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THE

TRANSACTIONS

OF

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES—VOLUME I.
ZOOLOGY.

EON DORN:

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:

SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXIXx.

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CONTENTS.

PART I.—SEptemser, 1875.

I. Onthe Morphology of the Skull in the Woodpeckers (Picidee) and Wrynecks (Yungide).
By Prof. W. K. Parker, “B.S. (Communicated by H. T. Stainton, Lsq., Sec.
MIE Te SOG!) (Elecegyk. Vin)et Yen at ee yn EOE BY Space ay

IT. On some Atlantic Crustacea from the ‘ Challenger’ Expedition. By Dr. R. v.
Witiemoes-Sunm, Naturalist to the Expedition. (Communicated by Professor
Wayyiin DHOMSON,, 27-12... G8.) . (Plates WRX TDD) ee 28

III. On the Structure and Systematic Position of Stephanoscyphus mirabilis, the Type of
a new Order of Hydrozoa. By Professor Atuman, ID., LL.D., F.RS., §c.,
NEGCS ETAT SOGNe a (lab eNO cate A get eS eee oo Wo) we og op Wl wa MO!

PART IJ.—Drcremper, 1875.

IV. On the external Anatomy of Tanais vittatus, occurring with Limnoria and Chelura
‘terebrans im excavated Pier-wood. By Joun Denis Macponatp, JLD., F.B.S.,
Staff-Surgeon R.N., Assistant Professor of Naval Hygiene, Netley Medical School.
(Communicated by W. T. Tatseuton Dyur, IA., F.L.S.) (Plate XV.). . 67

V. On Valencinia Armandi, a new Nemertean. By W.C. M‘Intosu, ILD., F. RSF,
iS CPAATONON Taliare ret ie nen (ah tert te Met Me cn Me a ae ar at mete eee he

L ty J
PART Mii =juny, 1876:

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. Kelaart,
F.L.S., at Trincomalee, and partly by Dr. Collingwood, F.L.S., in the Eastern
Seas. By Dr. Cutupert Cottinewoop. (Plates XVII.—XIX.) . . page 83

VII. On the Structure and Development of the Bird’s Skull. By W. K. ParKxsr,
HBS Holes 166) (bart Us) ((Rlates XGXeSXeXevail)| 2). 7 et

PART [V.—Janvary, 1877.

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups. By
Prof. J. O. Wxstwoop, U/.4., F.L.S., fe. (Plates XXVITI-XXXIII.). . 155

IX. On a new Genus of Trematoda, and some new or little-known Parasitic Hirudinei.
By Joann Denis Macponaup, ID., F.R.S., Deputy Inspector-General R.N.,
Professor of Naval Hygiene Army Medical School. (Communicated by G. EB.
DOBSON, Al MB: EES.) 1( Plate eXeXox Vi) see

X. On the Osteology and Pterylosis of the Spoon-billed Sandpiper (Kurynorhynchus
pygmeeus, Linn.). By Joun AnDERSON, ID., PRS, LS. ZS. S§¢.,
Curator of Indian Museum and Professor of Comparative Anatomy at Medical
College, Calcutta. (Plate XXXV. and 2 Woodeuts.). ..... =. . 213

PART V.—Ocroper, 1877.

XI. On Steere’s Sponge, a new Genus of the Hexactinellid Group of the Spongida. By
James Moris, ILD., LS, .GS., ae (Plates XXXVI. & XXXVII. and 2
WVOOd CUES Ait Meier aeon hae ce. ear wie eh tee Sy oo nae eae en

XII. Notes upon the Oxystomatous Crustacea. By Epwarv J. Mizrs, £.L.S., #.Z.S8.,
Assistant in the Zoological Department, British Museum. (Plates XXXVIII—
2.4) eee ae Ce rn Cee rein MC NG ee Vg ge BRS

XIII. On the Domestic Pig of Prehistoric times in Britain, and on the mutual Relations
of this Variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andamanensis, and
Sus barbatus. By GrorGEe Roxiirston, I.D., F.R.S., F.L.S., Linacre Professor
of Anatomy ond Physiology, Oxford. (Plates XLIXLIII.) . .. . . 251

beaer J

XIV. On two new Forms of Deep-sea Ascidians, obtained during the Voyage of H.ILS.
‘Challenger. By H. N. Moseuey, .A., F.RS., Fellow of Exeter College,
Oxford. (Communicated by Dr. J. Muniz, F.Z.8.) (Plate XLIV.) . page 287

XV. On new Forms of Actiniaria dredged in the Deep Sea; with a Description of certain
Pelagic Surface-swimming Species. By H. N. Mosetey, W_4., F.R.S., Fellow of
Exeter College, Oxford, late Naturalist on board H.U.S. ‘ Challenger. (Com-
municated by Dr. J. Murie, F.L.S.) (Plate XLV.) . . . . . . . . 295

_ PART VI.—Novemser, 1877.

XVI. On the Birds collected by Professor J. B. Steere in the Philippine Archipelago.
By R. Bowpier Suarer, /.L.S., 2.Z.S., e., Senior Assistant Zoological Depart-
ment, British Museum. (Plates XLVI-LIV.‘and 2 Woodcuts.) . . . . 3807

PART VII.—Avetst, 1878.

XVII. On the Male Genital Armature in the European Rhopalocera. By ¥. BUCHANAN
Wisin I De hei Ss, so) (Plates UV-Vis a se DT

XVIII. Worphology of the Mammalian Ossicula auditis. By ALBAN H. G. Doran,
F.R.C.S. (Communicated by Professor Firowrr, F.R.S., FDS. &c. (Plates
AV AINTATESOXCIE ES Jarek Se Sameer tala he es PR oem Tia bee eat See eit

XIX. On the Annelida obtained during the cruise of H.U.S. ‘Valorous’ to Davis Strait
im 1875. By W. Carmicuart M‘Intosu, ID., LL.D., P.R.SS. L. & #., FLS.,
Sex (Rlateniaxevy and: 2 Woodeuts:)i 28 5 Yeah e eae a 2 ws & 499

XX. On the Sacral Plexus and Sacral Vertebre of Lizards and other Vertebrata. By
St. GzorcE Mrvart, Sec. Linn. Soc., and the Rev. Ropert Cxuarxe, F.L.S.
(Ristes avy, &UxeVE andi 9: Woodeuts.)) 2 5 9 20. 2 a «o/s 518

PART VIII.—Junz, 1879.

XXI. The Butterflies of Malacca. By ARTHUR GARDINER ButiER, /.LS., F.Z.S8.,
Senior Assistant in the Zoological Department, British Museum. (Plates LXVIIT.
£5, Tike TIN) SONS) AE CoC ch Me ee an ord ta eae ON a ee a A Re ee RE

brevet

XXII. On certain Organs of the Cidaride. By Cuares Stewart, Hsq., F.L.S.,
U.R.C.S., Lecturer on Comparative Anatomy, St. Thomas’s Hospital. (Plate
TX.) Retest) OR Til! she) oo SOE ee Menge Sena ao

XXIII. On Hypsiprimnodon, Ramsay, a Genus indicative of a distinct Family (Pleopo-
didze), in the Diprotodont Section of the Marsupialia. By Professor Owry, C.B.,
F.R.S., F.L.S., &c., Superintendent of the Natural-History Departments in the
British Museum. (Plates LXXI. & LXXII. with 8 Woodcuts.) , . . . 578

XXIV. Descriptions of some minute Hymenopterous Insects. By Prof. J. O. WEstwoop,
MAR Hh itaS.. ces, \(Rlate XX.) oe egeeeiediae os): (cee

Divers ee ei eee ke RS ies gE a ele

ERRATA.

Page 59, bottom line, for loupe read pocket-lens
», 240, line 15 from top, for WV. plicata read NV. sinuata
» 066, ,, 5 from top, for Albisara read Abisara
» 068, ,, 18 from bottom, for Tagisdes read Tagiades

THE
TRANSACTIONS

OF

THE LINNEAN SOCIETY

OF

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART. THE FIRST.

LON DION:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE ;

AND BY LONGMANS, GREEN, READER, AND DYER, PATHERNOSTER=-ROW.

M.DCCC.LXXV.

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CONTENTS.

PART I.—1875.

I. On the Morphology of the Skull in the Woodpeckers (Picide) and Wrynecks
(Yungide). By W. K. Parker, Hsg., B.S. (Communicated by H. 'T. Statyton,
BE SG MISCO USS e AT NT a Pha adc ckeeaunes one) ess) a Neal wah Oe pace!

II. On some Atlantic Crustacea from the ‘ Challenger’ Expedition. By Dr. R. v.
Witiemors-Sunm, Naturalist to the Expedition. (Communicated by Prof. WYvILLE
PET ORESONE Die Hrs scale Se) ead iin oe Mi eA a Rl Un Oe Gee aeb

Ill. On the Structure and Systematic Position of Stephanoscyphus mirabilis, the Type
of anew Order of Hydrozoa. By Prof. Attman, U.D., LL.D., F.RS., &e., Pres.
RET ES OOM tiheaate Iie oye dined careoniet sb you ist heet pexh OW Le deck Paeces Ol

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Wa aye CaN ASA OO 4 nt Le ORE ea con \ oh
La

TRANSACTIONS

OF

THE LINNEAN SOCIETY.

I. On the Morphology of the Skull in the Woodpeckers (Picidee) and Wrynecks
(Yungide). By W. K. Parxsr, Esq., F.R.S. (Communicated by H. T. Statnton,
Esq., Sec. L.S8.)

(Plates I.-V.) ,
Read April 2nd, 1874.

INTRODUCTION.

MopERN zoologists, when at fault in their taxonomy, not knowing where to place
some perplexing specimen or type, begin to look to the working morphologist for
counsel and help in their difficulties.

These workers are merely separated for convenience’ sake: a thorough zoologist finds
no time for embryology ; an embryologist is most grateful to the skilled and accom-
plished classifier who arranges the various members of each faunal group ready to
his hand.

Hach kind of labourer has the greatest need of the results brought out by the other:
the patient dissector waits for the treasures supplied to him by the more mercurial
taxonomist ; whilst he, in turn, profits by the work of one to whom a single type may
serve for the labour of a year or more. Yet both are learning to look beneath the
surface of things, a growing knowledge of the types showing both that close kinship is
often marked by great difference in outward form, and that it is easy to be beguiled by
the external likeness of forms—isomorphic, indeed, but far apart zoologically. Never-
theless, on the whole, the keen eye of the zoological expert seldom errs in the grouping
of forms, even by their outward characters alone: but there are types that will baffle
all that skill; and then other counsel has to be called in.

Embryology, however, does not exist as a handmaid to zoology; its aims are higher
by far than that; and as for the zoologists proper, they exist for the morphologists, and
not the morphologists for them. ‘The familiar term “ zygodactyie” for birds with a
certain form of foot, has been very useful; and yet how much ignorance it may be made
to hide! It seems to be something when one knows that a certain bird belongs to that
- group; and yet a Cuckoo, a Parrot, and a Woodpecker come none the nearer each other
zoologically by the possession of that kind of foot.

SECOND SERIES.—ZOOLOGY, VOL. I. B

2 MR. W. K. PARKER ON THE MORPHOLOGY OF

The Woodpeckers, like the Parrots, form a well-defined group—too well defined for
the morphologist, who is pained for the paleontological losses the bird-fauna has sus-
tained, judging from analogy that the mixed forms are lost which would have yoked
the most divergent groups together. In the late geological period we have merely
the twigs of a few branches of the great genealogical bird-tree; the main branches and
the trunk are buried in the strata. Yet not the less does the morphologist seek in the
early stages of the young and embryo bird for the lineaments of the old feathered
fathers of the existent types. Hidden from the eye of the mere taxonomist, they yet
may be revealed to him, in some degree, by embryology; but birds cannot be inter-
preted morphologically, irrelatively to other vertebrata; judged merely by themselves,
and compared among themselves only, they present enigmas at every turn.

We all know now how peculiar is that type of ornithic face which is found in Passe-
rine birds, and is called by Professor Huxley the “Aigithognathous”’ palate. Nitzsch de-
scribed it; and I have long been familiar with its more marked peculiarities; but its
morphological importance dawned upon me when I saw that the parts of that complex
face, so conjugated and so metamorphosed, were really built up of elements which had
their true counterparts or “symmorphs”’ in the snake. But the snake does but repeat
these parts from the Amphibia; and the Amphibia borrow them from the cartilaginous
Fishes, amongst the lowest of which, namely the Lamprey, may be found the fullest
development, both morphologically and functionally, of cartilages that form the sub-
stratum of the most peculiar part of a Sparrow’s face.

Considered ornithologically, the ‘“ Picide”’ are full of interest; but looked at from a
higher stand-point they are almost of greater importance to the morphologist than the
Passerinz, having for the most part the same curious repetition of parts borrowed from
below, but also retaining more evident marks of their ascent from the quasi-pupal and
quasi-larval forms of the groups beneath.

The Woodpeckers have been designated “‘Celeomorphe” by Professor Huxley (Proce.
Zool. Soc. April 1867); but although the “ Picidee”’ and “ Yungide”’ are thus named as
to their general form, yet that author has not given a term for their peculiar facial
structure. With clearest insight, however, they are spoken of as “ not Desmognathous,
the palate of these birds exhibiting rather a degradation and simplification of the
Agithognathous structure” (p. 468). The fact is they are like early embryos of the
Passerine, in their palatal region arrested at a most simple and Lacertian stage,
whilst in other respects they are metamorphosed and specialized beyond any other
kind of birds. As far as their upper face is concerned, their arrested ‘“ maxillo-
palatines,”’ symmetrical “ vomers,” ‘“ septo-maxillaries,” and feebly developed turbinal
scrolls, entitle them to a name which shall be a memorial of their Lacertian facial
morphology. I therefore propose to call them the ‘‘ Saurognathe.”

On the Structure and Development of the Face in the “ Picidee (Celeomorphe,” H..,
*“ Saurognathe,” P.).
The necessity for a morphological study of birds becomes manifest when it is re-
membered that Coracias was for a very long time classed with the “ Corvide,” and that,

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 3

until the appearance of Professor Huxley’s paper, the best ornithologists classed birds of
the Cuckoo-form, together with the Woodpeckers and Wrynecks, as “ Picarie.’ So
much as the author of that paper made out of the palate of the Pici has been of great
value to me. My own study of the structure of these birds, both of hard and soft parts,
dates back to the early part of the year 1843; the unpublished results of that labour, in
the form of minutely careful drawings, are still of considerable value for reference.
More recently, before my friend’s researches were made, I had from time to time taken
up my preparations of the British species for the sake of comparing them with the skulls
of other ornithic types, and for their own elucidation. This was never fully satisfactory ;
and even with the help of Professor Huxley’s work, until very lately they continued to
be enigmatic.

In addition to my older material, including some nestling Wrynecks in spirits, the
opportunity to dissect three young green Woodpeckers, and to inspect and figure the
skulls of three southern species *, has resulted in the figures and descriptions which
I now offer. In the first species I shall describe the whole face, but in the others merely
the preeoral structures. This is in consonance with other researches of the same kind in
Avian families.

Morphologically considered, the palate is composed of just so much of the vertebrate
face as exists in front of the oral (double) cleft, namely :—the trabecule cranii, and the
parts developed around them in front and above; and below and behind those foremost
structures, so much of the mandibular arch as develops quasi-independently in the
maxillo-palatine lobe the pedate process of the mandibular arch.

These proper palatine structures form a secondary arch in front of the mouth, distinct
enough in the higher types; but in cartilaginous fishes, amphibia, &c. it shows its true
nature, in the former as a pedate spur of cartilage, growing forwards from the primary
apex of the visceral rod, and in the latter either as a fore-growing spur, as in the
** Urodela,” or as a conjugational band tying together the trabecula and the mandible, as
in the “ Anura.” ;

Thus in the large series of types which I have determined to compare together, it has
seemed fit to me to take a very small territory; yet that territory contains parts that
have undergone the greatest amount of metamorphosis of any in the whole body of a
high and noble vertebrate, and moreover being, in the bird, the skeletal framework of the
whole upper face, these parts are, as it were, an index of the amount of speciali-
zation undergone by any particular type—the ruling determining structures that lead to
all, and really demand all, the changes that take place in the rest of the organism. This
is especially explained for the benefit of those who will accuse me, and have already
accused Professor Huxley, of taking a narrow view of the Bird-types—touching with
the point of a needle some little tract, but unacquainted with and not able to appre-
ciate the Bird as a whole. This we calmly but indignantly deny; and the allegation
is the more ungrateful in that we do not come as those who would have dominion
over the faith of the mere zoologists, but as helpers of their work, letting in light from
a new chink.

* My helpers in this matter are my kind friends Messrs. Murie, Bartlett, Dines, and Salvin.
B2

4 MR. W. K. PARKER ON THE MORPHOLOGY OF

On the Structure and Development of the Skull in Picus (Gecinus) viridis.

The extraordinary complexity of the palatine structures in the Green Woodpecker can
only be understood by reference to the unfused elements in the fledgling (Plate I, fig. 2,
for the palate as a whole, and Plate I. figs. 3-8, and Plate IT. figs. 5-7, showing details
and sections). In the palatal view of the young bird the nasal structures are purposely
left out for the better display of the facial bones—that part of the lateral ethmoid which
belongs to the 1st preoral, the ‘“‘ os uncinatum,’ being coloured, as it is an essentially
upper palatine element.

Beginning at the apical region of the trabeculz, we see rudimentary basipterygoids
set on the side of the basifacial region, where it begins to form the rounded “ rostrum”
(Plate I. fig. 2, d.pg). This parasphenoidal under-balk sharpens in front, and ends
behind the free fore edge of the mesoethmoid. The premaxillaries have together a high
triangular outline, and the outer bar (dentary portion) ends behind by lying for some dis-
tance on the dentary region of the maxillary ; so that whilst many birds are isomorphic
of the typical “‘ Teleostei’’ in having an internally placed “os mystaceum,” this bird
exposes as large a maaillary margin as the salmon. ‘The palatine processes of the pree-
maxillaries equal the solid part in length, and in breadth they equal the outer or
dentary bar. Their position also is somewhat analogous; for they pass on the iside of
the “ preepalatine,” whereas in many Passerines they pass on the outside. This reversed
situation of processes is seen again in certain delicate Australian Passerines, namely
Acanthorhynchus and Ptilotis. 'The nasal processes of the preemaxillaries are broad,
flat, and fibrous, and they show a median suture late in life. The “os uncinatum” (0. w)
is part of the “ ectoethmoid,’ and becomes as large as its distinct symmorph in those
Parrots which have an imperfect suborbital bar.

The symmetrical ‘“vomers” are not at first easy to find at this stage, being very
small; but by removing the palatine they can be seen as small styles, attached to the tips
of the long mesopterygoid processes (Plate III. figs 5 & 6, v, m.pg). The symmetrical
“ septo-maxillaries ” (s.mx) are also to be found, by careful search, on the inside of the
palatine bars; one was seen on the right, and two on the left side. The pterygoids (pg)
are of moderate length, and they have a long styloid “ mesopterygoid process ”’ ( Plate ITT.
figs, 5 & 6, pg, m.pg); on their upper surface the pterygoids have a free spur, pointing in
the same direction as the mesopterygoid: this is very long in the old bird (Plate III.
fig..7, pg); it is peculiar to the “ Picide.” The “ epipterygoid”’ hook is aborted; and
the mesopterygoid not being differentiated, there is then an evident arrest of metamor-
phosis in this proximal pier segment of the pterygopalatine arcade. The prepalatine
bar is normally long and slender; it is closely wedged in between the maxillary style
and the retral palatine process of the preemaxillary. At the middle this narrow part
ceases ; the broad posterior half is gently suleate along its middle; and the inner portion
sends forward a styloid interpalatine spur (Plate I. fig. 2, pa, i.pa). The “ interpalatine
edges”’ of the two bones are almost parallel from the spur to the end of the base. The
broad part begins to narrow at its outer edge midway ; and forming an obtuse angle there,
the transpalatine is aborted. The upper edges meet at the mid line; and thus the nasal

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 5

canal is three fourths closed. The commissure above is peculiar: each bone has a car-
tilaginous selvedge; these meet together, coalesce, and in front endostosis takes place,
so that a dagger-shaped wedge of bone divides the sides of this arch. This is the “ medio-
palatine” (Plate I. fig. 2, and Plate IIT. figs.5 & 6, m.pa). The “ ethmo-palatine”
region is premorse, having no long spur as in most birds. The secondary parts of the
palatine arch equal in interest those formed directly in the primary bar: these are the
maxillary, palato-maxillary, and jugal. The maxillary (Plate I. fig. 2, and Plate III:
fig. 6, mx) is most developed in this species; but the maxillo-palatine plate is merely a
small suboval outgrowth, joined to the main bar by a wide isthmus; it is convex above
and concave below.

This process is far apart from its fellow, and grows out of the narrow maxillary bar
nearer to the posterior than the anterior end; both of these regions are long narrow
styles; and the bone grows outwards most where it gives off the inner plate or “ maxillo-
palatine.” The essential nature of the maxillary is well seen here as a long bony
bar applied to the outside of the second preeoral arch—long, because of the very pro-
gnathous face of the bird; in this case its palatine portion is scarcely more developed
than in the Teleostean fish. The posterior splint, the jugal, is of the normal size, and, as
in many of the arboreal birds, has no separate quadrato-jugal attached to it. Besides the
« maxillo-palatine ” process, there is, as in certain of the “‘ Coracomorphe,” Cardinals,
Buntings, &c., a “ palato-maxillary ” bone. This, however, gives a peculiar asymmetry
to these birds; for it only appears on the /ef¢ side (Plate I. fig. 2, p.mx): in the young
bird it is a small wedge attached to the inner side of the main maxillary, some distance
in front of the “ maxillo-palatine process.” Thus we see that the single mammalian
maxillary may be represented by ¢hree bones in the bird: the “ postmaxillary,” which
I first observed in the young Emu, is about as common as the distinct palato-maxillary ;
the “ septo-maxillary’’ belongs to the first preeoral arch *.

The newly fledged Green Woodpecker presents an excellent subject for determining
the morphological meaning of the parts composing the nasal labyrinth. These have
always to be worked out with the preoral arches, with which they are even more inti-
mately blended, by metamorphosis, than the postoral with the ear-sac. A vertical section
of the head, made a little to the left of the axial cartilage, is very instructive (Plate I.
fig. 3). This section takes in the whole length of the trabecular arch from its apex,
now involved in the basisphenoidal mass (6.s), to its azygous extremity, the shrunken
preenasal (p.m). The “ parasphenoid”’ has its “rostrum”” still distinct ; but the broad
hinder portion has been ingrafted on the apices of the trabecule and the squared end
of the “investing mass,” to form that composite cranio-facial bone, the “ basisphenoid.”’
The bony matter has reached to the ‘“ optic commissure” and the sides of the deeply
cupped “ sella turcica.”” A line drawn obliquely through the middle of the circular
optic foramen (2) and the oval interorbital space (é.0. s) would exactly divide the facial
from the cranial elements, so soon, by metamorphosis, wrought into a common mags,

* In the “ Fowl” paper {note at bottom of p. 773) I have spoken of the “ septo-maxillary” as “distinct in no
bird.” Only within the last few years have I found the true centre; and it is very commonly present, not, however,
in the subject of that paper, the Fowl.

6 MR. W. K. PARKER ON THE MORPHOLOGY OF

and only partly redivided by this same interorbital window. Here turns up the ex-
planation as to why the anterior sphenoid (p.s) should be perched up high above the
ethmoidal and basisphenoidal base-line. This part, with its arrested “ orbito-sphenoidal ”
lips, is at present unossified ; it has, afterwards at least, three centres—one for the body,
and a large pair for the wings; and it ends above in the peaked extremity of the frontal
region of the mesoethmoid (m.eth). This latter region is very large; but its lower half
is borrowed territory, belonging to the subjacent portion of the long trabecular com-
missure with its ascending crest; four fifths of this composite plate is already ossified,
about to become the largest mesoethmoid (relatively) in the vertebrate series: I speak
of that of Birds generally. In the adult the double nasal sac is separated for the most
part by the merest film of membrane (Plate IT. fig. 5, c. fm) : this fenestrate space has com-
menced as a “‘ notch” by the absorption of part of the trabecular tract (Plate I. fig. 3
c.f.n), both base and crest. This is the beginning of the notch which at once divides
the first facial arch into a proximal and a distal portion, and also separates the middle
wall of the nasal labyrinth into a vestibular and a true sensory region. The septum nasi
{s.n) is a thinnish continuous plate of cartilage, terminated by the stunted preenasal
above, and by the thick projecting trabecular cornua below (fig. 3, p.n, c.tv). An outer
side view of the same part of the face (fig. 4) shows that the great median ethmoidal
ossification (“ pars perpendicularis”’) is growing outwards into the alee, or upper part of
the “ectoethmoid,’ whilst the lower part (“‘ pars plana’’) has its own ossification
(p-e; p-p)-

The angle below the centre for the pars plana is the substratum that should form the
“os uncinatum ;”’ the mass in front of this part, and below the superior turbinal (w. <b),
is the nasal gland (m.g); the “ aliseptal”’ (a/.s), or root of the inferior turbinal, is seen
above; in front of this the “ alinasal”’ fold (al.2) ; below this the nasal floor (x. f), and
altogether in front the pedate trabecular cornu (c. ¢7). All these parts are brought into
daylight by transverse sections (Plate I. figs. 5-8). In the foremost of these (fig. 5) the
recurrent trabecular horns (c. ¢) are shown as two thick lobular masses of cartilage, with
a binding isthmus formed between the ends of the “ cornua.” This is very similar to
what is seen in an ordinary Mammal, e.g. the Pig. Hedging in the tuberous mass we
see the nasal processes of the preemaxillaries (nv. px), the dentary part (d. px), the fore
shaft of the maxillary (mx), the palatine spur of the preemaxillary (p. px), and outside
this the preepalatine bar (p.pa). In the next section (fig. 6) the same bony sections are
shown, with the exception of the palatine portion of the premaxillary. The “ nasal
nerves’ (7.7%) come into view here below the fore part of the septum (s. ), and at the top
this wall has grown into a pair of roof-plates (aliseptals), which develop into a large
thick half-scroll, the “ alinasal turbinal” (al. t+). Outside these valvular growths
there is an infero-lateral development of cartilage, which at first is a wall and then a floor
(n. w,.f). In the succeeding section (fig. 7) new parts come into view: the two arms
of the nasal bone (m); the interpalatine spur (i.pa); and the fore part of the “inferior
turbinal ” (7. ¢b). The section of this arrested scroll is hammer-shaped, an outgrowth
being developed inside as well as outside the root; the outer coil is the rudiment of the
scroll, which in Rhea and Tinamus is coiled upon itself three times (“Ostrich skull,”

THE SKULL IN THE WOODPECKERS AND WRYNECKS. i

' plate x. figs. 14 & 15, and plate xv. fig. 7. ¢b); in the Fowl (“ Fowl’s skull,” plate Ixxxvi.
figs. 4—6, 7. tb) it has two coils. In the same figure (7) the alinasal turbinal (a/. ¢0) is
seen to form one nearly perfect coil. It is somewhat more like the inferior turbinal of
ordinary birds. In these things the Woodpecker is but little more specialized than a
Lizard or Sea-Turtle.

Behind, the inferior turbinal (fig. 8) is continuous above with the aliethmoid (a/.e),

which infolds itself, is pinched in and crenate, and becomes a rudimentary superior tur-
binal. This will be better seen in the thoroughly ossified parts of the adult (Plate V.
fig. 2, w. tb): where the inferior turbinal ends, running into the front of the antorbital
plate, there a rudimentary “middle turbinal”’ is seen (ibid. m. ¢b). In the young bird
this section (Plate I. fig. 8) shows the termination of the nasal floor (n.f), curving
upwards to the partially ossified “ perpendicular plate” (p.e) ; the nasal nerves (n. 7),
the palatine (pa), the nasals, frontals, and nasal processes of the preemaxillaries (n, f, n.p2)
are also cut through, and a good view is obtained of the extent and relations of the
“‘ maxillo-palatine processes” (mwx.p). The condition of the nasal floor in this section
is preparatory to a remarkable metamorphosis, to be described hereafter. The adult skull
and face (Plate II.) can now be interpreted, notwithstanding the mixture there is in it
of arrest and of an unusual degree of metamorphosis.
_ The palate (figs. 1 & 2) is a remarkable piece of basketwork; and as in this species
ossification is intense, the nasal labyrinth brings in its many bony parts to add to the
complexity. The preoral facial bones are thin and splintery, but are, notwithstanding,
very strong. The strongest part is the solid fore end of the preemaxillary; yet it is
quite hollow, forming scarcely any diploé except short side props at the junction of the
lateral with the lower plate. The head is the hammer, and the bony beak the helve, to
this curious combination of axe and chisel, the dense horny sheath forming an apt “suc-
cedaneum ” for steel. The bindings on and interlacements of the facial plates and bars
have their meaning in this, that they carry off the jar from the brain of this plumy artisan.
Taking the parts in detail, we find that the “ basipterygoid processes” (b.pg) are arrested,
and yet the pterygoids cleave close to the converging cranio-facial base.

The cleanly cut groove formed by the pterygo-palatine bars glides smoothly under the
rounded “ parasphenoidal rostrum,” (7. 6.8); this is a synovial joint : the rostrum ends in
a free style behind the oblique fore edge of the ethmoid, which spreads into two small
wings (figs. 1, 2, 3, 7.0.s, p.e). The “notch” from being small and kidney-shaped
(Plate I. fig. 3, ¢.f. 7), has grown into a very large nearly closed fenestra, which runs
forward to near the end of the septum. Thus the septum nasi (Plate IT. fig. 3, s.z) is
reduced to a sharp bony keel running down a short way below the double sac; and the
trabecular remnant, with its stunted przenasal beak (p.2), has an equally low crest.
One third of the trabecular bar is absorbed entirely behind; and scarcely half the
remainder is ossified by its centre; the rest is a continued growth of bony matter from
the “ septum” and the “ trabecular cornua” (fig. 1, s.”,¢.¢r). The “notch” forming
the cranio-facial hinge does not run through; it is arrested earlier than in the Gal-
linaceous birds (“ Fowl’s Skull,” plate Ixxxvi. fig. 14); and being anchylosed to the
ethmoid, the face sits more stiffly on the head than in most birds. The palatine process of

8 MR. W. K. PARKER ON THE MORPHOLOGY OF

the preemaxillary (figs. 1, 2,3, p.pa) comes, besides, under the fore part of the septum with
its recurrent wings; it has anchylosed with the inner edge of the long lathy pre-
palatine bars (p.pa). There is also a slender ‘“‘ median palatine process” to the pre-
maxillaries (m.p. px); in the specimens before me it is distinct from the nasal septum ;
but in another of my preparations, now in the College of Surgeons, the two parts are
anchylosed. The dentary region of the premaxillary overlaps, and is completely fused
with the maxillary; the nasal processes (fig. 5, 2.px) are well anchylosed to their nasal
and frontal surroundings, with a small part of their own median suture evident.

It is in the intermediate splints, however, rather than in the parasphenoid and pre-
maxillaries, that the peculiarity of the Woodpecker’s palate is most remarkable; these
are the “ vomers”’ and “ septo-maxillaries.”

It is evident that Professor Huxley had not appreciated the true nature of the latter
bones in the Reptilian skull in his ‘Elements ;’ for both in the Lizard and the Snake
(fig. 90, p. 225) they are described as ossifications in the internasal cartilage. Such
ossification I have never seen; and the bones figured are what I at first called in the
Reptile “ praevomers,”’ and now call “ septo-maxillaries ” *.

Seen from below (Plate IT. figs. 1 & 2, v.s.mz), the vomerine bones are attached loosely,
behind, to the premorse “ethmo-palatine”’ (e.pa), but more strongly to the inner
edge of the “ maxillo-palatine” plate (ma.p) and of the alinasal and inferior turbinals
(al.tb, 7. tb). Their position is vertical, their convex face within towards the septum
nasi, and their concave face towards the turbinal scrolls.

The inner view (fig. 3) shows them edgewise; but they are also shown as dissected
away, with the inner edge of the “ maxillo-palatine ” in figs. 4 & 5: here the piece of
the maxillary is thrown back to display the other pieces. The palatine bones and
maxillo-palatine plates lie directly beneath the vomerine series ; in figs. 2 & 6 the palatine
is cut away to display them better. The main septo-maxillary (s.ma%) is a conchoidal
bone, equal in size to the broadest, inturned part of the maxillary just adjacent. It has
a strong keel on its septal or convex surface ; and its hinder margin is broadly transverse,
ragged at its angles, and has its edge thickened on the inner surface, like the edge of a
bivalve shell. The lower edge of the bone is attached to the maxillo-palatine; and its
narrowed anterior dentate margin carries a ragged wedge of bone, the “ anterior septo-
maxillary” (s.ma’). Between the dorsal keel and the uncinate posterior angle there is a
rough wedge of bone one fourth its size: this is the main “vomer”’ (v’); it is sharp in
front and notched behind. In the ligament which attaches the main vomerine piece to
stunted “‘ ethmo-palatine” edge (e.pa), there is another, much smaller, irregularly oblong
bony centre. ‘his is the posterior vomer (v”). The large conchoidal “ septo-maxillary ”
is a compound bone, as I shall soon show. These four pairs of bones do not, however,
exhaust the vomerine series; for the tract of ligamentous tissue hedging in the great
*‘notch’’ below, from the mesoethmoid to the septum nasi, has in it two “ median septo-

* My first term is perfectly correct if the facial arches are looked upon as a series of rib-like hoops growing down-
ward from the head. These bones appear to have been confused with the Mammalian “ cornets inférieurs ” by most
authors. Professor Owen (‘ Report on Archetype,’ 1846, p. 293) speaks of them in Reptiles as “‘ The small and in-
constant ossifications in the capsule of the organ of smell.”

THE SKULL IN THE WOODPECKERS AND: WRYNECKS. 9

maxillaries,” small ovate-oblong patches of bone (figs. 1, 2, 5, m. s.mx). A single bone in
this region is not uncommon in Birds, as I have already shown. These apparently
azygous bones will be explained by a younger specimen.

In this species (the Green Woodpecker) the main septo-maxillary piece rivals its
symmorph in many of the Lizard kind; and when all the details of this instructive but
perplexing skull are mastered, I am confident that the term “saurognathous” for this
kind of palate will not be thought inappropriate.

The nasal labyrinth is so intimately connected with the trabecular arch that its
description may well come in here. I have already shown its structure in the July bird,
and also the relation of the median part to the first arch.

In this species the turbinals and al nasi are endosteally ossified; the alee project
strongly in front, and in infolding themselves form a remarkable boss on each side,
which is strongly attached to the fore part of the septum nasi by continuous ossification
(figs. 1, 2, 5, al.n, c.tr). These bosses are the trabecular cornua. The “alinasal
turbinal,” coiled once upon itself (Plate V. figs. 6 & 7, al. tb), is of enormous size; the
“inferior turbinal,”’ equally simple (Plate V. figs. 7 & 8, 2. ¢), is one fourth less. In a
front view of the antorbitals and perpendicular ethmoid (Plate V. fig. 2, p. e) we see the
“rostrum ” of the “ parasphenoid,” the fore end of the median ethmoid, flat-faced below,
and growing into the aliethmoid above the elegant ear-shaped “superior turbinals”’
(wu. tb), the rudiments of the “middle” (m. tb), and scars where the inferior turbinals
(«. tb) have been broken away. The parts are intensely anchylosed ; but we can discover
the passages for the first and nasal branch of the fifth nerve (1, 5), the great size of the
“ ectoethmoid,”’ mounting to the top above, and sending backward a large “ uncinate”
process below. The frontal bones (/), strongly roofing in the forehead, are thrown into
a solid ridge by the tips of the ‘ cornua majora” (4r. 1), which turn to the right side,
and lie in an obliquely directed smooth groove.

The second, or pterygo-palatine arch (figs. 1, 2, 4) has a short posterior segment, or
pier, the pterygoid (pg). This bone is broad, inwardly bent, and bifurcate (Plate
III. fig. 7, pg); for it sends upwards and forwards a process, or muscular attachment,
behind the true mesopterygoid process (m.pg): the latter does not become segmented
off; and hence this bespeaks less metamorphosis than what obtains in a Passerine bird.
There is no “epipterygoid’’ process. The long, flat, elastic palatines (pa), are anchy-
losed to the dentary and palatine portions of the preemaxillary, but not to any part of the
maxilla or their own splints. The “transpalatine” angle is aborted and obtuse; the
inferior surface of the broad part of the bone is gently sulcate along its middle. This
broad part is of great extent, and ends in the slender “interpalatine” spine (¢.pa),
which looks towards the free retral end of the palatine plate of the premaxillary, but does
not in this instance help it to enclose an oval space; it may do so in very old birds.

The postpalatine region is narrow, the bones attenuating suddenly ; and the “ ethmo-
palatine” lamine (e.pa) are of very small extent, and have no spine, or only a rudiment
for attachment to the vomerine ligament ; the plate is fenestrate on the right side. The
commissure is finished by the endosteal mediopalatine (fig. 2, m.pa), a lozenge-shaped
bone, dagger-like behind. The palatine arch is seen to be fenced on its outer side by a

SECOND SERIES.—ZOOLUGY, VOL. I. Cc

10 MR. W. K. PARKER ON THE MORPHOLOGY OF

bony lath, which runs from the body of the preemaxillary to the cup in the outer face of
the quadrate (fig. 4, q). The main part of this outer bar is the maxillary; the terminal
style is the jugal (see also Plate I. fig. 2, mx, j). The ear-shaped “maxillo-palatine”’ is
scooped above, where the ‘‘inturned nasal wall” (Plate I. fig. 8) and alinasal turbinal
(al. tb) rest upon it. Opposite the palato-pramaxillary hook we see a smallish, oval,
bony plate, on the left side of the palate—the palato-maxillary (Plate II. figs. 1, 2,
p.m).

In this species I cannot pass by the postoral arches without some notice, notwith-
standing the greater interest of the preorals. The structure and development of the
mandible (Plate V. fig. 1) corresponds very exactly with that of the Crow-form; the
internal angular process is strong and recurved, and the posterior process short and
broad. The five pairs of splints, and one pair of endosteal bones (‘‘ articulo-Meckelian ”’),
form but one V-shaped bone in the adult, with but little trace of sutures, and with no
fenestra. The ossified “symphysis” is shortish; and that end of the bone is formed on
the model of the upper beak.

The stapedial apex of the largely aborted second postoral arch has some peculiarities of
importance. The ¢rue stapedial, or periotic portion (Plate I. fig. 11), is rather large and
roughly oval, the side towards the “‘ opisthotic” bar separating the fenestra ovalis from
the fenestra rotunda being straightest. .

The capitular portion of the arch, continuously ossified with the base, is the flattish
*““mediostapedial’’ (m.s¢); a bony rod from this bar runs down the anterior “ infra-
stapedial”” bar (z.s¢), bringing to mind the small bone in the “‘stapedius” muscle of
the mammal; it is its symmorph. 'This double ‘“infrastapedial” is new to me; it ends
below in a spatulate stylohyal (st.h), which has a proximal ossicle just below the infra-
stapedial fenestra. The extrastapedial (e.s¢) is faleate and broad-backed ; the “‘supra-
stapedial”’ (s.st) is small; and from it and from the proximal end of the extrastapedial
a fibrous fan arises, which supplements the small “tubercular” head of this facial rod.
The proximal portion of the large “ stapedial fenestra” (f) is hidden in this view ; but it
scoops the faleate ‘‘ extrastapedial”’ beneath its thick outer or back part.

The rest of the arch, save its merest extremity, is membranous; and the terminal
rudiments (“‘cornua minora”’) have coalesced. These (Plate I. figs. 9,10, c.h) form,
even in my youngest specimen, a spearhead-shaped piece, notched deeply above, and
less below; for there the coalescence of the two ossified rods is greatest. The upper sur-
face is flat, the lower subcarinate; the whole piece is covered with a barbed horny sheath
(Plate V. fig. 1, ¢.h). The rest of the hyoidean skeleton is a most exquisitely elastic
structure of thread-like bones, ossified, at first, endosteally (Plate I. figs. 10, 11, 6.07).

The basihyal (Plate V. fig. 1, 6.67) is 13 inch in length, the proximal part of the next arch
(1st branchial) (67. 1) 1 inch 2 lines, and the distal portion is about 43 inches in length;
each ramus of the 3rd postoral is therefore about 6 inches long. There isno “ urohyal.”’
In this the Woodpeckers agree with the Gannet, Baleniceps, Rhea, and some other
“‘ Microglosse.”

Besides the smooth, shallowish, double groove on the upper part of the head, there are
to be noticed the feather-pits with which this skull is dinted. The whole skull is very

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 1a

strong, and the orbits well marked; the interorbital space is small; the lachrymal is a
little bone attached to the nasal. The exoccipital and basitemporal together, with the
help of about three tympanics, that soon coalesce, form the most curious tympanum seen
in the class; it is like a cowrie shell: the Duck comes nearest to the Pici in this
respect. Even with materials at hand, I cannot go further into the skull, as the fore
Jace is at present my proper field.

Since the above description of the skull of the fledglings and the adult Green Wood-
pecker were written, Samuel Whitbread, Esq., F.R.S., has sent* me a first-autwmmn spe-
cimen of this kind; and this stage happily bridges over the great distance between the
fledgling and the old bird. Its description may well come in after that of the adult
skull; for it serves as a most perfect key to the mysteriously complex basketwork of the
old bird’s palate, and also sheds a clear light upon the embryo-passerine characters seen in
these birds.

The progress of anchylosis through the five additional months has been intense, and
many new parts have appeared. This is in conformity with what was observed in the
Fowl, many of the metamorphic changes taking place during the first year, although so
much had been gone through in the egg. |

The anterior trabecular splints, the preemaxillaries (Plate IT. fig. 6, px), have coalesced
with the palatines and maxillaries; and the appearance of free ends to their palatine
processes, well seen in Picus minor (Plate IV. fig. 1, p.sp.x), does not appear to be the
ease here, the retral snags being due in the Green Woodpecker to additional ossicles.
All the three membrane bones that lie near the septum, on each side, belong. to the
vomerine series (vomers and “ septo-maxillaries”’); and those which had appeared on the
fledgling (Plate I. fig. 2, v, s.mx) have now attained their fullest free development.
Yet I cannot find the “posterior vomer” of the old bird here (Plate IT. figs. 2-5, x”).
The series of “ mediotrabecular splints” forms a long chain, reaching from the anterior
end of the przenasal style (p.m) to the “ ethmo-palatine laminz.”

‘Beginning at the hinder extremity of the series, we have the true symmorphs of the
vomerine crura of the Passerine bird (the “shoulders” of the compound bone I shall
describe presently); and only on the right side do I see any attempt at cutting off the
“posterior vomer”’ (Plate IT. figs. 6 & 8 v”).

These “ free vomerine crura”’ are long, thin, half-coiled bones, convex outside, concave
within, thick below, sharp and serrate above. They do not reach their proper fastening-
point (the ethmo-palatines) behind, having become dislocated by growth (see Plate IIT.
figs. 5 & 6, v, for their primordial “ Rhynchosaurian ” position) f.

Attached to the fore end of the vomer by fibrous tissue, is a somewhat slenderer
needle-shaped bone (Plate IT. figs. 6 & 8, sma’). This is the hindermost of the “ septo-
maxillary chain ;’”’ it undergirds the “inturned alinasal lamine ” (¢. a. 1), to which it is
strongly tied by fibrous tissue. A similar bone, but still slenderer, is attached to the

* This bird was sent from Southill (Biggleswade) on the 1st of November, 1872; it was a young of the first
autumn, and therefore half a year old.
+ Compare those figures of the youngest Green Woodpecker’s vomer with Dr. Giinther’s figures and descriptions
of Hatteria (Phil. Trans. 1867, part ii. plate i. fig. 2, p. 5).
c 2

12 MR. W. K. PARKER ON THE MORPHOLOGY OF

front of this (s.ma*); this reaches to, and is bound under, the trabecular cornu (c. ¢r)
The retral growths of the alinasal folds have also two short irregularly oval patches of
fibrous bone, the 3rd and 2nd “septo-maxillaries”’ (s.ma’*, s.ma’); and these have
running in front of them the first of the septo-maxillary series (s.ma'); it is a frail,
slightly inbent style, running as far forward as the preenasal (fig. 3, pz). The ossicles
at the mid line have a character which was not suspected in the examination of the
adult; and they were not discernible in the fledglings; the section of these (Plate L.
figs. 7 & 8) showed no submesial cartilage inside the inturned alinasal lamina (¢. a/, 1.w).
Here, however late and abortively developed, we have the “labial cartilages” ossified,
though independently of the other vomerine bones (vomers and “ septo-maxillaries ”),
and very unsymmetrical in shape. The /eft bone (J. c’) is hindermost; it is tuberculate
behind, broad and grooved below in the middle, and styloid in front; this styloid part is
strongly bound beneath the right piece (J. c'). This latter is a sessile leaf of bone, on a
long fore-and-aft style, the foremost style being tied beneath the true septal bone or
trabecular ossicle (s.”), In the embryo of the Linnet (to be described in another paper)
are to be seen the labial cartilages of the Passerine bird, which so soon coalesce, and then
send backward exogenous “crura;’’ here, in the Woodpecker, they are abortively developed,
come nearer and nearer to the mid line (see in the old bird, Plate IT. figs. 1, 2, 3, s. 2, s.mza),
and appear like mere patches of fibrous bone. Almost all the elements of the complex
highly metamorphosed Passerine vomer keep apart in the Woodpecker, but not al/; the
5th septo-maxillary coalesces with the ossified inturned lamina (Plate II. figs. 2 & 5,
s.mx,%. al); and that part of the nasal capsule, contrary to all experience, detaches
itself from the rest, and becomes the scooped part of the complex 5th septo-maxillary of
the old bird. In the Passerines this part of the nasal capsule coalesces with, and often
becomes largely ossified by, the contiguous vomerine bones. In the old bird (Plate IT.
figs. 1-5) the Ist, 2nd, & 3rd septo-maxillaries coalesce with the contiguous parts of
the preemaxillary and infero-mesial parts of the nasal capsule. In the subadult bird
(Plate IL.) the huge, simple “alinasal turbinals” (a/. 70) are ossifying below; the “in-
ferior turbinal” (7. ¢b) is almost entirely soft. The inferior part of the labyrinth, in
front, has the very constant endosteal patch of bone; it has already ossified the long
“preenasal”’ style (which afterwards becomes blunt, Plate II. fig. 1, p.m); laterally
(fig. 7) it is affecting the “ trabecular cornua” (ce. ¢7); and behind it runs along the fore -
half of the septum below ; the rest of the septum below, or rather its trabecular addition,
has the usual infero-posterior osseous centre (¢7, s. 2), which is attached to the right ossified
labial cartilage. Binding the “inturned lamina,” below is the sessile, leafy ‘‘ maxillo-pala-
tine” plate; and undergirding this is the contiguous part of the palatine (pa), whose
interpalatine process (ipa), far forwards in position, ties itself to the outer and under
surface of the 4th and 5th septo-maxillaries. The rest of the parts of this most re-
markably complex facial basketwork have been described already; and the reader will
now be able to follow me whilst I give the details of other species of Woodpeckers.

The first of these exhibits the “ vomerine cartilages,” but unossified, and is only to be
understood by references to the condition of these parts in various Passerine birds. These
references will be given in due time and place.

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 13

On the Palatal Structures of Hemilophus fulvus.

