, ‘On ne voit ni cellules columnaires a Ja périphérie, ni cellules polygonales au 
centre; il n’y a que des apparences. Et ces apparences sont dues vraisemblablement 


* [R. Fusari (“ Contribuzione allo Studio dello Sviluppo delle Capsule surrenali e del Simpatico nel Pollo e 
nei Mammiferi,” Arch. per le Scienze med., Torino, 1892, vol. xvi. no. 14, pp. 249-301, tay. iv.-vii.), from 
investigations on mammals, maintains that the interrenal body is not homologous with any part of the supra- 
renal capsule, but with a certain adipose tissue found round the suprarenals in some mammals. 

On the other hand, M. Braun (“ Bau und Entwickelung der Nebennieren bei Reptilien,” Arbeit. a. d. zool.- 
zoot. Inst. Wiirz., Band y. pp. 1-30, Taf. i., ji., 1882), R. Semon (“* Studien iiber den Bauplan des Urogenital- 
systems der Wirbelthiere (Ichtiophis), Jena. Zeitschr., 1891, Band xxvi. pp. 89-203), and F. van Wyhe (“ Ueber 
Mesodermsegmente des Rumpfes und die Entwickelung des Excretionsystems bei Selachiern,” Arch. f. mikr. 
Anat, Band xxxiii. 1889, pp. 461-516, Taf. xxx.—xxxii.) favour Balfour’s views. 

Valenti (Atti della Soc, Toscana di Scienze nat. 1889, Pisa, vol. x. tay. x) believed the suprarenal capsule to 
be a rudimentary organ.—-S. V., 10. 1. 97.] * See footnote to p, 49. 


SUPRARENAL BODIES IN FISHES. 49 


aux contours des mailles de la trame conjonctive des corps. Les dissociations nous ont 
également donné des résultats négatifs.” He doubts their analogy to the suprarenal 
bodies of Mammalia, and describes their structure thus :—‘ De la fine membrane qui les 
enveloppe partent des filets qui vont former dans son intérieur, de concert avec ceux 
qui naissent de l’anneau conjonctif entourant Vartériole, une sorte de réticulum trés 
compliqué, 4 mailles inégales, dans lesquelles se trouvent un nombre considérable de 
noyaux ovales. Chacun de ces noyaux, pourvu de plusieurs petits corps trés réfringents, 
parait simplement plongé dans le protoplasme granuleux; on pourrait peut-étre 
supposer qu'il appartient 4 une cellule sans membrane d’enveloppe dont le proto- 
plasme granuleux se fusionne avec celui des cellules voisines. De cette fusion 
résulterait une masse protoplasmique unique contenant tous les noyaux et remplissant 
Jes mailles du réticulum. Enfin des ganglions et des cellules sympathiques se trouvent 
assez souvent plongés dans le parenchyme du corps, et, 4 sa surface, circulent sans 
pénetrer, du moins en apparence, dans sa substance, et sans émettre de rameaux, des 
filets nerveux appartenant également au systéme sympathique.” 

Passing on to the interrenal, Chevrel says, “A Vétat frais, les éléments propres de ce 
corps disparaissent sous une couche de globules clairs, légérement teintés, qui lui 
donnent une couleur jaunatre particuliére.” Leydig thought these globules to be fat. 
Balfour, on the contrary, thought they were not fat. Chevrel is of the latter opinion, 
since ether does not dissolve them, nor osmic acid blacken them to any extent. He 
proceeds to describe their microscopical structure as being much the same as that of the 
suprarenals. ‘The chief differences, according to him, are—(1) Inequality in size of 
nuclei (10 w in “inter,” 9 in “supra”). (2) Absence in the suprarenal of the clear 
globules which the interrenal contains in abundance. (3) A distinct division some- 
times, in the case of the interrenal, into “capsules ou vésicules nettement marquee.” 
(4) A less abundant vascular supply in the interrenal. 

Chevrel does not think there is any very direct connection between the interrenal 
and the suprarenals, and notes that the “interrenal” has no relations with the 
sympathetic. He does not decide whether the interrenal body is of a different order 
of structure from the suprarenals. As to the connection between the suprarenals and 
the small sympathetic ganglia, he states, ‘‘ This connection in the case of the smaller 
bodies is not so frequent as Leydig, Semper, and Balfour imagined.” 

Chevrel has also more recently (4) given an account of the sympathetic system in 
the Sturgeon. He gives a drawing of the relations of the suprarenal bodies to 
the sympathetic nervous system, and also a very unsatisfactory representation of the 
microscopical structure of the ganoid suprarenal }. 

1 [Since this was written, the papers of V. Diamare (“ Ricerche intorno all’ Organo interrenale degli Elasmo- 
branchi col ai Corpuscoli di Stannius dei Teleostei” etc., Mem. matem.-fis. ser. 3, t. x. 3 tay., 1896) and 


Pettit (Thése, ‘ Recherches sur les capsules surrénales,’ Paris, 1896) have appeared. 
These will be briefly referred to again, as occasion requires, in later footnotes.—S. V., 10. 1. 97.] 


VoL. XIv.—Part 11. No. 2.—April, 1897. H 


MR. SWALE VINCENT ON THE 


III. Crassiriep List oF SPECIES EXAMINED. 
j. ELASMOBRANCHII. 
A. SELACHOIDEI. 
Seyllidee: Seyllium canicula. 
53 catulus. 


Spinacide: Acanthias vulyaris. 
Rhinide : Rhina squatina. 


B. Bavorpet. 
Rajide: Raja batis. 
5, clavata. 
» maculata. 


ii HOLOCEPHALA. 


Chimeeride: Chimera monstrosa. 


iii. GANOIDEI. 

A. CHonprostrEr, 

Spatularide : Polyodon folium. 
Acipenseride : Acipenser sturio. 
iv. TELEOSTEI. 
A. PLecroGNaTHt. 

a. Sclerodermi. 


Balistide: Balistes maculatus. 


6, Gymnodontes. 
Molide: Orthagoriscus mola. 
Tetrodontide: Tetrodon nigropunctatus. 


B. Puysostomt. 
Mureenidee: Anguiila anguilla. 
Conger conger. 
Clupeidze : Clupea harengus. 
Esocide : Esox lucius. 
Salmonide : Salmo salar. 
bye PNULCDs 
Cyprinidee : Leuciscus rutilis. 
x” cephalus. 
35 vulgaris. 


C. ANACANTHINI. 
Gadidee: Gadus morrhua. 
»  @glefinus. 
»  merlangus. 
3  ©vtrens. 


SUPRARENAL BODIES IN FISHES. 5 


Merluccius vulgaris. 

Motella tricirrhata. 

Molva vulgaris. 
Pleuronectidz : Hippoglossus vulgaris. 


Pleuronectes flesus. 
5 limanda. 
= platessa. 


Solea vulgaris. 
Hippoglossoides limandoides. 
Zeugopterus. 

Rhombus levis. 


D, Acanruorrert. 
a. Pharyngognathi. 
Coris pulcherrima. 
6. Acanthopterygii. 
Percide: Gastrosteus spinachia. 
Perca fluviatilis. 
Labraz lupus. 
Mullide: Mudllus barbatus. 
Sparide: Pagellus centrodontus. 
Cantharus griseus. 
Triglide: Cottus gobio. 
Trigla pini. 
»  lyra. 
Scomberide : Scomber scomber. 
Zeus faber. 
Blennidz : Anarrhichas lupus. 
Pediculati: Lophius piscatorius. 
c. Acanthopterygii mugiliformes, 
Mugilide : Mugil capito. 
d. Acanthopterygii gobiiformes. 
Discoboli: Cyclopterus lumpus. - 


v. DIPNOI. 


Lepidosiren paradoxa (L. appendiculata, Ehlers *). 
Protopterus annectens, 


IV. Gross ANATOMY. 
1. ELASMOBRANCHIL. 


In this order there are two distinct sets of bodies which have borne the name of 
“suprarenals.” One of these is a more or less rod-shaped structure, unpaired in some 


‘ [See, however, Lankester, “ Ou Lepidosiren and Protopterus,” Trans. Zool. Soc. vol. xiy. part i., April 1896. 
eae iV. 10, 1. 97.] 
H2 


52 MR. SWALE VINCENT ON THE 


genera, as Scyllium, paired in others, as Raja, which lies in the region of the hinder 
end of the kidney—in the middle line, as a rule, in the first case; on the inner and 
hinder border of the kidney in the second (PI. IX. figs. 1, 2, 5, & 4, Pl. X. figs. 4, 6, 
& 7, i7.). This structure was called by Balfour the ‘ interrenal” body, and has since 
usually borne this name. ‘The other set of structures is a segmentally arranged series 
of paired bodies, placed on the intercostal or parietal branches of the aorta, and 
extending in many cases throughout the whole length of the abdominal cavity (Pl. LX. 
figs. 1-4; Pl. X. figs. 5, 6, & 7, aw.h., & s.r.). 


a. SELACHOIDEI and BATOIDEIL. 


I have examined the following seven species, and in some cases several individuals of 
the same species :— 


Scyllium canicula. Raja batis. 
xs catulus. », clavata, 
Acanthias vulgaris. »» maculata, 


Rhina squatina. 

The Seqmentally arranged Bodies——These are disposed in much the same manner in 
all Plagiostomes. Their general arrangement has been already weil described by 
Leydig, Semper, and Balfour; and Chevrel has given a description of their anatomical 
connections with the sympathetic. It is therefore only necessary for me to give a 
brief account of their usual anatomy, emphasizing such points as may seem to need it. 

These bodies are placed for the most part in a very definitely segmental manner 
(Pl. IX. fig. 2). The anterior pair are always elongated and equal in size to several of 
the following bodies (Pls. IX. & X. figs. 1-7, ax.h.); as a matter of fact they seem to 
correspond usually to three or four segments. ‘They are arranged on branches of the 
aorta, the intercostal arterioles (Pl. IX. figs. 1 & 4; Pl. X. figs. 5 & 6), and extend on 
each side of the vertebral column from the front part of the sinus of Monro to a variable 
distance posterior to the origin of the posterior cardinal veins. The anterior pair and 
some of those bodies which immediately follow these on either side are placed in the 
cavity of the sinus, and therefore during life are bathed in its blood (Pls. IX. & X. 
figs. 1-7, ax.h., s.7.). They usually more or less completely surround the artery with 
their substance, and with the aid of a lens one can nearly always see the artery, vein, 
and a fine nerve distributed to each segmental body. ‘The first pair are placed on or 
near the axillary arteries, hence their old name “Axillary Hearts.” An arteriole usually 
communicates between the axillary and the succeeding branch of the aorta, and runs 
through the whole length of the body (Pl. IX. figs. 2a & 4). 

In the anterior part of the abdominal cavity these bodies have no relation to the 
kidney-substance, but in the region of this organ they become more or less imbedded 
in it, Where the renal substance is distinctly segmented they are often placed deep 
down in the grooves which separate the segmental portions, and are often in this way 
almost completely hidden. ‘They usually overlap the region of the interrenal, and often 


SUPRARENAL BODIES IN FISHES. 53 


extend nearly to the hinder end of the kidney in company with the interrenal (Pl. IX. 
figsc 6c 4) Pili XX. figs. 08 7; 37)" nother casese (PI) EX) figsy dee 3); PIX. 
fig. 6, s.r.) they cannot be traced very far behind the anterior extremity of the inter- 
renal. When the segmentally arranged bodies extend very far back they have a great 
tendency to get less and less distinct and more fragmentary in appearance. 

Their relation to the sympathetic is made evident in a general way by ordinary care 
in dissection, but the precise details of this relationship are not at all easy to make 
out. Chevrel has, however, described this part of the subject with such care that little 
need be said; but as very little besides his work has ever been done upon this subject, 
it may be as well to call attention to some points. . It is a great mistake to suppose 
that the sympathetic ganglia are all, or indeed mostly, wrapped up in the same sheath 
as the segmental suprarenal bodies. This is undoubtedly true in many cases, perhaps 
in all, of the first pair, but it is not nearly so common with the posterior ones. Indeed, 
without in any way questioning the value of Balfour's developmental researches, I am 
inclined to think that the connection between the sympathetic nervous system and 
these bodies has been overstated. They are intimately involved in the sympathetic 
plexuses, and often have tiny ganglia very close to them; but in the adult, at any rate, 
whateyer their developmental relations may be, it can, in my opinion, not be truly said 
that they are an integral part of the sympathetic nervous system. 

The Interrenal Body.—Vhis body, as we have seen, was the structure to which the 
name “‘suprarenal” was first applied. After the discovery of the “ paired suprarenals ” 
of Leydig, a new name was required for the old body. ‘This was supplied by Balfour, 
who called it “‘interrenal.” 

The interrenal body is an “ochre-yellow” rod-shaped structure, paired in the Rays, 
unpaired in the Sharks, lying usually in the region of the posterior part of the kidney, 
but sometimes extending as far forward as its anterior extremity (Pls. IX. & X. 
figs. 1-7, 7.r.). It bears a striking resemblance in its colour, general appearance, and 
relations to the kidney, to the suprarenals of Amphibia and Reptilia. 

The unpaired interrenal of the Dog-fishes (Pl. IX. figs. 2, 3, & 4; Pl. X. fig. 5, 7.7.) 
lies between the inferior surface of the dorsal aorta and the superior surface of the 
unpaired caudal vein, or, as Chevrel prefers to call it, the “ interrenal” vein. 

The paired interrenal of the Skates (Pl. X. figs. 6 & 7,7.) lies on the median 
side of the ureter and on a superior level, so that it often lies on the dorso-internal 


edge of each kidney adjoining the middle line. 
In addition to this one often finds slight streaks or dots of the same characteristic 


yellow colour in other parts of the kidney '. 


1 This may be compared on the one hand with the multiple suprarenals of the Sturgeon, and on the other 
with the frequency of “ accessory ” suprarenals in Teleostsand in Mammals, The accessory bodies in Mammals 
are said to consist of cortex only, and it may be as well to note here that my investigations lead me to 
conclude that interrenal, suprarenals in Teleosts and Ganoids, and cortex in Mammals are all strictly homo- 


logous with one another. 


54 MR. SWALE VINCENT ON THE 


The interrenal is generally much thicker behind than in front (Pl. IX. figs. 1, 3, 
& 4); in fact, while it is posteriorly often markedly bulbous, it usually tapers off 
anteriorly into a broken line of extreme tenuity. It diminishes in diameter, however, 
not regularly, but is often more or less moniliform; the constrictions may be 
complete here and there, and at the anterior extremity one often finds several small 
pieces of interrenal separated by a considerable interval from the rest of the body 
(Pl. IX. figs. 1, 3,& 4; Pl. X. figs. 5 & 7, a.ir.). 

In the Batoidei the interrenal is never quite symmetrical; nearly always the body of 
one side extends further forward than that of the other, and in one case, in Raja batis 
(Pl. X. fig. 6, 7.7.), I found a complete and comparatively thick bridge connecting the 
interrenal of one side with that of the other. ‘This might perhaps be looked upon as 
a single body behind, which became divided in front; in this instance the organ 
extended further forward on the right side, and further backward on the left. 

In some cases (e. g., Acanthias) the unpaired interrenal body appears to be placed 
not quite mesially, but is laid on the median surface of one or other kidney (Pl. X. 
fig. 5, i.r.). 

Occasionally (aja maculata) the anterior piece of the interrenal of each side, which 
is cut off by more or less of an interval from the rest of the body, shows curious claw- 
like processes on its outer side running in the kidney-substance towards the ureter 
(BIEXG figs ai a-2/72): 

Balfour quotes Semper as describing an anatomical connection between the inter- 
renal and the paired bodies. Balfour questions this, and I am convinced from the 
examination of several fresh specimens, both of Scylliwm and Raja, that there is never 
any such connection or continuity. Zhe two kinds of structure are always totally 
independent and unconnected. I have mentioned in the historical section Balfour’s 
surmises with regard to the existence of a third kind of body in relation to the kidney 
in Scyllium, and I have intimated that masses of adenoid tissue were to be expected, 
and were frequently found in various situations in and about the kidney. I have, 
nevertheless, carefully searched for anything which might rank as a “third kind of 
body.” I have been able to examine a number of fair-sized perfectly fresh specimens, 
and I have failed entirely to find anything of the kind. So that we may consider that 
we have to limit the morphological and physiological problem, difficult as it is, to the 
relationships and significance of the paired segmental bodies and the “ ochre-yellow 
strips” or interrenal bodies. The problem which naturally presents itself is: —Which 
of these two structures (if not both) corresponds to the suprarenal capsules of Teleosts 
and higher Vertebrates? For an account of the discussion see the historical section 
above. My own view will be stated after I have given a description of the histology. 

In the above account I have treated the Sharks and Rays together, because there is 
not sufficient difference between the general arrangement in the two cases to make a 
separate description necessary. 


SUPRARENAL BODIES IN FISHES. 55 


6. HOLOCEPHALA. 


In Chimera monstrosa (Pl. IX. fig. 1) from the examination of four well-preserved 
specimens, I find the suprarenals arranged almost exactly after the type of the 
Plagiostomes. There is the same interrenal, unpaired as in the Dog-fishes !, enlarged 
and rounded posteriorly and broken up at its anterior end. ‘There is also the same 
arrangement of the segmental bodies, the only noticeable difference being that the 
anterior pair, instead of being elongated and irregular, are smooth and regular oval in 
shape (PI1.1X. fig. 1, av.h.)?._ They are, nevertheless, many times larger than any of 
the other bodies of the same series. 


2. GANOIDEI. 

Among Ganoids I have only been able to obtain representatives of two families of 
the Chondrostei, viz., Acipenser sturio and Polyodon folium. The Polyodon was a 
spirit-specimen in a bad state of preservation, and I could find nothing in the way of 
suprarenals, so that I am limited to the Sturgeon for information about this order. 
The Sturgeon is practically the only Ganoid it is possible to examine in anything like 
a fresh state in this country. I have been able to obtain and have carefully examined 
two specimens whose tissues were, to all intents and purposes, in a living condition. 
The first specimen was 1:65 metre in length, with a kidney 63 centim. long; the 
second was 2°14 metres, with a kidney of 1 metre. 

The suprarenals in the Sturgeon are “ ochre-yellow” bodies of precisely the same 
tint as the interrenal in Elasmobranchs. ‘They vary extremely in size and shape, and 
are scattered in a more or less irregular manner throughout the substance of the 
kidney (Pl. X. fig. 8, s.). They are for the most part finely lobulated, almost 
coarsely granular to the naked eye, and many of the larger ones have processes or 
claws extending out in various directions into the kidney-substance. A certain 
number of the larger bodies are visible on the surface of the kidney, or revealed with 
very little dissection. A still larger number come into view on slitting up the 
posterior cardinal sinus, since they lie in abundance in its walls and in the immediate 
neighbourhood. A large residue, including the majority of the smaller bodies, are 
only revealed by digging away the kidney-substance in various parts. 

I find the larger bodies placed anteriorly (Pl. X. fig. 8); and in this respect my 
observations differ from those of Leydig, who says that the larger ones are posterior. 
It is quite possible that the arrangement differs in different specimens. I found none 
whatever in the hinder fifth of the kidney, and by far the larger number, at least of 
bodies of any size, were in both my specimens in the anterior seventh, 7. ¢., just behind 
the lymphatic head-kidney (Pl. X. fig. 8, 4.4.). Thus the most striking feature 
about their arrangement is their extreme anterior position as compared with Teleosts. 

1 Leydig describes a paired “ suprarenal,” but although my specimens were old, yet I am convinced that 


there was a single median interrenal in each case. 
2 [This appearance may possibly be due in part to the effect of long preservation. | 


56 MR. SWALE VINCENT ON THE 


Many of the suprarenals are of exceeding minuteness, even less than 1 mm. in 
diameter, and these are seen in large numbers from the interior of the posterior 
cardinal sinus, whence they appear like “ tubercles” in the wall of this vessel ; indeed 
the appearance of these tiny bodies as seen through the lining epithelium of the vein 
is almost identical with the caseous nodules of pathology. In the two specimens I 
have examined the large suprarenals were arranged in two very irregularly symmetrical 
groups just behind each “ head-kidney.” 1 Each group consisted of about a dozen, 
and all of them were in close proximity to the venous sinus (PI. X. fig. 8). 

Some of the suprarenals lying on the posterior surface of the kidney were flattened, 
almost wafer-like. 

The bodies I have seen varied in size from about 1 mm. in diameter to something 
under 1 cm., but measurements are difficult as their shape is often so irregular, They 
have a decidedly fatty appearance to the naked eye, and on being placed for a few 
hours in a 5 per cent. solution of osmic acid they become quite black, so that the 
presumption is that they contain some material of a fatty nature. 

I have not been able to make out any very intimate relations between the 
suprarenals and the sympathetic nervous system in the Sturgeon, but I find that in 
many cases the bodies are placed on blood-vessels which are running in the kidney- 
substance. 

3. TELEOSTEI. 
a. PLECTOGNATHI. 


In the tribe Sclerodermi I have only been able to obtain a small spirit-specimen of 
Balistes maculatus. It was not in a good state of preservation, and the results were 
negative. 

In the Gymnodontes I have examined the Sun-fish (Orthagoriscus mola) as a 
representative of the Molide and Tetroyon nigropunctatus among the Tetrodontide. 
The last was an ill-preserved specimen and nothing was found. The following is the 
arrangement in the Sun-fish :— 

Mo.ipa#.—In a specimen of Orthagoriscus mola (Pl. X. fig. 9, s.7.), 70 cm. in 
length, with kidneys of 20 cm. in length, I found one suprarenal only, which appeared 
to belong to the right side (Pl. X. fig. 9, s.r.). It was kidney-shaped, whitish, and 
fatty-looking, but of firm texture, and was about 1 em. long by about 5 mm. broad. 


6. PHYSOSTOMI. 


Muranip#.—In this family I have investigated both the Conger and the Silver Eel. 
There are two suprarenals in both species; these are in Conger conger (Pl. X. fig. 10, 
s.r.) triangular with rounded corners, in Anguilla anguilla round or oval in shape. 
Their position is peculiar in that they lie very anteriorly, being not far behind the 
point where the two crura of the kidney unite (Pl. X. fig. 10, s.r.). They are situated 


* [In some cases they extend into the lymphatic tissue—S. V., 10. 1. 97.] 


SUPRARENAL BODIES IN FISHES, 57 


on the ventral surface of the kidney, and are usually closely approximated to the middle 
line. They are only slightly imbedded in the substance of the kidney. The following 
may be taken as the typical arrangement in the Murenidze :— 

The suprarenals are about 1-5 mm. in diameter (of course they may be much larger 
in Conger). They are triangular with rounded corners; there is one on each side of 
the middle line and they are nearly in contact. They are on the ventral surface of 
the kidney, just behind the junction of the crura, and both are on the same level. 
The bodies are smooth, pale pink, and only slightly imbedded in the kidney. 

Crureip2.—In Clupea harenqus (Pl. X. fig. 11) I have found two unequal supra- 
renals placed on the spinal surface of the kidney at about the mid-point of its length. 

Esocip#.—I have examined two examples of Esox lucius (Pl. XI. fig. 12). The 
first specimen was 50 cm. in length, the second 57, both being young specimens. 

In the first (Pl. XI. fig. 12) the suprarenals were two in number, one on each side. 
They were imbedded in the substance of the kidney, nearer the spinal than the ventral 
surface, and placed distinctly in advance of the mid-point of the kidney length, so as 
to be about in the middle point of the length of the body. The bodies are thus 
even more anterior than in the eels. They were each about 5:25 mm. long by 2mm. 
broad. ‘Their surface was distinctly lobulated (Pl. XI. fig. 12, s.r.). 

In the second their arrangement appeared at first sight to be identical with that of 
the first, but, on closer examination, the left was seen to be represented by two bodies, 
and these were placed slightly in advance of the right one. They were, like those 
above described, imbedded in the kidney-substance. But in this specimen there were 
tiny “ accessory ” bodies, some two or three in number, in different parts of the kidney. 
I have not found, however, anything approaching the condition described by Stannius, 
in which the posterior half of the kidney is studded with suprarenal bodies. Very 
possibly this appearance was pathological. 

SaLmonip#.—In a specimen of Salmo salar, 83 cm. long, I found five suprarenals 
(Pl. XI. fig. 13, s.r.). There were three on the left side and two on the right. None 
of them were visible on the ventral surface of the kidney, as they were placed on the 
lateral or spinal surfaces. The bodies belonging to the right side were situated 
respectively 20-5 and 20 cm. from the hinder end of the kidney, being placed almost 
in the middle point of its length. Those of the left side were placed at unequal 
intervals opposite them. The right suprarenals are respectively 5 mm. and 3 mm. in 
diameter. The larger is anterior and slightly kidney-shaped. The largest on the left 
side is directly opposite the larger on the right ; it is 7 mm. by 3°5 mm. and distinctly 
kidney-shaped. Ata short distance in front of this is a body about half its size and 
likewise kidney-shaped. At a further interval behind it lies an elongated oval one. 
The suprarenals are very pale pink in colour, almost white, and have a fatty 
appearance. They do not project appreciably from the surface of the kidney. They 
are distinctly lobulated and marbled with blood-vessels. 

vou. XIv.— Part mI. No. 3.—April, 1897. I 


58 MR. SWALE VINCENT ON THE 


In another Salmon, a young fish only 47 cm. in length, I found three suprarenals, 
placed, as in the former specimen, at about the middle point of the length of the 
kidney and on the spinal surface. There were two chief ones, right and left respec- 
tively, situated each about 1 mm. from the middle line, the left one being about 3 mm. 
by 1:5, the right about 5 by 1:5, but irregular in shape. About 1 cm. anterior to 
these two is another about 1 mm. by ‘5 mm. on the left side. 

In a specimen of Salmo trutia, 58 cm. long, with a kidney of 26 cm., I found the 
suprarenals seven in number. ‘There were six on the spinal and one on the ventral 
surface, all small. They were pale pink, roundish or oval bodies, and were partly 
imbedded in the substance of the kidney. This fact, coupled with the presence of one 
on the ventral surface, suggested to me that some might be completely buried in the 
substance of the kidney. This, however, I found not to be the case. The most 
anterior one was very small, and was situated 10 cm. from the anterior end of the 
kidney; the rest were scattered throughout the succeeding 4:25 cm. The one on the 
ventral surface was 12:5 cm. from the anterior extremity of the kidney. None of the 
bodies were more than 3 mm. in diameter. 

In Osmerus eperlanus (Pl. XI. fig. 14) the suprarenals were represented by two 
miliary bodies about the size of a pin’s head, projecting from the hinder end of the 
kidney, placed one behind the other (s.7.). 

Among the Cyprinide I have examined :— 

Leuciscus rutilis. (Pl. XI. fig. 15.) Leuciscus vulgaris. (Pl. XI. fig. 17.) 
£ cephalus. (Pl. XI. fig. 16.) 

The suprarenals are usually two; but in a specimen of L. vulgaris there were four 
(Pl. XI. fig. 17, s.r.). They are usually rounded in this family. 

As to position, they are usually in the posterior fourth of the kidney, but they may 
reach as far forward as the junction of the posterior with the middle third. They are 
usually placed one on each side of the middle line, but are often very close to it. 
They are on the spinal surface or the edge of the kidney. 

In the fishes of this family I have examined the bodies have never been more than about 
2 mm. in diameter, and they are often buried to some extent in the kidney-substance. 


ce. ANACANTHINI. 
Gapip#.—In this family I have investigated :— 


Gadus morrhua. (Pl. XI. fig. 18.) Merluccius vulgaris. (Pl. XI. fig. 20.) 
» e@eglefinus. (Pl. XI. fig. 19.) Molva vulgaris. (Pl. XI. fig. 21.) 
»  merlangus. Motella tricirrhata. 
» -virens. 


In the Gadide there is considerable variation in number and size of the suprarenals, 
although their shape is nearly always rounded. 


As for number, two must be regarded as the rule, but there is even more variation 


SUPRARENAL BODIES IN FISHES, 59 


in this respect than in the Pleuronectide (g. v.). Thus, in a specimen of Gadus 
morrhua I have found as many as five; in G. merlangus I found only one. I have 
occasionally found one also in Molva vulgaris. In Merluccius vulgaris I have found in 
one case as many as five; in this case the suprarenal of the left side was represented 
by four small bodies instead of one larger one (Pl. XI. fig. 20, s.r.), 

Their position is usually one on each side, but, as in other families, the right and 
left are rarely quite on the same level, one being usually anterior or posterior to the 
other. They also vary as to their relation to the middle line, some being more lateral, 
others more central. 

The suprarenals in this family are almost always visible on the ventral surface of the 
kidney, though they may (either one or both) be lateralized, or even exceptionally 
may be partially on the spinal surface. They are usually more or less imbedded in 
the kidney-substance. 

The bodies appear to be situated always in the region of the posterior third of the 
length of the kidney, though they sometimes approach very near the anterior limit of 
this distance. 

The size varies from 1 mm. to 1 cm. in longest diameter in the fishes I haye 
examined. 

In one specimen of Gadus morrhuwa the arrangement was peculiar. The suprarenals 
were five in number. The tail of the kidney projected 3 cm. under the hemal arch. 
Four of the bodies were under the arch on the ventral surface of the kidney; they 
were compressed into angular form owing to their position. There were two large 
ones, the anterior being 1 cm. by ‘5 em. (irregular oblong), the posterior being 
rhomboidal and having about the same greatest dimensions. There were in addition 
two quite small bodies placed between the two larger ones. The fifth one was on the 
spinal surface of the kidney, just anterior to the hemal arch. It is about 1 cm. by 
‘5 cm., oval, and appears divided into two parts on the surface by a median longi- 
tudinal depression. 

The typical arrangement of the suprarenals in the Gadide may be stated thus :— 

The organs are rounded and about 5 mm. in diameter. There is one on each side 
of the middle line on the ventral surface of the kidney, but one is more lateral than 
the other. They are placed at about one quarter the length of the kidney from its 
hinder end, and one is slightly anterior to the other. They are smooth and partly 
imbedded in the kidney-substance, and are pale pink in colour. 

PLEvRONECTIDA.—I have been able to obtain and examine :— 


Pleuronectes flesus. (Pl. XI. fig. 22.) Hippoglossoides limandoides, (Pl. X11. fig. 26.) 
5 limanda, (P1. XI. fig, 23.) Solea vulgaris, (Pl. XII. fig. 27.) 
- platessa. (Pl. XI. fig. 24.) Rhombus levis. (Pl. XII. fig. 28.) 
Hippoglossus vulgaris, (Pl. XII. fig. 25.) Zeugopterus. 


In Pleuronectide there is considerable variation in the number, form, size, and 
12 


60 _ MR. SWALE VINCENT ON THE 


position of the suprarenal bodies; the variation in each of these respects affects not 
only the different species, but even different individuals of the same species. 

With regard to their number this may be stated as usually two. Thus there are 
most often two in Hippoglossus vulyaris, Plewronectes flesus, P. limanda, and Solea 
vulgaris. But I have found one only in some specimens of P. limanda and Hippo- 
glossoides limandoides, while I have found three in one specimen of P. flesus. [In 
this case, however, the two of one side were close together, and were, taken together, 
not larger than the one of the opposite side. } 

Their shape is nearly always rounded, but in P. flesus one was oval. 

As to their position, the bodies appear to be affected to some extent by the 
characteristic asymmetry of the family, though sometimes they may be arranged 
symmetrically. There is usually one belonging to each side, but sometimes one is 
placed behind the other in the middle line (Pl. XII. fig. 26, s.r.). This was found, 
é. g.,in P. flesus and Solea vulgaris. Occasionally they are placed abreast, but both 
on one side of the median line; this occurred in P. limanda. Often the body of one 
side is decidedly more anterior or posterior than that of the other. Again, sometimes 
they are close together in the middle line (Pl. XI. fig. 24); at other times separated 
by the whole breadth of the kidney. 

The suprarenals are always enclosed in the capsule of the kidney; but appear never 
to be imbedded in the kidney-substance in this family. They are always near the 
posterior end of the kidney, and never further forward than the junction of the 
posterior with the middle third of the kidney. 

The size of the capsules in the Pleuronectidz varies in specimens I have seen from 
‘5 mm. to 1:75 cm. in diameter. They do not appear to vary very directly in 
proportion to the size of the individual fish, but they are larger in the larger species. 

The typical arrangement of the suprarenals in the Pleuronectide may be described 
as follows :— 

The organs are about ‘75 cm. in diameter and rounded. ‘There is one on each side 
of the median line on the spinal surface of the kidney, but one is nearer to the middle 
line than the other. They are situated about one-quarter the length of the kidney 
from its posterior extremity, and one is placed rather in front of the other. They are 
smooth and free on the kidney-surface, and are pale pink or glistening white in aspect. 


d. ACANTHOPTERI. 


In the Pharyngognathi I have only been able to obtain a spirit-specimen of Coris 
pulcherrima. It was not in a good state of preservation and no suprarenals were found. 

In the Acanthopterygii I have been enabled to investigate representatives of several 
families. 

PrrciD&.—In two specimens of Perch (Perca fluviatilis, Pl. XII. fig. 29) I have 
found only one suprarenal body to be present in each case. In both it was about 


SUPRARENAL BODIES IN FISHES. 61 


8 mm. from the hinder end of the kidney on its spinal surface, and was about 1 mm. 
in diametez. 

Gastrosteus spinachia was too small for anything to be seen with certainty. 

In a specimen of Labrax lupus 70 cm. in length, with a kidney of 30 cm., I found 
two suprarenals placed one immediately behind the other on the spinal surface. The 
anterior one is 5 mm. by 24 mm.; the posterior one is about half the size of the anterior. 

MuLuip%.—Two specimens of Mullus barbatus (Pl. XII. figs. 30 & 30 A) have 
been examined. In one, 24 cm. long, there were three suprarenals present ; in the 
other, 42 cm. long, there were five. 

In the first specimen these were crowded together in the middle line on the spinal 
surface about 3:25 cm. from the hinder end of the kidney (Pl. XII. fig. 30, s.7.). 

In the second one (Pl. XII. fig. 80 4) there were two ‘“‘ chief” bodies, right and left, 
situated a few mm. behind the hinder end of the kidney, attached to the kidney and 
to the intestine and body-wall by fibrous strands. The left one has an “accessory ” 
body just behind it, and there are two small ones on the ventral surface of the left tail 
of the kidney (Pl. XII. fig. 304, s.r.). The right chief body is 4 mm. by 3 in 
diameter, the left 5 by 2. 

It is interesting to note, in relation with the abundance of suprarenal tissue, that 
this fish is of a red colour, and has an intensely pigmented, almost black, peritoneum. 

Sparip#.—In the common Sea-Bream (Pagellus centrodontus) I have found two 
suprarenals, 2 mm. in diameter, and very pale pink in colour, situated 1 cm. from the 
hinder end of the kidney on its spinal surface (Pl. XII. fig. 31). In Cantharus 
griseus almost exactly the same arrangement was found. 

Trietip#.—In this family I have examined Cottus gobio (Pl. XII. fig. 32), Trigla 
pini (Pl. XII. fig. 33), and 7. lyra (Pl. XII. fig. 34). The rule as to number is the 
same as in Teleosts generally, 7. ¢., that there are two; but in one specimen of Trigla 
pint I have found one only, in the middle line, on the spinal surface, about 8 mm. 
from the hinder end of the kidney. 

Their shape is rounded, and their position as a rule bi-lateral; but in Trigla lyra I 
have found two, placed one behind the other, both to the left of the left edge of the 
kidney near its hinder extremity (Pl. XII. fig. 34, s.r-.). 

Except in this one example of Z. lyra I have always found them on the spinal 
surface of the kidney in this family. 

They are of a very pale pink colour, and are free on the surface of the kidney, 
sometimes attached to the body-wall by bands of fibrous tissue. In size they vary 
between | and 2 mm. 

ScomBrip2.—In this family I have been able to examine Scomber scomber and Zeus 
faber (Pl. XII. figs. 35 & 36). 

In the Mackerel (Pl. XII. fig. 35) I have found the suprarenal bodies on the 
ventral aspect of the kidney, about 4°5 cm. from its posterior extremity, almost 


62 MR. SWALE VINCENT ON THE 


completely concealed, however, by kidney-substance. The body of the right side was 
rounded and about 2 mm. in diameter, the left was about 3mm. by 15 mm. In 
another specimen, however, I found them placed on the dorsal aspect of the kidney at 
about the middle of its length, the right being slightly smaller than, and somewhat 
anterior to, the left one. 

Frey says that in this fish the suprarenals are placed one behind the other. This is 
certainly not the rule, but it may be so in some cases. 

I have examined several specimens of John Dory (Pl. XII. fig. 36). There were 
usually two suprarenals, one on each side of the middle line, but occasionally the body 
of one or other side has been represented by two smaller bodies. Sometimes they are 
placed side by side, sometimes one behind the other. They are always near the 
junction of the posterior crura of the kidney. They are mostly white or slightly 
iridescent and smooth, being enclosed in a firm fibrous capsule. They are rounded and 
situated on the spinal surface of the kidney. This is noteworthy, as I have always 
found them on the ventral surface in Scomber. Another point of distinction between 
the two genera is that in the John Dory they are not imbedded in the kidney- 
substance ; they always lie within the posterior quarter of the kidney. 

BLenniD#.—In a specimen of Annarhichas lupus (Pl. XII. fig. 37), 115°5 cm. in 
length, I found two suprarenals, situate 5 cm. from the hinder end of the kidney. 
The left was on the ventral surface of the kidney, nearly round, and 5 mm. in 
diameter. The right was on the dorsal or spinal surface, and, being the larger, its 
central point was posterior to that of the left, as their anterior ends were on the same 
level. This right body was oval in shape and 1 cm. by ‘5 cm. The organs were 
partly imbedded in the kidney-substance. 

PepicutattiIn the Angler (Lophius piscatorius, Pl. XII. fig. 38), in a fish of 
54 cm. in length, I found five suprarenals. The chief pair were situated on the 
ventral surface of the kidney, about 1 cm. from its hinder end. The right was 3 mm. 
in diameter, the left 3°5 mm. They were of a pale flesh-colour and marbled with 
very fine blood-vessels, 

In addition to these two, there was on the right side a body about 1:25 mm. in 
diameter, just at the hinder extremity of the kidney, where the ureter springs from its 
substance ; again, just anterior to it is another, about one-third its size. On the left 
side there is also a second body, about 1°5 mm. by ‘75 mm., situated midway between 
the ‘‘ chief” body and the end of the kidney. 

The larger suprarenals were more or less imbedded in the substance of the kidney. 

Mvaitipa.—In Mugil capito (Pl. XIII. fig. 39), 55 em. long, I found one suprarenal 
capsule of large size (6 mm. by 3). It was placed about 4 mm. from the posterior 
extremity of the kidney, and, being imbedded in the kidney-substance, its large size 
enabled it to be seen either from the spinal or the ventral surface (Pl. XIII. fig. 39). 

In another example, 35 cm. long, there were two suprarenal bodies, on the spinal 


SUPRARENAL BODIES IN FISHES. 65 


surface, at the extreme hinder end of the kidney. They were packed close together in the 
middle line, the right one being about 2°5 mm. by 2, and the left being about 2 by 1. 

Here, as well as in Mullus barbatus, we have a proportionately large amount of 
suprarenal tissue associated with a deeply pigmented peritoneum. 

Discosout.—lI have only so far been able to examine one specimen of Cyclopterus 
lumpus (Pl. XIII. fig. 40). This fish was 46-25 em. in length. The suprarenals were 
situated on the spinal surface of the kidney. The central point of the left was 3 cm. 
from the hinder end of the kidney, that of the right was 3°75 cm. distant. The right 
body was rounded and about 4:5 mm. in diameter, the left was oval (5 mm. by 2°5 mm ). 

It will have been seen from the above account that in the Teleosts J find suprarenal 
bodies in all fresh specimens examined. There are usually paired, round or oval, 
pale pink bodies, placed on the spinal or ventral surface of the kidney. They are near 
the posterior extremity of the renal mass and are either free on its surface or more or 
less imbedded in its substance. 


4, DIPNOI. 

Nothing is known of the suprarenal capsules in this order of fishes. At any rate 
they are not described in Lepidosiren (g)1; in Ceratodus (8) and in Protopterus their 
presence is exceedingly doubtful. However, what has to be said on this head will be 
stated below in the histological section (V.). 


Haying thus reviewed the gross anatomy of the suprarenal bodies in the orders I 
have examined, a few general remarks must be here interpolated. Being unable to 
obtain fresh specimens of any of the Dipnoi, I will leave this order quite out of the 
question?. In Elasmobranchs, Ganoids, and Teleosts, suprarenals have long been 
described, but never very systematically or connectedly. This I have endeavoured to 
do as far as possible in the above account. That there was need of this is clear from 
the fact that some zoologists of standing still seem largely to ignore their existence. 
Thus Beard (2), in his very interesting paper, says :—“ Not one of the least brilliant 
of Mihalkovics’ discoveries is that of the relationship between the reproductive gland 
and the suprarenal bodies ?. Mihalkovics showed that the non-nervous part of these 
organs is the most anterior portion of the reproductive gland, and that in those animals 
in which suprarenals occur the germinal cells which give rise to them have under- 
gone degeneration and have got separated off from the rest of the germinal epithelium 
as a mass of cells remaining in a so-called embryonic state.” This may be true enough, 

1 See also Ehlers, E., “ Zur Kenntnis der Eingenweide yon Lepidosiren,” Vorliiufige Mittheilung, Nachrichten 
der k. Gesellschaft der Wissenschaften zu Gottingen, Math.-physik. Klasse, 1895, Nr. 1. 


2 See footnote 2, page 73. 
3 Tt is noteworthy in this relation that in the Reptilia the suprarenal body is most often in very close 


relationship with the reproductive gland. The same applies to birds. 
4 [It may be observed, by the way, that the medulla is probably not ‘‘ nervous,” and that the cortex is certainly 


not * embryonic.”—S. V., 10. 1. 97.] * 


64 MR. SWALE VINCENT ON THE 


but Beard goes on: “ It isa significant fact, hitherto, I believe, unnoticed, that suprarenals 
are found only in those forms in which a reduction in number of ripe sexual elements 
required has taken place”}, In regard to this I can only state that suprarenals (one 
or both parts) are probably universally present throughout Elasmobranchs, Ganoids, 
and Teleosts. Further, this writer observes: ‘“ In the present state of our knowledge 
of these bodies, I do not wish to lay too much stress on the point of existence or non- 
existence of suprarenals in any order of Vertebrates.” But he adds: “If Rabl’s ‘ tree’ 
be correct in respect of the ancestry of Ganoids from Sharks... ..I can conceive 
that the non-eaistence of suprarenals in the former group! would be a fact which our 
author might find difficulty in accounting for—unless he ignored it entirely!” 

Suprarenals non-existent in Ganoids! Either Beard has quite overlooked their 
presence in the Sturgeon, or he has some view of his own as to the significance of those 
yellow bodies which are scattered throughout the kidney. If the latter were the case 
one would have surely expected him, in dealing with the subject, to note the fact. 

Further on in the same paper our author gives a “ tree,” in which it is stated that 
suprarenals are totally absent not only in Ganoids and Marsipobranchs, but also in 
Teleosts?. It would be interesting to know what he calls those little bodies I have 
represented in my plates. Later on he says: “It would doubtless be interesting to find 
some traces of suprarenal bodies in the Dipnoi.” It will be seen from what I say on 
this subject under the head of histology, that I am decidedly of the opinion that in all 
probability there are suprarenals of some sort in the Dipnoi ®. 

With regard to the occurrence of suprarenals in Teleosts, in addition to the species 
in which I have described them above, there are mentioned many other species in 
which other writers have found them. Thus Stannius and Ecker, also Hyrtl, state that 
they are present in several species I have not been able to obtain. M*‘Kenzie, too (15), 
describes them in Amiurus catus. So that, although of course I cannot affirm from my 
own observations that these bodies are universally present in this order, I should, from 
a joint consideration of my own and previous researches, consider this to be more than 
probable. Jf suprarenals are not present in all Teleosts, in which species are they absent? 


V. Histonoey. 
1, ELASMOBRANCHII. 

My results in this department appear to differ in many respects from those of Leydig, 
Semper, and Balfour. They agree fairly well in some respects with those of Chevrel, 
but there are important points of difference, and, besides, Chevrel’s histological drawings 
are so imperfect that little can be learnt from them. 

’ The italics are mine.—S. V. 

* This is not stated in so many words, but the “tree” has inscribed upon it, ‘‘Suprarenals totally absent on 


this side”; and “ this side” includes the orders named above. 
* [These have since been discovered by Petitt (Joc. cit.).—S. V., 10. 1, 97.] 


SUPRARENAL BODIES IN FISHES, 65 


ParReD “SUPRARENAL”’ BopiEs.— With regard to the segmentally-arranged bodies, there 
is not a great deal new to be said. But it is certain that in all cases the structure is very 
different from that which Balfour described and depicted. Chevrel explains Balfour’s 
appearances as due to the action of reagents, and I am satisfied, from the examination 
of many individuals and many species with various modes of preparation, that there is 
never any such arrangement of a cortex of columnar and a medulla of polyhedral cells 
(Pl. XIII. figs. 41 & 42). 

The structure of these bodies is far from easy to describe, and this I find, notwith- 
standing that my material was perfectly fresh and my preparations mostly very satis- 
factory. The organs are surrounded with a tolerably firm fibrous capsule of a thickness 
about 4-7 w (Pl. XIII. fig. 41, ¢.), which sends in septa (s.), which in some parts are 
distributed almost as regularly as in a Mammalian lymphatic gland. These septa 
rapidly break up into an irregular fibrous meshwork which is distributed throughout 
the body (Pl. XIII. fig. 41, sér.). 

The parenchyma of these bodies is not arranged in any definite acini or alveoli, thus 
marking them off distinctly from the suprarenals in Teleosts, Ganoids, and the interrenal 
body in Elasmobranchs (q. v.) and indeed from suprarenal capsules in other Vertebrata. 

There are, as a rule, no definite cell-outlines to be made out, and the main part of 
the parenchyma appears to consist of an irregular or wavy fibrous stroma (Pl. XIII. 
fig. 41, str.), with protoplasm (pr.) and scattered nuclei of round or oval sbape (n.). 
The protoplasm is often granular, sometimes finely, sometimes coarsely. The nuclei 
are of different characters, sometimes showing nuclear figures, at other times devoid of 
them, sometimes large and faintly stained, sometimes small and darkly stained. They 
vary in diameter from 5 to 9 u. However unsatisfactory this may seem, it is all that 
can be ascertained, and it is difficult to imagine how former observers such as Leydig 
and Balfour could have described them as having definite lobules and cell-outlines. 

But there are definite cells in some parts (Pl. XIII. fig. 42, p.c.) quite apart from 
the easily recognizable nerve-ganglion cells. They have, so far as I know, not been 
previously described. ‘They are mostly triangular or multipolar in shape (Pl. XIII. 
fig. 42, p.c.) and of a uniform sepia-brown tint, and they contain large, very darkly 
stained, round nuclei. It is difficult to state their size, as their shape is so irregular ; 
they vary, however, in their greatest lengths from 10-30 4; the nucleus is usually 
about 6-8 » in diameter. ‘These cells are found particularly in the more central 
parts of the anterior paired bodies, but their distribution is irregular. The brown 
coloration of these cells is possibly due to the Miiller’s fluid in which the material 
was hardened. But, even if this be the case, the appearances indicate a difference in 
chemical reaction, and therefore in physiological import. ‘These cells appear in some 
places to communicate together by their processes, but whether this communication is 
real or apparent I cannot be absolutely certain. Is it possible that these cells have 
anything to do with nerve-cells ? 

VOL, x1v.— Part 11. No. 4,—April, 1897. K 


66 MR. SWALE VINCENT ON THE 


There is no essential difference in structure between the more anterior and the more 
posterior paired bodies. But, as Balfour has pointed out, the ganglionic development 
is more abundant in the case of the anterior ones. The presence of nerve-cells in the 
substance of the body itself is also more frequent in the case of the anterior suprarenals ; 
in fact, nerve-cells are almost absent from the interior of the most posterior bodies, 
which consist entirely of suprarenal tissue proper. 

The anterior pair or “axillary hearts,” however, from their size and extensive 
nervous connections, require a separate description. 

A longitudinal section of the “axillary heart” of Scylliwm canicula, taken some- 
where near its median part, shows that the body consists of an external and an internal 
portion. The internal part is characterized by the above-described brown pigmented 
cells1. The external part on one side is composed to a great extent of large nerve-cells 
with nerve-fibres running longitudinally. The nerve-cells are on an average 55 in 
diameter, but some of the oval ones may have a length of more than twice this number. 
This formation extends for about two-thirds of the length of the gland. On the opposite 
side and at the ends their external part is composed of the tissue which makes up the 
bulk of the segmental bodies, viz. a very fine but irregular fibrous matrix enclosing 
protoplasm with oval nuclei without definite cell-outlines *. 

At about a third of the length of the gland from the anterior end, a large group of 
nerve-cells occupies the central part of the structure, and scattered nerve-cells are 
distributed in other parts of the organ. 

In some instances (e. g. axillary hearts of Acanthias vulgaris) there are to be seen some 
very large nuclei, twice or thrice the usual dimensions, and in addition one sees nuclei 
of varying sizes down to the very smallest. 

This is not intended to be an exhaustive account of the histology of these bodies, 
and with one more remark I will conclude for the present what I have to say about 
them. I believe that many of the fibrils in the paired suprarenals (with the exception, 
perhaps, of the axillary hearts), which Balfour took for nerve-fibres, are in reality 
nothing more than connective tissue, which, as we have seen, runs in an irregular 
manner throughout the parenchyma. 

TyTerRENAL Bopy.—I may as well state at the outset that I agree with the view that 
this body corresponds with the cortical part of the suprarenal in Amphibians, Reptiles, 
Birds, and Mammals. I am further of opinion that it is essentially a secreting-gland. 
These conclusions have been forced upon me by its very close resemblance in structure 
to the cortex of the suprarenal capsules of higher Vertebrates, and by its strikingly 

1 The appearance of these pigment-cells is quite different from those which are found in various organs and 
tissues in fishes. There are in most cases no irregular masses or granules, but the whole cell, which has clear 
and definite outlines, is of a uniform brown tint. In some few cases, however, the cells appear to be of the 
ordinary coarsely granular type. 


Since the above was written, by careful examination of a good preparation with oblique light, I have 
succeeded in making out the cell-gutlines in the “ axillary hearts ” of Scylliwm canicula. 


SUPRARENAL BODIES IN FISHES. 67 


glandular arrangement into alveoli and its markedly granular protoplasm. The cell- 
outlines, too, are clearly seen in the majority of instances (Pl. XIII. fig. 43, al.w., e.c.). 
The alveoli appear to be arranged, in some parts at any rate, in a more or less radiating 
manner round large veins or venous sinuses (Pl. XIII. fig. 43, v.s.). 

A very striking feature in most of my sections is the presence of structures which 
remind one of the ‘“‘demilunes”” in mammalian mucous glands (Pl. XIII. fig. 43, d.c.). 
Their presence is quite clear and unmistakable, but I cannot guess at their significance !. 

The general appearance under the microscope reminds one very forcibly of cortical 
suprarenal tissue in the Mammalia. The interior of the organ is made up of lobules 
or alveoli, each enclosed in a delicate but distinct fibrous capsule about 2m in 
thickness. These alveoli are of various sizes, more or less oblong in form, about 
50 » thick and reaching 140-150 » in length, so far as I can judge by my sections; 
in many cases fibrous tissue appears to divide the individual cells one from another. 
The cells vary much in shape and size, the majority being elongated; some of the 
longest of these are 30-50 4 in length and reach quite across the thickness of an 
alveolus. The nuclei of the cells have an average diameter of 104. The above 
description of the interrenal is taken from a specimen of Raja clavata. The interrenal 
of Scylliwum has much the same structure, but the alveoli are not perhaps quite so 
clearly marked out. ‘This applies to the Sharks generally. 

In Chimera the anterior pair of the suprarenal bodies (“ axillary hearts ”), although 
long preserved in spirit, presented appearances which are worth recording. It is 
somewhat easier to imagine definite cells in the case of Chimera than in the Elasmo- 
branchs, for although there are no distinct cell-walls to be seen, yet the protoplasm 
appears to be gathered in separate portions of varying shape round the nuclei. These 
are of very different sizes and shapes. It is noticeable that there seems to be no 
ganglion of any great size in immediate connection with the body, and undoubted 
nerve-cells in its substance are almost totally absent. 

The interrenal of Chimera shows on microscopic examination a distinct alveolar 
arrangement as contrasted with the absence of such an arrangement in the paired 
suprarenals. This corresponds with what is found in the Sharks and Rays. 

With regard to the true nature of these two kinds of body in Elasmobranchs, and 
the question as to their relations to the suprarenal bodies in the higher Vertebrates, 
there are one or two points to be noticed. 

1. The interrenal of the Cartilaginous Fishes bears a very striking external resemblance 
to the structure which, in Amphibians and Reptiles, is always admitted to be a supra- 
renal. Its colour, its position, and its proneness to become broken up at its anterior 
end are all points of resemblance. 


1 The drawing (Pl. XIII. fig. 43) represents the appearances after hardening, double-staining in bulk, and 
saturating with paraffin, When examined fresh, the structure is largely concealed by the abundance of fatty- 


looking globules. 
K2 


68 MR. SWALE VINCENT ON THE 


2. The interrenal, in its histological features, very closely resembles the suprarenals 
of Teleosts, and in some cases its structure is almost identical with that of the cortex 
of certain Mammalian capsules. 

3. The paired bodies, as we have seen, have not such a definitely glandular 
appearance. They consist simply of a fibrous stroma, protoplasm, nuclei of various 
characters1, and nervous elements. 

Although the question is still very doubtful, and though I have not attempted any 
investigation of the development, I am decidedly inclined to the view that if one and 
not both of these bodies corresponds to the suprarenals of the higher Vertebrata, this 
one is the interrenal and not the paired suprarenal. And I am led to this view from 
consideration of its general appearance and position as well as its minute structure 2 

I think there can no longer be any doubt, at all events, that the paired suprarenals 
and the interrenals are totally distinct structures and almost certainly have quite 
different functions. 

On the other hand, one is much more tempted to believe that the interrenal 
corresponds to the cortex and the paired suprarenals to the medulla of the Mammalian 
capsules, and this hypothesis would, so far as I can see, accord fairly well with known 
facts. But I should hesitate to give myself over entirely to this view until the 
development of the suprarenals in Mammals has been put upon a more satisfactory 
basis. Besides, even if this hypothesis were correct, it would not be by any means a 
final solution of the difficulty. For the questions naturally arise, What is the 
significance of this dual origin in Elasmobranchs? What is the essential nature 
of the interrenal and the segmental bodies? Balfour and others have proved 
conclusively a very intimate structural and developmental connection between the 
sympathetic chain and the segmental bodies. But, in effect, what does this mean? 
The segmental bodies are not in their essence nervous structures, and we know of 
nothing elsewhere which illustrates or throws any light on this curious development 
of glandular material in connection with the nervous system. 

Again, I am not sure that the paired segmental bodies are not more intimately related 
to the blood-vascular than to the nervous system. We have seen that each one is 
placed around an arteriole, and many of them are immersed in the blood of the cardinal 
sinus. Further, it occurs to me that, after all, their connection with the sympathetic 
system may be in a certain sense accidental: 7. e. they are abundantly supplied by the 
sympathetic as are other abdominal organs ; but that, owing to their position, close to 


1 See footnote 2, page 66. 

2 Since the aboye was written, and as the result of further investigations throughout Vertebrates, I have 
elsewhere emphasized the view that the suprarenals of Mammals are to be regarded as compound glands derived 
from these two organs in Elasmobranch Fishes, and haye suggested that each portion probably has a distinct 
function. The function of the medulla is becoming tolerably well understood. The function of the cortex is 
still unknown (Oliver and Schiifer, Proc. Physiological Society, 1895; Swale Vincent, Proc. Birm. Nat. Hist. & 
Phil. Soc. 1896, vol. x. pt. i.; Birm, Med. Review, Aug. 1896; also Anat. Anz. xiii. Bd. Nr. 1 & 2, 1897). 


SUPRARENAL BODIES IN FISHES. 69 


the main chain of the sympathetic, they have become connected with it in a remark- 
ably intimate manner !. 


2. GANOIDEI. 


STRUCTURE OF THE SUPRARENAL Bopres or THE SrurcEoN.—I have examined some of 
these bodies after putting them quite fresh into 1°/, osmic acid for twenty-four 
hours, and then cutting in gum. They were stained quite black to the naked eye, and 
on cutting and examining microscopically their alveolar structure was obvious. 

If examined immediately in Farrant’s fluid, the alveoli appeared to be filled with 
mulberry-masses of material most probably of a fatty nature, as they had taken a deep 
brown stain with osmic acid, so that the cell-outlines were not to be made out. 

If, however, the sections were passed through turpentine or Canada balsam, the 
structure was beautifully shown (Pl. XIV. fig. 44). The rounded or elongated oval 
alveoli (50-60 » in diameter, or even 100 p long by about 60» broad) are bounded by 
bold thick walls, averaging 3 » in thickness (a/.w.), and the cell-outlines were admirably 
preserved (wv). ‘The preponderating shape of the cells was round or oval, and in some 
parts they are seen to overlap, as the section was thick enough to contain several layers (2). 
In other parts the cells are more polyhedral or irregular. Like the alveoli, they vary 
somewhat in size; their average diameter is about 20. The nuclei (n.) are deeply 
stained and somewhat irregular in shape, having a diameter of 3-6 ». The protoplasm 
is very finely granular as a rule, occasionally more coarsely granular 2. There are small 
herve-ganglia in connection with some of the bodies. 


* As for the meaning of the two kinds of body present in the Elasmobranchs and of one only in Teleostei 
and Ganoids there are two alternative theories. 

One is that the two kinds of structure are both represented in the higher Vertebrates, but that the paired 
bodies have disappeared in Teleosts. 

The other view is that the paired bodies in Elasmobranchs are the remains of former more important bodies, 
while in higher forms they disappear altogether. In favour of this it may be urged that their nerve-supply is 
quite disproportionate to their size and apparent importance, and possibly represents the persistence of a nerye- 
supply suited to a former larger structure. 

As to which of these is the correct view I cannot at present be certain, but on the whole perhaps the usual 
view is the most probable, viz., that in Elasmobranchs the paired suprarenals and the interrenal correspond 
respectively to the medulla and the cortex of Amphibians, Reptiles, Birds, and Mammals. 

There is not much difficulty in recognizing that the interrenal of the Cartilaginous Fishes corresponds to the 
cortex of higher Vertebrates, but it is far from easy to imagine that the masses of protoplasm with scattered 
nuclei which constitute the paired bodies in Elasmobranchs are really homologous with the branched granular 
cells of the medulla of higher Vertebrates. 

* When the suprarenals of the Sturgeon are hardened, double-stained in bulk, and cut in paraffin, they do 
not show nearly such a distinct alveolar arrangement as do those treated fresh with osmic acid. The larger 
oyal nuclei show a very distinct nuclear network and a large darkly-stained nucleolus centrally placed. In 
other cases only dark nuclear granules of various sizes are to be made out. The cells appear for the most part 
made up of a variable number (5-6) of vesicles, from which evidently the fat has been dissolved by this mode 
of preparation. The cells of the body have shrunk to some extent, so as to leave spaces of variable extent 


between them. 


70 MR. SWALE VINCENT ON THE 


I have no doubt, from the above structure, that these bodies are representatives of 
the suprarenals in Ganoids, and in my opinion they correspond to the cortical portion 
in higher Vertebrates. 

3. TELEOSTEI. 

The minute structure of the suprarenal bodies does not exhibit any wide variation 
throughout the different families of Teleosts. Between the individual members of the 
families the differences are of course still less. There are, however, such differences, 
and these appear to consist chiefly of variation in amount of fibrous tissue, variation 
in blood-vascular supply, and in the shape of the alveolar compartments into which 
they are all divided. 

The organs are surrounded by a capsule of very varying thickness (in the species I 
examined from 4 to 70 ), in close connection with which are sometimes found sympathetic 
nerve-fibres and small ganglia, and blood-vessels. Outside the capsule there is usually 
more or less adenoid tissue. In the majority of instances this is nothing more than 
a local increase, in the neighbourhood of the suprarenals, of the ordinary renal inter- 
tubular adenoid tissue. 

The capsule is always thicker throughout the area which is in contact with the 
kidney. This is probably to be explained by the fact that we have a double layer, 
consisting of the capsule of the suprarenal and the capsule of the kidney, fused 
together at this part. There is never any direct anatomical connection between the 
parenchyma of the suprarenals and kidneys, and, so far as I have been able to discern, 
nothing to suggest that the suprarenals are parts of the kidney. 

The fibrous capsule sends in trabecule, which divide and subdivide at first in a 
rather irregular manner; but ultimately they form in all cases very regular alveoli, 
having about the same average dimensions as those of the interrenal of the Elasmo- 
branchs (g. v.), strikingly suggestive of those of secreting-glands, and these alveoli are, 
as a rule, completely filled with the suprarenal cells, which are much smaller than 
those of the interrenal of Elasmobranchs. But in some species (Conger conger, 
Anguilla anguilla, and Salmo trutta) there is a more or less empty space in the centre 
of each alveolus (Pl. XIV. figs. 45 & 46, e.sp.). These spaces are, however, never 
complete; one finds scattered cells, nuclei, and shreds of protoplasm, which suggest 
that this space does not exist in life. Most probably the central cells of each alveolus 
are more loosely connected together than the rest, or possibly they suffer a very rapid 
post-mortem change!. In by far the majority of Teleosts a very regular row of nuclei 
can be traced round the circumference of each alveolus, and in many cases their cell- 

1 [Pettit (Joc. cit.) has laid great stress on this peculiar structure of the alveoli in the Eel, looking upon it as 
the fundamental type of a suprarenal gland. He finds a true physiological compensatory hypertrophy of one 
gland after removal of the other, and urges that the body is a true secreting-gland. But he seems to be totally 
unaware that the known suprarenals of the Eel and other Teleosts correspond to the cortex only of the 


Mammalian gland, and that all we know about the function of the suprarenals so far is confined to the 
medulla.—S. V., 10. 1. 97.] 


SUPRARENAL BODIES IN FISHES. 71 


outlines can also be perceived. In some cases, as in the Trout, the alveoli are very 
distinctly marked, and have every appearance of being not closed vesicles, but in 
communication one with another. 

The essential suprarenal cells I find to be of two kinds. The first are irregular 
branched cells, most often tending towards a spindle-shape, about 7-8» in length, 
with very large prominent nuclei (Pl. XIV. figs. 48 & 49). These nuclei are often as 
large in diameter as the widest part of the cell (7. ¢., about 3:5 ») (Pl. XIV. fig. 48), 
and appear to bulge it out at that point. In the nucleus can be seen a number of 
small dark nucleoli, and often a distinct nuclear network. The protoplasm of these 
cells is distinctly granular. 

The second kind of cell is usually irregularly triangular, having generally a greatest 
length of about 6°5 or 7, with a small dark nucleus not more than 2:5 » in diameter 
(Pl. XIV. fig. 49). 

These two kinds of cells are readily distinguished, and both are usually present in 
any given specimen, but in very varying proportions. 

I have doubted whether these appearances might be simply due to treatment or to 
accidental modes of staining, but I believe they depend in some way on the functional 
activity (for the suprarenals are not mere embryonic remains), stage of development, or 
the age of the fish, as their relative occurrence is found to differ very widely even 
among closely allied species. Thus in Gadus eglefinus I found nearly all the cells had 
large round clear nuclei and nuclear figures, while in Merluccius vulgaris the cells 
appeared almost all to have small dark nuclei!. ‘This point I hope to have the 
opportunity of investigating by taking series of specimens of different ages, and after 
different modes of feeding, starvation, poisoning, &c. At present I must leave their 
true nature quite an open question. 

There is no distinction to be made out between cortex and medulla in this order of 
Jishes, or, rather (as appears probable by comparison with other groups of Vertebrates), 
the suprarenal consists entirely of cortex *. 

I have made careful preparations of the suprarenals of all the species whose gross 
anatomy is given above, but, as I have said, these do not offer any great variety among 
themselves. It may, however, be useful to indicate briefly the kind of variation one 
finds in the different families by a few illustrative extracts from my laboratory note- 
book. Perhaps one of the most characteristic appearances is found in the Murenide. 
In this family (Pl. XIV. figs. 45 & 46) the alveoli are rounded, and contain cells poly- 
hedral or angular in shape. These are arranged for the most part in a single tier 
around the circumference of each alveolus, leaving an irregularly-shaped cavity in the 

? T have even found that in one specimen of Mullus barbatus there was a yast majority of one kind of 
cell, and in another specimen of the same fish a corresponding preponderance of the other kind. 

2 [Diamare (Joc. cit.) has arrived at the same conclusions as myself as regards the homology of the interrenal 
body in Elasmobranchs with the suprarenal bodies of Teleostei. It follows that the representative of the supra- 
renal medulla is absent in Teleostean fishes.—S. V., 10. 1. 97.] 


72 MR, SWALE VINCENT ON THE 


centre (Pl. XIV. fig. 46, ¢.sp.). This appearance is described above, and some expla- 
nation of it has been offered. The alveolar walls are permeated by capillary blood- 
vessels (Pl. XIV. fig. 46, cap.). 

In the Pleuronectide the appearance under a low power is very like that of a 
lymphatic gland, but under a high power an alveolar arrangement can always be seen, 
and the nuclei are seen to belong to more or less closely-packed cells of the two 
varieties described above. In Plewronectes limanda distinct acini are often seen almost 
like those of the mammalian pancreas, filled with cells of spindle-shape, with large round 
nuclei and very distinct nuclear figures. There is a tendency towards a clear space in 
the central part of each. In Rhombus levis the acini are quite filled with cells, which 
are more rounded than in other members of this family, and there is generally more 
cell-protoplasm. 

In the Gadide the individual members do not differ very widely from each other. 
In Gadus morrhua the cells are pear-shaped, oval, rounded or irregular, and in the 
specimens I have examined nearly all have large round figured nuclei. In Gadus 
eglefinus the appearance is peculiar: the acini appear to be occupied by scarcely 
anything more than free nuclei with nuclear figures, the cell-protoplasm being scanty 
and irregularly branched. Molva vulgaris shows the acinar arrangement very well, 
with a very regular ring of nuclei round the outside of each alveolus. 

Pl. XIV. fig. 47 represents the appearances in the suprarenal of the Wolf-fish 
(Anarrhichas lupus). The alveoli are elongated, and there is an incomplete central 
space (Pl. XIV. fig. 47, ¢.sp.). Thus it is possible to consider that there are two chief 
types in the alveolar arrangements of teleostean suprarenals: one, as in the Murenide, 
is not unremindful of the testis in some animals (cf. Mihalkovics, 14), while the other 
is more suggestive of the mammalian pancreas. 

The suprarenals of the Sunfish (Orthagoriscus mola) have a very peculiar structure, 
in that the acini are very long and tubular. 


4. DIPNOI. 

In Protopterus annectens, Parker (17) describes “around the kidney, but more parti- 
cularly along its dorsal and outer sides, masses of brown cells, which in appearance 
remind one of the adrenal bodies of Amphibia,” and he suggests the enquiry “ whether 
they or the lymphoid cells which give rise to them have anything to do with the 
adrenals,” 

I have examined this point with some care, and I have been able to verify in every 
detail Parker’s account of this lymphoid tissue, both round the kidneys and along the 
alimentary canal, and forming the substance of the spleen. This I was enabled to do 
by means of two specimens of Protopterus which Prof. Parker was kind enough to 
send me. 

By the kindness of Prof, Bridge, I have also been able to examine the kidneys and 


SUPRARENAL BODIES IN FISHES. 73 


surrounding tissue of Lepidosiren. The specimen was a long-preserved one, and L 
could make out nothing with the naked eye. On cutting sections, however, the same 
pigment-patches were obvious in the perirenal tissue as in Protopterus; if anything, 
these were more abundant in Lepidosiren. Here, too, the resemblance to suprarenals 
was greater than in Protopterus, for I found not only small patches or individual 
pigment-cells, but large rounded accumulations, which represent not continuous 
columns, but spherical masses, thus approximating anatomically to adrenals. 

This tissue is of such great interest from many standpoints that I hope before long 
to be able to give a fuller description of it in another place. I am persuaded that it is 
really a “ large-celled adenoid tissue,” and some of the chief varieties of leucocytes can 
be recognized in some of my sections. But the appearance in places is so very glan- 
dular that Prof. Schafer, who has been kind enough to look at some of my slides, said 
he should be very much inclined to think it might be suprarenal tissue, if a material 
so very like it were not found in other places. In fact, the intestinal walls show 
abundance of the same structure, and the spleen is very little different 1. 

I have carefully compared both these with the pigment-patches in other parts, and 
find that they are quite comparable to these, so that, on the whole, I am inclined to 
the view that they have nothing to do with the adrenals. 

Nevertheless, from @ priori considerations, I believe that adrenals of some sort are 
almost certainly present in the Dipnoi. These fishes closely approach the Amphibians 
in many respects, and I am persuaded that could one obtain perfectly fresh specimens of 
large size, suprarenals of a type resembling that of the Amphibians would be found 2. 


My investigations upon the structure of the suprarenal capsules in Fishes all point to 
their being blood-vascular secreting-glands of two distinct kinds, and this coincides 
entirely with the most recent views as to the function of the Mammalian organs. 


VI. Tue RELATION OF THE SUPRARENAL Bopiges TO THE HEap-Kipyey. 
1. Historical. 


Balfour (23 and 24) first clearly made out that in Actpenser and Lepidosteus the 
anterior dilatations of each kidney consist not of kidney-substance, but of lymphatic 
tissue. He says, “ Zhus the whole of that part of the apparent kidney in front of the 


1 But, after all, it may be that there is no “‘great gulf” fixed between the leucocytes of adenoid tissue 
and the cells of a secreting-gland. I have recently seen a preparation from a case of Paget’s disease of the 
nipple in the human subject which appeared to show a transition between the epithelial cells of the ducts and 
ordinary leucocytes. See also Beard (Anat. Anz. ix. Band, Nr. 15, p. 481, and his references to Kélliker and 
A. Prenant). 

* [Since the above has been in type, Petitt (Joc. cit. pp. 67-68) has claimed to have found the suprarenals in 
Protopterus. He says that in general form and relations they resemble those of the Teleostei, while in minute 
anatomy they are rather like those of Batrachians. But he gives no histological details, and says nothing 
about cortex and medulla.—S. V., 10. 1. 97.] 

VoL, XIV.—ParT 1. No. 5.—April, 1897. L 


74 MR. SWALE VINCENT ON THE 


ureter, including the whole of the so-called head-kidney, is simply a great mass of 
lymphatic tissue, and does not contain a single uriniferous tubule or Malpighian body.” 
He next proceeded to examine Hsou lucius, Osmerus eperlanus, Anguilla anquilla, and 
Lophius piscatorius among Teleosts, and found pretty much the same condition as in 
Ganoids. He does not speak very positively about the Angler, but believes that 
tubules were present in all parts of the kidney. He sums up “that the pronephros, 
though found in the larve or embryos of almost all the Ichthyopsida, except the Elasmo- 
branchii, is always a purely larval organ, which never constitutes an active part of the 
excretory system in the adult state.” Balfour describes the lymphatic tissue of the 
so-called head-kidney as formed of trabecular work and cells resembling a lymphatic 
gland. This tissue is very vascular, with a regular plexus of very large capillaries, 
which appear to have distinct walls, and which pour their blood into the posterior 
cardinal vein as it passes through the organ. With regard to the function of this 
tissue Balfour suggests that this is either the formation of lymph-corpuscles or of 
blood-corpuscles, and he was inclined provisionally to regard it as a lymphatic gland. 

Parker (Brit. Assoc. Reports, 1882) stated that in many instances the mesonephros 
has grown forward in front of the air-bladder, and taken the place of the pronephros. 

In studying Dactylopterus volitans, Calderwood (25) found the pronephros entirely 
separated from the body-kidney, and situated anterior to the abdominal cavity in the 
same transverse plane as the heart. On section this head-kidney appeared to bea 
functional kidney, only it did not contain so many tubules as the body-kidney. In 
Cyclopterus lumpus he finds that not till the fish has become sexually mature does its 
head-kidney commence to degenerate. Calderwood concludes that in adult Teleosteans 
the renal function is performed in some instances by the body-kidney only, in others 
by the head-kidney only, and in others, probably a very limited number, by both 
the body- and head-kidneys. Besides Dactylopterus he is aware of only one instance 
where the head-kidney is described as possessing tubules and Malpighian bodies, 
viz. Fierasfer (Emery 26 and 27)}. 

Weldon (35 and 36), in a paper on the head-kidney of Bdellostoma, describes a 
lobulated glandular body lying in front of the secreting part of the kidney, which he 
suggests is “a part of the embryonic kidney, modified in connection with the needs of 
the animal to perform some unknown function in the elaboration or purification of 
the blood.” Then he instances Balfour’s description of the lymphatic head-kidney in 
Teleosts and Ganoids as being a further illustration of a modification of a part of the 
embryonic kidney into an organ like a lymphatic gland. Next he suggests that in 
higher Vertebrates the suprarenal capsules are similarly modified portions of the 
primitive kidney. He continues, “Jn Teleostei suprarenals are, at all events, frequently 
absent”; or, as I would suggest, they are represented by the greatly-metamorphosed 

+ In a later paper (Scientif. Trans. Roy. Dublin Society, vol. v. (ser. ii.) ix. 1895, Survey of Fishing- 


grounds, &c.) Holt and Calderwood add to this list Mora mediterranea and all the species of Macruride. 
* The italics are mine.—S. Y. 


SUPRARENAL BODIES IN FISHES. 75 


head-kidney described by Balfour. In other cases where suprarenals have been detected ', 
they have always been attached to the surface of the kidney.” Later on he talks about 
“the very general absence of suprarenals as separate structures in Teleosteans” 1, as if 
their lying upon the surface of the kidney abolished them as ‘‘separate structures”! 
More recently, Miss Kirkaldy (33) has expressed much the same views as Weldon. 
She remarks, “It may therefore be concluded that the pronephros in Myzine 
represents the mesoblastic part of the suprarenal bodies, which have been shown by 
Prof. Weldon to be derived from the anterior part of the mesonephros in the 
higher Vertebrata.” 

I have already shown conclusively that suprarenals are not frequently absent, but 
are probably always present. It only remains to deal with their supposed relation to 
the head-kidney. Weldon’s teaching seems to have found pretty general acceptance. 
Thus Auld? says, referring to the lymphatic tissue of the head-kidney, “ Now in 
Teleosteans and Teleosteoid Ganoids which possess this tissue no suprarenal organs are 
found ', and hence it is to be concluded with Balfour (!) that they are represented by 
this tissue.” Balfour, at any rate in the paper quoted, says nothing of the kind; but 
the comparative portion of Auld’s paper is of no importance, except as illustrating the 
injurious influence of Weldon’s inaccuracy. 

It will have been noticed that although many writers have declared suprarenals to 
be often absent in Teleosts, none of them have stated definitely in what species we 
may look for them in vain. The above quotations imply a very general belief some- 
thing to this effect: that suprarenal bodies are rare phenomena in Teleosts and 
Ganoids, and that when present there is no lymphatic head-kidney, while when they 
are absent their place is taken by this structure. 

The following quotation from Wiedersheim (Lehrbuch der vergl. Anat. der 
Wirbelthiere, 2° Aufl., Jena, 1886) shows the need for clearing up this point :— 

“Bei Teleostiern sind die Nebennieren nicht tiberall in klarer und tiberzeugender 
Weise nachgewiesen, wo dies aber der Fall ist, handelt es sich, wie friiher schon 
angedeutet wurde, um Beziehungen zu der in lymphoides (adenoides) Gewebe umge- 
wandelten Kopfniere. (Dies gilt nach W. Weldon auch fiir die Cyclostomen | Bdello- 
stoma Forsteri}.) In andern Fallen, aber, sind sie enge mit der Niere verbunden.” 

The plain facts of the case are quite the contrary, and are these :— 

Suprarenal capsules are present in certainly the majority of Teleosts and Ganoids, 
and I believe in all. Further, as far as I know at present, the purely lymphatic head- 
kidney is present in all Ganoids and in all Teleosts except Lophius, Dactylopterus, 
Fierasfer, and, as will be stated below, Orthagoriscus mola °. 


1 The italics are mine.—S. V. ? Brit. Med. Journ., Oct. 6th. 1894. 
* And, according to Holt and Calderwood, in Mora mediterranea and the Macruride. 


L2 


76 MR. SWALE VINCENT ON THE 


2. The Nature of the Intertubular Material in the Kidneys. 


The lymphatic head-kidney is not in any true sense a very specialized portion of the 
kidney. By “lymphoid” tissue we mean a variety of retiform connective tissue, in 
which the meshes of the network are filled up to a large extent with lymph-corpuscles. 
These differ from the white corpuscles of the blood in that their nuclei show a net- 
work. Some are as large as white blood-corpuscles, others (the majority) have a 
minimum of protoplasm, and appear as almost free nuclei. This structure is found 
typically in a mammalian lymphatic gland. 

Now, on examining the kidney of any fish, the tubules appear to be separated by a 
variable, often a large, amount of intertubular material, which presents all the features 
given above characteristic of lymphatic or adenoid tissue. This appearance presents a 
marked contrast to the compact mass of tubules and Malpighian bodies one sees in a 
mammalian kidney. The kidney of the frog appears to be intermediate between the 
mammalian and fish’s kidney, in respect of the amount of this intertubular adenoid tissue. 

This tissue varies in amount in different regions of the kidney, but on the whole 
forms a fairly even bed for the tubules. In Teleosts, as one approaches nearer and 
nearer to the anterior end of the organ, the amount of adenoid tissue becomes greater 
and greater until in most cases the extreme anterior extremity of the apparent kidney 
consists entirely of adenoid tissue. 


3. Histology of the Head-Kidney. 


SrructurRE oF THE Heap-Kipney or THE Line (Molva vulgaris)—The tissue was 
hardened in spirit, stained in bulk with hematoxylin and eosin, and imbedded in 
paraffin; sections were then cut with the rocking microtome. 

On examination with a low power, the organ is seen to be obviously a lymphoid 
structure. Not a trace of kidney-tubules or Malpighian bodies is to be found in any 
part of the section. The organ is enclosed in a very definite fibrous capsule. There 
are large blood-vessels ! in abundance full of red blood-corpuscles. 

With a high power the substance of the head-kidney is seen to be made up of a 
delicate reticulum enclosing in its meshes closely-packed leucocytes with many red 
cells apparently free in the reticular spaces, as well as in the large capillaries. In 
addition, there are many hexagonal crystals of a red colour, most probably oxy- 
hemoglobin *, and masses of dark pigment-crystals of acicular form, evidently some 
derivative of hemoglobin. 

I have found practically the same structure in the head-kidneys of the Pleuronectide, 
Gadide, and in fact in all Teleosts I have examined. I have only found this lymphatic 


* And, in some cases, blood-sinuses (vide infra). 

* The crystals of oxy-hzemoglobin obtained from the blood of the Squirrel and Hamster (Halliburton) are 
described as of this shape, also those from the Mouse (Bojanowski) and Horse (Hiifner and Biicheler). It is 
tolerably certain that these crystals are of the same character. 


SUPRARENAL BODIES IN FISHES. 77 


head-kidney absent in Zophius piscatorius and Orthagoriscus mola. Other authors 
have added some few more species, see above, also summary below. A few detailed 
" illustrations will suffice. 

Cyclopterus lumpus.—I have particularly examined the kidney of this species. It 
will be seen (Pl. XIII. fig. 40) that it has a peculiar shape. 

At its anterior extremity are two dark red spherical masses of tissue, only attached 
to the rest of the kidney by connective tissue. On microscopical examination these 
are found to consist of an extremely vascular lymphoid tissue, with no trace of urinary 
tubules or Malpighian bodies !. 

The divided portion of the kidney on each side is represented by two very thick 
masses nearly half the total length of the kidney (fig. 40). 

I find that almost the anterior third of this part, as well as the dark red bodies in 
front, is composed entirely of lymphoid tissue. As one approaches the junction of the 
anterior and middle thirds of this part of the kidney, one or two tubules make their 
appearance in the central portion of the section. In the middle third tubules are 
more frequent, while in the posterior third the appearance is almost that of “ body- 
kidney ” generally. 

So that we have in Cyclopterus a well-developed lymphoid head-kidney, with an 
anterior part completely constricted off. This constriction may possibly throw some 
light upon other masses of lymphoid tissue which one finds in the neighbourhood of 
the kidney in Elasmobranchs as well as in Teleosts. 

With regard to Lophius piscatorius, which was supposed by Hyrtl to have only a 
head-kidney, I find with Balfour tubules and Malpighian. bodies in all parts of the 
kidney-mass except the extreme anterior end, where Malpighian bodies are absent. 
Here, too, there is more adenoid tissue than elsewhere, and it appears more probable 
that the whole kidney is contracted longitudinally, and that the lymphoid remains of 
the pronephros have been encroached upon by the mesonephros than that the whole 
of the ordinary excretory organ has been lost, and the pronephros has remained as the 
functional kidney. 

In Anguilla anguilla I found in a specimen 68 em. long, with a total length of 
kidney of 25 cm., that exactly the anterior three-quarters of the ununited portion of 
kidney on each side was entirely lymphoid, 7.¢., half the total length of the kidney. 
At the mid-point of the kidney-length tubules begin to appear and rapidly increase in 
number till the normal secreting structure is reached. 

In the Sturgeon the whole of the part /.&. (Pl. X. fig. 8) as far as the dotted line is 
purely lymphoid. ‘There is also a small area of lymphatic tissue at the extreme hinder 
end of the kidney. 

In the Sunfish all parts of the kidney contain urinary tubules, even the extreme 


1 [This structure corresponds to that of a ‘* heemo-lymph gland ” (see Vincent and Harrison, Journ. of Anat. 
and Phys., Jan. 1897).—S. V., 10. 1. 97.] 


78 MR. SWALE VINCENT ON THE 


anterior end which reaches far into the head. So that this species must be added to 
the list of those which have no purely lymphatic head-kidney. 


4, Function of the Lymphatic Head-Kidney. 


I am convinced that the appearances above described (blood-corpuscles free in the 
tissue spaces, crystals of oxy-hemoglobin and other derivatives of hemoglobin) point 
to a blood-destroying function of the lymphoid anterior end of the kidney, and very 
possibly also of the rest of the intertubular material, and this appears not unreasonable 
since lymphatic glands in Man can probably carry on this function. ‘They appear to 
do so, at all events after removal of the spleen, and it is not conceivable that an organ 
should be able to take on in an emergency duties which it had never performed to the 
slightest extent previously !. 


5. Relation of the Suprarenals to the Head-Kidney. 


I have already sufficiently disproved the idea that where one of these is present the 
other is absent. I have further, by careful measurements, attempted to make out if 
there were any inverse ratio between them as to respective bulk, but in this I have 
failed entirely. So that I am forced to conclude that there is no anatomical relationship 
whatever between them. 

Physiologically one cannot be so certain, but it seems very likely that the head- 
kidney functionates as a lymphatic gland, while the suprarenals, in all probability, are 
secreting-glands, which minister to the needs of the blood, just as in the higher Verte- 
brates. Functional relationship is, then, equally improbable. 

Grosglik (29) considers that the head-kidney of adult Teleosts consists of two parts, 
the degenerated pronephros and the cortical part of the suprarenals, and that the 
known suprarenals of Teleosts correspond to the medullary substance in the Amniota”. 

It is certainly curious that we find the lymphatic head-kidney just in those cases 
where only one portion of the suprarenal appears to be represented. But, in my 
opinion, the part which is not represented in Teleosts and Ganoids is the medulla and 
not the cortex, so that, if the head-kidney has anything to do with the suprarenals at 
all, I should expect that it would represent the medulla. But, satisfactory as such a 
conclusion would be, I cannot find any grounds for it whatever. There is nothing in 
the structure of the degenerated pronephros which suggests any connection with supra 
renal structure of either kind. 

The question as to the physiological correspondence of the suprarenal in Fishes to 
the organs in higher Vertebrates, I hope to have the opportunity of settling at an early 
date by means of direct experiment ?. 


* [It has been mentioned above that Balfour considered the function to be that of formation of red 
corpuscles. This view has also been held by Emery (26), and Bizzozero and Torre, Mem. Accad. Lincei Roma, 
vol. xvii, 1883-84.—S. V., 10. 1. 97.] 

* An t, Anz, xiii. Bd. Nr. 1 & 2 (1897). 


SUPRARENAL BODIES IN FISHES. 79 


VII. Summary ann Conciustons. 
A. Gross Anatomy. 


My investigations in this department have led me to conclude that :— 

1. Suprarenals are almost certainly present in all Elasmobranchii, Holocephala, 
Ganoidei, and Teleostei, and very probably in Dipnoi also 1, 

2. In Elasmobranchs the interrenal body is totally distinct and separate from the 
segmental bodies, and there is no kind of connection between them. 

3. There is no “third kind of body” in relation to the kidneys, as Balfour surmised 
there might be. 

4. The yellow bodies scattered in the Sturgeon’s kidney are the true suprarenals in 
Ganoids, and correspond to the interrenal of Elasmobranchs and the suprarenals of 
Teleosts. 

B. Histology. 

The following points in histology are either new or have not been sufficiently 
emphasized by previous observers :-— 

1. In Elasmobranchs the interrenal body consists of definite alveoli, containing cells 
with large nuclei, and curious structures resembling “ demilune” cells of mucus-glands. 
This structure indicates its “secretory” nature and its analogy to the suprarenals of 
Teleosts and the cortical part of the suprarenals of the higher Vertebrates. 

2. The segmentally arranged bodies in Elasmobranchs have no cortex and medulla, no 
definite alveoli, and cell-outlines seen only with difficulty. There are, however, in some 
places branched pigment-cells which appear to communicate together by their processes. 
The nuclei are of very varying size. 

3. These segmental bodies, having such a different structure from the interrenal, are 
probably different also both morphologically and physiologically 2. 

4. That the fibrils which Balfour observed in the substance of the paired bodies were 
probably many of them only connective tissue, that the significance of the relations of 
the paired suprarenals to the sympathetic has been much overstated, and that their 
relation to the vascular system is probably much more important. 

5. The suprarenals in the Sturgeon have a structure which is definitely alveolar and 
cellular—analogous to the interrenal of Elasmobranchs and the suprarenats of Teleosts. 

6. The interrenal of Elasmobranchs and the suprarenals of Ganoids and Teleosts are 
in their essence ‘ secreting-glands,” as the Mammalian organ is now believed to be. 

7. The segmental suprarenals of Elasmobranchs are also secreting-glands, though 
probably of a different nature. 


C. Head-Kidney. 


1. That all Teleosts and Ganoids, as far as is known at present (eacept Lophius, 
Dactylopterus, and Fierasfer, Orthagoriscus mola, Mora mediterranea, and all the 
species of Macruride), have a “ lymphatic head-kidney.” 


1 See footnote 2, page 73. * See Anat, Anz. xiii, Bd, Nr, 1 & 2 (1897), 


80 MR. SWALE VINCENT ON THE 


2. That this head-kidney is a lymphoid organ with a probable blood-destroying 
function. 

3. That the kidney of fishes consists of two totally distinct parts—(1) The secreting- 
tubules and Malpighian bodies, and (2) an adenoid intertubular material. 

4. That the head-kidney in adult Teleosts is not a “specialised” portion of the 
primitive kidney, but simply an increase, at the anterior end, of the lymphatic tissue 
which exists between the tubules throughout the kidney. 

5. That there is no anatomical or physiological relationship of any kind between the 
suprarenals and the head-kidney. 


VIII. Brstiocraray. 


A. On the Suprarenal Bodies. 


1. Batrour, F.M. A Monograph on the Development of the Elasmobranch Fishes. London, 1878. 

2. Bearp, J. The Inter-relationships of the Ichthyopsida. Anat. Anz., V. Jahrgang, 1890. 

3. Cuevret, R. Sur |’Anatomie du Systéme nerveux grand sympathique des Elasmobranches et 
des Poissons Osseaux. Archives de Zoologie expér. et gén. 2° série, tome v. bis (1887), 
Supplémentaire. 

4. Cuevret, R. Recherches anatomiques sur le systéme nerveux grand sympathique de 
YEsturgeon. Arch. de Zool. Expér. et gén. 3° série, tome 11. 1894. 

Also a paper bearing the same title in the C. R. de Acad. des Sciences, tome cxvii. no. 18, 
2° semestre, p. 441. 

5. Esertu, —. Article “Suprarenal Capsules ” in Stricker’s ‘ Handbuch,’ 1871. 

6. Ecxrr, —. Der feinere Bau der Nebennieren, etc. Braunschweig, 1846. 

7. Frey, H. Article “ Suprarenal Capsules ” in Todd’s ‘ Cyclopzedia,’ vol. iv. (1852). 

8. Gtintuer, A. Description of Ceratodus. Phil. Trans. vol. clxi. 1871. 

g. Hyatt, J. Lepidosiren paradora. Abhandl. d. Bohm. Gesellsch. Prag, 1845. 

10. Hyrtt, J. Das uropoetische System der Knockenfische. Sitz. Wiener Akad. 1851. 

11. Leypic, F. Beitrage zur mikros. Anat. etc. der Rochen und Haie. Leip. 1852. 

12, Lrypic, F, Anatomisch-histologisch. Untersuchungen iiber Fische u. Reptilien. Berlin, 1853. 
13. Leypic, F. Zur Anatomie und Histologie der Chimera monstrosa. Miiller’s Archiv, 1851. 


14. MrAtxovies, —. Entwick. d. Harn. u. Geschlechtsapp. der Amniota. ui, Die Geschlechts- 
driisen. Internat. Monatschr. f. Anat. etc. Bd. ii. 1885, pp. 389-402. 
15. M‘Kenziz, —. Contributions to the Anatomy of Amiurus. Proc. Canad. Inst. of Toronto, 


n. s. vol. ii. no. 8, 1884. 

16. Muxurr, J. Vergleichende Anatomie der Myxinoiden. Schluss. Abhandl. d. Akad. der 
Wissenschaften zu Berlin, 1843, S. 113. 

17. Parxer, W. N. On the Anatomy and Physiology of Protopterus annectens. Trans. Roy. I. 
Acad. vol. xxx. part iii. 1892. 


18, Rerzius, —. Observationes in Anatomiam Chondropterygiorum. Lund, 1819. 4to. 
19. Semper, C. Urogenitalsystem d. Plagiostomen. Arb. zool.-zoot. Inst. z. Wiirzburg, vol. 11. 
1875. 


20. Stannivs, H., in Miiller’s Archiv, 1839, 8. 97 ff. 
21. Srannius, H. Vergleichende Anatomie. Berlin, 1846. 
22, Srannius, H., u. vy. Srrsotp. Handbuch der Zootomie. 2% Aufl, 1854. 


23. 


24. 


25; 


26. 


27h 
28. 


29. 
30. 


31. 
32. 
33- 
34. 
35: 


36. 


SUPRARENAL BODIES IN FISHES. 81 


B. Head-Kidney. 


Batrour, F. M. The Pronephros of Teleosteans and Ganoids. Brit. Assoc. Reports, 1881, 
p. 721. 

Batrour, F.M. On the Nature of the Organ in adult Teleosteans and Ganoids which is 
usually regarded as the Head-Kidney or Pronephros. Q. J. M. S. vol. xxii. p. 12 
(Jan. 1882). 

Catprrwoop, W.L. The Head-Kidney of Teleostean Fishes. Journ. of the Mar. Biol. Assoc. 
of United Kingdom, N. §., vol. ii. no. 1 (May 1891). 

Emery, C. Zur Morphologie der Kopfniere der Teleostei. Zool. Anz. vil. pp. 742-744. 
(Atti dell’ Accademia dei Lincei, 1882.) 

Emery, C. Le specie del genere Fierasfer nel Golfo di Napoli. Leip. 1881. 

Fiztp, H. H. Development of Pronephros and Segmental Duct in Amphibia. Bull. Mus. 
Comp. Zool. xxi. (1891) pp. 201-340 (8 pls.). 

Groseiix, —. Zur Morphologie der Kopfniere der Fische. Zool. Anz. 1885. 

Janostx, —. Bemerkungen iiber die Entwickelung der Nebennieren. Arch. f. mikr. Anat. 
Band xxii. pp. 738-746, Taf. xxvii. (1883). 

Juncersren, H. F. E. Development of Kidneys in Sturgeon. Zool. Anz. xvi. (1893), 
pp. 464-467, 469-472 (1 fig.). 

Juncursren, H. F. E. Development of Kidneys in Amia calva. Zool. Anz. xvii. (1894), 
pp- 246-252 (5 figs.). 

Kirxatpy, J. W. On the Head-Kidney of Myzxine. Q. J. M. S. vol. xxxv. (Jan. 1894), 
p- 353. 

Semon, R. Relation of Mesonephros to Pronephros and Suprarenal Bodies. Anat. Anz. v. 
(1890), pp. 455-482 (8 figs.). 

Wetpon, W. F. R. On the Head-Kidney of Bdellostoma, &e. Q. J. M.S. xxiv. p. 171 
(April 1884). 

Wetpon, W. F. R. Suprarenal Bodies of Vertebrata. Q. J. M.S. vol. xxv. p. 127. 


EXPLANATION OF THE PLATES. 


Reference-letters to figs. 1-40 (Plates [X.—XIII.). 


a.a., axillary artery; a.i.r., anterior broken-off portions of the interrenal body ; 


ao., aorta; aa@.h., anterior pair of suprarenal bodies; f-¢., bands of fibrous tissue ; 
h.a., anterior limit of hemal arch; h.k., head-kidney; 7.a., intercostal arteries ; 
ir., interrenal body; &., kidney; /.%., lobe of kidney-substance ; x., nerves 
running through deep fissures in head-kidney; @., cesophagus cut across; 
s.r, suprarenal bodies; sy., main chain of the sympathetic; sy.g., sympathetic 
ganglion; sy.pl., sympathetic plexus ; wr., ureter. 


Figs. 1-40 represent the outlines of the kidneys in the several species examined, 


with the suprarenals and interrenals shaded in. In the Teleosts (figs. 9-40) 
the suprarenals when on the ventral surface are drawn in plain lines, when 
on the spinal surface in dotted lines. 


VOL. XIV.—PaART 111. No. 6.—April, 1897. M 


Fig. 


MR. SWALE VINCENT ON THE 


PLATE IX. 


. 1. Ventral view of the kidney, suprarenals, and interrenal of Chimera monstrosa, 


showing the smooth, oval, anterior pair of suprarenal bodies. 1. 

2. Dissection of Scyllium canicula (young female specimen) giving a ventral view 
of suprarenals and interrenal. The parovarium has been dissected away. 
This drawing may be taken as a typical representation of these bodies in 
Elasmobranchs. ‘lhe connections with the sympathetic are indicated to 
some extent in the anterior part of the figure. The suprarenals were 
displayed by Semper’s chromic acid method. x 4. 


ig. 2 A. From the same preparation as the last. First three suprarenals of each 


side. xX 2. 


. 3. Suprarenals of Scyllium canicula brought out by Chevrel’s osmic acid method. 


Their irregular outline is well shown, and many of the sympathetic fibrils 
are stained black. x 1. 


. 4. Ventral view of kidneys, suprarenals, and interrenal of Scylliwm catulus. 


This drawing shows fairly well the relations to the sympathetic. There is a 
large plexus anterior to and outside the axillary hearts, with occasional 
ganglia. x 4. 


PLATE X. 


g. 5, Suprarenals, &c., of Acanthias vulgaris, shows sympathetic relations in anterior 


part. The interrenal is not quite in the middle line, but is laid on the 
surface of the left kidney. x 1. 


. 6. Ventral view of kidneys, &c., of Raja batis. ‘This drawing represents a not 


unusual condition in the Rays, in which there is a bridge-like communica- 
tion between the interrenals of the two sides. The sympathetic is shown to 
some extent about the middle of the left kidney. x lI. 


. 7. Same view in Raja maculata. The anterior separated portions of the inter- 


renals show processes or claws on their outer sides. 1. 


. 8. Ventral view of kidney and suprarenals of Acipenser sturio. Part of the 


kidney-substance has been dissected away so as to display the suprarenal 
bodies distributed throughout the organ. xX 75. 


. 9. Ventral view of kidneys and suprarenal of Orthagoriscus mola. The kidneys 


are a very peculiar shape. ‘There is one suprarenal attached by fibrous 
bands behind the right kidney. x i. 


. 10. Kidney and suprarenals of Conger conger, suprarenals on ventral surface of 


kidney. Xx +. 


. 11. Clupea harengus, suprarenals on spinal surface of kidney. X 1. 


Dw pw bl lM bv 


LW bo 
Or 


(or) 


co oO 


an 


wo op OF OF OF DO bo bb 
o 


i) bo 


wo 
ho 


ig. 40. 


a 
mm op DW 1 OS © oo 


~J 


SUPRARENAL BODIES IN FISHES. 


PLATE XI. 


Esox lucius. Suprarenals in kidney-substance. 4. 


. Salmo salar. Five suprarenals on spinal surface of kidney. > 4. 

. Osmerus eperlanus. Suprarenals projecting behind hinder end of kidney. 
. Leuciscus rutilis. Suprarenals on ventral surface. x 1. 

. Leuciscus cephalus. Suprarenals on ventral surface. 1. 

ig. 17. 


. Gadus morrhua. Suprarenals on ventral surface. 4. 


Leuciscus vulgaris. Suprarenals on spinal surface. x 1. 


Gadus eglefinus. Suprarenals on ventral surface. 1. 
Merluccius vulgaris. Suprarenals on ventral surface. x 


bole 


Molva vulgaris. Suprarenals on ventral surface. x 4. 
Pleuronectes flesus. Suprarenals on ventral surface.  X 1. 
Pleuronectes limanda. Suprarenals on spinal surface. x 1. 
Pleuronectes platessa. Suprarenals on spinal surface. x 1. 


PLATE XII. 


5. Hippoglossus vulgaris. Suprarenals on spinal surface. xX 4. 


83 


adle 


Hippoglossoides limandoides. Suprarenals projecting from hinder end of 


kidney. x 1. 
Solea vulgaris. Suprarenals on spinal surface.  X 1. 


. Rhombus levis. Suprarenals on spinal surface. 1. 

. Perca fluviatilis. One suprarenal on spinal surface. X 1. 

. Mullus barbatus. Suprarenals on spinal surface.  X I. 

0a. Another specimen, showing hinder end of kidney and five suprarenals. 
. Pagellus centrodontus. Suprarenals on spinal surface. X 1. 

. Cottus gobio. Suprarenals on spinal surface. 1. 


Trigla pini. One suprarenal on spinal surface. 1. 


ail 


Trigia lyra. Two suprarenals off left side of tail of kidney, attached by bands 


of fibrous tissue. > 1. 
Scomber scomber. Suprarenals on the ventral surface of the kidney. xX 


. Zeus faber. Suprarenals on spinal surface of the kidney. X 1. 
. Anarrhichas lupus. Left suprarenal on ventral surface of kidney, right one 


on spinal surface. x 4. 
Notice peculiar shape of the kidney-masses.  X 1. 


PLATE XIII. 


seen from either surface.  X 4. 
Cyclopterus lumpus. Suprarenals on spinal surface of kidney. X 


bole 


bole 


. Lophius piscatorius. Five suprarenals on ventral surface of the kidneys. 


. Mugil capito. One large suprarenal in substance of tail of kidney; could be 


84 


ad. 


vy 


Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


ON THE SUPRARENAL BODIES IN FISHES. 


Reference-letters to figs. 41-49 (Plates XIII., XIV.). 


adenoid tissue of kidney between the tubules; al.w., walls of alveoli; Jid.c., 
blood-corpuscles; ¢., capsule ; cap., capillary blood-vessels ; ¢.sp., central space in 
alveoli; d.c., cells resembling ‘“‘ demilune ” cells; ¢@.c., elongated cells; ., nuclei; 
n.c., nerve-cell; n.net., nuclear network; n/., nucleoli; p.c., branched pigment- 
cells; pr., granular protoplasm; s., septa; s#r., fibrous stroma; ¢., tubules of 
kidney ; v.s., venous sinuses; 2, cells seen to be overlapping. 


41. Section of a suprarenal body of Acanthias vulgaris, showing the capsule, 
trabecule, stroma, protoplasm, and various-sized nuclei. Zeiss H. immers. 
E. P. 2. Drawn with Zeiss’s camera lucida. 

42. Section of a suprarenal body of Scyllium canicula, from about the middle 
region of the abdominal cavity. In this section are seen a large nerve-cell 
and several branched pigment-cells. Zeiss H. immers. E. P. 2. Camera 
lucida. 


. 43. Section of the interrenal body of Raja clavata, showing alveoli of various 


shapes and sizes, filled with cells, many of them elongated. Zeiss 


apochrom., 3," water immers. Camera lucida. 


PLATE XIV. 


44, Section of a suprarenal body of Acipenser sturio. ‘The body was put into 
osmic acid about 12 hours after death, and sections were cut with the 
freezing microtome on the following day. ‘The alveolar arrangement is well 
seen and the cell-outlines are admirably preserved. Zeiss H. immers. 
E. P. 2. Camera lucida. 

45, Section through a portion of the kidney and the two suprarenals of Conger 
conger, showing the renal intertubular material, the low power appearance 
of the suprarenals, and their connexions with the kidney. x 70. 

46. Portion of suprarenal of Conger from same slide as preceding, showing the 
alveoli containing a regular tier of irregular cubical cells, and central spaces 
containing nuclei and shreds of cells. Zeiss apochrom. ;'y" water immers, 
Camera lucida. 


. 47. Section of suprarenal of Anarrhichas lupus, showing a curious serpentine 


arrangement of the alveoli. Zeiss j'y" apochrom. water immers. Camera 


lucida. 


. 48. Separate cells of suprarenal of Pleuronectes limanda. Zeiss apochrom. y'y" 


water immers. Compens. 0:18. 


. 49. Separate cells of suprarenal of Mullus barbatus. Shows the two kinds of 


cells. Same power as fig, 48. 


rans. Zo0t. Soc Vt AN FE IX. 


Geo West & Sons ump. 


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V. A Supplementary Report on the Crustaceans of the Group Myodocopa obtained 
during the ‘Challenger’ Expedition, with Notes on other new or imperfectly known 
Species. By G. Srrwarvson Bravy, M.D., LL.D., F.R.S. 


Received August 28, 1895, read December 3, 1895. 


[PLates XV.-XVII.] 


SINCE the publication of my Report on the Ostracoda of the ‘Challenger ’ 
Expedition I have received from Dr. John Murray and Professor G. O. Sars further 
specimens which were met with during the examination of other groups of Crustacea. 
All of them are pelagic species belonging to the group Myodocopa, and it is with 
Dr. Murray’s approval that I record them here. In addition to the specimens so 
obtained, I am in possession of other undescribed or imperfectly known forms which 
have come into my hands at various times and which are of sufficient interest to make 
their further description desirable. Among these are some collected by my brother, 
the late Dr. H. B. Brady, F.R.S., among the South Sea Islands, and which were 
described by me a few years ago! in the ‘Transactions’ of the Royal Society of 
Edinburgh. At that time, however, I was unacquainted with the soft parts of many 
of the species, and was able to describe the shells only. Some of these descriptions I 
am now able to improve by the addition of anatomical details derived from the 
examination of macerated specimens. 


OSTRACODA., 
Section UWYODOCOPA, 
Family CYPRIDINIDZ. 


Genus CYCLASTEROPE, noy. gen. 


The shell is more nearly spherical than is usual in Asterope. Frontal tentacle stout, 
3-(or 2?)jointed. he first joint of the mandibular foot has a falcate masticatory 
process as in Asterope, but much more elaborately spinous; the second joint, instead of 
being produced backwards in an angular process, bears on its distal margin a large 
tongue-like appendage which extends as far as the extremity of the following joint. 
The last limb (vermiform foot) is very profusely armed with sete, many of the 
segments bearing two or three on each lateral margin. In other respects the anatomy 
is that of Asterope. 

1 “On Ostracoda collected by H. B. Brady, Esq., LL.D., F,R.S., in the South Sea Islands,” Transactions of 
the Royal Society of Edinburgh, vol. xxxv. part 11 (1888). 

VOL. XIV.—PART 111. No. 7.—April, 1897. N 


86 DR. G. 8. BRADY ON MYODOCOPA OBTAINED 


CYCLASTEROPE HENDERSONI, n. sp. (Plate XV. figs. 1-12.) 

Shell of the female (figs. 1, 2) very tumid, subspherical: seen from the side broadly 
ovate; greatest height situated near the middle and equal to two-thirds of the length ; 
anterior extremity somewhat narrowed in comparison with the posterior, which is 
broad and well rounded; the beak narrow, curved, and sharply pointed, notch very 
narrow; dorsal margin boldly arched, but towards the front flattened and rather steeply 
sloping; ventral margin evenly and gently convex. Seen from above the outline is 
subovate, wide in the middle and but slightly tapered towards the extremities, width 
nearly equal to the height; lateral margins strongly convex, anterior extremity rather 
abruptly narrowed, obtuse and slightly produced, posterior rounded. Surface of the 
shell smooth or but very slightly punctated; the anterior margin clothed with very 
small fine hairs; colour yellowish. Length 7 millim. Male unknown. 

Frontal tentacle (fig. 8) stout, two-jointed, the basal joint short ; apical joint twice as 
long, subclavate, constricted at the base, and slightly also in the middle, whence it 
tapers gradually to the extremity. Secondary branch of the antenna (fig. 4) two- 
jointed, geniculated, basal joint bearing several marginal sete, distal joint one long 
ringed seta. The basal joint of the mandibular foot bears a falciform masticatory 
process the convex side of which is, towards the apex, divided into numerous very 
irregular processes (figs. 6-8), those at the apex being slender and falcate, the others 
broader, tooth-like, and marginally denticulated: below this series of processes the 
margin of the organ forms a flattened laminar flange which bears several transverse 
rows of spinules. The last pair of limbs (“‘ vermiform feet”) take the form, usual in 
Cypridinide, of flexuous hollow cylinders containing in the interior a spirally coiled 
chitinous thread: the free extremity (fig. 9) has four sets—two on each side—of spine- 
like teeth, and bears a vast number (about two hundred and fifty) of fine barbed set 
(fig. 10), which spring irregularly from the margins of the segments and are of various 
lengths, a single segment frequently having on each side three set of different sizes, 
others only one or two. 

Each caudal lamina bears eight slender curved ungues, which increase progressively 
in length from the first to the last, the first four, however, being distinctly smaller and 
less robust than the rest (fig. 11); the concave margins of the ungues finely pectinated 
with lancet-shaped spinules (fig. 12). In the intervals between the third, fourth, fifth, 
and sixth ungues are attached small marginal sete about one-third as long as the 
ungues themselves: the abdominal margin in front of the ungues is densely hispid. 

The eyes consist of subreniform black pigment-masses about which numerous 
simple convex lenses are clustered in regular rows; a delicate capsule invests the 
whole structure, and is continued over a peduncle in which are contained muscles for 
the movement of the eye as well as numerous nucleated (nerve ?) cells. The whole 
organ lies loosely within the shell and has no attachment to the body of the animal 


DURING THE ‘ CHALLENGER’ EXPEDITION. 87 


except by its peduncle. ‘There are seven pairs of laminar branchiz, and in all essential 
particulars the anatomy is that of Asterope. 

I have pleasure in naming this fine species after Mr. Henderson of the Christian 
College, Madras, to whom I am indebted for numerous specimens. 


The species was dredged by Mr. Henderson in Madras Harbour in a depth of 4-5 
fathoms. 


CYCLASTEROPE ORBICULARIS, n. sp. (Plate XV. figs. 13-19.) 


Female.—Shell rather thin, subspherical (figs. 18, 14): seen from the side almost 
circular, height equal to five-sixths of the length; beak short and sharp; rostral notch 
narrow and moderately deep, situated quite in the middle of the anterior margin; seen 
from above the outline is regularly ovate, tapering a little towards the posterior 
extremity, which is evenly rounded off, and more considerably to the front, which is 
somewhat obtuse. Anterior margins of the shell fringed with short fine hairs ; surface 
smooth; colour light brown. Frontal tentacle (fig. 15) three-jointed; the second joint 
short, last joint elongated, conical. Secondary branch of the antenna three-jointed 
(fig. 16); first joint broad and fringed below with numerous fine hairs, second much 
more slender and bearing a single short marginal seta, third very small and ending in 
a single long ringed seta. The mandibular foot (fig. 17) is very similar to that of 
C. hendersoni, except that the falciform process (which having been broken was only 
partially seen) seems to be much smaller and less elaborately armed. Caudal lamin 
(fig. 19) broad and short, bearing eleven marginal ungues, the first eight of which are 
merely slender closely-set curved sete, nearly equal in length and slightly spinulose ; 
the remaining three are very stout, strongly curved, and bear numerous marginal teeth, 
the first of them about as long as the preceding sete, the second twice, and the third 
fully three times as long; abdominal margin in front of the lamina densely hairy. 
Length of the shell 2°7 millim. 

I have seen only one specimen of this species. It bears the inscription “ Valparaiso,” 
without any further particulars; and I am not sure whether I had it from the 

Challenger’ or from some other source. 


Genus CypripinA, Milne-Edwards. 


CYPRIDINA ALBOMACULATA, Baird. 
1880. Cypridina albomaculata, Baird, “ Note upon the Genus Cypridina, Milne-Edwards,” Proc. 
Zool. Soc. Lond., Annulosa, p. 201, pl. Ixxi. figs. 1, la, 10, le, 1d. 
The type of this species is in the British Museum, and is stated in Dr. Baird’s 
description to be from “Swan River,—from the collection of Mr. Cuming.” The 
Nz 


88 DR. G. S. BRADY ON MYODOCOPA OBTAINED 


single specimen which I possess was taken many years ago by Dr. Coppinger, of 
H.M.S. ‘ Alert,’ in Dundas Strait, N.W. Australia. 


CYPRIDINA CASTANEA, n. sp. (Plate XVI. figs. 1-4.) 


Male.—Shell corneous, flexible, dark brown, as are also the limbs of the animal. 
The rostrum is bordered by a thin scale-like flange, which is strengthened below by 
several stout spines; the convexity of the sub-rostral cleft has a fringe of numerous 
sleuder spines, which are long in the middle of the series and gradually smaller 
towards the ends; the shell-surface near the rostral angle has also an elevated hump to 
which are attached a number (about fifteen) of long, rigid sete (fg. 1). Masticatory 
process of the mandibular foot bifid and bearing a long lateral seta (fig. 2). The 
antennules have two long apical sete and several small sensory ones, and to two of the 
smaller of these are attached circular, saucer-shaped disks. The distal extremity of the 
first maxilla (fig. 3) bears numerous ciliated sete, and four very stout spine-like teeth 
which have spinulose margins. Caudal lamine (fig. 4) provided with nine equidistant, 
slender, curved, and denticulated ungues, the last two of which are very long, the rest 
gradually decreasing in length to the first. Length of the shell about 7 millim. 

One example only of this species was taken by the ‘ Challenger’ Expedition in 
a “deep haul” from the Atlantic, lat. 52° 41’ N., long. 36° 6’ W., depth 1675 
fathoms, 6th May, 1876. The specimen was much damaged, and neither the shell nor 
the soft parts were in a condition to be accurately examined. Provisionally, however, 
it may be referred to the genus Cypridina, though some of the characters show an 
approach to Philomedes. 


CyPRIDINA BAIRDI, G. S. Brady. (Plate XVI. figs. 22, 23.) 


1865, Cypridina bairdii, G. 8S. Brady, “ On new or imperfectly known species of Marine Ostracoda,” 
Trans. Zool. Soc. vol. v. p. 387, pl. Ixii. figs. 7 a-m. 

Of this species I have seen no specimens other than those on which the original 
description was founded. The squamous outgrowths of the shell in the neighbourhood 
of the beak (figs. 22, 23) are, however, somewhat similar to those of C. castanea, with 
which the soft parts also agree so far as they have been examined. It is probable 
that a fuller knowledge of the two forms may show other points of resemblance. 

C. bairdii was taken in the tow-net, off Pescadores, China, by the late Mr. A, Adams. 


Cypripina Japonica, G, S. Brady. 
1865. Cypridina japonica, G. 8. Brady, loc. cit. p. 386, pl. Ixii. figs. 8 a—d. 


This would appear, from the imperfectly preserved specimen in my collection, to be 
rightly referred to the genus Cypridina. 
Collected in Japan by Mr. A, Adams. 


DURING THE *‘ CHALLENGER’ EXPEDITION. 89 


Cypripina (?) sLoneata, G. S. Brady. 
1865. Cypridina elongata, G. S. Brady, loc. cit. p. 386, pl. Ixii. figs. 9 a-d. 


Respecting this species I am able to add nothing to the short description already 
published. ; 


(?) Cypripina PunctatTa, Dana. (Plate XVI. figs. 5-9.) 
1852. Cypridina punctata, Dana, Crustacea of U.S. Exploring Exped. p. 1293, pl. ix. figs. 2 a, b. 


Three specimens only of a species closely similar to, if not identical with, Dana’s 
C. punctata were taken by the ‘Challenger’ Expedition at some point, not accurately 
noted, in the Eastern Archipelago. All the three were in very bad condition—many 
of the limbs broken or absent. In no case have I been able to find the slightest trace 
of the vermiform limb. 

Shell extremely thin and membranaceous: seen from the side (fig. 5) subovate, 
greatest height in the middle and equal to somewhat more than half the length, dorsal 
and ventral margins evenly arched, the dorsal having the larger curve; anterior 
extremity obliquely subtruncate, with a short wide beak, above which is a shallow 
sinuation, subrostral sinus narrow; posterior extremity produced below the middle 
into a rather wide, obtusely angular process, which is continuous with the ventral 
margin below, but is separated from the dorsal margin by an abrupt sinuation ; seen 
from above the outline is ovate, with produced subacute extremities. The mandibular 
foot terminates in three short, equal ungues (fig. 7), which are slightly swollen at their 
bases; there is also a single slender curved seta, considerably shorter than the ungues; 
the masticatory process (fig. 8) is slightly bifid at the apex. Caudal lamine (fig. 9) 
having eight marginal ungues, the first three extremely small and weak, the rest 
successively increasing in length to the last; the fifth, sixth, and seventh ungues are 
not articulated to the limb but are simply processes of it. Length of the shell 
1°5 millim. 


CypRIDINA DAN&, G. S. Brady. (Plate XVI. fig. 24.) 

1880. Cypridina dane, Brady, Report on the Ostracoda of the ‘ Challenger’ Expedition, p. 156, 
pl. xxxvi. figs. 2 a-d. 

The head of the vermiform appendage, here figured, is peculiar, the upper portion 
being massive, strongly curved, and armed with a series of marginal teeth; the lower 
portion much smaller, and consisting of a simple, almost rectangularly curved process, 
wide at the base but slender beyond the middle. 


CyPRIDINA (2) ARMATA, n. sp. (Plate XV. figs. 20, 21.) 


Shell thin and membranaceous: seen from the side oblong, subovate, greatest height 
situated in the middle and equal to two-thirds of the length; dorsal and ventral 


90 DR. G. S. BRADY ON MYODOCOPA OBTAINED 


margins evenly arched; beak wide, obtusely pointed, notch deeply excavated ; posterior 
extremity wide, produced in the middle into a broad, bifid prominence. Seen from 
above the outline is subpentagonal, with parallel and almost straight lateral margins, 
which in front of the middle converge sharply towards the subacute anterior extremity; 
posterior extremity very wide, scarcely narrower than the widest part of the shell, 
truncated, the outer angles forming two backward-pointing spines, obtusely mucronate 
in the middle. Surface of the shell smooth ; behind the middle, on the lateral aspect 
of each valve, there is a very large, acutely lancet-shaped spine which points directly 
backwards, its point reaching almost as far as the hinder border of the shell; just 
within the dorsal margin, above the base of the larger spine, is a much smaller one of 
similar form, but pointing upwards. Colour brown, the spines almost black at their 
apices. Length 1:4 millim. 

Animal unknown. 

One specimen from a depth of 7 fathoms, Flinders Passage, Sept. 8th, 1874— 
‘ Challenger’ Expedition. 


CypripINA (?) squamosa, n. sp. (Plate XVI. figs. 10-12.) 


Shell very thin and membranaceous: seen from the side almost circular (fig. 10), 
the height equal to about three-fourths of the length; anterior extremity broadly 
rounded, the beak wide and almost rectangular, notch deep and narrow; posterior 
extremity rounded above the middle, below the middle forming a wide truncate 
prominence ; dorsal and ventral margins boldly convex. Shell marked throughout 
with a reticulated pattern as of imbricated scales (fig. 11). Caudal lamine (fig. 12) 
bearing only three ungues, which are strongly curved, the first being very small, the second 
about thrice as long, and the third twice as long as the second. Length 0:9 millim. 

One specimen only of this species has been noticed; it occurred in a gathering from 
the Eastern Archipelago the precise locality of which was not stated. Only fragments 
of the soft parts were recognizable, and its position as a Cypridina must be looked upon 
as provisional only. 


Genus Puitomepes, Lilljeborg. 


PHILOMEDES AGILIS, Thomson. (Plate XVI. figs. 13-16.) 

1878. Philomedes agilis, Thomson, “On the New Zealand Entomostraca,” Trans. New Zealand 
Institute, vol. xi. p. 257, pl. xi. fig. C. 8a-e, D. 1 a-g. 

Male.—Shell, seen from the side (fig. 13), oblong, ovate, greatest height in the 
middle and equal to about two-thirds of the length, beak obtuse and only slightly 
produced, notch very shallow, anterior extremity narrow and somewhat angular, 
posterior broad and well rounded; dorsal and ventral margins equally arched; the 
dorsal margin, especially in its posterior half, irregularly emarginate and presenting an 


DURING THE ‘ CHALLENGER’ EXPEDITION. 91 


abruptly undulated outline. Seen from above, the outline is compressed, subclavate, 
sides nearly parallel but slightly converging towards the obtusely-pointed front ; 
posterior extremity wide and subtruncate, greatest width considerably less than half 
the length. Surface of the shell covered with numerous subrotund pits and sparingly 
setose. Colour light brown. Eyes large and black. Length 1-9 millim. 

Secondary branch of the antenna (fig. 14) three-jointed and strongly prehensile ; 
basal joint short and bearing several sete, one of which is long and plumose; second 
joint much elongated, and having two long sete in the middle of the external margin; 
third joint also long, curved, with a denticulated external margin, a single long seta at 
the base, its apex rounded, crenulated, and slightly split up in a multifid fashion. 

Apex of the vermiform appendage (fig. 15) having its upper process armed with five 
slender curved spines; lower portion rounded, obtuse, and toothless. Caudal lamine 
(fig. 16) bearing three principal ungues and six very small spine-like sete: the first 
five setee are quite minute, then follows a strong curved unguis of moderate size; a 
sixth small seta intervening between it and the following unguis, which is twice as long 
as the preceding; the last unguis is still longer and, like its neighbour, has a spinulose 
margin. 

Female.—* Valves somewhat larger and much more circular in outline, with the beak 
small and very slightly produced; oral notch nearly rectangular; height about three- 
fourths of the length ; easily distinguished externally from the male by the small size 
of the eye-spot. Eyes reddish.” 

I am indebted to Professor G. M. Thomson, of Otago, for specimens of this 
interesting species, which was taken by him “swimming actively in the rock-pools 
on the Taieri Beach.” ‘The specimens sent to me were all males. I have therefore 
quoted Prof. Thomson’s description of the female shell. 


PHILOMEDES CORRUGATA, n. sp. (Plate XVI. figs. 17-21.) 


Shell very tumid: seen from the side (fig. 17) subrhomboidal, highest about the 
middle, height equal to two-thirds of the length; the dorsal and ventral margins 
markedly convex ; anterior extremity narrowed and rather oblique, beak blunt and not 
very prominent, notch very shallow; posterior extremity evenly rounded. Seen from 
above (fig. 18), the outline is subpentagonal, greatest width situated behind the middle 
and equal to four-fifths of the length; the anterior margin is rounded off but extremely 
broad and almost subtruncate, the posterior margin slopes very abruptly and with an 
irregularly sinuous curve from the widest point, and terminates in a very broad obtusely 
rounded median process; the lateral margins converge very slightly and in sinuous 
fashion towards the wide anterior extremity. The surface of the shell is much 
wrinkled and pitted, and slightly hispid with short distant hairs. Colour reddish 
brown. Length 1:4 millim. 

The sculpturing and general outline of the shell as shown in the lateral view (fig. 17) 


92 DR. G. S. BRADY ON MYODOCOPA OBTAINED 


are possibly to some extent incorrect owing to shrivelling of the specimen when taken 
out of the spirit. The soft parts of the animal, so far as they can be made out, do not 
differ materially from those generally belonging to the genus. 

The first three natatory sete of the female antenna are marginally spinous as in 
Pleoschisma (see figs. 3, 4, Pl. XVII.); so also is the first seta of the male antenna. The 
prehensile branch of the male antenna (fig. 19) is irregularly dentated on both margins. 
Caudal lamine (fig. 20) armed with seven marginal sete, four of which are unguiform 
and much stouter than the others; of these ungues the first is very short, the second 
about twice as long, the third twice as long as the second, and the fourth slightly 
longer than the third; three very short, slender, and nearly equal sete are placed in 
front of the first unguis and in the spaces behind the first and second and second and 
third ungues. 

Hab, One specimen (a male) of this species was found in a tow-net gathering from a 
depth of 2-10 fathoms, off Port Jackson, Australia; another (female) from a depth of 
7 fathoms in Flinders Passage, Australia. 


Genus Pieoscuisma, G. S. Brady. 
PLEOSCHISMA MOROIDES, G. 8. Brady. (Plate XVII, figs. 1-11.) 


1888. Pleoschisma moroides, G. 8. Brady, ‘On Ostracoda collected by H. B. Brady, Esq., LL.D., 
F.R.S., in the South Sea Islands,” Trans. Roy. Soc. Edinb. vol. xxxy, p. 514, pl. i. figs. 23, 24, 

* Shell, seen from the side, subcircular, height equal to three-fourths of the length. 
Anterior extremity wide, feebly rounded, almost flat, notch obsolete; posterior narrower, 
rounded, slightly sinuated above and below; dorsal and ventral margins moderately 
convex. Seen from above, broadly ovate, nearly equal in width throughout; extre- 
mities broad and rounded, the anterior rather the narrower of the two; lateral margins 
moderately arcuate; width equal to four-sevenths of the length. Surface of the shell 
minutely punctated, and in old specimens raised into circular bosses; colour dark 
brown. Length 1:2 mm.” 

Antennules, in both sexes, very similar to those of Philomedes: the sensory setiferous 
seta attached to the antepenultimate joint in the male is dilated at the base, and is 
setiferous on the outer side only (fig. 1). The antenna of the male is larger than that 
of the female, and the dilated basal joint is closely and delicately striated (fig. 2); the 
first and third joints of the natatory branch are very long, the relative lengths of the 


é 56 3 1.2.3.4.5.6.7.8.9 
nine jomts being approximately as follows: ;3~{yy-3-9 9» 97; the secondary 


a 


branch is strongly prehensile, its terminal joint curved, its concave margin deeply and 
irregularly indented, and its apex minutely crenulated (fig. 5); each of these two joints 
bears a single seta; the much smaller basal joint has four or five small cilia. In the 
female (fig. 3) the basal joint is not at all striated, and the relative lengths of joints of 


DURING THE ‘ CHALLENGER’ EXPEDITION. 95 


2 5.6 : 
the natatory branch (fig. 3) are as follows: ese? «Seif St Oo peices ateehed 


to the second, third, fourth, and fifth joints are extremely short, equal in length, and 
are armed with short stout spines along their outer margins (figs. 3, 4); the sete of 
the last four joints are from three to four times as long as the preceding and are not 
spinous ; secondary branch simple, curved, one-jointed. The mandibular foot is four- 
jointed, and is in the male (fig. 7) devoid of any masticatory process, but bears 
numerous simple sete; in the female the first joint bears a bifid process, and the 
second has, in addition to several simple sete, a small bisetose finger-like process at its 
distal end (fig. 8). The first maxilla (fig. 6) consists of one stout principal lobe with 
four smaller lobes, all of which are setiferous, most of the stouter sete having one of 
the margins strongly denticulated: the principal lobe is two-jointed, the last joint 
small and giving attachment to about six apical setee, which are graduated in size from 
the first to the last and bear strong marginal teeth. The vermiform limb has a single 
claw-like process at its apex. Caudal lamine (fig. 10) short; the first three marginal 
appendages consisting of very short slender sete, the fourth an unguiform spine about 
twice as long as the preceding setze and very much stouter; then follows another small 
seta, then a second stout unguis about twice as long as the first, and lastly a third 
unguis which is more than twice as long as the second. Eyes small, deep red, the 
lenses irregularly placed and of very unequal size; in the male subreniform (fig. 11), in 
the female smaller and globose. 

My first description of this genus was deficient in any account of the soft parts of 
the animal, but I have recently by careful maceration of some of the dried shells been 
able to obtain preparations which show, though imperfectly, some of the more 
important structures.’ These are certainly sufficiently characteristic, and even apart 
from the shell would constitute a valid generic distinction. 

The specimens were obtained from various localities among the South Sea Islands. 


Genus SARSIELLA, Norman. 


SARSIELLA scULPTA, G. S. Brady. (Plate XVII. figs. 12, 13.) 

1888. Sarsiella sculpta, G. S. Brady, “ Ostracoda collected in the South Sea Islands,” Trans. Roy. 
Soe. Edinb. vol. xxxv. p. 517, pl. 1. figs. 17-20. 

The original description of this species was taken from the shell only, but I have 
recently succeeded in finding remains of the soft parts in a specimen taken by the 
: Challenger, and from it I have here figured the end of the vermiform foot and one 
of the caudal lamine. These, it will be seen, exactly agree with Sarsied/a. 

The ‘Challenger’ specimen was taken at a depth of 7 fathoms in Flinders 


Passage. 
vou. xiv.—Ppart 11. No. 8.—Apri/, 1897. 


94 DR. G. 8S. BRADY ON MYODOCOPA OBTAINED 


SARSIELLA SIMPLEX, G. S. Brady. (Plate XVII. figs. 16-19 
1888. Sarsiella simplex, G. S. Brady, “ Ostracoda collected in the South Sea Islands,” Trans. 
Roy. Soc. Edinb. vol. xxxv. p. 516, pl. iv. figs. 15, 16. 
One specimen was taken by the ‘Challenger’ Expedition in the tow-net off Cape 
Howe, Australia, at night. 


SaRSIELLA RoBusTA (G. S. Brady). (Plate XVII. figs. 14, 15.) 


1888. Pleoschisma robusta , G. 8S. Brady, “Ostracoda collected in the South Sea Islands,” 
Trans. Roy. Soc. Edinb. vol. xxxv. p. 518, pl. iv. figs. 13, 14. 

Further examination of the dried shells of this species has revealed portions of the 
limbs, which show that it must be referred to the genus Sarsiella. The antennules 
(fig. 14) are provided with the dense fascicle of hairs which has been described by 
Dr. G. W. Miiller as characteristic of the male Sarsiella, and also by Dr. Norman and 
myself under the genus Vematohamma. 


Genus Eurypy vs, G. 8. Brady. 


EuRYPYLUS PEeTRosuUS, G. S. Brady. (Plate XVI. figs. 25, 26.) 
1870. Eurypylus petrosus, G. S. Brady, Les Fonds de la Mer, tome i. p. 141, pl. xviii. figs. 1, 2. 


The type specimen of this species remains as yet the only one which has been seen. 
It was described from the shell only, and inasmuch as other very nearly allied forms 
belonging to the genera Sarsiella and Pleoschisma have since been published, I 
thought it well to sacrifice the shell for the sake of examining the contained animal, of 
which, unfortunately, I succeeded in obtaining fragments only. ‘The mandibular foot 
(fig. 25) and caudal lamina (fig. 26) are here figured. The former, it will be seen, is 
very similar to that of Sarsiel/a; the latter to that of Pleoschisma. The differences are 
in fact quite sufficient to indicate three distinct genera. 

The type of Ewrypylus was taken off St. Vincent (Cape Verd). 


Family HALOCYPRID. 


Of many of the species belonging to this group I have seen only imperfect examples, 
and cannot, therefore, figure or describe them satisfactorily. They have, however, been 
fully illustrated by Dr. Claus in his works on the Atlantic and Mediterranean Halo- 
cypride. Some of the new genera proposed by Dr. Claus seem to me to be based on 
trivial and insufficient characters, and are adopted here only as a temporary expedient. 

The most interesting point in connexion with the specimens here noted is the con- 
firmation which they afford to the generally accepted idea of the almost cosmopolitan 
distribution of many pelagic animals. With one exception they are referable to species 
already described by Dr. Claus from the North Atlantic and Mediterranean, whereas 
the ‘ Challenger’ specimens are mostly from the Pacific and Australasian areas. 


DURING THE ‘ CHALLENGER’ EXPEDITION. 95 


Genus Concnacra, Dana (in part). 


CoNCH@CIA BISPINOSA, Claus. 


1890. Conchecia bispinosa, Claus, Die Gattungen und Arten der mediterranen und atlantischen 
Halocypriden, p. 10. 
1891. 5 ss Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 
p. 59, pl. v., pl. vi. fig. 1, pl. viii. figs. 7, 8. 
Taken in the tow-net during the ‘Challenger’ Expedition: lat. 32° 41’ N., long. 
36° 6’ W.; and in lat. 36° 32’ S., long. 132° 52’ W. 


Genus ParaconcuaciA, Claus. 


PARACONCHG@CIA OBLONGA, Claus. (Plate XVII. figs. 20, 21.) 


1890. Paraconchecia oblonga, Claus, Die Gattung. und Arten der medit. und atlant. Halocypr. p. 13. 
1891. 55 = Claus, Die Halocypr. des atlant. Oceans und Mittelmeeres, p. 63, 
pl. viii. figs. 10, 11, pl. ix. 

Hab. Off Kandabu, Fiji; and in lat. 46° 46’ S., long. 45° 31’ E. (St. 146, ‘ Chal- 
lenger’). The specimens described by Dr. Claus were from lat. 37° 45! N., long. 13° 
38’ W., depth 1500 metres; and from lat. 34° 18’ N., long. 15° 34’ W., depth 1000 
metres. Ihave no record of the depth at which the ‘Challenger’ specimens were 
taken. 


Genus Concn@certa, Claus. 


CoNcH@cETTA AcUMINATA (?), Claus. 
1890. Conchecetta acuminata, Claus, Die Gattung. und Arten der medit. und atlant. Halocypr. p. 16. 
1891. 53 a Claus, Die Halocypr. des atlant. Oceans und Mittelmeeres, p. 67 
pl. xill., xiv. 
A single specimen, apparently belonging to the young of this species, was found in a 
‘Challenger’ gathering from the neighbourhood of the Philippine Islands. 


Genus CoNcH@cILLA, Claus. 


CoNCH@CILLA DAPHNOIDES, Claus. 


1890. Conchecilla daphnoides, Claus, Die Gattungen und Arten der mediterranen und atlantischen 
Halocypriden, p. 18. 
1891. a ee Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 


p. 68, pl. xv. 
One specimen from a gathering made by the ‘ Challenger’ Expedition off Kandabu, 
Fiji. 


02 


96 DR. G. 8. BRADY ON MYODOCOPA OBTAINED 


Genus Concuacissa, Claus. 
ConcHacissa ivpricata, G. S. Brady. 
1880. Halocypris imbricata, G. S. Brady, Report on the Ostracoda of the ‘ Challenger ? Expedition, 
p- 167. 
1890. Conchecissa armata, Claus, Die Gattungen und Arten der mediterranen und atlantischen 
Halocypriden, pp. 19, 20. 
1891. Ba » Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 
p- 70, pl. xvi., xvii., xvill. 
This is referred doubtfully by Dr. Claus to his more recently described species, 
Conchexcissa armata, and, notwithstanding the slight discrepancies referred to by that 
author, I can scarcely doubt that the two are identical. 


Genus PsrupoconcHaecla, Claus. 
PSEUDOCONCHGCIA SERRULATA, Claus. (Plate XVII. figs. 22-24.) 
1874. Conchecia serrulata, Claus, Die Familie der Halocypriden, p. 61, pl. 1. figs. 2-7, 9-11, pl. 11. 
figs. 12, 13, 17, 19. 
1880. Halocypris atlantica, Brady, Report on the Ostracoda of the ‘ Challenger ’ Expedition, p. 164, 
pl. x1. figs. 1-15, pl. xli. figs. 11, 12. 
1890. Pseudoconchecia serrulata, Claus, Die Gattungen und Arten der mediterranen und atlant- 
ischen Halocypriden, p. 20. 
1891. 5 3 Claus, Die Halocypriden des atlantischen Oceans und Mittel- 
meeres, p. 72, pl. xix. figs. 1-14, pl. xxii. figs. 1-13. 

The reference of this species in the Report of the ‘Challenger Expedition’ to 
Halocypris atlantica, Lubbock, can scarcely be maintained, though the figures and 
description there given are, I think, correct so far as they go. ‘The descriptions, both 
of Dana and Sir John Lubbock, are so concise and vague that it is impossible to say 
certainly to what they refer. There can, however, be no doubt that Pseudoconchecia 
serrulata, Claus, is identical with Halocypris atlantica of the ‘Challenger’ Report. 
Some characters which were passed without notice in that Report I figure here—the 
hooked appendage of the right and left male antenna (figs. 25, 24) and the swimming- 
setee of the same limb (fig. 22). These sete are figured in order to show their narrowly 
lanceolated or spathulate extremities. All the setee seem to end in this way, but the 
two springing from the apical joint much more distinctly so than the rest : in fact, it is 
not easy to see the slightly dilated apices of the lateral sete except by rotating them 
so as to get a front view. But although it was in P. serru/ata that I first noticed the 
dilated apices, I now find on closer investigation that the antennal sete of other 
species (perhaps, indeed, of all the Halocypride) have a similar structure. This point 
is noted (though not figured) by Prof. Claus in his definition of the subfamily 
Concheecine. There seems to be no such structure in the sete of the Cypridinide. 

P. serrulata occurred in many of the * Challenger’ gatherings. ‘Those which I have 


DURING THE ‘ CHALLENGER’ EXPEDITION. 97 


been able to verify since recognizing the identity of the specimens are as follows :— 
Surface-net between Api and Cape York and off Port Jackson; in tow-net at 30 fathoms, 
South Atlantic, lat 42° 32’ S., long. 56° 27’ W.; and in lat. 35° 41’ N., long. 159° 
AV’ E. 
Genus MicroconcHactr, Claus. 
Microconcu@cta claus (G. O. Sars). 
1887. Halocypris clausii, G. O. Sars, Nye Bidrag til Kundskaben om Middelhavets Invertebrat- 
fauna, iv. Ostracoda Mediterranea, p. 87, pl. xi. figs. 7-10, pl. xiv. 
figs. 6-18. 
1890. rp 5 Claus, Die Gattungen und Arten der mediterranen und atlantischen 
Halocypriden, p. 22. 
1891. Mikroconchecia clausii, Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 
p: 73, pl. xx. 
The only ‘ Challenger’ gathering in which this species had been detected was from 
the surface-net off Kandabu, Fiji, where it occurred in considerable abundance. 


Genus Hatocyrris, Dana. 
HALocyrris concHa, Claus. 
1874. Halocypris concha, Claus, Die Familie der Halocypriden, pl. ii. figs. 20-25, pl. iii. figs. 26- 


35. 

1891. E » Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 
p- 77, pl. viii. fig. 12, pl. xi. figs. 6, 7, pl. xxii. figs. 1-12, pl. xxvi. 
fig. 11. 


In a ‘Challenger’ surface-net gathering from the North Atlantic: lat. 26° ZN. 


99° OF 


long. 33° 37’ W. 


Hatocypris PELAGICA, Claus. 


1890. Halocypris pelagica, Claus, Die Gattungen und Arten der Halocypriden, p. 25. 
1891. a ms Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 


p- 78, pl. xxi. figs. 1-11. 
In two surface-net gatherings from the ‘Challenger’: lat. 35° 18’ N., long. 
144° 8’ FE. ; and lat. 35° 41 N., long. 157° 41’ E. 


Genus Hanocypria, Claus. 


HAtLocypera GLoBosa, Claus. 

1890. Halocypria globosa, Claus, Die Gattungen und Arten der mediterranen und atlantischen 
Halocypriden, p. 25. 

Claus, Die Halocypriden des atlantischen Oceans und Mittelmeeres, 
p- 79, pl. xxii. figs. 13-18. 


1891. =o 5 


In the surface-net between Api and Cape York, and in lat. 36° 32’ S., long. 
132° 52’ W. (‘ Challenger’). 


98 


o 
5: 


Fig. 


Fig 


Fig. 
Fig. 


Me 
2. 
3. 
4, 
5. 
6. 


DR. G. 8S. BRADY ON MYODOCOPA OBTAINED 


EXPLANATION OF THE PLATES. 


PLATE XV. 


Cyclasterope hendersoni, 2 (p. 86). 


Shell seen from right spay 

x8. 
55 above, 

Frontal tentacle. 

Secondary branch of antenna. 

Mandibular foot. 

Falciform process of mandible. 


Figs. 7, 8. End and side of falciform process, more highly magnified. 
Fig. 9. End of vermiform limb. 


Fig. 
Fig. 
Fig. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 


Fig. 
Fig. 


pay 


H oO Lo 


10 
11 


12. 


13. 
14. 
16. 
16. 
alii 
18. 
19, 


20. 
21. 


. Segments of the same with sete. 
. Caudal lamina. 
% spines of unguis. 


Cyclasterope orbicularis, 2 (p. 87). 

Outline of shell seen from left side, 
5 above, } 20 

Frontal tentacle. 
Secondary branch of antenna. 
Mandibular foot. 
Maxilla of first pair: (@) terminal process of same. 
Caudal lamina. 


Cypridina (?) armata (p. 89). 
Shell seen from left side, 
} x40, 


35 above, 


PLATE XVI. 


Cypridina castanea, ? (p. 88). 


Supero-anteal margin of left valve. 
Mandibular foot. 

Masticatory lobe of first maxilla. 
Caudal lamina. 


Fig. 


DURING THE ‘ CHALLENGER’ EXPEDITION. 


Cypridina punctata (p. 89). 


. Outline of shell, x 40. 


af rostrum of right valve. 


. Masticatory process of the same. 
. Caudal lamina. 


. 24, 


5, 
g. 26. 


3) 
6. 
ig. 7. End of mandibular foot. 
8 
9 


Cypridina (%) squamosa (p. 90). 


. Shell seen from right side, x 50. 
. Portion of shell with sculpture. 
. Caudal lamina. 


Philomedes agilis, 2 (p. 90). 


. Shell seen from left side, x 40. 
. Secondary branch of antenna. 

. Apex of vermiform foot. 

. Caudal lamina. 


Philomedes corrugata (p. 91). 


. Shell of female seen from right side, } 40 


above (Flinders Passage), 


99 


. Secondary branch of antenna, ¢ . 
. Caudal lamina. 
. Outline of shell (Port Jackson). 


Cypridina bairdi (p. 88). 


. Outline of right valve in region of notch. 


23. 


Outline of left valve. 


Cypridina dane (p. 89). 


End of vermiform foot. 


Eurypylus petrosus (p. 94). 


Mandibular foot. 
Caudal lamina. 


99 


100 


a 


POD MON anFf oe Dd 


DR. G. 8. BRADY ON ‘CHALLENGER’ MYODOCOPA. 


PLATE XVII. 


Pleoschisma moroides (p. 92). 


. Apical joints of antennule, ¢. 
. Antenna, ¢ (imperfect). 


Phe 

92 

», one of the short sete more highly magnified. 
- secondary branch, 3. 


. First maxilla. 

. Mandible, ¢. 

. Mandible, ¢. 

. End of vermiform foot. 
. Caudal lamina. 

- Hye, . 


Sarsiella sculpta (p. 93). 


2. End of vermiform foot. 
. Caudal lamina. 


Sarsiella robusta, & (p. 94). 


. Antennule. 
. Secondary branch of antenna. 


Sarsiella simplex, 2 (p. 94). 


. Shell seen from right side, 

" 3 } x 80. 
se 

. First maxilla. 

. Caudal lamina. 


es above, 


Paraconchecia oblonga, 3 (p. 95). 


. Shell seen from left side, x 40. 
. Anterior margin of shell. 


Pseudoconchecia serrulata, 3 (p. 96). 


. Natatory branch of antenna. 
. Secondary branch of right antenna. 


3 left A 


\ 


. a) 
~ y 


EP ST 
SE T a 


Ga Brady del 
MP Parker hth 


CRUSTACEANS OF THE GROUP MYODOCOPA, 


Trans. Zoot. Foc. Vol XW FE XVI. 


—_ 


G.8.Brady del Geo West & Sons imp. 


CRUSTACEANS OF THE GROUP MYODOCOPA. 


MP Parker lith. 


rans. Loot Soc Wot AN G4 XVIL 


Geo West & Sons unp. 


G.3.Brady del. 
M.P. Parker lith. 


CRUSTACEANS OF THE GROUP MYODOCOPA. 


BO APR 1897 


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CONTENTS. 


1V. Contributions to the Comparative Anatomy and Histology of the Suprarenal 
Capsules.—The Suprarenal Bodies in Fishes, and their Relation to the so-called 
Head-Kidney. By Swate Vincent, M.B.Lond., Demonstrator of Physiology 
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ENG Xe Vs) steee ecee . . page 41 
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VI. A Revision of the Oriental Hesperiide. By H. J. Ewes, F.RS., F.L.S., F.Z.S8., 
and James Epwarps, /.E.S. 


Received April 14, 1896, read June 2, 1896. 
[Puates XVIII.-XXVII.j 


ALTHOUGH Mr. F. Moore had previously described many genera and species of 
Hesperiide, the majority of which are sound and well characterized, yet this large and 
difficult group of Butterflies was, perhaps, less known than any other in the Old 
World before Mr. de Nicéville and Captain Watson began to pay attention to it; 
and until Captain Watson’s paper on the classification of the family in the Proceedings 
of this Society for 1893 laid down a fair basis for its arrangement, Elwes had not 
attempted to arrange his own collection, which contains a far greater number of 
species and specimens from the Oriental region than that of the British Museum, 
on which Watson based his work. 

On proceeding to do so, Elwes found numerous species which Watson had not seen, 
and which required a detailed examination in order to decide their specific and generic 
position. This he would not have been able to undertake without the able assistance of 
Mr. Edwards, who has given all his spare time for nearly three years to the task of working 
out the large mass of material which has passed through their hands during that period, 
including much necessary dissection of the genitalia and the comparison of their 
generic characters with those given by Watson. How far the characters drawn from 
a careful examination of the male genitalia can be properly used in deciding questions 
of specific identity or distinction is a point which by some entomologists has been 
questioned; and a very considerable practice in making this examination, and great 
experience in estimating the value of the characters observed, are necessary in order 
to form an opinion on the subject. But those who have most experience and who 
have devoted most time to this study are, we think, practically agreed that these 
characters are of great importance and assistance, and the question has been so well 
stated by Messrs. Godman and Salvin in the ‘Biologia Centrali-Americana,’ 
Rhopalocera, vol. ii. p. 245, that we cannot do better than quote their remarks, in 
which we fully concur. 

They say :—“ As in the foregoing families, we have examined the male secondary 
sexual organs and find an almost endless variety of structure. We have not ventured 


1 The Editor of the Society’s Publications wishes it to be understood that he is not responsible for the 
nomenclature used in the present memoir, which deviates in several points from that habitually employed by 
the Society.—P. L. 8. 

VOL. xIv.—Part Iv. No. 1.—October, 1897. iz 


an MUS 
a he 


102 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


to use these characters for building up any system of classification, as we consider 
the time is hardly come for their adoption for any such purpose. But we have found 
them invaluable in deciding points of specific distinction where the external characters 
are not clearly defined. In some cases a slight external feature distinguishes wo 
forms, which is sometimes greatly strengthened by a marked divergence in the male 
genitalia, and sometimes no difference can be traced in these organs. In the former 
case a specific difference between two similar forms can safely be conceded, while in 
the latter the opposite course should be followed.” 

As regards the arrangement of genera, we have adopted that proposed by Watson in 
his last paper, “A Key to the Asiatic Genera of the Hesperiide,” published in the 
Journal of the Bombay Natural History Society for 1895, which in some respects 
is an improvement on his earlier paper before referred to. We have been compelled 
to propose a few new genera and some seventy species; and though probably there 
are still many additions and corrections to be made to the list, especially in the Malay 
islands and Indo-Chinese countries, yet we have spared no efforts to procure for 
examination all the material which Elwes’s collection did not previously contain. In 
doing so we have to acknowledge the very great assistance which has been afforded 
us by Mr. L. de Nicéville and Captain Watson in India, who have sent us advance 
copies of all their papers and descriptions. In Europe Dr. Georg Semper, of Altona, 
has been good enough to lend us authentic specimens of all the species included 
in his great work ‘ Die Schmetterlinge der Philippinischen Inseln.’ Dr. O. Staudinger 
has sent us a large number of specimens from Borneo and other Malayan islands, 
the novelties amongst which he has kindly permitted us to describe. Herr P. C. 
T. Snellen, of Rotterdam, has been equally liberal with specimens from Java and 
Sumatra. M. Charles Oberthiir has enabled us to examine some of the new and 
rare forms in his magnificent collection from China, and Mr. Leech has also been 
most obliging in lending types described in his ‘ Butterflies of China &c.’ and other 
specimens for examination. The Hon. Walter Rothschild kindly allowed us to select 
from and describe several species of interest in his collection; and we are indebted 
to Mr. O. Salvin and Sir G. F. Hampson for advice and assistance in many points 
of difficulty and for the loan of specimens. Col. Swinhoe has also lent us many of his 
types and allowed us to describe new species in his collection; and so far as we 
know there is no collection containing many types to which we have not had access, 
except that of M. Mabille, which we have not visited!. Some species which have 
been described by the latter and by Plétz we have, in common with Captain Watson, 
failed to identify, and we have possibly overlooked a few others ; yet we think that the 
material at our disposal has been hitherto unrivalled, and the localities given for 


1 Since the aboye was written M. Mabille has been kind enough to lend us a number of his types, which we 
have been able to identify with certainty and put in their proper order ; so we do not think that there can exist 
in Europe, at present, any important sources of information in this family of which we have not availed 
ourselyes, 


A REVISION OF THE ORIENTAL HESPERIIDA. 103 


each species may be relied upon as those of specimens actually in coll. Elwes or 
examined personally by us. We have not, as a rule, quoted localities given by other 
authors, because in some cases it is doubtful to what species they refer. 

As regards the respective share which the authors haye had in this paper, we think it 
best to say that Edwards alone is responsible for the whole of the drawings and dissections 
of genitalia and for the conclusions derived from them; he has arranged in the form 
of analytical tables such of the differential characters of the several species as he found 
to be diagnostic in the greatest degree, and he has had the principal share in the 
descriptions of genera and species. Elwes alone is responsible for all the localities, 
geographical distribution, and questions of specific distinction arising from their 
consideration. 