This Celebesian species, from Mr. Salvin’s collection, is about the size of our native
Gecinus viridis; it presents some modifications of great interest. In many respects the
description of the last species would serve for this; but the practised eye soon detects a
number of modifications. The rudimentary basipterygoids (Plate III. fig. 1, b.pg) are
sharper and more divergent; the part answering to the “os uncinatum” (0.2) has less
suborbital development, but is more distinct in outline. The anterior border of the
“«meso-ethmoid ” is somewhat sharper than in G'ecinus; the nasal septum and the rest of
the labyrinth are ossified to nearly the same degree. The three palatal bands of the pre-
maxillary (fig. 1 pa) are all fused together by bony deposit; and this excess of ossifica-
tion has filled in the angle between the side of the palatine and the end of the pre-
maxillary palatine plate. Opposite the sharp interpalatine spurs (¢.pa) there is a
mass of cartilage, not seen, in this form at least, in any dissection of mine, in the other
“ Picide.” This is evidently two flat plates that have coalesced at the mid line (fig. 2,
l. €), and then, becoming carinate below, send forwards a blunt, and backwards a sharp
spur. The septo-maxillary bones are strongly attached to, but are not grafted upon, this
cartilage. Here we have the vomerine cartilages of the Wren, developed only on their
front portion, not metamorphosed into part of the vomerine bar, but distinct as in the
Rhea. As to the morphological meaning of this double cartilage, I am fairly satisfied
that it answers to a pair of upper labials. In front of the shoulders of this cartilage
there is, on each side. a falciform septo-maxillary; it reaches to where the premax-
illary style has coalesced with the inner edge of the palatine. Behind the broad end of
the right-hand style there are four bony grains; on the /eft side there is one larger and
one smaller grain. All these are strongly tied to the interpalatine style, to the labial
cartilage, and to the rounded fore end of each true “vomer.” These latter bones (v)
reach from the septo-maxillary grains to the short snags of the “ ethmo-palatine” (e.pa).
They are bowed inwards, as in the embryo of the Passerines, have a thick lower and a
crested upper edge; they are free in front, but anchylosed behind.

Between these, behind, I find one “median septo-maxillary’’ (m. s.mx) attached to
the base of the free edge of the ethmoid. It is probable that the larger posterior septo-
maxillary on the left side was once three grains of bones, and that each vomer ossified
from-two points, as in Gecinus. Anyhow this is as curious a piece of facial morphology
as any to be seen in the whole range of the vertebrata, and makes the observer very
cautious of predicting what he shall or shall not find. The palatine arch is, on the
whole, so closely like that of the green species, that it requires no fresh description; the
small “ palato-maxillary ” (p.mz) on the /ef¢ side is very similar to that of Gecinus.

On the Palatal Structures of Picus analis.

Mr. Salvin’s rich Neotropical collection has supplied me with a Javan Woodpecker
intermediate in size between our native P. minor and P. major. In Picus analis the
fore edge of the “‘basitemporal”’ plate (Plate ITI. fig. 3, 0.¢) is elegantly trilobate, the
outer enlargements being partly due to the presence of rudimentary basipterygoids.

14 MR. W. K. PARKER ON THE MORPHOLOGY OF

The “rostrum” of the “ parasphenoid”’ (pa.s) is rather short, and glides in the pterygo-
palatine groove. Its pointed end comes short of the anterior sharp edge of the ‘‘ meso-
ethmoid.”

The nasal septum (s. 7), ‘‘ anterior and inferior turbinals” (al.td, 7. tb) are well ossified,
as in Hemilophus ; these are only faintly indicated in the figure, to give prominence to
the facial arches.

The median palatine part of the preemaxillary (px) lies above the lateral palatine
plates (p.pz), so as to form a rounded sulcus; the latter have their retral tips anchy-
losed to the vomerine bar. The dentary regions of the preemaxillary (d. px) run con-
tinuously into the maxillary bar (mz).

The palatine process of the preemaxillary is connected with the stunted feeble ethmo-
palatine plate (e.pa) by a delicate, sinuous, rugose band of bone, which gradually broadens
backwards (figs. 3 & 4, p.px, s.mx,v). Here, whatever number of ossicles existed in the
growing bird, in the adult are all undistinguishable; the only distinction between septo-
maxillary and vomer is in certain irregularities in this frail bony bar. Even here a
“median septo-maxillary”’ (m. s.mx) bounds the lower outline of the “hinge:” it is
fixed at the septal end.

The pterygoids (pq), each with its large muscle-snag and scythe-shaped “‘ mesoptery-
goid”’ process, are quite normal, as also are the palatines, which, however, have deve-
loped a gnawed transpalatine rudiment (¢.pa). The “interpalatine” spurs have the
same irregular eroded appearance; the ‘“‘mediopalatine” lozenge (m.pa) is imbedded
in more cartilage than in Hemilophus.

The “maxillo-palatine” processes (ma.p) form merely a small subtriangular lip to the
inner edge of the main maxillary bar (mx), and look broader than they are, because of the
outbowing of the bone on the dentary edge, where the angle of the mouth is. Small as
the ingrowth is, yet it is scooped below, to form an air-passage to the body of the bone.
All trace of suture in the jugal bar is gone, as in the other species. Here we come
athwart the most developed specimen of the odd “palato-maxillary ” bone (p.mz) ; it is
a wedge-shaped plate, strongly attached to the overlying “alinasal wall,” with its
enclosed turbinal. The “os uncinatum”’ (fig. 3, 0. uw) has a semidistinct appearance,
being well marked out at the angle of the large well-ossified “pars plana.” The la-
crymal is, as in Gecinus and Hemilophus, a smallish semioval plate, which becomes
anchylosed to the hinder edge of the descending crus of the nasal, as in Thrushes. tp

On the Palate of Picus major.

The description of P. analis may serve also for this for the most part; yet this is a
stouter kind, and the palatal bars are straighter and stronger (Plate I. fig. 1). The
specimen from which the drawing was taken, the gift of Dr. Murie, was rather young,
and it had lost the “ palato-maxillary” and the “mediopalatine” by maceration ; its
less-developed condition gives a softer state of the ossifying nasal labyrinth. Yet
already the vomerine series had become fused together and to the preemaxillary and
palatine bones (s.ma, v, p.pa, e.pa). This species has a free “interpalatine” spur,

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 15

and a more decided “transpalatine” angle; and the “os uncinatum’’ is much more
distinct than in P. analis; their “ maxillo-palatine” lobes are about equal. i

The main body of the palatine is more deeply grooved than in Gecinus; in this it
agrees with P. analis; the “ethmo-palatine” hook shows some distinctness from the
end of the vomer. The absence of a “median septo-maxillary” is probably due to
maceration.

On the Palate of Picus minor vel striolatus.

Some years ago Mr. Bartlett put into my hands two ripe nestlings of this small
species; these were immediately dissected, and have been greatly valued because of the
distinctness of their sutures.

A palatal view of one of these skulls (Plate IV. fig. 1) well serves to illustrate the
structure in Picus proper (Plate IV. figs. 1 & 2). In front of the very wide “ basi-
temporal” plate, with its median beak, we see a pair of very arrested “ basipterygoids”’
(d.pg) on the wide-rooted, short, sharp-pointed “parasphenoid” (pas) which underlies
the massive “‘meso-ethmoid.” In front of the “hinge” the nasal septum (s. 2) is still
cartilaginous to a great extent, although a bony tract from the trabecular cornu has run
backwards beneath it. The fore part of the nasal labyrinth is still cartilaginous ; but the
“pars plana” behind has coalesced with the “meso-ethmoid.” The premaxillaries are
shorter here, and their palatine processes (p. px) curve inwards at their end, thus free-
ing themselves from the inner edge of the palatine. Immediately above the styloid
“interpalatine” processes*(7.pa) there are seen three “septo-maxillaries” (figs. 1 & 2,
s.mx), one on the right, larger, and two on the left, lesser ossicles. Their clavate ends
are attached to the clubbed fore end of the subarcuate vomers, which are suddenly fili-
form, and then expand into a twisted crest, where they join behind to the broad,
eroded, inturned “ethmo-palatines” (e.pa,v). The palatines are very broad and thin,
and have an obtuse “transpalatine” angle (¢.pa). Their upper plates are uniting by
synchondrosis ; and a small “ medio-palatine ” centre has appeared on the left side, in front
(m.pa). The terminal part of the palatine is sharp and notched, and is overlain by
the large falciform “‘ mesopterygoid process.

The pterygoids (pg) are more slender than usual; and the muscle-snag above is short
at present. The “os uncinatum” (0. w) is represented as yet by the unossified condition
of the “ecto-ethmoidal” angle. A small cuneiform “ palato-maxillary” (p.pr) has
appeared in the angle between the palatine and maxillary ; this latter bone sends inwards
a small subtriangular “maxillo-palatine” lip (mx.p), scarcely larger than, and not so
distinct as, the smallest process of the kind seen in the Bird Class, namely, in Wenura
superba. I may mention the large “orbito-sphenoids” seen on each side in these
specimens, as they indicate a more general character of the bird than we have in the
Crow form, in the higher types of which, at least, these do not appear.

The lacrymal is, at present, a very small, short thread of bone behind the nasal.

The jugals (7) are quite distinct from the zygomatic process of the maxillary.

16 MR. W. K. PARKER ON THE MORPHOLOGY OF

On the Palate of Picumnus minutus.

This Woodpecker, from Bahia, Brazil, is about the size of our native Golden-crested
Wren; it leads on to the lesser group, or the Wrynecks, having soft tail-quills. In
several respects it is of extreme interest, especially as relating to the Passerines of its
own geographical area, namely those low forms of the Western Notogea the “ Cotin-
gide” and “ Formicariide.” J have had to work out the parts of this bird’s palate
from the fractured skull in a dry skin, kindly given me by Osbert Salvin, Esq. Labour of
this kind was never more richly rewarded.

The nasal septum (Plate V. figs. 3 & 6, s.) is semiosseous; the rest of the labyrinth
is principally persistent cartilage. The upper anterior and posterior regions of the
septum are all occupied with one continuous bony tract, which also runs into the ale
of the lower and trabecular region.

Between and behind these wings there are three centres of bone, the last of them
being the familiar postero-inferior bone of the septum, a subterminal trabecular centre.
The cranio-facial “ notch ” is not perfect, and the ossified septum and meso-ethmoid (p.e)
are articulated together.

A distinct ‘“ septo-maxillary ’”’ (s.mx) could be seen on the left side, and in front of it
a thickening of the same nature. The right bone had coalesced with the inner edge of
the palatine, that part which receives increment from the palatal process of the pre-
maxillary.

The vomers (figs. 4, 5, v) retain their embryonic (“ Rhynchosaurian’’) condition, as in
the young of Gecinus viridis (Plate ITI. figs. 5, 6, v), being arcuate “horns” attached
to the mesopterygoid processes of the pterygoid (pg). These latter processes are normal,
but shorter than in our native species; and the rest of the bone is similar to that of
Picus minor, but stouter.

The palatines, as seen from below (fig. 3, pa, pt.pa), struck me at once as being sin-
cular, the postpalatine flanges not being bevelled off, as in the other kinds, but largely
developed as in the ‘“‘ Formicariide”’ and “ Cotingide”’ amongst the Passerines of South
America*, and notably also in the Australian Lyre-bird (lenura superba). In these birds,
however, Thamnophilus, Pachyramphus, &c. these side walls of the posterior narial tunnel
are continuous with the rest of the palatal ossification; but in Picwmnus (Plate V. fig. 5)
they are distinct laminee of bone, each with a down-growing ala.

These “postpalatine” bones end in front behind the “interpalatine’’ spurs, and
below the “ ethmo-palatine” laminee, which here, as in the Green Woodpecker, are feebly
developed. ‘The interpalatine spurs are well developed, and are more normally placed,
further backwards, than in the “ Picide” generally. The rest of the palatine is quite
like that of its congeners. The maxillaries (mx) have a very rudimentary maxillo-
palatine process (ma.p); and the separate “ palato-maxillary ” (p.mzx) on the eft side is
represented by an enlargement of the inner edge of the bone, which was once, most pro-
bably, quite separate.

* T cannot give illustrations of these Passerine structures here ; they are freely given in the first part of my paper
on the “ Aigithognathe,” about to appear in the ‘ Zoological Transactions.’

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 17

_ The jugals (7) are slender, save at the quadrate end, where they are very solid and
inhooked.
Altogether this small far-western type is extremely instructive, and helps to lead the
imagination down to extinct types in which the characters of the Hemipod, the Low
Passerine, and the Woodpecker were existent in one generalized form—a form and a type
only a step or two above the raft-breasted Ostrich tribe,

On the Development of the Face in the Subfamily Yungide: example
Yunx torquilla.

We now come to the feeblest members of the “ Celeomorphe ;” yet the Wrynecks
are stout little birds, with the same restless energetic character as the larger types.

The possession of several nestlings of this form, the gift of Mr. Dines, has enabled me
to work at a still earlier condition than that of the youngest specimen of Gecinus
viridis.

The palate (Plate IV. fig. 3) shows that these birds are true Woodpeckers, and have
no special affinity to the Crow and its allies.

Here we have the same form of occipital condyle (0. ¢), the same massive broad “ basi-
temporal”’ plate (4.2), the rudimentary “ basipterygoids”’ (b.pg), and, what is very instruc-
tive, an explanation of the very remarkable cowrie-shaped tympanum seen in those types.

The “ basitemporal”’ forms a large retral hook where it joins the “exoccipital” (e.0),
and is then roundly scooped at its edge. Here the lobular growths of the adult are seen
to be due to the presence, of a ‘‘ tympanic chain;” three of these bones are now to be
counted on each side (Plate IT. fig. 1, as compared with Plate IV. fig. 3, ty).

In Yunex the “ parasphenoidal rostrum” (pa.s) is much longer than in Pécus, and the
face much shorter, especially the preemaxillary (px). This is most remarkable when
compared with Gecinus and Hemilophus: something is due, however, to age; for these
specimens are very young.

The solid part of the premaxillary is very short indeed; but the three main bars
are long and extremely slender (d.px, p.px); they have the same relations as in Picus.
The nasal septum, with its thick trabecular base, is still unossified, and indeed scarcely
hardens at all in this type, nor do the turbinals. Here the preenasal cartilage (fig. 3, p.2)
is still large.

There are no septo-maxillaries at this stage; nor am I aware of their existence in the
adult*. The “ vomers,” however, are very noticeable. At first I expected to find them
partly preformed in hyaline cartilage, as in the ‘‘ Coracomorphe,”’ especially as Hemi-
lophus retains in its adult state its large or prevomerine labial plate. It is not so; they
arise, as in Gecinus viridis, in a band of soft connective web on each side of a remarkable
recess in the mucous membrane of the nose, the azygous counterpart of those paired.
recesses called in Man the “ sphenoidal sinuses” (Plate IV. fig. 11, v, m.sph).

Below the vomers is the palatal mucous membrane (Plate IV. figs. 10, 11, v). Seen
from below (fig. 4, v) they are remarkably like those of a young Passerine, being pointed

es The adult skull may be seen in the macerated condition in the College of Surgeons ; it was prepared by me with

the other bones, and drawn thirty-two years ago.
SECOND SERIES.— ZOOLOGY, VOL. I. D

18 MR. W. K. PARKER ON THE MORPHOLOGY OF

behind, broad and obliquely premorse in front. Outside they are overlapped by the long
ethmo-palatal style (e.pa), whilst the filiform very elongated mesopterygoid processes
(m.pg) nearly reach them behind. Here the “ parasphenoidal rostrum” (pa.s) ends,
emarginate and toothed, whilst in front the vomerine pieces are tied to the nasal wall by
a band of membrane continuous with that which forms a commissure to the two bones.
In front of the vomers this membrane thickens at the mid line, being scooped submesially
for nervesand vessels. This thick alate tract of fibrous tissue, however, is the symmorph
of the cartilaginous mass in Hemilophus (Plate III. fig. 2, 7. c); and the fibrous bands in
which the vomers are developed are the degraded counterparts of the free labial cartilages
of the Rhea, the Hemipod, and the “Coracomorph.” The whole “ectoethmoid” (p.p)
is still soft; but the “ trabeculo-palatine”’ conjugational lobe is marked out (0.w).

The inbent clinging pterygoids are thoroughly Picine, with their exceedingly large and
long mesopterygoid (m.pg), and having no “ epipterygoid”’ spur. They are underbound
by the filiform out-turned ends of the palatines (figs. 3 & 4, pa), which are slightly
grooved in their gradually broadening wider part. This region forms a very obtuse
angle, the Passerine transpalate being quite gone. They differ in this from Picus.

The interpalatine and ethmo-palatine styles are equally long and slender (i.pa, e.pa) ;
and the prepalatine bar is filiform and of great length. I have not been able to find a
*‘ mediopalatine.” The maxillaries (mx) have an extremely small palatine lip (mz.p)
opposite the thick angular process; the jugal process is very long, and is separate
from the jugal (j). The palatine edge of the maxillary is more abortively developed on
the left than on the right side; but the deficiency is supplemented by the palato
maxillary ossicle (p.mz).

A series of sectional views will illustrate the relations of the nasal sacs to the rest of the
face. The first of these (Plate IV. fig. 5) is through the body and fore part of the palatine
process of the preemaxillary and the preenasal cartilage (pa, p.px, pn). The next (fig. 6)
cuts through the fore part of the alinasal cartilage (al.7), the alinasal turbinal (al.¢0), the
fore part of the septum with its trabecular base (s. 2), and the related parts of the facial
bars (.px, pa, mx, p.px). The next section (fig. 7) shows that the alinasal turbinal
(al.tb) is coiled upon itself once and a half, whereas it is scarcely coiled once in
Gecinus (Plate I. fig. 7, al.tb). In this section the upper and lateral parts of the pre-
maxillary, the maxillaries, the two crura of the nasals, and the preepalatine bar are cut
through: here the septum (s. 7) is at its greatest depth. Further back (fig. 8), behind
the maxillary angle, the inferior turbinals have come into view (i. td). They do not finish
one coil, and are scarcely more developed than in Gecinus (Plate II. figs. 7 & 8,7. 6). The
“ alinasal turbinal”’ (ad. tb) is there also behind its root; it has lost most of its coil, but
has developed an inner outgrowth. This section is through the notched part of the
cranio-facial axis. Still further back, behind the alinasal turbinal, and behind the notch,
we come to the thick-topped shelving “ mesoethmoid” (p.e), with its bony matter; the
alee above are the “ ali-ethmoids” (a/.e). Here on each side of the lower nasal nerves
there is a large flap of the nasal fioor, which reappears behind. The “ali-ethmoids’’ grow
down and form a subvertical wall, from the lower part of which the posterior extremity
of the inferior turbinal is growing. Above this there is a new scroll. All these scrolls

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 19

begin above; the one in front of this is the inturned and flattened fold which represents
the upper turbinal (w. ¢b). Here the inferior turbinal (i. 42) sends out an inner out-
growth like the one in front of it.

The middle turbinal is a mere ridge in the front of the pars plana, where the inferior
turbinal ends (see in Gecinus, Plate V. fig. 2, m.¢b). When the razor has cut through
the thickest part of the rapidly ossifying “‘ mesoethmoid,” we get a view of the orbital face
of each “ pars plana” (Plate IV. fig. 10, pe, pp) with its conjugational angle corresponding
to the ‘os uncinatum.”’ It is in this section that we get a view of the ‘‘ median sphenoidal
sinus” (m.sph) and the two vomers; (see enlarged view, fig. 11) in this region; both
endostosis and ectostosis are at work, hardening the “ perpendicular plate,’’ which grows
outward into the “ ali-ethmoids,” which, infolding themselves and becoming pinched
at the fold, give us ultimately the ear-shaped wavy crenate upper turbinal (compare fig. 9,
with Plate V. fig. 2, w. tb).

Already I find a small styloid lacrymal behind the nasal crus, and a distinct centre in
the orbito-sphenoidal region.

Summary of characters of the ‘‘Saurognathous”’ type of Palate.

A. Trabecular Arch.

1. Retention and ossification of trabecular cornua, and of basitrabecular (przenasal)
rostrum.

2. Addition thereto, in certain types, of a pair of “ labial” cartilages, which do not
form the foundation of an oxhead-shaped compound vomer as in the Passerine.

3. Detachment, in Gecinus, at least, of part of the nasal wall, after the ingrafting on
it of a septo-maxillary.

4.. Great number and bilateral independence of the vomerine series of bones, some
of which are azygous.

B. Palatine Arch.

1. Absence of a distinct mesopterygoid, the process answering thereto being very
long, and in the embyo reaching the vomer of that side, as in the New-
Zealand Lizard Hatteria.

2. A dagger-shaped basipalatine between the right and left bones.

3. Absence of a distinct “ transpalatine.”

4, Presence of a postpalatine bone in the most generalized form (namely, Picumnus).

5. Absence of epipterygoid hook, and presence of a large spur for muscular attach-
ment on the pterygoid.

6. Abortive development of maxillo-palatine plates, and presence of a distinct palato-
maxillary bone on the left side only.

C. Hyoidean Arch.

1. Abortive development and early fusion of the ceratohyals.
2. Double “ infrastapedial ” and ossified stylohyal.
p2

20 MR. W. K. PARKER ON THE MORPHOLOGY OF

D. Third postoral arch.

Length and tenuity of basal piece and upper and lower pieces of the cornua caynes
hyals, or 1st cerato- and epibranchials).

E. Large size, but great simplicity, of nasal turbinal folds.

F. Cowrie-shaped tympanic cavity,

Concluding Remarks.

The “ Celeomorphe ” of Huxley form a most natural and well-defined group—a group
equal, zoologically, to the Pigeons or the Parrots. Evidently this differentiation has
taken place through the gradual extinction, during long secular periods, of conjugational
types more generalized than those now extant. Iam not without evidence as to the
truth of this view; but it must be discussed elsewhere ; yet proofs will be given in proper
time and place.

Those who are familiar with the Lacertian type of skull will agree with me as to the
propriety of such a morphological term for the Woodpeckers and Wrynecks as ‘“ Sau-
rognathe.” It will exactly fill up the lacuna left by my friend in the masterly paper
already referred to. The view there expressed that these birds have a Passerine foun-
dation, but that they are somewhat abortively developed, arrested one way and wonder-
fully specialized in another, will be seen to be the exact truth of the matter.

Corresponding with what I am showing in papers for the ‘ Zoological Transactions,’
with regard to the “ Aigithognathe (Passerinz,’ “ Coracomorphe”’), the most embry-
onic and least specialized Woodpecker is a South-American form, namely Picwmnus.

We have seen how numerous the bony centres are in this group; and it is not easy to
account for this peculiarity. They are very numerous in all birds, but least so in the
Thrushes (“Turdidz’’); whilst in the “ Coracomorphz” generally there is an average
development as to numbers. There is some reason for these differences ; but it is hidden
amongst the extinct forms, both warm- and cold-blooded.

Teleologically, the “ Picidze’”’ cannot possibly benefit by the numerous small bones
that havea temporarily separate existence in their palate, many of these being mere grains,
and others most feeble splints.

The general agreement of the palate of Picuwmnus with that of the last of the Rhyn-
chosaurian Lizards (namely Hatteria) is very suggestive. In the dow South-American
Passerinz, the “‘ Formicariide”’ and the “Cotingide,” the essentially Reptilian face
shows itself most clearly. .

But that which characterizes the whole of the ‘‘Celeomorphze”’ is the want of fusion
of the parts of the palate at the mid line. This is evidently an ancient reptilian
“atavism.” This must be admitted in a metaphorical sense, if not in an actual: it is a
remarkable verisimilitude, if not a real historic truth.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.

THE SKULL IN THE WOODPECKERS AND WRYNECKS.

DESCRIPTION OF THE PLATES.

Puate I.

1. Palate of Picus major, adult. x 4 diameters.

2. Palate of Gecinus viridis, oldest fledgling. x 24 diameters.

3. Vertical section of skull of G. viridis, younger fledgling. x 24 diameters.
4. Side view of face of ditto. x 24 diameters.

5.
6
7
8
9

Ist section of ditto. x 5 diameters.

. 2nd section of ditto. x 5 diameters.
. 8rd section of ditto: x 5 diameters.
. 4th section of ditto. x 5 diameters.
. Part of tongue of youngest Gecinus (lower view). x 10 diameters.

10. Part of ditto (upper view). x 10 diameters.
11. Stapes of adult Gecinus. x 74 diameters.

mn Oo BB & OD

Puate II.

Gecinus viridis (continued).

. Palate of adult. x 24 diameters.
. Part of same. x 5 diameters.

Septum nasi. x 4 diameters.

. Vomerine bones (outer view). x 4 diameters.
. Vomerine bones (inner view) x 4 diameters.

Palate of specimen six months old. x 4 diameters.

. Part of nasal capsule of ditto. x 4 diameters.
. Part of fig. 6. x 8 diameters.

Prats II.

. Palate of Hemilophus fulvus (adult). x 24 diameters.

. Part of same. x 5 diameters.

. Palate of Picus analis (adult). x 5 diameters.

. Part of same. x 10 diameters.

. Part of palate of Gecinus viridis (older fledgling) from above. x 43 diameters.
. Ditto (younger fledgling). x 44 diameters.

. Pterygoid and part of palate of adult Gecinus. x 24 diameters.

Puate IV.

. Palate of Picus minor (fledgling). x 6 diameters.

. Part of same. x 12 diameters.

. Palate of Yuna torquilla (nestling). x 54 diameters.
. Part of same. x 11 diameters.

21

ON THE SKULL IN THE WOODPECKERS AND WRYNECKS.

5. Ist of a series of vertically transverse sections of face of Yunx torquilla. x 8 diameters.
6. 2nd section of ditto. x 8 diameters.

7.
8
2)

3rd section of ditto. x 8 diameters.

. 4th section of ditto. x 8 diameters.
. 5th section of ditto. x 8 diameters.
. 10. 6th section of ditto. x 8 diameters.
.11. Part of same. x 24 diameters.

PuatTe V.

. Side view of skull of Gecinus viridis. x 2 diameters.

. Fore view of ethmoid of ditto. x 2 diameters.

. Palate of Picumnus minutus (from below). x 10 diameters.

. Part of ditto (from above). x 10 diameters.

. Part of dito (side view). x 10 diameters.

. Side view of septum nasi of Picumnus minutus. x 10 diameters.

. Diagram of visceral arches and sense-capsules in the embryo of an aérial bird at the middle of

incubation :—o/, nasal sac; e, eye; au, auditory sac; tr, trabecula; p.n, prenasal; c. tr, tra-
becular cornu; pa, palatal; pg, pterygoid; g, quadratum; mk, Meckel’s cartilage ; st, stipes ;
5, 5th nerve; s.st, suprastapedial; e.st, extrastapedial ; i.st, infrastapedial ; st.h, stylohyal ;
c.hy, ceratohyal; bh, basihyal; uh, urohyal; br. 1, 1st branchial.

Trans.Linw Soc.,SER.2.ZooL.Vor.1 Tas. |.

WK. P. ddl.ad nat; GWeet lth,

WWest &Co. imp.

a Traws.Liny.Soc.,Ser.2. Zoot Von.1. Tas. II.

\, WE Padel.ad nat; GWest lth.
j ff . 2 . «
7 Gecinus viridis.

W.West &Co. imp.

‘ Trans.Linw. Soc. Ser. 2. Zoou.Vou..Tas. 3.

WK. P. del. ad nat. C.West ith.

' ines Ae &. Hemilophus fulvus. Figs 3,4. Picus analis, Figs 5-7. Gecinus viridis.

W. West & C? imp!

Tears) Lin Soe. SER 2.Zoor. Vow.|. Tas 4

a Cana

SST
RAE LINSS
7

WEP. del. ad nat.G.West lith

j Figs 1,4. Picus mmor. Fig? 3—11. Yumx torquilla. WiWerhe: C2 eue

Trans. Linw.Soc.Ser.2.Zoor Vor. Tas. 5.

WK PR. del ad nat G. West lith

: - Oe ER : f W. West & C° imp
Fige1,2a@- Gecinus viridis. Figs 3—6. Picumnus minutus.

[ 23 |

II. On some Atlantic Crustacea from the ‘ Challenger’ Expedition. By Dr. R. v.
WILLEMOES-SuuM, Naturalist to the Expedition. Communicated by Prof. WYVILLE
THomson, F.L.S. & L.S.

(Plates VI-XIII.)

Read May 7th, 1874.

AMONG the great quantity of deep-sea Crustacea which have been. brought up during
our cruise in the Atlantic, either by the dredge or the trawl, I have selected the most
interesting ones for immediate description. Besides, readers will find in this paper the
description of both sexes of an interesting Nebalia from the shallow water of the
Bermudas, some remarks on the structure of Cystisoma, and a note on the develop-
ment of a Land-Crab from the Cape-Verds. Some of the deep-sea forms, the Wille-
moesie (Deidamie), Astacus, Zaleucus, and Gnathophausia, have been already figured in
‘Nature,’ in the “* Notes from the ‘ Challenger’”’ published by Prof. Wyville Thomson ;
and a short description, taken from my first notes on the subject, has been added. In
this paper it is now intended to give a more detailed account of their structure and their
systematic position. The paper is divided into the following seven parts :—

I. On a blind deep-sea Tanaid.
II. On Cystisoma Neptunus (Thawmops pellucida).
III. On a Nebalia from the Bermudas.
IV. On some genera of Schizopods with a free dorsal shield.
V. On the development of a Land-Crab.
VI. On a blind deep-sea Astacus.
VII. On Willemoesia (Grote), a deep-sea Decapod allied to Eryon.

I have only worked out such species as, from their structure and zoological affinities,
present a greater interest or belong to groups about which we had a satisfactory lite-
rature on board. Should, nevertheless, a want of literary information be remarked in
some point or other, the reader is asked kindly to remember that this paper was worked
out on board a ship cruising in the Atlantic.

H.M.S. ‘ Challenger,’
Cape of Good Hope, November 1873.

I. ON A BLIND DEEP-SEA TANAID. (PI. XII. figs. 1-9.)

During our cruise in the Atlantic, members of the Anisopoda tribe were found twice.
First, on the 2nd of May a real Tanais was got from a depth of 1700 fathoms off the
North-American coast; I cannot determine whether this animal belongs to a known
species or not. Later, when cruising among the Azorean islands San Miguel and Santa

24 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

Maria, we got from a depth of 1000 fathoms another Anisopod, which belongs to the
interesting genus Apseudes, according to Mr. C. Spence Bate* the only Isopod in which
the antennal scale, so common in the Macrura, is present on the lower antenne, and, as
we may add, one of the few in which there are two flagella in the anterior antenna.
In the species of Apseudes hitherto known the eyes are pedunculated: in this deep-sea
animal there are no eyes at all; hence we might call it 4. ceca. It has a length of
6 millims., the females having the same length as the males.

The cephalothorax, to which is attached the first pair of gnathopods, has a spiny
rostrum and two prominent spines on each side. The first antenne (fig. 1, a’, and fig. 2)
show a very strong funiculus and two flagella, one of which consists of twenty-four, the
other of eight joints. The longer is double the length of the smaller one.

- The exterior or second antennee (fig. 1, a’, and fig. 3) have a smaller flagellum, con-
sisting of eight joints and an antennal scale articulating with the first joint of the
funiculus (fig. 3, s). The mandibul have a palpus consisting of several joints.

The maxilliped (fig. 9) has strong tufts of hairs on the last three joints. The two
enathopoda are very different in form: the first one (fig. 4) is short, and terminates
in a strong recurved chela; while the second is much longer (fig. 5), and has enlarged
and flattened joints with strong spines, as may also be seen in the other species of
Apseudes. The pereiopoda are strong but not very much enlarged appendages. We
have drawn the first and second as well as the top of the fourth; the third is terminated
by a bifid claw, as is the sixth in A. falpa, Mont., which we have not drawn, as only
once could we clearly get sight of it. When afterwards it was tried again to get a view
of the two claws, it was impossible to separate them from each other.

The first five segments of the pleon are short, and possess small biramous pleopoda.
The last segment is elongated, as in 4. talpa, and shows at the top two pairs of caudal
sete.

In the females there are breeding-lamelle at the base of the second gnathopoda and
the first two pereiopoda, which are wanting in the males. -This is the only difference
which I could find out between the two sexes.

Most of the specimens which we got were washed out from the mud which the. dredge
had brought up. We did not meet with this Crustacean again.

_ This is not the first instance of blind Isopods being described, as some subterranean +
and parasitical { forms are well known to have no eyes, or to lose them, as Anceus does,
when getting old. There is also a whole family, the Munopside, which is blind, though
most of its members live in moderate depths. A very large member of this family
has also been got by us in the deep sea; it cannot, however, be described now, as the
necessary literature is not at hand.

II. ON Ovsrisoua Neprowvs (Taavmors petrrvcrpa). (Pl. XI. figs. 4-8.)
In a postscript to my paper on Thawmops pellucida (Phil. Trans. of the Royal So-
ciety, 1873) I have described the male of this remarkable Amphipod, and given some

* Facts and Arguments for Darwin, by Fritz Miller, translated by Dallas (London), 1869, p. 132.
+ Pitanethes albus and Lyphloniscus Steinir. ; + Bopyride and the female of Anceus.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 25

details on the structure of both sexes, which could only be discovered when it was
possible to dissect one of the specimens. I now have to publish some figures which
could not be given in my first paper, and add a few words to them. But before doing
this, I must declare that the name of Thaumops pellucida must be withdrawn, as this
animal has been already described under the name of Cystisoma Neptunus by Guérin-
Méneville*, from a single specimen caught in the Indian ocean. I see this from Spence
Bate’s ‘Catalogue of Amphipodous Crustacea’ +, which I have just received, and which
contains a copy of Guérin-Méneville’s rather bad figure and a short description of a
specimen, which probably is a male.

Guérin-Méneville (and Spence Bate?) regards Cystisoma as belonging to the Hyperide
family, and takes the antenne (which are in front of the head) as representing the
second pair. Why they should be homologous with these I do not know; nor do I think
they are; for in Phronimids, Oxycephalids, and Hyperids the second antenne have a
funiculus consisting of several (more than two) joints. Now in Cystisoma the antennze
have got two joints, which seem to represent the one a funiculus joint, and the other the
flagellum of the first antennze of Phronima.

Against a union of Cystisoma with the Hyperids may be advanced, besides the form
of the head (which is more Typhid-like) and the absence of the second antennz in both
sexes, the absence of a palpus on its mandible (Pl. XI. fig. 6). The palpus is always
present, according to Claus, in Hyperids, but is wanting in Phronimids. Besides, there
are so many peculiarities distinguishing Cystisoma, which I have enumerated in my
above-mentioned paper, that I do not think it possible to keep it in one of the hitherto
received families, but that now, when we know its structure better, we have to establish
for it a special family called Cystisomide.

The first maxilla has been described, and is now figured in Pl. XI. fig. 7. The second
maxilla could not be found, and is probably represented by an organ which might have
arisen from the union of the second maxille and the under lip (Pl. XI. fig. 8). This
organ (7) is attached to the third joint of the reduced maxillipedes, and surely acts phy-
siologically as an under lip. The maxillipedes are reduced and fused together so as to
form one organ consisting of two basal joints and two claws (fig. 8, ™), between which is
attached the organ which we suppose to represent the maxillz and the under lip.

The male differs by the absence of glands at the top of nearly all the appendages,
especially in the last pair of pereiopoda, which, according to this, have not the same
clumsy appearance as in the female. The two testes begin just behind the stomach
(fig. 5,7), and send vasa deferentia to the last segment of the pereion, where two simple
genital openings are to be seen between the last pair of legs (fig. 5, a g).

Probably (as in Phronima) the full-grown male is somewhat smaller than the female,
it seems that Cystisoma Neptunus can attain a very considerable size; for the last
and largest male which we got in the trawl has a length of 108 millims. It seems
that these oceanic amphipods have a very wide geographical distribution. Cystisoma,
as we have seen, was first described from the Indian Ocean; and Oxycephalus piscator is

* Guérin-Méneyille, in Revue Zoologique, July 1842, p, 214. 7 Page 311, pl. 1. fig. 7.
SECOND SERIES.—ZOOLOGY, VOL. I. £

26 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

mentioned as occurring in the same. Now we have found in the Atlantic not only four
specimens of Cystisoma, but also numerous specimens of two species of Oxycephalus, one
of which seems to me to be identical with O. piscator.

III. On a Nezzazrs FROM THE BERMUDAS. (Plate VI.)

During our stay in the Bermudas Mr. Murray collected one day some small Crustacea.
in Harrington Sound, a bay which only communicates with the sea through a narrow
passage. Among them I was very glad to find a Nebalia, and went then myself to the
bay, where this interesting little Crustacean was not rare under stones and corals. I
collected, perhaps, twenty females and two males, and was able to make some observa-
tions on them which, added to the embryological researches of Mecznikoff and the
anatomical remarks of Claus* on both sexes of the Mediterranean N. G'eoffroy?, will, I
think, enlarge somewhat our knowledge of these Crustacea, which for such a long time
have been considered to be Phyllopods, but, according to the above-named papers, are
more rightly placed among the Schizopods.

The female of this Nedalia, which I am going to call N. longipes, has a length of
6 millims., and is, when living, of a milk-white colour. The shell, posteriorly and above
somewhat flattened, has an oval form; it terminates anteriorly in a sharp and slender
rostrum, which covers the pedunculated eyes; the upper surfaces of these eyes are entirely
covered with small spines, so that the perception of the light is only possible through
the facets of the under surface. The shell does not quite extend so far backwards as in
N. Geoffroyi, where it reaches the fifth abdominal segment.

The anterior antenne (figs. 1, 2, a, and fig. 5) consist of four joints, the last of which is
a denticulated claw (fig. 5, /), From the base of this claw start two flagella (fig. 5, a and
b), the shorter of which is transformed into a large squamiform appendage, fringed with
hairs; the other one is more slender and densely covered with olfactory hairs.

The posterior antenne (fig. 7) consist of three joints only, the last of which is densely
coyered on its protuberances with tufts of hairs, especially at the point of insertion of the
flagellum, which itself is divided into nine imperfect joints marked by tufts of hair.

A comparison of the mandibula and the two mazille (figs. 8-10) with the corresponding
parts in V. Geoffroy: shows that there are only very slight differences between them. I
saw the palpus of the first maxilla (“ Putzfuss,” Claus) in the same position in the male
as in the female, z.e. bent backwards.

More important are the pectoral feet; for on their account especially Nebalia was
formerly supposed to be a true Phyllopod. If this Bermudan species had been the first
Nebalia which had been noticed, I do not think that this would have occurred ; for in the
feet of this one the phyllopodal characters are very little perceptible. In fig. 3 I have
figured a pectoral foot of NV. Jongipes, and in fig. 4, for comparison, one of NV. Geoffroyi
(taken from Claus’s paper); so that the corresponding parts will be easily found out.
The two-eared branchial appendage of the basal joint (figs. 3 & 4, K A) is very large in WV.
Geoffroyi, and in N. longipes only rudimentary. The appendage of the second joint has

* Von Siebold und Kolliker’s Zeitschrift fiir wissenschaftliche Zoologie, vol. xxii. p, 323.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 27

entirely lost its flattened form, and is a sort of leg, which, however, like the main branch,
is not jointed, but is densely covered with hairs on one side. When comparing such a foot
with the pereiopod of a true Schizopod, as, for example, Lophogaster (Sars, Beskrivelse
over Lophogaster typicus, tab. il. fig. 35), we see in both on the basal joint the bran-
chial appendages, which in Nebalia longipes are very simple, and in Lophogaster divided
and arborescent. From the second joint starts the palpus, which in our case has a
different shape from that of Lophogaster, but which morphologically is the same thing.
In the foot itself we find in Lophogaster well-defined joints and a terminating claw, in
N. Geoffroyt we still find the joints, but in our case we find no joints; in all three, how-
ever, it is not difficult to find out the corresponding parts.

The first four pairs of abdominal feet or pleopods (fig. 11) have a large basal joint den-
ticulated exteriorly, to which two rami are attached, the interior of which is longer and
has at its base a little appendage, which is by means of a denticulation at the top united
to a similar one of the other pleopod. It has been called by Claus, who found it in W.
Geoffroy, the retinaculum. Such appendages have been shown by Claus also in Stoma-
topods, and figured by Milne-Edwards from the Decapod genus Afya. The fifth and
sixth pair of pleopods resemble very much those in WV. Geoffroyi.

The two joints of the fwrca are covered at the top with tufts of long and dense hairs.

Among twenty to twenty-five females I discovered two or three males, easily recogni-
zable by their minute size and the enlarged flagellum of the anterior antenne, which here
are transformed into real prehensile organs. The males are only 8 millims. long, and
differ from the females also by being more slender, by their shorter pleopods and their
antennee, which are on the whole somewhat shorter and bigger. The pectoral feet are
exactly the same as in the females. In the males of WV. Geoffroyi the second pair of
antenne have an enormously long flagellum, something like the Cuma male; while in our
Nebalia the second antenna differs very little from that of the female, nor are the olfac-
tory hairs to be seen in much greater quantity, unless perhaps at the top and the outer
edge of the third joint. The first antenna (fig.6), however, is very different in the male ;
for its squamiform flagellum is greatly swollen up, and the joints are only to be discerned
as small protuberances with some hairs on them on one side. This conformation is
rather a phyllopodal one—so far as I can make out here, something like that found in
Estheria.

I found, like Claus, a ductus ejaculatorius directed towards the base of the eighth
pereiopod, but could not exactly see its opening.

This Nebalia differs from the two species hitherto known, by the spines on the upper
surface of its eyes, by the tuft of hairs at the top of the furca, by the form of its legs, in
which the phyllopodal character has been nearly entirely lost, and the schizopodal cha-
racter has been more approached than in any other species of this genus. in the male,
however, characters have been found which are wanting in the male of NV. Geoffroyi. We
see a modification of the first antenna reminding us of Astheria, but no prehensile organs
on any of the feet. This species therefore approaches, so far as the shape of its legs is
concerned, the higher Crustacea more than NV. Geoffroyi; but the prehensile organs of
its male are characters which show, again, that there remains a good deal of phyllopodal

ee

28 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

element in Nebalia. We have now, however, discovered another Schizopod, in which the
legs are entirely normal and the shield is phyllopodal (Gnathophausia), but evidently
not to be derived from the Hstheria-like shell of Nebalia, but from the carapace of Apus.
And in this same Gnathophausia there is an abdominal segment which shows that, now
being essentially a Schizopod, it is in genealogical connexion with forms which have had
more segments in the abdomen than the Schizopods have.

These two forms, though both Schizopods, show characters which connect the lower
Crustacea more and more with the higher ones. Perhaps the fossil forms might give some
further evidence. We may also hope to get more forms from the deep sea which will
enable us more and more to understand the curious genera which are known under the
names of Ceratiocaris, Dithyrocaris, &c.

IV. ON sOME GENERA OF SCHIZOPODS WITH A FREE DoRSAL SHIBLD.

1. On GNATHOPHAUSIA, a new Genus of Lophogastride. (Plates IX. & X. figs. 2-4.)

During our cruise from the Bermudas to the Azores, Madeira, Cape-Verds, and from
there to the Brazilian coast, three species of this new genus of Schizopods came up in
the trawl or in the dredge. One of these species (now probably figured in Professor
Thomson’s “‘ Notes from the ‘Challenger,’” in ‘Nature’) is very large, probably the largest
Schizopod which we know. Of this one, which I have called Gn. gigas, only one male
was caught; of another, smaller species, Gn. zoéa, two males and two females; and of the
third, Gn. gracilis, only one male. Of these I have dissected one female of Gn. zoéa. I
have also made drawings of the legs of Gu. gigus, which I could not dissect ; but finding
that they have a close resemblance to those of the former one, I shall not publish them.
A sketch of the oral apparatus, however, and of a maxilliped, of the larger species are
among my figures.
| GNATHOPHAUSIA *, n. gen.

1.. GNATHOPHAUSIA GIGAS, 0. Sp.

The specimen, a male, came up from a depth of 2200 fathoms in lat. 38° 22’ N. and
long. 37° 21' W., between the Bermudas and the Azores.

Ti has a length of 142 millims., and much resembles at first sight a large Caridid
shrimp ; but the abdomen is straight and not bent down by the enlargement of the third
pleon-segment, as is the case in the Caridide.

'' The Carapace.—Its anterior part extends as a triangular rostrum (85 millims. in
length), starting in the form of a high ridge from the midst of the hepatic region. From
its base to the top of the rostrum the dorsal shield has a length of 64 millims.; it is a
flexible parchment-like duplicature of the skin, covering entirely the lateral branchiz,
and touching with two lateral and backward-directed spines the middle of the second
abdominal ring. Its upper posterior edges, however, do not quite cover the last segment
of the pereion. In Lophogaster it leaves two segments entirely free, and its borders are
not united with the pereion. Here the carapace starts from the cephalic and the first
pereion segments as in Apus and Nebalia. It is in no connexion with the five posterior

* Tya0os and davotos.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 29

segments of the pereion. On its upper surface there are five longitudinal ridges and two
transverse sulci, which can best be understood from the figures (Pl. X. figs. 2 & 2 a),
These ridges divide the carapace into five regions, which we might call the regiones
cephalica, hepatica, cardiaca, branchialis, and marginalis. I think it unnecessary to give
for each the exact dimensions.

In front there is a cleft for the pedunculated eyes, above which there is a sharp spine ;
laterally, above the cleft, which is occupied especially by the second maxilla, there is
another spine. The lateral edges are more or less flattened, and form two “rounded
plates,” as Sars calls them in his paper on Lophogaster *, between which there is the
just-mentioned cleft. This can best be explained by Pl. X. fig. 8, where I have figured
the parts of the mouth and the upper lateral edges of the dorsal shield, seen from below.
There the upper plate (2), to which are attached the muscles of the oral apparatus, covers
to some extent the mandible; the lower plate (y) protects the palpus and the accessory
eye of the second maxilla from behind.

In Lophogaster there is also to be seen a rudiment of a rostrum and the longitudinal
ridge (Sars, fig. 1). The lateral spines are very much like those of Guathophausia; the
lateral edges of the shield, however, are not prolonged to such an extent backwards as in
our genus, and the posterior margin leaves, as has been mentioned already, two segments
entirely free.

The eyes issue from between the rostral portion of the shield and the base of the inner
antennz ; they have a length of 8 millims., and show two joints in their peduncle.

The «ner antenne consist of a short and stout funiculus and two flagella. The funi-
culus is four-jointed, and attains a third of the length of the squamiform appendage of
the outer antenna. It is twice the diameter of the eyes in width, and is densely covered
with hairs on its inner side. The exterior flagellum is very long (130 millims.), and flat-
tened at its base (2 millims.), but the interior flagellum rounded and much shorter (length
40 millims.). Underneath and behind the inner antennz is the large base of the squami-
form appendage, showing outside a strong spine. The appendage itself entirely covers the
funiculus, and has a length of 19 millims.: its inner border is even, but densely covered
with hairs; the outer one has at the base a rounded prominence, and further on four
sharp spines, in addition to the top spine. A longitudinal ridge (a trace of which may
also be seen in the cordiform appendage of Lophogaster—Sars, fig. 13 e) divides the
appendage into a larger and a smaller portion. Underneath this appendage we see a
three-jointed long funiculus of the exterior antenna (length 5 millims.), the flagellum of
which has the length of the shorter one of the inner antenna (40 millims.).

Oral apparatus. (Plate X. fig. 3.)

The upper lip is a large cordiform plate (63 millims. in length by 5 millims. in width),
the top of which reaches upwards to the first joint of the antenne interioris funiculus.
A short groove descends from its centre towards the mouth ; this organ is nearly of the
same shape here as in Lophagaster (Sars, fig. 17). Very Lophogaster-like are also the
strong mandibles (fig. 8, md), to which are attached two three-jointed palpi, the inner

* «Beskrivelse over Lophogaster typicus’ (Christiania, 1862), pl. i. figs. 3 & 4, and p. 2.

30 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

side of which is densely covered with hairs (fig. 3, mp). The palpi are situated on both
sides of the upper lip, and reach higher up than the funiculus of the exterior antenna.
The first pair of maxillz could not be inspected, as they are entirely covered by the second
pair, which, like the whole oral apparatus, have a strong resemblance to those of Lopho-
gaster (Sars, fig. 28, cd). All the parts mentioned there are also to be found in our case
(fig. 8, ma) :—on the inner side the two digitiform manducatory processes covered with
stiff hairs; close to them on both sides a flagellum (fig. 3, /), which is jointed in Gna-
thophausia; and on the outsides the palpus (fig. 3, p) in the form of a large leaf-like
appendage fringed with hairs and occupying, like a large cover, the whole space between
mandibles and maxillipeds. From the basal joint of these maxille start a pair of small
black pigmented accessory eyes, the peduncles of which have a length of 2 millims.
Their position and direction are such that they just peep out between the inferior mar-
gin of the dorsal shield and the gnathopods. A dissection of these accessory eyes is
of course impossible in our case; but I think one is right in supposing that they consist,
like the accessory eyes of the Euphausiide, of a lens, nervous rods, and a pigmented sub-
stance. They have been found in the Euphausiide on the basal joint of the pereiopods,
as well as of the pleopods, and in Thysanopoda norvegica even up to the number of eight
pairs. In Zophogaster there are none, nor have accessory eyes ever been observed on the
maxille of any animal.