With regard to the geographical area covered by the paper, though it relates 
specially to Asia, we have thought it best to take in the Hesperiide of Europe, 
because their male genitalia had not previously been extensively studied, and in some 
cases this study has led to striking results, but we have not included those of North 
America because they belong for the most part to the Neotropical region. We have 
not attempted to deal with the species occurring in the Malayan islands east of 
Wallace’s line, because, although a good many Indian forms occur in them, the Austro- 
Malayan element is probably predominant, and our knowledge of the Hesperiide of 
the New Guinea Region is too small to enable them to be profitably classified at 
present. 

In the descriptive portion of the paper we have used the Continental system of 
numbering the veins in preference to that adopted by many English authors, because it 
seems to us shorter and more convenient. We have adopted a few terms from 
Mr. Meyrick’s recent ‘ Handbook of British Lepidoptera’: thus the dorsum is the 
edge of the wing opposite to the costa, the termen the edge of the wing opposite to 
the base, and the tornus is the angle in which the dorsum joins the termen. ‘The 
transverse vein, sometimes called the upper, middle, and lower discocellular nervules, is 
that which forms the apical boundary of the discal cell. ‘The subcostal and median 
segments are the pieces of the subcostal and median veins which lie between the bases 
of veins 7-11 and 2-4 respectively; they are numbered from the base: thus the 
second median segment forms the base of cell 2 and so on. The simple vein nearest to 
the dorsum in both fore and hind wings is called vein la, and the second vein from 
the dorsum of the hind wing is called vein 1b. With the exception of the discal cell 
in each wing, which is referred to simply as the cell, the name of each cell is taken 
from that of the vein forming its lower boundary. 

With a few unimportant exceptions, this paper will be found to contain a concise 
statement of the diagnostic characters of all the species of Hesperiide hitherto 
described from the region treated of; although we have not given or quoted 
descriptions in full, except in the case of newly-described species and a few obscure 
ones which we have not seen. 

P2 


104 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Although the Hesperiide as a group are very well-marked and easy of recognition, 
their classification inter se is a matter of great difficulty, owing, mainly, to the paucity 
of index characters. Venation is probably of less assistance here than in any other 
group of Rhopalocera. The fore wing has a free yein near the costa, another near 
the dorsum, and a discal cell from which arise ten simple veins; the hind wing has 
a discal cell giving off six simple veins, a free vein near the costa, and two near the 
dorsum. ‘The modifications of this type are few in number and slight in character. 

Taking the group as a whole, we may divide off—First, Pyrrhopyge and its allies, 
a group characterized by the large blunt recurved club to the antenne and the cell 
of the fore wing more than two-thirds as long as the costa; and secondly, Ismene 
and its allies, a group characterized by the porrect and filiform third joint of the palpi. 
There then remains a vast number of species for the grouping of which no such 
satisfactory characters are available. They may, however, be divided into two series 
according to the position and direction of vein 5 in the fore wing, in which particular 
they may be regarded as conforming to one of two types: the first, exemplified by 
Thanaos tages, in which vein 5 of the fore wing is straight and therefore practically 
parallel to vein 6; and the second, exemplified by Augiades sylvanus, in which 
the basal third of vein 5 is decurved towards the origin of vein 4, and consequently 
recedes to that extent from vein 6. This basis of subdivision is not invariably well- 
marked, but that circumstance does not give rise to any real difficulty, because in 
doubtful cases the correct position of a given species is determinable from other 
considerations, 

All the specimens, dissections, and drawings of genitalia, including a great number 
not figured, are open to the examination of any naturalist who may be interested 
in the subject. 

We have not considered it necessary to give a full synonymy of the species dealt 
with, but only those references which are original or important. 


ORTHOPH@TUS. 


Pteroxys, Watson, P. Z.S. 1893, p. 29, nom. proce. Type phaneus, Hew. 
Orthophetus, Watson, Jour. As. Soc. Beng. 1895. 


In defining this genus Watson says “fore wing: ¢ with a costal fold”; this is true 
of lidderdali, Elwes, and lalita, Dohy., but not of phanwus, Hew., which, however, he 
gives as the type. In the latter species as well as lidderdali, Elwes, and Jalita, Dohy., 
the third joint of the palpi is concealed; but it would be better to extend the 
definition of the genus so as to include the Celenorrhinus omeia of Leech, a species 
which is very closely allied to the three former, but has the third joint of the palpi 
distinct ; there is no costal fold in the male of omeia, Leech. 

The species might be thus distinguished :— 


A REVISION OF THE ORIENTAL HESPERIIDA, 105 


1 (6). Third joint of palpi concealed. Hind wing below with a dark discal 


spot. 
2 (3). No costal fold in the male. Upperside fusco-fulvous. Postmedian 
series of spots on hind wing above small, black . . . . . . . phaneus, Hew. 


3 (2). Male with a costal fold. 
4 (5). No pale spots in cells 4 and 5 of fore wing above. Upperside bright 
fulvous. Postmedian series of spots on hind wing above black, 
withvorwithoutewbite centres!) =). 4) =) sl eee ene) Calta ohys 
5 (4). Fore wing above with pale spots in cells 1-8. Postmedian series of 
spots on hind wing above black, with yellow edges Elly A lidderdali, Elwes. 
6 (1). Third joint of palpi distinct. Hind wing below plain brown. No 
mostallielelimnitmale 5 5 6 a 6 5 55g 0 68 a 5 6 0 6 MMA IEE, 


!1 ORTHOPH@TUS PHANAUS. 
Eudamus phanwus, Hewitson, Descr. Hesp. p. 14 (1867). 
Casyapa phaneus, Distant, Rhop. Mal. p. 386, pl. xxxv. fig. 18 (1886) ; Watson, Hesp. Ind. p. 109 

(1891). 

In point of colour Distant’s figure represents 0. lalita rather than O. phancus, but 
the Perak specimens in my coll. agree with the Bornean type. 

Hab, Perak, Pulo Laut (Doherty); Labuan (coll. Stgr.); Selesseh, Sumatra 
(Martin in coll. Rothschild). 


! ORTHOPH@TUS LALITA. 


Erionota? lalita, Doherty, Jour. As. Soc. Beng. 1886, p. 263. 
Casyapa phaneus, Elwes, P. Z.S. 1892, p. 657. 


Until Mr. Edwards separated this from the Bornean species by the costal fold, Elwes 
had confused it with 0. phaneus. It is, as he pointed out, much yellower in colour, 
but has similar markings. It has only been taken, so far as we know, by Doherty 
in the Chittagong and Karen hills. 

Hab. Chittagong (Doherty); E. Pegu (Doherty). 


ORTHOPHETUS LIDDERDALI. (Plate XX. fig. 18, ¢.) 


Cheticnema? lidderdali, Elwes, Trans. Ent. Soc. Lond. 1888, p. 459. 
Casyapa lidderdali, Watson, Hesp. Ind. p. 109 (1891). 


The type in the British Museum remains unique ; but as Dr. Lidderdale’s collection 
was made at Buxa in Bhutan and Darjeeling it is almost certainly from one of those 
localities, probably the former. We append further particulars of this specimen, 
supplementary to the original description cited above. 

¢ witha costal fold. Upperside: fore wing warm brown, with ten ochreous hyaline 
patches placed as follows :—one across the end of the cell, one each in cells 2-8, and 


1 This sign is prefixed to the names of the species which are in Mr, Elwes’s collection. 


106 MESSRS. H, J. ELWES AND JAMES EDWARDS: 


two, one above the other, near the middle of cell 1 a@; an ochreous line along vein 12 
and also along the upper and lower margins of the cell, an ochreous line bordering 
cell 2 within from the base as far as the hyaline spot, and an ochreous streak in the 
base of cells 9 and 10; hind wing warm brown, basal two-thirds clothed with long 
brownish-yellow hairs, a postmedian series of eight elongate oval, velvety black, 
yellow-edged spots, placed as follows:—two in cell 14 and one in each of cells 2-7. 
Fringe of the fore wing brownish grey, of the hind wing yellowish grey. Thorax 
concolorous ; abdomen brown, hind margin of the segments narrowly pale Underside as 
above, save that there is on the hind wing a dusky oblong spot next the transverse vein. 
Hab. Bhutan? (Lidderdale). 


! ORTHOPH@TUS OMEIA, 
Celenorrhinus omeia, Leech, Butt. China &c. p. 572, pl. xxxviii. fig. 5, g (1894). 


Leech suggests that a new genus is necessary for this species; but, as it differs from 
Orthophatus chiefly by the form of the palpi, we do not agree with him. 
Hab. Omei-shan, W. China. 


CALLIANA. 
Calliana, Moore, P. Z. 8. 1878, p. 686; Watson, P. Z.S. 1893, p. 31. Type pieridoides, Moore. 
The single species in this genus is well distinguished by its facies. The antenne 
in the male are about half as long as the costa, the club slender, the apiculus gradual, 
about half as long as the club. 


CALLIANA PIERIDOIDES. 
Calliana pieridoides, Moore, P. Z. 8. 1878, p. 687, pl. xlv. fig. 2, ¢; de Nicéville, Jour. Bomb. 
Nat. Hist. Soc. 1891, p. 377, pl. G. fig. 25, 2. 


Hab. Khasia Hills (Hamilton) ; Margherita, Upper Assam (Doherty). 


CaPILA. 
Capila, Moore, P. Z. 8. 1865, p. 785; Watson, P. Z. 8. 1893, p. 30. Type jayadeva, Moore.* 


Moore founded this genus for his Zsmene jayadeva, but the insect which he described 
as the female of jayadeva is really the male of another species for which we adopt the 
name C. zennara, Moore, in accordance with a suggestion of Captain Watson. The 
females of these two species were the originals of the two sexes of Moore’s genus 
Pisola. 

We include here Rhopalocampta translucida, Leech, which, however, may be not 
congeneric with the other species. 

The distinctive characters of the species under consideration may be thus 
expressed :— 


A REVISION OF THE ORIENTAL HESPERIID®, 107 


1 (6). With a pencil of long hairs at the base of hind tibiz (7. e. males). 
2 (5). Dark brown above, with white rays in the cells of both wings. 
3 (4). Crown, thorax, and base of wings sordid orange . . . . . . . . jayadeva, Moore. 
4 (3). Crown, thorax, and base of wings sordid brown . . - . «+ . . zennara, Moore. 
5 (2). Dark brown above; disc of both wings tienearent sordid white, 

crossed by dark veins . . oy Poa xs * 2 ae 9 trensiucida, Leech, 
6 (1). No pencil of long hairs at base of hind Fe (i.e. ferales), 
7 (8). Hind wing above brown, with two grey lines in the outer half of each cell. jayadeva, Moore. 
Si(7)= Ebndiwine’above plam browm = . . =. . =: . «=. =. « « « sennara, Moore. 


The female of C. translucida, Leech, is unknown. 


! CAPILA JAYADEVA. 
Ismene jayadeva, Moore, Cat. Lep. E.I. C. i. p. 248 (1857 
Capila jayadeva, Moore, P. Z. 8. 1865, p. 785, pl. xlii. fig. 3g',49 ; Watson, Hesp. Ind. p. 25 (1891). 

Hab. Sikkim (Mller); Naga Hills, K. Pegu (Doherty). 

It will he impossible to work out the distribution of this species and the next 
correctly without an examination of the actual specimens referred to by various writers 
in their local lists, because the authors of such lists, in recording Capila jayadeva and 
Pisola zennara, have but rarely mentioned the sex of their specimens. 


!CAPILA ZENNARA. 
g =Capila jayadeva, Moore, ?, P. Z.S. 1865, p. 785 
2 =Pisola zennara, Moore, 3, t.c. p. 786. 


Hab. Sikkim (Moller, Elwes); E. Pegu (Doherty). 


CAPILA TRANSLUCIDA. 
Rhopalocampta translucida, Leech, Butt. China &e. p. 642, pl. xxxix. fig. 11, ¢ (1894). 


Hab. Omei-shan, W. China (Leech). 
We are indebted to Mr. Leech for the opportunity of examining his unique example 
of this species. 


CROSSIURA. 
Crossiura, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 189%, p. 350. Type pennicillatum, de Nicév. 


This genus, in our opinion, might well be merged in Orthophetus, where the single 
species would follow omeia, Leech. The fore wing in both sexes is banded as in the 
female of Capila, and there is in addition a series of small white spots, one in each of 
cells 4 to 8 or 5 to 8, independent of sex; the male has no costal fold, but the 
dorsum of the hind wing is folded under, and the tornus bears, near the end of 
vein 10, a tuft of long hairs, and there is also a fringe of long hairs to that part of the 


108 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


wing which lies below vein 14. Mr. de Nicéville (Jour. Bomb. Nat. Hist. Soc. 1892, 
p. 351) speaks of the anal angle (tornus) being fringed with long stout sete; this is 
misleading, as the hairs forming the tuft and fringe are not more bristle-like than 
those which form the tibial pencil in many species of Hesperiide. These observations 
are, in part, based on an examination of a male specimen of C. pennicillatum, de Nicév., 
ex coll. E. Swinhoe. 


CROsSIURA PENNICILLATUM. 
Crossiura pennicillatum, de Nicéville, t. c. p, 351, pl. J. fig. 1 g,2 9. 


Hab. Khasia Hills, where it seems rare or very local. 


ACHALARUS. 


Achalarus, Scudder, Syst. Rev. Am. Butt. p. 50 (1872); Watson, P.Z.S. 1893, p. 33. Type 
lycidas, Sm. Abb. 
Lobocla, Moore, Jour. As. Soc. Beng. 1884, p. 51. Type diana, Atkinson. 


Vein 5 in the hind wing is obsolete in /ycidas and most of the American species, 
but well-developed in didiana and all the Eastern species, as well as some of the 


American. 
The species known to us we distinguish as follows :— 


1 (2). Fore wing above: white discal band reaching vein 1 a, white 

spot in cell 3 filling up the base of that cell . . . . . Uliana, Atkinson. 
2 (1). Fore wing above: white discal band not reaching vein 1 a, white 

spot in cell 3 usually not filling up the base of that cell. 
3 (6). Fore wing above: white spot in cell 3 contiguous to that in 

cell 2. 
4 (5). Clothing of second joint of palpi grey. Hind wing below: 

terminal grey band irregular, not wide enough to reach 

the narrow subterminal pale band . . . . . . . . difasciatus, Brem. & Grey. 
5 (4). Clothing of second joint of palpi blackish. Terminal grey 

band on hind wing below suffused, broad, regular, reaching 

about half the length ofcell3. . . . «. «. . «. . « simplex, Leech. 
6 (8). Fore wing above: white spot in cell 8 not touching that 

in cell 2. 
7 (12). Dark spots on the hind wing below very prominent, much 

darker than the ground-colour. 
8 (11). Fore wing below: hyaline spot in cell 5 not touching that 

in cell 6. 
9 (10). Hind wing below: subterminal dark spot across cells 16 and 

2 as wide where it is cut short by vein 3 as where it is 

crossed by vein 2, the pale border of its inner edge 

straight, that of its outer edge sinuate, receding from the 

termen as it approaches vein 3. . . . . . . . . « proximus, Leech, 


A REVISION OF THE ORIENTAL HESPERIIDA, 109 


10 (9). Hind wing below: subterminal dark spot across cells 16 and 

2 ending in a point near the middle of vein 3, the pale 

border of its inner edge evenly convex, that of its outer 

edge nearly parallel with the termen and continuous with 

the pale outer border of the spots in cells 3 and 4-5 . . 1! frater, Ob. 
11 (8). Hyaline spots in cells 4-8 of fore wing forming a continuous 

SETICS eck iougen sr eer aCe i) | ues saree en Germanus: Ob: 
12 (7). Dark spots on hind wing below not very prominent, but little 

darker than the ground-colour. Otherwise very similar 

1 2b GRU ~~ 8 6 6 5 6 boo 6 o o 9 6 0 gH, Old 


! ACHALARUS LILIANA. 


Plesioneura liliana, Atkinson, P. Z. 8. 1871, p. 216, pl. xii. fig. 2. 
Lobocla liliana, Watson, Hesp. Ind. p. 157 (1891). 
Lobocla casyapa, Moore, Jour. As. Soc. Beng. 1884, p. 52; Watson, t.c. p. 158. 

Hab. Khasia (Hamilton); EK. Pegu (Doherty) ; N.W. Himalaya (Lang). 

A specimen of A. casyapa from Moore is marked with the locality Mynpoorie, but 
as this is on the plains of the N.W. Provinces, while Lang and Reed have both 
taken it in the Himalaya, I doubt its correctness. ‘This specimen, notwithstanding 
its widely separated habitat, is inseparable from liliana. 


! ACHALARUS BIFASCIATUS. 
Eudamus bifasciatus, Bremer & Grey, Schmett. N. China’s, p. 10, pl. iu. fig. 1 (1853) ; Oberthiir, 
Htud. d’Ent. xi. pl. vi. fig. 47 (1886). 
Goniloba bifasciatus, Ménétriés, Mus. Petr., Lep. i. pl. v. fig. 3 (1855). 
Achalarus bifasciatus, Leech, Butt. China &c. p. 560 (1894). 
Achalarus bifasciatus, var. contractus, Leech, |. c. pl. xxxvii. fig. 9, g. 


Hab. Central and Western China (Pratt); Korea (Leech); Amurland (Jankowsky). 


! ACHALARUS SIMPLEX. 


Eudamus simplex, Leech, Entom. xxiv. Suppl. p. 58 (June 1891). 
Eudamus gener, Oberthiir, Htud. d’Ent. xv. p. 18, pl. i. fig. 2 (July nec June 1891). 
Achalarus simplex, Leech, Butt. China &c. p. 561, pl. xxxviii. fig. 12, d (1894). 


Hab. Western China (Pratt). 


! ACHALARUS PROXIMUS. 


Eudamus proximus, Leech, Entom. xxiv. Suppl. p. 59 (June 1891). 
Achalarus proximus, Leech, Butt. China &e. p. 560, pl. xxxviii. fig. 7,3 (1894). 


Hab. 'Ta-tsien-lo, Moupin, W. China (coll. Oberthiir). 


* In A. prowimus the extreme base of cell 7 on the hind wing below is whitish ; this point is not indicated 
in the figure of A. frater, but it is not certain that it had not disappeared from the specimen figured as 
the result of rough handling. 

VOL. XIv.—ParT Iv. No. 2.—October, 1897. Q 


110 MESSRS. H. J, ELWES AND JAMES EDWARDS 


ACHALARUS FRATER. 
Eudamus frater, Oberthiir, Etud. d’Ent. xv. p. 18, pl. i. fig. 8 (July nec June 1891). 
Hab. Yunnan (Delavay, fide Oberthiir). 


! ACHALARUS GERMANUS. 
Eudamus germanus, Oberthiir, Etud. d’Ent. xi. p. 26, pl. vi. fig. 48 (1886). 
Achalarus germanus, Leech, Butt. China &e. p. 561 (1894). 

Hab. Ta-tsien-lo (Biet); Pu-tsu-fong, Wa-ssu-kow, 5000-8000 feet, June and July 
(fide Leech). 


1 ACHALARUS NEPOS. 
Eudamus nepos, Oberthiir, |. c. pl. vi. fig. 49. 
Achalarus nepos, Leech, 1. ¢. 


Hab. Ta-tsien-lo (Biet) ; Pu-tsu-fong, 8000 feet, July (Leech). 


HANTANA. 


Hantana, Moore, Lep. Cey. i. p. 179 (1881) ; Watson, Hesp. Ind. p. 144 (1891) ; id. P. Z. S. 1893, 
p- 87. Type infernus, Felder. 


The type of this monotypic genus is well distinguished among its allies by the 
dark purple-brown colour of the upperside and the absence of hyaline spots except 
four on the fore wing, namely, one in the cell next the apical fourth of its upper 
edge, and one each (small and roundish) in cells 6,7, and 8. ‘he hyaline spot in 
the cell of the fore wing has a tendency to disappear, and the base of cell 4 is twice 
as long as the base of cell 5. 


! HANTANA INFERNUS. 
Eudamus infernus, Felder, Verh. zool.-bot. Gesellsch. Wien, 1868, p. 283. 
Hantana infernus, Moore, Lep. Cey. i. p. 179, pl. Ixvii. fig. 6 (1881); Watson, Hesp. Ind. 
p. 145 (1891). 


Hab. Ceylon (Mackwood). 


CHARMION. 
Charmion, de Nicéyville, Jour. As. Soc. Beng. 1894, p. 48. Type ficulnea, Hew. 


Antenne more than half as long as costa, somewhat shorter in the female, club 
slender, apiculus robust, about half as long as the club. Palpi appressed, densely 
scaled, third joint concealed. Fore wing: vein 5 straight, nearer 6 than 4, vein 2 
from the basal fourth of the lower margin of the cell. No alar sex-mark in the male. 
Tibial epiphysis present. Species fuliginous brown, with an oblique white median 
band on fore wing, abbreviated at each end. 


A REVISION OF THE ORIENTAL HESPERIIDA, 111 


In C. ficulnea the hind tibie of the male are thickly fringed on their basal half, 
and have at the base a thick tuft of long ochreous hairs nearly as long as the joint itself. 


! CHARMION FICULNEA. 
Hesperia ficulnea, Hewitson, Descr. Hesp. p. 37 (1868). 
Plesioneura signata, Druce, P. Z. 8. 1878, p. 360, pl. xxxiii. fig. 8. 
Plesioneura leucographa, Plétz, Berl. ent. Zeits. xxvi. p. 262 (1882). 
Charmion ficulnea, de Nicéyille, Jour. As. Soc. Beng. 1894, p. 49. 
Hab. Perak (Doherty); Kina Balu, Borneo (Waterstradt); Sumatra (Martin in 
coll. Rothschild). 


CHARMION TOLA. 

Plesioneura tola, Hewitson, Ann. & Mag. Nat. Hist. ser. 5, vol. i. p. 840 (1878). 
Plesioneura zawi, Plotz, Berl. ent. Zeits. 1885, p. 225. 
Plastingia? plesioneure, Staudinger, Ex. Schmett. p. 299, pl. C, 9 (1888). 

This species is distinguished from the preceding by having a small roundish 
projection of the white band on the fore wing reaching about halfway across cell 1 a. 
Hewitson’s type isa male. The type of P. zawi, Plétz, lent to me by Dr. Staudinger 
is a female. 

Dr. Staudinger concurs in our opinion that his P. plesionewrw is synonymous 
with C. tola. 

Hab. Tondano (fide Hewitson) ; Minahassa, Celebes (fide Staudinger). 


CELHNORRHINUS. 


Celenorrhinus, Hiibu. Verz. p. 106 (1816); Watson, P. Z.S. 1893, p. 49. Type eligius, Cr. 
Gehlota, Doherty, Jour. As. Soc. Beng. 1889, p.131. Type sumitra, Moore. 

Species large and robust; dorsum of the fore wing usually longer than the termen. 
Fore wing with several white spots of which some form an oblique discal band, or 
with yellow spots forming an oblique discal band, or with a continuous yellow oblique 
discal band. Hind wing rounded, its termen as a rule not evidently excavated 
before the tornus, in several species much spotted with yellow. Antenne half as 
long as the costa or longer, club slender, apiculus gradual, the shaft sometimes pure 
white in front. Palpi appressed, third joint suberect. Vein 5 of the fore wing 
straight, arising as a rule distinctly nearer to vein 6 than to vein 4, but in flavocincta, 
de Nicév., intermediate between the two. Hind tibie with two pairs of spurs, and 
bearing a hair-pencil in the male. 

The different phases of this large genus are well exemplified by swmitra, Moore, 
leucocera, Koll., and aurivittata, Moore, respectively. 

Though de Nicéville so recently as 1889 gave an elaborate revision of this genus}, 

? Jour. Bomb. Nat. Hist. Soc. 1889, p. 177. 
Q2 


112 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


yet we have in many cases been obliged to dissent from the opinions he arrived at, 
probably because the material at our disposal is much more complete than his at 
the time he wrote. 

The following is a list of the species known to us; the value of the characters relied 
on for their discrimination has been carefully considered and supplemented by an 
examination of the male genitalia of numerous specimens :— 


1 (58) Hind wing below, not brown with a narrow yellow spot on 
the transverse vein, its fringes not pale unspotted yellow. 
). Fore wing with a pale spot in each of cells 6, 7, and 8. 
). Fore wing with a small pale spot near the base of cell 1 a’. 
13). Macular band on fore wing pure white. 
). Basal spot in cell 1a of fore wing yellow. 
). Male: shaft of antenne white in front throughout; hind 
wing distinctly produced on vein 16. Expanse 62 mm. pero, de Nicév. 
7 (6) .Male: shaft of antenne white in front only near the base 
of the club ; hind wing rounded, not produced on vein 1 6. 
Expanse49mm. . ... .- - +e « + « « pulomaya, Moore. 
8 (5). Basal spot in cell 1 @ of fore wing ie 
9 (10). Hind wing below with two well-defined yellow spots in the 
middle of the cell, the lower one fare to the base by a 


yellow streak . . . . maculosa, Feld. 
10 (9). Hind wing below with only one eayeile mein in the ‘middle of 
the cell. 
11 (12). Hind wing below with the pale spots orange-yellow ; fringe 
orange-yellow, scarcely interrupted with darker . . . aspersa, Leech. 
12 (11). Hind wing below with the pale spots cream-coloured ; 
frmge chequered ©. . = : . ee « + Consanguinea, Leech. 
13 (4). Macular band on fore wing above dirty Gite: Sua 
grey-brown, fringes chequered. . . . . . ambareesa, Moore. 


14 (8). Fore wing with no pale spot near the base of a la, 
15 (16). Hind wing below (except a brown terminal band) orange- 
yellow, with several (about 9) black spots . . . . . *flavocincta, de Nicév. 
16 (15). Hind wing not as in 15. 
17 (34). Fore wing above with the pale macular band pure white. 
18 (27). Fore wing above; extreme base of cell 3 not filled up 
with white. 
19 (26). Hind wing above not conspicuously clothed with brownish- 
yellow scales and hairs almost to the termen. 


1 Very rarely this spot is traceable in the female of lewcocera, Koll. 

2 This species will probably be found to belong to the group (par. 3 supra) which has a small pale spot near 
the base of cell 1a of the fore wing, as such a spot is indicated in a male specimen from the Mdller collection ; 
the point, however, is of no consequence, as the species is distinguished from ail its congeners by the pattern and 
coloration of the hind wing below. 


A REVISION OF THE ORIENTAL HESPERIIDA, 


20 (21). Pale spots on hind wing below large, numerous, and orange- 
yellow; shaft of antenne in the male not entirely 
WAMBO 5 6 Ge . ne 

21 (20). Pale spots on hind wing below neither eee numerous, nor 

orange-yellow. 

22 (25). Pale spots on hind wing below small, stramineous, sharply 
defined, variable in number, sometimes reduced to one 
only next the transverse vein. 

228 (22>). Shaft of antennz in the male spotted with white in front . 

22> (224). Shaft of antennz in the male entirely white in front. 

23 (24). Pale spot in cell 2 of fore wing as wide as or wider than 

high. Fore wing broader and less pointed . 

24 (23). Pale spot in cell 2 of fore wing at least one and a half times 
as high as wide, its inner and outer edges both straight. 
Fore wing narrower and more pointed, termen straight 
and forming an angle of about 110° with the dorsum 

25 (22). Pale spots on hind wing below suffused and indistinct, or 
entirely wanting. Shaft of antenne in the male not 

entirely white in front 


Hind wing above clothed Hrgushens with feowniat ellen 


6 (19). 

scales and hairs except a narrow terminal space . : 

27 (18). Fore wing above: extreme base of cell 3 filled up with 
white. 

28 (33). Fore wing above with one or two white spots in cell 1 a. 

29 (30). Upperside blackish brown; fringe of the hind wing brown, 
not chequered . . . Greta) eo oe oe 

30 (29). Fringe of the hind wing cieaeereall 

31 (82). Upperside grey-brown, white spot in cell 2 of fore wing 


subrhomboidal or irregular, always ee on the 
inner edge . : 
32 (31). Upperside blackish Bee eine pak in al 2 of fore 
wing irregularly roundish . eo Mer pe eC 
). Fore wing above without any white spot in cell 1 @ nor in 
cells 4 and 5 : eee dc - 
34 (17). Fore wing above with the ale feed vallowel ahite, 
yellow, or orange-yellow. 
35 (48). Pale spot in cell 1 @ of fore wing hyaline. 
36 (37). Pale spot in cell 1 a of fore wing reaching to vein 1 @ 
37 (36). Pale spot in cell la of fore wing not reaching to 
vein ] a. 
38 (47). Pale spot in cell 3 of fore wing filling up the extreme base 
of that cell. 
39 (46). Pale spot in cell 1 @ of fore wing subquadrate or irregular, 
joining that in cell 2 near its lower outer angle. 


33 (28 


113 


sumitra, Moore. 


maculicornis, n. sp. 


leucocera, Koll. 


leucocera var. angustipennis. 


spilothyrus, Feld. 


chamunda, Moore. 


balukinus, n. sp. 


nigricans, de Nicév. 


orbiferus, 1. sp. 


asmara, Butl. 


tibetana, Mab. 


50 (51). 


51 (50). 


. Pale spot in cell 2 


. Expanse 382 mm. 


MESSRS. 


. Pale spot in cell 1 a of fore wing not produced into a tooth 


at its muer lower angle. 
of fore wing reaching inwardly much 


beyond the base of vein 3. 


. Expanse 38 mm. Tegumen and clasp as in figures 1, 


la, 16, Plate XXII. a Esher 
Tegumen and ‘taeps as in figures 2, 


2a, Plate XXII. . 


. Pale spot in cell 2 of fore wing not cha poe 


beyond the base of vein 3. 


. Pale spot in cell 1 a of fore wing bites into a fies 


tooth at its lower inner angle 


. Pale spot in cell 1 a of fore wing small, punctiform, joming 


that in cell 2 near its half length 


. Pale spot in cell 3 of fore wing not filling up the extreme 


base of that cell . 1°. 2. °° 


. Pale spot in cell 1 a of fore wing opaque. 
. Fore wing with the pale band deep orange-yellow, 


practically unicolorous throughout, 7. e. the part in 
cells 2 and 3 and that across the cell are semitransparent, 
but do not differ appreciably in tint from the other 
parts. 

Fore wing: breadth of the pale band where it crosses the 
cell equal to about one seventh of the length of the 
costa, its outer edge regular, not broken ee the pro- 
jection of the pale spot in cell 3 . : 

Fore wing: breadth of the pale band where it crosses the 
cell equal to about one fifth of the length of the costa, 
its outer edge broken by the sate of the pale 
spot incell3 . 


. Fore wing with the pale band Pe ee in calle 2 ia 3 


and across the cell, distinctly stronger in colour in cell 
1 a and next the costa. 


4). Fringe of hind wing chequered . 
. Fringe of hind wing brown ee Fe HPC : 
. Fore wing above with a yellow oblique -bubinedian bail 


but no pale spot in cells 6, 7, or 8. Hind wing above 
brown. 


. Hind wing below brown, basal half yellow-brown by 


reason of a thick clothing of yellow scales, a feeble 
suffused pale spot next the transverse vein, and a 
suffused yellow spot at the apex of cell 1 b 


. Hind wing below plain brown : 0 0 
. Hind wing below brown, with a narrow A Silo ed on the 


transverse vein; fringe pale yellow, unspotted . 


H. J. ELWES AND JAMES EDWARDS: 


dhanada, Moore. 

andamanica, W.-M. &de Nicév. 
inequalis, n. sp. 

dentatus, n. sp. 
fulvescens, n. sp. 


saturatus, 0. sp. 


cameront, Dist. 


lativittus, n. sp. 


affinis, un. sp. 


aurivittata, Moore. 


ladana, Butl. 
batchianus, n. sp. 


badia, Hew. 


A REVISION OF THE ORIENTAL HESPERIIDA. 115 


! CELHZNORRHINUS PERO. 


Celenorrhinus pero, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1839, p. 183, pl. B. fig. 12, ¢; 
Elwes, P. Z.S. 1892, p. 660. 


Hab. Naga Hills (Doherty). 


! CELHNORRHINUS PULOMAYA. 


Plesioneura pulomaya, Moore, P. Z.S. 1865, p. 787. 
Celenorrhinus lucifera, Leech, Butt. China &c. p. 571, pl. xxxix. fig. 5, 2 (1893-4). 


Hab. Himalayas from Kangra to Sikkim, Naga Hills (Doherty); Moupin (fide 
Leech). 


! CELHNORRHINUS MACULOSA. 


Pterygospidea maculosa, Felder, Reise ‘ Novara,’ Lep. iii. p. 528, pl. lxxiii. fig. 7, § (1867). 
Celenorrhinus maculosa, Leech, Butt. China &c. p. 569, pl. xxxix. fig. 2, g. 


Hab. Central and Western China (Pratt). 


! CELHNORRHINUS ASPERSA. 
Celenorrhinus aspersa, Leech, Entomologist, xxiv. Suppl. p. 61 (1891); id. Butt. China &e. 
p- 571, pl. xxxix. fig. 4, g (1893-4). 
Celenorrhinus clitus, de Nicév. Jour. Bomb. Nat. Hist. Soc. 1891 (published 2 Feb. 1892), 
p. 878, pl. G. fig. 26, 3d ; Elwes, P. Z.S. 1892, p. 660. 
Hab. Naga Hills, Bernardmyo (Doherty) ; Western China (fide Leech.) The type 
of clitus is in coll. Elwes. 


! CELANORRHINUS CONSANGUINEA. 


Celenorrhinus consanguinea, Leeck, Entomologist, xxiv. Suppl. p. 61, 1891; id. Butt. China &e. 
p- 570, pl. xxxix. fig. 3, g. 


Hab. Central and Western China; Eastern Tibet (Pratt, Kricheldorf ). 


! CELZNORRHINUS AMBAREESA. 


Plesioneura ambareesa, Moore, P. Z. 8. 1865, p. 788; de Nicéville, Jour. As. Soc. Beng. vol. hi. 
p. 87, pl. x. fig. 9, 2 (1883). 
Hab. Western Ghats from Mahableshwar to Goa; Nilgiris (Swinhoe, Hampson). 


! CELHNORRHINUS FLAVOCINOTA. 
Plesioneura flavocincta, de Nicéville, P. Z.S. 1887, p. 464, pl. xl. fig. 9, 2. 
Hab. Sikkim (Knyvett). 


! CELHNORRHINUS SUMITRA. 
Plesioneura sumitra, Moore, P. Z.S. 1865, p. 787. 


116 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Celenorrhinus pyrrha, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 181, pl. B. fig. 11, ¢. 
Celenorrhinus patula, de Nicéville, t. c. p. 182, pl. B. fig. 4, 2. 

Celenorrhinus playifera, de Nicéville, t. c. p. 182, pl. B. fig. 18, 2. 

Celenorrhinus pluscula, Leech, Butt. China &e. p. 571, pl. xxxix. fig. 6, 9 (1893-4). 


Hab. Sikkim (Elwes) ; Naga Hills, Bernardmyo (Doherty) ; China ( fide Leech). 


! CELHNORRHINUS MACULICORNIS, n. sp. (Plate XVIII. fig. 8, ¢.) 

3. This species may be distinguished from C. lewcocera by reason that the 
upperside or front of the shaft of the antennz in the male is minutely and regularly 
spotted with black and white. This might be considered a trivial and insufficient 
character, were it not for the fact that the male of C. leucocera, whatever may be the 
state of the spotting of the wings, always has the upperside or front of the shaft of the 
antenne continuously pure white from the base to the apex. 

There are in coll. Elwes more than twenty males of C. leucocera from various 
localities, and we have examined at least as many more in other collections without 
finding any trace of a tendency to spotting on the front of the shaft of the antenne. 
The relationship of C. maculicornis to C. leucocera is quite different from that of 
C. leucocera var. angustipennis; the latter being separated from C. leucocera by 
differences in the spotting and shape of the wings, which are distinctly variable 
in G. leucocera; whilst the character by which we separate C. maculicornis is one 
which proves to be constant in C. leucocera. The male genitalia do not afford 
material for its separation from C. leucocera or C. sumitra. 

Hab. Shillong, Khasias, 6400 feet (Elwes); ? Kumaon? (fide Doncaster t Khasias). 

Described from two males in coll. Elwes. It seems probable that this species is 
passed over in collections as the female of C. lewcocera, which has brown antenne 
with a broad white band on the club. We do not know the female of C. maculi- 
cornis. 


! CELHNORRHINUS LEUCOCERA. 
Hesperia leucocera, Kollar, in Hiigel’s ‘ Kaschmir,’ vol. iv. p. 454, pl. xviii. figs. 3, 4, 9 (1848). 
Plesioneura putra, Moore, P. Z. 8. 1865, p. 788. 
P. leucocera, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1881, p. 257. 
P. munda, Moore, Jour. As. Soc. Beng. 1884, p. 48. 
P. leucocirca, Elwes, Trans, Ent. Soc. Lond. 1888, p. 462. 
Celenorrhinus putra, Watson, Hesp. Ind. p. 139 (1891). 
C. munda, Watson, l. c. 


Hab. N.W. Himalaya (Young, Hocking); Sikkim, Khasias (Elwes); Nagas, Burmah, 
Bali (Doherty); Andamans (de Roepstorff’); Nilgiris (Hampson); C. China (Pratt) ; 
Java (Fruhstorfer). 

A common and variable species. The type of putra, which is in the British Museum, 
is from Java. 


A REVISION OF THE ORIENTAL HESPERIiD/. 117 


Var. ANGUSTIPENNIS, nov. 

This form, which is apparently confined to W. Java, is readily distinguished by the 
characters given in the table above, and particularly by the greater length of the costa 
of the fore wing in proportion to the dorsum. 

Hab. Preanger, West Java, circa 5000 feet (Piepers); Sukabumi, 2000 feet 
(Fruhstorfer). 

Var. BRAHMAPUTRA, Stgr. MS. 

This also is a long-winged form, from Kina Balu, Borneo (Waterstradt), and wants 
the white spots in cells 4 and 5 on the fore wing above. ‘The hind wing above is very 
slightly spotted (in the male) or without spots (in the female). 

All the insular forms of lewcocera appear to be of a darker colour and less spotted 
on the hind wing above than those from Continental India. 


! CELH]NORRHINUS SPILOTHYRUS. 
Eudamus spilothyrus, Felder, Verh. zool.-bot. Gesellsch. Wien, 1868, p. 283, d. 
Plesioneura ruficornis, Mabille, Ann. Soc. Ent. Belg. xxi. p. 32 (1878). 
Plesioneura spilothyrus, Moore, Lep. Cey. vol. i. p. 179, pl. 67. figs. 4, 4a (1880-81). 
? (?var. dist.) Plesioneura fusca, Hampson, Jour. As. Soc. Beng. 1888, p. 367. 
Hab. Sava (Staudinger); N. Canara (Aitken) ; Malabar (Vidal) ; Nilgiris (Hampson) ; 
Ceylon (Green). 
We have received three specimens from Bali and one from Arjuno, Java, collected 
by Mr. Doherty, which may constitute a distinct local race; they are, however, only 
distinguishable from spilothyrus by the almost complete disappearance of the spots on 
the hind wing below, and agree exactly with a specimen of his P. ruficornis sent by 
M. Mabille; they have the costal spot of the fore wing white instead of yellow, thus 
resembling South Indian specimens (P. fusca, Hampson) in which this spot is usually 
but not always white, and not Ceylon specimens in which it seems to be always yellow. 


! CELA NORRHINUS CHAMUNDA. 
Plesioneura chamunda, Moore, P. Z. S. 1865, p. 788. 

Hab. Sikkim (Moller); Khasias (Hamilton); Nagas (Doherty); Shan Hills 
(Manders). 


! CELHNORRHINUS BALUKINUS, n. sp. (Plate XVIII. fig. 1, ¢.) 


Plesioneura balukina, Staudinger, in lilt. 


In addition to the characters laid down in the table above, this species differs 
from C. nigricans, de Nicév., as follows:—The white spot in cell 1a of the fore wing 
is irregularly roundish, not reaching halfway across the cell, and placed next the apex of 
the lower edge of the white spot in cell 2. On the underside the hyaline spot in 

VOL. XIV.—part Iv. No. 3.—October, 1897. R 


118 MESSRS. H,. J. ELWES AND JAMES EDWARDS: 


cell 1 a of the fore wing is attended on the outer side by two suffused whitish blotches 
divided by the intra-neural fold. The hind wing below is dark brown with somewhat 
of a purple shade, and there is a feeble suffused yellow spot next the transverse vein 
and a subterminal series of suffused indistinct yellow spots, of which the strongest 
are the two in cell 1 0. 

Expanse 323-3634 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from three males and two females ex coll. Staudinger. Males only 
are in coll. Elwes. 


! CELHENORRHINUS NIGRICANS. 
Plesioneura nigricans, de Nicéville, Jour. As. Soc. Beng. 1885, p. 123, pl. ii. fig. 6, 2. 
Hab. Mandi, N.W. Himalaya (Young); Sikkim (J/6ller) ; EK. Pegu (Doherty). 


CELEHNORRHINUS ORBIFERUS, n. sp. (Plate XVIII. fig. 2, 3.) 


g. Very near to C. nigricans, de Nicév., from which it differs in the following 
particulars :—Upperside darker brown, the white spot in cell 2 of the fore wing 
suborbicular, occupying the whole width of the cell; the white spot in cell 1a of 
the fore wing minute, punctiform, and placed near the apical fifth of vein 2. On 
the underside of the fore wing the white spot in cell 1 @ is larger than on the upperside, 
and accompanied by a suffused subquadrate whitish spot. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from one example ex coll. Staudinger. 


! CELANORRHINUS ASMARA. 

Plesioneura asmara, Butler, Trans. Linn. Soe. Lond., Zoology, ser. 2, vol. i. p. 556 (1877)%; 
Distant, Rhop. Mal. p. 400, pl. xxxv. fig. 28 (1886). 

Celenorrhinus consertus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 222, pl. E. fig. 12, 3. 

Celenorrhinus cacus, de Nicéville, t. c. p. 228, pl. HE. fig. 11, d ; Elwes, P. Z. S. 1892, p. 661. 

Plesioneura goto, Mabille, Ann. Soc. Ent. Belg. xxvii. p. lvi (1883). 

Notocrypta goto, Leech, Butt. China &e. p. 628, pl. xxxviii. fig. 4, ¢ (1894). 

Plesioneura palajava, Staudinger, Isis, ii. p. 156, pl. 11. fig. 10 (1889). 

Celenorrhinus palajava, Semper, Schmett. Philipp. p. 360 (1892), sec. spec. comm. 

Hab. EK. Pegu, Pulo Laut, Borneo (Doherty); Khasia Hills, Rangoon (fide de 
Nicéville); Malacca (Pinwill in B. M.); Japan? (fide Mabdille); Palawan (Platen) ; 
E. Java (Piepers). 

We have examined Staudinger’s type of palajava, which agrees with what Elwes 
followed de Niceville in calling C. cacus, but which agrees with an example from 
Malacea figured by Distant as asmara, Butl. We have seen Mabille’s type of 
P. goto, but strongly doubt the habitat given by him. 


A REVISION OF THE ORIENTAL HESPERIID. 119 


! CELZNORRHINUS TIBETANA. 


Pterygospidea tibetana, Mabille, Ann. Soc. Ent. Fr. ser. 5, vol. vi. p. liv (1876). 
Yotocrypta tibetana, Leech, Butt. China &e. p. 628, pl. xxxviii. fig. 6, ¢ (1894). 


Hab. West China (Pratt); Siau-lou (? Yunnan, fide Oberthiir). 


! CELHNORRHINUS DHANADA. (Plate XXII. figs. 1, 1a, 10.) 
Plesioneura dhanada, Moore, P. Z. 8. 1865, p. 789. 
Kerana dhanada, Watson, Hesp. Ind. p. 150 (1891) ; Hiwes, P. Z. S. 1892, p. 663. 

Hab. Sikkim (1/6ller). 

We have seen no specimen of this species except from Sikkim. It is therefore probable 
that so-called dhanada from the Khasias and Burmah in collections are really C. affinis 
(vide p. 121 post). 


! CELANORRHINUS ANDAMANICA. (Plate XXII. figs. 2, 2 a.) 
Plesioneura dan, var. andamanica, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1881, p. 257 ; 


id. op. cit. 1887, p. 391. 
Coladenia dan, var. andamanica, Watson, Hesp. Ind. p. 120. 


In the male of this species the lower lobe of the clasp is acute with a strong 
tooth near the middle of its upper edge, and the tegumen is bifid in its apica] third. 
Hab. Andamans (de Roepstorff); Cherra Punji (fide Swinhoe). 


CELZNORRHINUS INEQUALIS, n. sp. (Plate XVIII. fig. 3, 6.) 


3. Allied to C. saturatus, from which it differs as follows:—The band on the 
fore wing is narrower and paler, somewhat resembling that of C. tibetana, Mab., 
and the spots which connect the band with the costa are quite hyaline. ‘The pale spot 
in cell 3 fills up the base of that cell, the inner edge of the pale spot in cell 2 does 
not reach further inwards than the base of vein 5, and the pale spot in cell la, 
which joins the apical third of the lower edge of that in cell 2, is oblique, about 
one-half higher than wide, and has its outer edge almost continuous with that of 
the spot in cell 2. On the hind wing below there is a feeble suffused yellow spot 
next the transverse vein and the faint indication of a postmedian series of similar 
spots running parailel to the termen. The fringe of the hind wing is greyish yellow, 
slightly interrupted with brown next veins 2-4, and the short scales are brown. 

Expanse 36 mm. 

Hab. Gedeh, Java. 

Described from one example ex coll. Staudinger. 


CELANORRHINUS DENTATUS, n. sp. (Plate XVIII. fig. 4, ¢.) 


Differs from C. saturatus as follows :— 
$. Upperside browner, the hind wing therefore not exhibiting brown spots; the 
pale spot in cell 3 of the fore wing filling up the extreme base of that cell; the 
R2 


120 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


pale spot in cell la of the fore wing oblong, with a large tooth continuous with 
its inner edge jutting out at its inner lower angle. 

g. Similar to the male, but the hyaline spot in cell la of the fore wing below 
is attended on its outer side by a subquadrate yellow opaque spot. 

Expanse 38-39 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from one pair ex coll. Staudinger. 


CELEZNORRHINUS FULVESCENS, n. sp. (Plate XVIII. fig. 5, 3.) 

é. Differs from C. saturatus in the following particulars:—Rather smaller and 
decidedly brighter and more reddish yellow-brown in colour; the pale spot in cell 3 of 
the fore wing fills up the entire base of that cell, and the pale spot in cell 1 a of the 
fore wing is a mere point situate next the middle of the lower edge of the pale spot in 
cell 2 ; the latter character will serve to distinguish it from the variety of C. saturatus 
in which the pale spot in cell 3 of the fore wing fills up the base of that ceil. 

Expanse 33 mm. 
Hab. Kina Balu, Borneo ( Waterstradt). 
Described from one example ex coll. Staudinger. 


! CELANORRHINUS SATURATUS, n. sp. (Plate XVIII. fig. 6, ¢; Plate XXII. figs. 5, 5a.) 


3. Upperside warm brown, passing into yellow-brown on the hind wing by 
reason of the rather heavy clothing of dull yellow scales: fore wing with an 
oblique sordid yellow median macular: band composed of the following elements :— 
an opaque spot next the costa, a large oblong hyaline spot across the cell having 
its outer edge notched, a hyaline spot ocupying the basal third of cell 3 except 
the extreme base, a large hyaline spot occupying the middle third of cell 2 and 
usually in the form of a parallelogram, and a small roundish or subquadrate hyaline 
spot in cell 1 a, standing next to the apical third of the lower edge of the spot in 
cell 2; three small subequal yellowish hyaline spots in cells 6, 7, and 8, and 
sometimes a pale point near the middle of one or both of cells 4 and 5: hind 
wing with a suffused brown discal spot and a postmedian macular band of suffused 
brown spots running parallel with the termen. Underside: fore wing brown, a 
little paler along the dorsum; pale spots as on the upperside, but that next the 
costa stronger; sometimes there is a suffused pale subtornal spot: hind wing brown, 
with a feeble pattern of suffused dull yellow spots, of which one lies across the 
middle of the cell, one before the middle of cell 7, and the remainder form median 
and postmedian curved. series running parallel with the termen. Fringe of the 
fore wing brown, sometimes paler next cell la; of the hind wing brown, more or 
less chequered with yellowish grey. Antenne brown, more or less spotted with 
yellowish white and with a feeble yellowish ring next the base of the club. Body 
and legs concolorous with the wings. Second joint of palpi clothed with greyish- 


A REVISION OF THE ORIENTAL HESPERIID®. 121 


yellow hairs, with a few black ones intermixed. Lower lobe of clasp subtruncate, 
its upper edge simple; tegumen bifid to the middle. 

@. Similar to the male. 

Expanse 37 mm. 

Hab. Megamendong, Tjampea, Java (Piepers); Java (Fruhstorfer); Arjuno, Java, 
Bali (Doherty). 

Described from six males and one female in coll. Elwes. One of the six males 
has tle base of cell 3 entirely filled up with yellow. This species might be confused 
with large specimens of Coladenia dan, Fab., but the latter may be at once distin- 
guished by the porrect third joint of its palpi. 

Specimens of this species were sent by Herr Snellen as dhanada, Moore, and it is 
probable that the dhanada of the British Museum and some other collections is the 
same thing. 


| CELZNORRHINUS CAMERONI. 


Plesioneura cameroni, Distant, Ann. & Mag. Nat. Hist. ser. 5, vol. x. p. 248 (1882). 
Kerana aurivittata, var. cameroni, ibid. Rhop. Mal. p. 403, pl. xxxiv. fig. 19 (1886). 


Hab. Perak (Doherty); Malacca (ex coll. Stgr.). 


! CELENORRHINUS LATIVITIUS, n. sp. (Plate XVIII. fig. 7, 3.) 
Plesioneura lativitta, Staudinger, in litt. 

3. Resembles C. dentatus, but is rather smaller and darker and also differs from it 
in the following particulars :—Band on the fore wing deep golden yellow, practically of 
the same tint throughout, its inner edge straight and continuous from the costa to vein 2, 
its width where it crosses the cell equal to or exceeding one-fifth of the length of the 
costa; the pale spot in cell 1 @ of the fore wing triangular, adjoining the apical half 
of the lower edge of that in cell 2, its apex nearly touching vein 1 @, its outer edge 
irregular and attended by a more or less distinct suffused yellow spot; on the under- 
side of the fore wing the triangular yellow spot in cell 1@ and a subquadrate spot 
between it and the termen are plain and confluent, forming a large subtornal spot. On 
the upperside the surface of the triangular spot in cell 1a of the fore wing is similar to 
that of the spot in cell 2, but on the underside its surface is scaled as heavily as the 
rest of cell 1 a, but with yellow scales, 

2. Similar to the male, but a little larger and paler. 

Expanse 35-37 mm. 
Hab. Kina Balu, Borneo (Waterstradt). 
Described from one pair in coll. Elwes. Others are in Dr. Staudinger’s collection. 


! CELHZNORRHINUS AFFINIS, n. sp. (Plate XVIII. fig. 9, 9; Plate XXII. fig. 4.) 


Differs from dhanada, Moore, in the particulars set forth in the table of species. 
Having regard to the male genitalia, it is much more nearly allied to C. dhanada than 


122 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


to ©. aurivittata, and it is, probably, the species upon which Watson bases his 
assertion (Hesp. Ind. p. 149) that the fringe of the hind wing in aurivittata is not 
invariably brown. 

Hab. Khasias (Hamilton); E. Pegu (Doherty); ‘Tenasserim (Bingham). 


! CELANORRHINUS AURIVITTATA. (Plate XXII. figs. 5, 3 a.) 
Plesioneura aurivittata, Moore, P. Z. S. 1878, p. 848, pl. liii. fig. 2. 
Kerana aurivittata, Watson, Hesp. Ind. p. 149 (1891). 

The best points for distinguishing this species from dhanada, Moore, lie in the deep 
yellow spot near the tornus of the fore wing, which forms the end of the pale band, 
and the unchequered brown fringe of the hind wing ; in these respects it resembles the 
aurivittata var. cameroni of Distant, which, however, is easily distinguished by the 
difference in the pale band on the fore wing set forth in the foregoing table. Watson’s 
statement that the fringe of auwrivittata is not invariably brown on the hind wing 
leaves one in doubt whether the localities which he gives for that insect really refer 
to the aurivittata of Moore and of this paper or to C. affinis. 

Dr. Staudinger sends from Kina Balu, Borneo,a single female specimen, which differs 
from aurivittata as follows:—The pale spots in cells 6, 7, and 8 of the fore wing are 
unequal, that in cell 7 being at least twice as large as the minute punctiform ones in 
cells Gand 8; the pale spot in the cell of the fore wing reaches two millimetres further 
inward than that in cell 2, and the hind wing below has a natrow yellow spot next the 
transverse vein and a subterminal series of suffused indistinct yellow spots, of which the 
strongest are two near the apex of cell 1 0. 

Hab. Nagas, Upper Burmah (Doherty); Tavoy (Pitman); Andamans (fide Swinhoe). 


! CELANORRHINUS LADANA. 
Carystus ladana, Butler, Trans. Ent. Soc. Lond. 1870, p. 502; id. Lep. Exot. p. 170, pl. lix. fig. 1 
(1874). 
Hab. Perak (Doherty). 


CELANORRHINUS BATCHIANUS, n. sp. (Plate XVIII. fig. 10, ¢). 


3. Upperside dark brown: fore wing with a somewhat narrow pale yellow oblique 
band from the upper edge of the cel] to vein 2, of which latter it adjoins the preapical 
fourth; the inner edge of this band is practically regular. Underside similar to the 
upperside, but somewhat paler. Fringes brown. Antenne brown above, grey-brown 
beneath, the apiculus whitish or yellowish on the underside. Body and legs con- 
colorous with the wings. 

@. Similar to the male, but a little larger and paler, and having in cell 1a of the 
fore wing a triangular yellow spot adjoining the lower edge of the yellow spot in cell 
2 beyond its middle and reaching less than halfway across the cell. On the under- 


A REVISION OF THE ORIENTAL HESPERIID®. 123 


side of the fore wing there is a little yellow suffusion from the upper edge of the cell 
towards the costa. 

Expanse 40-43 mm. 

Hab. Batchian (fide Stgr.). 


Described from one pair ex coll. Staudinger. 


! CELEZNORRHINUS BADIA. 
Pterygospidea badia, Hewitson, Ann. & Mag. Nat. Hist. ser. 4, vol. xx. p. 322 (1877) ; id. Desc. 
Lep. Coll. Atk. p. 4 (1879). 
Plesioneura badia, de Nicéville, Jour. As. Soc. Beng. 1883, p- 88, pl. x. fig. 10, ¢. 
Hab. Sikkim (Mller). 


This seems an extremely rare and local species. 


ABRAXIMORPHA, gen. noy. 


This generic name is proposed for Pterygospidea davidii, Mab., a species winch is 
totally different in facies from any of the species properly referred to either of the 
genera in which it has hitherto been placed, namely, Pterygospidea and Celenorrhinus. 

Antenne, wing-shape, venation, and tibial spurs of Celenorrhinus. Palpi: second 
joint ascending, third well-developed, porrect. Fore wing above greyish black, with 
many irregular pure white spots; hind wing below pure white, with four irregular. 
transverse series of greyish-black spots—basal, antemedian, postmedian, and terminal. 
Front cox of the male with a long hair-pencil on the inner lower side; no hair-pencil 
on the hind tibie. 


! ABRAXIMORPHA DAVIDII. . 
Pterygospidea davidii, Mabille, Ann. Soc. Ent. Fr. 1876, p. liy. 
Celenorrhinus davidi, Leech, Butt. China &c. p. 572, pl. xxxix. fig. 9, 3. 
Hab. Central China (Pratt). 
SARANGESA. 
Sarangesa, Moore, Lep. Cey. 1. p. 176 (1881) ; Watson, Hesp. Ind. p. 53 (1891) ; id. P. Z. S. 1893, 
p. 48. Type purendra, Moore. 
Hyda, Mabille, Bull. Soc, Ent. Fr. (6) ix. p. elxxxiii (1889). Type micacea, Mab. 
Sape, Mabille, Bull. Soc. Ent. Belge, p. Ixvii (1891). Type lucidella, Mab, 
The slight arching of the lower margin of the cell of the fore wing between veins 
2 and 3 in Sarangesa, upon which Watson in his table relies for the separation of the 
former from Coladenia, Moore, is so slight as not to be readily appreciable. 
The following is a table of the species known to us :— 
1 (4). Hind wing below with small pale spots. 
2 (3). A large transparent pale spot in cell 2 of forewing . . . . . purendra, Moore. 
3i(2)Norsuch spot imcelll 2 of forewings... . . . .-. .-. . satt, de Nicev. 


124 MESSRS. H. J. ELWES AND JAMES EDWARDS; 


4 (1). No pale spots on hind wing below. 
5 (6). Hind wing below grey-brown, with three transverse series of 


darker spots’ . . . . dasahara, Moore. 


6 (5). Hind wing below for the most part pale bluish white . . . . . albicilia, Moore. 


!SARANGESA PURENDRA. (Plate XXII. figs. 6, 6 a.) 
Surangesa purendra, Moore, P.Z. 8. 1882, p. 262 ; Watson, Hesp. Ind. p. 54 (1891). 

Hab. Mandi, N.W. Himalaya (Young) ; ? Nilgiris (Roberts). 

As defined by the characters in the foregoing table this form seems a good species. 
We only know it certainly from the N.W. Himalaya, but I have a specimen believed to 
be from the Nilgiri Hills, whence dasahara is recorded by Sir G. F. Hampson. ; 


!SaRANGESA saTI. (Plate XXII. fig. 8.) 
Sarangesa sati, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 391, pl. G. fig. 37,3. 
Hab. Kutch; Rajputana. 


!SARANGESA DASAHARA. (Plate XXII. fig. 7.) 
Nisioniades dasahara, Moore, P. Z. 8. 1865, p. 787. 
Sarangesa dasahara, Watson, Hesp. Ind. p. 54 (1891). 

Hab. Kangra (Hocking); Mandi (Young); Sikkim (Méller); Nagas, E. Pegu 
(Doherty); Ganjam (Minchin); Bombay (Swinhoe); Canara (Aitken); Tenasserim 
(Bingham) ; Akyab (Adamson). 

This species and albicilia are remarkable for the possession, in the male, of a slightly 
curved horn on each side of the base of the terminal part of the tegumen; in purendra 
and sati this part is simple, not subtended by a horn on each side. 


!SARANGESA ALBICILIA. 


Sarangesa albicilia, Moore, Lep. Cey. i. p. 176, pl. Ixviii. figs. 5, 5 a (1881) ; Watson, Hesp. Ind. 
p- 55 (1891). 

Hab. Ceylon (Wade, Mackwood). 

This insect, in its typical state, is evidently a Ceylonese local form of S. dasahara, 
Moore; the male genitalia in each absolutely agree, and the only point of distinction is 
the development in albicilia of the bluish-white shade (with the co-extensive white 
fringe) on the hind wing below, which proceeds from the tornal region and spreads over 
the whole surface, except about the costal third. The two specimens on which 
Mr. Hampson includes a/bicilia in his Nilgiri list (Jour. As. Soc. Beng. 1888, p. 368), and 
of which he remarks that they differed from Ceylon specimens in being dusky instead of 
white on the underside of the hind wing, seem to have been intermediate between the two 
forms. The tendency to the development of the pale suffusion of the hind wing below 


A REVISION OF THE ORIENTAL HESPERIIDZ. 125 


is evident in an otherwise normal specimen from Akyab ; and Moore seems to have had 
in mind specimens similar to this when he wrote that albicilia “ differs from S. dasahara 
in the hind wing being more prominently white.” 


CoLADENIA. 


Coladenia, Moore, Lep. Cey. vol. i. p. 180 (1881); Watson, P. Z.S. 1893, p. 49. Type indrani, 
Moore. 


A genus of few species, placed together on account of their resemblance to “ Plesio- 
neura” indrant, Moore. The third joint of the palpi is porrect. 
The species known to us may be distinguished as follows :— 


Fore wing above with a black or denuded and therefore dark spot or spots 
near the basal third of cell 1 a, visible on one or both surfaces. 
Hind wing below with three or more black spots. 
Fore wing below with a more or less distinct yellow spot in the apex of 
cell 1 a. 
Pale spots in cells 6, 7, and 8 of fore ie subequal in size, their bases 
practically inline . . . ; indrant, Moore. 
Pale spots in cells 6 and 8 of fore wing leecee ‘se that j in cell 7, their 
bases nearly in line with the apex of the spot mcell7 . . . .  tissa, Moore. 
No such spot on the fore wing below. 
Termen of hind wing bluntly angulated between veins 2 and 4. 
Hyaline spot in cell 7 of fore wing well developed. Dark 
spots in cell 7 of hind wing below subequal in size . . . laxmi, de Nicéy. 
Termen of hind wing rounded or scarcely perceptibly produced nciueen 
veins 2 and 4. Hyaline spot proper to cell 7 of the fore wing 
obsolete or wanting. Distal dark spot in cell 7 of the hind wing 
below subquadrate, twice as large as the proximal one . . . sobrina, 0. sp. 
Hind wing below brown, with three transverse curved series of ated 
ochreous spots. . . . ; 0 : oe a dan eal: 
Fore wing above with no dark pee near the Space third of eal ib a. 
Hyaline spots on the fore wing pure white. 
Outermost row of dark spots on the hind wing passing through cells 4—5 
just before the middle. 


Upperside pale brown. Back of tegumen simple . . . . . . . agni, de Nicéy, 
Upperside dark umber-brown. Tegumen witha large dorsal crest. . agnioides, n. sp. 
Outermost row of dark spots on the hind wing passing through cells 4-5 
evidently beyond the middle =: 9. - . ©. . = =. = = = » » 49na, Semper. 


Hyaline spots on the fore wing sordid yellowish white. Outermost row of 
dark spots on the hind wing passing through cells 4—5 just before the 
middle. Upper edge of clasp with one small sharp triangular tooth 
EMP WOT IE “Son GG 8 5. Al deac GEG uwo DD 6 6 tH. faygrh ety. 


vou. XIv.—Part 1v. No. 4.—October, 1897. 8 


126 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! COLADENIA INDRANI. 


Plesioneura indrani, Moore, P. Z. S. 1865, p. 789. 
Coladenia indrani, Watson, Hesp. Ind. p. 118. 


Two specimens from North Canara (Aitken, 11.7.90 and 14.8.90) differ from the 
ordinary form in the umber-brown colour of the upperside, and in the absence of the 
subterminal series of pale markings, except the spot in cell 1 @ on the fore wing below. 
A specimen from the Nilgiris (Hampson, 8.9.84), which is probably of the same 
species as the specimens recorded by Hampson from the Nilgiris as C. tissa, is inter- 
mediate in appearance between the Canara specimens and the ordinary form. The 
male genitalia afford no means of distinction between the three forms. 

Hab. Sikkim (Moller); Burmah (Doherty); Tenasserim (Bingham); N. Canara 
(Aitken) ; Nilgiris (Hampson). 


COLADENIA TISSA. 
Coladenia tissa, Moore, Lep. Cey. i. p. 180, pl. Ixvii. fig. 6 (1881). 


This species, though otherwise closely resembling the preceding, and probably just 
as variable in colour, may be known in both sexes by the distinct displacement inwards 
of the middle one of the three pale spots which form the subapical series. The 
differences in the male genitalia of the two species are merely those of degree. 

Hab. Ceylon (Wade, Mackwood). 


!CoLADENIA LAXMI. (Plate XXII. fig. 10.) 
Plesioneura laxmi, de Nicéville, Jour. As. Soc. Beng. 1888, p. 290, pl. xiui. fig. 5, 2. 
? Netrocoryne atilia, Mabille, Le Naturaliste, 1888, p. 88. 
Plesioneura atilia, var. palawana, Staudinger, Iris, ii. pp. 156, 165, pl. 1. fig. 11 (1889). 
Tapena laxmi, de Nicéville, op. cit. 1891, p. 382, pl. G. fig. 28, g ; Watson, Hesp. Ind. p. 123 
(1891) ; Semper, Schmett. Philipp. p. 316 (1892). 


Hab. Perak (Doherty; ¢ type in coll. Elwes) ; Burmah (Bingham, fide de Nicéville) ; 
Luzon, S.E. Mindanao (Semper); Palawan (P/aten); Kina Balu (Waterstradt) ; 
Namoe Oekor, Sumatra (Martin). 


CoLADENIA SOBRINA, n. sp. (Plate XVIII. fig. 12, ¢.) 

g. Upperside: fore wing fuscous or mouse-grey, with a suffused dark eerie band 
and an oblique white macular median band from vein 2 to the costa, composed of the 
following hyaline spots:—one large and oblong near the middle of cell 2, one small 
and subquadrate near the base of cell 3, not filling up the base of that cell, one large 
and oblong across the cell, and one also oblong between the subcostal and the costa, 
two hyaline points placed obliquely, the lower innermost, beyond the middle of cell la, 
and a small hyaline spot in each of cells 6 and 8, that usually found in cell 7 obsolete 
or wanting; near the basal third of cell 1 @ a somewhat indistinct geminate dark spot or 


A REVISION OF THE ORIENTAL HESPERIIDA, 127 


pair of spots: hind wing fuscous, with antemedian and postmedian irregular curved 
series of blackish spots, and a feeble indication of a narrow suffused subterminal dark 
band from the costa as far as the middle. Underside: fore wing similar to the 
upperside, but somewhat paler and haying the dark spots near the basal third of 
cell 1a@ distinct: hind wing with a postmedian series of black spots corresponding with 
that on the upperside; a black spot in the upper distal angle of the cell, and two in 
cell 7, one near the base and the other, which is subquadrate and reaches across the 
cell, near the middle. Fringes dark fuscous, on the underside with a pale anteciliary 
line. Second joint of palpi clothed with black and pale grey scales intermixed. 
Antenne above dark brown, the apiculus and the underside of the shaft pale grey. 
Body and legs concolorous with the wings. 

Expanse 35 mm. 

Hab. Namoe Oekor, Sumatra (Martin). 

Described from a specimen ex coll. Rothschild. This species most nearly resembles 
C. laxméi in colour and markings, but is easily to be distinguished by its larger size and 
the rounded termen of the hind wing. 


!CoLADENIA DAN. (Plate XXII. figs. 9, 9 a.) 


Papilio dan, Fabricius, Mant. Ins. ii. p. 88 (1787). 

? Hesperia eacus, Latreille, Kne. Méth. ix. p. 738 (1823). : 

Coladenia dan, Distant, Rhop. Mal. p. 398, pl. xxxv. fig. 27 (1886) ; Watson, Hesp. Ind. p. 120 
(1891). 

? Coladenia dan, var. dea, Leech, Butt. China &c. p. 568, pl. xxxviii. fig. 10, g (1894). 

Hesperia fatih, Kollar, Hiigel’s Kaschmir, iv. p. 454, pl. xvii. figs. 5, 6 (1848). 

Coladenia fatih, Watson, t. c. p. 119. 

Watson (¢. ¢. p. 120) records Mr. de Nicéville’s opinion, contrary to that generally 
prevalent, that this species is distinct from C. fatih, Koll.; but he proceeds, “I am 
unable to say by what characteristic they are to be separated, though C. fati/) seems to 
be constantly larger.” ‘The latter view is carried out in Watson’s arrangement of the 
specimens in the British Museum, where the larger specimens are called fatih, Koll., 
and the smaller ones dan, Fab. 

Hab. Kangra (Hocking); Kulu (Young); Sikkim (J/6ller); Margherita (Doherty) ; 
Khasias (Hlwes); Bernardmyo, E. Pegu (Doherty); Akyab, Moulmeir (Adamson) ; 
Perak (Doherty); Java (Fruhstorfer); Sambawa (Doherty); Bali, low country 
(Doherty). 


'CoLADENIA AGNI. (Plate XXII. figs. 11, 11a.) 


Plesioneura agni, de Nicéville, Jour. As. Soc. Beng. 1883, p. 87, pl. x. fig. 4, 2. 
Tapena agni, Watson, Hesp. Ind. p. 122 (1891). 


Hab. Sikkim (Méller); E. Pegu (Doherty); Kina Balu, Borneo ( Waterstradt). 


sz 


128 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


!COLADENIA AGNIOIDES, n. sp. (Plate XVIIL fig. 11; Plate XXII. figs. 12, 12 a.) 


This species is distinguished from agnt in the first place by its dark umber-brown 
colour and the dark umber-brown fringe to cells 6 and 7 on the hind wings below; the 
pale fringe to these cells is not absolutely constant in agni, but is well marked in 4 
out of the 5 male specimens examined. The male genitalia also differ considerably : 
in agnioides there is a large dorsal crest on the tegumen and the upper lobe of the clasp 
is merely a very small pointed strap-shaped piece; in agni the back of the tegumen is 
simple and the upper lobe of the clasp is at least as wide as the lower lobe, rounded 
on the upper edge and triangularly pointed. 

The type in coll. Elwes was taken in the Naga Hills by Mr. Doherty. 


CoLADENIA IGNA. 
Tapena igna, Semper, Schmett. Philipp. p. 316 (1892). 


Hab. Luzon, E. Mindanao (Semper). 


CoLADENIA SEMPERI, 0. sp. 
Tapena lavmi, Semper, Schmett. Philipp. p. 316 (1892), in part. 


This species is founded on the male specimen from Camiguin de Mindanao referred 
to by Semper (J. c.) as differing from his other specimens of /aami in having the hyaline 
spots smaller and yellower, and the hind wing above browner, with the outer row of 
dark spots more distinct. Fortunately we have been able to examine the clasp-form of 
this specimen without dissection, and find that it differs from that of /aami in the 
particulars given in the table above. The angulation of the hind wing also is less 
evident than in laxmi. 

Hab. Philippines (Semper). 


CoLADENIA HAMILTONII. 
Coladenia hamiltonii, de Nicéville, Jour. As. Soc. Beng. 1888, p. 291, pl. xi. fig. 8, ¢ ; Watson, 
Hesp. Ind. p. 121 (1891). 


We transcribe the original description of this species, taken from a single specimen 
from Sylhet :— 

“ Male. Upperside : fore wing olive-greenish fuscous, with two very irregular broad 
discal black fascie joined in the middle; three most minute transparent subapical dots, 
the uppermost the largest, placed at the outer edge of the anterior portion of the outer 
black fascia; a very minute similar spot in the second median interspace; a very 
attenuated spot across the middle of the first median interspace, both placed on the 
outer black fascia; the inner margin somewhat broadly irrorated with greyish scales , 
a submarginal indistinct broad blackish fascia. Hind wing: ground-colour much as in 


A REVISION OF THE ORIENTAL HESPERIIDA. 129 


the fore wing, but the outer third of the wing irrorated with grey scales ; a recurved 
black macular decreasing band from the costa near the apex of the wing to the second 
median nervule; the discocellular nervules defined by a pale line. Underside: both 
wings vinous fuscous. Fore wing with the transparent spots as above. Hind wing 
with the disc irrorated with whitish ; the macular black band much as above; an ante- 
ciliary whitish line. Cilia fuscous. 

Expanse, 3, 1:6 inches.” 

The figure gives one the idea of a much-worn and possibly abnormal specimen, and 
it is remarkable that the species, if species it is, has never since turned up among the 
numerous collections recently sent from the Khasias. 


CoLADENIA BUCHANANII. 
Celenorrhinus buchananii, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 187, pl. B. fig. 2, 2. 


We transcribe the original description of this species, which was described from a 
single specimen from the Ruby Mine District, Upper Burmah, as we haye not seen a 
specimen :— 

« Female. Very closely allied to C. lazmi, mihi, from which it differs in its con- 
siderably larger size. Upperside: fore wing with the white discal band fully twice as 
wide, not divided into spots, extending uninterruptedly from the costa to the submedian 
nervure, its edges very irregular, its lower portion posterior to the first median nervule 
much narrower than the rest of the band: this species lacks the two obliquely placed 
black dots found near the base of the submedian interspace in C. /avnu. Hind wing, 
instead of possessing two parallel discal macular black bands, has a rounded black spot 
towards the end of the discoidal cell and a discal series of 6 black spots, of which the 
anterior one is round and well separated from the spot which follows it, the second 
spot is round, the next pair are the largest and elongated, and the last pair smaller but 
also elongated; cilia of hind wing anteriorly white, posteriorly dark brown. Under- 
side: both wings with the same differences as above, but all the spots of the hind wing 
more prominent.” 

Although this species is compared with C. laxmi it agrees better with C. agni in the 
shape of the hind wing. 


CoLADENIA VITREA. 
Coladenia vitrea, Leech, Butt. China &c. p. 569, pl. xl. fig. 15, 5 (1894). 


This species, which was described from a single specimen in the collection of Mr. H. 
Grose Smith, taken at Ta-tsien-lo, may be distinguished by the hyaline spots on the 
hind wing below: these, with the exception of one near the base of cell 7, form two 
irregular transverse series, and are placed as follows :—a subquadrate one near the 
middle of cell 7, one filling the basal third of cell 6, a large subquadrate oae occupying; 


130 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


the distal half of the cell, a narrow one across cell 1 6 before the middle, one in cells 
4-5 just beyond the transverse vein, about half as large as that in the cell, one filling 
the basal third of cell 3, an oblong one near the middle of cell 2, and a narrow one 
across cell 1 4 just beyond the middle. 


! COLADENIA MANIATA. 
Coladenia meniata, Oberthiir, Etudes d’Ent. xx. p. 42, pl. ix. fig. 164, g (1896). 


Closely allied to C. vitrea, Leech, but differs in the number and position of the 
hyaline spots in the hind wing below as follows :—the spot in cell 6 does not fill the 
apex of that cell, there is no spot in cells 4—5, nor across cell 16 before the middle. 
The termen of the hind wing also is more evidently produced near vein 4 than in 
C. vitrea. 

Hab. E. Tibet (fide Oberthiir). 

M. Oberthiir was good enough to give Elwes a specimen of the species taken at a 
place called Meenia, which he says is near Ta-tsien-lo; but, notwithstanding the points 
of difference above noted, we think it may prove to be only a variety of C. vitrea, 
which came from the same district. 


SATARUPA. 


Satarupa, Moore, P.Z.8. 1865, p. 780; Watson, Hesp. Ind. p. 87 (1891); id. P.Z.S. 1893, 
p. 46. Type gopala, Moore. 
Daimio, Murray, Ent. Mo. Mag. xi. p. 171 (1875) ; Watson, P. Z. S$. 1893, p.47. Type tethys, Mén. 


It would be better to place in this genus all the species which Watson puts into the 
genus Daimio. Murray used the latter name for tethys, Mén., but if such insects as 
sinica, Feld., phisara, Moore, and sambara, Moore, are associated with gopala, Moore, 
and nymphalis, Speyer, in one genus, as they well may be, then tethys, Mén., which is 
evidently congeneric, should be with them and the name Daimio becomes unnecessary. 

Watson says of Satarupa: “Closely allied to Daimio, from which it may be sepa- 
rated by the shape of the wings, especially of the hind wing, by the much greater 
length and more decided hook in the terminal portion of the antennal club, and by the 
scaling of the hind tibie of the male.” Of the points on which he relies to separate 
the genera, the greater length of the hind wing is only found in gopala, Moore; the 
more decided hook in the terminal portion of the antennal club is merely a matter of 
degree, and the scaling of the hind tibie of the male is not sufficiently uniform in the 
species to form a generic distinction. Of Satarwpa he says: “In the male the hind 
tibiz are fringed along their upper edge, and the inner side of the tibie is clothed with 
long, coarse, recumbent hairs,’”—thus omitting all mention of the tuft of long hairs which 
springs from the upperside of the base of the hind tibie in gopala, Moore, which he 


A REVISION OF THE ORIENTAL HESPERIIDZ, 


makes the type of the genus (nymphalis, Speyer, in which a similar structure occurs 


was, apparently, not before him), 
rupa, there is no trace of the tibial pencil in the male. 


In sambara, Moore, which he also places in Sata- 
It appears, therefore, that 


whilst the tibial pencil is always present in the males of Daimio, it is also found in two 
species of Satarupa (gopala, Moore, and nymphalis, Speyer) and absent from one 
(sambara, Moore). 

The genus Satarupa was used by Moore, its founder, in the extended sense in which 


it is here treated. 
The following is an analytical table of the species known to us :— 


12 (9) 
13 (4) 
14 (3) 
15 (16) 
16 (15) 
17 (2) 

(19) 


. Clothing of lower side of second joint of palpi yellow. 

. Pale band on hind wing above similar in colour in both sexes. 

. Pale band on hind wing above pure white or smoke-grey. 

. Pale band on hind wing above pure white. 

. Pale spot in cell of fore wing adjoining its upper edge or reaching 


across the cell. Species large; expanse 62-73 mm. 


. Pale spot in cell of fore wing subtriangular, adieming its upper edge 


but not reaching the lower 


. Pale spot in cell of fore wing eipreionn feneline quite across 


the cell 


. Pale spot in cell of fore wing ean next wife fase of call 3. Species 


small, not exceeding 48 mm. 


. White band on hind wing below complete, reaching the costa. 
. Pale spot in cell la of fore wing nearly twice as wide as that in 


cell 2. White band on hind wing much wider than the marginal 
dark band 


. Pale spot in cell 1a of tere wing Paakennal in mite to chat in cell 2 2. 


White band on hind wing not wider (sometimes ce than 
the marginal dark band 


. White band on hind wing below cut ee by vein 8, not ~aeankne 


the costa . 


. Pale band on hind wing shove smoke: -grey. Pale snot in ‘cell 2 


small, not reaching vein 2, in shape like the lower half of a crescent. 


. Pale band on hind wing above cream-coloured or sordid white. 
. Pale spot in cell of fore wing large, reaching from side to side 
. Pale spot in cell of fore wing small, next the base of cell 3, or absent. 
. Pale band on hind wing above bright yellow in the male, white in 


the female; pale band on hind wing below white or yellowish 
white in both sexes. 


. Hind wing above with an indistinct suffused pale subterminal line, 


the space between it and the pale discal band somewhat darker 
than the space between it and the termen, and broken up by 
slightly paler lines along the veins into a series of dark spots . 


gopala, Moore. 


nymphalis, Speyer. 


narada, Moore. 


diversa, Leech. 
dire, de Nicév. 
fumosa, 0. sp. 


bhagava, Moore. 
phisara, Moore. 


celebica, Feld. 


132 MESSRS. H, J. ELWES AND JAMES EDWARDS: 


9 (18). Hind wing above with a subterminal series of suffused spots (on the 
upperside brownish grey in the male and whitish in the female, on 
the underside white in both sexes), the space between it and the 
discal band not broken into spots by pale lines. ... . . . . corona, Stgr. 
(1). Clothing of lower side of second joint of palpi pure white. 
(26). No white spot in cell of fore wing. 
(25). Abdomen above wholly or in part white. 
5 (24). Abdomen above entirely white. Base of hind wing below bluish white. dohertyi, Watson. 
( 


24 (23). Abdomen above dark at base and apex, white in the middle. Base 

of hind wing below brown . .... =. =.=. =.=. =... +. sambara, Moore, 
25 (22). Abdomen above entirely brown . .. . Cah . . . affinis, Druce, 
26 (21). A large white spot reaching quite across cell of ‘fore wing. ‘Avdomnien 


dark with narrow pale rings or entirely dark, 
7 (28). White band on hind wing above very distinct . . . . .  sinica, Feld, 
28 (27). White band on hind wing above absent or but faintly ‘ndicaied . .  tethys, Mén. 


! SATARUPA GOPALA. 
Goniloba gopala, Moore, Cat. Lep. Mus. HI. C. i. p. 246. 
Satarupa gopala, Moore, P. Z. 8S. 1865, p. 780, pl. xli. fig. 1; Watson, Hesp. Ind. p. 90 (1891), 


Hab. Sikkim (Moller); Khasia (Hamilton). 


! SaraARUPA NYMPHALIS. 
Tagiades nymphalis, Speyer, Stett. ent. Zeit. xl. p. 348 (1879) ; Staudinger, Mém. sur Lép. iil. 
p- 153, pl. vii. fig. 4 (1887). 
Satarupa nymphalis, Leech, Butt. China &c. p. 562 (1894). 
Hab. Amurland (fide Speyer) ; Chefoo, North China (in B. I.) ; Central and Western 
China (Pratt). 


! SATARUPA NARADA. 
Satarupa narada, Moore, Jour. As. Soc. Beng. 1884, p. 51; Watson, Hesp. Ind. p. 89 (1891). 


Hab. Sikkim (Méller); E. Pegu (Doherty). 


!SATARUPA DIVERSA. 
Plerygospidea diversa, Leech, Entomologist, xxiii. p. 46 (1890). 
Daimio narada, var. diversa, Leech, Butt. China &c. p. 566, pl. xxxviii. fig. 14, g (1894). 


This insect is quite as distinct from S. narada, Moore, as are some of the admitted 
species inter se, and there seems to be no advantage in treating it as a variety of 
narada. 

Specimens from the Khasia Hills agree with those from Central China. We have 
not seen it from any intermediate locality. 


A REVISION OF THE ORIENTAL HESPERIIDA. 133 


! SATARUPA DIRA&. 


Daimio dire, de Nicéville, Jour, Bomb. Nat. Hist. Soc. 1895, p. 369, pl. Q. fig. 49, g. 
Tagiades graya, Staudinger, MS, 


Hab. Pulo Laut; Arjuno; Java; Bali (Doherty); N.E. Sumatra (fide de Nicéville). 

Abdomen black, hind margin of the segments narrowly white. 

A single female from Pulo Laut, which otherwise much resembles dire, has the 
abdomen white with the apical third black. 


SATARUPA FUMOSA, n. sp. (Plate XVIII. fig. 13, 3.) 


3. Upperside fuliginous brown: fore wing with seven white hyaline spots—one, 
the largest, in shape like the lower half of a crescent, in cell 2 next the base of vein 3 
and not quite reaching vein 2, one in cell 3 outwardly oblique from the base of vein 4, 
four small and punctiform (one near the middle of cells 5 and 8 and one near 
the basal third of cells 6 and 7), and one small and roundish near the apical fourth 
of the upper edge of the cell: hind wing with a broad discal band and a narrow 
subterminal macular band smoke-grey. Underside: fore wing as on the upperside and 
having two suffused subquadrate white spots divided by the intraneural fold near the 
apex of cell 1a: hind wing fuliginous brown, with bluish-white hairs near the base, a 
broad pure white discal-band from the dorsum as far as vein 7 and continued into ~ 
cell 7 by a suffused whitish spot near the middle of the lower edge of that cell, and a 
subterminal series of seven suffused sordid whitish spots, two in cell 16 and one in 
each of cells 2-6. Fringes and body above concolorous with the wings. Antenne 
blackish, apiculus grey beneath. Clothing of the second joint of the palpi and the 
breast yellow, of the legs sordid white. Abdomen beneath white. 

Expanse 35 mm. 

Hab. Sumatra (Martin). 

Described from one specimen in coll. Rothschild. 


!SATARUPA BHAGAVA. 


Satarupa bhagava, Moore, P.Z. 8. 1865, p. 781; Watson, Hesp. Ind. p. 88 (1891). 

Satarupa phisara, var.?, Elwes, P. Z.S. 1892, p. 655, 

Tagiades bhagava, var. andamanica, Wood-Mason & de Nicéville, Jour, As. Soc. Beng. 1881, 
p. 256, pl. iv. fig. 5, ¢; Watson, l.c. 


Hab. Bernardmyo, Burmah (Doherty) ; Tavoy (Zucker); Sikkim. 

We identify this species by Moore’s description alone. He gives N.E. Bengal as the 
habitat. It may occur in Sikkim, but we have no specimens from there. 

The var. andamanica, according to the figure, differs from the type in wanting the 
cream-coloured subquadrate spot near the middle of cell 1@ on the fore wing above 
and the cream-coloured streak below vem 1a@ by which the spot is continued to the 

VoL. XIv.—Part iv. No. 5.—October, 1897. T 


134 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


dorsum; it agrees with the type on the underside. Judging from the description, 
however, it is a mere trifling variety with no constant character and not worthy of a 
name. 


!SATARUPA PHISARA. . 


Satarupa phisara, Moore, Jour. As. Soc. Beng. 1884, p. 50; Wood-Mason & de Nicéville, op. cit. 
1886, p. 390, pl. xvii. fig. 4, ¢; Watson, Hesp. Ind. p. 89 (1891). 
Satarupa bhagava ?, de Nicéville, Jour. As. Soc. Beng. 1883, p. 90, pl. x. fig. 14, 2. 


Hab. Sikkim (M@ller); Khasia (Hamilton). 


! SATARUPA CELEBICA. 
Pterygospidea celebica, Felder, Reise Nov., Lep. iii. p. 528, pl. Ixxili. fig. 8, ¢ (1867). 
Pterygospidea permena, Hewitson, Descr. Hesp. p. 51 (1868). 


Hab. Sula Island (coll. Stgr.); Celebes ( fide Felder). 


! SATARUPA CORONA. 
Tagiades corona, Staudinger, MS. 


Satarupa corona, Semper, Schmett. Philipp. p. 306, pl. xlix. fig. 2, $ (1892). 
Hab. Davao, Mindanao (coll. Staudinger); Philippines (Semper). 


SATARUPA DOHERTYI. 
Satarupa dohertyi, Watson, P. Z.S. 1893, p. 46. 
Rather larger than S. sambara, with the postmedian row of black spots on the hind 


wing below more regular and freer from the dark terminal band. 


Hab. Kumaon ( fide Watson). 


SATARUPA SAMBARA. 
Goniloba sambara, Moore, Cat. Lep. Mus. E.I. C. i. p, 246 (1857). 
Satarupa sambara, Moore, P, Z.S. 1865, p. 781 ; Watson, Hesp., Ind. p. 89 (1891). 


Hab. Sikkim (Moller); Khasia (Hamilton). 


! SATARUPA AFFINIS. 


Satarupa affinis, Druce, P. Z. S. 1878, p. 360, pl. xxxiii. fig, 9. 
Satarupa affinis, var. cognata, Distant, Rhop. Mal. p. 385, pl. xxxy. fig. 17 (1886). 
Tagiades niphates, Weymer, Stett. ent. Zeit. vol. xlviii. p. 15, pl. i. fig. 5 (1887). 


This species is nearest to S. sambara. 


Hab. Kina Balu ( Waterstradt) ; Sumatra (fide de Nicéville); Perak (fide Distant) ; 
Preanger, W. Java (Piepers). 


A REVISION OF THE ORIENTAL HESPERIIDA, 135 


! SATARUPA SINICA. 


Pterygospidea sinica, Felder, Wien. ent. Mon. vi. p. 30 (1862). 

Pterygospidea moori, Mabille, Ann. Soc. Ent. Fr. 1876, p. clii; Alphéraky, Rom. Mém. sur Lép. 
vy. p. 122, pl. v. figs. 9 a, b (1889). 

Daimio felderi, Butler, Ann. & Mag. Nat. Hist. (5) vii. p. 140 (1881). 

Daimio sinica, Leech, Butt. China &e. p. 565 (1894), 


Hab. Central and Western China (Pratt, Leech). 

The insect here dealt with is the same as that figured by Alphéraky as cited above, 
but we have no means of deciding if it is really the true sinica of Felder. 

The sinica, Feld., of the British Museum collection, from North China, is a species 
with the clothing of the second joint of the palpi below yellow, the white band on 
the hind wing below reaching from the costa to the dorsum, and bearing a subquadrate 
brown spot near the apical third of cell 7; the fore wing above has a white band from 
the middle of the dorsum to the hyaline spot in cell 2 three-fourths as wide as that 
spot ; and the abdomen above is white with the apical fourth brown. 


SATARUPA TETHYS. 


Pyrgus tethys, Ménétriés, Cat. Mus. Petr. p. 126, pl. x. fig. 8 (1857). 
Daimio tethys, Pryer, Rhop. Nihon. p. 33, pl. x. fig. 6 6 (1889); Leech, Butt. China &e. p. 564 
(1894). 


Hab. Japan (Pryer, Leech); Amurland ; China (Pryer). 


ODINA. 


Odina, Mabille, Comptes Rend. Ent. Soc. Belg. p. exiii (1891); Watson, P. Z.S. 1893, p. 50. 
Type chrysomelena, Mab.,=hieroglyphica, Butl. 


Species orange-yellow, with blackish-brown markings, the dark pattern practically 
the same on both surfaces. Antenne more than half as long as costa, club gradual, 
apiculus acute, scarcely half as long as the club. Palpi ascending, third joint porrect, 
pointed in cuneiformis, Semper, cylindrical and subtruncate in decoraius, Hew. Fore 
wing: vein 5 a little nearer 6 than 4, vein 2 from the basal third of the lower edge ot 
the cell. 


Table of Species. 


1 (4), Dark markings forming an irregular network on the hind wing 

above. 
2 (3). Hind wing above with the large discal yellow spot triangular . . . hieroglyphica, Butl. 
3 (2). Hind wing above with the large discal yellow spot irregularly 

4-sided . . . . ; a0 Si . cuneiformis, Semper. 
A (1). Dark markings on the hind wing share an enteniediniis ee tcietia 

aud marginal row of distinct black spots . . . . . . . . . decoratus, Hew. 


T2 


136 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


ODINA HIEROGLYPHICA. 

Plastingia hieroglyphica, Butler, Trans. Ent. Soc. Lond. 1870, p. 511; id. Lep. Exot. p. 171, 
pl. lix. fig. 12 (1874) ; Dist. Rhop. Mal. p. 470, pl. xliv. fig. 25. 
Odina chrysomelena, Mabille, Comptes Rendus Soc. Ent. Belg. p. exiii (1891), fide Watson. 

Hind wing above orange-yellow, with a brown band from the middle of the costa to 
the tornus; this band gives off in cell 6 a Y-shaped branch to join the brown terminal 
band, and in cell 2 a simple branch to the same band, the large discal yellow spot is 
therefore triangular; the space between the lower edge of the cell and the dorsum 
brown, with two yellow spots, one near the middle of cell 14, and one near the apical 
third of cell la. Fore wing: yellow spots in cells 2 and 3 forming, when taken 
together, an oblique suboval spot divided by vein 3; that in cell 3 triangular, that in 
cell 2 not or scarcely reaching vein 2. These particulars are taken from the type 
specimen from Borneo in the collection of Messrs. Godman and Salvin. 

The 0. hieroglyphica of the British Museum collection has the hind wing above 
brown, with a large bright yellow spot near the middle of cell 6: this may be an 
extreme variation of the true 0. hieroglyphica, Butl., but we have had no opportunity 
to compare the two insects with the view of deciding this point. 

Hab. Labuan (coll. Staudinger); Perak (fide Distant); Borneo (Low in coll. 
Godman). 


ODINA CUNEIFORMIS, 

Plastingia cuneiformis, Semper, Schmett. Philipp. p. 314, pl. xlix. fig. 11, ¢ (1892). 

Hind wing above orange-yellow; the space between the lower edge of the cell and 
vein 2 and the dorsum brown, except in the apical fourth, and bearing a yellow streak 
in cell 1 a, and a roundish yellow spot near the middle of cell 14; from the middle of 
the costa to the base of vein 2 a straight brown band, which gives off in cell 6 an 
irregularly Y-shaped branch to join the brown terminal band, from the apex of vein 5 
to the apical third of vein 2 an irregular widening brown band, the large yellow discal 
spot is therefore irregularly four-sided. Fore wing: hyaline spots in cells 2 and 3 
forming a short regular yellow band from the middle of vein 2 to the base of vein 4. 

These particulars are taken from Herr Semper’s type specimen from Mindoro. 

Hab. Philippines (Semper). 


!ODINA DECORATUS. 
Eudamus decoratus, Hewitson, Descr. Hesp. p. 17 (1867). 
Pterygospidea decoratus, id. Bx. Butt. vol. v. pl. Pteryyospidea, fig. 2 (1878). 
Entheus bicolor, Oberthiir, Etudes d’Ent. xvii. p. 14, pl. iv. fig. 36, ¢ (1893). 
Hab. Sylhet, Java (fide Hewitson); Garo hills (fide de Nicéville); East Pegu 
(Doherty) ; Tonquin (coll. Oberthiir). 
We have not seen the female of this species, which seems to be unknown. 


A REVISION OF THE ORIENTAL HESPERIID®. 137 


ODINA ORTYGIA. 
Odina ortygia, de Nicéville, Jour. As. Soc. Beng. 1895, p. 531. 


Closely allied to hieroglyphica, Butl., from which it differs “on both surfaces in 
having all the black markings reduced by half, all the orange markings therefore 
greatly enlarged.” 

Expanse 363 mm. 

Hab. Daunat Range, Tenasserim, Burmah ( fide de Nicéville). 

Described from one male specimen. 


Darra. 
Darpa, Moore, P. Z. 8. 1865, p. 781; Watson, op. cit. 1893, p. 58. Type hanria, Moore. 


A genus of one species, distinguished as well by its facies as by its much dentate 
hind wing, which is produced into a strong tooth at the apex of each of veins 2, 3, 5, 
and 6. 


DARPA HANRIA. 
Darpa hanria, Moore, P, ZS. 1865, p. 781, pl. xlii. fig. 2. 


Hab, Sikkim (Moller). 


TAGIADES. 


Tagiades, Hiibner, Verz. p. 108 (1816); Watson, P. Z.S. 1893, p. 55. Type japetus, Cr. 
Pierygospidea, Wallgr. Rhop. Caffr. p. 53 (1857). Type flesus, Fab. 

A genus of soberly coloured species, brought together on account of more or less 
resemblance to Papilio japetus, Cr. In the more typical species the palpi are porrect, 
densely scaled, with the usual lateral rows of hairs rudimentary or absent, the last 
joint small and bluntly conical, and the lower edge of the cell of the fore wing is more 
or less arcuate between the bases of veins 2 and 3; but this latter is a very variable 
feature. Watson gives the hind tibie as “fringed” only, but the males of several 
species have penicillate hind tibiz, and in some the intermediate tibiz are penicillate 
also, 

The following is a table of the species known to us:— 


1 (48). Transverse vein of hind wing erect, 7. e. forming practically a right 
angle with vein 4. 

2 (89). Second joint of palpi densely scaled, when viewed from the side 
suboyate, nearly as wide as long, the lateral row of hairs 
rudimentary or absent. 

3 (26). No hyaline spot in cell 11 of fore wing. 


138 


iN 
na 
£3) 


NX & Ot 
=> 
or 
a 


9 (4). 


10 (23). 
11 (22). 
12 (17). 


13 (14). 


4 (18). 


15 (16). 


16 (15). 


U1) 


18 (21). 


i9 (20). 
20 (19). 


21 (18). 


22 (11). 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


. Hind wing above brown, no bluish-white pubescence near the 


termen. 


. Hind wing beneath brown or grey-brown 
. Hind wing beneath in greater part bluish white. 
. Expanse 40-47 mm. Fore wing beneath without a bluish-white 


suffused patch near the tornus 


. Expanse 49-55 mm. Fore wing beneath with a “uta white 


suffused patch near the tornus ee ois : 

Hind wing above with a white or bluish-white iatan on the lower 
half of the termen, or at least with bluish-white pubescence in 
the same region. 

Hyaline spots in cell of fore wing very small or absent in the male. 

Hind wing below with distinct black spots. 

Pale portion of hind wing above suffused with bluish white, or, if 
the tornal region of the termen is narrowly white, then the 
white passes into bluish white towards the base. 

Hind wing below with the dark costal portion suffused with the 
bluish white of the disc. Hyaline spots in cells 6, 7, and 8 of 
fore wing only in either sex . Béal niente 

Hind wing below with the dark costal nortan sharply separ: ated 
from the white discal portion, at least near the termen. 

Hind wing below : dark costal portion continued along the termen 
towards the tornus as an irregular line which is interrupted in 
cells 3 and 4. Hyaline spots in cell of fore wing wanting in 
both sexes . be seks. ack SOME SIMRO DY Yes cee a 

Hind wing below: dark costal portion continued along the termen 
towards the tornus as an uninterrupted row of spots. Hyaline 
spots in cell of fore wing visible in the male, well developed in 
the female . SathecUre Matt eacared) eles aH. 

Hind wing above: tornal region of the termen more or less 
broadly snow-white. 

Hind wing below: tornal region of the termen with an irregular 
row of 2 or more, more or less confluent, black spots, which is 
interrupted in cells 3 and 4. 

Hind wing below: no dark spot in the end of the cell 

Hind wing below: a dark spot in the end of the cell and one in 
each of cells 3-7, that in cell 7 feeble; in the female the dark 
spots in cells 3-6 are subequal in size and form a regular row . 

Hind wing below: termen with a black border from veins 14 to 
6 interrupted by the intra-neural folds, which appear as white 
lines parallel with the veins; a feeble dark spot in the end of 
the cell and another near the middle of cell 7; a well-defined 
black spot in each of cells 4—5 and 6, the former geminate . 

No well-defined black spots on hind wing below 


ravi, Moore. 


helferi, Feld. 


khasiane, Moore. 


alica, Moore. 


obscurus, Mab. 


japetus, Cr. 


gana, Mcore. 


karea, Mab. 


elegans, Mab, 
lavata, Butl, 


23 (10). 


24 (25). 


25 (24). 


26 (3). 


27 (88). 
28 (29). 
29 (28). 
30 (87). 
31 (36). 
32 (35). 


33 (84). 


4 (88). 


35 (32). 


6 (31). 


7 (80). 


8 (27). 


39 (2). 
40 (48). 
4] (42). 


2 (41). 


A REVISION OF THE ORIENTAL HESPERIIDA, 


Hyaline spots in cell of fore wing large and well defined in the 
male. 

Termen of hind wing below with a black or brown border from 
the tornus as far as vein 5 F he My 
Termen of hind wing below entirely white fron fie 4 tornus as far 

as vein 5 ae : 

Fore wing with a hyaline spat in teal ll. 

Hind wing below with distinct black spots. 

Hind wing above with a dark postmedian spot in cell 10 

No dark postmedian spot in cell 16 on hind wing above. 

Hind wing above with the dorsum white for at least half its length. 

Fore wing below without distinct short white streaks in cell 1a. 

Hind wing above: black terminal spots on veins 1 4 to 4 subequal 
in size, sometimes confluent. 

Tibial pencil in the male yellowish white ; clasp simple. A 
hyaline spot at least indicated in each of cells 2 and 3 of fore 
wing below, and generally two in the cell 5G 

Tibial pencil im the male brown; upper edge of the clasp near the 
apex with a long, curved, pointed horn, half as long as the 
clasp. Fore wing above sometimes with two white points 
placed one aboye the other near the apical third of cell 1 a, 
after the manner of 7. pralaya and T. trichoneura : 

Hind wing above: terminal black spot on vein 14 at least twice as 
large as those on veins 2, 3, and 4. No hyaline spot in cells 2 
and 3 of the fore wing and only one (the upper) in the cell. 

Fore wing below with two distinct short white streaks placed one 
above the other near the apical third of cell 1 a 

Hind wing above with the dorsum white for about one- Sfourth of 
its length ; : aes 

Hind wing below Puicaons Bron withion bas snot a the 
male with a suffused white streak in cell 14 and a feeble 
indication of a suffused terminal macular whitish band arising 
near the tornus and becoming obsolescent before vem 4; in 
the female the whitish markings are more extensive, occupying 
about half of the wing and passing into bluish white at the base, 
and there is a suffused white spot on the transverse vein 

Second joint of palpi laxly scaled. 

No hyaline spot in cell 10 of the fore wing. 

Termen of hind wing distinctly excavate in cells 4-5 and distinctly 
produced on yein 1 4. No yellow on the hind wing above 

Hind wing above with rather more than the terminal third bright 
yellow. Fore wing above with the apical fifth of cell 1a yellow, 
and with hyaline spots in each of cells 4-8, those in cells 6-8 


large and oblong . 


fitus, Plotz. 


pteria, Hew. 


menaka, Moore. 


atticus, Fab, 


sambavana, ni. sp. 


139 


waterstradti, n. sp. 


martinus, Plotz. 


Nana, . Sp. 


nestus, Feld. 


dealbata, Dist. 


princeps, Stgr. 


140 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


43 (40). Fore wing with hyaline spots in cells 10 and 11, those in cells 2 
and 3 linear and oblique. 


44. (45). Pale portion of hind wing below yellow . . . . . . . . « pralaya, Moore, 
45 (44). Pale portion of hind wing below white. 
46 (47). Pale portion of hind wing above yellow . . . - . .  trichoneura, Feld, 
47 (46). Pale portion of hind wing above fants white, oa vallew only 
AGUS OM 6 4 09 0 o o ¢ te See: Bee . . + trichoneuroides, Stgr. 


48 (1). Transverse vein of hind wing oblique, 2. e, forming an ebidee angle 

with vein 4. Second joint of palpi, seen from the side, twice 

as long as wide, the lateral row of hairs well developed. 
49 (50). Fore wing with no hyaline spots in cells 4-8. Hind wing with the 

subterminal series of dark spots in part merging in the dark 

tenmuinallibarid lspecine- meee : j - pinwilli Butl. 
50 (49). Fore wing with hyaline spots in calls 4-8 aad a hyaline cial in 

the base of cell 4 next vein 4. Hind wing with the postmedian 

series of dark spots free from the dark terminal band. . . . ¢abrica, Hew. 


! 'TAGIADES RAVI 

Pterygospidea ravi, Moore, P. Z.S. 1865, p. 779. 

Tagiades ravi, Distant, Rhop. Mal. p. 388, pl. xxxiv. fig. 1, g (1886). 
Distant’s figure (/. ¢.) is rather poor and represents 7’. helferi rather than 7. ravi. 
Hab. Tavoy (Pitman) ; Nicobar, Perak, Pulo Laut (Doherty) ; Nias (Modigliani). 


! TAGIADES HELFERI. 
Pterygospidea helferi, Felder, Verh. zool.-bot. Gesellsch, Wien, vol, xii. p. 483 (1862). 
Tagiades noctis, Stgr. in litt. 

Hab. Nicobar (de Roepstorff’); Camorta (Doherty); Borneo (coll. Stgr.). 


! TAGIADES KHASIANA. 
Tagiades khasiana, Moore, Jour. As. Soc. Beng. 1884, p. 51. 

Hab. Khasias (Hamilton); Nagas (Doherty); Akyab (Adamson); Tavoy (Pitman); East 
Pegu (Doherty) ; Andamans (de Roepstorff’). 


!'TAGIADES ALICA. 

Tagiades alica, Moore, P. Z. 8. 1877, p. 593, pl. lvii. fig. 2, g. 
Tagiades meetana, id. t. c. 1878, p. 842, pl. li. fig. 1. 

Specimens with both series of fringe-scales white in the tornal. region are alica, and 
those with the long fringe-scales brown and the short ones white are meetana; these 
differences, however, depend on the amount of bluish-white scaling on the hind wing 
above. 

Hab. N. Canara (Aitken); KE. Pegu (Doherty); Tavoy (Pitman); Andamans 
(de Roepstorff); Perak, Pulo Laut (Doherty). 


A REVISION OF THE ORIENTAL HESPERIID#. 14] 


! TAGIADES OBSCURUS. 


Tagiades obscurus, Mabille, Ann. Soc. Ent. Fr. ser. 5, vol. vi. p. 274 (1876); Wood-Mason & 
de Nicéville, Jour. As. Soc. Beng. 1886, p. 389, pl. xvii. fig. 9, 2. 
Tagiades distans, Moore, Lep. Cey. vol. i. p. 175, pl. Ixviii. figs. 1, 1 a@ (1880-81). 

Hab. Nilgiri hills (Hampson); Ceylon (Mackwood); Sava? (fide Mabille). 

Though I have not been able to compare Malayan with Ceylon specimens, yet the 
absence of the discal spots relied on by Moore for distinguisking his species is a 
character of no value, the two spots being sometimes absent in Ceylon specimens. 

Tt is possible that obscurus, Mab., has not been correctly identified and may be a 
synonym of japetus, in which case the name distans will stand. 


TAGIADES JAPETUS. 

Papilio japetus, Cramer, Pap. Exot. iy. pl. ecelxy. E, F (1782). 
Tagiades brasidas, Doherty, Jour. As. Soc. Beng. 1891, p. 195. 

Hab. Java (Piepers); Sambawa, Bali, Lombok (Doherty); Amboina (coll. Snellen) ; 
Philippines ? 

Though the characters given in the table to separate this from the last species are 
fairly constant, yet I have two females from the Philippines which have the hind wing 
below rather as in o/scurus. Neither of these species is recorded by Semper from the. 
Philippines. 


'TAGIADES GANA. 


Pterygospidea gana, Moore, P. Z. S. 1865, p. 180. 
Tagiades gana, Distant, Rhop. Mal. p. 388, pl. xxxiv. fig. 2, g (1886). 


Specimens from Java, Nias, Perak, and Pulo Laut are constantly smaller than those 
from Sikkim. 

Hab. Sikkim (Moller); Perak, Pulo Laut (Doherty); Java (Fruhstorfer); Andamans 
(de Roepstorff); Palawan (Platen in coll. Staudinger). 


! TAGIADES KAREA. 


Tagiades karea, Mabille, Comptes Rendus Soc. Ent. Belg, iv. no. 16, p. Ixxiii (1891); Semper, 
Schmett. Philipp. p. 308 (1892). 


Hab, Philippines, generally distributed (Semper). 


! TAGIADES ELEGANS. 


Tagiades elegans, Mabille, Bull. Soc. Ent. Fr. 1877, p. x1; Semper, Schmett. Philipp. p. 309, 
pl. xlix. fig. 4, g (1892). 


Hab. Luzon, E. Mindanao (Semper). 


VOL. XIv.— Part 1y. No. 6.—October, 1897. U 


142 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


!'TAGIADES LAVATA. 
Tagiades lavata, Butler, Trans. Linn. Soc. Lond. Zoology, ser. 2, vol. i. p. 557, pl. Ixix. fig 8 
(1877); Distant, Rhop. Mal. p. 389, pl. xxxiv. fig. 5. 
Hab. N. Borneo (Pryer); Kina Balu, Borneo (Waterstradt); Tavoy (Pitman) ; 
Bunguran, Natuna Island (Everett). 


!TacrapEs TITUS. (Plate XX. fig. 15, ¢.) 

Tagiades titus, Plétz, Jahrb. Nass. Ver. xxxvii. p. 46 (1884) ; Semper, Schmett. Philipp. p. 310 
(1892). 

Tagiades latreillei, Mabille, fide Semper. 

Tagiades japetus, var. latreillei, Staudinger, Iris, ii. p. 160 (1889) (fide Semper). 

Male. Hind wing below white, with a blackish border round the costa and termen, 
its inner edge suffused; a feebly indicated dark spot near the middle of cell 3,a 
geminate spot in cells 4-5 and a larger one in cell 6 blackish. In the female the 
hind wing below is paler and the pale discal portion is bluish white. 

Hab. Philippines (Semper); Palawan (Staudinger). 


TAGIADES PTERIA. 

Pterygospidea pteria, Hewitson, Descr. Hesp. p. 51 (1868) ; id. Exot. Butt. v., Péerygospidea, pl. 1. 
fig. 1 (1873). 
Tagiades pteria, Semper, Schmett. Philipp. p. 308 (1892). 

In this species the hind wing above resembles that of T. pralaya or T. trichoneura, 
save that the pale part is white or yellowish white instead of yellow; it is distinguished 
from these species by the small and roundish hyaline spots in cells 2 and 3 and the 
want of hyaline spots in cells 10 and 11 of the fore wing. 

Hab, E. Mindanao (Semper). 


!'TAGIADES MENAKA. 
Pterygospidea menaka, Moore, P. Z. 8. 1865, p. 778. 
Tagiades atticus, Leech, Butt. China &c. pl. xxxviii. fig. 18, (1892-94), nee Fabr. 

The dark spot near the apical third of cell 1% in the hind wing is absolutely 
diagnostic of this species. It does not appear that this species ever has more than 
one pale spot in the cell of the fore wing, but this character alone will not separate 
it from those specimens of 7. atticus which want the lowermost pale spot in the cell 
of the fore wing. 

Hab. N.W. Himalaya (Young); Nipal; Sikkim (Mller); Khasias; Nagas, E. Pegu 
(Doherty). 


A REVISION OF THE ORIENTAL HESPERIID/. 143 


!'TaGlADEs articus. (Plate XXII. fig. 13.) 
Hesperia atticus, Fabricius, Ent. Syst. iii., i. p. 839 (1793). 
Tagiades atticus, Moore, Lep. Cey. vol. i. p. 175, pl. Ixviti. fig. 2 (1880-81); Wood-Mason 

& de Nicéville, Jour. As. Soc. Beng. 1886, p. 389, pl. xvii. fig. 10, 2. 

Vagiades calligana, Butler, Trans. Linn. Soc. Lond. Zoology, ser. 2, vol. i. p. 556, pl. lxix. fig. 11. 
Tayiades atticus, var. calliyana, Distant, Rhop. Mal. p. 387, pl. xxxiv. fig. 6. 
Tagiades menaka, Staudinger, Iris, ii. p. 159 (1889). 
Tagiades litigiosa, Méschler, Verh. zool.-bot. Wien, xlviii. p. 230 (1878). 

Hab. Sikkim (Moller); Nagas, E. Pegu, Pule Laut (Doherty); Palawan (Platen) ; 
Java (coll. Stgr.). 

We are indebted to Dr. Staudinger for the opportunity of examining the type of 
litigiosa, Moschl., from ‘“ India ? Silhet.” 


!'TAGIADES WATERSTRADTI, 2. sp. (Plate XX. fig. 7,3.) 


3 2. Nearest to 7. atticus, Fab., from which it differs in the following points :— 
On the fore wing there is no hyaline spot in cells 2 and 5, and only one (the upper- 
most) in the cell, and on the hind wing above the black basal portion is more extensive, 
the dorsum being black for at least half its length, and the black terminal spot on 
yein | 0 is at least twice as large as those on veins 2, 3, and 4. 

Expanse 363-583 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from three specimens ex coll. Staudinger, one of which is now in coll. Elwes. 


! TAGIADES MARTINUS. 
Tagiades martinus, Plétz, Jahrb. Nass. Ver. xxxvil. p. 47 (1884) ; Semper, Schmett. Philipp. 
p. 309, pl. xlix. fig. 3, ¢ (1892). 
Hab. Amboina, Philippines, Aru Islands, Mysol (Semper); Celebes (coll. Stgr.). 


! TAGIADES SAMBAVANA, n. sp. (Plate XX. fig. 10,3; Plate XXII. fig. 14.) 


Very similar to 7. atticus, but on the hind wing below the black spot in the cell is 
wanting or very small, and on the upperside there are sometimes two white points near 
the apical third of cell 1a in the fore wing (as seen in 7. trichoneura and T. pralaya). 
The species is however well distinguished by the clasp-form of the male. In T. atticus 
the apical part of the clasp is triangular and its apex, instead of being deflexed and 
serrate as in 7’. menaka, is simple; in 7. sambavana the clasp has a short spiniform 
tooth near the middle of its upper edge, and beyond this, at a distance about equal to 
the distance of the short tooth from the base of the clasp, there arises a long curved 
acuminate horn, which is about equal in length to the greatest width of the clasp. 

Hab. Sambawa, Bali (Doherty). Described from eight males and one female in 


coll. Elwes. 
v2 


144 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


!'TAGIADES NANA, n. sp. (Plate XX. fig. 13, 3.) 

g. Upperside dark olive-brown, with a pure white patch next the tornus of the 
hind wing bearing two black spots. Fore wing with nine transparent white points 
placed as follows, namely, two near the apex of the cell, one near the basal third of 
cell 3, one near the middle of each of cells 4, 5, and 6, one near the basal third of cell 7, 
one near the middle of cell 8, and one near the apex of cell 11; fringe concolorous, 
Hind wing with a pure white patch reaching from the dorsum as far as vein 3, and 
one-fourth or one-fifth the length of the wing, its inner edge rather sharply defined ; 
a large spot at the apex of vein 10 and a smaller one at the apex of vein 2, black; 
fringe white from the tornus to vein 4, thence dark brown. Underside: fore wing a 
little paler than above: hind wing bluish white, costal region as far as vein 6 brown, 
the colours suffused ; a large black spot near the middle of cells 4—5, a smaller one in 
cell 6, and the indication of one or more dark spots nearer the base of the wing; 
on the pale part of the wing there is a fine black terminal line, which expands into 
triangular spots at the apices of veins 1, 2, and 3. Body above dark olive-brown, 
abdomen beneath bluish white. 

Expanse 52 mm. 

This, the smallest of the genus known to us, appears quite distinct; the type 
specimen is in my collection. 

Hab. Khasia hills (native collectors, fide Doncaster); island of Nias (in coll. 
Rothschild). 


TAGIADES TOBA. 
Tagiades toba, de Nicéville, Jour. Bomb, Nat. Hist. Soc. 1896, p. 19, pl. T. fig. 47, g. 


“Male. Upperside : both wings deep black. Fore wing with the following transparent 
white dots :—Two placed outwardly obliquely towards the outer end of the discoidal cell, 
the lower one sometimes missing ; a costal one placed between the costal nervure and 
first subcostal nervule; one in the second median interspace; and five subapical forming 
a perfect S-shaped figure. Cilia fuscous. Hind wing with the anal angle broadly, 
as far as the second median nervule pure white, bearing a large round black spot on 
the margin in the submedian interspace, and a small one in the first median interspace ; 
a fine black anteciliary line in the white area. Cilia anteriorly fuscous, posteriorly 
and along the abdominal margin pure white; very long at the anal angle. Underside: 
both wings dull black. Fore wing with the dots as on the upperside, a pale suffused 
twinned spot towards the outer angle in the submedian interspace. Hind wing almost 
entirely white, the costa and apex broadly, a narrow anteciliary line alone being of the 
dull black ground-colour; the posterior edge of the blackish area bearing four rounded 
deep black spots; the two black spots on the outer margin as on the upperside. 
Thorax and abdomen above black, but the latter tipped with white. Palpi beneath, 
thorax, legs, and abdomen beneath pure white.” 


A REVISION OF THE ORIENTAL HESPERIID 2. 145 


Expanse 34-35 mm. 

Hab. Battak Mountains, N.E. Sumatra (fide de Nicéville). 

After our description and figure of 7. nana were printed we received Mr. de Nicéville’s 
description and figure of his 7. toba, which, judging from the figure, appears to be the 
same as I’. nana, although the spots on the termen of the hind wing, which he describes 
as lying in the submedian and first median interspaces respectively, are represented as 
originating on the apices of the veins, and the figure of the hind wing below shows 
three simple veins between the median vein and the dorsum. 


!'TAGIADES Nestus. (Plate XX. fig. 12, ¢.) 
Pterygospidea nestus, Felder, Sitz. Ak. Wiss., math.-nat. Cl. xl. p. 461 (1860). 
Hab. Amboina (Felder); Batchian (fide Staudinger). 


! TAGIADES DEALBATA. 
Tagiades dealbata, Distant, Rhop. Mal. p. 388, pl. xxxv. fig. 25 (1886). 
Hab. Upper Assam, Perak, Pulo Laut (Doherty) ; Burmah (Adamson). 


! TAGIADES PRINCEPS. 
Tagiades princeps, Staudinger, in litt.; Semper, Schmett. Philipp. p. 307, pl. xlix. fig. 5, ¢ (1892), 
Hab, Mindanao (Semper). 


! TAGIADES PRALAYA. 
Pterygospidea pralaya, Moore, P, Z, 8, 1865, p, 779. 


In this species and the next the intermediate tibie of the males are penicillate. 


Hab. Sikkim (Moller); Khasias (Hamilton). 


! TAGIADES TRICHONEURA. 


Pterygospidea trichoneura, Felder, Wien, ent. Mon, yol. iv. p, 402 (1860); id. Reise Noy., Lep, 
vol. iii. pl. Ixxiii. figs, 14, 15 (1867). 
Tagiades trichoneura, var., Distant, Rhop. Mal. p. 389, pl. xxxiv. fig. 20 (1886). 
Hab. B. Pegu, Perak, Arjuno, Java (Doherty). The specimens from the last-named 
locality have the hind wing below of a deeper yellow. 


! Var. TRICHONEUROIDES, Ster. MS. 


Only differs in the particulars set forth in the table above, 
Hab. Kina Balu, Borneo ( Waterstradt). 


! TAGIADES PINWILLI. 


Plesioneura pinwilli, Butler, Trans. Linn. Soc. Lond., Zoology, ser. 2, vol. 1. p. 556, pl. lxviii. fig. 4 
(1877) ; Distant, Rhop. Mal. p. 400, pl. xxxv. fig. 29, g, 
Celenorrhinus pinwilli, Watson, Hesp. Ind. p, 144 (1891), 


146 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Hab. Margherita, Assam, E. Pegu (Doherty) ; Malacca (Pinwill); ‘Tameang Lajang, 
S.E. Borneo (in coll. Rothschild). 


TAGIADES TABRICA. 
Pterygospidea tabrica, Hewitson, Exot. Butt. v. pl. lix. fig. 8 (1873). 
Lelenorrhinus tabrica, Watson, Hesp. Ind. p. 143 (1891). 

Hab. Darjeeling (Roberts, fide Hewitson). 

Hewitson’s type in the British Museum isa male, and it is very remarkable that if 
it really came from Darjeeling no other example has since been obtained. 


TAPENA. 
Tapena, Moore, Lep. Cey. i. p. 181 (1881); Watson, P. Z. 8. 1893, p. 60. Type thwaitesi, Moore. 


A genus of few species closely allied to the next, but distinguished by the single 
angulation of the termen of the hind wing, which occurs at the apex of vein 3. 
The hyaline spots in the fore wing of the male in all the species known to us are 
present only in cells 6, 7, and 8, but there is sometimes an indication of two others 
in cells 10 and 11; im the hind wing there is sometimes one in the cell near the 
transverse vein. 

In the males the clasp is wider than usual, and the upper lobe is developed into one 
or more processes which, when in situ, lie upon the back of the tegumen and evidently 
effect the depression of that organ in proportion as the inner faces of the clasps 
approach each other. ‘This structure is also found in-the males of Ctenoptilum. 


The following is a table of the species at present known to us :— 


Males. 


1 (2). Upper lobe of clasp bifid at the apex oe Slee ese le eultey ee me wwartest. Moores 
2 (1). Upper lobe of clasp not bifid at the apex. 
3 (4). Upper lobe of clasp with three limbs, all of which are serrate. Similar 

to thwaitesi, but smaller and paler above . . .- - minuscula, n. sp. 
4 (3). Upper lobe of clasp with two limbs, of which the 168 er is serrate a 

the upper simple. Size of ¢hwaitesi, but upperside in the male 

nearly uniform dark umber-brown . . - - + + + + + + + hampsoni, 1. sp. 


''Tapena THWAITESI. (Plate XXII. fig. 15.) 
? Tapena thwaitesi, Moore, Lep. Cey. i. p. 181, pl. Ixvii. figs. 2, 2a, § (1881) ; Watson, Hesp. Ind. 
p. 122 (1891). 
Hab. Perak (Doherty); Selesseh, Sumatra (in coll. Rothschild); Kina Balu, Borneo 
(Waterstradt); Chindwin District, Burmah (Adamson). 
As we have not been able to examine the clasp-form of a Ceylon specimen of 
T. thwaitesi, it is possible that we have not correctly identified it. 


A REVISION OF THE ORIENTAL HESPERIIDA. 147 


!TAPENA MINUSCULA, n. sp. (Plate XVIII. fig. 18,¢; Plate XXII. fig. 17.) 
Tapena thwaitesi, Elwes, P. Z.S, 1892, p. 659, in part. 


Similar to 7. thwaitesi, but smaller and paler. This species is best characterized by 
the peculiar clasp-form, which is figured. 

Two males from Bernardmyo (Doherty) in Elwes’s collection are the only ones we 
have seen of this species. 


!TAPENA HAMPSONI, n. sp. (Plate XVIII. fig. 19,6; Plate XXII. fig. 16.) 


This species also is best characterized by the clasp-form of the male. It resembles 
T. thwaitest in size, but the upperside in the male is darker and consequently the dark 
markings are less evident than in that species. 

Female. Upperside grey-brown ; fore wing with hyaline white spots as follows :— 
An L-shaped one in the cell, an irregular one before the middle of cell 2, a punctiform 
one in the base of cell 3, one in each of cells 6, 7, and 8, one in cell 11, and the indication 
of one (on the underside) in the base of cell 10; hind wing with a subquadrate hyaline 
white spot next the upper outer angle of the cell. Fringes brown in both sexes. 

The hyaline spot in the cell of the hind wing is of no importance as a differential 
character, as it is present in some specimens and absent from others, both of this species 
and 7. minuscula. 

Hab. Nilgiris (Hampson), recorded as T. thwaitesi; N. Canara (Aitken). 


CTreNOPTILUM. 
Ctenoptilum, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 220; Watson, P. Z. S. 1893, p. 60. 
Type vasava, Moore. 

Club of antenne finely pointed: Hind wing biangulated, produced into a tooth at 
the apex of veins 4 and 7, bearing on the basal half a cluster of irregular hyaline 
spots. Male with a tibial pencil of the ordinary form. Watson says of this genus: 
“Vein 8 (of fore wing) usually given out beyond the end of cell—that is, veins 7 
and 8 anastomose for a portion of their basal length (this is not an invariable character ; 
the length of the anastomosis varies in different specimens, and occasionally, though 
very rarely, veins 7 and 8 are free for their entire length).” In ten specimens in coll. 
Elwes from Sikkim, East Pegu, Akyab, and Tenasserim respectively, and in eight 
specimens in coll. Leech from Kiukiang and Ningpo, vein 8 is quite free from vein 7 ; 
but in two specimens in coll. Elwes from Kiukiang, Central China, veins 7 and 8 
anastomose through their basal half. 

The species occurring in our region may be separated as follows :— 

White spot in cell 7 of fore wing reaching beyond that in cell 6. 
Expanse about 34 mm. Fore wing above without any short sub- 
terminal dark band near the apex of the costa. Lower lobe of 
clasp broadly triangular . . . . «. «. «+ + + « « + « vasava, Moore, 


148 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Expanse about 88 mm. Fore wing above with a straight dark band 
from the costa just before the apex as far as vein 4. Lower lobe 
of clasp long and narrow, rounded at the apex . . . . . . . chinensis, n. sp. 
White spot in cell 7 of fore wing not reaching beyond that in cell6 . . multiguttata, de Nicév. 


! COTENOPTILUM VASAVA. (Plate XXII. figs. 18, 18 a.) 

Achlyodes vasava, Moore, P. Z. 8S, 1865, p. 786. 

Ctenoptilum vasava, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 221. 
Hab. Sikkim (Mller); Khasias (Hamilton) ; E. Pegu (Doherty). 


! CTENOPTILUM CHINENSIS, n. sp. (Plate XXIII. figs. 19, 19 a.) 
Ctenoptilum vasava, Leech, Butt.-China &e. p. 575, pl. xli. fig. 18, g (1893-94). 

Hab. Ningpo, April (Leech); Kiukiang, May (Pratt). 

Differs from C. vasava, Moore, in its larger size and in having a dark subterminal band 
on the fere wing above from the costa just before the apex as far as vein 4, and also in 
the clasp-form, as may be seen from the figures. 


! CTENOPTILUM MULTIGUTTATA. 
Ctenoptilum multiguttata, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 221, pl. E. fig. 10, g. 
Hab. Akyab (Adamson); 'Tenasserim (Bingham). 


ODONTOPTILUM. 
Odontoptilum, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 217; Watson, P. Z. 8S. 1893, 
p- 61. Type sura, Moore. 

Tip of antenne blunt. Male with the fore cox heavily scaled and having on the 
inner side a thick tuft of long straight hairs, somewhat exceeding the joint in length 
and springing, for the most part, from the base of the joint. In the male genitalia 
both the clasps and the tegumen are unsymmetrical, the excess of development in 
both cases being on the right side. The angulation of the hind wing presents three 
forms: one represented by swra, Moore, and helias, Feld., a second represented by 
pygela, Hew., and a third represented by leptogramma, Hew. 

The species known to us we separate as follows :— 


1 (4). Termen of hind wing once angulated, at vein 7. 
2 (3). Male: long hairs clothing the apex of the scapule of varying lengths 

and somewhat spreading, not collected intoa pencil. . . . « sura, Moore. 
3 (2). Male: long hairs clothing the apex of the scapule subequal in length 


and collected into a distinct pencil lying on each side of the thorax . helias, Feld. 
4.(1). Termen of hind wing more than once angulated. 


5 (6). Termen of hind wing twice angulated, at veins 4 and 7 - pygela, Hew. 
6 (5). Termen of hind wing thrice angulated, at veins 3,6,and7 . . . . Jeptogramma, Hew. 


A REVISION OF THE ORIENTAL HESPERIIDA. 149 


! ODONTOPTILUM SURA. 
Achlyodes sura, Moore, P. Z.S. 1865, p. 786. 
Abaratha sura, Distant, Rhop. Mal. p. 390, pl. xxxiv. fig. 16, g (1886). 
Odontoptilum sura, Watson, Hesp. Ind. p. 105 (1891). 
? Pterygospidea angulata, Felder, Verh. zool.-bot. Gesellsch. Wien, vol. xii. p. 488 (1862) ; id. Reise 
Noy., Lep. iii. p. 529, pl. xxiii. figs. 10, 11, § (1867). 

We use the name of swra in preference to that of angulata, Feld., adopted by 
Watson, which has three years’ priority, because we cannot identify Felder’s species 
with any certainty. He gives the habitat of angulata as China and Luzon, and his 
figure represents a species with a strongly angled band on the fore wing above, such as 
we have never seen in swra. 

Hab. Mandi (Young); Sikkim (Méller); Nagas, Burmah, Bali, Sambawa (Doherty) ; 
Java (Fruhstorfer). 


! ODONTOPTILUM HELIAS. 

Pterygospidea helias, Felder, Reise Noy., Lep. iii. p. 529, pl. Ixxiii. figs. 12, 13 (1867). 
Odontoptilum helias, var. helisa, Semper, Schmett. Philipp. p. 311 (1892). 

Hab. Celebes (Felder) ; Luzon, Samar, Bohol, E. Mindanao (Semper). 

A male and female of the variety helisa, Stgr. in litt., from Luzon differ from 
typical helias in their smaller size and in having the pale preapical fourth of cell la 
on the fore wing below and a spot near the base of that cell clothed with bluish-white 
scales. The hair-pencil on the scapule of the male is brown, very pale in the basal 
half and dark in its apical half, whereas in helias the entire pencil is blackish brown. 


ODONTOPTILUM HYPERIDES. 

Abaratha hyperides, Doherty, Jour. As. Soc. Beng. 1891, p. 195. 

“Very near Pterygospidea helias, Feld., from the Celebes, but the bands and the 
discal hyaline spots of the fore wing are absent, and the apex of the hind wing is 
broadly dark. From A. swra it differs in the fore wing, which is almost uniform dark 
brown above and below. The hind wing is also less variegated above and below, 
the white area is larger and more uniform, the inner line of spots is obsolete, the outer 
united, and dark apically, nearly obliterated by white scales in the median spaces. 

*“ Hab. Sambawa. 

** Another species, more like A. angulatus, was found in Sumba, but no specimens 
have survived.” (Doherty, l. c.) 

We have not seen this species and do not know where the type is. 


! ODONTOPTILUM PYGELA. 
Pterygospidea pygela, Hewitson, Desc. Hesp. p. 53 (1868) ; id. Exot. Butt. vol. v., Péerygospidea, 
pl. i. fig. 3 (1873). 
Abaratha pygela, Distant, Rhop. Mal. p. 390, pl. xxxiv. fig. 18 (1886) 
Hab. Perak, BE. Pegu (Doherty); Palawan (Platen). 
VOL. XIv.— Part iv. No. 7.—October, 1897. x 


150 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! ODONTOPTILUM LEPTOGRAMMA. 
Pterygospidea leptogramma, Hewitson, Descr. Hesp. p. 53 (1868) ; id. Exot. Butt. vol. v., Ptery- 
gospidea, pl. i. fig. 4 (1873). 
Odontoptilum leptogramma, Semper, Schmett. Philipp. p. 311 (1892). 


Hab. Bohol, Camiguin de Mindanao, Panaon, Mindanao (Semper). 


CAPRONA. 
Caprona, Wallengren, Rhop. Caffr. p. 51 (1857); Watson, P. Z. 8. 1893, p. 62. Type pillaana, 
Wallgr. 
Abaratha, Moore, Lep. Cey. i. p. 181 (1881). Type ransonnettii, Feld. 

This genus is given by Watson as closely allied to the last; and, apparently, 
separated for the most part on account of the secondary male character, which he 
describes as a radiating tuft of hairs attached to the fore coxe. This radiating tuft 
of hairs we have not been able to see, but in ransonnettii, Feld., and syrichthus, Feld., 
the fore cox in the male are heavily scaled, as in Odontoptilum, and bear on the 
inner side a tuft of long straight hairs, which only differs from that found in Odonto- 
ptilum in that the hairs are black and more than twice as long as the coxa. The 
genus is further distinguished from its allies by the possession of two small hyaline 
spots near the middle of cell 1 in the fore wing in addition to the usual discal and 
subapical series, and the termen of the hind wing is more or less produced at the apex 
of veins 2, 3, and 6, besides the main angulations at veins 4 and 7. 

The diagnostic characters of the species here dealt with are expressed in the following 
table :— 


No terminal row of pale spots on the fore wing above. 
No hyaline spot in the cell of the fore wing near the middle. Right clasp 
elongate, bifid at the apex; left one simple . . . . . . « ransonnettii, Feld. 
With a hyaline spot in the cell of the fore wing near the miidles 
Hind wing below with a little white scaling near the base, and three series 
of short dark brown transverse lines—two in the basal series, two in 
the median, and five in the postmedian, Right clasp simple, left 


one dissimilar, suddenly narrowed at the apex . . . . . . . saraya, Dohy. 
Underside “frosted with white throughout, except very narrowly along 
the outer margins of both wings” . ..... =... - . alida, de Nicév. 
A complete terminal row of pale spots on the fore wing above . . . . . syrichthus, Feld. 


! CAPRONA RANSONNETTU. (Plate XXIII. figs. 20, 20a.) 


Pterygospidea ransonnettii, Felder, Verh. zool.-bot. Gesellsch. Wien, 1868, p. 284. 
Pterygospidea potiphera, Hewitson, Exot. Butt. vol. v., Pterygospidea, pl. 1. fig. 7 (1873). 
Abaratha ransonnettii, Moore, Lep. Cey. vol. i. p. 182, pl. xevii. fig. 1 (1881). 

Abaratha taylorii, de Nicéyille, Jour. As. Soc. Beng. 1883, p. 88, pl. x. fig. 13, g. 


A REVISION OF THE ORIENTAL HESPERIIDA, 151 


We have no hesitation in treating faylorii as a synonym, as we have Nilgiri specimens 
which agree with it precisely. 

Hab. Ceylon ; Nilgiri hills (Hampson); N. Canara (Aitken); Travancore, Orissa 
(fide de Nicéville) ; Khasia hills (Hamilton). 


! CAPRONA SARAYA. (Plate XVIII. fig. 20, ¢; Plate XXIII. figs. 21, 21 a.) 
? Abaratha saraya, Doherty, Jour. As. Soc. Beng. 1886, p. 138. 


The C. saraya of this paper is the same as that of the British Museum collection, 
where there are three specimens thus named, which had been originally labelled on 
the pin faylorii, de Nicév. These specimens all come from “ Futeh Khan’s Bungalow, 
Kootur, Chittarpahar: probably 3000 feet.” 

It is, however, doubtful if they are correctly identified ; because, according to the 
original description, the hind wing below in @. saraya has “a black transverse streak 
at the end of the cell, a fainter one near the base of the cell, and a circle of large and 
conspicuous black spots, nine in all, round the disc, whereof two are between the costal 
and the subcostal, and two between the median and submedian veins.” C. saraya was 
described from a single male. 

Hab. Kumaon, 5500 feet (Doherty). 


! CAPRONA ALIDA. 

Abaratha alida, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 394, pl. G. fig. 40, ¢. 

Fore wing with a small round white transparent spot about the middle of the 
discoidal cell. ‘ Underside: both wings thickly overlaid with large pure white scales, 
so that the dark ground-colour is entirely hidden except narrowly along the outer 
margins, the whole wing surface having the appearance of being thickly strewn with 
hoar-frost.”” 

Very similar in appearance to C. ransonnettii, but the latter wants the hyaline spot 
near the middle of the cell of the fore wing. 

Hab. Tilin Yaw, Upper Burmah (Watson); Upper Chindwin River (Adamson). 


! CAPRONA SYRICHTHUS. 
Pterygospidea syrichthus, Felder, Reise Nov., Lep. vol. iii. p. 530, pl. lxxii. figs. 22, 23 (1867). 
Abaratha syrichthus, Elwes, P. Z. S. 1892, p. 656, pl. xiii. fig. 2, var. ? 
Pyrgus agama, Moore, Cat. Lep. Mus. E.I. C. vol. i. p. 249, pl. vii. figs. 1 (larva), 1 @ (pupa) 

(1857). 

Caprona elwesi, Wats. in litt. 

The specimens from Bernardmyo, which Elwes figured as a variety of C. syrichthus, 
are remarkable for their small size; the smallest expands only 28 mm. As, however, 
there is no difference in the genitalia, we consider that the name of C. e/wesi given by 


Watson to this form cannot stand. 
x2 


152 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Hab. Bhamo, Burmah (Adamson) ; Shan hills (Manders); Burmah, Bali (Doherty) ; 
West Java. 


CARCHARODUS. 
Carcharodus, Hiibner, Verz. p. 110 (1816) ; Watson, P. Z. 8. 1893, p. 67. Type lavatere, Esp. 
Spilothyrus, Dup. Pap. Fr. Diurn., Suppl. p. 415 (1832). Type alcee, Esp. 


A genus well distinguished by its facies. Fore wing with a dark basal shade and 
hyaline spots. Termen of the hind wing crenulate. 
The following table includes all the known species :— 


Fore wing below with no tuft of hair in the male. 


Fore wing with the hyaline spot in cell 2 wider thanhigh . . . . . . Javatere, Esp. 
Fore wing with the hyaline spot in cell 2 higher than wide . . . . . . alcee, Esp. 
Fore wing below with a tuft of hairinthemale. . . . . Se eal ede dike Eabr. 


! CARCHARODUS LAVATERZ. 


Papilio lavatere, Esper, Schmett. i. 2, pl. Ixxxii. fig. 4 (1783); Hiibner, Eur. Schmett. i, 
figs. 454, 455 (1798-1803). 


Hab. Central Europe to Asia Minor. 


! CARCHARODUS ALCE. 


Papilio alcee, Esper, Schmett. i. 2, pl. li. fig. 8 (1780). 

Pyrgus dravira, Moore, P. Z. 8S. 1874, p. 576, pl. Ixvii. fig. 5; de Nicéville, Jour. As. Soc. 
Beng. 1883, p. 88, pl. x. fig. 5, 9. 

Carcharodus swinhoei, Watson, P. Z. 8. 1893, p. 68. 

There is no difference in the genitalia of the Western and Eastern forms, and we see 
no reason for separating them. It is true that the desert form swinhoei, Watson, 
found in Beluchistan, is much paler than the Himalayan form dravira, Moore’s 
figure of which is very bad, but we find similar variations in colour almost as striking 
amongst European specimens. 

Hab. Central and Southern Europe to Central Asia and N.W. Himalayas up to 
9000 feet. 


CARCHARODUS ALTHE. 
Papilio alihee, Hibner, Eur, Schmett. i. figs. 452, 453 (1798-1803). 
This species also varies from dark to light grey-brown. 


The eggs, larva, and pupa are described by Heinrich Gross (Stett. ent. Zeit. 1894, 
p. 77); the eggs were laid and the larva fed on Betoniea officinalis. 
Hab. Central and Southern Europe to Central Asia. 


(oN) 


A REVISION OF THE ORIENTAL HESPERIID@. 15 


GoMALIA. 
Gomalia, Moore, P. Z. S. 1879, p. 114; Watson, P. Z. S. 1893, p. 67. Type albofasciata, Moore. 
A genus of one or two small species differing from Hesperia in facies and the slender 
and straight club of the antenne. Male with a costal fold but no tibial hair- 
pencil. 


! GOMALIA ALBOFASCIATA. 

Gomalia albofasciata, Moore, P. Z. 8. 1879, p. 144; id. Lep. Cey. vol. i. p. 183, pl. xxi. fig. 7 
(1881). 

? Gomalia littoralis, Swinhoe, P. Z. 8. 1884, p. 513, pl. xlvii. fig. 4. 

Hab. 8. India (Hampson) ; Hambantota, Ceylon (Green); Karachi (fide Swinhoe) ; 
Quetta, Beluchistan (Zurray in coll. Swinhoe). ; 

We have not seen the type of /itioralis, but, judging from the description and figure, 
it is indistinguishable from a/bofasciata, like specimens from Quetta which we have 
examined. 


HESPERIA. 


Hesperia, Fab. Ent. Syst. iii. vol. i. p. 258 (1793); Watson, P. Z. S. 1893, p. 64. Type 
malve, Linn. 

Pyrgus, Hiibn. Verz. p. 109 (1816). Type syrichtus, Fab. 

Syrichthus, Boisd. Ieones, p. 230 (1832-83). Type proto, Esp 

Scelothriz, Rambur, Cat. Lép. And. 1. p. 63 (1858). Type carthami, Hiibn. 


A numerous group of closely allied species, which are associated together on account 
of their resemblance to the Papilio malvw of Linnzeus. According to Watson, it is 
separable from Carcharodus, Hiibn., by the even termen of the hind wing, from 
Gomalia, Moore, by the curved club to the antenne, and from Thanaos, Bdy., which 
has a comparatively slender antennal club, by the robust club of its antenne. The 
following pale spots on a darker ground constitute what may be considered as 
the normal pattern of the hind wing below, namely :—one occupying the whole or part 
only of cell 1a; three in cell 7, basal, medial, and apical; two in cell 6, basal and 
medial ; one near the base of the cell; a large geminate one next the transverse vein 
and occupying the base of cells 4 and 5, and another geminate one near the apex of the 
game cells; one near the apex of cell3; two in cell 2, basal and apical; and three, 
basal, medial, and apical, in cell 14. These pale spots fall into three irregular trans- 
verse series. ‘The modifications of this wing-pattern are for the most part trifling, but 
they sometimes afford useful distinguishing characters. 

The following table will serve to show what characters have been found most useful 
in the arrangement of the species of this difficult group in coll. Elwes; this arrange- 
ment is doubtless far from perfect, but it certainly gives a better result than any other 


154 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


with which we are acquainted. The species of the orbifer group are exceedingly 
closely allied; the male genitalia and the essential wing-pattern are practically 
identical, but the characters given below will separate the majority of specimens. 
Therapne, Rbr., put by Staudinger as a variety of sao, is just as distinct from the latter 


as are orbifer and ali. 


1 (66). Hind wing below dark, with a pale median band. 
2 (57). Such band macular, generally much broken. 
3 (32). Fore wing with a subterminal series of small pale spots. 
4 (5). Fore wing below with a distinct white spot (not merely a pale 
dash) near the base of the cell. Median pale band on hind 
wing below continuous from veins 2-7. . . . c .  galba, Fab. 
5 (4). Fore wing below with the basal pale marking in the cell (ibe 
present) a dash, not a distinct spot. 
6 (21). Hind wing below with the pale spot near the middle of cell 7 
continued in cell 8. 
7 (16). Hind wing below: middle spot in cell 7 not joined to that on 
the transverse vein, or at most only touching it in a point. 
8 (15). Hind wing below with the apical third of cell 8 dark. 
9 (10). Hind wing below sordid yellowish green, pale spot on the 
transverse vein with sharp projecting angles . . . . - .  orbifer, Hiibn. 
10 (9). Hind wing below tile-red (more or less bright) or pale red-brown. 
11 (12). Hind wing below with the pale spot on the transverse vein not 
produced inwardly ; es strongly re. with sordid 
yellowiie ae  ea : therapne, Rbr. 
12 (11). The pale spot on the transverse vein of hind wing ‘below ideedl 
both outwardly and inwardly. 
13 (14). Such spot and that in cell 14 but little intensified ; hind wing 


below tile-red .. . : : sao, Bergstr. 
14 (13). Such spots much snteeafiedd hee snes ; hind” wing below 

pale red-brown. . Joao a PTO: 
15 (8). Hind wing below with the sia shied of all 8 als Soe the CERCA NEES 


16 (7). Hind wing below with the middle spot in cell 7 broadly con- 
nected with that on the transverse vein by a triangular spot 
in the base of cell 6. 
7 (18). Hind wing above with no pale spot in the middle of cell 7. No 
costal foldin the male. . . phlomidis, H.-S. 
18 (17). Hind wing above with a pale erat in the “middle of far 7. 
Male with a costal fold. 
9 (20). No tibial hair-pencil in the male, but several short spines on 
the outer side of hind tibie . .... =.=. =~. =. ==: ~~. £eribrellum, Hvers. 
( (19). Male with a tibial hair-pencil” . . . . . - . + + + + cynare, Rbr. 
1 Fide Watson. There is no tale of this species m coll, Elwes, and only one female. The latter can only 
be separated from cribrellwm by its name-label. 


21 (6). 
22 (27). 
23 (26). 
24 (25). 
25 (24). 
26 (23). 
27 (22). 
28 (29). 
29 (28). 
30 (31). 


31 (30). 


32 (3). 
33 (36). 


34. (35). 
35 (34). 
36 (33). 


7 (50). 


8 (49). 
39 (40). 


A REVISION OF THE ORIENTAL HESPERIIDA. 155 


Hind wing below with the pale spot in the middle of cell 7 not 
continued in cell 8. 

Hind wing below with the outermost pale spot in cell 7 adjoining 
or near the termen. 

Hind wing below with the pale median band conspicuously 
broken inwardly at cell 7 

Species smaller (exp. 30-38 mm.) and paler. Pale spots on 
upperside relatively larger Sto Pets MBs get 

Species larger (exp. 40-45 mm.) and darker. Pale spots on 
upperside relatively smaller su ego : 

Hind wing below with the pale median band continuous and 
equally wide in cells 3 to 7 inclusive : ‘ 

Hind wing below with the outermost pale spot in cell 7 remote 
from the termen. 

Costal fold Uae apparently absent; no tibial hair- 
pencil 3 aed & 

Costal fold conspicuous ; no tibial hair- neil 

Hind wing below: inner edge of the outermost spot in cell 7 
in line with the transverse vein. Size generally smaller. 
Subapical notch of clasp with an erect tooth 

Hind wing below: inner edge of the outermost spot in cell 7 
not in line with the transverse vein. Size larger. Subapical 
notch of clasp without a tooth. 

No subterminal series of small pale spots on foes wing. 

With a tibial hair-pencil but no costal fold in the male. Hind 
wing below with the pale macular band ending in a spot at 
the apex of cell 1 0. 

Hind wing above usually without pale markings 

Hind wing above with conspicuous pale markings . c 

Tibial hair-pencil and costal fold both present in the male 
Hind wing below with the macular band ending in a spot in 
the middle of cell 1 0. 

Hind wing below with the spot on the transverse vein straight- 
sided within.? 

Hind wing below with the markings sharply defined. 

Hind wing below with an irregular continuous pale terminal 
band* . 


tessellum, Hiibn. 
gigas, Brem. 


nobilis, Stgr. 


poggei, Led. 


proto, Esper. 


staudingeri, Speyer. 


‘cashmirensis, Moore. 
‘alpina, Esch. 


carthami, Hubn. 


1 These two species are exceedingly closely allied. 


race of cashmirensis. 


2 This character is not absolute. 


It seems doubtful whether alpina is not a geographical 


Exceptions occur in H, malve and some others; but malve is a well- 


known species, and the other exceptions occur so sparingly that the practical utility of the character is not 


affected. 


3 This character is not absolutely diagnostic of carthami, but in practice it is found that its occurrence in 


other species is rare and obviously abnormal. 


156 


40 (39). 
4] (48). 
42 (47). 
3 (46). 
44 (45). 
45 (44). 


46 (43). 
47 (42). 


48 (41). 
49 (38). 


50 (37). 
1 (54). 


52 (53). 


53 (52). 
54. (51). 


8 (61). 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Hind wing below without an irrregular continuous pale 
terminal band. 

Apex of tegumen undivided. 

Tegumen without lateral horns. 

Apical lobe of clasp much expanded, broadly and evenly rounded 
in a semicircle. 

Fore wing below greenish grey-brown ; hind wing below with a 
pale dot normally present in cell 6 F : 

Fore wing below blackish brown; hind wing palow: stetally 
without a pale dot in cell 6 6 ie. 

Apical lobe of clasp not expanded in a semicircle . i 

Tegumen with a horn on each side, tip compressed. Clasp of 
the same type as zona . 

Apex of tegumen bipartite : a ho oo 

Markings of hind wing below absolescent ty reason of ehietal 
paleness. Tegumen with a branched horn on each side “ 

Hind wing below with the pale spot on the transverse vein 
produced inwardly towards the origin of vein 7.° 

Hind wing below with a pale dash in cell 1 } near the base. 

Fore wing above with a straight oblique pale macular band 
from the costa or the middle to the dorsum before the 
middle Bats 

No such band on ae wing above 2 cme ae 

Hind wing below with the pale basal sioaeleinged in ei 1 b aspot, 
not a dash, 


. Hind wing below pale sordid yellowish- or russet-green 
. Hind wing below dark (usually blackish) green 
. Hind wing below with the pale median band sharply aenaea, 


narrow, non-macular, and preceded by a broad irregular brown 
band, which bears a small pale spot near the basal third of 
cell 7; or if macular, then not interrupted, and preceded by 
a dark Y-shaped marking reaching quite across the wing and 
followed by a dark subterminal band. 

Hind wing below with the pale median band preceded by a 
large dark Y-shaped marking reaching quite across the wing 
and followed by a dark subterminal band. 


9 (60). No dark terminal band on hind wing below 


* We cannot distinguish with certainty between serratule and alveus, except by the male genitalia, 


‘serratule, Rbr. 


*speyeri, Stgr. 
*alveus, Hiibn. 


malvoides, 0. sp. 
malve, Linn. 


melotis, Dup. 


andromede, W\grn. 


*cacalie, Rbr. 


onopordi, Rbr. 
centauree, Rbr. 


bieti, Ob. 


If 


we make the presence or absence of pale markings on the hind wing aboye the basis of separation, we find 
that this character varies too much to be reliable ; if, on the other hand, we use the male genitalia, we get two 


forms which do not intergrade. 


types, although both kinds are known to occur in the same locality, as at Vernet and Mont Cenis, 
* Very near the preceding, and probably not distinct. 
* Vide note to paragraph 37. 
* This insect has the same type of clasp form as andromeda. 