There is only one pair of maxillipeds, as there is in Lophogaster; and in both the
enathopods are used for walking. On the large transverse basal joint of the maxillipeds
we have a long flattened palpus (Pl. IX. fig. 16, p), and somewhat further on a small
flagellum (fig. 16, f). The second and third joints of these maxillipeds are very short ;
the fourth is longer, the fifth very short again; and the sixth is the terminating claw.

Gnathopods, Pereiopods, and Branchie.

We have remarked already that the gnathopods do not essentially differ from the perei-
opods, and are used for walking. At the base of all of them, with the exception of the
last pair of pereiopods, are the arborescent gills, the branches of which are partly situated
underneath the lateral parts of the dorsal shield, partly projecting into the water. There
are, as far as I can see without dissecting or spoiling our specimen, three branches, one
of them on the sides, one between the legs, and a larger one leaning on the pectoral shield,
which is entirely covered by two rows of branchiz. In the whole the arrangement seems
to me to be exactly the same as in Lophogaster; and the pictures given by Sars (figs. 16,
35, 37, and 45) give also a very fair idea of the branchial appendages of Gnathophausia.

The gnathopods and pereiopods (Pl. IX. figs. 8-14 may be compared here, though these
figures are drawn from Gz. zoéa) consist each of seven joints, if one counts that basal one
to which the branchize are attached. To the second joint the palpi are attached, which
differ somewhat from those of Lophogaster; they are shorter, and in Gnathophausia
the ramus palpi rapidly decreases in width, and has a great resemblance to the ramus of
one of the pleopods. The palpus of the first pereiopod has a length of 13 millims. The
third and fourth joints are in all the legs very short, and do not offer any thing parti-
cular; but the fifth and sixth are very different from each other in different legs. In

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. al

the first gnathopod they are even, very strong, and somewhat curved inwards; there are
only hairs at the inner side, not on the lateral parts of the joints, which in the second
gnathopod have got three lines of hairs, and in the following ones five and six. In the
last three pereiopods there are not only lines of hairs on the lateral parts of these joints,
but their point of attachment is strongly marked in the chitinous substance, so that
these joints look as if they were divided into five and six subjoints. And the same thing
is the case throughout with the fifth jomt. Such subjointed pereiopods are also found
in Yysis, but not in any other genus of Schizopods.

In the gnathopods as well as in the pereiopods the last joint is a short claw—even in
_ the first pair of the former, where it is wanting in Lophogaster.

The Pleon and its Appendages.

The abdomen has a length of 78 millims. It is square-shaped; for its segments are
nearly as large as they are high. The second segment, for example, is 11 millims. in
height by 10 millims. in width. The edges of the segments are even; only laterally
there are squamiform processes, which cover each other, and each of which has a rounded
anterior and a sharpened posterior portion. The shortest segment is the fifth; the sixth
is much larger, and is divided by a deep sulcus into two parts, which look perfectly like
. two distinct segments. One is the more inclined at first to think so, as the lateral squa-
miform processes, which terminate all the preceding segments, are attached to the first
of these two subdivisions. In the second one there are only two short spines on each
side. Sars has described something very much like this in Lophoguster ; and as we now
know that Medalia, in which there are nine abdominal segments, has essentially schizo-
podal characters, while Gnathophausia in its unattached dorsal shield has decidedly a
phyllopodal character, this pseudo-amplification of the abdominal segments seems to me
to be exceedingly interesting, as it shows that Gnathophausia must be in genealogical
connexion with forms (or a descendant of forms) in which there are more than seven
segments in the pleon.

The telson has a length of 30 millims.; on its surface there is a longitudinal groove
bordered by two ridges; the lateral borders of the telson are denticulated; at its end
there are two spines, between which there is a semicircular cleft. The caudal appendages
are both jointed (Pl. X. fig. 2); the interior are somewhat shorter than the exterior ones ;
and both are shorter than the telson ; at their basal articulation there is a spine on both
sides.

The five pairs of pleopods consist of a pear-shaped basal joint, at the posterior side ot
which there is a protuberance; at their inner side they are covered with a fringe of hairs.
‘The two annulated rami do not present any thing particular. Appendages such as have
been found here in the Huphausiidz do not exist in Guathophausia.

Before fixing the generic and specific characters of this large species, [ shall now give
a shorter description of the smaller one (of which four specimens were got), and point out
especially the characters in which it differs from the larger one.

32, DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

2. GNATHOPHAUSIA ZOBA, 0. sp.

In its general appearance this species differs from Gn. gigas at once by the presence

of a long posterior spine at the dorsal shield.

The specimens, two males and two females, were got in very different places. (1)
June 30,a male was obtained off the Azores in a depth of 1000 fathoms; it has a length
of 49 millims., the rostrum being 18 millims. long. (2) August 25, in lat. 1° 47’ N., long.
24° 26' W.,a male came up from a depth of 1650 fathoms: length 58 millims., with a
rostrum of 17 millims. (8) August 26, a female from 1500 fathoms. in lat. 1° 22'N., long.
26° 36’ W.: length 81 millims., with a rostrum of 14 millims. (4) September 12, off Rio
San Francisco on the coast of Brazil, another female came up from 750 fathoms: length
313 (rostrum 7) millims. According to this the size of the males varies between 49 and
58 millims., and that of the females between 313 and 81.

The substance of the dorsal shield is flexible, like strong parchment ; it is as in Gn.
gigas a duplicature of the skin, adhering to the body only in the first segments, and in
no connexion with the five posterior segments of the pereion. A ridge running over the
whole length of the shield is continued into a long triangular rostrum, which, in the
males, is much longer (3 of the length of the whole animal) than in the females (+ of the
length). The posterior spine is a continuation of this ridge, and has in both sexes a length
of 6-8 millims. Both rostrum and posterior spine are triangular and armed with small
secondary spines, which in the former are to be seen on all sides, in the latter only on two
sides. The posterior spine reaches backward to the third abdominal segment; and all the
segments of the pereion are covered by the dorsal shield, with the exception of the lateral
parts of the last one.

The posterior angles of the shield are rounded. In front there is a sharp spine above
each eye, and another somewhat smaller one below it, just inside, at the place where we see
the round plate, to which are attached the muscles of the oral apparatus. A transverse
sulcus, a transverse ridge, and two longitudinal ridges (besides the larger central one)
divide the shield, as in the former species, into cephalic, hepatic, cardiac, branchial, and
marginal regions (Pl. X. fig. 4).

The inner or first antenne hardly differ from those of the preceding species. Their
funiculus only reaches to the middle of the squamiform appendage. The outer or second
antenne differ from those in Gn. gigas by the shape of their scale, in which only one
spine is visible (Pl. IX. fig. 2, a1), while in the larger species there are five spines.

The parts of the mouth in this species could be dissected and submitted to micro-
scopical examination, which showed that they are very much like those in Lophogaster.
The very strong and big mandibule (Pl. IX. fig. 3) have a palpus the second joint
of which is somewhat recurved. The third joint shows two kinds of hairs—very short
ones, which also Sars has figured in his genus, and longer ones at the top. ‘The wpper
lip has the same cordiform shape as in the preceding species; and the wader lip (Pl. IX.
fig. 4) is more clumsy than that of Lophogaster, and does not show so deep an incision,
but has the same small hairs at its anterior border. The first maxilla is distinguished
by very strong bristles in its inferior portion, and has a palpus which, among Schizopods,
has hitherto only been found in Nedalia (fig. 5, p); at the top of this palpus we find very

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 30

long hairs which by far surpass it in length. In the larger species I could not examine
the first maxilla; but it probably has got a palpus too. Lophogaster has not got it; and
it is very interesting to find it in Gnathophausia, as it is one more connecting link between
this genus and Nebalia. The second maville are large organs (fig. 6), covering the former
ones and exhibiting a flagellum, a two-jointed palpus, and an accessory eye. I tried
to make out the structure of this eye by hardening it and making it transparent, but did
not succeed ; all I saw was the cornea and a dark pigment inside. For him, however,
who has seen the same organ in the larger species, there is not the slightest doubt that
these sense-organs on the second maxille are the same things which have been called
accessory eyes in the Huphausiidee. They are present in all the specimens of the genus
Gnathophausia in the same place.

In the mawillipeds (fig. 7) of this species I could not find the palpus, which in the
larger is so clearly visible (fig. 16). Possibly it is wanting in the female; for I do not
think that it escaped my attention as I looked for it with the microscope.

The two gnathopoda (figs. 8 & 9) have got at their base breeding-lamelle and three
pairs of gills, one of which is situated on the ventral side, while two are laterally covered
by the shield. No subjoints can be seen in these first two legs, traces of them are to be
found in the first and second pereiopods (figs. 10 & 11); but they are only very clearly
visible on the third and fourth (figs. 12 & 13). In the last (in which I have drawn the
breeding-lamella, fig. 14) they are wanting, as are also the gills at its base. Among
Mysidee we have the same thing which we find now in Lophogastride, for these sub-
joints which are wanting in Siriella (as they here are in Lophogaster) are to be met with
in Mysis.

The abdomen of Gn. zoéa differs somewhat from that of the larger species. We do not
find here those squamiform prolongations, but have simply a few small spines at the
posterior and inferior angle of each segment. In the sixth we have the same peculiar
arrangement of its being divided by a deep ridge into two pseudo-segments which do not
articulate with each other. There is a spine bent backwards at the end of the first one,
and several small spines at the end of the second.

The ¢elson (8 millims. in length) terminates in a half-moon; the caudal appendages
are both jointed; the pleopoda are very slender (fig. 15), and show a small tubercle at
‘the inner posterior side.

The male differs from the female in this species by the length of the rostrum, which
in the former has one third of the length of the whole body, and in the latter only one
fifth. In the size of the inner antennz I see no great difference. Whether the male or
the female attains a larger size I cannot say; the largest specimen of G. zoéa which we
got was a female; besides, it is very easy to tell a male from a female by the want of
‘breeding-lamelle in the former.

3. GNATHOPHAUSIA GRACILIS, n. sp. (Pl. IX. fig. 1).

Only one male of this smallest Gnathophausia-species has been hitherto got; it came up
on the 26th of August, in lat. 1° 22’ N., long. 26° 36’ W., 170 miles east of St. Paul’s rocks;
from a depth of 1500 fathoms. It was on this same day that one of the specimens of Gz.

SECOND SERIES.—ZOOLOGY, VOL. I. F :

34 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

zoéa and the male of Petalophthalmus were got. The specimen has a length of 41 millims.,
with a carapace of 24 and a rostrum of 14 millims. The length of the rostrum (one third
of the whole length) and the absence of the breeding-lamelle show that it is a male.

From the two preceding species this little Lophogastrid differs by the presence of two
spines at the anterior and posterior lateral angles of the carapace and by the presence of
spines at the top of the first, second, and fifth abdominal rmg. From Gn. gigas it is
easily distinguished by the presence of the middle posterior spine of the carapace and —
the absence of the five spines on the lamellar appendage. There are also several spines
on the line which divides the carapace in its full length, which are absent in the other
species.

As in other respects it shows no characters which have not already been found in
the two preceding species, I refer to the figure (fig. 1), and shall not describe in detail
all its parts.

Probably the genus G'nathophausia is represented at the bottom of the deep sea by
many species; but I do not think that these species are very numerous in individuals ;
for the trawl always brings up many more Peneids and Caridids than Schizopods.

Colour, and Mode of Life.

All the Gnathophausias, when they came up, were bright red, a colour which is com-
mon to all deep-sea Crustacea, but which they very soon lose in alcohol.

Like Lophogaster, which was found by Sars at the bottom of the sea in 50-60 fathoms,
these Schizopods live probably near the bottom without ever coming to the surface
(except perhaps in the larval stage). They evidently live together with a rich fauna,
of which the trawl has given us some idea. As to the animals which live together with
the larger species, I am obliged to cite here those which were got the day before and the
day after the capture of Gi. gigas; for on that same day when it came up the trawl con-
tained nothing else. In the first of these stations (June 24, lat. 38° 3’ N., long. 39° 19!
W.) we found a depth of 2175 fathoms; and in the other one, two days afterwards (lat.
38° 25’ N., long. 35° 30’ W.), when we already approached the Azores, there was a depth
of 1675 fathoms. We got there several sponges, a Palythoa, a Pennatulid, a small hydroid
polyp, the Sea-urchin Salenia, several Asterids, some Bryozoa, one Annelid (Hermione), a
Pyenogonid * (Zetes, sp.?), and some blind Isopods, one of which belongs to the Munop-
side family.

' The smaller species were got in different localities and depths (750-1850 fathoms)—
near the Azores, not far from St. Paul’s rocks, and near the coast of Brazil. Together
with them were found many inhabitants of the deep sea, some of which I shall now men-
tion. We got there the sponges Hyalonema, Tisiphonia, and Halichondria, the celebrated
Uinbellularia grenlandica, Fungia symmetrica, and a Flabellum. Among the Hchino-
derms (which were very numerous) may be mentioned Chirodota and Synapta, a Bathy-
crinus, Astropecten, Astrogonium, Asteracanthion, and the Urchins Hehinocyamus and

* From several reasons I supposed, when beginning our operations, that we should get Pycnogonids in great
abundance and variety; but just the contrary was the case, as hitherto, after ten months’ deep-sea dredging, we have
only got from two to three Pycnogonids, belonging, as far as I can make out here, to Kroyer’s genus Zetes.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 35

Salenia. Among the worms, which, unfortunately, usually come up in single and, in most
cases, spoiled specimens, we found some Annelids, a long Nemertean, and a very large
Balanoglossus*. Crustacea there are many—Peneids, Caridids, a Pagurus, and the genera
Chalaraspis and Petalophthalmus (which are described in this paper). Only one Brachi-
opod and several genera of Gastropods (Dentaliwm, Mitra, Pleuronectia, Newra, Limopsis,
and Cardium) belong to this fauna. There were also several fishes, one of which (the
Lophioid Melanocetus) certainly lives at the bottom in great depths.

The Systematic Position

of our genus is easily fixed after what we have seen of its resemblances to Lophogaster ;
for, after all that has been said about it, nobody, I am sure, will doubt that Guathophausia
is a member of the family Lophogastride. Now fortunately Sars has so well discussed
the affinities of Lophogaster, that the systematic position of our genus is easily under-
stood when we say that it is a Lophogaster with a free dorsal shield (like an Apus), with
a palpus on its first maxilla (like Nebalia), with accessory eyes on its second maxilla
and with subjoints in its perelopods:(like Mysis). Gnathophausia, it is true, has many
peculiarities which are not found in the genus Lophogaster; but I must nevertheless
maintain that it is closely allied to that form. The gnathopoda and pereiopoda all having
the shape of legs, the three branchiz, two of which are laterally covered by the carapace
and one not, and the presence of breeding-lamellee on all the legs, are fundamental cha-
racters which, according to our present systematical arrangement of the Schizopods,
assign it its place among the Lophogastride. The subdivision of the sixth abdominal
segment, the shape of all the appendages and of the telson, are nearly exactly the same
in the two genera; but there are other characters which Lophogaster has not, characters
which one might call atavistic ones ; and these are (1) the looseness of the dorsal shield,
(2) the presence of a palpus maxille prime, (3) the subjoimts in the legs, and (4) the
accessory eyes on the second maxille.

The subjoimts in the pereiopods (3) have been hitherto only known as a peculiarity of
the genus Mysis, and the accessory eyes (4) as occurring in some genera of the Euphau-
slide family. We therefore find characters united in Gnathophausia which were only
known separate in the two other families of Schizopods. Hereafter, however, we shall
find in a Mysid a character which until now was supposed to characterize only the
Lophogastride, and shall see that these deep-sea forms of Schizopods combine in many
ways the characters of the shallow-water forms, the types of the hitherto acknowledged
families. But we shall aiso see that they can nevertheless be regarded as belonging to
those families (with the exception of one genus), as they have not a mere mixture of
characters, but are Mysids or Lophogastrids which possess some characters of the other
families.

* Two fragments of this interesting worm were got August 19, in lat. 5° 48’ N., long. 14° 20’ W.., off the west
coast of Africa, from a depth of 2500 fathoms. The whole animal had probably a length of 4 inches; the head and
a piece of the body, however, were the only things we got. These fragments were very remarkable for the vivid
colours which they displayed: the head was yellow, the collar bright red, and the body of a yellowish red. Of this,

the Neapolitan species and the one which I discovered in the sand near Copenhagen, in shallow water, show nothing,
as they are uniformly white; a priori one would have been inclined to suppose just the contrary.

F2

30 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

The palpus of the first maxilla (2) and the free dorsal shield (1) are both characters
which, among Schizopods, we only find in Nedalia, a genus which has hitherto had a very
isolated position, and was considered a connecting link between these and the Phyllopods.
In Plate VI. fig. 9, we have figured the first maxilla with its long palpus in our Nebalia
longipes. ‘The palpus there is a mere continuation of the maxilla; here, however (where
it has very little resemblance to the Nedalia palpus), it seems to be in articulation
with the maxilla. This is a character which I have hitherto only found in G'nathophausia,
and confirms my opinion that this genus is, next to Mebalia, the most aberrant one among
the Schizopods.

In Lophogaster as well as in Guathophausia the dorsal shield is of a parchment-like
consistency. In the former its posterior and lateral edges are free, but the rest of the
shield forms the integument of the pereion-segments. A free dorsal shield has hitherto
never been found in a true Schizopod, but is also peculiar to Nebalia. Now Nebalia is
considered to be an aberrant Schizopod with nine abdominal segments ; and it seems to
me to be very interesting for the explanation of the semidivision of the last pleon-segment
in Gnathophausia, that this genus has two other phyllopodal characters, in the configura-
tion of its dorsal shield and the presence of a palpus on the first maxilla; but in all the
main points Gnathophausia is much more a typical Schizopod than Nebalia, and, I think,
not even closely allied to it. Probably both derive their origin from a series of animals
which were neither Schizopods nor Phyllopods, and to which possibly belonged the fossil
forms Dithyrocaris, Ceratiocaris, &c. I do not know how far these forms are really allied
to Gnathophausia, as I have not the literature on them on board; but I strongly suspect
that also on them some light may be thrown by the discovery of this interesting genus.

The Characters of the Lophogastride,

therefore, ought to be modified in several points; the following characters ought to be
added to the family-description :—connexion of the shield with the segments of the pereion
complete or incomplete ; with or without accessory eyes on the second maxilla; with or
without a palpus on the first maxilla; and with or without subjoints in the pereiopods.
- Besides, all the family characters given by Sars are applicable to both genera.

Characters of the Genus and Species.

GNATHOPHAUSIA™, n. gen.

Carapace in no connexion with the five posterior segments of the pereion; a long
rostrum ; first antenne with a short and stout pedunculus and two flagella, a long and
a shorter Inner one; second antennee with an oblong squamiform appendage ; stout
mandibles, with a three-jointed palpus; a palpus on the first maxille; accessory eyes,
palpus, and flagellum on the second maxillze; maxilliped showing nothing particular.
First gnathopods terminated by a claw; all the gnathopods and pereiopods leg-like, the
latter showing subjoints on their penultimate joint. Six pairs of gills at the base of the
gnathopods and the four anterior pereipods ; two branches of the arborescent gills laterally

* From yvalos and gavavos.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 37

covered by the carapace, one branch turned inside between the legs and in free commu-
nication with the water. Well developed pleopods. Caudal appendages jointed.

1. GN. GIGAS, 0. sp.

Scale of the second antennz with five spines, not jointed; short spines at the lateral
angles of the dorsal shield, no spine in the midst of the upper posterior margin.
_ Length 142 millims. Colour red.

2. GN. ZOBA, 0. sp.
Scale of the second antennz with only one spine, jointed ; dorsal shield with a sharp
spine in the midst of its upper posterior margin, lateral posterior angles rounded.
Length 59 millims. Colour red.

3. GN. GRACILIS, n. sp.

Scale of the second antennz with one spine; dorsal shield with a spine in the middle
of its upper posterior margin, and two spines in its anterior as well as in its posterior
lateral angles.

Length 41 millims. Colour red.

2. On CHALARASPIS, the Type of a new Schizopod Family. (Plate VIII.)

This little Crustacean is the commonest Schizopod of the deep-sea fauna, and seems to
enjoy a very wide bathymetrical and geographical distribution ; for we got it from depths
of 350-2500 fathoms, off the west coast of Africa, as well as off the east coast of South
America, down to the 35th degree of southern latitude. Whenever in the mid-Atlantic
true deep-sea animals came up in the dredge or the trawl, I was sure to get at least a
fragment of this Crustacean. Unfortunately most of the specimens were spoiled ; only a
few males and females were in a good state of preservation. The animal is so soft that it
does not stand the long passage of the trawl through the water; and even if it is not
broken to pieces, its carapace is usually overturned. On two occasions, however, I got
specimens which are quite intact; and these, as well as the many fragments, furnished
ample opportunity of studying this genus in all its details.

The females of this shrimp have a length of 35 millims., the carapace having from
above a length of 8 millims. and laterally of 12 millims. The males are 37 millims. in
length, but somewhat more slender than the females; their dorsal shield has the same
length from above as that of the females, but is laterally somewhat shorter (114 millims.).

The carapace is very soft, especially in the males, and in connexion with only the first
segments of the pereion. It has a very short, rounded rostrum (fig. 1 a), and on the
frontal border a pair of short spines underneath the eyes. The lateral anterior border is
rounded in the female, and somewhat angular in the male. The posterior border of the
carapace shows a deep excision, especially in the male, so that its rounded lateral borders
cover a great deal more of the pereion-segments than the upper margin (fig. 1). The
carapace is divided by a longitudinal ridge and two transverse sulci into several regions,
which in old females are very well marked; in the males, however, they are not always
plainly visible.

38 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

Underneath the short rounded rostrum, between the eyes, there is a small tubercle
(fig. 1 a, 7) which in the males is somewhat more prominent than in the females.

The eyes have the same position as in Gnathophausia, and are 2 millims. in length.
They cover somewhat the first joint of the funiculus of the first antenne, which consists
of three joints, the last of which has a lateral prolongation (fig. 2). Between the third
joint and the insertion of the flagella a small subjoint is to be seen. The external fla-
gellum is much bigger than the internal. Hairs are in both sexes at the inner side of
the funiculus, the size of which is also very much the same in both. The scale of the
second antenna is jointed and ovoid. The slender funiculus has three joints and a
flagellum, which exceeds in length the whole animal.

The wpper lip (fig. 11) is cordiform, with the point directed upwards. The mandibule
(fig. 4) are not particularly strong, and have a three-jointed palpus, the second joint of
which is somewhat bent backwards, and the third rounded at the top.

The excision between the two pieces of the under lip (fig. 5) is not so deep as in Gna-
thophausia. On its anterior border we find the same hairs which may be seen in the
Lophogastride.

The first mazilla (fig. 6) has two manducatory processes: the upper one has many
strong teeth ; while the under one has only a few hairs, no denticulation.

The second maxille (fig. 7) have a very elongated flagellum (/1) and a small two-jointed
palpus (7).

The maxillipedes (fig. 8) have a very large palpus and a flagellum on their enlarged
basal joint. They are terminated by a small denticulated claw.

The gnathopoda and the first pereiopod in this genus (which hitherto has not shown so
very great differences from Lophogaster) are not leg-like, but have all the shortened mawil-
lipedal form and function (figs. 1,9, & 10). This is the same in both sexes. The branchize
and (in the female) the breeding-lamelle begin at the base of the first gnathopod. The
form of this appendage (fig. 9) can best be seen from the figure. The terminating claw is
not denticulated ; the second is somewhat longer than the first, but shows no peculiarities.

It is very astonishing that in both sexes of this genus the first pereiopod has the same
maxillipedal form and function as the gnathopoda. This is the case in no other Schizopod
family. Also the form of its various joints (fig. 10) is very much like that of the gnatho-
pod-joints.

Among the four remaining pereiopods we have again two very different forms: three
of them are enormously elongated (figs. 1 & 12; they have a length of 27 millims.) and
terminated by a strong and recurved claw, which has many spines at its inner side,
whereas the last pereiopod (fig. 1, p*, & fig. 18) does not end by a claw, but by a simple
rounded joint, is very much shorter than the three preceding leg-like pereiopods, and all
over covered with hairs.

There are arborescent branchie at the base of all the gnathopoda and pereiopoda with
the exception of the last one. ach pair of branchiz consists of three branches, two of
which are laterally covered by the carapace; the third branch is bent inside between the
appendages, and is accordingly in free communication with the water. In this respect
there is no difference between this genus and the Lophogastride.

CRUSTACEA FROM THE ‘CHALLENGER’ EXPEDITION. 39

There are seven pairs of breeding-lamelle at the base of the gnathopoda and pereiopoda,
increasing in size towards the end of the pereion. This also is quite the same thing as
has been seen in the Lophogastride.

The abdomen of Chalaraspis differs from that of the family just mentioned especially
by the absence of the pseudo-segment. The last segment of the pleon is elongated and
perfectly smooth.

The pleopods are very perfect; they have a strong basal joint and two rami. They
offer nothing particular either in the male or the female.

The posterior borders of the quadrangular ¢e/son show a slight denticulation (fig. 1,0) ;
the outer appendages of the caudal fin are jointed.

The males differ from the females only by being somewhat more slender and by the
absence of the breeding-lamelle. The small tubercle underneath the rostrum seems to
be more prominent in the males than in the females.

Systematic Position of the Genus.

Among the three families of Schizopods (Myside, Huphausiide, and Lophogastridz)
hitherto known, this new form no doubt approaches most the latter, with which it
has in common the position and shape,of the branchiz, the breeding-lamelle, and the
pleopods; but the presence of only four leg-like appendages, and the fact that four
pairs of appendages act here as maxillipedes, do not allow us to class it together with
them, as here both genera (Lophogaster and Gnathophausia) have got seven pairs of leg-
like appendages. It depends, of course, on what one considers the main point of view
from which a classification of the Schizopods ought to be made, whether from the form
and position of the branchize, or from the number of appendages which are transformed
to maxillipedes. I think the best plan is to carry on the hitherto accepted principle—to
establish different families for those groups in which there is a different number of leg-
like appendages. One should only deviate from this principle in case an animal should
be discovered which in every other respect is so closely allied to a certain group that the
conformation of the legs appears only as a very secondary acquisition ; but from what
we have hitherto seen of the steadiness in which the same number of mazxillipedal
appendages is kept in forms which differ so much from each other as Guathophausia
does from Lophogaster, or Petalophthalmus from Mysis, I doubt very much whether such
forms will ever be found.

For the genus Chalaraspis one ought therefore to establish a separate family, Chala-
raspidze, holding its place between the Euphausiidz and Lophogastride, and characterized
(until further discoveries show whether it has got several living members differing from
each other) by the

Characters of the Genus.
CHALARASPIS*, n. g.

Dorsal shield or carapace in no connexion with the posterior segments of the pereion ;
rostrum short, rounded ; eyes present; no accessory eyes ; first antennze having a stout -

* XaXapos and doris.

40 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

funiculus and two flagella; scale of the second antenne jointed; upper lip triangular ;
mandibule with a three-jointed palpus; no deep excision in the under lip. Second
process of the first maxilla without denticulation ;. second maxilla with an elongated
flagellum and a small palpus; maxillipedes terminated by a denticulated claw, with large
palpus and flagellum. Gnathopoda and first pereiopod used as maxillipedes, shortened
and recurved; second, third, and fourth pereiopod having nearly the length of the body,
terminated by a long denticulated claw ; last pair of pereiopods shorter, terminated by a
rounded joint, very hairy, and without branchie. Three branches of branchiz on the
base of the gnathopoda and first four pereiopoda, two of which are covered by the cara-
pace. Breeding-lamellze on all the gnathopoda and pereiopoda. No pseudo-segmentation
on the sixth segment of the pleon. External caudal appendages jointed.

CHALARASPIS UNGUICULATA, 0. sp.

Length 35-37 millims.

Mode of Life, and Colour.

We have already seen at the beginning of this description that this is the commonest
deep-sea Schizopod with as wide a geographical as bathymetrical distribution. It lives
together with the same fauna as Gnathophausia zoéa, which several times came up
accompanied by Chalaraspis. Its colour is a bright red.

3. On PETALOPHTHALMUS, a Mysidiform Schizopod. (Plate VII.)

In Prof. Wyville Thomson’s ‘Depths of the Sea,’ p. 176, there is a quotation from a
preliminary note of the Rev. A. Merle-Norman, according to which specimens of Ethusa
granulata, a brachyurous Decapod, taken in 110-370 fathoms, are apparently blind, but
have two remarkable spiny eye-stalks, with a smooth rounded termination, where the eye
itself is ordinarily situated. In other specimens of the same crab the rostrum disappears
entirely, and the two eye-stalks, approaching each other, assume its functions. In
accordance with the altered conditions of life, the eyes and their stalks seem to undergo
in Hpethusa the most extraordinary modifications.

The same thing which now-a-days happens to a crab living in different conditions of
life has evidently also happened (but in the course of ages probably) to a Schizopod, in
which we find at the top of the eye-stalks, instead of organs of vision, two flat spherical
terminations in which no trace of an eye (which one would expect to find in this place)
has been left. But this peculiarity in the Schizopod seems not to be a modification of
individuals exposed to certain conditions of life; for we got specimens from very different
depths (1590-2500 fathoms) and latitudes, from the mid-Atlantic near St. Paul’s Rocks
down to the southern regions of Tristan d’Acunha, in both sexes of which the conforma-
tion of the eye is exactly the same. Though in the whole these Schizopods approach the
eenus Mysis, we shall find that they have several peculiarities which do not allow us to
suppose that this is a Jfysis which, like the above- mentioned Lyethusa, has changed
when exposed to certain conditions of life, and of which specimens might exist now-

a-days in full possession of their organs of vision. Once, of course, this process must
have taken place, but probably at a time when such Schizopods as Gnathophausia and

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 41

Chalaraspis were the only representatives of the order, from which only in later periods
of the earth the well-known shallow-water members of the Schizopod families have
developed.

T have called the animal which I shall now describe Petalophthalmus, and the species
P. armiger. The first specimen, a female, was caught by the trawl in lat. 2° 25’ N.,
long. 20° 1’ W., about midway between Cape Palmas and St. Paul’s Rocks, from a depth
of 2500 fathoms. We afterwards got a male in lat. 1° 22' N., long. 26° 36’ W., about
170 miles east of St. Paul’s Rocks, from 1500 fathoms, and, finally, a female in lat.
35° 41' S., long. 20°55' W., from 100 fathoms, about 400 miles west of Inaccessible Island
(Tristan d’Acunha group). The male has a length of 37 millims., the largest female of
44 millims., and the smaller one of 30 millims. .

The carapace, soft and parchment-like, is in no connexion with five posterior segments
of the body. In the female, where it has not quite one half of the total length of the
animal (it is 18 millims. in length in a specimen of 44 millims.), it covers in its ordinary
position nearly all the segments of the pereion. It has a small rostrum (fig. 1), which
is very little prominent, a small spine at its anterior border underneath the eye, and
sharpened anterior angles. The posterior and inferior angles are rounded; and the sur-
face of the whole carapace is perfectly smooth. A slight transverse sulcus divides it into
an anterior and a posterior portion, the latter of which is by far the largest.

Underneath the rostrum are the eye-stalks, terminated by concave spherical plates
having a diameter of 3 millims.; they are simply a duplicature of chitinous matter. On
examining them under the microscope I could not find a trace of any eye-like structure.

The first antenne have a three-jointed funiculus (I consider the small joint in fig. 3,
which is not to be seen in the first antenna of the male, to be not a fourth, but an ex-
ternal pseudo-joint) and two flagella, the external of which is enlarged at its base and
covered with hairs at the inner side.

On the second antenna (fig. 4) there is a lamellar appendage (fig. 5) which has no
spines.

The labrum is subcordiform (fig. 6). The mandibule (fig. 7) have a very strong
manducatory portion, with many denticulations and stiff hairs. The second joint of their
palpus is enlarged; the third has a rounded point and many strong hairs at its inner side.
The under lip is somewhat like that of Chalaraspis, and bordered anteriorly with small
hairs. I could not draw it, as I saw only a part of it, the organ being broken when I
tried to take it out.

The first maxille have two processes, showing many hairs and a strong denticulation
(fig. 8); they have no palpus. The second mazille (fig. 9) have both palpus (p) and fla-
gellum (fi). The mawilliped (fig. 10) has a much more perfect palpus, quite close to its
flagellum, than Grathophausia has; besides, there is at the inner side of the third joint
one of those flat lamellar appendages which are known in the maxilliped and first ena-
thopod of Mysis and Siriella, and which in the male of our genus are indeed present
on both appendages, in the female, however, only on the maxilliped.

The first gnathopod (fig.11) has still maxillipedal functions, though it is rather elon-
gated; its penultimate joint, however, is enlarged and recurved, and terminated by a

SECOND SERIES.—ZOOLOGY, VOL. I. G

42, . DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC ~

rounded claw, which fits into an excision at its top. The second gnathopod (fig. 12) is
perfectly leg-like. Its tarsus, like that of the following pereiopoda, is not subjointed, as
in Wysis and Gnathophausia, though there are hairs at intervals on it as in those Crus-
taceans.

The five pereiopoda are all very much alike; only the fifth is somewhat shorter than
the rest. I have only drawn the third (fig. 18).

At the base of ad2 the gnathopoda and pereiopoda we find breeding-lamelle (fig. 1, and
figs. 12 & 18), the posterior ones nearly as long as the legs. This is a character not
found in any other Mysids, and, among all the Schizopods, hitherto only known in the
Lophogastridze.

The rings of the pleon are perfectly smooth, and do not present any thing particular.
The sixth segment is longer than the rest. The five pairs of pleopoda are only rudi-
mentary and exceedingly small (fig. 14); they consist of a small basal joint and only one
ramus. The last ones, especially the fifth, are longer than the first.

_ The ¢ail (fig. 1) is formed by the telson, which shows a little excision and two appen-
dages on each side, the exterior one of each pair jointed.

Branchie are entirely wanting.

The male differs very much from the female. Its soft and free carapace is much shorter,
not much more than + of the total length; for in a specimen of 37 millims. it has a
length of 10 millims. Thus the last two segments of the pereion are entirely uncovered ;
while several of the preceding ones are laterally uncovered, the upper part of the cara-
pace being much longer than its lateral parts.

Very extraordinary are the prehensile organs of the male—the first antenna, the pal-
pus mandibularis, the maxilliped, and the first gnathopod—altogether forming a for-
midable prehensile apparatus (fig. 2).

The funiculus of the first antenna is enlarged, and all the joints are elongated; it
has a length of 10 millims. Its flagella, however, are of ordinary size, the exterior
one being not enlarged at the base. The second antenne (fig. 2, a’) do not differ
from those of the female. The principal prehensile organ is the palpus mandibulze
(fig. 2, pm), the second and third joints of which are enormously enlarged, and the third
armed with strong sete. The maxilliped (fig. 15) and the first gnathopod (fig. 16) have
enlarged joints, and are terminated by strong claws. At their inner side is the lamellar
appendage (figs. 15 & 16, da), which we found in the female only in the maxilliped.

Unfortunately the only male which we caught has lost the second gnathopod and
the first pereiopod. The second pereiopod is terminated by a tuft of hairs instead of a
claw (fig. 2, prp?). The three last ones do not differ from those of the female, except by
being somewhat more slender.

In the male the pleopods are more perfect (fig. 17); for, besides being stronger, they
have two rami, one of which is broad and bordered at its rounded top with hairs, the other
pointed and nearly filiform. The last pleopoda are longer than the preceding ones.

The telson of the male (fig. 2@) does not show such a deep excision as that of the
female.

No traces of branchiz are to be seen.

CRUSTACEA FROM THE * CHALLENGER’ EXPEDITION. 43

Surely these characters of the Podophthalmus males ave quite exceptional among Schi-
zopods, and much more like what we find in certain ge in than what we should
expect to find among the higher Crustaceans. i

Systematic Position of the Genus.

-The six pairs of leg-like appendages, the rudimentary condition of the abdominal feet,
and the absence of the branchiz assign to this Schizopod its place among the Mysidee. It
differs from them only by the free dorsal shield and the presence of breeding-lamellz on
seven appendages, which it has in common with the Lophogastridee. It is also sur-
prising to find that a member of this latter family has a Mysid character (the subjoints
of the legs in Gnathophausia), which, however, is only a peculiarity of one of its generic
groups (genus Mysis and subgenera). It accordingly stands in the same relations to
Myside as Gnathophausia stands to Lophogastridee—there being in both a character
which shows some distant relationship to the other group, and both having a free cara-
pace, which is not to be found in the hitherto known Schizopods.

Among Myside there are two generic groups :—Wysis and the genera connected with
it, characterized by joints in the tarsal parts of the legs and the absence of branchial
appendages on the pleopoda; and S%riel/a, in which the legs are simple and terminated
by a claw, as in our case, but in which there are branchial appendages on the pleopoda,
which Petalophthalmus has not got. This genus therefore belongs to none of these
generic groups, and can only be united with the Myside if the family characters are so
far enlarged as to allow the entrance within it of an animal which has a free dorsal
shield and breeding-lamelle on seven appendages.

The eyes having disappeared, and the eye-stalks having assumed a spherical termi-
nation, are of course entirely secondary characters, and probably not even of generic
value.

Characters of the Genus.

PETALOPHTHALMUS*, nov. gen. 4. .

Carapace in no connexion with the five posterior segments of the pereion; rostrum
very short; three joints in the funiculus of the first antenne; scale of the second
antenna without spines; labrum subcordiform ; mandibule having a large manducatory
process and a three-jointed palpus. Very strong teeth and setze on both processes of the
first maxilla; second maxilla consisting of a manducatory portion (two elongated pro-
cesses and a larger lamellar one), a ciliated flagellum, and a palpus. Maxilliped in
both sexes with a lamellar appendage on one of the basal joints ; first gnathopod having
the same only in the male; in the female it is elongated, and its last two joints are
clumsy and recurved; second gnathopod and the five pereiopoda leg-like; pleopoda in
the female quite rudimentary, with only one ramus; in the male somewhat more per-
fect, with two rami; telson showing a slight excision at the top; outer appendages
jointed.

’ No branchiz on the pereion, nor any branchial appendages on the pleopoda of the
male. Seven pairs of breeding-lamelle at the base of the pectoral appendages.

* Tléradoy and d¢Oadpos.
G 2

4A, DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

The male differs from the female by the rudimentary condition of its carapace and the
metamorphosis of the first antennz, the palpus mandibularis, the maxilliped, and first
gnathopod into prehensile organs.

PETALOPHTHALMUS ARMIGER, N. Sp.
Eyes wanting. Eyestalks with spherical concave terminations at the place where one
would expect to find the eyes. Length 37-44 millims.

Habits of Life, and Colour.

Petalophthalmus is a member of that deep-sea fauna which we sketched when de-
scribing Gnathophausia zoéa, in which, however, it seems not to be common. Its colour
is a bright red.

We have now described (1) a lophogastriform genus (Guathophausia), with three
species, (2) a genus allied to the former, but forming a new family (Chalaraspis), and (3)
a Mysidiform genus (Petalophthalmus)—all of which have a free carapace and certain
peculiarities which show that their existence dates from a period in which the family
characters of Schizopods were not so sharply defined as they are now. They are pre-
cursors of the Myside and Lophogastride, and representatives of another family, having
characters which, among the Schizopods, are only shared by Nebalia, but in all other
respects entirely different from this genus. Now the idea presents itself whether it would
be advisable to bring all these Schizopods together into one group, the chief and only
universal character of which would be that they all have a free dorsal shield. We could
best describe this in the following synoptical way :—

(9 abdominal segments, palpus maxille prime ..................-. NEBALIA.
( Tlegs, subjoints, palpus maxille prime, supple-
Group I. | mentary eyes, Lophogastriform ............ GNATHOPHAUSIA.

Dorsal shield free. < :
(aleilomuael Seg 1) g legs, no subjoints, 7 breeding-lamelle, Mysi-

BEEN: COOH cconcconoas0oucc08 Pacgansodmeooas PETALOPHTHALMUS.
\. 4 legs, no subjoints, branchiz as in Lophogaster.. CHALARASPIS.
C (7 legs, no subjoints, no palpus maxille prime,
Group II. no supplementary eyes...............----- LopHogastER.
ove elite 228 7 abdominal seg- } 6 legs, with or without subjoints, 2 breeding-
tened to the )
ments. lena samemiaaie s ucian digit eee ee ore ooo Sole Mysis and Srrierra.

pereion.

8 legs (the last wanting or rudimentary), branchiz
| free, supplementary eyes...........-.....-. KUPHAUSTA.

This arrangement of the Schizopods would, no doubt, have a certain advantage, as it
prevents the destruction of the family characters of our old and well-known forms in
favour of a few deep-sea genera, all of which have a certain peculiarity which they share
only with Nebalia. But it is not a natural genealogical arrangement; for, from what
has been said, there is no doubt that Gnathophausia is, if 1 may use the expression, an
ancient Lophogaster, and Petalophthalmus an ancient Mysis. I therefore propose to

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 45

unite these with the families of the shallow-water forms, and would suggest for the
Schizopods the following systematic arrangement :—

9 abdominal segments, 8 legs ...........-.- 02 -eeeer creer eens I, NeparupZ.
Ghlogs cee opersnctsboieyceslsversats ckovche etevathu ay before IL. Mysinz.
: Sep Sk eectausiegsscvsrssveiai ness ler Syaueuaveos'cseiako eleyais Ill. EvpHavsiipz.
7 abdominal segments, A NEBR hte sisheccyoreiae forse celeaies sare wate ate IV. CaanarasPipz.
MUG RS rviareys ae cools eee Sera lan betel wae VY. LopHogasrripx.

In order to expose the genealogical connexion between the genera of these families,
it would be necessary to prepare a large table; but I think that their relations will be
well understood from what I have said about the subject in the preceding pages, as well
as from a comparison of my plates with the figures given on the other genera by Sars,
Claus, and other authors.

Two questions will arise, of course, from a comparison of the deep-sea forms with the
shallow-water forms:—(1) Are there no Schizopods in’ the depths which show the
ordinary conformation of a shield fastened to the pereion? and (2) As the Mysidee and
Lophogastride have relations of very extraordinary characters living in the deep sea,
are there no forms which represent the third of the hitherto acknowledged families—the
Euphausiide? These questions are partly answered by the discovery of a very large
member of this family, which came up in the trawl in lat. 35° 41’ S., long. 20° 55’ W..,
from a depth of 1900 fathoms. This animal is a male, has a length of 84 millims., and
belongs to Dana’s genus Euphausia; for its dorsal shield is perfectly fastened to the
pereion, and of the eight pairs of legs only six are developed, the last two being only
represented by the branchial appendages and the palpi. But, unlike all other species
of Huphausia hitherto known, it has no accessory eyes. There are very long olfac-
tory hairs on the enlarged first antennze, and the peculiar appendages to the pleo-
poda which characterize the males of this genus. I shall figure and describe this
species, which I intend to call Huphausia simplex, n. sp., when we come home, and
when it will be possible to compare it with the other species of the genus (which have
been described by Dana and Claus). From the discovery of this Huphausia, it appears
that not all the deep-sea Schizopods offer the said peculiarity, and that, among very
peculiar genera such as have been described above, a form is also to be found which
differs from the surface-Huphausie only by the absence of accessory eyes. ‘These, I
think, are to be considered a secondary adaptation to pelagic life; and therefore 2. sim-
plex is probably a more ancient form than #. Wiilleri, splendens, and superba. Probably,
however, this will not be the last discovery we make in this interesting group; and I
think it is very likely that a form will come up which is in the same relations to the
Huphausiide as Gnathophausia is to the Lophogastride, and Petalophthalmus to the
Mysidez. Then we should have made one step forward in the knowledge of Schizopodal
genealogy, a step which will probably lead to further discoveries concerning the rela-
tions of the Phyllopods to the Schizopods—of the lower to the higher crustaceans.
Perhaps it will now also be possible to understand certain fossil forms which seem to
be allied to Nebalia and to Gunathophausia.

46 . DR. R. V: WILLEMOES-SUHM ON SOME ATLANTIC,

V. On-THE DEVELOPMENT OF A LAND-CRAB.
(Plate XI. figs. = 3:

In most books on natural history we find concerning the land-crabs a remark that,
according to Mr. Westwood’s observations, they migrate to the sea when the time for de-
positing their eggs has come, and that the young ones, when coming out, have the same
form as their parents. The same thing isto be observed in Astacus fluviatilis; and the
knowledge of these two facts has been much in the way of the general acceptance of
Mr. Vaughan Thompson’s first observations on the Zoéa-stages of crabs. Also his state-
that he found Zoéa-brood in the eggs of a Gecarcinus has been doubted ; for, according to
Fritz Miller *, Mr. Bell considered himself justified in eliminating Thompson’s observa-
tion, because he could only have examined ovigerous females preserved in alcohol. The
following lines, however, will show that there is no reason to doubt Mr. Thompson’s
statement. Fritz Miller himself, though we owe him many observations on land-crabs,
says only that there is Zoéa-brood in Ocypoda and Gelasimus, but had no occasion to
observe the Gecarcinoids.

Watching the habits of these crabs is always attended with some difficulties, as De
of them are nocturnal. In St. Thomas I only succeeded in getting a few Gelasimi.
Only in the Bermudas did we get Gecarcinoids:—the large Cardisoma guanhumi,
which was caught by torch-light in the interior of the island, but all the specimens we
got were males ; besides, Mr. Moseley caught Gecarcinus lateralis and Ocypoda rhombea,
and I myself caught several crabs allied to Boscia. 1 watched there also for a long time
the lively Grapsus cruentatus, which lives in great quantities in Hungry Bay, among the
mangrove trees. It has entirely taken to terrestrial habits, makes deep holes and runs up
and down the mangrove trees. ‘he holes, however, it always makes so near the shore,
that it is sure to find water at a depth of 2 or 3 feet. What its mode of development
is I could not find out, as it was evidently not their breeding-season (June). Arriving
in the beginning of August at the Cape-Verd Islands, I found in St. Vincent many holes
on shore inhabited by Ocypoda hippeus, a crab running with marvellous velocity over the
sand, and very valuable to me, as it gave me ample opportunity of studying the interest-
ing entrance to the branchial cavity described by Fritz Miller. But also in these I could
discover no trace of eggs or of young. Holes: made by the same crab we found
again in the island of San Jago, where they are to be seen in abundance near the
houses of the little village which surrounds the old cathedral of Ribiera Grande. But
besides Ocypoda hippeus, we got in San Jago another large land-crab, caught by Mr.
Moseley’s seining-party at night in the bay of Porto Praya, a true Gecarcinoid, be-
longing to the same genus (Car disoma) to which the Bermuda ones belonged. Unfor-
tunately I cannot make out its specific name, as it is not mentioned in Milne-Edwards’s
‘ Histoire Naturelle des Crustacés’ +; but I suppose it is an animal well known to Euro-
pean carcinologists as inhabiting the Cape-Verd Islands, and, very likely, also the coast

* «Facts and Arguments for Darwin,’ by Fritz Miller (translated by W. 8. Dallas. London: 1869), p. 48.
+ Probably described in the same author’s ‘ Crustacés des fles du Cap Vert.’

CRUSTACEA FROM THE ‘ CHALLENGER” EXPEDITION. AT

of the African continent. This Cardisoma inhabits large holes all along the shore,
especially in the palm-groves on both sides of the town. But when I went there the next
day to catch more of them, the difficulties in getting them were too great; for the holes
were more than 8 feet deep. The one, however, which Mr. Moseley had caught, a
female, proved to be very interesting; for after having killed it in spirits, I found the
next-day that its abdominal feet were covered with strings of eggs, that most of these
eggs were empty, some of them containing an embryo, and that among the empty eggs
there was a good number of young animals newly come out. These young ones, how-
ever, were not like their mother, but Zoéas. The greater number of them had evidently
left the mother already; some were still to-be found among the empty capsules; and
others had not yet emerged. In fig. 3 Mr. Wild has drawn me sucha string of eggs
(each 0°42 millim. in diam., having the form of a grape), in which the berry-like
peduneulated eggs surround a common axis.