The male genitalia can always be referred with certainty to their respective 


A REVISION OF THE ORIENTAL HESPERIIDA, 157 


60 (59). Hind wing below with a dark terminal band separated from 
the dark subterminal band by a zigzag white line or row of 
irregular spots . . . oberthiri, Leech. 
1 (58). Hind wing below with the male median Gana nieeared ae a 
broad irregular brown band, which bears a small pale spot 
near the basal third of cell 7. 
62 (63). Hind wing above with a median row of pale spots. Fore wing 
below with the apex yellow-brown; pale spot near the apex 
of cell 5 oblique, distinct from that in cell 4. No sub- 
terminal pale band on hind wing below . . . . . . . zona, Mab. 
63 (62). Hind wing above with two rows of pale spots. 
64 (65). Fore wing below with the apex yellow-brown; the pale spot 
near the apex of cell 5 oblique, distinct from that in cell 4 . maculatus, Brem. & 
65 (64). Fore wing below with the apex white, with a short oblique [Grey. 
yellow-brown dash ; pale spots near the apex of cells 4 and 5 
coalescent, their inner edges erect . . . . . 6 thibetanus, Ob. 
66 (1). Hind wing below white, with two irregular or page-velloy anda 
narrowly margined with black. 
67 (68). Fore wing above with the pale spot in the cell higher than wide, 
generally constricted in the middle. Male with costal fold 
and tibial hair-pencil . . . . side, Esper. 
68 (67). Fore wing above with the pale spot in Ale cell vier his high, 
subrhomboidal. Neither costal fold nor tibial hair-pencil in 
WSR 5 9 oc 9 oo 8 op 6 Go of 816 o 6 5 og Witednin Shae, 


In the matter of synonymy we have restricted the references to those which appear 
to be necessary for the identification of the species under consideration. 


! HESPERIA GALBA. 
Hesperia galba, Fabricius, Ent. Syst. vol. iii. p. 852 (1793) ; Moore, Lep. Cey. vol. i. p. 183, pl. Ixxi. 
fig. 6 (1881). 
Pyrgus superna, Moore, P. Z. S. 1865, p. 792. 
Pyrgus evanidus, Butler, Ann, & Mag. Nat. Hist. ser. 5, vol. v. p. 223 (1880). 
Hesperia evanidus, Watson, Hesp. Ind. p. 156 (1891). 
Pyrgus zebra, Butler, op. cit. ser. 6, vol. i. p. 207 (1888). 
‘Hesperia zebra, Watson, Hesp. Ind, p. 156 (1891). 
Hesperia hellas, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 177, pl. B. fig. 9, g. 


I cannot see any good grounds for separating P. zebra=hellas (fide Watson) from 
galba. Though I have no Campbellpur specimens for comparison, yet the characters 
relied on by de Nicéville seem very variable. The species has a very wide range 
throughout the drier parts of India and extends from Burmah (Manders) to Aden, being 
apparently common in the foot-hills of the N.W. Himalaya. 


VOL. XIV.— Part Iv. No. 8.— October, 1897. Y 


158 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! HESPERIA ORBIFER. 
Papilio orbifer, Hiibner, Eur. Schmett. i. figs. 803-806 (1818-27). 
Hab. S. Europe to Turkestan and Amurland. 


! HESPERIA THERAPNE. 
Hesperia therapne, Rambur, Aun. Soc. Ent. Fr. 1882, pl. vii. fig. 4. 


Hab. Corsica, Sardinia. 


! HESPERIA SAO. 


Papilio sao, Bergstrasser, Nomencl. vol. ii. pl. xl. figs. 8, 9 (1779) ; Hitbner, Eur. Schmett. vol. i. 
figs. 471,472 (1798-1808). 


Hab. Mont Cenis, Valais, 2000 ft., Kreusnach, Briancon (Elwes); 8. Spain (Elwes). 


! HESPERIA ALI. 
Syrichthus ali, Oberthir, Etudes d’Ent. vi. p. 61, pl. ii. fig. 8. 
Hab. Batua, Constantine, Algeria (Elwes). 


! Hesperia GERON. (Plate XXIII. fig. 31.) 
Hesperia geron, Watson, P. Z. 8. 1893, p. 66. 


Though we have only one male of this species from Quetta, we think it is a distinct 
species. , 
Hab. Beluchistan (Watson); Shahrud, Persia (Zeller, fide Watson). 


! HESPERIA PHLOMIDIS. (Plate XXIII. fig. 30.) 
Hesperia phlomidis, Herrich-Schiaffer, Schmett. Eur, vol. i., Hesp. figs. 8, 9 (1845). 


Hab. Asia Minor, Greece, Caucasus. 


! HuSPERIA CRIBRELLUM. 


Hesperia cribrellum, Eversman, Bull. Mosc, 1841, p. 25; Freyer, Neuere Beitr. vol. iv. pl. eccxlix. 
fig. 1 (1840?) ; Herrich-Schaffer, Schmett. Eur. vol. i., Hesp. figs. 12, 13 (1845). 


Hab. S. Russia, West Asia, Turkestan, Upper Amur (fide Graeser). . 


! HESPERIA CYNARA. 


Hesperia cynare, Rambur, Faun. And. pl. viii. figs. 4,5 (1839); Herrich-Schiffer, Schmett. Eur, 
vol. i., Hesp. figs. 4-7 (1845). 


Hab. S. Russia to Turkestan. 


A REVISION OF THE ORIENTAL HESPERIIDA. 159 


! HESPERIA TESSELLUM. 


Papilio tessellum, Hiibner, Hur. Schmett. vol. i. figs. 469, 470 (1798-1803). 
Hesperia tessellum, Rambur, Faun. And. pl. viii. figs. 1, 2 (1839). 
Hesperia noma, Lederer, Verh. zool.-bot. Gesellsch. Wien, 1855, p. 1938, pl. i. fig. 7. 


Hab, 8. Russia, Armenia to Central Asia. 


! HESPERIA GIGAS. 
Pyrgus gigas, Bremer, Lep. Ost-Sib. p. 96, pl. viii. fig. 3 (1864). 
Hab. Amurland. 


! FIuSPERIA NOBILIS. 
Pyrgus nobilis, Staudinger. Stett. ent. Zeit. 1886, p. 255. 


Hab. Turkestan. 


! HESPERIA POGGEL. 
Hesperia poggei, Lederer, Wien. ent. Mon. 1858, p. 141. 
Hab. Syria, Armenia to Turkestan. 


! Hesperia proto. (Plate XXIII. fig. 22.) 
Papilio proto, Esper, Schmett. vol. i. p. 2, pl. exxiii. figs. 5, 6 (1806 ?) ; Hiibner, Eur. Schmett. 
vol. i. figs. 918-921 (1827-41). 
Syrichthus mohammed, Oberthiir, Bull. Ent. Soc. Fr. 1887, p. xlviii; id. Etudes d’Ent. xii. p. 23, 
pl. v. figs. 23a g,236 2 (1888). 


Hab. 8. Spain, Portugal to Syria and Armenia; Lambessa, Algeria (Bleuse). 


HESPERIA LEUZEA. 

Syrichthus leuzee, Oberthiir, Etudes d’Ent. vi. p. 60, pl. iii. fig. 10 (1881). 

This species, which was described from a single specimen from Boisduval’s collection, 
appears to be nearest to H. proto. It may be distinguished from that species, however, 
by reason that the pale subterminal band on the hind wing below, which in H. proto is 
‘at the most only represented by a row of small indistinct pale spots, is developed 
into a well-defined band about two-thirds as broad as the median band, 

Hab. Mascara, Algeria (fide Oberthiir). 


! HESPERIA STAUDINGERI. (Plate XXIII. fig. 23.) 


Syrichthus staudingeri, Speyer, Stett. ent. Zeit. 1879, p. 344. 
Pyrgus (Syrichthus) proteus, Staudinger, t. c. 1886, p. 253. 


Hab. Turkestan. 


160 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! HESPERIA CASHMIRENSIS. 


Pyrgus cashmirensis, Moore, P. Z. S. 1874, p. 274, pl. xliii. fig. 7. 
Hab. Kulu, N.W. Himalaya 12,000 ft. (Young), to Kashmir (Leech). 


! HESPERIA ALPINA. 
Hesperia alveus, var. alpina, Erschoft, Lep. Turk. p. 24, pl. ii. fig. 18 (1874). 


Hab. Turkestan (Haberhauer, &c.). 


! HESPERIA CARTHAMI. 
Papilio carthami, Hiibner, Eur. Schmett. vol. i. figs. 720, 723 (1803-18). 
Hesperia carthami, Rambur, Faun, And. pl. vii. fig. 8 (1839). 


Hab. Central and South Europe, Alps and Pyrenees to 5000 ft. (Elwes). 


! HESPERIA SERRATULHZ. (Plate XXIII. fig. 24.) 
Hesperia serratule, Rambur, Faun. And. pl. viii. fig. 9 (1839) ; Herrich-Schaffer, Schmett. Eur. 
vol. i., Hesp. figs. 18-20 (1846). 


Hab. Alps to 6000 feet, Central and South Europe to Caucasus, Amasia (Fuchs). 


! HESPERIA SPEYERI. (Plate XXIII. fig. 26.) 
Scelothrix speyeri, Staudinger, Rom. Mém. sur Lép. iii. p. 153, pl. viii. figs. 5 a, 6. 


Hab. Amurland (Dorries). 


! HesPerta ALVEUS. (Plate XXIII. figs. 25, 25 a.) 
Papilio alveus, Hiibner, Eur. Schmett. i. figs. 461-463 (1798-1803). 


Hab. Alps to 6000 feet, Central Europe to Amurland, Syria (Delagrange). 


! HESPERIA MALYOIDES, n. sp. (Plate XXIII. figs. 27, 27 a.) 


We propose this name for an insect, of which we have three specimens from 
Biarritz, taken by Elwes on July 25, 1887, and a single male from Granada (ibe), 
which has all the facies of H. malve but very distinct male genitalia. The differences 
in this respect will be apparent from the figures. The proportion of black hair-scales 
in the clothing of the second joint of the palpi appears to be considerably greater in 
H. maloe than in H. malvoides. 

We should not have ventured to separate this on genitalic characters alone but for 
the fact that the genitalia in six specimens of H. malvw from Denmark, Kreusnach, 
Rennes, Brittany, Stettin, and Brussa are all absolutely similar inter se, as are the three 
specimens above mentioned. Probably it will be found that H. malvoides has a wider 
range and other distinctive characters. 


A REVISION OF THE ORIENTAL HESPERIID A, 161 


! HESPERIA MALVA. (Plate XXIII. figs. 28, 28 a.) 


Papilio malve, Linneus, Faun. Suec. p. 285. n. 1081 (1761) ; id. Syst. Nat. i. 2, p. 795. n. 267 
(1767) ; Esper, Schmett. i. 1, pl. xxxvi. fig. 5 (1778). 

Papilio taras, Bergstrasser, Nomencl. vol. iv. pl. xci. figs. 5, 6 (1780). 

Hesperia taras, Meigen, Eur. Schmett. i. p. 61, pl. lv. figs. 3a, 6 (1830). 


Hab. Europe to Amurland. 


! HusPeRIA MELOTIS. (Plate XXIII. fig. 29.) 
Hesperia melotis, Duponchel, Lép. Fr., Suppl. i. pl. xlii. figs. 1, 2 (1832). 
Pyrgus hypoleucos, Lederer, Verh. zool.-bot. Gesellsch. Wien, 1855, p. 193, pl. i. fig. 8. 


Hab. Beirut, Syria (Zach). 


! HESPERIA ANDROMEDA. 
Syrichthus andromede, Wallengren, Vet.-Ak. Forh. 1853, p. 25; id. Lep. Scand., Rhop. p. 272 
(1857). 


Hab. Alps, 6000 ft. (Elwes); Norway (Schoyen). 


! HESPERIA CACALLA. 
Herperia cacalie, Rambur, Faun. And. pl. viii. figs. 6, 7 (1839) ; Herrich-Schiffer, Schmett. 
Eur. i., Hesp. figs. 23-25 (1846). 


Hab. Alps, 6000 feet (Hlwes). 


! HESPERIA ONOPORDI. (Plate XXIII. fig. 25 4.) 


Hesperia onopordi, Rambur, Faun. And. pl. viii. fig. 13 (1839). 
Scelothrix onopordi, Rambur, Cat. Lép. And. vol. i. p. 72 (1858). 


Hab. 8. Spain (Staudinger); Batna, Algeria (Ewes). 

This difficult species may be distinguished from H. alveus, of which it has been 
doubtfully treated as a synonym, by the pattern of the hind wing below, which in 
H. onopordi has the white spot in cells 4-5 projecting into the discoidal cell next the 

‘subcostal, whilst in H. alveus the spot in question has no such projection, and also 
by the difference in the form of the male genitalia. 


! HESPERIA CENTAUREA. 
Hesperia centauree, Rambur, Faun. And. pl. viii. fig. 10 (1839) ; Herrich-Schaffer, Schmett. 
Hur. 1., Hesp. figs. 1-3 (1845). 


Hab. Norway, 2000-6000 ft. (Elwes, Schoyen); Finland (Schilde) ; Labrador 
(Moeschler) ; Laggan, Alberta Terr., British N. America, 5000 ft. (Elwes). 


162 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! HESPERIA BIETI. 
Syrichthus bieti, Oberthir, Eitud. d’Ent. xi. p. 26, pl. vi. fig. 50 (1886). 
Hesperia bieti, Leech, Butt. China &e. p. 578 (1894). 

Hab. Western China (fide Leech); N.E. Tibet (fide Alphéraky). 


HiksPERIA OBERTHURI. 
Syrichthus oberthiiri, Leech, Entomologist, xxiv. Suppl. p. 59 (June 1891). 
Syrichthus delavayi, Oberthiir, Htud. d’Ent. xv. p. 20, pl. iii. fig. 31, (July 1891). 
Hesperia oberthiiri, Leech, Butt. China &e. p. 579, pl. xli. fig. 5,3. 
Hab. Western China (fide Leech) ; Yunnan (fide Oberthiir). 
We rely on Mr. Leech’s identification of H. delavayi with H. oberthiivi, not having 
seen the former. 


! HESPERIA ZONA. 
Scelothrix zona, Mabille, Ann. Soc. Ent. Fr. 1875, p. eexiv. 
Scelothriz (Pyrgus) alhistriga, Mabille, t. c. 1876, p. xxvii. 
Pyrgus sinicus, Butler, Ann. & Mag. Nat. Hist. ser. 4, vol. xix. p. 96 (1877). 
Syrichthus sinicus, Pryer, Rhop. Nihon. p. 35, pl. x. fig. 22 (1889). 
Hesperia zona, Leech, Butt. China &c. p. 577, pl. xli. figs. 1, 3 (1893-4). 

Hab. Japan, Shanghai (Pryer) ; Central China (Pratt); Korea (Leech). 

We rely on Mr. Leech’s identification of H. sinicus with H. zona, which is, we 
believe, correct. We do not feel equal confidence in the constancy of the characters 
by which he separates it from H. maculatus, especially as the differences in their 
genitalia are of degree only. 


! HESPERIA MACULATUS. 


Syrichthus maculatus, Bremer & Grey, Schmett. N. China’s, p. 11, pl. iii. fig. 6 (1853) ; Pryer, 
Rhop. Nihon. p. 35, pl. x. fig. 21 (1889). 

Pyrgus maculatus, Ménétriés, Cat. Mus. Petr, pl. v. fig. 5 (1855). 

Hesperia maculata, Leech, Butt. China, &e, p. 576, pl. xli. fig. 2, g (1893-4). 


Hab. Amuiland (Graeser) ; Askold, Shanghai (Pryer). 


! HESPERIA THIBETANUS. 


Syrichthus maculatus, var. thibetanus, Oberthiir, Etud. @’Ent. xv. p. 20, pl. iii. fig. 27 (1891). 
Hesperia thibetana, Leech, Butt. China &e. p. 578 (1894). 


Hab. Western China (fide Leech). 
We follow Mr. Leech in treating this as a distinct species, as our material is insufficient 
to decide the question ; the differences noted are not, however, very striking. 


! HESPERIA SIDA. 
Papilio side, Esper, Schmett. i. 2, pl. xc. fig. 3 (1784); Hiibner, Eur. Schmett. i. fig, 468 
(1798-1803). 
Hab. Alassio, N. Italy (Mrs. Nicholl) ; 8.E. Europe ; Asia Minor to Turkestan. 


A REVISION OF THE ORIENTAL HESPERIIDA, 163 


! HESPERIA ANTONIA. 
Syrichthus antonia, Speyer, Stett. ent. Zeit. 1879, p. 342. 
Hab. Turkestan, Amurland. 


'THANAOS. 
Thanaos, Boisd. Icones, 240 (1832-33) ; Watson, P. Z. S. 1893, p. 69. Type tages, Linn. 


This name is applied generically to certain species more or less resembling the 
Papilio tages of Linnzeus. 
We separate the species in coll. Elwes as follows :— 


Male with a costal fold. 
Hind wing above without yellow spots . ........ . . . tages, Linn. 
Hind wing above with yellow spots. 

Expanse about 85 mm. Fore wing below with the first row of pale spots 
beyond the end of the cell suffused and irregular. Fore wing above 
with the postmedian band broader, at least as broad as the width of 
cell 2. Tegumen with two pairs of lateral dorsal lobes, of which the 
pair nearest the apex are smaller, triangular, and simple, and the 
pair nearest the base are larger, ear-shaped, and rough with short 
AMES og 6 o & 9 6s 6 0 1a yop p Mo woe ON Gmina m, DotD Teton, 

Expanse about 30 mm: Fore wing below with the first row of pale 
spots beyond the end of the cell regular and well-defined. Fore 
wing above with the postmedian band narrower, in cell 2 about 
half as broad as the width of that cell. Tegumen wanting the pair 
of ear-shaped spinose lateral dorsallobes . . . .. =.=. =. . Ueechii,n.s. 

Male without a costal fold. 
Fore wing above with the pale spots in cells 7 and 8 well marked. Basal 


angle of the lower lobe of left clasp not produced intoahorn. . . . marloyi, Bdy. 
Fore wing above with the pale spots in cells 7 and 8 obsolete. Basal angle 
of the lower lobe of left clasp produced into a long curved horn . . . pelias, Leech. 


~ !'THANAOS TAGES. 

Papilio tages, Linneus, Faun. Suec. p. 286. n. 1082 (1761); id. Syst. Nat. 1. 2, p. 795. n. 268 
(1767) ; Esper, Schmett. i. 1, pl. xxiii. fig. 3 (1777) ; Hiibner, Eur. Schmett. 1. figs. 456, 457 
(1798-1803). 

Thanaos cervantes, Gras]. Ann. Soc. Ent. Fr. 1836, p. 558, pl. 17 8. figs. 1, 2. 

Nisoniades tages, var. sinina, Groum-Grshimailo, Hore Soc. Ent. Ross. xxv. p. 461 (1891) ; Leech, 
Butt. China &e. p. 661 (1894). 

? Thanaos popoviana, Nordm. Bull. Mose. 1851, ii. p. 443, pl. xii. figs. 3, 4, 


_ Hab. Europe to Amurland. 

Judging from a specimen from Koko-nor, Tibet, ex coll. Rothschild, the variety 
sinina has the ground-colour paler and greyer than is usual in 7’. tages proper, the 
terminal row of spots on both wings are white, the hind wing below has a well-marked 


164 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


postmedian band of small white spots and a narrow white spot on the transverse vein, 
and on the fore wing below there are two small suffused white spots near the apical 
third of cell 1 a, separated by the intraneural fold. Having regard to the relationship 
of 7. montanus and T. nigrescens, it is possible that sinina is really a distinct species 
from J. tages, but we have had no opportunity of comparing their male genitalia. 

T. cervantes, Grasl., is a larger and darker form, which prevails in Andalusia, 
but the male genitalia are identical with those of typical 7. tages, L. 


!THANAOS MonTANUS. (Plate XXIII. fig. 34.) 
Pyrgus montanus, Bremer, Bull. Acad. Petr. iii. p. 473 (1861) ; id. Lep. Ost-Sib. p. 31, pl. i1. fig. 4 
(1864). 
Thanaos rusticanus, Butler, Jour. Linn. Soc., Zool. ix. p. 58 (1866). 
Nisoniades montanus, Pryer, Rhop. Nihon. p. 35, pl. x. fig. 23 (1889). 
Thanaos montanus, Leech, Butt. China &c. p. 580. 


Hab. Amurland, Japan, China. 


! THANAOS LEECHII, n. sp. (Plate XXIII. fig. 35.) 
Thanaos montanus, var. nigrescens, Leech, Butt. China &c. p. 581, pl. xhi. fig. 2, g (1894). 


Distinguished from J. montanus by the characters given in the table above, of 
which the most constant are its smaller size and different male genitalia. The 
tegumen expands near the middle into two small, triangular, smooth, lateral dorsal 
lobes, as in 7. montanus, but these are not subtended, as in that species, by a pair of 
much larger spinose lobes. 

Hab. Wa-ssu-Kow, Ta-tsien-lu, W. China (fide Leech); Tchang-Kou, Tibet (fide 
Oberthiir). 


!THanaos MARLOYI. (Plate XXIII. figs. 32, 32 a.) 


Thanaos marloyi, Boisduval, Jeoues, pl. xivii. figs. 6, 7 (1832). 
Hesperia sericea, Freyer, Neuere Beitr. iii. pl. cexly. fig. 4 (1838) ; Herrich-Schiiffer, Schmett. 
Eur. i., Hesp. figs. 29, 30 (1846). 


Hab. S.E. Europe, Asia Minor, N. Persia, Syria. 


!THANAOS PELIAS: (Plate XXIII. figs. 53, 33 a.) 
Nisoniades pelias, Leech, Entomologist, xxiv. Suppl. p. 60 (June 1891). 


Nisoniades erebus, Groum-Grshimailo, Horz Soc. Ent. Ross. xxv. p. 461 (1891). 
Thanaos pelias, Leech, Butt. China &c. p. 581, pl. xli. fig. 3, g (1893-94). 


The male genitalia of 7. pelias, Leech, and 7. erebus, Gr.-Gr., are identical in form, 
as we find on examination of authentic specimens of each. 
Hab. W. China, E, Tibet (Pratt); N.E. Tibet (Growm-Grshimailo), 


A REVISION OF THE ORIENTAL HESPERIIDA, 165 


PAMPHILA. 

Pamphila, Fabricius, I]. Mag. vi. p. 287 (1807) ; Watson, P. Z. S. 1893, p. 89. Type palemon, Pall. 
Steropes, Boisduval, Voy. Astrol. p. 167 (1832), nom. preoce. 
Carterocephalus, Lederer, Verh. zool.-bot. Gesellsch. Wien, ii. p. 26 (1853). Type, palemon, Pall. 

A group of small species associated on account of their general resemblance to the 
Papilio palemon of Pallas. Palpi porrect. Antenne short, simply clavate. Fore 
wing: vein 2 arising about the middle of the lower edge of the cell; hind wing: 
cell 6 a little longer than cell 3. Hind tibiz more or less fringed (at least in the 
male), generally with but one pair of spurs, but with two pairs in P. abax, Ob. 

The species with which we are acquainted may be distinguished as follows :— 


1 (10). Hind wing below with no pale spot near the middle of cell 8. 

2 (9). Pale markings on the hind wing below non-metallic. 

3 (6). Fore wing above blackish brown with yellow spots. 

4 (5). Hind wing below with a subterminal series of pale spots. Hind 
tibize with one pair of spurs . . . Ponaeeor rc . . palemon, Pall. 

3 (4). No subterminal series of pale spots on aa wing below. cand 
Libizegwitbaiwospalrsiah spursiy | - et © i @ - - -)  ebaxr. Ob. 

6 (3). Fore wing above yellow with black spots. 


7 (8). Hind wing below with the pale spot on the transverse vein 


irregularly roundish, enclosing that vem. . . . . . . .  sélvius, Knoch. 
8 (7). Hind wing below with the pale spot on the transverse vein straight- 
sided within, bounded inwardly by that vein . .. . houangty, Ob. 


9 (2). Pale markings on the hind wing below metallic silvery white, and 
consisting of the following :—A streak along the upper edge of 
the cell from the base as far as the middle of cell 5, constricted 
or narrowly interrupted at its half-length, the lower edge of its 
outer half coalescing with a somewhat semi-circular spot in cell 4, 
a spot occupying the apical third of cell 6, a roundish spot near 
the base of cell 2, a clavate streak from the base to the middle 
of cell 1 4, and a submarginal series of oblong spots, placed one 
in each of cells 1 ¢, 2, and 3, sometimes feebly continued in 
cells4and5 ... . . . pulchra, Leech. 

10 (1). Hind wing below with a male: sig near nih riddle of call 8. 

11 (16). Pale markings of the upperside yellow or yellowish white.’ 

12 (13). Hind wing below with no pale spot near the apex of cells 2 and3; 
the dise of the wing occupied by a large subpyriform silvery 
spot, which arises near the middle of cell 7, and spreads outwards 
and downwards by way of the transverse vein ; a small pale spot 
near the basal third of the cell, and the indication of another 


near the middle of cell6. . . . . opis te - . avanti, de Nicév. 
13 (12). Hind wing below with a pale spot near ne apex of a of cells 
2 and 3. 


‘ See note to paragraph 16 post. 
VOL. XIV.—ParT Iv. No. 9.—October, 1897. Z 


166 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


14 (15). Fore wing below with a pale spot occupying the extreme base of 
cell 2, and another, at least twice as wide as high, near the 
middle of that cell . anes a 

15 (14). Fore wing below with no pale oe in ore bane of cell 2, but a 
subquadrate one before the middle of that cell . 

6 (11). Pale markings on upperside pure white." 

17 (18). Hind wing below with the pale spots near the apex of cells 6 and 7 
not connected with those near the apex of cells 2 and 3 by 
a pale band traversing cells 4 and 5. 

17 a (174). Fore wing above without white spots near the base ; hind wing 
above with two distinct unequal white discal spots c 

176 (17a). Fore wing above with two white spots near the base—one 
hyaline near the base of the cell, and the other opaque next the 
costa; hind wing above with a large white discal spot formed 
by the confluence of two unequal spots 2 

18 (17). Hind wing below with the pale spots near the apex of dais 6 
and 7 connected with those near the apex of cells 2 and 3 by a 
pale band traversing cells 4 and 5, and thus forming an 
irregular subterminal pale band. 

19 (20). Fore wing below with the pale spots in cells 1@ and 2 remote 
from that near the apical third of the cell . . . . . . . 

20 (19). Fore wing below with the pale spots in cells 1a and 2 joined to 
that near the apical third of the cell to form a continuous 
TACGUETU RING! Gy 6168 Gy C8 on Knees et) teede aot 


! PAMPHILA PALAMON. 
Papilio palemon, Pallas, Reise, vol. i. p. 471 (1771). 
Papilio paniscus, Fabricius, Syst. Ent. p. 531 (1775); Esper, Schmett. 
(1778 ?) ; i. 2, pl. xev. fig. 5 (1788). 


argyrostigma, Kversm. 


flavomaculatus, Ob. 


niveomaculatus, Ob. 


christophi, Gr.-Gr. 


dieckmanni, Graeser. 


micio, Ob. 


i. 1, pl. xxvii. fig. 2 


Papilio brontes, Wien. Verz. p. 160 (1776) ; Hiibner, Eur. Schmett. i. figs. 475, 476 (1798-1803). 


Hesperia mandan, Edwards, Proc, Ent. Soc. Philad. vol. 11. p. 20 (1863). 
Carterocephalus palemon, Bean, Can. Ent. xxv. p. 145 (1893). 


Hab. Europe to Amurland, N. America; Laggan, Alberta Terr., Brit. N. Amer., 


5000 ft. (Elwes); Nepigon, ‘Lake Superior (Fletcher). 


There is no doubt in our mind that the American form described as mandan is 


identical with the European. 


! PAMPHILA ABAX. 


Carterocephalus abax, Oberthiir, Eitud. d’Ent. xi. p. 27, pl. v. fig. 27 (1886). 


Pamphila abax, Leech, Butt. China &e. p. 587 (1893-4). 
Hab Ya-tsien-lo (Pratt); Moupin, E. Tibet (tricheldorf). 


1 What we identify as the female of niveomaculatus has the spots on the upperside yellowish instead of 


pure white. 


A REVISION OF THE ORIENTAL HESPERIIDA. 167 


! PAMPMILA SILVIUS. 
Papilio silvius, Knoch, Beitr. Ins. i. pl. v. figs. 1, 2 (1781) ; Esper, Schmett. i. 2, pl. Ixxx. figs. 5, 6 
(1782) ; Hiibner, Eur. Schmett. 1. figs. 477, 478 (1798-1803). 


Hab. N. Europe to Amurland (Graeser). 


! PAMPHILA HOUANGTY. 
Carterocephalus houangty, Oberthiir, Btud. d’Ent. xi. p. 27, pl. v. fig. 5 (1886). 
Pamphila houangty, Leech, Butt. China &e. p. 586 (1893-4). 


Hab. West China, Ta-tsien-lo (Pratt). 


! PAMPHILA PULCHRA. 
Pamphila pulchra, Leech, Entomologist, xxiv. Suppl. p. 59 (June 1891); id. Butt. China &e. 
p. 586, pl. xl. fig. 20, ¢ (1894). 
Carterocephalus ops, Groum-Grshimailo, Horce Ent. Soc. Ross. xxv. p. 460 (1891). 
Hab. Ta-tsien-lo, Wa-su-Kow, West China (Pratt); N.E. Tibet (Growm-Grshimailo, 


Jide Leech). 
We rely on Mr. Leech’s identification of C. ops, which we have not seen. 


! PAMPHILA AVANTI. 
Pamplula avanti, de Nicéville, Jour. As. Soc. Beng. 1886, p. 255, pl. xi. fig. 10, ¢; Watson, Hesp. 
Ind. p. 159. 


Type in coll. Elwes. 
Hab. Interior of Sikhim, circa 12,000 feet (White). 


! PAMPHILA ARGYROSTIGMA. 
Steropes argyrostigma, Eversmann, Bull. Mose. 1851, i. p. 624; Nordmann, 1. c. ii. p. 442, pl. xii. 
n figs. 1, 2 (1851). 
Carterocephalus argyrostigma, Alphéraky, Mém. sur Lép. v. p. 123 (1889). 
Pamphila argyrostigma, Leech, Butt. China &e. p. 585 (1893-4). 


Hab. Amurland (Eversmann) ; Mongolia (Potanine, fide Alphéraky). 


! PAMPHILA FLAVOMACULATUS. 
Carterocephalus flavomaculatus, Oberthiir, Etud. d’Ent. xi. p. 27, pl. ii. fig. 9 (1886). 
Pamphila flavomaculata, Leech, Butt. China &c. p. 587 (1893-4). 


Hab. West China (Pratt); E. Tibet (Biet). 


! PAMPHILA NIVEOMACULATUS. (Plate XIX. fig. 25, 2.) 
Carterocephalus niveomaculatus, Oberthir, Btud. d’Ent. xi. p. 27, pl. ii. fig. 8 (1886). 
Pamphila niveomaculata, Leech, Butt. China &e. p. 588 (1893-4). 

Hab. Ta-tsien-lo and E. Tibet (Biet); Koko-nor, Tibet (in coll. Rothschild). 

The female is not described or figured by M. Oberthiir, though he says that he has both 
sexes. ‘Two females from Koko-nor in coll. Rothschild differ in having the spots of the 
upperside yellowish instead of pure white, but we have little doubt that they are really 


females of this species. 
Z2 


168 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! PAMPHILA CHRISTOPHI. 
Carterocephalus christophi, Groum-Grshimailo, Hore Ent. Soc. Ross. xxv. p. 460 (1891). 
Pamphila niveomaculata, var. christophi, Leech, Butt. China &e. p. 588 (1893-4). 
Aubertia dulcis, Oberthiir, Etud. d’Ent. xx. p. 40, pl. ix. fig. 162 (1896). 

Hab. Sinin-Shan mountains (Growm-Grshimailo) ; Tibet (fide Oberthiir) 


! PAMPHILA DIECKMANNI. 
Carterocephalus dieckmanni, Graeser, Berl. ent. Zeits. 1888, p. 102. 
Carterocephalus gemmatus, Leech, Entomologist, xxiv. Suppl. p. 59 (June 1891). 
Carterocephalus demea, Oberthiir, Etud. d’Ent. xv. p. 19, pl. iii. fig. 24 (July 1891). 
Pamphila gemmata, Leech, Butt. China &c. p. 588 (1893-4). 

Hab. Wladiwostock, Amurland (Graeser) ; Ta-tsien-lo (Pratt); Lutschau (Dorries), 
Blagowetshensk (Dieckmann), Koslofska (Biikow in coll. Dieckmann); Koko-nor, Tibet 
(in coll. Rothschild). 

We have no hesitation in restoring Graeser’s name for this species, as we have 
examined three typical specimens kindly lent to us by Mr. Dieckmann. As a result 
of the reduction of the silvery markings on the hind wing below, a specimen from 
Koslofska, in coll. Dieckmann, has the roundish spot proper to the cell barely indicated. 


PAMPHILA MICIO. 
Carterocephalus micio, Oberthir, Etud. Ent. xv. p. 19, pl. iii. fig. 29 (1891). 
Pamphila micio, Leech, Butt. China &e. p. 589 (1894). 


Hab. Tse-kou, E. Tibet (fide Oberthiir). 


HETEROPTERUS. 
Heteropterus, Duméril, Zool. Anal. p. 271 (1806) ; Watson, P. Z. S. 1893, p. 89. Type morpheus, Pall. 
No hyaline spots on the wings. Antenne less than half as long as costa, simply ° 

clavate. Palpiporrect. Fore wingslongand narrow. Body longand slender. Vein 2 
of fore wing arising nearer to the base of the wing than to the end of the cell. No 
sex-mark on the wings of the male. 
1 (2). Hind wing below yellow, with twelve suboval black-margined white spots. 

Front tibiz with a short spur on the mmer side near the middle, hind 

tibize with two pairs of spurs. Tornus of hind wing evident . . . morpheus, Pall. 
2 (1). Hind wing below dull greenish yellow, without suboval white spots. 

Front tibiz simple, hind tibiz with only one pair of spurs. ‘Tornus of 

hind wing completely rounded off. 


3 (4). Hind wing below without silvery markings . . . - . . unicolor, Brem. 
4 (3). Hind wing below with a silvery middle stripe Gin the _ to the 
termen, and frequently another incell16 . . . . . . . . . ornatus, Brem. 


! HETEROPTERUS MORPHEUS. 
Papilio morpheus, Pallas, Reise, vol. i. p. 471 (1771). 


Papilio steropes, Wien. Verz. p. 160 (1776) ; Esper, Schmett. i. 1, pl. xli. fig. 1 (1778?) ; Htibner, 
Eur. Schmett. 1. figs. 473, 474 (1798-1803). 


A REVISION OF THE ORIENTAL HESPERIIDA. 169 


In the male the hind wing below has the outer half of the fringe brown, but in the 
female the same fringe is brown only in spots opposite the veins. 
Hab. Biarritz (Elwes); Europe ; Korea (Leech) ; Amurland (Graeser). 


! HETEROPTERUS UNICOLOR. 
Steropes unicolor, Bremer & Grey, Schmett. N. China’s, p. 10, pl. iii. fig. 3 (1853). 
Cyclopides ornatus, Bremer, Bull. Acad. Petr. iii. p. 473 (1861); id. Lep. Ost-Sib. p. 33, pl. ii. 

fig. 5 (1864). 

Cyclopides unicolor, var. ornatus, Stgr. Rom. Mém. vi. p. 209. 

Hab. Japan (Pryer); Central China (Pratt); Amurland ( fide Staudinger). 

Though ornatus and wnicolor have hitherto been treated as distinct species, I agree 
with Staudinger and Leech that they cannot be separated, on account of the inter- 
mediate forms. The genitalia offer no differences. 


DEJEANIA. 
Dejeania, Oberthiir, Etud. d’Ent. xx. p. 40 (1896). Type bicolor, Oberthiir. 

Antenne bluntly pointed, non-apiculate, half as long as the costa. Palpi porrect, 
second joint laxly scaled, third joint of moderate length. Fore wing: dorsum a littie 
longer than the termen, the latter evenly curved; vein 11 free, transverse vein erect, 
vein 5 straight, equidistant from veins 4 and 6, vein 2 from near the half-length of 
the cell, vein 3 arising twice as near to vein 4 as to vein 2. Hind wing about one- 
fourth longer than broad, termen evenly rounded, cell a little longer than half the 
wing, vein 2 from beyond the half-length of the cell, vein 3 arising much before the 
cell-end opposite to the origin of 7, vein 5 strong, transverse vein feeble but practically 
erect. Tibial epiphysis present, but very small. Hind tibiz in the male fringed, and 
bearing two pairs of spurs. 

Allied to Heterepterus, Dum., in venation, palpi, and antenne, but has the hind wing 
rounded at the apex instead of produced. 

The above particulars are taken from authentic specimens of Dejeania bicolor, Ob., a 
brown species with a broad oblique postmedian yellow band on the fore wing, bearing 
a narrow dark spot on the transverse vein, and a straight narrow yellow band on the 
hind wing below from the apex of vein | 4 to the apex of vein 7. 


DEJEANIA BICOLOR. 
Dejeunia bicolor, Oberthir, t. c. p. 40, pl. ix. fig. 163, 3. 
Hab. 'Tse-kou, Tibet (Dubernard) ; Sido-Léu ( fide Oberthiir). 


BaRacts. 
Baracus, Moore, Lep. Cey. i. p. 162 (1881) ; Watson, P. Z. S. 1893, p. 114. Type vittatus, Feld. 
Antenne about half as long as the costa, apiculus about as long as the diameter 
of the eye. Palpi ascending ; third joint porrect, prominent, about one-half as long as 


170 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


the diameter of the eye. Fore wing: vein 11 running closer to 12 than to 10, vein 2 
arising from the distal third of the lower edge of the cell; 3, 4, 5, and 6 almost equi- 
distant at the base, 6 and 7 from the same point. Hind tibie fringed and bearing two 
pairs of spurs. 

What may be regarded as the typical wing-pattern in this genus is peculiar: the 
pale markings on the fore wing above consist of four oblong spots placed in two pairs, 
one near the base of each of cells 2 and 3, and one near the base of each of cells 6 
and 7, and there is sometimes a pale spot or spots in cell l@; in the latter case the 
pale markings almost assume the form of a postmedian curved macular band, inter- 
rupted in cells 4 and 5. ‘The normal pattern is found in both sexes of B. subditus, 
Moore, and B. septentrionum, de Nicéy., but only in the female of B. vittatus, Felder. 

The species known to us are distinguished as follows :— 


Males. 


1. Upperside brown ; interno-basal half of the fore wing and the 
entire hind wing, except the costa and the termen, pale. . 2. 
Upperside brown; fore wing with two pairs of pale spots, one 
in each of cells 2 and 3, 6 and 7, and sometimes onein cellla. 3. 
2. Pale portion of upperside sordid creamy white . . . . . vittatus, Feld. 
Pale portion of upperside pale whitish blue . . . . ~ . plumbeolus, Feld. 
3. Hind wing below ochreous brown, with a pale stripe from the 
base of the wing as far as the half-length of cell 5, and a ter- 
minal series of pale streaks, one in each of cells 2, 3,4, and6. 4. 
Hind wing below at first sight appearing yellow, with many 
brown streaks, but really brown sprinkled with yellow scales, 
and having the veins, a spot occupying the basal half of 
cells 4-5, and sometimes continued to the base of the wing, 
and a subterminal series of spots consisting of one in each 
of cells 2-7, of which those in cells 2, 3, and 6 are about 


twice as long as the others, yellow. . . . . . hampsoni, n. sp. 
4, Expanse about 32 mm. Pale markings on the hind wing [de Nicéville. 
below but little paler than the ground-colour . . . . septentrionum, Wood-Mason & 


Expanse about 27 mm. Pale markings on the hind wing 
below much paler than the ground-colour, the stripe from 
the base of the wing to the middle of cell 5, and another 
incelleWG; whitish! oy) 6 se" «ca Meme oulim te aie) a mNSHOLICES Viaores 


! BARACUS VITTATUS. 


Isoteinon vitiatus, Felder, Verh. zool.-bot. Gesellsch. Wien, xii. p. 480 (1862). 
Baracus vittatus, Moore, Lep. Cey.i. p. 162, pl. lxix. figs. 1, 1 @ (1881) ; Watson, Hesp. Ind. p. 151 
(1891). 


Hab. Nuwara Eliya, Ceylon, 7000 feet (Elwes). 


A REVISION OF THE ORIENTAL HESPERIIDA. ill 


! BARACUS PLUMBEOLUS. 

Hesperia plumbeola, Felder, Reise Nov., Lep. iii. p. 519, pl. Ixxi. fig. 20, 3 (1867). 

In the female of this species the blue colour of the upperside is on the hind wing 
less extensive than in the male, and on the fore wing is almost confined to the dorsum 
below vein 1a, leaving the remainder of the wing brown, save for a few pale bluish 
scales near the base. 

Hab. Luzon, Philippines (Semper). 


! BARACUS SUBDITUS. 
Baracus subditus, Moore, P. Z. S. 1883, p. 584; Watson, Hesp. Ind. p. 151 (1891). 

Hab. Nilgiri hills (Hampson); Palnai hills (Castets). 

A specimen from the Nilgiris (Hampson), taken on the 14th October, has the pale 
markings on the hind wing below as feeble as they are in B. septentrionum. 

! BARACUS SEPTENTRIONUM. 
Baracus septentrionum, Wood-Mason & de Nicéyille, Jour. As. Soc. Beng. 1886, p. 379, pl. xviii. 

figs. 4, 4a, § ; Watson, Hesp. Ind. p. 151 (1891). 


Hab. Sikkim (Knyvett) ; Shan hills (Manders). 


! BARACUS HAMPSONI, n. sp. (Plate XVIII. fig. 21, ¢.) 

3. Upperside brown; fore wing with four yellow spots, one in each of cells 2, 3, 
6, and 7. Underside: fore wing dark brown from the dorsum as far as the upper edge’ 
of the cell, the outline of the dark space concave from the apex of vein 3 to a point 
near the middle of cell 6, the remainder of the wing yellow, the yellow spots in cells 6 
and 7 as on the upperside: hind wing at first sight appearing yellow, with many brown 
streaks, but really brown closely sprinkled with yellow scales, and having the veins, a 
spot occupying the basal half of cells 4-5 and sometimes continued to the base of the 
wing, and a subterminal series of spots consisting of a spot in each of cells 2-7, of 
which those in cells 2, 5, and 6 are about twice as long as the others, yellow; a pale 
anteciliary line on both wings. Fringes grey, chequered with darker, the short scales 
dark brown. 

2. Similar to the male, but slightly larger. 

Expanse 24—26 mm. 

Hab. N. Canara (Aitken). 


Described from five males and two females in coll. Elwes. 


ASTICTOPTERUS. 
Astictopterus, Felder, Wien. ent. Monatsb. iv. p. 401 (1860) ; Watson, P. Z.S. 1893, p. 114. 
Type jama, Feld. 
In structure of the antenne, palpi, and veins this genus does not differ essentially 
from Baracus, although vein 2 of the fore wing arises near the middle of the lower 
edge of the cell, and the last joint of the palpi is comparatively shorter. The facies of 


172 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


its members, however, is different, and the hind tibie are not fringed with long hairs 
as in Baracus. The species are dark brown above, with or without a few small white 
spots, which, when present, are placed near the base of cells 6 and 7 or 6,7 and 8, 
and more rarely there is another near the basal third of cell 3. 
The species known to us may be distinguished as follows :— 
1 (2). Fore wing below umber-brown, the costal and apical regions but little 
paler: male without hyaline spots; female usually with hyaline spots 
in cells 6, 7, and 8. Hind wing below umber-brown, with very ill- 
defined antemedian and postmedian macular dark bands . . . . .  olivascens, Moore. 
2 (1). Fore wing below fuliginous brown, the costal and apical regions pale 
reddish brown, the latter with some grey scaling next the termen 
from vein 8 to the apex: both sexes usually with hyaline spots in 
cells 6, 7, and 8, and in the female there is frequently a short hyaline 
spot near the basal third of cell 3. Hind wing below pale reddish 
brown, with antemedian and postmedian darker bands, of which the 
latter sometimes bears some spots of whitish scales . . . . . . Aenrici, Holland. 


! ASTICTOPTERUS OLIVASCENS. (Plate XVIII. fig. 16, 3.) 
Astictopterus olivascens, Moore, P. Z. S. 1878, p. 692 ; Watson, Hesp. Ind. p. 146 (1891). 

Hab. Khasia hills (Hamilton) ; Naga hills (Doherty) ; Shan hills (Manders); Sikkim 
(Métler); Buxar, Bhutan (in coll. Elwes); Perak (Doherty); W. Java (Piepers); 
Cherrapunji (in coll. Swinhoe). 


! ASTICTOPTERUS HENRICI. (Plate XVIII. fig. 17, 3.) 

Cyclopides henrici, Holland, Trans. Amer, Ent. Soe, xiv. p. 124, pl. 11. fig. 5 (1887). 
Astictopterus kada, Swinhoe, Trans. Ent. Soc. Lond. 1898, p. 328. 
Astictopterus olivascens, Leech, Butt. China &c. p. 629, pl. xhi. fig. 1 (1894). 

Hab. Khasia hills (Hamilton); Burmah (Adamson); E. Pegu (Doherty); Nilawa 
(Manders) ; W. China (Leech); Chia-ting-fu (Pratt). 

Chinese specimens are darker on the underside than Indian ones. Leech’s figure 
(l. ¢.) is not quite characteristic. 

Dr. Holland, to whom we had sent a transcript of our statement of the diagnostic 
characters of olivascens and henrici, as given in the preceding table (using the name 
kada for the latter) writes as follows :—“1 have complied with your request, and 
carefully examined my type of Astictopterus! (false Cyclopides) henrici. It agrees best 
and quite certainly with the description of sada, Swinhoe. I have not a particle of 
doubt that it is the latter.” 


ASTICTOPTERUS JAMA. 


Astictopterus jama, Felder, Wien. ent. Mon. iv. p. 401 (1860) ; Distant, Rhop. Mal. p. 401 (1886). 


Not having seen this species, we transcribe the original description :—“ Alis utrinque 
saturate fuscis, cervino atomatis. ¢. Coll. Felder. 4. pygmeo, Fabr., species hee in 


A REVISION OF THE ORIENTAL HESPERIIDA. 175 


India continenti etiam occurrens, similis, sed ale breviores et latiores palporumque 
articulus tertius minutus.” 

Hab. Malay Peninsula. 

Felder’s collection is now in the Museum of the Hon. Walter Rothschild at Tring, 
and Dr. Jordan, to whom we applied for an account of Felder’s type of this species, 
writes that the specimen which stands as Astictopterus jama in Felder’s collection is 
not that species, as it has a broad yellow band across the fore wing somewhat like 
sindu, and that there are a number of specimens from Malacca in the same drawer 
without names, of which one may be the type of jama. We do not know what insect 
was identified by Distant as jama, Feld., nor how Watson identified as jama, Feld., 
the insect from which his description (P.{Z. S. 1893, p. 114) of the genus Astictopterus 
was taken. 


APOSTICTOPTERUS. 
Apostictopterus, Leech, Butt. China &c. p. 630 (1894). Type fuliginosus, Leech. 

Antenne about half as long as costa, club gradual; apiculus robust. Palpi ascend- 
ing; third joint porrect, cylindrical, at least as long as half the diameter of the eye. 
Wings comparatively long and narrow: tip of fore wing truncate, 7.¢. the termen is 
nearly vertical from vein 5 to the costa, and inwardly oblique from vein 5 to vein 1a, 
with an obtuse but distinct angle at the end of vein 5; vein 2 from the basal third 
of the lower edge of the cell; veinda trifle nearer 4 than 6; vein 11 normal, 7. ¢. not 
in any way approaching vein 12. ‘Tibial epiphysis present, hind tibiee with two pairs 
of spurs. 

The foregoing characters are present in Mr. Leech’s unique type of Apostictopterus 
fuliginosus, which he has been so good as to entrust tous for examination. ‘The genus 
should stand next to Astictopterus. 


APOSTICTOPTERUS FULIGINOSUS. 
Apostictopterus fuliginosus, Leech, Butt. China &c. p. 631, pl. xxxviii. fig. 8, g (1894). 
Hab. Omei Shan, Western China ( fide Leech). 


SANCUS. 
Sancus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 395; Watson, P. Z. 8, 1893, p. 87. 
Type subfasciatus, Moore =pulligo, Mab. : 

Body and wings above entirely dark brown. Palpi appressed, 2nd joint densely 
scaled, 3rd joint almost concealed. Fore wing with vein 12 shortly touching vein 11 
near its base. Sex-mark of the male, when present, an oblong-oval patch of modified 
scales in the basal third of cell 1a on the fore wing below and touching the median 
and the base of vein 2; its presence is not always well indicated on the upper surface 
of the wing. The fore wing is rather narrow and feebly subtruncate at the apex, the 

VOL. Xiv-—part Iv. No. 10.—October, 1897. 2A 


174 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


termen between veins 5 and 6 reaching a little beyond the actual tip; hind wing 
rounded, its greatest length not exceeding that of the dorsum of the fore wing. 
Abdomen comparatively long and slender. On the hind wing below the common 
pattern consists of the following spots of pale scales, namely :—one in cell 7 erect from 
the base of vein 7, one near the middle of the upper margin of the cell, one in the 
basal third of cell 6, and four in a straight line almost parallel with the termen, 
one each in cells 2, 3, 4, 5; some of these spots are sometimes suppressed, and in some 
specimens the whole are scarcely discernible. 

Two species are mentioned by Watson (P. Z.S. 1893, p. 87), which may be 
distinguished as follows :— 


Sex-mark present in the male. Hind wing below with pale spots or bands, or 
both . ae ee ete asl ee Rg ee a Sea pulligo, Mab. 
Sex-mark wanting in the male. Hind wing below unmarked . . . . . . «. fuscula, Snell. 
Watson says of S. fuscula that the male mark is present, but inconspicuous, and he 
separates the species from S. pulligo by having the underside of the hind wing entirely 
unmarked, and by the beautiful plum-like bloom of the upperside, though this last 
character is only apparent in fresh specimens. 


! SANCUS PULLIGO. 


Tagiades pulligo, Mabille, Bull. Soc. Ent. Fr. 1876, p. xxvi; id. Ann. Soc. Ent. Fr, 1876, p. 272. 
Astictopterus subfasciatus, Moore, P. Z. 8. 1878, p. 842; Wood-Mason & de Nicéville, Jour, As. 
Soc. Beng. 1886, p. 380, pl. xviii. figs. 1, 1 a, ¢ ; Watson, Hesp, Ind, p. 148 (1891). 

Astictopterus ulunda, Staudinger, Iris, ii. p. 146 (1889). 
Sancus subfasciatus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 396. 
? Antigonus kethra, Plétz, Jahrb. Nass. Ver. 1884, p. 24. 
? Astictopterus kethra, Plétz, Stett. ent. Zeit. xlvii. p. 110 (1886). 
Sancus pulligo, Semper, Schmett. Philipp. p. 319, pl. xlix. fig. 1, ¢ (1892). 

Hab. Nilghiri hills (Hampson); Akyab, Moulmein (Adamson); Perak, Pulo Laut, 
Bali (Doherty); Java (Piepers) ; Borneo, Philippines (Semper) ; Palawan (Platen). 


SANCUS FUSCULA. 


Tagiades? fuscula, Snellen, Tijd. voor Ent. vol. xxi. p. 42, pl. ii. fig. 3 (1878) ; de Nicéville, Jour. 
Bomb. Nat. Hist. Soc. 1891, p. 397. 


Hab. S.W. Celebes (fide de Nicéville). 


KORUTHAIALOS. 
Koruthaialos, Watson, P. Z. S. 18938, p. 76, pl. ii. fig. 8. Type hector, Wats. 
A genus of few species, well exemplified by the Astictopterus xanites of Butler. 
Apart from facies, it differs from Suastus and Jambrix in the fact that vein 11 of the 
fore wing touches or anastomoses with vein 12 fora portion of its length. Perhaps the 


A REVISION OF THE ORIENTAL HESPERIIDA. 175 


most remarkable feature of the genus is the frenulum-like tuft of bristly hairs found at 
the base of the costa of the hind wing in the male. 
The following table deals only with the three species in coll. Elwes :— 


; Males. 

1 (4). Fore wing with a red band, at least on the underside. 
2 (3). Red band on fore wing below broader, its breadth near the : 

middle about equal to one-third of the length of the costa . wanites, Butl. 
3 (2). Red band on fore wing below narrower, its breadth near the 

middle about equal to one-fourth of the length of the costa. hector, Wats. 
4 (1). Fore wg without anyred band . . . .... =. . . butleri, W.-M. & de Nicév. 

With regard to K. wanites and K. hector, it is difficult to lay down any satisfactory 

character for the separation of these two species, if species they are. In their typical 
state they offer no difficulty, H. xanites being the larger insect, expanding about 
4] mm., with the red band on the fore wing above broad, and reaching from the middle 
of the costa to the tornus; whilst X. hector is smaller, expanding about 35 mm., with 
the red band on the fore wing above comparatively narrower and rarely reaching quite 
to the tornus, frequently abbreviated at each end, and sometimes entirely wanting. 
Pending the comparison of the male genitalia of typical specimens when opportunity 
offers, we find that the most constant character by which to separate the two forms lies 
in the relative breadth of the red band of the fore wing, as exhibited on the underside. 


! KORUTHAIALOS XANITES. 
Astictopterus xanites, Butler, Trans. Ent. Soc. Lond. 1870, p. 510; id. Trans. Linn. Zool. Soe. ser. 2, 
vol. i. p. 555, pl. lxix. fig. 7 (1877). 
Koruthaialos xanites, Watson, ‘P. Z. 8. 1893, p. 77. 
Hab. Borneo (fide Butler); Malacca (jide Watson); Kina Balu, Borneo (Water- 
stradt); Perak, Bali (Doherty). 


! KORUTHAIALOS HECTOR. 
Koruthaialos hector, Watson, P. Z. 8. 1893, p. 77. 
Astictopterus xanites, Distant, Rhop. Mal. p. 402, pl. xxxiv. fig. 28 (1886) ; Watson, Hesp. Ind. 

p. 145 (1891). 

Astictopterus xanites, var. palawites, Staudinger, Iris, 11. p. 148 (1889). 
Kerana gemmifer, Semper, Schmett. Phillipp. p. 318 (1892), sec. spec. comm. 

Hab. Luzon (fide Semper); Palawan (fide Staudinger) ; Burmah (Adamson, Doherty) ; 
Tenasserim (Bingham) ; Perak, Pulo Laut, Bali, Arjuno, Java (Doherty) ; Java (Piepers) ; 
Bunguran, Natuna Is. (Hverett); Palawan (in coll. Rothschild). 


! KORUTHAIALOS BUTLERI. 


Astictopterus butleri, Wood-Mason & de Nicéyille, Jour. As. Soc. Beng. 1883, p. 98, pl. x. 


fig. 3, g; ud. op. cit. 1886, p. 380. 
2a 2 


176 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Dark fuliginous brown with somewhat of a purple tinge, the latter most evident 
on hind wing below. Male with no trace of a pale band on either side of the 
fore wing; in the female! (fide de Nicéville) the fore wing has an obsolete oblique 
discal red band, which is sometimes quite absent from the upperside, but always 
distinctly traceable on the underside. ‘This species, though distinct enough to admit 
of its definition, is evidently but little removed from the preceding ; being, in fact, more 
remote from it in the matter of facies than it is in the form of the male genitalia. 

Hab. Sikkim (Moller); Upper Assam (Doherty); Khasias (Hamilton). 


KoRUTHAIALOS VERONES. 
Astictopterus verones, Hewitson, Ann. & Mag. Nat. Hist. (5) i. p. 341 (1878). 
Koruthaialos verones, de Nicéville & Martin, Jour. As. Soc. Beng. 1895, p. 534. 

“Both sides rufous brown. Underside of the anterior wing marked by a subapical 
rufous spot.” (Hewitson, l.c.) 

Hab. Sumatra ( fide Hewitson). 


KKORUTHAIALOS KERALA. 
Koruthaialos kerala, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1896, p. 20, pl. T. fig. 48, ¢. 


“ Male. Upperside, both wings shining fuscous. Fore wing with a broad oblique 
discal orange fascia exactly as in Kerana armatus, Druce, that is to say, the band is 
about twice as long as it is broad, and it does not quite reach the costa, the outer 
margin at the anal angle, or the inner margin; but it differs slightly in form from the 
band in that species. as its inner edge anteriorly is obliquely cut off instead of being 
continued straight to the margin. Hind wing unmarked. Underside: both wings 
paler than on the upperside, of a more sooty brown. Fore wing has the discal band 
rather broader than on the upperside, reaching the inner margin; it is paler, moreover, 
throughout, especially so posteriorly. Hind wing immaculate. Antenne black, the 
apex of the club paler beneath. Palpi blunt, the third joint hidden beneath the 
second.— Female exactly like the male.” 

Expanse, ¢, 44-45 mm.; 9, 44-54 mm. 

Hab. Perak, Malay Peninsula; Battak Mountain, N.E. Sumatra (fide de Nicéville). 

We have not seen this species. 


KORUTHAIALOS KOPHENE. 
Koruthaialos kophene, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1896, p. 21, pl. T. figs.49 5,509. 


«“ Female. Upperside, both wings sooty-brown. Fore wing with a broad oblique 
discal orange fascia (not as broad as in Celenorrhinus ladana, Butler, Kerana armatus, 
Druce, and Koruthaialos kerala, de Nicéville) commencing near the costa, ending 
near the submedian nervure, its edges irregular, crossed by the dark brown veins, 


1 We have never seen the female of this species. 


A REVISION OF THE ORIENTAL HESPERIID 2, 177 


bearing a dark brown line which defines the discocellular nervyules. Hind wing 
unmarked. Underside, both wings rather paler than above. Fore wing with the 
discal band broader and paler, especially posteriorly, than on the upperside, the 
discocellular dark line narrower and more prominent. Hind wing unmarked. Antenne 
black, the apex of the club beneath paler. Palpi blunt, the third joint hidden 
beneath the second.—WMale similar to the female but smaller, the orange fascia on 
both sides of the fore wing rather more obscure and narrower.” 

Expanse, ¢,40; 2, 45-525 mm. 

Hab. N.E. Sumatra, Central Java ( fide de Nicéville). 

We have not seen this species. It will be observed that both this species and the 
preceding are remarkable for the position of the third joint of the palpi; in all the 
Hesperiide known to us the third joint of the palpi proceeds from the distal end 
of the second joint and is not hidden beneath it. 


SuADA. 
Suada, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 370. Type swerga, de Nicéville. 
This genus is distinguished from Swastus by having vein 11 of the fore wing 


touching or approaching vein 12. 
The species known to us may be distinguished as follows :— 


1 (2). Hind wing above not marked with white. . . . . . . swerga,de Nicév. 
2 (1). Subtornal region of hind wing above in greater or less ant Bi pure ite 
8 (4). Fore wing fuligimous brown, without hyaline spots or, at most, with one 

or more of the subapical series. Hind wing below pure white, 

unspotted, narrowly brown along the costa. . . : . . . cataleucos, Stgr. 
4 (3). Fore wing above fuliginous brown, with three small feakne white spots 

on the disc, one each, subequal in size, in the cell and cell 2 next the 

base of vein 3, and one, smaller and triangular, in the base of 


cell 3. 
5 (6). Hind wing below white as far as vein 7, thence to the costa brown ; 
termen suffused with brown in the apex of cells land2 . . . . . albinus, Semper. 


6 (5). Hind wing below white as far as vein 7, thence to the costa brown, the 
white portion with some black spots, namely, one in the apex of the 
cell, one near the apical third and one at the apex of cell 1 4, both large 
(the former subquadrate), and followed by some smaller ones in cells 
2, 3, and 4, 5, the whole indicating a postmedian and terminal macular 
band respectively. . . 2 © + + + + + + 5 © + © « + « ScOpas, Stgr. 


! SUADA SWERGA. 
Hesperia swerga, de Nicéville, Jour. As. Soc. Beng. 1883, p. 89, pl. x. fig. 12, ¢. 
Suastus molleri, Moore, Jour. As. Soc. Beng. 1884, p. 49. 
Suada swerga, de Nicéville, Jour. Bomb. Nat. Hist. Soc, 1895, p. 372. 


Hab. Sikkim (Moller); Arjuno, Java (Doherty). 


178 MESSRS. H. J. ELWES AND JAMES EDWARDS 


!SuUADA CATALEUCOS. 


Heteropterus? cataleucos, Staudinger, Iris, u. p. 162, pl. i. fig. 13, g (1889). 
Hab. Pulo Laut, Borneo (Doherty); Palawan (Platen). 


SUADA ALBINUS. 
Suastus albinus, Semper, Schmett. Philipp. p. 299, pl. xlix. fig. 8, 9 (nee g) (1892). 

Hab. 8.W. Mindanao (Semper) ; Davao, Mindoro (Staudinger). 

Herr Semper is mistaken as to the sex of his single example of this species; it is 
undoubtedly a female. 


SUADA SCOPAS. 
Heteropterus ? (Steropes) scopas, Staudinger, Iris, ii. p. 161, pl. ii. fig. 12, ¢ (1889). 
Hab. Palawan (Platen). 


Distinguished from albinus by the smaller amount of white on the hind wing above 
and the black spots on the white portion of the hind wing below. 


SUASTUS. 
Suastus, Moore, Lep. Cey. vol. i. p. 168 (1881) ; Watson, P.Z. 8.1893, p. 75. Type gremius, Fab. 


No hyaline spot in cell 4 of fore wing. Third joint of palpi acicular, erect, one- 
third to one-half as long as the diameter of the eye. Fore wing: vein 11 not 
touching 12; vein 2 nearer the base of the wing than the end of the cell. 

We are acquainted with five species of this genus, distinguished as follows :— 


1 (8). Hind wing above not marked with white. 
2 (7). Hind wing below not sprinkled with purple scales. 
3 (6). Pale spot in cell lia of fore wing below not reaching the termen. 


4 (5). Dark spots on hind wing below black. Expanse about 38 mm.. . . gremius, Fab. 
5 (4). Dark spots on hind wing below but little darker than the ground- 
colour. Expanse about30mm. .. . 5 ; iauket ae wie eel, ube. 


6 (8). Pale spotin cell 1 a of fore wing below pure hte, and, with the exception 

of a small piece in the upper outer angle of that cell, oceupymg 

the entire space between the dorsum and vein 2 from the termen to 

the middle of the wing. Hind wing below brown, closely sprinkled 

with whitish-grey scales and bearing a few brown spots, of which 

two in cell 1 4, one near the base of vein 2, and the other near the 

apical third of the cell, one each in cells 2and 3, and one im the 

apex of the cell are the most evident; an anteciliary brown line; 

fringe white . . . : aS, (c 22 ee. deem wai negneus: Semper: 
7 (2). Hind wing below rathien thickly apeaikied with itepie scales on the 

basalitwo-thirds:. .” <-.) “.ciheucbiaaiee iene os uae ete Be) ae ee moEpUNCrUS Swine 


A REVISION OF THE ORIENTAL HESPERIID A. 179 


8 (1). Subtornal region of hind wing above in greater or less part pure 
white. Hind wing below white; costa brown as far as vein 7 ; a large 
subquadrate black spot beyond the middle of cell 14, a small one 
next the transverse vein, and one or two others . . . . . . . tripura, de Nicév. 
The Carystus telesinus of Mabille, which Semper includes in this genus, we place 
tentatively in Plastingia: it has not the facies of Swastus nor the long acicular third 
joint of the palpi characteristic of that genus. 


! SUASTUS GREMIUS. 
Hesperia gremius, Fabricius, Ent. Syst. Suppl. p. 483 (1798). 
Hesperia divodasa, Moore, P. Z. 8. 1865, p. 791. 
Hesperia subgrisea, Moore, P.Z.S. 1878, p. 689. 
Suastus gremius, Moore, Lep. Cey. vol. i. p. 168 (1881). 
Suastus subgrisea, Moore, l. ec. 


Hab. N.W. Himalayas (Young); Bangalore, Ganjam (Minchin) ; Sikkim (Moller) ; 
Khasia (Hlwes) ; Bombay (Swinhoe); Burmah (Adamson). 


SUASTUS CHILON. 
Suastus chilon, Doherty, Jour. As. Soc. Beng. 1891, p. 198. 


** Above, male all dark brown, no hyaline markings nor patches of lighter brown 
scales. Below, fore wing with a minute white dot discally in the lower median space, 
the subapical hyaline spots represented by two slight dark’streaks, the lower (in one 
specimen) containing a lighter dot. Hind wing nearly white (not grey as in 
S, gremius), the borders dark ; a conspicuous black cell-spot, and a row of black discal 
spots, six in one specimen, four in the other. The absence of hyaline spots 
distinguishes it from all others. Two males, Sumba coast.” (Doherty, 1. c.) 

We have not seen this species and do not know whether the types still exist. 


!Suastus sana. (Plate XVIII. fig. 15, 3.) 
Hesperia sala, Hewitson, Trans. Ent. Soc. Lond. 1866, p. 500; Wood-Mason & de Nicéyille, 


Jour. As. Soc. Beng. 1881, p. 259. 
Suastus aditus, Moore, Jour. As. Soc. Beng. 1884, p. 49. 


Hab. Sikkim (Moller); E. Pegu, Andamans (Doherty) ; Java (Piepers). 


! SUASTUS MIGREUS. 
Suastus migreus, Semper, Schmett. Philipp. p. 300, pl. xlix. fig. 9, g (nec ?) (1892). 


Hab. Luzon, Mindanao (Semper) ; Mindoro (coll. Staudinger). 

Herr Semper has been good enough to lend us the specimen figured in his work, 
which is certainly a male. The species is well characterized, by the large white patch 
in the tornus of the fore wing below. 


180 MESSRS. H. J. ELWES AND JAMES EDWARDS. 


!Suastus BipuNcTus. (Plate XVIII. fig. 14, ¢.) 
Suastus bipunctus, Swinhoe, Ann. & Mag. Nat. Hist. (6) v. p. 364 (1890). 
Suastus aditus, Hampson, Jour. As. Soc. Beng. 1888, p. 365. 


Hab. Nilgiri hills (Hampson). 


! SUASTUS TRIPURA. 
Tagiades tripura, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 392, pl. G. fig. 39, 2. 
Carystus albescens, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 51 (1893). 


Hab. Perak, Pulo Laut, Bali (Doherty) ; Java (Fruhstorfer). 


SUASTUS MINUTA. 


Tagiades minuta, Moore, Ann. & Mag. Nat. Hist. (4) xx. p. 343 (1877); id. Lep. Cey. 1. p. 176, 
pl. xviii. figs. 4, 4.@ (1881) ; Watson, Hesp. Ind. p. 96 (1891). 


Resembles S. ¢ripwra in general appearance, but has the hind wing above entirely 
brown. Hind wing below white, brown along the costa; an ill-defined brown spot 
near the apical third of cell 1 4, an indication of a smaller one in cell 2, and another 
next the transverse vein. 

Hab. Ceylon. 


SUASTUS ROBSONII. 
Suastus robsonii, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 372, pl. Q. fig. 50, g. 


Nearest to S. minuta. Hind wing below “(as seen under a strong magnifying- 
glass) overlaid throughout with dull ochreous scales; an elongated minute black 
spot at the end of the cell.” Described from a single example taken at Masuri, 
W. Himalayas, 12th July, 1892. 


SUASTUS PHIDITIA. 

Hesperia phiditia, Hewitson, Trans. Ent. Soc. Lond. ser. 3, vol. ii. p. 501 (1866). 
Suastus phiditia, de Nicéville & Martin, Jour. As. Soc. Beng. 1895, p. 535. 

This species, which we have not seen, has one large vitreous spot on the hind wing 
above, and the kind wing below “ochreous, crossed at the middle by a band of four 
brown spots” (rufo-ochraceis, fascia fuscd). 

Hab. Sumatra (fide Hewitson). 


TAMBRIX. 


Iambrix, Watson, P. Z. 8. 1893, p. 76, pl. iii. fig. 25. Type salsala, Moore. 


A genus established by Watson for the Misoniades salsala of Moore and its 
immediate allies. It differs from Swastus in facies and in the fact that vein 3 of the 
fore wing arises immediately before the end of the cell. The males of the three 


A REVISION OF THE ORIENTAL HESPERIIDZ. 181 


species placed in this genus by its author, together with one other, may be 
distinguished by the characters given below. In salsala and stellifer the white 
spots on the hind wing below are very inconstant in size and number, and so, ina 
lesser degree, are the white spots on the fore wing of the females of those species. 


Males. 


Fore wing above not banded with yellow-red. Hind wing below with 2—4 small 
snow-white spots (sometimes absent). No patch of androconia on the 
hind wing above. 
Fore. wing above with a sordid yellow macular band running obliquely 
outwards from the middle of the dorsum (sometimes indistinct, but 
always indicated by more or less distinct patches of golden-yellow scales). salsala, Moore. 
Fore wing above without any trace of a pale discal macular oblique band . ._ stellifer, Butl. 
Fore wing above with a broad yellow-red postmedian band, which is sometimes 
more or less abbreviated at each end. A large patch of androconia at 
the base of the hind wing above near the costa. 
Fore wing below: red band narrower, lying at a right angle to the dorsum 
and reaching about halfway across cell 3, its inner edge nearly straight, 
Mo, maclnMatl Ai iene GaCLOrwns Gall 6 5 o 6 6 8 a 0 56 0 5 6 6 pity ING, 
Fore wing below : red band broader, reaching to about the apical sixth of cell 8, 
its inner edge deeply mdented at the end of the cell, its outer edge 
forming an almost regular curve from the apical fifth of the costa to 


the tornus latifascia, n. sp. 


! TAMBRIX SALSALA. 
Nisoniades salsala, Moore, P. Z.S. 1865, p. 786. 

Astictopterus salsala, Distant, Rhop. Mal. p. 401, pl. xxxiv. fig. 21 (1886). 

In typical females of this species there is on the underside of the fore wing a round 
white spot at the end of the cell and a curved postmedian series of roundish white 
spots, placed one each in cells 2-8 inclusive, and decreasing in size towards the 
costa. 

Hab. Sikkim (Moller); Assam, Naga hills, East Pegu, Perak (Doherty); Burmah 
(Adamson); Ceylon; Ganjam (Minchin); West Java (Piepers); Kina Balu ( Waterstradt). 


! TAMBRIX STELLIFER. 

Astictopteryz stellifer, Butler, Trans. Linn. Soc. Lond., Zool. ser. 2, vol. 1. p. 555 (1877) ; Moore, 
Lep. Cey. vol. i. p. 163 (1881). 

This is a good and distinct species, notwithstanding all that has been written to the 
contrary. In typical females there is on the underside of the fore wing a round 
white spot in the cell and three other roundish white spots placed as follows, namely : 
one near the basal third of cell 2, one near the basal third of cell 5, and one near 

VOL. xIv.—Part Iv. No. 11.—October, 1897. 2B 


182 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


the middle of cell 5, thus forming an oblique series running outwardly towards the 
apex of the wing. 
Hab. Fast Pegu, Pulo Laut, Sumatra (Doherty); Nias (Modigliani); Java (Fruhstorfer). 


! JAMBRIX SINDU. 
Astictopterus sindu, Felder, Wien. ent. Mon. iv. p. 401 (1860); Distant, Rhop. Mal. p. 402, 
pl. xxxv. fig. 30 (1886). 
Astictopterus obliquans, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 51 (1893). 


Hab. Perak, Pulo Laut (Doherty); Labuan, Kina Balu, Borneo (Waterstradt) ; 
Padang, Sumatra (Sachs, Martin). 


! TAMBRIX LATIFASCIA, n. sp. (Plate XXI. fig. 9, ¢ .) 


3 2. Slightly larger than J. sindu, from which it is distinguished by the different 
shape and much greater breadth of the red band on the fore wing. These differences 
are detailed in the table above. The male genitalia are similar to those of J. sindu. 

Expanse, ¢ 26, 9 29 mm. 

Hab. Kinu Balu, Borneo (Waterstradt) ; Liwa, S.E. Sumatra (Doherty). 


Described from specimens ex coll. Staudinger, one of which is now in my collection. 


GE. 
Ge, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 373. Type geta, de Nicév. 

A genus, for the present, containing one species only, allied to Matapa, from which 
it is best distinguished by the sex-marks of the male. The latter are as follows :—On 
the fore wing above, near the base of cell 2, an oval depression fringed on its inner edge 
by a series of erecto-patent hair-scales; the latter are blackish, becoming pale at the 
base, and when viewed from above appear to entirely cover the oval depression, the 
long axis of which reaches quite across the cell; on the hind wing above a fringe 
of long hair-scales along the basal half of vein 7, directed obliquely outward and 
downward, and covering wholly or in part a pale patch of modified scales. 


! GE GETA. 
Ge geta, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 374, pl. Q. fig. 51, 3. 


The female is undescribed. 
Hab. Pulo Laut (Doherty); East Java (Piepers); Selesseh, Sumatra (Martin). 


TARACTROCERA. 
Taractrocera, Butler, Cat. Lep. Fabr. p. 279 (1869) ; Watson, P. Z.S. 1898, p. 94, pl. iu. fig. 20. 
Type mevius, Fab. 
A genus of small species, associated on account of their general resemblance to the 
Hesperia mevius of Fabricius, as figured by Butler (Cat. Lep. Fabr. pl. iii. fig. 13). 


A REVISION OF THE ORIENTAL HESPERIIDA, 


183 


They may be readily distinguished by the blunt club of the antennz, which is excavated 
on the upperside. 
We know nine species, which we distinguish as follows :— 


6 


ww oo 


5 (4). 
(1). 
7 (14). 
8 (11). 
9 (10). 


(or) 


10 (9). 
11 (8). 
12 (13). 
13 (12). 
14 (7). 


15 (16). 


16 (15). 


5). 


(6). Pale markings of the upperside white or whitish. 
(3). 
(2). 
(5) 


Hind wing below with the veins conspicuously pale . 

Hind wing below with the veins not conspicuously pale. 

Hind wing below with a whitish te from the base to the 
termen in cell 14 y : 

No whitish stripe in cell 1 0 of feed wing talon ; 

Pale markings of the upperside deep tawny- or epileealia, 

Hind wing below with a pale spot near the middle of cell 7 

Fore wing below: cell brown, with a yellow spot near the apex. 

Size generally larger. Pale spots of upperside smaller, those of 
the hind wing above remote . : F 

Size smaller. Pale spots of upperside Tateer, those EE He tend 
wing above contiguous, forming an irregular macular extra- 
discal band CROC PUM a ee aut eral aay Ne 

Fore wing below : cell brown, with at least the distal half yellow. 

Hind wing below: cell 7 black, with an oe yellow spot in 
the middle 

Hind wing below : basal third of bail 7 wal a subensarate spot 
beyond the middle of that cell yellow . 

Hind wing below with no pale spot in cell 7. 

Yellow band on hind wing above not recurved towards the costa, 
ending at vein 6; fore wing below with the cell brown nearly 
to the middle, thence yellow . : : 

Yellow band on hind wing above ferurved (ouands the conta, 
ending at vein 7; fore wing below with the cell brown, with a 
yellow spot near the apex . 


! TARACTROCERA MVIUS. 


Hesperia mevius, Fabricius, Ent. Syst. 1. p. 352 (1793). 
Taractrocera mevius, Butler, Cat. Lep. Fabr. p. 279, pl. ii. fig. 13 ; Moore, Lep. Cey. vol. i. 

p- 172, pl. lxx. fig. 5 (1880-81) ; Watson, Hesp. Ind. p. 62. 
Pamphila sagara, Moore, P. Z. S. 1865, p. 792. 


N.W. Himalayas (Young, Hocking); Ganjam, Calcutta (Minchin); Ceylon 
(Mackwood); Burmah (Watson). 


Hab. 


*mevius, Fab. 


danna, Moore. 


ardonia, Hew. 


ceramas, Hew. 


nicevillet, Wats. 


ziclea, Plotz. 


flavoides, Leech. 


archias, Feld. 


oberthiiri, nu. sp. 


? Specimens from Ceylon have the hind wing below sordid green and the veins less conspicuously pale, 


232 


184 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! TARACTROCERA DANNA. 


Pyrgus danna, Moore, Cat. Lep. E. I. C. i. p. 249. 
Pamphila danna, Moore, P.Z.S. 1865, p. 508, pl. xxx. fig. 8. 
Taractrocera danna, Watson, Hesp. Ind. p. 63. 


Hab. N.W. Himalaya, 5000-9000 feet (Young); Sikkim (Elwes). 


! TARACTROCERA ARDONIA. 
Ancyloxypha ardonia, Hewitson, Descr. Hesp. p. 45 (1868). 
Hab. Kina Balu, Borneo (Waterstradt), Pulo Laut (Doherty). 


! TARACTROCERA CERAMAS. (Plate XXI. fig. 24, ¢ .) 
Cyclopides ceramas, Hewitson, Descr. Hesp. p. 44 (1868). 
Taractrocera ceramas, Watson, Hesp. Ind. p. 63. 


Hab. N. Canara (Aitken); Travancore, Malabar; Nilgiris (Minchin). 


! TARACTROCERA NICEVILLEI. (Plate XXI. fig. 12, ¢.) 
Tapactracera nicevillei, Watson, P.Z.S. 1893, p. 95. 

The form of the male genitalia is similar in this species and the preceding, and the 
essential wing-pattern of the two is also similar, the difference in facies being caused 
by the greater development of the yellow spots in 7’. nicevillei. We have not, however, 


seen any intermediate specimens. 
Hab. Bombay (Swinhoe). 


!'TARACTROCERA ZICLEA. 
Thymelicus ziclea, Plotz, Stett. ent. Zeit. xlv. p. 289 (1884). 
Pamphila mesoides (luzonensis, Mab.), Staudinger, Inis, il. pp. 145, 165 (1889). 
Ampittia ziclea, Semper, Schmett. Philipp. p. 304. 
Ampittia mesoides, id. t.c. p. 358. 
Pamphila luzonensis, Mabille, sec. spec. comm. 

This species and the next are interesting as combining the facies of Telicota dara, 
Koll., with the generic characters of Zaractrocera. After examining a male and female 
of ziclea and a male of mwsoides from Semper, we are satisfied that they represent but 
one species. There are some small differences of degree in wing-pattern, but having 
regard to the latitude of similar variation which obtains in insects having a similar 
facies (e.g. Telicota dara, Koll.) these differences cannot be taken as satisfactorily 
characterizing a species. 

Hab. Luzon, Mindoro, Samar, Bohol, Cebu, Mindanao (Semper). 


! TARACTROCERA FLAVOIDES. 
Taractrocera flavoides, Leech, Butt. China, &c. p. 590, pl. xl. fig. 10 9, 11 g (1893-94). 


Hab. Omei-shan, Central China (Pratt); Moupin (ex coll. Oberthiir). 


A REVISION OF THE ORIENTAL HESPERIIDA. 185 


! TARACTROCERA ARCHIAS. 

Pamphila archias, Felder, Sitzb. Ak. Wiss. Math. Nat. Cl. xl. p. 462, sep. p. 15 (1860). 
Thymelicus nigrolimbatus, Snellen, Tijd. Ent. xix. p. 165, pl. vii. fig. 5 (1876). 
Telicota nigrolimbata, Distant, Rhop. Mal. p. 384, pl. xxxv. fig. 16 (1886). 

According to a specimen received from M. Mabille, this is the same as Pamphila 
dschalia, Plotz. 

In the Felder collection, under the name “ Pamphila archias,” are four specimens ; 
two of these are without locality-labels and may be disregarded, but the remaining 
two which are labelled in contemporary handwriting “ Amboina, Dolesch,” are 
without doubt the same as Thymelicus nigrolimbatus, Snellen. 

Hab. Java (Piepers); Sambawa, Bali, Arjuno, Java (Doherty). 


! TARACTROCERA OBERTHURI, 0. sp. 

9. Upperside similar to that of 7. ziclea, but having the yellow markings less 
extensive. Hind wing below sordid yellow, by reason of a dense clothing of sulphureous 
scales on a dark brown ground; a large oblong pale spot occupying about the middle 
third of cells 4—5, a roundish one in the cell near the base of vein 7, another near the 
basal third of cell 2, and a feeble one in the base of cell 6; cell 14 pale. 

Hab. Trichinopoly, 8. India (Castets). 

Described from one female lent to us by M. Charles Oberthiir. 


Ocuus. 
Ochus, de Nicéville, Jour. As. Soc. Beng. 1894, p.51. Type subvittatus, Moore. 


This is a genus established by Mr. de Nicéville for the Cyclopides subvittatus of 
Moore. It is mainly distinguished from Baracus (e. g. B. septentrionum and B. subditus) 
by having the base of cell 6 in the fore wing nearly as long as that of cell 5, and vein 
11 not approaching vein 12. 


OcHUS SUBVITTATUS. 
Cyclopides subvittatus, Moore, P. Z.S. 1878, p. 692; Wood-Mason & de Nicéville, Jour. As. Soc. 
Beng. 1886, p. 392, pl. xvii. figs. 6,6a ¢, x2; Watson, Hesp. Ind. p. 69 (1891). 
Cyclopides subradiatus, Moore, 1. c. p. 693. 


Hab. Sikkim, Khasia (Hlwes), Nagas (Doherty). 


AMPITTIA. 
Ampittia, Moore, Lep. Cey. i. p. 171 (1881) ; Watson, P. Z.S. 1893, p. 95. Type maro, Fab. 

A group comprising a few small species, dark brown with yellow spots, associated on 
account of their general resemblance to the Hesperia maro of Fabricius. In the latter 
there is no evident apiculus to the club of the antennz, although a small one is present 
in the other species. The males of those species possessing a sex-mark on the fore 
wing above have the “‘tuning-fork” arrangement of the subcostal veins in the hind 
wing. We distinguish the species known to us as follows :— 


186 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


1 (4). Fore wing above with a sex-mark in the male. 
2 (3). Exp.25mm. Male: fore wing below—cell yellow except at the base. maro, Fab. 
3 (2). Exp. 30-32 mm. Male: fore wing below—cell yellow, with a black 
streakein: themmiddles, 43 se s-wen eye Wen (cmy-nn meee Leo elueechs 
4 (1). No sex-mark on fore wing above in the male. 
5 (6). Fore wing more pointed ; termen straight, oblique. Dark markings 
on hind wing below consisting of small, scattered, black spots . maga, Leech. 
6 (5). Fore wing less pointed, termen evenly rounded. 
7 (8). Hind wing below yellow, with irregular pale brown markings forming 
three more or less distinct transverse series ; a large, oblong, pale 
spot occupying the basal half of cells 4-5. . «. . «© . . .  trimacula, Leech. 
8 (7). Hind wing below yellow-brown by reason of a close sprinkling of 
yellow scales on a brown ground, with three yellow spots near the 
base and an irregular subterminal and postmedian series of yellow 
spots, the largest member of the latter being a spot near the 
middle of cells 4-5 and occupying the entire width of those cells . delai-lama, Mab. 


! AMPITTIA MARO. 
Hesperia maro, Fabricius, Ent. Syst., Suppl. p. 432 (1798), ¢. 
Cyclopides camertes, Hewitson, Descr. Hesp. p. 43 (1868), ¢. 
Ampittia maro, Moore, Lep. Cey. i. p. 172, pl. Ixxi. figs. 1, 1 @ (1880-81) ; Watson, Hesp. Ind. 
p. 61 (1891). 
Telicota maro, Distant, Rhop. Mal. p. 383, pl. xxxv. fig. 14 g,15 2 (1886). 
Hab. N. Canara (Aitken) ; Ceylon (Mackwood); Nilgiris (Roberts); Calcutta (de Nicé- 
ville); Burmah, Tenasserim (Watson); Shanghai (Pryer) ; Bali (Doherty). 


AMPITTIA MAROIDES. 
Ampittia maroides, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1896, p. 190, pl. T. fig. 51, 3. 
Differs from A. maro ‘on both sides of the fore wing in lacking the chrome-yellow 
spot in the middle of the submedian interspace.” 
Expanse 25 mm. 
Hab. Daunat Range, Tenasserim (fide de Nicéville). 
We have not seen this species. 


! AMPITTIA VIRGATA. 

Pamphila virgata, Leech, Entomologist, xxii. p. 47 (1890). 

Padraona virgata, Leech, Butt. China, &c. p. 598, pl. xl. fig. 15, g (1886). 
Hab. Changyang, Central China (Pratt). 


! AMPITTIA MAGA. 

Pamphila maga, Leech, Entomologist, xxiii. p. 48 (1890). 

Padraona maga, Leech, Butt. China, &e. p. 599, pl. xl. fig 18, ¢ (1894). 
Hab. Ichang, Ningpo (Leech). 


A REVISION OF THE ORIENTAL HESPERIIDA, 187 


AMPITTIA TRIMACULA. 
Taractrocera trimacula, Leech, Entomologist, xxiv. Suppl. p. 60 (1891). 
Padraona trimacula, Leech, Butt. China, &c. p. 599, pl. xl. fig. 17, g (1894). 


Hab. Wa-su-kow, West China (fide Leech). 


! AMPITTIA DELAI-LAMA. 
Cyclopides delai-lama, Mabille, Ann. Soc. Ent. Fr. 1876, p. lvi. 
Taractrocera lyde, Leech, Entomologist, xxiv. Suppl. p. 60 (1891). 
Aeromachus delai-lama, Leech, Butt. China, &e. p. 620, pl. xl. fig. 16, g (1894). 


Hab. West China, E. Tibet (Pratt, Kricheldorf). 


AEROMACHUS. 


Aeromachus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 214. Type stigmata, Moore. 


This is a group of small dull-coloured species associated together on account of 
their resemblance to the Thanaos stigmata of Moore. ‘The distinctive wing-pattern is 
generally well marked, and consists of a postmedian and subterminal curved series of 
small pale spots running almost parallel to the termen of both fore and hind wings: it is 
best displayed on the underside, and the postmedian series on the fore wing is usually 
the most sharply defined. In kali, de Nicév., the most aberrant species known to us,. 
the essential pattern is traceable, although the pale spots are purple instead of whitish, 
and the sex-mark of the male is of the same kind as that found in inachus. 

We separate the species known to us as follows :— 


1 (22). Hind wing below without purple markings. 
2 (9). Sex-mark on the fore wing above in the male a seam passing 
obliquely outwards from the middle of vein 1 @ to the base of 
vein 3. 
3 (8). Fore wing below with two rows of pale spots—one postmedian, the 
other subterminal. 
4 (7). Hind wing below: space between the postmedian and subterminal 
pale macular bands darkened and divided into a series of spots 
by the pale veins. 
5 (6). Hind wing below with the subterminal row of dark spots greenish 
brown; markings in cell 7 scarcely developed . . . . . . stigmata, Moore. 
6 (5). Hind wing below with the subterminal row of dark spots velvety 
greenish-black ; markings in cell 7 strongly developed. Fore 
wing longer in proportion to its width, its termen more distinctly 
CULV EOe ee aM e ge, sk oe eR Ty SS ALT Ue yo Sm CIRENSTS YN. SDs 
7 (4). Hind wing below uniform pale greenish grey, the veins not 
evidently paler than the disc, the space between the postmedian 


188 


14 (15). 
15 (14). 
16 (13). 


17 (18). 


21 (20). 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


and subterminal pale macular bands not darker than the 


remainder ... - - - 


. Fore wing below with only one row of pale spots—the postmedian 


one—and that but feebly developed. Hind wing below grey- 
brown, closely sprinkled with yellow scales, with a feeble 
irregular postmedian band of pale spots 


. Sex-mark on the fore wing above in the male a small longitudinal 


fold of pale scales on vein 1 a a little beyond the middle, or none. 


. Hind wing below with the veins pale. Tegumen (viewed from 


above) slightly angularly widened on each side just below its 
rounded apex ; lower lobe of clasp broadly rounded, its margin 
serrate and continuous with the outline of the upper lobe 


. Hind wing below with the veins concolorous. 
. Fore wing broader and more triangular, the termen straighter and 


longer in proportion to the dorsum. 


. Fore wing below: postmedian and subterminal pale macular bands 


both well developed, usually complete from the dorsum to the 
costa. Tegumen (seen from above) with a small notch at each 
distal angle, its apex truncate or but slightly rounded between 
the notches. 
Hind wing below grey-brown; the postmedian series of pale spots 
not stronger and more evident than the subterminal series . 
Hind wing below greenish ochreous; the postmedian series of pale 


piceus, Leech. 


musca, Mab. 


inachus, Mén. 


discreta, Plotz. 


spots stronger and more evident than the subterminal series . jhora, de Nicév. 


Fore wing below: pale macular bands imperfect or absent, not in 
any case reaching the dorsum. 

Fore wing below with a spot of pale scales in the cell near its 
upper distal angle. Tegumen (seen from above) with a small 
rounded projection on each side near the apex, beyond which it 
is produced in a bluntly-rounded triangle 


. No such pale spot on the fore wing below. ‘iegumen as in 


dubius, n. sp. 


discrela and jhora’ . . . . - . + +s +s + « - + gavanicus, 0. sp. 


. Fore wing narrower and more oblong; termen more strongly 


curved, almost evenly curved from the dorsum to the costa. 


. Fore wing below with the pale postmedian macular band continuous 


from cell 2 to cell 7 or 8, and running almost paralle) with the 
termen; no pale spot in the cell near its upper distal angle. 
Tegumen parallel-sided in its apical half, evenly rounded at the 
apex; lower lobe of clasp not produced, broadly rounded, 
serrate and continuous with the outline of the upper lobe 

Fore wing below with the pale spots in two outwardly oblique 
series—one series in cells 5, 6, and 7 or 8, or 6, 7, and 8, the 
other series in cells 2 and 3; no pale spot in cell 4; a pale 
spot in the cell near its upper distal angle. Tegumen a little 
constricted on each side just below the apex, which is bluntly 


indistincta, Moore. 


A REVISION OF THE ORIENTAL HESPERIIDA, 189 


triangular; lower lobe of clasp acuminate, produced much 

above the level of the upper edge of the clasp . . . . . . manus, Leech. 
22 (1). Hind wing below with pale purple spots. 
23 (24). Upperside fuliginous brown. Sex-mark on the fore wing of the 

male a small longitudinal fold of pale scales on vein 1 a about 

the middle. Purple markings on the dise of hind wing below 

forming a large irregular macular patch having its outer edge 


perallelawithathewtexmenyecmy.) Ce einen ves eee ele) kali, de Nicév. 
24. (23). Upperside pale brown. Sex-mark on fore wing of the male aseam 
passing obliquely outwards from the middle of vein 1a to the base 
of vein 3. Purple markings on disc of hind wing below forming 
a continuous curved macular band from the costa as far as 


Vari IL) oe SURO od GL Se SMR ice NA a eT Tamia 


! AEROMACHUS sTiGMaTA. (Plate XIX. fig. 3, 3.) 
Thanaos stigmata, Moore, P. Z. S. 1878, p. 694. 
Aeromachus stigmata, Watson, Hesp. Ind. p. 68 (1891). 

Specimens from the N.W. Himalayas and some from Sikkim show a band of 
whitish spots across the fore wing and a discal spot, which are faint or absent on the 
upperside of Khasia and Naga hill specimens; the undersides also vary to some 
extent, but the genitalia seem to be identical in both forms. 

Hab. N.W. Himalayas (Young); Sikkim (Mller); Khasias (Hiwes); Naga hills: 
(Doherty). 


! AEROMACHUS CHINENSIS, n. sp. 
Aeromachus inachus, Leech, Butt. China, &c. p. 619, pl. xl. fig. 19, g¢ (1893-94), in part. (nec 
Ménétriés). 
This insect, which in some parts of Western China appears to represent A. stigmata, 
may be readily distinguished from that species by the characters given in the table 
above. The genitalia do not differ materially. The sex-mark in the two male 


specimens which we have seems less conspicuous, but their condition is not perfect 
Hab. Wa-shan, Ta-tsien-lu, West China (Pratt). 


! AEROMACHUS PICEUS. 
Aeromachus piceus, Leech, Butt. China, &c. p. 618, pl. xl. fig. 16, g (1893-94), 
Hab. Moupin, E. Tibet (fide Leech). 


! AEROMACHUS MUSCA. 

Pamphila? musca, Mabille, Bull. Soc. Ent. Fr. 1876, p. xxvi. 

Aeromachus musca, Semper, Schmett. Philipp. p. 305 (1892). 
A very distinct little species, of the size of nanus, Leech, or indistincta, Moore. 
Hab, Luzon, Siargao (Semper). 

VOL. XIv.— Part Iv. No. 12.—Octoter, 1897. ZN 


190 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! AEROMACHUS INACHUS. (Plate XIX. fig. 7; Plate XXIII. figs. 36, 36 a.) 

Pyrgus inachus, Ménétriés, Bull. Acad. Petr. xvii. p. 217 (1859) ; Schrenk’s Reisen, p. 46, pl. iv. 
fig. 2 (1859). 

Closely allied to stigmata, but separable by its somewhat paler colour and the 
different form of the sex-mark. 

It has the wing-shape of chinensis. 

We have not been able to identify this with Ménétriés’s type except by the locality ; 
but Amur specimens agree well with those from Japan. Mr. Leech has included under 
this name what we call chinensis, and we have specimens from his collection which 
were mixed with others from Western China but which may be from Japan. 

Hab, Yokohama (Mazley); Oiwake, Japan (Pryer); Amurland, West China (coll. 
Leech). 


! AEROMACHUS DISCRETA. (Plate XIX. fig. 6, 2.) 
? Apaustus discreta, Plotz, Berl. ent. Zeit. 1885, p. 232. 


Hab. Khasia, 5000 feet (Elwes); Bernardmyo, Burmah (Doherty) ; Battak Mountains, 
Sumatra (Martin). Cf. javanicus, post. 


! AEROMACHUS JHORA. (Plate XXIII. figs. 37, 37 a.) 
Thanaos jhora, de Nicéville, Jour. As. Soc. Beng. 1885, p. 122, pl. il. fig. 12, 3. 


Hab. Sikkim (Miller). 


! AEROMACHUS DUBIUS, n. sp. (Plate XIX. fig. 10, 3; Plate XXIII. fig. 39 a.) 


3. Upperside dark olive-brown, the postmedian series of spots on the fore wing 
very indistinct or wanting; sex-mark on the fore wing a small inconspicuous fold 
of raised scales in cell 1@next vein la@ near the middle. Underside: fore wing 
umber-brown, the apical and costal regions similar to the hind wing; a small pale 
spot near the upper distal angle of the cell; the postmedian series of pale spots 
represented by about six spots, which become gradually fainter in their course from 
cells 8-3; a pale anteciliary line: hind wing ochreous brown by reason of a close 
sprinkling of yellow scales on the umber-brown ground; the postmedian and sub- 
terminal series of pale spots very indistinct or wanting; a pale anteciliary line. 
Fringes greyish white, the short scales grey-brown. Antenne above blackish, 
minutely spotted with yellowish: beneath greyish yellow, the shaft spotted with 
black. Second joint of palpi clothed with yellow and black hairs intermixed. 
Body above concolorous with the wings, beneath with greyish-yellow pubescence. 
Legs with greyish-yellow scaling. 

Tegumen (seen from above) with a small rounded projection near the apex on. 
each side, beyond which it is produced in a bluntly rounded triangle. 


A REVISION OF THE ORIENTAL HESPERIIDA. 191 


@. Differs from the male in wanting the sex-mark and in having the pale spots 
proper to the underside more strongly developed, and consequently the postmedian 
series appears faintly on the costal portion of the upperside of the fore wing. 

Expanse 21-214 mm. 

Hab. Palnai hills (Castets) ; Peermaad, Travancore. 

Described from four males and one female. Of the species known to us this 
comes nearest to A. jhora, de Nicéy., from which, however, it differs in the form 
of the tegumen as well as in the spotting of the wings. 


! AEROMACHUS JAVANICUS, n. sp. (Plate XIX. fig. 24, ¢.) 


3. Upperside pale umber-brown without markings, a few greyish-yellow scales 
near the base of the fore wing towards the costa; on the hind wing a line of long 
grey hairs along the basal two-thirds of vein 14; sex-mark on the fore wing a 
small inconspicuous fold of raised scales in cell 1@ next vein la near the middle. 
Underside: fore wing pale umber-brown ; the apical region similar to the hind 
wing, and | aving the usual postmedian series of pale spots indicated by very indistinct 
pale spots in cells 5-7 or 5-8; a pale anteciliary line: hind wing grey-brown, with 
a very faint greenish tinge by reason of the close sprinkling of yellowish-grey 
scales on the pale brown ground; a pale anteciliary line; the usual postmedian 
and subterminal series of pale spots are only indicated by a few very indistinct pale 
spots. Fringes pale grey, the shorter scales grey-brown. Antenne above black, 
minutely spotted with yellowish; beneath greyish yellow, the shaft spotted with 
black. Second joint of palpi clothed with whitish-grey and black hairs intermixed. 
Body above concolorous with the wings, beneath with yellowish-grey pubescence. 
Legs with yellowish-grey scaling. 

Tegumen as in A. discreta and A. jhora. 

Expanse 21 mm. 

Hab. West Java. 

Described from a single male kindly sent to us by Herr P. C. T. Snellen with 
the name Apaustus discreta, Plotz. In general appearance it comes nearest to 
the insect from the Khasia hills and Burmah, which we identify as discreta, Plotz, 
originally described from India; but the latter species, in all the specimens which 
we have seen, has the pale pattern on the underside well marked and constant. 


! AFROMACHUS INDISTINCTA. (Plate XIX. fig. 11, ¢; Plate XXIII. fig. 39.) 


Thanaos indistincta, Moore, P. Z. 8. 1878, p. 694. 
Aeromachus indistincta, Watson, Hesp. Ind. p. 66 (1891). 


Hab. Nilgivi hills (Hampson); Tavoy (Tucker); Tounghoo, Burmah (Adamson) ; 


Bernardmyo (Doherty) ; ? Khasias. 
202 


192 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! AFROMACHUS NANUS. (Plate XXIII. figs. 38, 38 a.) 
Aeromachus nanus, Leech, Butt. China, &ec. p. 620, pl. xl, fig. 21, g (1893-94). 


Hab. Shanghai (Pryer) ; Ichang (Pratt). 


! AFROMACHUS KALI. 
Thanaos kali, de Nicéville, Jour, As. Soc. Beng. 1885, p. 123, pl. ii. fig. 3, g. 


The largest of the genus and a very distinct species. 
Hab. Sikkim (M@ller) ; Naga hills, Bernardmyo (Doherty). 


AEROMACHUS CATOCYANEA. 


Pamphila catocyanea, Mabille, Ann. Soc. Ent. Fr. 1876, p. lv. 
Aeromachus catocyaneus, Leech, Butt. China, &e. p. 618 (1894). 


We have relied on the drawing of Mabille’s type of this species referred to by 
Leech in separating this species, which we have not seen, but which seems nearest 
to A. kali. 

Hab. “ Thibet” (David). Type in Mus. Paris. 


SEBASTONYMA. 
Sebastonyma, Watson, P. Z. S. 1898, p. 81. Type dolopia, Hew. 


A monotypic genus closely allied to Halpe. Vein 5 of the fore wing is 
straight and arises very little nearer to vein 4 than to vein 6, and the sex-marks 
in the male consist of an infra-alar tuft on the fore wing and a large suboval 
patch of androconia near the base of the hind wing above, situate in the cell next its 
lower edge and occupying about one-half of its area. The sex-mark on the hind 
wing is not noticed in the original diagnosis of the genus. 


SEBASTONYMA DOLOPIA. 
Hesperia dolopia, Hewitson, Descr. Hesp. p. 27 (1868) ; id. Ex. Butt. v. pl. lv. figs. 60, 61 (1873). 
Halpe dolopia, Watson, Hesp. Ind. p. 74 (1891). 


Hab. Sikkim (Méller) ; Nagas, E. Pegu (Doherty); Khasias (Hamilton). 


PEDESTES. 
Pedestes, Watson, P. Z. S. 1893, p. 81. Type masuriensis, Moore. 
Watson made IJsoteinon masuriensis, Moore, the type of this genus, and he also 
included in it J. pandita, de Nicév. 


The male genitalia in these two species are remarkable for a certain amount 
of asymmetry and the possession of separate edeagus-quards, not part of the tegumen, 


A REVISION OF THE ORIENTAL HESPERIIDA. 193 


of which the right is larger than the left. The tegumen is symmetrical in both 
species, and the clasps are so in pandifa; but in the clasps of maswriensis the 
asymmetry is extreme. 
The species here included in the genus may be distinguished as follows :— 
1 (4). Fore wing with hyaline spots in cells 6, 7, 8. Hind wing 
below not marked with about ten black spots. 
2 (3). Upperside dark brown with a purple shade, pale spots on the 
fore wing pure white. Hind wing below grey-brown, generally 


with a minute pale spot in each of cells 2 and 3; fringe white. . masuriensis, Moore. 
3 (2). Upperside brown, pale spots on the fore wing sordid yellowish 
white. Hind wing below yellow-brown ; fringe brownish grey . pandita, de Nicéy. 


4 (1). Fore wing without hyaline spots in cells 6, 7, and 8, or with one in 

cell 6 only. Hind wing below marked with about ten black 

spots. 
5 (6). Fore wing with no pale spot in cell 6. Fringes of hind wing below 

pale, with dark spots next the veins. Male with the long hairs 

near the base of the hind wing above normal, not forming a tuft. 

Apex of tegumen (viewed from above) elongate-triangular, with 

a short horn on each side of the base. . . . . . . . . +. maculicornis, n. sp. 
6 (5). Fore wing with a pale spot in cell 6. Fringes of hind wing below 

pale with the basal half dark. Male with a distinct tuft of 

subequal Jong hairs near the base of hind wing above. Apex of 

tegumen (viewed from above) elongate-triangular, with an ear- 

like lobe on each side of the base . . . . . =.=. =. =. ~~ ~~ fuscicornis, n. sp. 


PEDESTES MASURIENSIS. (Plate XXIV. figs. 40, a, 4, c.) 
Isoteinon masuriensis, Moore, P. Z. S. 1878, p. 693; Watson, Hesp. Ind. p.79; Staudinger, Exot. 
Tagf. 1. p. 301, 1. pl. e. 
Hab. N.W. Himalaya (Young) ; Sikkim (Moller). 


PEDESTES PANDITA. (Plate XXIV. figs. 41, a, 0.) 
Isoteinon pandita, de Nicéville, Jour. As. Soc. Beng. 1885, p. 181, pl. 1. fig. 14, 2 ; Watson, 
Hesp. Ind. p. 81. 


Hab. Sikkim (Méller) ; Naga hills (Doherty). 


PEDESTES MACULICORNIs, n. sp. (Plate XVIII. fig. 23,4; Plate XXIV. fig. 42.) 


¢. Upperside dark umber-brown: fore wing with four yellowish-white hyaline 
spots—one large, occupying the prebasal fifth of cell 2, one occupying the prebasal 
fifth of cell 3, and two in the cell, one on the lower edge and next the base of 
cell 3 and one on the upper edge near the base of cell 9; a streak of golden-yellow 
scales along the costa from the base to near the middle and another along the 
upperside of vein la from the base to beyond the middle. Underside: fore wing 


194 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


with the discal area brown, apical area brownish grey; hyaline spots as on the 
upperside ; a large oblong suffused pale spot in cell 1a, and a small brown spot 
in each of cells 4-7: hind wing brownish grey, with nine roundish black spots 
placed as follows—one in the basal fourth of cell 7, one in the cell next the 
base of cell 6, one near the basal third of cell 14, one near the middle of cell 7, 
one in cell 6, one in cells 4-5, one each in cells 2 and 3, and one near the apical 
third of cell 14, the six last named forming a subterminal curved series; fringe pale, 
with dark spots next the veins. 

Body above concolorous with the wings. Palpi clothed with yellowish-grey scales 
intermixed with black ones. Antenne brown, spotted with white on the underside 
from the base to the club, which bears a white ring. 

g. Similar to the male. 

Expanse 37-38 mm. 

Described from three males and one female in coll. Elwes. 

Hab. Pulo Laut (Doherty). 


PEDESTES FUSCICORNIS, n. sp. (Plate XVIII. fig. 25,3; Plate XXIV. fig. 43.) 

g. Upperside dark umber-brown: fore wing with five yellowish-white hyaline 
spots—one, the largest, in cell 2 and reaching from the basal fifth to about the 
middle of that cell, one occupying the prebasal fourth of cell 3, one, punctiform, 
near the basal third of cell 6, and two in the cell, one on the lower edge and next 
the base of cell 3 and one on the upper edge near the base of cell 9; the wing- 
membrane on each side of vein 1@ from the base to the middle paler in colour 
and having a somewhat inflated appearance: hind wing with a tuft of subequal 
long brown hairs near the base and reaching nearly to the middle. Underside: 
fore wing brown, the hyaline spots as on the upperside; a large oblong suffused 
pale spot in cell la, and traces of small dark spots one each in cells 4-8, most 
distinct in cells 4 and 5: hind wing brown, with a distinct purple shade and nine 
black spots placed as follows :—one in the basal fourth of cell 7, one in the cell next 
the base of cell 6, one near the basal third of cell 10, one near the middle of cell 7, 
one in cell 6, one in cells 4—5, one each in cells 2 and 3, and one near the apical 
third of cell 1 4, the six last named forming a submarginal curved series ; fringe pale, 
the basal half (that is, the short scales) dark. 

Body above concolorous with the wings. Clothing of palpi above dark brown, 
beneath of yellowish-grey and brown scales intermixed in nearly equal numbers. 
Antenne brown, underside spotted with white near the base; club with a white ring. 

ixpanse 30-34 mm. 

Described from three male specimens in coll. Elwes, in one of which the purple 
shade is absent from the underside of the hind wing. 


Hab. Pulo Laut (Doherty). 


A REVISION OF THE ORIENTAL HESPERILIDA. 195 


LoPHOIDES. 


Lophoides, Watson, P. Z. S. 1893, p. 84. Type apis, de Nicé. 


Antenne two-thirds as long as the costa; club moderate, gradual; apiculus gradual. 
Palpi suberect, third joint short, almost concealed. Fore wing pointed, dorsum subequal 
in length to the termen; vein 4 straight, practically intermediate between veins 4 and 6; 
vein 2 from the basal third of the lower edge of the cell (further from the base of the 
wing in the female) ; base of cell 3 subequal in length to that of cell 4; hyaline spot in 
cell 4 reduced to a point or wanting. Hind wing with the termen evenly rounded, cell 
about half as long as the wing ; vein 3 arising immediately before the end of the cell, 
vein 2 just beyond the middle of the lower edge of the cell. 

In the male the dorsum of the fore wing has in its middle third a scanty fringe 
of long hairs pointing obliquely downward and outward; on the underside of the 
fore wing the middle third of the dorsum bears a thick fringe of long hairs pointing 
obliquely upward and outward ; on the upperside of the hind wing there is a patch of 
long recumbent hairs attached to the basal portion of vein 8; and the fringe of the 
hind wing next the tornus is much elongated. 

The first described species belonging to this genus was obscura, Distant, a species for 
which that author made the genus /sma; but as his account of that genus is both 
imperfect and inaccurate (he ascribes to the type a pyriform antennal club, which it 
certainly does not possess), his name must sink in favour of Lophoides, Watson, of which 
iapis, de Nicéy., is the type, and of which the diagnosis does include a statement of the 


differential characters of the type species. 


1 (8). With at least one cell-spot on the fore wing, that next the lower 
edge of the cell. 

2 (7). Fore wing with the pale spot in cell 2 small, higher than wide (at 
least in the male). 

3 (6). Hind wing below without any purplish suffusion on the basal half. 

4 (5). Hind wing witha sprinkling of yellowish-grey scales, sometimes by 
their absence in parts giving rise to the appearance of a very 
obscure dark spot next the transverse vein, and a postmedian series 
of four or five similar spots. Fore wing broader, termen subequal 
in length to the dorsum. Upper edge of lower lobe of clasp not 
serrate ; cedeagus-guards toothed on the edges. . . . . . . tapis, de Nicév. 

5 (4). Hind wing below with a feeble brown spot next the transverse vein 
and a postmedian series of four similar spots, one each in cells 2, 
3, 4-5, and 6 respectively. These spots are of purplish-brown 
scales and not merely bare patches of the ground-colour of the 


wing. Fore wing longer and narrower, termen longer than the 


dorsum obscura, Distant. 


196 MESSRS. H. J. ELWES AND JAMES EDWARDS : 


6 (3). Hind wing below with a feeble purple suffusion on its basal half. 
Upper edge of lower lobe of clasp serrate ; cedeagus-guards smooth 


ontheedges. . . c 2 : . purpurascens, 1. sp. 
7 (2). Fore wing with the pale Soot in een 2 eee, vie ‘Hart AEN Male 
genitalia as iniapis  . . : Peace Po phous speUulso viable 


8 (1). No cell-spot on the fore wing. Hind wing tele with two minute 
white points on the disc, one each in cells 2and3 .... . binotatus, n. sp. 


! LopHomEs IAPIS. (Plate XXIV. figs. 44, 44 a.) 


Isoteinon iapis, de Nicéville, Jour. Bom. Nat. Hist. Soc. 1890, p. 218, pl. E. fig. 9, 5; Watson, 
Hesp. Ind. p. 86. 


Hab. Johore, Malay Peninsula; Mergui Archipelago (fide de Nicéville) ; Pulo Laut, 
Borneo; Perak (Doherty); Banguey (coll. Staudinger). 


LOPHOIDES OBSCURA. 
Isma obscura, Distant, Rhop. Mal. p. 386, pl. xxxv. fig. 19 (1886). 


We are indebted to Messrs. Godman and Salvin for the opportunity to critically 
examine the type of this species, which is from Singapore ( Wallace). 


! LOPHOIDES PURPURASCENS, n. sp. (Plate XVIII. figs. 26 ¢, 279; Plate XXIV. 
figs. 45, 45 a.) 


Distinguished from Z. iapis by the purple suffusion of the basal half of the hind 
wing below. 

Expanse 31 mm. 

Hab. Pulo Laut, Borneo (Doherty). 

Described from three males and one female in coll. Elwes. 


! LopHoIDES vULSO, Mab. 
Pamphila vulso, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 55 (1893). 


Hab. West Java (Piepers); Java (Fruhstorfer); Bali (Doherty) ; Selesseh, Sumatra 
(Martin). 


LOPHOIDES BINOTATUS, n. sp. (Plate XVIII. fig. 28, ¢ .) 


°. Upperside dark brown : fore wing with five hyaline white spots, one about one and 
a half times as high as wide in cell 2, one less than half the size of the former in cell 3, 
one very small in cell 4, and two small and punctiform, one each in cells 6 and 7, anda 
small white spot next the upper edge of vein 1 @ near the middle ; hind wing with an 
indistinct pale point on the disc, coinciding with that in cell 3 on the underside. 
Underside brown: fore wing darker on the disc, with a suffused whitish spot near the 
middle of cell 1a, and the hyaline spots as on the upperside; hind wing with two 


A REVISION OF THE ORIENTAL HESPERIIDA. UN/ 


minute white points on the disc, one each in cells 2 and 3. Fringe of the fore wing 
grey-brown, a little paler next cell 1@; of the hind wing dark grey, with the short 
scales dark brown and a feeble indication of chequering. Antenne blackish, finely 
spotted with white on the underside; club white beneath, apiculus dark. Body above 
concolorous with the wings. Second joint of palpi clothed with yellowish and black 
hairs intermixed. Clothing of breast and legs brown. Abdomen beneath yellowish 
grey. 

Expanse 30 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 


Described from one example ex coll. Staudinger. 


HYaroris. 
Hyarotis, Moore, Lep. Cey. i. p. 174 (1881). Type adrastus, Cr. 
This is a monotypic genus established by Moore for Hesperia adrastus, Cr., an insect 
sufficiently distinguished by its facies. Vein 5 of the fore wing is straight throughout. 


HIYAROTIS ADRASTUS. 


Hesperia adrastus, Cramer, Pap. Ex. vol. iv. pl. cccxix. figs. F, G (1780). 
Plesioneura praba, Moore, P. Z. S. 1865, p. 790. 
Hesperia phenicis, Hewitson, Ex. Butt., Hesp. pl. iv. figs. 36, 37 (1869). : 
Hyarotis adrastus, Moore, Lep. Cey. vol. i. p. 174, pl. lxvii. figs. 5, 5 a (1881) ; Distant, Rhop. 
Mal. p. 397, pl. xxxiv. fig. 4 (1886) ; Watson, Hesp. Ind. p. 117. 
Hab. Kangra (Hocking); Sikkim (Moller); HE. Pegu (Doherty); Ceylon; Java 
(Fruhstorfer) ; Philippines (Semper); Palawan (Platen). 


IsorEINon. 
Isoteinon, Felder, Wien. ent. Monats. vi. p. 30 (1862); Watson, P.Z.S. 1893, p. 83. Type 
lamprospilus, Feld. 

Watson restricts the name Jsoteinon to the lamprospilus of Felder and diagnoses the 
genus by the direction of the third joint of the palpi, which is “ erect, reaching well 
above the vertex of the head, slender, obtusely conical.’ We have not seen any 
specimens of lamprospi/us, Feld., with palpi of this kind; for us, the third joint of the 
palpus is short, about equal in length to one-fourth of the diameter of the eye, erecto- 
patent, and not reaching to the level of the vertex; and this condition does not appear 
to be due to deflexion of the head in setting. ‘The basal portion of vein 5 of the 
fore wing recedes from vein 6. 


IsorEINON LAMPROSPILUS. 
Tsoteinon lamprospilus, Felder, Wien. ent. Mon. vi. p. 30 (1862) ; id. Reise Nov., Lep. iii. pl. Lxxiy. 
fig. 20 (1867) ; Leech, Butt. China, &e. p. 582 (1893-94). 
VOL. Xiv.—PartT tv. No. 13.—October, 1897. Dip 


198 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Pamphila vitrea, Murray, Ent. Mo, Mag. xi. p. 171 (1875). 
Pamphila lamprospilus, Pryer, Rhop. Nihon. p. 33, pl. x. fig. (1889). 


Hab. Japan (Pryer, Leech); Changyang, C. China (Pratt). 


Ipwon. 
Idmon, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 375. Type “wnicolor,” de Nicév. (nec Dist.) 

This is a genus made by Mr. de Nicéville for a species from Perak, of which the 
female, according to him, agrees very closely with Distant’s figure of Baoris unicolor. 
The latter is an entirely brown insect with an expanse of about 30 mm., and, as we have 
satisfied ourselves by an examination of the type kindly lent to us by Dr. Staudinger, is a 
true Parnara, and a male, not a female as supposed by Mr. de Nicéyille. In the genus 
Idmon, according to his description, “ the middle and lower discocellulars of the fore 
wing are very upright, slightly inwardly oblique only, both slightly concave, the middle 
a very little longer than the lower, the second median nervule arises close to the 
lower end of the cell, and the first median arises a little nearer to the base of the wing 
than to the lower end of the cell.” There is on the fore wing of the male a sex-mark, 
which is described by him first as “a sexual brand placed anterior to, but against the 
submedian nervure towards its base, this brand is narrow and raised,” and afterwards as 
“a narrow raised brand of modified scales lying alongside a portion of the submedian 
nervure towards its base on the upperside of the fore wing.” 

The hind tibia has “a tuft of hairs attached to its proximal and two pairs of spines 
on its distal end.” ‘This latter character is not found in any species of Hesperiide 
known to us; and as Mr. de Nicéville says that the female of his /dmon differs from the 
male only in the wings being broader and lacking the sexual brand on the fore wing, 
it would appear that there are two pairs of spines on the distal end of the hind tibia in 
both sexes. 

Mr. de Nicéville gives as the type of this genus an insect which he was unable 
to identify correctly without seeing, and his generic diagnosis relates to the species 
mentioned below, which we have never seen, and not to Baoris unicolor, Dist. 


IpMoNn UNICOLOR. 
Idmon unicolor, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 377, pl. Q. fig. 53, ¢. 


Hab. Perak ( fide de Nicéville). 


ARNETTA. 
Arnetta, Watson, P. Z. 8. 1893, p. 81. Type atkinsoni, Moore. 


Watson includes in this genus /soteinon atkinsoni, Moore, and J. vindhiana, Moore ; 
the former has veiu 5 in the fore wing straight and very little nearer to vein 4 than to 


A REVISION OF THE ORIENTAL HESPERIID. 199 


vein 6, and in the latter the basal portion of vein 5 in the fore wing recedes from 
vein 6, so that vein 5 arises much nearer to vein 4 to vein 6. ‘The palpi are porrect 
in both species. 

The two species may be distinguished as follows :— 


1 (2). Male with a tuft of hair near the middle of the dorsum of the 

fore wing below. Vein Ja of fore wing deflexed to the dorsum 

near the middle. Hind wing below with small pale spots . . . atkinsoni, Moore. 
2 (1). No such tuft in the male, vein 1 a of fore wing straight. Hind wing 

below divided into a pale basal and a dark apical area, the 

boundary between these being in a line from the apical third of 

vein 8 to the apical fourth of vein 16 and somewhat angulated 

rho) fore epee? Gl Beg none Som ea 6 6 6 ce 6 ONCHTTO, Milo: 


! ARNETTA ATKINSONI. 
Isoteinon atkinsoni, Moore, P. Z. 8. 1878, p. 693, pl. xlv. fig. 10; Elwes, Trans. Ent. Soe. Lond. 
1888, p. 455, pl. xi. fig. 9,¢; Watson, Hesp. Ind. p. 77, 
Isoteinon subtestaceus, Moore, t.c. p. 844; Watson, t.c. p. 78. 
Isoteinon khasianus, Moore, t. c. p. 693 ; Watson, t.c. p. 78. 


Hab. Sikkim (Elwes); Khasias (Hamilton); Nagas, Burmah (Doherty). 

Atkinsoni varies in the ground-colour of the hind wing below and the development 
of the pale spots thereon; in typical examples the latter are well marked, but they are 
sometimes entirely absent. and many intermediate states occur: the unspotted or faintly 
marked examples (subtestaceus, M.) were regarded by Moller as the spring brood, and 
Watson got atkinsoni only in August and sudbtestaceus only in April. Watson says 
(P. Z. 8. 1893, p. 82) that in the males of atkinsoni and subtestaceus there is attached 
along the basal half of the inner margin of the fore wing a tuft of longish hairs, which 
are turned up and spread out fanwise over the underside of the hind wing. In the 
male specimens of atkinsoni before us there is on the dorsum of the fore wing, 
from a point distant about one-fourth from the base of the wing to a point about the 
middlle of the dorsum, a fringe of long black hairs, which form a tuft pointing 
in the direction of the tornus and covering the middle portion of vein 1 a, which is 
deflexed so as to touch the dorsum, and is there thickened and channelled. 


! ARNETTA VINDHIANA. (Plate XVIII. fig. 24, 3.) 


Tsoteinon vindhiana, Moore, P. Z. S. 1883, p. 533; Watson, Hesp. Ind. p. $4. 
Isoteinon nilgiriana, Moore, |. c.; Watson, l. ec. 
Isoteinon modesta, Moore, t. c. p. 534; Watson, 1. c. 


Hab. Jubbulpore (fide Moore) ; Nilgiris (Hampson) ; Mhow, Matheran (jide 
Swinhoe); Travancore (in coll. Rothschild). 
2n2 


200 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Irs. 
Itys, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 377. Type iadera, de Nicé. 


The following are the chief characters given by Mr. de Nicéville for this genus and 
are taken from the male:—Antenne more than half as long as costa, club elongate 
slender, apiculus short. Palpi porrect, densely hairy, third joint almost concealed. 
Fore wing: dorsum considerably longer than the termen, vein 5 rather nearer 4 than 6, 
vein 2 a little nearer to the end of the cell than to the base of the wing; dorsum 
on the underside with a long tuft of hair-scales directed upwards and outwards. 
Hind wing much longer than broad. Abdomen slender, reaching to the tarsus of 
hind wing. Tibial epiphysis present. Hind “tibia with a single pair of long spines 
at its distal end.” 

Table of Species. 
] (2). Entire insect shining brownish fuscous. . . . . . . . iadera, de Nicéy. 
2 (1). Uppersidedark vandyke-brown suffused with purple. Fore wing 
below with four whitish hyaline spots, one each in cells 2, 
3, 6, and 7, and a geminate cell-spot which is twice as high 
as wide. Hind wing below vandyke-brown, suffused with 
purple, with indistinct traces of three dark bands, one sub- 
basal and two postmedian, and not more than four small 
ochreous-white spots, of which latter, according to the 
figure, there are two in cell 7, and one in the cell near its [& de Nicéville. 
upperoutersangle) 9-092 -) fh ney fee 0) a eee ecrosiielum, Wood-Mason. 


Irys IADERA. 
Itys iadera, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 379, pl. Q. fig. 52, ¢. 
Hab. Penang, N.E. Sumatra (de Nicéville). 


ITYs MICROSTICTUM. 


Isoteinon microstictum, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 385, pl. xvii. 
figs. 3 d,3a@ 9; Watson, Hesp. Ind. p. 82. 


Hab. Sileuri, Cachar (Wood-Mason & de Nicéviile). 


ZOGRAPHETUS. 
Zographetus, Watson, P. Z. 8. 1893, p. 84. Type satwa, de Nicév. 

Species of small size, without markings on the hind wing above; no hyaline spot in 
cell 4 of the fore wing; alar sex-mark when present not of the kind found in Halpe. 
Basal portion of vein 5 of the fore wing decurved. ‘The latter point is less evident ir. 
satwa than in ogygia and the other species here included in the genus. Antenne 
about half as long as the costa (somewhat exceeding that length in the male of ogygia), 
the club and apiculus moderate and gradual. Palpi ascending, third joint short. 
Hind tibiz with two pairs of spurs. 


1 (10). 
2 (5), 
13: 
4 (8) 
5 (2) 
6 (7) 
7 (6) 
8 (9) 
9 (8). 
10 (1). 


A REVISION OF THE ORIENTAL HESPERIIDA. 


The species known to us may be distinguished as follows :— 


The largest or only pale spot in the cell of the fore wing placed next 
the lower edge of the cell. 

Hind wing below yellow or greenish yellow in the basal half, purple 
or brownish purple in the terminal half. 

Ground-colour of the fore wing above plain brown. Fore wing in the 
male with a distinct ridge or fold on each side of the prebasal fourth 
of vein 1 @ and on each side of the basal third of vein 2. Hind wing 
below bright yellow, the termen broadly brownish purple 


. Ground-colour of the fore wing above brown with a dark purple 


shade. No sex-mark on the fore wing of the male. Hind wing 
below greenish yellow, the termen broadiy purple-brown . 


. Hind wing below not as in paragraph 2. 
. Male: fore wing above with a tuft of long hair-like scales near the 


middle of thedorsum. Hind wing below yellow; termen narrowly 
brown from the tornus as far as vein 7; a brown spot next the 
transverse vein, a smaller one in cell 5, and three small subcon- 
tiguous brown rings, one each in cells 16, 2, and 3 


. No tuft of long hair-scales on the dorsum of the fore wing in the male. 
. Hind wing below brown, with several suffused patches of yellow forming 


two series, a median and subterminal, the latter being the more 
distinct ; about six more or less indistinct suffused dusky spots, one 
near the middle of cell 7, one near the middle of the upper edge of 
the cell, and a postmedian series of about four commencing in 
cell 2 and running parallel to the termen oF 

Hind wing below red-brown, with about seven auffased at purple- 
brown spots, one near the middle of cell 7, one near the middle of 
the upper edge of the cell, and a postmedian series of five others 
placed one each in cells 16, 2, 3, 4-5, and 6, and ee ae 
to the termen 

The largest or only pale spot in ats all of the fare wing placed next 
the upper edge of the cell. Hind wing below yellow, with a nearly 
straight postmedian series of five oblong brown spots passing from 
just beyond the middle of vein 14 to just beyond the middle of 
yein 6, and a more or less distinct suffused dusky terminal band , 


ZOGRAPHETUS SATWA. 


201 


satwa, de Nicév. 


durga, Plotz. 


flavalum, de Nicév. 


ogygia, Hew. 


ogygioides, u. sp. 


auriferus, 1. Sp. 


Isoteinon satwa, de Nicéville, Jour. As. Soc. Beng. 1883, p. 86; Watson, Hesp. Ind. p. 79. 


Hab. Sikkim (Elwes); E. Pegu, Perak (Doherty). 


! ZOGRAPHETUS DURGA. 


Apaustus durga, Plotz, Stett. ent. Zeit. xlv. p. 153 (1884). 


TIsoteinon durga, Semper, Schmett. Philipp. p. 306, pl. xlix. fig. 10, § (1892). 


? See next page, under flavalum. 


202 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Resembles J. satwa, de Nicév., so closely that the differences only need be noticed. 
Fore wing with a dark purple shade throughout ; no pale spot in cell 7, no modification 
of veins 1 and 2 by way of sex-mark ; the yellow colour on the hind wing below more 
extensive but much less bright; club of antenne pure white beneath. 

Hab. Camaguin de Mindanao (Semper); Sambawa (Doherty). 


ZOGRAPHETUS FLAVIPENNIS. 

Tsoteinon flavipennis, de Nicéville, Jour. As. Soc. Beng. 1885, p. 122, pl. u. fig. 4, 9; Watson, 
Hesp. Ind. p. 81. 
Zographetus flavipennis, Watson, P. Z. 8. 1893, p. 85. 

The hind wing below is described as “ ferruginous-ochreous, glossed and marbled with 
purple,” and as having “a dark brown spot in the cell and a series of five or six 
similar spots placed around the cell.” ‘The figure shows the termen of the hind wing 
below rather broadly and suffusedly brown. 

Col. Swinhoe has kindly lent us an insect from Sikkim (Monro), labelled by Mr. de 
Nicéville “ Zsoteinon flavipennis, de Nicéville, 2 ,” which agrees fairly well with the 
figure above cited. The underside of the hind wing of this specimen is dull red- 
yellow, darker next the termen, with seven dark purple-brown spots, one near 
the middle of cell 7, one near the middle of the upper edge of the cell, and a 
postmedian series of five others placed one each in cells 10, 2, 5, 4-5, and 6, and 
running parallel to the termen. The fore wing is remarkable for the large size of the 
hyaline spot in cell 6, which is oblong and nearly as large as that in cell 3; the one in 
cell 7 is punctiform. In cell 1a@ on the fore wing above there is a yellowish-white spot 
next vein 1 @ beyond the middle. 

Hab. Buxa, Bhutan (Moti Ram); Sikkim (Méller); South Andaman Island (de 
Roepstorff). All fide de Nicéville. 


ZOGRAPHETUS FLAVALUM. 

Tsoteinon flavalum, de Nicéville, P. Z. 8. 1887, p. 463, pl. xl. fig. 10, ¢ ; Watson, Hesp. Ind. p. 83 
(1891). 

Hab. Silkkim. 

Having regard to the statement in the original description of this species that 
there are “no secondary male sexual characters,” it is well to note that in the 
type specimen (kindly lent to me by Mr. Rothschild) there is a tuft of long pale hair- 
scales occupying the middle third of the dorsum of the fore wing above directed very 
obliquely outward and upward and reaching as far as vein la. It is, however, just 
possible that this tuft may have been misplaced in setting the insect, and that its 
normal position is on the underside of the fore wing, as in some other species. 


A REVISION OF THE ORIENTAL HESPERIIDA. 203 


! ZOGRAPHETUS OGYGIA. 
Hesperia ogygia, Hew. Trans. Ent. Soc. Lond. ser. 3, vol. 11. p. 500 (1866). 
Hab. Pulo Laut (Doherty); Labuan (coll. Staudinger). 


ZOGRAPHETUS OGYGIOIDES, n. sp. (Plate XIX. fig. 9, 2.) 


¢. Upperside dark brown: fore wing with five yellowish-white spots placed as 
follows—one large and subquadrate in cell 2, one much smaller but also subquadrate 
in cell 8, an oblong one of moderate size near the apex of the lower edge of the cell, a 
punctiform one just above the one last named, and a small foundish one in cell 6; no 
evident sexual modification of veins | and 2. Underside: fore wing dark brown on the 
disc, paler along the dorsum, the extra-discal region concolorous with the hind wing, the 
pale spots as on the upperside ; hind wing red-brown, with about seven suffused dark 
purple-brown spots, one near the middle of cell 7, one near the middle of the upper edge 
of the cell, and a postmedian series of five others placed one each in cells 1 4, 2, 3, 4-5, 
and 6, and running parallel to the termen. Fringes whitish grey, the short scales 
grey-brown. Antenne dark brown, spotted with whitish beneath ; club white on the 
upperside, apiculus pale red beneath. Body and legs brown. Second joint of palpi 
clothed with greyish-yellow and black hairs intermixed. 

Expanse 27-28 mm. 

Hab. Kina Balu (Waterstradt); Banguey (coll. Stgr.). 

Described from three specimens, one from Gunong Ijan ex coll. Rothschild and two 
ex coll. Staudinger; one of the latter, that from Banguey, is a little smaller and paler 
than the other and wants the dark spot in cell 6 on the hind wing below. 


! ZOGRAPHETUS AURIFERUS, n. sp. (Plate XIX. fig. 138, ¢.) 

3. Upperside brown: fore wing with five white spots placed as follows—two 
oblong, one each near the base of cells 2 and 3, two very much smaller but also oblong, 
one each near the base of cells 6 and 7, and one, small and indistinct, near the apical 
third of the upper edge of the cell; no evident sexual modification of veins 1 and 2. 
Underside : fore wing brown, the extra-discal region more or less thickly clothed with 
yellow scales; the pale spots as on the upperside, a suffused and rather indistinct sub- 
terminal yellow macular band from the costa to vein 3: hind wing yellow, with a nearly 
straight postmedian series of five oblong brown spots passing from just beyond the 
middle of vein 14 to just beyond the middle of vein 6, and a more or less distinct 
suffused dusky terminal band. Fringes grey, the short scales brown. Antenne dark 
brown, finely spotted with yellowish white beneath and becoming entirely pale on the 
underside of the basal part of the club; apiculus pale red beneath. Body above brown. 
Second joint of palpi, breast, and legs clothed with yellow and black hairs intermixed. 
Abdomen beneath sordid yellowish white. 

Expanse 25-26 mm. 


204 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Hab. Island of Nias (Modigliani). 

Described from three specimens, one in coll. Elwes, one ex coll. Rothschild, and 
one ex coll. Staudinger; in the fore wing of the two latter the pale spots proper to 
cells 6 and 7 are wanting, and the pale cell-spot is only visible on the underside. 


ScoBURA, nom. nov. 


Isma, Watson, P. Z. S. 1893, p. 83, nec Distant, Rhop. Mal. p. 386 (1886). 


The species here dealt with under this genus are associated on account of their 
general resemblance to Hesperia cephala and H. bononia of Hewitson, and the main 
distinguishing feature common to them all is the presence of pale spots in both fore 
and hind wings; the palpi are ascending, with an inconspicuous third joint, and there is 
no sex-mark on the wings of the male. Antenne more than one-half, in some species 
two-thirds, as long as the costa; club slender, apiculus moderate and gradual. It 
comprises two natural sections, one consisting of cephala and cephaloides, and the other 
of feralia and its allies. 

Watson, in his Revision of the genera of Hesperiide, characterizes a genus which he 
calls Jsma, Distant, and gives obscura, Distant, as the type of it; but it appears from a 
statement of Mr. de Nicéville (Jour. Bomb. Nat. Hist. Soc. 1895, p. 378) that Watson’s 
diagnosis of the genus Jsma was drawn up from “ Hesperia” cephala, Hew., and we find 
that it does not agree with the type specimen of Jsma obscura, Dist. Under all the 
circumstances, it appears that the genus Jsma of Watson requires another name, and 
that of Scobwra has accordingly been applied to it. 

The following is a table of the species known to us :— 


1 (4). Fore wing: hyaline spots in cells 2 and 3, 2 and 4, or in cell 2 
only, not in cells 2,3, and 4, Hind wing below with the pale 
spots pure white and dark-edged. 

2 (3). Fore wing: no hyaline spot in cell 3. Hind wing below 
yellowish green or greyish green, with a large white spot near 
the base of cells 4-5, touching veins 4 and 6. 

a (26). Cell-spot of the fore wing not ae. the subcostal. Hind wing 


below yellowish green . . . . cephala, Hew. 
6 (2a). Cell-spot of the fore wing passing fre om the sectlaata to fhe Bubeoetall 
Hind wing below greyish green: . . . . . .. «. = . martini, n. sp. 


83 (2). Fore wing: no hyaline spot in cell 4. Hind wing below with the 

basal half yellow, the apical halt reddish brown ; a white spot 

about the middle of cell 14, and one each near the bases of 

cellsi23,,andsbe re) anes = - . . « » » cephaloides, de Nicéy. 
4 (1). Fore wing with hyaline spots in alls 2, 3, ane 4. 
5 (14). Pale spots on hind wing oblong or cuneate or both. 


A REVISION OF THE ORIENTAL HESPERIIDA. 205 


6 (7). Hmd wing below with two pale spots, one in cell 2 near the 
middle and the other in cell 3 near the base . . . . . . dononia, Hew. 
7 (6). Hind wing below with three contiguous pale spots, one in each 
of cells 2 and 3, and one in cells 4—5, and sometimes one or two 
others. 
8 (13). Fore wing with two long unequal cell-spots. 
9 (12). Hind wing below: pale spot in cell 2 subequal in length to or 
shorter than that in cell 3. 
10 (11). Pale spot in cell 1a of the fore wing above as wide or wider than 
high, not reaching vein 2. No pale spot in cell of hind wing 
below .. . eo aa . feralia, Hew. 
11 (10). Pale spot in cell la of fore wing hove neatly twice as rie as bowie, 
reaching quite across the cell. Hind wing below with a pale 
spot in the cel] next the base of cell 3. Lower lobe of clasp 
produced intoasharptriangle . . . . . . . . . . .  fenestrata, n. sp. 
12 (9). Hind wing below: pale spot in cell 2 about twice as long as that 
in cell 3, the cell generally with a pale spot next the base of 
cell 8. Pale spot in cell 1a of fore wing above as wide or wider 
than high, not reaching vein 2. Lower lobe of clasp not pro- 


ducedintoatriangle . ...... +. +. =.=. +. +. +. %arime, de Nicév. 
13 (8). Fore wing without any cell-spot . . . 5 9 8 0 8 6 ONMUID IT Te, Ra 
14 (5). Pale spots on the hind wing small and foanaiah 
15 (16). Hind wing: vein 2 arising near the half-length of the cell. . . dipunctata, n. sp. 
16 (15). Hind wing: vein 2 arising near the apical third of the cell . . wmbrosa, n. sp. 


! ScoBURA CEPHALA. 
Hesperia cephala, Hewitson, Ent. Mo. Mag. 1876, p. 152; id. Deser. Lep. Coll. Atk. p. 4 (1879). 
Isoteinon cephala, Elwes, Trans. Ent. Soc. Lond. 1888, p. 456, pl. xi. fig. 10 ¢; Watson, Hesp. 


Ind. p. 80. 
Isma isota, Swinhoe, Trans. Ent. Soc. Lond. 1893, p. 320. 


Hab. Sikkim (Mller); Burmah (Watson); E. Pegu (Doherty); Tavoy (Tucker) ; 
Shillong (fide Swinhoe). 

Specimens in which the pale spot is absent from cell 4 of the fore wing are not 
uncommon ; the type of J. ¢sota, Swinh., is one of these. 


ScoBURA MARTINI, n. sp. (Plate XVIII. fig. 22, 2.) 


9. Differs from S. cephala, Hew., in the following points:—The cell-spot of the 
fore wing reaches right across the cell and is rather more than half as long next the 
subcostal as it is next the median, and there is no white spot in cell 4; the hind wing 
below and the apical region of the fore wing pale greyish green, the former with a 
large hyaline spot across cells 4-5, and three small more or less distinct brown 

VoL. XIV.—ParT iv. No. 14.—October, 1897. on 


206 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


spots, one before the middle of cell 6, one before the middle of cell 2, and one beyond 
the middle of cell 1 8. 

Expanse 30 mm. 

Hab. Battak Mountains, Sumatra (Jartin). 

Described from a single specimen in coll. Rothschild. 


!ScoBURA CEPHALOIDES. 
Hesperia? cephaloides, de Nicéville, Jour. As. Soc. Beng. 1888, p. 288, pl. xill. fig. 4 3. 
Isoteinon cephaloides, Watson, Hesp. Ind. p. 80 (1891). 


Hab. Naga hills ; Bernardmyo, Burmah (Doherty). 


! ScOBURA BONONIA. 


Hesperia bononia, Hewitson, Desc. Hesp. p. 29 (1868) ; id. Exot. Butt. v., Hesp. pl. vii. figs. 75, 76 
(1876). 
Isma bononia, Distant, Rhop. Mal. p. 386, pl. xxxv. fig. 20 (1886). 


Hab. Pulo Laut (Doherty). 


! ScOBURA FERALIA. 
Hesperia feralia, Hewitson, Descr. Hesp. p. 31 (1868) ; id. Exot. Butt. iv., Hesp. pl. iv. fig. 32 (1869). 


Hab. Kina Balu, Borneo (Waterstradt); Pulo Laut (Doherty); West Java 
(Piepers). 


| ScoBURA FENESTRATA, n. sp. (Plate XIX. fig. 16, g ; Plate XXIV. fig. 46.) 


3. Closely allied to 8. inarime, de Nicéy., which it resembles in the possession of a 
hyaline spot in the cell of hind wing below, next the base of cell 3. It is distinguished, 
however, by the shape of the white spot which stands on vein 1 a of the fore wing above, 
which is oblong, nearly twice as high as wide, reaching from vein 1a to vein 2, and 
having its inner edge continuous with the inner edge of the white spot in cell 2. 

Hab. Pulo Laut (Doherty). Type in coll. Elwes. 


!ScoBURA INARIME. (Plate XXIV. fig. 47.) 


Isma inarime, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 391, pl. vi. fig. 38 ¢. 
Pamphila zetus, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 55 (1893). 


Hab. Perak; Pulo Laut (Doherty); Java (fide Mabille). Type in coll. Elwes. 


!ScoBURA CONCINNA, n. sp. (Plate XIX. fig. 12, 2 .) 

2. Upperside deep warm brown: fore wing with six irregular white spots, of which 
four placed one in each of cells 1 @ to 4 form an oblique series from just beyond the 
middle of vein 1a@ to cell 4, and two small oblong ones are placed one in each of cells 6 


A REVISION OF THE ORIENTAL HESPERIIDA. 207 


and 7: hind wing with three white spots on the disc, that in cell 2 oblong, that in cell 3 
cuneate, and that in cell 4 roundish. Underside : fore wing blackish-brown on the disc, 
closely sprinkled with greenish-yellow scales in the costal and apical regions ; pale spots 
as on the upperside save that the one in cell la@ is represented by a suffused white 
patch and a pale point is feebly indicated in cell 8: hind wing dull greyish green by 
reason of a close sprinkling of greenish-yellow scales on a brown ground, pale spots as 
on the upperside but feebly and irregularly dark-edged. Fringes pale grey, the short 
scales brown. Antenne above blackish, spotted with white beneath; club white on 
the underside. Second joint of the palpi clothed with greyish-yellow and black hairs 
intermixed. Body above and legs brown ; abdomen beneath and the tarsi yellowish grey. 

Expanse 31 mm. 

Hab. Pulo Laut, Borneo (Doherty). 

This species, which is described from a single exemple in coll. Elwes, is well 
distinguished from its allies by the want of pale spots in the cell of the fore wing. 


ScoBURA BIPUNCTATA, n. sp. (Plate XIX. fig. 2, 2.) 


?. Upperside ochreous brown: fore wing witha yellow spot near the middle of cell 
1q@ and seven hyaline spots—two small and remote in the cell, one in each of cells 2, 3, 
and 4 forming a rapidly decreasing series, and one in each of cells 6 and 7; hind wing 
with two small irregularly roundish hyaline spots on the disc, and one in each of cells 2 
and 3, Underside: fore wing brown on the disc, the dorsal region pale yellowish grey 
nearly to the base, the costal and apical regions thickly clothed with greenish-yellow 
scales, the hyaline spots as on the upperside: hind wing dull greenish-yellow by reason 
of the thick clothing of greenish-yellow scales on a brown ground, a median band of 
small irregular, bare, and therefore brown, spots running parallel to the termen, the 
hyaline spots in cells 2 and 3 not so well defined as on the upperside. Fringe of the 
fore wing yellowish grey indistinctly chequered; of the hind wing greyish yellow on the 
upperside, on the underside with the short scales and a few long ones next veins 2 and 
3, brown. Second joint of palpi clothed with greyish-yellow and black hairs inter- 
mixed. Antenne above brown, spotted beneath with yellowish white ; club white on the 
underside. Body above concolorous with the wings, abdomen beneath and legs paler. 

Expanse 28 mm. 

Hab. Palawan (Platen). 

Described from a single specimen ex coll. Staudinger. 


ScoBURA UMBROSA, n. sp. (Plate XIX. fig. 1, 2.) 


2. Upperside deep warm brown, dorsal region of the hind wing paler: fore wing 
with seven sordid white spots placed as follows :—one, roundish, near the middle of cell 
1 a@ next vein 1a, one, subquadrate, near the basal third of cell 2, one much smaller near 

252 


208 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


the basal third of cell 3, one, small and roundish, near the middle of cell 4, one in each 
of cells 6 and 7, and one, small and somewhat roundish, near the upper edge of the cell: 
hind wing with two small hyaline spots on the disc, one in each of cells 2 and 3, and 
sometimes with a third pale spot indicated by a feeble pale point in cell 5. Underside 
grey-brown, the disc of the fore wing darker towards the base; fore wing with the pale 
spots as on the upperside save that the one in cell 1 @ is larger and suffused; hind wing 
with the pale spots in cells 2and 3 as on the upperside and the pale point in cell 5 
distinct. Fringe of the fore wing brown, becoming a little paler towards the tornus, of 
the hind wing dark grey feebly chequered. Antenne above blackish, finely spotted 
with white on the underside, club yellowish white beneath. Second joint of palpi 
clothed with yellowish-grey and black hairs intermixed. Body and legs concolorous 
with the wings. 

Expanse 33 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from two specimens ex coll. Staudinger. 


Matapa. 
Matapa, Moore, Lep. Cey. i. p. 163 (1881) ; Watson, P. Z. S. 1893, p. 85. Type aria, Moore. 


A natural and easily recognized group of species, associated on account of their 
resemblance to the /smene aria of Moore, for which he subsequently established the 
present genus. Wings above and below brown unspotted; fore wing pointed, 
termen nearly straight, on the upperside in the male bearing a narrow curved impressed 
marking from the middle of vein 1 @ to the base of vein 3. Body stout ; antenne rather 
more than half as long as costa, Palpi appressed, very densely scaled, 3rd joint 
concealed. 

The males of the species in coll. Elwes may be distinguished by the characters given 
below :— 

1 (2). Fringe of hind wing below whitish grey, the basal half (i. e. the 
short scales) brown. . . Ba 8 ery Wiloloye ey 
2 (1). Fringe of hind wing below in — ai bright clio 
3 (6). Fore wing above with the sex-mark black. 
4. (5). Fore wing above with a purple shade in certain lights. Upper edge 


of lower lobe of clasp excavated . . . . . . . - + « « purpurascens, U. sp. 
5 (4). No purple shade on fore wing above. Upper ae of lower lobe of 
clasp convex, even ae 5 A 2) enone arn aNloore. 


6 (8). Fore wing above with the sex- aS petal grey. 
7 (8). Hind wing below with all the scales in the fringe to cells 1-3 bright 
Wellow sieeuicuse- sasivarna, Moore. 
8 (7). Hind wing below Sth ie short ie in the fringe to "palie 1. 3 fae 
the long ones yellow ic opto satater 6 ae 4 6. Cob CCl Mie 


A REVISION OF THE ORIENTAL HESPERIIDA, 209 


! MATAPA ARIA. 
Ismene aria, Moore, P. Z. 8. 1865, p. 784. 
Matapa aria, Moore, Lep. Cey. vol. i. p. 164, pl. Ixvi. figs. 1, 1 @ (1881); Distant, Rhop. Mal. 
p- 378, pl. xxxv. fig. 8 (1886) ; Watson, Hesp. Ind. p. 22. 


Hab. E. Pegu, Bali (Doherty); Andamans (de Roepstorff); Ganjam (Minchin) ; 
N. Canara (Aithen) ; Java (Piepers); Philippines (Semper) ; Palawan (Platen). 


! MATAPA PURPURASCENS, n. sp. (Plate XX. fig. 1, ¢ ; Plate XXIV. fig. 48.) 


3. Upperside brown, with a purple shade which is strongest on the apical half of the 
fore wing, sex-mark blackish. Underside brown, fore wing with the dorsum and apex 
suffusedly paler. Fringe of the fore wing whitish grey, of the hind wing deep yellow 
from the tornus about as far as vein 6, thence to the apex brown. Antenne brown, 
spotted with yellowish beneath, club yellowish on the underside. Clothing of palpi, 
body, and legs concolorous with the wings. 

Expanse 40 mm. 

Hab. Khasias (Hamilton) ; KE. Pegu (Doherty). 

Described from two males in coll. Elwes, where, however, there is no female which 
can be certainly identified as the female of this species, although there are two purpu- 
rascent females, one from Sikkim with the underside entirely pale grey-brown, and the 
other from Tavoy with the underside as in J. druna; both these females have the 
clothing of the apex of the abdomen yellow. 

This species is nearest to M. druna, Moore, from which it differs in having a purple 
shade on the fore wing above, and also in the clasp-form of the male. 

Doherty has sent from East Pegu no less than three species of this genus—purpu- 
rascens, sasivarna, and shalgrama, and also a single female which does not agree with 
any of those species ; it has not the purple shade on the upperside proper to purpu- 
rascens, the underside is too pale and the yellow clothing of the apex of the abdomen 
too extensive for sasivarna (of which, moreover, I have typical females from East Pegu), 
and it has not the pale red-brown underside nor the brown clothing to the apex of the 
abdomen proper to shalgrama; its underside is pale grey-brown, and the clothing of 
the abdomen beneath is orange-yellow through rather more than its apical third; 
the fringe of the hind wing is orange-yellow, passing into pale grey-brown next 
cells 4-8; the colour of the hind wing below is uniform and not paler in the basal 
third as usual in druna 2, and the pubescence of the hind part of the thorax and of 
the basal part of the hind wing above shows in certain lights a brilliant greenish-blue 


colour. 


210 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


!MarTapa DRuNA. (Plate XXIV. fig. 49.) 
Ismene druna, Moore, P. Z. S. 1865, p. 784, g ; Wood-Mason and de Nicéville, Jour. As. Soc. 


Beng. 1881, p. 255 92. 
Matapa druna, Watson, Hesp. Ind. p. 23. 


Hab. Sikkim (Miller); Andamans (de Roepstorff); Kina Balu ( Waterstradt). 


! MATAPA SASIVARNA. 


Tsmene sasivarna, Moore, P. Z. 8. 1865, p. 784. 
Matapa sasivarna, Watson, Hesp. Ind. p. 23. 


Hab. Sikkim (Méller) ; Khasias (Hamilton); E. Pegu, Perak, Pulo Laut (Doherty); 
Tavoy (Zucker). 


! MATAPA SHALGRAMA. 
Hesperia aria, Hewitson (nec Moore), Exot. Butt. vol. iv., Hesp. pl. iii. figs. 24, 25, 9 (1868) ; fide 
de Nicéville 2. c. post. 
Matapa shalgrama, de Nicéville, Jour. As. Soc. Beng. 1888, p. 85 ; Watson, Hesp. Ind. p. 24. 


Hab. Sikkim (Méller) ; E. Pegu, Pulo Laut, Bali (Doherty) ; Java (Fruhstorfer). 


SEPA. 


Sepa, de Nicéville, Jour. As. Soc. Beng. 1894, p. 50. Type cronus, De Nicév. 


This genus will include certain species distinguished from Parnara by having vein 5 
of the fore wing straight and practically intermediate between vein 4 and vein 6. 

Antenne more than half as long as costa, reaching nearly to the end of the cell, club 
gradual, apiculus about one fourth as long as the club. Palpi with the second joint 
densely scaled, third joint short, nearly or quite concealed. Fore wing moderately 
pointed, costa about one-fourth longer than the dorsum, termen moderately curved, a 
trifle shorter than the dorsum ; vein 5 straight, practically intermediate between vein 4 
and vein 6, base of cell 3 a little shorter than that of cell 4. Hind tibize with two 
pairs of spurs. 

The linear sex-mark on the fore wing in the first three species is most easily observed 
whilst the wing is wet with benzole. 


Males. 


1 (10). Fore wing with distinct hyaline spots. 

2 (7). Sex-mark on fore wing above linear, being merely a ridge forming 
the inner boundary of the hyaline spot in cell 2, and sometimes 
continued across ce! la. 

3 (6). Sex-mark reaching from vein 1 a to vein 3. 


A REVISION OF THE ORIENTAL HESPERIID A. 211 


4 (5). Sex-mark angulated near the middle, erect from near the middle of 
vein 1@ as far as vein 2, thence oblique to the base of vein 3. 
Hyaline spot in cell 2 of fore wing above linear, no white spot 
near the middle of cell la. Hind wing below with a pale dot in 
each of cells 2 and 4 and the indication of another next the 
transverse vein. Fringe of the hind wing gradually increasing in 
length from vein 2 to the tornus, where it is about three times as 
long as at vein2 5 > oo 6 0 0 0 CRON, Ole NOR, 
5 (4). Sex-mark continuous, Fahtly oblique ehvouetiout: Hyaline spot in 
cell 2 of the fore wing above oblong, rather more than twice as 
high as wide, a small white spot near the middle of cell la. Hind 
wing below with a small pale spot near the basal third of each of 
cells 2 and 3. Fringe of the hind wing normal or nearly so . . cicatrosa, n. sp. 
6 (8). Sex-mark between veins 2 and 3 only, not extended across cell 1a. 
A long-oval slightly raised space near the base of cell la and 
lying next to vein 1a, which is there slightly sinuate . . . miosticta, de Nicéy. 
7 (2). Sex-mark a long-oval raised space near the base of cell 1a and tite 
next to vein la, which is there sinuate. 
8 (9). Fore wing with two unequal cell-spots . . . . . . . . . « guttulifera, n. sp. 
9 (8). Fore wing without cell-spots . . . . cinnamomea, N. sp. 
10 (1). Pale spots absent from the fore wing or only sharply Bened on the 
underside. 
11 (14). Dorsum of the fore wing feebly sinuate and bearing on its middle 
third a thin fringe of long hairs directed obliquely outward and 
downward. 
12 (18). Fore wing with pale spots, feebly indicated on the upperside, sharply 
defined on the underside. Hind wing below with a pale point in 
each of cells 2 and 3 and sometimes the indication of another in 
Gall.G. 7 ase Dato GANG 0 6 6) 6 6 5 0 MOSM, i GD 
13 (12). Wings brown, enitelyy taenaeed Seas 6 o 6 oO 0 6 o GANT, Me Ro 
14 (11). Dorsum of the fore wing normal. Wings brown, entirely unspotted. noctis, Ster. 


SEPA CRONUS. 

Sepa cronus, de Nicéville, Jour. As, Soc. Beng. 1894, p. 50, pl. v. fig. 4 ¢. 

Hab. Battak Mountains, N.E. Sumatra (én coll. Martin); Kina Balu, Borneo 
(Waterstradt); Gunong Ijan (i coll. Rothschild), 

The following is the description of a female Sepa from Gunong [jan ex coll. 
Rothschild, which may or may not be the female of S. cronus, the male of which from 
the same locality is in the same collection. 

@. Upperside brown; fore wing with seven sordid white hyaline spots placed as 
foliows :—one, quadrate, across the basal third of cell 2, one, also quadrate, across the 
basal third of cell 3, one, small and narrow, before the middle of cell 4, one, small and 
roundish, in each of cells 6 and 7, and two in the cell, the upper one punctiform, the 


212 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


lower one larger and suboval. Underside brown; dorsal region of the fore wing as far 
as vein 2, except at the base and apex, suffusedly pale grey. Fringes concolorous with 
the wings. Antenne blackish, minutely spotted with white in front; club broadly 
white on the underside next the apiculus. Body above concolorous with the wings. 
Clothing of the second joint of palpi of whitish and black hairs intermixed, of the 
breast of yellowish and black hairs intermixed, of the legs brown. Abdomen beneath 
grey. 
Expanse 34 mm. 


SEPA CICATROSA, n. sp. (Plate XIX. fig. 4, ¢ .) 


3. Upperside rather pale brown: fore wing with two small indistinct white cell- 
spots, a round white spot in each of cells 6 and 7 and a minute white point in cell 8, 
an erect oblong white spot standing on vein 1a near the middle and reaching halfway 
across cell 1a, an oblong white spot standing on vein 2 before the middle and passing 
obliquely to the base of vein 3, a white spot near the basal third of cell 3, and a small 
linear erect white spot near the middle of cell 4, the inner edge of the white spots in 
cells la and 2 is bounded by a linear ridge of androconia: hind wing with a white 
point in each of cells 2 and 3. Underside paler than above ; the pale spots as on the 
upperside save that the one in cell 1a is represented by an oblong patch of white scales 
reaching quite across cell 1@ and similar in size and shape to that in cell 2. Fringes 
brown, that of the hind wing showing a tendency to elongation next the tornus. 
Antennz above blackish, spotted with white beneath, club whitish on the underside. 

Expanse 33 mm. 
Hab. Kina Balu, Borneo (Waterstradt). 
Described from one male in coll. Staudinger. 


!SEPA MIOSTICTA. 
Parnara miosticta, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 385, pl. G. fig. 31 ¢. 


Hab. Perak (Doherty). Type in coll. Elwes. 


SEPA GUTTULIFERA, n. sp. (Plate XIX. fig. 8, 2.) 


g. Upperside deep warm brown: fore wing with two unequal suboval yellowish- 
white cell-spots of which the lower one is much longer than the upper, a small 
yellowish-white spot in cell la next to vein 1a near the middle, a large subquadrate 
yellowish-white spot near the middle of cell 2, a small yellowish-white spot near the 
basal third of cell 3, and a small yellowish-white spot in each of cells 6 and7; near 
the base of cell 1a@ and next to vein la, which is there slightly sinuate, a long-oval raised 
space: hind wing sometimes with two pale dots, one in each of cells 2and 3. Underside 
paler brown than the upperside, the pale spots as on the upperside save that there is a 


A REVISION OF THE ORIENTAL HESPERIIDZ. 213 


very minute pale dot near the middle of cell 4, and that the pale spot in cell 1 a is 
represented by a whitish blotch near the middle of that cell. Fringe of the fore wing 
brown, of the hind wing paler, inclining to yellow-brown near the middle; the short 
scales brown. Antenne above blackish, spotted with white beneath; club whitish on 
the underside. Body and legs concolorous with the wings. Second joint of palpi 
clothed with greyish-yellow and black hairs intermixed. 

Expanse 38 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from two specimens in coll. Staudinger. 


SEPA CINNAMOMEA, n. sp. (Plate XIX. fig. 25, 3.) 


3. Upperside brown, inclining to yellow-brown; fringe of the hind wing yellow 
except at the apex: fore wing above with four hyaline spots—one large, about one-half 
higher than wide, across cell 2 near the middle, and one in each of cells 3, 6, and 7, that 
in cell 7 very minute ; a long-oval raised space near the base of cell 1 a and lying next 
to vein 1a, which is there slightly sinuate: hind wing above sometimes with the pale 
spots proper to the underside faintly showing through. Underside much paler than 
the upperside; fore wing with the pale spots as on the upperside : hind wing with three 
small pale discal spots, one near the basal third of each of cells 2 and 3 and one in 
cells 4-5, the latter sometimes only represented bya pale point near the basal third of 
cell5. Fringe of the fore wing grey-brown ; of the hind wing grey-brown in cells § and 7, 
the remainder yellow in fresh, yellowish grey in somewhat faded specimens. Antenne 
blackish above, spotted with white beneath, club whitish on the underside. Second 
joint of palpi clothed with yellow hairs, with black ones intermixed. Body beneath 
and tarsi yellowish. 

Expanse 32-34 mm. 

Hab. Wina Balu, Borneo (Waterstradt). 

Described from two specimens in coll. Staudinger. 


SEPA BISERIATA, 0. sp. 


Parnara? species? Semper, Schmett. Philipp. p. 299 (1892). 


3. Upperside warm brown: fore wing with four very indistinct pale spots, one near 
the upper outer angle of the cell and one in each of cells 2, 3, and 6; dorsum feebly 
sinuate and bearing on its middle third a fringe of long hairs directed obliquely outward 
and downward. Fore wing below rather paler than above ; middle third of the dorsum 
thickly fringed with long hairs directed obliquely upward and outward, of which those 
nearer the base are more obliquely placed than the remainder ; the pale spots placed 
as above, but sordid white in colour, that in cell 2 twice as high as wide and reaching quite 
across the cell just before the middle, the others small and roundish ; the middle of the 

VOL. XIV.—Part Iv. No. 15.— October, 1897. 2 


214 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


dorsum, where it is in great part covered by the upper series of the fringe of hair-scales, 
is pale grey: hind wing with two small, feeble, roundish, sordid white spots, one before 
the middle of each of cells 2 and 8. Fringe brownish grey, the short scales brown. 
Antenne, body, and legs concolorous with the wings, the clothing of the scapulie 
appearing metallic green in a strong light. 

Expanse 32 mm. 

Hab, Philippines (Semper). 

Described from Herr Semper’s single male specimen from Central Luzon and one 
from Mindoro in coll. Staudinger. The latter differs from the former only in having a 
small pale spot in each of cells 7 and 8 on the fore wing below. 


!SEpa cinta, n. sp. (Plate XIX. fig. 22, 3.) 


g. Upperside dark brown with a faint purple shade. Dorsum of the fore wing 
sinuate and bearing on its middle third a fringe of long hairs, which are directed 
obliquely outward and downward. Underside similar in colour; dorsal region of the 
fore wing pale brown as far as vein 2, the dorsum bearing on its middle third a thick 
fringe of long hairs directed obliquely upward and outward, of which those nearer the 
base are more obliquely placed than the remainder. Fringes brown. Antenne, body, 
and legs concolorous with the wings. 

@. Similar to the male, but a little larger and paler. 

Expanse, ¢ 352 mm., 9 35 mm. 

Described from a single pair from Pulo Laut (Doherty). 

Type in coll. Elwes. 


! SEPA NOCTIS. 
Pamphila noctis, Staudinger, Iris, ii. p. 143 (1889). 
Plesioneura dissimilis, Snellen, in litt. 
Pamphila perfusca, Mabille, Ann. Soe. Ent. Belg. xxxvii. p. 53 (1893). 


A little larger than the preceding species, with no purple tinge, and no infra-alar 
tuft on the fore wing of the male. 

Hab. Palawan, Mindanao (in coll. Staudinger); Pulo Laut (Doherty); W. Java 
(Piepers) ; Sumatra (Martin, in coll. Rothschild). 

Herr Snellen has kindly sent us a pair of his Plesionewra dissimilis. The male we are 
unable to distinguish from “ Pamphila” noctis, Stgr., of which we have several speci- 
mens, including one from Dr. Staudinger. The female is a brown insect with a broad 
white discal band on the fore wing between the upper edge of the cell and vein 2, 
composed of the following hyaline white spots, namely—a large oblong one across the 
cell near its apical third, a small one filling up the base of cell 3, and a large one, 
oblong with rounded angles, in cell 2. Save that the white band on the fore wing is cut 
short by vein 2 and there is no suffused white subtornal patch on the fore wing below. 


A REVISION OF THE ORIENTAL HESPERIIDZ. 215 


it agrees well with the description and figure of Notocrypta monteithi, Wood-Mason 
and de Nicéville (Jour. As. Soc. Beng. 1886, p. 391, pl. xviii. figs. 3,3a,9). As, 
however, Herr Snellen tells us that this female “is certainly the other sex [of his 
P. dissimilis|, being bred from the larva by Mr. Piepers,” we do not feel justified in 
placing it in the genus Votoerypta as we should have done. 

We have seen a female from North Borneo in coll. Rothschild which agrees with the 
description and figure of NV. monteithi, and is similar to Herr Snellen’s female. 


ACERBAS. 


Acerbas, de Nicéyille, Jour. Bomb. Nat. Hist. Soc. 1895, p. 381. Type anthea, Hew. 

This genus was established by Mr. de Nicéville for the Hesperia anthea of Hewitson. 
We associate with the latter species three others which in form, colour, and pattern 
resemble it more than they do any other species known to us; they differ, however, 
from anthea in wanting an alar sex-mark in the male, and from that species and each 
other in venation. 

We distinguish the species as follows :— 


1 (2) Cell-spot normally absent from fore wing. Cell of hind wing at 

least half as long as the wing. Disc of fore wing below in the 

male covered with modified scales. . .. . =... =... anthea, Hew. 
2 (1) Cell-spot normally present in the fore wing. Cell of hind wing 

distinctly less than half as long as the wing. No alar sex-mark 

in the male. 
3 (6). Base of cell 4 about one-fourth as long as the base of cell 3. Band 

on the hind wing below dull white. One pale spot in cell of 

fore wing near its upper edge. 
4 (5). White band on hind wing above evanescent towards the costa. Dark 

part of underside plain brown. Pale band on the hind wing 

below white throughout, a little contracted next the costa. . . martini, Dist. 
5 (4). White band on hind wing above broad, sharply defined, abruptly cut 

short at vein 6. Dark part of underside brown, with a distinct 

purple shade, which is strongest in the apical region of the fore 

wing. Pale band on hind wing below white, becoming yellow in 

cells 7 and 8, not contracted next the costa. . . . duris, Mab. 

6 (3). Base of cell 4 about half as long as the base of cell 3. ‘Band on 
hind wing below glistening white. Two pale spots in cell of 


foreswineye: es ies) fe aye e me nitidifasciata, n. sp. 


! ACERBAS ANTHEA. 


Hesperia anthea, Hewitson, Descr. Hesp. p. 29 (1868). 
Plesioneura? anthea, Distant, Rhop. Mal. p. 404, pl. xxxv. fig. 32 (1886). 
? Carystus tagiadoides, Mabille, sec. spec. comm. 


Hab. Pulo Laut (Doherty). 


216 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! ACERBAS MARTINI. (Plate XXIV. fig. 50.) 
Zea martini, Distant, Ann. & Mag. Nat. Hist. ser. 5, vol. xix. p. 274 (1887). 
Hab. Pulo Laut (Doherty). 


! ACERBAS DURIS. 

Carystus duris, Mabille, Comptes Rendus Soe. Ent. Belg. iii. no. 31, p. lix (1883). 
Lotongus duris, Semper, Schmett. Philipp. p. 289, pl. xlix. fig. 7, ? (1892). 
Carystus mabillei, Staudinger, MSS. 

Hab. Philippines (Semper); Kina Balu, Borneo (Waterstradt). 

Though the characters given in the table seem sufficient to separate the few specimens 
(only six in all) that we have seen of these two species, yet the occurrence of both of 
them in Borneo throws some doubt on the constancy of the differences, and we have 
seen no female of martini as yet. The female of duris is similar to the male. The 
male genitalia of the two forms do not afford material for their separation. 


! ACERBAS NITIDIFASCIATA, 0. sp. (Plate XX. fig. 9, ¢ ; Plate XXIV. fig. 51.) 


Besides the characters given above this species may be distinguished from 
A. martini by its larger size (expanse 47 mm.) and the broader white band on the 
hind wing below. ‘The latter at its greatest breadth extends from the cell at the level 
of the base of vein 2 as far as the apical third of cells 4-5; in A. martini this band at 
its greatest breadth extends from the cell at the level of the base of vein 2 only half- 
way across cells 4-5. In A. nitidifasciata the apex of the lower lobe of the clasp is 
simply rounded and serrate; in A. martini the same part is broadly truncate, and its 
inner angle is produced into a long strap-shaped lobe, rounded at the apex. 

Hab. Labuan (fide Staudinger) ; N. Borneo (Pryer); Pulo Laut (Doherty). 


PUDICITIA. 
Pudicitia, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 379. Type pholus, de Nicév. 

This genus has been established by Mr. de Nicéyille for his Parnara pholus, an insect 
of the size and shape of P. assamensis, Wood-Mason and de Nicéy., and haying deep 
yellow hyaline spots on both wings, those on the hind wing forming a transverse discal 
series. ‘The course of vein 6 in the fore wing is not stated in the description, but vein 
la in that wing has in the male “a narrow brand formed of modified black scales 
placed anteriorly against it along its basal second and third fifths.” 

Pudicitia is compared by its author with Erionota, Mab. 


PuDICITIA PHOLUS. 


Parnara pholus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 172, pl. B. fig. 3, 2. 
Pudicitia pholus, 10. op. cit. 1895, p. 381. 


Hab. Bhutan (Moller); Khasias (fide de Nicéville). 


A REVISION OF THE ORIENTAL HESPERIIDZ. 217 


ERIoNoTA. 

Erionota, Mabille, Ann. Soc. Ent. Belg. xxi. p. 34 (1878) ; Watson, P. Z.S. 1893, p. 86. Type 

thrax, Linn. 
Large species expanding 60 to 90 mm. Hind wing with vein 7 almost equidistant 
between 6 and 8, veins 2 and 3 normal in both sexes. 
We are acquainted with the three following species :-— 

1 (4). Pale spots on fore wing above yellow. 

2 (3). Fore wing above: pale spot in cell 3 remote from the others, situate 
near the middle of the cell, variable in shape, but never wider than 
high PON Ce ROR OMS holo phn Sg uch en CAC Abii, 

3 (2). Fore wing above : pale spot in cell 3 occupying the basal half of the 


cell with the exception of the extreme base, and therefore only 
separated from that in cell 2 byven3 . . . .- =. . =. . alexandra, Semper. 
4 (1). Pale spots on fore wing above pure white . . . . . . . . . grandis, Leech. 


! ERIONOTA THRAX. 


Papilio thrax, Linneus, Syst. Nat. i. 2, p. 794 (1767) ; Donovan, Ins. Ind. pl. xlix. fig. 2 (1800). 
Erionota thraz, Mabille, Ann. Soc. Ent. Belg. xxi. p. 35 (1878) ; Watson, Hesp. Ind. p. 107 (1891). 
Telegonus acroleucus, Wood-Mason & de Nicéville, Proc. As. Soc. Beng., August 1881, p. 143. 
Hesperia hiraca, Moore, Traus. Ent. Soc. Lond., September 1881, p- 313, ?. 
Hesperia acroleuca, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1881, p. 260. 
Telegonus lara, Swinhoe, Ann. & Mag. Nat. Hist. 1890, p. 365 (fide Watson). 
Erionota acroleuca, Watson, Hesp. Ind. p. 107 (1891). 
Hab. Sikkim (Moller); Khasias (Hamilton); Pulo Laut, Sambawa, Bali (Doherty) ; 
Nias (Modigliani) ; Andamans (de Roepstorf’) ; Philippines (Semper) ; Palawan (Platen). 
Two specimens of acroleuca, named by de Nicéyille, from the Andamans, seem to 
us inseparable from thraa, though they are conspicuously smaller. 


ERIONOTA ALEXANDRA. 
Erionota alexandra, Semper, Schmett. Philipp. p. 312 (1892). 


Hab. N.W. Luzon (Semper). 
The larva of this species is very differently coloured to that of EZ. thrax 


! ERIONOTA GRANDIS. 


Plesioneura grandis, Leech, Entomologist, xxii. p. 47 (1890). 
Hidari grandis, Leech, Butt. China &e. p, 633, pl. xxxix. fig. 13, g¢ (1894). 


Hab. West China (Pratt). 
GANGARA. 
Gangara, Moore, Lep. Cey. i. p. 164 (1881) ; Watson, P. Z.S. 1893, p. 86. Type thyrsis, Fab. 
This genus may be distinguished from Erionota by having veins 2 and 3 of the 


218 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


hind wing more or less swollen in their basal half in the males, as in Paduka; the 
underside, too, has some bluish-white scales in suffused patches, or a pink shade 


towards the base. 
The three species known to us are distinguished as follows :— 


1 (4). Fore wing above with pale spots in cells 6, 7, and 8. No 

yellowish-white spot in the base of cell 7 of hind wing below. ~ 
2 (3). Fore wing above: pale spot in cell 6 remote from that in cell 7, that 

in cell 8 oblong. Male: fore wing above with a vein-like ridge 

arising out of the upperside of vein 1 a in its apical fourth, and 

running parallel to it nearly to the base, and a ridge on each side 

of the basal half of vein 2; fore wing below with an obiong patch 

of yellow hairs near the denen, one . . thyrsis, Fab. 
3 (2). Fore wing above: pale spot in cell 6 close to ane in cell 7, that 3 in 

cell 8 punctiform. Hind wing below brown, with a pink shade 

near the base, and a sub-basal and median series of suffused 

oblong greenish-brown spots. . . . . 50 . sybirita, Hew. 
4 (1). Fore wing above without pale spots in cells 6, 7, or 8. Hind wing 

below with a yellowish-white spot near the base of cell 7. Male 

without a sex-mark on fore wing above, the tuft of hair near the 

dorsum of fore wing below much less evident than in thyrsis . sanguinocculus, Martin. 


! GANGARA THYRSIS. 
Papilio thyrsis, Fabricius, Syst. Ent. p. 582 (1775). 
Hesperia pandia, Moore (Horsf. & Moore), Cat. Lep. Mus. E.I. C. vol. i. p. 254, pl. vil. 
figs. 10 larva, 10 a pupa (1857) ; Moore, P. Z. 8. 1865, p. 790. 
Gangara thyrsis, Moore, Lep. Cey. vol. i. p. 165, pl. Ixvi. figs. 3, 3 a (1881) ; Distant, Rhop. Mal. 
p. 894, pl. xxxv. fig. 13 (1886). _ 
Hab. Babuyanes, Luzon, Mindanao (Semper); Palawan (Staudinger); Akyab 
(Adamson); E. Pegu, Pulo Laut, Bali (Doherty); Java (Piepers); Andamans 
(de Roepstorff). 


GANGARA SYBIRITA. 


Hesperia sybirita, Hewitson, Ann. & Mag. Nat. Hist. ser. 4, vol. xviii. p. 451 (1876). 
Hidari sybirita, Distant, Rhop. Mal. p. 395, pl. xxxv. fig. 24 (1886). 


We have only been able to examine a female of this species, but entertain no doubt 


as to its being properly placed in this genus. 
Hab. Malay Peninsula, Singapore (fide Distant) ; Borneo (Cator). 


!GANGARA SANGUINOCCULUS. (Plate XX. fig. 17.) 


Erionota sanguinocculus, Martin, Einige neue Tagschmetterlinge von Nordost-Sumatra, p. 5 
(Miinchen, pub. Gotteswinter 1895). 


3. Upperside brown: fore wing with three aureo-hyaline spots—one occupying the 


A REVISION OF THE ORIENTAL HESPERIID 2. 219 


apical third of the cell, one occupying cell 3 from the base to the middle, its outer 
edge obliquely concave and passing from the basal third of vein 4 to beyond the 
middle of vein 3, and one irregularly ovate and placed obliquely near the middle of 
cell 2, of which it occupies the entire width. Underside brown: fore wing with the 
hyaline spots as on the upper surface, cell 1 pale, except in its apical fifth, a suffused 
brownish-grey patch near the middle of the costa and a suffused patch of bluish-white 
scales reaching from the apex of the cell to the costa just before the apex of the wing ; 
hind wing with a small sharply defined cream-coloured spot near the base of cell 7, 
cell 1 sparsely sprinkled throughout with bluish-white scales, and on the disc some 
suffused patches of bluish-white scales indicating an irregular median and postmedian 
band. Fringes concolorous, a little paler near the tornus of each wing. Body, 
palpi, legs, and antenne brown, the latter brownish yellow on the underside of 
the club. 

@. Fore wing above with the hyaline spot in cell 3 completely fillmg up the base 
of that cell, and the hyaline spot in cell 2 forming a parallelogram, of which the 
upperside occupies the basal half of vein 3 and the lower side is separated from 
the base of vein 2 by two-fifths, and from the apex of vein 2 by one-fifth of the entire 
length of that vein. Otherwise like the male. 

Expanse 52-55 mm. 

Hab. Perak (Doherty); N.K. Sumatra (Martin). 

The above description was drawn up from a single pair in coll. Elwes long before 
Dr. Martin’s description appeared; and as the latter did not know the female, it has 
been allowed to stand. 


PADUKA. 


Paduka, Distant, Rhop. Mal. p. 375 (1886); Watson, P. Z.S. 1893, p. 85. Type glandulosa, 
Dist.,=/ebadea, Hew. 

Allied to Matapa; the sex-marks in the male constitute the main differences ; there 
is in that sex a large tomentose patch on the disc of the fore wing above, a tuft of 
long hairs on the dorsum of the same wing below, and a conspicuous seam on the 
basal half of veins 2 and 3 of the hind wing above. There is but one described 
species, lebadea, Hew., a large insect expanding 51-65 mm. ‘The male resembles a 
Matapa in the stout body, contour of wings, uniform brown coloration of the latter 
. above, and the yellow fringe to the hind wing; the female (which appears to be 
hitherto undescribed) exactly resembles the male on the underside, but has three deep 
yellow spots on the fore wing and otherwise closely resembles Hrionota thraz. 


! PADUKA LEBADEA. 


Hesperia lebadea, Hewitson, Exot. Butt. iv., Hesp. pl. ii. figs. 22, 23 (1868). 
Tsmene subfasciata, Moore, P, Z. 8. 1878, p. 686. 


220 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Matapa subfasciata, Moore, Lep. Cey. i. p. 164, pl. lxiv. figs. 3a, 6 (1881); Watson, Hesp. Ind. 
p. 24 (1891). 
Paduka glandulosa, Distant, Rhop. Mal. p. 376, pl. xxxv. fig. 5, g (1886). 


Hab. Perak, Pulo Laut (Doherty); Java (Fruhstorfer) ; Sikkim (Knyvett). 


WaATSONIA, gen. Noy. 


Antenne half as long as the costa; club moderate ; apiculus acuminate, bent almost 
at a right angle, two-thirds as long as the club. Palpi appressed, second joint densely 
scaled, third almost concealed. Fore wing: distal two-thirds of the costa straight, 
dorsum two-thirds as long as the costa, about one-sixth longer than the termen ; 
vein 5 straight, a trifle nearer to vein 4 than to vein 6, vein 12 approaching 11 in its 
apical fourth, base of cell 10 one-third longer than that of cell 9, of celi 8 about half 
that of cell 9, of cells 6 and 7 mere points, base of cell 3 about half as long as that 
of cell 4 and giving off a recurrent vein, that of cell 2 nearly or quite as long as the 
first median segment; transverse vein nearly parallel with the termen. Hind wing 
suborbicular, a little longer than broad; termen nearly straight between veins 1 
and 3; first and second subcostal and median segments subequal; cell less than half 
as long as the wing; transverse vein concave, slightly angulated at the origin of 
vein 5. No hyaline spots in either wing. ‘Tibial epiphysis present; hind tibie with 
two pairs of spurs. 

Differs from Kerana in the shorter antenne, straighter costa to the fore wing, the 
position of vein 5, and the shorter base to cell 3 in the same wing. 


| WATSONIA SWINHOEI, n. sp. (Plate XX. fig. 6, ¢ .) 

3. Upperside deep brown with a dull purple shade, except on the apical region of 
the fore wing. Underside brown: fore wing with the dorsum broadly pale brown and 
the apical region ochreous brown; hind wing with a faint purple shade. Fringes 
brown. Antenne, palpi, body, and legs concolorous with the wings, antenne a little 
paler in front. 

¢. Similar to the male, but a little larger and paler. 

Expanse, ¢ 47, 9 54 mm. 

Hab. Khasia hills (fide Swinhoe). 

Described from one pair given to me by Col. Swinhoe as a new genus and species. 
There are other specimens in Col. Swinhoe’s collection. 


KERANA. 
Kerana, Distant, Rhop. Mal. p. 402 (1886) ; Watson, P. Z.S. 18938, p. 115. Type armatus, Druce. 


No hyaline spots on either wing. Species brown or black above, generally with an 
oblique yellow or red-yellow band on the fore wing. Antenne more than half as long 


A REVISION OF THE ORIENTAL HESPERIIDA, 221 


as costa; club slender; apiculus acute. Palpi appressed, third joint almost concealed. 

Fore wing: dorsum longer than the termen, vein 5 nearer 4 than 6; first and second 

median segments subequal in length, the third about two-thirds as long as the second, 

vein 12 feebly sinuate towards vein 11. Hind wing: first median segment about 

three times as long as the second, the third about one-fourth as long as the second. 
The species known to us may be distinguished as follows :— 


1 (6). Fore wing above with a red-yellow or yellow band. 

2 (5). Pale band on fore wing above red-yellow, passing obliquely from the 
middle of the costa to the tornus.* 

3 (4). Male with a black patch of androconia near the base of the hind wing 
above, coinciding with the cell. Expanse50mm. . . . . . armata, Druce. 

4. (3). No sex-mark on hind wing above in the male. Hind wing above 
with a small spot of pale purple scales in each of cells 2, 3, 5, and 6. 
Expanse 32-34 mm, ; : 40 . - 

5 (2). Pale band on fore wing above yellow, very broad, passing from the 
dorsum towards the middle of the costa, where it is cut short by 
the upper edge of the cell, its outer edge convex, its inner edge 


gemmifer, Butl. 


passing obliquely from the apex towards the base and giving off 
along the dorsum a triangular tooth to the base of the wing . . fulgur, de Nicév. 
6 (1). Entirely fuliginous or olive-brown above... . . . . . . diocles, Moore. 


! KERANA ARMATA. 
Astictopterus armatus, Druce, P. Z.S. 1873, p. 359, pl. xxxii. fig. 7. 
Kerana armata, Distant, Rhop. Mal. p. 402, pl. xxxv. fig. 31 (1886). 


Hab. Perak, Pulo Laut (Doherty); Nias (Modigliant). 


! KERANA GEMMIFER. 
Astictopterus gemmifer, Butl. Trans. Linn, Soc. Lond., Zool. ser. 2, vol. i. p. 555 (1877). 
Kerana yemmifer, Distant, Rhop. Mal. p. 403, pl. xxxiv, fig. 29 (1886) ; Watson, Hesp. Ind. p. 149 
(1891). 
Hab. Perak, Pulo Laut (Doherty); Bunguran, Natuna Is. (Everett); Penrisen Mt. 
(in coll. Rothschild). 


KERANA FULGUR. 
Kerana fulgur, de Nicéville, Jour. As. Soc. Beng. 1894, p. 55, pl. i. fig.6, 9; id. Jour. Bomb. Nat. 
Hist. Soc. 1895, p. 383, pl. Q. fig. 54, 3. 
3. Upperside rich dark brown ; fore wing with a very broad yellow median band 
passing from the dorsum towards the middle of the costa and cut short by the upper 


1 In a pair of specimens of armata from the Island of Nias the red-yellow band of the fore wing above is 
entirely wanting in the male and faintly indicated in the female. 
vou. xiv.—Part Iv. No. 16.—October, 1897. 26 


222 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


edge of the cell; the outer edge of this band is nearly regularly convex, and its inner 
edge, which is straight and passes obliquely inwards and downwards, gives off along 
the dorsum a large triangular tooth to the base of the wing: hind wing with 
three divergent stripes of golden-yellow hairs from the base nearly to the middle. 
Underside paler brown than the upperside: fore wing with the yellow band as above, 
save that its inner boundary is wanting; an oblong brown spot from the middle of the 
base as far as vein 2, an oblong suffused red-yellow patch between the apical fourth of 
the cell and the costa, and a suffused yellow spot in the apex of the wing: hind wing 
with six pale bluish-purple spots placed as follows—three at equal distances in 
cell 1 b, one near the apical third of cell 2, one next the transverse vein, and one near 
the middle of the upper edge of the cell, and a few scales of a similar colour near the 
base of the costa. Antenne brown; apex of the club and the apiculus beneath yellow. 
Second joint of palpi clothed with yellow scales, intermixed with black ones. Thorax 
concolorous with the base of the wings. ‘The abdomen and legs are not in a condition 
for description. 

Expanse 42 mm. 

Hab. Borneo (in coll. Cator); Battak Mountains, Sumatra (Martin). 

The above description of the male of this insect had already been written from a 
specimen from Borneo in the collection of Mr. D. Cator when Mr. de Nicéville’s 
description appeared. The female we have not seen, but from Mr. de Nicéville’s 
description and figure we gather that the band on the fore wing in that sex differs in 
colour and shape from that of the male, being red-yellow and passing obliquely from 
the middle of the costa towards the tornus, which it does not quite reach. 


! KpRANA DIOCLES. 
Nisoniades diocles, Moore, P. Z. 8. 1865, p. 787. 
Kerana diocles, Distant, Rhop. Mal. p. 403, pl. xxxiv. fig. 8 (1886) ; Watson, Hesp. Ind. p. 148. 
Hab. Sikkim (Méller); Khasias (Hamilton); Tavoy (Tucker); Perak, Pulo Laut 
(Doherty) ; Java (Piepers); Nias (Modigliani). 


ANCISTROIDES. 


Ancistroides, Butler, Trans. Ent. Soc, Lond. 1874, p. 436; Watson, P. Z.8. 1893, p. 116. Type 
longicornis, Butler. ; 


Nearest allied to Kerana, but differs from it in the relative proportions of the third 
median segment in both wings; that of the fore wing being one-third as long as the 
second, that of the hind wing about one-fourth as long as the second. 


ANCISTROIDES OTHONIAS, (Plate XXI. fig. 18.) 
Astictopterus othonias, Hewitson, Ann. & Mag. Nat. Hist. ser. 5, i. p. 8342 (April 1878). 


Hab. Labuan, Kina Balu, Borneo (Waterstradt); N. Borneo (Pryer, in coll. 
Rothschild). 


bS 
bo 
ee) 


A REVISION OF THE ORIENTAL HESPERIIDA, 


PIRDANA. 
Pirdana, Distant, Rhop. Mal. p. 376 (1886) ; Watson, P. Z.S. 1893, p.116. Type hyela, Hewitson. 


Wings unspotted. Species plain brown above or with a blue, green, or purple shade 
from the base of the wings outwards; tornus of hind wing more or less broadly yellow. 
Antenne more than half as long as costa; club slender. Fore wing with tke termen 
as long as or longer than the dorsum, at least in the male, the former straight from 
the tornus to vein 5; vein 2 from the basal third of the lower edge of the cell, vein 5 
nearer 4 than 6. Hind wing: termen distinctly produced at vein 16; vein 2 from the 
apical third of cell. Hind tibize roughly scaled on the outer side, but not fringed with 
long hairs. 

We distinguish the species known to us as follows :— 


1 (6). With a sex-mark on the fore wing above in the male; antennz above 
brown. 

2 (5). Sex-mark on upperside of fore wing in the male consisting of a raised 
seam on each side of the basal third of vein 2 and the middle third 
of vein la. Hind wing below purplish brown with green veins, or 
washed with green. 

3 (4). Hind wing below purplish brown, the veins broadly and the dorsum 


pale green . . 3) ic Later gma meee aeeS eee Cla ewe 
4 (3). Hind wing below neanlish ineoan ea with green ; veins concolorous ; 
cells 1a and 16 purple, shct with green . . . 0 0 CMT, Se 


5 (2). Sex-mark on the upperside of the fore wing in the mules consisting of 
three oblong-oval patches of androconia, a large one near the base 
of cell 2, and two others much smaller, placed one above the other 


near the middle of cell 1a. Hind wing below plain brown . . . celsina, Feld. 
6 (1). No sex-mark on fore wing above in the male; antennz above in greater 
Partwhite) @) sees aes als = fy sp een tan Seemgera/ozcorn7s 0. SD. 


! PIRDANA HYELA. 


Hesperia hyela, Hewitson, Descr. Hesp. p. 23 (1867). 
Pirdana hyela, Distant, Rhop. Mal. p. 376, pl. xxxv. fig. 6, 9 (1886). 
Pirdana rudolphii, Elwes & de Nicéville, Jour. As. Soc. Beng. 1886, p. 438, pl. xx. fig. 6, 3. 

As the type specimen of P. rudolphii is now in the Calcutta Museum, we only know 
it from the plate above referred to, The only difference between the figures of 
rudolphii and hyela from Borneo is the smaller extent of yellow at the anal angle 
in the former. 

Hab. Perak, Pulo Laut (Doherty); Tenasserim (Bingham); Java (fide Hew.); 
Khasias (coll. Swinhoe). 


262 


224 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! PIRDANA DISTANTI. 
Pirdana distanti, Staudinger, Iris, ii. p. 141 (1889). 
Pirdana pavona, de Nicéville, Jour. As. Soc. Beng. 1895, p. 540. 

Hab. Malacca (fide Staudinger) ; Labuan (fide Staudinger, in coll. Elwes); E. Pegu, 
Perak (Doherty) ; ? Sikkim (Lang, fide de Nicéville); Buitenzorg, Java (Piepers). 

This species seems constantly distinct from hyela, though its geographical range 
covers that of hyela. It is probably the species referred to from Sikkim by de 
Nicéyille in our description of P. rudolphii. 


PIRDANA CELSINA. 
Hesperia celsina, Felder, Reise Nov., Lep. iii. p. 512, pl. Ixxi. fig. 12, 9 (1867). 
Pamphila celsina, Staudinger, Exot. Tagf. p. 298, pl. 99, 3 (1888). 
Tanyplera celsina, Semper, Schmett. Philipp. p. 294 (1892). 


Hab. Celebes ( fide Felder); Philippines ( fide Semper). 


PiRDANA ALBICORNIS, n. sp. (Plate XXI. fig. 14, 3.) 


3. Upperside: fore wing dark brown with a dull purple shade, glossed from the 
base almost to the middle with metallic greenish blue; hind wing dark brown as far 
as the apex of vein 4 and the apical fourth of the dorsum, thence bright yellow, the 
disc from the base glossed with metallic greenish blue. Underside verdigris-green ; 
disc of the fore wing dark purple, the dorsum of that wing broadly pale brown: 
hind wing with the bright yellow subtornal region as on the upperside ; cell 1 4 black, 
glossed with metallic greenish blue. Fringe of the fore wing grey-brown, of the hind 
wing yellow from the tornus as far as vein 4, thence somewhat dusky. Antenne in 
greater part white on the upperside, beneath blackish, with a white spot next the 
apiculus; club black; apiculus white. Clothing of second joint of palpi, breast, legs, 
and abdomen beneath yellow. Thorax above clothed with greyish-yellow and black 
hairs intermixed. Abdomen above brown. 

Expanse 42 mm. 

Hab. Kina Balu, Borneo ( Waterstradt). 

Described from one specimen in coll. Staudinger. 


PLASTINGIA. 
Plastingia, Butler, Ent. Mo. Mag. 1870, p. 95; Watson, P. Z. S. 1893, p. 118. Type flavescens, 
Felder, 
This genus is best distinguished by its facies, which is well exemplified in 
P. callineura, Felder. 
Antenne usually two-thirds as long as the costa; club slender; apiculus acute, 
recurved. Palpi appressed; third joint short, conical, almost concealed. Fore wing 


A REVISION OF THE ORIENTAL HESPERIIDA. 225 


pointed, termen straight, oblique; vein 5 nearer 4 than 6, vein 2 from the basal 
fourth of lower edge of the cell. Upperside brown, with pale spots in cells 2, 3, 
6, and 7, one or two near the apex of the cell on the fore wing, and sometimes one in 
cells 4 and 5; the disc of the hind wing is more or less extensively yellow. Epiphysis 
present ; hind tibiz fringed and bearing two pairs of spurs. 

There is some difference in the comparative length of the cell of the hind wing in the 
different species ; it islonger in P. tessellata, Hew., than in the others. In P. callineura, 
Feld., and its allies, the base of cell 5 in the fore wing is about one and a half times as 
long as that of cell 4; in P. tessellata, noemi, &c., the bases of cells 4 and 5 in the fore 
wing are subequal in length. 

The species known to us we distinguish as follows :— 


Males. 


1 (16). A hyaline spot in cell 4 of the fore wing. 
2 (11). Hind wing below with pale purple or bluish-white spots. 
2a (26). Termen of the hind wing broadly and decreasingly yellow from the 
dorsum as\farjas vem 3) % 4 <5 + « w+ = =) 4 os 9) vermiculata, Hew. 
26 (2a). Termen of the hind wing uot as above. 
8). Shaft of antenne entirely pale beneath. 
). Hyaline spots in cell of fore wing remote. Tegumen simple. 
(6). Hind wing below: veins rather broadly vermilion-red. Lower lobe 
of clasp produced into two strong teeth at apex . . . . callineura, Feld. 
6 (5). Hind wing below: veins yellow. Lower lobe of clasp prodncea at 
the apex into one strong tooth, the apical half of which is 
spiniform ., . . syeliettredies Shek oo oto sn 0 5, Lena, lala. 
7 (4). Hyaline spots in cell of fore wing epiitineat Tegumen with a short 
tooth on each side at the base . . . » + « + « Margherita, Dohy. 
8 (3). Shaft of antennze entirely blackish, club wale Genes 
9 (10). Fore wing above: basal three-fifths of cell 1a filled up with yellow, 
the yellow colour spreading to the dorsum at the base. Hind 
wing above: cell entirely pale, the dise of the wing yellow from 
the dorsum as far as vein 6, the dorsum broadly yellow as far as 
thentermen’ .: Wh oh ay ee Patera eines Meee ene rey ett Met et = @ Elen Ut 
10 (9). Fore wing above: an elongate triangular yellow spot in the basal 
haif of cell 1 a, its apex reaching the base of the wing on vein 1a. 
Hind wing above: upper half of the cell blackish, the yellow 
discal patch not reaching the dorsum, the latter dark brown 
THU ONAL lS ole Son 6 oo bp oe 6 a 0 on Ginger, Willey 
11 (2). No purple spots on hind wing below. 
12 (13). Fore wing with the basal two-thirds of cell 2 hyaline. Hind wing 
below red-yellow, with irregular black dashes; a broad black line 
in) the basal two-thirds oficell’7 5) 0 |) sw ee Quranttaea, n.Sp. 


15 (14). 


16 (1). 
17 (20). 
18 (19). 
19 (18). 
20 (17). 
21 (22). 


22 (21). 
23 (24). 
24. (23). 


25 (26). 


26 (25). 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


. Hyaline spot in cell 2 of fore wing much less than half as long as 


the cell. 


. Hind wing below brown, with the following pale yellow markings— 


a broad oblique postmedian band between veins 2 and 6, a broad 
stripe in cell 8, the apical third of cell 7 nearly as far as the termen, 
a stripe next the upper edge of the discoidal cell, a subterminal 
row of spots, one in each of cells 1 4 to 6, and three rays, of which 
the innermost is the shortest, next the dorsum soe hsha: 

Hind wing below yellowish green with feeble pale spots, one near 
the bases of each of cells 2, 8, and 4, and sometimes a small one 
near the middle of cell 5, that in cell 4 oblong and occupyimg 
about the basal third of the cell 

No hyaline spot in cell 4 of the fore wing. 

Hind wing below chequered throughout with pale spots. 

Hind wing below brown, chequered with greyish-yellow spots 

Hind wing below brown with dull silvery-white spots . 

Hind wing below not chequered. 

Hind wing below yellow, with the termen narrowly, the tornus 
broadly, the veins, a short line in the discoidal cell next the 
outer half of its upper edge, and a line in the basal half of cell 
1d, black wot aes 

Hind wing below: veins not black. 

Hind wing below unspotted, ochreous, blackish in cell 1 a 

Hind wing below with a postmedian band of blackish spots (some- 
times with pale centres) and one near the base of cell 7, one near 
the apex of the cell, and one near the base of cell 1 4. 

Expanse 34 mm. Fore wing above with the uppermost of the two 
pale spots in the cell small and roundish. Tegumen simple . 
Expanse 89-40 mm. Fore wing above with the uppermost of the 
two pale spots in the cell oblong, as large as that in cell 6. 

Tegumen with a decurved horn on each side about the middle 


! PLASTINGIA CALLINEURA. (Plate XXIV. fig. 52.) 
Hesperia callineura, Felder, Reise Nov., Lep. iii. p. 513, pl. Ixxi. figs. 9, 10 (1866). 


? Plastingia callineura, Distant, Rhop. Mal. p. 396, pl. xxxv. fig. 26 (1886); Watson, Hesp. Ind. 


p. 113 (1891). 


Distant’s description (/. ¢.) applies rather to the next species than to the true 
callineura of Felder, who correctly says of the hind wing below “ venis ferrugineo-rufo 


marginatis.” His plate, however, does not admit of identification with any of the 


species in coll, Elwes. 


Hab. Kina Balu, Borneo (Waterstradt) ; KE. Pegu, Perak (Doherty); Battak Moun- 


tains, Sumatra (Martin). . 


liburnia, Hew. 


telesinus, Mab. 
tessellata, Hew. 


naga, de Nicév. 


viburnia, Stgr. 


corissa, Hew. 


similis, n. sp. 


noémi, de Nicév. 


bo 
RS) 
4 


A REVISION OF THE ORIENTAL HESPERIIDA, 


! PLASTINGIA LaToIA. (Plate XXIV. fig. 43.) 

Hesperia latoia, Hewitson, Descr. Hesp. p. 34 (1868) ; id. Exot. Butt., v. Hesp. pl. vi. figs. 62, 63 
(1873). 
? Plastingia callineura et var. flavia, Staudinger, Iris, ii. p. 150 (1889). 

Distinguished from the preceding species by its slightly smaller size, the yellow 
veins on the hind wing below, and the single tooth into which the lower lobe of the 
clasp is produced. 

The variety flavia from Palawan is said to be distinguished from callineura by its 
smaller size; it may possibly be a distinct species, but we have had no opportunity of 
comparing its male genitalia with those of P. latoia. A female from Palawan, ex coll. 
Rothschild, expands 59 mm., and very possibly represents a distinct species; it does 
not agree with the same sex of either datoia or callineura. 

Hab. Pulo Laut (Doherty); Hast Java (coll. Piepers) ; Singapore (fide Hewitson) ; 
Selesseh, Sumatra (¢ coll. Rothschild). 


! PLASTINGIA MARGHERITA. (Plate XXIV. figs. 54, 54a.) 
Plastingia margherita, Doherty, Jour. As. Soe. Beng. 1889, p. 131, pl. x. fig. 5, do. 

Hab. Margherita, Upper Assam (Doherty); Naga hills (Doherty). 

As I have the type of this species, I am able to say that it seems as distinct a species 
as any in the genus, though closely allied to /atoia.-—H. J. E. 


! PLASTINGIA HELENA. 
Plastingia helena, Butler, Lep. Exot. p. 190, pl. Ixiv. fig. 3 (1874). 


Hab. Sarawak (Low, fide Butler); Pulo Laut (Doherty); Namoe Ockor, Sumatra 
(in coll. Rothschild). 


PLASTINGIA VERMICULATA. 
Hesperia vermiculata, Hewitson, Ann. & Mag. Nat. Hist. ser. 5, vol. i, p. 846 (1878). 
Plastingia vermiculata, de Nicéville, Jour. As. Soc. Beng. 1894, p. 56, pl. v. fig. 15, 3. 

This species is distinguished by having the termen of the hind wing broadly and 
decreasingly yellow from the tornus as far as vein 3. The hind wing below has several 


large bluish-white spots or dashes. 
Hab. Battak Mountains, Sumatra (Martin, fide de Nicéville). 


! PLASTINGIA FRUHSTORFERI. (Plate XIX. fig. 19, ¢; Plate XXIV. figs. 55, 55 a.) 
Plastingia fruhstorferi, Mabille, Ann, Soc, Ent. Belg. xxxvii. p. 52 (1893). 

Hab. Java (Fruhstorfer) ; Pulo Laut (Doher aw: Labuan (coll. Staudinger) ; Selesseh 
(im coll. Rothschild). 


228 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


! PLASTINGIA AURANTIACA, n. sp. (Plate XIX. fig. 17,  .) 


3. Upperside: fore wing blackish brown with five hyaline spots, one in each of cells 2, 
3, 4, 6, and 7, and another indicated by a small pale dash near the apex of the cell next 
its upper edge; an elongate rhomboidal spot in cell 1 @ almost coinciding in length with 
the hyaline spot in cell 2, an acuminate streak next the costa from the base of the wing 
as far as the apex of cell 10, and a narrow border to the hyaline spots bright orange- 
yellow: hind wing with the disc irregularly, and a streak along vein 14, orange-yellow. 
Underside: fore wing orange-yellow, with the costa and termen narrowly black; a 
subterminal band of blackish spots arising in a large subquadrate spot in the apex of 
cell 1 a, and decreasing in size towards the apex; a black streak from the end of the 
discoidal cell in part along the course of vein 5, and another in cell 8 continued 
towards the base along the upper edge of the discoidal cell; dorsum black as far as vein 
1a, and an oblong, black-edged, somewhat metallic pale spot in cell 1 @ from the base 
of vein 2 to the base of the wing: hind wing deep orange-yellow, the costa and termen 
narrowly black ; cell 1 @ and the lower half of cell 1 black, the former with an orange 
streak in its basal two-thirds; cell 1 orange-yellow ; disc of the wing with several 
irregular black dashes placed as follows, namely—two in cell 7, of which the basal one 
occupies two-thirds of the length of that cell, two in each of cells 2, 3, 4-5, and 6, one in 
the discoidal cell, and three in cell 1 4. Fringes below orange-yellow, the short scales 
on the fore wing blackish. Antenne blackish, the shaft yellow below from the base 
to the club. Clothing of palpi pale yellow, with a few black hairs intermixed. 
Clothing of body and legs orange-yellow. 

Expanse 34-36 mm. 

Hab. Pulo Laut (Doherty) ; Poeh, Sarawak (in coll. Rothschild). 

Described from two males in coll. Elwes, 


! PLASTINGIA LIBURNIA. 
Hesperia liburnia, Hewitson, Descr, Hesp. p. 33 (1868) ; id. Exot. Butt. v., Hesp. pl. vi. figs. 58, 59 
(1873). 
Plastingia liburnia, Semper, Schmett. Philipp. p. 813 (1892). 
Hab. Luzon, Mindoro, Camotes, Mindanao (Semper). 


PLASTINGIA TELESINUS. 


Carystus telesinus, Mabille, Pet. Nouv. Ent. ii, p. 205 (1878). 
Carystus lenas, Mabille, Comptes Rendus Soc. Ent. Belg. iv. p. exix (1891). 
Suastus telesonus, Semper, Schmett. Philipp. p. 800 (1892). 

Hab. Luzon, Bohol (Semper). 

This species seems to be most nearly allied to P. corissa, Hew., to which it bears 
considerable resemblance in the colour of the underside, but the latter species wants 
the pale spots on the hind wing below. 


A REVISION OF THE ORIENTAL HESPERIIDZ. 229 


! PLASTINGIA TESSELLATA. 
Hesperia tessellata, Hewitson, Trans. Ent. Soc. Lond. ser. 3, vol. ii. p- 494 (1866). 
Hesperia eulepis, Felder, Reise Nov., Lep. iii. p. 517, pl. Ixxii. fig. 12, g (1867). 
Plastingia tessellata, var. palawata, Staudinger, Iris, ii. p. 149 (1889). 
Hab. Celebes ( fide Hewitson) ; Palawan (fide Semper) ; Pulo Laut (Doherty) ; Battak 
Mountains, Sumatra (Martin). 


! PLASTINGIA NAGA. 


Hesperia? naga, de Nicéville, Jour. As. Soc. Beng. (1883) p. 89, pl. x. fig. 2, 2. 
Plastingia naga, Watson, Hesp. Ind. p. 115 (1891); Semper, Schmett. Philipp. p. 314 (1892). 


Hab. Sibsagar, Upper Assam (Peal, fide de Nicéville); E. Pegu (Doherty); Jaintia 
hills (coll. Swinhoe) ; Lawas, N. Borneo (Everett); Khasias (in coll. Rothschild) ; Battak 
Mountains, Sumatra (Martin) ; E. Mindanao (Semper). 

Watson (P. Z. 8. 1893, p. 118) puts this species as a synonym of tessellata, Hew., 
and as it resembles that species in all but the colour of the pale spots on the underside, 
we were, so long as we had seen female specimens only, inclined to adopt the same 
view, seeing that the type was a female; but having, through the kindness of Herr 
Semper, had the opportunity of examining one of his two males from E. Mindanao, 
which exactly agrees with the original description and figure as well as with female 
specimens in coll. Elwes, we are satisfied that it is a distinct species. It has been sent 
to us from Java by Staudinger as fessellata, Hew. 


PLASTINGIA VIBURNIA. 
Plastingia viburnia, Staudinger, in. litt.; Semper, Schmett. Philipp. p. 314 (1892). 

Of this very distinct species the female resembles the male, save that the pale 
markings on the upperside are yellowish white instead of deep yellow. 

Hab. Mindoro (Semper). 


! PLASTINGIA CORISSA. 
Hesperia corissa, Hewitson, Ann. & Mag. Nat. Hist. ser. 4, xvii. p. 455 (1876). 


Plastingia drancus, Plétz, Stett. ent. Zeit. xlv. p. 149 (1884). 
Tsoteinon indrasana, Elwes & de Nicéville, Jour. As. Soc. Beng. 1886, p. 441, pl. xx. fig. 5,2 (fide 


de Nicéville) ; Watson, Hesp. Ind. p. 86 (1891). 
Plastingia.latonia, Staudinger, MS. 

Hab. Kina Balu, Borneo (Waterstradt); Pulo Laut (Doherty); Battak Mountains, 
Sumatra (Martin). 

Mr. de Nicéville identifies J. indrasana, which he described in the joint paper on 
Tavoy butterflies, cited above, as J. corissa. As the type is now in the Calcutta Museum, 
we take his word for it, but the plate does not represent our specimens correctly. A 
female of P. latonia, Stgr., marked by Dr. Staudinger as “ typisch,” differs from the same 

vou. xiv.—Parr iv. No. 17.—Octaber, 1897. 20 


230 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


sex of P. corissa in being a little larger, and in having the hyaline spots in cells 6 and 7 of 
the fore wing above longer, an oblong hyaline spot in cell 8, and a small irregular yellow 
spot in cell 5 near the lower outer angle of the spot in cell 6. 


'Prastinera Nor. (Plate XXIV. figs. 57, 57 a.) 
Plastingia noémi, de Nicéville, Jour. As. Soc. Beng. 1885, p. 120, pl. ii. fig. 15, ¢. 
Hab. Sikkim (Mller, fide de Nicéville, Knyvett) ; Ei. Pegu (Doherty). 


!PLASTINGIA SIMILIS, n. sp. (Plate XIX. fig. 18, ¢; Plate XXIV. figs. 56, 56a.) 


Closely allied to P. noémi, de Nicév., from which it differs in its smaller size, the less 
conspicuous yellow streak along the basal half of the costa of the fore wing above, and 
in the particulars set forth in the above Table of Species. 

Expanse 54 mm. 

Hab. Pulo Laut (Doherty). 

Described from one male and two females in coll. Elwes. 


PLASTINGIA SUBMACULATA, 

Plastingia submaculata, Staudinger, Iris, ii. p. 149, pl. 11. fig. 8, ¢ (1889). 

Fore wing above with two cell-spots, a pale spot in each of cells 2 and 3, and a 
yellow streak from the base next the upper edge of vein 1a. Hind wing below yellow- 
green, the dorsum darker, three black spots near the base, and six others forming a 
postmedian curved series. 

Hab. Palawan (Platen). 

We have not seen this species, which may not be a Plastingia; the figure of the fore 
wing below does not agree with the author’s description. 


Loroneus. 
Lotongus, Distant, Rhop. Mal. p. 371 (1886) ; Watson, P. Z. S. 1893, p.121. Type calathus, Hew. 
Zea, Distant, t. c. pp. 369 & 377. Type mytheca, Hew. 
Zela, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 386. Type zeus, de Nicév. 
Zampa, de Nicéyille, t. c. p. 389. Type zenon, de Nicév. 

Antenne (in the male) about two-thirds as long as the costa; club slender ; apiculus 
acicular, reflexed, rather less than half as long as the club. Palpi appressed, densely 
scaled ; third joint minute, acicular or obtuse, almost concealed. Fore wing: termen 
(in the male) longer than the dorsum, vein 5 nearer 4 than 6, vein 2 from the basal 
third of the lower edge of the cell, no hyaline spot in cell 4; in calathus, zeus, and 
mytheca, the base of cell 3 is about twice as long as the base of cell 4, and in zenon, 
avesta, sarala, and excellens the base of cell 3 is about one-third longer than the base 
of cell 4. Hind wing: cell less than half.the length of the wing, termen very feebly 
excavated in cell 14. Epiphysis present; hind tibie fringed and bearing two pairs 
of spurs. 


A REVISION OF THE ORIENTAL HESPERIID®. 231 


We find that the following species fall within the above definition :— 


1 (10). Hind wing above plain brown. 
2 (9). Hind wing below brown, with or without a whitish border round the apex. 
3 (6). Cell-spot present on fore wing above. 
4 (5). No sex-mark on fore wing above in the male. Fringe of hind wing not 
yellow in the tornal region. Hind wing below frequently more or 
less broadly whitish round the apex . . . . . calathus, Hew. 
5 (4). Sex-mark on fore wing above in the male linear, blackish, erect fromm 
the middle of vein 1a to the basal third of vein 2, thence oblique to 
near the base of vein 3. Fringe of hind wing yellow in the tornal region. zeus, de Nicév. 
6 (8). No cell-spot on fore wing above. 
7 (8). Hind wing above in the male with a tuft of long hairs near the base 
completely covering the cell; the latter very short, about one-third 
as long as the wing, transverse vein suberect, veins 3, 4, and 6 
equidistant at the base and much thickened in their basal half. Hind 
wing below brown, yellowish next the transverse vein . . . zenon, de Nicév. 
8 (7). Hind wing above in the male normal; below brown, with a sharply 
defined yellow transverse band from the apical third of the costa to 
the basal third of the dorsum but interrupted in cell 1 4. No celi-spot 
on the fore wing above .. . . : P . . . dvesta, Hew. 
(2). Hind wing below brown, with a white Hetial band fork, the costa to 
vein 16, the breadth of this band equal to half the length of the wing. mytheca, Hew. 
10 (1). Hind wing above in part yellow. 
11 (12). Fore wing above with a cell-spot, no pale spots in cells 6,7, or 8. Hind 
wing below with a yellow band having the same direction as in avesta. sarala,de Nicév. 
12 (11). Fore wing above with a cell-spot and a pale spot in each of cells 6 and 7 
or 6,7, and 8. Hind wing below with a broad yellow band from the 
middle of the costa to the middle of cell 14, sometimes continued to 
the dorsum ov es Mob a olay eo en i won nenen es pits in) CH Siete 


! LoroNGUS CALATHUS. 


Eudamus calathus, Hewitson, Ann, & Mag. Nat. Hist. ser. 4, vol. xviii. p. 353 (1876). 
Lotongus calathus, Distant, Rhop. Mal. p. 371, pl. xxxiv. fig. 14 (1886). 
Lotongus maculatus, Distant, t.c. p. 372, pl. xxxv. fig. 1. 
Hesperia parthenope, Weymer, Stett. ent. Zeit. xlviii. p. 17, pl. ii. fig. 8, 2 (1887). 
Plesioneura aliena, Staudinger, Iris, 11. p. 155 (1889), sec. spec. typ. 
Lotongus parthenope, de Nicéyille, Jour. Bomb. Nat. Hist. Soc. 1892, p. 354, pl. J. figs. 49, 59. 
Proteides zalates, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 52 (1898). 
Proteides surus, Mabille, Bull. Soc. Hnt. Fr. 1895, p. lix. 
Hab. Nias (Modigliani); Palawan; Kina Balu, Borneo; Java (fide Staudinger) ; 
Selesseh, Battak Mountains, Sumatra (Martin). ; 
We have been obliged to unite the three supposed species of Lotongus upon which 


Mr. de Nicéville has written, we think, without sufficient material. We possess specimens 
2H 2 


232 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


of both sexes from Distant’s collection, with the label “ Malacca, Biggs.” The males are 
marked in his writing maculatus, the female calathus. A male from Pulo Laut agrees 
with these males. We have also three males and four females of parthenope, Weym.. 
from the typical locality Nias, which vary considerably inter se in the very characters by 
which alone they can be separated from the Malacca and Bornean specimens, namely, 
the size and number of the spots on the fore wing in the male, and in the female by 
the amount of white on the apex of the hind wing below, which is present in both 
sexes of calathus, but in some females only of parthenope. The Javan form figured by 
de Nicéville as parthenope 3 has only one spot in the cell, whilst two of our three 
males have two. The genitalia of the two forms are identical. 


! LoroNnGuUs ZEUS. 

Zela zeus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 388, pl. Q. fig. 57, g. 

We give a description of this species, which had been prepared before Mr. de Nicé- 
ville’s paper appeared. 

3. Upperside dark brown: fore wing with four hyaline whitish-yellow spots—one, 
about twice as high as wide, near the middle of cell 2, one, subquadrate, near the 
basal fourth of cell 3, one, oblong, near the base of cell 6, and one, longer than wide, 
in the cell, next the base of vein 3; sex-mark linear, blackish, erect from the middle of 
vein 1a to the basal third of vein 2, thence oblique to near the base of vein 3: hind 
wing with the basal two-thirds thickly clothed with long fuliginous hairs. Underside 
brown: hind wing and basal two-thirds of the costal region of fore wing bright yellow- 
brown, a small feeble yellow spot next the transverse vein of the hind wing. Fringes 
of the fore wing whitish grey; of the hind wing bright yellow in the tornal region, 
passing gradually into whitish grey towards the apex; short scales dark on both wings 
except on the brightest of the yellow part. Antenne brown; club slender, paler 
beneath; the apiculus equal in length to one-half of the clavate portion. Vein 2 of 
fore wing arising near the basal third of the cell. Body and legs concolorous with the 
upperside. 

Expanse 39-47 mm. 

Hab. Pulo Laut (Doherty); Mindoro (coll. Semper); Khasia hills (fide Swinhoe). 

This species is the type of Mr. de Nicéville’s genus Zelda. A male specimen from 
the Khasia hills, ex coll. Swinhoe, of which we have compared the genitalia, is larger 
than specimens from Pulo Laut, expanding 47 mm., and the pale spots on the fore 
wing are reduced in size, that proper to cell 6 being absent. ‘Type in coll. Elwes. 


! LoroNGUS ZENON. 
Zampa zenon, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 391, pl. Q. fig. 58, ¢. 


This species, of which the type is in coll. Elwes, is the type of Mr, de Nicéville’s 
genus Zampa. 


A REVISION OF THE ORIENTAL HESPERIID A. 233 


The curvature of the dorsum of the fore wing in the male is correlated to the supra- 
alar tuft on the hind wing, and is not peculiar to this species. Cf. Parnara oceia, 3. 
Hab. Pulo Laut (Doherty). 


! Loronaus AVESTA. 


Hesperia avesta, Hewitson, Descr. Hesp. p. 30 (1868). 
Lotongus avesta, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 383, pl. Q. fig. 56, 2. 
Pamphila tamiata, Stgr. MSS. 


Hab. Kina Balu, Borneo (Waterstr «dt); Pulo Laut (Doherty). 


! Loroneus MYTHECA. 
Hesperia mytheca, Hewitson, Ann. & Mag. Nat. Hist. ser. 4, vol. xix. p. 81 (1877). 
Zea mytheca, Distant, Rhop. Mal. p. 377, pl. xxxv. fig. 7 (1886). 


Hab. Perak (Doherty). 


! LoTONGUS SARALA. 
Parnara sarala, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, pl. B. fig. 6, 9; Watson, Hesp. 
Ind. p. 48. 


Hab. Khasia hills (Hamilton). 
Both sexes of this species, which do not differ, are in the Elwes collection. 


! LOTONGUS EXCELLENS. 
Proteides excellens, Staudinger, Iris, ii. p. 141, pl. 11. fig. 6 (1889). 
Hab. Pulo Laut (Doherty); Palawan (Platen, fide Stgr.) ; Sumatra (in coll. Rothschild). 


CRETEUS. 
Creteus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 385. Type cyrina, Hew. 

This is a genus erected by Mr. de Nicéville for the Hesperia cyrina of Hewitson, a 
species which is remarkable for the presence of hyaline spots on the hind wing, placed 
as follows :—one in the cell next the transverse vein, one in the basal third of each of 
cells 2 and 3, and one near the middle of each of cells 6 and 7. The hyaline spots on 
both wings are apt to be reduced in size, and in a specimen from Borneo in which this 
is the case the hyaline spot proper to cell 2 of the hind wing is wanting altogether. 
The sex-mark on the fore wing above in the male consists of a streak of modified scales 


occupying about the middle third of vein 2. 


! CRETEUS CYRINA. 
Hesperia cyrina, Hewitson, Ann, & Mag. Nat. Hist. ser. 4, vol. xviii. p. 450 (1876) ; Watson, 
Hesp. Ind. p. 160 (1891). 


234 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


Parnara parca, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 174, pl. B. fig. 10, 2. 
Pamphila meleagrina, Staudinger, MSS. 

Hab. Khasia (Hamilton); Kina Balu, Borneo (Waterstradt). 

All the spots on both the Bornean specimens in coll. Elwes are smaller than in the 
Khasia one, but there is no other difference. 


UNKANA. 


Unkana, Distant, Rhop. Mal. p. 369 (1886); Watson, P. Z. S. 1893, p. 123. Type batara, 
Moore, MS. 

A genus established by Distant for Zsmene batara, Moore, MS. 

Sexes dissimilar. Fore wing with the full complement of hyaline spots, 7. ¢. a cell- 
spot and one in each of cells 2 to 8 in both sexes. Antenne (in the male) more than half 
as long as costa; club slender ; apiculus acicular, about half as long as the club. 
Palpi appressed, densely scaled; third joint blunt, very short, almost concealed. Fore 
wing: termen in the male as long as or longer than the dorsum, cell 4 slightly narrowing 
to the base, vein 2 from the basal third of the lower edge of the cell, transverse vein 
oblique, base of cell 3 about twice as long as that of cell 4. Hind wing: termen 
angularly produced at vein 1, cell less than half the length of the wing. Epiphysis 
present; hind tibie fringed and bearing two pairs of spurs, the submedian pair small 
and appressed. 

Watson, in his ‘ Revision,’ places this genus (presumably after having examined the 
type species) in his Pamphiline, Section B, and Erionota in his Pamphiline, Section A. 
In his more recent paper, before referred to, he says that batara correctly belongs to 
the genus Hrionota, and sinks the genus Unkana accordingly ; but, since batara differs 
from thraz, the type of Zrionota, much more than thrax does from thyrsis, the type of 
Gangara, there seems to be, even on his own estimate of genera, no good reason for 
such a course. 

So far as we can judge, the Erionota (Casyapa) mabillei, Stgy., belongs here rather 
than to Erionota, and it is placed in this genus accordingly. 


! UNKANA ATTINA. 


Tsmene batara, Moore, MS.; Horsfield & Moore, Cat. Lep. H.I. C. i. p. 249 (1857), 3. 
Hesperia attina, Hewitson, Trans. Ent. Soc. Lond. ser. 3, vol. ii. p. 489 (1866), 2. 
Hesperia latreillei, Felder, Reise Nov., Lep. iii. p. 511, pl. Ixxi. fig. 8 (1866), 2. 
Goniloba cruda, Herrich-Schiaffer, Prod. Syst. Lep. iii. p. 75 (1869). 
Unkana batara, Distant, Rhop. Mal. p. 370, pl. xxxiv. fig. 11, f (1886). 
Unkana attina, Distant, t. ec. p. 371, pl. xxxiy. fig. 830, 9 ; Watson, Hesp. Ind. p. 4 (1891). 
Hab. Malacca (Biggs); Perak (Doherty); Moulmein (Adamson); Java (Fruhstorfer) 
Palawan (Platen); Mindanao (Semper). 


A REVISION OF THE ORIENTAL HESPERIIDA, 235 


UNKANA MABILLEL 
Erionota (Casyapa) mabillei, Staudinger, Iris, vol. ii. p. 135 (1889). 

The male differs from that sex of U. attina, Hew., on the upperside by the want of 
pale spots in cells 4 and 5 and the different shape and sordid yellow colour of the 
remaining spots, that in the cell being one and 4 half times as wide as high; the hind 
wing below is brown, with some dull red spots near the lower angle of the cell, a broad 
irregular dark brown band gradually narrowiny from the costa and almost reaching 
vein 14, and a large brown subtriangular patch next the base. Fringe of the fore 
wing whitish grey, the short scales brown; of the hind wing brown from the apex to 
vein 4, thence pale yellowish grey. 

These particulars are taken from an example, ex coll. Staudinger, in which the hind 
wing below is much rubbed. 

Hab. Labuan (coll. Stgr.); Palawan (Platen, fide Stgr.). 


Hipari. 

Hidari, Distant, Rhop. Mal. p. 395 (1886) ; Watson, P. Z.S. 1893, p.123. Type irava, Moore. 

Fore wing with the termen (in the male) as long as or longer than the dorsum; hind 
wing short, rounded; abdomen about one-fourth longer than hind wing. Antenne 
about half as long as costa; club slender; apiculus acute, less than half as long as 
club. Palpi appressed, second joint densely scaled, third joint very short, blunt, almost 
concealed. Fore wing: vein 2 from the basal third of the lower edge of the cell, trans- 
verse vein oblique, base of cell 4 about half as long as the base of cell 3. Hind wing 
with the cell about half as long as the wing. 

Distant includes in this genus three species, which we distinguish as follows :— 
1 (4). Fore wing above with the pale spots free. 
2 (8). Fore wimg above with a pale spot near the middle of cell 1a next vein la, 

but, normally, no pale spots in cells 7 and 8; hind wing below with a 


small pale spot in the cell, near the base of vein6 . . . . : irava, Moore. 
3 (2). Fore wing above with no pale spot in cell 1a, and a pale spot in cach of 
cells 6, 7, and 8; no pale spot in the cell on hind wing below. . . sybirita, Hew. 


4 (1). Fore wing above with the pale spots confluent and forming a broad 
irregular deep yellow band from the upper edge of the cellto vein 1 a. staudingeri, Dist. 


! HipaRi IRAVA. 
Hesperia irava, Moore; Horsfield & Moore, Cat. Lep. Mus. E.I. C. i. p. 254 (1857); Plotz,. Stett. 
ent. Zeit. xlii. p. 328 (1882). 
Hidari irava, Distant, Rhop. Mal. p. 395, pl. xxxiv. fig. 15, 2 (1886) ; Watson, Hesp. Ind. p. 112 
(1891). 
Hab. Perak, Bali (Doherty); Java (Piepers). 


236 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


HIIDARI DOESOENA. 


Hidaria doesoena, Martin, Binige neue Tagschmetterlinge von Nordost-Sumatra, p. 6 (1895). 


Differs from H. irava, Moore, in having the pale spot in cell 2 of the fore wing only 
separated from the cell-spot by the lower boundary of the cell, and the small pale spot in 
the cell of the hind wing below white instead of yellow. We have not seen this species. 

Hab. N.E. Sumatra (Martin). 


HIDARI SYBIRITA. 


Hesperia sybirita, Hewitson, Ann. & Mag. Nat. Hist. ser. 4, vol. xviii. p. 451 (1876). 
Hidari sybirita, Distant, Rhop. Mal. p. 395, pl. xxxv. fig. 24 (1886). 


Hab. Singapore (coll. Hewitson). 


! HIDARI STAUDINGERI. 
Hidari staudingeri, Distant, Rhop. Mal. p. 395, pl. xxxv. fig. 25 (1886). 


Hab. Perak (Doherty). 


HIDARI BHAWANI. 


Hidari bhawani, de Nicéville, Jour. As. Soc. Beng. 1888, p. 291, pl. xi. fig. 6, ¢; Watson, Hesp. 
Ind. p. 112 (1891). 


Resembles H. irava in the spotting of the fore wing above, but the hind wing below 
is pale ochreous, coarsely striated transversely with brown, and there is a brown streak 
near the costa and another next the dorsum. We have not seen this species. 

Hab. Arracan coast, Burma (Bingham, fide de Nicéville). 


EEtTIon. 
Eetion, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 395. Type elia, Hew. 


This is a genus erected by de Nicéville for the Hesperia elia of Hewitson, a species 
which is well distinguished from its immediate allies by the presence of hyaline spots 
on the disc of the hind wing; these form a transverse row and are placed one in each of 
cells 1b, 2, 3, and 4-5. ‘There is in the male a tuft of long hairs near the base of the 
dorsum of the fore wing below, and on the upperside of the fore wing a seam of 
modified scales passing obliquely inwards and downwards from the base of vein 38, 
nearly half of its length lying below vein 2. 


! EETION ELTA. 


Hesperia elia, Hewitson, Trans. Ent. Soe. Lond. ser. 3, vol. ii. p. 489 (1866). 
Unkana elia, Distant, Rhop. Mal. p. 370, pl. xxxiv. fig. 25 (1886). 


Hab. Singapore (Godfrey); Perak (Doherty). 


A REVISION OF THE ORIENTAL HESPERIIDA. 237 


PITHAURIA. 
Pithauria, Moore, P. Z. 8. 1878, p. 689; Watson, op. cit. 1893, p. 119. Type murdava, Moore. 
Pithauriopsis, Wood-Mason & de Nicéville, Jour, As. Soc. Beng. 1886, p. 387; Watson, 1. ec. 
Type aitchisoni, Wood-Mason & de Nicéy. 

Fore wing rather narrow and pointed ; thorax very stout ; abdomen conical, not longer 
than the dorsum of the hindwing. Antenne about half as long as costa; club slender; 
apiculus acicular, reflexed, at least one-half as long as the club. Palpi appressed, 
densely scaled, third joint almost concealed. Fore wing: no hyaline spot in cell 4, 
vein 2 arising near the half-length of the cell, vein 5 nearer 4 than 6. Hind wing: 
cell Jess than half as long as the wing, angulated at the origin of vein 7 and also at the 
origin of vein 2, vein 5 absent ; termen feebly curved, nearly straight between veins 1 a 
and 7; tornus not distinctly produced. Epiphysis present ; hind tibie with two pairs of 
spurs. Front coxe in the male densely clothed with long hair-like scales, and in the 
same sex the disc of the hind wing above is densely clothed with long hair-like scales. 

There are three species separable as follows :— 


Males. 
1 (4). No sex-mark on the fore wing above. 
2 (3). The long clothing of hind wing above not conspicuously pale. mwrdava, Moore. 
3 (2). The long clothing of hind wing above whitish grey . . . stramineipennis, de Nicéy. 
4. (1). A double sex-mark near the middle of cell 1 a on fore wing 
above, precisely similar to that found in the genus Halpe. aitchisoni, W.-M. & de Nicéy. 


! PITHAURIA MURDAVA. 
Ismene murdava, Moore, P. Z. 8. 1865, p. 784. 
Pithauria murdava, Moore, P. Z. 8S. 1878, p. 689, pl. xlv. fig. 13; Distant, Rhop. Mal. p. 378, 
pl. xxxv. fig. 9, d (1886) ; Watson, Hesp. Ind. p. 27 (1891). 

The female of this species has on the hind wing below a suffused subterminal pale 
band, and an oblong pale spot in each of cells 6 and 7, being thus distinguished 
from the female of P. stramineipennis, in which the hind wing below is uniform grey by 
reason of a sprinkling of whitish-grey scales on a pale greenish-brown ground ; in one 
of my two specimens of the latter there is a feeble pale spot near the basal third of 
cell 6 on the hind wing below. 

Hab. Sikkim (Moller); Khasia (Hamilton). 


! PITHAURIA STRAMINEIPENNIS. 


Pithauria stramineipennis, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 388, 
pl. xx. fig. 5, ¢d; Watson, Hesp. Ind. p. 27 (1891) ; Leech, Butt. China &c. p 631, pl. xli. 
fig. 19, d (1894). 
Hab. Sikkim (MGller) ; Nagas (Doherty) ; Tavoy (Tucker). 
Vou. XIv.—Part iv, No. 18.—October, 1897. 21 


238 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! PITHAURIA AITCHISONT. 
Pithauriopsis aitchisoni, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 387, pl. xv. 
fig. 4, ¢; Watson, Hesp. Ind. p. 28 (1891). 
Pamphila glauca, Staudinger, in litt. 

@. Hind wing above brown, without the grey clothing found in the male. Hind 
wing below red-brown, paler along the dorsum, with the following pure white markings, 
namely :—a streak from the base along cell 7 as far as its middle, a streak in the basal 
half of cell 6, but not reaching the base of that cell, a series of four small oblong 
spots, one near the middle of each of cells 2, 3, 4, and 5, and sometimes another beyond 
the middle of cell 16. Otherwise like the male. 

Described from a specimen from Kina Balu, Borneo, ex coll. Staudinger. 

Hab. Kina Balu, Borneo (Staudinger) ; Java (Piepers). 


Norocrypra. 
Plesioneura, Felder, Wien. ent. Monatsb. vi. p. 29 (1862), nom. preocc. Type curvifascia, Felder. 
Notocrypta, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 188; Watson, P. Z. 8S. 1893, 
p- 112. Type curvifascia, Felder. 

Antenne about two-thirds as long as costa, rather shorter in the female. Palpi 
ascending ; third joint porrect, almost concealed. Fore wing: vein 5 arising much 
nearer 4 than 6, and having a distinct upward curve, or straight and practically inter- 
mediate between veins 4 and 6; vein 2 from the basal third of the lower edge of the 
cell. Hind tibiz with two long pairs of spurs. 

Species blackish brown above, with white hyaline markings on the fore wing, of 
which the most prominent is a more or less complete broad band passing from the 
middle of the costa to the distal third of the dorsum. 

In ‘ Iris,’ vol. ii. p. 151, Dr. Staudinger attempts to show that signata, Druce (P. Z. 8. 
1873, p. 360, pl. xxxiii. fig. 8), is an aberration of alysos, Moore; but this view is 
scarcely tenable, because, apart from the differences in the venation in the two insects, 
the band on the fore wing in signata, Druce (recte Charmion ficulnea, Hew.), is cut 
short by vein 2, which is not the case in alysos, Moore (recte feisthamelit, Bdv.). 

The species dealt with in this paper may be distinguished as follows :— 


1 (6). Vein 5 in the fore wing curved, its basal portion receding 
from vein 6; yein 5 therefore arising much nearer to 
vein 4 than to vein 6. 
2 (5). Hind wing below without white spots. 
3 (4). General direction of the termen of the fore wing forming 
amore or less obtuse angle with the dorsum. Fore 
wing above with or without white spots besides those 
forming the band ......... . . © (feisthameli, Bdv. 


A REVISION OF THE ORIENTAL HESPERIIDA. 239 


4 (3). General direction of the termen of the fore wing forming 
a right angle with the dorsum. Fore wing above 


without white spots except those forming the band . quadrata, n. sp. 

5 (2). Hind wing below with an angular white spot near the 
distal third of the cell and another near the basal third [de Nicév. 
ofecelll2 ees 4. : . paralysos, Wood-Mason & 


6 (1). Vein 5 in the fore wing straight, practically intermediate 
between veins 4 and 6. 

7 (10). Base of cell 3 on the fore wing not filled up with white. 

8 (9). Basal fifth or fourth of hind wing below yellow. 


Subtornal blotch on fore wing below obsolete or 
basiflava, de Nicév. 


wanting PE er Mt ithe DERE ep acl eds) Ee 
9 (8). Hind wing below pale brown. Subtornal blotch on fore 
wing belowsordid white . ..... =. . . -. tornatd,n. sp. 
10 (7). Base of cell 3 on the fore wing filled up by a triangular [de Nicéy. 


white spot . monteitht, Wood-Mason & 


NovocrRyPTa FEISTHAMELIL. 
Thymele feisthamelii, Boisduval, Voy. Astr., Lép. p. 159, pl. ii. fig. 7 (1832). 
Plesioneura curvifascia, Felder, Wien. ent. Mon. vi. p. 29 (1862). 
Plesioneura alysos, Moore, P. Z. S. 1865, p. 789 ; id. Lep. Cey. i. p. 178, pl. Ixvii. figs. 3 ¢,3a Q, 
30 larva and pupa (1881) ; Distant, Rhop. Mal. p. 399, pl. xxxiv. fig. 7, g (1886). 
Plesioneura albifascia, Moore, P. Z. 8. 1878, p. 848, pl. lini. fig. 8, 3. 
Plesioneura restricta, Moore. Lep. Cey. i. p. 178 (1881); Wood-Mason & de Nicéville, Jour. As. 
Soe. Beng. 1887, p. 390, pl. xvii. fig. 5, ¢- 
Plesioneura voluw, Mabille, Ann. Soc. Ent. Belg. 1883, p. lvi. 
? Plesioneura clavata, Staudinger, Iris, ii. p. 153, pl. ii. fig. 9 (1889). 
Notocrypta alysos, Watson, Hesp. Ind. p. 126 (1891). 
Notocrypta albifascia, Watson, Hesp. Ind. p. 128 (1891). 
Notocrypta feisthameli, var. rectifascia, Leech, Butt. China, &e. p. 627, pl. xxxvin. fig. 2, g (1894). 
Notocrypta curvifascia, Leech, t. c. p. 626, pl. xxxviil. fig. 1, ¢. 
Notocrypta restricta, Leech, t. c. p. 627, pl. xxxviii. fig. 3, g ; Watson, Hesp. Ind. p. 128 (1891). 
Judging from the male genitalia it seems probable that there is but one species of 
this genus with the fringe of the hind wing grey-brown, the white band on fore wing 
above not cut short by vein 2, and having a dark sub-basal and median band, but no 


white spot on the hind wing below. ‘The oldest name given to an insect falling within 


this definition is that of feisthamelii, Bdv. The presence or absence of small white 
spots on the fore wing, an opaque white patch on the fore wing below continuing the 
white band to the costa, or a white dash in each of cells 9 and 10 on the fore wing 
below between the hyaline spot in cell 8 and the costa, are particulars not sufficiently 
constant to be of any use as specific characters. The form albifascia, Moore, which we 
have from Java and Pulo Laut, a comparatively small insect with no white markings 


except the rather narrow band on the fore wing above, is very different in appearance 
212 


240 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


from the much larger typical feisthamelii (which we have also from Jaya), with its broad 
white band on the fore wing and five white spots besides; but there is nothing in the 
male genitalia of the two forms to prevent us from regarding the former as a 
depauperate local form of the latter. 

The main differences in the several named forms consist in the greater or lesser 
number of white spots on the fore wing above, exclusive of those forming the discal 
band, and in a lesser degree the differences in the shape of the band itself and whether 
or not it is extended to the costa on the underside by a pale patch, Beyond those 
which form the discal band the largest number of white spots found on the fore wing 
above appears to be seven, namely, one in each of cells 3-9 inclusive ; all or any of these 
may be absent, and specimens might in this respect be grouped under three heads : 
(a) those with no white spots, except those forming the band ; (6) those with one spot, 
that in cell 4; and (c) those with more than one spot, 7. e. with any number from 2 to 7, 
and of these the specimens with five or six spots are perhaps the most numerous. Such 
a grouping, however, would be purely artificial and would answer no useful purpose. 

Though at first sight it might seem that we have united several forms more distinct 
inter se than others which we have previously treated as good species, yet a careful 
study of no less than 35 males and 26 females from all parts of India and the Malay 
Islands, representing all the named forms, together with the fact that the genitalia of 
all the specimens examined seem to be identical in structure, convinces us that it is 
impossible to separate them. ‘The principal points which have been relied on by 
authors are the number of the spots beyond the band on fore wing above, the form of 
the band, and whether or not it is extended to the costa below. 

P. albifascia, Moore, is a small form from Java, Borneo, and Bali, with a narrow 
band and usually no spots beyond the band. 

P. volux, Mab., of which we have seen the type in Dr. Staudinger’s collection, 
is also spotless. 

Mr. Leech, who has given much attention to these insects, divides the specimens 
found in the region treated of in his book as follows :— 


Pale band on fore wing below continued to the costa by a pale patch . . . . feisthameli, Bdy, 
Pale band not so continued. 
Fore wing below with a white dash in each of cells 9 and 10 between the hyaline 
Spotin cell’$ and the'costa- 9. 5 2 5 2s 5 ses ws Curvifascia, Held: 
Fore wing with no such dashes. . .-. . =... =. =. -+ = « « « « 7estricta, Moore, 


In this connection I may say that I have specimens from Jaya which combine the 
characters relied on by Mr. Leech for the separation of feisthameli and curvifascia. 

Hab. India, generally distributed. Burmah, Perak, Andamans, Bali (Doherty); Java 
(Piepers) ; Nias (Modigliant) ; Philippines (Semper) ; Borneo, Arjuno, Java (Doherty) ; 
China (Leech); Liu Kiu Islands (Pryer). 


A REVISION OF THE ORIENTAL HESPERIIDA. 241 


! NorocryPTA GUADRATA, n. sp. (Plate XX. fig. 3, 3.) 

3. Nearest to WV. feisthamelii, from which it differs in the general direction of the 
termen of the fore wing, which forms approximately a right angle with the dorsum, 
and in having the white band, which is cut short by vein 1 a and the upper edge of the 
cell, broader and more regular in outline ; there are usually no hyaline spots, except 
those forming the band, but this is not a feature of much importance, as one specimen 
has a hyaline point near the middle of cell 4. On the underside the band is continued 
to the costa by an opaque white patch. 

Expanse 363-384 mm. 

Hab. Kina Balu, Borneo ( Waterstradt), 

Described from two specimens in coll. Elwes received from Dr. Staudinger, who has 
others from the same locality. 


! NoTOcRYPTA PARALYSOS. 
Plesioneura paralysos, Wood-Mason & de Nicéville, Jour. As. Soc, Beng. 1881, p. 257. 
Notocrypta paralysos, Watson, Hesp. Ind. p. 127 (1891). 

Though this species has the genitalia characteristic of WV. feisthamelii, the pure white 
spots on the hind wing below, two in number, distinguish it with certainty. 

Hab. Andamans (de Roepstorff). 


! NOTOCRYPTA BASIFLAVA. 
Plesioneura basiflava, de Nicéville, Jour. As. Soc. Beng, 1888, p. 290, pl. xiii. fig. 7, 3. 
Notocrypta basiflava, Watson, Hesp. Ind. p. 130 (1891). 

Hab. Nilgiris (Hampson). 


NovocRyPTA INORNATA, n. sp. (Plate XXI. fig. 15, 3.) 

3. Upperside dark brown: fore wing with a large oblong cell-spot, notched on its 
outer edge, and a large irregular oblong spot occupying about the middle third of 
cell 2 pure white, and forming together an oblique abbreviated discal band; a white 
point near the basal third of cell 3, Underside of fore wing as the upperside, save 
that the outer half of cell 1a@ is almost entirely occupied by the white subtornal 
blotch. Body, antennz, and legs concolorous with the wings; club of the antennz 
whitish beneath. Second joint of palpi clothed with black and grey hairs intermixed. 

Expanse 41 mm. 

Hab. Kina Balu, Borneo (Waterstradt). 

Described from a single example in coll. Staudinger. 


NovocrYPra MONTEITHI, 
Plesioneura monteithi, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 391, pl. xviii. 
figs. 3, 3a, 2. 
Notocrypta monteithi, Watson, Hesp. Ind. p. 129 (1891). 
Carystus singularis, Mabille, Ann. Soc. Ent. Belg. xxxvii. p, 51 (1893). 


242 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


This species differs from any form of WV. feisthamelii in having on the fore wing a 
triangular spot, which occupies the basal third of cell 3, included in the white band. 
Very few specimens are known, all females. 

Hab. Irangmara, Cachar (Wood-Mason & de Nicéville); Sumatra (fide de Nicéville) ; 
Java ( fide Mabille) ; N. Borneo (Pryer, in coll. Rothschild). Cf. p. 215 ante. 


CERANE, gen. nov. 


Antenne two-thirds the length of the costa; club slender; apiculus about one-fourth 
as long as the club. Palpi laxly scaled, second joint ascending, third long, acicular, 
erect. Fore wing (male): costa one-third longer than the dorsum; termen straight 
from the tornus to vein 5, subequal in length to the dorsum, vein 5 with its basal 
portion receding from vein 6 and therefore arising evidently nearer to vein 4 than to 
vein 6. Species blackish brown above, with an oblique white abbreviated discal band 
between vein 2 and the upper edge of the cell. Hind tibize with two pairs of spurs. 

This genus contains at present the two species mentioned below, and as both of 
them exhibit the characters given above it cannot be said that one is more typical 
than the other. 

They may be thus distinguished inter se :— 

Subtornal blotch on fore wing below pale brown. Fore wing (male) apparently 


narrower, by reason that the angle between the dorsum and the straight 


part of the termen is about 115° ah agit \acut eo .0 neera, de Nicév. 
Subtornal blotch on fore wing below white, almost as pure as the discal band. 
Fore wing (male) apparently broader, by reason that the angle between the 


dorsum and the straight part of the termen is about 100°. . . . . . microthyrus, Mab. 


!CERANE NERA. 

Notocrypta neera, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 579, pl. G. fig. 27, °. 
Hab. Pulo Laut, Perak (Doherty) ; Kina Balu, Borneo ( Waterstradt). 
There is a specimen in the Hewitson collection from Singapore unnamed. 


CERANE MICROTHYRUS. 
Plesioneura microthyrus, Mabille, Comptes Rendus Soc. Ent. Belg. iii. no. 31, p. lvii (1883). 
Plesioneura mindorana, Staudinger, in litt. 
Notocrypta microthyrus, Semper, Schmett. Philipp. p. 317 (1892). 

Hab. Luzon, EK. Mindanao (Semper); Mindoro (coll. Staudinger). 

Herr Semper was mistaken in his statement that he had not the female of this 
species, the specimen submitted to us is undoubtedly of that sex. 

In this species the pale band on the fore wing below is continued to the costa, and 
there is a tendency to the acquisition of this character in some females of G2. newra. 


A REVISION OF THE ORIENTAL HESPERIIDA. 243 


UDASPES. 
Udaspes, Moore, Lep. Cey. i. p. 177 (1881) ; Watson, P.Z. S. 1893, p. 113. Type folus, Cr. 


Antenne about half as long as costa; club gradual; apiculus acute, about two-thirds 
as long as the club. Palpi porrect; third joint short, almost concealed. Fore wing: 
dorsum longer than the termen, the latter nearly evenly curved ; vein 5 with its basal 
portion receding from vein 6 and therefore arising much nearer 4 than 6; vein 2 
arising before the half-length of the cell, Hind wing nearly as broad as long ; 
evenly rounded; cell less than half as long as the wing. Hyaline spots present in both 
wings. ‘Tibial epiphysis present; hind tibie fringed and bearing two pairs of spurs. 
No alar sex-mark in the male. 

The two species known to us may be distinguished as follows :— 


termen 


Disc of the hind wing above white, completely surrounded by a broad dark brown 
border. Expanse about 42 mm. oP rer oe WD Cees Belted ate . folus, Cr. 
Hind wing above fuliginous brown, with a sharply defined rhomboidal white spot on 
the disc standing on vein 5 near the middle, and a white point just below it near 
the basal third of cell 3. Expanse about 33mm. . .... =... . .~ stellata, Ob. 


! UDASPES FOLUS. 
Papilio folus, Cramer, Pap. Exot. i. pl. Ixxiv. fig. 7 (1779). 
Udaspes folus, Moore, Lep. Cey. i. p. 177, pl. Ixviii. figs. 3, 3. @ (1881) ; Distant, Rhop. Mal. p. 398, 
pl. xxxiv. fig. 3 (1886) ; Watson, Hesp. Ind. p. 125 (1891). 
Hab. N.W. Himalaya (Young); Sikkim (Moller); Khasia (Elwes); Burmah 
(Watson, Doherty) ; Java (Piepers); Sambawa, Bali (Doherty). 


! UDASPES STELLATA. 
Plesioneura stellata, Oberthiir, Etudes d’Ent. xx. p. 41, pl. ix. fig. 165 (1896). 


Hab. Menia, ? prope Ta-tsien-lo, E. Tibet (coll. Oberthiir). 
I am indebted to M. Charles Oberthiir for a specimen of this distinct species, 
which he has recently received from his native collectors in Tibet. 


ACTINOR. 


Actinor, Watson, P. Z. S. 1893, p. 108. Type radians, Moore. 

This genus was erected by Watson for the Halpe radians of Moore, an insect 
combining with its own peculiar facies most of the other characters of Halpe, save that 
the sex-mark is absent in the male and vein 2 of the fore wing arises near the basal 
third of the cell. 

The pattern of the hind wing below is quite peculiar; the veins and two narrow 
irregular straight transverse bands are yellowish white, the antemedian band starts from 
-the basal third of vein 1 6 and passes over the base of vein 2 obliquely across the cell 


244 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


and is cut short by vein 8, the postmedian band starts from the apical fourth of 
vein 14 and is continued in an almost regular zigzag to the apex of vein 6; there is a 
pale spot near the base of cell 7. 


! ACTINOR RADIANS. 
Halpe radians, Moore, P. Z. 8. 1878, p. 690, pl. xlv. fig. 1; Watson, Hesp. Ind. p. 74 (1891). 


Hab. Mandi, N.W. Himalayas (Young). 


GEHENNA. 
Gehenna, Watson, P. Z. S. 1893, p. 108. Type abima, Hew. 


Distinguished from Halpe by the secondary sexual characters on the underside of the 
fore wing in the male, which consist of a thickening of the basal part of vein 2 anda 
portion of the lower edge of the cell near it, so as to form a swollen <, which is partly 
covered by a thick tuft of long black hairs springing from the dorsum near the base. 

Three species are known to us, and may be distinguished as follows :— 


1 (4). Male. Hind wing: vein 8 and the upper edge of the cell normal. 
2 (8). Cell-spot on the fore wing geminate; hind wing below with “five un- 
defined brown spots” (de Nicév.). . . . . + P65 0 6 oo Gm lel 
3 (2). One moderately large spot next the lower edge of the cell on the fore 
wing, sometimes with another punctiform one just above it. Hind 
wing below brown, more or less closely covered with greyish-yellow 
scales, except on a large spot near the end of the cell and a triangular 
postmedian spot ; the base of this brown triangle occupies about the 
basal half of vein 2, and its apex just reaches ven 6... . . angulifera, n. sp. 
4 (1). Male. Hind wing: vein 8 and the upper edge of the cell fein a 
“tuning-fork ” at their junction; upperside with a small oval oblique 
streak close to the base; underside dark reddish brown, immaculate, 
“frosted over with scattered golden-yellow scales” . . . . . . . gree, de Nicéy. 


GEHENNA ABIMA. 
Hesperia abima, Hew. Aun. & Mag. Nat. Hist. (4) xix. p. 83 (1877). 


Hab. Macassar (fide Hewitson). 


GEHENNA ANGULIFERA, n. sp. (Plate XIX. fig. 20, 3.) 

g. Upperside brown: fore wing with four or five sordid yellowish-white spots 
placed as follows :—one near the apex of the lower edge of the cell, sometimes with a 
punctiform one just above it, one about twice as high as wide in cell 2, one roundish 
or subquadrate near the basal third of cell 3, and one small and roundish in cell 6. 
Underside: fore wing as on the upperside, but paler along the dorsum, and with the 


A REVISION OF THE ORIENTAL HESPERIID#. 245 


extra-discal portion more or less closely sprinkled with greyish-yellow scales: hind 
wing brown, more or less closely sprinkled with greyish-yellow scales, except on a 
spot near the end of the cell, and a curved postmedian band, the base of which 
occupies about the basal half of vein 2, and its apex just reaches vein 6. Fringes 
dark grey, the short scales darker, especially on the hind wing. Antenne above 
dark brown, very finely spotted with white beneath, apiculus pale red on the 
underside. Body above brown ; clothing of the palpi, breast, and legs of yellow and 
black hairs intermixed; abdomen beneath blackish, hind margins of the segments with 
a narrow band of whitish-grey scales. 

Expanse 28 mm. 

Hab. Mindoro. 

Described from two specimens ex coll. Staudinger. 

According to a specimen received from M. Mabille, this is the same as his Pamphila 
rama; we have not, however, been able to find any reference to the publication of the 


latter name. 


GEHENNA GRAM. 
Gehenna gree, de Nicéville, Jour. Bomb, Nat. Hist. Soc. 1895, p. 399. 


Hab. N.E. Sumatra (fide de Nicéville). 


Curitua, Moore. 
Cupitha, Moore, Jour. As. Soc. Beng. 1884, pt. ii. p. 47; Watson, P. Z.S. 1893, p. 111. Type 
purreea, Moore. 

A genus of a single species, best distinguished by the secondary sexual characters of 
the male. The pattern of the upperside is essentially the same as that found in 
Telicota gola, Moore. Male: fore wing with the dorsum angularly produced just 
before the middle, on the underside with a tuft of long hairs on the dorsum between 
the base and the angulation ; hind wing above with an oval depression at the end of 
the cell filled with what appears to be a waxy matter, the middle of this depression is 
traversed by the confluent basal portion of veins 3 and 4, which thus form a loop 


above the lower margin of the cell. 


! CUPITHA PURREEA. 
Pamphila purreea, Moore, P. Z. S. 1877, p. 594, pl. lviii. fig. 10. 
Cupitha tympanifera, Moore, Jour. As. Soc. Beng. 1884, p. 48. 
Pamphila lycorias, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 54 (1893). 
Hab. Sikkim (Moller); E. Pegu, Bali (Doherty); Java (Fruhstorfer); Nias 
(Modigliani) ; Camiguin de Mindanao (Semper); Palawan (coll. Staudinger). 


VOL. XIV.—PART ivy. No. 19.—October, 1897. 2K 


246 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


AUGIADES. 
Augiades, Hiibn. Verz. p. 112 (1816) ; Watson, P. Z. 8. 1893, p. 103. Type sylvanus, Esper. 


Closely allied to Erynnis, the points of difference being practically those which 
separate the Papilio sylvanus of Esper from the Papilio comma of Linneus. In 


Augiades siva, Moore, and its immediate allies, hyaline spots are more or less strongly 
developed in the fore wing. 


Antenne rather more than half as long as costa; club elongate, oval; apiculus longer 
than in Hrynnis, in most species as long as the greatest width of the club. Third 
joint of palpi almost concealed, second joint laxly scaled. 

In seven species examined there occurs a marked peculiarity in the male genitalia ; 
the cedeagus has growing out of its lower side one or two large dissimilar horns, 
variously shaped and armed, sometimes nearly as large as the organ itself. 

The species in coll. Elwes are distinguished as follows :— 


1 (10). Hind wing below with the veins concolorous. 

2 (8). Fore wing below-with the blackish colour confined to the base and 
dorsum, bounded externally by the sex-mark, and above by 
vein 1 a ¢ 

3 (2). Fore wing below with the dete half eaied iy a  Sultiesed bieckioh 


patch, which is bounded above by the lower margin of the cell 
and vein 3. 


4 (7). Hind wing above with pale spots in cells 2-6. 
5 (6). Size larger. Fore wing below with a pale suffused spot near the 
middle of the dorsum. Cideagus with one horn apne 
6 (5). Size smaller. Fore wing below without a pale spot near the aneldie 
of the dorsum. (#deagus with two horns 
7 (4). Hind wing above with pale spots in cells 2, 3, and 6. 
8 (9). Sex-mark black. Fore wing below with the basal half of the costal 
region nearly concolorous with the ground-colour of the hind wing. 
Hind wing below with the pale spots silvery white, dark-edged .  siva, Moore. 
9 (8). Middle streak of the sex-mark whitish. Fore wing below with the 
basal half of the costal region reddish ochreous. Hind wing 
below with the pale spots neither silvery white nor dark-edged. 
9a (9b). Expanse 33-85 mm. Lower distal angle of the lower lobe of the 
clasp acute Tuc tice she ah" Sis) eee 
b (9a). Expanse 483-47 mm. Lower distal angle of the lower lobe of the 
clasp obtuse . : ; 
10 (1). Hind wing below with the veins Place 
1] (12). Size smaller. Fore wing less pointed, its underside with the fringe 
deep yellow, the short scales dark brown for a varying distance 
from the apex. Cdeagus with one horn erence rs 
12 (11). Size larger. Fore wing more pointed, its underside ith ite fringe 


yéllowish grey, passing into yellow near the tornus. (&deagus 
with two horns 


sylvanus, Esp. 


subhyalina, Brew. 


sylvanoides, Leech. 


brahma, Moore. 


crateis, Leech. 


ochracea, Brem. 


majuscula, . sp. 


A REVISION OF THE ORIENTAL HESPERIIDA, 247 


! AUGIADES SYLVANUS. 
Papilio sylvanus, Esper, Schmett. i. 1, pl. xxxvi. fig. 1 (1778?) ; Hiibner, Eur. Schmett. i. 
figs. 482-484. (1798-1803). 
Hesperia venata, Bremer & Grey, Schmett. N. China’s, p. 11, pl. iii. fig. 5 (1853) ; Ménétriés, Cat. 
Mus. Petr., Lep. i. pl. v. fig. 8 (1855). 
Pamphila selas, Mabille, Pet. Nouv. ii. p. 233 (1878). 
Pamphila herculea, Butler, Ann. & Mag. Nat. Hist. (5) vil. p. 140 (1881). 
Hesperia hyrcana, Christoph, Iris, vi. p. 87 (1893). 
Augiades sylvanus, Leech, Butt. China &e. p. 601 (1894). 
Hab. Kurope to Amurland, Korea (Leech); J apan (Pryer). 
A male and female in coll. Rothschild, from Koko-nor, Tibet, expand only 23 mm. 
and 25 mm. respectively. We have not had an opportunity of comparing the male 
genitalia of this small form with those of specimens of normal size. 


! AUGIADES SUBHYALINA. (Plate XXIV. fig. 58.) 


Hesperia subhyalina, Bremer & Grey, Schmett. N. China’s, p. 10, pl. iii. fig. 4 (1858). 
Pamphila subhyalina, Ménétriés, Cat. Mus. Petr., Lep. i. pl. v. fig. 7 (1855). 
Augiades subhyalina, Leech, Butt. China &e. p. 602, pl. xli. fig. 8, g (1893-94). 
Pamphila subhyatina, var. tibetana, Oberthiir, Htudes d’Ent. xi. p- 28, pl. vi. fig. 45 (1886). 
Hab. China (Pratt) ; Korea (Leech); ? Japan (fide Leech); Khasias (Hamilton, fide 
Doncaster). 
Leech states that he has specimens intermediate between A. tibetana and subhyalina. 
We have seen no specimens from Japan, but three males from the Khasia hills, 
procured from Mr, Doncaster, are undoubtedly subhyalina. 


! AUGIADES SYLVANoIDES. (Plate XXIV. fig. 59.) 
Augiades sylvanoides, Leech, Butt. China &c. p. 604, pl. xli. fig. 4, ¢ (1893-94). 
Hab. Ta-tsien-lo, Western China (Pratt). 


AUGIADES SIMILIS. 
Augiades similis, Leech, Butt. China &c. p. 605, pl. xli. fig. 6, ¢ (1893-94). 

Differs from sylvanoides, Leech, in the darker (brown, not yellowish-green) ground. 
colour and more distinct pale spots of the hind wing below. 


Hab. W. China (Leech). 


| AUGIADES SIVA. (Plate XIX. fig. 28, 3.) 
Pamphila siva, Moore, P. Z. 8. 1878, p. 692. 
Telicota siva, Watson, Hesp. Ind. p. 57. 


Hab. Khasias, 6000 ft. (Elwes); E. Pegu, Bernardmyo (Doherty). 
2x2 


248 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


AUGIADES BOUDDHA. 
Pamphila bouddha, Mabille, Ann. Soc. Ent. Fr. 1876, p. lvi. 
Augiades bouddha, Leech, Butt. China &e. p. 603, pl. xli. figs. 7 9, 14 g (1893-94). 

Closely resembles A. siva, Moore, but the spots on the hind wing are larger and 
more quadrate. 


Hab. Moupin (David). 


! AUGIADES BOUDDHA, var. CONSORS. 
Augiades bouddha, var. consors, Leech, t. c. p. 604, pl. xli. fig. 10, ¢. 

We believe that this insect, which comes from Moupin, is really the female of 
A, bouddha, and that the female specimen from Omei-shan, figured on Leech’s plate 
xli. fig. 7, belongs to another species. Mr, Leech himself observes that the Moupin 
female, the original of his plate xli. fig. 10, agrees better in the colour of the spots 
on the hind wing with the type male of bowddha figured by him than the Omei-shan 
female, the original of his plate xli. fig. 7; at the same time, the latter, which has 
the spots on the hind wing bright fulvous instead of white, agrees very well with 
Mabille’s description of bowddha, female. The specimens figured appear to be the 
only ones which Mr. Leech had seen, and we have seen no others. 


! AUGIADES BRAHMA. (Plate XXV. fig. 61 0.) 
Pamplila brahma, Moore, P. Z. 8. 1878, p. 691, pl. xlv. fig. 8. 
Telicota brahma, Watson, Hesp. Ind. p. 57. 

Hab. Masuri, N.W. Himalayas (Lang); Kumaon (Ramsay); Fort White, Chin 
hills, 7000 ft. (fide de Nicéville). 

The differences between A. siva and brahma, as given in the table, are well marked 
and constant; therefore the occurrence of the latter species in the Chin hills, where 
A, siva would be expected, is very curious. 


! AUGIADES CRATEIS. (Plate XXYV. fig. 61a.) 

Augiades crateis, Leech, Butt. China &. p. 603, pl. xli. figs. 9 9,11 ¢ (1893-94). 

Differs from A. brahma, Moore, in being larger, darker, and more strongly marked, 
especially on the hind wing below, and in the male genitalia; both have the whitish 
line on the black sex-mark in the male. 

Hab. Omei-shan, Chia-kou-ho (Leech). 


! AUGIADES ocHRACEA. (Plate XXIV. fig. 60.) 
Pamphila ochracea, Bremer, Bull. Acad. Pet. iii. p. 473 (1861) ; id. Lep. Ost-Sib. p. 33, pl. i. fig. 11 
(1864). 
Pamphila rickuchina, Butl. Cist. Ent. 11. p. 275 (1878). 
Augiades ochracea, Leech, Butt. China &e. p. 605 (1893-94). 


Hab. Japan (Pryer); Amurland (Graeser). 


A REVISION OF THE ORIENTAL HESPPRIIDA. 249 


! AUGIADES MAJUSCULA, n. sp. (Plate XIX. fig. 21, ¢; Plate XXV. fig. 61.) 


Resembles A. ochracea generally, and particularly in the black veining of the bind 
wing below, but is larger (exp. 41 mm.), and the fore wing has a much straighter 
termen, and therefore a more pointed appearance. The form of the cedeagus is quite 
distinct from that of A. ochracea, but is not to be distinguished from that of 
A, sylvanus. 

Described from two males from China (Pryer, in coll. Elwes). 

We should not have ventured to separate this species from ochracea if it had not 
been for the marked difference in the cedeagi, which are figured. Elwes obtained the 
types from a collection made many years ago by the late Mr. Pryer in China, mostly 
at Shanghai. He found specimens in Mr. Leech’s collection marked herculea, Butl., 
which appear to be the same species. 


TELICOTA. 


Telicota, Moore, Lep. Cey. i. p. 169 (1881) ; Watson, P. Z. S. 1893, p. 102. Type augias, Lin. 
Padraona, Moore, t.c. p. 170; Watson, t.c. p. 101. Type mesa, Moore,=dara, Koll. 


Under this generic name we bring together a number of species which agree well in 
general appearance and in all important points of structure. Antenne more than half 
as long as costa; club gradual, of moderate size ; apiculus as long as, or a little longer 
than the greatest width of the club. Palpi: second joint densely scaled, third joint 
short, conical, erecto-patent ; or second joint laxly scaled, third joint acicular, sub- 
erect, sometimes almost as long as one-third of the diameter of the eye. Wings 
without hyaline spots. Basal portion of vein 5 in the fore wing receding from vein 6, 
vein 5 therefore arising much nearer to vein 4 than to vein 6; vein 5 in the hind wing 
obsolete. Upperside dark brown or black, with a yellow (fulvous or stramineous) 
band on each wing ; that on the fore wing starts from the middle of the dorsum and 
runs obliquely outwards to join some pale spots in cells 6-8, it is sometimes broken 
into spots, and there is also sometimes a pale spot or spots in the cell, or an elongate . 
triangular pale spot next the costa from the base to beyond the middle; the band on 
the hind wing is submedian, and does not reach the costa or the dorsum, but is in 
nearly all the species reproduced, outlined in black, on the underside. ‘Tibial epiphysis 
present ; hind tibiz fringed with long hairs, and having two pairs of spurs. 

The species known to us we arrange as follows :— 


1 (16). Hind wing: termen distinctly excavate between veins 16 and 2 
or 16 and 3, the tornus therefore apparently shortly produced 
into a rounded lobe. 

2 (11). Male with an alar sex-mark. 

3 (6). Sex-mark on the fore wing: a streak of androconia reaching from 
the middle of vein 2 to the base of vein 4. (Texicors proper.) 


250 


4 (5). 


5 (4) 
6 (3) 
7 (8) 
8 (7) 
9 (10). 
10 (9). 
11 (2). 
12 (15). 
13 (14). 
14 (18). 
15 (12) 
16 (1) 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Fore wing above: lower outer angle of the yellow spots in cells 
2-4. narrowly produced along the contiguous vein nearly or quite 
to the termen; terminal dark band brown. Lower apical angle 
of clasp rounded off, or at most barely evident . 


. Fore wing above: lower outer angle of the yellow spots in one 


2-4 not, or but little, produced; terminal dark band black- 
brown. Lower apical angle of clasp produced into a triangular 
tooth, or, at least, strongly right-angled . 


. Sex-mark on the hind wing: a tuft of long hairs springing from an 


otherwise bare pale patch near the middle of cell 7 and the base 
of cell 6, and directed obliquely inwards and downwards. 


. Fore wing above: no yellow spot in cell 5. Fore wing below 


brown, the pale band distinct 


. Fore wing above with a suffused yellow poe near te middle of 


cell 5. 

Hind wing above bright fulvous as far as vein 7, thence to the 

costa warm dark brown : 

Hind wing above brown, with a beau uaunnedisa Helios aril 
which nearly reaches the termen and is cut short by vein 6 . 

Male without an alar sex-mark. 

Hind wing below with an irregular pale postmedian macular band 
outlined with black. 

Expanse of the male about 40 mm. Yellow band on fore wing 
narrower ; pale spot in cell 2 of fore wing above not reaching 
the base of that cell; fore wing below with the basal part of 
the cell blackish. Apex of the tegumen tridentate ‘ 

Expanse of the male about 44mm. Yellow band on fore wing 
above broader, basal two-thirds of cell 2 on the fore wing above 
entirely yellow. Fore wing below with the cell entirely yellow. 
Apex of tegumen bidentate 


. Hind wing below without distinct Ste Beenie brown 


passing into ochreous on the terminal half 


. Hind wing: termen practically straight (not excavate) plein 


veins 1 6 and 2 or 1 6 and 3; tornus not produced. 


. Fore wing above: cell 3 yellow at the extreme base and for about 


half its length. 


. Pale discal marking on hind wing below band-like. 
19 (20). 
20 (19). 
21 (18). 
22 (17). 
23 (28). 
24 (29). 


Size larger. Cell of fore wing entirely black . : 

Size smaller. Cell of fore wing with a yellow spot near the apex. 

Pale discal marking on bind wing below subovate . 

Fore wing above: cell 3 black at the extreme base. 

With a pale spot in each of cells 4 and 5 of the fore wing. 

Pale markings of the upperside deep yellow, sometimes inclining 
to falvous. 


augias, Linn. 


bambuse, Moore. 


prusias, Feld. 


insularis, n. sp. 


kuehni, Plotz. 


palmarum, Moore. 


augiades, Veld. 


simplex, n. sp. 


concinna, Nn. sp. 
gola, Moore. 
paragola, de Nicév. 


A REVISION OF THE ORIENTAL HESPERIID”. 251 


25 (26). Male with a sex-mark on the fore wing above. Fore wing above : 
pale band regular, its inner edge straight. Sex-mark consisting 
of a dull streak of androconia along the inner edge of the yellow 


band from the middle of vein 1a to the base of ven 4. . rectifasciata, n. sp. 
26 (25). No sex-mark on the fore wing above in the male; the pale band 

irregular, broken outwardly at cells 4-5 . . . ....~. dara, Koll. 
27 (24). Pale markings of the upperside cream-yellow . . . . . . dilutior, n. sp. 
28 (23). No pale spot in cells 4 and 5 of the forewing . . . . . . orphitus, Mab. 


!'TELICOTA AUGIAS. (Plate XXV. figs. 62, 62a.) 
Papilio augias, Linn. Syst. Nat. i. p. 794 (1767). 
Telicota augias, Distant, Rhop. Mal. p. 382, pl. xxxiv. fig. 23 (1886); Watson, Hesp. Ind. p- 55 (1891). 

The sex-mark in the male is subject to some variation; generally it is broad and 
continuous, sometimes it is broken up into three pieces, and in a specimen from Menado 
it is continuous but only about half as wide as usual and its black edging is very 
narrow and regular. 

Hab. Calcutta (Minchin); Akyab (Adamson); Poona (Swinhoe); Andamans, Perak, 
Pulo Laut, Sambawa (Doherty); Nias (Modigliani); Java (Fruhstorfer) ; Philippines 
(Semper); Hong-Kong ( Walker). 


!TELICOTA BAMBUSH. (Plate XXV. fig 63.) 


Pamphila bambuse, Moore, P. Z. S. 1878, p. 691, pl. xlv. figs. 11,12 ¢ 92. 
Pamphila pythias, Mabille, Pet. Nouv. ii. p. 234 (1878) ; id. Ann. Soc. Ent. Belg. xxi. p. (1878). 
Telicota bambuse, Distant, Rhop. Mal. p. 382, pl. xxxv. fig. 12 (1886) ; Watson, Hesp. Ind. p. 56(1891). 
Pamphila eurotas, Felder [Sitz. Ak. Wiss., Math.-nat. Cl. xl. p. 461, sep. p. 14 (1860)], 
is, Judging from a male specimen in the Felder Collection labelled in contemporary 
handwriting “ Hesp. ewrotas, Feld.,” ** Amboina,” mainly distinguished from 7. bambuse 
by having the postmedian yellow band on the fore wing above only half as wide, 
narrowed towards the apex and only reaching just beyond vein 5. 
Hab. Nilgiris (Hampson); Sikkim (Moller); Calcutta (Minchin); Assam, E. Pegu, 
Perak, Arjuno, Java, Bali (Doherty) ; Java (Fruhstorfer) ; Kina Balu ( Waterstradt) ; 
Mt. Mulu, N. Borneo (Hose). 


!I'sticora prusras. (Plate XIX. fig. 14, 3.) 
Pamphila prusias, Felder, Sitz. Ak. Wiss., Math,-nat. Cl. xlii. p. 44: (1861), 

In a female of this species sent by Semper the underside exactly agrees with that of 
the male, but is a little paler, thus differing from a specimen sent by Staudinger as the 
female of 7. prusias, in which the hind wing below bears a sharply defined postmedian 


macular pale band between veins 1 6 and 6. 
Hab. Celebes (coll. Stgr:) ; Palawan (Platen, Everett); Philippines (Semper). 


252 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


4 


! TELICOTA INSULARIS, n. sp. (Plate XIX. figs. 26 ¢, 272.) 

¢. Upperside: fore wing blackish brown ; a broad fulvous postmedian band from 
the dorsum nearly parallel with the termen as far as the end of the cell, recurved 
from thence to the costa, a streak along the basal half of the costa and two streaks 
in the cell, the upper one occupying its entire length, the lower one only half as 
long, also fulvous: hind wing fulvous, the costal region as far as vein 6 and a 
little beyond that vein on the inner two-thirds of the wing black ; a suffused anteciliary 
black line in cells 5-2, expanding in cell 10 into a suffused spot; cells 1 and la 
blackish ; a tuft of long hairs springing from an otherwise bare pale patch near the 
middle of cell 7 and the base of’ cell 6 and directed obliquely inwards and downwards. 
Underside: fore wing yellowish red-brown, passing into yellow on the dorsum; 
basal half as far upwards as the cell black, the black portion bearing an oblong bare 
pale spot from the dorsum as far as the base of vein 2; a fine anteciliary line and the 
veins blackish ; fringe deep yellow, the short scales blackish grey from the apex as 
far as vein 1a: hind wing yellowish red-brown, brownish yellow from vein 2 to the 
dorsum, a fine anteciliary black line from the apex to vein 2; fringe deep yellow. 
Palpi clothed with yellow scales intermixed with black. Clothing of body above 
fulvous. Antenne blackish, club pale beneath. 

@. Upperside: fore wing yellow-brown, pale markings as in the male but yellow rather 
than fulvous and the postmedian band narrower ; hind wing yellow-brown, with a broad 
suffused median yellow band passing from the apex of cells 4-5 towards the dorsum ; 
basal region clothed with long yellow hairs. Underside: fore wing as in the male, but 
the dark basal portion is blackish only and bears no pale patch, the dorsum as far 
as vein 1 @ dusky; hind wing as in the male, but less strongly tinged with red. 

A very distinct species, which comes nearest to 7. prusias. 

Expanse 42 mm. 

Hab. Pulo Laut (Doherty). Types in coll. Elwes. 


TELICOTA KUEHNI. 


Hesperia kuehni, Ploiz, Stett. ent. Zeit. xlvii. p. 101 (1886). 
Padraona kuehni, Semper, Schmett. Philipp. p. 302 (1892). 


Hab. Luzon (Semper); E. Celebes (in coll. Godman & Salvin). 


! TELICcOTA PALMARUM. (Plate XXYV. figs. 64, 64 a.) 


Pamphila palmarum, Moore, P. Z. 8. 1878, p. 690, pl. xlv. figs. 6, 7,¢ 2. 

Hesperia chrysozona, Plotz, Stett. ent. Zeit. xliv. p. 228 (1883). 

Pamphila augiades, var. bambuse, Staudinger, Iris, ii. pp. 144, 165 (1889). 

Padraona palmarum, Watson, Hesp. Ind. p. 60 (1891), 

Padraona chrysozona, Semper, Schmett. Philipp. p. 301, pl. xlix. figs. 13 g, 14 9, pl. B. 
fig. 11, larva (as Telicota bambuse) (1892). 


A REVISION OF THE ORIENTAL HESPERIIDA. 258 


Hab. Calcutta (de Nicéville); Upper Assam (Doherty); Khasias (Hamilton); Java 
(Piepers) ; Luzon, Mindoro, Negros, Bohol, Mindanao (Semper); Palawan, Bunguran, 
Natuna Is. (Lverett); Bukan, N. Borneo (im coll. Rothschild). * 


!'TeLtcoTA AvGIADES. (Plate XXV. figs. 65, 65 a.) 
Pamphila augiades, Felder, Sitzb. Ak. Wiss., Math.-nat. Cl. xl. p. 461 (1860). 
Hesperia augiades, Felder, Reise Nov., Lep. ii. pl. Ixxi. fig. 5 (1867). 
2 = Hesperia acalle, Hopf. Stett. ent. Zeit. 1874, p. 41. 
Hab. Amboyna (Felder); Batavia (ex coll. Snellen). We are unable to give any 
characters by which the females of 7. palmarum and T. augiades can be distinguished. 


TELICOTA SIMPLEX, n. sp. (Plate XIX. fig. 15, ¢ .) 


3. Upperside resembling that of 7. insularis, save that the yellow streaks next 
the costa of the fore wing are less strongly developed, and in this respect it resembles 
T. prusias ; no sexual tuft of hairs near the base of the costa of the hind wing. Fore 
wing below resembling that of 7. insularis, but wanting the pale denuded patch near 
the middle of cell 1 a, which is correlated with the tuft of hairs on the upperside of the 
hind wing. Hind wing below ochreous brown, passing into ochreous near the middle. 

Expanse 40 mm. 

Hab. N. Celebes (coll. Stgr.). Type in coll. Staudinger. 

Described from a single specimen from Minahassa sent by Staudinger as kuehni, 
Plétz. The latter species, however, according to a specimen received under that name 
from Semper, has in the male a sex-mark as in prusias and insularis, and the yellow 
colouring on the hind wing above is in the form of a distinct band, as stated in the 
original description. 


| TELICOTA CONCINNA, n. sp. (Plate XXI. fig. 20, g ; Plate XXV. figs. 67, a, b.) 


é. Differs from gola, Moore, in its larger size and the want of any yellow spot in 
the cell of the fore wing. Tegumen in the lateral aspect regularly decurved. In 
gola the tegumen in the lateral aspect is sinuate. 

Expanse 28 mm. 
Hab. Nilgivis (Davison, Hampson). 
Described from two specimens in coll. Elwes. 


| TeLicora GOLA. (Plate XXV. figs. 66, a, 0.) 


Padraona gola, Moore, P. Z. 8. 1877, pl. lviii. fig. 9, ¢ ; Leech, Butt. China, &. p. 598, pi. x 
fig. 12, ¢ (1892-94); Watson, Hesp. Ind. p. 59 (1891). 
Padraona goloides, Moore, Lep. Cey. i. p. 171, pl. Ixxi. figs. 3, 3. @ (1880-81) ; Watson, Hesp. Ind. 
p- 601 (1891). 
Telicota goloides, Distant, Rhop. Mal. p. 382, pl. xxxv. fig. 138, g (1886). 
VOL. X1v.—PaArT iv. No. 20.—October, 1897. 21 


254 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Pamphila akar, Mabille, Comptes Rendus Soc. Ent. Belg. iii. no. 31, pl. Ixxi. (1883). 
Pamphila goloides, var. akar, Staudinger, Iris, ii. p. 146 (1889). 
Padraona akar, Semper, Schmett. Philipp. p. 303, pl. xlix.fig. 16, ¢ (1892). 

Hab. Nilgiris (Hampson); Ceylon (Green); Khasias (Hamilton); Andamans, 
Perak, Java, Bali, Sambawa, Pulo Laut (Doherty); Luzon, Guimaras, Camotes, 
Mindanao (Semper); Palawan, Kina Balu, Borneo (Staudinger); Nias (Modigliani). 


TELICOTA PARAGOLA. 

Padraona paragola, de Nicéville, Jour, As. Soc. Beng. 1895, p. 546. 

Hab, N.E. Sumatra (de Nicéville & Martin); W. Borneo (ex coll. Staudinger). 

Size of 7. gola, but very distinct on the hind wing below, which is dark yellow-brow 
by reason of a sprinkling of yellow scales on a blackish ground and has a sharply 
defined subovate yellow discal patch. 

We know this species only from a specimen sent by Staudinger which agrees 
with de Nicéville’s description. 


! TELICOTA RECTIFASCIATA, n. sp. (Plate XXI. fig. 17,3 ; Plate XXYV. figs. 68, 68 a.) 


3. Differs from dara, Koll., in the following points :—Fore wing above : inner edge 
of the oblique pale band straight, bounded inwardly by a broad stripe of dull brown 
modified scales reaching from near the middle of vein 1 @ to the base of vein 4; in dara, 
Koll., the inner edge of the pale band is irregular because the bases of the spots in cells 
2 and 3 project inwardly out of line with the bases of the others forming the band. 

Expanse 28-31 mm. 

Described from three male specimens, one from Sikkim (J@/ler) and two from East 
Pegu (Doherty). We have not yet been able to identify the female of this species. 


!'TELICOTA DARA. (Plate XXV. figs. 69, 69 a.) 


Hesperia dara, Kollar, Hiigel’s Kaschmir, vol. iv. p. 455 (1848). 

Pamphila mesa, Moore, P.Z. 8. 1865, p. 509, pl. xxv. fig. 9. 

Pamphila flava, Murray, Ent. Mo. Mag. xii. p. 4 (1875). 

Pamphila nitida, Mabille, Pet. Nouv. ii. p. 114 (1877). 

Pamphila taxilus, Mabille, Ann. Soc. Ent. Belg. xxi. p. (1878). 

Pamphila trachala, Mabille, Pet. Nouv. ii. p. 237 (1878). 

Pamphila mesoides, Butler, Trans. Linn. Soc. Lond. ser. 2, Zool. vol. i. p. 554 (1879). 

Padraona mesoides, Moore, Lep. Cey.i. p. 171, pl. Ixxi. figs. 5, 5 a (1881). 

Telicota mesoides, Distant, Rhop. Mal. p. 883, pl. xxxiv. fig. 24 (1886). 

Padraona pseudomesa, Moore, Lep. Cey. i. p. 170 (1881). 

Padraona dara,Watson, Hesp. Ind, p. 57 (1891); Leech, Butt. China, &e. p. 596, pl. xl. figs. 18, 14, vars. 

Pamphila heterus, Mabille, Comptes Rendus Soe. Ent. Belg. iii. no. 31, p. Ixxii (1883); Staudinger, 
Tris, ii. p. 145 (1889). 

Padraona heterus, Semper, Schmett. Philipp. p. 303, pl. xlix. fig. 15, 2 (1892). 


A REVISION OF THE ORIENTAL HESPERIID. 255 


Pamphila zebra, Mabille, sec. spec. comm. 
? Pamphila sunias, Velder, Sitzb. Ak. Wiss., Math.-nat. Cl. xl. p. 462, sep. p. 15 (1860). 

‘This species varies much, in size, in the ground-colour of the hind wing below, 
and in the size of the spots which form the pale markings on the upperside. The pale 
spot in cell 6 on the hind wing below is absent or but feebly developed in certain males 
from Pulo Laut, Nias, Java, and Perak; in females from Pulo Laut and Nias, otherwise 
similar to the males last mentioned, this spot is present, but a female from Perak 
exactly resembles males from the same locality in wanting that spot. 

This species seems to have a wider range and to be more generally abundant than 
any other Eastern Hesperid. 

Elwes’s collection contains forty-six males and thirty-six females from almost every 
part of tropical Asia, including Japan. Edwards has examined the genitalia of no 
less than fifteen specimens from various localities and finds considerable variation 
in degree, but no differences which can be treated as specific. 

The five specimens now standing in the Felder collection as “ Pamphila sunias” 
clearly do not belong to the species described by Felder under that name, for he says 
of the hind wing below “atomis nigris fasciam cingentibus.” Of the specimens in 
question four are labelled “* Amboina” and one, a male, “ Aimboina, Doleschall.” The 
latter agrees exactly with a specimen from Celebes sent by Dr. Staudinger as Pamphila 
prusias, Felder ; the other three males are only distinguishable from the specimen last 
mentioned by the narrower pale band on the fore wing above (the pale spot in cell 2 
reaches to about the half length of vein 2); the remaining specimen, a female, appears 
to be properly associated with the narrow-banded males, as the band on the fore wing 
above is not so wide as one would expect to find in the female of 7. prusias having 
regard to the width of it in the male of that species. 

Hab. N.W. Himalaya (Young); Sikkim (Moller, Elwes) ; Khasias (Hlwes) ; Nagas, 
E. Pegu, Perak, Pulo Laut, Sambawa, Arjuno, Java, Bali (Doherty) ; Ceylon (Mackwood) ; 
Nias (Modigliani) ; Hong Kong (Pryer); Andamans (de Roepstorff’) ; Java (Piepers) ; 
Philippines (Semper); Kina Balu (Waterstradt) ; Japan (Pryer) ; C. China (Prait). 


! TELICOTA DILUTIOR, n. sp. (Plate XXI. fig. 21, 3.) 

3 2. Differs from dara, Koll., in the following points:—Pale markings of the 
upperside of a cream- or straw-yellow, the spots of which they are composed reduced 
in size ; ground-colour of the hind wing below dark yellowish green, owing to the sparse 
clothing of yellow scales on the black parts. 

Expanse 30-5] mm. 

This species seems constantly different from any form of dara. 

Described from several specimens, including three males and three females, from 
Pulo Laut (Doherty). ‘Types in coll. Elwes. 

Hab. Pulo Laut, Labuan, Kina Balu, Borneo (Waterstradt) ; N. Borneo (Kverett). 

2u 2 


256 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! 'TELICOTA ORPHITUS. 
Pamphila orphitus, Mabille, C. R. Ent. Belg. xxvii. p. Ixxiv (1883). 
Padraona pavor, de Nicéyille, Jour. As. Soc. Beng. 1894, p. 53, pl. iv. fig. 8, 5. 

In the male of this species there are three tracts of androconia on the fore wing 
above, namely, a streak in cell 1 @ next vein 1 @ and occupying about the middle third 
of the latter, a roundish patch near the middle of vein 2 divided by that vein into 
two nearly equal parts, and a short streak in cell 2 next the base of vein 3. 

Hab. N.E. Sumatra (Martin) ; Java ( fide Mabille). 


TELICOTA PHILZNUS. 

Pamphila philenus, Mabille, C. R. Ent. Belg. xxvii. p. Ixxii (1883). 

Somewhat smaller than 7. orphitus, Mab., which it resembles on the upperside, save 
that there are yellow spots in cells 4 and 5 of the fore wing and a yellow spot in cell 6 
of the hind wing, which is half as large as that in cells 4-5, of which it adjoins the 
outer half; the hind wing below is unmarked pale ochreous, and there is on vein 2 of 
the fore wing above near its middle an oblong patch of androconia bounded above and 
below by a slight folding of the wing-membrane. 


Hab. Malaysia (fide Mabiile). 


TELICOTA MARNAS. 
Pamphila marnas, Felder, Sitzb. Ak. Wiss., Math.-nat. Cl. xl. p. 462, sep. p. 15 (1860). 

The following particulars are taken from a male specimen in the Felder collection 
labelled in contemporary handwriting “ Hesp gnarnas, Feld.”, ‘‘ Amboina, Doleschall” :— 

3. Upperside: fore wing brown, with the basal half of the costa, a geminate cell- 
spot continued towards the base and having its upper part extended for about half its 
length beyond the lower part, a streak in cell la@ adjoining vein la as far as the 
sex-mark, a streak along the basal two-thirds of the dorsum, an oblique postmedian 
band from vein la@ to vein 4, a subquadrate spot near the apical third of cell 4, a 
smaller spot near the apical fourth of cell 5, and an oblong spot in each of cells 6, 7, 
and 8, deep yellow or fulvous; sex-mark an irregular line of pale brown androconia 
running from vein 1 @ near the middle to vein 4 near the base, and separated from the 
inner edge of the postmedian band by an irregular linear tract of brown; hind wing 
brown; a few scales and the pubescence near the base, a broad irregular postmedian 
band reaching from vein 14 to vein 6, and the indication of a spot near the apical 
third of cell 6 fulvous. Underside: fore wing as on the upperside save that the 
cell-spot is not continued towards the base, the streaks in cell 1 @ and next the dorsum 
are wanting, and the spots in cells 5 to 8 are more strongly developed; fringe fuscous, 
passing into yellow at the tornus ; hind wing brown, closely sprinkled with yellow scales, 
the pale markings as on the upperside save that there is a round yellow spot in the cell 
near the base of vein 6, and the pale spot in cell 6 is more strongly developed, reaching 
nearly to vein 7; fringe fuscous, yellow from the tornus to vein2. Wing-length 14 mm. 


A REVISION OF THE ORIENTAL HESPERIIDA, 257 


HALPE. 
Halpe, Moore, P. Z. 8. 1878, p. 689; Watson, op. cit. 1893, p. 108. 

Species of moderate or small size, brown or grey-brown above; fore wing with 
hyaline spots, but not one in cell 4; no hyaline spots in hind wing except in 
H. submacula, Leech. Antenne longer than half the costa; club slender; length 
of the apiculus about twice the width of the club. Third joint of palpi short, 
porrect, nearly concealed. Fore wing: basal portion of vein 5 receding from vein 6, 
vein 5 therefore arising much nearer to vein 4 than to vein 6; or vein 5 straight, 
arising very little nearer vein 4 than vein 6, its basal portion not receding from vein 6 ; 
vein 2 from near the half-length of the cell (except in separata, blanchardi, and 
subflava, where it arises from the apical third of the lower margin of the cell); 
termen generally shorter than the dorsum. Hind wing: vein 5 nearly obsolete; 
in the males vein 7 is abruptly bent upwards at the base, and at the same point 
there is a corresponding deflexion of the upper margin of the cell, so that the whole 
subcostal vein of the hind wing forms a figure resembling a tuning-fork; the latter 
structure is correlated to a discal sex-mark on the fore wing and is feund in a few other 
genera. ‘Tibial epiphysis present ; hind tibiz with two pairs of spurs. 

Sex-mark on the fore wing of the male a broad ridge of modified scales passing 
obliquely from below the basal third of vein 1 @ to the base of vein 5; in structure 
this sex-mark consists of two large corneous pits filled with yellow flocculent hairs 
and covered with broad modified scales; vein 1 @ is abruptly angulate before the 
middle. 

The sex-mark is wanting in H. astigmata, Swinh., and H. hieron, de Nicév. 

The following table includes all the species that we have been able to examine 
critically. The species of this genus can rarely be identified with certainty from 
figures :— 

Males. 
1 (58). Dise of hind wing above not occupied by a large suffused or well- 
defined yellow spot traversed by the brown veins. 
2 (57). Hind wing below without silvery stripes. 
3 (56). Clothing of breast and palpi not pure white. 
4 (28). Club of antenne banded with white on the upperside next to the 
apiculus. 
5 (22). Hind wing without hyaline spots. 
6 (9). Hind wing below with a sharply defined white median band 
running from the apex of cell 7 to the middle of cell 1 6, where 
it is abbreviated. 
(8). Fore wing: pale spot in cell 2 about one-third as long as that cell. zema, Hew 
8 (7). Forewing: pale spot in cell 2 about one-sixth as long as that cell. ormenes, Weym. 
9 (6). Hind wing below not as in paragraph 6. 
10 (13). Hind wing below with a pale spot near the base of cell 7. 


258 


1 (12). 


2 (11). 


13 (10). 
14 (15). 
15 (14). 


eS) 


16 (17). 
7 (16). 


pas 


8 (21). 
9 (20). 


20 (19). 


21 (18). 


22 (5). 


23 (4). 


24 (51). 
25 (50). 
26 (43). 


27 (30). 
28 (29). 


29 (28). 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Hind wing helow with a more or less prominent white macular 
median band; white spot at the base of cell 7 conspicuous . 
Hind wing below without a macular median band; white spot 
at the base of cell 7 inconspicuous ; a suffused stramineous spot 
near the apical third of cell 1 6, a minute pure white spot in each 
of cells 2, 3, 4, 5, and 6, those in cell 3, 4, and 5 very minute 

and sometimes wanting. No sex-mark on fore wing above 

No pale spot in the base of cell 7 on hind wing below. 

Fringe of hind wing pure white . 

Fringe of hind wing not pure white. 

Fore wing above: pale spot in the cell geminate : ; 

Fore wing above: pale spot in the cell single, next the upper bigs 
of the cell, at most with a punctiform pale spot below it. 

Fringe of the fore wing beneath whitish or chequered. 

Cell-spot in fore wing single, next the upper edge of the cell. 
Termen of the hind wing continuously rounded, the tornus 
consequently not pees to be produced; the fringe pale 
brownish grey Bre. Mer aR ae : 

Cell-spot in the fore wing saith a pale point next its deer eee 
Termen of the hind wing somewhat straightened between veins 
4 and 14, the tornus therefore appearing somewhat produced ; 
the fringe white with the short scales brown. ee 

Fringe of the fore wing beneath grey-brown, concolorous with 
the wg oP 28 

Hind wing with ites iyaline spots : 

Club of antennz not banded with white on the npperkide next 
the apiculus. 

Vein 2 of fore wing arising near the half-length of the cell. 

Hind wing below normally with markings. 

Pale pattern of hind wing below consisting of a straight oblique 
discal band and a more or less distinct subterminal band of 
suffused spots, the latter arising in a pale spot near the apical 
third of cell 1 and becoming gradually obsolescent towards 
the apex. This pattern is frequently almost obsolete, but it is 
always possible to trace its relationship to the typical pattern 
as displayed in a well-marked example of homolea. 

Hind wiug below : discal band sharply defined. 

Discal band on the hind wing below white, appearing on the 
upperside as an indistinct suffused pale patch : 
Diseal band on the hind wing below yellowish white, not 

appearing on the upperside 


. Hind wing below: discal band suffused or obacieeeene 

. Hind wing below with the pattern at least traceable. 

. Hind wing below with the basal half of cells 4-5 stramineous. 

. Size larger: a pale spot in cell 8 of fore wing above ; long hairs 


on the disc of hind wing above brown . 


cerata, Hew. 


astigmata, Swinh. 
insignis, Dist. 


hyrie, de Nicéy. 


kumara, de Nicéy. 


knyvetti, n. sp. 


fasciata, n. sp. 
submacula, Leech. 


moorei, Wats. 


ceylonica, Moore. 


nephele, Leech. 


34 (33). 
5 (32). 
36 (37). 
37 (36). 
38 (41). 
9 (40). 


40 (39). 


4] (38). 


42 (31). 


43 (26). 
44, (45). 
45 (44). 
46 (47). 
47 (46). 
48 (49). 

9 (48). 
50 (25). 
51 (24). 
52 (58). 
53 (52). 


54 (55). 


55 (54). 


A REVISION OF THE ORIENTAL HESPERIIDA. 259 


Size smaller: no pale spot in cell 8 of fore wing above; disc of 
hind wing above conspicuously clothed with long grey hairs. 
Hind wing below: basal half of cells 4-5 not stramineous, or if 

so the hmits of the pale colour not sharply defined. 

Expanse 36 mm.. 

Expanse about 30 mm. 

Fore wing: hyaline spot in cell 2 not crescent-shaped, generally 
oblong and straight-sided; ground-colour of the uuderside 
grey-brown. 

Fore wing below: fringe grey, not distinctly chequered ; side horns 
of tegumen falcate, pointed, reaching about halfway to the apex. 

Fore wing below : fringe whitish, distinctly chequered, at least 
next veins 2 and 3; side horns of tegumen nearly straight, 
reaching nearly to the apex, their apex rather meee 
truncate with the angles produced . es ; 

Hyaline spot in cell 2 of fore wing small and crescent- =ahiauedl. 
ground-colour of underside vinous brown a Vg 

Hind wing below brown with a sprinkling of grey re eenles, the 
latter so arranged as very faintly to indicate subterminal and 
discal bands. Apex of tegumen, in the dorsal aspect, consisting 
of two broadly faleate incurved connivent teeth om 4h 

No pale discal band on the hind wing below, the pale markings 
there consisting of more or less distinct spots. 

Hind wing below ochreous brown, with a conspicuous white 
punctiform spot in each of cells 2 and 3 . iy ish aa eels 

Hind wing below normally with a pale spot in each of cells 2,3, and 6. 

Hind wing below brown, more or less closely covered with 
sulphureous scales' and thus appearing yellow or greenish 
yellow with brown veins é 

Hind wing below with the veins concolorous. 

Hind wing below closely sprinkled with grey scales and thus 
appearing greenish brown ; a faint grey spot in cells 2, 3, and 6. 

Hind wing below closely sprinkled with ochreous scales and thus 
appearing greenish ochreous ; an indistinct pale spot in each of 
cells 2,3, and 6, and a suffused subterminal macular pale band. 

Hind wing below without markings ; 

Vein 2 of fore wing from the apical third of the hoe else of the cell 

Hind wing below dull brownish green without markings 

Hind wing below with markings. 

Hind wing below brown, closely covered with ochreous scales, 
which form an indistinct suffused subterminal macular band 
Hind wing below bright yellow, with a small black spot in each of 

cells 2, 3, and 7, and a blackish stripe im cell 16 


sulphurifera, H.-S. 


majuscula, i. sp. 


sikkima, Moore. 


homolea, Hew. 


brunnea, Moore. 


fusca, Elwes. 


sitala, de Nicév. 


varia, Murr. 
gupta, de Nicév. 
debilis, n. sp. 
aina, de Nicéy. 


separata, Moore. 


blanchardi, Mab. 


subflava, Leech. 


1 Sometimes there is little or no yellow scaling on the hind wing below, which is then greenish brown with 


three pale spots. 


260 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


56 (3). Clothing of breast and palpi pure white . . . . . . . . . albipectus, de Nicév. 
57 (2). Hind wing below with silvery markings, namely a spot in the 

base of cell 7, a stripe occupying the whole of cell 6, a stripe 

occupying the basal two-thirds of cell 3, and a small oblong 

spot near the apical third ofcell2. . ... . bivitta, Ob. 
58 (1). Dise of hind wing above with a large suffused or sab defined 

yellow patch traversed by the brown veins. 
59 (60). Basal portion of vein 5 in the fore wing receding from vein 6, 

vein 5 therefore arising much nearer to vein 4 than to vein 6. 

A sex-mark on the fore wing above . ... . . . . . decorata, Moore. 
60 (59). Vein 5 in the fore wing straight, arising very little nearer to vein 

4 than to vein 6, its basal portion not receding from vein 6. 

No sex-mark on the fore wing above. 
61 (62). Dise of hind wing above with one or more sharply defined yellow 

spots, of which that next the transverse vein is the largest and 


SURONSESta se nee : cares Wi aes masoni, Moore. 
62 (61). Disc of hind wing aioe allow ue colour cuties: te “tli 
brown colour of the termen . . .... . =. +... +. . honorei, de Nicév. 


! HALPE ZEMA. 
Hesperia zema, Hewitson, Ann. & Mag. Nat. Hist. (4) xix. p. 77 (1877). 
Halpe zema, Elwes, Trans. Ent. Soc. Lond. 1888, p. 455, pl. xi. fig. 7, ¢ ; Watson, Hesp. Ind. 

p. 74 (1891). 

Hesperia ormenes, Plotz, Stett. ent. Zeit. 1886, p. 92; Weymer, t. c. 1887, p. 16, pl. ii. fig. 6 

Of H. ormenes, Weymer, described from Nias, I have two doubtful specimens from 
Pulo Laut and one from Bali which differ from zema’only in the smaller size of 
the spots on fore wing. As, however, a Tavoy specimen is intermediate in this 
respect and a specimen in coll. Staudinger from Borneo is nearer to zema than to 
ormenes, we prefer to treat is as an inconstant variety of zema. 

Hab. Sikkim (Elwes, Moller); Nagas (Doherty); Atavan Valley, Tavoy (Zucker) ; 
Borneo (coll. Stgr.) ; Pulo Laut, Bali (Doherty). 


! HALPE CERATA. 
Hesperia cerata, Hewitson, Ent. Mo. Mag. 1876, p. 152. 
Halpe cerata, Elwes, Trans. Ent. Soc. Lond. 1888, p. 454, pl. xi. fig. 8, ¢ ; Watson, Hesp. Ind. p. 73. 


Hab. Sikkim (Moller); Bhamo, Burmah (in coll. Stgr.); ?Philippine Islands 
(Réssler, ex coll. Snellen, as sulphurifera, H.-S.). 


! HALPE ASTIGMATA. 
Parnara astigmata, Swinhoe, Ann. & Mag. Nat. Hist. (6) v. p. 8363 (1890). 

3. No sex-mark on the fore wing above. Upperside dark brown, with a purple 
shade on the fore wing; the latter with five hyaline spots, one, geminate, in the cell, 
and one each in cells 2, 3,6, and 7, the last-named punctiform. Underside dark 


A REVISION OF THE ORIENTAL HESPERIIDA, 261 


brown: fore wing with the hyaline spots as on the upperside and a very minute 
white spot in cell 8, a pale suffused macular subterminal band faintly indicated ; 
hind wing a little warmer in colour than the fore wing by reason of a sprinkling 
of yellowish-grey scales, an inconspicuous white spot at the base of cell 7, a suffused 
stramineous spot near the apical third of cell 1d, and a minute pure white spot 
in each of cells 2, 3, 4, 5, and 6, those in cells 3, 4, and 5 very minute and sometimes 
wanting, a pale suffused submarginal band faintly indicated. Fringes whitish 
grey, chequered with brown next the ends of the veins. Body and legs concolorous 
with the wings; palpi somewhat lighter, the second joint being clothed with ochreous 
and black scales intermixed. Antenne above dark brown, conspicuously banded 
with white next the base of the pale red apiculus, spotted with white in front; club 
beneath pale ochreous. 

The above description was written before we knew that the insect had been already 
described, and we therefore print it in the hope that it may prove useful to some 
who may not have ready access to the original description. 

Hab. Nilgiri hills (Hampson). 


HALPE INSIGNIS. 
Baoris ? insignis, Distant, Rhop. Mal. p. 391, pl. xxxv. fig. 22 (1886). 


Hab. Singapore (type) ; Tandjong Djatti, Sumatra (Martin). Type in coil. Rothschild 


! HALPE HYRIE. 
Halpe hyrie, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 388, pl. G. fig. 34, 3. 


Hab. Naga hills (Doherty). 


! FIALPE KUMARA. 
Halpe kumara, de Nicéville, Jour. As. Soc. Beng. 1885, p. 121, pl. ii. fig. 10, ¢ ; Watson, Hesp. 
Ind. p. 72. 


Hab. Sikkim (Moller). 


! HALPE KNYVETTI, n. sp. (Plate XXI. fig. 2, 3.) 

Besides the characters given in the table above this species has the hind wing 
below yellow-brown by reason of a close sprinkling of yellow scales on a brown 
ground: in one specimen there is a faint indication of a tendency to the pattern 
on the hind wing below found in H. homolea and its allies, in a suffused yellow 
spot near the apical third of cell 1d preceded by traces of a dark postmedian 
macular band; in the other specimen the hind wing below is somewhat paler along 
the course of vein 14, but is otherwise unmarked. H. knyvetti may, however, be 
distinguished from any member of the homolea group by the white band on the 
upperside of the club of its antenne next the apiculus. 

VOL. XIV.—PART Iv. No. 21.—October, 1897. 2M 


262 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Expanse 40 mm. 

Described from two males from Sikkim in Elwes collection. 

This species is undoubtedly close to kumara, of which I have but one specimen, but 
it seems sufficiently distinct. 


HAcre rasciata, n. sp. (Plate XXI. fig. 7, 2.) 