The Zoéa of Cardisoma (fig..1) leaves the eggs in a somewhat more advanced state
than the Zoéa of Carcinas menas. If we compare it with the figures given by Mr. Spence
Bate (‘Philosophical Transactions,’ 1858, pl. xl.), we find that it represents a middle
stage between the larva of Carcinus menas which has just left the egg and the one
after its first moult (figs. A and B). With the former it has in common the want of the
frontal spine, with the latter the presence of setze on most of the appendages, and the
more developed caudal fin.

The carapace is very globular, and its dorsal spine is only small. The frontal spine,
entirely wanting now, is probably only developed after the next moult. The eyes are
very large. I could not see the pedunculus; but as I had only dead animals before me,
which had been for some time in spirit, I do not deny that a very short pedunculus
might exist. The first antenna has a few more hairs than in the second stage of
Carcinus menas; but the second ones are very much like each other in both cases (a'
and @’, fig. 1). The parts of the mouth (m) had equally already got their hair. I had
some difficulty in getting a side view of them, and could have only isolated them with
very great difficulty, which was not worth while, as there was no chance of getting
later stages of this larva. The two gnathopoda have four long hairs in the last joint,
which in our case (gp' and gp’) are not so long, and not feathered, as they are in the
second Zoéa-stage of Carcinus menas. I have not seen any buds of the pereiopods
behind those two legs, which are present in the young Carcinus menas.

Each segment of the pleon presents some black pigment spots, some of which were
also-observed in the carapace. There are no appendages on the pleon. The last segment
is bifurcate; and at the inner side of the two terminal pieces six feathered setae may be
observed, very much like those in fig. 21” of Spence Bate’s larva.

I think there is no doubt that these Zoéas, as they partly had done already, leave the
mother and lead a pelagic life until they have undergone all their metamorphosis ; and
IT hope that in other parts of the world which we are about to visit, I shall be abie to
complete these observations on the propagation of the land-crabs, and to make out in
which genera there is a metamorphosis, and in which not.

I may add here an observation which I made some years ago in Italy, and which,

48 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

if not quite directly concerning our subject, at least touches it; for the animal I am
to speak about is the freshwater crab of Southern Europe, Zelphusa fluviatilis. In the
mountains above Spezzia I caught some specimens at a spring coming out from the Bocca
Lupara; and one of these being a female, carried with it nearly 150 young ones under
the abdomen, the smallest of which had a length of 3 millims. I think this shows clearly
that also Telphusa is, with Gecarcinus and Astacus, one of the few Decapods which
develop directly without metamorphosis.

VI. ON A BLIND DEEP-SEA ASTACUS.
(Plate X. fig. 1.)

In a very successful haul on the 15th of March near Sombrero Island, W. I., we got,
from a depth of 450 fathoms, a blind Astacus, one of the claws of which is developed to
an extraordinary extent. In the notes which I gave about it to Professor Thomson, I
have called this animal Astacus zaleucus ; for, strange as it may seem at first sight, there
is no reason for separating this animal from the genus Astacus, except perhaps the want
of the eye andits pedunculus. The male of Astacus zaleucus has a length of 110 millims.,
the cephalothorax having a length of 50, the abdomen of 60 millims.

The carapace is laterally very much compressed, and divided by a deep transverse
sulcus into two portions or regions. At the top of the anterior one is the rostrum,
having a length of 9 millims. Its borders have six small teeth. Before the rostrum there
are two large triangular patches, which are densely covered with short hairs; and these
continue from both sides towards the rostrum. On both sides of the anterior portion
there are besides several small spines, and two sulci, which establish again a small mar-
ginal and triangular subdivision. The posterior portion of the carapace can also be
subdivided, into a quadrangular upper portion and two lateral regions. The last segment
of the pereion is in articulation with the carapace, quite in the same way in which it is
in all Astaci,

On both sides of the rostrum there are two qaean places, where in other Species the
pedunculus of the eye is fastened. Here there is not a trace of it.

The four-jointed funiculus of the antenna interior bears two very hairy flagella, of the
length of 40 millims, The base of the antenna exterior terminates on one side in a sharp
spine. The squamiform appendage is somewhat curved outwards, reaches the middle of
the second funiculus-joint, which is rather long, and has at the inner side eight sharp
spines, The flagellum of the antenna exterior has a length of 130 millims. Underneath
its base the large opening of the ‘‘ green gland” is clearly to be seen.

The margin of the upper lip is somewhat tilted upwards, and beset with 6 small
spines. About the mandible I have nothing particular to say. The chewing-plates of
the first maxillee are very strong, and the palpi of the maxillipedes very large. They look
nearly like a ramus of one of the swimmerets. The palpi of the two gnathopods,
however, have the ordinary form. The second gnathopod has thrice the length of the
first one.

The pereiopods are all very hairy, especially their first two joints, which, with the

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. . 49

exception of the right claw, are all densely covered with hairs. To three pairs there are
chele; the fourth has a small recurved claw at the end; and in the fifth the sixth joint
is nothing but a very small stump. Very remarkable are the first pair of pereiopods; the
chela of the right one has a length of 100 millims., and that of the left one of only
48. Between the two chels, however, there is not only the difference in length, but
also in width, as the carpus in the right one is very much expanded, spiny, and
hairless; in the left one it is even and covered with hairs and spines. The two digiti
chele have large teeth at their inner side, 57 on the digitus mobilis, and 62 on the oppo-
site one; so that the claws of the larger pereiopod look like the jaws of some ferocious
reptile.

The genital openings are here at the base of the third pereiopod, where we also found
them in the male of Deidamia leptodactyla.

Between the last pairs of pereiopods there is a small triangular pectoral plate.

The pleon is very flat; and the sharp angles formed by a longitudinal line on both
sides give it nearly a square shape. Its lateral pieces do not terminate in sharp spines,
nor are they very large; the edges of these, as well as the whole surface of the third
and fourth segments, are covered by dense and short hairs. The telson is a square, as
it is in many Astaci; its lateral appendages are very large, and extend, like wings, to
both sides.

The pleopoda of the first, very short segment are styliform appendages; the other
pleopods have a very solid unjointed base and two broad rami, the interior of which is
longer than the exterior (fig.1q@). All these, as well as the caudal appendages, are
densely fringed with hairs.

, Characters of the Species.
ASTACUS ZALEUCUS, 0. Sp.

Hyes and eyestalks wanting entirely. Chelee of the first pereiopod unequally developed,
the right one having more than double the length of the left one. Hight spines at the
squamiform appendage of the outer antenna. Genital opening of the male at the base of
the third pair of pereiopods.

Those who consider the eyestalk an appendage of a separate segment of its own
would probably be inclined to make another genus of this form. But I think this could
be the only reason for separating it generically from Astacus, and for me not a sufficient
one. We find in Deidamia that the genital opening of the male is, in one species, at the
third pereiopod, and in another, undoubtedly belonging to the same genus, at the fifth.
I therefore think that also this cannot be a reason for separating it generically from
Astacus, in which genus this species will be the first one made known as an inhabitant of
the sea.

To the Astacus of the Mammoth-cave this crayfish has only a physiological resem-
blanee; for even in those species in which the eyes are abortive the stalks at least are
present. But we have another form, which we got in deep water near the Bermudas,
in which the eyes have the same hidden position as in A. pellucidus, to which it has in
all other respects an extreme likeness; only in the marine species the squamiform ap-

SECOND SERIES.—ZOOLOGY, VOL. I. H

50 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

pendage of the outer antenna is entirely wanting. The same thing is the case in a
crustacean recently described by Mr. Wood-Mason from deep water near the Andaman
Islands, which he calls Nephropsis Stewarti*. Iam inclined to think that the crusta-
cean which we got near the Bermudas belongs to the same genus; but as I have not
seen the plate which Mr. Mason has added to his original paper, I shall defer its descrip-
tion until we come back.

Colour, and Mode of Life.

A. zaleucus came up on the swabs of the dredge, together with the large chele of
another, smaller specimen, the body of which was lost. The one we got had the red
colour of all deep-sea Crustacea. It lived on a bottom of globigerine ooze, evidently
frequented by a great many animals. Several sponges and seven echinoderms (Ophio-
musium, Luidia, Archaster, Astrogoniun, Oidaris, and Hchinus), a Planularia, Mopsea,
and an Jsis, several Annelids and a Sipunculus, a Galathea, a Peneid, an Arcturus, and
the fine crustacean to which was given the name of Willemoesia crucifera—four Bryozoa,
a Dentalium, and many small shells—finally, a fish probably belonging to the genus
Chauliodus, were got in the same place with this extraordinary Astacus. Unfortunately
our stay in the West Indies was only a very short one. The few hauls, however, which
we had near Sombrero Island and St. Thomas showed us that a great quantity of new
and probably also interesting animals live there in moderate depths (800-400 fathoms).

VIL. ON Witrmorsra, A DEEP-SEA DECAPOD ALLIED TO Hrvon.
(Plate XII. fig. 10, and Plate XIII.)

In the “ Notes from the Challenger,” sent by Professor Wyville Thomson to ‘ Nature,’
two blind crustacea have been described, both of which were found on our cruise from
Teneriffe to St. Thomas. The larger one of these remarkable animals was called by me
Deidamia leptodactyla, obtained on the 4th of March, by the dredge, in a most perfect state,
from a depth of 1900 fathoms, in lat. 21° 38’ N., long. 44° 39° W. Another specimen
was caught in the trawl in lat. 35° 41’ S., and long. 20° 55’ W., about 400 miles W. of
Inaccessible Island (‘Tristan d’Acunha), from the same depth at which the first was got.
The second, smaller species, which I proposed to call D. crucifera, was got on the 15th of
March, close to Sombrero Island, West Indies, from a depth of 450 fathoms, and equally
well preserved. I shall now give a description of both of them, but especially of the
larger one, which may be considered the type of the genus, and of the smaller one as far
as it offers any differences of importance.

The generic name Deidamia, however, must be dropped, as (according to Mr. Grote,
who has done me the honour of proposing for it the name Willemoesia) it was given
by Dr. Clemens in 1859 to a genus of North-American Sphingidee.

1. WILLEMOESIA LEPTODACTYLA.
The body of this crayfish approaches in form closer to the well-known fossil qa
than any other crustacean known to me, the body being flattened above, and there being

* “On Nephropsis Stewarti, a new Genus and Species of Macrurous Crustaceans,” by James Wood-Mason, ‘ Annals
and Magazine of Natural History,’ No. 67, July 1878.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 51

no lateral compression as in Astacus or in Palinurus. The cephalothorax exceeds in
width half of its length; and the abdomen too is very flat. The whole animal, being a
male, has a length of 120 millims.* (Plate XIII. fig. 1.)

The cephalothoraz has a length of 60 millims. by 33 in width at its base. Its upper
lateral edges are very sharp, and formed by a series of lateral spines, those of the under-
side, however, rounded. The whole cephalothorax, owing to this configuration of the
carapace, is square, the upper surface being wider than the under one, and somewhat
convex, the side parts being not rounded as in Astacus but perfectly flattened ; also the
upper surface forms a square, the angles of which are terminated anteriorly by two
spines, and are posteriorly rounded. A line of spines runs longitudinally through the
midst of the carapace, and divides it into two portions, being terminated anteriorly by
a sharp spine. The lateral spiny edges are divided by small but clearly perceptible
fissures into four portions, the first of them containing 8, the second 5, the third 5,
and the last 6 larger spines. A deep transverse sulcus divides the carapace into an
anterior and a posterior portion, which by secondary elevated lines are again divided
into several regions, which we might call the stomachal, hepatic, lateral, cardiac, and
branchial regions, each of which will be easily seen in the drawing (fig. 1). The inner
borders of the carapace of the under surface are perfectly smooth; and the rounded exte-
rior edges of the same surface are covered by a fringe of long and beautiful hairs. The
whole carapace is covered by very small spines, all directed forward, giving it a certain
likeness in appearance to the well-known chagrin of the sharks.

The eyes are entirely wanting ; nor is there, as in Astacus zaleucus, any place left open
where you might expect to find them.

Immediately underneath the frontal edge of the carapace we find in a prominent place
the wing-like expansions of the joints, funiculi antennarum anteriorwm. They are (espe-
cially at the inner side) densely covered with hairs, and very closely approach each
other. The funiculus consists of three joints, and has a length of 10 millims. Its in-
ternal flagellum, consisting of very many rings, has a length of 65; while the external
is very short, being only 10 millims. in length. On the same line with the inner antennee
and close to them, are the outer ones, the funiculus of which is a little more elongated,
and consists equally of three joints (12 millims. in length). The squamiform appendage
is very small (8 millims. in length by 2 in width), lancet-shaped, and inserted at the base
of the first joint. The flagellum is 60 millims. in length.

The opening of the mouth is angular (fig. 4); and you find above its rejected border a
small tuft of hairs. I could not find any labrum. The mandibule, covering nearly the
whole opening with their strong manducatory processes, have behind these a strong and
hairy palpus (fig. 5, pm), and at their base the two slender organs (fig. 5, Jab) which repre-
sent the labiwm or under lip. The first maxilla (fig. 6 a) shows avery small rudiment of a
flagellum (1), which in the second mawilla (fig. 6 b) attains so considerable dimensions.

The maztluped has two manducatory processes (fig. 7), a palpus (fig. 7, p), and a
flagellum, which is divided into two portions, the upper one (jl’) being attached to
the palpus, and the under one to the base of the maxilliped (/?).

* This description refers to the larger of the two males procured.

52 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

The first guathopod (fig. 8), as well as the second (fig. 9), are terminated by a denticulated
claw, and covered by hairs at their inner side. It is quite certain that the gnathopods
(or the second and third maxillipeds) have no palpi.

The pereiopods (fig. 1) are all terminated by chelee, but of very different sizes. The length
of the first pair, enormously developed, is 155 millims., thus exceeding by 35 millims. the
length of the whole body. The basal joint (the coxa) is very large and flat, the second,
however, very thin at its articulation, bent somewhat backwards, and increasing in size
only at its top, where it articulates with the broad base of the third joint. This is the
longest one, being covered on its upper surface by a series of small spines, and terminated
at its top articulation by two recurved spines. The fourth joint is again very slender;
and so is also the delicate chela, the immovable finger of which is armed with a large
transverse spine. This first pair of pereiopods are perfectly smooth ; all the others, however,
are covered with hairs up to the top. They are much smaller and clumsier, the second
pair having only a length of 39 millims. At the coxal joint of the third pereiopod
are the genital openings.

As to the underpart of the cephalothorax, it may still be mentioned that there is no
distinct opening for the so-called green glands, that there is a strong spine between the two
maxillipeds above the entrance of the mouth, and that an elevated line, covered with
a fringe of hairs, runs over the sternal shield between the first pair of pereiopods.

The abdomen (pleon) has a length of 60 mm., and consists of six segments and the
telson. The top of the second to fifth segments is marked by a strong spine, which is
bent forward, and which in the first and sixth segments is only represented by a small ele-
vation. There are also in the first segment no squamiform lateral appendages, which in
the subsequent ones are very large, but decrease in size from the second to the sixth. All
of them are fringed with long hairs; and so are also the telson and the caudal appendages,
the form of which can best be seen from Mr. Wild’s excellent drawing.

The first pair of pleopods is transformed to styliform appendages (fig. 2), our specimens
being both males. The four subsequent ones have a length of 33 millims., and are evi-
dently very powerful organs of locomotion. They consist (fig. 3) of a very strong basal
joint, to which the external palpus is attached, of smaller second joint, which goes
laterally over into the second interior palpus, and of a very small and styliform third
joint (fig. 3, a)—quite the same arrangement which we find in W. erucifera and in an un-
described Palinurid from the deep sea near the Bermudas (fig. 12). All the four pleopods
are alike; and the posterior ones are very little smaller than the anterior ones.

Of the internal organization I can only say that the carapace is wonderfully trans-
parent for the size of the animal, and permits the heart to be seen, which is situated
just between the hepatic and stomachal regions.

2. WILLEMOESIA CRUCIFERA, n.sp. (Plate XII. fig. 10, and Plate XIII. figs 10 and 11.)

The species we have hitherto considered is very delicate, and has very long fringes
of hairs on the legs, the carapace, and the pleon. All this is less the case in the
smaller species, which is much more solid, not transparent at all, and in which the
fringes are not so large and beautiful. Nevertheless also this one is a very elegant

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 53

creature, which on the whole has a strong family likeness to W. leptodactyla. It has
a length of 42 millims. x

The form of the carapace is nearly the same as in the former species. It has a length
of 19 millims., by 18 of greatest width; at its base, however, it is only 10 millims. wide.
The denticulation of the edges is a much stronger one than in the preceding species;
there is a line of spines divided by two incisions into three divisions, the first of which
contains 7, the second 5, and the third 17 spines on each side, all bent forward and
somewhat upward. Two very prominent lines, one longitudinal, ending at the front with
a spine, and one transverse, form the figure of a cross; and following the lines of
caruncles, you might trace the same regions which we have described in the former
species, but which in this case are not so clearly visible. At the frontal border there are
two spines on the right side and four spines on the left side of the median spine. The
whole carapace is finely covered by very small spines, which look like very fine chagrin.

The interior antennz have several spines on the inner side of their three funiculus-
joints, but not such a wing-like expansion as we have observed in the former species.
The exterior flagellum has a length of 17, the interior of 7 millims. The exterior
antenna does not offer any thing particular.

The parts of the mouth have exactly the same form as in the preceding species ; and
the absence of palpi at the base of the gnathopods can also be stated in this case.

Of the five pairs of pereiopods only four are terminated by chele ; and this is the chief
difference from W. leptodactyla. The first pair also are proportionally much shorter,
(only 34 millims. long), and have no spine on the slender fingers of the chela. In the
third and fourth pairs of pereiopods the chele are much more slender than in the first.
The last one is terminated by a simple recurved claw. The last four pairs of pereiopods
are covered with hairs, but not so densely as in WV. leptodactyla. The genital opening
is, in this species, not on the coxal joint of the third pair of pereiopods, but, as is usually
the case, on that of the fifth.

The abdomen has a length of 23 millims. Its first segment bears one spine; and all
the following ones have two spines on a little prominence. Telsen, caudal appendages,
and lateral processes of the segment are fringed with hairs.

The first pair of pleopods is transformed into two styliform and flattened appendages.
In the second pair (fig. 10) we observe the peculiarity that the third styliform joint is
covered by another, smaller joint, the morphological value of which I do not understand.
In the other three pairs (fig. 11) this supernumerary joint is not attached to the third one ;
so that these are very much like the pleopods of W. leptodactyla.

Colour, and- Habits of Life.

Especially the larger of these two species was very beautiful when it came up; for it
was of a fine red, while the hairs bordering it were yellowish. The smaller one also was
red. This colour, however, rapidly disappeared when the specimens had been for a
certain time in spirit.

As was also the case with Gnathophausia, the larger species of the two was caught im
very deep, the smaller one in shallower water. W. leptodactyla was found on a plateau

54 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

which probably extends southwards from Greenland, and on which in the north are
situated the Azores, and in the south Tristan d’Acunha. Before getting to it from
Teneriffe we had the greatest depth of that section of the ocean, 3150 fathoms ; and after-
wards again we gradually came down to a depth of 3000 fathoms before getting into the
shallower water of the West Indies. On that plateau we got with the first specimen a
sponge, some Bryozoa, a Brachiopod, a Lamellibranchiate, a Gastropod, and a small
Shrimp, besides this magnificent Decapod, which was entangled in the swabs of the
dredge. With the second specimen a great many animals came up, among which I may
mention a Palythoa, a Fungia, and a Pennatulid. A Holothuria, an Archaster, and an
Urchin represented the Echinoderms. Crustacea were present in large numbers—espe-
cially Peneid and Caridid Shrimps and the Schizopods which I have called Petalophthat-
mus and Chalaraspis, together with a large species of Huphausia. The smaller species
was got near Sombrero Island, W.I., ina depth of 450 fathoms, together with Astacus
zaleucus ; and in describing this species I have already given an idea of the fauna which
lives in the same place with these crustacea.

Characters of the Genus, and Systematic Position.
WILLEMOESIA, Grote *.

Carapace large, flattened, and quadrangular. Anterior margin and lateral borders
denticulated, the latter divided by two incisions into three denticulated portions. A
longitudinal line divides the carapace into a right anda left portion. Hyes and eye-stalks
entirely wanting. Funiculus of the first antennze rather large, its inner flagellum longer
than the outer one. Small lamellar appendage at the base of the second attenne, the
flagellum of which has the same length as the longer one of the inner antennz. Man-
ducatory portion of the mandibula very strong. A large flagellum attached to the second
maxilla. The palpus and flagellum of the maxillipedes reach the base of the first
antennee. No palpi at the base of the gnathopoda (second and third maxillipeds). First
pair of pereiopoda very elongated, terminated by a slender pair of chele. Subsequent
pereiopoda much shorter.

Small styliform appendages at the inner side of the ramus interior of the pleopoda.
Abdomen flattened, divided by a longitudinal line of spines into aright anda left portion.
Telson rapidly decreasing in size, longer than the shield-like caudal appendages.

1. W. LEPTODACTYLA, 0. sp.

Lateral borders of the carapace not very much expanded. Denticulation not very deep.
First pair of pereiopoda longer than the body.. All the pereiopoda terminated by chele,
Length 120 millims.

2. W. CRUCIFERA, 0. sp.

Carapace with wing-like expanded lateral borders. Deep incisions and strong den-
ticulation at its lateral borders. First pair of pereiopoda shorter than the body. Only
four pereiopoda terminated by chele. Length 42 millims.

* Tn ‘Nature,’ October, 1873.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION. 55

Among the living Decapoda Macrura there is hardly a group with which Willemoesia
could be said to be very closely allied. Nearest to it are undoubtedly the Scyllarine ; but
these, like all the genera of the family Palinuride, differ from it in the absence of the
lamellar appendage of the second antenne, and in the presence of palpi at the base of
the gnathopoda, which, as we have seen, are wanting in this new genus. Nor can it, for
this latter reason, be referred to the Astacidee, with which it has in common the presence
of the antennal scale. ©

It is very astonishing, indeed, that, among all crustaceans known to us, Willemoesia
approaches most closely the fossil Eryontide. If we compare, for example, our figure
of W. crucifera (Plate XII. fig. 10) with a figure of Hryon arctiformis, and the description
of the “Tribu des Eryons,” given by Milne-Edwards* (and probably taken especially
from Desmarest’s ‘ Crustacés Fossiles’), we find most striking resemblances between the
two forms. In W. crucifera as well as in Eryon the carapace has nearly half the length of
the whole body; and in both forms its lateral borders are wing-like expansions which
are divided by two deep incisions into three portions. The anterior border of the carapace
is nearly straight in both forms.

Eryon was probably not blind ; for the eye-stalks have been found in several specimens.
Its antennz seem to be somewhat more reduced than in Willemoesia; but the second
pair of them has, according to Desmarest, “ une écaille assez large, ovoide et forte-
ment échancrée.” This is the chief difference between Hryon and the Palinuride, and
the same in which also Willemoesia differs from that group.

Milne-Edwards says nothing on the parts of the mouth; but according to Quenstedt
they had a very large mandibula, one of the teeth of which was preeminently strong.
This is very much like what we find in Willemoesia; but in the fossil genus palpi were
present at the base of the first and second gnathopods, which are wanting in the living
genus. ‘The first pair of pereiopoda is in both forms longer than the following ones, and
terminated by a pair of long and slender chelze. In Hryon three pairs of pereiopoda, in
W. leptodactyla five, and in W. erucifera four are terminated by chelee. The form of the
last pereiopod in Z#. arctiformis is exactly the same as in WV. crucifera; and the abdomen
of these two forms is, as the above-mentioned figures show, so very much alike in the
two forms, that, if the last pair of pereiopoda and the pleon of Hryon were presented to me
without my knowing to what they belonged, I should undoubtedly declare them to be
parts of the genus Willemoesia. There are the same line of spines at the top of the
rings, the same winglike expansions on both sides, and that characteristic “ nageoire
caudale, dont la lame médiane est pointue et les quatre lames latérales moins longues
que la médiane et hastiformes.”” Also the fine fringe of hairs which distinguishes the
caudal fin of Willemoesia is to be seen in the fossil crustacean.

Eryon differs from the living genus chiefly by the presence of eye-stalks and of palpi at
the base of the gnathopoda. According to Quenstedt the latter were observed only with
some difficulty ; and their presence seems not to be beyond all doubt. I shall only on my
return be able to look myself over the original specimens and papers, and then, I hope,
be able to give a more detailed account on the relations of Willemoesia to Eryon.

* Histoire Naturelle des Crustacés, tome ii. (Paris, 1837) p. 278.

56 DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

That they must be very close appears already from what we have said; and among the
Eryontidz this new deep-sea genus must take its place. The Eryontide, now consisting
of two genera, must either form a new family intermediate between the Astacide and
Palinuridee, or be united with the latter.

During our cruise in the Atlantic we got deep-sea animals (7. e. from more than 350
fathoms; stations where only Foraminifera-shells have been got are not included) from
67 stations. After every haul a list of these animals, as much specialized as our know-
ledge of the thing allowed us to do, has been put down by me in the so-called Station-
book. As it might be of some interest to know how often the different groups of animals
have been got, I have made the necessary additions, and shall terminate this paper by
giving the results as far as Crustacea are concerned. According to this,

Cimipediathavesbeenscot momma ilem na enn WUIGIES:
Ostracoda hen cn t.. Mecy tat ee ie mace oem RCL
STO MVOC. one | Gilkaniol eich. Surin womb candy 9 C HMNES.
Wsopoday, | ei oa) eu 26 FR el oe en
Cumiaced OU OA OD COT OREO Te
Stomatopoda. . ........ . J time.
Schizopoda, |. . . «°% 5 =. « =.» -12\times.
Decapoda Macrura - 4) so. 5 2 ss . 22-5
IDecapock lke, 5 8! 556 6 os eo Wily

EXPLANATION OF PLATES.

Prats, VI.
All the figures, with the exception of fig. 4, refer to Nebalia longipes.

Fig. 1. A female. Low power. a, antenna of the first pair; 6, antenna of the second pair; Oc, eye;
R, rostrum; M, stomach; D, intestine; F, joint of the furca.
Fig. 2. A male. Letters the same as in fig 1. d, palpus of the first maxilla (‘‘ Putzfuss” Claus) ;
G, ductus eiaculatorius.
_ Fig. 3, Leg of a female: a, basal joint with the branchial appendage (K, A) ; 8, second joint with lateral
appendage (NN, A); y, main branch.
Fig. 4. Phyllopodal foot of the female of N. Geoffroyi, copied from Claus. Letters the same as in fig. 3.
y—-v, Joints of the main branch.
5. First antenna of a female. &. fourth joint, a claw. a, External, 6, ternal flagellum.
6. First antenna of a male. Letters the same as in fig. 5.
Vig. 7. Second antenna of a female.
8. Mandibula; p, palpus
9. First maxilla; p, palpus of a female.
Tig. 10. Second maxilla
Fig. 11. One of the anterior pleopoda: a, basal joimt; @, ramus exterior ; y, ramus interior; 6, retina-
culum corresponding to the same on the other side.

or
<I

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION.

Fig. 12. Fifth

Fig. 18. Sixth } pleopod of a female.

Prats VII.

All the figures refer to Petalophthalmus armiger. All drawn with very low power,
except figs. 7-9.

Fig. 1. A female, x 2 nat. size. a’, first antenna; ap, appendix lamellaris; a, second antenna;
pm, palpus mandibule ; mp, maxilliped; gn, gnathopoda; prp, pereiopoda; Ja, breeding-
lamelle.

Fig. 1 a. Caudal fin of the same.

Fig. 2. A male, x 2 nat. size. a’, first antenna; ap, appendix lamellaris ; a*, second antenna; pm,
enlarged palpus mandibule; mp, maxilliped; gn’, first gnathopod; prp?, second pereiopod ;
prp’, fifth pereiopod.

Fig. 2 a. Caudal fin of the same.

Fig. 3. First antenna of the female.

Fig. 4. Second antenna of the same.

Fig. 5. Scale of the second antenna.

Fig. 6. Upper lip.

Fig. 7. Mandibula of the female: pm, palpus mandibule.

Fig. 8. First maxilla of the same.

Fig. 9. Second maxilla of the same: p, palpus; fl, flagellum. ;

Fig. 10. Maxilliped of the same: a/, appendix lamellaris ; p, palpus; fl, flagellum.
Fig. 11. First gnathopod of the same.

Fig. 12. Second gnathopod of the same.

Fig. 13. Third pereiopod of the same.

Fig. 14. Pleopoda 1-5 of the same.

Vig. 15. Maxilliped of the male: /a, lamellar appendage.

Fig. 16. First gnathopod of the same: Ja, lamellar appendage.

Fig. 17. First, fourth, and fifth pleopoda of the same.

Prave VIL.

All the figures refer to Chalaraspis ungwiculata.

Fig. 1. A male, x2 nat. size. a’, first antenna; a, second antenna; gu, gnathopoda; p, pereiopoda.

Fig. la. Head of the male, seen from above: sc, small tubercle beneath the rostrum and between the
eyes.

Fig. 16. Caudal fin of the same.

Fig. 2. First antenna of a female.

Fig. 3. Second autenna, with lamellar appendage of the same.
Fig. 4. Mandibula of the same.

Fig. 5. Under lip of the same.

Fig. 6. First maxilla of the same.

Fig. 7. Second maxilla of the same: p, palpus ; fl, flacellum.
Fig. 8. Maxilliped ofthe same: p, palpus; ji, flagellum.

Fig. 9. First gnathopod

5 ; : |of the same.
Fig. 10. First pereiopod

Fig. 11. Upper lip of the same.
SECOND SERIES.—ZOOLOGY, VOL. I.

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DR. R. V. WILLEMOES-SUHM ON SOME ATLANTIC

- 12. Fourth pereiopod i
. 13. Fifth pereiopod fof the same.
. 14, Anterior pleopod

Puate IX.
1. Gnathophausia gracilis, male, x 4 nat. size.

2. Second antenna of Gn. zoéa: al, appendix lamellaris ; fw, funiculus; jl, flagellum.
3. Mandibula of the same ; p, palpus.

. 4. Labium of the same.
. 5, Hirst maxilla of the same, higher power: p, palpus maxille.
ig. 6. Second maxilla of the same: fl, flagellum ; p, palpus; 0, accessory eye.

- Maxilhped of the same: jl, flagellum.

g. 15. One of the anterior pleopoda of the same.
g. 16. Maxilliped of Gn. gigas: fl, flagellum ; p, palpus.
g. 17. Outlines of the eye of the same.

PLATE X.

. 1. Astacus zaleucus, male, nat. size

g. la. One of its right pleopoda.

Nig. 2. Gnathophausia gigas, male, nat. size.

. 2a, The same, side view (the carapace is lifted somewhat to show that it is not connected with the

pereion), nat. size.

Hig. 3. Oral apparatus of the same: /, labrum; md, mandibula; mp, palpus mandibule ; p, palpus of the

second maxilla; jl, its flagellum; ac, accessory eyes on the second maxilla; basal joint of
the maxilliped with its palpus (the other joints of m have not been drawn, in order not to
confuse the figure) ; 2 and y, plates formed by the flattened inferior borders of the carapace.

4. Gnathophausia zoéa, male, x 4 nat. size.

Prats XI.

‘ig. 1. Zoéa of Cardisoma, sp.? from San Jago (Cape-Verd Islands). High power. a’, first, a?, second

antenna; m, parts of the mouth; gp, the gnathopoda.

». 2. Caudal fin of the same, seen from above. High power.
e. 3. A string of eggs and egg-capsules, partly containing zoéas, partly left by them. ‘Taken from

the abdominal feet of the mother. Low power.

lig. . Cystisoma neptunus (Thaumops pellucida), male. Nat. size.
ig. 5. Genital organs of a smaller male: A, segments of the percion; B, first segment of the pleon ;

t, testes; vd, vasa deferentia; ag, aperture genitales.

. 6, Mandibula of the same. High power.
7. Maxilla of the same.
8. Maxillipedes (m) and second maxilla, with (/) labium (?) of the same.

Pruate XII.

Vigs, 1-9 refer to Apseudes ceca.

Vig.

1. A female, x 4 nat. size: a, antenna; gn, gnathopoda.

Fig

1
(of)

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Fig.
Fig.
Fig.

CRUSTACEA FROM THE ‘ CHALLENGER’ EXPEDITION.

. 2, First antenna.

. 8. Second antenna: s, scale; f, flagellum.
. 4. First gnathopod.

. 5. Second gnathopod.

s. 6, 7. First and second pereiopods.

. 8. Last joints of the fourth pereiopod.
. 9. Maxilliped.
. 10. Willemoesia crucifera, x 4 nat. size.

Pruate XIII.

. Willemoesia leptodactyla, male, nat. size.
. First styliform pleopod.

. Upper border of the opening of the mouth.
. Mandibula, with the palpus (ym) and the left labium (Jad).
- 6a. First maxilla: fl, rudiment of a flagellum.

oF WOH

g. 66. Second maxilla: fi, flagellum.
g. 7. (First) maxilliped: p, palpus; fl, upper half of the flagellum ; fi”, under half.
g. 8. First gnathopod (second maxilliped).

9. Second gnathopod (third maxilliped).
10. Second left pleopod of W. crucifera, with two styliform appendages.
11. Third right pleopod of the same, with one styliform appendage.

- Second pleopod of the right side: a@, styliform appendage to the inner ramus.

12. Second right pleopod of an undescribed Palinurid from deep water near-the Bermudas.

Wherever the words “high power” are not added, a low power of a Hartnack’s microscope, or, in

case it was not possible to dissect the animal, a loupe has been used.

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[ 61 ]

III. On the Structure and Systematic Position of Stephanoscyphus mirabilis, the Type
of a new Order of Hydrozoa. By Prof. Attman, W.D., DL.D., FR.S., &c., Pres.
Linn. Soe.

(Plate XIV.)

Read November 19th, 1874.

WHILST engaged last May in zoological researches on the coast of Antibes, I was
struck by some small patches of one of the horny sponges which had attached them-
selves to stones at a slight distance below the surface of the sea, and which had arrested
my attention by what appeared to be unusually distinct and well-defined oscula. After
transferring some specimens to a phial of sea-water, I was astonished to find that from
each of the apparent oscula a beautiful crown of delicate tentacles soon began to develop
itself. The tentacles were quite like those of certain hydroids; and the appearance was
such as might easily have led to the belief that the anticipated form had at last been
found which would decide the zoological position of the sponges, and remove all doubt of
the hydrozoal structure recently assigned to them*. A closer examination, however,
showed that the orifices on the surface were not really oscula, and that the tentacles
proceeded, not from the sponge itself, but from a totally different organism, which lay
imbedded in its substance, and which sent tubular prolongations to the surface, where
they opened in the manner of genuine oscula.

It was further evident that the organism thus associated with the sponge was in the
highest degree interesting ; and a careful study of its structure made it plain that, while
in all essential points it possessed a true hydrozoal organization, it differed from all the
recognized orders of Hydrozoa so widely as to necessitate our assuming it as the type of
an entirely new one.

To this remarkable animal I have given the generic name of Stephanoscyphus. When
freed from the surrounding sponge, it is seen to be composed of an assemblage of zooids,
which are included in chitinous tubes, and associated into a composite colony by a common
basal plexus of tubes, the aspect of the whole being very much that of an ordinary hydroid
trophosome (figs. 2,3). In all the specimens examined the common plexus lay at the
base of the sponge, and the tubes given off from this penetrated the sponge-tissue, and,
rapidly increasing in width, ultimately reached the surface, on which they opened. The
tubes really extended beyond the general surface; but each carried up the sponge-sub-
stance with it, so that the orifices appeared elevated on the summit of conical papillee—
a condition very common in the oscula of sponges (fig. 1).

The distal extremity of the animal can extend itself beyond the safes of the chitinous
tubes, and can again withdraw itself far into the interior. When extended (fig. 3, a), it
displays a coronal of tentacles, which surround a wide terminal orifice. The tentacles
spring from the very margin of this orifice ; for there is nothing like a hypostome or pro-

* Haeckel, ‘ Kalkschwimme.’

62, PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS.

boscis, such as in the true hydroids we find intervening between the tentacles and the
mouth of the hydranth ; and it was the great width of the terminal orifice, with its mar-
ginal circlet of tentacles, so different from the narrow mouth of a hydroid elevated on
the summit of a hypostome, that first led me to doubt the justice of referring Stephano-
scyphus to the true hydroids. The appearance of the tentacular crown, indeed, was not
altogether unlike that of certain zoanthal Actinozoa; but all grounds for assigning to
Stephanoscyphus such actinozoal relations were removed by dissection.

The tentacles form a single circlet, which is composed of two closely approximated
alternating series of about eighteen tentacles in each, the outer series being slightly
shorter than the inner, and usually depressed in extension, while the inner series is held
more elevated. Not unfrequently the whole circlet may be seen bent back upon the
outer side of the chitinous tube (fig. 3, ¢). The tentacles are very contractile: when
fully extended they are long and filiform; but in certain states of contraction they
assume a very distinctly clavate form (fig. 8, 6). Their structure is entirely that of the
typical hydroid tentacle: they present the septate condition so characteristic of such
tentacles ; and they carry along their length, just as in most hydroids, somewhat spirally
arranged slightly elevated clusters of thread-cells (fig. 4). In complete retraction the
tentacles are not only greatly shortened, but are thrown inwards and entirely with-
drawn within the terminal orifice of the animal, which is then nearly or altogether
closed over them (fig. 3, d).

In the completely extended state of the animal, four longitudinal canals may be traced
from the base of the tentacular crown for some distance backwards in the walls of the
body ; but the general want of transparency, and the way in which the greater part of
the animal is concealed by the surrounding sponge, renders it impossible to obtain,
without dissection, a satisfactory view of the deeper-lying parts. Sections, however, were
easily made of specimens which had been preserved in spirits; and some further im-
portant points of structure were thus determined.

When a longitudinal section (fig. 5) of such specimens is made through the axis of
the body, it is seen that the terminal orifice opens into a wide thin-walled cavity, which
becomes continuous posteriorly with the canal which occupies the axis of the stem.
The contracted tentacles are now also brought into view with their extremities thrown
back, and with the whole tentacular crown entirely withdrawn into the cavity.

By the same section another very important piece of structure becomes apparent; for
‘it may be now seen that the knife has divided a circular canal which runs in the walls of
the body round the terminal orifice. This canal is very obvious; it is remarkable for its
width, and easily admits the passage into it of a needle. It supports the tentacular
circlet; and when a transverse section of the body is made along its course, it is seen to
be quite continuous and destitute of septa. It possesses a very distinct endodermal
lining.

With the circular canal the distal extremities of the longitndinal canals are in con-
nexion, and they probably open into it. ‘These canals can now be traced far back in the
walis of the body. They lie quite close to the inner surface of the walls ; and their sides
project into the cavity as four slightly elevated ridges. In the deeper parts of this

PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS. 63

cavity they present a somewhat lobed or pouched appearance; but beyond this they
become lost in the thickened walls of the body, so that their actual termination could
not be made out. Their walls are thick; and their lumen, which is very narrow, is
doubtless lined by an endoderm; but this could not be demonstrated with the same
distinctness as in the circular canal.

The walls of the cavity which occupies the axis of the narrow portion, or stem, are
much thicker than those of the wide distal chamber, and posteriorly encroach so much
upon the axile cavity as to reduce this to a very narrow canal.

Thread-cells exist throughout in the ectodermal layer; but these are everywhere
exceedingly minute. Those examined from the body-walls in spirit specimens were
nearly spherical, and under a 3‘; immersion-lens were seen to include a spirally coiled
filament ; they were about the gso9 of an inch in diameter.

The more posterior or proximal parts of the animal are fixed in the external chitinous
perisarcal tube, and possess no obvious contractility; but the more anterior or distal
portion is very contractile, and is quite free from the tube, so that it can move up and
down in it during extension and retraction, quite like a campanularian hydranth in its
hydrotheca.

The surrounding chitinous tube, indeed, increases so rapidly in width towards its
distal end as to resemble here the hydrotheca of a campanularian (fig. 3). It is marked
by annular ruge for some distance from its free extremity, and usually by fine longi-
tudinal striz ; but towards its attached end the marking disappears.

At rather irregular distances along its length its inner surface sends off remarkable
processes, which project far into the interior (fig. 5, a, a, and fig. 3, e). At four equi-
distant points, situated on the same level, a thin chitinous lamina separates from the
inner surface of the tube, and forms a hollow pyramidal process. The four processes
thus formed radiate towards the axis, pushing before them the soft parts, and forming a
stelliform constriction of the body (fig. 6). Between each of these four radiating pro-
cesses a much smaller one is sometimes present, thus raising to eight the number of pro-
cesses in a set, and indicating an increase in number by consecutive intercalations of
younger ones between those previously formed. These imperfect septa, notwithstanding
their striking appearance, do not seem to possess any important significance in the orga-
nization. They will, however, suggest a comparison with the tabule of the tabulate
corals, and may be regarded as offering some support to Agassiz’s view of the hydrozoal
nature of these animals. j

In none of the specimens examined could generative elements be found. It is almost
certain, however, that the absence of obvious generative elements is the result of the
immature condition of the reproductive organs. We should expect to find them in the
walls of the longitudinal canals; and I believe that the pouched condition observed in
these canals in the deeper part of their course is only an early state of the reproductive
system.

64 PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS.

General Remarks—Homology.

From the account now given of Stephanoscyphus mirabilis, it is evident that we have in
this remarkable animal a tue hydrozoal structure ; but while its apparent form and habit
are those of a hydroid trophosome, its organization is, in its essential features, that of a
hydroid planoblast or medusa. If we imagine a campanularian hydroid in which the
hydranths have been replaced by peculiarly modified medusze, capable, as in the hydranth,
of exsertion beyond the hydrothec, and of retraction within them, we shall form very
nearly a true conception of Stephanoscyphus. 'The wide thin-walled chamber into which
the terminal orifice opens would thus correspond to the umbrella of a medusa; and the
terminal orifice itself, though neither lithocysts nor ocelli appear to be developed
on its margin, would be homologous, not with the mouth of a hydranth, but with the
codonostome or umbrella-aperture of a medusa; while the tentacular crown would admit
of a direct comparison, not with the tentacular crown of a hydranth, but with the mar-
ginal tentacles of a medusa. The true mouth would thus have to be sought for at the
point where the wide distal cavity passes into the narrow proximal cavity which occupies
the axis of the stem. Then there are the four longitudinal canals running symmetrically
in the umbrella-walls, and the circular canal surrounding the codonostome, all so many
obvious features of medusal organization. It is true that no manubrium appears to be
developed, and the bottom of the umbrella-cavity seems to pass uninterruptedly into the
true gastric cavity (which is continued through the axis of the stem), neither is there a
velum developed at the codonostome, nor do the walls possess the gelatinous consistence
and transparency of the typical medusze; but these special modifications in no degree
invalidate the points of homology here insisted on. In the absence of a manubrium we
are reminded of the swimming-bells or nectocalices of the Siphonophora, which, though
closely resembling in all other respects a true hydroid medusa, and undoubtedly homo-
logous with it, are destitute of a manubrium.

If the views now advocated be correct, the wide cavity into which the terminal orifice
opens ought to have no endodermal lining: and this I believe to be really the fact; for a
very careful examination has failed to demonstrate more than a single layer in its walls.
On the contrary, the endoderm of the circular canal is very obvious; and though a
proper lining membrane cannot be distinctly recognized in the longitudinal canals, this
ought not, in an observation rendered difficult by the narrowness of these canals and by
the want of transparency of the tissues, to be urged as an argument against its presence.
On the other hand, the axile cavity of the stem appears to possess a well-developed endo-
derm, and the proper gastric cavity must be regarded as confined to this part.

The peculiar position assumed by the tentacles in extreme retraction is very different
from the condition of these parts in the retracted state of the hydranth in a hydroid
trophosome, and quite corresponds with that which they present in the greater number
of young medusiform planoblasts, in which before the planoblast becomes free the
marginal tentacles are thrown back, and lie entirely within the cavity of the umbrella.

1 have elsewhere * endeavoured to show that the radiating canals of a medusa are —
strictly homologous with the tentacles of a hydranth, and that, if the latter were con-

* Monograph of Gymnoblastic Hydroids.

PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS, 65

nected with one another by an extension of the ectoderm, we should have the tentacular
crown of the hydranth converted in nearly all essential points into the umbrella with the
radiating canals of a medusa.

Now in certain species of Hydroida (Laomedea flexuosa for example) the tentacles are
actually connected at their bases by an ectodermal web. Here, then, there is little
needed to render the parallelism complete but the presence of a circular canal. Ste-
phanoscyphus, however, supplies the missing element, and, by the addition of a well-
developed circular canal, beautifully confirms the view here taken of the homological
relations between hydranth and medusa.

Whether the relation between Stephanoscyphus mirabilis and the sponge with which
it is found associated is one of parasitism, or whether the association is merely acci-
dental, it is at present impossible to say. At all events, in no instance did I meet with
the Stephanoscyphus free from the accompanying sponge.

From the account now given it is obvious that Stephanoscyphus must form the type
of a distinct order of Hydrozoa. With the order Hydroida it possesses very decided
affinities, but it is nevertheless removed from the Hydroida by a distance at least as great
as that which separates these from the Siphonophora. For the new order I propose the
name of THECOMEDUS£; and if we bring together the characters given above, so as to
enable us to assign to Stephanoscyphus its true systematic position, we shall obtain
the following diagnostic category :—

Subkingdom CHLENTERATA.
Class HyDROZzOA.
Order THECOMEDUS&.

THECOMEDUS2.

Animal composite, zooids medusiform, with circular and radiating canals, and included
in a chitinous rooted perisare, which forms thecze, within which they are retractile.

Genus Stephanoscyphus.
Name from orépavoc, a crown, and sxioc, a cup, in allusion to the cup-like body with
its crown of marginal tentacles.
Gen. char.—Terminal orifice surrounded by a single circlet of simple filiform contractile
tentacula; four longitudinal canals; velum and manubrium undeveloped.
Generative elements formed in the walls of the longitudinal canals ?

Stephanoscyphus mirabilis.

Perisarcal tubes springing from a basal tubular plexus, and rising to a height of about
one third of an inch, simple, rapidly enlarging upwards, annulated for some distance
from their distal extremities ; tentacles about 36 in number.

Hatitat.—Immersed in the substance of certain corneous sponges.
Locality —Antibes, Mediterranean coast of France.—G. J. A.

SECOND SERIES.—ZOOLOGY, VOL. I. K

66 PROF. ALLMAN ON STEPHANOSCYPHUS MIRABILIS.

DESCRIPTION OF PLATE.
PuaTe XIV.
Fig. 1. Stephanoscyphus mirabilis, with its investing sponge : nat. size.

2. Perisarcal tubes of the same freed from the sponge: nat. size.

3. A group of zooids magnified. a, the animal fully exserted and the tentacles extended; 5, the
animal partially retracted ; c, the animal with the tentacles thrown back over the sides of the
tube; d, the animal completely retracted; e, an empty perisarcal tube: the internal chitinous
processes are seen through its walls.

4. Distal portion of a tentacle seen im optical longitudinal section : more highly magnified.

5. Longitudinal section through a perisarcal tube and the retracted animal, from a specimen preserved
in spirit: a, a,imternal chitious processes.

6. Transverse section of same through one of the verticils of chitinous processes.

7. Thread-cell from body.

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Trans. Linn. Soc. Ser 2. Zoou No.l Tas.

W. West & C° amp.

mirabilis.

Steph anos cyphus

GJ Allman del.
WH Wesley lth.

THE .

TRANSACTIONS

OF

THE LINNEAN SOCIETY

OF

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART THE SECOND.

PAO) NEDEOAN:
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M.DCCC.LXXV.

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IV. On the external Anatomy of Tanais vittatus, occurring with Limnoria and Chelura
terebrans in excavated Pier-wood. By JoHN Dents Macponatp, I.D., F.R.S.,
Staff-Surgeon R.N., Assistant Professor of Naval Hygiene, Netley Medical School.
(Communicated by W.'T. Tuisetton Dyer, WA, F.L.S.)

(Plate XV.)

Read Feb. 18th, 1875.

WHILE searching for Limnoria terebrans in excavated wood from N etley Hospital
Pier my attention was diverted to another little Crustacean, rather smaller in size, but
wearing the outward appearance of a chelate Decapod so completely that it was only by
counting its legs and more minute inspection of the leading features of its structure I
was enabled to determine it to be a veritable Isopod, though of an aberrant type.