?. Upperside dark brown; fore wing with five white spots, one quadrate in 
cell 2, one subquadrate or roundish in cell 5, one (subquadrate or roundish) 
in each of cells 6 and 7, and one very small and indistinct next the upper edge of the 
cell. Underside grey-brown: fore wing somewhat darker on the disc, the white 
spots as on the upperside and a subterminal row of indistinct suffused whitish 
spots; hind wing with a subterminal row of indistinct suffused whitish spots. Fringe 
of the fore wing grey-brown concolorous with the wing, of the hind wing similar 
but indistinctly spotted with whitish. Antennz above black with a white band 
next the apiculus, underside spotted with white. Body above dark brown, clothing 
of the breast and legs grey ; abdomen beneath blackish, hind margin of the segments 
with a narrow indistinct pale band. 

Expanse 33 mm. 

Hab. Kina Balu, Borneo (Waterstradt). 

Described from one example ex coll. Staudinger. Of the species known to us 
this species comes nearest to kumara and knyvetti, but it is in our judgment distinct 
from either. The close yellow scaling of the hind wing and the extra-discal portion 
of the fore wing on the underside so conspicuous in those two species is wanting 
in the specimen before us, but this may possibly be due to abrasion. 


HALPE LUCASIL. 


Hesperilla lucasii, Mabille, Ann. Soc. Ent. Fr, 1876, p. cliii. 
Halpe lucasi, Leech, Butt. China, &c. p. 624 (1894). 


We have not seen this species, but, according to Mr. Leech, it is allied to 
H. kumara, from which it differs in having a double spot in the cell of the 
fore wing, a more elongate hind wing, and a central series of four black spots on the 
hind wing below. 


Hab, Moupin (David). 


HALPE C&NIS. 
Halpe cenis, Leech, Butt. China, &c. p. 625, pl. xlii. fig. 16, ¢ (1894). 


Of this species, which was described from one male specimen taken in Western 
China at Chia-kou-ho in August, we transcribe the original description :— 

“Blackish brown, clothed about the base of primaries and disc of secondaries 
with fulvous hairs. Sexual brand well defined ; there are six white spots on primaries 


A REVISION OF THE ORIENTAL HESPERIIDA. 263 


placed as in H. varia, Murray, but that at end of cell is double. Under surface 
of primaries fuliginous, much suffused with ochreous brown about costa and apex ; 
spots as above: secondaries ochreous brown, with very faint indications of white 
central spots. Fringes whitish above, greyish beneath with faint chequering. 
Antenne black above, ringed beneath with yellow; club yellow beneath, tipped with 
castaneous. 

“ Expanse 40 millim.” 

The figure of this species greatly resembles H. kumara, but differs in its larger 
size and in having the hind wing narrowed towards the tornus. 


Hab. W. China (coll. Leech). 


! HALPE SUBMACULA. 
Halpe submacula, Leech, Entomologist, xxiii. p. 48 (1890); id. Butt. China, &. p. 622, pl. xii. 
fig. 13, d (1894). 
Hab. Central China (Pratt). 
This is a very distinct species on account of the three large spots on the hind wing 
above. 


HALPE BeTuRIA. (Plate XXI. fig. 13.) 
Hesperia beturia, Hewitson, Descr. Hesp. p. 36 (1868). 
Halpe beturia, Watson, P. Z.S. 1893, p. 110. 

According to Watson this species “differs from moorei in having only four spots 
on the upperside in the male, two discal and two subapical, and the fringe is 
unicolorous throughout; in what appears to be the female there is an additional 
minute subapical spot, a minute indistinct spot at upper angle of cell, and the usual 
spot peculiar to the female on the submedian. On the underside of the hind wing 
all the spots are much diffused and irrorated with yellow. It is also a considerably 
larger insect, the male expanding 42 mm. and the female 43 mm.” 

Hab. Celebes ( fide Hewitson). 

We take this opportunity of figuring the type specimen in the British Museum. 


! HALPE MOOREL. 


Halpe moorei, Watson, P. Z.S. 1893, p. 109. 
Halpe beturia, auct. nec Hew. 
Halpe teliga, Swinhoe, Trans. Ent. Soc. Lend. 1893, p. 326. 


Hab. Calcutta (de Nicéville); Khasia (fide Swinhoe); Trichinopoly (Castets) ; 
Burmah (Watson, Manders) ; Andamans (de Roepstorff'). 


! HALPE CEYLONICA. 
Halpe ceylonica, Moore, P. Z. 8. 1878, p. 690, pl. xlv. fig. 9. 


Hab. Nilgivi hills (Hampson) ; Ceylon (fide Moore). 
2M 2 


264 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


! HALPE NEPHELE. 
Halpe nephele, Leech, Butt. China, &c. p. 622, pl. xlii. fig. 15, ¢ (1894). 
Hab. Western China (Zeech). 


HALPE MAJUSCULA, n. sp. (Plate XXI. fig. 6, 3.) 


¢. Upperside brown: fore wing with the usual Halpe sex-mark and four sordid 
yellowish hyaline spots, one each near the base of cells 2 and 3, and one (small 
and roundish or subquadrate) in each of cells 6 and 7. Underside grey-brown : 
fore wing with the pale spots as on the upperside and anarrow feeble, suffused, 
macular subterminal band; hind wing with a broad suffused macular median band 
and a subterminal band of about four suffused pale spots, of which the one near the 
apical third of cell 14 is more than twice as large as the others. Fringe of the 
fore wing grey-brown, of the hind wing dark grey with the short scales grey-brown. 
Body above brown, breast and legs with grey clothing; abdomen beneath blackish, 
hind margins of the segments with a band of grey scales. Antenne above dark brown, 
spotted beneath with yellowish grey; club yellowish grey beneath ; apiculus pale red 
on the underside. 

This species is very like H. nephele, Leech, but the markings on the hind wing 
below are paler and more suffused. 

Expanse 536 mm. 

Hab. Minahassa, 

Described from one example ex coll. Staudinger. 


! HALPE SULPHURIFERA. 

Cobalus sulphurifera, Herrich-Schiaffer, Prod. Syst. Lep. 11. p. 82 (1867). 
? Hesperilla luteisquama, Mabille, Bull. Soc. Ent. Fr. 1876, p. excix. 
Halpe sulphurifera, Semper, Schmett. Philipp. p. 305 (1892). 

A specimen of Z. luteisquama sent by M. Mabille has cell 1 @ on the fore wing below 
entirely brown; in Philippine specimens of H. sulphurifera this cell is crossed near 
the middle by an oblique white spot, which reaches from vein 1a@ to vein 2, and is 
constricted near the middle in the female, but is less developed and scarcely reaches 
vein 2 in the male. It may be a distinct species. 

Hab. Luzon, Mindoro, Guimaras, Bohol, Leyte, Sulu I. (Semper); Moupin (fide 
Matille). 


! HALPE sIkKIMA. (Plate XXYV. fig. 70.) 


Halpe sikkima, Moore, P. Z.S. 1882, p. 407; Watson, Hesp. Ind. p. 70 (1891). 
Pamphila homolea, var. palawea, Staudinger, Iris, ii. pp. 144, 165 (1889). 
Halpe palawea, Semper, Schmett. Philipp. p. 358 (1892). 
Judging from a specimen sent by Semper palawea is certainly not separable as a 
species from stkkima, of which the type is in Elwes’s collection. 


A REVISION OF THE ORIENTAL HESPERIIDA, 265 


A specimen from Kina Balu, Borneo, sent by Staudinger. only differs from a 
specimen from East Pegu in wanting the pale spot in cell 7 of the fore wing. 
Hab. Sikkim (Llwes); Nagas, E. Pegu (Doherty); Palawan (Platen). 


! Haire yomonea. (Plate XXV. fig. 71 ) 

Hesperia homolea, Hewitson, Descr. Hesp. p. 29 (1868); Watson, Hesp. Ind. p. 71, note (1891). 
Halpe sikkima, Elwes, Trans. Ent. Soc. Lond. 1888, p- 453 (in part.), pl. xi. figs. 3, 4, g. 
Halpe aucma, Swinhoe, Trans. Ent. Soc. Lond. 1893, p- 325, sec. spec. typ. 

Halpe perara, id. 1. ¢., sec. spec. typ. 

Halpe marta, id. 1.c., sec. spec. typ. 

Halpe wantona, id. |. c., sec. spec. typ. 

Hab. Sikkim (Elwes); Nagas, E. Pegu, Pulo Laut, Perak, Bali (Doherty). 

In Watson and de Nicéville’s opinion this species is doubtrully distinct from 
Hf. sikkima, and the occurrence of both in Sikkim, Nagas, and Pegu would tend to 
confirm this opinion, but the genitalia of numerous specimens examined by 
Mr. Edwards prove the species to be distinct. 

We are indebted to Col. Swinhoe for the opportunity of examining the male 
genitalia of his four species mentioned above. 

A specimen in coll. Rothschild from Palawan (Hverett), expanding only 25 mm., 
appears to belong here; this may be the “ Pamphila homolea, Hew. ?, var.?palawea” 
of Dr. Staudinger (Iris, ii. p. 144, 1889), which he subsequently (¢. c. p. 165) elevated 
to the rank of a species; but it is to be remarked that whilst Dr. Staudinger’s insect 
is stated by him to be separable from homolea by the want of any hyaline cell-spot on 
the fore wing, the specimen before us has two cell-spots on the fore wing, of which the 
upper one only is truly hyaline, the lower one being visible on the underside only. 


HALPE HIERON. 
Halpe hieron, de Nicéville, Jour. As. Soc. Beng. 1894, p. 54, pl. iv. fig. 1, ¢. 

s. Upperside grey-brown; pale spots proper to the fore wing obsolete or absent; 
no sex-mark. Underside pale brownish grey, closely and irregularly clothed with 
whitish-grey scales so as to indicate a pale macular subterminal band on the fore wing 
and a pale postmedian band on the hind wing; a suffused whitish-grey spot in each 
of celis 2 and 3 of the fore wing. 

Expanse 29-31 mm. 

Hab. Bekantschan, N.E. Sumatra (Martin). 


HALPE BRUNNEA. 
Halpe brunnea, Moore, Lep. Cey. i. p. 174, pl. Ixx. figs. 4, 4.@, 2 (1880-81). 

g. Upperside dark vinous brown: fore wing with four small hyaline white spots— 
one crescent-shaped in cell 2, one much smaller and roundish in cell 3, oue punctiform 
in cell 6, and one very minute in cell 7. Underside pale vinous brown, disc of the 


266 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


fore wing darker; on the fore wing an indistinct subterminal pale macular band 
from the costa as far as vein 3, and an indication of a small whitish spot near the 
apical fourth of the upper edge of the cell; hind wing with the discal and subterminal 
bands as in homolea, save that the former is directed rather to a point beyond the 
apex of the wing than to the actual apex as in homolea. Fringe of the fore wing dark 
grey indistinctly chequered, of the hind wing dark grey with the short scales brown. 

2. Similar to the male, but having the pale cell-spot on the fore wing visible on 
the upperside. 

Hab. Ceylon. 

The above particulars are taken from a pair from Ceylon, ex coll. Rothschild. 
Moore’s figures are not characteristic. 


! HaPe Fusca. (Plate XXV. fig. 72.) 
Halpe fusca, Elwes, P. Z.S. 1892, p. 653, p 1. xlin. fig. J,. 
Hab. Bernardmyo, Burmah (Doherty). 


! HALPE SITALA, 
Halpe sitala, de Nicéville, Jour. As. Soc. Beng. 1885, p. 121, pl. ii. fig. 5, ¢ ; Watson, Hesp. Ind. 
p- 75 (1891). 
Hab. Nilgivis (Hampson); Arnamalai hills (Davison). 


HALPE LATRIS. 
Halpe latris, Leech, Butt. China, &c. p. 623, pl. xlil. fig. 17, g. 
Chiefly differs from H. sita/a in having the dorsum of the hind wing below blackish 


as far as vein | 0. 


Hab. Ta-tsien-lo (Leech). 


! HLALPE VARIA. 


Pamphila varia, Murray, Ent. Mo. Mag. xi. p. 172 (1875). 
Halpe varia, Leech, Butt. China, &c. p. 621, pl. xlii. fig. 18, d (1894). 


Hab. Japan (Pryer, Leech). 


! HALPE GUPTA. 


Halpe gupta, de Nicéyille, Jour. As. Soc. Beng. 1886, p. 255, pl. xi. fig. 1, ¢ ; Watson, Hesp. Ind. 
p. 73 (1891). 


Hab. Sikkim (Moller). 


HALPE DEBILIs, n. sp. (Plate XX1. figs. 1 3,5 2.) 
3. Upperside brown: fore wing with six sordid white hyaline spots—two oblong, 
one each in cells 2 and 3, three small and oblong, one each in cells 6,7, and 8; the 


A REVISION OF THE ORIENTAL HESPERIIDZ. 267 


cell-spots narrowly confluent and appearing as one spot much constricted in the 
middle. Underside greenish ochreous, the disc of the fore wing blackish ; hind wing 
with an indistinct whitish spot in each of cells 2, 38, and 6, and a pale suffused 
subterminal macular band. Fringe of the fore wing pale grey, becoming whitish 
towards the tornus, of the hind wing whitish with the short scales grey. Antenne 
blackish spotted with yellow in front; club and part of the shaft yellow beneath; 
apiculus pale red. Body above concolorous with the wings. Clothing of the second 
joint of palpi, breast, legs, and abdomen beneath greyish yellow. 

@. Similar to the male, but having on the fore wing a triangular white spot in 
cell 1@ next vein 1@ just beyond the middle, and the cell-spots broadly confluent and 
therefore forming one solid spot scarcely constricted in the middle. 

Expanse 32-54 mm. 

Hab. Khasia hills. 

Described from one pair ex coll. Rothschild. 


! HALPE AINA. 
Halpe aina, de Niceville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 176, pl. B. fig. 8, ; Watson, 
Hesp. Ind. p. 72 (1891). 


Hab. Sikkim (Moller). 


! HALPE SEPARATA. 
Halpe separata, Moore, P.Z. 8. 1882, p. 407 ; Elwes, Trans. Ent. Soc. Lond. 1888, p. 454, pl. xi. 
figs. 5 ¢, 6 2; Watson, Hesp. Ind. p. 71. 
Hab. Sikkim (Hlwes); Nagas (Doherty). 
Type in coll. Elwes. 


! HALPE BLANCHARDI. 


Hesperilla blanchardi, Mabille, Aun. Soc. Ent. Fr. 1876, p. cliii. 
Halpe blanchardi, Leech, Butt. China, &c. p. 625, pl. xli. fig. 19, ¢ (1894). 


Hab. West China (Leech). 


! HALPE SUBFLAVA. 
Halpe subflava, Leech, Butt. China, &c. p. 625, pl. xlii. fig. 21, ¢ (1894). 

A very distinct and well-named species on account of the clear yellow colour of the 
hind wing below. 

Hab, Western China (Leech). 

HALPE ALBIPECTUS. 
Halpe albipectus, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 389, pl. G. figs. 35 ¢, 36 2. 


Hab. Maingyé, Shan hills (Manders). 
g type in coll. Elwes. 


268 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


HALPE BIVI?TA. 


Pamphila bivitta, Ob. Etud. d’Ent. xi. p. 28 (1886). 
Pamphila albivitta, id. t. c. p. 88, pl. vi. fig. 46. 
Halpe bivitta, Leech, Butt. China, &e. p. 623 (1894). 


Hab. West China (Pratt). 


HALPE DECORATA. 
Halpe decorata, Moore, Lep. Cey.i. p. 178, pl. Ixxi. fig. 2 (1881) ; Watson, Hesp. Ind. p. 76 (1891). 


Hab. Avisawella, Ceylon (in coll. Rothschild). 

This is a true Halpe, which bears much resemblance on the upperside to H. honorei, 
de Nicév. ‘The hind wing below is chrome-yellow, with the termen narrowly and 
suffusedly pale brown, and with several small irregular pale brown spots placed as 
follows :—two beyond the middle of cell 16 divided by the intraneural fold, two in 
ceil 2 at equal distances from the base and apex of that cell and from each other, 
one near the basal third of cell 38, one near the middle of cells 4-5, one across the 
basal fourth of cell 6, and two in cell 7 at equal distances from the base and apex 
of that cell and from each other. These particulars are taken from a male specimen 
ex coll. Rothschild. J 


! HALPE MASONI. 


Pamphila masoni, Moore, P. Z.S, 1878, p. 842, pl. lii. fig. 5, 3. 
Isoteinon masoni, Elwes & de Nicéville, Jour. As. Soc. Beng. 1886, p. 442, pl. xx. fig. 4, ¢ ; Watson, 
Hesp. Ind. p. 85 (1891). 


Hab. Burmah (Watson, Adamson). 
! HALPE HONOREL. 
Halpe honorei, de Nicéville, P. Z.S. 1887, p. 464, pl. xl. fig. 8,9 ; Watson, Hesp. Ind. p. 75 (1891). 
Hab. Nilgiris (Hampson); 'Trichinopoly (Castets). 
HALPE ORNATA. 


Hesperia ornata, Felder, Reise Nov., Lep. iii. p. 515, pl. Ixxii. fig. 6, g (1867). 
Parnara ornata, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 382, pl. xviii. 
figs. 7, 7 a, ¢ ; Watson, Hesp. Ind. p. 88 (1891). 


We have not seen this species. 
Hab. Buitenzorg, Java (Helder); Doarband, Cachar (Wood -Mason & de Nicéville). 


ONRYZA. 
Onryza, Watson, P. Z. 8. 1893, p. 112, pl. ii. fig. 5, 


A monotypic genus, of which, through the kindness of Col. Adamson, we have been 
able to examine the type. It is distinguished from the masoni section of Halpe chiefly 


A REVISION OF THE ORIENTAL HESPERIIDA, 269 


by the sex-mark of the male, which is in the form of “a patch of long recumbent hairs 
on the upperside of the hind wing, attached along vein 8 from close to the base of 
the wing.” 


ONRYZA MEIKTILA. 
Parnara? meiktila, de Nicéyille, Jour. Bomb. Nat. Hist. Soc. 1891, p. 386, pl. G. fig. 32, g. 
Hab. Meiktila, Upper Burmah, Upper Chindwin, Feb. (Adamson); Poungadaw, 
Burmah (Watson); Ataran Valley, Burmah (Bingham, fide de Nicéville). 


Iron. 
Iton, de Nicéyille, Jour. Bomb, Nat. Hist. Soc. 1895, p. 399. Type semamora, Moore. 


This is a genus established by Mr. de Nicéville for semamora, Moore, and watsonii, 
de Nicév. Both these species have been placed in the genus Parnara; and if the form 
of their antenne, palpi, and wings, and the venation be compared with that of some 
species of Parnara, such as P. austeni, their relationship to that genus will at once be 
evident. The chief point of distinction lies in the pattern of the hind wing below, of 
which the main diagnostic feature is the fact that the discoidal cell is wholly white, and 
cells 4—5 and 6 are white except at the apex. The male of semamora has a patch of 
long upturned hairs near the middle of the dorsum of the fore wing below; the same 
sex of watsonii has no alar sex-mark. 


! [TON SEMAMORA. 


Hesperia semamora, Moore, P. Z. 8. 1865, p. 791. 
Hesperia barea, Wewitson, Trans. Ent. Soc. Lond. ser. 3, vol. ii. p. 490 (1866), fide de Nicéville. 
Parnara semamora, Watson, Hesp. Ind. p. 46 (1891). 

Hab. Sikkim (Miller, Gammie) ; Tilin Yaw, Burmah (Watson); Khasia (Hamilton) ; 


Pegu, Pulo Laut (Doherty). 


! Iron WATSONII. 

Parnara watsonii, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1890, p. 223; id. op. cit. 1891, p. 388, 
pl. G. fig. 29, ¢ ; Watson, Hesp. Ind. p. 46 (1891). 

Hab. East Pegu (Doherty); Tilin Yaw (Watson); Fort Stedman, Shan hills 
(Manders). 

We have seen, from the collection of Col. Adamson, an aberrant male specimen of 
this species taken at Kaléwa, on the Chindwin River, in October; on the upperside 
of the hind wing the white patch is quite as extensive as in some specimens of 
semamora, but the hind wing below, although very much abraded, still retains the 
characteristic pattern of watsoni; the clasp-form can, without dissection, be made out 
sufficiently well to show that in this respect it agrees very well with waésonii, and is 
absolutely distinct from semamora. : 

VOL. XIv.—ParT Iv. No. 22.—October, 1897. 2N 


270 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


PaRNARA. 


Parnara, Moore, Lep. Cey. i. p. 166 (1881) ; Watson, P. Z.S.1893, p.105. Type guttatus, Brem. 
Chapra, Moore, t. c. p. 169. Type mathias, Fab. 

Baoris, Moore, t. c. p. 165; Watson, t. c. p. 106. Type oceia, Hew. 

Caltoris, Swinhoe, Trans. Ent. Soc. Lond. 1893, p. 323. 


Body robust, fore wing pointed, upperside brown with a few white or yellowish- 
white spots. Antenne as long or longer than half the costa; club moderate; apiculus 
distinct, as long or longer than the greatest width of the club. Palpi: second joint 
densely scaled; third joint obtuse, very short, and almost concealed. Fore wing with 
a series of bare (and therefore hyaline) white spots, of which the position and full 
complement is as follows :—one each, decreasing in size, near the bases of cells 2, 3, 
and 4; one each, of nearly equal size, in cells 6, 7, and 8; and two near the apex of 
the cell; sometimes there is a white spot in cell 1 a, but this does not properly belong 
to the hyaline series, as it is not usually bare on the underside, but is represented 
there by a suffused whitish spot; it does not appear that there is ever a pale spot in 
cell 5, except in P. beavani and the female of P. assamensis; the pale spot in cell 4 is 
normally always present, its absence occurs most frequently in P. oceia; vein 5 arises 
much nearer 4 than 6, and curves upward from the base, and vein 2 arises at about the 
half-length of the cell. Hind wing: sometimes with a series of white spots, but 
without any pattern on the underside, save a row of spots: vein 2 from the apical 
fourth of the cell, vein 5 obsolete. Legs as in Gegenes. 

Zelleri, Led., tulsi, de Nicéy., and cwrulescens, Mab., are true Parnaras in venation. 
P. pugnans, de Nicév., has vein 2 of the fore wing arising from near the basal third of 
the cell. 

The generic term Parnara, as here used, includes Parnara proper, Chapra and Baoris, 
Moore, and Caltoris, Swinhoe. Distant and Watson use Baoris in the same sense, but 
as the first three names are synchronous in publication, and both Paoris and Chapra 
were definitely limited by their author to species possessing a certain kind of alar 
sex-mark in the male, we prefer the term Parnara on the ground that it was not so 
limited, and that it has been in use for the majority of the species for a long time. 
The presence or absence of some of the pale spots on the wings is a very useful 
character for the separation of species in this difficult genus, particularly when it is 
correlated with differences in the male genitalia. In the species of the oceia group, 
however, the number of the pale spots has proved perfectly unreliable as a distinctive 
character, but this fact does not affect the utility of the character where other species 
are concerned. It has not been found practicable to form any linear arrangement of 
the species which shall satisfactorily indicate their natural affinities inter se, but the 


consecutive arrangement here adopted will probably be found as useful as any other 
yet proposed. 


A KEVISION OF THE ORIENTAL HESPERIIDA. 


271 


The following table, which deals with male insects only, sets forth the characters by 


means of which we separate the species known to us:— 


10. 


Males. 


. Hind wing above with a tuft of long hair on the basal half of 


vein 6 covering a suboval patch of androconia. (Baoris.) 

Fore wing below with an upturned tuft of long hair near the 
middle of the dorsum 

Fore wing above with a linear Smee S€X- ee a fe pads 
of vein 1a to the basal third of vein 2. (Chapra.) . 


Wings without a sex-mark. (Caltoris.) 


. Hind wing below plain brown 


Hind wing below greenish yellow-brown ; 
Hind wing purple-brown, plain brown in cells 1 a an 1 7 


. Sex-mark of even width throughout 


Sex-mark constricted near the middle . 


. Hind wing below with three or more pale spots . 


Hind wing below unspotted 


. Hind wing above with white spots (forming a row), one in each 


of cells 2, 3, and 4, or 2, 3, 4, and 5 
Hind wing above with no such row of white spots . 


. Hind wing above with an ill-defined whitish spot in cells 6 and 3; 


hind wing below with the pale spot in cell 6 the most strongly 
developed. Fore wing with the pale spot in cell 2 subquad- 
rate. Size larger . WS Re eye Ole eee) 
Hind wing below with the 
pale spot in cell 6 not more strongly developed than the 


Hind wing above without pale sate. 

others. Fore wing generally with the pale spot in cell 2 
narrow and oblique, about twice as high as wide. Size 
smaller 


. Hind wing below hen a ne faa fGen the wiaale of ts 


costa to the dorsum just before the tornus A200 
Hind wing below with a white band of the kind last Sseitionel’ 


. Hind wing below not marked with purple or whitish purple . 


Hind wing below marked with purple or whitish purple . 


. Hind wing below with a very suffused whitish-purple median 


band . Dearie ore WE ce ov foe hoes) oh tS 
Hind wing below with a postmedian row of Seas spots in cells 
1lbto6 . uaa : otk Aa tc nore 
Hind wing below without manne or at most with one pale 
spot BM Bet a ou 
Hind wing below with more ete one walle deed pale spot . 


2. 
plebeia, de Nicév. 


3. 

Zo 

oceia, Hew. 
leechii, n. sp. 
simillima, n. sp. 
4. 

nascens, Leech. 
5. 

brunnea, Snell. 


sinensis, Mab. 
6. 


subochracea, Moore. 


mathias, Fab. 

8. 

hasoroides, nu. sp. 
10. 

9. 

tulsi, de Nicév. 


cerulescens, Mab. 


11. 
2. 


272 


11. 


12. 


14. 


15. 


16. 


17. 


18. 


9: 


20. 


21. 


22. 


MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Fore wing without cell-spots, or if one is present it is placed 
next the upper edge of the cell Astrea 

Fore wing with one or two cell-spots, if there is only one it is 
placed next the lower edge of the cell is eters 

Fore wing with a white spot in cell 1 a, next vein 1 @ just beyond 
the middle . é 

Fore wing without any such spot eee We 

Pale spot in cell 4 of fore wing above small and soandish 

Pale spot in cell 4 of fore wing above oblong, as long as that in 
GAB oo e200 Se CF Oe Oe ; 

Hind wing below Been Not more than one pale spot in 
cell 1 a of the fore wing F : 

Hind wing below with a minute whieh pet near the middle of 
cell 2. Two pale spots in cell 1a of the fore wing, one of 
which is punctiform and placed next to vein 2 

Fringes not yellow 

Fringes bright yellow 

Without a pale spot in cell 8 of ‘ie Gs wing SME : 

With a pale spot in cell 8 of the fore wing, and also in call la 
next vein 1 a just beyond the middle aks 

Hind wing below deep warm brown, darker than the reel 
region of the fore wing below . 

Not as above 5 ; rs : ; 

Fore wing below with the is spot near he mididle of call la 
obsolete or absent; the disc not much darker than the 
remainder 

Fore wing below with a anfioeed ipleeear Siceal witeh tea the 
base to the middle, the pale spot near the middle of cell la 
distinct, suffused . 

Expanse 46 mm. 

Expanse 37—41 mm. 5 5 Ree arar tier | nk fea na ee 

Hind wing above wit a more or les distinct row of pale spots 
near the middle 5 ¢ rule : 

Hind wing above without a row of vale sols near tiie meiteile 5 

Pale spots on the hind wing in a straight row, gradually decreas- 
ing in size from cells 2-5" 

Not as above 


12. 
14. 


philippina, H.-S. 
kumara, Moore. 
15. 


pugnans, de Nicév. 


16. 


bromus, Leech. 
Wis 

aurociliata, n. sp. 
18. 


pagana, de Nicéy. 
moolata, Moore. 
19. 


20. 


cahira, Moore. 
robusta, nu. sp. 
austeni, Moore. 


22. 
26. 


guttatus, Brem. 
23. 


* This is very characteristic of the majority of specimens, but in the smaller ones in which the pale spots on 
the hind wing are only feebly developed (bada, Moore) this character is not so obvious ; 
may always be distinguished from colaca and bevani by the absence of any trace of the pale spot near the 


middle of cell 1a on the fore wing above. Im all the forms the structure of the tegumen is characteristic of 


the species, 


these latter, however, 


A REVISION OF THE ORIENTAL HESPERIID®, 273 


23. Hind wing below: pale spots in cells 4-5 contiguous, but 


distinct, frequently there is also a pale spot in cell6 . . . 24, 
Hind wing below: pale spots in cells 4—5 coalescing to form one 
quadrate spot, no pale spotincell6 . . . . : 25. 
24, Hind wing below greenish ochreous, pale spot in cell 2 aabeanel 
in size to that in cell 4. Side lobes of tegumen spiniform . pellucida, Murr. 


Hind wing below russet-green, pale spot in cell 2 distinctly 
smaller than that in cell 4. Side lobes of tegumen capitate.  contigua, Mab. 
25. Fore wing above: pale spots yellowish white, the lower edge of 
that in cell 2 twice or nearly twice as long as its inner edge. 
Apex of the cedeagus (seen from below) produced on the left 
side into a long sharp horn, the right side spose . . . eltola, Hew. 
Fore wing above: pale spots white, the lower edge of that in 
cell 2 one and a half times or less than one and a half times 
as long as its inner edge. Apex of the cedeagus produced 
into two subequal spinose lobes . . . . . . . . . . discreta,n. sp. 
26. Hind wing below with no pale spot inthe cell . . . . Wie, 
Hind wing below with a large white spot in the cell near Ae 
apex of its upper edge. Species otherwise resembling 


pellucida, Murray. . . - . » . Jansonis, Butl. 
27. Hind wing below with the wale spots not Rae Hordes eds eae eeon 
Hind wing below with the pale spots black-bordered . . . . zelleri, Led. 
Pe}. TIP ANGS Mol; EES GY MN 5 6 5 6 5 5 6 5 6 5 o AY 
Expanse 46 mm.ormore . ..... . Gp nono, OCs 
29. Fore wing with a pale spot in cell 4 and also in cell Ss a 4 Sh 


Fore wing with no pale spot in cells 4 or 8. Hind wing 

with two hyaline points—one near the middle of each of cells 
etl os 6 6 6 6 6 6 Oo Oo me eo 6 6 5 6 WMT We Gy 

30. Fore wing below: no pale spot in cell 5, termen straight or 

feebly concave between veins 1a and 3. Tegumen with a 


long thin sharp dorsal horn reaching from the middle to 


the apex. colaca, Moore. 


Fore wing below: fequently, with a minute Hae knot in “al 5 
(always present in the female), termen evenly curved through- 
out. Tegumen with nosuch horn. . . 6 bevani,; Moore. 

31. Hind wing above with a small pure white spots in cell 4; ce 
on the fore wing pure white Bro yaldees 

Hind wing above with no such spot; spots on fhe fae wing 
sordidawhite: as ccs at a) cle Wa aia) Seb em louis cue) MCONUIClOm Elan. 


assamensis, W.-M.&deNicéy. 


‘ In specimens from Perak some ot the spots on the hind wing are wanting, but the species may always be 


known by the form of the tegumen. 


274 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


!PaRNARA OCEIA. (Plate XXV. figs. 74, a, 6, ¢.) 
Hesperia oceia, Hewitson, Descr. Hesp. p. 31 (1868) ; Wood-Mason & de Nicéville, Jour. As. Soc. 


Beng. 1881, p. 258. 

Hesperia farri, Moore, P. Z. 8. 1878, p. 688; Watson, Hesp. Ind. p. 44 (1891). 

Baoris oceia, Moore, Lep. Cey. i. p. 165 (1881); de Nicéville, Jour. As. Soc. Beng. 1883, p. 85, 
pl. x. fig. 11,9 ; Watson, Hesp. Ind. p. 29 (1891). 

Baoris penicillata, Moore, Lep. Cey. i. p. 166 (1881) ; Watson, t. c. p. 30. 

Baoris scopulifera, Moore, P. Z. S. 1883, p. 582. 

Baoris unicolor, Moore, P. Z. S. 1883, p. 533; nec Distant, Rhop. Mal. p. 381, pl. xxxv. fig. 11 


(1886). 
Baoris sikkimu, Swinhoe, Ann. & Mag. Nat. Hist. 1890, v. p. 362. 
Hab. Sikkim (Méller), Nagas (Doherty), Andamans (de Roepstorff'); Philippines 
(Semper), Palawan (Platen); Khasia hills (Swinhoe). 


! PARNARA LEECHI, n. sp. (Plate XXI. fig. 3, ¢; Plate XXYV. figs. 73, a, 6, ¢.) 
Baoris oceia, Leech, Butt. China, &c. p. 616, pl. xlii. fig. 6, 3. 


Differs from the preceding and following species in the colour of the hind wing 
below and in the clasp-form as figured. 

Though there is but a single specimen in Elwes’s collection in somewhat worn con- 
dition, yet we have examined several in Mr. Leech’s collection which confirm our belief 
that the true oceia does not occur in China. 

Hab. Central and Western China (Pratt) ; Foochow, Ningpo (Leech). 


! PARNARA SIMILLIMA, n. sp. (Plate XXI. fig. 22,¢; Plate XXVI. figs. 75, a, 8, c.) 


Though the slightly paler colour of the hind wing below from cell 16 to the dorsum 
distinguishes this species from any examples of oceta or leechi that we have seen, yet 
the marked difference of the genitalia as figured are the best proof of its specific 
difference. Described from one male in Elwes’s collection. 


Hab. Pulo Laut (Doherty). 


! PARNARA PLEBEIA. 


Parnara plebeia, de Nicéville, P. Z. S. 1887, p. 466, pl. xl. fig. 2,g; Watson, Hesp. Ind. p. 40 
(1891). 
Pamphila mormo, Mabille, Aun. Soc. Ent. Belg. xxvii. p. 53 (1893). 


The male of this species is readily distinguished by the tuft of long hair near the 
middle of the dorsum of the fore wing below, a feature not noted in the original 
description; the female may be separated from the same sex of austeni by the colour 


A REVISION OF THE ORIENTAL HESPERIIDA, 275 


of the hind wing below, which in plebeia is simple umber-brown, and in austeni 
ochreous brown. 

A male from Kina Balu, Borneo, ex col]. Staudinger, has the termen less oblique 
than other males in coll. Elwes, including one from Pulo Laut, and the white spots on 
the fore wing are reduced in size. 

Hab. Sikkim (Moller, Elwes); Pulo Laut (Doherty); Kina Balu (Waterstradt) ; 
Java (fide Mabille). 


! PARNARA BRUNNEA. 
Pamphila brunnea, Snellen, Tijd. Ent. xix. pl. vii. fig. 4 (1876). 


Chapra cere, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1891, p. 388, pl. G. fig. 33, g. 
Pamphila sodalis, Mabille, Ann. Soc. Ent. Belg. xxxvii. p. 53 (1893). 


Hab. W. Java (ex coll. Snellen) ; Pulo Laut, Bali (Doherty). 


! PARNARA SINENSIS. 
Gegenes sinensis, Mabille, Bull. Soc. Zool. Fr. 1877, p. 232. 
Chapra prominens, Moore, P. Z. 8. 1882, p. 261; Watson, Hesp. Ind. p. 33 (1891). 
Pamphila similis, Leech, Ent. xxiii. p. 48 (1890). 
Parnara sinensis, Leech, Butt. China, &e. p. 608, pl. xl. fig. 11, ¢ (1894). 


Hab. China (Pratt); N.W. Himalaya (Young); Sikkim, Khasia (Ziwes); Nagas, 
Bali (Doherty). 


! PARNARA SUBOCHRACEA. (Plate XXI. fig. 26,4; Plate XXVI. fig. 83.) 


Chapra subochracea, Moore, P. Z. S. 1878, p. 691; Watson, Hesp. Ind. p. 32 (1891). 
Chapra mathias, var., de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 176, pl. B. fig. 7, g. 
This species is undoubtedly distinct from P. mathias, Fab., differing both in the 
particulars given in the table above and the form of the male genitalia. 
Hab. Barrackpur, near Calcutta (Minchin), Nilgiris (Hampson), Cachar (fide de 
Nicéville), Peermaad, Travancore (ex Doncaster). 


! PARNARA MATHIAS. (Plate XXVI. fig. 84.) 


Hesperia mathias, Fabricius, Ent. Syst., Suppl. p. 433 (1798). 

Hesperia thraz, Lederer, Verh. zool.-bot. Ges. Wien, 1855, p. 194, pl. 1. figs. 9, 10. 

Chopra mathias, Moore, Lep. Cey. 1. p. 169, pl. Ixx. figs. 1, 1a (1881); Watson, Hesp. Ind. p. 31 
(1891). 

Baoris mathias, Distant, Rhop. Mal. p. 380, pl. xxxv. fig. 10 (1886). 

Hesperia agna, Moore, P. Z. S. 1865, p. 791. 

Chapra agna, Moore, Lep. Cey. i. p. 169 (1881); Watson, Hesp. Ind. p. 32 (1891). 

Hesperia chaya, Méore, P. Z. 8. 1865, p. 791. 

Baoris chaya, Distant, Rhop. Mal, p. 380, pl. xxxiv. fig. 9 (1886). 


276 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Pamphila mencia, Moore, Ann. & Mag. Nat. Hist. (4) xx. p. 52 (1877). 
Parnara mencia, Leech, Butt. China, &c. p. 607, pl. xlu. fig. 14, 3 (1894). 


This seems to be about the most widely distributed and generally common species in 
Asia, extending from Syria to the Liukiu Islands and Sambawa. It occurs all over 
India from Kashmir to Malabar, and from Bombay to Pegu and the Malay Peninsula. 
In addition to the above-named localities, I have it from Borneo, Bali, Japan, and China ; 
and Semper states that it is generally distributed in the Philippines. Holland also 
records it, as Baoris chaya, from Hainan. 


! PARNARA NASCENS. 
Parnara nascens, Leech, Butt. China, &c. p. 614, pl. xIn. fig. 8, § (1894). 

Differs from sinensis, Mab., in the following particulars:—Upperside darker 
brown, the pale spots less strongly developed, one only in cell of fore wing, that 
in the upper half, sex-mark of the male interrupted in the middle; underside dark 


brownish green. 
Hab. Central China (Pratt). 


!PARNARA PHILIPPINA. (Plate XXI. figs. 4¢, 89; Plate XXVI. figs. 85, a, b, ¢.) 


Cobalus philippina, Herrich-Schiffer, Prod. Syst. Lep. iii. p. 81 (1869). 
Hesperia seriata, Moore, P. Z. 8. 1878, p. 688. 

? Baoris seriata, Moore, Lep. Cey. i. p. 166, pl. Ixix. figs. 4, 4a (1881). 

Parnara seriata, Watson, Hesp. Ind. p. 42 (1891). 

Parnara philippina, Semper, Schmett. Philipp. p. 298, pl. xlix. fig. 12, g (1892). 

The type of Baoris seriata, Moore, in the British Museum from Ceylon is a female ; 
the wide range of the species suggests that it occurs in other localities, but we know it 
only from those here mentioned. Though Semper has identified P. philippina, H.-S., 
with P. seriata, Moore, yet we think that this identification must remain somewhat 
uncertain, and we have not been able to compare the Eastern specimens here treated 
of with the type. 

Hab. Ceylon (in Brit. Mus.) N. Canara (Aitken) ; Philippines (Semper); Amboina 
(Stgr.); Palawan (fide Stgr.), Sangir, Talaut (Doherty). 


! PARNARA KuMARA. (Plate XXVII. figs. 90, a, 6, ¢.) 
Hesperia kumara, Moore, P. Z. 8. 1878, p. 687. 
Baoris kumara, Moore, Lep. Cey. i. p. 166, pl. lxix. figs. 2, 2a (1881). 
Parnara kumara, Watson, Hesp. Ind. p. 41 (1891). 
Hab. Kina Balu, Borneo, Java (Staudinger) ; Sikkim (Méller) ; Nilgiris (Hampson) ; 
Arjuno, Java (Doherty). 


A REVISION OF THE ORIENTAL HESPERIIDZ. 277 


! PARNARA BROMUS. 
Parnara bromus, Leech, Butt. China, &e. p. 614, pl. xlii. fig. 10, ¢ (1894). 


We recently received from Dr. Staudinger two insects from Folo (tin the Philippine 
Islands), as Pamphila philippina ; of these the reputed male is a female of P. bromus, 
Leech, agreeing exactly with the description and figure cited above, and the female is 
a specimen of that sex of P. toona, Moore. 

I have great doubt as to whether P. bromus can be separated from P. philippina. 
Leech’s plate, which seems to represent a female and not a male, as stated, is 
indistinguishable from a female of philippina, from the Philippines, which I received 
from Dr. Staudinger. Semper, however, states that philippina never has a transparent 
spot in the middle cell of the fore wing, whilst Leech’s plate and one of my specimens 
show two.—H. J. E. 

The fact remains that in philippina the cell-spots of the fore wing are wanting, or, 
at most, feebly developed, and the hind wing below is unspotted, and in bromus there 
are two well-developed cell-spots visible on the upperside of the fore wing, and the 
hind wing below bears a small pale spot near the middle of cell 2. We have not seen 


any intermediate specimens. 


PARNARA CANARAICA. 
Parnara canaraica, Moore, P. Z. S. 1883, p. 534; Watson, Hesp. Ind. p. 42 (1891). 


3. Fore wing with eight white spots including two in the cell; hind wing above 
without markings; hind wing below with two discal white spots. 

@. Fore wing with ten pale spots, eight placed as in the male and two in cell 1a— 
one punctiform next vein 2 near the middle, and one triangular next vein 1 @ near the 
middle; hind wing above with three discal semi-diaphanous spots; hind wing below 
with four discal white spots and a fifth at the end of the cell. 

Expanse 38-47 mm. 

Hab. Canara (Ward). 


We have not seen this species, which bears some resemblance to P. bromus, Leech ; 
but that species has the two pale spots in cell la@ of the fore wing above in both 
sexes similar to those found in the female of P. canaraica, and the latter species is 
well distinguished by the five white spots on the hind wing below. 


! PARNARA PAGANA. 
Parnara pagana, de Nicéville, P. Z. S. 1887, p. 465, pl. xl. fig. 7, ¢; Watson, Hesp. Ind. p. 40 (1891). 


All the specimens of this species in Elwes’s collection have spots in cells 6, 7, and 8. 

Hab. Sikkim (Méller); Pegu (Doherty); Tenasserim, Borneo (coll. Stgr.); Selesseh, 
Sumatra (in coll. Rothschild). 

VOL. X1V.—Part Iv. No, 28.—October, 1897. 20 


278 MESSRS. H. J. ELUWES AND JAMES EDWARDS: 


!PaRNARA MOOLATA. (Plate XXVI. figs. 86, a, 6, c.) 
Hesperia moolata, Moore, P. Z. 8S. 1878, p. 848. 
Pamphila dravida, Mabille, Pet. Nouy. i. p. 242 (1878). 
Baoris moolata, Distant, Rhop. Mal. p. 379, pl. xxxiv. fig. 10, ¢ (1886). 
Parnara moolata, Watson, Hesp. Ind. p. 42 (1891). 
Parnara cahira, Semper, Schmett. Philipp. p. 298, see. spec. comm. (nec Moore). 
Caltoris onchisa, 2, Swinhoe, Trans. Ent. Soc. Lond. 1893, p. 323, sec. spec. typ. 


Hab. Khasia (Hamilton); Perak, Pulo Laut (Doherty); West Java (coll. Snellen) ; 
Philippines (Semper); Palawan, Kina Balu, Borneo (Staudinger); Selesseh, Sumatra 
(in coll. Rothschild). 


! PARNARA AUROCILIATA, n. sp. (Plate XXI. fig. 23,g; Plate XXVI. figs. 87, a, 6, ¢.) 

A species at once distinguished by its bright yellow fringes. 

3. Upperside deep warm brown: fore wing above with seven yellowish-white hyaline 
spots—two in the cell, of which the upper is the smaller, one in each of cells 2, 3, 4, 6, 
and 7, the three last-named punctiform, and a small yellowish-white spot next vein 1@ 
near the middle. Underside clear rich brown, somewhat darker in the basal region of 
the fore wing ; pale spots as on the upperside, but the spot in cell 1 @ suffused. Fringes 
bright chrome-yellow, brown uext cell 7 of the fore wing and cells 4-7 of the hind 
wing, the short scales brown throughout. The punctiform hyaline spots on the fore 
wing have a tendency to disappear. 

Expanse 45-48 mm. 

Hab. Sikkim (Moller). Type in coll. Elwes. 

We separate this species from P. pagana and P. kumara with some doubt, but the 
genitalia are different from those of both those species. 


! PARNARA CAHIRA. (Plate XXI. fig. 25, 2 ; Plate XXVII. figs. 91, a, b,c.) 


Hesperia cahira, Moore, P. Z.S. 1877, p. 593, pl. lviii. fig. 8. 
Parnara cahira, Watson, Hesp. Ind. p. 48. 


Moore’s figure above cited does not agree with his description. It is said by 
Wood-Mason and de Nicéville (Jour. As. Soc. Beng. 1881, p. 258) that the female of 
oceia, Hew., was described by Moore as that of his cahira; but it does not appear 
certain that these authors examined the type of cahira. 

Hab. Kast Pegu (Doherty); Andamans (de Roepstorff’); Nias (Modigliani) ; China 
(fide Staudinger). 

Mr. de Nicéville (‘ List of the Butterflies of Sumatra,” Jour. As. Soc. Beng. 1898, 
p- 550) states that he has from Sumatra specimens which agree with Moore’s figure 
and description of cahira, also specimens which agree with Moore’s description and 


A REVISION OF THE ORIENTAL HESPERIIDZ. 279 


Elwes’s woodcut of qusteni, also specimens agreeing with Moore’s description of 
moolata, and specimens agreeing with Moore’s figure and description of Awmara; and 
he goes on to say that all his Sumatran specimens appear to him to represent one 
species, and that it may be subsequently found on an examination of the prehensores of 
the male that some of these species may be valid. We give below a table of characters 
drawn from the male genitalia sufficient for the separation of these species by any 
person of moderate experience in the examination of such objects. ‘The figures will 
assist in the elucidation of these characters. 


A. Lower distal angle of the lower lobe of clasp produced much beyond the upper 
angle in a long nearly straight lobe; the upper distal angle produced into a 
recurved lobe. . . . : . ~ 7 = = « “moolata, 
B. Lower distal angle of the ibe er sabe of clea foutided! not steanred: 
a. The two lobes forming the apex of the tegumen (seen from beneath) broad, flat, 
and subtruncate, their apices in the vertical aspect triangular. . . . - austeni. 
b. The two lobes forming the apex of the tegumen (seen from beneath) nance, 
pointed, and separate at the apex . . . Jeu . kumara, 
c. The two lobes forming the apex of the tegumen (geen fiom iheneath) Gather 
broad, curved, and connivent atthe apex . - . . =. =.=. . -. = - . cahira. 


We also repeat here the external characters by which these four species may be 
distinguished inter se :— ; 


Males. 
1 (2). Fore wing without cell-spots, or if one is present it is pe next the subcostal ; 
no white spot near the middle of vem la... . : . kumara. 


2 (1). Fore wing with one or two cell-spots; if there is hina one it is placeainne next the 
median. 
3 (4). Fore wing above with a white spot, sometimes reduced to a point, near the 
middle of vein la. Fore wing below with a suffused blackish discal patch 
from the base to the middle; the pale spot near the middle of cell 1 a distinct, 
suffused. Hind wing halos ochreous brown . . . 6 6) 6b Se one CHAE 
4 (3). No white spot near the middle of vein 1 @ on the fore wing ei dbote! 
5 (6). Underside: apical region of the fore wing paler than the disc of the hind wing, 
the latter deep warm (vinous) brown. Lower distal angle of the lower lobe 
oisthe\claspmuchyproduced ayy sans enn: : : : moolata. 
6 (5). Underside: apical region of the fore wing éouioalordus with the ae of the 
hind wing, the latter reddish yellow-brown, Lower distal angle of the lower 
loberotithe clasp rounded ey eiee arm ain ee ew enn) Pe CLUSL ETT. 


With regard to the females of these species an exact knowledge can only be obtained 
from specimens taken in cop., but no real difficulty arises except as to the females of 
202 


280 MESSRS. H. J. ELXWES AND JAMES EDWARDS: 


P. kumaraand P. austeni. Judging from the material in coll. Elwes, it appears probable 
that of those females in which the cell-spots are absent from the fore wing, those which 
have a white point in cell 1 @ near the middle of vein 2 should be referred to P. austeni, 
and those in which this spot is wanting to P. kumara. ‘The female of P. moolata 
differs from the male in having a white spot near the middle of vein 1 @ in the fore 
wing, and the female of P. cahira does not differ from its male in the number of white 
spots on the fore wing. 


! PARNARA ROBUSTA, n. sp. (Plate XXI. fig. 19, g ; Plate XXVII. figs. 88, a, 4, c.) 

3. Differs from P. cahira in its larger size, the want of a large suffused blackish 
patch on the disc and a pale suffused spot near the middle of cell 1 @ on the fore wing 
below, the browner and less ochreous colour of the hind wing below, and the clasp- 
form, as may be seen from the figures. 

Expanse 46 mm. 

Hab. Sangir (Doherty). 


Described from a single specimen in coll. Elwes. 


! PARNARA AUSTEN. (Plate XXVII. figs. 89, a, 8, c.) 
Baoris austeni, Moore, P.Z.S. 1883, p. 533. 
Parnara austeni, Elwes, Trans. Ent. Soc. Lond. 1888, p. 448, fig. 3; Leech, Butt. China &c. 
p- 613, pl. xl. fig. 7, d (1894); Watson, Hesp. Ind. p. 43 (1891). 
Caltoris onchisa, 3 , Swinhoe, Trans. Ent. Soc. Lond. 1893, p. 328, sec. spec. typ. 


The female of this species has no pale spots in the cell of the fore wing, a character 
which it shares with the females of P. plebeia, P. kumara, and P. rhypara. 

Hab. Sikkim (Moller); Khasia (Hamilton) ; Coonoor, Nilgiris (Davison, Hampson) ; 
Ceylon (Green); Andamans (de Roepstorff); Kweichow, China (coll. Leech). 

We are indebted to Col. Swinhoe for the loan of his types of Cultoris onchisa and 
permission to dissect the genitalia of the male. Having availed ourselves of this 
permission we are satisfied that the male is P. austeni; the female specimen, however, 
belongs, as before pointed out, to P. moolata. The pale subapical fascia on the 
fore wing below in the specimens appears to us to be the result of accident rather 
than normal wing-pattern. 


! PARNARA conJuNCTA. (Plate XXVII. figs. 92, a, 6, c.) 


Goniloba conjuncta, Herrich-Schiffer, Prod, Syst. Lep. iii. p. 75 (1869), fide Snellen. 

Gegenes javana, Mabille, Bull. Soc. Zool. Fr. 1877, p. 232. 

Hesperia narooa, Moore, P. Z. 8S. 1878, p. 687, pl. xlv. fig. 4. 

Parnara narooa, Moore, Lep. Cey. i. p. 167, pl. Ixix. figs. 3 a, b (1881) ; Watson, Hesp. Ind. p. 39 
(1891). 

Baoris narooa, Distant, Rhop. Mal. p. 380, pl. xxxiv. fig. 12, 2 (1886). 


A REVISION OF THE ORIENTAL HESPERIID. 28k 


Though we have no specimens from India or Ceylon, yet we think the plute in ‘ Lep. 
Ceylon’ sufficiently identifies the species, which has been hitherto known as narooa, Moore. 

Hab. Java (Piepers) ; Nias (Modigliani) ; Perak (Doherty) ; Kina Balu (Waterstradt) ; 
Sumatra (im coll. Stgr.); Philippines (Semper). 


! PARNARA ASSAMENSIS. 


Parnara assamensis, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1882, p. 65; id. op. cit. 
1886, p. 382, pl. xvii. figs. 5, 5a, d, pl. xvii. figs. 7, 7a, 2; Watson, Hesp. Ind. p. 37 (1891). 


Hab. Sikkim (Elwes) ; Khasia (Hamilton); Nagas (Doherty). 


! PaRNARA GUTTATUS. (Plate XXVI. fig. 76). 
Eudamus guttatus, Bremer & Grey, Schmett. N. China’s, p. 10, pl. iii. fig. 2 (1853). 
Goniloba guttatus, Ménétriés, Cat. Mus. Petr. Lep. i. pl. v. fig. 4 (1855). 
Hesperia fortunei, Felder, Verh. zool.-bot. Ges. Wien, xii. p. 489 (1862); id. Reise Noy., Lep. 
pl. Ixxii. fig. 11 (1866). 
Pamphila mangala, Moore, P. Z.S. 1865, p. 792. 
Hesperia bada, Moore, P. Z.S. 1878, p. 688. 
Parnara bada, Moore, Lep. Cey. i. p. 167, pl. Ixx. figs. 2, 2a (1881). 
Hesperia nondoa, Plétz, Stett. ent. Zeit. 1886, p. 97, fide Fruhstorfer. 
Parnara guttata, Elwes, Trans. Ent. Soc. Lond. 1888, p. 445 ; Watson, Hesp. Ind. p. 34 (1891). 


Hab. India, generally distributed, China, Japan (Leech, Pryer); Liu Kiu Islands 
(Pryer); Perak, Pulo Laut, Bali (Doherty); Java (Fruhstorfer); Kina Balu, Borneo 
( Waterstradt). 

We have two females from Java sent by Herr Fruhstorfer under the name Hesperia 
nondoa ; these correspond with males from Pulo Laut and Perak which are undoubtedly 
P. guttatus; the underside of all these specimens is darker and the arrangement of the 
pale spots on the hind wing is less evidently rectilinear than is usual in P. guttatus, 
and the cell-spots in the fore wing are entirely wanting. Further, we have received 
seven specimens from the island of Bali, collected by Doherty, which vary very much 
in the number and position of the spots on the hind wing, one of them being without 
these spots either above or below. We were inclined to treat these specimens as 
belonging to a distinct species, but an examination of the genitalia goes to show that 
they are only extreme forms of guttatus. 


| PARNARA CONTIGUA. (Plate X XVI. figs. 78, 78 a.) 


Pamphila contigua, Mabille, Bull. Soc. Zool. Fr. 1877, p. 232 (note). 
Hesperia toona, Moore, P. Z. 8. 1878, p. 689, 3. 
Parnara toona, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 383, 2; Watson, 


Hesp. Ind. p. 45. 
Pamphila scortea, Mabille, Ann. Soc. Ent. Belg. xxxvil. p. 53 (1893), 


This species has wrongly been placed as a synonym of P. pellucida, from which it differs 


282 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


in the darker ground-colour and smaller spots of the hind wing below, and also in the 
form of the male genitalia. We have examined Mabille’s types of contigua and scortea. 

Hab. Sikkim (Moller); Khasia (Elwes); Nagas, Pegu, Perak, Pulo Laut, Sambawa, 
Bali (Doherty) ; Hong Kong (Pryer). 


! PARNARA ELTOLA. (Plate XXVI. figs. 79, a, 6.) 
Hesperia eltola, Hewitson, Ex. Butt. iv., Hesp. pl. iv. fig. 40 (1869). 
Parnara eltola, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 384, pl. xviii. 
figs. 6, 6a, 3; Watson, Hesp. Ind. p. 45 (1891) ; Leech, Butt. China &c. p. 613 (1894). 
This species is distinguished from the next by the yellower colour of the pale spots 
on the upperside, which is most marked in the males, and by having the apex of the 
cedeagus, as seen from below, produced on the left side into a long sharp horn, and 
the right side spinose. 


Hab. Sikkim (Elwes); Nagas (Doherty). 

! PARNARA DISCRETA, n. sp. (Plate XXI. fig. 16, ¢ ; Plate XXVI. figs. 80, a, 6.) 

Closely allied to P. eltola, Hew., from which it differs in the following particulars :— 

é. Pale spots on fore wing above sordid white, that in cell 2 shorter in proportion, 
its inner and outer edges nearly parallel; ground-colour of the underside and the 
fringes, especially those of the hind wing, paler. Apex of the cedeagus produced into 
two subequal spinose horns, side lobes of the tegumen narrow and simple. 

?. Pale spot in cell 2 of fore wing comparatively shorter ; underside, especially of 
hind wing, paler. 

Hab. Khasias (Elwes); Sikkim (Moller); Tenasserim (bingham); E. Pegu, 
Bernardmyo, Burmah (Doherty). 

This species may be distinguished from P. e/tola (with which it is probably mixed 
in collections) at first sight by the purer white of the pale spots on the upperside ; 
and the difference in the male genitalia affords conclusive evidence of its distinctness. 


PARNARA PELLUCIDA. (Plate XXVI. figs. 77, 77 a.) 


Pamphila pellucida, Murray, Ent. Mo. Mag. xi. p. 172 (1875). 
Thymelicus pellucida, Staudinger, Rom. Mém. sur Lép. iii. p. 152, pl. viii. fig. 3 (1887). 
Parnara pellucida, Leech, Butt. China &c. p. 611 (1894). 
The ground-colour of the hind wing below is not sufficiently bright in Staudinger’s 
figure above cited. 


Hab. Japan (Pryer, Leech); Ussuri, Amurland (Dérries, fide Stgr.). 
PARNARA JANSONIS. 


Pamphaila jansonis, Butler, Cist. Ent. ii. p. 284 (1878). 
Parnara jansonis, Leech, Butt. China &c. p. 612, pl. xlii. fig. 12, ¢ (1894). 


Hab. Japan. (Pryer). 


A REVISION OF THE ORIENTAL HESPERIDA. 283 


PARNARA BIPUNCTATA, n. sp. (Plate XIX. fig. 5, ¢.) 

3g. Upperside: fore wing brown, heavily clothed with ochreous hair-scales towards 
the base ; six sordid whitish hyaline spots—two remote in the cell, one large, pentagonal, 
in cell 2, one, smaller and subquadrate, in cell 3, and one, minute and punctiform, in 
each of cells 6 and 7; an elongate triangular yellow spot next vein l1@ beyond the 
middle: hind wing ochreous by reason of the hair-scales; the costa brown as far as 
vein 6, and the termen narrowly and suffusedly brown; two pale points on the disc, one 
near the middle of each of cells 2 and 8. Underside ochreous green, disc of the fore 
wing brown: fore wing with the pale spot next vein 1 @ larger and more suffused than 
on the upperside, the other spots as on the upperside; hind wing with the two 
pale points as on the upperside. Fringes brown, becoming ochreous grey towards 
the tornus of each wing. Antenne blackish, spotted with yellowish on the underside ; 
club yellowish beneath. Second joint of palpi clothed with ochreous and black hairs 
intermixed. Body and legs ochreous brown. 

Expanse 363 mm. 

Hab. Batchian. 

Described from one specimen ex coll. Staudinger. 

At first sight this species bears some resemblance to P. contigua, Mab., from which, 


however, it is abundantly distinct. 


! PaRNARA CoLAcA. (Plate XXVI. fig. 81.) 
Hesperia colaca, Moore, P. Z. S. 1877, p. 594, pl. lviii. fig. 7. 
Parnara colaca, Blwes, Trans. Ent. Soc. Lond. 1888, p. 446, fig. 1; Watson, Hesp. Ind. p. 36 
(1891). 
Parnara cingala, Moore, Lep. Cey. i. p. 167, pl. Ixx. figs. 3a, 3 6 (1881). 
Hab. Sikkim (Mller); Ranchi (Irvine); Bangalore (Minchin); Trichinopoly 
(Castets); Andamans, Bali (Doherty); Nias (Modigliani) ; Ceylon ( Jide Moore). 


! PaRNARA BEVANI. (Plate XXVI. fig. 82.) 


Hesperia bevani, Moore, P. Z. 8. 1878, p. 688. 
Parnara beavani, Elwes, Trans. Ent. Soc. Lond. 1888, p. 447, fig. 2. 
Parnara bevani, Watson, Hesp. Ind. p. 36 (1891). 
Parnara thyone, Leech, Butt. China &c. p. 610, pl. xlii. fig. 4, 3 (1894). 

A specimen of P. thyone from Moupin, presented by Mr. Leech, seems inseparable 
from P. bevant. re 

Hab. N.W. Himalayas (Hocking, Young); Sikkim, Khasias, Bombay (Elwes) ; 
Nagas (Doherty); Kina Balu (Waterstradt, in coll. Stgr.); Moupin, E. ‘Tibet 
(Kricheldorf). 


284 MESSRS. H. J, ELWES AND JAMES EDWARDS: 


PaRNARA FLEXILIS. 
Isoteinon flewilis, Swinhoe, P. Z.S. 1885, p. 147, pl. ix. figs. 9,10; Watson, Hesp. Ind. p. 85 (1891). 

@. Size of P. colaca or P. bevant. Upperside brown: fore wing with the hyaline 
spots small, two in the cell, one next the middle of vein 1 a, one in each of cells 2, 3, 6, 7, 
and 8, that in cell 4 only indicated on the underside. Hind wing below brown, clothed 
with grey scales so as to give the appearance of a dark subterminal band. Fringes 
brownish grey, the short scales whitish grey. 

Hab. Poona. 

These particulars are taken from the type specimen in the British Museum. This 
is, we think, a very doubtful species. 


! PARNARA ZELLERI. 
Hesperia zelleri, Lederer, Verh. zool.-bot. Ges. Wien, 1855, p. 194. 

A little known, though very distinct species, easily distinguished by the black-bordered 
pale spots on hind wing below. 

Hab. Beyrout, Syria (Zach) ; Esmir, Busagha, near Tangier, Morocco (Walker). 


! PARNARA TULSI. 


Parnara tulsi, de Nicéville, Jour. As. Soc. Beng. 1883, p. 86, pl. x. fig. 1, ¢ ; Watson, Hesp. Ind. 
p. 44 (1891). 
Hesperia jolanda, Plotz, Stett. ent. Zeit. xlvii. p. 95 (1886) (fide Mabille). 
Hab. Sikkim (Moller); Sava (Fruhstorfer). 
Though this species does not seem to have been found in any part of Burmah or the 
Malay peninsula, I have a specimen from Java which is identical. 


! PARNARA CAERULESCENS. 


Pamphila cerulescens, Mabille, Ann. Soc. Ent. Fr. 1876, p. ly. 
Parnara cerulescens, Leech, Butt. China &e. p. 615, pl. xlii. fig. 9, 3. 


Hab. Wa-ssu-kow, Ta-tsien-lo (ex coll. Leech & Oberthiir). 


! PARNARA PUGNANS. 
Parnara pugnans, de Nicéyille, Jour. Bomb. Nat. Hist. Soc. 1891, p. 384, pl. G. fig. 30, 2. 


Hab. Pulo Laut (Doherty) ; Nias (Modigliani). 


PaRNARA HASOROIDES, n. sp. (Plate XXI. fig. 11, ¢.) 


é 2. Upperside brown. Underside brown: the hind wing with a broad white 
band from the middle of the costa to the dorsum just before the tornus, but interrupted 
in cell 16; the longer portion of this band is distinctly contracted at each end in the 
male, less distinctly so in the female. Antenne brown, the club yellowish beneath 


A REVISION OF THE ORIENTAL HESPERIIDZ. 285 


Fringes brown. Body above brown. Clothing of the second joint of palpi, breast, and 
legs dark grey. Abdomen beneath brown. 

Expanse 45-48 mm. 

Hab. Halmaheira; Batchian (ex coll. Stgr.). 

Described from one pair ex coll. Staudinger. 

The band on the hind wing below resembles that found in Hasora hurama, Butl., 
save that in the latter it is not interrupted in cell 1 0. 


PARNARA UMA. 

Parnara uma, de Nicéville, Jour. As. Soc. Beng. 1888, p. 592, pl. xiii. fig. 9, 9 ; Watson, Hesp. 
Tnd. p. 38 (1891). 

?. Upperside brown : fore wing with a geminate cell-spot and five others, one in each 
of cells 2, 3, 6, 7, and 8, white. Hind wing below vinous brown, with a broad silvery- 
white subcostal streak, adjoining at its apex the last of a postmedian series of silvery- 
white subquadrate spots which commences in cell 1 3. 

Hab. Karen hills, Burma. 

The single specimen from which this species was described appears to be still 
unique. 


PARNARA PHILOTAS. 
Baoris (Parnara) philotas, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 402, pl. Q. fig. 60, 

Upperside brown: fore wing and hind wing each with two hyaline white spots 
one in each of cells 2 and 3. Underside “ thickly covered with ‘old-gold’ coloured 
scales”; dorsum of the fore wing broadly dark purplish brown ; pale spots as on the 
upperside. 

Expanse 30 mm. 

Hab. Travancore (Ferguson) ; North Canara (Bel/). 

We doubt whether this insect belongs to the genus Parnara. The figure shows the 
colour of the underside to be dull greenish ochreous. 


PARNARA UNICOLOR. 
Baoris unicolor, Distant, Rhop. Mal. p. 381, pl. xxxv. fig. 11, g (1886). 

Hab. Malacca (Eichhorn) 

Through the kindness of Dr. Staudinger we have had the opportunity of making 
a critical examination of the unique type of this species. The specimen is a male and 
a true Parnara in venation and in the number and position of the tibial spurs. Ai 
the base of the fore wing above, between the dorsum and the submedian, there is a long 
raised space about equal in length to one-fifth of the dorsum; this is probably a sex- 
mark and confined to the male. In the fore wing the first median segment is about 
two and a half times as long as the second, which is a little more than twice as long as 

VOL. XIV.—PakT Iv. No. 24.—October, 1897. uP 


286 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


the third; the transverse vein is oblique, and vein 5 arises much nearer 4 than 6 and 
curves upwards from its base. The palpi and the apices of the antenne are now 
wanting, but there is one hind leg still left which has two pairs of spurs in the usual 
position. The specimen is in a worn condition. 

Mr. de Nicéville named this species, which he had not seen, as the type of his genus 
Idmon (vide p. 198, ante), but his generic description does not apply to it. 


PARNARA DISTICTUS. 

Baoris distictus, Holland, Trans. Amer. Ent. Soc. xiv. p. 123, pl. il. fig. 4 (1887). 

Hab. Hainan (Henry, fide Holland). 

“ Male. Upper surface uniformly olive-brown, cilia pale cinereous. Primaries with 
two small, triangular, semi-diaphanous white spots beyond the middle of the wing and 
between the submedian nervules. Underside lighter than the upper, with a hoary 
lustre, especially on the posteriors. A few scarcely visible light scaleless spots may be 
detected adjacent to the two white spots of the primaries, which reappear on the under- 
side. A curved submarginal row of obscure brown spots is found upon the secondaries. 
Underside of palpi and head white. Expanse of wings 13 inches. Type in coll. Holland.” 

Not having seen or been able to identify this species, we transcribe the original 
description. Its distinctive characters appear to be a subterminal series of obscure 
brown spots on the hind wing below, and the “ underside of palpi and head white.” 


GEGENES. 


Gegenes, Hiibn. Verz. p. 107 (1816) ; Watson, P. Z.S. 1893, p. 104. Type pyymeus, Hiibn. 
Philoodus, Rambur, Faun. Ent. Andal. ii. p. 808 (1840). Type nostrodamus, Fab. 

A genus resuscitated by Watson for the reception of the Papilio nostrodamus of 
Fabricius and two or three other species; it would be more convenient to treat it asa 
section of Parnara, in which it would be well distinguished by its short antenne. 

Antenne about one-third as long as the costa ; club elongate ovate ; apiculus scarcely 
evident. Fore wing with vein 2 from the half-length of the cell in both sexes, vein 5 
much nearer 4 than 6. Hind wing feebly excavated between veins 14 and 2, vein 5 
obsolete, vein 2 from apical fourth of cell. Tibial epiphysis present. Hind tibie 
with two pairs of spurs. 


GEGENES NOSTRODAMUS. 
Papilio nostrodamus, Fabricius, Ent. Syst. 1. p. 323 (1793). 
Papilio pygmeus, Hiibner, Eur. Schmett. 1. figs. 458-460 (1798-1803). 
Hesperia karsana, Moore, P. Z. 8. 1874, p. 576, pl. Ixvu. fig. 6. 
Chapra nostrodamus, Watson, Hesp. Ind. p. 33 (1891). 
Hab. Gibraltar (Walker); Biskra, Algeria (Elwes); Beyrout (Zach); Quetta 
(Swinhoe) ; Samarkand (Haderhauer); N.W. Himalaya (Young). 


A REVISION OF THE ORIENTAL HESPERIIDA. 28 


“I 


ERYNNIS. 
Erynnis, Schrank, Fauna Boica, ii. p. 157 (1801) ; Watson, P. Z. S. 1893, p.99. Type comma, Linn. 
Ocytes, Scudder, Syst. Rev. p. 55 (1872). Type metea, Scudd. 

The species of this genus are best known by their resemblance to the Papilio comma 
of Linneus. Antenne about half as long as the costa; club abrupt, subovoid ; apiculus 
minute. Third joint of palpi almost concealed, second joint densely scaled. Erynnis 
comma, Linn., varies much in the details of the wing-pattern, but the essential pattern 
on the hind wing below remains the same, even in those specimens which at first sight 
seem to be the most distinct. Several American forms of this species have been investi- 
gated in great detail by Scudder (Mem. Bost. Soc. Nat. Hist. ii. pp. 841-353, pls. x., xi.), 
but the differences in the male genitalia which he figures appear to us, for the most 
part, mere differences of degree. 


! ERYNNIS COMMA. 
Papilio comma, Linneus, Faun. Suec. p. 285 (1761); Esper, Schmett. i. pl. xxiii. figs. 1 a, b 
(1777) ; Hiibn. Eur. Schmett. i. figs. 479-481 (1798-1803). 
Hab. Kurope to Amurland. 
The following varieties of LE. comma may be treated as more or less constant local 
races, but we have not seen sufficient examples to enable us to form an opinion as to 
how far their distinctions are really constant. 


! ERYNNIS COMMA, var. CATENA. 
Hesperia comma, var. catena, Staudinger, Stett. ent. Zeit. 1861, p. 357. 
Hab. Lapland (Staudinger). 


ERYNNIS COMMA, var. DIMILA. 
Pamphila dimila, Moore, P. Z. S. 1874, p. 576; de Nicéville, Jour. Bomb. Nat. Hist. Soe. 1892, 
p- 355, pl. J. fig. 9, 9 ; Watson, Hesp. Ind. p. 160. 
Erynnis comma, var. dimila, Leech, Butt. China, &c. p. 595, pl. xli. fig. 12, ¢ (1892-94). 
Erynnis dimila, de Nicéville, Jour. As. Soc. Beng. 1894, p. 52, pl. i. fig. 7, 


Hab. N.W. Himalayas (Hellard) ; Ta-tsien-lo ( fide Leech). 


! ERYNNIS COMMA, Var. FLORINDA. 


Pamphila florinda, Butler, Cist. Ent. i. p. 285 (1878). 
Erynnis comma, var. florinda, Leech, Butt. China, &c. p. 594, pl. xli. fig. 17, ¢ (1892-94). 

Hab. Japan (Pryer). 

I have no specimens of comma from Amurland, and therefore cannot say whether the 
form described by Staudinger [Rom. Mém. sur Lép. vi. p. 211 (1892)] as var. repugnans is 
a well-marked form or not; but I think that when he speaks of florinda as more probably 
a var. of sylvanus and not of comma he cannot have seen the true florinda, which is clearly 
more nearly allied to comma than to sy/vanus, which also occurs in Japan.—H. J. E, 
Dp 2, 


a ~ 


288 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


ADOPHA. 
Adopea, Billberg, Enum. Ins. p. 81 (1820); Watson, P. Z. S. 1893, p. 98. Type thaumas, Hufn. 
Pelion, Kirby, List Brit. Rhop. (1858). Type thaumas, Hufn. 

A small group of species associated together on account of their resemblance, greater 
or less in degree, to the Papilio thaumas of Hufnagel (linea, Wien. Verz.). Antenne 
short, generally less than half as long as the costa; club elongate, gradual, not apiculate. 
Third joint of palpi acicular, suberect, nearly as long as half the diameter of the eye. 

The species are distinguished as follows :— 


Males. 
). Veins of hind wing below concolorous. 
(9). With a sex-mark on fore wing above. 
(8). Sex-mark continuous, leaving the extreme base of cell 2 free. 
(5). Fore wing below with the black patch in the base of cell 1a sharply 
defined and confined to the basal third of the wmg . . . . . thaumas, Hufn. 

5 (4). Fore wing below with the black patch in the base of cell 1 a apeeuaiee 

along the dorsum or almost obsolete. Hind wing below with no 

distinct stripe of the same colour as the disc of fore wing in cell 1 0. 
6 (7). Costa of the fore wing one-third longer than the dorsum. Hind wing 

below and apical area of fore wing below sordid greenish yellow, 

dise of fore wing below rusty yellow . . . Soo oe 6 pre leak 
7 (6). Costa of the fore wing one-fourth longer than the fe, Hind 

wing below and fore wing below, except a broad space along the 


dorsum, rust-yellow or greenish yellow . . . .. . . . « act@on, Rott. 
8 (3). Sex-mark interrupted, one end of the long piece filling the base of cell 2. lineola, Ochs. 
9 (2). No sex-mark on fore wingabove ....... +... . « stigma, Stgr. 
10 (1). Veins of hind wing below black. 
11 (16). Sex-mark wanting. 
12 (15). Fore wing above: the dark outline of veins 6-10 more or less spread- 


ing and confluent near their bases to form a dark patch beyond the 
end of the cell, basal third of the wing more or less dark. 
13 (14). Upperside without purple reflection; veins on the underside less 


broadly black . . . . eee te ol secs Selec) Une ea emmney warica= brent 
14 (13). Upperside with a purple iecian veins on the underside more 

broadly black . . . . tenebrosa, Leech. 
5 (12). Fore wing above: dark outline os veins 6- 10 a iranenoat, abel 

third of the wing concolorous. . . .... «=... =~. «. mervulata, Mab. 
16 (11). Sex-mark present. Fore wing above: basal third concolorous, dark 

marginal baud narrow. 9. . . . + « © 6 6 a 8 8 «leona, Butle 


! ADOPAA THAUMAS. 
Papilio thaw, as, Hufnagel, Berl. Mag. ii. p. 62 (1776) ; Esper, Schmett. i. 1, pl. xxxvi. figs. 2, 3 
(17782), 3. 2, pl. xeviii. figs. 5-10 (1790?). 
Papilio linea, Wien. Verz. p. 160 (1776) ; Hiibner, Eur. Schmett. i. figs. 485-487 (1798-1803). 
9. Papilio venula, Hubner, Eur. Schmett. i. figs. 666-669 (1803-18). 


Hab. Europe to Asia Minor, 


A REVISION OF THE ORIENTAL HESPERIIDA. 289 


! ADOPAHA HYRAX. 
Hesperia hyrax, Lederer, Wien. ent. Mon. y. p. 149, pl. i. fig. 6 (1861). 
Hab. Amasia, Asia Minor, Syria. 


! ADOPAA ACTON. 


Papilio acteon, Rottemburg, Naturf. vi. p. 30 (1775); Esper, Schmett. i. 1, pl. xxxvi. fig. 4 
(1778 ?) ; Hiibner, Eur. Schmett. i. figs. 488-490 (1798-1803). 


Hab. S. and C. Europe; Asia Minor; N. Africa; Canaries (Zeech); Samarkand 
(Haberhauer). 


ADOPA HAMZA. 

Hesperia hamza, Oberthiir, Etudes d’Ent. i. p. 28, pl. iii. figs. 2 a-c (1876). 

Very near to A. actwon, from which it appears to be best distinguished by the more 
fulvous colour of the upperside and the want of the suffused pale curved macular band 
which is seen just beyond the end of the cell on the fore wing of A. acteon. 

Hab. Oran (Oberthiir). 

We have not any specimens which we can identify from Oberthiir’s plate. A male 
from Lambessa in the Province of Constantine appears to us to be inseparable from 
A. lineola. 


! ADOP#A LINEOLA. 


Papilio lineola, Ochsenheimer, Schmett. Ent. i. 2, p. 230 (1808). 
Papilio virgula, Hiibner, Eur. Schmett. i. figs. 660-663 (1803-18). 


Hab. Europe to Amurland; Algeria (Oberthiir). 


! ADOPAA STIGMA. 
Thymelicus (Hesperia) stiyma, Staudinger, Stett. ent. Zeit. xlvii. p. 252 (1886). 

This species belongs to the sylvatica group, but the veins on the underside are not 
black ; the fore wing above has the dark terminal band sharply defined, the basal third 


concolorous, and the transverse vein rather conspicuously dark. 
Hab. Turkestan. 


! ADOPASA SYLVATICA. 


Pamphila sylvatica, Bremer, Bull. Acad, Petr. iii. p. 474 (1861) ; id. Lep. Ost-Sib. p. 34, pl. iii. 
fig. 10 (1864). 
Adopea sylvatica, Leech, Butt. China &c. p. 591, pl. xl. figs. 5g, 89 (1892-94). 
Hab. Amurland, Japan (Pryer); Korea (Leech). 


! ADOPAA TENEBROSA. 
Adopea tenebrosa, Leech, Butt. China &c. p. 591, pl. xl. figs. 6g, 92 (1892-94). 
The difference between this species and A. sylvatica are but trifling, and the genitalia 


290 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


afford no distinctive characters; as, however, it can be separated by its darker colour 
from A. sylvatica, we retain the name for the Chinese form. 
Hab. Kiukiang (Pratt). 


ADOPA NERVULATA. 


Pamphila nervulata, Mabille, Ann. Soc. Ent. Fr. 1876, p. lvi. 
Adopea nervulata, Leech, Butt. China &e. p. 592, pl. xl. fig. 3, g (1892-94). 


The insect here dealt with is that described and figured by Leech ; whether it is the 
true nervulata of Mabille remains doubtful, since the specimens do not possess the 
“striga anticarum sexuali nigra filiformi” mentioned in the original description. 


Hab. Moupin ( fide Mabille) ; Ta-Chien-lu, Wa-ssu-kow (fide Leech). 


! ADOPAIA LEONINA. 


Pamphila leonina, Butler, Cist. Ent. 11. p. 286 (1878). 


Thymelicus leonina, Staudinger, Mém. sur Lép. ii. p. 151, pl. viii. fig. 2, (1887), vi. p. 210 
(1892). 


Adopea leonina, Leech, Butt. China &c. p. 592, pl. xl. figs. 1 & 4g, 2? (1892-94). 
Hab. Japan (Pryer); Amurland ( fide Staudinger). 


ADOPAA ASTIGMATA, Leech. 
Adopea leonina, var. astigmata, Leech, Butt. China &e. p. 593, pl. xl. fig. 7,g (1892-94). 


Differs from typical Jeonina in wanting a sex-mark in the male and being rather 
more suffused with black. The male genitalia are similar to those of A. Jeonina, but 
so also are those of A. tenebrosa. 

Hab. Chang-yang, C. China (Pratt). 


ISMENE. 


Ismene, Swainson, Zool. Ill. i. pl. xvi. (1820-21) ; Moore, Lep. Cey. i. p. 157 (1881) ; Watson, 


Hesp. Ind. p. 9 (1891); id. P. Z. S. 1893, p. 125, pl. i. figs. 14-16, pl. ii. figs. 11, 12, pl. iii. 
fie. 18. 


This genus and the four which succeed it form a natural group well distinguished by 
the form of the palpi, which have the second joint appressed and densely scaled, and 
the third joint porrect, long, and slender, about half as long as the diameter of the eye. 
In most of the species of this genus the club of the antennz is as long as or longer than 
the shaft. Vein 5 in the fore wing straight, its base equidistant from veins 4 and 6 or 
(in mahintha and ‘tonis) a trifle nearer to vein 6 than to vein 4. Vein 3 of hind wing 
arising close to the end of the cell. 

The species known to us may be distinguished as follows :— 


10 (5). 
11 (4). 


A REVISION OF THE ORIENTAL HESPERIIDA. 


Males. 


. Fringe of hind wing orange-red, more particularly near the tornus. 
. Fore wing above with a sex-mark. 

. Sex-mark deep black, sharply defined. 

- Vein 1 a of fore wing distorted next the sex-mark. 

). Hind wing: vein 6 approaching vein 5 in a deep curve. 

. Costa of the hind wing folded over on to the upper surface of the 


apex. 


. Expanse 48 mm. Long hairs clothing dise of thorax and base of 


hind wing above brilliant greenish blue . . . 


. Expanse 66 mm. Long hairs clothing dise of thorax aad ase of 


hind wing above brown . . its 


. Costa of the hind wing not falda over on to heat upper sites of 


WHEN) Ga oe fot ot Souy hoe 
Hind wing: vein 6 straight . 
Vein | a of fore wing straight. 


13). Hind wing below : no straight pale band from the costa. Sex-mark 


running from the middle of vein 1 a to the base of vein 3, and 
composed of four short irregular stripes, of which the two in 
cell 1 @ are confluent 


. Hind wing below with a whitish postinedian bard running 


straight from the apical third of the costa to the apical fourth 
of vein 1 6. Sex-mark continuous, more than twice as high as 
wide, running from the middle of vein 1 @ to the base of vein 3. 


. Sex-mark fuliginous, suffused. 
. Fore wing below with a sharply defined whitish spot in the cell 


near its apical third, and a curved series of suffused pale spots, 
one each in cells 4-8 


subapical curved series of suffused pale spots 


. Sex-mark wanting. 

. Hind wing below not green, with black longitudinal lines. 

. Clothing of the front of palpi orange-yellow. 

. Expanse70 mm. Cell 6 of hind wing above concolorous : 
. Expanse 57 mm. Cell 6 of hind an above whitish, except at the 


base and apex . 


. Clothing of front of Pee grey. Cell 6 of hind wing above 


concolorous 


. Hind wing below eindrabieenten with fine eee aerial lines, 


for the most part two in each cell, equidistant from each other 
and from the veins. 


. Fore wing above plain brown, except a pale streak uext the basal 


half of the costa . 


291 


edipodea, Sw. 
edipus, Stgr. 


ataphus, Wats. 
tuckeri, n. sp. 


mahintha, Moore. 


ionis, de Nicéy. 


; Be 6 7 - + + Jaina, Moore. 
. No sharply defined whitish spot in the zal of fore wing teins nor 


fergusonii, de Nicév. 


etelka, Hew. 


harisa, Moore, 


anadi, de Nicéy. 


vasutana, Moore. 


292 . MESSRS. H. J. ELWES AND JAMES EDWARDS: 


5 (24). Fore wing above fulvous, passing into brown beyond the middle ; 
basal half of veins 1 a, 2, and 3 (except the extreme base of 
vein 1 a) broadly margined with black, the confluent bases of 
veins 7-I] also black . . . - . « » - + «~~. ' « septentrionis, Feld. 
26 (1). No orange-red in fringe of hind wing. 
27 (32). Hind below green or whitish green, the veins broadly bordered 
with black. Tibial pencil yellow or pale. 
28 (29). Hind wing below uniformly striped with black and green 
throughout . . . . . + » amara, Moore. 
29 (28). Hind wing with a broad palle ee fieanoks the ans 
0 (31). This stripe divided beyond the transverse vein by the dark borders 
of vein 5. Fore wing above with the basal half of cell 2 grey- 
brown, a little paler than the apical half. . . . .. . gomata, Moore. 
31 (30). This stripe continued to the termen, the dark borders to vein 5 
only indicated at the extreme apex. Fore wing above with the 
basal half of cell 2 sordid yellowish white . . . . . «= dara, Leech. 
32 (27). Hind wing below pale brown. Tibial pencil dark Giown . . . aquilina, Speyer. 


! ISMENE GEDIPODEA. 
Ismene edipodea, Swainson, Zool. Ill. i. pl. xvi. (1820-21). 
We have only a single male of this species from Java, and rely on Mr. Watson for its 


identification with the description of Swainson. 
Hab. Java (Piepers); Sumatra (fide de Nicéville); Palawan (in coll. Rothschild). 


! ISMENE G@DIPUS. 

Ismene edipus, Staudinger, MS. 

We have a single specimen of this species sent by Dr. Staudinger with the name of 
adipus. ‘The species does not appear to have been described, but it is distinguished 
from cdipodea by its much larger size, and the fact that the long hairs clothing 
the centre of the thorax and the base of the hind wing above are brown instead of 
greenish blue. 

Expanse 66 mm. 

Hab. Sula Island ( fide Staudinger). 


! ISMENE ATAPHUS. 


Ismene ataphus, Watson, P. Z. 8. 1898, p. 126. 
Ismene edipodea, Moore, Lep. Cey. i. p. 158, pl. lxiv. figs. 2a, 6 (1881) ; Watson, Hesp. Ind. p. 10 
(1891). 
The female of this species resembles the male, but wants the black patch near the 
base of the fore wing above, and the long hairs clothing the base of the wings show, in 
certain aspects, a brilliant greenish-blue colour. 


Hab, Kangra (Hocking) ; Sikkim (Moller) ; Khasia (Hamilton) ; E. Pegu (Doherty). 


A REVISION OF THE ORIENTAL HESPERIIDA. 293 


! ISMENE TUCKERI, n. sp. (Plate XX. fig. 4, ¢.) 

g. Not distinguishable on the upperside from the male of J. ataphus, Wats., but 
differs from that species in the following particulars:—Fore wing below having 
the yellow tinge replaced by greenish. Hind wing, vein 8 evenly curved throughout, 
vein 6 straight; underside pale greenish brown with pale green streaks very narrowly 
edged with purple, the latter placed as follows: two in cell 10, one in each of cells 
2 to 8, and one near the middle of the cell. 

Expanse 44 mm. 

Hab. Tavoy (Tucker). 

Described from one example in coll. Elwes, taken in Feb. 1892. 


! ISMENE JAINA. 
Ismene jaina, Moore, P. Z. S. 1865, p. 782 ; Watson, Hesp. Ind. p. 10 (1891). 


Hab. Sikkim (MGiller); Khasia (Hamilton); E. Pegu (Doherty). 


! ISMENE FERGUSONII. 
Tsmene fergusonii, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1892, p. 345, pl. J. fig. 6, g. 

Hab. Nilgiri hills (Hampson); N. Canara (coll. Swinhoe). 

Female resembling the male, save that the sex-mark is wanting and the long hairs 
clothing the disc of the thorax and the base of the wings are bluish grey. 


! ISMENE ETELKA. 
Tsmene etelka, Hewitson, Exot. Butt. iv., Ismene, figs. 14, 15 (1867). 


This species is easily distinguished in the male from L. fergusonii by the absence 
of the sexual patch on the fore wing and the much deeper and more extended 
orange-red markings below. The female on the upperside is brown with a purple 
shade, passing into greenish blue towards the base of the wings; on the fore wing 
below there is a suffused postmedian macular pale band, and the disc of the wing has 
a deep purple shade. 

Hab. Kina Balu (Waterstradt); Singapore (coll. Staudinger); Lawas, N. Borneo 
(Everett). 


! ISMENE HARISA. 
Ismene harisa, Moore, P. Z. 8. 1865, p. 782. 
Chouaspes harisa, de Nicéville, Jour. As. Soc. Beng. 1883, p. 84, pl. x. fig. 8, ¢ ; Distant, Rhop. 
Mal. p. 378, pl. xxxiv. fig. 22, ¢ (1886); Watson, Hesp. Ind. p. 6 (1891). 
Female brown above, with a metallic greenish-blue shade from the base of the wings, 
VOL. XIV.—PART Iv. No. 25.—October, 1897. 2Q 


294 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


but not covering the apical area of the fore wing or the terminal area of the hind 
wing. 

Herr Snellen sent a female specimen from Java, not distinguishable from this 
species, with the name consobrina, Plotz. 

Hab. Sikkim (Mller); Khasia (Hamilton) ; Nagas, Arjuno, Java (Doherty); Burmah 
(Watson); West Java (Piepers). 


!ISMENE ANADI. 
Choaspes anadi, de Nicéville, Jour. As. Soc. Beng. 1883, p. 83, pl. x. fig. 6, ¢ ; Watson, Hesp 
Ind. p. 7 (1891). 
The female of this species is said to differ from the male only in being larger and 
darker, the upperside of the hind wing concolorous with the rest of the wiug, not 
broadly pale ochreous as in the male; this is remarkable when we consider the close 


relationship of this species to J. harisa, which has a dissimilar female. 
Hab. Sikkim (M6ller); Khasia (Hamilton). 


! ISMENE VASUTANA. 


Ismene vasutana, Moore, P. Z.S. 1865, p. 782. 
Choaspes vasutana, Watson, Hesp. Ind. p. 8 (1891). 

This species is remarkable amongst its congeners for the greater or less development 
of hyaline spots in the fore wing; the full complement of these is one in each of cells 2 
and 3, but in the more usual state of the male that in cell 3 only is visible on the 
upperside, that in cell 2 being traceable on the underside only or wanting altogether ; 
sometimes these spots are quite absent from the upperside and their presence is only 
indicated by a pale dot near the base of cell 3 on the underside. 

It appears from Mr. de Nicéville’s remarks (Jour. Bomb. Nat. Hist. Soc. 1895, 
p- 404) that a somewhat similar development of these pale spots occurs in 7. mahintha, 
Moore. 

The female is brown above, with a pale greyish-green shade from the base of the 
wings, passing into purple near the middle of the fore wing and about the apical third 
of the hind wing; the hyaline spots on the fore wing are well developed. 

Hab. Sikkim (Moller); Nagas (Doherty). 


! ISMENE SEPTENTRIONIS. 
Ismene septentrionis, Felder, Reise Noy., Lep. iii. p. 525, pl. Ixxiii. fig. 3 (1867); Leech, Butt. 
China, &e. p. 636 (1894). 
Ismene striata, Hewitson, Exot. Butt. iv., Ismene, pl. i. figs. 6, 7 (1867). 
Hab. Shanghai (Dr. Muirhead, fide Felder) ; China (fide Hewitson); Siao-Lou 
(ex coll. Oberthiir). 


A REVISION OF THE ORIENTAL HESPERIID#, 


i) 
oo 
Or 


! ISMENE MAHINTHA. 

Ismene mahintha, Moore, P. Z.S. 1874, p. 575, pl. xlvii. fig. 4, ¢ ; Watson, Hesp. Ind. p. 11 
(1891). 

A female of this species is recorded by Wood-Mason and de Nicéville (Jour. As. Soc. 
Beng. 1886, p. 378) as taken at Silcuri, Cachar, on 7th June; but this sex does not 
appear to have been separately described and we have never seen it. 

Hab. Bernardmyo, Burmah (Doherty). 


! ISMENE IONIS. 
Ismene ionis, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 403, pl. Q. fig. 61, ¢. 
Hab. Sambawa (Doherty); West Java (Fruhstorfer, fide de Nicéville). 


! ISMENE AMARA. 

Ismene amara, Moore, P. Z. 8. 1865, p. 783. 
Choaspes amara, Watson, Hesp. Ind. p. 8 (1891). 

The female of this species usually closely resembles the male; but a specimen from 
the Naga hills has the upperside a little darker than usual, and the veins of the fore 
wing bordered with dark purple, broadly for some distance from the base, the purple 
colour gradually narrowing and not reaching the termen. 


Hab. Sikkim (Moller); Khasia (Hamilton); Nagas (Doherty). 


! ISMENE GOMATA. 
Ismene gomata, Moore, P. Z. 8. 1865, p. 783, 3. 
Ismene lorquini, Mabille, Bull. Soc. Ent. Fr. 1876, p. x; id. Ann. Soc. Ent. Fr. 1876, p. 266 (jide 
Semper). 
Choaspes gomata, de Nicéville, Jour. As. Soc. Beng. 1883, p. 83, pl. x. fig. 7, ? ; Watson, Hesp. 
Ind. p. 7 (1891). 

Female with the upperside purple-brown with a metallic bluish-green shade, which 
is strongest towards the base of the wings; underside as in the male, save that the 
ground-colour of the fore wing is dark purple-brown and there is in each of cells 2 
and 3 a suffused oblong pale spot. 

A female specimen kindly sent by Herr G. Semper as the Choaspes lorquini 
of his work only differs from a female of J. gomata from Java in having the suffused 
oblong pale spots in cells 2 and 3 of the fore wing below visible also on the upperside. 

Hab. Sikkim (Mller); Java (Piepers) ; Philippines (Semper). 

!IsMENE LARA. (Plate XX. fig. 14, 2.) 

Tsmene gomata, var. lara, Leech, Butt. China, &e. p. 634, pl. xxxix. fig. 12 (1894). 

With regard to specific validity this insect stands in the same relation to J. gomata, 

Moore, as J. fergusonii, de Nicév., does to I. jaina, Moore. 


Although the sexes resemble the respective sexes of J. gomata, both may be 
2Q2 


296 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


distinguished from that species by their slightly larger size and the much greater 
development of the pale markings on both upper and under sides. 
Hab. Western China (Pratt). 


! ISMENE AQUILINA. 
Ismene aquilina, Speyer, Stett. ent. Zeit. 1879, p. 346; Staudinger, Rom. Mém. sur Lép. vi. 
p. 214 (1892) ; Leech, Butt. China, &e. p. 635 (1894). 
Ismene jankowskii, Oberthiir, Etud. d’Ent. y. p. 23, pl. i. fig. 2 (1880). 
Proteides chryseglia, Butler, P. Z. S. 1881, p. 856. 


According to the figure the female of J. jankowskii, Ob. (the only sex figured), differs 
from females in Elwes’s collection from Yesso in having the spots on the fore wing above 
not so pale and the pale postmedian band continued towards the dorsum by two spots in 
cell 1a divided by the intraneural fold, instead of stopping short at vein 2; but we 
agree with Staudinger in considering this a synomym of aquilina. 

Hab. Japan (Pryer); Amur (Christoph). 


HAsora. 


Hasora, Moore, Lep. Cey. i. p. 159 (1881) ; Watson, P Z S. 1893, p. 127. Type dadra, Moore. 
Parata, Moore, t. c. p. 160. Type chromus, Cr. 

Vein 1a of fore wing angularly bent near the base in both sexes. Tornus of hind 
wing distinctly produced. Vein 2 of fore wing arising nearer the base in the male 
than in the female. Pale spots on the fore wing better developed in the female than 
in the male, and sometimes present in that sex only. Hind tibie in the male fringed. 

The species known to us we distingush as folliows :— 


1 (16). Hind wing below without a pale band from the costa towards 
the tornus. 
2 (13). Fringe of the hind wing not yellow in the tornal region. 
3 (8). Hind wing below with a pale spot in the cell, next the 
transyerse vein. 
4 (7). Termen of hind wing distinctly concave from vein 3 to vein 1 8, 
the tornus therefore distinctly produced. 
5 (6). Hind wing below: pale cell-spot small and roundish, less than 
half as wide as the cell. . “ineisy = go limbus “eee .oaaraMloore; 
6 (5). Hind wing below: pale cell-spot large and subquadrate, 
nearly as wide as the cell, the pale spot in cell 1 4 correlatively 
larger SSR eA Se . gneus, Plotz. 
7 (4). Termen of hind wing straight from vein 3 to vein 14; the hind 
wing therefore simply angulated at the apex of vem 15 . . anura, de Nicéy. 
8 (3). No pale spot in the cell on hind wing below. 
9 (12). Sex-mark on the fore wing of the male a broad seam of modified 
scales passing from the dorsum near the middle to the base 
of vein 3. 


10 (11). 


15 (14). 


16 (1). 
17 (24). 
, 

20). 


20 (19). 


0 (25). 


A REVISION OF THE ORIENTAL HESPERIID#A, 


Underside brown: fore wing with a large suffused subtriangular 
dark purple patch adjoming the costa from the middle to 
about the apical sixth ; hind wing with a suffused dark purple 
median band from the costa towards the dorsum 


. Purple suffusion of underside much less evident than in celenus, 


almost absent . 


. No sex-mark on the fore wing of ‘ie sane tind wing below 


~ in the male pale purple, with a pale cell-spot and postmedian 
band faintly indicated; in the female the purple is replaced 
ly TAB EEEN 6 5 5 5 5 6 - Hoe a We 


. Fringe of the hind wing yellow in ilte ‘ovnal region. 
. Hind wing below sharply divided just beyond the middle into 


a dark basal and a paler terminal portion, the latter chrome- 
yellow from the tornus and passing gradually into pale brown 
WGP USING 6 9 c . 

Hind wing below brown, ih. a deen black onl ante receded 
by a pale ray in cell 1a and an oblong pale yellow spot in 
el) 5 5 o 5 o © nas Ob ethos SOs ORION Mc 

Hind wing below with a pale hand from the costa towards the 
tornus. 

Pale band on hind wing below interrupted near vein 1 0. 

No sex-mark on fere wing above in the male. 

No pale point in cell 6 of the fore wing. Pale band on hind wing 
below narrow bluish white, both its edges somewhat suffused. 

A pale point in cell 6 of fore wing. Pale band on hind wing 
below with its inner edge usually more sharply defined than 
Wa G@mneG5 6 6 4 30 6 0 00 a oD Oo 6 


. Male with a sex-mark on fore wing above. 
. Expanse 39-48 mm. Tegumen with two pairs of long curved 


horns 


. Expanse 46-50 mm. Teeuitiet without eae ; ae 

. Pale band on hind wing below not interrupted near vein 1 6. 

. Pale band on hind wing below white. 

. Hind wing below from the base to the white band purple, more 


or less tinged with green. 


. Fore wing in the male with a white spot in cell 6. Pale band 


on hind wing below broad, suffused, tinged with purple on 
the edges and near the dorsum, Sreee the latter in its 


preapical fourth, . . . vB 


. No white spot in cell 6 of the fore 5 wing of the ‘Bal : : 
. Hind wing below from the base to the white band plain brown, 


the white band constricted near vein 8 to about half of its 
previous width, its outer edge straight . 
Pale band on hind wing below yellow. 


celenus, Cr. 


simplicissima, Mab. 


violacea, Stgr. 


myra, Hew. 


mUS, 0. SP. 


mestissima, Mao. 


chabrona, Plotz. 


chromus, Cr. 
inermis, 0. sp. 


proximata, Stgr. 
proxissima, D. sp. 


borneensis, n. sp. 


297 


298 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


31 (32). Hind wing above brown, with a broad yellow median band from 
the costa to the tornus. The pale cell-spot and those in cells 
2 and 8 on the fore wing above sharply defined hyaline and 
(CONMOMOUS)., ch sik eh, s, Seamer Areal wet ewe eee chuza, Hew. 
32 (81). Hind wing above yellow, with a broad brown band along the 
costa and termen, receding a little from the latter between 
veins 7 and1%. The pale cell-spot and those in cells 2 and3 
of the fore wing above suffused, contiguous, not denuded of 


coloured(scales 15. as: ooelaueey Here oes Le peo mace gediog-atlew: 


! HASORA BADRA. 
Goniloba badra, Moore, P.Z.S. 1865, p. 778. 
Hasora badra, Moore, Lep. Cey. i. p. 159, pl. Ixv. figs. 4, 4a (1881); Watson, Hesp. Ind. 
p- 12 (1891). 
Ismene quadripunctata, Mabille, sec. spec. comm. 
Hab. Sikkim (Moller); Khasia (Hamilton); Akyab (Adamson); Tavoy (Tucker) ; 
Nias (Modigliani) ; Pulo Laut, Bali (Doherty). 


! HASORA GNABUS. 


Ismene gneus, Plétz, Stett. ent. Zeit. xlv. p. 58 (1884). 
Ismene badra, var. celebica, Staudinger, Iris, ii. p. 188 (1889). 
Hasora gneus, Semper, Schmett. Philipp. p. 290 (1892). 

Hab. Mindanao (Semper); Mindoro, Palawan (Staudinger). 

We accept Semper’s identification of this form. It is near to badra, but the absence 
of purple gloss on the underside and conspicuously larger spot at base of hind wing 
below, which seems constant, appear to justify its separation. 


!HASORA ANURA. 
Hasora anura, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p.170, pl. B. figs.5 ¢,1 9 ; 
Watson, Hesp. Ind. p. 12 (1891) ; Leech, Butt. China, &c. p. 639, pl. xxxix. fig. 10, 9 (1894). 


Hab. Sikkim (Méller); Moupin (Kricheldorf). 


HAsorA HADRIA. 
Hasora hadria, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1889, p. 172. 
Hasora badra, Distant, Rhop. Mal. p. 374, pl. xxxv. fig. 3, g (1886), fide de Nicéville. 
We have not seen this species, and therefore transcribe the author’s account of it :— 
“ ? HuspeRIA BADRA, Butler (nec Moore), Trans. Linn. Soc. Lond., Zoology, second 
series, vol. i. p. 554. n. 3 (1877); Hasora badra, Distant (nec Moore), Rhop. Malay. 
p. 374. n. 1, pl. xxxv. fig. 3, male (1886). 
‘ Habitat. Perak, ? Malacca. 
““Expanse: ¢, 2°] inches. 


A REVISION OF THE ORIENTAL HESPERIID@. 299 


“Description: Male. Upperside both wings as in H. anura, mihi. Fore wing 
lacking the subapical yellow dot (this, however, is a trivial character). Underside 
both wings dull brown, not slightly glossed with purple as in H. anura, or strongly so 
as in [7. badra, Moore. Hind wing with a small anal lobe bearing a black patch, in 
H. anura there is no black patch or anal lobe, in H. badra both are large. ‘This 
species is probably variable with regard to the presence or absence of a white or 
greyish spot in the cell of the hind wing on the underside, and a white or greyish 
streak above the anal angle, as in the two allied species above named; Mr. Distant 
describing a ‘ var.’ of this species as lacking these characters. I have not figured this 
species, as Mr. Distant has done so in his ‘ Rhopalocera Malayana. Ihave described 
it from a single male from Perak in the collection of the Indian Museum, Calcutta, 
which Mr. Distant ticketed ‘ Hasora badra,’ Moore {var.).” 

This is, to us, a doubtful species, as the description is contradictory. First 
we read, ‘upperside both wings as in H. anura,” and then a few lines afterwards 
“hind wing with a small anal lobe bearing a black patch, in H. anura there is no 
black patch or anal lobe.” It is true that the species is said to be represented by 
Distant’s figure, but there is no more anal lobe in that figure than in H. anura (of 
which we have specimens), and certainly no black patch near the tornus of the hind 


wing below. 


! HASORA CELANUS. 
Papilio celenus, Cram. Pap. Exot. iv. p. 393, A, B. 
Hasora celenus, de Nicéville, Jour. As. Soc. Beng. 1895, p. 554. 


Hab. Amboina, Sumatra (fide de Niceville). 


! HASORA SIMPLICISSIMA. 
Ismene simplicissima, Mabille, Bull. Soc. Ent. Fr. sér. 5, vol. vi. p. xxv (1876) ; Staudinger, Iris, ii. 


p. 138 (1889). 
Hasora simplicissima, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 405, pl. Q. 


figs. 62 g, 63 @. 
Ismene philetas, Plétz, Stett. ent. Zeit. xlv. p. 56 (1884). 
Parata simplicissima, Semper, Schmett. Philipp. p. 292 (1892). 
Hab. Philippines (Semper); Palawan (Platen, fide Stgr.); Burmah, Sumatra (fide de 
Nicéville) ; Bali (Doherty). 


! HasoRA VIOLACEA. 

Tsmene violaceus, Staudinger, MS. ? 

On the upperside the female of this species is brown with a violet-purple shade, which 
is strongest in the apical region of the fore wing and on the lobe of the hind wing. 
‘This purple shade is absent or but faintly indicated in the male. 

Hab. Amboina (coll. Stg’.). 


800 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


HASORA MAISTISSIMA. 
Tsmene mestissima, Mabille, Bull. Soc. Ent. Fr. 1876, p. xxv; id. Ann. Soc. Ent. Fr. 1876, p. 263; 
Staudinger, Iris, ii. p. 136 (1889). 
Hab. Mindanao (Semper); Palawan (Platen, fide Stgr.); Kina Balu (Waterstradt). 


HASORA VITTA. 
Hesperia vitta, Butl. Trans. Ent. Soc. 1870, p. 498; id. Lep. Ex. pl. lxix. fig. 9; ¢f. de Nicéville 
Jour. Bomb. Nat. Hist. Soc. 1895, p. 408. 
? Hasora vitta, Swinh. Trans. Ent. Soc. 1893, p. 329. 


Hab. Sarawak (Low, fide Butler). 

We have never seen a specimen of this species, which is described and figured as 
without any apical spot on the fore wing. We have seen a Bornean specimen from 
Staudinger’s collection which has this spot, which is constant in chabrona from all 
localities. ‘There is no other character by which we can distinguish vitta, Butler, as 
described, and we are therefore unable to say whether it is a good species confined to 
Borneo, as de Nicéville suggests, or a mere aberration of chabrona. 


! HASORA CHABRONA. 
Ismene chabrona, Plétz, Stett. ent. Zeit. xlv. p. 56 (1884). 
Hasora vitta, Distant (nec Butl.), Rhop. Mal. p. 375, pl. xxxv. fig. 4, ¢ (1886) ; Semper, Schmett. 
Philipp. p. 291 (1892), sec. spec. comm. 
Hasora chabrona, de Nicéville, Jour. Bomb. Nat. Hist. Soc. 1895, p. 406. 
Hab. Sikkim (Méller); Assam, E. Pegu (Doherty); Andamans (de Roepstor(f); 
Java (Piepers); Kina Balu (Waterstradt). 


HAsoRA COULTERI. 

Hasora coulteri, Wood-Mason & de Nicéville, Jour. As. Soc. Beng. 1886, p. 378, pl. xviii. 
figs. 8 ¢, 8a, 8b 9; Watson, Hesp. Ind. p. 14 (1891); de Nicéville, Jour. Bomb. Nat. 
Hist. Soc. 1895, pp. 407, 408. 

We have not seen this species. It ‘“‘is very closely allied to H. chabrona ; the females 
of the two species may be distinguished on the underside of the hind wing by both 
the edges of the discal white band being sharply defined in H. coulteri, much blurred 
in H. chabrona. The female of H. coulteri possesses the subapical spot to the fore 
wing, which is lacking in the male, and by the absence of which, together with the 
discal band, it is distinguished from the same sex of H. chabrona.” The male has not 
the male mark as in chromus, the upperside of the fore wing being “ without spots, 
but with three ill-defined discal bands composed of modified scales arranged along each 
side of the submedian nervure, and of the first and second median nervules, and 
probably concealed by sete in the living insect.” 

Hab. Sileuri, Cachar (Wood-Mason & de Nicéville). 


A REVISION OF THE ORIENTAL HESPERIIDA. 301 


Hasora coromus. (Plate XXVII. figs. 93, 938 a.) 

Papilio chromus, Cramer, Pap. Exot. iii. pl. eclxxxiv. fig. E, ¢ (1782). 

Tsmene malayana, Velder, Wien. ent. Mon. iv. p. 401 (1860) ; id. Reise Nov., Lep. iii. pl. lxxii. fig. 15 
(1866). 

Parata chromus, Moore, Lep. Cey. i. p. 161, pl. Ixv. figs. 1 a, 6 (1881); Watson, Hesp. Ind. p. 16 
(1891). 

Parata malayana, Watson, t. ec. p. 18. 

Parata alexis, Moore, |. c. figs. 2 a, 25; Watson, t. c. p. 17. 

Hasora chromus, Leech, Butt. China, &c. p. 638, pl. xxxix. fig. 7 (1894). 

‘This species varies much in size ; the smallest specimen I have expands 39 millim. The 
pale band on the hind wing below varies in width and shape; its inner edge is always 
straight, but its outer edge may be either straight, when the band is narrow (about 
1 mm. wide ora mere line), or sinuate, when the band is about 3 mm. wide in 
its widest part. In Sikkim specimens the brown ground-colour of the hind wing 
below is washed with metallic purple inclining to green, and in specimens from the 
Andamans the green colour predominates over the purple, except in the space beyond 
the pale band (these latter seem to be the true malayana, Feld.) ; in some specimens 
the greenish-purple colour is scarcely visible in the space beyond the pale band; these 
differences, however, are not correlated with differences in locality and cannot usefully 
be made the basis for grouping specimens. 

Hab. Sikkim (Maller); Khasia (Elwes); Burmah (Doherty); Andamans (de Roep- 
storff) ; Ceylon, Bangalore (Minchin) ; Pulo Laut (Doherty) ; Java (Piepers). 


! HASORA INERMIS, n. sp. (Plate XX. fig. 16; Plate X XVII. figs. 94, 94 a.) 


This species closely resembles H. chromus, from which it differs by its larger size 
and the different form of the male genitalia. ‘The male has the sex-mark on the fore 
wing above and wants the pale point in cell 6 of that wing; the female resembles 
that sex of H. chabrona in having on the fore wing a somewhat crescent-shaped 
whitish hyaline spot in each of cells 2 and 3, and a whitish point near the basal third 
of cell 6. 

Expanse 46-50 mm. 

Hab. Liu Kiu Islands (Pryer). 

Described from one pair in coll. Elwes. The species can only be distinguished with 
certainty by the male genitalia. 


HASORA PROXIMATA. 


Ismene proximata, Staudinger, Iris, 11. p. 137 (1889). 
Hasora proximata, Semper, Schmett. Philipp. p. 291, pl. xlix. fig. 6, 2 (1852). 


Hab. Mindanao (Semper); Palawan, Celebes (Platen, fide Stgr.). 
von. XIv.—Part Iv. No. 26.—October, 1897. 2R 


302 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


HAsoRA PROXISSIMA, n. sp. (Plate XXI. fig. 10, 3.) 

g. Upperside brown. Underside: fore wing brown, paler along the dorsum, basal 
third of the dise thickly clothed with coarse darker scales, a purple streak next the 
upper edge of the cell and a curved series of suffused purple spots beyond the end of 
the cell from the costa to cell 3: hind wing with a white band from just before the 
apex of the costa to the dorsum just before the tornus, from the base to the band 
purple-brown shot with green, beyond the band plain brown; the inner edge of the 
band is evenly curved (the convexity towards the base of the wing) from the costa to 
vein 1 6, whence it runs straight to the dorsum and is tinged with purple and green, 
the outer edge is nearly straight from the costa to vein 1 6, where it is angulated. 

Expanse 41 mm. 

Hab. Mindoro (coll. Stgr.). 

Described from one specimen ex coll. Staudinger. 

H. proximata and H. proxissima belong to a group, not represented in our region by 
any species, which we may call the hurama group, because they agree with that species from 
North Australia in having the band on the hind wing below continuous from the costa to 
the dorsum, whilst in all the specimens in Elwes’s coll. of chabrona (10) and chromus (30) 
this band, though very variable in breadth, coloration, and distinctness, is interrupted 
near vein 1 by the black subtornal patch. The differences between proaissima and 
proaimata are precisely those between vitta, Butl., and chabrona, Plotz, namely the 
presence or absence of the apical spot on the fore wing. ‘This may or may not bea 
constant character, but it is constant in the only species of which we have a good series, 
namely chabrona. All these four species want the sex-mark which is so conspicuous in 
H. chromus, and the round spot at base of hind wing below which distinguishes the 
badra group. 


! HASORA BORNEENSIS, n. sp. (Plate XX. figs. 8 g, 112.) 


3. Upperside brown, the dorsal half of the hind wing yellow-brown, by reason of 
the thick clothing of paler hair-scales. Underside: fore wing brown, with a suffused 
whitish > with its apex to the termen between veins | @ and 3, transverse vein indis- 
tinctly pale, and a suffused indistinct macular pale band from beyond the end of the cell 
towards the tornus as faras cell 2: hind wing brown, with a cream-white band from the 
preapical fifth of vein 8 to the dorsum just before the tornus, the outer edge of this band 
is straight from vein 8 to the apical seventh of vein 14, where the short piece which runs 
to the dorsum forms an obtuse angle with the remainder, its inner edge is almost straight 
from vein 8 to the base of vein 3, whence it is irregularly curved outward, and the width 
of the band is thus reduced to about one-half of its previous width; tornal region somewhat 
darker than the remainder of the wing, but without any black patch. Fringes pale 
grey, becoming paler towards the tornus of each wing; the short scales grey-brown. 
Antenne brown, the club and apiculus reddish beneath. Body above concolorous with 


A REVISION OF THE ORIENTAL HESPERIIDA, 303 


the wings. Second joint of palpi clothed with grey and black hairs intermixed. 
‘Clothing of breast and legs brown, of the abdomen beneath ochreous. 

?. Upperside: fore wing brown, with six yellowish-white hyaline spots—one about 
as high as wide across the apical third of the cell, its outer edge notched, one about 
one and a half times as high as wide lying obliquely across cell 2, its outer edge concave, 
one about one and a half times as high as wide with its outer edge concave across cell 3, 
and three others small and oblong or suboval, one in each of cells 6, 7, and 8 ; hind 
wing dark brown, the dorsal half thickly clothed with bright ochreous hair-scales. 
Underside : fore wing brown, the costal region as far as the end of the cell paler ; 
the hyaline spots as on the upperside, a subtriangular white spot next the upperside of 
vein 1 a beyond the middle and sometimes a minute suffused white spot in cell 1 @ next 
the lower outer angle of the hyaline spot in cell 2. Otherwise as in the male. 

Expanse 46-50 mm. 

Hab. Kina Balu, Borneo (Waterstradt). 

Described from two pairs ex coll. Staudinger. 


! HASORA MYRA. 

Ismene myra, Hewitson, Exot. Butt. iv., Ismene, pl. i. fig. 8, 5 (1867). 

We give a full description of both sexes of this species; the female doés not appear 
to have been described. 

3. Upperside: fore wing warm brown; hind wing brown from the costa about 
as far as vein 3, thence to the apex ochre-yellow ; pubescence of both wings yellow- 
brown. Underside: fore wing grey-brown, suffusedly paler along the dorsum, beyond 
the end of the cell, and at the apex: hind wing dark brown from the base to beyond 
the middle, the limit of that colour well defined and passing from the costa at the 
level of the apical fourth of vein 6 in a straight line to the apical third of vein 14, 
where it is broken by a pale streak along the course of the vein last-named and then 
passes to the dorsum; the space beyond this dark basal shade is pale brown in the 
apical region and ochre-yellow in the tornal region, those colours passing gradually the 
one into the other. Fringe of the fore wing brown, a little paler towards the tornal 
angle; of the hind wing brown at the apex, passing gradually into ochre-yellow as the 
tornal angle is approached. Clothing of body and legs yellow-brown, paler on the 
abdomen. 

9. Fore wing similar to that of the male but with hyaline spots placed as follows :— 
a small triangular one in the cell near the base of vein 3, one in each of cells 2 and 3, 
narrow oblique and nearly or quite reaching from side to side of those cells, and one 
(small and roundish) in each of cells6 and 7. Hind wing similar to that of the male, but 
with the yellow portion more extensive and the limits of the brown and yellow colours 
more sharply defined. 

Expanse, ¢ 46 mm., 2 52 mm. 

Hab. Java (Piepers). 

2R2 


804 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


! Hasora Mts, n. sp. (Plate XX. figs. 2 5, 5 2.) 

3. Upperside brown, inclining to yellow-brown towards the dorsum of the hind 
wing; subtornal region of the termen of the hind wing blackish. Underside: paler 
than the upperside and more decidedly yellow-brown, the middle of the dorsal region 
of the fore wing dull ochreous; tornus of the hind wing with a large deep black oblong 
patch from the dorsum nearly to vein 2, preceded by an oblong yellow patch near the 
apical third of cell 14 and a yellow ray in cell la. Fringe of the fore wing grey-brown ; 
of the hind wing grey-brown from the apex to near vein 5, thence gradually becoming 
yellow. Antenne blackish, a little paler on the underside. Body above concolorous 
with the wings. Clothing of second joint of palpi yellow, the lateral bristles black, 
that of the underside and legs dull ochreous. 

¢. Hind wing above blackish brown, the dorsal half thickly clothed with ochreous 
hair-scales, otherwise like the male. 

Expanse 363-383 mm. 

Hab. Kina Balu (Waterstradt). 

Described from a pair sent by Dr. Staudinger now in Elwes’s collection. He has 
other similar specimens, and we have seen in M. Oberthiir’s collection from Perak what 
may be the same species. 


! HAsoRa CHUZA. 
Ismene chuza, Hewitson, Exot. Butt. iv., Ismene, pl. i. fig. 4 (1867). 
Choaspes chuza, Distant, Rhop. Mal. p. 373, pl. xxxiv. fig. 27 (1886). 

Hab. Nagas, Upper Burmah, Pulo Laut (Doherty); S.E. Borneo (coll. Schénberg) ; 
Java (Piepers). 


! HASORA SAIDA. 


Tsmene saida, Hewitson, l. c. fig. 5, 2. 
Parata saida, Semper, Schmett. Philipp. p. 293 (1892). 
Parata gentiana, Semper, /. c., sec. spec. comm. 

Hab. Luzon, Samar, Bohol, Mindanao (Semper); Philippines (Hewitson). 

In the male the spots forming the pale discal band on the fore wing above are pale 
yellow, opaque, confluent, and suffused at the edges; in the female they are whitish 
hyaline, angular, and contiguous but sharply defined. Felder’s description of his 
Ismene gentiana (Reise Nov., Lep. iii. p. 527) agrees well with this insect, but not so 
his figures (¢. ¢. pl. Ixxii. figs. 18, 19). 


BIBasis. 


Bibasis, Moore, Lep. Ceyl. i. p. 160 (1881) ; Watson, P. Z.S. 1893, p. 128. Type sena, Moore. 


Fore wing: vein 1@ not distorted near the base. Hind tibie in the male thickly 
scaled above and bearing a pencil of hairs as in Zsmene. 


A REVISION OF THE ORIENTAL HESPERIIDA. 305 


The clothing of large flattened scales, which imparts to the hind tibie of the male 
in this genus and Jsmene their fusiform shape, really forms a case in which the tibial 
pencil is enclosed. 

We distinguish three species of this genus, of which the diagnostic characters are as 
follows :— 

1 (4). Pubescence of hind wing above lighter or darker yellow-brown. Upper 


edge of clasp with a spinose lobe near the base. 
2 (3). Outer edge of the pale band on the hind wing below suffused and 


becoming purplish . . . : cee : . sena, Moore. 
3 (2). Outer edge of the pale band on the isd wing elo as shar aly defied as 
theamneredee mse amen : Ct 277), 07,772 25 SEES De 
4 (1). Pubescence of the hind wing above ioright rust- yallow. No spinose lobe 
on the upper edge of the claspmearthe base . . . . . . . . . sambavana,n. sp. 


! BIBASIS SENA. 
Goniloba sena, Moore, P. Z. S. 1865, p. 778. 
Bibasis sena, Moore, Lep. Cey. i. p. 160, pl. lxv. figs. 3, 3@ (1881) ; Watson, Hesp. Ind. p. 15 
(1891). 


Hab. Sikkim (Moller) ; Khasias (Hamilton) ; Nagas (Doherty) ; N. Canara (Aitken). 


! BIBASIS UNIFORMIS, n. sp. (Plate X XVII. fig. 95.) 


Ismene sena, var. palawana, Staudinger, Iris, ii. p. 1389 (1889). 
Bibasis sena, var. palawana, Semper, Schmett. Philipp. p. 292 (1892). 


Differs from B. sena in having the outer edge of the white band on the hind wing 
below as sharply defined as the inner edge. Specimens from Java and Kina Balu, 
Borneo, have more or less whitish-purple suffusion, sometimes forming one or two spots 
near the end of the cell on the fore wing below, as in B. sena; this is entirely wanting 
in var. palawana, Stgr., which also has the pale subdorsal patch on the fore wing 
below pure white. 

Hab. Java (ex coll. Snellen); Kina Balu (Waterstradt); Palawan (fide Staudinger) ; 
Luzon (fide Semper). 

The male genitalia of Javan and Bornean specimens agree with those of B. sena, of 


which this is, doubtless, an insular form. 


! BIBASIS SAMBAVANA, n. sp. (Plate X XVII. fig. 96.) 

The single male example of this species is, unfortunately, not in a condition for 
detailed description, but as it, nevertheless, presents diagnostic characters it has been 
thought well to point these out. It appears probable that in the colour and pattern of 
the underside it will prove to resemble B. sena rather than B. uniformis; but, however 
this may be, it is readily distinguished from either by the bright rust-yellow hairy 


306 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


clothing of the inner part of the hind wing above, which in B. sena and B. uniformis 
is olive, and the different clasp-form. The latter will be best appreciated from the 
figures. 

Hab. Sambawa (Doherty). Type in coll. Elwes. 


BapDaMIA. 
Badamia, Moore, Lep. Cey. i. p. 156 (1881); Watson, P. Z.S. 1893, p. 128. Type exclamationis, 
Fab. 

Fore wing very narrow, its greatest width equal to three-fourths of the length of the 
termen ; hind wing with the termen deeply, almost semicircularly, concave from vein 3 
to vein 14. Vein 2 of fore wing arising near the base of the cellin both sexes. Vein 
3 of hind wing arising well before the end of cell. Hind tibie in the male fringed. 


BADAMIA EXCLAMATIONIS. 
Papilio exclamationis, Fabricius, Syst. Ent. p. 530 (1775). 
Papilio ladon, Cramer, Pap. Ex. iii. pl. eclxxxiv. fig. C, 2. 
Ismene thymbron, Felder, Sitzb. Ak. Wiss. Math.-nat. Cl. xl. p. 461, sep. p. 14 (1860). 
Badamia exclamationis, Moore, Lep. Cey. i. p. 157, pl. xvi. figs. 2, a, 6 (1881) ; Watson, Hesp. Ind. 
p. 3 (1891). 

The specimen in coll. Felder labelled in contemporary handwriting “ Zsmene thymbron, 
Feld.,” ““ Amboina, Doleschall,” is a male of B. exclamationis, only differing from Indian 
examples in the slightly warmer tint of the upperside. 

Hab. N.W. Himalaya (Young) ; Sikkim, Khasia (Elwes); Andamans (de Roepstorff’) ; 
Burmah, Bali, Pulo-Laut (Doherty). 


RHOPALOCAMPTA. 
Rhopalocampta, Wallengren, Rhop. Caffr. p. 4; Watson, P. Z.S. 1893, p.129. Type forestan, Cr. 
Choaspes, Moore, Lep. Cey. i. p. 158 (1881). Type benjamini, Guér. 

Watson diagnoses this genus by the absence of vein 5 in the hind wing, but this vein, 
though perhaps less evident than in the allied genera, is present in the three species 
which we possess The hind tibie in the male bear a long pencil the distal half of 
which is held close to the tibia by a thick fringe of hair-like scales. 

The species known to us we distinguish as follows :— 


1 (4). Hind wing above with the subtornal angulation black, more or less 
widely margined with yellow. 

2 (8). Hind wing shorter, vein 14 subequal in length to the dorsum of the 
fore wing. Pale space near the tornus of the hind wing below 
Ofange-Tedi. | moma ciaye ee te earch © a ee eee eae 

3 (2). Hind wing longer, vein 1 4 about one-fifth longer than the dorsum of 
the fore wing. Pale space near the tornus of the hind wing below 
lemon=yellow.— <) .< sia. a+- Settee eeeas Ses are ener Enum SC/OWD/ Ure wD ISLanEs 


benjamini, Guér. 


A REVISION OF THE ORIENTAL HESPERIID., 307 


4 (1). Hind wing above with the subtornal angulation and the adjacent region 

entirely yellow. 
5 (6). Upperside dark brown with a dull purple shade, pubescence of thorax 

above dull green; hind wing narrower, tornus more produced . . subcaudata, Feld. 
6 (5). Upperside brown with brilliant purple reflections, passing imto pale 

green towards the base of the wings, pubescence of thorax above 

bluish grey ; hind wing broader, its tornus less produced . . . . renidens, Mab. 


! RHOPALOCAMPTA BENJAMINI. 


Thymele benjamini, Guérin, Delessert’s Souv. Voy. Ind. ii. p. 79, pl. xxi. figs. 2, 2a (1843). 

Choaspes benjanini, Moore, Lep. Cey. i. p. 159, pl. Ixiv. figs. 1, a, b (1881); Watson, Hesp. Ind. 
p. 5 (1891). : 

Rhopalocampta benjamini, Leech, Butt. China &e. p. 641 (1894). 

As a rule the sexes may be distinguished by colour, the males being dark greenish 
and the females bluish at the base of the wings and on the body, but one Japanese 
male has the tint of the female. 

Hab. Kumaon (Ramsay); Sikkim (Moller); Khasia (Hamilton) ; Nagas (Doherty) ; 
Nilgiris (Morris) ; Ceylon (Green) ; Japan (Pryer, Leech); Moupin (Kricheldorf ). 


! RHOPALOCAMPTA CRAWFURDI. 


Choaspes crawfurdi, Distant, Rhop. Mal. p. 372, pl. xxxiv. fig. 26 (1886). 
Choaspes electra, Stgr. MSS. 


Hab. Province Wellesley (Distant); Perak, Pulo Laut (Doherty) ; Borneo (ex coll. 
Stgr.); Lawas, N. Borneo, April (Everett). 


! RHOPALOCAMPTA SUBCAUDATA. 

Ismene subcaudata, Felder, Reise Noy., Lep. iii. p. 526, pl. Ixxii. figs. 20, 21 (1867). 
Hab. Java (Piepers); Bali (Doherty). 
RHOPALOCAMPTA RENIDENS. 


Ismene renidens, Mabille, Compte Rendus Soc. Ent. Belg. iv. no. 16, p. Ixxviil (1891). 
Choaspes renidens, Semper, Schmett. Philipp. p. 289 (1892). : 


Hab. Philippines (Semper). 


EXPLANATION OF THE PLATES. 
PLATE XVIII. 


Fig. 1. Celenorrhinus balukinus, nu. sp., 3: p- Ts 
Fig. 2. 2 orbiferus, n. sp., 6: p- 118. 
Fig. 3. a inequalis, n. sp.. ¢: p. 119. 
Fig. 4 3 dentatus, n. sp.. ¢: p- 119. 
Fig. 5 ss fulvescens, n. sp., 6: p. 120. 


308 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


Fig. 6 Celenorrhinus saturatus, n. sp., 6: p. 120. 


Higa is 26 lativittus, n. sp. d: p. 121. 
Fig. 8. . maculicornis, n. sp., 6: p. 116. 
Hiswao: 33 affinis, n. sp., 2: p. 121. 

Fig. 10. 43 batchianus, n. sp., g: p. 122. 
Fig. 11. Coladenia agnioides, n. sp., 3d: p. 128. 

Fig. 12. . sobrina, n. sp.. ¢: p. 126. 


Fig. 13. Satarupa fumosa, n. sp., ¢: p. 133. 

Fig. 14. Suastus bipunctus, Swinh., 3: p. 180. 

Fig. 15. ,, sala, Hew., x3, ¢: p. 179. 

Fig. 16. Astictopterus olivascens, Moore, ¢: p. 172. 
=A henrici, Holland, 3: p. 172. 

. Tapena minuscula, n. sp., ¢: p. 14%. 

19. ,, ~=— hampsoni, n. sp., 6: p. 147. 

. Caprona saraya, Doherty, 3: p. 161. 

21. Baracus hampsoni, 3: p. 171. 

22. Scobura martini, n. sp., 2: p. 205. 

23. Pedestes maculicornis, n. sp., 6: p. 193. 

24. Arnetta vindhiana, Moore, g: p. 199. 

25. Pedestes fuscicornis, n. sp., d: p. 194. 

ig. 26. Lophoides purpurascens, n. sp., 36: p. 196. 

Fig. 27. 3; A 9: p. 196. 

Fig. 28. is binotatus, n. sp., 2: p. 196. 


coco) 
ie ee ein 
ug de a 
= 
coo™N 


dq oq 0g 
bo 
S 


de de dq do 


PLATE XIX. 
Fig. 1. Scobura umbrosa, n. sp., 2: p. 207. 
1, 4 »  bipunctata,n. sp., 2: p. 207. 
Fig. 3. Aeromachus stigmata, Moore, ¢, X%: p. 189. 
Fig. 4. Sepa cicatrosa, n. sp., 3: p. 212. 
Fig. 5. Parnara bipunctata, n. sp., ¢: p. 283. 
Fig. 6. Aeromachus discreta, Plotz, 6, X2: p. 190. 
Fig: 7. = imachus. Mén., 6, X$: p. 190. 
Fig. 8. Sepa guttulifera, nu. sp., ¢: p. 212. 
Fig. 9. Zographetus ogygioides, n. sp., ¢: p. 203. 
Fig. 10. Aeromachus dubius, n. sp.. 6, X23: p. 190. 
Fig. 11. a indistincta, Moore, 6, X#: p. 191. 
Fig. 12. Seobura concinna, n. sp., 2: p. 206. 
Fig. 13. Zographetus auriferus, n. sp.. ¢: p. 208. 
Fig. 14. Velicota prusias, Feld., 3: p. 251. 
Fig. 15. . simplex, n. sp. 3: p. 253. 


A REVISION OF THE ORIENTAL HESPERIIDA. 309 


Fig. 16. Scobura fenestrata, n. sp., 3: p. 206. 
Fig. 17. Plastingia aurantiaca, n. sp., 5: p. 228. 
Fig. 18. Se similis, n. sp., ¢: p. 230. 


Fig. 19. »  fruhstorferi, Mab., o: p. 227. 
Fig. 20. Gehenna angulifera, n. sp., 5: p. 244. 


a 


. Augiades majuscula, n. sp., 3: p. 249. 


9 
Fig. 22. Sepa ciliata, n. sp., $: p. 214. 
Fig. 23. Pamphila niveomaculatus, Ob., 2, * 3: p. 167. 
Fig. 24. Aeromachus javanicus, n. sp., 6, X$: p. 191. 
Fig. 25. Sepa cinnamomea, n. sp., o: p. 213. 
Fig. 25. Telicota insularis, n. sp., ¢: p. 252. 
Fig. 27. & sy 2: p. 252. 
Fig. 28. Augiades siva, Moore, 3: p. 247. 
PLATE XX. 
Fig. 1. Matapa purpurascens, n. sp., $: p. 209. 
Fig. 2. Hasora mus, n. sp., o: p. 304. 
Fig. 3. Notocrypta quadrata, n. sp., 5: p. 241. 
Fig. 4. Ismene tuckeri, n. sp., 3: p. 293. 
Fig. 5. Hasora mus, n. sp., 2: p. 304. 
Fig. 6. Watsonia swinhoei, n. sp., 3: p. 220. 
Fig. 7. Tagiades waterstradti, n. sp., 5: p. 143. 
Fig. 8. Hasora borneensis, n. sp., 5: p. 302. 
Fig. 9. Acerbas nitidifasciata, n. sp., $: p. 216. 
Fig. 10. Tagiades sambavana, n. sp., 3: p. 143. 
Fig. 11. Hasora borneensis, n. sp., 2: p. 302. 
Fig. 12. Tagiades nestus, Feld., ¢: p. 145. 
Fig. 13. » nana, n. sp., o: p. 144. 
Fig. 14. Ismene lara, Leech, 2: p. 295. 
Fig. 15. Tagiades titus, Plotz, 3: p. 142. 
Fig. 16. Hasora inermis, n. sp., o: p. 301. 
Fig. 17. Gangara sanguinocculus, Martin, ¢: p, 218. 
Fig. 18. Orthophetus lidderdali, Elwes, 3: p. 105. 
PLATE XXI. 
Fig. 1. Halpe debilis, n. sp., 3: p. 266. 
Fig. 2. ,, knyvetti, n. sp., o: p. 261. 
Fig. 3. Parnara leechii, n. sp., $: p. 274. 
Fig. 4. »  philippina, H.-S., 5: p. 276. 
Fig. 5. LHalpe debilis,n. sp., 2: p. 266. 
Fig. 6. ,,  majuscula, n. sp., $: p. 264. 


VoL. XIv.—PArT Iv. No. 27.—October, 1897. 2s 


310 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


7. Halpe fasciata, n. sp., 2: p. 262. 
Fig. 8. Parnara philippina, H.-S., 9: p. 276. 
9. Lambrix latifascia, n. sp., 3: p 
Fig. 10. Hasora proxissima, n. sp., 3: p. 302. 

Fig. 11. Parnara hasoroides, n. sp., 5: p. 284. 
Fig. 12. Taractrocera nicevillei, Wats., 3: p. 184. 
Fig. 13. Halpe beturia, Hew. (type): p. 263. 
Fig. 14. Pirdana albicornis, n. sp., g: p. 224. 
Fig. 15. Notocrypta inornata, n. sp., 5: p. 241. 
Fig. 16. Parnara disereta, n. sp., 3: p. 282. 

Fig. 17. Telicota rectifasciata, n. sp., 5: p. 254. 
Fig. 18. Ancistroides othonias, Hew.: p. 222. 
Fig. 19. Parnara robusta, n. sp., 5: p. 280. 

Fig. 20. Telicota concinna, n. sp., S$: p. 253. 
Fig. 21. »  dilutior, n. sp., $: p. 255. 

Fig. 22. Parnara simillima, n. sp., 3: p. 274. 
Fig. 23. 5 aurocitiata, n. sp., 3: p. 278. 
Fig. 24. Taractrocera ceramas, Hew., 3: p. 184. 
Fig. 25. Parnara cahira, Moore, 2: p. 278. 
Fig. 26. »  subochracea, Moore, 3: p. 275. 


PLATE XXII. 
Fig. 1. Celenorrhinus dhanada, Moore; dorsal aspect of the tegumen: p. 119. 
Fig. la. Do.; lateral aspect of the tegumen. 
Fig. 146. Do.; inner face of left clasp. 
Fig. 2. Celenorrhinus andamanica, Wood-Mason & de Nicéy.; dorsal aspect of the 
tegumen: p. 119. 
Fig. 2a. Do.; inner face of left clasp. 
Fig. 3. Celenorrhinus aurovittata, Moore ; dorsal aspect of the tegumen: p. 122. 
Fig. 3a. Do.; inner face of left clasp. 
Fig. 4. Celenorrhinus affinis, n. sp.; inner face of left clasp: p. 121. 
Fig. 5. Celenorrhinus saturatus, n. sp.; dorsal aspect of the tegumen: p. 120. 
Fig. 5a. Do.; inner face of left clasp. 
Fig. 6. Sarangesa purendra, Moore ; dorsal aspect of the tegumen: p. 124. 
Fig. 6a. Do.; inner face of left clasp. 
Fig. 7. Sarangesa dasahara, Moore; dorsal aspect of the tegumen: p. 124. 
Fig. 8. Sarangesa sati, de Nicéy.; dorsal aspect of the tegumen: p. 124. 
Fig. 9. Coladenia dan, Fab.; dorsal aspect of the tegumen: p. 127. 
Fig. 9a. Do.; inner face of left clasp. 
Fig. 10. Coladenia laxmi, de Nicéy.; inner face of left clasp: p. 126. 
Fig. 11. Coladenia agni, de Nicév.; inner face of left clasp: p. 127. 


A REVISION OF THE ORIENTAL HESPERIIDA., 311 


g. lla. Coladenia agni, de Nicév. ; lateral aspect of the tegumen. 

. 12. Coladenia agnioides, n. sp.; inner face of left clasp: p. 128. 

g. 12a. Do.; lateral aspect of the tegumen. 

g. 13. Tagiades atticus, Fab.; inner face of left clasp: p. 142. 

ig. 14. Tagiades sambavana, n. sp.; inner face of left clasp: p. 143. 

. 15. Tepena thwaitesi, Moore ; inner face of left clasp and dorsal aspect of apex 


more enlarged: p. 146. 


. 16. Tapena hampsoni, n. sp.; inner face of left clasp: p. 147. 
ig. 17. Tapena minuscula, n. sp.; inner face of left clasp: p. 147. 
ig. 18. Ctenoptilum vasava, Moore ; inner face of left clasp: p. 148. 
. 18a. Do.; dorsal aspect of left clasp. 


PLATE XXIII. 


. 19. Ctenoptilum chinensis, n. sp. ; inner face of left clasp: p. 148. 

. 19a. Do.; dorsal aspect of left clasp. 

ig. 20. Caprona ransonnettii, Feld. ; inner face of left clasp: p, 150. 

. 20a. Do.; inner face of right clasp. 

‘ig. 21. Caprona saraya, Doh. ; inner face of left clasp: p. 151. 

. 21a. Do.; inner face of right clasp. 

. 22. Hesperia proto, Esp.; inner face of left clasp: p. 159. 

g. 23. Hesperia staudingeri, Spey. ; iner face of left clasp: p. 159. 

. 24. Hesperia serratulw, H.-S.; inner face of left clasp: p. 160. 

. 25. Hesperia alveus, Hiibn.; inner face of right clasp: p. 160. 

. 25a. Do.; lateral aspect of the tegumen. 

. 25 6. Hesperia onopordi, Ramb.; lateral aspect of the tegumen : p. 161. 
ig. 26. Hesperia speyeri, Frey. ; inner face of left clasp: p. 160. 

x. 27. Hesperia malvoides, n. sp.; dorsal aspect of the tegumen: p. 160. 
. 27a. Do.; inner face of left clasp. 


28. Hesperia malve, Linn.; dorsal aspect of the tegumen: p. 161. 


. 28a. Do.; inner face of left clasp. 

. 29. Hesperia melotis, Dup.; inner face of right clasp: p. 161. 

x, 30. Hesperia phlomidis, H.-S.; inner face of left clasp: p. 158. 

g. 31. Hesperia geron, Wats.; inner face of right clasp: p. 158. 

ge. 32. Thanaos marloyi, Bday. ; inner face of left clasp: p. 164. 

ge, 32a. Do.; inner face of right clasp. 

g. 33. Thanaos pelias, Leech ; inner face of left clasp: p. 164. 

ig. 33a. Do.; inner face of right clasp. 

eg. 34. Thanaos montanus, Brem.; dorsal aspect of the tegumen: p. 164. 
g. 35. Thanaos leechii, n. sp.; dorsal aspect of the tegumen: p. 164. 
ig. 86. Aeromachus inachus, Mén. ; inner face of left clasp: p. 190. 
g. 36a. Do.; dorsal aspect of the tegumen. 


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MESSRS, H. J. ELWES AND JAMES EDWARDS: 


. 37. Aeromachus jhora, de Nicéy.; inner face of left clasp: p. 190. 

ig. 37a. Do.; dorsal aspect of the tegumen. 

ig. 38. Aeromachus nanus, Leech ; inner face of left clasp: p. 192, 

g. 38a. Do.; dorsal aspect of the tegumen. 

ig. 39. Aeromachus indistincta, Moore; dorsal aspect of the tegumen: p. 191. 
ig. 39a. Aeromachus dubius, n. sp.; dorsal aspect of the tegumen: p. 190. 


PLATE XXIV. 


. 40. Pedestes masuriensis, Moore; inner face of left clasp: p. 193. 
ig. 40a. Do.; inner face of right clasp. 
g. 40 6. Do.; dorsal aspect of the tegumen. 
ge. 40 ¢. Do. ; edeagus with the guards detached. 
. 41. Pedestes pandita, de Nicév.; inner face of right clasp: p. 195. 
. 41a. Do.; dorsal aspect of the tegumen. 
eg. 416. Do.; cedeagus-guards. 
ig. 42. Pedestes maculicornis, n. sp.; dorsal aspect of the tegumen: p. 193. 
. 43. Pedestes fuscicornis, n. sp.; dorsal aspect of the tegumen: p. 194. 
. 44. Lophoides iapis, de Nicév.; inner face of left clasp: p. 196. 
. 44a. Do.; ventral aspect of the cedeagus. 
. 45. Lophoides purpurascens, nu. sp.; inner face of left clasp: p. 196. 
. 45a. Do.; ventral aspect of the cedeagus. 
. 46. Scobura fenestrata, n. sp.; inner face of left clasp: p. 206. 
. 47. Scobura inarime, de Nicéy.; inner face of left clasp: p. 206. 
ig. 48. Matapa purpurascens, n. sp.; inner face of right clasp: p. 209. 
. 49. Matapa druna, Moore ; inner face of right clasp: p. 210. 


50. Acerbas martini, Dist. ; inner face of left clasp: p. 216. 
51. Acerbas nitidifasciata, n. sp.; inner face of left clasp: p. 216. 


. 52. Plastingia callineura, Feld. ; inner face of left clasp: p. 226. 


55. Plastingia latoia, Hew. ; inner face of left clasp: p. 227. 

54. Plastingia margherita, Doh.; ventral aspect of the tegumen: p. 22 
54a. Do.; inner face of left clasp. 

55. Plastingia fruhstorferi, Mab.; ventral aspect of the tegumen: p. 227. 
55a, Do.; inner face of left clasp. 

56. Plastingia similis, n. sp. ; dorsal aspect of the tegumen: p. 230. 

56a. Do.; inner face of left clasp. 

57. Plastingia noemi, de Nicéy.; dorsal aspect of the tegumen: p. 230. 
d7 a. Do.; inner face of left clasp. 


. 58. Augiades subhyalina, Brem. ; ventral aspect of the edeagus: p. 247, 
. 59. Augiades sylvanoides, Leech; ventral aspect of the ceedeagus: p. 247. 
. 60. Augiades ochracea, Brem.; ventral aspect of the ceedeagus, with additional 


aspect of branch more enlarged: p. 248. 


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A REVISION OF THE ORIENTAL HESPERIID. 313 


PLATE XXY. 


ig. 61. Augiades majuscula, n. sp.; ventral aspect of the cedeagus: p. 248. 
ig. 61 a. Augiades crateis, Leech ; inner face of right clasp: p. 248. 

g. 616. Augiades brahma, Moore; inner face of right clasp: p. 248. 

. 62. Telicota augias, Linn. ; inner face of right clasp: p. 251. 

. 62a. Do.; dorsal aspect of the tegumen. 

. 63. Telicota bambuse, Moore ; inner face of right clasp: p. 251. 

g. 64. Telicota palmarum, Moore ; dorsal aspect of the tegumen : p. 252. 
. 64a. Do.; inner face of right clasp. 

. 65, Telicota augiades, Feld.; dorsal aspect of the tegumen: p. 253. 

. 65a. Do.; inner face of right clasp. 

. 66. Telicota gola, Moore; dorsal aspect of the tegumen: p. 255, 
. 66a. Do.; lateral aspect of the tegumen. 


66 6. Do.; inner face of left clasp. 


. 67. Telicota concinna, n. sp.; dorsal aspect of the tegumen: p. 255. 

. 67a. Do.; lateral aspect of the tegumen. 

. 67 6. Do.; inner face of left clasp. 

. 68. Lelicota rectifasciata, n. sp.; dorsal aspect of the tegumen: p. 254. 


68a. Do.; inner face of left clasp. 


. 69. Telicota dara, Koll. ; dorsal aspect of the tegumen: p. 254. 

. 69a. Do.; inner face of right clasp. 

. 70. Halpe sikkima, Moore ; ventral aspect of the tegumen: p. 264. 
. 71. Halpe homolea, Hew.; ventral aspect of the tegumen: p. 265. 
. 72. Halpe fusca, Elwes ; ventral aspect of the tegumen: p. 266. 

. 73. Parnara leechii, n. sp.; ner face of left clasp: p. 274. 

g. 75a. Do.; ventral aspect of the apex of the tegumen. 

g. 73 b. Do.; dorsal aspect of the apex of the tegumen. 

. 73. ¢. Do.; lateral aspect of the tegumen. 

. 74. Parnara oceia, Hew.; inner face of left clasp: p. 274. 

. 74a. Do.; ventral aspect of the apex of the tegumen. 

. 746. Do.; dorsal surface of the apex of the tegumen. 

g. 74. Do.; lateral aspect of the tegumen. 


PLATE XXVI. 


. 75. Parnara simillima, n. sp. ; inner face of left clasp: p. 274. 

. 75a. Do.; ventral aspect of the apex of the tegumen. 

. 756. Do.; dorsal aspect of the apex of the tegumen. 

.75¢. Do.; lateral aspect of the tegumen. 

. 76. Parnara guttatus, Brem.; Jateral aspect of the tegumen: p. 281. 
77. Parnara pellucida, Murr.; ventral aspect of the tegumen: p. 282. 
. 77a. Do.; lateral aspect of the tegumen. 


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. 78. Parnara contiqua, Mab. ; ventral aspect of the tegumen: p. 281. 
. 78a. Do.; lateral aspect of the tegumen. 
. 79. Parnara eltola, Hew. ; dorsal aspect of the tegumen : p. 282. 
ig. 79a. 1)o.; inner face of left clasp. 
. 796. Do.; ventral aspect of the cedeagus. 
g. 80. Parnara discreta, n. sp.; dorsal aspect of the tegumen: p. 282. 
. 80a. Do.; inner face of left clasp. 
. 80 6. Do.:; ventral aspect of the cedeagus. 
ig. 81. Parnara colaca, Moore; lateral aspect of the tegumen: p. 283. 
. 82. Parnara bevani, Moore; lateral aspect of the tegumen: p. 283. 
g. 83. Parnara subochracea, Moore ; ventral aspect of the apex of the tegumen : p. 275. 
. 84. Parnara mathias, Fab. ; ventral aspect of the apex of the tegumen: p. 275. 
. 85. Parnara philippina, H.-S. ; ventral aspect of the apex of the tegumen: p. 276. 
. 85a. Do.; dorsal aspect of the apex of the tegumen. 
. 85 6. Do.; lateral aspect of the tegumen. 
. 85 ce. Do.; inner face of left clasp. 
. 86. Parnara moolata, Moore; ventral aspect of the apex of the tegumen: p. 278. 
. 86a. Do.; dorsal aspect of the apex of the tegumen. 
. 866. Do.; lateral aspect of the tegumen. 
. 86¢. Do.; inner face of left clasp. 
. 87. Parnara aurociliata, n. sp.; ventral aspect of the tegumen : p. 278. 
. 87a. Do.; dorsal aspect of the tegumen. 
g. 87 6. Do.; lateral aspect of the tegumen. 

. 87 ¢. Do.; inner face of left clasp. 


PLATE XXVII. 


. 88. Parnara robusta, n. sp. ; ventral aspect of the tegumen: p. 280. 
. 88a. Do.; dorsal aspect of the apex of the tegumen. 
. §8 6. Do.; lateral aspect of the tegumen. 


88 ¢. Do.; inner face of left clasp. 


. 89. Parnara austeni, Moore ; ventral aspect of the apex of the tegumen: p. 280. 


89a. Do.; dorsal aspect of the apex of the tegumen. 
89 b. Do.; lateral aspect of the tezumen. 
89 ¢. Do.; inner face of left clasp. 


90. Parnara kumara, Moore; ventral aspect of the apex of the tegumen: p. 276. 


90a. Do.; dorsal aspect of the apex of the tegumen. 
906. Do.; lateral aspect of the tegumen. 


. 90 ¢. Do.; inner face of left clasp. 
g. 91. Parnara cahira, Moore ; ventral aspect of the apex of the tegumen: p. 278. 
. 91a. Do.; dorsal aspect of the apex of the tegumen. 
g. 916. Do.; lateral aspect of the tegumen. 

. 91¢. Do.; inner face of left clasp. 


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Fig. 92@. Do.; dorsal aspect of the apex of the tegumen. 
Fig. 926. Do.; lateral aspect of the tegumen. 


g. 92.c. Do.; inner face of left clasp. 

. 93. Hasora chromus, Cr.; dorsal aspect of the tegumen: p. 301. 

. 93a. Do.; inner face of left clasp. 

. 94. Hasora inermis, u. sp.; dorsal aspect of the tegumen: p. 301. 
. 94a. Do.; inner face of left clasp. 

ig. 95. Bibasis uniformis, n. sp.; inner face of left clasp: p. 305. 

ig. 96. Bibasis sambavana, n. sp.; inner face of left clasp: p. 305. 


INDEX. 


[Generic Names with initial capitals. Synonyms in italics. | 


Abaratha, 150. 

abax (Pamphila), 165, 166. 
abima (Gehenna), 244, 
Abraximorpha, 123. 

acalle (Hesperia), 253. 
Acerbas, 215, 

Achalarus, 108. 

acroleucus (Telegonus), 217. 
actzeon (Adopeea), 288, 289. 
Actinor, 245. 

aditus (Suastus), 179. 
Adopea, 288. 

adrastus (Hyarotis), 197. 
Aeromachus, 187. 


affinis (Celenorrhinus), 114, 121. 


affinis (Satarupa), 132, 134, 
agama (Pyrgus), 151. 

agna (Hesperia), 275. 

agni (Coladenia), 125, 127. 
agnioides (Coladenia), 125, 128. 
aina (Halpe), 259, 267. 
aitchisoni (Pithauria), 237, 238. 
akar (Pamphila), 254. 

albescens (Carystus), 180. 
albicilia (Sarangesa), 124. 
albicornis (Pirdana), 223, 224. 
albifascia (Plesioneura), 239. 
albinus (Suada), 177, 178. 
albipectus (Halpe), 260, 267. 


albistriga (Scelothrix), 162. 
albivitta (Pamphila), 268. 
albofascia (Gomalia), 153. 
alcese (Carcharodus), 152. 
alexandra (Erionota), 217. 
aleais (Parata), 301. 

ali (Hesperia), 154, 158. 
alica (Tagiades), 138, 140. 
alida (Caprona), 150, 151. 
aliena (Plesioneura), 231. 
alpina (Hesperia), 155, 160. 
althew (Carcharodus), 152, 
alveus (Hesperia), 156, 160. 
alysos (Plesioneura), 239. 
amara (Ismene), 292, 295. 


ambareesa (Celenorrhinus), 112, 115, 


Ampittia, 185. 
anadi (Ismene), 291, 294. 
Ancistroides, 222. 


andamanica (Celnorrhinus), 114, 119. 


andamanica, var. (Tagiades), 133. 
andromedz (Hesperia), 156, 161. 
angulata (Pterygospidea), 149. 
angulifera (Gehenna), 244, 


angustipennis, var. (Celenorrhinus), 113, 117. 


anthea (Acerbas), 215. 
antonia (Hesperia), 157, 163. 
anura (Hasora), 296, 298, 
Apostictopterus, 173, 


315 


. 92. Parnara conjuncta, H.-S.; ventral aspect of the apex of the tegumen: p. 280. 


316 MESSRS, H. J. ELWES AND JAMES EDWARDS: 


aquilina (Ismene), 292, 296. 
archias (Taractrocera), 183, 185. 
ardonia ('aractrocera), 184, 184. 
argyrostigma (Pamphila), 166, 167. 
aria (Matapa), 208, 209. 

armata (Kerana), 221. 

Arnetta, 198. 

asmara (Celenorrhinus), 113, 118. 
aspersa (Celeenorrhinus), 112, 115. 
assamensis (Parnara), 273, 281. 
Astictopterus, 171. 

astigmata (Adopza), 290, 
astigmata (Halpe), 258, 260. 
ataphus (Ismene), 291, 292. 

atila (Netrocoryne), 126. 

atkinsoni (Arnetta), 199. 

atticus (Tagiades), 139, 143. 
attina (Unkana), 234. 

aucma (Halpe), 265. 

Augiades, 246, 

augiades (Telicota), 250, 253. 
augias (Telicota), 250, 251. 
aurantiaca (Plastingia), 225, 228. 
auriferus (Zographetus), 201, 203. 
aurivittata (Celenorrhinus), 114, 122. 
aurociliata (Parnara), 272, 278. 
austeni (Parnara), 272, 280. 
avanti (Pamphila), 165, 167. 
avesta (Lotongus), 231, 233. 


bada (Hesperia), 281. 

Badamia, 306. 

badia (Celenorrhinus), 114, 123. 
badra (Hasora), 296, 298, 

balukinus (Celenorrhinus), 113, 117. 
bambusz (Telicota), 250, 251. 

Baoris, 270. 

Baracus, 169. 

barea (Hesperia), 269. 

basiflava (Notocrypta), 239, 241. 
batara (Ismene), 234. 

batchianus (Celenorrhinus), 114, 122. 
benjamini (Rhopalocampta), 306, 307. 
beturia (Halpe), 263. 

bevani (Parnara), 273, 283. 

bhagava (Satarupa), 131, 133. 
bhawani (Hidari), 236. 

Bibasis, 304. 


bicolor (Dejeania), 169. 

bicolor (Entheus), 136. 

bieti (Hesperia), 156, 162. 
bifasciatus (Achalarus), 108, 109. 
binotatus (Lophoides), 196. 
bipunctata (Parnara), 273, 283. 
bipunctata (Scobura), 205, 207. 
bipunctas (Suastus), 178, 180. 
biseriata (Sepa), 211, 213. 
bivitta (Halpe), 260, 268. 
blanchardi (Halpe), 256, 267. _ 
bononia (Scobura), 205, 206. 
borneensis (Hasora), 297, 302. 
bouddha (Augiades), 248. 
brahma (Augiades), 246, 248, 
brahmaputra, var. (Celenorrhinus), 117. 
brasidas (Tagiades), 141. 
bromus (Parnara), 272, 277. 
brontes (Papilio), 166. 

brunnea (Halpe), 259, 265. 
brunnea (Panara), 271, 275. 
buchananii (Coladenia), 129, 
butleri (Koruthaialos), 175. 


cacalize (Hesperia), 156, 161. 
cacus (Celenorrhinus), 118. 
czenis (Halpe), 262. 

cere (Chapra), 275. 

cxrulescens (Parnara), 271, 284. 
cahira (Parnara), 272, 278. 
cahira (Parnara), 278. 

cealathus (Lotongus), 231. 
Calliana, 106. 

calligana (Tagiades), 143. 
callineura (Plastingia), 225, 226. 
Caltoris, 270. 

cameroni (Celewnorrhinus), 114, 121. 
camertes (Cyclopides), 186. 
canaraica (Parnara), 277. 
Capila, 106. 

Caprona, 150. 

Carcharodus, 152. 
Carterocephalus, 165. 

carthami (Hesperia), 155, 160. 
cashmirensis (Hesperia), 155, 160. 
casyapa (Lobocla), 109. 
cataleucos (Suada), 177, 178. 
catena, var, (Erynnis), 287. 


A REVISION 


catocyanea (Acromachus), 189, 192. 
Celenorrhinus, 111. 

celzenus (Hasora), 297, 299. 
celebica (Satarupa), 131, 134. 
celebica, var. (Ismene), 298. 
celsina (Pirdana), 223, 224, 
centaures (Hesperia), 156, 161. 
cephala (Scobura), 204. 
cephaloides (Scobura), 204, 206. 
ceramas (Taractrocera), 183, 184. 
cerata (Halpe), 258, 260. 
cervantes (Thanaos), 163. 
ceylonica (Halpe), 258, 263. 
chabrona (Hasora), 297, 300. 
chamunda (Celsenorrhinus), 113, 117. 
Chapra, 270. 

Charmion, 110. 

chaya (Hesperia), 275. 

chilon (Suastus), 179. 

chinensis (Aeromachus), 187, 189. 
chinensis (Ctenoptilum), 148. 
Choaspes, 306. 

christophi (Pamphila), 166, 168. 
chromus (Hasora), 297, 301. 
chrysceglia (Proteides), 296. 
chrysomelena (Odina), 136. 
chrysozona (Hesperia), 252. 

chuza (Hasora), 298, 304. 
cicatrosa (Sepa), 211, 212. 

ciliata (Sepa), 211, 214. 

cingala (Parnara), 283. 
cinnamomea (Sepa), 211, 213. 
clavata (Plesioneura), 239. 

clitus (Celeenorrhinus), 115. 
cognata, var. (Satarupa), 134. 
colaca (Parnara), 273, 283. 
Coladenia, 125. 

comma (Erynnis), 287. 

concinna (Scobura), 205, 206. 
concinna (Telicota), 250, 253. 
conjuncta (Parnara), 273, 280. 
consanguinea (Celenorrhinus), 112, 115. 
consertus (Celenorrhinus), 118. 
consors, var. (Augiades), 248. 
contigua (Parnara), 273, 281. 
corissa (Plastingia), 226, 229. 
corona (Satarupa), 132, 134. 
coulteri (Hasora), 300. 


VoL. XIv.—Part 1v. No. 28.—October, 1897. 


OF THE ORIENTAL HESPERIIDZ. 317 


crateis (Augiades), 246, 248. 
crawfurdi (Rhopalocampta), 306, 307. 
Creteus, 233. 

cribrellum (Hesperia), 154, 158. 
cronus (Sepa), 211. 

Crossiura, 107. 

cruda (Goniloba), 234. 
Ctenoptilum, 147. 

cuneiformis (Odina), 135, 136. 
Cupitha, 245. 

curvifascia (Plesioneura), 239. 
cynare (Hesperia), 154, 158. 
cyrina (Creteus), 233. 


Daimio, 130. 

dan (Coladenia), 125, 127. 
danna (Taractrocera), 183, 184. 
dara (Telicota), 251, 254. 
Darpa, 137. 

dasahara (Sarangesa), 124. 
davidii (Abraximorpha), 123. 
dea, var. (Coladenia), 127. 
dealbata (Tagiades), 139, 145. 
debilis (Halpe), 259, 266. 
decorata (Halpe), 260, 268. 
decoratus (Odina), 135, 136. 
Dejeania, 169. 

delai-lama (Ampittia), 186, 187. 
delavayi (Syrichthus), 162. 
demea (Carterocephalus), 168. 
dentatus (Celenorrhinus), 114, 119. 
dhanada (Celenorrhinus), 114, 119. 
dieckmanni (Pamphila), 166, 168. 
dilutior (Telicota), 251, 255, 
dimila, var. (Erynnis), 287. 
diocles (Kerana), 221, 222. 

dire (Satarupa), 131, 133. 
discreta (Aeromachus), 188, 190. 
discreta (Parnara), 273, 282. 
dissimilis (Plesioneura), 214. 
distans (Tagiades), 141. 

distanti (Pirdana), 223, 224, 
distictus (Parnara), 286. 

diversa (Satarupa), 131, 132. 
divodasa (Hesperia), 179. 
doesoena (Hidari), 236. 

dohertyi (Satarupa), 132, 134. 
dolopia (Sebastonyma), 192: 


bo 
| 


318 MESSRS. H. J. ELWES AND JAMES EDWARDS. 


drancus (Plastingia), 229. 
dravida (Pumphila), 278. 
dravira (Pyrgus), 152. 

druna (Matapa), 208, 210. 
dschalia (Pamphila), 185. 
dubius (Aeromachus), 188, 190. 
dulcis (Aubertia), 168. 

durga (Zographetus), 201. 

duris (Acerbas), 215, 216. 


eacus (Hesperia), 127. 

Eetion, 236. 

electra (Choaspes), 307. 
elegans (Tagiades), 138, 141. 
elia (Hetion), 236. 

eltola (Parnara), 273, 282. 
elwesi (Caprona), B51. 

erebus (Nisoniades), 164. 
Erionota, 217. 

Erynnis, 287. 

etelka (Ismene), 291. 

evlepis (Hesperia), 229. 
eurotas (Pamphila), 251. 
evawidus (Pyrgus), 157. 
excellens (Lotongus), 231, 233. 
exclamationis (Badamia), 306. 


farri (Hesperia), 274. 

fasciata (Halpe), 258, 262. 

fatih (Hesperia), 127. 
feisthamelti (Notocrypta), 238, 239. 
feldert (Daimio), 135. 

fenestrata (Seobura), 205, 206. 
feralia (Scobura), 205, 206. 
fergusonii (Ismene), 291, 293. 
ficulnea (Charmion), 111. 

flava (Pamphila), 254. 

flavalum (Zographetus), 201, 202. 
Slavia, var. (Plastingi), 227. 
flavipennis (Zographetus), 202. 


flavocineta (Celenorrhinus), 112, 115. 


flayoides (Taractrocera), 183, 184. 


flavomaculatus (Pamphila), 166, 167. 


flexilis (Parnara), 284, 
florinda, var. (Erynnis), 287. 
folus (Udaspes), 243. 
fortunet (Hesperia), 281, 
frater (Achalarus), 109, 110. 


fruhstorferi (Plastingia), 225, 227. 
fulgur (Kerana), 221], 

fuliginosus (Apostictopterus), 173. 
fulvescens (Celzenorrhinus), 114, 120. 
fumosa (Satarupa), 131, 133. 

fusca (Halpe), 259, 266. 

fusca (Plesioneura), 117. 

fuscicornis (Pedestes), 193, 194. 
fuscula (Sancus), 174. 


galba (Hesperia), 154, 157. 
gana (Tagiades), 138, 141. 
Gangara, 217. 

Ge, 182. 

Gegenes, 286. 

Gehenna, 244. 

Gehlota, 111. 

gemmatus (Carterocephalus), 168. 
gemmifer (Kerana), 221. 
gener (Eudamus), 109. 
gentiana (Parata), 304. 
germanus (Achalarus), 109, 110. 
geron (Hesperia), 154, 158. 
geta (Ge), 182. 

gigas (Hesperia), 155, 159. 
glandulosa (Paduka), 220. 
glauca (Pamphila), 238. 
gneus (Hasora), 296, 298. 
gola (Telicota), 250, 253. 
goloides (Padraona), 253. 
Gomalia, 153. 

gomata (Ismene), 292, 295. 
gopala (Satarupa), 131, 132. 
goto (Plesioneura), 118. 
gree (Gehenna), 244, 245. 
grandis (Erionota), 217. 
graya (Tagiades), 133. 
gremius (Suastus), 178, 179. 
gupta (Halpe), 259, 266. 
guttatus (Parnara), 272, 281. 
guttulifera (Sepa), 211, 212. 


hadria (Hasora), 298. 

Halpe, 257. 

hamiltonii (Coladenia), 128. 
hampsoni (Baracus), 170, 171. 
hampsoni (Tapena), 146, 147. 
hamza (Adopea), 289. 


A REVISION OF THE ORIENTAL HESPERUDZ. 


hanria (Darpa), 137. 

Hantana, 110. 

hasoroides (Parnara), 271, 284. 
hector (Koruthaialos), 175. 
helena (Plastingia), 225, 227. 
helferi (Tagiades), 138. 

helias (Odontoptilum), 148, 149. 
helisa, var. (Odontoptilum), 149. 
hellas (Hesperia), 157. 

henrici (Astictopterus), 172. 
herculea (Pamphila), 247. 
Hesperia, 153. 

heterus (Pamphila), 254. 
Heteropterus, 168. 

Hidari, 235, 

hieroglyphica (Odina), 135, 136. 
hieron (Halpe), 265. 

hiraca (Hesperia), 217. 

homolea (Halpe), 259, 265. 
honorei (Halpe), 260, 268. 
houangty (Pamphila), 165, 167. 
Hyarotis, 197. 

Hyda, 123. 

hyela (Pirdana), 223. 

hyperides (Odontoptilum), 149. 
hypoleucos (Pyrgus), 161. 

hyrax (Adopia), 288, 289. 
hyrcana (Hesperia), 247. 

hyrie (Halpe), 258. 


iadera (Itys), 200. 

Tambrix, 180. 

iapis (Lophoides), 195, 196. 
Idmon, 198. 

igna (Coladenia), 125, 128. 
inachus (Aeromachus), 189, 190. 


inzequalis (Celenorrhinus), 114, 119. 


inarime (Scobura), 205, 206. 
indistincta (Aeromachus), 188, 191. 
jndrani (Coladenia), 125, 126. 
indrasana (Isoteinon), 229. 
inermis (Hasora), 297, 301. 
infernus (Hantana), 110. 
jnornata (Notocrypta), 239, 241. 
insignis (Halpe), 258, 261. 
insularis (Telicota), 250, 252. 
jionis (Ismene), 291, 295, 

irava (Hidari), 235, 


Isma, 204. 
Ismene, 290. 
isota (Isma), 205. 
Isoteinon, 197. 
Iton, 269, 

Itys, =00. 


jaina (Ismene), 291, 293, 

jama (Astictopterus), 172, 
jenkowskii (Ismene), 296. 

jansonis (Parnara), 273, 282. 
Japetus (Tagiades), 138, 141. 
javana (Gegenes), 280. 

Javanicus (Aeromachus), 188, 191. 
jayadeva (Capila), 107, 

jhora (Aeromachus), 181, 190. 
jolandu (Hesperia), 284. 


kada (Astictopterus), 172. 
kali (Aeromachus), 189, 192. 
karea (Tagiades), 138, 141. 
karsana (Hesperia), 286. 
kerala (Koruthaialos), 176. 
Kerana, 220. 

kethra (Astictopterus), 174. 
khasiana (Tagiades), 138, 140, 
khasianus (Isoteinon), 199, 
knyyetti (Halpe), 2538, 261. 
kophene (Koruthaialos), 176, 
Koruthaialos, 176. 

kuehni (Telicota), 250, 252. 
kumara (Halpe), 258, 261. 
kumara (Parnara), 272, 276. 


ladana (Celzenorrhinus), 114, 122. 
ladon (Papilio), 306. 

lenas (Carystus), 228. 

lalita (Orthophetus), 105. 
lamprospilus (Isoteinon), 197. 
lara (Ismene), 292, 295. 

lura (Telegonus), 217. 

latifascia (Iambrix), 181, 182, 
lativittus (Celenorrhinus), 114, 121. 
latoia (Plastingia), 225, 227. 
latonia (Plastingia), 229. 

latreille: (Hesperia), 234, 

latreilles (Tagiades), 142. 

latris (Halpe), 266, 


320 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


lavata (Tagiades), 138, 142. 
layaterse (Carcharodus), 152. 
laxmi (Coladenia), 125, 126. 
lebadea (Paduka), 219. 

leechii (Parnara), 271, 274. 
leonina (Adopxa), 288, 290. 
leptogramma (Odontoptilum), 148, 150. 
leucoeera (Celienorrhinus), 113, 116. 
leucocirca (Plesioneura), 116. 
leucographa (Plesioneura), 111. 
leuzew (Hesperia), 159. 
liburnia (Plastingia), 226, 228. 
lidderdali (Orthopheetus), 105, 
liliana (Achalarus), 108, 109. 
linea (Papilio), 288. 

lineola (Adopxa), 288, 289. 
litigiosa (Tagiades), 143. 
littoralis (Gomalia), 153. 
Lobocla, 108. 

Lophoides, 195. 

lorquint (Ismene), 295. 
Lotongus, 230. 

lucasii (Halpe), 262. 

lucifera (Celenorrhinus), 115. 
luteisquama (Hesperilla), 264. 
luzonensis (Pamphila), 184. 
lycorias (Pamphila), 245. 

lyde (Taractrocera), 187. 


mabillei (Carystus), 216. 

mabillei (Unkana), 235. 

maculatus (Hesperia), 157, 162. 
maculatus (Lotongus), 231. 
maculicornis (Celenorrhinus), 113, 116. 
maculicornis (Pedestes), 193. 
maculosa (Celenorrhinus), 112, 115. 
meniata (Coladenia), 130. 

musa (Pamphila), 254. 

maesoiles (Anpittia), 184. 

meesoides (Pamphila), 254. 
mestissima (Hasora), 297, 300. 
mevius (Taractrocera), 183. 

maga (Ampittia), 186. 

mahintha (Ismene), 291, 295. 
majuscula (Augiades), 246, 249. 
majuscula (Halpe), 259, 264. 
malayana (Ismene), 301, 


malye (Hesperia), 156, 161. 
mandan (Hesperia), 166. 
mangala (Pamphila), 281. 
margherita (Plustingia), 225, 227. 
marloyi (Thanaos), 163, 164. 
marnas (Telicota), 256. 

maro (Ampittia), 186. 
maroides (Ampittia), 186. 
marta (Halpe), 265. 

martini (Acerbas), 215, 216. 
martini (Scobura), 204, 205. 
martinus (Tagiades), 139, 143. 
masoni (Halpe), 260, 268. 
masuriensis (Pedestes), 193. 
Matapa, 208. 

mathias (Parnara), 271, 275. 
meetana (Tagiades), 140. 
meiktila (Onryza), 269. 
meleagrina (Pamphila), 234. 
melotis (Hesperia), 156, 161. 
menaka (Tagiades), 139, 142. 
mencia (Pamphila), 276. 

micio (Pamphila), 166, 168. 
microstictum (Itys), 200. 
microthyrus (Crane), 242. 
migreus (Suastus), 178, 179. 
mindorana (Plesioneura), 242. 
minuscula (Tapena), 146, 147. 
minuta (Suastus), 180. 
miosticta (Sepa), 211, 212. 
modestu (Isoteinon), 199. 
mohammed (Syrichthus), 159. 
mélleri (Suastus), 177. 
montanus (Thanaos), 163, 164. 
monteithi (Notocrypta), 239, 241. 
moolata (Parnara), 272, 278. 
moorei (Halpe), 258, 263. 
moori (Pterygospidea), 135. 
mormo (Pamphila), 274. 
morpheus (Heteropterus), 168. 
multiguttata (Ctenoptilum), 148. 
munda (Plesioneura), 116. 
murdava (Pithauria), 237. 

mus (Hasora), 297, 304. 
musca (Aeromachus), 188, 189. 
myra (Hasora), 297, 303. 
mytheca (Lotongus), 231, 233. 


A REVISION OF THE ORIENTAL HESPERIID, 


naga (Plastingia), 226, 229, | 
nana (Tagiades), 139, 144. | 
nanus (Aeromachus), 189, 192. 
narada (Satarupa), 131, 132. 

narooa (Hesperia), 280. 

nascens (Parnara), 271, 276, 

nezra ((Hrane), 242. 

nephele (Halpe), 258, 264. 

nepos (Achalarus), 109, 110. 

neryulata (Adopwa), 288, 290. 

nestus (Tagiades), 139, 145. 

nicevillei (Taractrocera), 183, 184. 

nigrescens, var. (I'hanaos), 164, 

nigricans (Celzenorrhinus), 113, 118. 

nigrolimbatus (Thymelicus), 185. 

nilgiriana (Isoteinon), 199. 

niphates (Tagiades), 154. 

nitida (Pamphila), 254. 
nitidifasciata (Acerbas), 215, 216. 
niyeomaculatus (Pamphila), 166, 167. 
nobilis (Hesperia), 155, 159. 

noctis (Sepa), 211, 214. 

noctis (Tagiades), 140. 

noémi (Plastingia), 226, 230. 

noma (Hesperia), 159. 

nondou (Hesperia), 281. 

nostrodamus (Gegenes), 286. 
Notocrypta, 238. 

nymphalis (Satarupa), 131, 132. 


oberthiiri (Hesperia), 157, 162. 
oberthiiri (Taractrocera), 183, 185. 
obliquans (Astictopterus), 182. 
obscura (Lophoides), 195, 196. 
obscurus (Tagiades), 138, 141. 
oceia (Parnara), 271, 274. 
ochracea (Augiades), 246, 248. 
Ocytes, 287. 

Odina, 135. 

Odontoptilum, 148. 

cedipodea (Ismene), 291, 292. 
cedipus (Ismene), 291, 292. 
(rane, 242. 

ogygia (Zographetus), 201, 203. 
ogygioides (Zographetus), 201, 203. 
olivascens (Astictopterus), 172. 


omeia (Orthopheetus), 105, 106. 


321 


onchisa (Caltoris), 278, 250. 
onopordi (Hesperia), 156, 161. 
Onryza, 268, 

ops (Carterocephalus), 167. 
orbifer (Hesperia), 154, 158. 
orbiferus (Celznorrhinus), 113, 118. 
ormenes (Halpe), 257, 260. 
ornata (Halpe), 268, - 

ornatus (Heteropterus), 168, 169. 
orphitus (Telicota), 251, 256. 
Orthophcetus, 104, 

ortygia (Odina), 137. 

othonias (Ancistroides), 222. 


Padraona, 249. 

Paduka, 219. 

pagana (Parnara), 272; 277. 
paleemon (Pamphila), 165, 166. 
palajava (Plesioneura), 118. 
palawana, yar. (Ismene), 305. 
palawata, var. (Plastingia), 229. 
palawea (Halpe), 264. 
palmarum (Telicota), 250, 252. 
Pamphila, 165. 

pandia (Hesperia), 218. 
pandita (Pedestes), 193. 
paniscus (Papilio), 166. - 
paragola (Telicota), 250, 254, 
paralysos (Notocrypta), 239, 241. 
Parata, 296. 

parca (Parnara), 234. 

Parnara, 270. 

parthenope (Hesperia), 231. 
patula (Celenorrhinus), 116. 
pavona (Pirdana), 224, 

pavor (Padraona), 256. - 
Pedestes, 192. . 
pelias (Thanaos), 163, 164. 
Pelion, 288. 

pellucida (Parnara), 273, 282. 
penicillata (Baoris), 274. 
pennicillatum (Crossiura), 108. 
perara (Halpe), 265. 

perfusca (Pamphila), 214. 
permena (Pterygospidea), 134. 
pero (Celxenorrhinus), 112, 115. 
phanzus (Orthopheetus), 105. 


322 MESSRS. H. J. ELWES AND JAMES EDWARDS: 


phiditia (Suastus), 180. 
philenus (Telicota), 256. 
philetas (Ismene), 299. 
philippina (Parnara), 272, 276. 
Philoodus, 286. 

philotas (Parnara), 285. 
phisara (Satarupa), 131, 134. 
phlomidis (Hesperia), 154, 158. 
phenicis (Hesperia), 197. 
pholus (Pudicitia), 216, 

piceus (Aeromachus), 188, 189. 
pieridoides (Calliana), 106. 
pinwilli (Tagiades), 140, 145. 
Pirdana, 223. 

Pithauria, 237. 

Pithauriopsis, 237. 

plagifera (Celenorrhinus), 116. 
Plastingia, 224. 

plebeia (Parnara), 271, 274- 
Plesionewra, 238. 

plesionewree (Plastingia 2), 113. 
plumbeolus (Baraeus), 170, 171. 
pluscula (Celenorrhinus), 116. 
poggei (Hesperia), 155, 159. 
popoviane (TRaraos), 163- 
preba (Plesioneura), 197. 
pralaya (Tagiades), 140, 145- 
princeps (Tagiades), 139, 145_ 
prominens (Chapra), 275. 
proteus (Pyrgus), 159. 

" proto (Hesperia), 155, 159. 
proximata (Hasoza), 297, 301. 
proximus (Achalarus), 108, 109. 
proxissima (Hasora), 297, 302. 
prusias (Telicota), 250, 251. 
pseudomesa (Padvaona), 254. 
pteria (Tagiades), 139, 142. 
Pteroxys, 104. 

Pterygospidea, 137. 

Pudicitia, 216. 

pugnans (Parnara), 272, 284. 
pulchra (Pamphila), 165, 167. 
pulligo (Sancus). 174. 

pulomaya (Celencrrhinus), 112, 115, 
purendra (Sarangesa), 123, 124. 
purpurascens (Lophoides), 196. 
purpurascens (Matapa), 208, 209, 


purreea (Cupitha), 245. 

putra (Plesionewra), 116. 

pygela (Odontoptilum), 148, 149, 
pygmeus (Papilio), 286. 

Pyrgus, 153. 

pyrrha (Celenorrhinus), 116. 
pythias (Pamphila), 251. 


quadrata (Notocrypta), 239, 241. 
quadripunctata (Ismene), 298. 


radians (Actinor), 244. 
ransonnettii (Caprona), 150. 

ravi (Tagiades), 138, 140. 
rectifascia, var. (Notocrypta), 239. 
rectifasciata (Telicota), 251, 254. 
renidens (Khopalocampta), 307. 
restricta (Plesioneura), 239. 
Rhopalocampta, 306. 

rickuchina (Pamphila), 248. 
robsonii (Suastus), 180. 

robusta (Parnara), 272, 280, 
rudolphii (Pirdana), 223. 
ruficornis (Plesioneura), 117. 
rusticanus (Thanaos), 164, 


sagara (Pamphila), 183. 

saida (Hasora), 298, 304. 

sala (Suastus), 178, 179. 
salsala (Iambrix), 181. 
sambara (Satarupa), 132, 134. 
sambavana (Bibasis), 305, 
sambayana (Tagiades), 139, 143. 
Sancus, 173. 

sanguinocculus (Gangara), 218. 
sao (Hesperia), 154, 158. 
Sape, 123. 

sarala (Lotongus), 231, 233. 
Sarangesa, 123. 

saraya (Caprona), 150, 151. 
sasivarna (Matapa), 208, 210, 
Satarupa, 130. 

sati (Sarangesa), 123, 124. 
saturatus (Celenorrhinus), 114, 120, 
satwa (Zographetus), 201. 
Scelothrix, 153. 

Scobura, 204, 


A REVISION OF THE ORIENTAL HESPERIIDA. 


scopas (Suada), 177, 178. 
scopulifera (Baoris), 274. 

scortea (Pamphila), 281. 
Sebastonyma, 192. 

selas (Pamphila), 247. 
semamora (Iton), 269. 

semperi (Coladenia), 125, 128. 
sena (Bibasis), 305. 

Sepa, 210. 

separata (Halpe), 259, 267. 
septentrionis (Ismene), 292, 294. 
septentrionum (Baracus), 170, 171. 
seriata (Hesperia), 276. 

sericea (Hesperia), 164. 
serratule (Hesperia), 156, 160. 
shalgrama (Matapa), 208, 210. 
side (Hesperia), 157, 162. 
signata (Plesioneura), 111. 
sikkima (Baoris), 274. 

sikkima (Halpe), 259, 264. 
silyius (Pamphila), 165, 167. 
similis (Augiades), 247. 

similis (Pamphila), 275. 

similis (Plastingia), 226, 230. 
simillima (Parnara), 271, 274. 
simplex (Achalarus), 108, 109. 
simplex (Telicota), 250, 253. 
simplicissima (Hasora), 297, 299. 
sindu (Iambrix), 181, 182. 
sinensis (Parnara), 271, 275. 
singularis (Carystus), 241. 
sinica (Satarupa), 132, 135. 
sinicus (Pyrgus), 162. 

sinina, var. (Nisoniades), 163. 
sitala (Halpe), 259, 266. 

siva (Augiades), 246, 247. 
sobrina (Coladenia), 125, 126. 
sodalis (Pamphila), 275. 

speyeri (Hesperia), 156, 160. 
Spilothyrus, 152. 

spilothyrus (Celenorrhinus), 113, 117. 
staudingeri (Hesperia), 155, 159. 
staudingeri (Hidari), 235, £36, 
stellata (Udaspes), 243, 

stellifer (Iambrix), 181. 
Steropes, 165. 

steropes (Papilio), 168. 


| 


stigma (Adopza), 288, 289. 
stigmata (Aeromachus), 187, 189. 
stramineipennis (Pithauria), 237. 
striata (Ismene), 294. 

Suada, 177. 

Suastus, 178. 

subcaudata (Rhopalocampta), 307. 
subditus (Baracus), 170, 171. 
subfasciatus (Astictopterus), 174. 
subfasciatus (Ismene), 219. 
subflava (Halpe), 259, 267. 
subgrisca (Hesperia), 179. 
subhyalina (Augiades), 246, 247. 
submacula (Halpe), 258, 263. 
submaculata (Plastingia), 230. 
subochracea (Parnara), 271, 275. 
subradiatus (Cyclopides), 185. 
subtesiaceus (Isoteinon), 199. 
subvittatus (Ochus), 185. 
sulphurifera (Halpe), 259, 264. 
sumitra (Celenorrhinus), 113, 115. 
sunias (Pamphila), 255. 

superna (Pyrgus), 157. 

sura (Odontoptilum), 148, 149, 
surus (Proteides), 231. 

swerga (Suada), 177. 

swinhoet (Carcharodus), 152. 
swinhoei (Watsonia), 220. 
sybirita (Gangara), 218. 

sybirita (Hidari), 235, 236. 
sylvanoides (Augiades), 246, 247. 
sylvanus (Augiades), 246, 247. 
sylvatica (Adopma), 288, 289. 
Syrichthus, 153. 

syrichthus (Caprona), 150, 151. 


tabrica (Tagiades), 140, 146. 
“tages (Thanaos), 163. 
Tagiades, 137. 

tagiadoides (Curystus), 215. 
tamiata (Pamphila), 233. 
Tapena, 146. 

Taractrocera, 182. 

taras (Hesperia), 161. 
tavilus (Pamphila), 254. 
telesinus (Plastingia), 226, 228. 
Telicota, 249. 


323 


524 A REVISION OF THE ORIENTAL HESPERIIDA, 


teliga (Halpe), 263. 

tenebrosa (Adopzea), 288, 289. 
tesscllata (Plastingia), 226, 229. 
tessellum (Hesperia), 155, 159. 
tethys (Satarupa), 132, 135. 
Thanaos, 163. 

thaumas (Adopza), 288. 
therapne (Hesperia), 154, 158. 
thibetanus (Hesperia), 157, 162. 
thrax (Erionota), 217. 

thrax (Hesperia), 275. 

thwaitesi (Tapena), 146. 
thymbron (Ismene), 306. 

thyone (Parnara), 282. 

thyrsis (Gangara), 218. 

tibetana (Celenorrhinus), 113, 119. 
tibetana, var. (Pamphila), 247. 
tissa (Coladenia), 125, 126. 

titus (Tagiades), 139, 142. 

toba (Tagiades), 144. 

tola (Charmion), 111. 

toona (Hesperia), 281. 

trachala (Pamphila), 254. 
translucida (Capila), 107. 
trichoneura (Tagiades), 140, 145. 
trichoneuroides (Tagiades), 140, 145. 
trimacula (Ampittia), 186, 187. 
tripura (Suastus), 179, 180. 
tuckeri (Ismene), 291, 293. 

tulsi (Parnara.), 271, 284. 
tympanifera (Cupitha), 245. 


Udaspes, 243, 

ulunda (Astictopterus), 174. 

uma (Parnara), 285. 

umbrosa (Scobura), 205, 207. 
unicolor (Baoris), 274, 

unicolor (Heteropterus), 168, 169. 
unicolor (Idmon), 198. 

unicolor (Parnara), 255. 
uniformis (Bibasis), 305. 

Unkana, 254. 


varia (Halpe), 259, 266. 

yasava (Ctenoptilum), 147, 148. 
vasutana (Ismene), 291, 294. 
venata (Hesperia), 247. 

venula (Papilio), 288. 
yermiculata (Plastingia), 225, 227. 
verones (Koruthaialos), 176. 
viburnia (Plastingia), 226, 229. 
vindhiana (Arnetta), 199. 
violacea (Hasora), 297, 299. 
virgata (Ampittia), 186. 
virgula (Papilio), 289. 

yitrea (Coladenia), 129. 

vitrea (Pamphila), 198. 

vitta (Hasora), 300. 

vittatus (Baracus), 170. 

volua (Plesioneura), 239. 

yulso (Lophoides), 196. 


wantona (Halpe), 265. 
waterstradti (Tagiades), 139, 143. 
Watsonia, 220. 

watsonii (Iton), 269. 


xanites (Koruthaialos), 175. 


zalates (Proteides), 231. 
Zampa, 230. 

zawi (Plesioneura), 111. 

Zea, 230. 

zebra (Pamphila), 254. 

zebra (Pyrgus), 157. 

Zela, 230. 

zelleri (Parnara), 273, 284. 
zema (Halpe), 257, 260. 
zennara (Capila), 107. 

zenon (Lotongus), 231, 232. 
zetus (Pamphila), 206. 

zeus (Lotongus), 231, 232. 
ziclea (Taractrocera), 183, 184. 
Zographetus, 200. " 
zona (Hesperia), 157, 162. 


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CONTENTS.. 


VI. A Revision of the Oriental Hesperiide. By H. J. Ewes, F.R.S., F.LS., F-.Z.S., 
and James Epwarps, F.E.S. (Plates XVIII-XXVIL) . . . . page 101 


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ges26nu 


VII. On the Morphology of the Skull in the, Paraguayan Lepidosiren and in other 
Dipnoids. By Professor T. W. Briven, Sc.D., F.Z.S., Mason College, Birmingham. 


Received May 10, 1897, read June 1, 1897. 


[Puates XXVIII. & XXIX.] 


ContTEnts. Page 

eelntroduchonie yy-lis itis PRE iit CADIPICROACNG Hi tos OLAS 325 

II. Description of the Skull of the Paraguayan Lepidosiren .............. 327 
III. Revision and Comparison of the Structure of the Skull in Ceratodus, 

IAM MAR EMO SG TENSIKAD. Loogsasoouepos so cuctun sb50000050 350 

IV. The Structure of the Skull in Fossil Dipnoi.....................-.. 366 

Y. References to previous Literature ...........505..222000-s erences 373 

Vile Explanationxofiiey bl ates mernctrtesetsases stair nacre yells ate eret- Tee 375 


I. Introduction. 


THE skull which forms the subject of the following description belonged to one of 
the numerous specimens of Lepidosiren collected by the German traveller Dr. Bohls 
in the region of the Upper Paraguay River, and brought to Kurope in 1894. As to 
the specific identity of the Paraguayan specimens with the Amazonian Lepidosiren 
paradoza, it is at present premature to express a decided opinion, but on the evidence 
so far available I see no reason to question the probability that the two are referable 
to the same species. In any case, as Lankester [20] has shown, the distinction which 
Ehlers [8] has attempted to draw between his Paraguayan species L. articulata and 
L. paradoxa, as regards the segmentation or non-segmentation of the axial cartilage 
of the pectoral and pelvic fins, seems to have no foundation in fact. 

In order to facilitate the comparison of my Paraguayan specimen with others 
obtained from the same region, and with the Lepidosiren paradoxa of the Amazons, 
the following measurements were made, and are here expressed in terms of the ratios 
adopted by Lankester [20, p. 15] :— 


Total length in | Head-length | Ratio of total Ratio of inter- Ratio of post- Ratio of cervico- 
centimetres. unit, length. membral length. pelvic length. dorsal length. 


75 1 4:51 3:17 27 


VoL. x1v.— Part Vv. No. 1.—February, 1898. ZU 


326 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


It may be remarked that the ratios exhibit a general agreement with those of the 
larger of the two specimens in the Oxford University Museum [Lankester, /. c. p. 15], 
except that their relative values are smaller, a fact due to the greater value of the 
unit of measurement, that is, the head-length, in the former as compared with the 
latter specimen. For the rest it may be said that my specimen had the usual black 
colour of the Paraguayan specimens, and in the character of its skin-areas and 
denticulated scales closely resembled the specimen described and figured by Lankester. 
The cloaca was situated to the left of the medio-ventral line, and both the pectoral 
and pelvic fins were supported by an axial skeleton in the form of an obviously- 
segmented rod of cartilage. Exclusive of the cranial rib, the specimen had 54 pairs of 
ribs, or one pair fewer than the specimens described by Bischoff and Hyrtl. Lastly, 
it may be added that the specimen was a male, with the characteristic series of 
villous processes along the postero-medial margins of its pelvic fins. 

The first detailed account of the structure of the skull of Lepidosiren paradora, and 
certainly the most important, is that given by Bischoff in his well-known and copiously- 
illustrated memoir [2] published in 1840. This was followed a few years later (1845) 
by Hyrtl’s monograph [18], which, however, contained but a single figure of the skull, 
viz. a lateral view. Hyrtl corrected one or two errors in Bischoff’s memoir, and 
slightly modified the nomenclature of certain of the cranial elements, but otherwise 
added little to our knowledge of the structure of the skull; indeed, as he himself says, 
«Die Schiidelknochen wurden von Bischoff so vollstindig abgehandelt, dass ich mich 
hier nur in eine Aufzihlung, nicht in eine Beschreibung derselben einzulassen brauche ” 
(1. c. p. 613). Briihl [4] two years later (1847) reproduced several of Bischoff’s 
figures, and suggested certain modifications in the names of some of the cranial bones, 
but in other respects his description is obviously based on the work of his distinguished 
predecessor, 

The figures given in Bischoff’s memoir, although for the most part accurate, 
nevertheless justify the criticism that neither the shape nor the sutural boundaries 
of the various bones are always represented with sufficient clearness, and are further 
defective in that sufficient attention has not been given to the structure of the 
chondrocranium. In several points the text also needs revision, more particularly in 
the light of modern researches in cranial morphology. 

Of other Dipnoids, the cranial anatomy of Protopterus has perhaps been the more 
fully treated. The first account by Owen [25] was very imperfect, but was revised and 
certainly improved in his ‘ Comparative Anatomy and Physiology of Vertebrates ’ [26]. 
Peters [34], Huxley [14], and Cobbold [6] have also contributed to our knowledge of 
the skull of this Dipnoid. It is to Wiedersheim [41], however, that we are indebted 
for the best account of the skull of Protopterus, and his admirable paper has the 
additional merit of being illustrated by excellent and accurate figures. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 327 


The skull of the remaining Dipnoid, Ceratodus, was first described and figured by 
Giinther [11], whose account was subsequently revised, and also amplified in certain 
particulars, in an eminently suggestive paper by Huxley [15]. 

The object of the present communication is (a) to place in the hands of morpho- 
logists the results of a detailed investigation into the structure of the skull of the 
Paraguayan Lepidosiren, illustrated by accurate and carefully-drawn figures; (6) to 
revise the accounts of the skulls of Protopterus and Ceratodus given by preceding 
writers; and (¢) to institute a more detailed comparison of the differences and 
resemblances between the various Dipnoid skulls than has yet been made. 

I desire also to express my thanks to the Council of the Royal Society for a grant 
from the Research Fund in aid of this and other investigations. 


II. Description of the Skull of the Paraguayan Lepidosiren. 


In somewhat striking contrast to the relatively short and bluntly-conical shape of 
the head, the skull, when stripped of its investing skin and powerful muscles, appears 
relatively longer and more sharply conical in contour. The narrowest portion of the 
skull is the central region, or that part which lies directly behind the eyes, expanding 
anteriorly in the nasal region, and widéning even more behind in the auditory and 
laterally-deflected suspensorial regions. Viewed laterally the skull presents a strikingly 
carnivorous appearance, largely due to the existence of a prominent sagittal crest along” 
the medio-dorsal line of the fronto-parietal bone, and to the backwardly-projecting 
“lambdoid” margin in which that bone terminates dorsad to the occipital plane. 

Of the various cranial bones the fronto-parietal (the parieto-frontal of Bischoff and 
Hyrtl; in Protopterus, the parietal of Owen, Peters, and Cobbold, and the “ Fronto- 
parietale ” of Wiedersheim) (Pl. XXVIII. figs. 1,2, and 4; Pl. XXIX. figs. 13-19 fp.), 
trom the share which it takes in the formation of the roof and side-walls of the cranial 
cavity, is perhaps the most important. In shape each lateral half of the bone is 
somewhat triangular (fig. 1), the broad bases of the two halves meeting in the medio- 
dorsal line, and there forming a strong longitudinal sagittal crest or ridge (figs. 1, 2, 
and 4, sg.c.) which serves for the origin of the more superficial portion of the temporal 
muscle. In no other Fish with which I am acquainted are the temporal muscles so 
powerfully developed in proportion to the size of the skull as in Lepidosiren, and in 
none do these muscles extend so far on to the dorsal surface of the skull. It is to 
the exceptional development of the temporal and masseter muscles that the bluntly- 
conical shape of the head is mainly due, since they are principally responsible for the 
thickening of what, so far as the skull alone is concerned, would otherwise be almost 
the narrowest portion of the head. From the sagittal crest the lateral portions of 


the fronto-parietal extend downward and outward, and form a gable-roof for all that 
20 2 


528 PROF, T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


portion of the skull which lies posterior to the nasal capsules, varying, however, in 
their relations to the cranial cavity and to the chondrocranium in different regions. 
Thus, in the anterior part of their extent (figs. 1, 4, and 15), the lateral portions of 
the bone, after forming the roof of the cranial cavity, curve sharply downward to the 
inner side of the trabecular cartilage, and by it are separated from the palato-pterygoid 
bone (fig. 15), eventually terminating on each side in an obliquely-fissured and some- 
what irregular inferior margin (fig. 4), without, however, quite meeting the cartila- 
ginous plate which, in front of the parasphenoid, forms the cranial floor. More 
posteriorly the fronto-parietal, as it descends from the roof into the side-walls of the 
cranial cavity, meets on each side the upturned lateral margin of the parasphenoid 
(figs. 4, 16, and 17) in a broadly V-shaped suture, which is best seen in a vertical 
longitudinal section of the skull (fig. 4), but is hidden externally by the trabecular 
cartilage (fig. 1, ¢7.c.). More posteriorly still (figs. 17 and 18) each lateral portion of 
the bone is continued outward from the cranial roof, externally to the cartilaginous 
auditory capsule, and thence is prolonged downward and outward in close relation 
with the outer surface of the suspensorial cartilage, extending almost as far as the 
articular surface for the mandible ; eventually the bone tapers to a contracted apex, 
which is wedged in between the hinder part of the palato-pterygoid bone and the 
squamosal, articulating with the superior margin of the former and the anterior borde 

of the latter (fig. 1). Behind the auditory capsule (figs. 1 and 19) the fronto-parietal 
overlies the cartilage of the supraoccipital region, and its lateral margins, converging 
from below upward and backward, project beyond the chondrocranium, and, after 
overlapping the two exoccipital bones and the first neural arch, terminate dorsally in a 
backwardly-projecting conical process (figs. 1 and 2). The converging lateral margins 
of the bone simulate the appearance of the characteristic “lambdoid” crest of the 
carnivorous Mammalia (fig. 2, /d.c.), and serve for the insertion of a portion of the 
lateral musculature of the trunk. From what has been stated as to the extent and 
relations of the fronto-parietal bone, it is obvious that it forms not merely the roof 
but also the lateral walls of all that section of the cranial cavity which lies between 
the mesethmoid region anteriorly and the auditory capsule posteriorly, and further, 
that in addition to strengthening the cranial roof in the auditory and post-auditory 
regions of the skull, it also contributes to the rigidity of the articular condyle for the 
lower jaw by investing the outer surface of the suspensorial cartilage. 

Anteriorly to the fronto-parietal, and resting on the cartilaginous internasal septum, 
and also extending laterally so as to partially invest the fenestrated dorsal walls of the 
olfactory capsules, is a relatively thick and somewhat triangular bone (Pl. XXVIII. 
figs. 1, 2, and 4; Pl, X XIX. figs. 11 and 12, d.e.), The broad hinder margin of the 
bone is connected by a tough fibrous tissue, and not by suture, with the anterior edge 
of the fronto-parietal (fig. 4), and from this point the bone gradually contracts to a 
blunt apex which terminates a little posterior to the laterally-diverging cornua 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 329 


trabecule and the base of the prenasal process (fig. 2)1.. The bone has been termed 
“premaxilla” by Natterer [24], Bischoff, and Hyrtl, apparently from its relations to 
what these writers considered to be premaxillary teeth, and it is obvious that it is the 
homologue of an almost precisely similar bone which in Protopterus is the conjoined 
premaxilla and nasal of Owen [26], the ethmoid of Peters and Rose [36], and the 
nasal of Cobbold, Miall [22], Wiedersheim, and Huxley [14]. In Ceratodus the 
equivalent bone has been regarded by both Giinther and Huxley as an “ ethmoid.” 
That the bone is not a “ premaxilla” is proved by the fact that the latter element is 
invariably developed in front of the most anterior portion of the chondrocranium, 
which is certainly not the case with the bone in question, while the so-called “ pre- 
maxillary teeth ” are without doubt the representatives of the vomerine teeth of other 
Fishes and of Amphibia. The possibility that the bone may represent a pair of con- 
joined nasals cannot be so easily rejected, although, so far as I am aware, the fusion 
of two such elements to form a median nasal bone is without precedent in any other 
Fishes. For the term “ ethmoid” much more may be said, but in the application of 
this name a distinction must be drawn between the characteristic mesethmoid of 
Teleosts, which is always an ossification of the mesethmoid cartilage, and the “ supra- 
ethmoid” (Parker), which is a dermal bone situated directly beneath the superficial 
skin, and altogether external to the cartilage. From its position, external to the 
cartilage and immediately beneath the skin, it may be concluded that the Dipnoid 
bone is not a mesethmoid element, while it is obvious that it is in every way the exact 
counterpart of the bone which in some Teleosts (e. g. Salmo) has been termed “ supra- 
ethmoid” by Parker [27], and in such Fishes exists in conjunction with ordinary paired 
nasals. For these reasons it seems preferable to regard the Dipnoid bone as a “ dermal 
ethmoid.” It may be mentioned that a similar median bone, with essentially similar 
relations to the nasal region of the skull, exists in Polypterus and in many fossil 
Fishes, such as, for example, the Arthrodira (e. g. Coccosteus) and the Paleoniscide. 
Two singular bones, which for the present will be referred to as ‘“ supraorbital ” 
elements, take origin from near the anterior margin of the fronto-parietal, and thence 
arch upward and backward nearly to the hinder end of the skull, lying immediately 
beneath the external skin and dorsad to the fronto-parietal, from which they are 
widely separated by the great temporal and masseter muscles (Pl. XXVIII. figs. 1, 2, 
and 4, ec.e.). At its anterior end (fig. 2) each bone is expanded laterally so that the 
two are only slightly separated from each other, while the outer margin forms the 
dorsal boundary of the orbit. At this point also (fig. 1) each is horizontally forked 
in such a way as to clip a backwardly-projecting process derived from the contiguous 


1 The transverse suture described and figured by Bischoff (1. c. tab. iii. fig. 4) as extending across this bone, 
and the curiously angular relations of the two portions, are obviously the result of an accidental fracture, as 
Hyrtl (J. c.) pointed out. 


530 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


portion of the palato-pterygoid bone (figs. 1 and 2, p.pt.a'.), and to this process, as 
well as to the fibrous connection between the dermal ethmoid and the fronto-parietal, 
the anterior portion of the bone is firmly connected by ligamentous fibres. The 
extremity of the outer division of the fork approaches closely to the posterior margin 
of the cartilaginous olfactory capsule, but does not in any way contribute to the 
formation of a hinder wall to the capsule. Posteriorly, the bones contract to the 
condition of slender, laterally-compressed rods, and at the same time become widely 
separated from each other, eventually terminating behind in pointed extremities. 

The identification of these bones, which are present in all existing Dipnoi, and have, 
on the whole, similar relations to other cranial structures as already described for 
Lepidosiren, has given rise to much divergence of opinion. Bischoff (/. c.) termed 
them “jugal” bones (‘ Jochbeine?”), although it is evident that he entertained some 
doubt as to the accuracy of his interpretation. By Hyrtl (J. ¢.) they were named super- 
ciliary bones (“ Superciliarknochen ”). In Protopterus the homologous structures have 
been termed “ Frontale, incl. Frontale post” (Peters, J. ¢.), “ frontals” (Cobbold, J. ¢.), 
‘“super-temporal ” (Owen, J. ¢.), “ supraorbital ” (Huxley, /. ¢.), ‘*tendon-bones” 
(*‘Sehnenbeine,” Wiedersheim, J. ¢.), and in Ceratodus “ os frontale ” (Giinther, J. ¢.), 
‘¢ frontal,” Miall (/. c.), and “inner lateral bones,” but probably representing the 
frontals of Polypterus (Huxley, J. ¢.). The term “jugal” is obviously inappropriate, 
as Cobbold pointed out, and “superciliary” or “supraorbital ” are purely descriptive 
names based on the relation of the anterior portion of each bone to the orbital cavity. 
It is also clear that the bones in question have nothing in common with the bones 
ordinarily termed “ frontals” in other Fishes, inasmuch as, except at their anterior 
extremities, they are widely separated from the proper cranial roof by the whole thick- 
ness powerful jaw-muscles, and lie directly beneath the dermis and scales of the super- 
ficial skin. On the other hand, the term ‘‘ Seknenbein,” applied by Wiedersheim to 
the essentially similar bone in Protopterus, is somewhat novel, and on this point his 
own words may be quoted. After referring to the position of the two bones dorsad 
to the proper cranial roof, Wiedersheim says:—‘ Am frischen Praparat ist der 
ganze Zwischenraum von dem zum Processus coronoideus mandibule ziehenden 
M. temporalis ausgefiillt, und mit der gewaltigen Entwickelung des genannten Kau- 
muskels bringe ich auch die Entstehung der beiden sonderbaren Knochenlamellen 
in Verbindung, d. h. ich halte sie fiir in Folge des Muskelzuges entstandene Ossifica- 
tionszonen in der Fascia temporalis resp. in der fast den ganzen Kopf einhiillenden 
subcutaneus Fascie tiberhaupt. Mit dem bis jetzt dafiir gebrauchten Namen ‘ Supra- 
orbitalknochen’ ist nichts erklart, ja der Name is schon deswegen nicht passend, weil 
sie sich weit tiber das Gebiet der Augenhdhle hinaus erstrecken” (/.¢. pp. 46-47). 
According to Wiedersheim, therefore, the bones are to be regarded as ossifications in 
the superficial temporal fasciz of the head, and presumably on that account are not to 
be compared with any of the ordinary cranial bones of other Fishes. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 331 


The origin of the temporal fasciee and their relations to the bones, muscles, and 
other structures of the head in Lepidosiren may be briefly described. The superficial 
stratum of each of the great dorso-lateral trunk-muscles passes into a thin fibrous 
fascia which, after extending over the temporo-masseter muscles, blends anteriorly 
with a fibrous ring encircling the orbit, and is also attached to the posterior curvature 
of the antorbital cartilage, finally becoming continuous in front with the fibrous 
investment of the nasal capsules. Inferiorly, the fascia is continuous with the tough 
fibrous tissue investing the outer surface of the hinder portion of the mandible. 
Dorsally, the fascia becomes continuous with the external margin of the supraorbital 
bone of its side, and, reappearing on the inner margin, becomes, in turn, continuous 
with the inner margin of the corresponding bone of the opposite side of the 
skull 

From the relations of the two supraorbital bones to the temporal fasciz it would 
seem at first sight that the suggestion of Wiedersheim in the case of Protopterus is 
equally applicable to Lepidosiren. When, however, the position and relations of the 
equivalent bones in Ceratodus are carefully examined, it is not quite so clear that the 
name “ Sehnenbein ” is wholly accurate in either instance. 

For the hinder part of its extent each supraorbital bone in the latter Dipnoid is a 
thin but relatively much wider plate than in either Protopterus or Lepidosiren, and 
becomes almost fibrous at its irregular free posterior margin, where it is continuous with 
the equivalent of the more posterior portion of the temporal fascia. In this region the 
bone lies externally to the jaw-muscles, but, unlike its representative in other Dipnoi, 
is connected with its fellow through the intervention of a similarly-situated bony lamina 
which is the “ postero-median ” bone of Huxley [15, fig. 7 B], and the “ scleroparietal ” 
of Giinther [11, pl. xxxiv. fig. 4,a]. More anteriorly, as it passes over the orbit, the 
bone thickens considerably, but nevertheless so far retains the position and relations of 
the supraorbital bone of Lepidosiren and Protopterus. Instead, however, of terminating 
in the orbital region, the bone extends forward, closely investing the dorsal wall of the 
hinder part of the cartilaginous nasal capsule, and at the same time suturally articulates 
internally with the outer margin of the dermal ethmoid (Pl. XXIX. fig. 20 ec.e.). 
The preorbital portion of the bone forms moreover a characteristic descending 
process which, passing downward between the orbit and the nasal capsule, becomes 
greatly thickened laterally 1, and also closely applied to the lateral wall of the chondro- 
cranium in the region of the mesethmoid. Finally, the process terminates inferiorly 
in a sutural articulation with the dorsal edge of an ascending lamina derived from the 
dentigerous palatine portion of the palato-pterygoid bone (fig. 20, ppt.a.). It is 
obvious, therefore, that the preorbital section of the bone forms not only a partial 
roof, but in addition a posterior wall to the olfactory capsule, and at the same time 


1 The centre of this part of the bone contains a cavity filled with fatty connective tissue. 


332 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


constitutes the anterior boundary of the orbit. Comparison of the preorbital portion 
of the supraorbital bone of Ceratodus with the bone which in certain Amphibia has 
been termed “ external ectethmoid” by Parker [30, p. 172; also 28] and “ prefrontal” 
by Wiedersheim [42, 43] proves that the two are essentially similar in position and in 
their relations to contiguous cranial structures. Although absent in the Perenni- 
branchiate Urodela, an ‘external ectethmoid” in varying degrees of development 
is present in all Caducibranchiata (Derotremata) (e.g. Amphiuma, Cryptobranchus, 
Menopoma), and generally also in both the Mecodont (e.g. Salamandra, Triton) and 
Lechriodont (e.g. Ellipsoglossa, Ranodon, Amblystoma, Desmognathus, Spelerpes) 
sections of the Salamandrida (Myctodera). In all the Urodela in which it is present, 
the “ ectethmoid ” or “ prefrontal” is a membrane-bone in relation with the antero- 
external extremity of the frontal of its side, partially investing the hinder part of 
the roof of the nasal capsule, and also, by its extension downward towards the palatine 
region, forming the boundary between the orbital and nasal regions of the skull}. 
In several instances the bone extends to a varying distance backward over the orbit, 
the roof of which it helps to form. ‘The similarity of the two bones in all essentials 
is so striking that it is difficult to avoid the conclusion that the “ supraorbital” of 
Ceratodus and the “external ectethmoid” or “ prefrontal” of the Urodele Amphibia 
are homologous structures. In Protopterus and Lepidosiren the relations of the 
so-called “supraorbital” bones are admittedly somewhat different. In these genera 
each bone forms a roof to the orbital cavity, and even extends slightly downward in 
front of it, but certainly not so far as to enable it to form a posterior wall, and still 
less a partial roof, to the nasal capsule. On the contrary, the bones seem to have been 
displaced dorsally and backward, as it were, by the upward and backward growth 
of an ascending process from the palato-pterygoid. Nevertheless, it is scarcely possible 
to doubt that the “supraorbital” bones are homologous cranial elements in all three 
genera and also with the Amphibian ectethmoid, although it may be admitted that 
in Protopterus and Lepidosiren the homology is less obvious at first sight, owing to the 
secondary displacement of the bones by the ascending process of the palato-pterygoids, 
and might not even have occurred to any one but for the fact that Ceratodus, in retaining 
the primitive relations of these bones, affords the necessary clue to their real nature. 
There is one feature, however, in which the Dipnoid “ external ectethmoid ” differs 
from its Amphibian representative, and that is the extension of the bone backward to 
the occipital region of the skull, but for this difference I am inclined to think thatjthe 
exceptional development of the Temporalis muscle is wholly responsible. In Lepidosiren 
this muscle begins posteriorly as a cranial extension of the deeper stratum of the 
dorso-lateral trunk-musculature. On the lateral surface of the hinder part of the skull 


‘ A similar bone (“prefrontal”) is also present in the Labyrinthodont Amphibia [Stegocephala], and in 
Reptilia. 


IN THE PARAGUAYAN LEPIDOSIREN, ETO. 333 


it becomes greatly thickened by the addition of fibres having their origin from the outer 
surface of the fronto-parietal bone, including its sagittal crest; from the median vertical 
fibrous sheet which continues the crest as far externally as the superficial temporal fascia, 
and aloue in this region separates the temporal muscles of opposite sides of the head ; and 
also from the outer margin and ventral surface of the hinder two-thirds, or postorbital 
portion, of the external ectethmoid. From this extensive origin the Temporalis runs 
obliquely downward and forward, anteriorly and externally to the much less 
developed Masseter muscle, to its insertion into the coronoid process of the mandible. 
These facts suggest that the unusual backward extension of the external ectethmoid 
is due to the necessity of providing an additional surface for the origin of an excep- 
tionally developed muscle. It is not only possible, but even probable, that the hinder 
or postorbital section of each ectethmoid owes its existence to the ossification of the 
temporal fascia investing this muscle, and to this extent it is possible to agree with 
Wiedersheim that the bone is a ‘“Sehnenbein.” Nevertheless, and for the reasons 
mentioned above, it may be maintained that the orbital and preorbital portions of the 
bone represent a true “ external ectethmoid ” element. 

The identification of the “supraorbital” bones with the Amphibian “ external 
ectethmoids” removes almost the only difficulty in the way of the comparison of the 
cranial bones of the Dipnoi with those of other Vertebrata. Parker’s term “ external 
ectethmoid ” is not, perhaps, the most suitable name for the bones under discussion ; 
dermal ectethmoids, or, in order to distinguish these paired bones from the median 
dermal ethmoid and the true mesethmoid bone, dermal lateral ethmoids, are much 
more convenient terms. 

Underlying the base of the cranium, and extending backward from a point at some 
distance behind the symphysis of the palato-pterygoid bones to the level of the first 
neural arch, is the well-developed parasphenoid (“ Keilbein ” of Bischoff; in Protopterus 
the “ Basilare” of Peters, “‘sphenoid” of Cobbold, “‘ basioccipito-sphenoid ” of Owen, 
and the “ parasphenoid” of Huxley and Wiedersheim ; in Ceratodus the ‘ basal” bone 
of Ginther, and ‘“‘ parasphenoid ” of Huxley), (PJ. XXVIII. figs. 3 and 4; Pl. XXIX. 
figs. 16-19, ps.). Viewed from below (fig. 3) the bone appears somewhat spatulate 
in shape, the anterior half widening considerably, aud having its lateral margins 
deflected and at the same time closely applied to the inner surfaces of the palato- 
pterygoid bones, more particularly where the latter bones curve downward to form the 
articular condyles for the lower jaw. Anteriorly, the bone terminates at some distance 
behind the palato-pterygoid symphysis, not, however, as in Protopterus, by an abruptly- 
truncated transverse margin, but by a contracted and somewhat conical extremity. 
From the auditory region backward the bone gradually narrows, and by lateral 
compression becomes greatly thickened, eventually terminating in an almost pointed 
extremity ventrad to the basal cartilage of the occipital region. The share taken by 
the parasphenoid in conjunction with the fronto-parietal in forming the side-walls 

Vou. xIv.— Part v. No. 2.—February, 1898. 2X 


334 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


of the central portion of the cranial cavity has already been mentioned (see fig. 4). 
Reference to transverse sections of the skull (figs. 16 and 17) will show that in 
this region each lateral margin of the parasphenoid divides into an ascending lamina 
which meets the descending portion of the fronto-parietal internally to the trabecular 
cartilage, and a deflected lamina which, as mentioned above, overlaps the inner surface 
of the palato-pterygoid bone of its side. These sections also prove that in the same 
region the parasphenoid alone forms the floor of the cranial cavity, the cartilaginous 
basis cranii being here entirely absent. Perhaps the two most remarkable features 
in connection with the parasphenoid of Lepidosiren are the share which, with the 
fronto-parietal, it takes in forming the lateral walls of the cranial cavity, and the 
somewhat abrupt termination of the bone anteriorly, so that a considerable area of the 
cartilaginous basis cranii is freely exposed between its anterior margin and the palato- 
pterygoid symphysis. 

The bone termed “ palatine” by Bischoff and Hyrtl in Lepidosiren—in Protopterus 
considered as equivalent to the maxilla, pterygoid and palatine (Owen), or termed 
“ Gaumenkieferbein ” (Peters), “palatine” (Cobbold), ‘“ Pterygo-palatinum” or 
“ palato-pterygoid ” (Wiedersheim and Huxley); and in Ceratodus “ pterygo-palatine ” 
(Giinther, Huxley)—is one of the most characteristic elements in the Dipnoid skull 
(Pl. XXVIII. figs. 1-4; Pl. XXTX. figs. 12-17, p.pt.). The bone seems to represent 
a pterygoid element ossified continuously with a “ tooth-bone ” formed by the fusion of 
the strongly-developed palatal teeth, and may therefore rightly be regarded as a 
palato-pterygoid. 

Posteriorly the bone makes its appearance at a puimt immediately anterior to the 
foramen for the exit of the hyomandibular division ot the Facial nerve in the form of 
a relatively wide thin plate, closely applied to the inner surface of the suspensorial 
cartilage, and overlapped along its inner and dorsal margin by the deflected lateral 
edge of the parasphenoid (fig. 3). ‘Traced downward towards the articular extremity 
of the suspensorial cartilage the bone thickens somewhat, and forms the inner margin 
of the articular surface for the lower jaw (figs. 3 and 4). From this point the palato- 
pterygoid loses its lamellar character and becomes an exceptionally strong, laterally- 
compressed bone, arching upward and forward towards the nasal capsule. In the latter 
part of its extent it is in relation by its dorsal margin with the band-like trabecular 
cartilage (fig. 1), and by it is separated from the sutural union of the descending lamina 
of the fronto-parietal and the parasphenoid (fig. 16). In front of the anterior termi- 
nation of the latter bone the palato-pterygoid skirts the lateral margin of the cartila- 
ginous basis cranii (fig. 15). On reaching the hinder boundary of the nasal capsule, 
each palato-pterygoid begins to curve obliquely inward and forward, and is finally united 
with its fellow in an elongated and exceptionally massive symphysis which is interposed 
between the two nasal capsules, and dorsally is in contact with their roof in the median 
plane (figs. 5, 4, and 12). The extraordinary development of the symphysial portions 


» 


N 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 335 


of the two bones is doubtlessly associated with their function as basal supports for the 


powerful palatal teeth, but the effect produced on the nasal region of the skull is 
remarkable, and will be referred to subsequently. As described by Wiedersheim in 
Protopterus, the symphysial extremity of each bone is produced into three processes 
which, when viewed ventrally (fig. 3), appear to radiate outward from the median line 
of the symphysis. Of these processes, the hindermost and strongest is nearly transverse, 
or at most has but a slight backward inclination, and, projecting outward, terminates in 
a pointed extremity (figs. 1-3 and 13, at. p.). This is the equivalent of the ‘* Processus 
antorbitalis ” of Wiedersheim, and, in conjunction with the body of the bone, supports 
ventrally the basal portion of the posterior and largest of the three plate-like palatine 
teeth (fig. 3, p.p.t.). The central process is transversely disposed and supports the 
much smaller central tooth (m.p.t.), while the third is directed obliquely outward and 
forward, and forms the basis of the anterior and smallest of the palatal teeth (a.p.t.). 
In addition to the three tooth-supporting processes, the anterior portion of each palato- 
pterygoid gives off from its dorsal border a stout conical process, corresponding to 
the ‘Processus ascendens ” of Wiedersheim, which is directed obliquely upward and 
backward, and terminates in a pointed extremity projecting a little beyond the level 
of the cranial roof (figs. 1 and 5, p.pt.a.). The process extends so far backward 
as to overlap the anterior margin of the fronto-parietal (fig. 1), and fill up what would 
otherwise be an oblique notch or fissure between the lateral, or descending, and the 
dorsal portions of that bone, and therefore, as seen in figs. 4 and 13, contributes to 
the formation of the outer wall of the extreme anterior section of the cranial cavity 
and of the olfactory fossa of its side. Dorsally, the two processes are separated from each 
other by the anterior extremity -of the fronto-parietal (figs. 2 and 5), and each is over- 
lapped, and partially hidden from view, by the expanded anterior section of the dermal 
ectethmoid. On the outer surface of each process there is a strong lateral ridge, 
coincident in direction with the inclination of the process itself, but terminating above 
in a free projecting extremity, which is clipped by the bifurcate outer margin of 
the anterior extremity of the ectethmoid (figs. 1, 2, 5, and 13, p.pt.a’.). 

The squamosal bone (‘Quadrate,” Bischoff and Hyrtl; in Protopterus, the “ Quadrate,” 
Peters, “ tympanic,” Owen, ‘‘ zygomatic” or ‘“ jugal,” Cobbold, “ squamosal,” Huxley and 
Wiedersheim ; in Ceratodus, the “ squamosal” of Giinther and Huxley) invests the outer 
surface of the hinder part of the suspensorial cartilage (Pl. XXVIII. figs. land 2; 
Pl. XXIX. figs. 17 and 18, sg.). For the dorsal two-thirds of its extent it is a relatively 


thin plate, somewhat ovate in shape, its posterior margin coinciding with the hinder 


edge of the suspensorial cartilage (fig. 1). Inferiorly, the bone is constricted to a 


neck-like portion, but, again expanding, terminates in a rounded margin, which forms 
the outer surface of the articular condyle for the lower jaw. For a portion of its 
extent the anterior border of the bone overlaps the ventral termination of that portion of 


the fronto-parietal which extends on to the outer surface of the suspensorium (fig. 1). 
2x2 


336 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


The chondrocranium of Lepidosiren presents several interesting features, and for 
convenience in description may be divided into (1) an occipito-periotic region, including 
also the suspensorium ; (2) a central trabecular portion ; and (3) an anterior or ethmo- 
nasal region. 

The occipito-periotic region consists of a thick basal cartilage underlying the foramen 
magnum and extending in the floor of the cranial cavity as far forward as the middle 
of the periotic capsule, where it terminates somewhat abruptly on the cranial surface 
of the parasphenoid by a well-defined transverse margin, immediately behind the 
position occupied by the pituitary body (Pl. XXVIII. fig. 4, and Pl. XXIX. fig. 19). 
A slight concavity on the upper surface of the parasphenoid is the only indication of a 
pituitary fossa (Pl. XXIX. fig. 18). Into the axis of the basal cartilage (fig. 4) 
the filiform intracranial portion of the notochord (no.) is prolonged, and may be 
traced in sections nearly as far forward as the cartilage itself extends. On its ventral 
surface the basal cartilage is in close relation with the hinder part of the parasphenoid, 
while behind, where it becomes continuous with the chordal sheath, the cartilage helps 
to support dorsally and laterally the bases of a pair of ossified “ basi-dorsals ” forming 
the lateral elements of the first neural arch (Pl. XXVIII. figs. 1 and 4, na.). 
Anteriorly to the “basi-dorsals” are the two exoccipital bones (figs. 1 and 4; 
Pl. XXIX. fig. 19, e0.), each of which consists of an inwardly-curved basal plate 
resting on the upper surface of the basal cartilage, but separated mesially from its 
fellow by an intervening tract of the same cartilage. Each bone then curves upward 
(fig. 4), forming the side-walls of the hindermost section of the cranial cavity, 
and terminates dorsally in a thick plate of cartilage which occupies the supraoccipital 
region, beneath the hinder part of the fronto-parietal bone, and forms the actual roof 
of the cranial cavity in this region (fig. 19), In a transverse section of the cranium 
taken through the two exoccipitals (fig. 19), it is seen that the dorsal cartilage extends 
downward for some distance on each side into the substance of the exoccipital, 
dividing the latter for a portion of its extent into an outer lamina of bone, which 
dorsally abuts against the outer margin of the fronto-parietal, and an inner lamina 
ending above in the dorsal cartilage. ach occipital (fig. 4) is deeply constricted in 
the centre, owing to the presence of a deep notch in its anterior and posterior margins, 
the anterior notch forming the posterior boundary of the foramen for the Vagus nerve 
(x.), the posterior ivansmitting the roots of the Hypoglossal or first spinal nerve 
(sp.n'). 

The basal cartilage and the cartilage of the supraoccipital region, though otherwise 
distinct, become continuous in front of the foramina for the Vagus nerves with the 
laterally-bulging cartilage of the periotic capsule, and through the latter with the 
proximal portion of the suspensorial cartilage (fig. 4). 

Externally and dorsally (fig. 1), the periotic capsules are completely hidden from 
view by the lateral extension of the fronto-parietal, except for a narrow tract on each 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 337 


side immediately above the upper extremity of the squamosal. Ina ventral view, 
however, each capsule is well seen between the foramina for the Facialis and Vagus 
nerves, the rounded prominence which it presents in this region corresponding to the 
outer wall of the recess for the sacculus (fig. 3) 

In a vertical longitudinal section of the skull (Pl. XXVIII. figs. 4 and 6) the cavity 
of the periotic capsule appears as a relatively spacious lateral diverticulum of the 
cranial cavity, the inner wall of the capsule, which in Elasmobranchs and Amphibians 
separates the two cavities, having completely atrophied in Lepidosiren, as in Ceratodus 
and Protopterus, and also in all existing Ganoids and Teleosts. The cavity of the 
capsule exhibits a division into two deep recesses, of which the more external and 
dorsal lodges the ‘‘ pars superior” (utriculus) (fig. 6, wt.r.) of the auditory organ, 
while the more internal and inferior contains the “ pars inferior” or sacculus. The 
numerous foramina perforating the periotic capsule will be referred to subsequently. 

The suspensorium consists of a somewhat triangular lamina of cartilage, the broad 
base of which is continuous dorsally with the outer wall of the auditory capsule and 
with the hinder portion of the trabecular cartilage, and thence is continued obliquely 
downward and forward to the articular condyle for the lower jaw (Pl. XXVIII. 
fig. 5, and Pl. XXIX. fig. 18, a7.c.). So complete is the investment of its external 
surface by the fronto-parietal and squamosal bones that only a small portion of the 
cartilage is visible just above the mandibular articulation (Pl. XXVIII. fig. 1). 
Ventrally, however, the suspensorial cartilage (Pl. XXVIII. fig. 3) may be seen 
anteriorly and externally to the foramen for the exit of the hyomandibular branch of 
the Facial nerve, but even here the cartilage is extensively invested by the hinder part 
of the palato-pterygoid. The posterior margin of the suspensorial cartilage is deeply 
emarginate (fig. 5), so that in a ventral view the inner surface of the squamosal is 
partially exposed, and in the same view a rounded prominence near the hinder margin 
of the cartilage indicates the point of attachment of the hyoid arch. A charac- 
teristic foramen termed by Hyrtl (/. c.) the “ Schlafengrube ” perforates the suspensorial 
cartilage at its junction with the periotic capsule (vl. XXVIII. figs. 1 and 2, ¢.f.; 
Pl. XXIX. fig. 18), and in a ventral view is visible near the base of the skull (fig. 3). 

The suspensorial cartilage of Lepidosiren must be regarded as the metapterygo- 
quadrate or proximal portion of the mandibular arch, the equivalent of the palato- 
pterygoid cartilage of the more typically autostylic skulls of Chimera and Ceratodus, 
having undergone complete atrophy or become replaced by the palato-pterygoid bone. 
It is worthy of note that the suspensorium makes a much more open angle with the 
fore part of the basicranial axis than in Protopterus or in Ceratodus, and the effect of 
this on the curvature of the palato-pterygoid bone is such that, in passing from the 
articular end of the suspensorium to the nasal region, the bone describes a segment of 
a larger circle in the former Dipnoid than in either of its congeners. 

The pulley-like articular condyle for the mandible (Pl. XXVIII. figs. 1, 2, 5 and 4, 


338 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


ar.c.) is concave from side to side and convex from before backward. It is formed in 
the centre by the distal extremity of the suspensorial cartilage, strengthened on its 
inner surface by the hinder part of the palato-pterygoid bone, and externally by the 
inferior extremity of the squamosal. No trace of a true quadrate ossification could be 
detected. 

The trabecular region of the chondrocranium may be considered to begin as an 
extension forward of the cartilage of the auditory capsules in the form of two parallel 
cartilaginous rods or plates, situated one on each side of the anterior half of the cranial 
cavity. At its origin from the periotic capsule each trabecular rod is also continuous 
with the anterior portion of the suspensorial cartilage, and forms a relatively thick plate 
of cartilage, invested externally by the lateral extension of the fronto-parietal, and 
separated from the cranial cavity by the ascending lamina of the parasphenoid, the 
descending plate of the same bone closely investing its inferior margin (Pl. XXIX. 
fig. 17, t7.c.). More anteriorly the rod contracts, and, owing to the diminished lateral 
growth of the fronto-parietal, now becomes visible in a side view of the skull, 
externally and parallel to the line of junction of the latter bone with the parasphenoid 
(Pl. XXVIII. fig. 1, and Pl. XXIX. fig. 16, #.c.)1. More anteriorly still the 
trabecular rods blend inferiorly with the cartilaginous lamina which, in front of the 
termination of the parasphenoid, alone forms the basis cranii, and, at the same time 
increasing somewhat in vertical extent, become overlapped internally by the lateral 
portions of the fronto-parietal, and partially also externally by the palato-pterygoids 
(Pl. X XIX. fig. 15). On approaching the nasal capsules, each trabecular cartilage 
detaches itself from the basis cranii, and, still further contracting, assumes the condition 
of a slender laterally-compressed rod, which curves downward across the outer surface 
of the fronto-parietal and palato-pterygoid bones (Pl. XXVIII. fig. 1, and Pl. XXIX. 
figs. 13 and 14, an.p.), and then passes ventrally between the antorbital process of 
the last-mentioned bone and the hinder wall of the nasal capsule. The cartilage now 
becomes very slender, and, after giving off a short anteriorly-directed process (fig. 1) 
below and parallel to the outer margin of the nasal capsule, enters the posterior 
margin of the upper labial fold of its side, and thence describes a bold curve back- 
ward beneath the eye. Kventually the rod curves downward and slightly forward, 


* A similar reduction of the central or interorbital region of the chondrocranium to the quasi-primitive 
condition of two laterally-situated cartilaginous trabecular rods occurs also in seyeral Urodele Amphibia, such 
as, for example, Menobranchus (Huxley, 17, and Wiedersheim, 43) and Amphiuma (Wiedersheim, 1.c.). The 
resemblance of the latter to Lepidosiren is also heightened by the fact that in both cases the cranial roof-bones 
suturally articulate on each side with the parasphenoid to form the lateral walls of the cranial cavity. 
Comparison of transverse sections through the cranium of Lepidosiren (Pl. XXIX. figs. 16 and 17) with 
similar sections of, for example, the skull of Menobranchus (Wiedersheim, figs. 40 and 42) will sufficiently 
illustrate this point. The occurrence of similar parallel modifications in the Ophidian skull is sufficient to 
prove that, however striking may be the structural resemblances between the Amphibian and Dipnoid skulls 
in these respects, they are nevertheless homoplastic in their nature. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 339 


and finally terminates in the inferior margin of the labial fold (Pl. XXVIII. figs. 1-4, 
an.p'). Although regarded by both Bischoff (/. c.) and Hyrtl (/.c¢.) as pertaining to 
the system of labial cartilages, there can, I think, be little doubt that, from the point 
of its separation from the cartilaginous basis cranii, each of these singular rods 
represents an “antorbital process,”1 almost identical in its mode of origin and 
relations with a homologous cartilaginous process which exists in many Elasmobranchs 
(the ‘“‘seitlicher Vorsatz der Ethmoidal-Region” of Gegenbaur, 10, or “antorbital 
or ethmo-palatine process” of Parker, 29), and more especially in the Urodele 
Amphibia (“antorbital process ” of Wiedersheim, 43, or “ ethmo-palatine cartilage” of 
Parker, 28, 30). Ordinarily in both Elasmobranchs and Urodela the process is 
either directed outward at right angles to the axis of the skull, or inclined slightly 
backward, but it is interesting to note that in some Urodela (e.g. Menopoma and 
Siredon) the antorbital process is directed forward as in Lepidosiren, parallel to the 
outer margin of the nasal capsule, with which it may even fuse anteriorly (Menopoma), 
or, as in Siredon, remain distinct (Wiedersheim, 43). 

The singular development of the antorbital process in Zepidosiren is evidently 
associated with its function as a skeletal support for the posterior portion of the 
overlapping and somewhat pendulous upper labial fold of this Dipnoid. It may 
be mentioned that besides its skeletal support the labial fold is provided with 
special muscles, either as derivatives from the anterior portion of the contiguous 
temporal muscle, or arising independently from the temporal fascia (Hyrtl, /.c.), and 
also with an abundant nerve-supply by the ramus buccalis of the Facialis nerve. 

After the separation and divergence of the two antorbital cartilages, the cartilaginous 
basis cranii becomes pushed upward, as it were, by the palato-pterygoid symphysis, and, 
rapidly contracting in width, fuses with the nasal roof in the median line, dorsad to the 
symphysis and beneath the dermal ethmoid (fig. 4). Immediately posterior to the 
palatine symphysis a vertical mesethmoid cartilage divides the much-contracted anterior 
section of the cranial cavity into two short, laterally-situated, tubular passages for the 
transmission of the Olfactory nerves to the nasal sacs (fig. 4, and P]. XXIX. fig. 13, 
ms.é.). Dorsally and posteriorly the mesethmoid cartilage is prolonged into a median 
styliform process (Pl. XXVIII. fig. 5, and Pl. XXIX. fig. 14, st.p.) which extends 
backward for a short distance in the cranial roof internally to the fronto-parietal bone, 
and is presumably a remnant of the more extensive cartilaginous cranial roof of the 
embryo. Inferiorly, the mesethmoid cartilage is coincident with a small oval vacuity 
in the basicranial cartilage, through which it may be seen, in a ventral view of the skull, 
immediately behind the palatine symphysis (Pl. XXVIII. fig. 3, and Pl. XXIX. fig. 13, 
b.c.v.). Anteriorly to this vacuity the vertical extent of the mesethmoid becomes 
rapidly diminished by the upward growth of the symphysis, and finally, after blending 


* Wiedersheim (J. ¢.) was the first to regard the representatives of these cartilages mm Protopterus from 
this point of view. 


340 PROF, T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


below with the basicranial cartilage, becomes continuous in the median line with the 
cartilage of the nasal roof. ‘The tubular passage on each side of the mesethmoid cartilage 
is limited externally by the ascending process of the palato-pterygoid, and not, as in 
Ceratodus, by an extension of the lateral wall of the chondrocranium, and communi- 
cates anteriorly with the nasal sac of its side dorsad to the palatine symphysis. 

The internasal septum (Pl. XXVIII. fig. 4, and Pl. XXIX. fig. 11, i.m.s.) begins as 
an inconspicuous inwardly-projecting ridge from the median line of the cartilaginous 
nasal roof, but is separated from the mesethmoid cartilage by a deep arch-like notch 
(fig. 4) resulting from the upward growth of the palatine symphysis. Anteriorly to the 
symphysis the septum increases in vertical extent and also thickens somewhat, 
supporting on its ventral margin the two acutely-conical vomerine teeth (Pl. XXVIII. 
figs. 3 and 4; Pl. XXIX. fig. 11, vo.t.). In front of the anterior boundary of the nasal 
capsules the internasal septum projects into the upper lip in the form of a short 
prenasal process terminating in two short laterally-directed cornua (Pl. XXVIII. 
figs. 2, 3, and 5, pn.p.). Laterally, the septum is prolonged outward into two thin 
lamine of cartilage with deflected and slightly thickened external margins, which form 
the roof and outer walls of the dorsally-convex and ventrally-concave nasal capsules 
(Pl. XXVIII. figs. 1, 2, and 5, 2.c.). Dorsally, the internasal septum, and to some 
extent also the roof of each capsule, are invested by the dermal ethmoid (fig. 2). 

As in Ceratodus and Protopterus, the continuity of each nasal roof is interrupted by 
a series of vacuities which in Lepidosiren are elongated and somewhat oval in shape, 
transversely disposed, and in the fresh specimen filled in by fibrous membrane!. At 
the anterior margin of each nasal capsule the cartilage is somewhat thicker than else- 
where, and laterally projects outward in the form of a thickened and slightly recurved 
process (figs. 1, 2, 5, ¢.c.). From their position, and their relations to the anterior 
boundary of the nasal capsules, I am inclined to regard these cartilages as representing 
the persistent trabecular cornua of the embryonic skull; in any case, they closely 
resemble the well-known trabecular cornua which Parker [31] has described in such 
Anurous Amphibia as, for example, Bufo ornatus. By Rose [36], on the contrary, the 
equivalent cartilages in Protopterus are regarded as representing a pair of “ upper 
labials,” but on what grounds it is difficult to see. With the exception of a thin 
rod of cartilage (Pl. XXVIII. figs. 3 and 12, sn.p.), which extends inward from 
the deflected outer margin of each nasal capsule and passes between the ventrally- 
situated anterior and posterior narial apertures to a fibrous attachment to the 

* With the exception of the Dipnoi, this curious fenestration of the nasal roof occurs in no other Vertebrates 
except certain Urodele Amphibia [Wiedersheim 43, H. H. Wilder 44], and affords another instance of the 
many homoplastic modifications which are to be noticed in the two groups. 

[Since the above footnote was written, I have been informed by Professor Howes that the nasal capsules of 


a skull of Cestracion philippi, in the Museum of the Royal College of Science at South Kensington, exhibit 
feeble, but nevertheless unmistakable, indications of fenestration. | 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 341 


internasal septum, the two capsules are entirely devoid of any cartilaginous floor, 
These rods may be termed the subnasal cartilages. 

The most noteworthy feature in connection with the olfactory region of the skull of 
Lepidosiren is the effect produced on the chondrocranial elements by the relatively 
massive development of the palatine symphysis. Not only has the dorsal growth of the 
symphysis completely interrupted the direct continuity of the internasal septum and 
mesethmoid cartilage, so that the symphysial portions of the palatines (Pl. XXVIII. 
fig. 4, and Pl. XXIX. fig. 12, p.sy.) are in actual contact with the hinder part of the 
nasal roof in the median line (fig. 12), but, by its extension anteriorly and mesially 
between the two nasal sacs for nearly two-thirds of their extent, the symphysis, and not 
the true internasal septum, forms the actual median division between the two capsules, 
Reference to fig. 12 will sufficiently illustrate these modifications and also show how, in 
consequence of its forward extension, the symphysis reduces the vertical dimension 
of each capsule, while at the same time contributing to the support of the nasal 
floor. 

As will be shown later on, the accounts given by different writers as to the existence 
of upper labial cartilages in the Dipnoi are not always easy to reconcile one with 
another or with actual facts, and not infrequently they are mutually contradictory. 

In Lepidosiren the structures erroneously regarded as labial cartilages by Bischoff 
and Hyrtl are without doubt antorbital cartilages, formed as lateral outgrowths from 
the anterior trabecular region of the chondrocranium, between the orbital and nasal 
regions, and, as already mentioned, may be considered as the equivalents of similarly- 
situated cartilages in the Elasmobranchs and Urodele Amphibians. The true upper 
labials were overlooked by both of these writers, but there can be no question as to 
their existence. Attached by ligament to the posterior margin of each nasal capsule, 
near its junction with the internasal septum, there is a slender cartilaginous filament 
which passes directly downward, immediately anterior to the initial descending portion 
of the antorbital process, and ends in the fibrous posterior wall of the nasal capsule 
(Pl. XXVIII. figs. 1, 3, and 5, w./.c.). These filaments represent a single pair of upper 
labial cartilages, and are the only structures which can be recognized as such in 
Lepidosiren. 

The anterior section of the cranial cavity for a short distance behind the mesethmoid 
cartilage is completely filled by a dense mass of extremely tough fibrous tissue, con- 
tinuous dorsally with the fibrous connection between the dermal ethmoid and the 
fronto-parietal, and also filling up the two olfactory passages. The hinder face of 
this fibrous plug is slightly hollowed out for the reception of the anterior extremity 
of the brain. The mass is also traversed by the two Olfactory nerves as they pass 
forward to enter the two olfactory passages, and into it projects the median styliform 
process from the mesethmoid cartilage (Pl. XXIX. fig. 14). 

The lower jaw of Lepidosiren is very similar to that of Protopterus, as described by 
VoL. XIvV.—Pari V. No. 3.—February, 1898. ZY 


342 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


Wiedersheim (J. ¢.), and each ramus consists of articular, coronoid, and dentigerous 
regions, and a persistent and curiously-modified Meckelian cartilage (Pl. XXVIII. 
figs. 7 and 8, and Pl. XXIX. fig. 9). 

Meckel’s cartilage consists of a thick proximal portion (fig. 8, mk.c.), deeply 
excavated posteriorly so as to form a concave, transversely-disposed, articular surface 
for the suspensorium, and of a thin band-like extension of the foregoing, which makes 
its appearance on the outer surface of the ramus in front of the angular bone (fig. 7), 
and thence runs forward in a groove near the lower border of the splenial (sp.) as far 
as the mandibular symphysis, where it becames continuous with its fellow of the 
opposite side in an expanded symphysial plate (fig. 9). In the angle between the 
anterior and central of the three cutting tooth-plates with which each mandibular 
ramus is furnished, the Meckelian cartilage gives off a vertically-disposed ascending 
process (fig. 7). Three similar, but shorter, processes are also given off from the upper 
margin of the symphysial plate, of which two are supero-lateral and one central, and 
lie in the angle which the anterior tooth-plate forms with its fellow of the opposite 
ramus (fig. 9). A short cartilaginous nodule may also be found between the central 
and posterior tooth-plates, but is connected with the Meckelian cartilage only by 
fibrous tissue (fig. 7). Morphologically, it is possible that the symphysial plate and 
its supero-lateral processes may represent lower labials, as Rose (J. ¢.) has suggested in 
the case of Protopterus; but at present no valid reason has been assigned for 
regarding them as other than expansions or outgrowths from the Meckelian cartilage 
for the support of certain folds of mucous membrane which occupy the outer angles of 
the tooth-plates, and, as Rose (/. c.) himself as shown, are important factors in the 
process of tooth-regeneration and growth. 

Two bones only are represented in each half of the mandible. Of these, the splenial 
(in Protopterus, the ‘“‘dentary” of Owen, Peters, and Miall, and the “articulare” of 
Wiedersheim ; in Ceratodus, the “ dentary ” of Giinther, and the “ splenial” of Huxley) 
is by far the larger, and forms nearly the whole of the bony portion of each ramus, 
retaining, however, its primary and normal position in relation with the inner surface 
of Meckel’s cartilage (figs. 7-9, sp.). Posteriorly and dorsally it rises into an unusually 
strong coronoid process (cor.p.) for the insertion of the powerful temporal and 
masseter muscles, and anteriorly articulates and partially fuses with its fellow in an 
exceptionally massive symphysis (fig. 8, m.sy.), which, internally as well as externally, 
is produced into a strong “spina mentalis” (fig. 8, s.m.i.; fig. 9, s.m.e.). For the 
anterior half of its extent the bone supports dorsally the three splenial or mandibular 
tooth-plates (fig. 7, a.m.t., m.m.t., p.m.t.). The second bone is the “angular” (in 
Protopterus the “articular” of Owen, the ‘‘dentale externum” of Wiedersheim, and 
the “angular ” of Peters and Miall; in Ceratodus, the “‘ articulary ” of Giinther, and the 
“angular” of Huxley). It is a relatively small bone (figs. 7, 9, av.), commencing at 
the extreme angle of the jaw and thence extending for a short distance forward, external 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 343 


to the proximal portion of Meckel’s cartilage, ultimately terminating at the point 
where the latter makes its appearance on the outer surface of the ramus (fig. 7). There 
is no representative in Lepidosiren of the bone which in Ceratodus has been identified 
by Huxley (/. c.) as a dentary element. 

The hyoid arch consists of but a single element on each side—the equivalent of the 
cerato-hyal of other Fishes. It is a somewhat curved, relatively thick bar, cylindrical 
in the middle, laterally compressed proximally, but club-shaped inferiorly, where it is 
somewhat loosely connected by ligament with its fellow (Pl. XXVIII. figs. 1, 3, and 4, 
c.h.). Dorsally, the proximal extremity of the cerato-hyal is connected with the skull 
by a strong hyo-suspensorial ligament (figs. 3, 4, hy.s.l.), which is partly inserted into 
a rounded prominence on the inner surface of the suspensorial cartilage near its hinder 
margin, and is partly continued as a broad band of fibres to the ventral surface of the 
parasphenoid. A strong hyomandibular ligament also takes origin from the outer 
surface of the proximal end of the cerato-hyal, and thence passes obliquely downward 
to an insertion into the angle of the lower jaw. Structurally, the cerato-hyal consists 
of an axial cartilaginous portion, invested, except at its proximal and distal extremities, 
by a relatively thin sheath of superficial bone. ‘here is no trace of any structures 
corresponding to the hyomandibular, or basi- or glosso-hyal, cartilages of Ceratodus. 

A curious patch of cartilage is attached to the anterior surface of the distal portion 
of the cerato-hyal, external to the osseous sheath, and quite distinct from the cartilage 
of the distal extremity (Pl. XXVIII. fig. 1). As to the significance of this cartilage I 
can express no opinion beyond suggesting the possibility that it may be the remnant of 
a cartilaginous hyoidean ray, and may therefore be included in the category of those 
cartilages (vestigial hyoidean rays) which will be subsequently described in connection 
with the opercular and interopercular bones. The existence of several similar patches 
of cartilage in relation with the external surfaces of the ossified portions of the hyo- 
mandibular and cerato-hyal elements of Acipenser has already been noticed by Parker 
and Howes [32a], by whom the cartilages in question were regarded as “rudimentary 
branchial rays” (/. ¢. p. 174). The presence of such nodules or patches of cartilage 
in connection with doth hyoidean segments in this Ganoid affords a further resemblance 
to the hyoidean rays of Klasmobranchs, and strengthens the above suggestion as to the 
real nature of the still more reduced and vestigial rays of Lepidosiren. 

The opercular bones are represented in Lepidosiren by two elements—an operculum 
and an interoperculum (‘“ Kiemendeckelstiicke,” Bischoff; “ Opercularknochen,” Hyrtl ; 
in Protopterus, “ Kiemendeckelstiicke,” Peters; “ opercularia,” Wiedersheim). The 
operculum (in Protopterus, “ preoperculum,” Owen; ‘“ symplectic,” Cobbold ; in 
Ceratodus, “ opercular,” Giinther and Huxley) is a relatively slender bone, situated 
immediately behind and parallel to the posterior margin of the squamosal (Pl. XXVIII. 
fig. 1, op.). Its inferior extremity forms a small disc-like base, and is closely attached 
by ligament to the hinder border of the squamosal, a little dorsad to the articular 

2y2 


844 PROF, T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


extremity of the latter bone. The interoperculum (in Protopterus, “branchiostegal,” 
Qwen, “preoperculum,” Cobbold; in Ceratodus, “ suboperculum,” Giinther, “ inter- 
operculum,” Huxley) is also a somewhat slender bone, shorter than the operculum, 
with a hooked ventral or distal extremity, and situated directly behind and parallel to 
the last-mentioned element (fig. 1, 7.op.). The operculum is attached by strong liga- 
mentous fibres to the hinder margin of the squamosal and the suspensorial cartilage, 
while the interoperculum is similarly connected with the outer surface of the proximal 
portion of the cerato-hyal, with the operculum, and also with the contiguous margins of 
the squamosal and the suspensorial cartilage. A strong but nevertheless slender liga- 
ment extends also from the ventral extremity of the interoperculum obliquely down- 
ward and forward to its insertion into the inferior margin of the mandible. 

The interesting series of cartilages first described by Huxley [15] in connection with 
the inner surface and distal extremity of the operculum and interoperculum of 
Ceratodus are also represented in Lepidosiren, although apparently overlooked both by 
Bischoff and Hyrt]. Closely applied to the inner surface of the interoperculum, and 
considerably wider than the bone itself, is a thin lamina of cartilage (Pl. XXVIII. 
figs. 1 and 4, v.hy.r.), the upper extremity of which is segmented off from the rest as 
an apical piece and projects somewhat beyond the extremity of the bone. In the 
operculum, however, the only indication of these structures is in the form of a thin 
nodule of cartilage attached to the inner surface of the upper extremity of the bone and 
projecting slightly beyond it (fig. 1, v.Ay.r.). As Huxley (/. ¢.) has suggested in the 
case of Ceratodus, these cartilages may be regarded as vestiges of the cartilaginous 
hyoidean rays of Elasmobranchs, and in the latter group are often arranged in two 
series, a dorsal series attached to the hinder margin of the hyomandibular and repre- 
sented in the Dipnoi by the opercular cartilages, and a ventral series connected with 
the cerato-hyal, and having as their Dipnoid equivalents the interopercular cartilages. 
The presence of vestigial cartilaginous hyoidean rays in conjunction with bony 
opercular elements seems to be peculiar to the Dipnoi; at any rate, after a careful 
search I have failed to find any indication of their existence in those Fishes in which one 
might with some reason expect to find them, viz. :—Acipenser, Polyodon, Polypterus, 
and Amia. It is interesting to notice, however, that in Polyodon [3] the bony oper- 
culum and interoperculum still retain the rayed character of their cartilaginous 
predecessors. 

The branchial arches of Lepidosiren have been briefly described, but not figured, by 
both Bischoff and Hyrtl. They are five in number (Pl. XXIX. fig. 10), and separate 
four linear branchial clefts. On each side the arches consist of a series of slender, 
unsegmented, cartilaginous rods, distinct from one another and from their fellows of 
the opposite side, and situated in the walls of the pharynx immediately external to the 
pharyngeal mucous membrane. Dorsally and ventrally their extremities are con- 
nected by a fibrous band; but, apart from the fact that the dorsal band is loosely 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. ; 545 


connected with the fibrous tissue of the base of the skull, no special ligamentous 
attachments to the latter could be perceived. ‘The first arch, which, in consequence 
of the suppression of the hyoidean cleft, is in close proximity to the inner surface of 
the hinder margin of the suspensorial cartilage, dorsad to the upper extremity of the 
cerato-hyal, is the longest of the series, but very slender, The second is much 
stouter, but shorter, slightly expanded at its ventral extremity, and bifurcate and 
grooved at its dorsal end for the reception of the second aortic arch, The third is 
somewhat more slender than the second, but thicker than the first or fourth, and is 
neither grooved nor forked dorsally. The fourth and fifth resemble the third, but are 
very slender, the fifth being much the shortest of the series, No trace of the two 
basibranchial elements, described by Huxley (J. ¢.) as present in Ceratodus, or of the 
epibranchial elements described by Ridewood [35 a] as existing in connection with the 
second and third branchial arches in Protopterus, could be detected in Lepidosiren. 

It may be mentioned that, in consequence of the suppression of the hyoidean or hyo- 
branchial cleft, the hyoidean hemibranch really projects into the cleft separating the 
first and second branchial arches. 

The characteristic “cranial rib”’ of other Dipnoi is present also in Lepidosiren, and 
was erroneously designated by Bischoff (/. ¢.) “ Suspensorium der Schulter ” (Pl. XXVIII. 
figs. 1-4, ¢.r.). Its dorsal extremity is furnished with two rounded cartilaginous con- 
dyles for articulation with a suitably-modified concavity in the chondrocranial cartilage 
between the Vagus foramen in front and above, the exoccipital behind, and the lateral 
edge of the parasphenoid below. Its extremities are cartilaginous, but the rest 
of its extent consists of an axial core of cartilage invested by a relatively thick 
bony sheath. Morphologically, it may be considered to represent the costal element 
pertaining to the first neural arch. 

The attachment of the dorsal extremity of each lateral half of the pectoral girdle 
to the skull is effected by a stout ligament which extends from the postero-lateral 
portion of the cranium, behind and alittle dorsad to the Vagus foramen, and is inserted 
into the upper extremity of the supraclavicle. There is apparently no representative 
of the “ post-temporal ” of Protopterus and Ceratodus. 

The only account of the cranial nerves of Lepidosiren (L. paradowa, Fitz.) is that 
given by Hyrtl (/. ¢.), whose description, in so far as the course and distribution of the 
larger branches are concerned, is in the main accurate, although characterized by the 
defect that the author has not always been successful in discriminating between the 
branches of the Facialis and those of the Trigeminus. In order to ascertain their 
relations to the various cranial foramina, a careful investigation of the peripheral dis- 
tribution of the various cranial nerves of the Paraguayan Lepidosiren was made, with the 
result that my own observations proved to be in fairly close agreement with those pre- 
viously recorded by Hyrtl, and more especially with the account given of the cranial 
nerves of Protopterus annectens in a recent and admirable paper by Pinkus [35]. In 


846 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


the present communication it is intended to refer only to those cranial nerves or their 
branches which are related to cranial foramina, or to certain definite regions of the 
skull. ‘The nomenclature of the nerves will be that adopted by Pinkus in the case of 
Protopterus. 

The Olfactory Nerve [i.]. In passing from the olfactory lobes to the nasal sacs these 
nerves traverse the olfactory passage on either side of the mesethmoid cartilage 
(Pl. XXIX. figs. 13, 14, and 15, i.), and reach their destination after passing forward 
dorsad to the palatine symphysis. 

The Optic Nerve [ii.] This nerve leaves the cranial cavity through a linear cleft 
between the ascending process of the palato-pterygoid and the anterior margin of the 
lateral plate of the fronto-parietal, which is apparently a remnant of the much more 
extensive antero-lateral cranial vacuity of Protopterus, and in the latter Dipnoid also 
transmits the Optic nerve (Pl. XXVIII. figs. 1 and 4, i1.). 

With regard to the nerves supplying the muscles of the eye-ball, viz. the Motor 
Oculi, the Patheticus, and the Abducens, the earlier writers, among whom may be 
mentioned Owen, Hyrtl, and Humphry, failed to find any evidence of their existence 
in the particular Dipnoi they examined. Wiedersheim [41] was apparently the first 
to discover the Motor Oculi nerve in Protopterus, although at the time he was unable 
to state with certainty whether it arose independently from the brain or was a branch 
of the Trigeminus. More recently Pinkus (/. c.) proved that in the same Dipnoid all 
three nerves are present, and not only determined their precise origin from the brain 
but also their distribution in the eye-muscles. Similarly with the muscles themselves, 
Owen [25] asserted that no such muscles were present in Protopterus, and hence the 
absence of their nerves. Hyrtl (7. ¢.) described four recti in Lepidosiren, but failed to 
find the two obliqui. Humphry [13] also found the four recti in Protopterus, but 
no oblique muscles. Pinkus (J. ¢.), on the contrary, found that the latter Dipnoid not 
only possessed the usual recti, but superior and inferior oblique muscles in addition. 
It is therefore extremely probable that all. Dipnoi possess a complete series of eye- 
muscles and also a Motor Oculi, a Patheticus, and an Abducens for their innervation, 
as in the generality of Vertebrates. In the Paraguayan Lepidosiren there is no doubt 
as to the existence of the usual six eye-muscles, but, with the exception of the Motor 
Oculi, I was unable to detect any trace of their nerves. The analogy of Protopterus 
suggests, however, that my failure to discover the Patheticus and Abducens was 
probably due to the extreme tenuity and fineness of these nerves rather than to their 
absence. Unfortunately, lack of the necessary additional material prevented me from 
resorting to the only satisfactory method of determining with absolute certainty the 
presence or absence of the nerves in question, viz. by sections of the head and brain. 

The Motor Oculi (iii.)—The third cranial nerve traverses one of the oblique fissures 
in the ventral margin of the fronto-parietal (Pl. XXVIII. fig. 4, iii.), about midway 
between the foramen for the Optic nerve and that for the ophthalmic branch of the 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 347 


Trigeminus (v.'). Externally, the nerve issues from the cranial cavity immediately 
dorsad to the trabecular cartilage, and thence passes forward to the usual eye-muscles 
(Pl. XXVIII. fig. 1, iii.). 

The Trigeminus (v.).—The various divisions of the Fifth and Seventh cranial nerves 
and certain of their principal branches are related to a considerable number of 
diversely-situated cranial foramina. With the exception of the ramus ophthalmicus 
of the Fifth nerve, their primary branches may almost be said to leave the cranial cavity 
by a single large foramen leading into a lateral diverticulum of the cavity situated at 
the junction of the trabecular and periotic cartilages, and bounded anteriorly by the 
former and posteriorly by the latter cartilage. This diverticulum will in future be 
referred to as the Gasserian recess. The separation and divergence of the primary 
branches of these nerves lead to the formation of a number of closely-related foramina 
of various sizes, either at the outer extremity of the recess or in its margins and floor. 

The first division of the Trigeminus, or the ramus ophthalmicus, leaves the cranial ~ 
cavity by a separate foramen in the hinder part of the trabecular cartilage, immediately 
behind the junction of the parasphenoid and fronto-parietal bones (Pl. XXVIII. 
fig. 6, v.'), and, after a short course obliquely outward and forward, emerges on the 
lateral surface of the skull a little dorsad to the first appearance of the trabecular 
cartilage externally (Pl. XXVIII. fig. 1, v.'). From this point the nerve passes directly 
forward parallel to the trabecular cartilage and dorsad to the eye. One branch of its 
ramus ophthalmicus profundus traverses a foramen in the fibrous tissue of the posterior 
wall of the nasal capsule, situated immediately anterior to the root of the upper labial 
cartilage (fig. 1), and then runs forward between the cartilaginous roof of the capsule 
and the membranous wall of the enclosed olfactory sac; eventually the nerve issues 
from the nasal capsule through a minute foramen in the extreme anterior part of its 
roof (Pl. XXVIII. figs. 2 and 5, y.'p.), and is then joined by another branch of the 
same nerve, which, in its course to the junction, has remained external to the nasal 
roof and parallel to the outer margin of the dermal ethmoid. ‘The ramus maxillaris 
and the ramus mandibularis emerge through a single large foramen in the outer 
extremity of the Gasserian recess (Pl. XXVIII. fig. 6, v."), which is apparent externally 
in the angle between the dorsal extremity of the squamosal and the hinder margin of 
the suspensorial portion of the fronto-parietal (Pl. XXVIII. figs. 1 and 2, y.”). The 
foramen does not open directly on to the lateral surface of the skull, but into the 
“ Schlafengrube” (figs. 1 and 2, and Pl. XXIX. fig. 18, t.f. and v.”), aud through the 
latter communicates with the exterior. The nasal branch of the ramus maxillaris 
traverses a foramen (fig. 1) between the root of the upper labial cartilage and the 
adjacent portion of the antorbital process, and then enters the nasal capsule. In its 
course downward to the lower jaw the ramus mandibularis runs parallel to the 
anterior margin of the squamosal and in contact with the outer surface of that bone. 
A small branch of this nerve disappears through a minute foramen on the outer surface 


348 PROF, T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


of the mandible, between the angular and splenial bones (Pl. XXVIII. fig. 7, v.!"). 
Humphry [13] has described a similar branch in Protopterus, and also states that it 
perforates the lower jaw, but the statement is contradicted by Pinkus (/. c.). Such a 
nerve certainly exists in Lepidosiren and behaves as described above. 

The Facialis (vii.)—The superior palatine branch of this nerve leaves the Gasserian 
recess by a very small foramen in its floor (Pl. XXVIII. fig. 6, vii’). After entering the 
foramen the nerve pursues a forward course, perforating the trabecular cartilage, and 
then runs between the cartilage and the palato-pterygoid and parasphenoid, near the 
sutural line of the two bones; eventually, the nerve emerges through a small aperture 
in the angle between the antero-lateral margin of the parasphenoid and the inner 
border of the palato-pterygoid (Pl. XXVIII. fig. 3, vii.’), and is distributed to the oral 
mucous membrane. One or two branches of the nerve can be traced forward, dorsad 
to the palatine symphysis and directly beneath the upwardly-tilted basicranial cartilage, 
and appear to supply the vertical folds of mucous membrane in the angles of the 
palatine tooth-plates. 

The * nervus lateralis facialis ” (Pinkus, /. c.) leaves the Gasserian recess by the same 
foramen which transmits the maxillary and mandibular divisions of the Trigeminus 
(Pl. XXVIII. fig. 6, v."). Soon after emerging from the “ Schlafengrube ” 
(Pl. XXVIII. figs. 1 and 2, ¢.f.) the nerve divides into a ramus buccalis and a ramus 
ophthalmicus superficialis, the latter accompanying one of the branches of the ramus 
ophthalmicus profundus of the Trigeminus over the dorsal surface of the nasal capsule 
to the skin of the snout. 

The ramus hyomandibularis traverses a relatively large foramen in the postero- 
inferior wall of the Gasserian recess (Pl. XXVIII. fig. 6, vii.). After perforating the 
periotic capsule the foramen opens into the ‘‘ Schlafengrube,” and through it the 
ramus hyomandibularis reaches the inner or under surface of the suspensorial cartilage 
(Pl. XXVIII. figs. 3 and 4, Pl. XXIX. fig. 18, vii.), where it divides into opercular, 
hyoidean, and external and internal mandibular branches. At the emarginate posterior 
border of the suspensorial cartilage (fig. 3, vii.) the ramus mandibularis internus 
passes forward between the cartilage internally and the contracted inferior portion of 
the squamosal externally, and reappears on the outer surface of the suspensorium a 
little above its articular condyle (Pl. XXVIII. figs. 1 and 2, vii.”). At this point the 
nerve is joined by a branch of the ramus mandibularis of the Trigeminus, and then 
extends obliquely downward and forward to the outer surface of the lower jaw. ‘The 
corresponding nerve in Ceratodus (Van Wijhe, 40) and also in Protopterus (Humphry, 
1. ¢., Pinkus, J. ¢.) behaves in an almost precisely similar fashion. 

The communicating branch between the Facialis and Vagus nerves emerges from the 
Gasserian recess through the same foramen which also transmits the maxillary and 
mandibular divisions of the fifth nerve and the nervus lateralis facialis (figs. 1 and 2, 
y."). In its course backward to join the great lateral branch of the Vagus, the ramus 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 349 


communicans occupies a slight groove on the exposed outer surface of the periotic 
capsule at its junction with the proximal border of the suspensorium, and between the 
dorsal extremity of the squamosal and the lateral margin of the fronto-parietal (fig. 1). 
When viewed from the interior of the cranial cavity (Pl. XX VIIL. fig. 6) four foramina 
may be seen in the cartilage of the periotic capsule, of which two are dorsally situated 
and lie close together, just within the cavity of the capsule. The other two may be 
seen on opposite sides of the utricular recess, near its anterior and posterior lips. The 
dorsally-situated foramina (fig. 6, v.s.c.) are the extremities of the tubular canals in 
which the anterior and posterior vertical semicircular canals are lodged, while each of 
the remaining foramina (/.s.c.) transmits the ampullary end of a vertical canal and, in 
addition, one extremity of the horizontal semicircular canal. 

The Glossopharyngeus (ix.).—From the cranial cavity the foramen for this nerve is 
seen just within the hinder margin of the utricular recess (fig. 6, ix.). It perforates 
the posterior portion of the periotic capsule and becomes visible externally just in front 
of the aperture for the Vagus (P]. XXVIII. figs. 1 and 3, ix.). 

The Vagus (x.)—The foramen for this nerve is bounded anteriorly by the periotic 
cartilage, and behind by an emargination in the anterior edge of the exoccipital bone 
(Pl. XXVIII. figs. 4 and 6, x.). The depression into which the cartilage opens 
externally lodges the extraordinarily large Vagus ganglion (Pl. XXVIII. figs. 1 
and 3, X.). 

The Anterior Spinal Nerves.—As correctly described by Hyrtl (/. c.), the dorsal and 
yentral roots of the first spinal nerve (Hypoglossal) emerge through a deep but narrow 
notch in the hinder border of the exoccipital (Pl. XXVIII. figs. 1 and 4, sp.n.), but 
the roots of the second nerve, instead of traversing a foramen in the exoccipital as in 
Protopterus, perforate the fibrous wall of the neural canal between that bone and the 
first neural arch (figs. 1 and 4). The main trunks of the two nerves pass downward 
and a little forward, and unite to form the brachial nerve externally to the proximal 
third of the “cranial rib.” 

Wiedersheim figures and describes in Protopterus (1. c. fig. 10) a branch of the Vagus 
as fusing with the main trunk of the first spinal nerve (Hypoglossal), dorsad to the 
junction of the latter with the second and third spinal nerves to form the brachial 
nerve. The existence of this singular anastomosis has also been affirmed by Iversen 
[19] and more recently by Newton Parker [33]. On the other hand, it is expressly 
stated by Pinkus (/. ¢.): “ Vaguselemente habe ich in den Plexus brachialis nicht 
iibertreten sehen, im Gegensatz zu den Angaben von Wiedersheim, Iversen, und Parker, 
und in Uebereinstimmung mit dem Befund bei Lepidosiren paradoxa” (p. 331). 

With reference to Lepidosiren it may be stated that although the brachial nerve 
contains no ‘‘ Vaguselemente,” yet a connection does undoubtedly exist between a 
branch of the Vagus and the Hypoglossal. Near the point of junction of the Hypo- 
glossal with the second spinal nerve to form the brachial nerve the former gives off a 

voL. XIv.— Part V. No. 4.—February, 1898. ZZ 


350 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


fine branch, which, after a short course obliquely downward and forward across the 
outer surface of the “cranial rib,” joins a slender branch of the Vagus. The latter 
nerve evidently corresponds to a similar nerve in Protopterus, described by Pinkus (J. ¢. 
p- 325) as being peripherally distributed to the tongue-muscles and to the musculature 
of the pharyngeal wall. Hence it follows that, although a ramus communicans does 
connect the Hypoglossal with the Vagus-group of nerves, yet the brachial nerve 
contains no Vagus fibres; and further, we have the interesting possibility that the 
fibres of the Hypoglossal contribute both to the formation of the brachial plexus and 
to the innervation of the muscles of the tongue. 

The Larger Blood-vessels and the Cranial Foramina.—An examination of the relations 
of some of the larger blood-vessels to the cranial foramina yielded results substantially 
agreeing with the previously recorded statements of Hyrtl (J. ¢.). 

The superior jugular or anterior cardinal vein emerges from the cranial cavity 
through the Vagus foramen (Pl. XXVIII. fig. 1, x.), and is at once joined by the 
inferior or external jugular vein. 

The internal carotid artery perforates the base of the skull near the outer margin of 
the parasphenoid (Pl. XXVIII. fig. 3, in.c.), and ultimately enters the cranial cavity 
through a small foramen near the inferior margin of the Gasserian recess (Pl. XXVIII. 
fig. 6, in.c.). After its origin from the first aortic arch the external carotid traverses 
the “Schlafengrube,” and, in company with the mandibular and maxillary branches of 
the Trigeminus nerve, appears on the lateral surface of the skull. It then breaks up 
into several branches for the supply of the temporal and masseter muscles and the 
orbital region, and also sends a considerable branch to the lower jaw, which in its 
course accompanies the ramus mandibularis of the Fifth nerve. 


III. Revision and Comparison of the Structure of the Skull in Ceratodus, 
Protopterus, and Lepidosiren. 


A. Ceratodus. 


In comparing the cranial characters of the three genera it will be advantageous and 
certainly convenient to begin with the skull of Ceratodus. For the necessary data I 
shall rely principally on the accounts of Giinther and Huxley, but in certain minor 
details on the results of my own observations. 

The most obviously distinctive feature in the skull of Ceratodus is the retention 
throughout life of a complete chondrocranium, forming the continuous dorsal, lateral, 
and ventral walls of the cranial cavity. To this may be added the fact that the 
ethmoidal and gasal regions are better developed and more normal than in any other 
Dipnoi. By way of illustration, as well as for the sake of comparison with other types, 
a brief description of the ethmo-nasal region may be given. tae 

The anterior section of the cranial cavity is continuous with two tubular olfactory 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 351 


passages, situated immediately dorsad to the palato-pterygoid symphysis and provided 
with complete cartilaginous walls, continuous behind with the rest of the chondro- 
cranium. Mesially, the two passages are separated by a thick vertical mesethmoid 
cartilage (Pl. XXIX. fig. 20, ms.e.). More anteriorly the floor and outer wall of each 
olfactory passage terminate somewhat abruptly, leaving a large foramen for the entrance 
of an Olfactory nerve into the nasal sac of its side, but the mesethmoid cartilage and the 
overlying chondrocranial roof are directly continuous with the internasal septum and 
with the cartilaginous lamine which grow out on each side from the dorsal edge of 
the septum to form the irregularly-fenestrated roof of the nasal capsules. The more 
normal condition of the mesethmoid region in Ceratodus may be associated with the 
relative thinness of the symphysial portions of the two palato-pterygoid bones, so that 
the vertical constriction of this region by the excessive thickening of the symphysis, 
which is so characteristic a feature in Lepidosiren, does not take place, and, in con- 
sequence, the mesethmoid cartilage and internasal septum are freely continuous, and 
the palatine symphysis remains widely separated from the chondrocranial roof. Com- 
parison of figs. 12 and 20 (Pl. XXIX.), representing equivalent sections through the 
symphysis in Ceratodus and Lepidosiren, will sufficiently illustrate these remarks. 

The roof of the two nasal sacs terminates somewhat in front of the internasal septum 
in a thin, marginally-rounded lamina of cartilage (Huxley, /. c. fig. 7, a.), without 
presenting any recognizable indications of trabecular cornua, and without exhibiting so 
much as a vestige of the bicornuate prenasal process of other Dipnoi. 

The presence of representatives of the antorbital cartilages of other Dipnoi is 
doubtful. The central trabecular region of the chondrocranium is directly continuous 
with the cartilaginous side-walls of the mesethmoid region, internally to the junction 
of the preorbital portion of the dermal ectethmoid and the ascending process of the 
palato-pterygoid ; and if, as Rose [36] suggests, the hinder upper labial cartilages of 
Huxley are really the equivalents of antorbital processes, it is obvious that they have 
lost their primitive continuity posteriorly with the trabecular cartilage (Pl. X XIX. 
fig. 20, an.p.). 

The characteristic plug of dense fibrous tissue which in Lepidosiren fills up the 
anterior section of the cranial cavity is entirely absent in Ceratodus, and the anterior 
extremity of the brain is in close relation with the hinder edge of the mesethmoid 
cartilage. 

Posteriorly, the separation between the skull and the vertebral column is less evident 
than in either Lepidostren or Protopterus, inasmuch as at least two pairs of “ basi- 
dorsals,” representing the lateral elements of two neural arches, are partially confluent 
with one another and also with the cartilage of the exoccipital region of the skull. 

The suspensorium approximates more closely to the typical autostylic or Chimeroid 
condition than in any other existing Dipnoid. The metapterygo-quadrate cartilage is 
very strongly developed, especially behind, where it is prolonged into a thin but wide 

2242 


352 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


cartilaginous plate which, on each side, helps to roof in the branchial apparatus, while 
a narrow strip of cartilage extending forward from the distal end of the suspensorium 
along the dorsal border of the palato-pterygoid bone as far as the ethmoidal region, 
and continuous dorsally with the lateral margin of the cartilaginous basis cranii, may 
be regarded as representing a palato-pterygoid cartilage. Further, it may be affirmed 
that the condyle for the lower jaw is somewhat more anteriorly situated than in 
Lepidosiren, or, in other words, the suspensorium of Ceratodus is inclined forward to a 
greater extent and makes a more acute angle with the fore part of the basicranial axis. 

The identification of upper labial cartilages in Ceratodus is by no means easy to 
determine satisfactorily.  Giinther (J. ¢.) apparently overlooked the existence of 
possible representatives of these structures, but subsequently two pairs of cartilages 
were discovered by Huxley (J. ¢.), the position and relations of which he thus 
describes :—‘ One of them lies in the roof of the mouth, just in front, and on the inner 
side, of the posterior nasal aperture. It is fixed to the mesethmoid cartilage (m.c.) by 
fibrous bands, and is broader behind than in front. The inner edge of this cartilage is 
concave, the outer convex, and it has a nearly horizontal direction. The second 
cartilage [J. c. figs. 4, 5, 7, 2] is stouter, and lies behind, and on the outer side of, the 
posterior nasal aperture. Its dorsal end is attached to the base of the skull and the 
anterior part of the palato-pterygoid cartilage, just above the middle of the palatine 
tooth. It then descends into the upper lip, near the angle of the mouth” (J. c. pp. 32- 
33). Huxley’s description of the first of these cartilages, which provisionally may be 
called an anterior upper labial, is quite accurate, except that its mesial attachment is 
rather to the internasal septum than to the mesethmoid cartilage; and it may also be 
added that the outer extremity of the cartilage extends into the horizontal fibrous 
septum between the two narial apertures. On the other hand, his description of the 
“posterior upper labial” is not quite correct, inasmuch as the dorsal attachment of 
the cartilage is neither to the base of the skull nor to the anterior part of the palato- 
quadrate cartilage ; on the contrary, the dorsal or proximal extremity of the cartilage 
is connected by ligament with the fibrous suture between the ascending process of the 
palato-pterygoid bone and the preorbital portion of the ectethmoid, and hence by the 
sutural union of these bones the cartilage is widely separated from the chondrocranial 
portion of the skull. It is, in fact, somewhat difficult to be quite sure that Huxley's 
posterior labials are not the equivalents of antorbital cartilages, such as are present in 
Lepidosiren and Protopterus, but otherwise wholly absent in Ceratodus, and, indeed, 
have been so considered by Rose [36], whose view is corroborated by the fact that each 
cartilage extends downward into the upper lip, near the angle of the mouth. At the 
same time, it is evident that these cartilages differ from the antorbital processes of 
other Dipnoi in not being continuous posteriorly with the trabecular region of the 
chondrocranium, although it is at least possible that this want of continuity may be 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 353 


the result of the exceptional forward and downward growth of the dermal ectethmoid 
in front of the eye. It is also equally clear that the cartilages in question do not 
precisely agree in position with the upper labials of Lepidosiren, inasmuch as they have 
no ligamentous connection with the hinder margin of the nasal roof. Considered 
either as antorbital processes or as labial cartilages, there can be no doubt that their 
position and relations have been greatly modified by the extensive preorbital growth of 
the dermal ectethmoid, which seems to have dissociated them, as it were, from their 
primitive relations to the chondrocranium or to the nasal capsules, as the case may be, 
and in the absence of developmental data it seems impossible to decide as to the real 
nature of these structures, The anterior upper labials of Huxley have certainly nothing 
in common with the undoubted upper labials of other Dipnoi, since they lie anteriorly 
to the posterior nasal aperture, practically, in the floor of the nasal sac between the 
two narial openings—that is to say, in the position occupied by the inner half of the 
subnasal cartilage in Zepidosiren; and if they are rightly to be regarded as upper 
labials, it is evident that they must represent an additional pair of these structures 
which have no counterpart in any other Dipnoi. For my own part, I am inclined 
to believe that these cartilages owe their existence to the dismemberment of a 
subnasal cartilage, primitively continuous with the outer margin of the cartilaginous 
nasal roof, and therefore cannot be true labials. ‘There are several facts which seem 
to be in harmony with this suggestion. In the first place, the so-called anterior labials 
of Ceratodus and the subnasal cartilages of Lepidosiren occupy precisely similar positions 
in the floor of each nasal sac between the anterior and posterior nasal apertures, and 
both are attached, internally or mesially, to the internasal septum by fibrous tissue. 
Further, the outer margin of each nasal capsule in Ceratodus gives off a short lateral 
process (Huxley, /. ¢. p. 37, fig. 7), which coincides externally with the position of the 
‘anterior labial” internally and ventrally, and may possibly represent the outer portion 
of a transversely-divided subnasal cartilage. Lastly, the subnasal cartilage of Lepido- 
siren is somewhat constricted at one point—a fact which may perhaps be taken to 
suggest the possibility of its inner portion becoming segmented off from the rest as an 
independent cartilage similar to the supposed anterior upper labial of Ceratodus. 

The only endochondrial bones in Ceratodus are the two exoccipitals. They have 
been correctly described by Huxley as two hollow cones of bone embedded in the side- 
walls of the skull near its junction with the vertebral column. Lach is said to be 
“wider above and externally than below and internally, where it lies above the 
notochord ” [7. ¢. p. 38]. I may add that the bones are so thickly invested externally 
by cartilage as to be completely hidden in a lateral view of the skull, and for this 
reason in all probability they escaped the notice of Giinther. 

With the exception of a fronto-parietal bone, which is entirely absent, and the 
presence of a “scleroparietal” (Giinther, /. ¢.), and also of a series of four or five 


854 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


circumorbital ossicles, the skull-bones of Ceratodus correspond with those of Lepido- 
siren, although differing somewhat from them in their regional extension and their 
precise relations to the chondrocranium. 

The dermal ethmoid is a relatively larger bone than in other Dipnoi, and extends so 
far posteriorly as to project beyond the mesethmoid region, and by its sutural union 
with the “ scleroparietal” (Giinther) widely separates the two ectethmoids. The bone 
identified by me as the equivalent of the Amphibian dermal ectethmoid is a much larger 
and more important element than in Lepidosiren, and has already been sufficiently 
described (p. 831). The most noticeable feature about it is the downward growth and 
lateral thickening of its preorbital portion, so that a conspicuous bony antorbital protu- 
berance separates the nasal from the orbital region of the skull. The mesial area on the 
dorsal surface of the skull, between the two dermal ectethmoids, which in Lepidosiren 
is occupied by an extension of the temporal fasciz, is filled up in Ceratodus by a thin, 
even partially transparent lamina of bone—the “ scleroparietal ” of Giinther, or the 
postero-median bone of Huxley. The squamosal bone consists of two distinct but 
nevertheless continuous portions, the representatives of Huxley’s “ preopercular” 
and “proper squamosal” divisions. The first of these overlies the suspensorial 
cartilage and is a relatively narrow bone, evidently the equivalent of the entire 
squamosal of Lepidosiren. The second portion is an extension of the dorsal extremity 
of the former in the shape of an expanded, fan-like, but thin plate of bone, 
situated externally to the jaw-muscles, and articulating superiorly by an extensive 
suture with the outer margin of the postorbital portion of the dermal ectethmoid 
and with one of the circumorbital ossicles. The hinder portions of the dermal 
ectethmoids, the “scleroparietal,” and the dorsal portions of the squamosals all 
combine to form the eminently characteristic supracranial roof. As regards the mode 
of development of these various factors, there is to my mind little doubt that they are 
ossifications of the temporal fasciz. The parasphenoid is a purely ichthyic bone, and 
in no way contributes to the formation of the lateral walls of the cranial cavity. Itisa 
relatively much larger bone than in other Dipnoi, extending so far anteriorly as slightly 
to overlap the palatine symphysis and entirely hide from view the extensive area of 
basicranial cartilage which is freely exposed behind the symphysis in Lepidosiren, 
while laterally it slightly overlaps the inner surfaces of the two palato-pterygoid bones. 
Posteriorly the bone extends for a relatively greater distance beneath the notochord 
than in other Dipnoi, reaching to about the origin of the third pair of ribs. 

The palato-pterygoid bones are less massive structures, and their symphysial 
portions, although disposed much as in Lepidosiren and overlapped by the nasal sacs, 
are but moderately thick plates. Owing to the smaller size of the tooth-plates, there 
are no conspicuous lateral projections from the palatine bones for their support, and 
there is no obvious representative of the “processus antorbitalis ” of Lepidosiren. A 
feebly-developed ascending process is present, and articulates with a descending plate 
derived from the preorbital portion of the dermal ectethmoid, the two processes being 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 355 


closely applied to the lateral chondrocranial wall of the mesethmoid region. Perhaps 
the only other noteworthy feature is the more vertical position which each bone 
assumes in passing from the distal end of the suspensorial cartilage to the symphysis 
anteriorly—a peculiarity which evidently owes its explanation to greater forward 
inclination of the suspensorium itself. 

The opercular and interopercular bones and their cartilaginous vestigial hyoidean 
rays are essentially similar to those of other Dipnoi, but on the whole are perhaps 
better developed. 

The three suborbital bones described by Huxley were represented in the specimen 
examined by myself by four 1 somewhat irregularly-shaped and tubular ossicles which 
lodged the infraorbital division of the sensory canals of the head. There is a similar 
but much larger bone situated somewhat above and behind the orbit, articulating 
internally with the supraorbital margin of the dermal ectethmoid and behind with the 
lamellar portion of the squamosal, where it is also in fibrous connection with the 
hindermost of the infraorbital series. This bone is likewise traversed by a tubular 
canal, and, as it transmits the supraorbital sensory canal, may perhaps be considered 
as the remains of a supraorbital series of ossicles. The latter bone is apparently the 
one figured by Huxley (/. c. fig. 7, p. 37), and by him described as one of the com- 
ponent elements of the supracranial roof. Jt has also been figured by Miall [22] as a 
“ postorbital ” bone. 

The lower jaw is very unlike that of other Dipnoi. It is relatively much more 
slender and has little more than a rudiment of a coronoid process. The largest bones 
are those described by Huxley as the splenial and the angular, but while the former 
is wholly restricted to the inner surface of the ramus, the latter forms the outer surface 
and extends from the articular extremity nearly to the mental symphysis of the two 
splenial elements. The two dentary plates described by Huxley are peculiar to 
Ceratodus. Anteriorly to the symphysis the two Meckelian cartilages unite in a 
torwardly-projecting concave lamella, regarded by Giinther as representing ‘lower 
labial cartilages,” but doubtfully so considered by Huxley. There is no trace of the 
vertically-disposed processes which in Lepidosiren are derived from the Meckelian 
cartilages and support the folds of mucous membrane projecting into the angles 
between the mandibular tooth-plates. 

The hyoid arch is much more complete than in any other Dipnoid, and was originally 
described by Giinther (/. ¢.) as consisting of a pair of basihyals and a glossohyal. 
Ridewood [35 q@] subsequently showed that the paired basihyals are really hypohyals, 
and the “glossohyal” a true basihyal. In addition to these elements there is also a 
variously-shaped cartilage attached by ligament to the inner and hinder margin of each 
suspensorial cartilage, dorsad to the upper extremity of the cerato-hyal, and considered 
by its discoverer, Huxley (/. ¢.), and also by Gadow (ga) and Ridewood (J. ¢.), to 
represent a hyomandibular element. On the authority of Ridewood (J. ¢.) it would 


» Traquair (37, p. 9) found five suborbital bones in a specimen which he examined. 


356 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


appear that the ventral or symplectic extremity of the hyomandibular element may be 
segmented off as a separate cartilage, and further, that an additional nodule of cartilage, 
which is occasionally found in the hyo-suspensorial ligament, “ may possibly have the 
value of an interhyal” (J. c. p. 636). No trace of any patches of cartilage or 
“vestigial hyoidean rays” could be detected in connection with the cerato-hyal in 
the only specimen of Ceratodus which I have had the opportunity of examining. 

Ceratodus has five branchial arches, with a rudiment of a sixth if Huxley has 
correctly interpreted the nature of a small nodule of cartilage at the ventral extremity 
of the fifth arch. ‘The first four are bisegmental, consisting of a “long ventral and a 
short dorsal piece of cartilage ”; the last, of a single cartilage only. Two small cartilages 
situated in the median and ventral line, between the ventral extremities of the branchial 
arches, have been considered by Huxley to represent ‘“‘ anterior and posterior meso- 
branchials” (/. c. p. 27). 

There is a well-developed hyoidean cleft between the hyoid arch and the first 
branchial arch. 

A representative of the ‘‘ cranial rib” of other Dipnoi is present, but its relations to 
the ossified “ basi-dorsals ” of the first neural arch are precisely those of the succeeding 
ribs to their basi-dorsals, 

The attachment of the dorsal extremity of each lateral half of the pectoral girdle to 
the skull is effected by “a thin broad lamella of an obliquely ovate shape, entirely 
ossified” (Giinther, 7. ¢. p. 531), and directed obliquely backward and outward. It 
is directly connected by ligament at its upper or inner extremity with the lateral wall 
of the chondrocranium, between the Vagus foramen and the exoccipital, and distally 
also by ligament with the supraclavicle of Parker. The bone has been termed “ supra- 
scapula” by Giinther, and its equivalent in Protopterus ‘“‘ suprascapula” (Peters, /. ¢.) 
and “ post-temporal” (Parker, 32). 


B. Protopterus. 

With the exception of minor structural features, and certain modifications of 
nomenclature, the data necessary for the comparison of the skull of this Dipnoid with 
those of Ceratodus and Lepidosiren have been principally taken from Wiedersheim’s 
paper (J. ¢.). 

Owing to the development of a large fronto-parietal bone and the share which it 
takes in forming the roof, and also, in conjunction with the parasphenoid, the lateral 
walls of the cranial cavity, the chondrocranium has largely atrophied, leaving, neverthe- 
less, an occipito-periotic portion and an ethmo-nasal region, connected by a pair of 
laterally-situated and relatively narrow, band-like, trabecular cartilages, which in the 
middle portion of the skull are the only remains of the complete chondrocranium of 
Ceratodus. The occipito-periotic cartilage is somewhat more extensive than in Lepido- 
siren, the cartilage of the supraoccipital region extending on the inner surface of the 


a 


IN THE PARAGUAYAN LEP{DOSIREN, ETC. 557 


fronto-parietal as far forward as the anterior limit of the periotic capsules, while the 
exoccipital bones are invested externally, but not internally, by cartilage. In fact the 
chondrocranium is complete dorsally and laterally in the periotic and post-auditory 
regions, and would also be so ventrally were it not that the basal cartilage ceases a little 
behind the middle of the periotic region. The trabecular cartilages are similar to 
those of Lepidosiren, but of somewhat greater vertical extent in the region immediately 
anterior to the periotic capsules, and for a short distance behind the junction of the 
fronto-parietal and parasphenoid they even contribute to the formation of the lateral 
walls of the cranial cayity (Wiedersheim, /. c. Taf. ii. fig. 5), Precisely as in Lepido- 
siren, the trabecular cartilages become continuous in front of the anterior termination 
of the parasphenoid with the persistent and upwardly-deflected basicranial cartilage of 
this region ; but, again separating from it anteriorly, they diverge outward and down- 
ward across the outer surface of the palato-pterygoid bone in the form of extremely 
slender antorbital cartilages. The latter curve downward and backward into the 
hinder margin of the upper labial fold and there terminate, but without extending 
forward into its inferior border, or giving off anteriorly-directed processes. The 
continuity of the trabecular and antorbital cartilages has been described by Wieders- 
heim (/. ¢.), although, curiously enough, it is not represented in any of his figures. 
Peters (/. ¢.), however, was the first to show the relations of the two structures in his 
figure of the skull of a Protopterus from Zambesi, and indeed gives a more accurate 
representation of the shape and curvature of the antorbital process than any other 
figure with which I am acquainted. 

From the point where the basicranial cartilage begins to be deflected upward by 
the palato-pterygoid symphysis the ethmo-nasal portion of the chondrocranium differs 
but little from that of Zepidosiren, and the figures given of this region in the latter 
Dipnoid are, in the main, equally applicable to Protopterus. Trabecular cornua, 
subnasal cartilages, and a bilobed prenasal process are present, and in every respect are 
almost identical with the corresponding structures in Lepidosiren. In both genera the 
fenestration of the roof of the two nasal capsules is regular and almost symmetrical. 
The presence of subnasal cartilages was overlooked by Wiedersheim, although they had 
previously been figured by Peters (J. ¢. Taf. ii. fig. 5, /'), whose accuracy in this respect 
J am able to confirm. I notice that Wiedersheim (/. ¢. Taf, ii. fig. 5) represents that 
portion of the internasal septum which carries the two vomerine teeth as segmented off 
from the hinder part of the septum and separated from it by a curved vertical suture, 
and a statement to the same effect occurs in the text (/. ¢. p. 52). With reference to 
this point I can only say that in a skull which I have had the opportunity of examining 
no such division could be detected, and the septum was continuous throughout its 
extent. The same figure is also slightly inaccurate in not representing the mesethmoid 
cartilage which, as in Lepidosiren, is situated immediately behind the palato-pterygoid 
symphysis. 

VOL, XIv.—Part Vv. No. 5.—February, 1898. 3A 


358 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


Although previously unnoticed, or at all events unrecorded 1, a well-marked basi- 
cranial vacuity or cleft is present, and may be seen in a ventral view of the skull close 
to the hinder margin of the palatine symphysis. It is oval in shape, and filled up by a 
thick fibrous sheet continuous internally with the fibrous mass which occupies the 
ethmoidal portion of the cranial cavity. 

From a comparison of the figure of a plastic model of the nasal region given by Rose 
(7. c. fig. 4) with a similar model of the same region figured by himself, Pinkus 
(1. c. fig. 7) infers that the cartilaginous lamine which separate the oval vacuities in 
the nasal roof increase in number with the growth of the fish and still further 
subdivide pre-existing vacuities. 

No mention is made of the existence of upper labial cartilages in Protopterus either 
by Owen or Cobbold, or even in the excellent and much more recent paper by Wieders- 
heim. According to Huxley [16] there are ‘“‘two upper labial cartilages—one 
fibro-cartilaginous immediately behind the anterior narial aperture, and the other 
behind the posterior narial opening. These answer to the upper labial cartilages of 
Chimera and Cestracion” (1. c. p. 181). Rose (J. c.) describes two pairs of upper 
labials, an anterior pair represented by the terminal lobes of the prenasal process, and 
a posterior pair which evidently correspond to the thickened and laterally-recurved 
anterior margins of the nasal capsules in Zepidosiren. Peters (J. c.) has also figured, 
but without describing, two pairs of upper labials. 

Huxley’s account is not very clear, but from his statement of their position I think 
there can be no doubt that his anterior labials are really the subnasal cartilages, 
and not true labials at all, while his description of the posterior labials leaves one in 
doubt whether he is referring to the antorbital cartilages or to the equivalents of the 
undoubted upper labials of Lepidosiren. The identification of upper labials given by 
Rose is certainly based on insufficient evidence, inasmuch as there is at present no 
reason to believe that the cartilages which he terms labials have developed indepen- 
dently of the nasal region of the chondrocranium, and I have already given reasons for 
the view that his posterior labials are persistent trabecular cornua, such as not 
infrequently form the anterior walls of the nasal capsules in many Anurous Amphibia. 
The posterior labials of Peters are really the antorbital processes, but from their 
position it would seem probable that the structures figured by him (/. ¢.) as anterior 
labials are the representatives in Protopterus of the single pair of upper labials of 
Lepidosiren: 

With the object of clearing up the obscurity as to the existence of upper labials in 
Protopterus, | made a careful examination of a skull of this Dipnoid, with the result 
that there is no doubt that Protopterus and Lepidosiren are in close agreement so far 


‘ A linear shading in one of Peters’s figures (J. c. Taf. ii. fig. 8):may be intended to represent this vacuity, 
but no further reference is made to it, either in the figure or in the text. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 309 


as these structures are concerned. Protopterus, in fact, has but a single pair of upper 
labial cartilages, which, in position and relations, are identical with those of Lepidosiren, 
and also with the so-called “anterior labials” correctly figured by Peters sixty years 
ago, but apparently overlooked by every subsequent writer. 

The suspensorial cartilage of Protopterus differs from that of Ceratodus in the 
restricted antero-posterior dimension of its proximal portion, which does not therefore 
arch backward over the branchial apparatus to the extent it does in the latter genus, and 
also in the absence of any representative of a palato-pterygoid cartilage. The cartilage 
figured by Wiedersheim (/. ¢. Taf. ii. fig. 3, tr.) fringing the dorsal border of each 
palato-pterygoid bone, and rightly named by him the “ pars trabecularis cranii,” lies 
above the level of the cranial floor, and is therefore a remnant of the lateral chondro- 
cranial wall and not a palato-pterygoid cartilage. The suppression of the latter 
cartilage is perhaps to be associated with the increased relative thickness of the 
palato-pterygoid bone, and its more intimate relations with the lateral margins of the 
basis cranii. On the other hand, Protopterus agrees with Ceratodus and differs from 
Lepidosiren in the degree of inclination of the suspensorium to the basicranial axis. 

The two exoccipitals of Protopterus are peculiar in that they meet and fuse dorsally 
beneath the cartilage of the supraoccipital region. Wiedersheim regards them as 
representing a ‘‘supraoccipitale,’ but the resemblance of the lateral halves of the bone 
to the paired bones of Lepidosiren, both in shape and in their relations to the foramina 
for the exit of the Vagus and Hypoglossal nerves, is sufficiently close to justify one in 
regarding them as a pair of confluent exoccipital elements. 

The remaining bones of the skull are on the whole very similar to those of Lepidosiren. 
As compared with Ceratodus the most striking differences relate to the presence of a 
fronto-parietal forming a complete gable-roof to the cranial cavity, except for a limited 
extent between its anterior margin and the dermal ethmoid, and also, in conjunction 
with the marginal ascending plates of the parasphenoid, contributing to form the bony 
side-walls of the central region of the same cavity. It may be remarked, however, that 
in the suspensorial region the fronto-parietal is restricted to the cranial roof, and does 
not in any way invest the outer surface of the periotic capsule or the suspensorial 
cartilage, or hide in an external view the continuity of either with the trabecular 
cartilage. To these features may be added the absence of a complete supracranial 
roof, due to the want of a “scleroparietal” and of the superficial lamellar plates of 
the squamosal, combined with the relative narrowness of the dermal ectethmoids. The 
characteristic preorbital or nasal section of the dermal ectethmoid of Ceratodus is 
wholly wanting in Protopterus, the bone being displaced upward and backward by 
the growth of an exceptionally large ascending process from the palato-pterygoid bone. 
Nevertheless, the dermal ectethmoids are much better developed than in Lepidosiren, 
and, in the absence of a “ scleroparietal,” their expanded anterior portions meet in a 


median suture dorsad to the fronto-parietal. 
3A 2 


360 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


A noteworthy feature in Protopterus is the existence of extensive lateral vacuities in 
the side-walls of the anterior section of the cranial cavity, between the anterior margins 
of the lateral plates of the fronto-parietal bone and the ascending processes of the 
palato-pterygoids, which are continuous dorsally with a fissure between the fronto- 
parietal and the dermal ethmoid. These vacuities are filled up by a tough fibrous 
membrane, continuous internally with a dense mass of fibrous tissue previously 
mentioned as obliterating the cranial cavity in this region, while the dorsal fissure is 
closed by the fibrous tissue which extends between the contiguous margins of the 
fronto-parietal and the dermal ethmoid, and is also continuous internally with the 
cranial mass of the same tissue. 

The parasphenoid is much more restricted in its relations to the basis cranii than in 
Ceratodus, inasmuch as it terminates somewhat abruptly by a nearly straight transverse 
margin at some distance behind the palatine symphysis, and only very slightly projects 
backward beneath the post-cranial section of the notochord. In the share which it 
takes in forming the lateral walls of the cranial cavity, and in its relation to the 
trabecular cartilages and the palato-pterygoid bones, the parasphenoid differs but little 
from its homologue in Lepidosiren. 

The palato-pterygoids are also essentially similar to those of Lepidosiren, especially 
in the modifications produced in the ethmo-nasal region of the chondrocranium by the 
massive development of their symphysis. They have well-marked lateral processes for 
the partial support of the two anterior tooth-plates, and a strong “ antorbital process” 
for the posterior plate. There are also stout ascending processes which, in addition to 
carrying the dermal ectethmoids, form the outer walls of the olfactory fosse anteriorly 
to the lateral cranial vacuities. 

There is a ring of fibrous tissue encircling the orbit, but no circumorbital bones are 
developed in it. The opercular and interopercular elements are essentially similar to 
those of Ceratodus, but relatively smaller. 

Vestigial hyoidean rays have not previously been recorded as existing in Protopterus, 
but there is no doubt as to their existence. The opercular cartilage closely 
resembles the corresponding piece in Ceratodus (Huxley, /. c. p. 38). It is horseshoe- 
shaped and closely applied to the inner surface of the upper third of the operculum. 
The interopercular cartilage, on the other hand, is essentially similar to its representative 
in Lepidosiren. 

The hyoid arch is a much simpler structure than in Ceratodus, and, as in Lepidosiren, 
there is no trace of hyomandibular, hypohyal, or basihyal elements. As in the latter 
Dipnoid, there is a patch of cartilage (vestigial hyoidean ray) on the outer surface of 
the distal portion of each cerato-hyal. No mention of this cartilage is made by 
Wiedersheim, although it is apparently figured by Peters (JU. ¢. Taf. ii. fig. 2), without, 
however, being indicated by reference-letters or referred to in the text. 

As regards the precise number of branchial arches in Protopterus, there is some 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 361 


discrepancy in the statements of different writers. According to Owen [25], Peters 
[34], McDonnell [21], and Wiedersheim [41], there are six branchial arches. Newton 
Parker [33] states, however, that there are but five, and is apparently doubtful as to 
the existence of the arch which Wiedersheim (J. ¢. p. 56, fig. 8) figures and describes 
as situated in front of the first cleft and forming the first of the series of six. ‘There 
is also some obscurity as to the existence of a hyobranchial or hyoidean cleft. Neither 
Owen nor Peters is explicit on this point, although from their statements as to the 
position and relations of the gill-filaments and branchial clefts the absence of a hyoidean 
cleft may perhaps be inferred. Wiedersheim figures (J. c. fig. 8) the first cleft as 
situated between his first and second branchial arches, but there is no special reference 
in the text to the presence or absence of a hyobranchial cleft. On the other hand, 
according to Newton Parker (/. ¢. pp. 161-162), a hyobranchial cleft is present, 
and has in relation with its anterior wall the hyoidean hemibranch or “ opercular 
gill.” 

On examining the branchial region of a specimen of Protopterus 31 cm. in length, I 
had no difficulty in detecting the presence of an extremely slender, unsegmented, 
cartilaginous filament, 7 mm. in length and rather less than 1 mm. wide, situated 
immediately in front of the first branchial cleft and forming its anterior boundary. 
The mucous membrane covering the posterior margin of the filament carried a series of 
minute tooth-like projections, similar to the single row of the second arch and to the 
double series found on the anterior and posterior margins of the succeeding arches, with 
the exception of the last, which has but a single row. The cartilage occupies a position 
precisely similar to the first branchial arch in Lepidosiren, being in close relation with 
the hinder margin of the suspensorial cartilage, dorsad and posterior to the proximal 
extremity of the hyoid arch. It is scarcely open to doubt that this slender cartilaginous 
rod represents the first of a series of six branchial arches, and, as there is certainly no 
cleft anterior to it, the conclusion that the hyobranchial cleft has undergone total 
suppression necessarily follows. It may be mentioned that Pinkus (J. c. p. 318) records 
the presence of a small piece of cartilage embedded in the oral mucous membrane 
opposite the first gill-arch. The cartilage is apparently very small, being only 1:08 mm. 
in length in a specimen 14 em. long, and 2:36 mm. in one 17 cm. in length. “Sein Bau 
entspricht genau dem der iibrigen Kiemenbogen, nur dass es bloss eine, und zwar 
mediale, Reihe zahnartiger Zacken triigt, deren die iibrigen Bogen je zwei, eine mediale 
und eine laterale, besitzen.” After discussing the relations of the cartilage to the 
branches of the Glossopharyngeal and Vagus nerves, Pinkus concludes :—* Ich rechne 
diesen rudimentiiren Kiemenbogen bei der Zihlung nicht mit, sondern zihle, der 
Innervation und dem iiblichen Schema entsprechend, den darauf folgenden Bogen als 
ersten Kiemenbogen. Demnach liegt die erste Kiemenspalte zwischen diesem Kiemen- 
bogen und dem Hyoid, die zweite Kiemensplate zwischen diesem Kiemenbogen und 
dem zweiten Kiemenbogen” (p. 318). Iam not quite certain that the cartilage Pinkus 


562 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


describes is identical with the one which, in agreement with Wiedersheim, I have 
described as existing in front of the first cleft, although it is extremely probable that 
such is the case; but I entertain no doubt whatever that the latter cartilage is 
the first branchial arch and that the hyobranchial cleft has been completely suppressed, 
and further that, as in Lepidosiren, the hyoidean hemibranch or pseudobranch is in 
relation with the anterior wall of the cleft between the first and second branchial 
arches. 