It proved to be conformable with the characters of Tanais Cavolini (Edw.) = Crossurus
vittatus (Rathke), which was originally found in the Gulf of Naples, the habitat assigned
to it by Milne-Edwards; but it has since been shown to have a wider range. Its occur-
rence under the above-mentioned circumstances is worth recording ; for I have also found
it in perforated and decayed wood from the pier at Alum Bay, Isle of Wight; and I have
no doubt that it would be captured in other localities if it were sought for in the burrows
of Limnoria.

Quite exceptionally, in the Tanaidee the head and first thoracic segment coalesce above
to form a perfect little carapace or cephalothorax, which, in the little creature under
consideration, is somewhat triangular in form, with an obtuse but distinct rostrum in
front, from beneath which the two pairs of antenne project forwards; and immediately
to the right and left of these are the orbital notches for the eyes.

These latter organs are very prominent, in articulation with the carapace, and all but
peduneculated. They are conical in form, with the apex directed forwards, and rather
more convex or rounded on the outer than on the inner side. Round lenticular cornes,
not unlike those of Limnoria, but much smaller, are recognized by their brilliancy, and
the relief of a central mass of black pigment radiating amongst the little nervous cups in
connexion with them.

Having thus commenced with the eyes, I shall describe the members of the succeeding
segments seriatim, which will render their homological relations more intelligible.

The two pairs of antenne (figs. 1, 2, & 3) are nearly of equal length, rather shorter
than the cephalothorax, and disposed in two lines, one above another. The superior
antennee (a, figs. 2 & 3), which are much stouter and somewhat longer than the inferior,
are each composed of three joints, successively diminishing in their proportions, and
setaceous at their distalends. Several bristles of a ligulate form (fig. 2, e), with transverse
strize near their base, occur amongst the simple ones in the tuft at the tip of the terminal
segment. A labyrinth-like excavation at the upper and inner side of the base of the first

SECOND SERIES.—ZOOLOGY, VOL. I. L

68 DR. J. D. MACDONALD ON TANAIS VITTATUS.

articulation (fig. 2,¢) represents the so-called olfactory organ of the Decapod. The second
or inferior pair of antenneze are narrower than the superior, and composed of five joints,
of which the first, third, and fifth are shorter than the second and fourth; and the ulti-
mate joint is tipped with a brush of long and simple hairs, springing from what would
appear to be an aborted multiarticulate filament.

The oral organs consist of a spatulate upper lip, fringed with short hairs (fig. 6, a), a
pair of two-branched mandibles without palps (0), and articulated laterally with the buccal
frame, two pairs of maxille (¢ & d), and a pair of five-jointed jaw-feet (/f), inclosing, at
their base, two membraneous plates, forming a lower lip (¢).

The internal maxille (¢) are in the form of rounded membraneous lamelle, fringed
with fine hairs, and presenting a Single-jointed palpiform process near their summit
externally.

The external maxillee(d), on the contrary, are strong and sabre-shaped, with a spiny
and setaceous extremity. ‘Two oval leaflets (d’), connected with these internally, form a
tongue-like organ in the mouth; and at their base externally, but concealed by the cara-
pace, is a conical appendage (d”) or flagellum, with long diverging hairs at the tip. In
the possession of this appendage, as Fritz Miller has pointed out, the Tanaidee make a
still further approach to the Decapoda. Thus, in his ‘ Facts and Arguments for Darwin’*,
he remarks, “ Whilst in all other Oniscoida the abdominal feet serve for respiration,
these in our cheliferous Isopod (Zanais dubiws(?) Kr.), are solely motory organs, into
which no blood-corpuscle ever enters ; and the chief seat of respiration is, as in the Zoez,
in the lateral parts of the carapace, which are abundantly traversed by currents of blood,
and beneath which a constant stream of water passes, maintained, as in the Zoez and the
adult Decapod, by an appendage of the second pair of maxille, which is wanting in all
other Edriophthalmata.” In passing, I may mention that the figure of Tanais dubws, so
called, loc. cit., is rather more conformable to the genus Leptochelia. In this connexion
Bate and Westwood’s beautiful work on the British sessile-eyed Crustacea may be con-
sulted with advantage. But to return to the subject, the joints of the jaw-fect (f) are
furnished with rigid comb-like bristles, arising from the inner border of the third and
fourth, and the free extremity of the fifth. The lower lip (e) is also fringed with fine
short hairs.

Following the jaw-feet just noticed are the seven pairs of legs common to all the Kdri
ophthalmata, the first two and last five respectively corresponding with the two external
‘pairs of jaw-feet, and the five pairs of legs appertaining to the thorax of the Decapod.

The two first limbs (fig. 7) are strongly chelate, and many times thicker than the
ambulatory legs succeeding them. The fixed claw presents a trenchant upper border,
more prominent in the middle, with which, at least in the female, the movable claw cor-
responds more or less closely. In the male, however, (fig. 15) both claws are more
arcuate. As in the beak of the Crossbill, the opposable tips of the claws are curved
with sufficient obliquity to cross over one another, the index taking the outer side. They
are of a rich reddish amber tint, similar to that of the dental processes of the mandibles.

* Translated by W. 8. Dallas, F.L.S8.

DR. J. D. MACDONALD ON TANAIS VITTATUS. 69

The first or basal portions of these limbs form two oval projections beneath the thorax ;
the second is a small triangular piece, with which the third is so connected as to present
a rounded projection posteriorly, while it extends forwards to articulate with the manus.
In the female (figs. 1 & 7) eight or nine stout bristles spring in a row at the outer side
of the base of the cutting-border of the manus, while in the male they are interrupted in
the middle. Other bristles occur near the articulation, 7. e. one on the inner and several
on the outer side.

As already mentioned, the first thoracic ring having coalesced with the head, the six
following pairs of legs arise from the six apparent thoracic segments. ‘The first visible
segment, distinct from the carapace, must, therefore, carry the second pair of legs. These
have long curved thigh-pieces, with which the succeeding joints of the limbs are so arti-
culated as to permit them to play their legitimate part as external jaw-feet, curving
inwards and forwards round the basal prominence of the first pair. Their claws (fig. 8)
are long, simpie, styliform, and but slightly curved; they are themselves rather longer
than the other legs.

The claws of the third and fourth pairs (figs. 9 & 10) are simply uncinate, while those
of the last three pairs (figs. 11, 12, & 13), which are curved forwards, are also combed
on either side, though this is not quite so apparent on the inner side in the last pair.
Instead of it, however, they present a close row of stiff serrated hairs at the extremity of
the penultimate joint, which is moreover roughened with rasping tubercles (13 a), quite
like those which beset the whole surface of the body in Limnoria. In all the legs, with
the exception of first and second pairs, the third segment presents a row of stout spines on
either side of its articulation with the fourth.

The abdomen consists of five segments, the first three of which bear the respiratory
lamelle and swimmerets together. The fourth is much smaller than those preceding it,
and without appendage. The fifth is rather more than twice the length of the fourth,
broadly shield-shaped, with four rigid hairs projecting backwards from the middle of
its posterior border. Besides the swimmerets there is a fourth pair of appendages
(figs. 1, 4, 5, & 14), which are rather conspicuous on account of their projection beyond
the terminal segment. They have a simple folded lamella on the inner side, and much
resemble a pair of three-jointed antennz, with stout diverging hairs at the distal end of
each joint.

The inner ovate and acute leaflets of the abdominal appendages (fig. 4, 6) lie flat, and
are apparently motionless ; while the outer ones, which are rather semicircular (a), cross
over them superficially towards opposite sides. Both borders of the deep leaflets and the
outer border of the superficial ones, as well as that of the broad propodia, are beautifully
fringed with long flat acute and feathery setze (d). Similar feathery hairs forma trans-
verse linear band or fascicle upon the dorsum of the first and second segments.

The generative pores lie side by side near the middle of the sternal plate, between the
two last thoracic limbs.

The female (fig. 1) carries her eggs in a membraneous expansion or saccule under the
thorax ; and, from having observed the chele and other anatomical characters of the
adult in the young soon after leaving the egg, I think it may be concluded that there is

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V. On Valencinia Armandi*, a new Nemertean.
By W.C. M‘Intosu, U.D., F.R.S.H., BLS.

(Plate XVI.)
Read June 17th, 1875.

WHILE searching for a rare Annelid (Mea mirabilis) that oceurs near low water in
the extensive waste of sand at Southport, a slender thread, which glistened in the October
sun like gossamer, attracted notice in the upturned sand. This was the proboscis of the
ruptured Nemertean, which exists in considerable abundance in the sand amongst the
tubes of Lanice conchilega, associated with Amphiporus lactifloreus, A. hastatus, Micrura
Jusca, and other forms usually characteristic of the laminarian and coralline regions of
the north. It appears to live in a burrow in the sand, and readily takes to a case or tube
in confinement. A

The body is 7 or 8 inches in length, and not much thicker than a stout thread. It is
rounded in front and in the middle, but becomes extremely flattened towards the tail,
which terminates in a somewhat lanceolate point. The anterior part of the body is
whitish, the posterior pale buff, while the tail is translucent. The head is rounded in
front, flattened, and slightly enlarged behind the tip. There is no trace of eyes, fissure,
or furrow. On the whole the Nemertean very much resembled Carinella linearis, from
Lochmaddy; and it was placed in spirit under this impression. More careful scrutiny,
however, reveals that it properly comes near the genus Valencinia, De Quatrefages,
char. emend., the only British example of which (Valencinia lineiformis) was dredged in
numbers amongst the shell-gravel of Bressay Sound, in Shetland. The Zetlandic species
is larger, and has an evident row of eyes on each side of the head, besides other points of
distinction, which will by-and-by be alluded to.

In the structure of the cutaneous system Valencinia Armandi agrees with its allies.
The entire surface is clothed with cilia; and the cutis has the usual cellulo-granular
appearance (fig. 9, @), with numerous vertical streaks. The latter are not conspicuous in
the Zetlandic species (fig. 10, a), which has a minutely cellulo-granular cutis, very easily
removed in the preparations. In the new form the cutis is bounded externally by a some-
what definite edge, which probably represents a rudimentary cuticle. In the next layer
(4, figs. 9 & 10) a considerable difference exists between the two species. In Valencinia
Armandi the basis-layer is very thin, while in /. lineiformis it is dense. In the snout
of the former the cutis is proportionally thicker, but the basis-layer is not increased—
another contrast; for the latter is especially developed on the snout in the Zetlandic
species. In VY. Armandi there is an increase of this layer opposite the nerve-cords. In
connexion with the cutaneous system, it may be mentioned that, in a large Canadian

* Named after M. Armand de Quatrefages, who long ago did such good work in the group.

74. W. C. MIINTOSH ON VALENCINIA ARMANDI.

Carmella, fibres from the elaborate muscular system of the snout pass from the trans-
verse bands into the cutaneous layer; so that, with the arrangement of the basis-layer, the
tissue has a characteristic arborescent appearance.

Underneath the basis-layer is a well-marked belt of circular muscular fibres. Towards
the tip of the snout these fibres are not developed as an external coat, probably because
the complex system of internal muscles, hereafter to be described, renders their presence
less necessary. Immediately behind the anterior border of the snout (fig. 1) a differen-
tiation of the cutaneous stroma occurs. At first, in front of the proboscidian aperture,
a single distinct transverse band appears, with numerous vascular meshes beneath it,
such probably representing the area of communication between the lateral trunks. Two
transverse muscular bands, however, as soon as the proboscidian canal is reached (fig. 2,
e & e’), pass from side to side, leaving a considerable space between them; and this is
occupied by vertical bundles, which enclose various meshes, the chief being in the centre
for the proboscidian canal, while several smaller vascular spaces exist on each side.
Very shortly, however, as we proceed backward, greater differentiation occurs; for the
transverse bands increase in strength, and numerous longitudinal muscular fibres appear
above and beneath their outer borders. A transverse section, therefore, in this region
(fig. 2) shows beneath the cutis (a) the pale and comparatively thin basis-layer (0). The
two great transverse muscular bands (e & e’) pass right across, forming a kind of ellipse,
each spreading out so as almost to meet its fellow at the circumference on each side, and
perforated here and there by the vertical bands. Numerous vertical bands extend
from the superior and inferior borders of the ellipse to the circumference, often in an
arborescent manner, and become lost at the basis-layer. The spaces thus left between
the outer edge of the ellipse and the latter are filled with powerful bundles of longitudinal
muscular fibres (7d). The area of the ellipse, again, presents the canal for the proboscis
(ao) in the centre, with its pale glandular lining internally, and externally its strong coat
of longitudinal muscular fibres. This canal is supported on each side by vertical mus-
cular bands, whose fibres, spreading out at each transverse muscle, pass to the circum-
ference above and beneath. Two bundles of fibres are especially conspicuous on each
side. Then come a great vascular space and a small one between it and the circumfer-
ence. Between these spaces and at the outer edge similar vertical bands of fibres occur,
with numerous longitudinal fasciculi. Along the inferior border of the superior trans-
verse muscle and the superior border of the inferior are many longitudinal bundles, form-

‘ing a somewhat regularly diminishing series towards the central line. Proceeding back-
ward, it is found that the chief changes in the great transverse bands are their diminu-
tion in length, increase in bulk, and more evident divergence at the extremities, so that
their lateral fibres almost decussate. The superior and inferior vertical bands are also
more boldly developed. There are still two vascular tubes on each side of the probosci-
dian canal, a larger and a smaller. By-and-by there is a tendency to further differentia-
tion in the proboscidian and in the vascular area, as shown by the greater development
of a lateral vertical band, composed internally of longitudinal, and externally of vertical
fibres. This becomes marked as soon as the anterior border of the ganglia and the turn-
ing over of the proboscidian sheath are reached, while only a single vascular channel is"

W. C. M‘SINTOSH ON VALENCINIA ARMANDI. 75

present on each side. A thin ring of circular muscular fibres can now be traced all
round the united area of the proboscis and the vascular canals, the great transverse
bands being inflected at each side where the circular fibres pass. Between the united
area and the circumference of the body we have now the ever-increasing ganglia, which
rest inferiorly on the outer division of the inferior transverse band, and are bounded
superiorly by the outer fibres of the upper transverse band. Externally they apparently
abut on the basis-layer. About this region a curious maculation of the part within the
basis-layer is observed, especially superiorly—a feature due to the presence of two or three
narrow longitudinal separate layers of muscular fibres, which form a complete investment.
Just in front of the mouth they form a rather thick compound layer, best marked supe-
riorly and inferiorly; but the advent of the oral aperture separates the latter into two
great masses. Some circular fibres also appear to be present at the basis-layer, but they
are much less conspicuous than the longitudinal. A section made through the ganglia,
almost in the plane of both commissures (fig. 3), shows three rows of these interrupted
muscles (d’) superiorly and inferiorly; but the lateral regions are less distinct. The
ganglia (/) surround the united area of the proboscidian and vascular systems; and, besides,
their other investments have increased in complexity. The external circular and internal
longitudinal fibres of the area still form a complete ring; but there is also an inferior
appendix in the shape of a canal (s) on each side, separated from the great vascular tunnel
by a powerful transverse band (m). From the centre of the latter superiorly springs a
process ( p), which separates the great vascular tubes, and terminates above in the circular
coat of the proboscidian chamber(ao). Then, shortly afterwards, a series of longitudinal
fibres appear in the isthmus between the vascular channels, and run round the inner
circular coat, which further presents on its inner surface a complete lining of longitu-
dinal fibres. The two vascular canals appended to the inferior border of the transverse
band are soon separated by a broad transverse belt, and by-and-by are obliterated by the
mouth. The broad transverse belt just mentioned spreads out on each side underneath
the diminishing ganglia; and some of the inferior fibres join the external longitudinal
series of separate muscles. The only representative of the great longitudinal muscular
layer afterwards so highly developed is a narrow arch (fig. 4, d), extending from the upper
end of each great vascular channel, and meeting at the raphe in the dorsal median line.
This, however, soon increases in thickness, and presents two channels (fig. 4, vs) between
its inner surface and the proboscidian sheath; while behind the mouth it quite encircles
the body. The circular muscular coat (c) likewise completes the circuit of the body
immediately after the mouth. The separated layer of longitudinal fibres (d’) outside
the circular layer soon breaks up into small fasciculi, which, in transverse section, form
a dotted belt outside the nerve-cords.

There is thus a very characteristic difference between this form and Valencinia linei-
formis or Carinella. In V. linetformis the snout, though very much longer in front of
the mouth, is less complex, there is only one vascular channel (of large size) on each
side of the proboscidian canal, and the ganglia have no longitudinal fibres between them
and the basis-layer. From without inwards there are the cutis, basis-layer, and circular
_ muscular fibres, while the rest of the area, with the exception of the three channels just
SECOND SERIES.—ZOOLOGY, VOL. I. M

76 W.C. M‘INTOSH ON VALENCINIA ARMANDI.

mentioned, is filled up by the great longitudinal muscular layer. The nerves lie in the
substance of the latter at once, whereas in the new form the nerves are placed quite out-
side the great longitudinal coat, and the whole region is much more complex. Ina
large Canadian example of Carinella*, again, the nerves lie outside all the muscular
layers, but the complexity of the muscular system of the snout is equally wonderful.
The tip is formed of an interwoven series of longitudinal, vertical, and transverse
fibres—which lie within the circular coat, and interdigitate with certain decussating
fibres of the latter at its inner margin. A very symmetrical arrangement of this
kind occurs in the middle line at the tip in front of the proboscidian aperture. At first
the snout is quite filled up with these elaborate muscles, with the exception of the pro-
boscidian canal; then a series of vascular spaces form above the latter, and range
themselves on each side, while a great longitudinal muscular mass occurs inferiorly.
Strong vertical bands on each side of the proboscidian canal, and transverse muscles
below it, map out the region very effectively. Thus at the ganglia a transverse section
shows within the cutaneous layer the circular muscular coat, with it decussating fibres
dorsally and ventrally. The great longitudinal coat forms a complete circuit, pierced
here and there by the vertical and other fibres. There are apparently two great vascular
channels above the great transverse muscular band on each side of the proboscidian canal ;
but the septa may only indicate temporary division of a single channel. Another large
canal lies in the centre below the transverse band; but the occurrence of the mouth soon
obliterates the latter. It is probable that all the canals communicate (as in the Anopla),
and the circulating medium is represented in the preparations by a minutely granular
substance.

The occurrence of the special layers of separated longitudinal fibres in the new form is
therefore a feature of interest; and since they are confined to the head and cesophageal
region of the body, it is probable they aid the animal in its progress amongst the sand.

Within the great longitudinal muscular layer of Valencinia Armandi is superiorly the
cireular muscular coat of the proboscidian sheath, which divides opposite the upper part
of the nerve-trunks (fig. 5, os), and sends a strong band along the ventral surface of the
cesophageal region of the alimentary canal. Between the latter band and the chief longi-
tudinal layer is the great vascular channel (v) on each side.

The external circular muscular layer (¢) shows, some distance behind the mouth, a
tendency to form superiorly two distinct inner bands (fig. 5, ce), extending from one
nerve-cord to the other; and a little further backward the differentiation is observed all
round the body. The circular coat also increases in thickness; but little further change
ensues in this layer in its course backward, except the alteration of its relations to tke
nerve-trunks, which pass from its outer border into the great longitudinal layer.

The great longitudinal muscular layer forms a continuous investment, indented by the
two small vascular canals (vs) superiorly, and the great vascular channels (v) inferiorly
on its inner side, and by the great nerve-trunks externally. As we proceed backward a
tendency to enlargement is noticed at the regions above and below the latter, the coat

* Dredged by Mr. Whiteaves at various points in the Gulf of St. Lawrence.

W. C. M‘INTOSH ON VALENCINIA ARMANDI. vi

generally gets thicker, the median raphe superiorly and inferiorly more distinct, and the
fasciculi more regularly arranged. Then the nerve presses more and more on the circular
coat, penetrates it, indents the longitudinal layer deeply, and finally is included in its
outline (fig. 6, 7), though not at first surrounded by its fasciculi. The next step (which
oecurs a little further backward) is the increase in the relative thickness of this coat
on the disappearance of the great internal circular layer.. The dorsal region dimi-
nishes (in transverse section) to a blunt point at the raphe (fig. 7, 7), forms a consider-
able enlargement at the side of the proboscidian sheath, becomes broad and prominent
internally near the lateral nerve-trunk (which it now encloses), contracts a little again
below the nerve, and increases largely towards the inner edge of the vascular canal (this
being its thickest part), and, lastly, diminishes rapidly towards the ventral raphe. The
arrangement of the fasciculi has now become very regular, viz. in the form of long
rows running, chiefly in pairs, from without inward (fig. 9, d), and which fairly indicate
a stage in the penniform condition of the muscles, observed, for instance (first by M.
Marion), in Amphiporus spectabilis, De Quatref. (Drepanophorus, Hubrecht).

In Valencinia lineiformis the arrangement of the muscular fasciculi in the same region
(from the great vascular trunk downward) takes the form of long narrow rows
(fig. 10, d), quite different in appearance from those in the present form. Dorsally the
rows in VY. Armand: are for the most part directed downward and outward on each side
(fig. 7), then downward and inward, nearly horizontal above the nerve-cord, irregularly
arranged round the latter, and with the chief mass external to the trunk, then upward
and inward below the nerve, and upward and outward from near the vascular canal to the
ventral raphe. The whole field, therefore, has a striking appearance; and the arrange-
ment continues, with little change, to the posterior region of the body. At the tail,
again, the layer becomes somewhat thinner, and the diminution of the proboscidian
sheath causes a greater proportional increase near the dorsal median line, while the nerve-
trunks have the fasciculi only on their inner side.

To comprehend the next muscular layer (the inner circular iayer, fig. 6, 0s), it is
necessary to examine the commencement of the proboscidian sheath. Just in front of
the mouth the sheath (fig. 4, a0) is complete—that is, has externally a strong layer of
circular fibres, and internally a coat of more slenderly developed longitudinal fibres. The
inferior part of the sheath rests (over the mouth) on a series of longitudinal muscular
fibres (z). Behind the mouth a very noteworthy change rapidly takes place, viz. the
gradual absoption of the contiguous portion of the proboscidian sheath into the upper
wall of the digestive chamber, and the envelopment of both by the strong inner circular
layer. The latter is first formed by a splitting of the outer (circular) fibres of the pro-
boscidian sheath (fig. 5, os); and while the coat remaining between the sheath and the
digestive canal gradually diminishes, this new circular belt soon envelops the united
area in a dense muscular mass. Indeed so little of the thin partition between the
sheath and the digestive wall is visible, that at first sight it seems as if both chambers .
floated in a common cavity. After a considerable course this special inner circular layer
diminishes, the proboscidian sheath gets thicker, and its proper layers more evident, and
finally the former disappears, while the latter (fig. 7, a0) continues to the posterior end

M 2

78 W. C. M‘INTOSH ON VALENCINIA ARMANDI.

of the worm. The greatly developed layer of circular muscular fibres must exercise a
special influence on this region of the alimentary canal, probably as an cesophageal con-
strictor. The arrangement is one common in the Carinellide.

The proboscidian sheath (a 0) commences, as usual, at the incurvation in front of the
ganglia, and, as formerly described, becomes more or less atrophied behind the mouth,
while the great inner circular layer developed from its outer fibres acts as the chief
constrictor for this chamber as well as the alimentary canal. At first (figs. 3 & 4, ao) it
clearly possesses two muscular layers, an external circular and internal longitudinal ; but
after the change behind the mouth is completely established, all that can be noticed
inferiorly is a circular band in the tissue of the upper layer of the digestive chamber. A
reversion to the primary condition ensues on the diminution of the inner circular
muscular layer, the proboscidian sheath striking off from the latter and the alimentary
wall as a special tube, which is continued to the posterior end of the worm, where,
however, it forms an extremely small chamber (fig. 8,a0). In Carinella annulata it is
not continued to the posterior region of the body, and, as formerly observed, absence
or diminution of the proboscidian chamber (é.e. the cavity within the sheath) in the
Lineidz and Carinellide is accompanied by greatly enlarged lateral vessels.

The proboscis (figs. 4 & 7, A) agrees with that of the Carinellide ; but its very minute
size prevented an accurate discrimination of the external region into two layers. It had
the aspect of an elastic coat with external circular fibres. A reexamination of more
favourable examples of the proboscides of the Carinellidee has enabled me to correct an
error into which I fell on a former occasion, in stating that the external coat consisted
of elastic, and the next of circular muscular fibres. The latter are external, and the
structureless elastic internal. The next layer in Valencinia Armandi consists of a
powerful longitudinal muscular coat ; then comes a considerable circular muscular layer,
upon which the glandular lining of the organ rests. The glandular papille, as usual,
contain many rod-like bodies. The proboscis terminates posteriorly in a ribbon of lon-
gitudinal fibres.

Digestive System.—The mouth (in transverse section, fig. 3, w) forms a small lon-
gitudinal slit a little behind the ganglia, and is much less prominent than in /. linei-
formis, as well as more anterior in position. It leads into the glandular cesophageal
region, which soon becomes clasped by the powerful internal circular muscular layer
formerly alluded to (figs. 5 & 6,7). The walls of this region are thinner, but of firmer
consistence than the succeeding portion, which commences near the termination of the
special circular muscular layer. In transverse section (fig. 7, 7’) the densest masses of
the refracting (fatty ?) granules, which form the chief portion of the wall, lie on each side
above the nerve-trunks, while a layer beneath the proboscidian sheath joins them supe-
riorly, and a V-shaped bend ventrally completes the circuit. Towards the posterior end
of the body (fig. 8, 7’) the canal assumes the form of an ellipse in transverse section, the

- walls are lax, translucent, and more finely granular, and the cilia appear to be longer.
It is held in position in this region by circular fibres and vertical bands.

Nervous System.—There is nothing special in the structure of the ganglia, which, near

their commissures, are seen in fig.3, h; but the presence of a special longitudinal series

W. C. M‘INTOSH ON VALENCINIA ARMANDI. 79

of separated muscular fasciculi (d’) between them and the basis-layer is a new feature in
the Carinellide. In regard to the nerve-trunks on each side, it is an important fact that
while (in common with the ganglia) they lie clearly outside the proper circular and lon-
gitudinal muscular layers of the body-wall anteriorly, they gradually perforate both in
their progress backward to take up a position in the middle of the longitudinal layer.
Towards the tail the nerve-trunk again gets to the outer border of the latter, and almost
touches the circular coat (fig. 8, 2); but it does not seem to get quite clear of the lon-
gitudinal fibres, some of which form a thin line externally.

In regard to the vascular system, the general features are those of the Carinellid.
There are two great longitudinal trunks (v, v) ; placed within the longitudinal muscular
layer of the body. In the csophageal region (figs. 5 & 6) the trunks agree with
the arrangement in the common Carimella—that is, are situated outside the special
inner circular muscular layer. In the large Canadian Carimella, again, they lie within
this special layer, between it and the wall of the cesophageal region of the alimentary
tract. Anteriorly the trunks terminate in the new species in the vascular channels on
each side of the proboscidian canal, those of opposite sides: communicating in front. The
arrangement, indeed, forms a kind of vascular mesh-work towards the tip of the snout;
and the contents probably exercise an important influence on the mobile tissues of the
region when driven forward by the contraction of the great lateral trunks. At the
mouth (fig. 4) the latter lie nearly opposite the nerve-trunks on each side internally ; in
the anterior region of the body each vessel descends to a lower plane (fig. 6), while pos-
teriorly (fig. 8) the distance between the nerve and vessel greatly increases, the latter
having advanced inward almost to the wall of the centrally situated alimentary canal.

Two other accessory systems yet remain to be described. The first of these consists of
a vessel with highly contractile walls (va, fig. 5), which lies anteriorly within the pro-
boscidian sheath, just above the point where the wall of the digestive canal meets it on
each side. It begins immediately after the mouth, and extends backward only a short
distance ; but its connexions anteriorly and posteriorly have not yet been ascertained.
There can be little doubt about the nature of this structure, which is quite different
from the collections of proboscidian corpuscles and coagulated fluid often seen in similar
positions. The wall of each vessel is definitely formed of a hyaline contractile substance,
in which numerous granules are imbedded. The physiological importance of this arrange-
ment is apparent, and it is especially interesting since the discovery of a regular series of
canals into the cavity of the proboscidian sheath of Amphiporus spectabilis (Drepano-
phorus, Hubrecht). The second system consists in the establishment of a channel on
each side underneath the great longitudinal muscular layer (vs, figs. 4, 5, & 6), as soon
as the upper arch of the proboscidian sheath is formed. These channels assume con-
siderable dimensions, but do not appear to be very contractile, since they have only
a hyaline film all round, and thus stand in marked contrast to the great vascular
channels inferiorly, or even the intra-proboscidian trunks above mentioned. ‘They have
a lengthened course, but disappear towards the posterior end of the body.

None’ of the examples contained reproductive organs; so that this department in
the group is still in need of further investigation. In the large Carinella from Canada.

80 W.C. M‘SINTOSH ON VALENCINIA ARMANDI.

the ovaries are filled with large ova, and stretch from a little below the nerve-trunks on
each side upward to the dorsal region of the great longitudinal muscular coat, and
encroach considerably on the alimentary chamber behind the proboscidian sheath,

EXPLANATION OF PLATE.
Prats XVI.

The following letters are used to designate the respective parts throughout the series :—

a. Cutis.

b. Basis-layer of the cutaneous system.

c. Cireular muscular coat of the body.

d. Longitudinal muscular layer of the body.
ao. Proboscidian sheath.

d’. Special layer of separated longitudinal muscles anteriorly.
Ah. Ganglia.

j. Wsophageal region of the digestive system.

j.. Alimentary canal proper.

mn. lateral nerves.

r. Median raphe.

v. Great longitudinal vascular trunks.

w. Mouth.

A. Proboscis.

Fig. 1. Transverse (vertical) section near the tip of the snout of Valencinia Armandi: v, v, vascular

meshes by which the great trunks unite. x 210 diam.

Fig. 2. Transverse section of the snout behind the former. The two horizontal bands of fibres (e and e’)
have now attained considerable development ; and numerous vertical fibres pass from their sur-
faces to the circumference: ao, canal for the proboscis in the snout. Numerous longitudinal
muscular bundles (d) occur along the inner faces of the horizontal bands and in the forks of the
vertical series. x 210 diam. i

ig. 3. Transverse (and somewhat flattened) section just in front of the mouth, through the ganglia (A).
The separated series of longitudinal muscles (d’) are very conspicuous in this region, though a
process of the mouth (w) interrupts the circuit inferiorly. The proboscidian sheath (ao) is some-
what flattened out. The two accessory vascular meshes, s, s, are probably homologous with
those found in a similar situation in the typical Anopla; m, powerful transverse muscular band,
separating the chief from the accessory vascular channels ; m’, ring of circular and longitudinal
fibres surrounding the united area of the proboscidian and the vascular systems. In this pre-
paration the proboscidian sheath is differentiated from the general investment of the united area

les]
=
Q

very distinctly. x 90 diam.

Fig. 4. Similar section through the mouth. The separated series of longitudinal muscles (d’) are still
very evident. They form aconsiderable mass on each side of the mouth inferiorly. The basis-
layer (b) of the cutaneous system is greatly developed im this region. The proboscidian sheath
is wholly independent of the digestive system, a special band of longitudinal fibres (z) besides
the sheath proper and other tissues intervening. The commencement of the great longitudinal

W. C. M‘IINTOSH ON VALENCINIA ARMANDI. 81

layer of the body-wall is seen at d; vs, vs, the two dorsal accessory vascular channels between the
proboscidian sheath and the longitudinal muscular coat. x 90 diam.

.5. Section through the anterior esophageal region, showing the intraproboscidian vessel, va. The
proboscis, A, has been cut obliquely. The dorsal accessory vascular channels, vs, are very
distinct. The cutaneous basis-layer, 6; shows traces of the separated series of longitudinal fibres,
but they are insignificant. At os is a slight inflection of the inner circular coat, indicating the
point of separation of the thin layer still representing the proboscidian sheath. x 90 diam.

Fig. 6. Section through the internal circular muscular layer, os, in its complete condition. x 90 diam.

Fig. 7. Transverse section behind the esophageal region, and after the disappearance of the special inner
circular muscular layer (os, fig. 6). The great development of the longitudinal muscular layer of
the body is apparent, as well as the regularity of its arrangement; o/, inner (longitudinal) fibres
of the proboscidian sheath ; oc, outer (circular) of the same. x 90 diam.

Fig. 8. Section about one fifth of an inch from the tip of the tail, somewhat flattened by compression. The
proboscidian sheath (ao) is now very minute. The great longitudinal muscular layer (d@)
presents no regular arrangement of its fasciculi; and the lateral nerve-cord has almost reached
the circular coat (ce) exteriorly. The area on each side of the now attenuated alimentary

Fj

a
Q

So]

canal is filled up by a translucent fibrous stroma, which in this preparation has been much
stretched, especially on the left side. x90 diam.
Fig. 9. Portion of the body-wall towards the middle of V. Armandi, from the great lateral vessel outwards,
showing the arrangement of the muscular fasciculi of the longitudimal layer. x 350diam.
Fig. 10. Section of a similar portion in Valencinia lineiformis. The great thickness of the basis-layer is
conspicuous. The arrangement of the longitudinal muscular fasciculi also contrasts with the
former. x 350 diam.

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SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART THE THIRD.

1 OMNGDE OR INE:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET
SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXVI.

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CONTENTS.

PART III.—Jouty, 1876.

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. KELAART,
F.L.S., at Trincomalee, and partly by Dr. Couuinewood, F.L.S., m the Hastern
Sco o Gera COULINGWOOD) |). eey ia eles Sereda oss ey) ePAgserss

VII. On the Structure and Development of the Bird’s Skull. By W. K. ParKer,
ERAS LS Gio (UR ALbe DI) is oe Pe a ee Me eg SOD

[ 83 ]

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. Kevaart,
F.L.S., at Trincomalee, and partly by Dr. Couinewoop, E.LLS., in the Eastern
Seas. By Dr. CoLLINGwoop.

(Plates XVII. to XIX.)

Read March 18th, 1875.

Convenrs.
I. Prefatory and introductory. IV. Description of twelve new species of marine Pla-
II. Characters of two new genera of Kuryleptide. narians collected by Dr. Collingwood, and deli-
III. Synopsis of the genera and species described and neated in the accompanying figures.
figured in the present paper. Y. Dr. Kelaart’s descriptions of his figures of marine
Planarians collected in Ceylon.
J.

THE species of Planarians which I am about to lay before the Society were, as far
as I was concerned, the fruit of many searches upon rocky shores of the coast of China,
Borneo, Singapore, &c., and represent a considerable amount of labour near low-
water mark in turning over stones, pieces of coral, &c. in such situations. The little
animals in question, although extremely interesting, are all more or less minute; and it
requires not only a sharp but an educated eye to discover them; for so delicate are they,
and often so thin in texture, that out of the water, and adhering to a more or less muddy
or weed-grown stone, it is by no means easy to detect them. It must not be under-
stood, however, that these searches were unrewarded except by the comparatively small
number of specimens or new species here recorded. Many other animals of interest
divided the attention, as, for instance, nudibranchiate Mollusca, of which a number
of new species await the notice of the Society, and other no less interesting creatures.
The Planarians in many respects rival the Nudibranchs in gracefulness of form and
beauty of colouring; but they are even more delicate than the latter. When a small
spot of colour upon the under surface of a stone betrays the presence of one of these
little animals, a camel’s-hair brush is necessary to dislodge it safely ; and a bottle of salt
water must be at hand, into which to transfer the prize.

It may be thought that a dozen species is but a small number to reward one’s labour
on coasts which are supposed to abound with unknown forms of animal life. But
a good deal of experience in shore-hunting on our own coasts prepared me for the
fact, which I found amply corroborated on foreign shores, that we must not look for
a large number of species at any one locality, unless the same locality is examined many
times, and at different seasons of the year—conditions not always, or, indeed, often
practicable. These little organisms appear to be sporadically distributed over wide
areas. ‘They are seldom met with gregarious; and three or four species captured upon
any given occasion might be reckoned a prize more than ordinarily satisfactory.

SECOND SERIES.—ZOOLOGY, VOL. I. N

.

84: DR. COLLINGWOOD ON THIRTY-ONE

Although, however, I never lost an opportunity of examining a shore when I could do
so towards low tide, the opportunities for such rambles were necessarily limited.

And yet, although they appeared to be so sparsely distributed, a fact would come to
light every now and then which seemed to point out that they were very abundant in
the aggregate. For instance, a single visit to a certain spot would perhaps result in the
discovery of some minute organism identical with one found on another occasion at a
point separated perhaps by a hundred miles from the first-mentioned locality, or, as was
the case in some instances, by the whole extent of the China Sea. How was it that
I should stumble upon such an inconspicuous organism a second time at a spot so
distant from the first, unless they were abundantly distributed through the inter-
mediate spaces? Doubtless an explorer at the same places would still find many new
forms, and leave yet many more for a successor.

With regard to the Planarians collected by the late Dr. Kelaart, and included in this
collection, it is necessary that I should say a few words concerning how they came into
my possession. The late Mr. Albany Hancock, of Newecastle-on-Tyne, hearing from me,
in the course of correspondence, that I had collected some marine Planarians which
appeared to be new, informed me that he had in his possession a number of drawings
of similar animals made by Dr. Kelaart (formerly a Fellow of this Society) from speci-
mens collected by himself in Ceylon, and principally at Trmcomalee. These drawings
were briefly characterized by him in a paper “ On some new and little-known Ceylon
Invertebrata,’ published in the Journal of the Ceylon branch of the Royal Asiatic
Society. Part of this paper was printed in the ‘ Annals of Natural History’ for 1859;
but the portion relating to the Planarians was not so published; nor, need I say, were
the drawings published at all. Dr. Kelaart issued a ‘ Prodromus Faune Zeylanice,’ a
portion of which is in the Society’s Library, and which contains the description of the
Planarians here figured ; but I have nowhere met in the literature of the Planarians
with any reference to either of these publications, which appear to be unknown.
Mr. Hancock therefore suggested that if I could publish Kelaart’s species along with my
own, they would, as he expressed it, ‘‘ make a very interesting monograph on the
subject—would, in fact, make a valuable contribution towards the history of these little-
known animals of the eastern seas.”’ He therefore sent me the drawings, which, re-
garding them as a testament jointly from Dr. Kelaart and Mr. Hancock, both deceased,
I have accordingly submitted to a careful examination, and done my best to distribute
them according to the genera and species of the best classification. I may say that,
with Mr. Hancock, I consider these drawings are sufficient for identification; but they
are unaccompanied by details, and the descriptions of them are brief. This is to be
regretted, since the value of such drawings depends no less upon the accuracy of the
verbal descriptions taken from the living animals than upon the correctness and minute-
ness with which they are depicted. Unfortunately the animals are so extremely
perishable and delicate, that no means yet devised can possibly exhibit them after
death in any thing approaching to their natural appearance or beauty of form and
brilliancy of colouring. Good drawings and accurate descriptions, therefore, are of the
utmost value and importance; and either one without the other is consequently of

SPECIES OF MARINE PLANARIANS. 85

impaired value. Of my own drawings I can, of course, vouch for the faithfulness. They
were always taken from the living animal while in health and activity ; and any thing
striking and remarkable could not fail to be noticed and recorded.

The term Planaria, which was at first applied to these animals as a generic name for
them all, has been long since restricted to a small division of them which is not marine.
Linnzeus, Pallas, Lamarck, and Cuvier placed them among parenchymatous Entozoa ;
but the investigations of Dugés, von Baer, Quatrefages, and others have by degrees raised
them to their proper and natural position. This is now assigned to them under the
Turbellarian Annelids. The genera described by Ehrenberg and De Quatrefages out of
the old genus Planaria have in turn for the most part been split up or given place to
more recent classification. Ihave endeavoured to assign positions to the present species
under the arrangement of Diesing (‘ Systema Helminthum’), as revised by W. Stimp-
son in his ‘‘ Prodromus descriptionis animalium evertebratorum” of the South-Pacific
Federal Expedition, under Capt. Jno. Rodgers (Proc. Acad. Nat. Scien. Philad. 1857,
p- 19 &c.). Stimpson enumerates thirty-eight genera of Turbellaria Dendroccela (Max
Schultze, Wiegm. Arch. 1849, p. 292), including several new genera resulting from that
expedition. Of these thirty-eight genera, eight are fluviatile, four terricolous, one inhabits
brackish water, and oneis pelagic. The remaining twenty-four consist of ordinary marine
genera, under which group the present species would be all included. I have ventured
to constitute a new genus of two of the Ceylon species, from a character which does not
appear to be shared with any of those previously described. Inasmuch, however, as all
the Ceylon species have already been named by their collector and delineator, I shall
of course adopt his specific terms, only inserting them in what appears to me to be their
proper generic group.

The marine Turbellaria Dendroceela are for the most part flat, thin, soft, and delicate
animals, remarkable for their beautiful colours and often graceful forms and movements.
They have usually a pair of tentacles, these occasionally being mere folds or ear-like projec-
tions of the anterior margin (Huryleptidze)—sometimes true tentacles, situated occipitally
(Stylochidz), or even dorsally, in which case they are retractile (Planoceridz). In some
families, however, tentacles are entirely absent (Typhloleptidee and Leptoplanidee). The
margin is usually ample, and much folded or puckered, which is very noticeable when
the animal is at rest; but when in motion, this ample pallium is gracefully waved
in a vertical direction, and aids greatly such species as are natatory in swimming,
which they effect with ease and rapidity. They possess either two genital apertures (sub-
tribe Digonopora), as is the case with all the present species, or only a single one (sub-
tribe Monogonopora), as is the case with the true Planariade. The mouth is variously
situated on the median line of the under surface, and is in turn antecentral, subcentral,
or postcentral; and the dendriform gastrovascular ramifications are often more or less
visible upon the under surface.

One of their most peculiar characteristics consists of the eye-spots, small black specks
of varying size and position, sometimes clustered into congeries of a symmetrical form,
and sometimes scattered over the anterior and marginal portions of the upper surface, or
on the tentacles. These eye-spots, in some cases at least, consist of a highly refracting

N2

86 DR. COLLINGWOOD ON THIRTY-ONE

spherical body (crystalline), surrounded with pigment; and they are always situated in a
part which is free from the usual pigmentary matter of the animal, and where the skin
is reduced to a minimum of thickness (Baer). In any case they can but very imperfectly
perform the function of vision; and in one family (‘Typhloleptide) they appear to be
altogether absent *. Nevertheless these animals all seem to avoid the light, by concealing
themselves under stones and in situations where no light can reach them.

Their ova are deposited in thin flakes upon sea-weeds and stones, and in appearance are
not unlike those of the Nudibranchs, for which at first sight they might be mistaken.
They resemble these mollusks also in their habit of floating, back downwards, upon the
surface of the water. Nor do the resemblances end here ; for some genera of the Hury-
leptidee are provided with numerous papillze, more or less covering the whole upper sur-
face, after the manner of an Holis, which character earned for them the generic name
of Holidiceros from De Quatrefages.

It has often been said, and is copied from one book into another, that these animals
have the power of spontaneous fission, by means of which two individuals are produced
from one. I can only say that, of the numerous specimens I have met with, kept alive,
and carefully watched, I never saw any thing like this process take place in any instance.
Still I cannot assert that it does not happen.

It is worthy of remark that among the 19 species of Dr. Kelaart, no less than 11
appear to be referable to the genus Hurylepta (Hemp. and Ehrenb.), while of my own 12
species only one is of that genus. Mot one of Kelaart’s claims rank under Quatrefages’
old genus Proceros, in which one fourth of my own species oceur. In each collection is
found a new genus of Euryleptidz ; and the whole of the 31 species are included under
one subtribe—Digonopora, or monecious Turbellaria.

If.

(Characters of two new genera of Burylepide.)

TURBELLARIA DENDROCGLA.

Genus ACANTHOZOON, Coll.

Caput subdiscretum, tentaculis parvis approximatis. Corpus supra spinulis brevibus nigris ubique
imstructum.
Thysanozoon,

ACGANTHOZOON, Proceros,
Hurylepta.
Genus SPHYNGICEPS, Coll.

Corpus leve, caput discretum, tentaculis magnis subdistantibus ; ocelli occipitales et capitales.

Proceros,
SPHYNGICEPS,
Eurylepta.

* With regard to the eye-spots, structurally and as supposed organs of vision, consult H. N. Moseley’s remarks
thereon, ‘‘ Anat. and Histol. Land-Planarians of Ceylon,” Proe. Roy. Soc. 1873, p. 172 &e.

SPECIES OF MARINE PLANARIANS. 87

TE
(Synopsis of the Genera and Species described and figured in the present Paper.)

[Note—Diesing’s main divisions were founded upon the presence or absence of tentacles ; but Stimp-
son pertinently remarks that such a division separates allied forms, such as Stylochus and Leptoplana,
while widely separated ones, such as Hurylepta and Planaria, were approximated. In view of this I
therefore follow Stimpson in foundmg the subtribes upon the presence or absence of a second genital
aperture, instead of the mere presence or absence of tentacular appendages.—C. C.]

TURBELLARIA DENDROCGLA.
(Tubus alimentarius dendritice ramosus.)
Subtribus I. Digonovora. Two genital apertures.

(All the species are included in this subtribe.)

Family EURYLEPTID&.

Genus THYSANOZOON, Grube.

1. Thysanozoon Alderi, Coll. Labuan.
2. Thysanozoon Allmanni, Col/. Singapore.
3. Thysanozoon auropunctatum, Kel. Aripo.

Genus AcANTHOZOON, Col'., n.g.

4, Acanthozoon armatum, Kel. Ceylon.
5. Acanthozoon papilio, Kel. Ceylon.

Genus SpHyNneicers, Coll., n. g.

6. Sphyngiceps lacteus, Coll. Singapore.

Genus Proceros, Quatref.

me

. Proceros concinnus, Coll. Labuan.
. Proceros Hancockanus, Coll. Singapore.
. Proceros Buskiu, Coll. Singapore.

<> OO

Genus Hunyuepra, Hemp. & Ehren.

10. Eurylepta fusca, Kel. Ceylon.

11. Eurylepta atraviridis, Kel. Ceylon.
12. Hurylepta undulata, Kel. Ceylon.
13. Hurylepta violacea, Kel. Ceylon.
14. Eurylepta dulcis, Kel.

15. Eurylepta purpurea, Kel. Ceylon.
16. Eurylepta viridis, Kel. Ceylon.
17. Eurylepta affinis, Kel. Ceylon.

18. Eurylepta cerebralis, Kel. Ceylon.
19. Eurylepta striata, Kel. Ceylon.
20. Eurylepta zeylanica, Kel. Ceylon.
21. Eurylepta Kelaartu, Coll. Simgapore.

88

DR. COLLINGWOOD ON THIRTY-ONE

Family TY PHLOLEPTID &.

Genus TYPHLOLEPTA, Oerst.

. Typhlolepta Byerleyana, Coll. Pulo Barundum, west coast of Borneo.

Family LEPTOPLANIDA.

Genus CenTRostomum, Diesing.

. Centrostomum ocellatum, Kel. Ceylon.
. Centrostomum punctatum, Kel. Ceylon.

Genus EiasmopEs, Le Conte.

. Elasmodes obtusus, Coll. Singapore.

Genus LepropLana, Hemp. et Ehr.

. Leptoplana patellensis, Co//. Simon’s Bay.
. Leptoplana aurantiaca, Coll, Singapore.

Family STYLOCHID &.
Genus STYLOCHOPLANA, Stimpson.

. Stylochoplana elegans, Kel. Ceylon.
. Stylochoplana meleagrina, Kel. Ceylon.

Genus StyLocuopsis, Stimpson.

. Stylochopsis malayensis, Coll. Pulo Barundum, west coast of Borneo.

Family PLANOCERIDA.

Genus PLanocera, Blainv.

. Planocera thesea, Kel. Ceylon.

IV.

(Descriptions of the Marine Planarians collected in the Eastern Seas by Dr. Colling-

wood, F.L.S., and delineated in the accompanying figures.)

1. Thysanozoon Alder.

Length 2+ inches; breadth 14 inch.

Body thin, with very irregular margin, amply folded and puckered.
Upper surface of a general light brownish colour, with a narrow, pale external margin,

within which is a broad, black border, somewhat shaded and marbled. Down the
median line for about three quarters of its extent runs an irregular, black marbling ;
a faint marbled pattern of pale brown is diffused over the general surface; and a
lens discloses also a fine ramification of a darker tinge throughout.