In Wiedersheim’s description all the remaining five branchial arches are represented 
as simple, non-jointed, and very slender rods of cartilage. ‘The second arch is the 
longest, but is scarcely thicker than the others, neither is its dorsal end bifurcate or 
grooved. The remaining arches gradually decrease in length, the sixth being much 
the shortest of the series. According to Ridewood (J. ¢.), on the contrary, the second 
and third branchial arches differ from all the others, and resemble the branchial arches 
of Ceratodus, in that each is provided with a dorsal or “epibranchial” segment in 
addition to the relatively much longer ventral or cerato-branchial portion which, 
apparently, is alone represented in Wiedersheim’s figures. Meso- or basi-branchial 
cartilages appear to be wholly absent in Protopterus. 

The lower jaw is strikingly different from that of Ceratodus. The two rami are 
relatively shorter and more strongly developed, with a remarkably high coronoid 
process, and uniting anteriorly in a massive symphysis. As in Lepidosiren, practically 
the whole of the osseous portion of each ramus is formed by the splenial, the dentary 
being absent altogether, and the angular reduced to an insignificant splint on the outer 
surface of the articular extremity. The Meckelian cartilages unite anteriorly in a 
vertically-disposed symphysial plate, terminating above in a median and two lateral 
processes. According to Wiedersheim (J. ¢c.), a short ascending process is given off 
from each cartilage and occupies the angle between the anterior and central tooth- 
plates. I notice, however, that in one of Peters’s figures of his Zambesi Protopterus 
the Meckelian cartilage is represented as giving off an additional process behind the 
one referred to above, and, moreover, both are represented as if they were suturally 
distinct from the cartilage itself (J. c. Taf. ii. fig. 5). The additional process is 
apparently represented by a free nodule of cartilage in Lepidosiren (Pl. XXVIII. 
fig. 7). 

In Protopterus the “ cranial rib”? has become displaced from its normal relations to 
the elements of the first neural arch, and moved forward to an articulation with the 
chondrocranium between the Vagus foramen and the exoccipital. 

The foramina for the major divisions of the cranial nerves have the same general 
arrangement as in Lepidosiren. It need only be mentioned that each Optic nerve 
escapes through a small foramen in the fibrous sheet which closes the lateral cranial 
vacuity of its side, and that the foramen for the Motor Oculi perforates the side-wall 
of the skull at some distance behind the Optic nerve, but anteriorly to the aperture for 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 363 


the ramus ophthalmicus of the Fifth nerve. The roots of the first two spinal nerves 
are transmitted through two separate foramina in the exoccipital bone. 

The dorsal extremity of the pectoral girdle is connected with the occipital region of 
the skull by a stout fibrous band, in which is embedded what Parker [32] describes as 
‘a thin, subfalcate, subcutaneous bone” (J. c. p. 21), and which he designates the “ post- 
temporal.” The bone is relatively much smaller than in Ceratodus, and has apparently 
lost its direct articulation both with the skull and with the supraclavicle. 


C. Lepidosiren. 

In the main the skull of Lepidosiren closely resembles that of Protopterus. The 
most important of its structural peculiarities may be briefly summarized as follows :— 

The chondrocranial portion of the skull exhibits further indications of atrophy, 
especially in the occipito-periotic region. The cartilaginous roof, which in Protopterus 
extends as far forward as the anterior limit of the auditory capsules, has receded 
somewhat in Lepidosiren and is now restricted to the supraoccipital region. The 
exoccipitals are invested neither externally nor internally by cartilage, and hence the 
complete occipital ring of cartilage in Protopterus becomes divided in Lepidosiren into 
a dorsal and a basal plate, which are only connected through their continuity anteriorly 
with the periotic capsules. The trabecular cartilages are considerably reduced in 
thickness and in height, and are now represented by relatively slender, laterally- 
compressed bands, slightly increasing in vertical extent as they become continuous 
with the periotic and suspensorial cartilages, but only to a very slight extent do they 
become visible from the interior of the cranial cavity. ‘The styliform process of the 
mesethnoid cartilage is also shorter than in Protopterus. On the other hand, the 
antorbital cartilages are somewhat better developed, and, in addition to supporting the 
inferior and hinder margin of the upper labial folds, are prolonged into anteriorly- 
directed processes. 

The suspensorial cartilage is essentially similar in structure to that of Protopterus, 
except for the emargination of that part of its posterior margin which is covered 
externally by the squamosal. Undoubtedly its most interesting feature is the rotation 
backward of its articular extremity to a greater extent than in any other Dipnoid, and 
hence its forward inclination is less, and the angle which it makes with the fore part 
of the basicranial axis greater, than in either Ceratodus or Protopterus. The effect of 
this recession of the condylar extremity of the suspensorium on the palato-pterygoid 
bone is that the latter is not only of greater relative length, but deviates more from 
the vertical, and describes a segment of a larger circle in passing forward to its 
symphysial termination, than in either of the other two genera. 

The two exoccipital bones are distinct, but nevertheless closely approximated both 
dorsally and ventrally. 


364 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


The fronto-parietal is a relatively much larger bone in every dimension than in 
Protopterus. In the anterior part of its extent not only does the bone completely 
invest the dorsal surface of the cranium, but through the extension of its lateral or 
descending plates as far forward as the ascending processes of the palato-pterygoids, 
and posteriorly as far back as the periotic capsules, the lateral walls of the entire pre- 
auditory section of the cranial cavity are formed wholly by bone. Hence it follows 
that the lateral cranial vacuities of Protopterus have no existence in Lepidosiren, 
or at most are only represented on each side by a narrow suture, or rather fissure, 
between the ascending process of the palato-pterygoid and the anterior and lateral 
margins of the fronto-parietal, while the more or less extensive area of trabecular 
cartilage which, in a vertical longitudinal section of the skull of Protopterus, is visible 
in front of the periotic capsule, becomes entirely concealed from view in Lepidosiren. 
More posteriorly, in the suspensorial region, the fronto-parietal of Protopterus is 
confined to the cranial roof, and, except for its partial investment by the squamosal, 
the whole extent of the suspensorial cartilage and its continuity with the periotic 
capsule and the trabecular cartilage are clearly visible in an external and lateral view. 
In Lepidosiren, on the contrary, a lateral and downward extension of the bone almost 
completely invests the outer surfaces of the periotic and suspensorial cartilages, and, in 
a similar view, effectually hides the continuity of the latter with the trabecular 
cartilage. The sagittal crest is somewhat better developed in Lepidosiren than in 
Protopterus, and to this may be added a “lambdoid” ridge, which, owing to the 
relative narrowness of the hinder portion of the fronto-parietal, is scarcely indicated in 
the latter Dipnoid. 

The dermal ectethmoids are scarcely so well developed as in Protopterus. Their 
anterior portions are much narrower and fail to meet in a median suture: hence the 
fronto-parietal and its characteristic crest may be seen between them in a dorsal view 
of the skull. (Compare Pl. XXVIII. fig. 2, and Wiedersheim, J. c. Taf. ii. fig. 1.) 

Lepidosiren has but five simple, unsegmented, branchial arches, the sixth arch, like 
the fifth cleft, having been suppressed. As compared with Protopterus, a noticeable 
feature is the exceptional thickness of the second arch, and its dorsal cleft or groove 
for an aortic arch. 

As previously mentioned, the position and relations of the foramina for the exit of 
the cranial nerves are nearly identical in the two genera. It may be pointed out that 
the Optic nerves in Lepidosiren escape through the two clefts or fissures which 
represent the anterior cranial vacuities of Protopterus, and further that in the former 
Dipnoid only the roots of the first spinal nerve (Hypoglossal) perforate the exoccipital, 
the second pair simply traversing the fibrous tissue which intervenes between that bone 
and the first neural arch. 


. The distinct epibranchial elements of the second and third arches in Protopterus are wholly unrepresented 
in Lepidosiren 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 365 


The connection of the pectoral girdle with the postero-lateral regions of the skull is 
entirely by ligament, the “ post-temporal” bone of Ceratodus and Protopterus being 
wholly unrepresented. 

From what has been stated as to the structural differences and resemblances between 
the skulls of the three genera of existing Dipnoi it may be inferred that, while all 
three conform to the same fundamental type of structure, and at the same time exhibit 
successive stages of modification in the same general direction, the skull of Ceratodus 
is by far the most generalized and primitive ; and further, that of the two remaining 
genera the skull of Lepidosiren represents but a slightly more specialized type when 
compared with the skull of Protopterus. In support of the first statement it is only 
necessary to recall the more complete development of the chondrocranium in Ceratodus, 
especially in the ethmo-nasal, interorbitai, and palato-pterygoid regions, the absence of 
a fronto-parietal bone, the feeble development of the exoccipitals, the segmentation of 
the branchial arches, the more complete hyoid arch with its hyomandibular, hypohyal, 
and basihyal elements, the presence of a dentary splint in the lower jaw, and also 
the retention of a series of suborbital ossicles. On the other hand, Protopterus and 
Lepidosiren differ from Ceratodus and agree with each other in the extensive atrophy 
of the central region of the chondrocranium, the absorption of the palato-pterygoid 
cartilage, the characteristic constriction of the nasal from the ethmoidal region through 
the massive growth of the palatine symphysis, the development of a fronto-parietal 
bone, and the increased size and importance of the exoccipitals. It is none the less 
obvious, however, that in both genera specialization due to the increased development 
of certain cranial structures has proceeded side by side with the reduction or suppression 
of others. Of the latter process we have examples in such modifications as the atrophy 
of the dentary element of the lower jaw and of the suborbital bones, and the reduction 
in the hyoid arch by the loss of its hyomandibular, hypohyal, and basihyal elements, 
while the substitution of lungs for gills as the more important respiratory organs is 
clearly responsible for the reduction in size and simplification of the branchial arches, 
the closure of the hyoidean cleft, and the relatively smaller size of the opercular bones. 
Finally, the slightly more pronounced specialization of the skull of Lepidosiren is 
evidenced by the further atrophy of the chondrocranium and the compensating increase 
in the size of the fronto-parietal and in the extent of its relations to the cranial and 
suspensorial regions. To. these features may be added the increased backward rotation 
of the suspensorial cartilage, the somewhat greater development of the antorbital 
cartilages, and the total suppression of the sixth branchial arch and of the epibranchial 
elements of the second and third arches. Briefly, it may be said that if the skull of 
Ceratodus be taken to represent a relatively early larval stage, the skulls of Protopterus 
and Lepidosiren are comparable to two immediately succeeding and very much later 
stages, while, with one or two exceptions, the differences between the two latter 

you. XIv.—PartT Vv. No. 6.—February, 1898. 3B 


366 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


genera are much the same in nature and extent as those which distinguish the skull 
of first- and second-year Frogs. 

It cannot, however, be too strongly emphasized that in any comparison of the skulls 
of Protopterus and Lepidosiren it is important that due regard should be paid to the 
size and age of the specimens upon which such comparison is based. According to 
Ginther [12, p. 357] Protopterus sometimes attains a length of six feet, whereas most 
of the specimens which have reached Europe, and furnished the needful material for 
the hitherto-published papers on the structure of the skull, seem to have been com- 
paratively small and relatively young examples in which it is improbable that the skull 
had attained its final and complete development. Hence it is by no means improbable 
that, if comparison were made with larger and older specimens of Protopterus, some at 
least of the cranial modifications in Lepidosiren, which have been mentioned as 
distinctive of the genus, would either disappear or become so far softened down as to 
lose much of whatever importance they now seem to possess. ‘That this is probable is 
evident from the fact that certain of the more obvious differences in the cranial structure 
of the two genera are precisely those which might easily be bridged over by older 
specimens of Protopterus. On this point it is only necessary to refer to the greater 
development of the fronto-parietal, the further retrogression of the chondrocranium, 
and the greater backward rotation of the suspensorium in Lepidosiren. 


IV. The Structure of the Skull in Fossil Dipnoi. 


When the range of comparison is so far extended as to include the extinct Dipnoid 
families, the Dipteride, Phaneropleuride, and Ctenodontide!, it is obvious that, while 
the fossil types exhibit a general agreement with existing Dipnoi in the main outlines 
of their cranial structure, they differ from the latter, as well as from one another, in 
several striking and significant features. Broadly speaking, it may be affirmed that 
all known fossil Dipnoi agree with their living representatives and differ from all 
other Fishes in possessing the following combination of cranial characters :— 


(A.) Complete and typical autostylism. 

(B.) The presence of characteristic triturating palatal teeth, supported by palato- 
pterygoid bones, symphysially united beneath the ethmoidal region of the 
skull, and probably associated with the absence of premaxille and maxille 
in the upper jaw. 


1 Thave here followed the classification of the fossil Dipnoi as given by Smith Woodward [45]. Traquair 
[39 has suggested the separation of Wronemus from the Phaneropleuride as the type of a distinct family 
[ Uronemidz], and the union of the remaining genus Phaneropleuron with the Dipteridx and Ctenodontide in one 
comprehensive family of Ctenodontide. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 367 


(C.) The great development of the splenial bones of the lower jaw for the sole 
support of mandibular teeth, similar to the palatal teeth of the upper jaw, 
and their union anteriorly in a stout mental symphysis, to which may be 
added the absence of dentigerous dentary elements. 

(D.) The presence of only two opercular bones—an operculum and an interoper- 
culum—and, unless represented by the squamosal, the absence of a distinct 
preopercular element. 

Apparently the only constant cranial character in which all fossil Dipnoi differ 
from existing Dipnoi is the multiplicity and almost Acipenseroid arrangement of their 
cranial roofing-bones!. These bones seem to be most numerous in Dipterus (D. valen- 
ciennesi). A medio-dorsal longitudinal series can be distinguished, flanked on each 
side by several rows of longitudinally-disposed lateral plates. The inner lateral row 
of plates on each side tend to meet in pairs between the median plates, so as to 
interrupt the sutural continuity of the latter. A similar tendency to alternation may 
be also exhibited by the lateral series of plates, but their arrangement is far from 
regular, and is not always symmetrical on opposite sides of the cranial roof. There 
is a general similarity to Dipterus in the disposition of the cranial plates in other 
extinct Dipnoi, and it is worthy of note that in Ctenodus the plates become somewhat 
reduced in number, and two of the median series, the most anterior and the most 
posterior, become so far enlarged as to be distinguishable as a dermal ethmoid 
and a dermo-supraoccipital [ Fritsch, 9]. But, except in the most general manner, it is 
practically impossible to correlate any of these plates with the numerically-reduced 
and more definitely-disposed cranial roof-bones of the average Ganoid or Teleostean 
skull. To what extent the dermal plates are represented in living Dipnoi it is 
extremely difficult to say. Dollo [7] regards the sparsely-represented dermal cranial 
bones of existing types as the remains of the much more numerous plates of the fossil 
Dipnoi. If this be so, the most anterior of the medio-dorsal series of plates in the 
latter, and a solitary pair of the lateral plates in the orbital or ethmoidal region, may 
be represented in Ceratodus, Protopterus, and Lepidosiren by the median dermal 
ethmoid and the paired dermal ectethmoids or lateral ethmoids respectively. On the 
other hand, if the “scleroparietal” of Ceratodus is a ‘“‘ tendon-bone,” it probably has 
no counterpart in any fossil Dipnoid; and as the fronto-parietal of Protopterus and 
Lepidosiren is situated wholly internal to the jaw-muscles, which could scarcely 
have been the case with any of the cranial plates of Dipterus and its allies, the same 
conclusion may be suggested with regard to this bone. 

There are also other differences in cranial structure, which, if not distinctive of all 
fossil Dipnoi, are nevertheless characteristic of particular families or genera, and, so far 
as our knowledge at present extends, the more important of these may be briefly reviewed. 


' See remarks by Smith Woodward on Ctenodus [45, p. 253]. 
3B2 


368 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


i. The existence of endochondrial ossifications.—There is not wanting evidence that 
in some fossil forms the cartilaginous elements of the skull must have been replaced 
by bone to an extent which has no parallel in existing types. Thus, Traquair, who 
was the first to demonstrate the autostylic character of the skull in the Dipteridz and 
thereby conclusively prove the accuracy of the earlier suggestion of Giinther [11] as to 
the relationship of this family to the Dipnoi, in his important paper on the skull of 
Dipterus [37], remarks “that the chondrocranium was very much more extensively 
ossified than that of Ceratodus; in fact, its side-walls were entirely occupied by 
bone apparently as far as the interorbital region.” ‘The bony matter surrounding 
the foramen magnum may be held to represent the exoccipitals, and in front of 
it on each side the walls of the otic region are distinctly ossified—though, from 
the abraded condition of this part in all the skulls, it is hardly possible to trace 
any sutural lines marking off distinct osseous elements” (/. c. p. 5). There is also said 
to be a distinct quadrate provided with a facet for articulation with the mandible. 
“In fact, we have here before us, as nearly as possible, a counterpart in bone of the 
suspensorial cartilage of Ceratodus” (1. c. p. 6). Further, in the same Dipnoid there 
is an ossified articular element in the lower jaw (Traquair, /. c.), in addition to the usual 
angular and splenial bones. 

There is no satisfactory evidence of the existence of endochondrial bones in any other 
extinct Dipnoi, with the possible exception of the bone termed “ Gehérkapsel ” in 
Ctenodus (Fritsch, 1. ¢.), and doubtfully regarded as a periotic ossification. In the 
latter Dipnoid, according to Fritsch, there is an ossified cerato-hyal and a basihyal 
and, in addition, certain bones which he conjectures may represent the cerato-branchial 
elements of some of the branchial arches. 

ii. The presence of additional dermal bones in connection with the upper and lower 
jaws.—Of these we have examples in the median dentary at the mandibular (splenial) 
symphysis of Dipterus (Traquair, J. c.), and the premaxille and maxille which support 
the pointed conical “‘ marginal” teeth of Phaneropleuron (Traquair, 38), and, in addition, 
the several bones which, in the lower jaw of the same Dipnoid, carry similar teeth. 
The presence of ‘‘ marginal teeth” and supporting dermal bones in this genus must, 
however, be regarded as at least open to doubt, although usually cited as one of the 
distinctive characters of the Phaneropleuride. 

In a paper subsequent to the one quoted above, Traquair [39] remarks of Phanero- 
pleuron andersoni:—* As for the conical teeth described by Huxley, I have satisfied 
myself that they are merely tne outer aenticles of ctenodont plates. Whiteaves’s 
statement that in Scawmenacia curta ‘both the upper and under jaws are armed with 
smooth, conical, and somewhat compressed teeth,’ I have never been able to confirm— 
at least, if marginal teeth are hereby meant” (J. c. p. 264). 

ili. The presence of jugal plates—Dipterus has two pairs of such plates—an anterior 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 369 


and a posterior pair. Jugal plates are also present in Phaneropleuron, but are said to 
be absent in Ctenodus }. 

iv. The presence of circumorbital ossicles—In Dipterus there is series of bony plates 
encircling the orbit and covering the cheek, and also forming the upper border of the 
hinder part of the mouth [Traquair, 37]. According to Fritsch [/. c.], cireumorbital 
ossicles are probably present in Ctenodus, but of their existence in Phaneropleuron I 
have been unable to find any evidence. 

v. The squamosal bone.—This element is certainly present in Ctenodus, and has been 
described and figured by both Miall [23] and Fritsch (/. ¢.) as being very similar to its 
homologue in Ceratodus. As to the character or even the presence of this bone in 
the Dipteride and Phaneropleuride I can find no evidence. 

vi. The parasphenoid.—The parasphenoid of Dipterus and Ctenodus is known, but 
not, so far as I am aware, in Phaneropleuron. It has been described and figured in 
the first-mentioned genus by Traquair [37], and in Ctenodus by Barkas [1], Miall [23], 
and Fritsch (/. ¢.). In both Dipterus and Ctenodus the bone extends as far forward 
as the palatine symphysis, thus filling up the whole space bounded anteriorly and 
laterally by the palato-pterygoid bones, and in this respect these genera resemble 
Ceratodus, while differing from Protopterus and Lepidosiren. In Dipterus the para- 
sphenoid seems not to have extended beyond the posterior limit of the skull, but 
in Ctenodus, as Barkas (/. ¢. p. 51) pointed out, the exceptional length of the bone 
suggests that it extended for some distance beneath the vertebral column, thus 
affording an additional point of agreement to the many which are apparent when the 
skulls of Cfenodus and Ceratodus are compared. 

vii. The opercular bones.—There is perhaps some variation in the relative size of the 
opercular bones in different extinct Dipnoi, but it is certain that they are usually 
much larger than in any existing members of the group. 

viii. The presence-of ganoin.—As an investment to more or fewer of the skull-bones, 
the presence of ganoin is most evident on the outer surface of the cranial roofing-bones, 
the opercular bones, and the lower jaw of the Dipteride. In Ctenodus and Phanero- 
pleuron this characteristic investment is either entirely absent or restricted to fewer 
bones, at all events so far as the cranial elements are concerned. 

As to the question whether the fossil Dipnoi are to be regarded as more primitive 
than the living Dipnoi there is considerable difference of opinion. According to 
Smith Woodward [45], the former are more specialized than any existing Dipnoi, 
and the more generalized types have alone survived to represent the group at 
the present day. In an elaborate and suggestive essay ‘Sur la Phylogénie des 
Dipneustes,’ Dollo [7] has arrived at exactly opposite conclusions. According to his 
views, Dipterus (D. valenciennesi) is to be considered the most primitive Dipnoid, the 


1 Traquair [39, p. 265] has expressed a doubt as to the assumed absence of jugal plates in this genus. 


370 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


remaining genera—Scaumenacia, Phaneropleuron, Uronemus, and Ctenodus, in the 
order mentioned—representing successive grades of modification in the same general 
direction, and culminating in the living genera Ceratodus, Protopterus, and 
Lepidosiren as the final terms of an evolutionary series. It is not suggested that the 
sequence of the genera represents descent in a direct line, but rather that the genera 
are to be regarded as lateral offshoots of the main line of descent, while at the same 
time illustrating the general direction of the evolution of the group. Commencing 
with Dipterus valenciennesi, the principal modifications in cranial structure which the 
different genera exhibit in ascending order are (a) reduction in the number of the 
cranial dermal bones; (4) the gradual loss of their investment of ganoin; (¢) the 
suppression of the jugal plates; and (d) reduction in the size and importance of the 
opercular elements ?. 

Viewed from the standpoint of cranial structure alone, there are grave difficulties 
which militate against, the acceptance of at least one, and that perhaps one of the 
most important, of Dollo’s conclusions. So far from Dipterus being the most primitive 
of known Dipnoi, it is obvious, as Traquair [37] has pointed out, that in some of its 
cranial features this genus has attained a higher grade of specialization than any 
existing Dipnoid. In proof of this it is only necessary to refer to the more extensive 
ossification of the chondrocranium, the presence of a quadrate bone as the functional 
suspensorium, and the existence of an articular element in the lower jaw, all of which 
are features without parallel in any of the living genera. If we admit the accuracy 
of Dollo’s view, that Dipterus represents the most primitive type of the Dipnoid stock, 
it will be necessary to assume the possibility of an ossified skull so far degenerating 
as to lose almost all trace of endochondrial ossification, and secondarily revert to 
the condition of a skull so completely cartilaginous, and so primitive in other respects, 
as that exhibited by the living Ceratodus. So far as 1am aware, there is no evidence 
to justify belief in such a possibility. 

As to the relations of the fossil Dipnoi to one another and to their living allies, it is 
difficult to arrive at any satisfactory conclusion from the facts of cranial structure 
alone, and the difficulty is increased by the want of definite and precise information on 
many important points in connection with the fossil Dipnoi, more particularly as to 
the condition of the chondrocranial portion of the skull. On the evidence at present 
available it seems probable that Ceratodus is the most primitive of known Dipnoi, and 
that Protopterus and Lepidosiren are the specialized and direct descendants of some 
Ceratodus-like ancestor. Of the fossil Dipnoi, Ctenodus certainly, and possibly 
also Phaneropleuron, approach more closely to Ceratodus than do Dipterus and its 
allies. The last-mentioned Dipnoi, on the contrary, seem to represent a divergent 


and terminal branch of the Dipnoid stem, and to include the most highly-specialized 
examples of the group. 


» Tt must be mentioned that Dollo’s comparison is not restricted to the skull, but includes also the mesial 
and paired fins, the scales and other structures of the various fossil and living genera. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 571 


In the present communication it is not intended to refer to the obscure problem of 
the phylogenetic origin of the Dipnoi themselves. An exhaustive and suggestive 
discussion of this question is to be found in the previously-cited paper by Dollo. ‘Uhere 
is, however, one point which may be referred to as having a direct bearing on this 
problem. 

In common with several other morphologists, Dollo regards the Crossopterygii as the 
ancestral stock from which the Dipnoi have been derived, and bases his conclusion on 
the prevalence in certain fossil Dipnoi of characters common to the two groups, such 
as, for example, the existence of lobate paired fins, the presence of jugal plates, the 
ventral position of the narial apertures (in some Crossopterygii), and the ganoid 
investment of certain of the cranial bones. The obvious difficulty presented by 
the hyostylism of the Crossopterygii, and the pronounced autostylism of the Dipnoi, is 
met by the bold suggestion that the latter is an adaptive modification associated with 
the development of the peculiar Dipnoid type of dentition, which has also conditioned 
the suppression of the usual dentigerous bones, the premaxille, the maxille, and the 
dentaries. In support of this suggestion, Dollo emphasizes the association of auto- 
stylism with massive palatal teeth in the Holocephala and the incipient autostylism of 
Cestracion with the cochliodont dentition of that Selachian. 

With regard to Dollo’s theory of the adaptive character of autostylism, and its 
independent acquisition by widely different groups of Fishes, it may be remarked that 
the autostylism of the Cyclostomes and Amphibia still remains as a difficulty in the 
way of its acceptance, inasmuch as in neither group can the prevalence of autostylism 
be explained as due to the peculiarities of an exceptionally-developed dentition. It is 
nevertheless quite possible, and in my opinion even probable, that other factors besides 
peculiarities of dentition may condition the evolution of autostylism, and one of these 
may be the development of a suctorial mouth supported by a more or less elaborate 
system of labial cartilages. In the latter case the advantage derivable from the fixation 
by autostylism of the primary skeletal elements of the upper jaw may conceivably 
be quite as great as that gained in the case of a massive dentition. But, apart from 
the reasons advanced by Dollo, there are other considerations which suggest the 
derivation of the autostylism of the Dipnoi from a hyostylic type of skull. 

As shown by Huxley [15], the upper extremity of the hyoid arch (cerato-hyal) in 
Ceratodus is connected with the suspensorial cartilage through the intervention of a 
cartilaginous element which he regards as the representative of the hyomandibular 
bone or cartilage of other Fishes. A similar interpretation may perhaps be assigned to 
the upper division of the hyoid arch in Chimera, which, like the hyomandibular of 
Elasmobranchs, carries the dorsal series of hyoidean rays. Assuming the correctness 
of this interpretation of the nature of these cartilages, and that they are really homo- 
dynamous with the hyomandibular element cf the majority of Fishes, an obvious 
inference is at once suggested. In practically all existing Elasmobranchs and 
Teleostomi, or at all events with only a single exception (Notidanus), the hyo- 


372 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


mandibular forms the common suspensorium of the mandibular and hyoid arches, 
and is associated with a hyostylic condition of the skull. The small size of the 
hyomandibular element in Ceratodus may be due to its reduction to the condition of a 
purely vestigial structure, but none the less does the noteworthy fact of its retention in 
the most primitive of existing Dipnoi suggest that it may be regarded as a remnant 
of the primitive hyostylic condition of the skull in the ancestors of the Dipnoit. A 
similar significance may also be attached to the hyomandibular of Chimera, at all 
events to the extent of suggesting the origin of the Holocephala from hyostylic 
ancestors. For these reasons I am inclined to agree with Dollo that autostylism is a 
purely adaptive modification, and may occur independently in diverse groups of Fishes 
wherever any advantage is to be gained from the fixation by fusion to the skull of the 
primitive elements of the upper jaw (palato-quadrate cartilage) for the purpose of 
providing the needful support for a massive and peculiar dentition, or even, as I have 
suggested above, for a system of labial cartilages in a suctorial mouth. 

While agreeing with Dollo as to the adaptive character of autostylism, I may add 
that this does not necessarily imply agreement with his views as to the phylogenetic 
origin of the Dipnoi from the Crossopterygii. The theory is eminently controversial, 
and, although the author has much to say in its favour, there are nevertheless, in 
my opinion, weighty objections to its implicit acceptance, as well as no less cogent 
reasons for assigning to the Dipnoi a more remote ancestry. My object in referring 
to this question is rather to point out that autostylism in itself must not necessarily 
be regarded as an indication of genetic affinity. The necessity for caution in this 
respect is illustrated by certain recent views as to the supposed affinity of the 
Arthrodira (Coccosteus, Dinichthys, and their allies) to the Dipnoi. Thus, Smith 
Woodward [45], in discussing the affinities of the latter group, remarks: “ the evidence 
in favour of the autostylic character of the Coccostean Fishes has now accumulated to 
such an extent that we venture to regard them as an order of Dipnoi” (/. c. p. xxi). 
It is quite possible that further research may confirm the accuracy of these views, but 
for the present I venture to think that their ultimate proof must rest on other grounds 
than the occurrence of autostylism. 


’ With reference to this point the following remarks by Gadow [9 a, p. 459] may be quoted: “It is highly 
probable that in the ancestral Dipnoi the hyomandibula was much larger, and that it was already, as in 
Teleosteans, broken up into a proximal cranial persisting part, and into a distal or symplectic element, which 
later on, when the hyostylic support of the jaws was superseded, either persisted [ Proteus, &c.], or ultimately 
became lost [ Ceratodus, Salamandrina].” 


(Sy) 
=I 
(sh) 


IN THE PARAGUAYAN LEPIDOSIREN, ETC. 


V. REFERENCES TO PREVIOUS LITERATURE. 


1. Barkas, J. W.—On the Sphenoid, Cranial Bones, Operculum, and supposed Ear-bones of 
Ctenodus. Jour. Roy. Soc. New South Wales, 1877, p. 51. 

2. Biscuorr, T. H. W.—Lepidosiren paradoxa, anatomisch untersucht und _beschrieben. 
Leipzig, 1840. ‘Transl. in Ann. d. Se. Nat. vol. xiv. 1840, p. 116. 

3. Bripez, T, W.—On the Osteology of Polyodon folium. Phil. Trans. Roy. Soc. Lond. vol. 169, 

1878, p. 683 
4. Brinu, C. B.—Anfangsgriinde der vergleichenden Anatomie aller Thierklassen. Wien, 1847, 
p- 216, Atlas, tab. xviii. 

. Burcxuarpt, R.—Das Centralnervensystem von Protopterus annectens. Berlin, 1892. 

. Cossorp, T. S.—On the Cranial Bones of Lepidosiren annectens. Proc. Zool. Soc. Lond. 
1862, p. 129. 

7. Dotto, Lovis.—Sur la Phylogénie des Dipneustes. Bull. de la Société Belge de Géologie, 
tom. ix. 1895, p. 79. 

8. Enters, E.—On Lepidosiren paradoxa, Fitzinger, and Lepidosiren arcuata, sp. n., from 
Paraguay. Ann. & Mag. Nat. Hist. ser. 6, vol. xiv. 1894, p.1. (Translated from the 
Nachrichten der k. Gesellschaft der Wissenschaften zu Gottingen, 1894.) 

g. Frrrscu, A.—Fauna der Gaskohle und der Kalksteine der Permformation Bohmens, Bd. ii. 
Heft 3, p. 56. 

ga. Gavow, H.—On the Modifications of the First and Second Visceral Arches, with special 
reference to the Homologies of the Auditory Ossicles. Phil. Trans. Roy. Soc. Lond. 
vol. 179 B, 1888, p. 451. ; 

10. GecenBaur, C.—Untersuchungen zur vergleichenden Anatomie der Wirbelthiere. Leipzig, 
1872. 

11. Ginraer, A.—Description of Ceratodus, a genus of Ganoid Fishes recently discovered 
in rivers of Queensland, Australia. Phil Trans. Roy. Soc. Lond. vol. 161, 1871, 
p. 511. 

12. Inpem.—The Study of Fishes. Edinburgh, 1880. 

13. Humpury, G. M.—The Muscles of Lepidosiren annectens, with the Cranial Nerves. Jour. 
Anat. & Phys. vol. vi. 1872, p. 253. 

14. Huxiny, T. H.—Manual of the Anatomy of Vertebrated Animals. London, 1871, pp. 168- 
170. 

15. Inpm.—Contributions to Morphology.—Ichthyopsida. No. 1. On Ceratodus fosteri, with 
Observations on the Classification of Fishes. Proc. Zool. Soc. Lond. 1876, p. 24. 

16, Inem.—On the Position of the Anterior Nasal Apertures in Lepidosiren (Protopterus annectens). 
Proc. Zool. Soe. Lond. 1876, p. 180. 

17. (pem.—On Menobranchus lateralis. Proc. Zool. Soe. Lond. 1874, p. 186. 

18, Hyrrn, J.—Lepidosiren paradoxa. Monographie. Abhand. d. Bohm. Gesellsch. Bd. i. 1845, 
p: 605. 

19. Iversen, M.—Bemerkungen iiber die dorsalen Wurzeln des Nervus hypoglossus. Ber. der 
Naturforsch. Gesellsch. zu Freiburg, i. B. 18938, Bd. vii. Heft 2. 

20. Lanxester, E. Ray.—On the Lepidosiren of Paraguay and on the External Characters of 
Lepidosiren and Protopterus. Trans. Zool. Soc. Lond. vol. xiv. pt. 1. 1896, p. 11. 
VOL. XIV.—PaRT v. No. 7.—February, 1898. 3 ¢ 


An 


574 PROF. T. W. BRIDGE ON THE MORPHOLOGY OF THE SKULL 


21. McDonnett, R.—Observations on the Habits and Anatomy of the Lepidosiren annectens, 
Roy. Dublin Soe. Jour. ii. 1858-59, p. 388 ; also Nat. Hist. Review, Lond. vol. vii. 1860. 


p: 93. 
22. Miaru, L. C_—Monograph of the Sirenoid and Crossopterygian Ganoids. Palwontogr. Soc. 
Lond. 1878. 


23. Ipem.—On some Bones of Ctenodus. Proc. Geolog. & Polyt. Soc. W. Riding, Yorkshire, 
vol, vii. p. 289. 

24. Narrerrr, J.—Lepidosiren paradoxa, Ann. d. Wiener Museums, Bd. ii. 1839, p. 167, tab. x. 

25. Owrn, Ricuarp.—Description of the Lepidostren annectens. Trans. Linn. Soc. Lond. 
vol. xviii. 1839, p. 327. 

26. Ipem.—Comparative Anatomy and Physiology of Vertebrates, vol. i. 1866, p. 82. 

27. Parker, W. K.—On the Structure and Development of the Skull in the Salmon (Salmo salar, 
L.). Phil. Trans. Roy. So¢. Lond. vol. 163, 1873, p. 95. 

28. Ipzem.—On the Structure and Development of the Skull in the Urodelous Amphibia.—Part TI. 
Phil. Trans. Roy. Soc. Lond. vol. 167, 1876, p. 529. 

29. Ipem.—On the Structure and Development of the Skull in the Sharks and Skates. ‘Trans. 
Zool. Soc. Lond. vol. x. 1878, p. 189. 

30. Inem.—On the Morphology of the Skull in the Amphibia Urodela. ‘Trans. Linn. Soc. Lond. 
Series 2, Zoology, vol. ii. 1879, p. 165. 

31. Ipzm.—On the Structure and Development of the Skull in the Batrachia—Part. III. Phil. 
Trans. Roy. Soc. Lond. vol. 172, 1881, p. 1. 

32. Iprm.—Structure and Development of the Shoulder-girdle and Sternum in the Vertebrata. 
Ray Society, 1868. 

32a. Inpm.—On the Structure and Development of the Skull in Sturgeons (Acipenser ruthenus 
and A. sturio). Phil. Trans. Roy. Soc. vol. 173, 1882, p. 139. 

33. Parker, W. N.—On the Anatomy and Physiology of Protopterus annectens. Trans. Roy. Ivish 
Academy, vol. xxx. 1892, p. 109. 

34. Permrs, W.--Ueber einem dem Lepidosiren annectens verwandten Fisch yon Quellimane. 
Archiv fiir Anatomie, Phys., und wissenschaftliche Medicin [Johannes Miiller], 1845, 
pp- 1-14. 

35. Pinxus, F.—Die Hirnnerven des Protopterus annectens. Abdruck aus den morphologischen 
Arbeiten, Bd. iv. Heft ii. p. 275. 

35 4. Riprwoop, W. G.—On the Hyoid Arch of Ceratodus. Proc. Zool. Soc. Lond. 1894, p. 632. 

36. Rose, Cari.—Ueber Zahnbau und Zahnwechsel der Dipnoer. Anatom. Anzeiger, Jena, 1892, 
p- 821. 

37. Traquair, R. H.—On the Genera Dipterus, Sedg. & Murch., Paledaphus, Van Beneden und 
De Koninck, Holodus, Pander, and Cheirodus, M‘Coy. Ann. & Mag. Nat. Hist. ser. 5 
vol. 11. 1878, p. 1. 

38. Ipem.—On Phaneropleuron andersoni [Huxley], and Uronemus lobatus [Agassiz]. Jour. 
Geolog. Soe, Ireland, n. s. vol. iii, 1871, p. 41. 

39. Inem.—Notes on the Devonian Fishes of Campbelltown and Scaumenac Bay in Canada, No. 3. 
Gecl. Mag. 1893, p. 262. 

40. Van Wisner, J. W.—Ueber das Visceralskelet und die Nerven des Kopfes der Ganoiden und 
von Ceratodus. Niederlindisches Archiv fiir Zoologie, Bd. v. Heft 3, 1882, p. 207. 

41. WiepEeksuem, R.—Das Skelet und Nervensystem yon Lepidosiren annectens {Protopterus 
ang.] Morphologische Studien, Heft i. Jena, 1880, p. 45. 


42. Wirprrsueim, R.—Salamandrina perspicillata und Geotriton fuscus. 


IN THE PARAGUAYAN LEPIDOSIREN, ETC, 


43. Ippm.—Das Kopfskelet der Urodelen. 
44. Wiper, H. H.—Die Nasengegend von Menopoma alleghaniense und Amphiuma tridactylum, 
Zool. Jahrbiicher, Bd. v. Heft 2, 1892. 
45. Woopwarp, A. S.—British Museum Catalogue of Fossil Fishes, pt. ii. 1891. 


The cartilaginous portions of the skull are tinted blue. 
figures are of the natural size. 


Fig. 


bo 


have been removed. 


ig. 3. Ventral view. 


Leipzig, 1877. 


PLATE XXVIII. 


Lateral view of the skull of the Paraguayan Lepidosiren. 
View of dorsal surface. 


VI. EXPLANATION OF THE PLATES. 


1875. 


oo 
=I 
or 


Unless otherwise stated, the 


The lettering is uniform in both Plates. 


The cerato-hyal, cranial rib, and interoperculum of the left side 


is not shown. 
4, Vertical longitudinal section of the skull. 


pterygoid and fronto-parietal bones. 


+I 


. 8. Internal lateral view. 


External lateral view of the lower jaw. 


PLATE XXIX. 


. 9. External view of the mandibular symphysis. 


10. Branchial arches of the left side. 
Figs. 11-19. Tranverse sections through the skull at the various points indicated by 


dotted lines in fig. 21. 

. 11. Transverse section through A-—B in fig. 
ze 
Boley 
14. 
ig. 15. 
x NG: 
fig. Li. 
ig. 18. 
Fig. 19. 
Fig. 20. Transverse section through the nasal region of Ceratodus fosteri. 


” 


” 


C-D 
E-F 
G-H. 
I-J 
{—L 
M-N 
O-P 
Q-R 


3° 


bb) 


21. 


. 6. View of the periotic capsule from the interior of the cranial cavity. 


The vestigial hyoidean ray of the right interoperculum 


. 5. Dorsal view of the nasal region and the anterior portions of the palato- 
The ectethmoids have been removed. 


Fig. 21. Outline lateral view of the skull of the Paraguayan Lepidosiren to show the 


planes in which the transverse sections (figs. 11-19) have been taken. 
ectethmoids, opercular bones, and the hyoid arch have been removed. 


The 


376 ON THE SKULL IN THE PARAGUAYAN LEPIDOSIREN, ETC. 
REFERENCES TO LETTERING. 
a.m.t. Anterior mandibular tooth-plate. n.sp. Neural spine. 
an, Angular. op. Operculum. 
an.p. Antorbital process. p.c. Periotic capsule. 
an.p'. Labial portion of antorbital process. p-m.t. Posterior mandibular tooth. 
a.p.t. Anterior palatal tooth. pn.p. Prenasal process. 
ar, Articular surface of mandible. ppt. Palaio-pterygoid bone. 
ar.c. Articular extremity of suspensorial p-p.t. Posterior palatal tooth. 
cartilage. ppt.a. Ascending process of the palato-ptery- 
at.p. Antorbital process of the palato-ptery- goid. 
goid, p-pt.a'. Spur on the ascending process of the 
6.c. Cartilaginous “ basis cranii.”’ palato-pterygoid, to which the ect- 
b.c.v. Basicranial vacuity or cleft. ethmoid is attached. 
c.h. Cerato-hyal. ps. Parasphenoid. 
cor.p. Coronoid process of the mandible. p.t. Palatal tooth (Ceratodus). 
cr. “ Cranial rib.’” p-sy. Palatal symphysis. 
d.e. Dermal ethmoid. s.m.e. Spina mentalis externa. 
ec.e. Dermal ectethmoid. s.m.i. Spina mentalis interna, 
eo. Exoccipital. sn.p. Subnasal cartilage. 
jf-p. Fronto-parietal. sp. Splenial. 
hy.s.l. Hyo-suspensorial ligament. sg. Squamosal. 
i.n.s. Internasal septum. sg.c. Sagittal crest. 
i.op. Interoperculum. s.r. Saccular recess. 
Id.c. “ Lambdoid crest.” st.p. Styliform process. 
mk.c. Meckelian cartilage. sus.c. Suspensorial cartilage. 
m.m.t. Middle mandibular tooth, tr.c, Trabecular cornu. 
m.p.t. Middle palatal tooth. tr.c. Trabecular cartilage. 
ms.e. Mesethmoid cartilage. u.l.c. Upper labial cartilage. 
m.sy. Mandibular symphysis. ut.r. Utricular recess. 
n.a. First neural arch, v.hy.r. Vestigial hyoidean rays. 
n.c. Nasal capsule. vo.t. Vomerine teeth. 
no. Notochord. 1,2,... First, second, .., Branchial arches. 
n.s. Nasal sac. 
Foramina. 
3. Olfactory nerve. vii. Superior palatine branch of the Facialis. 
ii, Optic nerve. vii’. Ramus mandibularis internus (Facialis). 
i Motor oculi. ix. Glossopharyngeal. 
v Ramus ophthalmicusof the Trigeminus. x. Vagus, and internal jugal or anterior 
Vv Ramus maxillaris and the ramus man- cardinal vein. 
dibularis of the Trigeminus, the h.s.c. Ampullary extremity of the vertical 
Nervus Lateralis Facialis, and the semicircular canal and the horizontal 
ramus communicans between the canal. 
Facial nerve and the lateral branch in.c. Internal carotid artery. 
; of the Vagus. sp.n'. Dorsal and ventral roots of first spinal 
vp. Branch of ramus ophthalmicus pro- nerve (Hypoglossal). 
E fundus (Trigeminus). sp.n°. Dorsal and ventral roots of the second 
v". Branch of ramus mandibularis (Tri- spinal nerve, 
_ . geminus). tf. Temporal foramen (“Schlafengrube”). 
vil, Ramus hyomandibularis (Facialis). v.s.c. Vertical semicircular canals. 


replace Col AN of CINE 


\.-----. p.pe. 


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a 


p= =. ----= * 


MP Parker chromo hth. 


West, Newrnan_imp 


Skull of Lepidosiren. 


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4 


rans Loot oo. DIL II POLE 


3 / 
amet mort 4 
1 i 


ppt te. aa oe aS 
peta ¢ ; 
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F 


TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON 
(continued). 


VOLUME XII. (1886-1890, containing 65 Plates) . 


Parrl. (1886, containing 6 Plates) 


» 


» 


” 


”» 


PE] 


VOLUME XIII. (1891-1895, containing 62 Plates) . 


2. 


6S) Ont Ott eee 


10, 


(1886, containing 7 Plates) 
(1886, containing 2 Plates) 
(1886, containing 12 Plates) 
(1886, containing 5 Plates) 
(1887, containing 7 Plates) 
(1888, containing 8 Plates) 
. (1889, containing 3 Plates) 
. (1889, containing 7 Plates) 


Parrl. (1891, containing 6 Plates) 


(1891, containing 6 Plates) 


. (1891, containing 6 Plates) 
. (1892, containing 1 Plate). 
. (1893, containing 9 Plates) 
. (1893, containing 4 Plates) 


. (1893, containing 6 Plates) 


. (1894, containing 6 Plates) 
. (1894, containing 6 Plates) 


. (1895, containing 5 Plates) 


. (1895, containing 7 Plates and 


VOLUME XIV. 

Parr l. (1896, containing 2 Plates) 
2. (1896, containing 6 Plates) 
8. (1897, coutaining 9 Plates) 
4, (1897, containing 10 Plates) 
5. (1898, containing 2 Plates) 


Pes 


(1890, containing 8 Plates and 


» 


Title and Index) ,, 


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Title and Index) ,, 


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CONTENTS. 


VII. On the Morphology of the Skull in the Paraguayan Lepidosiren and in other 
Dipnoids. By Professor T. W. Briven, Sc.D. F.ZS., Mason College, 


Birmingham. (Plates XXVIII. & XXIX.) . . . . . . « « page 325, 


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VIII. On the Mammals obtained by Mr. John Whitehead during his recent Expedition 
to the Philippines. By OuprietD THomas. With Mield-notes by the Collector. 


Received May 19, 1897, read June 15, 1897. 


[Puares XXX.-XXXVI.] 


Mr. JOHN WHITEHEAD, whose exploration of Mount Kina Balu has already 
rendered him famous as a collector, has during the last three years been engaged in the 
exploration of the islands of the Philippine group, partly at his own expense, and 
partly at that of the “ subscribers to the Whitehead Fund,” to whose generosity the 
National Museum owes the donation of the whole of the specimens that the subscribers 
had a claim to under Mr. Whitehead’s agreement with them. 

As the exploration has been so remarkably successful, it is only fitting that an 
acknowledgment of their generosity should be appended to this account of the 
Mammals obtained during the expedition. ‘Their names are as follows :—Messrs. 
Matthew, James, and Andrew Arthur, the Duke of Bedford, Major Cooper Cooper, 
the late Mr. Alexander Dennistoun, Mr. John Dennistoun, the late Lady Huntingtower, 
the late Mr. Henry Seebohm, Mr. J. G. Sandeman, and Mr. J. T. Thomasson. 

The Philippine Islands, however rich in birds, have always previously been looked 
upon as a group very poor in Mammals, especially in comparison with the rich faunas 
of the other islands of the East Indian Archipelago. ‘This poverty was particularly 
evident in regard to really peculiar indigenous Mammals; for, with the exception of 
Phieomys cwmingi, scarcely a Mammal was known from the group other than 
members of widely-distributed genera, of which the Philippine species were either 
identical with or closely allied to Palawan, Bornean, or Celebean forms, 

Little, therefore, could have been expected from the expedition further than the 
discovery of a few fresh species of genera known to inhabit the group, and this, so far 
as regards the islands other than Luzon, is just what has occurred. But in the great 
northern island of the group Mr. Whitehead has made a most wonderful and un- 
expected discovery, that of a new and peculiar Mammal-fauna inhabiting the Luzon 
highlands, and, so far as is yet known, mostly isolated on a small plateau on the 
top of Monte Data, in the centre of Northern Luzon, at an altitude of from 7000 to 
8000 feet. 

VOL, X1V.— ArT vi. No. 1.—J/une, 1898. 5D 


378 MR. OLDFIELD THOMAS ON MAMMALS 


The plateau itself, as will be seen by Mr. Whitehead’s notes below, is of extremely 
small size, but in spite of this fact he obtained there specimens of the following 
remarkable series of animals :— 


Crocidura grayi. 

Felis domestica (feral). 

Paradoxurus philippinensis. 

Celenomys silaceus. New genus and species. 
Chrotomys whiteheadi. New genus and species. 
Rhynchomys soricoides. New genus and species. 
Phieomys pallidus, 

Mus everetti. 

Mus luzonicus. New species. 

Mus decumanus (feral). 

Mus chrysocomus. 

Mus ephippium negrinus. 

Batomys granti. New genus and species. 
Carpomys melanurus and C. pheurus. New genus and two new species. 
Crateromys schadenbergi. 


In addition to these, Crunomys fallax, a new genus and species, was obtained in the 
district of Isabella, east of Monte Data. 

Therefore no less than six new genera and eight new species were discovered in the 
island, a proportion of novelty that has perhaps never been equalled in the history of 
Mammal-collecting. 

Besides these new forms, Mr. Whitehead discovered a new genus and species of Bat 
in Mindoro and several additional species of Bats and Rodents in that and other 
islands. 

What are the true affinities of the isolated fauna of Luzon is a question that is 
not easy to answer, for the representative forms are mostly so peculiar as to render 
their zoological relationships more or less doubtful. On the whole, the connections, 
such as they are, seem to be partly with Celebes and partly with the Australian region. 
Thus Rhynchomys seems to have its nearest ally (Hchiothrix) in Celebes; Mus chryso- 
comus is actually a Celebean species; while Chrotomys, Celenomys, and Crunomys 
belong to a subfamily, the Hydromyine, hitherto known only from Australia and New 
Guinea. Finally. Crateromus seems to have its nearest ally in Lenomys from Celebes, 
and in another new genus not yet described that occurs in New Guinea. 

On the other hand, Ph/womys is so isolated that I can make no suggestions as to 
what is its nearest ally, and Carpomys and Batomys belong to a group of arboreal 
genera scattered over the oriental part of the East Indian Archipelago. This group 
of genera may possibly either haye a definite alliance one to tbe other, independent 


FROM THE PHILIPPINE ISLANDS. 379 


of Mus, or may be isolated survivors of an older murine fauna, of which Mus has 
now gained the dominant position, or finally may all be independent offshoots of the 
same central genus. Probably the second of these hypotheses approaches nearest 
to the truth, although one or two of the less differentiated genera, such as Vandeleuria, 
may have arisen in the third way. 

In any case, Science is to be congratulated on the wonderful series of new forms 
which Mr. Whitehead’s exploration of the little plateau of Monte Data has placed at 
her service, and I feel sure that, when they are studied by someone better able than 
I am to make out their complicated relationships, our general knowledge of geo- 
graphical and phylogenetic evolution will be by their aid materially increased. 

The following are Mr. Whitehead’s general notes on the collection :— 

“The Mammals were collected during a period of three years spent in the Philippine 
Islands. As my time was occupied chiefly with ornithology and no special effort was 
made to collect Mammals, the results may be looked upon as fairly satisfactory. 

“ The largest collection, and at the same time the most interesting, was formed in 
the highlands of the Province of Lepanto, North Luzon, chiefly on Monte Data, 
a table-topped mountain of from 7000 to 8000 feet in altitude. A few specimens come 
from Cape Engaifio, the most northern point of East Luzon, and a few Rats and a new 
Bat from the highlands of Mindoro. In Negros very few Mammals were met with. 
In Samar several interesting forms were obtained, including Tarsius philippensis, but, 
with the exception of a new Pigmy Squirrel, all well known to naturalists. : 

“Monte Data, my chief collecting-ground, is inhabited by a peculiar wild tribe of 
Malays, not of Negrito stock, who call themselves Igorrotes. I found them very pleasant 
savages, and, fortunately for me, they knew the value of coins. By purchasing all the 
animals brought to my camp, I soon had quite thirty Igorrote collectors hard at work 
most of the day with their little terriers, digging out Rats and snaring the larger 
mammals and birds. The various small Rats they brought in alive—-and often had 
their hands much bitten—as I refused to deal in dead specimens, the skulls being 
generally smashed to bits. In less than a month’s time the tent was festooned with 
rat-skins hanging up to dry, my collection consisting of over one hundred specimens. 
A perusal of the list (see p. 378) will give the reader an idea of the Rodents obtained 
on this mountain. 

“The table-top of Monte Data is perhaps over three miles long, by one mile (or more 
in places) broad. The vegetation consists chiefly of oaks and pines, all well clothed 
in lichen and other parasitic plants, but the undergrowth of bamboo, fern, and raspberry 
is very thick. ‘The ground is much burrowed by Rats; and I may mention that the 
largest known species of Old-World Scops Owl comes from this place, showing that 
the food-supply is both abundant and nutritious. In Mindoro I was most unfortunate, 
visiting that island in the wet season. We were unable to leave our tent for days 

3D 2 


380 MR. OLDFIELD THOMAS ON MAMMALS 


together, and during three months only some five days were fine. A variety of the 
Common Rat was soon attracted to our camp, where it became quite a nuisance. 
In Negros my camp was also infested with Rats, many of which we trapped. 

“The distribution of Mammalia throughout the Philippine Archipelago is most 
interesting, but the larger islands are by no means thoroughly explored—more 
especially Mindoro, Mindanao, and the Pacific coast of North and Central Luzon. 

“In the larger islands of Luzon, Mindoro, Panay, Negros, and Cebu, we find neither 
Tarsius, Galeopithecus, nor Sciurus; but all these Bornean genera are found in Samar 
and Leite. arsius is wanting from Mindanao and Bohol, but when the larger island 
is explored it will doubtless be met with there also. 

“In Luzon and Mindoro no indigenous Felis has yet been discovered ; this genus 
occurring in Panay, Negros, and Cebu. Though it has not yet been obtained in 
Mindanao, I expect it will be eventually discovered in that great island. 

“Tt is possible that /elis does not occur in Samar and Leite, as these islands are 
much more to the east and may have missed the migration—as apparently the Negros, 
Panay, and Cebu group have missed that of Sciwrus and other genera. If Felis is 
confined to the Negros group, it seems probable that Man was the agent of intro- 
duction of this Bornean animal. 

“Tjuzon has many wonderful Rodents peculiar to it, notably such genera as Phlwomys 
and Orateromys. In the island of Marinduque, Ph/womys also occurs; but this island, 
by its birds alone, is really a part of Luzon, from which island it is separated by a strait 
equal, however, to that which separates Luzon from Mindoro. 

“Mindoro is remarkable for its Tamarau (Bubalus mindorensis), an animal perhaps 
more nearly allied to the Anoa of Celebes than to any other. It is interesting to 
notice the absence of such Bornean genera as Tupaia, Mydaus, Aretictis, Hystrix, and 
Sciurus, which are found in Palawan and the Calamianes, but have never been able to 
cross into Mindoro. ‘There are, however, several Palawan birds in Mindoro which are 
not met with in any other of the true Philippine Islands. 

. Tn the west-central islands Panay, Negros, and Cebu, we find a paucity of Mammals, 
giving one the idea that any land-connection with Mindanao must have been either 
very ancient or of brief duration; while in the east-central islands Samar, Leite, and 
Bohol, we meet with several genera in common with Mindanao and Borneo. It is 
perhaps possible to state with some certainty that the true Philippine group has 
received no Mammals from Borneo vid Palawan, but several genera from Borneo 
and perhaps Celebes wid Mindanao, which have been unable to spread further north 
than Samar; and at that period of migration there was no land-connection with 
North-west Mindanao and the Negros group. Luzon probably received its peculiar 
Rodents vid Formosa, and they were unable to spread beyond that island; but at 
present the highlands of Formosa are a terra incognita. 


FROM THE PHILIPPINE ISLANDS. 581 


“There are, however, four genera of Mammalia which are dispersed throughout the 
entire Philippine Archipelago, viz. Macacus, Paradoxurus, Viverra, and Sus, all of 
which are found in Borneo and Palawan, and all of which are carried about by man; 
for to me it seems impossible to account in any other way for such a general distribution 
of these four genera, while so many other genera are so strangely and strictly 
distributed. Cervus is also found in many of the Philippines, but its exact distribution 
is probably unknown, neither have I heard of Palawan as a locality; it is also an 
animal much carried about by man. Cervus and Sus are also able, and doubtless do 
increase their distribution by swimming from island to island ; nearly every small island 
off the coast of the large islands being inhabited by Sus. On the top of Monte Data 
there were small herds of semi-wild pigs belonging to the Igorrotes, and doubtless 
many must revert to their wild state. Therefore I am much inclined to look upon 
man as the chief agent in the distribution of the Pig. Deer, of course, are conveyed 
everywhere and put down with the idea of affording future sport.” 


Macacus cyNnomo.eus (Linn.). 

a, d. Barit, Abra Dist., N. Luzon, Nov. 11, 1894. 

“The Long-tailed Green Monkey is common throughout the Philippine group. 
It, as is usual with the various members of this great family, does much harm among 
the crops planted by man. In North Luzon monkeys infest the forests in the neigh- 
bourhood of native plantations, especially those of maize and sweet potatoes; in Samar 
the rice-fields had to be carefully guarded from their attacks. From the sea-coast to 
the tops of the mountains the Chongo is ubiquitous. In North Luzon small bands 
frequented the flat summit of Monte Data, where during the winter months the tem- 
perature is as low as 28° Fahr. In Mindoro and Negros we also noticed it at 6000 feet. 

“ Distribution. Found commonly throughout the Philippine Islands, including 
Palawan. 

“Native Tagalo name, ‘Chongo,’”—J. W. 


TARSIUS PHILIPPENSIS Mey. 

Tarsius philippensis Meyer, Abh. Mus. Dresd. 1894-95, no. 1, p. 1 (1894), 

a, 6. Yg.g, adult 2. Samar, June 16, 1896. 

Dr. Meyer makes a primary character of the asserted nakedness of the tarsi in the 
Philippine Zarsius, but both these specimens, which may be looked upon as topotypes, 
instead of having “ tarsi denudati ” (“ vollkommen nackt”), should rather be described, 
like Dr. Meyer’s 7’. sangirensis 1, as “ tarsis fere nudis.” ‘The exact differences between 
the last-named and the Philippine Tarsius are not stated by Dr. Meyer when carefully 
explaining why it is distinct from 7. fuscus. 


' Abh, Mus. Dresd. 1896-97, no. 1, p. 9 (1896), 


382 MR, OLDFIELD THOMAS ON MAMMALS 


“This remarkable mammal is found in the islands of Samar and Leite, where it is 
called by the Bisayas ‘ Magou.’ So far as I am aware, it has not been obtained in 
Luzon or Mindoro to the north, or in Masbate, Cebu, Negros, or Panay, islands to the 
west and north-west of Samar. It probably occurs in the great island of Mindanao, 
and perhaps in Bohol, to the south of Leite. 

“Tn habits the ‘ Magou’ is nocturnal, as the enormous owl-like eyes would lead one 
to suppose ; it frequents abandoned clearings, where the new growth has sprung up to 
a height of some twenty feet, and in Samar, where the ground is also thickly covered 
with ferns and other plants to a height of some three feet. In such places this little 
animal easily conceals itself during the day. I had the good fortune to see a 
‘Magou’ in such a locality one day in Samar. The Yarsius was clinging to the stem 
of a small tree just above the fern-growth, with its peculiar hands round the tree; it 
was awake and intently watching my movements, and permitted me to approach as 
close as I wished: when, doubtless, at the least sudden movement of my hands it 
would have jumped to the ground and made off in the thick undergrowth. During 
the night the ‘ Magou’ is very active, and may often be heard, in localities where 
they are numerous, uttering a peculiar squeak like a monkey. From its habit of 
feeding only on insects, this animal has a strong Bat-like smell. 

“Tn Samar, where at different times I kept several ‘ Magous’ alive, I found them 
very docile and easily managed during the day. hey fed freely off grasshoppers, 
sitting on their haunches on my hand. When offered an insect, the ‘ Magou’ would 
stare for a short time with its most wonderful eyes, then slowly bend forward and 
with a sudden dash would seize the insect with both hands and instantly carry it to 
its mouth, shutting its eyes and screwing up its tiny face in a most whimsical fashion. 
The grasshopper was then quickly passed through the sharp little teeth, the kicking 
legs being held with both hands. When the insect was beyond further mischief, the 
large eyes of the ‘ Magou’ would open, and the legs and wings were then bitten off, 
while the rest of the body was thoroughly masticated. My captives would also drink 
fresh milk from a spoon. After the sun had set this little animal became most 
difficult to manage, escaping when possible, and making tremendous jumps from chair 
to chair. When on the floor it bounded about like a miniature kangaroo, travelling 
about the room on its hind legs with the tail stretched out and curved upward, 
uttering peculiar shrill monkey-like squeaks, and biting quite viciously when the 
opportunity offered. During the day the pupil of the eye becomes so contracted that 
it appears only as a fine line, but after dark it is so expanded as to fill up most of 
the iris. 

‘The popular native idea is that the ‘ Magou’ feeds on charcoal, the reason for 
this being that the animal is generally found after the old plantations have been cut 
down and burnt, the ‘Magou’ doubtless having returned to its old haunts from 


FROM THE PHILIPPINE ISLANDS. 385 


which it had been driven by the woodcutters. This delusion is fatal to all captured 
‘Magous,’ as they are immediately put on a diet of charcoal, and therefore soon 
starve to death.”—J. W. 


Preropus JuBaTuSs Eschsch. 

a,b,c. 3ad.sk. S2. Barit, Abra Dist., N. Luzon, Noy. 1894. 

These specimens, practically topotypes of the species, which was described from 
Manila, haye the brilliant golden napes and apparently all the other characters 
described by Prof. Elliot as diagnostic of his Pt. auri-nuchalis’. It seems probable, 
therefore, that this latter name should be considered as a synonym of Pt. jubatus, 
of which the range no doubt extends over the whole of the Philippines. 

“This large Fruit-Bat was in immense numbers in the Province of Abra, N. Luzon, 
where it had taken possession of a long, low range of hills, well covered with forest. 
Just at sunset these Bats issued from their roosting-place in thousands towards all 
points of the compass. Numbers of those that passed the Abra river dipped to drink 
in the stream, but seemed afraid, making often several attempts before they dared to 
come low enough to touch the water. On the sea-coast also the large Fruit-Bats 
often dip to drink in the sea on calm evenings. This Bat has a peculiar, though not 
disagreeable, odour. The wings are quite sticky to the touch. Met with in North- 
Central Luzon.”—J. W. 


PTEROPUS VAMPYRUS (Linn.). 

a, b. Verac, Catanduanes Island, Sept. 1894. 

This species occurs in every collection made in the Philippines, and is evidently 
common throughout the archipelago. 

“Tn the island of Samar we obtained several examples, which were unfortunately 
burnt with my collection on the s.s. ‘Weyland.’ In Samar this Bat was found 
roosting during the day in the mangrove-swamps in great numbers. 

“*My specimens were obtained in the islands of Catanduanes, South Luzon, and 


Samar.”—J. W. 


XANTHARPYIA AMPLEXICAUDATA (Geoffr.). 
a,b, $ 2. Highlands of Benguet, Luzon, 5000 feet, Feb. 24, 1894. 


These specimens represent Gray’s “ Eleutherura philippinensis,” from Manila. 
Although stated to have been received from Gould, no doubt the type of that form 
was originally obtained by Cuming. 


’ Field Col. Mus. Publ. vol. i. p. 77 (1896), 


584 MR. OLDFIELD THOMAS ON MAMMALS 


Genus Harpyionycreris ‘Thos. 
Harpyionycteris Thos., Aun. Mag. N. H. (6) xvii. p. 243 (1896). 

Index with a claw. Wings from the sides of the hairy back, inserted behind at the 
junction of the first and second toes. No tail. Hind limbs apparently very short. 
Interfemoral membrane obsolete, buried in thick fur. 

Dentition.—I. ;4,, C. 4, P. 3, M.4x2=28 or 30. 

Teeth (Plate XX XV. figs. 1-4). Upper incisors large, touching each other and the 
canines; shaped, when viewed in front, almost like those of Desmodus, each with a 
long oblique cusp touching its fellow in the middle line of the skull, but in section 
each is broadly triangular, with a broad posterior basal ledge. Canines with a large 
posterior secondary cusp, about half as high as the main cusp, and with a broad 
postero-internal basal ledge, but no additional internal cusps ; its direction much more 
slanting forward than usual, as is the lower canine also, so that the two cross each 
other nearly at right angles, instead of being approximately parallel. First two 
premolars about as in Cynopterus. Molars oblong in section and of a peculiar cuspidate 
character, the lateral longitudinal walls to the usual median groove broken up into 
several minute cusps, none of which are at all specially lengthened. Below, the 
incisors are practically obsolete, being minute and almost crowded out! by the large 
canines, which touch each other in the middle line, and have each an antero-internal 
and a postero-external secondary cusp and a broad posterior ledge. 

It is difficult to say with certainty to what previously known genus this remarkable 
form is most nearly allied. Its peculiar canines to a certain extent recall those of 
Harpyia, but this resemblance may be either accidental or due to their common 
descent from the (presumably) cuspidate-toothed ancestors of the Pteropodide?. On 
the whole it may be most conveniently placed near Xantharpyia and Boneia, with 
which it shares certain external characters, an indical claw, and the cheek-tooth 
formula of P. 3, M. 3; but the unique incisors, the short bi- and tricuspidate canines, 
and the multicuspidate molars separate it widely even from: these, and render it one 
of the most isolated of all the genera of the group. Its skull and dentition are 
figured on Plate XX XV. figs. 1-4. 


HARPYIONYCTERIS WHITEHEAD! Thos, (Plate XXX. fig. 1.) 

Size about as in Xantharpyia ampleaicaudata. Fur soft, close and woolly, especially 
posteriorly. General colour of the fur all over, above and below, a uniform chocolate- 
brown, a little darker on the face, and a little lighter on the nape and shoulders. 


' In the single type-specimen one lower incisor only is present, the other haying fallen. 
* See P. Z.8. 1888, p. 473. 


FROM THE PHILIPPINE ISLANDS. 385 


Wing-membranes dark, with a few whitish spots scattered about them. Lars of 
medium length, rounded at their tips. Fur of the back extending thinly on to the 
forearms, and covering the hind limbs densely down to the roots of the claws. Inter- 
femoral membrane barely a tenth of an inch wide, wholly buried in the fur. 

Dimensions of the type (an adult skin of-doubtful sex) :— 

Forearm 84 millim. (=3°3 inches); head and body 140; ear 17; index-finger and 
claw 60; third finger, metacarpal 59, first phalanx 44, second phalanx 54. 

Skull; basal length 37-5; greatest breadth 23-8; interorbital breadth, tip to tip 
of postorbital processes, 6-9. Front of canine to back of m.? 17. 

Hab. Mindoro, alt. 5000 feet. Dec. 1895. 

“This interesting new Fruit-Bat was shot by me in the highlands of Mindoro at 
an altitude of 5000 feet. It was flying round some high trees at dusk, at which 
time I generally sat out near my camp on the look-out for nocturnal birds. The 
specimen, when shot, fell into some tangled undergrowth, and it was only after a careful 
search with a lamp that my servant found it. 

“« Distribution. Mindoro, 5000 feet.”—J. W. 


CARPONYCTERIS AUSTRALIS Pet. 
a.9. Negros. 
“Obtained a short way up the Canloan volcano.’—J. W. 


HIPPosIDERUS DIADEMA Geoffr. 


a. Manitoc, Albay, 8.E. Luzon, Aug. 1894. 
6. Catanduanes, Sept. 24, 1896. 


PIPISTRELLUS IMBRICATUS (Horsf.). 
a. 9. Manila. 


A young individual, apparently of this rare species. 
“ Picked up in a dying state on the side-walk in Manila.”—J. W. 


Myotis MAcRorarsus (Waterh.). 

a,ad. al. 9. Manila, May 20,1876. Presented by Mr. Whitehead. 

This Bat was originally discovered by Cuming, and no other specimen has been 
received by the British Museum until now. I fail to see, either in the fresh specimen 
or in the type, that the wing-membrane is attached to the body much nearer to the spine 
than is usual, a character on which Dobson lays some stress. The black claws of the 
type, also specially mentioned by him, may have betn caused by some fluid in which 
the specimen had been put, for Mr. Whitehead’s fresh specimen, unquestionably 
identical specifically, has the claws of the normal pale colour. 

** Brought to me by some boys in Manila.”—J. W. 

VOL. XIV.—PaRrT vi. No. 2.—June, 1898. 35 


386 MR. OLDFIELD THOMAS ON MAMMALS 


KerivouLa WHITEHEADI Thos. 
Kerivoula whiteheadi Thos. Ann. Mag. N. H. (6) xiv. p. 460 (1894). 
a. &. Molino, Isabella, N.E. Luzon, May 1894. Type. Presented by Mr. Whitehead. 


Size and proportions about as in If. hardwickei, but the ears are slightly longer and 
the lower legs shorter. Upper surface of “wing-membranes to a line drawn from the 
elbow to the foot, whole of interfemoral membrane except the terminal half-inch, and 
surface of lower limbs to feet, thinly but distinctly clothed with long orange-coloured 
hairs, these parts in K. hardwickei being practically naked. Forearm, carpus, and 
index also thinly clothed. Hinder edge of interfemoral with a few short hairs along it, 
scarcely forming a fringe. 

Colour above rufous-orange, the slaty bases to the hairs showing through, below 
dark slaty, the lighter tips scarcely affecting the general dark tone. 

Upper inner incisors slender, with a distinct posterior secondary cusp, to the tip of 
which the unicuspid outer incisor just reaches. Other teeth apparently as in 
K. hardwickei. 

Dimensions of the type (an adult male in alcohol) :— 

Forearm 32 millim. (=1'25 inch). : 

Head and body 39 millim.; tail 39; head 16; ear from notch 13:5; tip to tip of 
ears across head 28°5 ; length of index 31:5; third finger (exclusive of cartilaginous 
tip) 61, fifth finger 47 ; lower leg 16-2; hind foot without claws 8. 

Hab. Isabella, N.E. Luzon. 

Type. B.M. 94. 10. 9. 2. 

This species is undoubtedly very close to K. hardwicket, but may be distinguished by 
its hairy interfemoral and by the different structure of its upper incisors. It may be 
noted that a Mindanao specimen of the older known species shows no approximation to 
K. whiteheadi. 


MINIOPTERUS SCHREIBERSI PUSILLUS Dobs. 
a, b, Barit, Abra, Luzon. Presented by Mr. Whitehead. 


“Captured in a butterfly-net, while chasing each other round my room.”—J. W. 


GALEOPITHECUS PHILIPPINENSIS Waterh. 

a. do. Samar, June 10, 1896. 

‘Fairly common in Samar and Leite, and on the small islands between; I have also 
seen dozens of skins from the island of Bohol. Several Spaniards do quite a trade in 
the skins of this Lemur, which are of all shades of brown, grey, and even bright yellow. 
Generally beautifully mottled, but at times quite unmarked. 

“The Flying Lemur passes the day in sleep, clinging to the trunk of some large tree 


FROM THE PHILIPPINE ISLANDS. 387 


—and doubtless the coloration of the tree-bark is selected to match the fur by the 
resting animal, for I have shot in Malacca grey specimens on grey-barked trees. 
“The ‘Caguang’ of the Bisayas.’—J. W. 


Crocipura (CRroc.) Grayr Dobs. 

Crocidura (Croc.) grayi Dobs. Ann. Mag. N. H. (6) vi. p. 494 (1890). 

a. Benguet, Luzon, Feb. 1894. Presented by Mr. Whitehead. 

b. Monte Data, Feb. 1895. 

This Shrew was described by Dr. Dobson from two specimens in the British Museum 
that had been received from the Zoological Society’s old collection, and had been 
obtained by Mr. H. Cuming. Although merely labelled “ Philippines,” they were 
most probably from Manila. 

Luzon also contains a member of the subgenus Pachyura, examples of which in the 
British Museum have been labelled by Dr. Dobson as C. murina. Probably they 
represent Peters’s C. luzoniensis 1. 


Friis MiInuta Temm. 
a. Negros. 


“This handsome little Cat is apparently found only in the islands of Panay, Negros, 
and Cebu; but as it also occurs in the great continental island of Borneo, doubtless 
it will some day be found in Mindanao. One of my hunters declared that he shot at a 
Wild Cat in Samar among some rough broken-up limestone, into which the wounded 
animal unfortunately disappeared. I think we may say for certain that this Cat does 
not occur in Luzon, which is so well cultivated that it could scarcely have escaped 
detection. In Mindoro it might be possible for this animal to have escaped detection, 
as the island is perhaps, after Mindanao, the wildest and most densely covered with 
forest of the whole group. 

“In Negros, where we obtained a specimen of Felis minuta, the animal 
frequented the sugar-plantations, where it finds an abundance of rats. During 
harvesting operations this Cat is often captured by the natives, who form a ring round 
the last patch of standing cane. One of my collectors said that he saw this animal as 
high as 6000 feet, on Canloan volcano. 

“ Distribution, Panay, Negros, and Cebu.”—J. W. 


Feis DoMestica L. 


Reference has already been made? to what appears to be a feral Domestic Cat 
obtained by Mr. Whitehead on Monte Data. Mr. Whitehead’s own notes on the 
subject are as follows :— 


1 MB. Ak, Berl. 1870, p. 595. 
* Ann, Mag. N, H. (6) xviii. p. 245 (1896). 


Lo 


35 


388 MR, OLDFIELD THOMAS ON MAMMALS 


“In North Luzon we obtained a very large specimen of a Wild Cat, on the mountains 
at an altitude of 7000 feet. This animal, I am told, isa feral race of the Domestic Cat, 
Felis domestica, but it is unlike any Cat that exists in the native villages of to-day, being 
nearly double the size of any Igorrote Cat, and tabby marked, on a rather sandy 
ground. My friend Mr. A. H. Everett, however, informs me that he obtained a Wild 
Cat very like it in Celebes, which turned out to be an offspring of some escaped Domestic 
Cat.”—J. W. 


VIVERRA TANGALUNGA Gray. 

a. Cape Engaiio, N. Luzon, May 17, 1895. 

“We met with this beautifully marked Musang at Cape Engafio, the most northern 
point of East Luzon. One of the specimens obtained is much more clearly marked 
than the other, and also slightly larger. This Musang was also snared by the natives. 
In habits it resembles Paradoxurus, both being decidedly nocturnal and expert tree- 
climbers. 

“ Distribution. Found in all the larger islands of the Philippines, including Palawan 
(Bourns and Worcester).”—J. W. 


PARADOXURUS PHILIPPINENSIS Jourd. 


a. 3. la Trinidad, Benguet Dist., N. Luzon, Feb. 8, 1894. 

6. 2. Monte Data, Lepanto, N. Luzon, Feb. 1895. 

“Common throughout North Luzon, especially in the high mountains, where mela- 
nistic forms seem to occur on an average of one to two with brown ones. ‘The 
Musang is easily secured by the Igorrote hunter, by setting springes in the narrow 
mountain pathways, the space on each side of the snare being carefully stopped, 
forcing a passing animal to walk over the trap, which generally nooses it by one of the 
fore-paws. In these mountain-paths will be noticed the numerous excreta of this 
animal, which are often composed of the seeds of small forest fruits; but if a coffee- 
plantation be in the vicinity the excreta are made up of coffee-stones, the pulpy 
encasement of the coffee-pip being very sweet. The Musang is, as might be expected, 
a great enemy to all sorts of poultry, killing simply for amusement after hunger has 
been satisfied. Met with in North Luzon from the coast up to 8000 feet. 

“ Distribution. Found in all the larger islands of the Philippines, including Palawan.” 
—J. W. 


ScIURUS SAMARENSIS Steere. 
a, Samar, June 6, 1896. 


The figure given by Dr. Meyer ' of this species is evidently very much over-coloured, 


* Abh, Mus. Dresd. 1896-97, no. 6. p. 29, pl. xi. fig. 2 (1896), 


FROM THE PHILIPPINE ISLANDS. 389 


as neither Mr. Whitehead’s specimen nor one of Steere’s co-types in the British 
Museum has feet anything like so strikingly black as is there shown. 

The British Museum possesses examples of three species of middle-sized Squirrels 
from the Philippines—%S. steerei Giinth., of Palawan and Balabac, S. philippinensis 
Waterh., of which, besides the much-deteriorated type from ‘‘ Mindanao,” Mr. Everett 
has sent examples from Zamboanga and Basilan, and S. samarensis Steere, of 
Samar. Whether, as the localities would indicate, S. mindanensis Steere (S. cagsi, 
Mey.) is synonymous with 8. philippinensis, or is most closely allied to S. samarensis, 
I am not at present able to determine. 

“Met with both in Samar and Leite, but by no means common, being difficult to 
see or shoot owing to the great height of the forest trees in these islands. 

“The ‘ Alalaksing’ of the Bisayas.”—J. W. 


NANNOSCIURUS SAMARICUS sp. n.! (Plate XXX. fig. 2.) 
a. 2. Samar, June 30, 1896. Type. 
Allied to WV. concinnus Thos., but greyer and less rufous. Two premolars present in 


the adult. 

Size and general characters very much as in WV. concinnus. Fur, however, much 
shorter and more velvet-like, the hairs about 5 millim. Jong on the back. General 
colour of head and body finely grizzled olive-grey, with only a faint tinge of rufous on 
the back, thus contrasting with the broadly rufous-washed WV. concinnus. - Under 
surface rather thinly haired, dirty greyish, not defined on the sides. Limbs dusky, 
upper sides of hands and feet dusky grizzled grey, a few orange-tipped hairs on the 
digits. Characters of sole-pads apparently much as in WV. coneinnus. Tail similar to 
that of the allied species, but the rufous rings on the hairs are less developed, and the 
black ones more, su that the general result is darker. 

Skull apparently very similar to that of the allied species, but the nasals are some- 
what narrower. 

Two upper premolars present, the anterior minute, styliform, circular, the posterior 
considerably larger, but still much smaller than m.!. Molars all much more rounded 
than in WV. concinnus, their transverse scarcely exceeding their longitudinal diameter. 

Dimensions of the type, an adult female, in skin :— 

Head and body 88 willim.; tail, without hair 69, with hair 94; hind foot 
(moistened) 25:2. 

Skull: greatest breadth 162; nasals, length 7°7, breadth 3-1; interorbital breadth 10 ; 
tip to tip of postorbital processes 12°6 ; diastema 6; length of cheek, with series (p.* 
to m.*) 4:1, of three molars only 3:0. Lower jaw: condyle to incisor tip 18°2; bone 


only 15°5, 


‘ See preliminary diagnosis in Minutes of P. Z. S. for June 15, 1897 (published June 19). 


390 MR. OLDFIELD THOMAS ON MAMMALS 


This little Squirrel is perhaps merely the representative of WV. concinnus in Samar, 
as it seems probable that there are really two premolars in that animal as in the other 
Malayan Wannosciuri. The original specimen was described by me as having only one 
premolar; but this latter proves on further examination! to be the milk-premolar, 
a fact which renders it rather uncertain whether the adult may not have the additional 
anterior premolar generally present. 

Apart from this question, WN. samaricus may be readily distinguished from LV. con- 
cinnus by its longer fur, much more rufous coloration, and rounder molars. 

“Like the last species, but less often observed.”—J. W. 


CELEZNOMYS, g. n.” 


Colour normal. External form as in Chrotomys. 

Skull (Pl. XXXV. fig. 12) broad and strong, evenly rounded, without ridges, very 
wedge-shaped in lateral view, owing to the great height of the brain-case, and the 
uniform way in which the fronto-nasal and palatal profiles approach each other 
anteriorly. Nasals short, not overhanging the incisors. Brain-case smooth and 
rounded. Interparietal strap-like, fairly well developed. Anteorbital foramen little 
expanded above, the front edge of its outer plate vertical, not produced forward. 
Palatal foramina very small. A distinct incisive fissure? present, nearly half the size 
of one of the palatal foramina. Posterior nares broad. Lower edge of mandible 
peculiarly flattened just behind the symphysis, and pierced with a large number of 
minute foramina, Coronoid processes long, strongly curved backward. 

Teeth. Incisors much thrown forward, simple, rounded and bevelled in front in a 
manuer similar to that found in Lophuromys. Molars 2 (Pl. XXXYV. fig. 11), in 
essential structure like the anterior two of Chrotomys (see below), but the ridges and 
crests less sharp, although this may be (indeed probably is) due to wear, a point 
which cannot be settled until young examples are examined. No trace of a third 
molar either above or below. 

Type. C. silaceus Thos. 

This genus, although it has the same reduced number of teeth as Hydromys and 
Xeromys, is no doubt really most closely allied to Chrotomys, to which, both in external 
form and in the general shape of the skull, it presents considerable resemblance. Still, 
besides the absence of m.°, it may be distinguished by its normal coloration, longer 
and narrower brain-case, and larger interparietal. 


1 Cf. Forsyth-Major, P. Z. 8. 1893, pl. xi. fig. 7. 

* ceXawés, dark-coloured ; in contradistinction to Chrotomys, derived from xpws, colour, in allusion to the 
striking coloration of Chrotomys whiteheadi. 

*° By this term I refer to a small mesial opening present, in a great many different forms, between the two 


premaxille, just behind the incisors. 


FROM THE PHILIPPINE ISLANDS. 591 


The suppression of m.* in Celenomys is an interesting sign of its relationship to the 
Australian members of the subfamily, Hydromys and Xeromys, both ot which have only 
two molars, while the other two Philippine genera, Chrotomys and Crunomys, have the 
normal Murine number of tliree molars. 


CELANOMYS SILACEUS (Thos.) (Plate XX XI. fig. 1.) 
Xeromys (?) silaceus Thos. Ann. Mag. N. H. (6) xvi. p. 161 (1895). 

a, b. Monte Data, Feb. 1895. 

Size of 1 common Rat. Fur soft, close and velvety, hairs on posterior back about 
10-12 millim. in length. General colour uniform slaty grey, very finely grizzled with 
whitish, but so finely as scarcely to affect the general grey tone. Sides of muzzle nearly 
black. Under surface rather paler than the back, not sharply defined, the hairs slaty 
grey basally, washed with buffy white terminally. Eyes small, not black-ringed. Lars 
short, uniform greyish. Hands and feet as far as the metapodials dark grey, the 
digits whitish or flesh-coloured. ‘Tail rather shorter than the body without the head, 
thinly haired, brown above basally, whitish below and at the tip. 

Skull as already described. 

Dimensions of the type ( 3 ) taken in skin :— 

Head and body (probably rather stretched) 195 millim.; tail 110; hind foot 
(moistened) 33°4. 

Skull, see p. 396. 

Hab. Monte Data, Lepanto, N. Luzon, 8000 feet. 

«This curious Mammal at first sight might easily be confounded with Rhynchomys 
soricoides, and, like that animal, was also obtained on the table-topped summit of 
Monte Data. It seems rare, only two specimens having been snared in some five weeks. 
The skull and teeth, instead of being frail as in Rhynchomys, are powerful, and much 
more nearly allied to Chrotomys. ‘The eye is small as in Rhynchomys, and the outward 
appearance quite as Shrew-like. The habits of this peculiar Mammal I am quite 
unable even to guess at. 

“ Distribution. High mountains of Central Northern Luzon.’—J. W. 


CHROTOMYS. 


Chrotomys Thos. Ann. Mag. N. H. (6) xvi. (1895) p. 161. 

Colour abnormal among Muride, the back prominently striped. Form suited for a 
terrestrial, not aquatic life. Size about as in the common Rat. Fur soft and straight. 
Muzzle apparently not cleft. Eyes rather small. Ears well developed. ‘Tail rather 
short, thinly haired, scaly. Pollex with a rounded nail; other digits, including hallux, 
with well-developed, little-curved claws. 


92 MR. OLDFIELD THOMAS ON MAMMALS 


O> 


Skull (P1. XXXV. fig. 9) in general form not unlike that of Celwnomys, but even 
more wedge-shaped owing to its greater height posteriorly. Nasals short, their anterior 
end level with the middle of the incisive fissure. Interorbital region similarly rounded 
and unridged. Brain-case broader and shorter, so that its breadth is equal to its 
length. Interparietal very small, a mere narrow transverse slip. Anterior edge of 
zygoma-plate slightly concave, the plate little developed. Incisive fissure large, quite 
half as large as one of the palatal foramina, which are. as usual in this group, very 
small. Posterior nares large and open, the hinder edge of the palate level with the 
posterior lamina of m.*. Pterygoids large, projecting downward considerably below 
the level either of the molars or bulle. Lower jaw as in Celwnomys. 

Teeth. Incisors pale yellow, thrown forward, simple, rounded in front. Molars 3 
(Pl. XXXYV. fig. 8), the anterior two very similar in structure to those of Xeromys 
(figured P. Z.S. 1889, pl. xxix. fig. 10), but m.! has its middle lamina simpler (more as 
in Hydromys) and its posterior lamina is almost obsolete, while m.? has its posterior 
supplementary cusp more definitely postero-external, the difference in position being 
no doubt due to the presence of the additional molar behind. V/.° quite small, 
transversely or obliquely oval in section. In size m.? and m.? together are barely two 
thirds the length of m.?. 

Below, m., is of the most ultra-hydromyine character, without any of the suppressed 
cuspidation of the anterior margin found in Xeromys, and even without the supple- 
mentary postero-external cusp found in both the Australian genera. /.,asin XYeromys. 
M., nearly circular, about one-sixth the size of m.,, slightly larger than m.*. 


CHROTOMYS WHITEHEADI Thos. (Plate XXXII.) 
a-d. Monte Data, Lepanto, 8000 feet, Feb. 1895. 


Size of Mus rattus. Fur soft and thick, but not specially long. General colour 
greyish brown, tending in some specimens to rufous; a well-defined buff or orange 
line extending from between the eyes down the back nearly to the tail, shown up on 
each side by a broad shining black band. Under surface dull slaty buff, not sharply 
defined on the sides. Top of muzzle dark brown, continuous with the dark edgings 
to the central yellow band. Ears of medium length, fairly covered with minute hairs, 
uniformly blackish brown. Metapodials shining grey, digits nearly naked, whitish. 
Tail short, slender, about half the length of the head and body, thinly hairy, brownish 
black above, rather paler below, extreme tip whitish. 

Skull and teeth as already described. 

Dimensions of the type, an adult male, measured in skin :— 

Head and body 196 millim. ; tail 111; hind foot (moistened) 35. 

Dimensions of skull, see p. 395. 

Type. B.M. 95. 8. 2. 19. 


FROM THE PHILIPPINE ISLANDS. 393 


Owing to the remarkable modification in its colour, quite unique among Muride, 
this animal may be looked upon as one of the most striking of all Mr. Whitehead’s 
discoveries. Scientifically, it shares with Celenomys and Crunomys the interest 
attaching to the occurrence of the subfamily Hydromyinze away from the Australian 
region, to which the only two previously known genera are confined. No member of 
the group has as yet been found in any of the intervening islands, although it is possible 
that when the higher mountains of the archipelago are more thoroughly explored other 
forms referable to the subfamily will also be found to occur there. 

“This handsome Rat was obtained on the summit of Monte Data. It is said by the 
natives to feed on sweet potatoes and grass, and to frequent the neighbourhood of their 
plantations. Chrotomys is also met with at almost the sea-level, as I saw in Manila a 
specimen obtained in the Forest of Tarlac in Central Luzon to the north of that city. 

“ Distribution. Probably throughout Luzon.’”—J. W. 


Crunomys!, g. n. 

Exteinal characters, apparently much as in Yeromys, though the number of mamme 
and sole-pads cannot at present be determined. Far thickly mingled with spines. Ears 
short and rounded, Hallux with a claw. ‘Tail rather short, thinly haired, apparently 
flattened at end, but this appearance may be simply due to contraction in drying. 

Skull (Pl. XXXV. fig. 6) with the peculiar shape characteristic of many Water- 
Rodents, such as Hydromys, Ichthyomys, and others; low, flattened, its frontal profile 
concave. Nasals long, overhanging the incisors in front. Interorbital region broad, 
its edges with scarcely a trace of beading. Interparietal large. General shape 
of anteorbital foramina almost exactly as in Chrotomys, the outer plate not produced 
forward. Incisive fissure minute. Anterior palatine foramina short. Posterior edge 
of palate just level with the hinder edge of m.?. 

Molars (Pl. XXXYV. fig. 5) much worn in the only specimen, so that it is difficult to 
make out their exact structure. It is, however, clear that they are more murine in 
structure than is the case with the other members of the Hydromyine; in m.! the 
anterior lamina is oblique just as in the other genera of the Hydromyine, but in other 
respects might almost be that of Mus itself. J£.? is also very murine, having a small 
antero-internal cusp, a long middle lamina, and a mesial circular one posteriorly ; m.? is 
subcircular, with a small antero-internal cusp. Below, on the other hand, the teeth are 
not unlike those of Chrotomys, except that m., is bilaminate as in Mus, a difference that 
one would expect to occur owing to the greater development of this tooth in Crunomys, 

Type. Crunomys fallax. 

This genus is most interesting from an evolutionary point of view, for it adds 


1 kpovsos, a well-spring; xpovra/, torrents or streams. 
VoL. XIV.—Part vi, No. 3.—June, 1898. 3F 


594 MR. OLDFIELD THOMAS ON MAMMALS 


another to the links that connect the aberrant Hydromys with the true Murine, and is 
indeed the last link needed. For we may take five main characters as distinguishing 
Hydromys from an ordinary Mus, viz.: (1) aquatic form; (2) flattened skull; (3) 
reduced plate to zygoma-root; (4) two molars only; and (5) peculiar molar structure. 
The first discovered linking genus, Xeromys, was murine as to 1, 2, and 3, hydromyine 
as to4and 5; then came Chrotomys, murine as to 1, 2, and 4, hydromyine as to 3 and 5. 
Celenomys, described above, is like Chrotomys, but also hydromyine as to £; and now 
comes Cyrunomys, murine as to 1, 4, and to a certain extent the highly important 5 
(molar structure), but with the hydromyine 2 and 3, in addition to the short palatal 
foramina found in all the genera mentioned. 

Like all annectant genera, Crunomys is most difficult to place satisfactorily in the 
system, and it js only with much hesitation that I have included it in the Hydromyine, 
a position which will have to be revised when specimens showing the unworn dentition, 
the mammary formula, and other characters are available for examination. 


The following is a rough synopsis of the genera now considered to belong to the 


Hydromyine? :— 
Molars 3. 
Aquatic. Skull flattened; frontal profile concave . . . . . . . I. Hydromys. 
Terrestrial. Skull rounded ; frontal profile normal ‘ 
Outer wall of anteorbital foramen slightly projected forward. . . 2. Xeromys. 
Outer wall of anteorbital foramen not projected forward . . . . 3. Celenomys. 


Molars 3. 
Molars strictly hydromyine in structure. Back striped. Fur soft. 
MADRE SONeG WOSOREM yo > G6 oO oem oo co } 
Molars more murine. Back unstriped. Furspiny. Semi-aquatic . . 5. Crunomys. 


A. Chrotomys. 


The first two are Australian, the last three Philippine. 


- CRUNOMYS FALLAX, sp. n.? (Plate XX XIII. fig. 1.) 


Size about as in Xeromys myoides. Fur short and close, profusely mixed with 
flattened spines; neither hairs nor spines longer than about 6 mm. on the back. 
General colour pale greyish, lined with yellowish on the back. Dorsal spines white, 
darkening to black at their tips. Belly dirty greyish white, not sharply detined, the 
hairs slaty basally, dull whitish terminally. Sides of muzzle brown. Whiskers 
numerous, long, mixed black and white. Ears short, uniformly brown. Hands and 
feet greyish brown on the metapodials, lightening to white on the digits; fifth hind toe 


1 Since this paper was read an additional genus, Leptomys, has been described from New Guinea (Ann. 
Mus. Genoy. (2) xvii. 1897). It has 3 molars, like Chrotomys and Crunomys. 
* See preliminary diagnosis in Minutes of P. Z. 8. for June 15, 1897 (published June 19). 


FROM THE PHILIPPINE ISLANDS. 395 


reaching to the end of the Ist phalanx of the fourth. ‘Tail about the length of the 
body without the head, uniformly short-haired, black, rather lighter along the middle of 
its under surface. 


Skull-dimensions (in millim.) of Rhynchomys and Hydromyine, all from type 


specimens.+ 
Rhynchomys | Celenomys Chrotomys Crunonys 

soricoides, silaceus. a fallax. 

Paceeneiie west cer eee taf eee 44 34 87-5 (aeanee 
Bast amslene aye vetseecs cpeaciere rage ed teanee men GeiiacscaNaserecs as 41:5 31:7 35 205.) 
Greatest breadth ‘yin. rccterusarevecuts, aise sve ateces, di totj crsiel oie 19:5 19 21 (¢.)) 121 
Nasal selene titacncrtsatensecevneperesus crecacte rie aie venir 20 12 13:2 9:2 
PD LORCUL Hay eccaera tiesto satire ieretenere tier 38 3:3 41 2°8 
mferorbital ibreadthyey. scsrsacererietont eter isicierstemers 6-9 69 6°38 4-6 
Mnterpariotall len thie craaisyara «icisictcrvcrelei neues a careie nace 3:2 2°8 22, — 
fe Dreadthom vans nvatarseyamemtavretetustaxeterecesrsnste 10°5 86 75 9-1 
Length of anterior zygoma-root .......-..sesesevese 25 3°6 2-9 iey 

alate enotnamomehenselionwenrn reenact titi 245 71 20 11 

IDEN ihc oe Gtk REET Mee 5 ito beon ke APRN caten canter 16:8 13-1 15°2 72 
Anterior palatine foramina, length .................. 6°5 35 Ad 35 
aS 5 preadbhy72y ele Rrrteloie actors 2-1 16 21 15 
Thength of upper molar'series ........624-0..s0ss0de0s — -- 52 3°6 
of dor Clee COMO! A BotaAsanacascad ones 2:5 3°78 48 372 
Lower jaw, condyle to incisor-tip................++05 34 — 31-2 16°5 
is on evoniliy; Ms ya 'sroierysd-vogrecate mais ak ainke s 30 "  21°5 25°71 13-7 
i height, coronoid to angle ................ 9°7 10 151 —_— 
FA % | SUMMIT oe, Gado sososoooc abe 33 4 5 31 


Skull as already described. 

Dimensions of the type, measured in skin, and all merely approxiniate ; — 

Head and body 105 millim.; tail 79; hind foot 23; ear 10. 

Skull, see above. 

Hab. Isabella, Central N. Luzon. Alt. 1000 feet. Coll. May 1894. 

Type. B.M. 97. 4.8.4. Presented by Mr, Whitehead. . 

This little animal might readily be, and indeed for some time was, taken ay a species 


‘ A few of these measurements differ to a minute extent from those previously published, these latter 
haying been taken before the skulls were perfectly cleaned. The present measurements may be considered 


as the more correct. 
3F2 


9 


396 MR. OLDFIELD THOMAS ON MAMMALS 


of Wus allied to the group of IZ. ephippium, of which specimens are often found with 
more or less spinous fur. Mr. Whitchead, however, from the character of the place 
where he took it, thought it would prove to be a peculiar form, and his opinion has 
been most fully confirmed by an examination of its skull. 

The following are Mr. Whitehead’s notes on its capture :— 

“In one of my wanderings through the parched-up forests of Isabella (in North- 
Central Luzon) I noticed a small red kingfisher (Ceya melanura) fly into the scrub near 
a small stream. Having only a large gun with me, I sent my servant back to the 
village for a small collecting-gun. While seated beside the stream, a small mouse 
was observed among some large stones on the opposite side, busily searching after 
food. I opened oue of my 16-bore cartridges and picked out all the shot (No. 6) but 
four or five pellets, and luckily killed the small animal without much damage being 
done. Being sure, from its peculiar habits, that it must be something interesting, I 
carefully skinned it and sent it home, and am now rewarded by the addition of another 
new generic form to this already interesting collection.” 


Rayncuomys Thos. 
Rhynchomys Thos. Ann. Mag. N. H. (6) xvi. p. 160 (1895). 


Form rather Shrew-like. Muzzle enormously elongate. Feet normally murine, 
pollex with a broad nail. ‘Taii Rat-like, scaly, thinly haired. 

Skull (Pl. XX XY. fig. 10) of very peculiar shape, the brain-case broad, smooth and 
rounded, and the muzzle narrow and much elongated. Nasals long and narrow, 
terminating behind at the same level as the premaxille; viewed in profile they show 
a curious rise at their anterior extremity, the general frontal profile being practically 
an even slope from the crown to a point at the end of the anterior third of the nasals, 
and then bending upward again, as shown in the figure. Interorbital region smooth, 
evenly rounded, quite unridged, Interparietal large, transversely oblong, unusually 
variable as to its exact shape. Anteorbital foramen typically murine in essentials, but 
its outer wall very narrow and much slanted backward, so that the anterior edge of 
the upper root—the bridge—is actually posterior to the hinder edge of the lower 
root. In these respects it recalls the S. American Oxymycterus. Zygomata slender, 
low, and little sloped vertically, Palate long and narrow; a distinct incisive fissure 
present ; palatal foramen of normal size, but comparatively far forward in the skull, so 
that their posterior end is nearly their full length in front of the molars. On each 
side, just in front of m.1, there is a distinct raised ridge about a couple of millimetres 
long, and there is a somewhat similar ridge behind the last molar in the lower jaw; 
these ridges are very possibly used to supplement the minute teeth in eating. Posterior 
bony palate broad and produced far back; posterior nares rather narrow; internal 


FROM THE PHILIPPINE ISLANDS. 397 


pterygoid processes large, triangular, projecting far downward, in fact below the level 
of the tips of the incisors; external processes practically or quite obsolete, so that 
there are in this animal no enclosed pterygoid fosse. Bulle small, but not of abnormal 
structure. 

Lowei jaw exceedingly low, slender, and little curved. Coronoid processes very fine, 
slanted backwards. 

Teeth (Pl. XXXY. fig. 7) extraordinarily reduced, the dental armature in 
Rhynchomys being less in proportion to the size of the animal than in any other 
Rodent, perhaps even—apart from the Cetacea—than in any other toothed mammal. 
Incisors white above, pale yellow below; the upper ones not grooved, very short, 
narrow, slender, and forming the are of a very small circle, so that their roots come 
opposite the anterior end of the palatal foramina, and the chord of thie circle they 
describe is barely more than a third of the diastema. Molars 3, so minute that it is 
difficult to understand of what use they can be to the animal; m.! oval, flat-crowned, or 
with low indistinct cusps, but without quite young specimens it is impossible to make 
out for certain whether there is any true cuspidate structure; m.? about half the size 
of m.! and a shorter oval in outline. 

Lower incisors very slender, and, owing to the oblique set of their enamel-covered 
faces, they wear to an unusually fine point, as sharp as a needle. Lower molars 
very similar both in size and shape to those of the upper jaw. In one specimen 
the posterior lower tooth is altogether absent on both sides, so that there is only one 
molar present; this fact shows strikingly the tendency there is to a progressive 
reduction of the molar teeth. 

Type. Rhynchomys soricoides. 

This most remarkable genus, by its peculiar Shrew-like appearance, reduced teeth, 
elongate muzzle, and other characters, seems at first sight perfectly isolated from any 
other known group, and in my recent classification of Rodents it was made the type of 
a subfamily of the Muride, the Rhynchomyine!. But among the Rodents then 
included in the Murinee—on account of its strictly murine molars—there occurs the 
genus LHchiothrix?, a native of Celebes, which also has an elongated snout. This 
animal, of which the Museum now possesses two perfect specimens from N. Celebes, 


* PZ. S. 18965 p. 1017. 

? Described by Gray, P. Z. 8. 1867, its correct locality determined by Jentink, Notes Leyd. Mus. v. p. 177 
(1883); renamed by me Craurothriv, Ann. & Mag. N. H.[6] xviii. p. 246 (1896). As I have now joined those 
who think that names should be retained as originally spelt, whether classically right or wrong (except in the 
case of obvious misprints), | am now prepared to consider that Peters’s Bchinothria of 1853 does not preoccupy 
Gray’s Echiothrix of 1867, and therefore again recognize the latter term. Those who are not of this opinion 
must call it Craurothriz. That the missing out of the letter is not a misprint is shown by Gray haying 
written on the type skin what appears to be “ Mchithrix,” might be “ Echiothrix,” but is certainly not 
‘* Behinothria.” 


398 MR. OLDFIELD THOMAS ON MAMMALS 


collected by Mr. Charles Hose, I have carefully compared with Rhynchomys, and have 
come to the conclusion that, in spite cf the absence of any tendency towards a 
reduction in the dentition, there is a genuine relationship between the two forms. 
In the Celebean animal the general shape of the skull is very similar to that 
found in Rhynchomys: the peculiar anterior nasal bulging is present; the brain- 
case is similarly smooth and rounded; the supraorbital and temporal ridges, 
although present, are very small; the zygomatic root is slightly slanted back ; and the 
posterior palatal region is strikingly similar to that of Rhynchomys both in the breadth 
and shape of the posterior nares, and the entire suppression of the external pterygoids. 
The incisors again—or at least the upper ones!—in size, proportions, and position are 
more like those of Rhynchomys than of ordinary murines. 

On the other hand, the molars of Echiothriz ave absolutely murine, and show no 
trace of reduction or any other peculiarity. The third molar is, of course, present 
above and below, and is of full murine proportions. 

On the whole it seems probable that we have in Hchiothriz a form which bears to 
Rhynchomys very much the relation that Crunomys does to ydromys, being, as in that 
case, the first commencement of a line of modification which culminates in a genus 
sufficiently distinct to demand subfamily separation from the main trunk of the 
Murine. If this be true, it would then probably be best to include all the members 
of the diverging branch within the special subfamily, even if nearer to the trunk than 
to the extremity, and I would therefore suggest, as in the case of Crunomys, that 
Echiothrix should be transferred to the Rhynchomyine, a name which would be 
particularly suitable owing to the long snout being the most obvious character that 
the two genera have in common. 

It is, of course, just possible that when unworn teeth of Rhynchomys are examined 
they will show a structure quite incompatible with the view that this form is related to 
Echiothrix, but it seems to me that the many cranial characters which the two forms 
have in common render this possibility very unlikely. 


RuyNcwomys soricowes Thos. (Plate XXXI. fig. 2.) 


Size of a common Rat. Fur thick, close, and velvety, about 14 or 15 millim. long on 
the back. General colour dark olivaceous grey, becoming more yellowish in old age. 
Under surface dirty grey, not sharply defined, but becoming lighter and more sharply 
defined in old examples; a white patch sometimes present on the throat or chest. 


1 The lower incisors of Echiothriv are perfectly unique in being widely separated from each other terminally, 
so that, being also very long, their tips bite up on each side of the upper incisors, which project down between 
them. How far up they actually go in life on the sides of the muzzle cannot be determined without 
the examination of fresh or spirit specimens, but their splay is sufficient for the whole muzzle to close down 
between them. 


FROM THE PHILIPPINE ISLANDS. 599 


Sides of snout obscure whitish, top blackish. Eyes small, not noticeably ringed. Ears 
rather large, thinly haired, the anterior half of their outer and posterior half of their 
inner surfaces blackish. Wrists and metacarpals brown above, digits whitish or flesh- 
coloured. Hind feet similarly coloured. ‘Tail shorter than head and body, very finely 
ringed, clothed with short hairs, not pencilled terminally, blackish above, scarcely 
paler below, the extreme tip white in most specimens. 

Skull and teeth as above described. 

Dimensions of the type, measured in skin ( ¢ ):— 

Head and body 215 millim.; tail 146; hind foot (moistened) 41. 

Skull, see p. 399. 

Hab. Monte Data, 8000 feet. 

The following are Mr. Whitehead’s notes on this most peculiar animal. It is 
unfortunate that he has no positive knowledge of its habits or food, as its anomalous 
dentition is certain to be correlated with some food very unusual among Muride; very 
possibly, as Mr. Whitehead suggests, it eats caterpillars or worms, for it is difficult to 
imagine any vegetable food for which its reduced dentition and Shrew-like snout would 
be at all suitable :— 

“This interesting Shrew-Rat was obtained on the summit of Monte Data, where only 
five specimens were snared. I am unfortunately unable to give any account of the 
habits of this extraordinary mammal. ‘The Igorrotes told me that it lives on grass, 
which is probably untrue, the teeth apparently being quite unfitted for such food ;. 
insects and worms are probably the diet suited to such rudimentary molars. he eye 
is, comparatively speaking, small, which leads me to believe that Rhynchomys is & 
dinrnal-feeding Rat, like the true Shrews. 

“ Distribution. High mountains of Central Northern Luzon.” 


Putaomys PALLipus Nehring. 

a. 6. la Trinidad, Benguet Dist., N. Luzon, Feb. 9, 1894. 

b,c. 6 9. Cape Engafio, Lepanto, N. Luzon, May 1895. 

d. Monte Data, Luzon, Feb. 1895. 

The specimens sent by Mr. Whitehead all belong to the larger soft-haired form to 
which Dr. Nehring applied the name of P. cwmingi, var. pallidus, but which appears 
to me to be sufficiently distinct to demand specific recognition. 

When Dr. Nehring first suggested the name, Dr. Meyer considered him wrong in 
doing so, and, with some whitish and piebald specimens before him, quoted a letter of 
mine, informing him that the original series of P. cumingi also contained both black 
and piebald specimens, and that therefore the species was to be regarded merely as a 
very variable one. On now looking again at the original specimens in the Museum 
collection, I find, to my surprise, that there is among them a bad, but perfectly 


400 MR. OLDFIELD THOMAS ON MAMMALS 


typical, specimen of P. pallidus, received from Mr. Cuming in 1853, some time 
after Mr. Waterhouse described P. cumingi, of which it was noted at the time 
to be a “variety.” This specimen is, of course, that referred to in my letter to 
Dr. Meyer, it not having been up till now distinguished from the typical dark-coloured 
P. cuming?. 

P. pallidus differs from P. cumingi in its larger size, longer and much softer fur, 
and paler colour. It is, however, very variable in colour, as has been described by 
Dr. Meyer on his specimens, and as those of Mr. Whitehead confirm. One of the 
latter even has no dark saddle-mark, a characteristic that seems to be nearly invariably 
present. In the skulls also there is an astonishing degree of variability in the size and 
shape of the interparietal bone, a variability I have never seen equalled elsewhere. 
But I have quite failed to divide the forms into two or more races, as the characters 
drawn from the interparietal run altogether at cross purposes to those drawn from the 
external ones. 

“This splendid Rodent, larger and more powerful even than Crateromys schadenbergi, 
is, on the high mountains of North-west Luzon, much rarer than that species. In six 
months I obtained only four specimens, all of which were captured by the Igorrotes, 
aided by their dogs. This Rat, they told me, lived in old tree-trunks, and one specimen 
was slightly singed, having been smoked out of a hole in an old tree. The Phlwomys 
is also found on the coast-level, two of my specimens having been shot at Cape Engafo 
as they were ascending trees in the early morning. The Engafo pair have much 
shorter fur and are browner underneath than those obtained in the higher altitudes, 
but still show the same black markings cn face and shoulders; two of the highland 
specimens are without black markings, but are undersized and probably immature. It 
is possible that Phlwomys pallidus is a grey variety of P. cumingi, which is a brown- 
coloured animal, as we find three distinct varieties of C. schadenbergt. 

“ A grey Phleomys occurs in the island of Marinduque to the S.W. of Luzon, The 
specimen I saw was in a kerosene-oil tin on a steamer in which I was a passenger. 
This animal had a white face like those just mentioned from Lepanto. 

“ Distribution. Luzon and Marinduque. 

“ Tgorrote name, ‘ Eut-eut.’”—J. W. 


Mus EVERETT Giinth. 

a-c. 6. Monte Data, 7500 feet, Feb. 1895. 

This fine Rat was hitherto known only from a single specimen, the type, now in the 
British Museum, and Mr. Whitehead’s beautiful skins are therefore particularly 
acceptable. 

“Much commoner than the next species, which is found in the same locality,’— 
J. W. 


FROM THE PHILIPPINE ISLANDS. 401 


Mus uuzonicus Thos. 

Mus luzonicus Thos. Ann. Mag. N. H. (6) xvi. p. 163 (1895). 

a,b. 2. Monte Data, Lepanto, Luzon, 8000 feet, Feb. 1895. a, type. 

c. Yg. al. Lepanto Highlands, Luzon. Presented by Mr. Whitehead. 

Allied to, and of about the same size and dorsal colour as, the last species. Fur 
much longer and softer, the wool-hairs about 20 millim. long on the back, and the 
longer hairs from 30 to 40. General colour coarsely grizzled brown, resulting from a 
mixture of buffy yellow and black; the wool-hairs dark slaty basally, their tips for 
4 or 5 millim. buff, the long hairs black, but some of them with their extreme tips 
whitish. Under surface dull slaty buff, not defined on the sides; the hairs slaty 
basally, buff terminally. Head clearer greyish, owing to the tips of the shorter hairs 
being rather whitish than yellow. Eyes with an indistinct blackish ring, most marked 
posteriorly above. ars of medium length, very thinly haired, their backs blackish, 
finely edged with white. Upper surface of hands and feet hoary, some of the hairs 
blackish, and others (the majority) silvery white. ‘ail rather shorter than in Mus 
everetti, well haired, though not pencilled, coarsely scaled (scales 8 or 9 to the cm.,), its 
proximal half or two-thirds black above, paler below, its distal portion white all round. 

Skull (Pl. XXXVI. fig. 4) markedly distinguished from that of I. everetti, and 
perhaps from all other Rats of so great a size, by the reduction of the supraorbital 
ridges, which merely form a fine beading along the edges of the frontal, and practically - 
disappear halfway along the parietals. Brain-case smooth, round, and swollen; and 
this character is present all over the skull, which is unusually smooth and without 
ridges and angles. Posterior nares broad and open, the palatal edge opposite the 
hinder margin of m2 Bulle smaller than in MZ. everetti. 

Incisors yellow, not the dark orange of WV. everetti. Molars broader than in that 
animal, the Jaminze more simply transverse, and the outer cusp of each lamina less 
distinctly defined from the middle cusp. 

Dimensions of type ( 2 ) measured in skin :— 

Head and body 240 millim.; tail imperfect (of another specimen 200); hind foot 
(moistened) 47. 

Dimensions of skull of type, see p. 404. Another specimen has a basilar length of 
44 millim. by a greatest breadth of 28°6. 

Hab. Monte Data, Luzon. 

“Scarce on Monte Data, where only four specimens were obtained.’—J. W. 

It is curious that two large Rats of the group with white-tipped tails should inhabit 
the Data plateau; but, like as they are in size and colour, there can be no question 
that they are of perfectly distinct species. 


[s\) 
Q 


VoL. XIV.—ParT vi. No. 4.—June, 1898. 


402 MR. OLDFIELD THOMAS ON MAMMALS 


Mus DECUMANUS Pall., var. 
a. Moute Data, Luzon, Feb. 1895. 


This isa Rat so similar to some of the forms of Mus decumanus that, like the 
Felis domestica above referred to, I can only suppose it to be the slightly modified 
descendant of introduced examples. 


Mus rartus L., var. 


a. 2. Negros, 6500 feet. 

6. 3. Mindoro, coast-level, Dec. 1895. 

The single specimen of the Mus rattus group from Negros seems sufficiently like the 
Bornean variety to be provisionally referred to it. In many ways it has more the 
aspect of some of the Indian forms of the species, such as WM. rattus rufescens, than 
any other Philippine or Bornean Rat that I have seen. 

A coast-level example from Mindoro may also be placed here. Its differences from 
the highland Mus mindorensis are very striking. 


MUS MINDORENSIS sp. n.! 
a-e. 5 sks. Monte Dulangan, Mindoro, 5000 feet, Dec. 1895. 


A Rat of the group of Mus rattus, apparently forming a peculiar insular race. 

Size of Mus rattus or rather smaller. Fur straight, sleek, and shining. General 
colour very dark as compared with the ordinary eastern forms of the group, Mus 
neglectus, &c.; back a dark finely grizzled brown, the grizzling much finer than usual. 
The light colour in the grizzling is a deep orange, becoming rather more yellowish on 
the sides. Under surface whitish or dirty slaty grey, not defined from the upper 
colour, and not unlike in tone that of typical house-haunting specimens of Mus 
musculus. Face uniformly dark like the body, hairs round base of ears behind nearly 
black. ars rather short, almost naked, the hairs so minute that a lens is needed to 
see them at all. Hands and feet blackish above, the digits scarcely paler. Tail 
decidedly shorter than the head and body, smooth, very thinly haired, almost naked, 
finely scaled (about 10 rings to the cm.), uniformly black above and below. 

Skull very uniform in character throughout the series. Brain-case rounded, swollen. 
Supraorbital edges with the usual ridges rather weakly developed, and scarcely to be 
distinguished on the posterior half of the parietals. Interparietal large, its anterior 
edge slightly curved forward. Palatal foramina large and well open, reaching pos- 
teriorly just to the level of the front edge of the anterior root of m.1. Posterior edge 
of palate broad, squarish. Bulle rather smaller than in typical Mus rattus. 

Dimensions of the type, an adult male in skin :— 

Head and body 190 millim.; tail 163; hind foot (moistened) 32°5. 


‘ Preliminary diagnosis in Minutes of P. Z. 8. for June 15, 1897 (published June 19), 


FROM THE PHILIPPINE ISLANDS. 405 


Dimensions of skull, see next page. 
Type. B.M. 97. 3. 1. 4. 


This Rat is one of the group allied to Mus rattus, so widely distributed over the 
‘ast Indian Archipelago. The Bornean examples of the group I have provisionally 
termed J. neglectus, Jent., and have hitherto also used this name for Philippine 
specimens. ‘The five highland Mindoro skins before me are, however, so uniformly 
different from any other specimens seen that they evidently ought to have a distinctive 
name. 

On the other hand, as already noticed, a coast-level specimen from Mindoro is in 
no way separable from ordinary Philippine examples of M. neglectus. No doubt the 
highland forms are more or less indigenous, while those from the coast have been more 
lately introduced. 

‘“‘T obtained several specimens of a variety of Mus rattus as high as 5000 feet in 
the forests of Mount Dulangan, Mindoro, and also on the Canloan volcano in Negros 
at an altitude of over 6000 feet. Like all the forms of Mus rattus, they were a great 
nuisance, entering my tent at night and biting holes in my rice-bags, often running 
over my body. 

“The specimens from the two islands differ slightly in outward appearance of the 
fur. The Mindoro Rat is peculiar in being of a much darker brown on the back, and 
the belly is mouse-grey. The fur is fine and short, and the tail is nearly black. The 
Negros specimens, on the other hand, are more common looking, sandy brown on the 
back, with the underparts nearly white; the fur is also much longer, and the tail grey. 
Mus rattus seems to turn up in some form or other over the whole world, especially on 
high mountains.’—J. W. 


Mus curysocomus Hoffm. 
a. 3g. Monte Data, Lepanto, 8000 feet, Feb. 1895. 


This interesting species, which differs from almost every other member of the 
genus in the entire absence of sharp supraorbital edges or ridges, has hitherto been 
recorded only from Celebes. ‘The present specimens, however, seem to agree closely 
both with Herr Hoffmann’s description and figure, and also with the notes which, 
by the kindness of Dr. Meyer, I was allowed to take on the typical specimen when in 
Dresden. 

“Common in the potato-fields on the top of Monte Data.”—J. W. 


MUS EPHIPPIUM NEGRINUS subsp. n. 

a,b. 62. Negros, 6600 feet. a, type. 

c. ¢. Monte Data, Luzon, 8000 feet, Feb. 1895. 

Similar in essential characters to the small, coarse-haired, brownish or rufous animal 
3G 2 


404 MR. OLDFIELD THOMAS ON MAMMALS 


known as I. ephippium, Jent., but rather larger, much longer and softer furred, and 
more greyish smoky in colour. 

Fur long and soft, the wool-hairs about 15 and the longer hairs 18 millim. in length 
on the back. General colour dark smoky grey, almost blackish along the middle of the 
back, lightening to buffy or yellowish on the sides, Belly not sharply defined, the hairs 
slaty at the base, yellowish white at the tip. Hands and feet silvery whitish above. 
Tail nearly as long as the head and body, uniformly brownish, or slightly paler below. 

Dimensions of the type, an adult male, in skin :— 

Head and body (apparently much stretched) 155 millim.; tail 135; hind foot 
(moistened) 26°5, 

Skull, see below. 

‘This is evidently an insular highland form of the common little Rat spread over the 
Malay Archipelago, to which I have generally applied the name of Mus ephippium, 
but which will perhaps be found to grade into the earlier described Mus concolor, Bly. 
In any case, however, the highland form now described seems worthy of subspecific 
distinction. 

“Common among the Igorrote sweet-potato fields on the top of Monte Data.”— 
A Nie 


Skull-dimensions (in millim.) of Species of Mus, Batomys, and Carpomys. 


Mus | 
Mus lu- | Mus min- | ephippium| Batomys | Carpomys | Carpomys 
zonicus. | dorensis, | negrinus, | grant, a le pheurus. 
Raa oughh Hie te te tas Matera hye +c 44 36 | Chambas | 40-5. | 3038 | 86 
Basilard engi Mee. ote «0. bt iasvere. Sere isia 40-2 32°8 | tip 30°3.), 37 36:3 33 
Groenbeat bread ubiarere liye teytal-v-ese ecru aet- tae rel 25°7 19-5 16 22:2 24:5 23 
Nasals; Men eth v.08 sc. sueeitt «cs cofareiwie nb ans ei 19:8 145 13-1 19°5 16 14:5 
nape DL CACtiten ere: phate aroreaber eaves regret cone 59 4-7 3:6 5 5:6 46 
Imterorbitall breadth) eer en seeenenceeiec 7 6-1 5:2 55 5:2 61 
Imferpantetals enmity mic ispeiis oisreyenie aie 53 5-6 — 5 6-1 4:8 
& preadthin saat eaten 11°6 ila _— 9-6 11 13 
Anterior zygoma-root, length.............. 6-1 38 3°6 45 51 3:8 
Palate, length from henselion ............ 23 19 16 19 19 16-4 
MD issteM as. pict ec eetacck Meee ee ee 13°5 ilalo2) SEA 12:8 12 11 
Anterior palatine foramina, length.......... | 32 ii 6°5 8:5 78 TEE 
: : combined breadth. 3 2-6 25 31 2-7 28 
Length of upper molar series .............. | 93 6°5 5 78 88 6-1 
Lower jaw, condyle to incisor-tip .......... 33°2 26 22 30°2 29°5 26°6 
i: ONCOL Sinai cctv sce tia orca Rete rors 31 23-2 19-7 26-7 27 24 


FROM THE PHILIPPINE ISLANDS. 405 


Batomys Thos. 
Batomys Thos. Aun, Mag. N. H. (6) xvi. p. 162 (1895). 

General external form very much as in Carponys, but with a shorter, though 
similarly hairy, tail. Hyes surrounded by a distinct naked, or at least very finely 
haired, ring, a peculiarity which forms one of the readiest means of distinguishing 
Batomys from Carpomys externally. Fore feet rather elongated; pollex with a nail. 
Hind feet broad ; sole-pads as usual six in number, but all very large, and both the 
fourth (hallucal) and fifth (usually small and rounded) elongated like the sixth; pads 
not striated. Whole of heel hairy to the level of the hinder end of the last sole-pad. 
Fifth hind toe reaching to the base of the third phalanx of the fourth; hallux just to 
the base of the second toe. 

Skull (Pl. XXXVI. fig. 8) more elongate and murine than that of Carpomys, in 
general outline not unlike that of Eliomys quercinus. Brain-case small, face compara- 
tively long. Interorbital space rather narrow, its edges with only the slightest 
indication of ridges. Interparietal fairly large. Anterior edge of zygoma-root not 
projected forwards. Anterior palatine foramina large. Bulle small. 

Incisors narrow, smooth in front. Molars (Pl. XXXVI. fig. 5) in their pattern like 
those of Mus, not of Carpomys, but instead of being distinctly brachyodont, as are 
those of nearly all other Murines, they are more or less hypsodont, the crown at least 
as high above the bifurcation of the roots as it is broad. Molar laminz, as in Mus, 
3—2—2; transverse, not oblique; m.? and m.? with well-defined antero-internal 
supplementary cusps. JZ, and m.y also with distinct posterior mesial supplementary 
cusps. 

This genus, although with a striking external resemblance to Carpomys, is really 
more nearly allied to Mus, as its elongate skull and the pattern of its molars indicate. 
Its curious bare eyelids and hypsodont molars are, however, characters in which it is 
different from all the other Eastern arboreal genera. 


Batomys eranti Thos. (Plate XX XIII. fig. 2.) 


Batomys granti Thos, t. c. p. 162. 
a—-c, 2 adult and 1 immature. Monte Data, Feb. 1895. 


Size of a large Rat. Fur thick, close, and rather coarse. General colour coarsely 
grizzled fulvous and black all over above, the face, however, more greyish; posterior 
back and rump tending more towards rufous. Ears of medium length, more thinly 
haired than in Carpomys, their backs black or dark brown, Under surface dirty buff, 
not sharply defined; the bases of the hairs slate-colour throughout, though an 
indistinct whitish mesial line is sometimes present. Metacarpals and metatarsals 
brownish mesially, whitish laterally and on the digits. Tail thickly and uniformly 


406 MR. OLDFIELD THOMAS ON MAMMALS 


clothed (except for its body-furred basal half-inch) with dark brown or black hairs, 
some 7 to 9 millim. in length, the scales quite hidden. 

Skuil and teeth as already described. Palatine foramina just reaching backward to 
the front edge of m.1; palate ending behind opposite the posterior lamina of m.’. 

Dimensions of the type, an adult male in skin :— 

Head and body 204 millim. ; tail (doubtfully perfect) 121; hind foot (moistened) 35:5. 

Dimensions of skull, see p. 404. 

Hab. Plateau of Monte Data. 

Type. B.M. 95. 8. 2. 15. 

This interesting animal, which, with very much the general appearance of Carpomys 
melanurus, is entirely different in essential characters, 1 have named in honour of 
my friend and colleague Mr. W. R. Ogilvie Grant, by whom all the business matters 
connected with Mr. Whitehead’s expedition were managed, and who has himself 
worked out and described the magnificent collection of birds, which contained examples 
of no less than fifty new species. 

“This interesting new Rat was captured for me by the Igorrotes, with the aid 
of their small terriers ; it seems rare, only three specimens being obtained, Batomys 
granti is at first sight the same animal as the next species, Carpomys melanurus, but 
has a bare ring round the eye, and when alive is easily separated from Carpomys 
by this character. The two animals, however, which externally appear almost 
identical, have the teeth so different that they have been separated by Mr. Thomas 
into different genera. Found at 7000 feet on Monte Data. 

“ Distribution. Highlands of Central Northern Luzon.”—J. W. 


Carpomys Thos. 
Carpomys Thos. Ann. Mag. N. H. (6) xvi. p. 161 (1895). 

Form more or less as in such arboreal Murines as Hapalomys and Pithechirus, Fur 
thick and woolly. Pollex with a large nail; other digits, including the non-opposable 
hallux, with claws. Tail long, well haired. Mammez 0—2=4. 

Skull (Pl. XXXVI. figs. 6 and 7) with a large rounded brain-case and short face. 
Supraorbital region without sharp ridges or overhanging ledges. Interparietal large ; 
zygoma-root as in Crateromys, i.e. without any forwardly-projecting plate, the front edge 
vertical or even concave. Anterior palatine foramina fairly long. Bulle small. 

Teeth (Pl. XXXVI. fig. 3). Incisors smooth in front. First and second upper 
molars, as compared with those of Mus, each with an additional lamina, formed 
apparently by the normal posterior lamina being doubled round on itself. The last 
molar is normal, so that the laminar formula is 4—3—2. Both m.? and m.° have well- 
marked antero-internal supplementary cusps. In the lower jaw m., has an additional 
lamina in front, and both it and m., have well-marked posterior supplementary cusps, 
while the last-named has in addition an antero-external one. 


FROM THE PHILIPPINE LSLANDS. 407 


It is difficult to decide what are the exact relationships of Carpomys, and it can only 
be said that it adds one more to the list of Oriental genera of Muride modified for an 
arboreal life, such as Hapalomys, Pithechirus, Chiropodomys, and Vandeleuria. 

This new genus contains two handsome Dormouse-like species with long hairy tails 
and fluffy fur. Both are evidently of arboreal habits. 

With a certain superficial resemblance to each other, the two species of Carpomys 
may be readily distinguished by their differently-coloured tails, the extension of the 
body-fur on to that organ in C. melanurus, and by the very much larger teeth, both 
absolutely and proportionally, of the same species. 


CarRpoMys MELANURUS Thos. (Plate XXXIV. fig. 2. 
Carpomys melanurus Thos. Ann. Mag. N. H. (6) xvi. p. 162 (1895). 

a-d. 3 ad. & 1 yg. sks., g 2. Monte Data, 7000-8000 feet, Feb. 1895. 

Size about as in Mus rattus. Fur soft, thick, and woolly. General colour deep 
fulvous, coarsely lined with black. Under surface and inner sides of limbs dull 
yellowish white, the bases of the hairs slate. Ears of medium size, well haired, dark 
brown, nearly black. Limbs to wrists and ankles furred and coloured like body. 
Metapodials brown mesially, laterally and on the digits white. ‘Tail longer than head 
and body, its basal inch or two thickly furry like the body, and of the same colour ; 
the rest closely covered with shining black hairs, some 5 to 7 millim. in length, entirely 
hiding the scales ; not specially tufted at tip. 

Skull (Pl. XXXVI. fig. 6) with the nasals broad in front, abruptly narrowing back- 
ward. Interorbital region narrow, broader in front than behind, and the traces of ridges 
mounting on to the top, and approaching each other to within 2 millim. in the middle 
line. Palatal foramina parallel-sided, attaining at once their greatest width anteriorly. 
Palate ending opposite the front edge of m.°. 

Teeth broad and heavy, Incisors broad, slightly flattened in front in old specimens ; 
dark yellow above, rather more whitish below. Molars (Pl. XXXVI. fig. 3) very 
broad and large (see skull-measurements), their combined length exceeding that of the 
palatal foramina. 

Dimensions of the type, an adult male, in skin :— 

Head and body 197 millim.; tail 211; hind foot (moistened) 34:2. 

Skull, see p. 404. 

Type. B.M. 95. 8. 2. 12. 

“The black-tailed Carpomys differs much from the next species, C. pheurus, both 
in size, colour, and length of fur; in fact it has externally the appearance: of Batomys. 
On Monte Data, where both these new torms were obtained, it was more numerous 
than either Batomys or the next species. 

* Distribution. Highlands of Central Northern Luzon.’—J. W. 


408 MR, OLDFIELD THOMAS ON MAMMALS 


Carpomys PHxzURUS Thos. (Plate XXXIV. fig. 1.) 
a—-c. 3. ad. sks.,g¢ ¢. Monte Data, 7000-8000 feet, Feb. 1895. 


Size rather less than in C. melanurus. Quality of fur and general colour almost 
exactly as in that species. Ears rather smaller, less thickly hairy, and not prominently 
black. Belly-hairs dull buffy white to their roots, not slaty basally. Tail with the 
body-fur not extending on to its base more than in ordinary Rats, more thinly haired 
than in C. melanurus, so that the scales, which are very small, running about 15 to the 
centimetre, are visible through the hairs; in colour it is uniformly dark brown, 
occasionally approaching black, but never the deep shining black of C. melanurus. 

Skull (Pl. XXXVI. fig. 7), as compared with that of C. melanurus, with the nasals 
less expanded anteriorly and less abruptly tapering posteriorly. Interorbital space 
comparatively broad and parallel-sided, the rudimentary ridges not approaching each 
other on the top. Palatal foramina pointed in front, gradually broadening backward. 
Palate ending opposite to front of m.°. 

Teeth light and delicate. Incisors comparatively narrow. Molats, in marked 
contrast to those of C. melanurus, quite small in proportion to the size of the animal, 
but of the same essential structure. 

Dimensions of the type, an adult male in skin :— 

Head and body (stretched) 195 millim. ; tail 178; hind foot (moistened) 31. 

Skull, see p. 404. 

Type. B.M. 95. 8. 2. 14. 

“The brown-tailed Carpomys was somewhat rare on Monte Data. The Igorrotes 
used to hunt for the various Rats on Monte Data in small parties accompanied by their 
dogs, and spent most of the day at this—to them—curious occupation. ‘The animals 
that I saw captured were dug out from among the roots of trees by the aid of spears 
and choppers. ‘The flat table-top of Monte Data is much burrowed by various species 
of Rodents ; the Igorrotes, unlike the Kina Balu Dusans, not trapping Rats for their 
food. 

“ Distribution. Highlands of Central Northern Luzon.”—J. W. 


CRATEROMYS. 
Crateromys Thos. Ann. Mag. N. H. (6) xvi. p. 163 (1895). 

Size very large; general form not unlike that of Phlwomys. Claws smaller and tail 
bushier than in that genus. 

Skull, in a very general way, not unlike that of a gigantic Neotoma, strikingly 
different from that of Phlwomys. Muzzle slender. Zygomata squarely and boldly 
expanded. Interorbital region narrow, narrowing backward, edged with distinct but 
not exaggerated ridges, which pass backward on to the parietal and interparietal 
bones, and show no tendency to overhang the temporal fosse. Interparietal large, 


FROM THE PHILIPPINE ISLANDS. 409 


Palatal foramina long. Outer and inner pterygoids well developed. Bulle very small, 
though more inflated than in Phlwomys. 

Incisors not large in proportion to the size of the animal, flat in front. Molars 
(Pl. XXXVI. fig. 2) large and heavy, separated in the middle line by a distance less 
than their breadth; rather hypsodont, though less so than in Batomys; their pattern, 
while in the number of lamine and cusps essentially as in Mus, yet peculiar on account 
of the diminution or suppression of the external and the great development of the 
internal cusp of each lamina. On this account the longitudinal groove between the 
inner and middle cusps, in which the inner cusp-row of the lower molars works, 
approaches the centre of the tooth-row, instead of being close to its inner edge. As 
an accompaniment to this development of the inner cusp of each lamina, the point 
of separation between it and the centre cusp is marked by a sharp and deep infolding 
of the anterior enamel wall of the lamina; this notch is so deep in many cases as 
almost to cut the lamina in two. Below, the two halves of each lamina are strongly 
bent backward, so as to form a sharp angle with each other in the middle line. 
M., and m., with well-developed supplementary posterior cusps; m., with its posterior 
lamina sharply notched in behind, so as to give it a very definite cordate shape. 

Altogether the molars have a general resemblance to those of the remarkable Mus 
meyeri, Jentink, an animal which (as may be seen from the footnote +) I think should 
also form a peculiar genus. 


CRATEROMYS SCHADENBERGI (Mey.). (Plate XXXVI. fig. 2.) 


Phleomys (?) schadenbergi Mey. Abh. Mus. Dresd. 1894-5, no. 6 (1895). 
Crateromys schadenbergi id. op. cit. 1896-7, no. 6, p. 32, pl. xiii. figs. 3-6 (skull), xiv. (animal) (1896). 


a-c. Monte Data, Feb. 1895. 


This fine animal was first discovered by Dr. Schadenberg, but it is to Mr. Whitehead 
that our chief knowledge of it is due, as the former’s specimen was only a skin without 


' Lrnomys g. n. 

Form Rat-like. Feet short and broad ; pollex forming a large rounded projection of the hand, on the top of 
which the small nail is placed; hallux short, not opposable, its terminal pad large, covering nearly the whole 
of its under surface, its claw shorter, blunter, and more curved downward than those of the other digits ; 
palmar and plantar pads all very large. 

Molars (Pl. XXXVI. fig. 1) very large, the space between them less than their breadth. All three 
cusps of each lamina very strongly defined, the points of junction on each side of the central cusp marked 
anteriorly by a notch, and posteriorly by a backward projection of the enamel. .* and m.’ have, besides the 
usual antero-internal supplementary cusp, another one to balance it antero-externally, while the latter tooth 
has also a mesial supplementary cusp posteriorly. Lower molars very like those of Crrateromys, but m., has 
two, and m., has one supplementary external cusp. These characters may be seen in the figures of the skull 
quoted below and in that of the teeth on Pl. XXXYI. 

Type. Mus meyeri, Jent. N. L. M. i. p. 12 (1878); Cat. Ost. Leyd. Mus. (M. P.-B. ix.) p. 211, pl. vii. 
figs. 5-8 (1887) ; Hoffmann, Abh. Mus. Dresd. 1887, no. 3, fig. 2. 


VOL. XIV.—Part VI. No. 5.-—June, 1893. 3 


410 MR. OLDFIELD THOMAS ON MAMMALS 


a skull, which Dr. Meyer placed, not unnaturally, in the genus Phlwomys. On the 
arrival of Mr. Whitehead’s series, the form was generically separated by myself, 
and a little later Dr. Meyer published a second account of the animal, with coloured 
figures. No further description of it is therefore necessary. 

“Schadenberg’s great Rat seems to be fairly common among the high mountains of 
Central N.W. Luzon. Like most Rodents, it is of nocturnal habits, and therefore the 
domestic economy of this Rat, or perhaps Squirrel-Rat, is difficult to describe. The 
Igorrotes, however, captured a number of specimens for me, some, they said, from holes 
in trees, others from holes among the tree-roots ; they described the animal as feeding 
on fruits up in the trees, and not on the fallen ones. As this Rat was nearly always 
brought to me alive, 1 often allowed it to climb the pine-trees, which it did with 
perfect ease. In the day these animals tried to hide from the sun as much as possible, 
and I formed an opinion that they were dull and inoffensive creatures, until one day, 
directly an Igorrote opened the basket in which he carried the captured Rat, the 
animal sprang out, and was back in the basket again in a second, but the Igorrote’s 
thumb had the top nearly bitten off. The cry of the Crateromys is a curious * Thewo 
thewo thewd,’ uttered so shrilly that the notes might proceed from some of the 
peculiar forest insects. 

“Generally speaking, Crateromys is jet-black ; about 30 per cent, are of a beautiful 
white-grey, and some 15 per cent. piebald, black and white. This distribution of 
colouring has nothing to do with age, as both grey and black young ones were 
obtained. ‘Che Igorrote name for this curious mammal is ‘ Bi-ut.’ 

“ Distribution. High mountains of Central Northern Luzon.”—J, W. 


BUBALUS MINDORENSIS Heude. 

Bubalus mindorensis Heude, Mém, Hist. Soc. Chin. ii. p. 50 (1888); Meyer, Abh. Mus, Dresd. 

_ 1896-7, no. 6, p. 12 (1896). 

Probubalus mindorensis Steere, P. Z. 8. 1888, p. 415. 

“ This interesting little Bovine is not uncommon in the huge virgin forests that cover 
nearly the entire island of Mindoro. It is, however, difficult to hunt the animal 
successfully, unless a number of beaters, accompanied by good dogs, are employed. I 
foolishly followed a professional (!) native hunter about for several days; but, although 
we found a number of fresh tracks, we never saw a sign of a ‘Tamarau.’ The 
‘'Tamarau,’ as the natives name this animal, is also found high up on the mountains. 
I have seen regular tunnelled pathways through the thick bamboo undergrowth which 
covers the mountain-sides above 6000 feet. But the animal is so small that one has 
to bend double or go on one’s hands and knees, making it quite impossible to follow up 
the tracks. On moonlight nights the ‘Tamarau’ might be heard bellowing on the 
mountain-side, generally far away and above my mountain-camp. The aboriginals of 
Mindoro told me that they never attack the ‘'Tamarau,’ being too much afraid of it ; 


FROM THE PHILIPPINE ISLANDS. 411 


the only reduction of its numbers is caused by a few sporting Spaniards and one or 


two professional Indian hunters. 
‘« Distribution. The island of Mindoro.”—J. W. 


SUS CELEBENSIS PHILIPPINENSIS Nehr. 

a, Head-skin and skull, g. Cape Engaiio, N. Luzon. Presented by Mr. Whitehead. 

“This Pig may be said to be ubiquitous throughout the whole Philippine group, 
passing the entire day in seclusion in the forests, and sallying forth at night into the 
maize- and rice-fields, where it does much damage. 

- “ Native name ‘ Babui.’”—J. W. 


EXPLANATION OF THE PLATES. 


PLATE XXX, 
Fig. 1. Harpyionycteris whiteheadi (p. 384). 
. Nannosciurus samaricus (p. 389). 


= 
59 
Lo 


PLATE XXXI. 


Fig. 1. Celenomys silaceus (p. 391). 
Fig. 2. Rhynchomys soricoides (p. 398). 


PLATE XXXII. 
Chrotomys whiteheadi (p. 392). 


PLATE XXXIII. 


Fig. 1. Crunomys fallax (p. 394). 
Fig. 2. Batomys granti (p. 405). 


PLATE XXXIV. 
Fig. 1. Carpomys pheurus (p. 408). 
Fig. 2. 5 melanurus (p. 407). 
PLATE XXXvV. 
Skulls and Teeth of Philippine Mammals. 
Figs. 1-4. Harpyionycteris whiteheadi (p. 384), fig. 2 nat. size, figs. 1, 3, and 4 twice 
nat. size. 


Fig. 5. Crunomys fallax (p. 5394), upper and lower molars, much magnified. 
Hig. 6: a x skull, nat. size and twice nat. size. 


412 


ON MAMMALS FROM THE PHILIPPINE ISLANDS. 


Fig. 7. Rhynchomys soricoides (p. 398), upper and lower molars, much magnified. 
Figs. 8, 9. Chrotomys whiteheadi (p. 292), molars, magnified, and skull nat. size. 

Fig. 10. Rhynchomys soricoides (p. 398), skull, nat. size. 

Figs. 11, 12. Celenomys silaceus (p. 391), molars, magnified, and skull, nat. size. 


Fig. 
Fig. 2 


Fig. : 


5 


Fig. 


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Fig. oO. 
Fig. 


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Fig. 
Fig. 


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8. 


PLATE XXXVI. 
Skulls and Teeth of Philippine Mammals. 


1. Lenomys meyert (p. 409), upper and lower molar teeth, much magnified. 

2. Crateromys schadenbergi (p. 409), upper and lower molar teeth, much magnified. 
3. Carpomys melanurus (p. 407), upper and lower molar teeth, much magnified. 
4, 
5 
6 
7 


Mus luzonicus (p. 401), skull, nat. size. 
Batomys granti (p. 405), upper and lower molar teeth, much magnified. 


. Carpomys melanurus (p. 407), skull, nat. size. 


* pheurus (p. 408), skull, nat. size. 
Batomys granti (p. 405), skull, nat. size. 


All the enlarged figures of the molar teeth, upper and lower, are of those of the 
right side; the figures of the upper molars are placed on the left, and those of the 
lower on the right. 


Trans. Took. he. Vol. NWIG 2 OG 


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SKULLS AND TEBTH OF PHILIPPINE MAMMALS 


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CARPOMYS MELANURUS CARPOMYS PHAURUS 


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SKULLS AND TEETH OF PHILIPPINE MAMMALS 


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CONTENTS. 


VIII. On the Mammals obtained by Mr. John Whitehead during his recent Expedi- 
tion to the Philippines. By Otnriztp Tuomas. With Field-notes by the 
Collector. (Plates XXX-KXXVI.) . ....... =. . page 377 


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poraraea] 


IX. On the Lepidosiren of the Amazons; being Notes on five Specimens obtained 
between 1895-97, and Remarks upon an Example living in the Paré Museum. 
By Dr. Emm A. Gortpi, C..Z.S., Director of the Paré Museum. 


Received September 16, 1897, read December 14, 1897. 


[Pirates XXXVII. & XXXVIII.] 


ConTENTSs. Page 

ii, ihimwihnnia oapgnobvadnosaddposucdduoses oO KoOn HOO nODOCOTCD 413 

Il. Geographical Distribution ...........2 ee cesses eee eee ee eens 414 
IM, Sex and Measurements «2... .02200 cece ce cece wee ee cree neers 414 
IV. Situation of the Anus, and external Colour .........+..++s0+-e0es 415 
VY. Segmentation of Limb-axis 2.1... ..: cece cere eee teen ees 416 
VI. System of Lateral Lines...........0c eee e cece cece cece cee eeeee 416 
VII. Popular Names of Lepidosiren and its Habits in a free State ........ 417 
VIII. Habits of Lepidosiren in Captivity .......... se eeeeee eee nee 418 
Explanation of the Plates .......+.csesese reese senses eteees 419 


I, InrRopuction. 


THE undeniable deficiency of original investigations upon the biology of the Amazonian 
Lepidosiren, made by naturalists and scientific travellers in the native country of this 
interesting Dipnoan fish, will, I think, sufficiently justify the following communication. 
Sufficient proof of this deficiency is found in the recent memoir by Prof. E. Ray 
Lankester 1, in which the following passage occurs (p. 19):—“.... We have no drawing 
or record of freshly-killed or living specimens of Lepidosiren.” On the other hand, a 
considerable number of articles and notes have been published during the last few 
years by myself and others relating to the general features of the natural history of 
Lepidosiren and its allies, and to the history, description, and number of specimens 
existing in the museums of the Old and New Worlds, so that I may take it for granted 
that these particulars are known to the public, and that I may be allowed to restrict 
myself to such a short account of the subject as is indicated by the title of this paper. 

The rediscovery of the Amazonian Lepidosiren, together with the elucidation of its 
habitat, distribution, and mode of life, formed one of my principal projects from the 
time I assumed my present position (1894). I will not here repeat the details of my 
efforts; it will be sufficient to state that I organized a methodical propaganda all over 

1 «On the Lepidosiren of Paraguay, and on the External Characters of Lepidosiren and Protopterus,” 
Trans. Zool. Soc. Lond. xiv. p. 11 (1896). 

VOL. XIV.—PART vil. No. 1.—August, 1898. 31 


414 DR. E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS. 


the native country of Lepidosiren, and that these efforts were at last successful. Three 
years’ continuance of them put into my hands five specimens of the celebrated Dipnoan, 
one of which was living—material more important than all that was then existing else- 
where in the scientific world. 


II. GEOGRAPHICAL DISTRIBUTION. 


My original supposition that Lepidosiren paradoxa would probably exist and be 
found all over Amazonia, and that the geographical distribution would be much larger 
than was generally admitted, proved to be right. My first specimen I got from the 
Atlantic coast of the island of Marajé 1, certainly a somewhat remarkable locality. A 
second small specimen was brought to me from Santarem, the exact locality being 
Igarapé Ayayd, Fazenda Taperinha, in the immediate neighbourhood of that town ®. 
Two larger specimens I got afterwards from Obydos, the exact locality being Parana 
de Baixo, likewise not very far from Santarem. One of these specimens, captured 
with a fishing-hook, lived for several hours, and a telegram from Obydos, sent by 
Senador M. F. Machado (whose interest and help in this and similar ichthyological 
matters deserve public acknowledgment), made me hope to receive it alive by the next 
steamer of the Amazon Company. However, this hope was not fulfilled; the specimen 
died, but was saved as a spirit-specimen. From Obydos finally I received a fifth 
specimen, captured by Deputado Lourenzo Valente do Conto. This example has lived 
now nearly four months in the Para Museum, and seems to be quite well, being treated 
with great attention and care. 

In order to bring up to date my sketch of the distribution of the Amazonian 
Lepidosiren, as recently published by me 4, it is necessary to add the localities Santarem 
and Obydos. This done, it will easily be seen at a glance that the localities form almost 
a continuous chain along the main stream, entering even some of the more important 
affluents (the Ucayale, Madeira, Rio Negro, Tapaj6z) and the Atlantic side of Marajo. 
Thus my original supposition has already assumed the shape of a positive fact. 


III. Sex anp MEASUREMENTS. 


Four of my specimens I judge to be females. Designating the specimens, in the 
succession and order above indicated, with the letters A to EK, they would be the indi- 
viduals A, B, D, E. If, on the one hand, the question seems somewhat problematical 


1 The specimen is now in the British Museum. The details of its capture are recorded in ‘Nature,’ 
vol. liv. p. 270 (1896). 

2 In September 1896 I visited the spot where it was obtained. It was a papyrus-meadow, submersed for 
several months every year. I made a tolerable photograph of the spot where the specimen had been captured. 

3 T am indebted for this specimen to Dr. Augusto Olympio, formerly ‘ Deputado,’ and now Director of Public 
Instruction of the State of Pard (January 18th, 1897). 

“ Boletim do Museu Paracnse, i. p. 442 (Oct. 1896). 


DR. E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS. 415 


in regard to the individual B, in which small size and youth may perhaps hide the 
characteristic exterior sexual signs, on the other hand, there is no doubt that specimen 
A from Marajé Island (now in the possession of the British Museum) is an adult 
female, the ovaries having been rendered visible by a section. Individual C, from 
Obydos, is evidently a male, as it shows the villi distinctly on the left abdominal fin. 
The right abdominal fin exists only in the form of a short prominence or stump, 
having been probably bitten away by some carnivorous fish. 

We have, therefore, a numerical proportion of 4 females to 1 male. This may be 
purely casual, but the possibility cannot be denied that there may exist in this Dipnoan 
a real predominance of females over males. Unfortunately I do not possess any 
information upon this point as regards the large collection of Paraguayan Lepidosiren 
recently made by Dr. Bohls and sent to Europe. 

Concerning the measurements of my five Amazonian Lepidosirens, I have drawn up 
the following table :— 


A. 59 cm. D. 70 cm. 
B. 39 cm. E. 60-70 cm. [approxim. ]. 
C. 59 cm. 


Individual B, from Santarem, is only 32 mm. in dorso-ventral diameter; individual 
C 49 mm., having a circumference of 14:8 cm.; individual D is 70 mm. in dorso- 
ventral diameter. 


LV. SITvaTIon oF THE ANUS, AND EXTERNAL CoLour. 


The asymmetrical position of the anus (on the left side, near the pelvic fin), already 
noticed by the original describers of Lepidosiren paradoxa, is repeated again on all my 
five recent individuals. 

As regards the external colour of Lepidosiren, it seems to me that the difference 
between dark and brown individuals ought to be considered as a secondary matter, 
often merely produced by the effects of alcohol, and not peculiar to the living animal. 
This is shown by my four spirit-specimens A-D. The specimens A and B are now 
decidedly and uniformly dark, the former nearly black, the latter of a slaty colour. 
Individual C is now dark above and brownish below; on the individual D brownish 
spots, and more or less extended patches of the same colour, are still to be observed on 
the dorsal side. I am personally convinced that the brownish colour is, as a rule, 
predominant in the living and fresh animal. The picture of our living Lepidosiren 
exhibited before this meeting of the Zoological Society has been executed with great 
care, and is highly instructive upon this point. 

The following passage in Prof. Lankester’s memoir (Trans. Zool. Soe. vol. xiv. p. 24) 
refers to this subject :— The most striking and uniform difference which I have observed 


is the colour: the Paraguayan specimens are black, the Brazilian specimens are clear 
312 


416 DR. E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS., 


brown. There is, however, little doubt that this is largely, if not altogether, due to the 
longer action of alcohol and sunlight upon the Brazilian specimens.” But this should 
be taken in connection with my observations on the subject taken in loco and from 
fresh material. 

YV. SEGMENTATION OF LIMB-AXIS. 