The whole upper surface is studded with small papillee of a conical form, the foot-

stalks of which are pale, and the distal extremities orange. Many of these papille

SPECIES OF MARINE PLANARIANS. 89

arise from an elevated white spot or tubercle, such tubercles producing only one
papilla each; and other papille exist upon the black margin, as well as on the
general surface.
Under surface whitish, edged with black, the part answering to the black marbled line
on the dorsum being here opaque white.
‘Head blackish, angular, raised somewhat above the general plane of the body, flexible,
and having two projecting angles or folded tentacles.

This was a very beautiful and striking animal, looking as if studded with golden beads.
it swam freely, with a graceful vertical vermicular movement of the ample sides.

One was found under stones about two feet under water at low tide, upon a reef of
the island of Labuan, coast of Borneo, on August 22. I have great gratification in
naming it after the late Mr. Joshua Alder, of Newcastle, a gentleman with whom I was
long in correspondence, and from whom, though we never met, I always received
uniform kindness and urbanity.

Fig. 1: a, general appearance of upper surface; 6, head and tentacles; c, portion of upper surface,
enlarged, showing papille; d, a papilla; e, a papilla-bearing tubercle.

2. Thysanozoon Allmani, Coll.
Length 23/5 inches ; breadth ? inch.

Body translucent, papillose.

Upper surface light brown, becoming darker towards the margin, and with an irregular
edging of opaque white all round, excepting the head. An elevated ridge runs
along the median line of the dorsum.

‘The whole upper surface is covered with clavate and pointed papille, of a deep
brown colour, and varying in size, the smallest being the lightest-coloured and most
numerously clustered, and occurring along the median ridge.

Under surface grey, darkening to deep brown at the sides, and edged with opaque
white.

Head with two long tentacles, often thrown back, and presenting the appearance of
hare’s ears. Tentacles dark brown, tipped with white. Twominute white tentacles
are situated in front of the head, beneath the hare-like ones.

Hye-spots situated in a light-coloured spot immediately. posterior to the head, in a double
cluster, consisting of two small crescentic patches of minute black spots.

A splendid species, the edges much corrugated, unless the animal is in full movement,
when the folds almost disappear. It swims freely, with a graceful serpentine movement,
and crawls with moderate rapidity and with an undulating movement. When at. rest
the animal assumes a nearly circular form, with numerous marginal folds.

Two specimens found at Singapore, west of the harbour, under pieces of dead coral,
on the beach between tide-marks, Nov. 22nd.

I have much pleasure in connecting this species with the name of the President of this
Society, so well known for his researches among the Invertebrate division of animais.
Fig. 2: a, animal in movement (dorsal surface) ; 6, papille, large and small, magnified; c, eye-spots

(magnified).

90 DR. COLLINGWOOD ON THIRTY-ONE

3. Sphyngiceps lacteus, Coll.

Length 4% inch ; breadth { inch.

Body graceful in form, semitransparent.

Upper surface cream-coloured, irregularly spotted with sparse and minute black dots,
and having faint marbling on either side of the median ridge, which is very con-:
spicuous. Margin irregularly blotched with red, and the whole body edged with a
narrow black line.

Under surface whitish, edged with red blotches and a black streak as on the upper side,
but somewhat less distinct. A broad streak of white occupies the anterior third
of the median line.

Head small and narrow, but rendered very conspicuous at times by its being raised up,
so that the median ridge is thus very much elevated in front. Head furnished
with two folded tentacles.

Eye-spots round, immediately posterior to the head, which has in front of it a pair of
larger single spots.

This is a remarkably graceful little animal, crawls with great rapidity, and with a
gliding movement, then suddenly throws its head back, and, leaving the side of the vessel,
swims with a graceful serpentine movement, sometimes very fast ; floats also on its back
on the surface of the water. It has the habit of throwing back its head very far, and
remaining in that position (rampant, as it were), at which time the head and anterior
portion of the body become narrow and tubular, giving the appearance of a neck. When
at rest it expands laterally, becoming in shape as broad as long.

One specimen found under a coral block, west of Singapore Harbour, at low water, on
Noy. 22nd.

Fig. 3: a, animal in motion; 0, head, enlarged, showing eye-spots.

4. Proceros concinnus, Coll.
Length 2 inch.

Body narrow, entire, without marginal foldings.

Upper surface cream-colour, approaching to yellow, with an edging of blue all round,
composed of small and larger spots running into one another. A similar blue streak
runs along the median line from a little behind the head to some distance from the
posterior extremity, through about three quarters the length.

Under surface cream-colour, similarly edged with blue, a brownish streak running
through the whole median line, the arbusculiform alimentary tube of a straw-
colour, occupying the anterior third. .

Head with two folded tentacles.

Eye-specks conglomerated in a small round spot midway between the head and the blue
median line.

A very pretty and lively species, particularly fond of floating on its back or crawling
rapidly along. I obtained one specimen on a reef at Labuan, near low-water mark, and

SPECIES OF MARINE PLANARIANS. 91

a second at Pulo Daak, a small island between Labuan and the mainland of Borneo,
August 25th.

Fig. 4: a, animal extended, upper surface ; 4, animal extended, under surface; c, eye-spots.

5. Proceros Hancockanus, Coll.
Length 1,45 inch; breadth ;8; inch.

Body velvety, opaque.

Upper surface of a deep velvety brown, edged with a double margin of equal widths, the
inner deep orange, the outer opaque white. Along the centre of the back was a
slightly elevated ridge.

Under surface grey, darkening towards the sides.

Head small, tentacles simply folded, long and graceful, the orange margin disappearing,
and the white alone being prseent.

Hye-specks in an oval elongated cluster, immediately posterior to the head.

A very beautiful species, no less from its rich colour than from its active movements.
Swimming by an undulating movement, it also floated on its back on the surface of the
water, after the manner of the Nudibranchiata. It crawled slowly, and with an undu-
lating motion. In texture it appeared somewhat translucent, and when suddenly dis-
turbed became much paler, as though by a sudden contraction of the tissues. In one
specimen a temporary slit appeared, occupying the middle third of the dorsal ridge,
rounded anteriorly, wedge-shaped posteriorly, through which slit the internal organiza-
tion was clearly visible. The slit remained open one day, and afterwards closed. The
intestines appeared sometimes as though protruding through this slit, which was not
exhibited by a second specimen. On placing one of these animals in glycerine, it imme-
diately discharged a quantity of a brownish-coloured fluid, and contracted considerably
in bulk.

Two specimens were obtained on succeeding days from among stones and old coral
blocks, between tide-marks, west of Singapore Harbour, Nov. 21st.

This species I have much pleasure in connecting with the name of the late Mr. Albany
Hancock, the coadjutor of Alder, from whom, although we never met, I received many

| kindly letters up to near the time of his lamented decease.

Fig. 5 : a, the animal in motion, exhibiting the open dorsal fissure; 4, the cluster of eye-specks.

6. Proceros Buskii, Coll.
Length 4 inch; breadth } inch.
Body opaque, flat, smooth.
Upper surface rich velvety olive-green, edged with pale yellow. :
Under surface dark grey, the dendritic marking whitish, and occupying the anterior half
of the median line.
Head with two folded earlike antennal projections.
Eye-spots in a circular cluster, difficult to detect owing to the dark colour of the animal,
and situated upon a ridge formed by the elevation of the antennal head.
SECOND SERIES.—ZOOLOGY, VOL. I. fc)

92 DR. COLLINGWOOD ON THIRTY-ONE

A very handsome active species, crawls rapidly, at the same time throwing back its
head, and floats, back downwards, on the surface of the water. One specimen only, found
under a stone, and upon a small grey incrusting sponge, between tide-marks in Singa-
pore Harbour, west of the town, December 3rd.

I have named this species after my friend Mr. Busk, than whom no one has been
more assiduous or successful in his investigations among the lower forms of Inver-
tebrata.

Fig. 6: a, animal at rest, dorsal surface; 6, head.

7. Eurylepta Kelaartu, Coll.
Length ;45 inch; breadth + inch.
Body small, smooth, thin.
Upper surface mottled dark purple, the mesial line presenting a slightly elevated ridge
of a darker colour.
Under surface the same colour as the upper, but paler.
Head small, with minute ear-like processes.
Hye-spots minute, roundish.

An actively crawling creature, occasionally throws its head back. Two specimens
were found in Singapore Harbour, west of the town, under stones, and apparently feeding
upon a small incrusting sponge. One of these specimens was of a lighter lake colour.
December 3rd.

As out of the 19 species delineated by Dr. Kelaart I have referred no less than 11 to
the genus Hurylepta, while of my own 12 this is the sole one of that genus, I have
dedicated it to him—a compliment to his memory which his industry in this as well as
in other branches of zoology well deserves.

Fig. 7: a, animal in motion; 0, eye-spot, magnified.

8. Typhlolepta Byerleyana, Coll.
Length ? inch ; breadth 2 inch.

Body smooth, thin, and the lateral parts very ample and puckered.

Upper surface beautifully marbled with light-brown rings, including roundish spaces of
a whitish colour, smaller rings being between the interstices of the larger; most
crowded and darkest in colour along the median line, paler and more delicate
towards the sides.

Under surface of a pale grey, the dendritic marking in the centre of an opaque white.

This very beautiful species I obtained from under a piece of coral on Pulo Barundum,
off the west coast of Borneo. Not having a brush at hand, I had great difficulty in dis-
lodging it from the crevices. Its movements were very contorted, and it did not exhibit
much activity. One specimen only was procured, October 6th.

I have named this elegant species after my friend Mr. J. Byerley, F.L.S., of Seacombe,
Cheshire, a gentleman who in the hardly-spared intervals of active practice has ex-

SPECIES OF MARINE PLANARIANS. 93

hibited the greatest interest in zoological science, and with great industry and labour
compiled the ‘ Fauna of Liverpool.’

Fig. 8. General appearance of upper surface.

9. Hlasmodes obtusus, Coll.
Length -§;; breadth + inch.

Body thin, delicate, smooth, semitransparent, dendritic.

Upper surface pale brown, with a dark, shaded brown streak along the middle half of the
median line, from which radiate delicate pale brown markings, which fade as they
reach the margin.

Under surface pale, and exhibiting the radiating markings, though fainter.

Head indistinct.

Eye-spots two, somewhat crescentic, situated upon a round, white, transparent space
immediately in front of the median dark brown streak.

A very lively, active, and graceful animal. When in motion and fully extended it has
a broad or truncated appearance anteriorly. It crawls rapidly, floats feet uppermost,
and swims gracefully and rapidly by a vertical movement of the sides of the body, which
is performed by quick and sudden jerks.

One specimen was obtained in Singapore Harbour, west of the town, from beneath a
stone between tide-marks, December 3rd.

Fig. 9. Upper surface of the animal.

10. Leptoplana patellensis, Coll.
Length $ an inch.

Body entire, opaque.

Upper surface cream-colour, smooth, beautifully mottled with rich light brown. A
ridge runs along the median dorsal line, irregularly marked with a darker brown,
from which the general mottling radiates to the margin, where it is palest.

Under surface of an opaque whitish colour, the dendritic marking occupying the middle
third, of an opaque white, and surrounded with dots of the same.

Eye-spots irregular and indistinct, consisting of an oval ring at the anterior part of the
median line, and on either side an irregular patch, that on the left roundish, and on
the right crescentic. ‘These spots when magnified appear roundish, but not circular,
and do not present any regular figure.

This animal moves with a leech-like motion, fixing itself by its anterior and posterior
ends alternately. It showed no inclination to swim or float like most of the Planarians,
nor even to leave the bottom of the vessel of water in which it was contained.

Two specimens were obtained from under the mantle of a large limpet (Patella oculus),
which I knocked off with my foot from the granite boulders of Simon’s Bay, Cape of
Good Hope, May 23rd.

Fig. 10: a, upper surface, animal in motion ; 0, head, enlarged, showing arrangement of eye-spots.

0 2

94 DR. COLLINGWOOD ON THIRTY-ONE

11. Leptoplana aurantiaca, Coll.
Length ;%; inch; breadth ¢ inch.

Body slender, semitransparent, entire.

Upper surface, general colour orange-chrome, with a median ridge of a pinkish colour.
From this ridge radiate a number of minute dendritic processes of a bright chrome-
colour, which approach the margin, where the body becomes perfectly translucent.
Sparse white spots are scattered irregularly over the general surface.

Under surface similar to the upper, only paler, as though from the colour being seen
through the semitransparent body.

Head. The anterior portion of the body is without a distinct head or tentacles, but
notched, and apparently folded; on the left side of the notch appeared a tentaculi-
form process tipped with a black spot, and having also two or three small black
specks in its neighbourhood.

Hiye-specks in a hippocrepiform congeries immediately anterior to the pik median line;
the spots few, and larger than usual compared with the smallness of the animal.

This species moves somewhat slowly, crawling like a slug, and also swims or floats
upon its back, like the nudibranchs, upon the surface of the water.
I obtained several specimens between tide-marks under stone westward of Singapore

Harbour in November. |

_ Fig. 11. Upper surface of the animal.

12. Stylochopsis malayensis, Coll.
Length 13 inch; breadth 3 inch.
Body smooth, folds ample; general colour a rich, velvety, deep brownish black, with a

narrow border of deep chrome, external to which is a second narrow edge of dull
white.

Under surface nearly as dark as the upper, with a central irregular line of rose-colour.
Tentacles large, separate, anterior, supporting ocelli.

One specimen found under a coral block on Pulo Barundum, west coast of Borneo,
between tide-marks, Oct. 6.

Fig. 12. Upper surface of the animal.
Vie

(Dr. Kelaart’s description of his figures of Marine Planarians taken at Ceylon, all of
which, except two, he included under the genus Planaria. They are therefore, in

this list, referred to their proper genera.)
Genus THysANOozoon, Grube.

18. Thysanozoon auropunctatum, Kel.
A large species.

Upper surface, a vich violet brown, darker in the centre, and edged all round with a

‘SPECIES OF MARINE PLANARIANS. 95

border of pure white. Thickly studded with papille, small and conical, the bases of
which are black, the apices golden yellow, and the intermediate band white.
Under surface pale purple, very dark towards the margin all round, but having the
narrow white border as above. :
Head furnished with two small rudimentary tentacles. Mouth situated between the
middle and anterior third.

Found at Aripo, 28th February.

Fic. 13: a, upper surface of animal; 0, papill, magnified.

Genus ACANTHOZOON, Coll., n. g. (see p. 86).

14. Acanthozoon armatum, Kel.
Length 14 inch; breadth 14 inch.
Upper surface of a dark purple colour, covered with short black spines.
Under surface pale purple, smooth.
Tentacles folded, but somewhat distinctly formed.
Fig. 14. Upper surface. j
15. Acanthozoon papilio, Kel.
Length about 1 inch.
Upper surface yellow, covered with small black spines; margin whitish.
Under surface pale yellow.
Tentacles as in last species, black, tipped with white.
Looks very like a butterfly moving in the water.

Fig. 15. Upper surface.
Genus EuryLerta, Hemp. et Ehr.

16. Eurylepta fusca, Kel.
Length 13 inch.
Upper surface dusky brown.
Under surface paler brown.
Tentacles inconspicuous and approximated.
Fig. 16. Upper surface.
17. Eurylepta atraviridis, Kel.
Upper surface dark mottled green, with a darker broad streak through the whole median
line. An edging of pale green runs all round.
Tentacles like the last.
Fig. 17. Upper surface.
18. Hurylepta undulata, Kel.
Length 2 inches.
Upper surface pale yellow, with undulating lines and spots of purplish brown, producing
a marbled appearance. Margin and median line purplish.
Tentacles rudimentary.

Vig. 18. Upper surface.

96 DR. COLLINGWOOD ON THIRTY-ONE

19. Hurylepta violacea, Kel.
Length 1} inch; breadth ? inch.
Upper surface violet purple, edged with bright yellow; median line yellowish.
Under surface rose-coloured.
Tentacles rudimentary.
Ova yellowish.

This beautiful species, in a quiescent state, resembles some variety of Pansy (Kel.). It
appears to be nearly allied to Planaria zebra, Leuck. See Riippell’s Atlas, t. 3. f. 1a, 6.

Fig. 19: a, upper surface; 4, under surface.

20. Eurylepta dulcis, Kel.
Length. 1 inch.

Upper surface light green, minutely spotted with reddish brown; margin white; median
line brown.

Tentacles rudimentary.

Fig. 20. Upper surface.

21. Hurylepta purpurea, Kel.
Length about 14 inch.

Upper surface of a beautiful purple colour.
Under surface paler purple, darkening towards the margin.
Tentacles very small and rudimentary.
Fig. 21. Upper surface.
22. Hurylepta viridis, Kel.

Length about 1} inch.
Upper surface green, spotted with brown; margin darkly grizzled brown.
Under surface paler green. ;
Tentacles rudimentary, small, brown.

This species appears to be nearly allied to Planaria limbata, Leuck. See Riippell’s
Atlas, t. 3. f. 4.—C. C.

Fig. 22. Upper surface.
23. Hurylepta affinis, Kel.
Upper surface purple, with yellow border.
Tentacles very small and rudimentary.
Fig. 23. Upper surface.
24, Hurylepta cerebralis, Kel.
Length 53 inches; breadth 3 inches..
Upper surface of a yellowish brown colour, and minutely streaked with fine wavy brown
lines; border ample, edged with black and streaked with white.
Under surface of a delicate salmon-colour, with a narrow blackish border.
Head with rudimentary tentacles, formed by two folds of the margin; mouth large,
placed on the anterior third of the lower surface; lips white.
Ova greenish white.

SPECIES OF MARINE PLANARIANS. 97

This was the largest specimen observed. Its colour and the ample foldings of the
margin call to mind the appearance of convoluted brain-substance.
Fig. 24. Upper surface.
25. Eurylepta striata, Kel.
Length 24 inches.
Upper surface brownish purple, streaked with brown; marginal folds ample, and edged
with a narrow border of dark brown.
Under surface pale orange-brown, darker towards the margin, and edged with a narrow
border of brown.
Fig. 25. Upper surface.
26. Eurylepta zeylanica, Kel.
Length 23 inches; breadth 13 inch.
Upper surface dark purplish chocolate-brown ; margin crenated, white, with an inner
border of orange, and another thin one of black.
Under surface paler.
Ova white.
Apparently allied to Hurylepta interrupta, Stimp. (Proc. Ac. Se. Phil. 1857, p. 26).—
Cre:

Fig. 26. Upper surface.
Genus CENTROSTOMUM, Dies.

27. Centrostomum ocellatum, Kel.

Length 2 inches.
Animal gelatinous.
Upper surface pale yellowish brown, with dark brown ocellated spots.
Under surface pale buff.
Tentacles none.
Mouth near the centre.
Hye-spots occipital.
Ova white.

Of this and the next species Dr. Kelaart made a new genus, which he called Penula,
the principal character of which was the absence of any form of tentacles. But this
character is not that of a genus, but of a subtribe, called by Diesing Aceridec ; and the
families Typhloleptide, Leptoplanide, &c. are therein included.—C. C.

Fig. 27. Upper surface.

28. Centrostomum punctatum, Kel.
Length 12 inch.
Upper surface white, shaded and minutely punctated with reddish brown.
Under surface very delicate white, clouded with faint reddish brown.

Fig. 28. Upper surface.

These two species (particularly the first) seem to bear a great resemblance to the Pla-
naria gigas, from the Red Sea, figured in Riippell’s Atlas, t. 3. f. 5.—C. C.

98 , DR. COLLINGWOOD ON MARINE PLANARIANS.

Genus STYLOCHOPLANA, Stim. -

29. Stylochoplana elegans, Kel.
Length 1 inch.
Upper surface pale yellow, shaded with greenish brown, and dotted with black; margin
black, lined with orange.
Under surface whitish, with brownish margin.
Tentacles two, small, reddish, tipped with red, occipital.

Vig. 29. Upper surface.

30. Stylochoplana meleagrina, Kel.
Length 12 inch.

Upper surface sixiped with broad streaks, the inner pure white, the others light purplish
and whitish ; the median line brownish red, edged with black ; the margin waved and
edged narrowly with black.

Tentacles small, oval, occipital. There are also two linear appendages on the occipital

region above the eye-spots.

Fig. 30: a, upper surface ; b, under surface.

Genus PLANOCERA (Blainville).

31. Planocera thesea, Kel.
Length 1} inch.
Upper surface chocolate-brown, edged with yellow.
Under surface pale purple, darker towards the margin.
Tentacles white, tipped with red, rising from depressions or cups placed near the middle
third of the body.
Mouth central.

Fig. 31. Upper surface.

| EprrortaAL ADDENDUM, with EXPLANATION of Puates XVII. to XIX.

The figures im the three Plates are consecutively numbered, and correspond with the numbers and

references appended at the close of each description of the species in the text. The original drawings of
figs. 1 to 12 (Pl. XVII. and part of Pl. XVIII.) were designed by Dr. Cuthbert Collingwood ; those of figs.
13 to 81 (part of Pl. XVIII. and the whole of Pl. XIX.) are by Dr. Kelaart. Owing to the protracted
absence of Dr. Collingwood abroad, the proof sheets have not had the advantage of his revision. This
has unfortunately led to a discrepancy in the lettering in the plates. It was originally intended that the
author’s own newly described species should be distinguished by Arabic, and Dr. Kelaart’s by Roman
numbers; but the artist has not carried out this minor matter of detail. The lettering im the text has
not been altered to correspond with the artistic error in the plates, which were printed aff before the

correction of proof sheets was undertaken.

Trans. Linn. Soc, Ser. 2. Zoon Vou.l'Tas. 17.

=
b
C Collingwood delt G. French Angas lith W. West&C° imp

New Species of Planaria.

Trans Linn Soc Szr.2 Zoor Vou | Tas 18.

om

L

¢. Collingwood. delt
Geo French Angos lth

New species of Planaria.

W.West & C2 ump

}

Trans. Low.Soc.Ser 2.Zoon Vou. 1. Tas 19.

-

Dr Feloort deleod not W West & Co ump
Geo French Angas ith.

New species of Planaria.

ou

[22 all

VII. On the Structure and Development of the Bird's Skull. By W. K. PARKER,
THES, LM beSe, Cfo (Ceehen 18)

(Plates XX.-X XVII.)
Read December 16th, 1875.

INTRODUCTORY REMARKS.

IN a paper of mine on the Development of the Bird’s Skull (Phil. Trans. 1869, pp. 755-
_ 807), a single species, the Common Fowl, was used as the subject; this was worked out
to as great an extent as was possible to me at the time.

Year by year since then steady work at other types of the Vertebrata has, by letting
in fresh light, shown me some serious deficiencies in that paper, and my mind has not
been able to rest without an attempt to superadd something of value to the older piece
of research.

I may at once remark that, criticising this my own work, I find that the eye and the
hand had done their work better than the md; the plates are clearer and truer to
nature than the descriptions. :

The Memoir which followed that on the Skull of the Fowl treated of this part in the
Common Frog, a type so diverse from that of the Vertebrata generally as to mislead me
somewhat in my comparisons, and to make me doubt the correctness of my earlier
descriptions. Further researches, however, have set my mind at ease on several points,
notably those on the Salmon’s Skull (Phil. Trans. 1878, pp. 95-145, plates i—viii.).

There was another reason, however, why it was desirable to supplement the paper on
the Skull of the common Gallinaceous bird, namely, that it is itself a very simple semi-
struthious type, and conveys but an imperfect idea of the remarkable metamorphoses the
Reptilian type of skull can undergo when subjected to the intense life-energy of a high-
class perching or climbing bird.

A third reason has acted as a propelling force in this matter: this was a strong desire
to clothe with flesh and form the masterly skeleton paper on the “ Classification of
Birds” given by Professor Huxley to the Zoological Society a few years ago (Proc. Zool.
Soe. 1867, pp. 415-472).

In that paper the palatal structures are chosen as the ground of classification; and
although, to a hasty observer, these might seem to be a mere fragment of the whole bird,
yet they are, indeed, the parts to which all others follow suit—the upper jaw and palate,
the first and part of the second facial arch. To these parts every thing else in the Bird is
correlated ; they rule, as it were, the whole economy of the Bird.

Renewed researches, which have been zealously followed up by me for some years, on
the structure and development of the “ trabeculee cranii,” “ pterygo-palatine arcades,”
and nasal labyrinth, have yielded results that seem to me to be of great value.

SECOND SERIES.—ZOOLOGY, VOL. I. P

100 MR. W. K. PARKER ON THE STRUCTURE

In this newer work the forms of the Fish and the Reptile have been before my eyes
night and day. These have been to me as the larval and pupal stages through which
may be traced the changing materials out of which that vertebrate imago, the Bird,
is constructed, its hot blood infusing new life and producing unlooked-for trans-
formations. ;

Now that cranial morphology is beginning to embody itself in a clear and intel-
ligible shape, it may be well to explain the meaning of certain terms that will of neces-
sity be in frequent use.

The term metamorphosis I use in exactly the same manner and for the same purposes
as the entomologist, namely, as describing the whole series of changes undergone by the
skull or skeleton throughout the life-history of the individual.

The borrowed term-isomorphic is used, in a zoological sense, for very similar types in
quite distinct groups: the Sun-birds (‘‘ Nectariniidee””) may be said to be the isomorphs
of the Humming-birds (“ Trochilidee ”’).

The term symmorphic may be used for any morphological part or parts in one type
which agree in their structure and development with those of some other types; thus
the “ Desmognathous” palate of the Parrot and that of the Goose are symmorphic; such
agreeing structures are also said to be homologous or representative.

Professor Huxley, in his paper (op. cit. p. 454), places the Goatsuckers and Humming-
birds with the Passerine types, supposing them to have the true “ Mgithognathous ”
palate. This is an error: they are Insessorial Schizognaths.

The Woodpeckers (“ Picidee,”’ p. 448) had this service done to them, namely, their vomer
was shown to be double, and their maxillaries to have a very rudimentary palatine plate.
Thus they could not be treated of as belonging to the Cuckoo tribe (“‘ Coceygomorphee”’).

The palatal structures (trabecule, pterygo-palatines, and nasal capsules) of the Wood-
peckers show that these birds can be classified with none of the groups established by
Professor Huxley, as their face is intensely Lacertian and even Ophidian. I have there-
fore called this group of birds the “ Saurognathe” (Trans. Linn. Soe., Zool. pt. 1,
1875, pp. 1-22, Plates I.—V.).

That peculiarity of the Bird’s face which has been termed “ Aigithognathous ” will be
explained. It exists in all the “ Coracomorphe”’ and, besides these, in the Swifts, or
“‘ Cypselomorphe ”’ (op. cit. p. 468).

The importance of the fact that a low, superstruthious bird like the Hemipod should
offer itself as a stock (“phylum”) to the half myriad of known Passerines cannot be
easily overrated *.

My principal reason for describing the types of skull presented here is not zoological
but morphological. I wish, in some degree, to fill up that which remains over of cranial

* On the distribution of “ Alectoromorphe,” see Huxley, Proc. Zool. Soc. 1868, pp. 294-319. ‘The Tinamous
stand with one foot in that group and with the other over the struthious border, and the larger “ Cracide” (Craa
globicera) and “* Megapodide ” (Megacephalon maleo) are “ Desmognathous” (see Huxley “‘ On Classification,” p. 423,
for a figure of the former). Thus the Fowl tribe sweeps over a huge zoological space, immediately overlying the
stratum occupied by the “ Ratite.” Amongst the ancient extinct forms of this group we must look in imagination
for the forefathers of the Birds of Prey, the Climbers, and the Perchers.

AND DEVELOPMENT OF THE BIRD’S SKULL. 101

morphology after that of the Fowl has been exhausted, so that the student’s conception
of the Bird’s skull may be richer and more in accordance with the fulness of Nature.

I shall take the present opportunity to give figures of skulls that contain elements
frequently spoken of in other papers by me, but not illustrated, such as the “ inter-
palatines’’ and “ mesopterygoids.” I shall also have to add other pieces, with their
names, namely, the “ medio-palatine”’ and the “ palato-maxillary.”’

Processes that grow in a secondary manner out of the simple facial bars, for union of
bar with bar, serially, are called here “‘ conjugational processes ;” the union of the right
and left bars of one arch at the mid line is called a “‘ commissure.”

Further Researches on the Chick's Skull.

My first stage (Phil. Trans. 1869, plate Ixxxi. figs. 1 & 2) is rather wnderdated as the
fourth day ; those given are of the fifth, and those soon to be described run nearly to the
sixth day. The second stage (figs. 3-13) should be put as dating from the sixth and
seventh days of incubation.

One of the first points to be determined in the morphology of the skull is the structure
of the “ investing mass” of the notochord and the “ rafters of the cranium” (“ tra-
beculze cranii”’), the first pair of facial bars.

The figures given in my former paper show the distinctness of these parts much more
definitely than I supposed at the time. These I now proceed to explam. At page 758
I speak of the ends of the investing mass as forming the cartilaginous “ lingul sphenoi-
dales,” thus confusing the free truncated end of the investing mass (plate Ixxxi.
fig. 2, 1.g), which looks forwards and outwards, with the free tops of the trabecular bars,
which are better seen in fig. 8, and in plate Ixxxii. figs. 1 & 3, l.g.

The explanation of these parts was made by the exquisitely lucid structures of the
embryo Salmon ; and now, if these figures of the Fowl be compared with those, it will be
seen that the trabecular apices are at first embraced by the squared out-turned ends of
the investing mass, and that in the next and third stages the curved trabecular tips,
where they surround the pituitary body, open out like the blades of a pair of callipers,
and growing more distinct and solid, form, in the third stage (plate Ixxxii. figs. 1,3, & 4,
lg), the free tongue-like cartilages, the meaning of which I for a long while failed
to see.

The first figure in the present paper (Plate XX. fig. 1, é, pm) ciears up another very
different point, namely, the formation of the “ prenasal rostrum,” or azygous con-
tinuation of the distal part of the trabecular arch.

In my first paper, on the “ Skull in the Ostrich tribe” (Phil. Trans. 1866, plate vii. p.x,
p- 122), the prenasal cartilage is described as being formed by confluence of the “ tra-
beculz ;” but in the second paper (p. 759) [have corrected this statement, but not without
falling into another error, namely, that of supposing the “ale nasi’’ to be formed directly
out of the “cornua trabecule.’” The anomalous position of the parts is caused by the
“© mesocephalic flexure,” the trabecule being positively hooked backwards.

The ‘alinasal, aliseptal, and the aliethmoidal cartilages are regions of membrane

which chondrify later than, but not separately from, the trabecule, the common
P2

102 MR. W. K. PARKER ON THE STRUCTURE

descending nasal septum becoming solid at its sides continuously with the crest that
grows upwards from each trabecula. The broadest part of the “trabecular commissure ”
(Fowl’s Skull, plate Ixxxi. figs. 1 & 2, tr) develops into two important structures,
namely, the subnasal floor (plate Ixxxii. fig. 11, s.v./) and the conjugational process,
which corresponds with the “ ethmo-palatine,”’ or orbital plate of the palatine. The sub-
nasal or “ supervomerine”’ lamine are of small extent and temporary in the Fowl
(plate Ixxxii. fig. 11, s.v./); and the conjugational process does not develop into a
distant structure, but remains as part of the antorbital wall, its infero-external angle
(plate Ixxxiil. fig. 5, p.p). In many birds, however, it hecomes separately ossified as the
“os uncinatum” of Magnus, and takes its place as a kind of transverse bone in the
“« Musophagide ” and other “‘ Coccygomorphee ”*.

In the first of my new illustrations of the Chick’s Skull (Plate XX. fig. 1, tr, pn),
the down-bent skull of a chick, intermediate between the first and second stages of my
former paper, has been sliced off in an obliquely transverse manner, cutting away the
fore part of the projecting optic lobes (C’) and thalamencephalon (C’’), and removing
most of the rudimentary hemispheres (C”). Underpropping the primary vesicle (C’’)
are two thick cartilages, which are severed at their commissure ; these are the trabeculez
at their bend backwards. From this position they send downwards an azygous sharp
keel. This is the nasal cartilage; it looks backwards.

In the next stage (“ Fowl’s Skull,” plate Ixxxi. fig. 3, p.m) it is still retral, but has
become thick and rounded.

In another section (fig. 2), taken at a different angle to the axis of the head and seen
from its front face and not from behind, as in the last figure, the trabecular knuckle
has been cut off, and with it the prenasal cartilage. Here the under-placed nasal sacs
(ns) are exposed, and the hooked trabeculze are cut through so as to expose them in
section, both in the meso-ethmoidal region above, and under the outer nostrils below.

In this section the fore part of the eyeballs (e) and the hemispheres (C”) are cut
through ; the floor of the nose shows cartilage in it only where the cornua trabeculz are
cut through; the prenasal has been pared off. In this rudimentary nasal labyrinth
rudiments of the superior and inferior turbinals (wf, i¢b) are seen, and the membrane
infolding them is the early condition of the aliethmoidal and aliseptal cartilages. The
eyeball is seen to rest on a very neatly formed ledge; this is the jugo-maxillary eleva-
tion; and on the side of this is a lesser ridge, which contains the second preoral bar,
the palato-pterygoid.

This palatine bar (Pa) is composed at present of indifferent tissue; it is triangular in
section, and the ascending crest is its ethmo-palatine region ; the conjugational process
is correlated to that of the trabeculz.

Tn a somewhat more advanced skull, sixth day (figs. 3 & 4), the eyeballs have been cut
through at their middle and the hemispheres at their hindermost part. This belongs toa

* Sce Dr. Reinhardt’s paper, “Om en hidtil ukjendt Knogle i Hovedskallen hos Turakoerne.” Copenhagen, 1871.
In my paper on the “ Ostrich’s Skull” (p. 158), I referred the “ os uncinatum” to the ethmoidal category ; but no
explanation of its true meaning was had until I saw the secondary bud growing from the outside of each trabecular
bar in the Salmon. :

AND DEVELOPMENT OF THE BIRD’S SKULL. 103

head that had begun to straighten, and nearly comes up to the earliest part of my second
stage (“ Fowl’s Skull,” plate lxxxi. fig. 3).

The interorbital septum (pe) has here been cut through in the anterior sphenoidal
region, and, as in the Mammal, the sphenoidal sinuses (sps) are exposed ; they have not,
however, any retral continuation of the nasal wall in a cartilaginous state, and they are
very narrow. Below, thestrongly compressed trabeculee (¢r) have coalesced, and are now
composed of hyaline cartilage; although the commissure is finished below, the crests of
the trabeculee are distinct. These solidify more tardily ; and their conversion into true
cartilage taking place at the same time as the cranial floor with its double descending
keel, there is at present no line of demarcation between these regions.

Afterward (see “ Fowl’s Skull,” plate Ixxxii. figs. 4, 5, & 11, p.s, 0.s) the mem-
branous interorbital space (é.0.s) redivides these dissimilar tracts, leaving for some
extent a descending presphenoidal keel and an ascending trabecular crest.

The orbito-sphenoids (figs. 3 & 4, os) are small, and lessen rather than increase in size.
The palatines are here narrowing towards the pterygoid region.

The next section (fig. 5) is through the optic lobes (C’), the pituitary body (py), and
the back part of the eyeball.

The manner in which the face is tucked under the head at this stage makes a most
remarkable display of parts possible in one section: these are the apices of the tra-
becular (¢7) and pterygo-palatine arches ( pg), and at the same time the distal extremities
of the first and second postorals (mk, ch, b.hy). The internal carotid arteries (éc) are
seen piercing the fibrous pituitary floor mesiad of the ends of the ‘trabeculz, the
actual apices of which look downwards in the figure. On each side of the high and
narrow mouth-roof (m) are seen nearly the whole of the pterygoids (pg); their “ trabecular
process,” or epipterygoid apex, looks a little inwards below, and is seen enclosed in the
rudiment of a “ soft palate.”

The end of each “ sphenoidal sinus” (sps) is here seen to reach to the side of the sella
turcica. The floor of the mouth (m) contains laterally the Meckelian rods (mk), supra-
mesially the tips of the small distal cerato-hyals (ch), and below the basi-hyal (b.hy).
Behind the optic lobes and eyeballs the head has become very narrow. The next section
(fig. 6), which has been made through the fore part of the auditory capsules, shows
this.

The notochord (ve) lies between the two halves of the investing mass (iv), which has
completely coalesced with the periotic cartilage. This section shows the cochlear cavities
(cl) and those of the anterior semicircular canals (asc) with their ampulle. Below the
cochlee are seen the internal carotids (ic), and towards the mid line the submucous
tissue is of great thickness; it forms the nidus in which those remarkable ornithic
detachments of the parasphenoid, the basitemporals (0¢), are developed. All but the
fore ends of the mandibular rods are here shown, and also their piers, the quadrates ;
mesiad of these are seen the basi- and cerato-hyals (bhy, chy).

With regard to the parasphenoid, although it is developed as an azygous bone ante-
riorly, yet the thick bed of stroma which forms its mother-substance is a symmetrical

104 MR. W. K. PARKER ON THE STRUCTURE

growth from the two sides (see fig. 4), when these masses are approximating below the
coalesced trabeculz.

Before dismissing the chick’s skull I have to speak of the posterior conjugational pro-
cesses connecting the trabeculze with the next facial bar, the pterygo-palatine.

These parts, the basipterygoids with their articular plate and the corresponding
structures in the pterygoids, are thoroughly worked out in my paper on the “ Fowl’s
Skull” (plate Ixxxiii. p. 781); but the general morphology, their symmorphism with the
anterior conjugationals of the same arch, was not clearly seen.

Before closing this supplemental description of the fowl’s skull, I may remark that
the fenestra which severs the perpendicular ethmoid from the nasal septum (‘“ Fowl’s
Skull,” plate Ixxxiii. fig. 4, ¢fic) is the redifferentiation of the trabecule from their
surroundings.

In many birds another fenestra, besides the “hinge” and the interorbital space, is
found in the nasal septum; and in the Goose tribe a fourth appears between the upper
turbinals; to this is added, in Pelecanoides urinatriz, or Petrel, a fifth large fenestra,
close behind the antorbitals.

In the adult Swift (Cypselus apus) there are fowr membranous fenestre in the inter-
orbital space, instead of two as in Pelecanoides, and one as in Birds generally. This
gradual fretting-away of growth-tracts which at one stage of metamorphosis obscure
essentially distinct parts is of extreme interest; the highest and, as it were, the #mago

forms show this most, notwithstanding that they use up and mix together in many ways
parts that remain thoroughly distinct in types that may be compared to the pupe and
larve. As the two preoral arches are intimately interblended, not only with each other
but also with the nasal labyrinth and the cranium, it may be well to set down fairly what
splints or secondary bones become applied to or grafted on these arches.

To the hinder pair, the “ palato-pterygoids,”’ secondary developments of the mandible
or first postoral, there belong the maxillaries, palato-maxillaries, interpalatines, jugals,
and quadrato-jugals.

The trabecular arch draws to itself, as subcutaneous or submucous bones, the follow-
ing—namely, the premaxillaries, septo-maxillaries, vomer or vomers, and the para-
sphenoid ; the “ os uncinatum”’ is an endosteal part.

The splints of the mandibular arch were described in my former paper. The tympanic
chain of the Bird is a masked splint-series to the segmented and metamorphosed apex of
the hyoid arch.

On the Structure and Development of the Facial Arches in the Passerine types,
“‘ Coracomorphee ’’ (Huxley).

In the paper already referred to, Professor Huxley has sought to group the flying
birds in accordance with the spirit of Humboldt’s plant-groups in his ‘ Views of Nature.’
The group under notice, then, is his “ Coracomorphe,” or Crow form (p. 469); this is
so large that (as my friend Mr. Osbert Salvin informs me) whilst the birds already known
and described are about ten thousand, half, or well nigh half, of them belong to this one
group. Now the type the skull of which I have already worked out, namely that of

AND DEVELOPMENT OF THE BIRD’S SKULL. 105

the Fowl, stands as the head of another group, the “ Alectoromorphe;” and this
represents also a broad and goodly family of forms. In another paper, ‘“‘ On the Classi-
fication and Distribution of the Alectoromorphze”’ (Proc. Zocl. Soc. 1868, pp. 294-319),
our author has broken this fine family into fragments, separating (see pp. 302-304)
not only the Pigeons (‘ Peristeromorphe”’), which are “ Altrices’’ and have tender
young, but the Sand-Grouse also, as the ‘“‘ Pteroclomorphe,”’ and the Hemipods as “ Tur-
nicimorphe.” As they stand at present, no cutting and contriving will make the
lesser groups, such as the ‘“ Charadriomorphe,”’ ‘‘ Pteroclomorphe,”’ and the like,
correspond to or in anywise be the equivalents of the great army comprised under the
Crow form. Much as I value my friend’s paper, I do but consider it in the light of a
sign-post to guide others who are footing it on the same road, and not as a finished work
to rest in.

In one of the most outlying of the Crow forms, the House-Martin (Chelidon urbica),
I have been able to demonstrate the second morphological stage of the second postoral
arch. I did not succeed in this point in the Fowl; but now that it is done it will do
duty for the whole of the “ Sauropsida.”

The metamorphosis of this, as of the other arches, has been fully worked out in the
“ Tchthyopsida,’” in my researches on the skull of the Frog and the Salmon; but
neither of these throw any light upon the long-stalked, trifoliate stapes of the Reptile
and the Bird. Professor Huxley, in his paper ‘‘ On the Representatives of the Malleus
and Incus” (Proc. Zool. Soc. 1869, pp. 391-407), says (p. 398), “the suprastapedial
cartilage”’ [he is describing the parts in Sphenodon] “turns out to be nothing more
than the proximal end of the hyoidean arch, while the stapes and its appendages are
exclusively related to this arch, and have nothing whatever to do with the mandibular
arch.”

True! and yet he does not say how much of the stapes belongs to the auditory capsule.
In the Urodela the stapes is cut out of that capsule, like a dung, and in Batrachia the
head of the upper segment of the hyoid arch articulates with the auditory stapes, its
uppermost part being segmented off as a “ pars orbicularis.” In the Mammalia the
Batrachia are closely followed (Phil. Trans. 1874, plates xxvi—xxxvii.). In my former
paper on the Fowl’s head (plate Ixxxi. figs. 5, 9, & 10, st) I showed, in the second stage,
a thick somewhat flattened club of cartilage, the knobbed end of which fitted into the
fenestra ovalis, whilst the other or distal incurved end was trifoliate, the upper process
being the “ suprastapedial ”’ rudiment, the lower the “infrastapedial,’”’ and the longer,
bowed leaf the “‘ extrastapedial.”

In a young Have-Swallow, at about the same astel, the upper end was seen to be
continuous with the periotic capsule im front of its fusion with the exoccipital cartilage,
and above the outer end of the rudimentary cochlea. This stapedial bar (Plate XX.
fig. 7, mst) has the characteristic curve of the summit of the facial arches, and at this
stage has coalesced with the auditory capsule as is the wont of these bars (see the
quadrate in continuity with the periotic cartilage, as figured in Huxley’s ‘ Elements,’
p- 138, fig. 57, F* Qu). This is undoubtedly the second or coalescing stage of the
second postoral; it has, however, only coalesced by the strong posterior head; the

106 MR. W. K. PARKER ON THE STRUCTURE

aborted anterior head, comparable to the “ tubercle” of a rib, and answering to the
short crus of the incus, has been left free. The anterior head, or “‘ suprastapedial” (s.s¢),
is, however, continuous permanently with the auditory capsule in Sphenodon (Huxley,
«On Malleus and Incus,” p. 397, fig. 4, S.S¢), and also, as I find, in the Bull-frog (Rana
pipiens). Notwithstanding that in Sphenodon the posterior head is free, and the anterior
is fixed, contrary to what is here shown in the Chelidon, yet in these two, the adult
Lizard and the embryo Bird, the “ extrastapedial”’ (Plate XX. fig. 7, est, and Huxley
op. cit. fig. 4, H.St) has a similar shape; even the foramen seen in Sphenodon (a) soon
appears in the Bird. The “ infrastapedial” (Plate XX. fig. 7, ist) of this young bird
is longer than I found it in the Fowl, even after the oval tract of ear-wall had come
away with its “ mediostapedial”’ stalk. This is not all; for its pointed apex is con-
tinuous with a band of cells of indifferent tissue which runs downwards towards the
entering carotid artery at the skull-base.

In Sphenodon (op. cit. fig. 4, Sth) the scooped “ extrastapedial ” passes directly down-
wards into a bar which is labelled Sth (stylo-hyal); but in the Crocodile (op. cit.
figs. 1 & 2, Sth) the stylo-hyal is a short club of cartilage, flattish, with its broad end
upwards, looking backwards, and having a clear membranous space between it and the
“ extrastapedial.”

The stylo-hyal of Chelidon answers to this description exactly ; and if the reader will
turn to Professor Huxley’s figure in this part of the Fowl (op. cié. fig.5, A, B, I.St) he
will see that it is there, but that it has become confluent with the infrastapedial. In
figs. 5,9, & 10, st, of my former paper, this part is not shown; but in a chick one day
older (fig. 14, i.s¢) it is certainly figured, but not distinct. Not having seen it in the
earlier stage, I could not choose but find it in the next, as it had then coalesced with
the “ infrastapedial.’”” In certain birds this part ossifies; and this fact led me to call it
the “stylo-hyal” *. The distal part only of the rest of the arch is chondrified, namely,
the small cerato-hyal (chy), which, with its fellow, lies in the substance of the tongue.
The basal element, “ basi-urohyal” (bhy, 667), is a stout wedge, which narrows, and
then is flat and emarginate behind. The so-called “thyro-hyals”” (= 1st branchials, br’)
are very large, the upper or proximal piece embracing the occiput. In this illustra-
tion the lingual skeleton is drawn as flattened out in the minute preparation, so as to
display all the parts at once.

I take this opportunity further to illustrate Professor Huxley’s account of these parts
in his invaluable paper (op. cit. p. 398), and first give the condition of these parts in a
fledgling Crow (Corvus corone). The membrana tympani (fig. 8, mfy) is shown from
the inner side, with the stapedial and mandibular structures; the articular end of the
lower jaw is shown slightly dislocated, and the ‘“ siphonium”’ is cut through.

This tube, the siphoniwm of Nitzsch, conveys air, not from the quadrate to the lower
jaw (see Huxley, op. cit. p. 391), as in the Crocodile, but it directly opens into the
tympanic cavity in the Birdt.

* Full often have I argued with the author of the paper just quoted that this dilated, decurved, sometimes bony
extremity of the bird’s “ infrastapedial” was the true “ stylo-hyal;” but he always turned his deaf ear towards
me. Jam silent now; but “ he will rise up at the voice of the bird,” so sharp and clear.

+ The dissection here figured was made to dispel a doubt of Professor Huxley’s as to the correctness of Nitzsch’s

AND DEVELOPMENT OF THE BIRD’S SKULL. 107

In the adult of this species, the Carrion-Crow, the siphonium is enringed by the
lowest and largest of the seven bones that form the curious tympanic chain. Figures of
these have been given in my paper on the “ Coracomorphe ”’ in the ‘ Transactions’ of the
Zoological Society.

The stapedial plate (st), turned a little forward for display, is continuous, as bone,
with the mediostapedial (mst), and then come the outspread leaflets of cartilage. The
* extrastapedial” (est) is sickle-shaped and broad-backed, its thick convex back being
strongly connected with the inner face of the membrani tympani (mty); and the fibres
of the perichondrial coat are interwoven with those of the drum-skin itself, and also meet,
from without, those of the tendon of the “ tensor tympani” muscle (stm).

The tongue-shaped rudiment of the suprastapedial (sst) lies in the base of a two-
winged membrane, the fibrous supplement of this part.

The cartilaginous strap running downwards from the extrastapedial to the entering
carotid (ic) is the infrastapedial (ist) and stylo-hyal (sth) in one band. The fore end of
the mandible (ms, d) shows that, as in the Fish, the mandibular rods do not meet to
form a “mentum” in these prognathous types; they keep distinct, and each receives
its own dentary.

The form and disposition of these parts in the adult are shown in fig. 9. They are
those of the Daw (Corvus monedula); the references are the same as in fig. 8.

In the Crows of the southern world there are many important differences of structure
from what is seen in the northern species: notably, the condition of the stylo-hyal is
instructive. In the Piping Crow (Gymnorhina tibicen) the general structure corresponds
very exactly with that of Corvus; but the stylo-hyal (figs. 10 & 11, sth) ends below in a
cheese-knife-shaped plate, which lies upon the outer edge of the basitemporal, and is
well ossified. Slender as in the mediostapedial shaft (mst), it is pneumatic and com-
municates with the tympanic cavity by both proximal and distal passages.

Being desirous that this, together with the paper on the Fow/’s Skull, should enable
the student to form an accurate conception of the skull of the Carimate Bird, as
distinguished not only from that of other classes, but also from that of the “ Ratite,’” or
Struthious order, I now proceed to show some of the most important modifications of the
ornithic face. Happily for me, I have at hand a strong and sharply expressed outline of
this subject ; and now it will be easy to make a more extended study and comparison of the
facial morphology of the higher birds (see Huxley’s “ Classification of Birds,’’ Proc. Zool.
Soc. 1867, pp. 415-472).

My former papers illustrate the “ Deomaeeinttous” face as seen in the raft-breasted
Ostrich (Huxley, op. cit. p. 425) and the keel-breasted Tinamou (Phil. Trans. 1866, plates
ix.—xv. pp. 118-183) ; the “ Schizognathous”’ face (Huxley, op. cit. p. 426), as seen in the
Common Fowl (Phil. Trans. 1869, plates Ixxxi—Ixxxvii. pp. 755-837); and the “ Aigi-
thognathee ” have been illustrated in the Zoological Transactions for 1875 *.

T return now to the Sparrow-faced birds, “ Mgithognathee”’ (see Huxley, op. cit. p. 450).

description, as that made the Bird differ from the Crocodile in the disposition of the pneumatic duct. It does differ,
as I have shown above.

* See Part I., in vol. ix., plates 654-62. Part II. will soon appear.
SECOND SERIES.—ZOOLOGY, VOL. I. Q

108 MR. W. K. PARKER ON THE STRUCTURE

The development of the preoral arches and nasal capsule in the “ Fringillide ” (which
in the person of the Sparrow gives name to the group) will show the peculiarities of this
type, a type which takes in all Professor Huxley’s “ Coracomorphz” (op. cit. p. 469),
the Swifts amongst his “ Cypselomorphe ” (p. 468), and the Hemipods also, which lie
down lowly, in an ornithic stratwm immediately over the “ Tinamide”’ *.

In embryos of the Brown Linnet (Linota cannabina), at about the end of the fifth
day of incubation, I find the facial arches and nasal capsules (Plate XXI. figs. 1 & 2)
in a very instructive condition. Nearly the whole of the long “ trabecular commissure ”
is shown, the bar being severed in the prepituitary region, where the ethmoid and basi-
sphenoid meet; the prenasal rostrum (pz) is in full size, as the continuation of the
great cartilaginous balk.

This balk is wasting between the ethmoidal and septal regions, preparatory to its
peculiar ornithic segmentation.

The open or naturally cloven state of the palate is here shown; for the folds of the
nasal labyrinth are well seen below, and the “ median nares” (mm) are large and
patulous. Neither the true olfactory nor the inferior turbinal regions are displayed in
this figure, only the immensely developed alinasal structures, so very secondary in
importance and size in the Mammal. The trabecule being bowed and folded over in
front send backwards, on each side, a flap of cartilage; this may be called the “ recur-
rent fold” (rec). These two flaps form a commissure below the fore end of the
septum nasi; and this becoming ossified from the alee, sometimes appears as an anterior
nasal bridge, a desmognathic band close behind the fore beak (e. g. in Dicholophus).

The posterior of the three folds, the inturned lamina (éa/), seen in the preparation, is
the swelling alinasal wall passing into the floor of the nasal vestibule. Here it is far in
front of the maxillo-palatine hook (map); but after a short while it will lie on this bony
bar, and become intimately connected with it by the binding fibres belonging to each
structure. Between the plaits already described is seen a middle cartilage; this is an
outgrowth of the wall or posterior plate. It is the alinasal turbinal, and ends in a
double free edge.

Although the premaxillaries (px) and maxillaries (mx) are already well developed, yet
one does not see here the broad forked vomer of this type. Looking, however, carefully
between the wasting middle bar and the inturned alinasal wall (¢a/), there is to be seen
at this stage a small oval cartilage; as this appears to me to be as yet undescribed, I
propose to call it the ‘‘ vomerine cartilage” (vc). It ossifies endosteally, and has even
now a small core of osteoblasts with the finest deposit of calcareous matter. One of these
cartilages is more highly magnified in fig. 2, and also with it part of the nasal wall, ali-
nasal turbinal, and maxillo-palatine.

This cartilage is small here in the Linnet, but very large, as I shall soon show, in
other “ Hgithognathe ;” and as it forms the mother patch for each vomerine moiety, it

* In the paper under notice (p. 459) Professor Huxley placed the Hemipods among the Fowls, ‘“ Alectoro-
morphe ;” but in another equally important contribution to Ornithology, namely, his “ Classification and Distribution
of the Alectoromorphe ” (Proc. Zool. Soc. 1868, pp. 294-319), he places the Hemipods apart, haying gained a clearer
insight into their peculiar characters (see p. 804). They are now his “ Turnicimorphe.”

AND DEVELOPMENT OF THE BIRD’S SKULL. 109

is of great interest, and isnew to me. If any morphologist knows what it answers to in
the lower vertebrate types it were well that he should explain it *.

As the metamorphosis of these parts is very rapid, it is necessary to examine embryos
not more than a day or so in advance of or behind each other. Inno longer a period, what
is displayed in fig. 1 is exchanged for what I have shown in fig.3. Here the small osseo-
cartilaginous nuclei have developed into the large ox-faced vomer, with its broad anterior
part grafted on each side upon the alinasal wall, which turns directly inwards, and
spreads in a pedate form to apply itself to the vomer. On one side of the V-shaped
emargination the vomerine piece is fenestrate; the bony matter here becomes detached,
and an additional bone formed. ‘This additional bone is seen in the fledgling (see it in
the young Sparrow, fig. 4, smx). Not only does the vomer grow into the nasal wall, but
the alinasal turbinal (fig. 3, atb) also; hence the huge size of the upper vomerine groove,
which is so striking in the Crow tribe (“ Corvide” proper); for the thick common end
of the wall and the turbinal approach the septum nasi, and, rising high on each side,
embrace the base of the septum. Another noteworthy characteristic of the Passerine
face is the elbowed form of the palatines near their hinder third; this out-bent part does
not ossify ‘‘ by first intention,” but lingers until it is solid hyaline cartilage, and then
receives its own endosteal centre. This new bone is an arrested “ transverse’ element,
my “ transpalatine”’ (¢pa).

That part of the Passerine palatine which is arrested from forming a bony palatal floor
generally exists as a free fore-looking spine. This is well seen in the Sparrow (fig. 4,
ipa); it is the “ interpalatine process.” In Ducks and Swans this is a separate knuckle
of bone; and in looking over a large number of Starlings’ skulls, I find it in certain of
them (see Plate XX. fig. 12, ipa).

Other parts of the Passerine face must be noticed.; but I turn now to the most striking
condition of the vomer I have hitherto found in the group: it is seen in the native
Wren (Troglodytes vulgaris). When the whole palate of the bird is viewed (Plate X XI.
fig. 5, v), the vomer is seen to be as large, relatively, as in the “ Ratite;”’ but its
anterior third is composed of true cartilage. In a spirit-specimen, by careful dissection,
I have been able to make out what is shown in fig. 6 on a larger scale. Here one of the
palatines (pa) and both of the maxillo-palatine hooks are removed; they are, however,
indicated by a dotted outline.

* Finding the development of the Passerine “vomer” so totally unlike what I always supposed, namely, that it is
an endosteal ossification of cartilage, my mind cast about for some symmorph in the cold-blooded types and in the
Reptilian birds (Ratite). I had not far to seek; for lying unexplained in my memory were certain structures in the
Snakes and also in the struthious Rhea, which I had figured years ago (Phil. Trans. 1866, plate x. fig. 14, the alate
cartilages above v), but which had hitherto resisted all efforts at explanation. On reexamination of these parts in the
Rhea chick, 1 find that they are free cartilages in the mazillo-palatine region, and attached to those processes and to
the fore forks of the double vomer by fibrous tissue. In the Snake the nasal glands and their bony capsules, the
vomers and septo-mawillaries (P), are flanked on their outside by an arcuato-spatulate flap of cartilage, continuous
by a narrow stalk with the recurrent lamine, or separate from that process, both states occurring in the same skull.
As in the furculum of the same bird (Linota, see “‘ Shoulder-girdle and Sternum,” plate xv. figs. 12-15), the Linnet’s
““yomer ” is a morphological compound, and the counterparts of its component elements are persistently separate in
the cold-blooded types.

Q2

110 MR. W. K. PARKER ON THE STRUCTURE

Part of the dialate septal base (¢r) and part of the nasal labyrinth of one side is here
shown. We here see that the trabeculze have only partially lost their flatness, and form
a, partial subnasal floor, and that the alinasal wall (mv), after giving off the “ alinasal”
turbinal (at), ends externally in a blunt process, and internally in an incurved bifurcate
rod (ial).

The forks of this rod are continuous with the corresponding vomerine half ; the halves
of the vomer are joined by a long commissure into a flat bone, notched at both ends; the
long posterior ends articulate with the ethmo-palatines, and the shorter anterior forks pass
into a cartilage shaped like a breast-plate. ‘This plate, being gently emarginate behind,
forms a fenestra with the help of the bony vomer; it narrows gently forwards, and ends
in a pair of straight styles.

This plate is nothing more or less than a forward continuation of the vomerine
cartilages (vc), the unossified parts being bridged over by a commissure. For relative
size, these vomerine cartilages * have their rivals only in the Snake, the Rhea, the
Hemipod, and in certain low-typed Passerines of the southern world (Notogea), namely,
Aneretes parulus, Pipra auricapilla, and in that remarkable type Pachyrhamphus.

Other characters in the Wren’s face are of importance, as illustrating what is com-
monly found in the numerous related forms.

The lacrymal (fig. 7, 7), often absent, is here very small; the lateral ethmoid (e¢h, pe),
on the other hand, is very large, and shows itself on the upper view ; its inferior angle is
separately ossified as an “‘ os uncinatum”’ (oz), or palato-trabecular conjugational bone.

Between and below the eyes the trabecular crested bar (¢) has been freely differentiated
from the ethmo-presphenoidal bar (pe), and the long overlapping process of the pterygoid
has become segmented into a distinct “‘ mesopterygoid ” (ms.pg), soon, however, to
ankylose with the palatine. In this rather young bird the transpalatine (fig. 5, tpa) was
scarcely confluent with the broad part of the long and slender palatine (pa). The deli-
cate pterygoids (fig. 5, pg) here are more like the epipterygoid of the Lizard than its ptery-
goid ; but the terminal part (apex of the bar) is alone upturned. This upturned part is
very long in some “ Coracomorphe ”—e. g. Coccothraustes, one of the most specialized of
the conirostral division of the group. In certain subfamilies of the “ Coracomorphe,”
although the maxillo-palatine processes do not solder the upper jaws together, yet the
palate is made more solid by means of an additional bony wedge, the “ palato-maxillary ;”
this grows in between the prepalatine bar and the body of the maxillary. In the “Car-
dinalide”’ (Plate XX. fig. 18, pa.mx) it is a large and thick bone; it is somewhat
smaller in the Buntings (Linberiza, Plectrophanes, Phrygilus), and also somewhat smaller
still in the “ Ieteridze,” “ Sylvicolidee”” (Mniotilta, Dendreca, Trichas, Chlorophanes),
and in the “ Tanagridee”’ (Zanagra, Prionocheilus, Stephanophorus).

That curious bone, the ‘‘ os uncinatum,” the matrix of which was first developed as a
conjugational bud from the trabecula, is very large in the Grosbeak ( Coccothraustes), as
in its climbing isomorphs the Parrots. The “posterior conjugational processes,” “ basi-
pterygoids,” and their counterparts on the pterygoid bones are early arrested in the

* Tf the Wren had possessed a “ recurrent alinasal,” like the long forked one of Vireosylvia olivacea, the two
subseptal floors would have reached each other.

AND DEVELOPMENT OF THE BIRD’S SKULL. aal

*« Coracomorphe ;” spiny rudiments, however, are not seldom seen, especially in southern
forms, as, for instance, Gymnorhina, Homorus, Dendrocolaptes, but most clearly of all
in a form. allied to the Sun-birds, namely Anthreptes malaccensis.

I have purposely mentioned these modifications of the face in the very numerous types
of the Crow form, and if it had been possible I should have sought to make the “ Agi-
thognathe”’ run parallel with the ‘“ Coracomorphe ;” but Nature herself forbids this.
Yet it is some gain to have been able, as Professor Huxley has done, to take in a lump
nearly half of the myriad of known birds, to show that all these are ‘ Aigithognathous,”
and to callthem by the name of the commonest and yet one of the highest of the group,
namely, the Crow. Here the zoological group cannot be made to hold all the mor-
phologically agreeing types, the ““symmorphs.”’ This will be felt still more strongly in
the next group to be considered, the “‘ Coccygomorphe”’ (Huxley, op. cit. p. 466),
which, with an extension of its boundaries to be proposed here, will contain “ Aigi-
thognathe,” “ Schizognathee,”’ and “ Desmognathee.”’

To make the matter a little clearer, it is necessary to show that there are serena
varieties or degrees in the morphological types of the ornithic palate, as designated by
Professor Huxley. In the “ Agithognathous” palate I find three varieties, namely :—

a. Incomplete.

6. Complete: var. 1.

¢. Complete: var. 2.

d. Compound.

a. Incomplete “ Megithognathism” occurs in the “Turnicimorphs,” Hemipodius or
Turniz. Here the vomerine cartilages are very large and incompletely ossified, and the
broad double vomer has a septo-maxillary at each angle; but these bones are only
strongly tied to the “ alinasal” cartilage, and do not graft themselves upon it.

b. Complete: var. 1. This occurs in some of the lowest harsh-voiced ‘ Coraco-
morphe.’ The vomers are developed in large “ vomerine cartilages,” which they often
only partially ossify ; but these osseous tracts are distinct from those of the often bony
alinasal walls and turbinals: a small septo-maxillary on each side generally appears,
limpet-like, on the angle of the alinasal inturned cartilage, but does not run into it.
This is well seen in Pachyrhamphus, Pipra, and Thamnophilus.

c. Complete: var. 2. This occurs in an-immense group, comprising most of the
“Coracomorphe”’ and also the Swifts (“ Cypselidee’’); here the vomerine bones are
grafted upon the nasal wall, and thus the bird loses its primary schizognathism,

d. Compound. 'This type occurs where, in a perfectly egithognathous face, desmo-
gnathism is produced by ankylosis of the inner edge of the maxillaries with a highly
ossified alinasal wall and nasal septum.—EHxamples, Gymnorhina tibicen, Paradisea
papuana, Artamus leucorhinus. Of this type a feebler form is produced when the
maxillaries only coalesce with the ossified alinasal wall, as in Dendrocolaptes albicollis,
Thamnophilus doliatus, and Phytotoma rara.

For Professor Huxley’s account of the Desmognathous palate, I refer to his paper
(p. 485). Ihave one remark to make, namely, that the vomer is very broad antero-
inferiorly (as in the Chelonia) in the most perfectly desmognathous Raptorial palate,

112 MR. W. K. PARKER ON THE STRUCTURE

that of the “ Falconide,’’ also that the vomer is not small in the “ Ardeide”’ and
Spoonbill, but very large; it is primarily double in them, but mostly single in the “Coc-
cygomorphe ”’ (when present) and in the “ Chenomorphe.”’ In the gallinaceous “ Schizo-
enathee’’ the vomer is azygous ; in the charadrians it is double.

The varieties of the Desmognathous palate are four, namely :—

a. Direct, as in Falcons and Geese, when the maxillo-palatines meet below at the
mid line, as in the mammal. Two subvarieties of this form occur—in the Falcon, where
the nasal septum is ankylosed to the hard palate, and in the Goose, where it remains
free:

b. Indirect. This is very common, and is best seen in Eagles, Vultures, and Owls;
the maxillo-palatines are ankylosed to the nasal septum by their inner margin, but are
separated from each other by a chink. This is well seen also in the fledgling of the
Falcon, which is ixdirectly desmognathous at that early stage.

c. Imperfectly direct. This is where the maxillo-palatine plates are united by har-
mony-sutures, and not by coalescence. Example, Dicholophus cristatus.

d. Imperfectly indirect. Here the maxillo-palatine plates are closely articulated with
and separated by the median septo-maxillary, but there is no ankylosis. Hxample,
Megalema asiatica.

e. Double desmognathism. <A fifth variety may be added in such a case as Podargus,
where the palatines as well as the maxillaries largely coalesce below; to aless extent this
is seen in the gigantic species of the Hornbills, e. g. Buceros birostris (see Huxley, op.
cit. p. 446, fig. 28, PZ).

Having thus analyzed their morphological characters, we shall be better able to deter-
mine their zoological value. Although my proper business in this paper is the deter-
mination of purely morphological meanings, yet I venture to offer these to the zoologist
embarrassed with the dazzling variety of types.

On the Structure and Development of the Face im Cuculine Birds.

After the many hundreds of the “ Coracomorphe”’ have been set aside, there yet
remains a large residuum of high-class “ Arboreal birds.”

Some of these are easy to cull from the rest, as they “pass under the hands of him
that telleth them,” without any confusion. Such are the Pigeons; such also are the
Parrots; and the two “ Families” of the Woodpeckers stand apart. A legion, however,
of other forms follow; these, for convenience’ sake, may take their group-name from the
Cuckoo, e. g. “ Coccygomorphe.”

Professor Huxley’s ‘‘ Cypselomorphe”’ (op. cit. p. 468) is his worst group; and he
has evidently tied its subdivisions together under some degree of misconception of their
palatal morphology. rochilus has not “a true Passerine vomer” (op. cit. p. 454), and
neither this bird nor the Goatsucker is Mgithognathous : Cypselus is, however ; and yet,
lying close by the true Swallows (“Hirundinidz”’), it differs in many most important
particulars from them, having no singing-muscles, no czca coli, and is possessed of
organs of flight very similar to those of the Humming-bird.

In fundamental cranio-facial morphology the Swifts are most Coracomorphous, and

AND DEVELOPMENT OF THE BIRD’S SKULL. 113

the Goatsuckers least: the Goatsucker, agreeing in very important characters with
Podargus and Steatornis on the one hand, and with the Trogons on the other, lies alto-
gether on a lower and more embryonic level than the “ Coracomorphe,”’ especially those
to which the Swifts are allied. These other ‘“‘ Families” may take their place for the
present with the other equally discordant “‘ Coccygomorphe,” and the term “ Cypse-
lomorphz’’ be allowed to drop. The Swift, so far as its cranio-facial morphology is
concerned, is an ‘“‘ Aigithognath,” and I have treated of it in a paper which has been
read at the Zoological Society. The Goatsucker and Humming-bird come under the
plan of the present memoir.

On the Structure of the Face in Caprimulgus europzus.

The moth-shaped mouth of the Fern-Owl is roofed in in a manner peculiar to itself;
but its gaping face is developed by a process in nowise different from that which operates
in. its nearer or more distant relations.

In the basal figure (Plate XXI. fig. 8) the trabecular structures are largely hidden
by the palatines ; and to understand this most frog-like of all the skulls of the “‘ Carinatze,”’
the student must examine the actual object.

Bony matter is here spun to its uttermost degree of fineness, and the resulting sub-
stance is something more like feather than bone. The whole shape is flattened and
outspread, and, as in the Batrachia, the originally flat broad trabecule have retained
much of their primary form. In front, however (Plate XXI. fig. 8, and Plate XXII.
fig. 1, sv), they are narrowed, and quite absorbed anteriorly, as in all other birds; at
least, the azygous prenasal outgrowth is early absorbed. Behind, the bony trumpets
that ‘grow from the ossifying apices of the trabecule spread far apart, forming the
“anterior tympanic recesses” (atr), and enclosing the V-shaped double Hustachian
tube (ez).

The trabeculz, as they surround and converge in front of the pituitary body, are con-
verted into a broad and winged mass of bone by the parasphenoid (pas), and at each
side of the common Eustachian opening there are two “ conjugational processes ”’
(fig. 8, bpg). ‘The lower of these is the “‘ basipterygoid ” proper, and it carries the free
cartilaginous plate for pterygoidean hingement; this flat broad process looks forward.
Above this is a retral hook, which merely holds a ligament.

As in the Frog, the trabecular and parasphenoidal structures project but little below
the lateral ethmoids (Plate X XI. fig. 8, pp), so that each pars plana seems to grow out
from the median, beam-like, large-winged outgrowths: they have carried the “ palatine
conjugational process” (ow) with them. This does not ossify into an “ os uncinatum,” as
is common in the “ Cuculinz,” but remains persistently soft as hyaline cartilage.

In front, between these ecto-ethmoidal wings, the broad parasphenoid and the trabecular
base of the meso-ethmoid each separately taper to a point; and here the first facial arch
has part of its long commissure absorbed to form the cranio-facial hinge.

In the pin-feathered nestling the trabecular base of the septum nasi (Plate XXII.
fig. 1, sv) is rather solid, but the azygous rostrum has been already absorbed. In the
adult (Plate XXTI. fig. 8, sz) the “ posterior septal bone” is rather solid than spongy, and

114 MR. W. K. PARKER ON THE STRUCTURE ©

its edges can be seen in an upper view (Plate X XI. fig. 10) projecting beyond the nasal
processes of the premaxillaries and upper spur of the nasals, and in the lower view
(fig. 8) above and in front of the vomer. But the rest of the trabecular part of the
septum nasi is absorbed, and only its nasal part is retained; it is bony, and the bony
matter passes a short distance into the paired “ale” (figs. 8 & 10). As to the secondary
or splint-bones of the trabecular arch, I have already spoken of the parasphenoid: the
others are, in this case, the premaxillaries and the vomers. The former (fig. 8, px)
appear very small on the lower surface, but above (fig. 10) each half is seen to be equal to
the corresponding maxillary, and to overlap it, as in birds generally and Teleostean Fishes.
The dentary margin, however, does not reach nearly so far back as in birds generally, and
this double bone is ornithically very small.

The other double bone isthe vomer. This is a long bar of spongy bone, rather bluntly
pointed at the fore end and sharp behind (figs. 8 & 9, v); it is quite distinct from the
overlying ethmo-palatal bars in newly fledged birds (fig. 9); but in the adult (fig. 8) it
becomes a derived commissural band to the palatine arch, as in a large number of birds,
notably the “ Coracomorphe.”

Flat above, where the palatal bars lie upon it (fig. 9, v, epa), the vomer is carinate below
throughout its entire length (fig. 8). In the younger (pin-feathered) fledglings I found
the vomers distinct throughout their whole length (Plate XXII. fig. 1, v). They were
long styles, approximating very closely in front, and gently diverging behind. They
were seen to have been formed by ossification of a fibrous and not a cartilaginous tract.
In more mature fledglings the fore ends had united (Plate XXI. fig. 11, v), and the
hinder ends were still more divergent. These again approximate, ankylosis takes place
thoroughly fore and aft, and the inferior surface ossifying, the interpalatine “ raphe ”
acquires its long keel. The fore end of the vomer in the adult (fig. 8) becomes rounded,
losing the beaked form it had in the newly fledged bird (fig. 9), and at this part below
the maxillo-palatine hooks (map) are notched on their inner margin to articulate with
this extremity of the vomer. A character like this gives the palate a very ‘“ Aigitho-
enathous”’ appearance, as this curious articulation, which bridges over the chink on each
side, is often seen in southern “ Coracomorphe,” both those of low and also of a high
type, as in Grallaria, Thamnophilus, Dicrurus, Dendrocolaptes, Homorus, and G'ymno-
rhina. But I have already said that these vomers are membrane bones; and I here show,
in three stages, the longish ligament which binds the vomer to the internal alinasal
wall with its long worm-like turbinal (Plate XXII. fig. 1, and Plate XXT. figs. 9 & 11,
aln,v). In the second of these three stages (fig. 11), the inturned alinasal wall is seen
to lie on the broad maxillo-palatine ; but it stops far short of the vomer, which also fails
to spread its shoulders towards the cartilage.

The palatine arch of the Fern-Owl equally departs from the Coracomorphous type, and
cleaves to what is signally Cwculine—I mean, in the broad sense of that word.

It has been already shown that the distinctively Passerine or Corvine character of the
palatine is the metamorphosis of the primary bar into fowr pairs of true morphological
endoskeletal elements, to say nothing of the secondary bones. But in Caprimulgus, my
youngest specimen (Plate XXIT. fig. 1) only shows a trace of true cartilage on the edge

AND DEVELOPMENT OF THE BIRD’S SKULL. 115

of the overgrown “ transverse”’ region; and the first and only palatine centre runs riot
over both morphological territories. Also in the hinder division of the arch-moiety the
pterygoids correspond to those of the young nestling, but not to those of the fledgling Pas-_
serine; for each pterygoid sends a spike-shaped process over the palatine; but this does
not become a “‘mesopterygoid ” bone (Plate XXI. fig. 11, and Plate XXII. fig. 1, pg).

The lower fore-looking spines of the palatines do not increase much in size—compare
the “ interpalatine”’ of the young (Plate XXII. fig. 1, ipa) with those of the adult
(Plate XXI. fig. 8). But the upper or orbital processes, the ethmo-palatine bars, grow
exceedingly, and meet by suture at the mid line (compare Plate XXTI. figs. 8, 9, & 11,
epa). This apposition is a step towards a true facial “‘ commissure.” The palatines are
elegantly bowed outwards from each other behind this tract, and then form a complete
commissure, like that of the trabecule (Plate XXI. figs. 8 £9). By the time this bird
is fully fledged the posterior palatal region is joined together at the mid line by a
cartilage common to both sides, and this cartilage becomes ossified by two endosteal
centres ; these I propose to call the anterior and posterior “ medio-palatines ” (Plate X XI.
figs. 8 & 9, mpa). Now the morphologist, at first ‘rounding about in darkness,” feels
himself safely within the great Cuculine territory, knowing that there and amongst the
** Celeomorphous ”’ tribes (Picidz) this structure is alone to be found *.

With regard to the articulation of the pterygoids with the basipterygoid processes
(bpg), Caprimulgus is countenanced by many types within the great Cuculine circle.
I see them everywhere in smaller or larger rudiments; but they are perfect in a very
near relation, the Oil-bird (Steatornis), and also in the Trogons and Humming-birds.

There are but two pairs of palatine splints, the maxillaries and jugo-quadrato-jugals
(me, 7); the latter are mere curved needles of bone; the former are elegantly ornithic pieces,
with, however, characters that are peculiar to this bird. In the young (Plate XXI.
fig. 11, and Plate XXIT. fig. 1, mx) the maxillary differs but little from that of Struthio
camelus (** Ostrich Skull,” plate viii. fig. 2, and Huxley, “Classification of Birds,”
p. 420, fig. 1, xp); for the palatal plate is rough, large, and 4-sided, and the cuneiform
body of the bone is very similar with its zygomatic process. In the adult, however
(Plate XXI. fig. 8), the change is as though the most delicate chiselling had been
fining it down into a very Passerine element ; the maxillo-palatine hooks especially might
belong to a southern Crow. The bone has its own caprimulgine diagnostic in an
f-shaped fore-turned spur, which is applied, for defence, to the nasal wall on each side
(Plate XXI. fig. 10, mp.mx). The figures show how thoroughly non-passerine is the
nasal labyrinth (Plate XXI. figs. 10 & 11, and Plate XXII. fig. 1, aln, als, atb, itb);
the upper view (fig. 11) shows, through the transparent tubular nostril-walls, the ali-
nasal turbinal (a/b), which is exposed in the inferior view of the younger specimen.” The
inferior turbinal (2¢b) is similarly displayed in the two figures, its line of insertion being
seen from above, and its anterior half is shown in the lower view (Plate XXII. fig. 1, 7¢6).
I have already described the relation of the nasal wall to the vomer, and have only to

* T am well aware that in the larger Gralle, Mycterta, Baleniceps, Ciconia, &c., and the “ Totipalmatz,” the
palatines form a bony “ commissure ;” but there is no key-stone in their palate. In the Pelicans this common bar
sends upwards a crest which is the symmorph of the trabecular crest.

SECOND SERIES.— ZOOLOGY, VOL. I. R

116 MR. W. K. PARKER ON THE STRUCTURE

add that the upper and middle turbinals are not more developed than in the common
Fowl. The small nasal septum (sz), diminishing forwards, is the only part ossified in
front of the hinge, the bone running a little into the alee ; and also in the true olfactory
region the root of the simple aliethmoid alcoil is alone ossified ; and this growth comes
directly from the perpendicular wall, or meso-ethmoid. The lachrymal (fig. 8, /) is a large
spongy bone, broad in the roof of the skull, pedate where it binds upon the zygoma,
and it entirely excludes the lateral ethmoid from the upper and outer regions of the face.
This is a Cuculine character.

On the Structure and Development of the Face in the Humming-birds
(** Trochilide’’).

Professor Huxley (op. cit. p. 454) says “ Trochilus has the true Passerine yomer, with
its broad and truncated anterior and deeply cleft posterior end. I have not yet been
able to obtain a perfectly satisfactory view of the structure and arrangement of the
palatine bones in the Humming birds.” That same paper contains several figures drawn
from his friend’s preparations ; and I wish those which could have been supplied to him
of the Humming-bird’s skull had been used by him also; they would have prevented the
error contained in the above quotation. I have been long familiar with these parts in
the old and nestling bird, in specimens from Barbadoes (? Lampornis); and now I am
able to add the structure of the largest known species, from the Andes of Peru (Patagona
gigas) *.

It might have been supposed that the Humming-bird would have been especially
related to its “ Tenuirostral’? isomorphs the Sun-birds (“ Nectariniide”’) and their
allies; but I find nothing in these, except extreme prognathism, which indicates any
special relationship. Having carefully dissected and drawn several scores of Passerine
types, to say nothing of the Cuculines over the border, it seems to me not altogether a
difficult task to weigh and measure the relationships of these bright “ minims” of
vertebrate life. No isomorphism apparent in types that grow from the uppermost twigs
of the life-tree should satisfy the ornithologist in matters of affinity; down by the
root must morphological observation be commenced. I had hoped to find the Trochiline
congeners in southern-world forms, such as Nectarophilus and Anthreptes, from Celebes,
or in the Australian genera Acanthorhynchus and Ptilotis; but such is not the case.
Morphologically—I do not say zoologically—I am nearer my starting-point down among
the superstruthious Hemipods, and the Passerines that have sprung up from that low
level, such as Pipra, Pachyrhamphus, and Thamnophilus.

All this is said in relation to Passerine modification, as such; not as seeking to lower
the idea of the specialization of the Trochiline type, which is without rival. These
zoological ideas force themselves forwards; but my work is with the morphology of the
region of the face.

* The old and young specimens came to me, the former in spirits, the latter dried in a letter, from my friend the

Rev. Jno. Philp, for several years missionary in that island. The gigantic form, also a spirit specimen, belongs to my
excellent co-worker in this field, Professor Alfred Newton, F.R.S.

AND DEVELOPMENT OF THE BIRD’S SKULL. 117

The forth-drawing (prognathism) of the facial bars has been carried to its uttermost
extent in the Humming-bird; but the secondary parts of the trabecular arch occupy two
thirds of the length anteriorly, their endoskeletal axis or prenasal rostrum early under-
going absorption. Behind, the apices of this arch are buried and lost in the outworks
of the tympanic cavity; and below, just in front of this part, the great splint, the para-
sphenoid, marks the bowed converging bars (Plate XXII. figs. 2,3, 4, 8, pas). Here, just
outside the common Hustachian meatus (ew), the trabeculae have sent out conjuga-
tional spurs, with their obliquely fore-looking facets for the pterygoids.

This high form gives us the extreme of a series commenced in the Struthionidz, where
the hinder end of the pterygoid is wedged in in the Emu between the quadrate and
basipterygoid process. Hven in some “ Ratitee” (see “ Ostrich Skull,” plates vii—xiv.)
the articulation is further forwards, whilst the series is obtained by passing to the
Tinamou (plate xv.), the Syrrhaptes (Trans. Zool. Soc. 1863, vol. v. pl. xxxvi. fig. 1);
then come the Plovers, Pigeons, Owls, &c., where it is in the middle; the “ Alectoro-
morphze”’ and ‘‘ Chenomorphe,’’ where it is at the anterior third; and, lastly, here in the
* Trochilide”’ it is quite at the fore end. Of necessity, this relative motion, as it were>
of the palatine on the trabecular arch is attended with shortening of the basipterygoid
processes as we pass along the series from the Emu to the Humming-bird. The tra-
becular base of the meso-ethmoid (Plate XXII. fig. 3, pas) is short, broad, pointed, and
keeled in front; and the hinge between this part and the septum nasi is finished by
uncleft cartilage above (Plate XXII. fig. 4, eth, sn). The remainder of the trabecular bar
is one with a most small ¢rwe nasal septum; there is a larger posterior and a smaller
anterior osseous centre in this small cartilaginous keel (fig. 4, sv). There are several
secondary bones applied to the trabecular arch besides the hinder plate, or parasphenoid.
In front the premaxillaries (pr) take the lead; they are here at their highest develop-
ment; they reign in this type, especially, and all things else in the organism are in con-
formity with them.

In the old bird (fig. 2) the premaxillaries are three fourths the length of the face; in
the nestling (fig. 8) they are only two thirds; so that it is evident that this excessive
prognathism takes place by the working of a general morphological law.

The boundaries of this double bone are best seen in the young (fig. 8); the dentary
processes (dpa) nearly reach the maxillo-palatines (map) behind, and the palatine pro-
cesses (figs. 8 & 9, ppx) are the twentieth of an inch shorter; as usual, the nasal pro-
cesses (fig. 5, wpa) are strongly grafted on and ankylosed to the forehead. Laterally
the upper and lower portions of the premaxillaries are thinned out extremely, and present.
ragged edges towards each other: this can be seen in the lower view (fig. 2). The halves
of the vomer are thoroughly fused together in my young specimen (figs. 8 & 9); they show
no sign of being formed on any thing else than a tract of fibrous tissue, and their likeness
and dissimilarity to the true Passerine vomer is well shown.

The long anterior spike in which the coalesced pieces terminate in front would not of
itself exclude them from the “ Aigithognathe ;” in the typical Chough (Pregilus graculus),
and in the aberrant Lyre-bird (Wenura superba), the double vomer is blunt-pointed in

front ; but in them, supero-laterally, the scars are seen in the dry skull where the alinasal
R 2

118 MR. W. K. PARKER ON THE STRUCTURE

cartilage has been macerated away. Besides the absence of the vomerine cartilages, we
have, then, in the “ Trochilide,” a lax relation of the vomer to the alinasal wall (see
figs. 2, 3; 8, 9, v, ial), the two being connected by a fibrous ligament. But the bone is
well shouldered, and above the broadish part the additional bones are seen, namely, the
‘* septo-maxillaries”’ (figs. 2-4, sma), so that the morphological elements are all here
except the pair of vomerine cartilages ; yet the metamorphosis of the parts is arrested
and the elements remain distinct. Besides the paired septo-maxillaries (smz), there is an
odd one, lying on the vomer, below the septum nasi, and in front of the small pair of centres
(fig. 4, jma’). These three,bones coexist in certain Aigithognathee, as Gymnorhina, where
they all coalesce with the vomer, and Homorus, where the median piece remains distinct:
This azygous piece I often find when the vomer itself is azygous, as in Anser palustris,
Psophia crepitans, Circus cyaneus, Haliastur indus, &c.; it is related, morphologically,
to the base of the cranio-facial notch, and is distinct from the centre which ossifies the
postero-inferior part of the septum nasi. Thus the unspecialized submucous plates seen
in the vomerine region of the Sturgeon reappear in their old profusion in these nobler
types, but are specialized to the highest degree.

The next arch, the pterygo-palatine, has as much and as little of the Passerine type as
the one described. ‘The palatines themselves are of great length (figs. 8 & 9, pa), and
are extremely slender, especially the distal part; they thicken gradually to the widest
part, and then gently taper off to their posterior end. On the inside of the widest part
each bone sends forward a long interpalatine style (¢pa) ; and on the outside a wart-like
protuberance represents the transpalatine of the “ Algithognathe.” This is only seen
in the adult, and isa mere periosteal outgrowth (figs. 3 & 4, ¢pa).

In some low-type Passerines, such as Pipra and Pachyrhamphus, the transp2latine is not
much larger; in the young Humming-bird the palatines are as much deficient here as
those of the Hemipods. The ethmo-palatine lamina (figs. 9 & 10, epa) is very much
stunted, the reverse of what obtains in the Swift and Goatsucker, where it is of extreme
length; and in them the vomerine crura articulate and then coalesce with these lamine,
whereas they remain distinct in the “ Trochilide.” The short stoutish pterygoids (pq)
are scooped in front, where they rest upon the end of the palatines ; in the young (fig. 10,
pg, mspg) the mesopterygoid piece is nearly distinct from the body of the bone; it
becomes quite separate, and then coalesces with the palatine. Both palatines and
pterygoids are faceted with cartilage to glide upon the short, stout, flat-faced basi-
pterygoids (bpg). Hach pterygoid is sharp-edged both above and below, and above the
shallow cup for the pterygoid there is scarcely any epipterygoid process (epg). In this
these birds are contrary to the “‘ Aigithognathe.” In the young (figs. 8 & 9, mx) the
maxillaries are clearly traceable; the broad flat part is about equal in length to the
long terminal spike, which is equal to that of the palatine; already, the posterior style
(zygomatic process) has coalesced witb the long jugal (j). The dentary edge is quite
overlapped by the premaxillary, opposite the end of which we see the characteristic
ornithie process, the “ maxillo-palatine.”

These outgrowths from the maxillaries are of great interest to the morphologist ; here,
in the Humming-bird, they show relationship as by the most delicate test. In the young

AND DEVELOPMENT OF THE BIRD’S SKULL. 119

(figs. 8 & 9, map) they agree in general with those of young Coracomorphe and Hemipods,
but in particular with both young and adult of that family of birds which has most
similarity to them in general bodily structure, namely Swifts (‘‘ Cypselide”). The
process itself is at first a mere hook, moderately thick, rather long, curving at first
inwards, and then directly backwards. This perfectly retrorsal rod runs directly along
the inside of the palatine; in the Swift it clears itself of the palatine, and is altogether
more flattened and sickle-shaped.

The maxillo-palatine of the ‘“Trochilidz’’ becomes more and more underlaid by the
palatine (figs. 2 & 3), so that a lower view shows little of them, and the specimen has to
be carefully studied in oblique aspects before its form and relations can be understood
(fig. 4, pa, mxp). Then it is seen that the little rod has become a flat thin plate, with a
terminal hook, and with its lower edge entirely ankylosed tothe palatine. This is similar
to the condition of things found in Cranes and Plovers, and totally unlike any thing seen
in the “ Agithognathe.” As far as my memory serves me, I have never seen any
ankylosis of these parts in the ‘‘ Coracomorphe,” not even in such highly ossified faces
as those of Phytotoma, Coccothraustes, Gymnorhina, or Artamus; the palatines, even
in these, keep scrupulously apart from the overlying maxillo-palatines.

The huge nasal labyrinth is much more in conformity with its counterparts in the Pas-
serine than the facial bars. The alee nasi (alm), like those of the Hemipods, are large
mussel-like valves, and have this special character, namely, that the widest part of the
nostril is behind; the teleological meaning of this. structure “lies at the door,” like the
other specializations of this type.

Unlike the generality of the small and delicate species of the “‘Coracomorphe,” such
as Acanthorhynchus, Nectarophilus, Phylloscopus, Troglodytes, Chelidon, &c., the nasal
outworks and walls are extensively ossified. The posterior septal bone is present, and
has a smaller ossicle in front of it (fig. 4, sv), and the anterior alar region (figs. 2-6, aln)
is ossified right and left in the recurrent lamina (7cc) and in the free valves; in the
latter there is a margin of cartilage left, and this has a selvedge of bone formed by a
distinct centre on each side. ‘This is the sickle-shape counterpart of the annular alinasal
ossicle of certain Parrots (Melopsittacus undulatus &c.). The large alinasal turbinals
(ath) are ossified to a considerable extent, the inferior turbinals (ib) scarcely at all.
The anterior nasal floor is but little developed; yet it sends inwards its inturned lamina
(ial), a broad unossified flap, to be tied to the vomer by a broad ligament (éal, v). The
true olfactory region (ecto- and meso-ethmoids) is of great size, and is densely ossified :
thoroughly Passerine, it has completely aborted the lacrymals, and appears on each side
of the frontals above (figs. 5 & 6, eeth), whilst below it forms most extensive antorbital
wings (pp), the suborbital part of which, reaching two thirds of the way towards the
quadrate, has no separate ossification of the ‘‘ os uncinatum.”

The skull is depressed: grooved above symmetrically for the long hyoid cornua
(8rd arch), it has its ethmo-presphenoidal plate (ye) solid, as in the Carp, and non-
fenestrate. The orbito-sphenoidal lips are narrow, and the orbital plates of the frontal
very little developed, so that the tilted floor of the cranium is largely membranous. The

120 MR. W. K. PARKER ON THE STRUCTURE

pier (quadrate) and proximal part of the mandible is much like that of the Swift. The
symphysis is long, and the distal part a mere copy of the upper jaw.

The 2nd and 3rd postoral arches are merely outdrawn modifications of the normally
ornithic hyoid bone, much less specialized than in the Woodpeckers. The true cerato-
hyals (fig. 7, chy) only extend for one fourth up the interior of the double horny sheath ; but
another fourth is filled, first with fibrous and then with fatty tissue. A short tract of
each bar is ossified, and all but the small bilobate end of the middle, or basi-urohyal bar
(bhy, bbr). The proximal part of the 3rd postoral (67) is only about one fourth the
length of the flat distal piece, which reaches, when the tongue is retracted, as far as to
the premaxillary apices: both are fully ossified.

Any one who will repeat these observations will find that the “ Trochilide”’ have no
right within the Passerine territory, and that they have at length diverged very widely
from the most striking tsomorphs within that range. The ‘‘ Coccygomorphe,” if they
be considered broadly, will be seen to be their truest relations; they are not, however, a
normal branch, but should be looked upon as forming a very distinct secondary leader,
arising close to the root of this widely ramifying group. It is well worthy of remark that
that Caprimulgine outlier of the “ Aigithognathe,” the Swift, comes very much nearer
the Humming-bird than the “ Nectariniide,” birds so mimetic and isomorphic of these
gorgeous “‘ vertebrated insects.”

On the Structure of the Face in Scythrops novee-hollandiz.

It would be difficult to find a more central form than Scythrops for the whole of the
* Coccygomorphe.” It appears to be equally allied to the Hornbills (“ Buceride’’),
the Toucans (“ Rhamphastide”’), the Plantain-eaters (“ Musophagidze”’), and the true
Cuckoos (“ Cuculidee’’). This paper, however, is given, not so much for the elucidation
of a difficult taxonomy, as for the purpose of exhibiting the more striking modifications
of the face and nasal organs in highly specialized birds. Ankylosis is in this case carried
to its uttermost limits as far as is consistent with the well-being of the type; yet the
regions can easily be determined. Rudiments of basipterygoid processes (Plate XXIII.
fig. 1, bpg) remain on each side of the subcarinate posterior part of the basifacial bar;
the bar becomes rounded and synovial where it glides along the common palatine groove
(fig. 2, pe, pa). In front of the hinge the bony septum nasi becomes lost in the delicate
diploé that, sponge-like, fills the. large upper jaw. Besides the huge premaxillaries,
which have melted into the common mass of the upper face, there is a well-marked
vomer (figs. 1 & 2, v), which appears as a thickish subcarinate plate of bone between the
maxillo-palatine wedge and the approximated interpalatine lips. This lesser bony wedge
thickens again above, and then thins out to articulate by fibrous tissue with a supero-
posterior ossicle. The vomer of Scythrops is an azygous bone, of the Chelonian type,
such as is also seen in Raptorial Birds, and in Mammalia of the Cat tribe ; but the hinder
part has been detached, and has coalesced with the palatines (ptpa, v).

There is, however, another element of the trabecular arch in this bird, which makes it
one of great interest to the morphologist; this is the os uncinatum. Notwithstanding
the normally large size of the lacrymal (fig. 3, 7), the ecto-ethmoid, or pars plana (pp),

AND DEVELOPMENT OF THE BIRD’S SKULL. 121

is a great squarish plate of bone; as in Caprimulgus, the infero-external angle of this
plate has its ossification stopped short, leaving that tract which primarily grew out of
the trabecule to conjugate itself with its counterpart process on the palatine bar. This
yoke-band is proximally broad and triangular (figs. 1 & 3, ow), and the apex of the
triangle, which alone is ossified, grows into two long and slender lashes of cartilage, one
of which runs directly across to its endoskeletal symmorph, tying itself to the very apex
of the ethmo-palatine crest (fig. 3, epa, ow), whilst the other runs forwards and down-
wards, and is attached to the upper surface of the palatine, near its outer edge and
opposite the fore margin of the maxillo-palatine floor (fig. 1, ow, mxp). The strong
palatine arch has, notwithstanding its intense specialization, attained the very form
which the foremost (trabecular) arch assumes as soon as it can be seen in a vertebrate
embryo as clouded bands of granular bioplasm. Here we have symmetrical sigmoid
bands, which behind (at their apex) are strongly hooked upwards and inwards; these
bands approximate, coalesce, and then diverge to their free flat anterior ends. Whilst
undergoing this junction the trabeculz form not only a commissure but a crest ; here, in
the palatine arch of Scythrops (figs. 1-3, pa, epa, pg, mspg), each bar grows upwards at
the commissure, which forms a semicylindrical trough above (fig. 2), on each side of which
the ethmo-palatine processes (trabeculo-palatine conjugational outgrowths) project.

Each pterygoid (pg) expands in a very ichthyic manner on the fore edge of the
quadrate, the expansion being the posterior margin of the subterminal part of the bar ;
the real apex is the epipterygoid hook (epg), which is as well developed here as in the
Finches. Each pterygoid is a strong bone, sharply keeled both above and below; leafy
above and hooked below, where it articulates with the palatine. It has evidently yielded
a mesopterygoid element (mspg) to the upper cartilage of the palatine behind.

The postpalatine region is vertical, the prepalatine broadly horizontal; the middle
part is mixed. The upper edges have united by ankylosis, not, however, without a
key-stone, a detached postvomerine piece, and perhaps also a medio-palatine (figs.
1 & 2, v, pa).

The deep thick-edged interpalatine lips, where they embrace the exposed vomerine
plate, nearly meet (fig. 1, ipa), so that altogether the lower or hard palate is of great
extent and complexity. By far the greater part of the hard palate is due to the larger
growth of the median and lateral palatine portions of the overgrown premaxillaries
(ppzx); but there is no little of the maxillary element besides. The proximal or lateral
portion of the maxillo-palatine plates (figs. 2 & 3, mxp) is spongy and of great width ;
it rises (fig. 3) up in front of the ethmo-palatine (fig. 3, epa); but the heart-shaped
median portion, which belongs to both sides, fills in half the space between the median
palatine process of the premaxillary and the gap formed by the diverging palatines. My
figure (fig. 1) does not show more than the inferior surface of this common median
maxillo-palatine mass ; but above, in front of the vertical vomer, it swells out into a very
elegant bulb of spongy bone, which is exactly heart-shaped, the base looking upwards.
The ankylosed ‘“‘jugal” bar (j) is strong, and greatly inhooked behind. The small
ossified alee nasi with their bony rudiment of a turbinal (fig. 3, alm, atb), form a nostril
that looks downwards and backwards. The whole labyrinth is huddled into a small

122 MR. W. K. PARKER ON THE STRUCTURE

space between the nostril and antorbital, and the short inferior turbinal is ossified. The
ecto-ethmoid (fig. 3, pp) is entirely excluded from the frontal region.

On the Structure of the Skull in Megaleema asiatica.

Having described a good representative Cuculine face, I take as my next instance that
of a bird which has a doubtful and mixed morphology. The physiognomy of the living
bird corresponds with this; for the most trained eye, seeing it for the first time, is
arrested by a sort of play and interchange of features, now looking like a relative of the
Cuckoo, and anon seeming to belong to the Southern Shrikes or Crows (e.g. Vanga,
Coronica, Gymnorhina).