I have already had occasion to state my opinion upon this question in my letter to 
Dr. A. Giinther, published in ‘Nature’ (liv. p. 270). In the first individual sent 
to me from Marajé, I discovered that the segmentation, considered by Prof. Ehlers to 
constitute a peculiarity of his Paraguayan Lepidosiren articulata, exists also in the 
Amazonian Lepidosiren, as clearly shown by Prof. Lankester in the above-mentioned 
memoir. The segmentation of the cartilaginous limb-axis evidently escaped the notice 
of the first describer Bischoff, but this oversight does not materially detract from the 
value of the excellent memoir upon Lepidosiren issued in 1840 by the celebrated 
anatomist of Vienna. 

VI. System or Lateran Lines. 


It seems that nobody has added anything more to the knowledge of the lateral-line 
system since Bischoff’s memoir, and that anatomist only mentioned and figured the 
lateral line of the head-region of Lepidosiren. The recent figures of Lepidosiren and 
Protopterus contained in Prof. Lankester’s memoir show what I consider to be a 
median lateral line on each side of the body; but in the text of his article no allusion 
or description is found referring to this point. Although surprising, however, this 
circumstance may be easily understood. Dead specimens of Lepidosiren generally 
become rapidly dark and the lateral lines cannot be so distinctly perceived, as I have 
had occasion to verify in my own spirit-specimens. Fig. 2, Pl. XXXVIL., taken from 
our living Amazonian Lepidosiren, shows most distinctly six lateral lines—that is, 
three on each side of the body, running longitudinally and in a straight line from 


head to tail. 
a. Median line. 


&. Dorsal line. 
c. Ventral line. 


The best-developed line, being nearly uninterrupted, is the median, giving out from 
time to time small vertical branches, generally two together, towards the dorsal line. 
The dorsal] line is less developed, being composed of longer or shorter, straight or 
curved marks, the longitudinal and lineiform arrangement of which is recognized 
without difficulty despite the numerous interruptions. It seems that these marks are 
small lateral branches, similar to those of the median line, but directed downward in 
the places where the dorsal line remains better preserved. Very well developed is also 
the ventral line, emitting vertical branches downward, nearly reaching the true yentral 
margin and almost touching the corresponding branches of the other side. When our 


DR. E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS. 417 


living Lepidosiren exposes to view the ventral side, the appearance may very well be 
compared to that of a rope-ladder. Probably the same relation exists between the 
dorsal and median lines of both sides of the body. 

Our figure (Pl. XXXVII. fig. 2) gives an adequate though not diagrammatic 
reproduction of the lateral-line system of Lepidosiren. Whether or to what degree 
these lateral branchings may be related to a metameric arrangement-I cannot say. 

In any case it is here shown that Lepidosiren possesses a complicated system of 
lateral lines, hitherto unknown—a system which I do not know to exist in the 
same high development in any other living species of fish. It would be very 
advisable to pay special attention to this point when examining specimens of the 
African Protopterus and the Australian Ceratodus. 


VII. Poputar NAMEs or LEPIDOSIREN AND ITS Hapits IN A FREE STATE. 


Between Obydos and Para it seems that the popular names most frequently used to 
designate the Lepidosiren paradowa are “ tariira-boia ” and “ piraruci-boia ”; sometimes 
the abridged forms “boia” or “cobra” are also employed. ‘he first name is 
composed of “ tariira,” the Amazonian term for the freshwater fish which is called 
“trahira” or “traira” in South Brazil (Macrodon trahira). ‘The first half of the 
second name consists of the native name for Arapaima gigas. Senador Machado 
informs me that these trivial names have their origin in the fear of the native 
fishermen, who state that our Dipnoan bites severely, and are convinced that it is 
poisonous. 

As yet we know but very little about the habits of Lepidosiren in a natural state. 
From the information kindly given to me verbally and in letters by Senador Machado and 
Civil Engineer Vicente Chermont de Miranda, I may extract the following :—Lepidosiren 
paradoxa is an inhabitant of submersed regions, where the water is shallow and does 
not possess a depth of many metres. Both my informants believe it to be more common 
than generally supposed, but the only chance of obtaining it is during the Amazonian 
summer, 7. é. the dry season. When the water nearly disappears and only small pools 
are left in the natural holes and depressions, the Lepidosiren is found left in them. 
As it is fond of disturbing the water of these pools, beating it with its tail and making 
violent evolutions with its eel-like body, the search for it then becomes easier, and 
only in such localities is the opportunity afforded of securing living specimens intact. 
The locality in which the first Marajé specimen was caught (Fazenda Dunas), and 
which I visited in September 1896, was in such a condition'. I saw the pool between 


 Senhor Vicente Ch. de Miranda informs me that in December 1894 he observed on his possessions on the 
Atlantic side of Marajo, in the locality called “ Rego do Jacaré-magro,” 23 kilometres distant from the coast, 
two specimens of “an unknown fish,” which he recognized afterwards as identical with the Lepidosiren sent 
to me. One was 41 centimetres long, the other 27 centimetres; the former was no doubt a female. 


418 DR, E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS. 


the roots of some papyrus-tufts, not more than a step distant from a long ditch, which 
at the midsummer period of my visit had sufficient water for a boat. 

The question arises what becomes of the Lepidosirens when even these pools dry 
up. ‘There is very little doubt that they hide in the lower regions of the mud, and 
thus survive the drying of the pools. ‘This is also the opinion of my above- 
mentioned informants, and I agree with it entirely, the more so as it establishes a 
parallel with the summer lethargy of the African Protopterus annectens. My two 
informants intend to clear up the matter, and to dig out the torpid Lepidosirens from 
the dried-up pools. 

The fact that two of my specimens from Obydos had been caught with a hook 
baited with fish certainly proves, as Senador Machado writes, that Lepidosiren paradoxa 
occasionally subsists on this kind of food. 


VIII. Hasits of Leripostren in Captivity. 


On the 27th April, 1897, our establishment received the news that Senhor Deputado 
Lourenzo V. do Conto had brought from Obydos a rare living fish, which was destined 
for the Museum. Sending one of the native keepers of our modest Zoological Garden 
for it, I asked him on his return what kind of fish he had brought. He replied a 
“poraqué” (Gymnotus electricus). Thinking that the creature was really what he 
said, and somewhat surprised that so common a fish as Gymnotus electricus should be 
spoken of as “a rare fish” worthy of being carried from Obydos, while plenty of them 
existed all round the city of Parad in every ygap6 pool, I ordered it to be put in the 
aquarium together with our other living Gymnoti. There the captive remained, 
through this singular mistake, for nearly a month. What was my surprise when one 
day we discovered that the supposed “ poraqué ” was in reality a Lepidosiren paradoaa! 
The prisoner’s condition was at once ameliorated by removing him to a separate 
aquarium in the Museum-building. Since that time we have had the specimen con- 
stantly under our personal observation, and it seems to be thriving and in good health. 

The specimen arrived with a large wound on the left side, nearly in the middle of 
the body. I do not know the nature and cause of the wound, but I have the pleasure 
of stating that it is now smaller, and that probably in a few weeks it will have perfectly 
healed. The tail also shows the results of some former mutilations, which is 
frequently the case with nearly all creatures cohabiting with such voracious colleagues 
as the “piranhas” (Piraya) and “‘jacarés” are. It is an interesting fact that our 
Lepidosiren lived nearly a month together with a number of large and smail 
Gymnoti without the slightest sign of having been injured by these companions, which 
certainly cannot be called agreeable. Nevertheless I have no intention of repeating 
the experiment. 

Our Lepidosiren now lives in a large glass aquarium, such as is commonly used in 
biological stations. The bottom has a stratum (1 decimetre high) of mud, leaves, and 


DR. E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS. 419 


other vegetable matter. A vigorous growth of alge helps to increase this stratum. 
The remainder is filled with fresh water, which is renewed every fourth day. As food, 
small pieces of meat, fish, ampullarias, crustaceans, as well as mandioca-roots, are 
constantly offered to it, but up to this time no one has seen our Lepidosiren actually 
take any of these things. It is, however, in a better state of nutrition than when it 
it arrived ; it is decidedly fat and round. ‘Therefore it eats something, and it probably 
does so when it burrows half the length of its body in the mud, as frequently seen. 

For respiration it comes from time to time to the surface of the water, and puts out 
a portion of the head. ‘The operation lasts several seconds. In the large aquarium 
this act is only repeated at intervals of several hours, but when placed in a smaller 
one—such as I used for my photographs of the living animal—the respirations at the 
surface are much more frequent, the intervals being perhaps of a quarter of an hour’s 
duration. The respiration is sometimes singularly prolonged. When descending, a 
series of air-bubbles is generally expelled from the branchial aperture on each side of 
the anterior part of the body. The whole body is covered with a viscous or gluey 
substance, which fills the water with whitish flakes, when the Dipnoan executes more 
rapid and violent evolutions (see my photographs, Pl. XX XVIIL.). 

During the day and when undisturbed our Lepidosiren is a quiet and passive 
creature, not changing its curled position for hours. Only once we have noticed it 
attempt to bite the finger of the keeper. It remains generally indifferent even when 
small living animals are offered it with the pincers. When disturbed it executes most: 
elegant and varied evolutions, and shows itself to be a first-class swimmer, at least as 
good as Anguilla and Murena. 

I had the rare pleasure of exhibiting the living Lepidosiren paradoxa at a public 
meeting of our Museum Society in Para on the night of June 3rd, 1897, which was 
honoured with the presence of the State’s Governor, Dr. Paes de Carvalho, a distin- 
guished physician, and of a large number of persons interested in our scientific work. 
Some days afterwards I received from Dr. O’Connor, of Oxley, Queensland, the news 
that living specimens of Ceratodus (the Dipnoan of the opposite side of the southern 
hemisphere) had been caught for the first time in Australia. 

Pard, August 26th, 1897. 


EXPLANATION OF THE PLATES. 


PLATE XXXVII. 


Fig. 1. Lepidosiren paradora, half natural size. 

Fig. 2. Outline of the body, showing the lateral lines. 

Fig. 3. Head seen from below. 

Fig. 4. A portion of the body, side view, in order to show the exact aspect of the 


420 DR. E. A. GOELDI ON THE LEPIDOSIREN OF THE AMAZONS. 


’ 


“lozenge-shaped areas” on the living animal. The lateral lines of this 
portion are accurately represented. 


Fig. 5. Lepidosiren paradowa during respiration on the surface. 


PLATE XXXVIII. 


Figs. 6-13. Evolutions of the living Lepidosiren paradowa in a small aquarium (from 
instantaneous photographs taken by myself). Figs. 6 and 7 show the 
Dipnoan in the act of respiration on the surface. 


Ve 


— 


—_ 
a 5 


CLLLE wz MEd 


P.J.Smit hth 
LE PIDOSIR§ 


Trams, Tod, Jee. Ve AW IC AXAVIL 


7 ah Ser er ee OT eee 
Re et ee fg 


Ss ee oe ee a 


aon 2 SN 


~ PARADOXA. 


Mintern Bros imp. 


Trans. Dock. Foo PANN IU, XXXVUL 


PJ.Smit. lith Mintern Bros imp 


EVOLUTIONS OF THE LIVING LEPIDOSIREN PARADOXA. 


bo 42h--] 


X. On a Collection of Fishes from the Rio Jurua, Brazil. 
By G. A. Bounencer, F.R.S., F.Z.S. 


Received December 8th, 1897, read February lst, 1898. 


[PLates XX XIX.-XLII.] 


THE Collection dealt with in this paper, and acquired by the Trustees of the 
British Museum, was made, in July 1897, by Dr. J. Bach, of La Plata, in the Rio 
Jurua, an affluent of the Amazons, the Fish-fauna of which had not been previously 
explored. The list here given will fill up an important gap in our knowledge of the 
distribution of South-American Fishes. As many as nine new species were 
discovered by Dr. Bach in the comparatively short time he was able to devote to this 
exploration. No doubt a more extensive collection in the same river, which I believe 
it is his intention to make in the near future, if circumstances permit, would 
result in very numerous additions to this list, which evidently contains but a small 
proportion of the representatives of this important class of Vertebrates in the Rio 


Jurua, 
SCLENID. 
1. Sci#Nna Amazonica Cast. 
PLEURONECTID. 
2. SoLzA FISCHER Stdr. 
3. SOLEOTALPA UNICOLOR Gthr. 
SILURIDZ. 


4. Sorvupim Lima Bl. Schn. 


5, PLATYSTOMA JURUENSE, sp. n. (Plate XXXIX.) 

Upper jaw projecting very slightly beyond the lower. Bands of small, equal, 
villiform teeth in jaws and palate; vomerine band as broad as premaxillary, single, 
notched in the middle, very narrowly separated from the much narrower palatine 
bands. Depth of body 5% times in total length, length of head 33 times. Head 13 
as long as broad, its greatest width 12 width of mouth; fontanelle very small; eye 

VOL, XIvV.—PART Vil. No, 2.— August, 1898. 3K 


499 MR. G. A. BOULENGER ON FISHES 


extremely small, its diameter 4 times in interorbital width, a little nearer opercular 
border than end of snout; occipital process narrow, not reaching basal bone of dorsal 
spine; maxillary barbel extending nearly to extremity of ventral; inner mandibular 
barbel extending to middle of pectoral, outer to middle of ventral. Branchiostegal 
rays 11. Dorsal I 6, originating at equal distance from the end of the snout and the 
middle of the adipose fin; spine rather feeble, not serrated, ? length of head. Adi- 
pose fin as long as base of dorsal or anal. Pectoral not reaching ventral ; spine feebly 
serrated, 2 length of head. Anal 14. Caudal deeply forked, both lobes produced 
into long filaments. Yellowish brown, with 9 equidistant dark brown cross-bands ; 
head dark brown above. 

Total length 190 millim, 

A single specimen. 


6. PimeELopus MAcuLaTus Lacép. 

7. PimeLopus Mopestus Gthr. 

8, CENTROMOCHLUS HECKELI Filippi. 
9. CrTopsis caNDIRU Ag. 


10. OxyDORAS STENOPELTIS Kner. 


11. OXYDORAS TRIMACULATUS, sp. n. (Plate XL. fig. 1.) 


No teeth in the upper jaw. Depth of body 43 times in total length, length of head 
32 times. Snout compressed, pointed, covered with skin ; posterior nostril close to the 
eye; diameter of eye nearly equal to length of snout, 3 times in length of head, 
12 interorbital width; cheeks and opercles covered with skin; skull striated above, 
granulate on the sides; fontanelle not produced as a groove posteriorly; bases of 
the six barbels united by the fold of the lower jaw; maxillary barbels branched, 
extending to base of pectoral spine; mandibular barbels short. Gill-cleft extending 
to below posterior border of eye. Humeral process striated, broader than and half as 
long as pectoral spine, obliquely truncated posteriorly. Pectoral spine as long as or 
slightly longer than dorsal, 13 length of head, extending to middle of ventral, very 
strongly serrated, especially on the inner side. Dorsal 15; spine feebly serrated in 
front, strongly behind, equally distant from the end of the snout and the adipose fin. 
Adipose fin half as long as base of anal. Anal 13. No shields between the dorsal 
fins or on the belly. Lateral shields 31-32, nearly half as deep as the body, with 
serrated border and moderately strong hooked spines. Caudal deeply bifurcate. 
Yellowish ; a black spot on the dorsal, involving the base of the spine and of the first 


FROM THE RIO JURUA. 423 


three rays; a small, elongate, horizontal black spot at the base of each lobe of the 
caudal. 

Total length 62 millim. 

Three specimens. 

Closely allied to O. stenopeltis Kner, which differs in the presence of an 
occipito-nuchal groove, the smaller eye, the presence of dorsal shields, and the absence 
of black spots on the fins. 


12. OxyDORAS TRACHYPARIA, n. sp. (Plate XL. fig. 2.) 


No teeth in the upper jaw. Depth of body equal to length of head, 33 times in total 
length. Snout rounded, rugose except in the internarial space on each side ; posterior 
nostril close to the eye; diameter of eye nearly equal to length of snout, rather more 
than 4 length of head, equal to interorbital width ; preeopercle, subopercle, and opercle 
bony, rugose ; cranial bones granulate; fontanelle not produced as a groove posteriorly ; 
barbels not fringed (or, rather, maxillary barbels with a single basal barb), their bases 
united by a fold of the lower jaw; maxillary barbel not reaching opercular cleft ; 
mandibular barbels short.  Gill-cleft extending to below posterior border of eye. 
Humeral process granulate, broader than and half as long as pectoral spine, obliquely 
truncated posteriorly. Pectoral spine as long as dorsal, as long as head, strongly 
serrated on the inner side, rather feebly on the outer. Dorsal I 6; spine rather 
strongly serrated on both sides, a little nearer the adipose fin than the end of the 
snout. Adipose fin half as long as anal. Anal 13. No shields between the dorsal 
fins or on the belly. Lateral shields 33-34, nearly half as deep as the body, with 
serrated border and rather feeble hooked spines. Caudal deeply bifurcate. Pale olive 
above, whitish on the sides and beneath ; fins white. 

Total length 93 millim. 

Two specimens. 

This species, which further demonstrates the untenability of the genus Hemidoras as 
distinct from Oxydoras, stands nearest to H. natterert Stdr., which differs in the fringed 
maxillary barbels, the more forward position of the posterior dorsal, and the smaller 
lateral shields. 


13. OxyporAS BACHI, sp. n. (Plate XL. fig. 3.) 

No teeth in the upper jaw. Depth of body equal to length of head, 33 times in total 
length. Snout obtusely pointed, covered with skin ; posterior nostril slightly nearer 
the eye than the anterior nostril; diameter of eye $ length of snout, ¢ length of 
head, 2 interorbital width ; cheek and opercle covered with skin ; skull striated above, 
granulate on the sides; fontanelle not produced as a groove posteriorly ; barbels not 


fringed, their bases united by a fold of the lower jaw; maxillary barbels just reaching 
3K 2 


424 MR. G. A. BOULENGER ON FISHES 


opercular cleft ; mandibular barbels short. Gill-cleft extending to below centre of eye. 
Humeral process striated, twice as broad and half as long as pectoral spine, obliquely 
truncated posteriorly. Pectoral spine longer than dorsal, a little longer than head, 
strongly serrated on both sides. Dorsal I 5; spine rather feebly serrated on both 
sides, a little shorter than the head, a little nearer the adipose fin than the end of the 
snout. Adipose fin not half as long as anal. Anal 11. No shields between the 
dorsal fins or on the belly. Lateral shields 30, 4 depth of body, with serrated border 
and moderately strong hooked spines. Caudal deeply bifurcate, with rounded lobes. 
Pale olive above, white below ; fins white. 

Total length 90 millim. 

A single specimen. 

Closely allied to 0. brevis Kner, which is distinguished by the longer dorsal spine, 
the narrower humeral process, the longer adipose dorsal, and the number (13-14) of 
anal rays. 


14. Oxyporas ELoNGATUS, sp. n. (Plate XL. fig. 4.) 


No teeth in the upper jaw. Depth of body 5 times in total length, length of head 
34 times. Snout obtusely pointed, covered with skin; posterior nostril twice as distant 
from the anterior as from the eye; diameter of eye nearly twice in length of snout, 43 
times in length of head, 1? in interorbital width; cheeks and opercles covered with 
skin ; cranial bones granulate ; fontanelle not produced as a groove posteriorly ; barbels 
not branched, their bases united by the fold of the lower jaw; maxillary barbels 
extending to opercular cleft; mandibular barbels'short. Gill-cleft extending to below 
posterior border of eye. Humeral process granulate and striated, twice as broad and 
half as long as pectoral spine, tapering to a point posteriorly. Pectoral spine as long as 
dorsal, nearly as long as head, extending to base of ventrals, strongly serrated on the 
inner side, more feebly on the outer. Dorsal I 6; spine feebly serrated in front, more 
strongly behind, equally distant from the end of the snout and the adipose fin. 
Adipose fin not half as long as base of anal. Anal 12. No shields between the dorsal 
fins or on the belly. Lateral shields 33, small, pluricuspid, with rather feeble hooked 
spines. Caudal deeply bifurcate. Olive above, white below; lateral shields and fins 
orange. 

Total length 105 millim. 

A single specimen. 

Resembles most O. humeralis Kner, which has minute teeth in the upper jaw, and 
much larger lateral shields, their depth being about 2 that of the body and their 
border bearing more numerous small spines. 


15. CaLLicHTHys arMATUS Gthr. 


16. PLEcostomus EMARGINATUS C. & Y. 


FROM THE RIO JURUA. 425 


17. Cua&TosToMuS BACHI, sp.n. (Plate XLI. fig. 1.) 

About 16 teeth on each side in each jaw. Depth of body 44 times in total length, 
length of head 3 times. Head very slightly longer than broad, entirely rough with 
small spines; snout rounded; diameter of eye 6 times in length of head, 3 times in 
length of snout, 34 times in interorbital width ; no postorbital groove; longest erectile 
preopercular spines as long as diameter of eye; barbel minute, hardly } diameter of 
eye. Throat and belly covered with small rough shields. Dorsal I 7; first ray a 
little shorter than head. Pectoral spine as long as head, covered with small spines, 
reaching middle of ventral. Ventral I 5, % length of head. Anal I 4. Caudal 
obliquely truncated, lower ray 14 as long as upper. Shields on body rough with small 
spinules, without keels, 26 in a longitudinal series. Pale olive, with rather indistinct, 
large, rounded, darker spots; caudal with dark cross-bars. 

Total length 110 millim. 

A single specimen. 

Nearest allied to Ch. ologospilus Gthr., which differs in the forked caudal fin and 
the naked belly. 


18. HyportopoMA GUENTHER! Blgr. 
19. LortcaRia RosTRATA Spix. 

20. LoRICARIA FILAMENTOSA Stdr. 
21. LorRICARIA ACIPENSERINA Kner. 


22, ACESYRA GLADIUS, sp.n. (Plate XLI. fig. 2.) 

Length of head 34 times in total length. Snout exceedingly long and narrow, 
sword-shaped, the pre-oral part as long as the rest of the head, the length of the 
rostrum 14 times its least width ; no bristles on the head ; eye small, its diameter 34 
times in interorbital width; upper angle of gill-opening nearer origin of dorsal than 
tip of snout. Dorsal I 6; first ray 4 length of head. Pectoral not reaching ventral. 
Anal 1 5; first ray 2 length of head. Caudal deeply emarginate, with the outer rays 
produced into very long filaments. Body moderately depressed; 7 scutes between 
occiput and dorsal, the second and third more than twice as broad as long; 3 series of 
yentral scutes, 6 on each side between pectoral and ventral. ‘Tail strongly depressed. 
32 scutes along each side, those on the body forming two obscure keels coalescing a 
little behind the vertical of the anal. Uniform olive-grey ; fins white, pectorals and 
ventrals with grey bars, caudal with a black band along the upper lobe. 

Total length 215 millim. 

Two specimens. 

Closely allied to A. oayrhyncha Kner. Distinguished by the absence of bristles on 


the snout and the broader dorsal scutes. 


426 MR. G. A. BOULENGER ON FISHES 


23. STEGOPHILUS NEMURUS Gthr. 


24, VANDELLIA crrRHOSsA C. & V. 


See remarks in P. Z. S. 1897, pp. 901 and 902. 

The following is a description of the four specimens in Dr. Bach’s collection :— 

Depth of body 7 to 10 times in total length, length of head 8 to 10 times. Head 
much depressed, nearly as long as broad; snout rounded, as long as the diameter of 
the eye, which is 33 times in length of head ; interorbital space one half to two thirds 
diameter of eye; mouth inferior; posterior nostrils between the eyes; maxillary barbel 
nearly half length of head; opercle and preopercle each with a bundle of erectile 
spines. Body slightly compressed. Pectoral as long as head minus snout. Dorsal 9, 
in the posterior fourth of the body. Anal 8-9, originating below middle of dorsal. 
Caudal truncated. Caudal peduncle twice as long as deep. Uniform white. 

Total length 62 millim, 


CHARACINID. 
25. CURIMATUS DoBULA Gthr. 


26. CURIMATUS ALBURNUS M. & T. 

27. TETRAGONOPTERUS MULTIRADIATUS Stdr. 
28. TETRAGONOPTERUS ORBICULARIS C. & V. 
29. TETRAGONOPTERUS MACULATUS L. 

30. CHIRODON ALBURNUS Gthr. 

51. CHALCINUS NEMATURUS Kner. 

52. GASTROPELECUS STELLATUS Kner. 

35. GASTROPELECUS PECTOROSUS Garm. 

34. ANACYRTUS KNERII Stdr. 

35. ANACYRTUS LIMZSQuAMIS Cope. 

36. ANACYRTUS AFFINIS Gthr. 

37. CYNODON PECTORALIS Gthr. 


38. CyNoDON VULPINUs Spix.” 
39. SERRASALMO PIRAYA Cuy. 


40. MYLETEs aLBIscoPus Cope. 


FROM THE RIO JURUA. 427 


CLUPEID. 
41. CETENGRAULIS JURUENSIS, sp.n. (Plate XLI. fig. 3.) 


Depth of body 5 times in total length, length of head 3} times. Snout very strongly 
projecting, a little shorter than diameter of eye, which is 5} times in length of head 
and 14 in interorbital width; jaws toothless; maxillary extending to articulation 
of mandible; cleft of mouth nearly } Jength of head. Géill-rakers long, finely denti- 
culate, about 40 on lower part of anterior arch. Dorsal 13, originating at equal 
distance from end of the snout and base of caudal. Pectoral a little more than 4 
length of head, extending a little beyond base of ventral. Anal 23, originating below 
posterior rays of dorsal. Caudal peduncle twice as long as deep. Caudal deeply 
forked. Scales 38 in a longitudinal series, 10 in a transverse series. Olive above, 
silvery on the sides and below; snout blackish above; fins pale orange, caudal rays 
blackish at the end. 

Total length 140 millim. 

A single specimen. 

This fish, the first freshwater representative of the genus Cetengraulis, is nearest 
to C. edentulus Cuy., which differs in the deeper body. 


42, PRISTIGASTER CAYANUS Cuv. 


OSTEOGLOSSID. 
43, ARAPAIMA GIGAS Cuy. 


Dr. Bach has shown me the photograph of a large specimen taken by him. 


GYMNOTID. 


44, STERNARCHUS NATTERERI Stdr. 
45, STERNARCHUS MACROLEPIS Stdr. 
46. STERNARCHUS OXYRHYNCHUS M. & T. 


47. STERNARCHUS TAMANDUA, sp. n. (Plate XLII.) 


Snout produced into a long, nearly straight tube, the length of which equals 4 times 
its least depth ; mouth very small, with several rows of minute teeth ; eye extremely 
minute, a little nearer the opercular cleft than the end of the snout. Depth of body 
half length of head. A very strongly developed adipose fin runs along the whole 
length of the body, from which it is easily detached. Pectoral 3 length of head. 
Vent under the chin. Anal 220, originating a little in advance of gill-opening, longest 
rays rather more than 4 depth of body. Scales very small, larger on the upper half of the 
body than on the lower; lat. 1. 85. ‘The tail, in the unique specimen, has been injured 
during life, and bears a short, regenerated caudal fin. Uniform yellowish white. 

Total length 400 millim. 


425 ON FISHES FROM THE RIO JURUA. 


‘This fish is very rare in the Jurua River, and but a single specimen could be procured 


by Dr. Bach. 
It is a most remarkable form, differing very considerably from any of the species 
with which we are acquainted. 


48. RHAMPHICHTHYS BLOcHI Kaup. 


49, SrEATOGENYS ELEGANS Stdr. 


‘The presence of a filament of adipose tissue, similar to the dorsal fin of Sternarchus, 
in a groove along each side of the mental region, to which attention has been drawn by 
Steindachner in describing his Rhamphichthys elegans, warrants, in my opinion, the 
establishment of a new genus, for which I propose the name Steatogenys. 


50. STERNOPYGUS VIRESCENS Val. 


TETRODONTID 2. 


51. T'erropon psirracus Bl. Schn. 


EXPLANATION OF THE PLATES. 
PLATE XXXIX. 


Platystoma juruense, p. 421, with upper and lower views of head 
and outline of premaxillary and palatal teeth. 


PLATE XL. 
Fig. 1. Oxydoras trimaculatus, p. 422. 
Fig. 2. Oxydoras trachyparia, p. 423. 
Fig. 3. Oxydoras bachi, p. 423. 
Fig. 4. Oxydoras elongatus, p. 424. 


PLATE XLI. 
Fig. 1. Chetostomus bachi, p. 425. 
Fig. 2. Acestra gladius, p. 425. 
Fig. 3. Cetengraulis juruensis, p. 427. 


PLATE XLII. 
Sternarchus tamandua, p. 427, with upper view of head. 


VTC, XXXTX, 


Ad 


oo. Ve 


0b, 


> 


Tra 7, 


PLATYSTOMA JURUENSE. Mintern Bros.imp. 


J.Green del etlith. 


J. Green delet lith MinternBros imp 
1.0XYDORAS TRIMACULATUS. 2.0.TRACHYPARIA. 3.0.BACHI. 4.0.ELONGATUS. 


“SISNANUNC STINVEONALHO' 8 SNIGVIS VEULS HOV Z THOVE SNWOLSOLWHODT 
* dha sorg ute yarpy “WAT 92 [ep Weare 


TW OOAIN O00 PB GS OL 


Trans. Lob, Soe VLANG¢, XLIT. 


rm Bro 


Minte 


del.et hth. 


~ Green 


STERNARCHUS TAMANDUA. 


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IX. On the Lepidosiren of the Amazons; being Notes on five Specimens obtained between 


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XI. On new or imperfectly-known Species of Ostracoda, chiefly from New Zealand. 
By G. Stewarpson Brapy, M.D., LL.D., D.Se., FBS. 


Received February 8, 1898, read March 15, 1898. 


[Prares XLIII-~XLVII.] 


FOr the opportunity of describing the following species of Ostracoda Iam indebted to 
Mr. G. M. Thomson of Dunedin, New Zealand, and to Dr. Meinert and Dr. H. J. Hansen 
of the Museum of Zoology, Copenhagen. Mr. Thomson’s specimens are the result of his 
own labours in the pursuit of natural history at Dunedin, Brighton, and other places 
in New Zealand; those sent to me from Copenhagen belong to the museum of that 
city, and were dredged in Lyttelton Harbour and Akaroa Harbour by Mr. H. Suter. 
One very interesting species (Eupathistoma natans) is from an altogether different 
geographical area, having been taken in the Bay of Bengal; for specimens of this 
species I have to thank my friend Mr. I. C. Thompson, F.L.S., of Liverpool. 

Though, except in the case of Hupathistoma, there is nothing very strikingly new in 
the morphology of the species here noticed, there are many points of interest as showing 
deviations from the usual types of structure; for example, the setose armature of the 
first pair of antenne in Trachyleberis scabrocuneata, the peculiar hairy cushions covering 
the valves of Sarsiella, the adventitious concretionary nodules found on the limbs of 
Philomedes sculpta, and the shell-glands and ducts of Philomedes flexilis. 

The species described or noticed in the present paper are as follows :— 


Asterope australis, G. 8. Brady, Otago, Akaroa, Lyttelton: p. 431. 
quadrata, sp. nov., Lyttelton Harbour: p. 432. 

os grisea, sp. nov., Akaroa Harbour: p. 482. 
Cyclasterope zealandica (Baird), Lyttelton Harbour: p. 433. 
ovulum, sp. nov., Stewart Island: p. 432. 

of tenera, sp. nov., Lyttelton Harbour: p. 433. 
Philomedes agilis, Thomson, Otago Harbour: p. 434. 
sculpta, sp. noy., Otago Harbour: p. 484. 

35 flewilis, sp. noy., Lyttelton Harbour, Akaroa Harbour: p. 435. 
Eupathistoma natans, gen. & sp. noy., Bay of Bengal: p. 437. 
Sarsiella hanseni, sp. noy., Lyttelton Harbour: p. 438. 
hispida, sp. noy., Akaroa Harbour : p. 489. 


2” 


” 


Ee, 


2) 
Cypris viridis, Thomson, Dunedin: p. 440. 
Cyprinotus flavescens, sp. noy., Dunedin: -p. 440. 
VOL. XIV.—PART vil. No. 1.-—December, 1898. 


(J) 
ial 


430 DR. G. 8S. BRADY ON NEW OR 


Loxoconcha punctata, Thomson, Otago, Brighton: p. 441. 
Xestoleberis luxata, sp. noy., Lyttelton Harbour: p. 441. 
# olivacea, sp. nov., Brighton: p. 442. 
#6 compressa, sp. nov., Brighton: p. 442. 
Cythere brunnea, sp. nov., Lyttelton Harbour; p. 442. 
ss innominata, nom, nov., Brighton: p. 443. 
3 truncula, sp. nov., Lyttelton Harbour: p. 444. 
Trachyleberis scabrocuneata, G. 8. Brady, Lyttelton Harbour: p. 444. 
Cytherideis nove-zealandie, sp. noy., Lyttelton Harbour: p. 446. 
Cytherella eburnea, sp. noy., Lyttelton Harbour: p. 447, 


In addition to these, there have been described the following New-Zealand species. 
which are unknown to me :— 


Cypris nove-zealandie, Baird. 
5, ciliata, Thomson. 
»  Uttoralis, Thomson. 
Cythere atra, Thomson. 


The memoirs quoted in this paper are the following. ‘They are indicated in the 
synonymy by the numbers attached to them in the list :— 


Barrp, W. (1).—Descriptions of several New Species of Entomostraca. (Proceedings of the Zoolo- 
gical Society of London, part xviii. Annulosa, plates xvil., xviii.) 1850. 
Brapy, G.S. (1).—A Monograph of the recent British Ostracoda. (Transactions of the Linnean 
Society, vol. xxvi.) 1868. 
+ (2).—The Voyage of H.M.S. ‘Challenger.’ Report on the Ostracoda. 1880. 
53 (3).—Notes on Entomostraca collected by Mr. A. Haly in Ceylon. (Journal of the 
Linnean Society, Zoology, vol. xix.) 1885. 
is (4).—On Ostracoda collected by H. B. Brady, Esq., LL.D., F.R.S., in the South-Sea 
Islands. (Transactions of the Royal Society of Edinburgh, vol. xxxv. part ii.) 
1888. 
5 (5).—A Supplementary Report on the Crustaceans of the group Myodocopa obtained 
during the ‘ Challenger’ Expedition, with Notes on other new or imperfectly- 
known species. (Transactions of the Zoological Society of London, vol. xiv. 
part ii.) 1897. 
Brapy, G. 8., & Norman, A. M. (1).—A Monograph of the Marine and Fresh-water Ostracoda of 
the North Atlantic and of North-west Europe.—Part ii. Myodocopa, Cladocopa, 
and Platycopa. (Transactions of the Royal Dublin Society, vol. v., ser. ii.) 
1896. 
Dana, J. D. (1).—Crustacea of the United States Exploring Expedition. 1852. 
Jonas, T, R. (1)—A Monograph of the Tertiary Entomostraca of England. (Paleontographical 
Society.) 1856. 
Miter, G. W. (1).—Die Ostracoden des Golfes von Neapel und der angrenzenden Meeres- 
abschnitte. 1894. 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 431 


Norman, A. M. (1).—Last Report on Dredging among the Shetland Isles. (British Association 
Reports.) 1868. 
Puiuirri, A. (1).—Zoologische Bemerkungen in Arch. Naturg. 6 Jahrg. 1840. 
Sars, G. O. (1).—Nye Bidrag til Kundskaben om Middelhavets Invertebratfauna. (Archiv for 
Mathematik og Naturvidenskab.) 1887. 
(2).—On some Freshwater Ostracoda and Copepoda raised from dried Australian Mud. 
(Christiania Videnskabs-Selskabs Forhandlinger, no. 8.) 1889. 
Tuomson, G. M. (1).—On the New Zealand Entomostraca. (Transactions of the New Zealand 
Institute, vol. xi.) 1878. 


2 


Genus AsteropE Philippi. 
Asterope Philippi (1), 1840; G. O. Sars (1), p. 11; Brady & Norman (1), p. 629. 
Cylindroleberis Brady (1), p. 465; G. W. Miiller (1), p. 216. 
Cypridina Auctorum. 


ASTEROPE AUSTRALIS Brady. (Plate XLIII. figs. 1-8.) 
Asterope australis Brady (4), p. 515, pl. iv. figs. 1, 2. 

Shell of the male, seen from the side (fig. 1), ovate, height equal to more than half 
the length, anterior extremity evenly rounded, with a short, wide beak and a rather 
wide but shallow antennal notch; posterior extremity evenly rounded; dorsal margin 
very slightly but evenly arcuate, ventral almost straight. Seen from above (fig. 2), the 
outline is elongate-ovate, widest in the middle, much more than twice as long as 
broad ; extremities broadly rounded, lateral margins gently and evenly curved. Surface 
of the shell smooth, destitute of hairs or sculpture, excepting at the posterior extremity, 
which bears two tufts of very fine hairs. Secondary branch of the antenna (fig. 5) with 
a large and very crooked terminal unguis, which bears near its base a very long and 
stout seta, the last joint (dactylon) having two much shorter sete. Post-abdomen 
(fig. 8) armed, on each lamina, with a series of seven ungues, which progressively 
increase in size and amount of curvature from the first to the last, the larger of the 
series very finely ciliated on the concave margin. Length of the shell 1:6 mm, Colour 
(of spirit-specimens) yellowish white; the black eye-spot very conspicuous. 

The shell of the female (figs. 3, 4) is somewhat smaller than that of the male, and 
seen laterally is higher behind than in front; seen from above it is subacuminate in 
front and narrowly rounded behind; the posterior extremity has no setose tufts. The 
secondary branch of the antenna consists of a simple cylindrical one-jointed stem, 
which ends in a single long seta (fig. 6). Length 1-4 mm. 

Hab. Males, taken abundantly in the surface-net, Otago Harbour; a single female 
specimen in a dredging from Akaroa Harbour, depth 6 fathoms, and Lyttelton Harbour, 
1-5 fathoms. The Otago gathering consisted almost entirely of males, the lesser 
swimming-power of the females doubtless keeping most of them at or near the bottom. 


The shell in most specimens is flexible and submembranaceous, but occasionally hard 
3L 2 


432 DR. G. S. BRADY ON NEW OR 


and calcareous, the soft ones probably imperfectly matured after moulting. The types 
of the species were described by me from specimens taken amongst the South-Sea 
Islands, and do not differ from the New Zealand examples except in being rather 


larger. 


ASTEROPE QUADRATA, sp. n. (Plate XLV. figs. 17-21.) 


In general appearance this is very like A. australis, but it is somewhat smaller, and seen 
dorsally has much more obtuse extremities, having almost parallel lateral margins; the 
inner antennal branch of the male (fig. 19) is very broadly clawed, and the larger ungues 
of the post-abdominal lamine in the female (fig. 21) are not only finely pectinated, but 
bear a few longer interspersed setae—about five on the last and three on the penultimate 
unguis; in the male, however, of which only one mutilated specimen was seen, there 
is only a simple pectination. Shell extremely thin and membranous. Length 1:3 mm. 

Hab. Lyttelton Harbour, dredged in a depth of 1-5 fathoms. 


ASTEROPE GRISEA, sp. n. (Plate XLIII. figs. 9-14.) 


Shell of the female, seen from the side, elongated, elliptical (fig. 9), height scarcely 
as much as half the length, and nearly equal throughout; extremities evenly rounded, 
the posterior rather the narrower of the two; dorsal and ventral margins evenly and 
very slightly curvate. Seen from above (fig. 10) the outline is narrowly ovate, the width 
equal to two-fifths of the length, slightly tapered towards the front, broader and 
rounded off behind. Shell chitinous, flexible, perfectly smooth, colour greyish. 
Length 2°8 mm. Frontal tentacle cylindrical, its terminal joint slightly dilated at 
the base; secondary branch of the antenna (fig. 12) small, simple, with a small apical 
seta; spines of the post-abdominal lamine (fig. 14) long, slender, very delicately ciliated. 
Claw of the secondary antennal branch in the male (fig. 11) much thickened and 
dilated at its base. 

Hab. Akaroa Harbour; many specimens dredged in a depth of 6 fathoms. Only one 
male could be found, and this was a good deal damaged; its only marked peculiarity, 
so far as could be made out, was the extreme length of the antennular sete. 


Genus CyciasTERore G. S. Brady. 
Cyclasterope Brady (5), p. 85. 


CYCLASTEROPE OVULUM, sp. nu. (Plate XLIII. figs. 24-30.) 


Shell of the female (figs. 24, 25), seen from the side, nearly circular, slightly narrowed 
and produced in front, antennal notch small and shallow; seen from above, ovate, widest 
in the middle, twice as long as broad, subacuminate in front, narrowly rounded behind. 
Shell dense and calcareous, smooth, finely punctate. Length 8 mm., height 6°6 mm. 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 433 


Frontal tentacle (fig. 27) three-jointed, the median joint very short and constricted at 
the base, distal joint conical, with a much-attenuated apex; secondary branch of the 
antenna (fig. 26) three-jointed, cylindrical, ending in a long seta, the two basal joints 
bearing a few marginal hairs; teeth of the bifid extremity of the vermiform limb 
(fig. 28) corrugated, their apices formed by a small spine, which is overhung by a 
knobbed columnar process: abdomen fringed distally with closely-set hairs; post- 
abdominal lamine (fig. 29) bearing three very stout, blunt, and strongly curved distal 
ungues, which bear two marginal rows of closely-set lancet-shaped spinules (fig. 30) ; 
in front of the ungues is a series of about nine long ringed and pinnated sete. Male 
unknown. 

Hab. Rock-pools, Stewart Island. 

This fine species closely resembles in external aspect C. orbicularis, of which a 
single specimen from Valparaiso was described by me (/oc. cit.), but C. ovudum is much 
larger, and differs in many points as regards the soft parts. 


CYCLASLEROPE TENERA, sp.n. (Plate XLIV. figs. 27-29.) 


Female. Shell, seen from the side (fig. 27), almost circular, slightly narrower towards 
the front, length about one-seventh greater than the height, very thin, membranous, 
and slightly wrinkled; anterior and inferior margins beset with distant, rigid hairs. 
Secondary antennal branch (fig. 28) simple, three-jointed, with a long apical seta; 
faleate process of the mandible bearing numerous marginal spines and teeth; . 
the post-abdominal claws are three in number (fig. 29), very unequal in length, and 
armed with closely-set short marginal spinules. Length 1:5 mm. 

Hab. Lyttelton Harbour, 1-5 fathoms. One specimen only. 


CYCLASTEROPE ZEALANDICA (Baird). (Plate XLIII. figs. 15-23.) 
Cypridina zealandica Baird (1), p. 257, pl. xvii. figs. 11-13. 

Shell of the female (figs. 15, 16), seen from the side, very broadly ovate, length 
about one-fourth greater than the height, the outline forming a perfect oval except at 
the antennal notch, which is small and shallow; seen from above, ovate, compressed, 
greatest width in the middle, and equal to half the length; extremities rather wide and 
evenly rounded, slightly emarginate at the contact of the valves. Shell calcareous, 
finely punctate and, towards the anterior extremity, faintly wrinkled in a transverse 
direction (fig. 23). Length 5 mm., height 4 mm. Secondary branch of the antenna 
(fig. 19) cylindrical, three-jointed, geniculated, the first joint bearing a tuft of marginal 
setze near its extremity, second joint with three small marginal sete externally and a 
long seta on its internal distal angle, last joint obliquely truncate and mucronate at 
its apex; ungues of the post-abdominal lamin (fig. 21) stout, subequal, and strongly 


434 DR. G. S. BRADY ON NEW OR 


curved, bordered with rows of lanceolate spinules (fig. 22), which, towards the bases of 
the ungues, have also intermediate smaller spinules; in front of the ungues a row of 
about ten flexuous ringed sete, which gradually decrease in length until they give place 
to a fringe of densely-set fine hairs. Length 5 mm., height 4 mm. 

Shell of the male (fig. 17), seen from the side, subovate, somewhat narrowed towards 
the front, greatest height situated in the middle and equal to nearly three-fourths of the 
length; anterior extremity rounded, beak sharp and curved, antennal notch wide ; 
posterior extremity subtruncate, rounded off below, abruptly angulated at its junctiow 
with the dorsal margin, which is almost straight for the greater part of its length, but 
is well rounded in front, and slopes steeply behind to join the posterior border ; surface 
minutely and closely punctate, fringed at the two extremities with long, fine hairs. 
Length 6:25 mm., height 4°5 mm. Terminal joint of the secondary antennal branch 
(fig. 18) forming a strong claw, very stout and constricted near the middle; penultimate 
joint armed on the opposing margin with a fascicle of very thick, rigid sete. 

Hab, Dredged in Lyttelton Harbour, 2-5 fathoms. Only few specimens, the males 
badly mauled, and their limbs clogged with mud so as to be viewed with difficulty. 

Dr. Baird’s figures and description agree well with these specimens, and his notice 
of the wrinkled shell-surface confirms the impression that this is the form to which 
he refers. 


Genus Puitomenpss Lilljeborg. 


PHILOMEDES AGILIs Thomson. 
Philomedes agilis Thomson (1), p. 257, pl. xi. figs. C. 8 a-e, D. 1 a-g; G.S. Brady (5), p. 90, 
pl. xvi. figs. 13-16. 


Taken in the surface-net, Otago Harbour. Males only. 


PHILOMEDES SCULPTA, sp.n. (Plate XLIV. figs. 15-20.) 


Shell of the male (fig. 15), seen from the side, elongated, subrhomboidal, highest in 
the middle and narrowing gradually to the extremities, height equal to half the length ; 
anterior extremity produced to a subacute point, notch obsolete ; posterior extremity 
produced below the middle into a wide, wedge-shaped, obtusely-pointed beak, above 
which it is rather deeply sinuated; dorsal margin moderately and evenly curved 
throughout its whole length, ventral almost straight in the middle and bent sharply 
upward towards each extremity. Seen from above (fig. 16), the outline is compressed, 
subovate, greatest width in front of the middle, and equal to less than half the length ; 
anterior extremity wide, subtruncate, mucronate in the middle, and emarginate towards 
the sides; posterior narrower, bluntly pointed, lateral margins convex and irregularly 
sinuous. Surface of the shell undulated, covered with irregular, small, polygonal 
pittings, and raised into two principal longitudinal ribs which join near the front, 
forming a single short median ridge, but remain separate behind, running into a 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 435 


marginal ridge; the spaces between, and less conspicuously outside of, these ribs are 
marked with smaller, curved, anastomosing ridges; the anterior margin round about 
the antennal notch (fig. 17) has a thin laminated flange marked by delicate transverse 
lines. Secondary branch of the antenna (fig. 18) geniculated between the second and 
third joints; first joint very short, bearing two short marginal sete, second and third 
very long and nearly equal, second with three long sete on its outer margin, third 
rugose with a number of wart-like protuberances on its opposing surface, and bearing 
a single seta near the distal and proximal ends respectively; extremity blunt and 
slightly furrowed longitudinally; post-abdominal lamin (fig. 19) armed with three 
principal ungues and seven smaller ones, the first of the smaller series situated between 
the second and third larger ones. Length 2°6 mm. 

Hab. Taken abundantly in the surface-net, Otago Harbour. Males only. 

An interesting peculiarity of this species consists in its tendency to develop calcareous 
concretions on the setz of the antenne and antennules, and sometimes in other situations. 
These concretions are extremely dense and dark-coloured, and when broken present a 
radiated crystalline appearance. ‘Treated with a weak acid, they are seen under the 
microscope to effervesce freely, and in time to disappear almost entirely, so that they 
are probably composed chiefly or altogether of calcium carbonate. Two or more sete 
are sometimes immovably soldered together by a concretion of this kind, which must, 
one would think, materially interfere with the locomotion of the animal. Though 
P. sculpta is especially liable to these concretions, they are sometimes met with in 
other species; in P. sculpta, however, full-grown specimens seem rarely to be free from 
them. Though in this gathering the majority of examples had shells of a flexible 
or membranous kind, some few were distinctly dense and calcareous. These differences 
may perhaps depend upon the lapse of time in relation to exuviation; but I am 
disposed to think that in the case of flexible shells there is often a larger development 
of the concretions above referred to, in which case it may be allowable to look upon 
them as pathological products which have withdrawn the lime otherwise available for 
shell-formation. A broken fragment of one of the nodules is shown in fig. 20. It 
is just possible that a species described by me from one dried shell under the name 
Streptoleberis crenulata may be identical with P. sculpta, but this I cannot decide 
with any certainty (see 5, p. 515, pl. iv. figs. 3, 4). 


PHILOMEDES FLEXILIS, sp. n. (Pl. XLIV. figs. 1-14; Plate XLV. figs. 16, 16.) 


Shell of the female (Pl. XLIV. figs. 1,2; Pl. XLV. figs. 15, 16), seen from the side, 
irregularly lozenge-shaped, widest in the middle, greatest width equal to about two- 
thirds of the length; anterior extremity somewhat produced and narrowed, beak 
inconspicuous, antennal notch shallow; posterior extremity narrowed, running out 
below the middle into a wide, blunt, and massive tuberosity; dorsal margins boldly 
arched (in old specimens rugged and ending in a tuberous projection); ventral 
margin moderately convex. Seen from above, the outline is irregularly polygonal, 


436 DR. G. S. BRADY ON NEW OR 


with wide extremities and subparallel sides, the laterally-produced rostrum and 
the posterior beak forming large terminal protuberances. Except in old specimens 
(Pl. XLIV. figs. 1, 2) the shell is flexible and membranaceous, and covered with 
rounded or subangular pittings (fig. 12); each valve bears three flexuous longitudinal 
ribs which, viewed dorsally, stand out very conspicuously as irregular translucent 
flanges on each lateral margin; rostrum and adjoining shell-margin bordered with a thin, 
semitransparent, radially-striated lamina; posterior extremity fringed with a few very 
small recurved hairs. Just within the anterior margin, and below the antennal sinus, 
a small patch of the shell is marked with a series of about ten parallel striz (fig. 13). 
Frontal tentacle (fig. 4) slender, filiform, sharply pointed, its median portion divided 
into about sixteen very small joints, base bulbously dilated. Secondary branch of the 
antenna (fig. 6) composed of a single (?) curved, sickle-shaped joint, which bears on its 
outer edge, near the base, three short sete, near the middle one extremely long plumose 
seta, and at the blunt apex a short flexuous seta. The sete of the swimming-branch, in 
small specimens (fig. 7), are short and non-plumose, but in fully-grown ones longer and 
plumose. ‘The principal chewing segment of the second maxilla (fig. 8) is in the form 
of a blunt, broad-ended lobe, with one broad tooth-like process at its inner end, and two 
similar but larger processes at the outer end; these are sometimes, though not always, 
of a deep purple colour; spines of the vermiform foot (fig. 9) with very thick 
peduncles; post-abdominal ungues about ten in number, progressively increasing in 
length from the first, which is extremely small (fig. 10) ; the seventh, ninth, and tenth 
ungues are stout, and bear rather stout and short marginal teeth ; the eighth and all 
the other ungues are more slender and only feebly ciliated. Eyes usually wanting, 
but sometimes well developed, deeply pigmented and distinctly visible through the 
shell (fig. 11). On the inner surface of the rostrum lies a convoluted “ shell-gland ” 
which seems to communicate with a nipple-like tubular prominence opening near 
the margin of the shell (fig. 14). Length 2-2°3 mm. The shell of the male is much 
‘narrower and more elongate (fig. 3), but in other respects agrees with that of the 
female; the eyes are well developed (fig. 4), and the secondary antennal branch (fig. 5) 
is very similar to that of P. sculpta. 

Hab. The specimens from which the description is drawn up were taken numerously 
by the dredge in depths of 1-5 fathoms in Lyttelton Harbour. Others which I refer 
to the same species occurred in a dredging from 6 fathoms in Akaroa Harbour. These, 
however, differ slightly from the types in having generally an almost smooth shell with 
little or no trace of ribs, though in not a few specimens the ribs are quite apparent, 
and are in character like those of the types. 

The great majority of specimens possess a quite flexible and membranous shell, but 
two or three (probably very old individuals) occurred in which it had become dense 
and calcareous, the various processes and ridges being at the same time strongly 
developed (figs. 1, 2). In some examples, which I suppose to be immature, the 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 457 


antennal sete are very short and non-plumose, like those of G. W. Miiller’s genus 
Pseudophilomedes, but in most cases they are long and strongly plumed. A peculiarity 
which I do not understand is the presence in a very few female specimens of well- 
developed eyes, organs of which in most cases I have been able to find no trace; another 
unintelligible point is the red coloration, in some specimens, of the teeth of the second 
maxilla. Of course it is possible that two distinct species may be mixed up in my 
description, though I scarcely think that is the case. I am not aware that the shell- 
gland has been previously noticed, and it is only in two or three specimens that I have 
found it; the duct which I have figured is probably connected with the gland, though I 
have not certainly made out the connection; the structure is probably an excretory one 
and homologous with the green gland of higher crustacea. 


Genus EvpaTHiIstoMa !, gen. nov. 


Shell membranous, in shape not unlike Philomedes. Antennules and antenne nearly 
alike in both sexes, and similar to those of Cypridina, except that the antennules of 
the male have no suckers; margins of the mouth provided with three pairs of finger-like 
sensory processes; mandibles and maxillz as in Cypridina; terminal armature of the 
vermiform foot one-sided, consisting of several unequal, slender lashes, the longest of 
which are longer than the diameter of the limb ; post-abdomen as usual in Cyp.*.inide. 

In general character this genus is intermediate between Cypridina and Philomedes, 
but the sensory mouth-organs are very remarkable, and seem to have no parallel | 
among other Myodocopa. One pair, at least, of these appendages has an arrange- 
ment of olfactory (2) filaments exactly like that found in the antennules of the 
Daphniade. 


EUPATHISTOMA NATANS, sp.n. (Plate XLIV. figs. 21-26.) 


Shell seen from the side (fig. 21) elongated, subrhomboidal, greatest height equal to 
more than half the length; anterior extremity obliquely truncated and slightly concave, 
distinctly angulated at its dorsal end, very acutely and prominently angular below, 
where it merges in a wide, well-rounded antennal sinus; posterior extremity produced 
below the middle into a wide, bluntly-rounded beak ; dorsal margin almost flat in the 
middle, sloping rather steeply to the anterior extremity, and with a steeper curve 
behind to its junction with the posterior beak, ventral margin rather boldly convex, 
more fully rounded in front than behind; seen from above (fig. 22), evenly ovate, 
twice as long as broad, mucronate behind, narrowly rounded in front. Length 
2mm. Male and female nearly alike. The sensory mouth-appendages consist of 
three pairs of finger-like or conical processes (fig. 24), one pair of which (@) are simple 


1 eizaOis, Sensitive; orduu, a mouth. . 


VOL. XIV.—Part vil. No. 2.—December, 1898. 3M 


438 DR. G. S. BRADY ON NEW OR 


cones with terminal rosettes of short olfactory (?) setee, a second pair (J) consisting each 
of two coalescent cones, a third (c) longer and finger-like with crenated margins. The 
mandible (fig. 23) has four long terminal ungues and a short seta; its chewing-lobe 
(fig. 23 a) is setose and ends in two slender mucrones; and the antepenultimate joint 
bears, like Cypridina, a short apical process with a lancet-shaped point and two lateral 
sete (fig. 230). Lateral sete of the vermiform limb few, generally not more than 
eight; terminal sete long, filiform, not at all tooth-like, unilateral (fig. 25). Post- 
abdominal lamine (fig. 26) bearing nine pectinated marginal ungues, which are 
progressively longer from first to last. 

Hab. Taken abundantly in the surface-net in the Bay of Bengal, lat. 14° N. For 
these specimens I am indebted to my friend Mr. I. C. Thompson, F.L.S., of Liverpool. 


Genus SarsteLLa Norman, 1869. 


Sarsiella Norman (1), p. 293; Brady & Norman (1) (?), p. 677. 
Nematohamma Brady & Norman (1) (2), p. 680; G. W. Miiller (1), p. 213. 


SARSIELLA HANSENI, sp. n. (Plate XLV. figs. 1-12.) 


Shell of the female, seen from the side (fig. 1), subquadrate, height equal to four- 
fifths of the length; anterior extremity very wide, truncate, slightly irregular, but 
almost straight, posterior narrower, deeply excavated, and bounded above and below 
by two prominences, the lower of which is much the largest and forms a stout sub- 
triangular beak; dorsal margin slightly arcuate, somewhat sinuous, highest in the 
middle, whence it slopes very gently towards the front, but steeply and with a distinct 
curve backward, ventral margin more strongly convex and sinuated in front of the 
posterior beak. Seen from above (fig. 2), the outline is elongated and subquadrangular, 
of nearly equal width throughout, width equal to more than half the length; anterior 
extremity wide, flattened, but deeply emarginate in the middle, posterior almost as 
wide as the greatest width of the shell, abruptly truncated and slightly prominent in 
the middle; lateral margins subparallel, very slightly divergent from behind forward, 
almost rectangular at their junction with the posterior extremity, but rounded off in 
front. The substance of the shell is flexible and submembranaceous, closely pitted 
with small impressed puncta and irregularly waved and ribbed; the anterior and inferior 
margins form a continuous elevated ridge; two longitudinal ribs run parallel to, but 
at a considerable distance within, the dorsal and ventral margins; there is a flexuous 
central rib between these two, rising near the front of the valve, and becoming lost 
behind the middle; between this and the dorsal rib is another rib which begins with 
a sharp bend in front of the middle, and, gradually becoming stronger, passes back- 
vard and ends in a curve a little within the posterior margin. Between these principal 
ribs the valves are irregularly corrugated; all the ribs are beset with long and rather 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 439 


coarse hairs (fig. 3), and the dorsal surface bears patches of very short, rigid, and 
closely-set hairs (figs. 2, 11), which have dilated subglobular or crutch-shaped apices, 
Length 1:05, height *85 mm. Shell of the male (fig. 4) elongated, not unlike that of 
Philomedes, nearly twice as long as high; surface-markings scarcely so pronounced as 
those of the female. Length 1:1 mm. The limbs of the female differ scarcely at 
all from those of S. capswla, which has been fully described by G. O. Sars and 
G. W. Miller. Post-abdominal laminz narrow, its four marginal ungues very slender, 
and increasing progressively in length from the first to the last, which is at least four 
times as long as the first (fig. 10), their inner margins sparingly spinulose. The 
mandible of the male (fig. 8) differs from that of the female in being more profusely 
setiferous, and in having only one, instead of three, terminal ungues; the antennule 
(fig. 5) has on the penultimate joint a cushion-like prominence which bears a dense 
brush of innumerable long fine hairs; extremity of the vermiform limb truncated, and 
- without the pairs of hooks which are found in the female. Copulative organ (fig. 12) 
ending in two stout and blunt chitinous hooks. Secondary branch of the antenna 
(fig. 6) three-jointed, geniculated ; inner margin of the second joint bearing at the base 
two very long and stout spine-like sete; terminal joint slightly thickened at its base, 
blunt and angularly bent. Post-abdomen like that of the female. 

Hab. Plentiful in a dredging, from 1-5 fathoms, in Lyttelton Harbour. 

The extremely hirsute character of the shell and the peculiar cushion-like patches of 
knobbed hairs, forming a sort of interrupted nimbus round the edges when seen 
dorsally, are very characteristic. But the surface-sculpture is not easily seen while 
the shell is immersed, and when removed from liquid it very speedily shrivels and 
loses its proper markings. The genus Nematohamma Brady & Norman was founded 
on male specimens of a Sarsiella, the relation of which to the female form was 
then unknown. 


SARSIELLA HISPIDA, sp.n. (Plate XLV. figs. 13, 14.) 


Shell membranous, thin and flexible, seen from the side subrhomboidal (fig. 13), 
height equal to two-thirds of the length; anterior extremity narrowed, rounded; 
posterior much wider, subtruncate, sloping very steeply, and terminating in a long 
conical beak; dorsal margin almost flat in the middle, whence it slopes gently toward 
the front; near its hinder end it is distinctly sinuated and terminates in an abrupt 
angle; ventral margin boldly arcuate, with a slight sinuation (antennal notch) near 
the anterior extremity; seen from above (fig. 14), subquadrate, oblong, sides nearly 
parallel, but indented near the middle, obtusely rounded in front, rectangularly 
truncated behind, with a large median mucronate process. Shell densely hispid, and 
on the dorsal aspect bearing patches of short, rigid, clubbed hairs; surface of the 
valves flexuously ribbed in a longitudinal direction. Length 1-5 mm. 

Hab. Akaroa Harbour; dredged in a depth of 6 fathoms. 

3M Z 


440 DR. G. S. BRADY ON NEW OR 


The test of this species is so very thin that it is impossible to examine it in the dry 
condition owing to its rapid shrivelling, and the markings are very difficult to make out 
accurately in fluid; the disposition of the ridges seems, however, to be very much like 
that of S. hansent. 

Genus Cypris O. F. Miiller. 

Cypris viripis Thomson. (Plate XLV. figs. 22-24.) 
Cypris viridis Thomson (1), p. 253, pl. xi. fig. A. 2a-g. 

Shell of the female short and very tumid; seen from the side (fig. 22) reniform, very 
slightly depressed in front, height equal to nearly two-thirds of the length ; extremities 
well rounded, dorsal margin boldly and evenly arched, ventral slightly sinuated in the 
middle ; seen from above (fig. 23), very broadly ovate, greatest width behind the middle 
and equal to about three-fourths of the length, anterior extremity abruptly tapered 
and subacute, posterior broadly rounded ; lateral margins strongly arcuate, converging 
steeply towards the front. Shell-surface finely punctated and covered with short, 
rather rigid hairs; the anterior and posterior extremities bordered by a narrow, 
pellucid flange, which is fringed with a single series of very fine hairs; behind this 
flange is a thicker band marked by irregular transverse glandular (!) streaks; left valve 
overlapping the right on the dorsal and posterior aspects. Colour an opaque dark 
green. Length-9mm. Post-abdominal rami (fig. 24) very small and slender; terminal 
seta as long as the ramus itself, lateral seta about half as long and attached very near 
the apex; the minor apical seta is extremely minute. 

Hab. “In pools about Dunedin and Taieri Plan” (@. M. Thomson). 


Genus Cyprinotus G. 8. Brady. 
Cyprinotus Brady (3), p. 301; G. O. Sars (2), p. 5 (separate copy). 


CYPRINOTUS FLAVESCENS, sp. n. (Plate XLVI. figs. 28, 29.) 


Shell, seen from the side (fig. 28), reniform, highest in the middle, height equal to 
rather more than half the length; extremities rounded, anterior wide and almost 
subtruncate, posterior narrower and slightly depressed; dorsal margin forming an 
evenly flattened arch, ventral gently sinuated towards the front; seen from above 
(fig. 29) the outline is compressed, ovate, widest in the middle, more than twice as 
long as broad, gradually tapered towards the acuminate anterior extremity, narrowly 
rounded behind; surface of the valves quite smooth. Colour yellowish. Length 
2-2 mm. 

Hab. Dunedin district (exact locality lost). 

The soft parts of the animal agree accurately with those of the typical Cyprinotus, 
but there is not so much inequality of the valves as in the better-known species. 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 44] 


Genus Loxoconcua G. O. Sars, 
Loxoconcna punctata Thomson. (Plate XLVI. figs. 5-5.) 
Loxoconcha punctata Thomson (1), p. 255, pl. x1. fig. B. 3 a-k. 


Shell of the female, seen from the side (fig. 3), subrhomboidal, height equal to more 
than two-thirds of the length; extremities obliquely rounded, the anterior sloping 
steeply above the middle and rounded off ventrally, posterior forming a bold curve 
from below to the dorsum, where it ends in an abrupt angle; dorsal margin forming 
a flattened arch with an excavation where it joins the posterior extremity, ventral very 
slightly sinuated in the middle; seen from above (fig. 14), lozenge-shaped, twice as long 
as broad, widest in the middle, whence it tapers rather suddenly to the extremities, 
which are subacuminate and nearly equal. Surface of the shell smooth, bearing 
numerous small papille; valves flattened out at the extremities, forming a bordering 
flange which is continued, though feebly, along the ventral margin. Colour generally 
greyish, but variable, as is also the texture of the shell. Length 57 mm. The shell 
of the male is rather larger (‘7 mm.), and has the characteristic shape of that sex in 
Loxoconcha. 

Hab. Otago Harbour, among seaweed and in rock-pools, Brighton, New Zealand. 

This species has been described and figured by Mr. G. M. Thomson (Joc. cit.); but, 
as the ‘ Transactions’ of the New Zealand Institute are not generally accessible, I here 
re-figure the female. 


Genus XustToLeBeris G. O. Sars. 
XESTOLEBERIS LUXATA, sp. n. (Plate XLVI. figs. 20-27.) 


Shell of the female compressed, subreniform, left valve much larger than the right, 
and overlapping everywhere except on the ventral margin (figs. 20, 21); seen from the 
side, nearly twice as long as high, highest behind the middle, depressed in front; 
anterior extremity narrow, rounded but somewhat flattened, posterior wide and boldly 
rounded; dorsal margin forming a continuous curve, flattened toward the front, but 
boldly rounded behind, ventral sinuated in front of the middle; seen from above, 
compressed, ovate, twice as long as broad, widest behind the middle, tapering gently 
towards the front, well rounded behind; shell-surface smooth, marked with numerous 
very small circular papille. Colour yellowish white. Length ‘6mm. Antennules 
(fig. 24) six-jointed, bearing two sete on the second joint, one on the third, three on 
the fourth, one on the fifth, and three on the sixth, all of them short; the mandibular 
branchia (fig. 26) consists of only two sete, and the animal is, in other respects, similar 
to the typical Xestoleberis. 

Hab. On alge in Lyttelton Harbour, New Zealand. 

In most species of Xestoleberis there is some inequality of the valves, but in this 
species it is more pronounced than in anv other known to me. 


449 DR. G. S. BRADY ON NEW OR 


XESTOLEBERIS OLIVACEA, sp. n. (Plate XLVI. figs. 6, 7.) 

Shell, seen from the side (fig. 6), subreniform, greatest height situated in the middle 
and equal to two-thirds of the length; extremities well rounded; dorsal margin boldly 
and evenly arched, sloping steeply behind, more gradually in front, ventral rather 
deeply sinuated in front, boldly arcuate behind, where it forms a curve continuous 
with that of the posterior extremity; seen from above (fig. 7), ovate, nearly twice as 
long as broad, widest in the middle, lateral margins evenly and boldly arcuate; 
anterior extremity subacuminate, posterior narrowly rounded. Shell-surface perfectly 
smooth and without markings, excepting a dark eye-spot. Colour dark olive. Length 
55 mm. 

Hab. Rock-pools, Brighton, New Zealand. 


XESTOLEBERIS COMPRESSA, Sp. n. (Plate XLVI. figs. 10-19.) 


Shell, seen from the side (fig. 10), reniform, highest in the middle, height equal to 
more than half the length; anterior extremity obliquely rounded, posterior boldly and 
evenly rounded; dorsal margin boldly arched, ventral rather deeply sinuated in the 
middle; seen from above (fig. 11), compressed, ovate, fully twice as long as broad, 
acuminate in front, rounded behind. Shell-surface smooth, marked with distant, very 
minute papille. Colour yellowish. Length ‘46mm. Antennules (fig. 13) six-jointed, 
third, fourth, and fifth joints each with a single strong apical spine and a slender seta, 
sixth joint with two sete; second joint fringed externally with fine, short hairs; in 
other respects like typical Xestoleberis. 

Hab. Rock-pools, Brighton, New Zealand. 

G. W. Miiller notes that in many, if not in all, species of Xestoleberis the shell bears 
a crescentic pellucid patch behind the eyes, and of this he gives several figures. I 
have not been able to find this patch in any of the species here described, though in 
X. compressa there is, at any rate in some specimens (fig. 10), a similar mark in front 
of the eyes. But the shell in this species and in others of the same genus is liable to 
the presence of similar irregular spots on various parts. 


Genus Cyruere O. F. Miiller. 
CYTHERE BRUNNEA, sp. n. (Plate XLVII. figs. 8-15.) 


Male. Shell, seen from the side (fig. 8), subreniform, greatest height in front of the 
middle, and equal to more than half the length; anterior extremity wide, rather 
obliquely rounded, forming a very wide band or fillet, which ceases above at the 
highest point of the valve, but is continued in a narrower form along the ventral 
margin, and as far as the postero-dorsal angle; posterior extremity much narrower 
than the anterior, subtruncate, rounded off below, almost angular above; dorsal 
margin slightly arcuate, sloping continuously and evenly from the front backward ; 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 448 


ventral deeply sinuated in front of the middle, behind which it is slightly convex ; 
seen from above, the outline is narrowly ovate, nearly thrice as long as broad 
(fig. 9), rounded behind and tapered gradually to the obtusely-pointed anterior 
extremity. Shell-surface covered with closely-set angular pittings, except on the 
marginal flange (fig. 10), the flange itself marked with closely-set, fine, radiating 
lines (lacune?) on its broad inner zone, and on the narrower outer zone with 
distant short hairs. Length ‘66 mm. Antennules (fig. 11) six-jointed, each of the 
last four joints bearing a stout apical spine, the last three having also one or two 
setee which are not very much longer than the spines; urticating seta of the antenna 
slightiy longer than the limb; poison-gland large and lobose. Mandible (fig. 12) stout, 
broad, and many-toothed; palp three-jointed, the long middle joint being again imper- 
fectly divided into three, terminal joint much narrower than the rest, finger-like, and 
bearing three broad apical sete and two short hairs; the long median joint has an 
apical brush of seven long hairs arising from a short external process; its inner 
margin has three very stout, curved, plumose sete, that of the distal segment being 
much longer than the other two; the middle of the outer margin has a fascicle of 
very short, fine hairs; the short basal joint bears two very small, plumose branchial (?) 
setee. Three pairs of feet, sparingly setiferous (figs. 13, 14), with strong curved ungues. 
Copulative organs complex (fig. 15), enclosing a coiled spermatic duct, with two stout 
plumose setz on the free margin. Female unknown. 

Hab. A few specimens taken in Lyttelton Harbour among alge. 

The antenne of this species with their long and stout urticating setz, and the 
mandible-palps with their slender terminal joint and stout, falcate sete, correspond 
closely with the same organs as seen in Cythere albomaculata Baird, and in no other 
species with which I am acquainted. 


CYTHERE INNOMINATA, nom. nov. (Plate XLVI. figs. 1, 2.) 
Cythere truncata’ Thomson (1), p. 254, pl. xi. fig. C. 2 a-c. 


Shell, seen laterally (fig. 1), subquadrate, highest in front, greatest height much more 
than half the length; anterior extremity broadly rounded, irregularly crenulated or 
almost dentated; posterior narrow, rounded off above and below, subtruncate below 
the middle, but deeply excavated above; dorsal margin sloping gently and in a slightly 
sinuous line from the front ; ventral nearly straight behind its junction with the anterior 
encircling fillet, gently upcurved toward the posterior extremity; seen from above 
(fig. 2), elongated, compressed, widest behind, nearly thrice as long as broad, lateral 
margins converging gradually, but with irregular sinuations from near the hinder end 
to the front, which is obtusely pointed; from the widest point the sides converge 


This specific name has been already used by Reuss for a Tertiary species. I therefore propose the new 
name innominata. 


444 DR. G. S. BRADY ON NEW OR 


abruptly to the posterior extremity, which is narrow and truncated; surface of the 
shell irregularly rugose, the ridges taking a transverse direction at the posterior end ; 
a conspicuous polished tubercle on the anterior hinge. Length 1:3 mm. 

Hab. In rock-pools, Brighton, New Zealand. ‘Two or three specimens only. 

The shell is somewhat like in general appearance to G. W. Miiller’s genus Para- 
cytheridea, but, so far as I have been able to observe, it does not agree with that genus 
in minor details. The types described by Mr. Thomson were got among alge in 
Otago Harbour. 


CYTHERE TRUNCULA, sp. n. (Plate XLVII. figs. 16, 17.) 

Shell, seen from the side (fig. 16), subquadrangular, greatest height near the front, 
and equal to about two-thirds of the length; anterior extremity obliquely and not very 
strongly rounded; posterior very wide, almost rectangularly truncated, produced below 
the middle, where it bears three short, stout, and blunt teeth; dorsal margin sloping 
rather steeply and irregularly from the front backward; ventral almost straight for 
about three-fourths of its length, where it forms a prominent abrupt angle, and thence 
curves upward to the posterior extremity; seen from above, the outline is lozenge- 
shaped or subhexagonal (fig. 17), widest in the middle, the width being nearly equal 
to the height ; extremities wide, truncated, and irregularly emarginate; lateral margins 
strongly arcuate, prominent in the middle, and converging rather steeply to each 
extremity. Surface of the shell covered with large irregularly-angulated fosse ; a large 
polished tubercle over the anterior hinge; anterior margin fringed with coarse hairs. 
Length ‘75 mm. 

Hab. Among weeds in Lyttelton Harbour. One specimen only. 


Genus TRACHYLEBERIS !, gen. nov. 


Shell beset with prominent nodules or spines. Antennules six-jointed, devoid of 
spines, but bearing numerous slender setz; antennz four-jointed, that of the female 
with a very short, faleate, urticating seta, which is absent from that of the male; 
mandible-palp with a small branchial plate bearing three upwardly-directed filaments, 
a very short one directed downward, and one very minute, horizontal ray. The 
remaining limbs as in Cythere; in front of the first pair of legs in the male a pair of 
one-jointed setiferous appendages. 


TRACHYLEBERIS SCABROCUNEATA (G. S. Brady). (Plate XLVII. figs. 1-7, 18-25.) 
Cythere scabrocuneata Brady (2), p. 103, pl. xvii. figs. 5 a—f; pl. xxiii. figs. 2 a-c. 

Shell of the male (figs. 1, 2), seen from the side, elongated, subquadrate, greatest 
height near the front, and equal to rather less than half the length ; anterior extremity 


" rpaxus, rough ; )e/Iypis, a shell. 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 445 


forming a steep curve above the middle, below the middle well rounded and broken up 
into a series of short blunt teeth; posterior extremity irregularly emarginate, narrowed 
and wedge-shaped, most prominent below the middle; dorsal margin slightly sloping 
from before backward, and broken into an irregular jagged line; ventral almost 
straight, with an upward bend behind. Seen from above, the outline is compressed, 
subovate, more than twice as long as broad, the whole circumference excessively 
Jagged and uneven, tapered at the extremities, which are obtuse, deeply emarginate in 
front; surface covered with large irregularly-rounded nodules, and beset with coarse, 
rigid hairs. A large polished tubercle over the anterior hinge; hingement of two 
terminal teeth on the right corresponding with depressions on the left valve. Length 
11 mm. Shell of the female shorter and more tumid. Antennules (figs. 18, 19) six- 
jointed, the last three joints bearing each three or four sete, each of which is about as 
long as the last four joints of the antennule; the second and third joints have one seta 
each, the first none; the last joint of the antenna (fig. 20) is small, and bears a long, slender 
unguis with two sete; the penultimate joint has three fascicles of long sete, and, near 
its base, a brush of small, short hairs; second joint small, bearing a single long seta 
and a few small hairs; first joint (in the female) with a short, strongly-curved, urticating 
seta—none in the male. Mandible wide and many-toothed; palp (fig. 3) four-jointed, 
its last joint rather shorter and much more slender than the preceding ; branchial plate 
(fig. 21) five-rayed; precrural appendages (fig. 25) elongated, quadrilateral, apices 
rectangularly truncate, and bearing numerous very fine long hairs; legs similar to 
those of the Cytheride generally (figs. 22-24), their terminal claws very long, slender, 
and moderately curved; the second joint of the last leg in the male (fig. 24) bears four 
small, but dense, fascicles of hairs; that of the female (fig. 4) has only one. Post- 
abdomen in the female (fig. 5) forming two trisetose lobes. 

Hab. Dredged abundantly in Lyttelton Harbour, depth 1-5 fathoms. 

This species was first described from specimens (shells only) taken during the cruise 
of the ‘Challenger’ in various depths, down as far as 40 fathoms: the localities were 
Kast Moncceur Island, Bass’s Straits; Inland Sea, Japan; and Wellington Harbour, 
New Zealand. ‘The very long and slender sete of the antennules—as long as those of 
many Cypride (Bairdia, Macrocypris, Paracypris, &c.)—seem to entitle this species to 
a separate generic position, and further distinctive characters are found in the 
peculiarities of the mandibular branchial plate, the urticating antennal sete, and 
the setose armature of the third pair of legs in the male. 


~ 


VOL. XIV.— PART vu. No. 3.—December, 1898. 


446 DR. G. 8S. BRADY ON NEW OR 


Genus CYTHERIDEIS Jones. 
Cytherideis T. R. Jones (1), p. 46; G. S. Brady (1), p. 454; Brady & Norman (1), p. 226. 


CYTHERIDEIS NOVA-ZEALANDIA, sp. n. (Figs. 1-4). 

Shell, seen from the side (fig. 1), suboval, elongated, three times as long as broad; 
extremities obliquely rounded, the anterior bordered with a narrow flange; dorsal 
margin quite straight, ventral straight, but indented at its junction with the anterior 
border; seen from above (fig. 2), ovate, more than thrice as long as broad, sides only 
slightly arcuate, anterior extremity subacuminate, posterior rounded, right valve over- 
lapped by the left; surface smooth. Antennule (fig. 3) six-jointed, bearing several 


Fig. 1. Fig. 2. 


Cytherideis nove-zealandic. 
Fig. 1. Shell, seen from left side, x 55. 
Fig. 2. 5 » above, x 55. 


Fig. 3. Fig. 4. 


Oytherideis nove-zealandic. 
Fig. 3. Antennule, x 210. 
Fig. 4. Mandible and palp, x 210. 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 447 


long sete, which are crowded together on the last four joints; mandible slender and 
feebly toothed (fig. 4); palp very long and slender, destitute of branchial rays, basal 
joint bearing one extremely long and stout, falcate, plumose seta, and two much 
shorter, simple sete; penultimate joint with two, last joint with three, small apical 


sete. Length -88 mm. 
Hab. One specimen only in a dredging from Lyttelton Harbour, 1-5 fathoms. 


Family CYTHERELLIDZ Sars. 


Genus CYTHERELLA Jones. 


CYTHERELLA EBURNEA, sp.n. (Plate XLVI. figs. 8, 9.) 

Shell, seen from the side (fig. 8), subelliptical, about twice as long as high: 
extremities well rounded, dorsal and ventral margins nearly straight. Seen from above 
(fig. 9), the outline is club-shaped, widest near the posterior extremity, width equal te 
more than one-third of the length; anterior extremity very obtusely rounded, posterior 
much wider, subtruncate, with rounded angles; lateral margins converging slightly 
from behind to the anterior extremity. Surface of the valves quite smooth and 
polished. Colour grey. Length -9 mm. 

Hab. Lyttelton Harbour, among weeds. One specimen only found. 


EXPLANATION OF THE PLATES. 


PLATE XLITI. 
Asterope australis, p. 431. 


Fig. 1. Shell of male, seen from left side, ] 
Fig. 2 5 zs above, 
: 40. 
Fig. Shell of female, seen from right side, € “a 
35 55 above, } 


. Inner branch of antenna, ¢ ‘| x 210. 


” 2° b 


. Process of mandibular foot, x 180. 
. Post-abdominal lamina, x 150. 


re} 
og 
Con SO OP 09 bt 


Asterope grisea, p. 432. 


Fig. 9. Shell of female, seen from right side, | x 25 


Fig. 10. a a above, J 
3Nn 2 


DR. G. S. BRADY ON NEW OR 


. Inner branch of antenna, ¢, X 100. 


” ” 2 > x 84. 


. Terminal teeth of vermiform limb, x 210. 
. Post-abdominal lamina, x 84. 


Cyclasterope zealandica, p. 433. 


. Shell of female, seen from right wa ie 


” ” above, 


. Shell of male, seen from right side, x 8. 
. Inner branch of antenna, ¢ | a 


tb} ihe} ’ 


. A terminal tooth of vermiform limb, x 540. 


91. Post-abdominal lamina, x 25. 


. Teeth at base of unguis of same, X 250. 
. Sculpture of anterior margin of shell, x 25. 


Cyclasterope ovulum, p. 432. 


. Shell of female, seen from left av x 6 


below, 


» 2»? 


. Inner branch of antenna, 2, x 40. 

. Frontal tentacle, x 40. 

. Terminal teeth of vermiform foot, x 250. 
. Post-abdominal lamina, x 25. 

. Marginal teeth of caudal ungues, x 250. 


PLATE XLIV. 
Philomedes flexilis, p. 435. 


” 3” below, 


. Old shell of female, seen from right side, 
| X 25 


. Shell of male from right side, 
. Eyes and frontal tentacle, ¢, X 84. 
. Inner branch, antenna, ¢, X 40. 


Led ” 2 id x 40. 


. Outer branch, antenna, 2, xX 84. 

. Principal tooth of second maxilla, x 210. 
. End of vermiform limb, x 210. 

. Post-abdominal lamina, x 84. 

. Hye of female, x 210. 


Fig. 12. 
Fig. 13. 
Fig. 14. 


Fig. 15. 


5 


Fig. 16. 
Fig, U7. 
Fig. 18. 
Fig. 19. 
Fig. 20. 


Fig. 21. 
Fig. 22. 
Fig. 23. 


Fig. 24. 
Fig. 25. 
Fig. 26. 


Fig. 27. 
Fig. 28. 
Fig. 29. 


al 
Cr) 
CO WD Or OD PO 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 


Portion of lining membrane of shell, x 84. 
Rostrum and laminar border, x 60. 
Shell-gland, x 200. 


Philomedes sculpta, p. 434. 


Shell of male, seen from left side, | 
Xx 20. 
” ” above, J 
Rostrum and marginal lamina, x 40. 
Inner branch of antenna, ¢, x 110. 
Post-abdominal lamina, x 84. 


Fragment of concretion, x 84. 


Eupathistoma natans, p. 437. 

Shell of male, seen from right side, 

5 “ below, 
Mandible, x 84. 

a, 6. Processes of same, X 210. 

Oral sense-organs, x 210. 
End of vermiform foot, x 210. 
Post-abdominal lamina, x 84. 


| x 26 


Cyclasterope tenera, p. 433. 


Right valve of lamina, seen from inside, X 30. 
Inner branch of antenna, < 84. 
Post-abdominal lamina, x 84. 


PLATE XLV. 


Sarsiella hansen, p. 438. 


. Shell of female, seen from right side, } 


35 5 above, | 


50. 


. Left valve, seen as transparent object, 
. Shell of male, seen from left side, 
. Antennule of male, x 84. 

. Inner branch of antenna, ¢, X 210. 

. Mandible of female. 


35 male, 


. End of vermiform limb, x 210. 

. Post-abdominal lamina, x 120. 

. Hairs from dorsal cushion, x 300. 
. Copulative organ, ¢, X 210. 


449 


450 DR. G. S. BRADY ON NEW OR 


Sarsiella hispida, p. 439. 


Fig. 13. Shell of female, from left side, x 50. 
Fig. 14. % $5 above, x 40. 


Philomedes flexilis, p. 435. 


Fig. 15. Shell of female, from right side, x 39. 
Fig. 16. os dorsal view (rather oblique), x 35. 


Asterope quadrata, p. 432. 
Fig. 17. Outline of shell ( ¢ ), seen from left side, 


Fig. 18. - * above, J ee 
Fig. 19. Inner branch of antenna, 3, x 210. 
Fig. 20. 55 Oe <All 


Fig. 21. Post-abdominal lamina, 2, x 210. 


Cypris viridis, p. 440. 
Fig. 22. Shell of female, seen from left fa x 50 
Fig. 23. a ef above, ; 
Fig. 24. Post-abdominal ramus, x 210. 


PLATE XLVI. 


Cythere innominata, p. 443. 


Fig. 1. Shell, seen from right side, 
| x 8a. 
Fig. 2. 5 A above, 


Loxoconcha punctata, p. 441. 
Fig. 3. Shell of female, seen from right side, 
: | x a4 
Fig. 4. a 3 above, 
Fig. 5. Portion of shell-margin, x 210. 


Xestoleberis olivacea, p. 442. 
Fig. 6. Shell of female, seen from left side, 
} x 84 
Fig. 7. Ap 55 above, 


Cytherella eburnea, p. 447. 
Fig. 8. Shell, seen from left side, 50 
Bigaeeg: 5 i above, } a 


Fig. 


Fig. 


. 28. 
» 29, 


Noor ond eH 


. Shell of male, seen from left side, 


IMPERFECTLY-KNOWN SPECIES OF OSTRACODA 


Xestoleberis compressa, p. 442. 


. Shell, seen from right el 84 


s 5 above, 


. Portion of shell-margin, x 210. 
. Antennule of female, 
. Antenna of female, | 
. Mandible and palp of female, x 210. 
. Foot of second pair, x 210. 


x 210. 


35 third pair, x 210. 


. Post-abdomen, x 250. 
. Copulative organ, ¢, X 250. 


Xestoleberis luxata, p. 441 


x 5 above, 
PA i behind, 


. Shell, seen from right side, 
| ante 


. Muscle-spots, x 210. 

. Antennule, x 210. 

. Antenna, x 210. 

. Mandible and palp, x 210. 

. Post-abdomen and third foot, x 210. 


Cyprinotus flavescens, p. 440. 


Shell, seen from right side, ze 
| x 20. 


6 “ above, 
PLATE XLVII. 


Trachyleberis scabrocuneata, p. 444. 


| x 53. 


) ” above, 


. Mandible and palp, x 210. 

. Foot of third pair, 2, x 120. 

. Post-abdomen, 2, X 210. 

. Copulative organ, ¢, X 100. 

. Portion of shell-margin, anterior, x 84. 


Cythere brunnea, p. 442. 


. Shell of female, seen from left side, 
| x $4. 


35 F above, 


45] 


452 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 
Fig. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


10. 
11. 
12. 
13. 
14, 
16. 


16. 
ie 


18. 
19. 
20. 
21. 
22. 
23. 
24, 
26, 


NEW OR IMPERFECTLY-KNOWN SPECIES OF OSTRACODA. 


Portion of shell-margin, x 250. 
Antennule, x 210. 
Mandible and palp, x 210. 
Foot of first pair, xX 210. 

» third pair, x 210. 
Copulative organ, ¢, X 210. 


Cythere truncula, p. 444. 


Shell, seen from left side, 
above, 


| x 84 


” bbs 


Trachyleberis scabrocuneata, p. 444. 


Antennule, x 110. 
Last four joints of antennule, x 250. 
Antenna of female, x 180. 
Branchial plate of mandible, x 210. 
Leg of first pair, ¢, x 210. 

» first pair, 29, X 210. 

» third pair, ¢, x 110. 
Precrural appendage, 2, X 210. 


Trans Loot. Joc Vib XN FLX 


leo Vout & Sona imp 
GS Brady del. 
Parker & Pereylith. 


OSTRACODA FROM NEW ZEALAND. 


an, Zoot. Jor. Vo XN TE XLV 


Wi 


e 


Vout & Sons imp 


Geo 


G 8.Brady del 
Parker & Perey lith 


OSTRACODA FROM NEW ZEALAND. 


Trans. Zoot. Soc. Yok XN. FEXLN. 


MOTT 


l 


Geo West & Sonaimp. 


GS Brady del 
Parker @ Pereylith. 
OSTRACODA FROM NEW ZEALAND. 


Geo Woot Sons imp 


Parker & Pereyhth. 


GS. Bradydel 


OSTRACODA FROM NEW ZEALAND, 


Trans. Zool. Soc. Yo XV ZF XIVIL 


Geo. West & Sons imp. 


GS. Brady del 
Parker & Percy lith 


OSTRACODA FROM NEW ZEALAND. 


453 


LIST OF THE PAPERS CONTAINED IN VOL. XIV. 


Bourencer, G. A., F.R.S., F.Z.S. 

On a Nothosaurian Reptile from the Trias 
of Lombardy, apparently referable to 
LUGS ok osoodocaeoadoscanoe 

On a Collection of Fishes from the Rio 
LEON socooc snob none Mogan aeaC 


On a Collection of Fishes from the Rio 
SAN UA MB LAAT  cctestays where. «fern ale win aie date 


Bravy, G. Srrwarpson, M.D., LL.D., F.R.S. 

A Supplementary Report on the Crustacea 
of the Group Myodocopa obtained during 
the ‘Challenger’ Expedition, with Notes 
on other new or imperfectly-known 
Species 


On new or imperfectly-known Species of 
Ostracoda, chiefly from New Zealand .. 


Bripven, T. W., Sce.D., F.Z.S., Mason College, 
Birmingham. 

On the Morphology of the Skull in the 

Paraguayan Lepidosiren and in other 

Dipnoids 


Ewes, H. J., F.R.S., F.Z.S., and Epwarps, 
James, F.E.S. 
A Revision of the Oriental Hesperiidae 


VOL. X1V.—ParT vill. No. 4.—December, 1898. 


Page 


325 


. 101 | 


| Goro, Dr. Eant A., C.M.Z.8., Director of the 
| Fara Museum. 
On the Lepidosiren of the Amazons ; being 
Notes on five Specimens obtained be- 
tween 1895-97, and Remarks upon an 
Example living in the Pari Museum .. 
| KH. Ray, M.A., F.RS., F.ZS., 
| Linacre Professor of Comparative Ana- 
tomy in the University of Oxford. 
On the Lepidosiren of Paraguay, and on 
the external characters of Lepidosiren 
and Protopterus 


| 
LANKESTER, 


| Tomas, Oxprienp, F.Z.S. 

On the Mammals obtained by Mr. John 
Whitehead during his recent Expe- 
dition to the Philippines ; with Field- 
notes by the Collector 


Vincent, Swartz, M.B.Lond., Demonstrator of 
Physiology and Assistant Lecturer on 
Histology, Mason College, Birmingham. 

| Contributions to the Comparative Anatomy 

| and Histology of the Suprarenal Cap- 
| sules.—The Suprarenal Bodies in Fishes, 
| and their Relation to the so-called 

Head-Kidney 


| 
| 


30 


Paga 


413 


11 


377 


INDEX OF SPECIES, 


EI¢., IN VOL. XIV. 


Abkaratha alida, 151. 
hyperides, 149. 
—— pygela, 149. 


ransonnettiz, 150. 


saraya, 151. 

sura, 149. 

—— syrichthus, 151. 

taylorii, 150, 151. 
Abraximorpha davidii, 123. 
Acanthias vulgaris, 50, 52, 66, 82, 84. 
Acanthopsis tenia, 45, 


Acara bimaculata, 26. 


tetramerus, 26. 

Acerbas anthea, 215. 

duris, 215, 216. 

martini, 215, 216, 312. 
nitidifasciata, 215, 216, 309, 312. 
Acestra gladius, 425, 428. 

oxyrhyncha, 34, 425. 

Achalarus bifasciatus, 108, 109. 

, var. contractus, 109. 


casyapa, 109. 

frater, 109, 110. 
germanus, 109, 110. 
liliana, 108, 109. 
nepos, 109, 110. 
proximus, 108, 109, 
simplex, 108, 109. 
Achlyodes sura, 149. 
vasava, 148. 
Acipenser sturio, 50, 55, 82. 
Actinor radians, 243, 244, 
Adopea actewon, 288, 289. 
astigmata, 290, 
hamza, 289. 

hyrax, 288, 289. 
leonina, 288, 290. 

, var. astigmata, 290. 


Adopea lineola, 288, 289. 
nervulata, 288, 290. 

stigma, 288, 289. 

sylvatica, 288, 289, 290. 
tenebrosa, 288, 289, 290. 
— thaumas, 288. 

Aeromachus catocyanea, 189, 192. 
chinensis, 187, 189, 190. 
— discreta, 188, 190, 191, 308. 
—— dubius, 188, 190, 308, 312. 
—— inachus, 187, 188, 190, 308, 311. 
—— indistincta, 188, 191, 308, 312. 
—— javanicus, 188, 191, 308. 
—— jhora, 188, 190, 191, 312. 
kali, 187, 189, 192. 

—— musca, 188, 189. 

—— nanus, 189, 192, 312. 

—— piceus, 188, 189. 

stigmata, 187, 189, 308. 
Ageniosus brevifilis, 27. 

Alligator sclerops, 12. 


Amiurus catus, 64. 
Ammodytes tobianus, 45, 46. 
Ampittia delai-lama, 186, 187. 
—— mesoides, 184. 

—— maga, 186. 

—— maro, 186. 

—— muroides, 186. 

— trimacula, 186, 187. 
—- virgata, 186. 

ziclea, 184. 

Anacyrtus affinis, 426. 
kenerit, 426. 


—— limesquamis, 426. 


macrolepis, 39, 
—— microlepis, 36. 
prognathus, 36, 39. 


Anarosaurus pumilo, 10. 


456 INDEX OF 

Anarrhichas lupus, 51, 62, 72, 83, 84. | 

Anatomy (Comparative) of the Suprarenal Capsules, | 
Contributions to the, by Swale Vincent, 41-84. | 

Ancistroides longicornis, 222. 

othonias, 222, 310. 

Ancylowypha ardonia, 184. 

Anguilla anguilla, 50, 56, 70, 74, 77. 

Antigonus kethra, 174. 

Aphaustus discreta, 190, 191. 

durga, 201. 


Apostictopterus fuliginosus, 173. 
Arapaima gigas, 417, 427. 
Arnetta atkinsoni, 198, 199. 
—— vindhiana, 199, 308. | 
Asterope australis, 429, 431, 447. 
—— gresia, 429, 432, 447. 
quadrata, 429, 432, 450. 
Astictopterus armatus, 221. 
butleri, 175. 

gemmifer, 221. 

henrici, 171, 308. 

jama, 171, 172, 173. 

kada, 172. 

kethra, 174. 

olivascens, 172, 308. 
othonias, 222. 

pygmeus, 172. 

salsala, 181, 182. 

sindu, 182. 

subfasciatus, 174. 

ulunda, 174. 

verones, 176. 


wanites, 175. 


, var, palawites, 175. 
Astictopteryx stellifer, 181. 
Aubertia dulcis, 168. 
Auchenipterus galeatus, 28. 
Auyiades bouddha, 248. 

, var. consors, 248. 
brahma, 246, 248, 313. 

—— cruteis, 246, 248, 313. 
herculea, 249. 

—— majuscula, 246, 249, 309, 313. 
—--— ochracea, 246, 248, 249, 312. 
similis, 247. 

siva, 246, 247, 248, 309. 
subhyalina, 246, 247, 312. 
—— sylvanoides, 246, 247, 312. 


SPECIES, 


Augiades sylvanus, 104, 246, 247, 24), 
Badamia exclamationis, 306. 
Balistes maculatus, 50. 


| Baoris austeni, 280. 


chaya, 275, 279. 

distinctus, 286. 

insignis, 261. 

kumara, 276. 

moolata, 278. 

narooa, 280. 

oceia, 274. 

penicillata, 274. 

scopulifera, 274. 

seriata, 276. 

sikkima, 274. 

unicolor, 198, 274, 285. 
—— (Parnara) philotas, 285, 
Baracus hampsoni, 170, 171, 308. 
plumbeolus, 170, 171. 
septentrionum, 170, 171, 185. 
—- subditus, 170, 171, 185. 
vittatus, 169, 170. 

Batomys granti, 378, 404, 405, 406, 411, 412. 
Bdellostoma forster, 75. 

Belone teniata, 37. 


vulgaris, 45. 

Bibasis sambavana, 305, 315. 

sena, 304, 305, 306. 

, var. palawana, 305. 

uniformis, 305, 306, 315. 

Boulenger, G. A. On a Nothosaurian Reptile from 
the Trias of Lombardy, apparently referable to 


Lariosaurus, 1-10. 

On a Collection of Fishes from the Lio 
Paraguay, 25-39. 

On a Collection of Fishes from the Rio 
Jurua, Brazil, 421-428. 
Brachychaleinus retrospina, 36. 


Brady, G. Stewardson. On new or imperfectly- 
known Species of Ostracoda, chiefly from New 
Zealand, 429-452. 

Bridge, T. W. On the Morphology of the Skull in 
the Paraguayan Lepidosiren and in other 
Dipnoids, 325-376. 

Buhalus mindorensis, 380, 41. 

Bufo ornatus, 340. 

Bunocephalus iheringii, 34. 

Calliana pieridoides, 106. 


INDEX OF 


Callichthys armatus, 424. 
—— asper, 29. 
longifilis, 29. 
—— paleatus, 29. 
pectoralis, 29, 38. 
thoracatus, 29. 
Caltornis onchisa, 278, 280. 
Cantharus griseus, 51, 61. 


Capila jayadeva, 106, 107. 
translucida, 107. 
zennara, 106, 107. 
Caprona alida, 150, 151. 
—— elwesi, 151. 


—— ransonneitii, 150, 311. 

saraya, 150, 151, 308, 311. 

—— syrichthus, 150, 51: 

Carcharodus alcee, 152. 

althee, 152. 

lavatere, 152. 

swinhoci, 152. 

Carpomys melanurus, 378, 404, 406, 407, 408, 411, 
412, 

—— pheurus, 378, 404, 408, 411, 412. 

Carponycteris australis, 385. 


Carterocephalus abax, 166. 
argyrostigma, 167. 
christophi, 168. 
demea, 168. 
dieckmanni, 168. 
flavomaculatus, 167. 
gemmatus, 168. 


houangty, 167. 
micio, 168. 


niveomaculatus, 167. 


—— ops, 167. | 


palemon, 166. 
Carystus albescens, 180. 
duris, 216. 

— ladana, 122. 
— lenas, 228. 
mabillet, 216. 
—— singularis, 241. 


tagiadoides, 215. 
—— telesinus, 228. 
Casyapa lidderdali, 105. 


phanceus, 105, 
Celenomys, gen. noy., 390. 
silaceus, 378, 390, 391, 411, 412. 


Cf 


SPECIES, 45 


Celeenorrhinus affinis, 114, 119, 121, 122, 308, 310. 
ambareesa, 112, 115. 
andamanica, 114, 119, 310. 
asmara, 113, 118. 

aspersa, 112, 1165. 

—— awurivittata, 111, 114, 122, 310. 
, yar. cameront, 122. 
badia, 114, 125. 

balukinus, 113, 117, 307. 
batchianus, 114, 122, 308. 
—— buchananii, 129. 


—— cacus, 118. 
cameroni, 114, 121. 


| —— chamunda, 113, 117. 


—— clitus, 115. 
consanguinea, 112, 115. 


consertus, 118. 

dentatus, 114, 119, 121, 307. 
dhanada, 114, 119, 121, 310. 
eligius, 111. 

flavocincta, 112, 115. 
fulvescens, 114, 120, 307. 
goto, 118. 

inequalis, 114, 119, 307. 
ladana, 114, 122, 176. 
lativittus, 114, 121, 308. 
— leuweocera, 111, 112, 113, 116. 
, var. angustipennis, 113, 116, 117. 


, var. brahmaputra, 117. 
—— lucifera, 116. 

maculicornis, 113, 116, 308. 
maculosa, 112, 115. 

munda, 116. 

migricans, 113, 117, 118. 
omeia, 105, 106. 

orbiferus, 113, 118, 307. 
palajava, 118. 

patula, 116. 

pero, 112, 115. 

—— pinwilli, 145. 


plagifera, 116. 

—— pluscula, 116. 

—— pulomaya, 112, 115. 

—— putra, 116. 

—— pyrrha, 116. 

saturatus, 114, 119, 120, 308, 310. 
spilothyrus, 113, 117. 

sumitra, 111, 113, 115, 116. 


458 INDEX OF SPECIES. 


Celenorrhinus tabrica, 146, Coladenia dan, var. andamanica, 119. 
tibetana, 113, 119. , var. dea, 127. 
Centromochlus heckelit, 422. Fatih, 127. 

Ceratodus, revision and comparison of the structure hamiltonii, 128. 

of the skull in, 350-356, —— igna, 125, 128. 

forsteri, 11, 20, 21, 375. indrani, 125, 126. 

Cestracion philippi, 340. lawmi, 125, 126, 127, 128, 129, 310. 
meeniata, 130, 

sempert, 125, 128. 

sobrina, 125, 126, 308. 

tissa, 125, 126. 

vitrea, 129, 130. 


Cetengraulis juruensis, 427, 428, 
Cetopsis candiru, 422. 
Uhetienema lidderdali, 105, 
Cheetostomus aculeatus, 31. 

bachi, 425, 428. 


cirrhosus, 31. Conger conger, 50, 56, 70, 82, 84. 
gigas, 30, 39. Coris pulcherrima, 51, 60. 
leucostictus, 31. Cottus gobio, 51, 61, 83. 
ologospilus, 425. quadricornis, 44, 
Chaleinus nematurus, 36, 426, Crateromys schadenbergi, 378, 400, 409, 412. 
paranensis, 36. Crenicichla johanna, 26. 
Chapra agna, 275. saxatilis, 26. 
cere, 275. Creteus cyrina, 233. 
—— mathias, 275. Crocidura grayt, 378, 387. 


—— nostrodamus, 286. 


luzoniensis, 387. 
—— prominens, 275, 


murina, 387. 


subochracea, 275. Crossiura pennicillatum, 107, 108. 
Charmion ficulnea, 110, 111, 238. Crunomys, gen. nov., 393. 

tota, 111. fallax, 378, 393, 394, 411. 
Chimera monstrosa, 45, 50, 55, 80, 82. Crustacea, supplementary report on the Myodocopa 
Chirodon alburnus, 426. obtained during the ‘ Challenger’ Expedition, 
Chironectes punctatus, 44. by G. S. Brady, 85-100. 
Choaspes amara, 295. Ctenoptilum chinensis, 148, 311. 


—— anadi, 294. 
—— benjamini, 307. 


multiguttata, 148. 
vasava, 147, 148, 311. 


— chuza, 304. Cupitha purreea, 245. 

—— crawfurdi, 307. ——. tympanifera, 245, 
—— electra, 307. Curimatus alburnus, 34, 426. 
—— harisa, 293. dobula, 426. 


renidens, 307. 
vasutana, 294. 


latior, 34. 

rutiloides, 34, 

Chrotomys whiteheadi, 378, 390, 392, 411, 412. Cyclasterope ovulum, 429, 432, 448. 
Clupea harengus, 46, 50, 57, 82. tenera, 429, 433, 449. 
nilotica, 44. zealandica, 429, 433, 448. 


Cobalus philippina, 276. Cyclopides camertes, 186. 
sulphurifera, 264. ceramas, 184, 

Cobitis fossilis, 45. —— delai-lama, 187. 

Coladenia agni, 125, 128, 129, 310, 311. —— henrici, 172. 
agnioides, 125, 128, 308, 311. — ornatus, 169. 

-—— buchananii, 129. —— subradiatus, 185. 


dan, 125, 127, 310. —— subvittatus, 185. 


INDEX OF 


Cyclopides unicolor, var. ornatus, 169, 
Cyclopterus lumpus, 51, 63, 74, 77, 83. 
Cynodon pectoralis, 426. 


vulpinus, 426. 

Cypridina zealandica, 433. 
Cyprinotus flavescens, 429, 440, 451, 
Cypris ciliata, 430. 

-—— littoralis, 430. 
nove-zealandie, 430. 
viridis, 429, 440, 450. 
Cythere albomaculata, 443. 

— atra, 430. 

— brunnea, 430, 442, 451. 
— scabrocuneata, 444. 

— thomsoni, 430, 443, 450. 
truncula, 430, 444, 452. 
Cytherella eburnea, 430, 447, 450. 
Cytherideis nove-zealandie, 430, 446. 
Dactylopterus volitans, 74. 

Daimio dire, 133. 

felderi, 135. 

sinica, 135. 

tithys, 135. 

Darpa hanria, 137. 


Dejeania bicolor, 169. 

Diagramma punctatum, 44. 

Diodon novemmaculatum, 45. 

Dipnoi, the structure of the skull in fossil, 366-372. 

Dipterus valenciennesi, 367, 369, 370. 

Doras costatus, 28. 

—— maculatus, 28. 

weddeliz, 28. 

Echeneis remora, 45. 

Edwards, James, and Elwes, H. J, A Revision of 
the Oriental Hesperiide, 101-324, 

Eetion elia, 236. 

Eleutherura philippinensis, 383. 


Eliomys quercinus, 405. 

Elops salmoneus, 45. 

Elwes, H. J., and Edwards, James. A Revision of 
the Oriental Hesperiide, 101-324. 

Engraulis olidus, 37. 

Entheus bicolor, 136. 

Evionota acroleuca, 217. 

-—— alexandra, 217. 

—— grandis, 217. 

lalita, 105. 

sanguwinocculus, 218. 


SPECIES. 459 
Erionota thrax, 217. 

—— (Casyapa) mabillet, 234, 235. 
Erynnis comma, 287. 

-—— -——, var. catena, 287. 

, var. dimila, 287. 

, var. florinda, 287. 
—— —., var. repugnans, 287. 
dimila, 287. 

Esococtus eariliens, 45. 

Esow lucius, 45, 57, 74, 83. 
Euanemus nigripinnis, 28, 38. 
Eudamus bifasciatus, 109. 

calathus, 231. 

decoratus, 136. 

Frater, 110. 

gener, 109. 


germanus, 110. 
guttatus, 281, 
—— infernus, 110. 


nepos, 110. 


phaneus, 105. 


proximus, 109. 


simplex, 109. 

-—— spilothyrus, 117. 

Eupathistoma, gen. nov., 437. 

natans, 429, 437, 449. 

Felis domestica, 378, 387, 388. 

—— minuta, 387. 

Fishes from the Rio Jurua, by G. A. Boulenger, 
421-428, 

Fishes from the Rio Paraguay, by G. A. Boulenger, 
25-39. 

Fishes, the Suprarenal Bodies in, and their relation 
to the so-called Head-Kidney, by Swale Vin- 
cent, 41-84. 

Gadus eglefinus, 50, 58, 71, 72, 83. 

barbatus, 45. 

merlangus, 50, 58, 59. 

morrhua, 50, 58, 59, 72, 83. 

virens, 50, 58. 

Galeopithecus philippinensis, 386. 


Gangura sanguinocculus, 218, 309. 


sybirita, 218. 

thyrsis, 217, 218. 
Gastropelecus pectorosus, 426, 
stellatus, 36, 426. 
Gastrosteus spinachia, 51, 61. 
Ge geta, 182. 


460 


Gegenes javana, 280. 


nostrodamus, 286. 


pygmeus, 286. 

Gehenna abima, 244. 

angulifera, 244, 309. 

gree, 244, 245, 

Geophagus duodecimspinosus, 26, 38. 

Goeldi, Emil A. On the Lepidosiren of the Amazons, 
413-420. 

Gomalia albofasciata, 153. 

littoralis, 153. 

Goniloba badra, 298. 

conjuncta, 280. 

cruda, 234. 

—— gopala, 132. 

guttatus, 281, 

sambara, 134. 

senda, 305. 

Gymnotus electricus, 418. 


Halpe acuma, 265. 

—— aina, 259, 267. 
albipectus, 260, 267. 
astigmata, 257, 258, 260. 
beturia, 263, 310. 

—— bivitia, 260, 268. 

—— blanchardi, 257, 259, 267. 
brunnea, 259, 265. 

cenis, 262. 

cerata, 258, 260. 
ceylonica, 258, 263. 
debilis, 259, 266, 309. 
decorata, 268. 


—— dolopia, 192, 

—— fasciata, 258, 262, 310. 

—— fusca, 259, 266, 313. 

gupta, 259, 266. 

hieron, 257, 261, 265. 
homolea, 259, 265, 266, 313. 
honoret, 260, 268. 

—— hyrie, 258, 261. 

insignis, 258, 261. 

—— knyvetti, 258, 261, 262, 309. 
kumara, 258, 261, 262. 
latris, 266. 

—— lucasii, 262. 

majuscula, 259, 264, 309. 
marta, 265. 


INDEX OF 


SPECIES. 


Halpe masoni, 260, 268, 
—— moorei, 258, 263. 
— nephale, 258, 264. 
—— ormenes, 257. 


ornata, 268. 
—— palawea, 264, 


perara, 265. 

radians, 243, 244. 

separata, 257, 259, 267. 
sikkima, 259, 264, 265, 313. 
sitala, 259, 266. 

subflava, 257, 259, 267. 
submacula, 257, 258, 263. 
sulphurifera, 259, 264. 
teliga, 263. 

varia, 259, 263, 266. 
wantona, 265. 

zema, 257, 260. 

Hantana infernus, 110. 
Harpyionycteris whiteheadi, 384, 411. 
Hasora anura, 296, 298, 299. 
badra, 296, 298, 299. 
borneensis, 297, 302, 309. 
celenus, 297, 299. 

chabrona, 297, 300, 302. 
chromus, 297, 300, 301, 302, 315. 
chuza, 298, 304. 

coulteri, 300. 

—— gneus, 296, 298. 

hadria, 298. 

inermis, 297, 301, 309, 315. 
meestissima, 297, 300. 

mus, 297, 304, 309. 

myra, 297, 303. 

—— pro«wimata, 297, 301, 302. 
— prowissima, 297, 302, 310. 
saida, 298, 304. 

—— simplicissima, 297, 299. 
—— violacea, 297, 299. 

vitta, 300, 302. 
Head-Kidney, function of the lymphatic, 78. 


, histology of the, 76. 

, nature of the intertubular material in the, 
76. 

, relation of the suprarenal bodies to the, 73. 


, relation of the suprarenals to the, 78. 
Hemiodus microlepis, 34. 


INDEX OF SPECIES. 461 


Hemiodus semiteniatus, 34. Hesperia ewlepis, 229. 
farri, 274. 

fateli, 127. 

acroleuca, 217. —— feralia, 206. 

—— adrastus, 197. —— ficulnea, 111. 
fortunei, 281. 

galba, 154, 157. 
geron, 154, 158, 311. 


Henisorubim platyrhynchus, 27. 


Hesperia abima, 244. 


—— agna, 275. 

ali, 154, 158. 
—— alpina, 155, 160. 
—— alveus, 156, 160, 161, 311. gigas, 155, 159. 
, var. alpina, 160. gremius, 179. 
andromede, 156, 161. — hamza, 289. 
anthea, 215. hellas, 157. 
— antonia, 157, 163. hiraca, 217. 


—— aria, 210. —— homolea, 265. 
— atticus, 143. —— hyela, 223. 
— attina, 234. —— hyracx, 289. 
avesta, 233. —— hyreana, 247. 
bada, 281. trava, 235. 
barea, 269. jolanda, 284. 
beturia, 263. — karsana, 286. 
bevani, 283. —— kwehni, 252. 
—— bieti, 156, 162. — kumara, 276. 
— hononia, 204, 206. —— latoia, 227. 


cacalie, 156, 161. 
cahira, 278. 


latreilla, 234. 
lebadea, 219. 


callineura, 226. -—— leucocera, 116. 
carthami, 153, 155, 160. — leuzee, 159. 
cashmirensis, 155, 160. liburnia, 228. 


celsina, 224, —— maculatus, 157, 162. 
centauree, 156, 161. meevins, 182, 183. 

— cephala, 204, 205. malue, 153, 155, 156, 160, 161, 311. 
— cephaloides, 206. —— malvoides, 156, 160, 311. 
cerata, 260. mandan, 166. 

chaya, 275. mare, 185, 186. 
chrysoxona, 252. mathias, 275. 

— colaca, 283. — melotis, 156, 161, 311. 
commu, var. catena, 287. —— moolata, 278. 

mytheca, 233. 

naga, 229. 


corissa, 229. 
cribrellum, 154, 158. 
cynare, 154, 158. 


narova, 280. 
nobilts, 155, 159. 


cyrina, 233. 


dara, 254. —— noma, 159. 
divodasa, 179. —— nondoa, 281. 
— dolopia, 192. —— oberthiiri, 157, 162. 
eacus, 127. oceta, 274. 
elia, 236. ogygia, 203. 
eltola, 282. —— onopordi, 156, 161, 311. 


VOL. XIV.—PART vill. No. 5.—December, 1898. 3P 


462 


Hesperia orbifer, 154, 158. 
ormenes, 260. 

ornata, 268. 

—— pandia, 218. 

—— phiditia, 180. 

phlomides, 154, 158, 311. 
pheenicis, 197. 

plumbeola, 171. 

pogget, 155, 159. 

—— proto, 153, 155, 159, 311. 
sala, 179. 

sao, 154, 158. 

semamora, 269. 


—— seriata, 276. 

—— sericea, 164. 

—— serratule, 156, 160, 311. 
—— side, 157, 162. 

—— speyeri, 156, 160, 311. 
—— staudingeri, 155, 159, 311. 
—— subgrisea, 179. 

—— subhyalina, 247. 

swerga, 177. 

—— sybirita, 218, 236. 

—— syrichtus, 153. 

taras, 161. 

tessellata, 229. 

tessellum, 155, 159. 
therapne, 154, 158. 
thibetanus, 157, 162.. 
—— toona, 281. 


—— venata, 247. 

—— vermiculata, 227. 

— vitia, 300. 

—— zebra, 157. 

zellert, 284. 

zema, 260. 

zona, 157, 162. 

Hesperiide, a revision of the Oriental, by H. J. 
Elwes and James Edwards, 101-324 

Hesperilla blanchardi, 267. 


lucaszi, 262. 


luteisquama, 264. 
Heteropterus cataleucos, 178. 


morpheus, 168. 


ornatus, 168. 


unicolor, 168. 
—— (Steropes) scopas, 178. 


INDEX OF 


SPECIES. 


Hidari bhawani, 236. 


| —— doesoena, 236. 


irava, 235, 236. 

staudingeri, 235, 236. 

sybirita, 218, 235, 236. 

Hippoglossoides limandoides, 51, 59, 60, 83. 

Hippoglossus vulgaris, 51, 59, 60, 83. 

Hipposiderus diadema, 385. 

Histology of the Suprarenal Capsules, Contributions 
to the, by Swale Vincent, 41-84. 

Hyarotis adrastus, 197. 


Hypoptopoma bilobatum, 31. 


guentheri, 31, 425. 

— steindachneri, 31. 

—— thoracatum, 31. 

Lambria latifascia, 181, 182, 310. 
salsala, 181. 

sindu, 181, 182. 

stellifer, 181. 

Idmon unicolor, 198. 


Isma bononia, 206. 


inarime, 206. 
isota, 205. 
obscura, 196, 204. 


' Ismene amara, 292, 295. 


anadi, 291, 294, 

—— aquilina, 292, 296. 
—— aria, 209. 

—— ataphus, 292, 293. 
— batara, 234. 

, var. celebica, 298. 
— chabrona, 300. 

— chuza, 304. 

—— druna, 210. 

etelka, 291, 293. 
Fergusonii, 291, 293, 295. 
—— gneus, 298. 

gomata, 292, 295. 

, var. lara, 295. 
—— harisa, 291, 293, 294. . 
wonis, 290, 291, 295. 
—— jaina, 291, 293, 295. 
—— junkowskii, 296. 

—— jayadeva, 106, 107. 
lara, 292, 295, 309. 
lorquini, 295. 


muestissima, 300. 


Ismene mahintha, 290, 291, 294, 295. 


malayana, 301. 
—~ murdava, 237. 
— myra, 303. 

—— edipodea, 291, 292. 
—— edipus, 291, 292. 
philetas, 299. 
proximata, 301. 


—— quadripunctata, 298. 
—— renidens, 307. 

—— saida, 304. 

—— sasivarna, 210. 


—— sena, var. palawana, 305. 


—— septentrionis, 292, 294. 
—— simplicissima, 299. 
—— striata, 294. 

—— subcaudata, 307. 

—— subfasciata, 219. 

—— thymbron, 306. 

—— tuckeri, 291, 293, 309. 
—— vasutana, 291, 294. 
violaceus, 299. 


Tsoteinon atkinsont, 199. 


cephala, 205. 
cephaloides, 206. 
corissa, 229. 
— durga, 201. 
flavalum, 202. 
flavipennis, 202. 
indrasana, 229. 


— khasianus, 199. 
—— lamprospilus, 197. 
—— masoni, 268. 

-—— masuriensis, 192, 193. 
—— microstictum, 200. 
—— modesta, 199. 

—— nilgiriana, 199. 
—— pundita, 192, 193, 
—— satwa, 201, 202. 
—— subtestaceus, 199. 
—— vindhiana, 198, 199. 
—— vittatus, 170. 

Ttox semamora, 269. 

— watsonii, 269. 

Itys iadera, 200. 

—— microstictum, 200. 


Kerana armata, 176, 220, 221. 


INDEX OF SPECIES, 463 


| Kerana aurivittata, 122. 


, var. cameroni, 121. 

— dhanada, 119. 

— diocles, 221, 222. 

— fulgur, 221. 

gemmifer, 175, 221. 

Kerivoula hardwickei, 386. 

whiteheadi, 386. 

Koruthaialos butlert, 175. 

hector, 175. 

kerala, 176. 

Icophene, 176. 

verones, 176. 

wanites, 174, 175. 

Labrax lupus, 51. 

Lankester, E. Ray. On the Lepidosuen of Paraguay , 
and on the external characters of Lepidosiren 
and Protopterus, 11-24. 

Lariosaurus balsami, 1, 2, 3, 8, 10. 


Lenomys, gen. nov., 409. 

—— meyert, 409, 412. 

Lepidosiren, comparison of the measurements of the 
Brazilian and Paraguayan specimens of, 24, 

——, morphology of the skull in the Paraguayan 

and in other Dipnoids, 325-376. : 

, revision. and comparison of the structure of 

the sknll in, 363-366. 

of the Amazons, by Emil A. Goeldi, 413-420, 

, anus of, 415. 

, colour of, 415. 

-—, geographical distribution of, 414. 


, habits of, in captivity, 418. 
——, measurements of, 415. 
——. popular names of, 417. 


, segmentation of limb-axis of, 416. 


, sex and measurements of, 414, 
, system of lateral lines of, 416. 
of Paraguay, and on the external characters 


of Lepidosiren and Protopterus, by E. Ray 
Lankester, 11—24. 

appendiculata, 51. 

articulata, 13, 21, 24, 325, 416. 

—— gigliolana, 23. 

paradoxa, 11, 13, 14, 24, 51, 80, 325, 326, 
345, 349, 

Leporina eques, 34. 


striatus, 34. 


BP 2 


464 INDEX OF 

Leuciseus cephalus, 50, 58, 83. 

rutilis, 50, 58, 83. 

vulgaris, 50, 58, 83. 

Lombardy, on a Nothosaurian Reptile from the 
Trias of, by G. A. Boulenger, 1-10. 

Lophius piscatorius, 44, 51, 62, 74, 77, 83. 

Lophoides binotatus, 196, 308. 

—— iapis, 195, 196, 312. 

—— obscura, 195, 196. 

purpurasceus, 196, 308, 312. 

vulso, 196. 

Loricaria acipenserina, 425. 


anus, 33. 
apeltogaster, 33, 39. 
evansi, 33. 

—— filamentosa, 33, 425. 
—- labialis, 32, 39. 
— lamina, 34. 

lata, 33. 

macrodon, 33. 
maculata, 33. 

—— nudiventris, 33. 
parva, 32, 39. 
—— rostrata, 32, 425, 


spixi, 33. 

Lota vulgaris, 45. 

Lotongus avesta, 230, 231, 233. 
calathus, 230, 231, 232. 
duris, 216. 

excellens, 230, 231, 233. 
maculatus, 231, 232. 

mytheca, 230, 231, 233. 

—— parthenope, 231, 232. 

sarala, 230, 231, 233. 

—— zenon, 230, 231, 232. 

zeus, 230, 231, 232. 
Loawoconcha punctata, 430, 441, 450. 
Macacus cynomolgus, 381. 

Macrodon trahira, 417. 
Macromerosaurus plinw, 1. 

Matapa aria, 208, 209. 

druna, 208, 209, 210, 312. 
—— purpurascens, 208, 209, 309, 312, 
—— sasivarna, 208, 209, 210. 
—— shalgrana, 208, 209, 210. 
subfasciata, 220. 

Merluccius vulgaris, 51, 58, 59, 71, 83. 
Mesonauta insignis, 26. 


SPECIES. 


Mesosaurus tenuidens, 4. 

Miniopterus schreibersi pusillus, 386. 
Molva vulgaris, 51, 58, 59, 72, 76, 83. 
Mora mediterranea, 74, 75. 

Motella mustela, 45. 

—— tricirrhata, 51, 58. 

Mugil capito, 51, 62, 83. 

Mullus barbatus, 51, 61, 63, 71, 83, 84. 
Mus chrysocomus, 378, 403. 
decumanus, 378, 402. 

ephippium negrinus, 378, 403, 404. 
everetti, 378, 400, 401. 

luzonicus, 378, 401, 404, 412. 
mayeri, 409, 

mindorensis, 402, 404. 

neglectus, 402, 403, 

rattus, 402, 403. 

rufescens, 402. 


Myletes albiscopus, 426. 
—— asterias, 37. 


brachypomus, 37. 


duriventris, 37. 

hypsauchen, 37. 

Myodocopa, supplementary report on the, obtained 
during the ‘ Challenger’ Expedition, by G. 8S. 
Brady, 85-100. 

Myotis macrotarsus, 385. 


Nannosciurus concinnus, 389, 390. 
samaricus, 389, 390,411. 
Nanostomus lateralis, 34. 


Netrocoryne atilia, 126. 

Neusticosaurus pusillus, 6, 7, 10. 

New Zealand, on new or imperfectly-known species 
of Ostracoda from, by G. 8. Brady, 429-452. 

Nisioniades dasahara, 124, 125. 

diocles, 222. 

erebus, 164. 

montanus, 164. 


pelias, 164. 
salsala, 180,181. 
tages, var. sinina, 163. 


Normyrus oxyrhynchus, 44. 

Nothosaurian Reptile from the Trias of Lombardy, 
apparently referable to Lariosaurus, by G, A. 
Boulenger, 1-10. 

Nothosaurians, References to the Literature on, 9. 

Notocrypta albifascia, 239, 240, 

alysos, 238, 239. 


INDEX OF SPECIES. 


Notocrypta basiflava, 239, 241. 
curvifascia, 238, 239, 240, 
feisthamelii, 238, 239, 240, 242. 
, var. rectifasciata, 239, 
goto, 118. 

inornata, 239, 241, 310. 
microthyrus, 242. 

monteithi, 215, 239, 241. 
neera, 242, 


paralysos, 239, 241, 
—— quadrata, 239, 241, 309. 
restricta, 239, 240, 

—— tibetana, 119. 

volux, 240. 

Ochus subvittatus, 185. 

Odina chrysomelena, 135, 136. 
—— cuneiformis, 135, 136. 
decoratus, 135, 136. 
hieroglyphica, 135, 136. 


ortygia, 137. 
Odontoptilum helias, 148, 149. 
, var. helisa, 149. 


—— hyperides, 149. 


leptogramma, 148, 150. 
pygela, 148, 149. 
sura, 148, 149. 

Grane microthyrus, 242, 


neeera, 242. 

Onryza meiktila, 269. 

Orthagoriscus mola, 50, 56, 72, 77, 82. 

Orthophetus lalita, 104, 105. 

lidderdali, 104, 105, 309. 

omeia, 104, 105, 106, 

phaneus, 104, 105. 

Osmerus eperlanus, 58, 74, 83. 

Ostracoda, on new or imperfectly-known Species of, 
by G. Stewardson Brady, 429-452, 

Otocinclus affinis, 32. 

Oxydoras bachi, 423, 428. 

brevis, 424. 

—— eigenmanni, 28, 38. 

— elongatus, 424, 428. 

—— humeralis, 424. 


stenopeltis, 422. 
trachyparia, 423, 428. 
trimaculatus, 422, 428. 
Pachypleura edwardsti, 2, 7, 9. 
Pachyurus schomburgki, 25, 


465 


Padraoma chrysozona, 252. 


goloides, 253. 


maga, 186. 


palmarum, 252. 


por, 256. 

— trimacula, 187. 

virgata, 186. 

Paduka glandulosa, 219, 220. 
lebadea, 219. 

Pagellus centrodontus, 51, 61, 83. 
Pamphila abax, 165, 166 
akar, 254, 

albivitta, 268. 

archias, 185. 
argyrostigma, 166, 167. 


augiades, 253. 


, var. bLambuse, 252. 
avanti, 165, 167. 

—— bambuse, 251. 

bivitta, 268. 

bouddha, 248. 

brahma, 248. 

—— brunnea, 275. 


cerulescens, 284. 


catocyanea, 192. 
—— celsina, 224. 
christophi, 166, 168, 
contigua, 281. 

danna, 184. 

— dieckmanni, 166, 168. 
dravida, 278. 
dschalia, 185. 
eurotas, 251. 

flava, 254. 
flavomaculatus, 166, 167. 
florinda, 237. 
gemmata, 168. 


glauca, 238. 
goloides, var. akar, 254, 
herculea, 247. 


heterus, 254. 


homolea, var. palawea, 264, 
houangty, 165, 167. 


jansonis, 282. 
—— lamprospilus, 198. 
—— leonina, 290. 
luzonensis, 184, 


lycorias, 245. 


466 


Pamphila mesa, 254. 
meesoides, 184, 254. 
— maga, 186. 
mangala, 281. 
marnas, 256. 


masoni, 268, 


—— meleagrina, 234. 


mencia, 276. 
—— micio, 166, 168. 
mormo, 274, 
museca, 189. 


nervulata, 290. 

nitida, 254. 

niveomaculatus, 166, 167, 309. 
, var. christophi, 168. 


noctis, 214. 
ochracea, 248. 
orphitus, 256. 
palemon, 165, 166. 
palmarum, 252, 
pellucida, 282. 
— philenus, 256. 


—— prusias, 251. 
—— pulchra, 165, 167. 
purreed, 245, 


—_ pythias, 251. 
rama, 245, 
rickuchina, 248, 
sagara, 183. 


— scortea, 281. 

selas, 247. 

silutus, 165, 167. 

—- similis, 275. 

siva, 247. 

—— sodalis, 275. 

—— subhyalina, 247. 

, var. tibetana, 247. 
—— sunias, 255. 


—— sylvatica, 289. 
tamiata, 238. 
tavilis, 254, 
— trachala, 254. 
—— varia, 266. 


—— virgata, 186. 
—— vitrea, 198. 
-— vulso, 196. 
— zebra, 255. 
—— zetus, 206. 


INDEX OF 


SPECIES. 


Papilio acteon, 289. 


alcece, 152. 
althew, 152. 
alveus, 160. 
augias, 251. 
brontes, 166. 
carthami, 160. 
celenus, 299. 
chromus, 301. 
comma, 246, 287. 
exclamationis, 306. 
folus, 243. 
japetus, 137, 141, 
ladon, 306. 
lavatere, 152. 
—— linea, 288. 

— lineola, 289. 
—— malve, 153, 161. 
morpheus, 168. 


—— nostrodamus, 286. 
—— orbifer, 158. 
palemon, 165, 166. 
paniscus, 166, 
proto, 159, 

—— pygmeus, 286, 
—— sao, 159. 

—— side, 162. 

—— silvius, 167. 


—— steropes, 168. 

—— sylvanus, 246, 247. 
—— tages, 163, 

—— taras, 161. 

—— tessellum, 159. 
— thaumas, 288. 
thrax, 217. 


.— thyrsis, 218. 


venula, 288. 


virgula, 289. 
Paradoxurus philippinensis, 378, 388. 
Parata chromus, 301. 


gentiana, 304. 

—— malayana, 301. 

saida, 304. 

simplicissima, 299. 

Parnara assamensis, 216, 270, 273, 281. 
astigmata, 260. 

aurociliata, 272, 278, 310, 314. 
austeni, 269, 272, 279, 280, 314. 


INDEX OF 


Parnara bada, 281. 

— bevani, 270, 272, 273, 283, 314. 
bipunctata, 273, 283. 

—— bromus, 272, 277. 

—— brunnea, 271, 275. 

—— cerulescens, 270, 271, 284. 

cahira, 272, 278, 279, 280, 310, 314. 
canaraica, 277. 


cingala, 283. 

colaca, 272, 273, 283, 314. 
conjuncta, 273, 280, 315. 
contigua, 273, 281, 282, 314. 
—— discreta, 273, 282, 310, 314. 
distinctus, 286. 

— eltola, 273, 282, 314. 
flewilis, 284. 

—— guttatus, 270, 272, 281, 313. 
— hasoroides, 271, 284, 310. 
jansonis, 273, 282. 

kumara, 272, 276, 279, 280, 314. 
leechit, 271, 274, 309, 313. 
mathias, 270, 271, 275, 314. 
meiktila, 269. 


mencia, 276. 


—— miosticta, 212. 

— moolata, 272, 278, 279, 280, 314. 
narosa, 280. 

nascens, 271, 276. 

oceia, 233, 270, 271, 274, 318. 
— ornata, 268. 


pagana, 272, 277. 


parca, 234, 

pellucida, 273, 281, 282, 313. 

— philippina, 272, 276, 277, 309, 310, 314. 
—— philotas, 285. 


—- pholus, 216. 

plebeia, 271, 274. 

pugnans, 270, 272, 284. 
robusta, 272, 280, 310, 314. 
semamora, 269. 


—— seriata, 276. 

— simillima, 271, 274, 310, 313. 
sinensis, 271, 275, 276. 
subochracea, 271, 275, 310, 314. 
—— thyone, 283. 

toona, 281. 

—— tulsi, 270, 271, 284. 


uma, 285. 


SPECIES. 467 


Parnara unicolor, 285. 


watsonii, 269. 

zelleri, 273, 284. 

Parodon affinis, 34. 

Parthanosaurus zitteli, 10. 

Pedestes fuscicornis, 193, 194, 308, 312. 

maculicornis, 193, 308, 312. 

masuriensis, 192, 193, 312. 

pandita, 193, 312. 

Perea fluviatilis, 51, 60, 83. 

Phaneropleuron andersoni, 368. 

Philippines, on the Mammals obtained by Mr, John 
Whitehead during his recent Expedition to the, 
by Oldfield Thomas, 377-412. 

Philomedes agilis, 429, 434. 

flewilis, 429, 435, 448, 450. 

sculpta, 429, 434, 449. 

Phlaeomys cumingz, 400. 

, var. pallidus, 399. 

pallidus, 378, 399, 400. 

schadenbergi, 409. 

Phycis blennoides, 45. 

Pimelodus albicans, 27. 

bayard, 44. 

cottoides, 27. 


gracilis, 27. 


labrosus, 27. 


lateristriga, 27. 


maculatus, 27, 422. 
modestus, 422. 


pati, 27. 


—— pirinampus, 27. 


platanus, 27. 
Pipistrellus imbricatus, 385. 
Pirdana albicornis, 223, 224, 310. 
celsina, 223, 224. 
distanti, 223, 224, 

—— hyela, 223. 

pavonu, 224, 

rudolphii, 223, 224. 
Pisola zennara, 107. 
Pithauria aitchisoni, 237, 238. 
murdava, 237. 


stramineipennis, 237. 

Pithauriopsis aitchisont, 238. 

Plagioscion ternetzt, 25, 38. 

Plastingia aurantiaca, 225, 228, 309. 
callineura, 224, 225, 226, 227, 312. 


468 


Plastingia callineura, var. flavia, 227. 


corissa, 226, 228, 229, 230. 
cunetformis, 136. 


drancus, 229. 
—— flavescens, 224. 


— helena, 225, 227. 

—— hieroglyphica, 136. 

latoia, 225, 227, 229, 312. 
liburnia, 226, 228. 

-— margherita, 225, 227, 312. 
—— naga, 226, 229. 

noemi, 225, 226, 230, 312. 
—— plesioneure, 111. 

—— similis, 226, 230, 309, 312. 
submaculata, 230. 

— telesinus, 226, 228. 
tessellata, 225, 226, 229. 

, var. palawata, 229, 
vermiculata, 225, 227. 
viburnia, 226, 229. 
Platessa passer, 45. 

Platystoma juruense, 421, 428. 
orbignianum, 27. 


Plecostomus cochliodon, 30. 


emarginatus, 424, 
francisci, 30. 
ternetzi, 30, 38. 
vermicularis, 30. 


Plesioneura agni, 127. 
albifascia, 239, 240. 
— aliena, 231. 


— ambareesa, 115. 

—— alysos, 239. 

anthea, 215. 

— asmara, 118. 

— atilia, var. palawana, 126. 
aurivittata, 122, 

badia, 123. 

balukina, 117. 

basiflava, 241. 

— cameron, 121. 

chamunda, 117. 

clavata, 239. 

curvifascia, 239, 

dan, var. andamanica, 119. 
— dhanada, 119. 

—— dissimilis, 214, 215, 


fruhstorferi, 225, 227, 309, 312. 


INDEX OF 


SPECIES. 


Plesioneura flavocincta, 115. 
fusca, 117. 

goto, 118, 

grandis, 217. 

indrani, 126. 

lativitta, 121. 
leucocera, 116. 


leucocirca, 116. 


—— leucographa, 111, 


microthyrus, 242, 
mindorana, 242. 
monteithi, 241, 
munda, 116. 
nigricans, 118, 


palajava, 118. 
—— paralysos, 241, 


pinwilli, 145. 


praba, 197. 


pulomaya, 115. 


putra, 116. 

restricta, 239. 

ruficornis, 117. 

signata, 111. 

spilothyrus, 117. 

stellata, 243. 

sumitra, 115. 

tola, 111. 

volux, 239, 240. 

zawz, 111. 

Pleuronectes flesus, 51, 59, 60, 83. 

limanda, 51, 59, 60, 72, 83, 84. 
platessa, 51, 59, 83. 

Polyodon folium, 50, 55. 

Pristigaster cayanus, 427. 

Probubalus mindorensis, 410. 

Proteides chrysaglia, 296. 

surus, 231. 

zalates, 231. 

Protopterus, revision and comparison of the structure 
of the skull in, 356-363. 

amphibius, 13. 

annectens, 11, 18, 14, 18, 19, 20, 21, 51, 72, 
80, 345, 418. 

, external gills of, 18. 

, measurements of specimens of, 16. 


Pteropus auri-nuchalis, 383. 
jubatus, 383. 
vampyrus, 383. 


INDEX OF 


Pterygospidea angulata, 149. 
badia, 123. 

celebica, 134. 

davidii, 123. 

decoratus, 136. 
diversa, 132. 

gana, 141. 

helferi, 140. 

helias, 149. 
leptogramma, 150. 


maculosa, 115. 
— menaka, 142. 
—— moori, 135. 
nestus, 145. 


—— permena, 134. 


potiphera, 150. 


pralaya, 145. 


pteria, 142. 
—— pygela, 149. 


ransonnettii, 150. 

—— sinica, 135. 

—— syrichthus, 151. 

tabrica, 146. 

— trichoneura, 145. 

Pudicitia pholus, 216. 

Pyrgus agama, 151. } 


cashmirensis, 160. 
danna, 184. 
dravira, 152. 
evenidus, 157. 
—— gigas, 159. 


hypoleucos, 161. 
anachus, 19\). 
maculatus, 162. 
montanus, 164. 
nobilis, 159. 
sinicus, 162. 


superna, 157. 

tithys, 135. 

zebra, 157. 

—— (Syrichthus) proteus, 159. 

Pyrrhulina semifasciata, 34. 

Raja butis, 44, 50, 52, 82. 

clavata, 44, 50, 52, 67, 84. 

—— fullonica, 44. 

maculata, 50, 52, 54, 82. 

Reptile, on a Nothosaurian, from the Trias of Lom- 
bardy, by G. A. Boulenger, 1-10. 


VOL. X1V.—PART vill. No. 6.—December, 1898. 


SPECIES. 469 


Rhamphichthys blochit, 38, 425. 
brevirostris, 38. 


elegans, 428. 

Rhina squatina, 50, 52. 
Rhinodoras Ieneri, 29. 
Rhombus levis, 51, 59, 72, 83. 
nuchus, 45. 


Rhopalocampta benjamini, 306, 307. 

— crawfurdi, 306, 307. 

forestan, 306. 

renidens, 307. 

—— subcaudata, 307. 

translucida, 106, 107. 

Rhynchomys soricoides, 378, 391, 397, 398, 411, 
412. 

Rio Jurua, on a Collection of Fishes from the, by 
G. A. Boulenger, 421-428, 

Rio Paraguay, on a Collection of Fishes from the, by 
G. A. Boulenger, 25-39. 


| Salminus maaillosus, 37. 
Salmo farto, 44. 


salar, 50, 57, 83. 

trutta, 50, 58, 70. 
Sancus fuscula, 174. 

-—— pulligo, 173, 174. 
subfasciatus, 173, 174. 
Saprolegnia ferox, 18. 
Sarangesa albicilia, 124, 125. 
— dasahara, 124, 310. 
—— purendra, 123, 124, 310. 
sati, 128, 124, 310. 
Sarsiella hansent, 429, 438, 449. 
hispida, 429, 439, 450. 
Satarupa affinis, 131, 134. 

, var. cognata, 134. 


| —— bhagava, 131, 133, 134. 


celebica, 131, 134. 

corona, 132, 134. 

dire, 131, 133. 

diversa, 131, 132. 

dohertyi, 132, 134. 

— fumosa, 131, 133, 308. 
gopala, 130, 181, 132. 
narada, 131, 132. 

— nymphalis, 130, 131, 132. 
—— phasara, 130, 131, 133, 194. 
sambara, 130, 131, 132, 134. 
sinica, 130, 132, 135. 


3Q 


470 


Sutarupa tithys, 130, 131, 135. 
Saurus lacerta, 44. 
Scaumenacia curta, 368, 
Scelothrix speyeri, 160. 


onopordi, 161. 


zona, 162. 

—— (Pyrgus) albistriga, 162. 
Schilbe mystus, 44. 

Sciades pictus, 27. 

Sciena amazonica, 421. 

—— surinamensis, 25. 
Sciurus cagsi, 389. 
mindanensis, 389. 


—— philippinensis, 389, 
samarensis, 388, 389. 

steerei, 389. 

Scobura bipunctata, 205, 207, 308. 
—— bononia, 205, 206. 

cephala, 204, 205. 
cephaloides, 204, 206. 
concinna, 205, 206, 308. 
fenestrata, 205, 206, 309, 312. 
feralia, 205, 206. 

inarime, 205, 206, 312. 
—— martini, 204, 308. 

umbrosa, 205, 207, 308. 
Scomber scomber, 51, 61, 83. 


Scyllium canicula, 50, 52, 66, 82, 84. 


catulus, 50, 52, 82. 
Sebastonyma dolopia, 192. 

Sepa biseriata, 211, 213. 
cicatrosa, 211, 212, 308. 
— ciliata, 211, 214, 309. 
— cinnamomea, 211, 213, 309. 
cronus, 210, 211. 

—— quttulifera, 211, 212, 308. 
miosticta, 211, 212. 

noctis, 211, 214. 
Serrasalmo humeralis, 37. 


piraya, 426. 


serrulatus, 37. 


spilopleura, 37. 

Silurus glanis, 44. 
Simosaurus pusillus, 10. 
Solea fischeri. 421. 

—-— jenynsii, 26. 

vulgaris, 51, 59, 60, 83. 


INDEX OF SPECIES. 


Soleotalpa unicolor, 421. 
Sorumbim lima, 26, 421. 
Squalus acanthias, 44. 
—— glaucus, 44. 
Steatogenys elegans, 428. 
Stegophilus nemurus, 426. 
Sternarchus albifrons, 37. 


macrolepis, 427. 
natterert, 427. 


oxyrhynchus, 427. 
— tamandua, 427, 428. 
Sternopygus carapus, 38. 
virescens, 38, 428. 
Steropes argyrostigma, 167. 


unicolor, 169. 


Streptoleberis crenulata, 435. 

Sturgeon, structure of the suprarenal bodies of the, 
69. 

Suada albinus, 177, 178. 

— — cataleucos, 177, 178. 

scopas, 177, 178. 

swerga, 177. 

Suastus aditus, 179, 180. 

albinus, 178. 

bipunctus, 178, 180, 308. 

chilon, 179. 

gremius, 178, 179. 

—— migreus, 178, 179. 

molleri, 177. 

—— phiditia, 180. 

robsonii, 180. 

sala, 179, 308. 

subgrisea, 179. 

telesinus, 228. 

tripura, 179, 180. 

Suprarenal Bodies in Fishes, contributions to the 


Comparative Anatomy and Histology of the, by 
Swale Vincent, 41-84. 
Sus celebensis philippinensis, 411. 
Symbranchus marmoratus, 38, 45. 
Syrichthus ali, 158. 
andromeda, 161. 
—— antonia, 163. 
biett, 162. 
delavayi, 162. 
leuzew, 159. 
maculatus, 162. 


Syrichthus maculatus, var. thibetanus, 162. 


mohammed, 159. 
oherthiiri, 162. 
sinicus, 162. 


— staudingeri, 159. 

Teniura dumerilit, 38. 

Tagiades alica, 138, 140. 
atticus, 189, 142, 143, 311. 
, var. calligana, 143. 


—— bhagava, var. andamanica, 133. 


brasidas, 141. 
—— calligana, 143. 


corona, 134, 

dealbata, 139, 145. 
—— distans, 141. 

elegans, 138, 141. 

—— fuscula, 174. 

gana, 138, 141. 

—— graya, 133. 

helferi, 138, 140. 
japetus, 137, 138, 141. 
—— —., var. latreillei, 142. 
karea, 138, 141. 

—— khasiana, 138, 140. 
latreillei, 142. 

levata, 138, 142. 

— litigiosa, 143. 

—— martinus, 139, 143. 
meetana, 140. 

menaka, 139, 142, 143. 
nana, 139, 144, 145, 309: 
—— nestus, 139, 145, 309. 
—— niphates, 134. 

noctis, 140. 

—— nymphalis, 132. 
obscurus, 138, 141. 
—— pinwilli, 140, 145. 
pralaya, 140, 143, 145. 
princeps, 139, 145. 
pteria, 139, 142. 
pulligo, 174. 

ravi, 138, 140. 


tabrica, 140, 146. 

titus, 189, 142, 309. 

— toba, 144, 145. 

trichoneura, 140, 148, 145. 


sambavana, 139, 143, 309, 311.. 


INDEX OF SPECIES. 


| Lagiades trichoneuroides, 140, 145. 


waterstradti, 139, 143, 309. 
Tanyptera celsina, 224. 


Tapena agni, 127. 

hampsoni, 146, 147, 308, 311. 
—— wna, 128. 

— laxmi, 126, 128. 

minuscula, 146, 147, 308, 311. 
—— thwaitesi, 146, 147, 311. 
Taractrocera archias, 183, 185. 


| -—— ardonia, 183, 184. 


ceramas, 183, 184, 310. 
danna, 183, 184. 

—— flavoides, 183, 184. 
—— lyde, 187. 


—— mevius, 182, 183. 


nicevillet, 183, 184, 310. 
oberthiiri, 183, 185. 
trimaculata, 187. 

ziclea, 183, 184, 185. 
Tarsius fuscus, 381. 
phalippensis, 379, 381. 


sangirensis, 331. 

Telegonus acroleucus, 217. 

lara, 217. 

Telicota augiades, 250, 253, 313. 
augias, 249, 240, 251, 313. 
bambuse, 250, 251, 313. 
brahma, 248. 

concinna, 250, 253, 310, 313. 
—— dara, 184, 249, 251, 254, 313. 


| —_ dilutior, 251, 255, 310. 


gola, 250, 253, 313. 
insularis, 250, 252, 253, 309. 


| ___. kwehni, 250, 252, 253. 


meesa, 249. 


—— mara, 186. 
marnas, 256. 


—— nigrolimbata, 185. 
orphilus, 251, 256. 


| —— palmarum, 250, 252, 253, 313. 
| —— paragola, 250, 254. 


—— philenus, 256. 
—— prusias, 250, 251, 253, 308. 


simplex, 250, 253, 308. 
siva, 247. 


rectifasciata, 251, 254, 310, 313. 


471 


472 INDEX OF 


Tetragonopterus abramis, 35. 


agassizit, 35. 


argenteus, 35. 


dichrourus, 39. 
lacustris, 35. 


lineatus, 35. 
maculatus, 426. 
multiradiatus, 35, 426. 
- orbicularis, 426. 
ternetzi, 35, 39. 


ulreyi, 35, 39. 
‘etrodon nigropunctatus, 50, 56. 


psittacus, 428. 

Thanaos cervantes, 163, 164. 
erebus, 164. 

indistincta, 191. 

jhora, 190. 

-— kali, 192. 

leechii, 163, 164, 311. 
marloyi, 163, 164, 311. 
montanus, 163, 164, 311. 
—— ——.,, var. nigrescens, 164. 


nigrescens, 164. 
—— pelias, 163, 164, 311. 
popoviana, 163. 


rusticanus, 164. 

sinina, 164. 

stigmata, 187, 189. 

tages, 104, 163, 164. 

Thomas, Oldfield, on the Mammals obtained by Mr. 
John Whitehead during his recent Expedition 
to the Philippines, 377-412. 

Thymale feisthamelii, 238. 

Thymelicus leonina, 290. 


nigrolimbatus, 185. 
pellucida, 282. 

ziclea, 184. 

—— (Hesperia) stigma, 289. 


Tracheluopterus coriaceus, 28. 


SPECIES. 


Trachyleberis, gen, nov., 444. 

scabrocuneata, 429, 4380, 444, 451. 

Trichomycterus brasihensis, 34. 

Trigla lyra, 51, 61, 83. 

—— pini, 51, 61, 83. 

Udaspes folus, 243. 

stellata, 243. 

Onkana attina, 234, 235, 

batara, 284. 

elia, 236. 

mabillei, 235. 

Uranoscopus scaber, 44. 

Vandellia cirrhosa, 426. 

Vincent, Swale. 
Anatomy and Histology of the Suprarenal Cap- 


Contributions to the Comparative 


sules.—The Suprarenal Bodies in Fishes, and 
their Relation to the so-called Head-Kidney, 
41-84. 

Viverra tangalunga, 388. 

Watsonia swinhoer, 220, 309. 

Whitehead, John, on the Mammals obtained by, 
during his recent Expedition to the Philippines, 
by Oldfield Thomas, 377-412. 

Xantharpyia amplexicaudata, 383, 384. 

Xeromys silaceus, 391. 

Xestoleberis compressa, 430, 442, 451. 

— luaata, 430, 441, 452. 

olivacea, 430, 442, 450, 

Xiphorhamphus ferox, 37. 

Zea martini, 216. 

Zeugopterus sp., 51, 59. 

Zeus faber, 51, 61, 83. 

Zographetus auriferus, 201, 203, 308. 

durga, 201. 

flavalum, 201, 202. 

SJlavipennis, 202. 

ogygia, 200, 201, 203. 

ogygioides, 201, 203, 308. 

satwa, 200, 201, 202. 


END OF VOLUME XIV. 


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CONTENTS. 


XI. On new or imperfectly-known Species of Ostracoda, chiefly from New Zealand. 
By G. Srewarvson Bravy, WD., LL.D., D.Se., PRS. (Plates XLIII- 


POCTEV ALS) Stes ve Nan Pig pR ae Pana Weecdtag 1 arp ae eae 
List of the Papers contained in Vol. XIV. . . 1. 2. «1 uss se. 468 
Tndex: of Species, Se. in Vol: RIV ..5 0 Pa Re Fe ene ae, oe 


Titlepage and Contents to Vol. XIV. 


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