Seen from below (Plate X XIII. fig. 4), the imperfectly “ desmognathous” and sezmi-
passerine characters are seen at one view. ‘The bony structures have the same Ouwl-like
spongy character as is general in the Cuculine birds; in the ‘“ Coracomorphe ” they are
more fibrous and stronger; they are more cleanly built.

In Megalema the basipterygoids are quite aborted, and the basifacial beam is
rounded to its end, the parasphenoidal rostrum ending below the hinge as a blunt spur.
In front of the hinge there is a solid postseptal bone, which ossifies that part of the
trabecular bar; for the rest, the septum nasi is aborted to a great degree, yet the recur-
rent alinasal folds send backward a style of bone which reaches almost to the vomerine
forks. The nasal labyrinth has its scrolls stunted ; those of the “ale” are partly ossified,
and the alinasal wall has become thick bone externally (ain); in front this bar divides
the external space into an anterior ¢rwe and a posterior false narial opening, as in many
Cuculines.

The spongy premaxillaries are remarkable in having their thick, hollow, rod-like
palatine processes free posteriorly (fig. 4, ppx). The dentary termination is equally
thick, but its lower edge is sharp and dentate.

The os uncinatum is not distinct from the pars plana (figs. 4 & 5, ow, pp), but it has
retained its primary embryonic shape as a somewhat hooked triangular flap. The
vomerine series of trabecular splints is extremely interesting. There is no evidence of a
cartilaginous origin for the evidently double vomer (v); but its explanation must be
sought for in aseries of forms outside the Cuculine boundary, as well as near the border,
namely, in the Humming-bird. As in Gymnorhina, Homorus, and Trochilus there are
jiwe vomerine bones, two vomerine moieties, forming a most elegant Y-shaped bone, a
pair of septo-maxillaries, and a median piece (fig. 4, v, sma, sma’). When the septo-
maxillaries are well developed, as in the ‘‘ Ophidia,” they stretch themselves beyond the
vomers, fore and aft, as is well seen in the Cobra (Naja tripudians); but when they are
feebly developed, a bony rudiment may appear in any part of the tract occupied by them
in their larger growth.

Thus in the Sparrow (Plate X XI. fig. 4, smx) they appear outside the broad shoulder
of the vomer ; in the Humming-bird in front of the broadest part; and here, in MWegalema,
behind, answering to that terminal spike of the bone in the Cobra which articulates
with the ethmo-palatine crest. I am satisfied that this is the true interpretation of the
posterior pair of vomerine bones in Megalema; for the bone does not form a palatine com-

AND DEVELOPMENT OF THE BIRD’S SKULL. 123

nmussure, and the medio-palatine, which occurs there, is azygous. The median septo-
maxillary turns up again in the Woodpecker; it has been already described in the
Humming-bird (Plate XXII. figs. 3 & 4, sma’); it exists free in the Coracomorphous
genus Homorus, and ankylosed to the vomer in Gymnorhina. For a vomer like the
one figured here (fig. 4, 7) we must look to some Celebesian Passerines. I have it in
Nectarophilus Grayi and in Anthreptes malaccensis; especially characteristic is that of
the former *.

But the vomer in these Passerines is confluent by its forks with the inturned alinasal
wall; in Megalema the arms of the Y form a neat joint, apparently without a cavity
(Plate XXIII. fig. 4, v, map), with the corresponding maxillo-palatine plates. The nearest
relation of this type known to me at present is the Roller (Coracias garrula) : this is one
of Cuvier’s “‘ Passerine ;” but it is far less passerine than Megalema, and has a chelonian
azygous vomer. ‘The narrowish, parallel, neat palatines are very much like those of the
Roller, but they are narrower, have a less distinct inner keel, and have distinct interpalatine
processes (pa); the rudimentary transpalatine angle (¢pa) is nearly equal in the two
kinds. The large mesopterygoid (fig.5, mspg) is semidistinct, and has not united with the
palatines in this specimen. In Coracias these parts are quite passerine, the mesopterygoid
having united with the palatine on one side, the left, whilst on the other all these are
fused together. In Coracias the epipterygoid process is abortively developed; in
Megalema (fig. 4, epg) it is an elegant rounded lobe. In Coracias the maxillo-palatines
are large, and have a complete ankylosis with each other; in Megalema (fig. 4, mxp)
they are smaller and do not meet, being kept apart by the “ median septo-maxillary ”’
(smz’).

In Coracias the lacrymal has its largest development and the ecto-ethmoid the least ;
in Megalema these things are reversed, and the lacrymal (fig. 5, 7) is a mere point,
whilst the ecto-ethmoids (eeth, pp) are like those cf the Passerines and Hemipods, being
largely exposed in the frontal region. In the structure of the nostrils the two types
agree; but conformably with its more intensely cuculine character, the Roller has a
very massive, spongy, osseous nasal septum ; it also has very large but non-functional basi-
pterygoids.

The characteristic modifications of Coracias are exaggerated in Hurystomus, save that
its basipterygoids are aborted. In no Owls that I have seen does the bone become so
frail a sponge as in the enormous lacrymals of that type; the ecto-ethmoids are in it
very minute flaps, the outer part of which is not ossified, as in Caprimulgus. Hurystomus
is a Roller, making no inconsiderable approaches to the Owl-like nocturnal Cuculines
next to be noticed.

The first of these is the Guacharo, or “ Oil-bird” (Steatornis caripensis, Humb. ;
Podargus, Cuv., Temm.).

The high arch both of beak and skull makes this bird’s head most completely iso-
morphic of that of the Owl; its basipterygoids are well developed (of enormous breadth,
the articular facet being oblong rather than oval), thus agreeing with the Ow/ and Fern-

'* The figures and descriptions of these forms, and of many others referred to in this paper, will soon appear in
the ‘ Zoological Transactions.’

SECOND SERIES.—ZOOLOGY, VOL. I. s

124 MR. W. K. PARKER ON THE STRUCTURE

Owl at once, and departing from Podargus ; Steatornis also agrees with Caprimulgus in
having the proximal part of the large flat antorbital (ecto-ethmoid) ossified. I mention
these things here to distinguish this type from the next to be described; a full account
of it must be reserved for another time and place.

On the Structure of the Face in ine Giant Goatsucker (Podargas).

Having been led up to Podargus by aseries of marginal and of central Cuculine forms,
J am anxious in this paper to give the most intensely modified form of face that has as
yet come under my notice. A face and skull are present here which, like that of the
equally specialized skull of the Crocodile, give us a prolepsis of that of the Mammal,
with its hard palate, the last degree of desmognathism having been attained.

This apparently ancient type, perhaps an early tertiary or even secondary waif, is also,
like the Oil-bird, an isomorph of the Owl, but not to the same degree. The huge upper
face, mimetic of the succulent leaf of an Aloe or a Mesembryanthemum, is mainly com-
posed of the premaxillaries and maxillaries (Plate XXIII. fig. 6). The figures already
given of the small and somewhat Passerine Fern-Owl (Plate XXI. figs. 8-11 and
Plate XXII. fig. 1) will help the student to understand the development of this thick,
arched, scooped, and although apparently solid, yet in reality spongy face. From the
terminal hook of the neb to the frontal region above, and to the truncated palatines below,
there is no trace of a suture. To make the division below the fore face on the one hand,
and the hind face and skull on the other, more complete, the Jugal process of the maxil-
lary is absorbed, and the jugo-quadrato-jugal bar articulates with the maxillary imside
its dentary angle. Here, then, the only movement possible is at the cranio-facial
hinge; for the pterygoids overlap and are tightly strapped to the palatines. The great
main hinge is behind, namely, that of the quadrate upon the auditory wall (fis. 7, q).
The motion here is almost as limited as in the claw-hinge of a Lobster.

A rudiment of the basipterygoid is seen as a ridge, somewhat pointed in its middle,
which runs from the optic foramen below, and is soon indistinct. Between these ridges
the basifacial bar is subcarinate, and then flattens out, running into short bony wings,
which are the proximal parts of the ecto-ethmoids; these are squarish flat cartilages
(Plate XXIII. fig. 7, pp). The septum nasi (fig. 7) is flat, bony, and notched behind,
below the groove for the nasal nerve; the hinge between the septum and the meso-
ethmoid (fig. 7, pe) is perfect. ‘The fore part of the median ethmoid is slightly keeled
between the antorbitals; its line is forwards and upwards equally. The ale nasi and
all the turbinals are unossified, and are extremely small and feeble; for the descending
crus of each nasal, and the face-plate of the maxillaries, straighten the nasal cavities into
two widish tubes. The posterior trabecular splint, the parasphenoid, does not project in
front, and is very short; the great anterior splints (premaxillaries) have few rivals in
relative size. With the nasal labyrinth so aborted the vomerine bones were likely to be
feeble. A flat band of fibrous tissue runs froma very soft bar of bone in front (v) to the
ethmo-palatine hooks behind (fig. 9, epa). Below, in front, we see this spongy wedge ;
it is higher at its smooth lower face than the great maxillo-palatine plate (map) to which
it is ankylosed. Above, in the end of the membranous vomerine band, there is a small

AND DEVELOPMENT OF THE BIRD’S SKULL. 125

ossicle, which lies between the ethmo-palatine hooks (fig. 9, v*, epa); this is a minute
posterior vomer, this element being double, as in Scythrops and other “ Cuculine.”

Caprimulgus is very passerine in having a huge double (symmetrical) vomer (Plate
XXII. fig. 1, v); but Steatornis comes much nearer Podargus, and has an azygous vomer,
of a clubbed shape in front, articulating with the palatine commissure. But the flat
anterior foot of the Chelonian vomer is again, as in Podargus, represented by a small
bony wedge in the cleft behind the maxillo-palatine synostosis.

The elegant leafy palatine arch is as fraught with interest as any other part of this
bird. The long prepalatine bar is aborted (compare Plate XXI. fig. 8, prpa, with
Plate XXIII. fig. 6, prpa); the broad truncated ends of these, the shortest of ornithic
palatines, are joined to the great fore jaw by a half-ankylosed dentate suture. For
short distance they are distinct, but they soon converge; these interpalatine plates are
ankylosed together, and form a bony raphe below to this superaddition to the bird’s
hard palate. This line of union is half the length of the palatines; a rounded notch
separates the extreme ends of the bars. The transpalatine process is triangular, with a
rounded point; it is not one third the relative size of this part in Caprimulgus; in
Steatornis it is entirely gone, as in many Cuculines. In that intermediate Goat-
sucker the interpalatine processes are rounded lobes, that grow near to each other in a
horizontal manner, but do not meet; whilst in Caprimulgus (Plate XXI. fig. 8, ipa) the
interpalatine ridges run parallel a long way, a short distance apart, and end in a delicate
spur. In Steatornis the ethmo-palatal commissure shows no medio-palatal keystone as
in Caprimulgus (Plate XXI. fig. 9, mpa); but in Podargus, although this tract is very
short, only half the extent of the lower hard floor, yet here, again, we have the two
azygous medio-palatines, the undoubted symmorphs of those I have described in the
Fern-Owl (Plate XXI. fig. 9, mpa, and Plate XXIII. fig. 8, mpa). In Steatornis the
palatine commissure is longer than in either the Giant or the Common Goatsucker, more
like that of Scythrops (Plate XXIII. fig. 1, ipa); the posterior notch is similarly small.

In all these thin types the apex of the palatine arch, or epipterygoid, is aborted, and in
Podargus and Caprimulgus this bone is very spongy (Plate XXIII. figs. 6 & 7, pg,
and Plate XXI. fig. 8, pg); in Steatornis it is less spongy and more crested. In this
latter bird the facet for the basipterygoid has an overlapping process, by which it lies
over as well as against the basal facet; in Caprimulgus there is a slight rudiment
of this structure; in Podargus there is no joint at this part. All these three types
agree in being embryonic, as regards the mesopterygoid. This spur does not segment
off in either; in the old Fern-Owl it has become aborted; the young agrees with
Steatornis and Podargus (Plate XXI. fig. 11, and Plate XXIII. fig. 7, pg); it lies ina
depression and is strongly bound to the supero-median surface of the palatine. I find
no defined os uncinatum in either the Fern-Owl or Podargus; but in Steatornis it is as
well developed as in Scythrops (Plate XXIII. fig. 1, ow), and has two cartilaginous tapes
that tie it in front and behind.

The lacrymal is large in the Fern-Owl (Plate XXI. fig. 8, 7); in Podargus it seems to
be absent; but a small ossicle may have coalesced with the supero-external angle of the
nasals, in front of the narrow frontals, as is the case with a larger piece in Steatornis.

$2

126 MR. W. K. PARKER ON THE STRUCTURE

The face of Steatornis is much like that of the Eared Owl (Asio otws), but the ale nasi are
much more ossified; they are quite soft in the other two, and ouly the fore edge is left
cartilaginous ; its septum nasi is also bony.

This agreement and disagreement, alternately, of these three Goatsuckers suggests the
probability that they each represent what were once large “ subfamilies;” but their
characters, by their intense inosculation, show that they belong to one highly natural
group.

The stapes of Podargus presents a very curious and instructive example of this hyootic
structure. The otic portion, or basal plug (figs. 10 & 11, st), and the medio-stapedial bar
(mst) are quite normal. The “suprastapedial”’ (ss¢) is large and unossified, and so
also is the infrastapedial (ifst), which ends below, continuously, in a spatulate recurved
stylo-hyal. The thing, however, to be noticed is the huge roughly spoon-shaped extra-
stapedial (est), all bony, convex outside, concavo-carinate within, and pneumatic; the
upper margin is evenly sinuous, the lower and the end sublobate; there is a fenestra
between it and the falcate suprastapedial (fig. 11). A ligament is seen growing from
the top of the extrastapedial spoon; and another larger ligament ties the supra-
stapedial to the prootic wall. The cartilage of the supra- and infrastapedial is con-
tinuous; the line of the original arch or bar is from the mediostapedial to the infra-
stapedial, whilst the suprastapedial becomes a second head; the extrastapedial is a
conjugational outgrowth. The bifurcation of the rod above produces processes which
are the isomorphs of the capitular and tubercular processes of a true rib; the supra-
stapedial resembles a tuberculum, and the mediostapedial a capitulum; the shaft, as we
saw in the Martin, chondrifies in patches, the backwardly bent stylo-hyal reuniting
with the upper piece by a later chondrification of the infrastapedial tract.

On the Structure of the Face in some other “‘ Coccygomorphe.”

In the Hornbills, the secondary growths of the trabecular and palatine arches, more
especially the former, acquire the uttermost degree of prognathic development consistent
with the mechanism of a bird’s skeleton as such. The result, taking in the correlated
modifications throughout the rest of the skeleton of these birds, is a form which makes the
conception of the living Péerodactyle an easy effort of the fancy. Pneumatic overgrowths,
of such a size as we see in the larger species, intensify the grotesqueness of the vreature ;
for the aneurismal bursting of the bony walls by the imprisoned breath has resulted in
their huge air-chambers, walled in merely by the horny epiderm and its filmy quick.
For a figure of the palate of Buceros bicornis the reader is referred to Professor Huxley’s
paper (op. cit. p. 446, fig. 28). Here the bony bar between the apertures marked a is the
foot of the Chelonian vomer, separated from its posterior portion by a membranous
space, and ankylosed to its surroundings, namely, the maxillo-palatines and the coalesced
interpalatines, behind; for these larger species have a palatinal hard palate. The styloid
posterior part is ankylosed with the palatines at their upper or ¢7we commissure. I see a
similar structure in B. ruficollis, Temm.; but the interpalatines are imperfect, and the
form of the vomerine foot is not so easily traceable. In this species the notch which
forms the cranio-facial hinge is reconverted into a fenestra by a tuberous growth of bone,

AND DEVELOPMENT OF THE BIRD’S SKULL. 127

a median septo-maxillary, which runs from the parasphenoidal beak to the bony septum
nasi, above the posterior vomer. The os uncinatum is not distinct in these Hornbills,
but the ecto-ethmoid terminates in a large pedate process which binds upon the zygoma ;
their lacrymal is aborted, and the large lateral ethmoid comes to the outside and top of
the face, as in Megalema and the Passerinee. With the complete ankylosis of the pala-
tines at their commissure, nothing can be learned from the adult as to the existence of a
medio-palatine keystone.

In the most exquisite of all vertebrate skulls, that of the Toucan (Ramphastos toco),
the structure largely agrees with the “ Buceride.”” The ecto-ethmoid projects as in them
and the Passerine, and the os uncinatum is represented by the pedate angle of that bony
plate. I see a small lozenge-shaped subcarinate vomer below the level of, but acting as
a secondary keystone to, the palatine commissure; this, however, is only the posterior
vomer. The anterior piece is a broad bifid ragged wedge of bone, with its narrow angular
end dovetailed in between the palatines, immediately above the small spiny semidistinct
interpalatines; its position is transversely vertical, and its size ten times that of the
hinder piece. The notch is complete that forms the wide hinge between this marvellous
face and skull; it is 14 inch across, and the beak of the parasphenoid is but little behind
a descending plate of bone, the sharply bifid middle portion of which is the posterior
angle of the nasal septum. The anterior vomer reaches upwards to these three bony
spikes by its ragged split upper end. The nostrils are nearly half an inch apart on the
upper surface, close in front of the hinge, and overshadowed by the rounded posterior
margin of the elevated beak. The thoroughly ossified ale nasi enclose a transversely
oval nostril; the other parts of the nasal labyrinth are all ossified, and the inferior
turbinals are exquisite little scrolls of scabrous bone, almost pisiform, and vertically
placed, the end of the scroll being seen from delow instead of in front as in other birds.
Between the splintery prepalatines and the maxillary synostosis there is a rather large
oval space; behind, the palatines are fused together.

The transpalatine region, with its retral spike, is something between what we see in
Megalema (Plate XXIII. fig. 4) and in Dacelo (Huxley, op. cit. p. 447, fig. 29, Pl). The
basipterygoids are small rudiments; the pterygoids are spongy, and have a small epi-
pterygoid lobe. This short ¢itial description may help the student to decipher the
morphology of this culmination of the “ Coceygomorphx.” TI have just referred to Pro-
fessor Huxley’s good woodcut ofthe palate of Dacelo ; in my specimens of the “ Alcedinze,”’
namely, Alcedo ispida, Upupa epops, Todus viridis, I see no vomer. These skulls are
thoroughly ‘‘ Desmognathous,” and are especially interesting as to their palatines, in
which the transpalatine sends backwards a spike, equal, in Alcedo, to the prepalatine
style in front ; in the young Hoopoe the transpalatine is obliquely set on to the palatine
as a distinct bone.

Inthe “ Musophagidee ” the os uncinatum attains its fullest development as an anterior
transverse bone (see Dr. Reinhardt’s paper, already quoted). In a species of Corythata
(2? buffoni) there are two medio-palatines and a posterior vomer; these lie behind the
short sutural commissure, in fibrous tissue ; for, posteriorly, the rounded palatines are
‘wide apart, exposing the parasphenoid.

128 MR. W. K. PARKER ON THE STRUCTURE

The foremost medio-palatine is a minute style, having the equally minute styliform
vomer beneath it; the second medio-palatine is a mere grain of bone *.

The nasal labyrinth is well developed in these birds, and is thoroughly ossified ; so that
the bony skull of no other bird is so useful to the student who would master this elegant
sense-organ.

On the Structure of the Skull in Birds of Prey (« Aetomorphe,” Huxley).

Both for this and for the Goose tribe I have sought to extend the lines that bound in
their “lot,” confident that in the Southern World we have extant, and, toa much greater
degree, in the buried Tertiary world there are extinct, many a type that cannot and could
not wear the ordinary ornithological harness. ‘The characters of an Hagle, a Falcon, or
an Owl may easily be set down; but when the scientific formalist comes athwart a bird
in which all these are shaken up together, like the lots in the helmet of Agamemnon,
then every thing breaks down, and his wire-drawn distinctions lose their value.

Yet these distinctions are not the less valuable, being applicable to the newest and most
specialized forms; but they melt away in certain types, just as the professional distinctions
of civilized life are seen to melt away when we contemplate the life and polity of
savage races.

I make these remarks in defence of the position in which it is here sought to place a
bird, the Cariama, the relationships of which seem to me to be self-evident, and which
has been looked at by most ornithologists through the coloured and refracting glasses of
their own arbitrary systems.

Having no misgivings as to the essentially Raptorial nature of the Cariama (Dicholo-
phus), I do the more boldly place it for comparison side by side with the two most
intensely specialized types in the whole group, viz. the Falcon and the Owl. Such a
collocation, however painful to the eyes of full-grown systematists, will, if its natural-
ness be proved, be of real value to the tentative worker, serving at least as a hint that
the strength of Nature will burst the new ropes we seek to bind her with, those bonds
being, indeed, as weak “as the tow when it smelleth the fire.”

On the Structure of the Skull and Face in Dicholophus cristatus.

The form of the whole head in this bird is Vulturine (Plate XXIV. fig. 2), as may be
seen at once if it be compared with that of Gyps fulvus (fig. 1) t-

* It is not always possible to be absolutely certain as to whether any given grain of bone belongs to the hinder
part of the yomerine region or to the palatine commissure.

+ How the eyes of naturalists have been holden against seeing the truth here I cannot understand ; the legs of the
bird, so thin and long, have robbed it of its aquiline glory. A few years since, showing two of these birds in the
gardens of the Zoological Society to my brother, a man of like tastes with myself, he at once recognized the Cariama
as a kind of Hawk, whilst it crouched upon the perch. Another individual was standing in the enclosure in
front of the aviary: “ What bird is that ?” I immediately asked. ‘I cannot say,” was the answer.

Capt. Burton (‘ Exploration of the Highlands of the Brazil, London, 1869, vol. i. p. 57) compares this bird with
the Secretary (Gypogeranus), and says that it has the same serpent-eating tastes (see also vol. ii. p. 18).

The keeper of the ayiaries where the Cariamas are kept informed me that they eat mice and sparrows, beating
them about before they kill them. On July 31st, 1872, I watched both D. cristatus and D. burmeistert dust them-

AND DEVELOPMENT OF THE BIRD’S SKULL. 129

The first and most notable difference between the skull of the Cariama and that of
Gyps fulous is that it is generally feebler (Plate XXIV. figs. 1& 2: fig. 1, Gyps; fig. 2,
Dicholophus), and its ossification is arrested; but it has the elongated vulturine face ;
whilst Gypogeranus comes close to the Harriers, Kites, and Buzzards.

The first and leading characteristic of the skull in this type is its palate, which cor-
responds to that of a young Falcon (Plate XXIV. figs. 3 & 4), and gives me that modi-
fication of the Desmognathous skull which I have termed (supra, p. 112) “ imperfectly
direct,”” the maxillo-palatine plates (mxp) being united by harmony-suture and not by
coalescence.

How this skull has been taken for one of the “ Schizognathous”’ type, and put with
that of Cranes and Bustards, I cannot understand. In respect of its feebly ossified nasal
labyrinth, this form approaches and agrees with other South-American types, namely

e “ Cathartide ;” and the skull of Sarcoramphus has greatly assisted me in working
this kind out; but there is no kind of Raptorial skull that it is not, more or less, in
harmony with, in one point or another; and the whole group of the Aetomorphe may
in some long past epoch have lain in the loins of a bird but little different from the
Cariama.

There are three main varieties in the general bony characters of the Raptores : in the
Cariama the skeleton has a fibrous lightness and delicacy, in the Owls a soft spongy
condition for the most part, the long bones being dense as to their walls; but these are
of extreme tenuity. In the Falcons the bones, everywhere, are almost like ivory in their
density, reminding one of those of Serpents.

The skeleton of the Cariama has very little in common with that of the Crane tribe ;
it is as delicate, relatively, as that of that lonely waif of a subfamily of the “ Gruine,”
the Sun-Bittern (Hurypyga helias), and still more like that of another subextinct type
of ancient and generalized Cranes, namely the Kagu (Rhinochetus). But the Crane
with which one would naturally compare it is the Trumpeter (Psophia); nothing, how-
ever, can be more fatal to the view of its being a Gruine bird than such a comparison :
I would rather make room for it amongst the Gallinaceous birds than see it so mis-
placed. In the genealogical life-tree of the families of Birds, some grow near to each
other, and have, as it were, their branches forming an acute angle with their leader, as
in the Lombardy poplar; whilst others, as in the black poplar, grow out at all angles,
most of which are wide and divergent.

If we take the ‘“‘ Coracomorphe,” they do so cling to each other, diverging very
gently; but the “ Aetomorphe” delight in wide and wild divergence; and although
manifestly ‘“‘ of one blood,” they yet have acquired the greatest variety in their speciali-
zations; but the Falcon is “ brother to the Owl.”

Notwithstanding the enfeebled length of the beak of the Cariama, the bones are
thoroughly fused together. The broad basitemporals (fig. 3, 0¢) end in front, as in the

selves by rolling exactly like the Common Fowl: this is in harmony with the fact that they impinge very closely
on the gallinaceous circle. They are Raptores with a very gallinaceous aspect; whilst the vulterine Guinea-hen and
the Brush-Turkey are Fowls with a very vulturine aspect.

130 MR. W. K. PARKER ON THE STRUCTURE

Owl (fig. 5), in two pouting lips; they underlie the common Eustachian opening (ew);
outside and in front of this emargination the basipterygoids (pg) are seen projecting
from those wings of the parasphenoid that form the “ anterior tympanic recess.” The
narrowed part of the parasphenoid (pas) is relatively slender, and the pterygo-palatine
bones undergird it for most of its extent. The great anterior palatine interspace is large,
but it is lessened in the dry skull by an ossification which has affected the retral part of
the trabeculee and al nasi (fig. 3, alz). A similar bony bridge is formed in Sarco-
ramphus papa by the maxillo-palatines where they join the nasal septum.

Most of the nasal capsule, as in pluvialine and gallinaceous birds, is unossified. The
vomer (fig. 3, v) is of a type rare in birds; it is thoroughly Chelonian, more so than in
any other bird save the Falcon (fig. 4, 7), although some approach is made to this in the
Sacred [bis and the Tree-Duck (Dendrocygna).

As in the other Raptores, it is truly azygous, so also it is in the Gallinacez; but in the
Cranes and all pluvialine birds it is composed of long lathy pieces, that converge and
ankylose, only keeping apart behind.

In the Chelonia the azygous knife-shaped vomer spreads out behind and above to
articulate with the orbital plates of the palatines, whilst in front it descends and helps
the maxillaries to form an anterior hard palate. Just so does this bone behave in the
Falcon and Cariama. In the Green Turtle (Chelone midas), however, the palatines have
a small palatine lamina joining this fore foot of the vomer; this part is arrested in the
Cariama and the Falcon (figs. 3 & 4, prpa’). In a side view the vomer of the Cariama
(fig. 3%, v) is seen to be very high in the middle, low and flat in front, and gradually to
lessen behind; the middle part is deficient of bone for a large space. This vomerine
fenestra is repeated in the large Falcons (Plate XXV. fig. 12). The articulation of
the vomer with the ascending or orbital plates of the palatines is through the medium
of the keystone-piece of that arch, the medio-palatine (mpa). This bone is of great
interest; hitherto I have only found it in Birds, and amongst them most developed in
Woodpeckers (Trans. Linn. Soc. ser. 2, Plate II. p.9). But it has just been shown to
exist very commonly in Cuculine types.

There is no abrupt line of demarkation between this ossicle and the paired meso-
pterygoids; and all the gradations are seen in the Raptores, where these intercalary
pieces are very variously developed. In the Cariama it is a roughish wedge of bone,
rounded behind, pointed in front, and wedged in between the right and left ethmo-
palatines laterally ; vertically is jammed in between the parasphenoid and vomer. There
is one ossicle here in the young of Falco tinnunculus (fig. 4, mspg); but this is the
apex of a right mesopterygoid, the left being suppressed. In Sarcoramphus (fig. 19)
it is more truly azygous; in Neophron (Plate XXV. figs. 14.&15) it is a notable right
mesopterygoid; but in that bird the mesopterygoid appears at both ends of its region
(fig. 16), a little grain of bone occurring at the junction of the pterygoid on the
same side.

In Helotarsus ecaudatus there is one small bone (a right mesopterygoid) in front
(Plate XXV. fig. 9, mspg); but certain Owls (Strix stridula, Plate XXV. fig. 20, and

AND DEVELOPMENT OF THE BIRD’S SKULL. 131

Ketupa ceylonensis, Plate XXV. fig. 11) come the nearest to Dicholophus in respect of
this bone and its relation to the vomer; but in these birds the vomer has no fore foot *.

The prepalatine bars (prpa) are long and slender; the transpalatine is rounded off
(¢pa), and passes into a short postpalatine (ptpa), on the inner margin of which the sub-
mesial muscular ridge ends ; the whole form and manner of these parts is Falconine, but
drawn out. The upper part, or ethmo-palatine (epa), meets its fellow process for some
extent in the middle, the fore end diverging for the medio-palatine keystone {mpa), and
the hinder for articulation with the diverging pterygoids (pg). Threads of bone represent
the interpalatine (¢pa), which, if developed, would have given a desmognathous palate,
bound together in the same manner, and composed of the same parts, as that of the
Green Turtle, which has the anterior vomerine foot inwedged between rudimentary
palatine plates of the maxillaries and palatines. This Reptile stands halfway between
the Lizard, with its highly schizognathous face, and the Crocodile, in which desmognathism
is carried to its uttermost, as in the Mammalian genus Wyrmecophaga. Walf the
charm of this research would vanish if we could possibly forget that the bird groups are
luxuriant outgrowths of the reptilian understock, and that their fiery nature once lay hid
in the stone-cold Reptile.

Some of the Raptorial characters of the Cariama are exaggerated. The shortness of
the pterygoids (pg) is greater than in the Owl and Falcon; they answer, however,
exactly in character to those of the latter, having no articular junction with the basi-
pterygoid processes (dpg), these being arrested. Their epipterygoid process (epg) is a
hooked flange. The splints of the face are confluent, the marginal line being continuous
from the neb to the quadrate hinge; the hinder jugal bar is slender. About two fifths
of the premaxillary mass is continuous in the fore palate; the palatine processes of the
premaxillaries are long, slender, and lie outside the prepalatine laths. At the middle
these large bones are again united by a commissure, namely, the alinasal bridge (adn).

But the most remarkable part of the face is the maxillo-palatine floor, which is
relatively larger than that of almost any bird; the processes forming this hard palate
are flat ear-shaped lobes, which unite at the mid line by harmony. They are pointed
behind to let in the foot of the vomer, and broad and notched in front, behind the large
anterior palatine space. These plates have the thin fibrous character of the main part of
the skeleton, instead of being dense as in the Falcons, or spongy as in the Owls. The
skulls of those types when young (figs. 4 & 5) are exactly correspondent in their essential
morphology to that of the Cariama, being of the “ imperfectly direct’ desmognathous
type.

The general form of the skull seen from above, with its alate lacrymals, is evidently
an enfeeblement of the vulturine type (fig. 2) as compared with that of Gyps fulvus
(fig. 1); and what arrest and feebleness there is belongs not to the Crane, with its dense
heavy skull and square-browed lacrymals, but is something intermediate between that

* T have the more carefully gone over these details because of the instructive fact that the structure of the Cariama
is best explained by that of the various Rapiorial subfamilies, and is not illustrated by members of the family to
which it has been supposed to belong. My notes on the soft parts of this bird, and the observations and comparisons
I have made of the rest of its skeleton, all tell the same tale.

SECOND SERIES.— ZOOLOGY, VOL. I. T

1382 MR. W. K. PARKER ON THE STRUCTURE

of the Fowl and the common Vulture. In the Crane, a truly pluvialine bird, the maxillo-
palatines project but little mesiad of the prepalatine bar, the huge almost struthious
double vomer being wholly displayed in the upturned palate.

Structurally, then, the Cariama is seen to have the remotest relation to the Pluvialine
type; and if the details just given be weighed and measured, they will show that its
Raptorial characters vastly preponderate ; that it is evidently a branch of the same stock
as the true Gallinacez, especially the Megapods and Curassows, and that it does not
belong to any extant type of Raptorial bird; but as the Palamedea is a Chenomorph, so
is the Cariama an Aetomorph, not holding out the flag of any one party of that common-
wealth, but being, as it were, a representative of the old fathers of the race, whose sons
have acquired such sharp characters of dissimilarity. It contains in itself not merely
the attributes of a single subfamily, as Hawk, Falcon, Owl, Vulture, but “all these in
their pregnant causes mixed.”

There is a bone on each side in the face of the Cariama which is not seen well in the
palatine view; this is the os uncinatum (Plate XXIV. fig. 2a, ow); it is well seen
in the side view.

As I have shown in the former part of this paper, this is common in the Cuculines ; in
Dicholophus it is similar to but larger than its counterpart in the Gulls (Plate XX VII.
fig. 11, ow). It has a like development in the Alabatross, and is abortively seen in Uria
troile and Alcatorda. I find it inno other Raptorial bird, nor in any of the Grallee that
lie on the same morphological level as the Cariama, and although not belonging to its
group have yet some affinity with it.

Here the bone is a notable style perched upon the zygoma, and underpropping the
lacrymal (/); it is, however, connected with the feebly developed pars plana (pp), the
free tongue-shaped part of which is unossified as in the pluvialine types.

This is the ornithic representative of the cartilaginous bar which yokes the trabecula
to the palatine in the Frog, and therefore of great interest morphologically.

On the Structure of the Skull and Face in the Falcon (Falco tinnunculus).

Having treasured up from time to time every obtainable stage of young birds, I am
able to compare the adult Cariama’s skuli with the immature skull (that of fledglings)
of the Falcon, the Hawk, and the Owl, the proper morphological counterparts of the
skull of such a root-form as the one just described *.

I trust to the trained eye of the ornithologist to see at once the essential agreement of
this Falconine palate (fig. 4) with that of the Cariama (fig. 3). Nature, anti-Procrustean
in all her works, has developed one skull into a form which is feline in its shortness,
massiveness, and strength, whilst the other is outdrawn, and, as it were, of a vulpine
shape ; yet the “habit” of both types is manifestly one. The difference depicted is not
the full measure of what would have been seen if an adult Kestrel had been selected for
comparison.

* The same method with regard to the relation of the Plovers to the Gulls has appeared to be fruitful to my

accomplished fellow workers, Professors Huxley and Newton; and their coinciding judgment has been to me in no
little degree determinative of renewed labour in this field.

AND DEVELOPMENT OF THE BIRD’S SKULL. 133

The transverse basitemporals (fig. 4, b¢) have joined their wings beneath the common
Eustachian opening (ew). Above this point the parasphenoid (pas) is seen with small
basipterygoid knobs (dpg), and above these and the basitemporals expands on each side
into a large trumpet of bone, whose mouth opens into the labyrinthic drum-cavity.
These trumpets form the “anterior tympanic recesses,” and are lipped by one or more
additional (tympanic) ossifications.

The fore part of the parasphenoid has the shape of a planter’s dibble, and ends, blunt-
pointed, above the palatine commissure, carrying the hinge mesethmoid on its grooved
upper face. The premaxillaries form a triangle whose sides are but little longer than
their (ideal) base from side to side, and after forming themselves separately on the
spatulate prenasal cartilage, combine at the mid line, not only above but also below,
aborting and taking the place of their model, growing even there much more solid than
their displaced prototype.

This is the most Chelonian of all the birds’ skulls examined by me; and knowing full
well that the bird’s skull, after it has fused its trabecule and acquired a cartilaginous
consistency, takes on immediately the most unmistakable Rhynchosaurian characters, I
am in nowise surprised to see the Faleon (which shoots upward so high above the
Chelonian) in its ascent passing close to and retaining much of the likeness of that
less specialized and much less metamorphosed type. ‘That the study of the modifications
produced by metamorphosis of the primordial face is of the utmost importance to
taxonomists is evident; and the vomer alone is, in this respect, worth more than all
Mr. Garrod’s muscles put together. In the Reptilian group the Chelonia are alone in
possessing an azygous vomer; and only a minority of the bird class have the same
character. Here, in the Falcon, this bone is not shaped like the proper ploughshare,
but like that less common implement the breast-plough, for sod-turning. The blade of
this bone is roundly three-toothed, its shaft flat vertically, down-bent in front and lifted
behind, and the handle is thick and forked. The blade is wedged behind in the maxillo-
palatine harmony, and between the ear-shaped maxillo-palatine lobes. The shaft divides
the median nares, right and left; the two-toothed handle bites the palatine commissural
plates ; it is fastened there by a wedge of bone, which has been driven in, as it were, on
the right side. This wedge is the right mesopterygoid (mspg). In this type I have
found no other vomerine bones.

The pterygoids (fig. 4, pg) are even less specialized at this stage than those of the
Cariama; they are larger relatively, have a less distinct auricular process (epg), and
their abortively developed apex only appears on the right side as the mesopterygoid wedge
(mspq). |

The only difference between the palatines and those of Dicholophus is their greater
shortness; also whilst the inner ridges end in the Falcon as a short interpalatine spur
(ipa), those of the other bird are ragged and formless. In both cases the transpalatine
region (tp) is evenly rounded off, but the outer lobe is scooped below for the palatine
muscles. In the jugum of the Cariama the elements once combined ; but I suppose these
were then the jugal process of the maxillary, the jugal itself, and the quadrato-jugal.
These two latter are not separate, as far as I can see, in the Falcon, which thus

T 2

134 MR. W. K. PARKER ON THE STRUCTURE

agrees with a large number of the “ Aves aérize,”” whereas in Hawks and Owls they are
both present. But the most striking part of their palatine outworks is the huge
maxillary (ma); it is, relatively to the premaxillary, as extensively developed as in the
Chelonia, although the dentary process of the latter bone overlaps it as in osseous Fishes,
and, indeed, in birds generally.

These parts are builded together in a manner which would please the eye of an
architect.

But the whole meaning of these parts can only be seen by the study of sections made
in young skulls. In a nestling younger than the one from which the palatal view was
taken, I have been able to decipher all those parts of the face in which the nasal capsules
have their furniture disposed amongst and within the foremost facial bars.

In this type the nasal labyrinth enfolds itself and encloses its passages with solid car-
tilage to a very unusual extent. Here (Plate X XV. fig. 1, em) the external nasal opening
is not a long slit, but a neat round opening; the rim is wrought, like the rim of a
cup, and is almost closed by a finger-shaped projection from the alinasal turbinal (a7).
This process looks outwards and backwards, and only leaves around it an oval chink.
The whole alinasal region (figs. 1-8, alm) is formed into a pair of oblong oval pouches,
like two carpels of an apocarpous fruit dehiscing; seen from below, there seem to he
two cells left out of four; and the still persistent lanceolate prenasal cartilage (pz)
running from the base of the septum (sz) carries out still further the similitude, it being
in the likeness of the fruit-axis. Behind, at this most bulbous part, these nasal pouches
are complete tubes, the floor having coalesced with the middle wall (fig. 3, adn, sm); this
is shown in section (fig. 5), also the apposition of those parts further forwards (fig. 6).

When the septum is removed, to show the inner face of the turbinals, it is seen that
the alinasal turbinal (fig. 2, a¢b) is a large leaf of cartilage, hollow outside and bulging
within (see also sections, figs. 5 & 6). This leaf is attached to the outer wall by its
upper edge, and is also confluent behind with the bulging floor, where that part rests on
the nasal nerves (x). Over the hinder third the true “inferior turbinal” arises; this
is at first a two-winged crest (fig. 5), but further backwards acquires the true type of the
organ, as seen in the “Carinate” generally; it has nearly three coils (see fig. 2, ib).
The middle turbinal is merely represented by an irregular ridge on the fore face of the
antorbital plate, and the upper turbinal by the coiling, on itself, of the aliethmoidal
plates, exactly as in the Common Fowl (Phil. Trans.).

- The septum nasi is connected by an uncleft isthmus of cartilage (fig. 4, sv) with the
perpendicular ethmoid; it is unusually deep and large for a bird (figs. 4-6). There is
one septal fenestra (sf) in addition to the “notch” between the septum and the ethmoid ;
this window is situate close behind the insertion of the alinasal bag. The hinder inser-
tion of that bag is much thickened, and at this part the nasal branch of the orbito-nasal
nerve (fig. 4, nm) bores its way through the cartilage, piercing the alinasal, and bur-
rowing in the septal cartilages; it soon gets below the septum (see figs. 2, 4, 5). Now
if these sections be compared with the palatal view (Plate XXIV. fig. 4), and these by the
reader with an adult skull before him, it will be easy to comprehend the compact and
complex desmognathism of this type, a type having the most highly ossified and, in many

AND DEVELOPMENT OF THE BIRD’S SKULL. 135

respects, the most specialized skull to be seen amongst the Raptores. The broad foot of
the vomer (Plate XXV. fig. 5, v) fits like a keystone between the large maxillo-palatine
slabs (map); and these are bound, externally, by the dentary plate and angle of the pre-
maxillaries (dpx). In front of the vomer (Plate XXIV. fig. 4) the maxillo-palatine plates
meet, and afterwards coalesce. Ultimately, when all these structures, without and
within, have coalesced by extended ossification, all but the vomerine undersetter, then
the lateral halves of the face are indissolubly bound together, and form a structure most
compact and strong.

On the Structure of the Skull in the Sparrow-Hawk (Accipiter nisus)*.

In greater feebleness of general structure the skull of this type is intermediate between
that of the Falcon and the Cariama; but the leanings of the Hawk are not so much
towards the latter as in the direction of that gigantic Vultwrine Harrier the Secretary
_ Bird (Gypogeranus). The skull of a partially fledged nestling yielded the results given
in Plate XXV. figs. 7&8. Here we see at once things that agree better with the skull
of the Cariama, and also things that are less in conformity with it than those just
described in the Falcon: whilst the Cariama agrees with the Falcons, Hagles, and the
more Aquiline types of Vultures in having the basipterygoids early arrested, the
Sparrow-Hawk shows them as strong short spurs in the young (Plate XXV. fig. 7, bpg),
and retains them as mammillary elevations in the adult. They are still more developéd
in the Harrier (Circus), more still in the Brahmany Kite (Haliastur indus), and perfectly
functional in the Secretary. In Accipiter the parasphenoid (pas) is stout, and bevelled
on each side by the under-gliding pterygo-palatines. The next median splint, the vomer
(v), is truly azygous, and is a long knife-blade at present; in the young it is neither bi-
dentate behind nor pedate in front; it alters but little afterwards, and agrees in this
respect with the whole Accipitrine division of the group.

The desmognathism is perfect, although, in the young bird, it is “ imperfectly direct,”
as in the adult Cariama. Already the septum nasi is fast fusing with the solid fore end
of the premaxillaries; and, behind, its nerve-bridges and connexion with the ale nasi
make it a fit wedge to fill up the space in front between the large lunate maxillo-
palatines (mxp), which, lying back to back, of necessity leave an open space in front for
the septum, and behind for the vomer. The premaxillaries form a less solid mass than
in the Falcon ; but this type is much more orthognathous than the Cariama. Whilst the
Hawk agrees with the gallinaceous types in having a truly azygous vomer, it differs from
them, and is in harmony with the huge groups of the Passerines and Pluvialines, in having
a large mesopterygoid wedge differentiated from the fore end of the proper pterygoid (figs.
7 & 8, mspg, pg). As usual, the transpalatine (¢pa) is rounded off, more developed than
in the Pluvialine types, but less than in the Passerines. The ethmo-palatine plates (epa)
are thin, curved, pointed laminz, embracing the head of the vomer; the interpalatine
ridge (ipa) has no free spur. ‘he prepalatine bars (prpa) reach as far forwards, or
nearly, as the trabecular axis of the premaxillaries ; they are thin laths of bone.

' * For further details on this type of skull, see M. Micr. Journ. 1873, plates v. & vi. pp. 45-50. In these plates.
for ep read op.

136 MR. W. K. PARKER ON THE STRUCTURE

Behind we see that the large mesopterygoids (mpg) appear at the mid line, between
the postpalatine processes (ptya). Their general shape, as sharp wedges, is seen in the
side view (fig. 8). The pterygoids (pg) differ but little from those of the Cariama, being
short, flattened out on the inside in front; whilst behind they send up a very rudi-
mentary epipterygoid plate (epg). The extent of the median part of the maxillaries, the
maxillo-palatine plates (map), is very large, equal indeed, fore and aft, to what is seen in
an ordinary mammal. They are spongy, have a somewhat lunate or, rather, ear-shaped
form, and their hinder part forms a large free lobe, curving outwards.

In addition to the jugal process of the maxillary, there are two slender styles on each
side, a jugal (j) and a quadrato-jugal (qj), and not merely one bone, as in the Falcon.

On the Structure of the Face in Buzzards, Kites, and Harriers.

Several subfamilies of the lesser rapacious birds range themselves under the Accipitrine
type as a distinct form, and inferior to the Falconine. Their cranio-facial structures
agree very closely with what is seen in the Eagles and the Aquiline Vultures.

The condition of the facial parts in a recently fledged Common Buzzard (Buteo vul-
garis) is very instructive, and is representative of what may be found in a large number
of Old-World Rapaces, excluding, of course, the culminating forms, the Falcons.

A large interorbital fenestra (Plate XXIV. fig. 9, tof) separates the ethmo-presphe-
noidal bar above from the ethmo-basisphenoidal bar below (pe, bs), and the flat roof of
the great ethmoid, covered by the frontals and nasals (f,7), is rapidly becoming bony
from the huge median centre (pe). This roof sends backwards the normally ornithic
spike over the olfactory nerve (1), which grooves the bony plate on each side. The
coiled aliethmoid (ale), the representative of the upper turbinal mass of the mammal,
is thus partly ossified.

The flat antorbital plate, or pars plana (pp), which represents the middle turbinal,
is already ossified, and has coalesced with the perpendicular plate or meso-ethmoid (pe) ;
it is roughly pentagonal in form.

But this orbito-nasal partition is a downward continuation of the ala growing from
the roof, the aliethmoid‘(ale); the two are connected by a narrowish band of carti-
lage: this band is separately ossified, forming an “upper ectethmoid” bone, as in the
Passerinze (Oorvus). It lies under the roof, however, and does not stand out as a
prefrontal mass, as in the Crow and its congeners.

- The nasal nerve runs into the nose from the orbit, inside this bar, skirting the upper
turbinal, and then sends most of its substance forwards and downwards (see dotted
line in the figure) to the base of the septum nasi.

The notch which divides the true olfactory region from the septum nasi (sz) is very
large, runs obliquely forwards and upwards, and ends a little distance from the top;
hence this cranio-facial cleft is imperfect. The septum itself, only partially ossified as
yet, is differentiated into a nasal or trabecular, and an upper or true alinasal region,
by a large, oval, internasal fenestra (inf). Below and in front of this space the septum
is ossified as one large bone (/so), whilst the upper part is soft, except at the roof,
behind; there is at that part an upper septo-nasal bone (uso). The large lower bone

AND DEVELOPMENT OF THE BIRD’S SKULL. 137

is burrowed by the nasal nerve; and above and in front of this burrowing there is a
transverse septal knob, where the inturned alinasal fold joins on to the septum. On
each side the large, spongy, pneumatic maxillo-palatines (map) flank the septum, and
have the fore end of the knife-shaped vomer (v) between them. The palatines (pa), the
mesopterygoids (mspg), and the pterygoids (pg) conform very closely to those of the
Sparrow-Hawk.

In the Harrier (Circus cyaneus) and in the Brahmany Kite (Halastur indus) I find
a separate, thin, “median septo-maxillary,” attached, vertically, to the upper edge of
the vomer. It is very possible that this state of things may be merely the appearance
of a second vomer; I am not certain on this point; parts primarily symmetrical do at
times become azygous or median by the overgrowth of one and the arrest of the other,
the stronger bone gaining the mid line, and dragging the feebler bone after it.

These feebler types have, as a rule, a flat superorbital on the free end of the long
superorbital process of the lacrymal. In the Falcon the ossification is continuous
along their frowning brow.

The reader is referred to Prof. Huxley’s “Classification of Birds” (Proc. Zool. Soc.
1867, p. 442, fig. 24) for the palatal view of the Secretary’s skull: that figure is defi-
cient ; it should show a small knife-shaped vomer, like that of a Fowl. The whole skull
of that bird comes very near to that of the Brahmany Kite (Haliastur mdus) and to
that of the Egyptian Kite (Zlanus ceruleus). Its general skeleton is hugely modified
on account of its stilted Crane-like stature; legs of any indifferency would have asked
for none of those specializations of shoulder and hip-girdle which we see in that type.

On the Palatine Bones in Eagles and Vultures.

I shall merely note a few things to be seen in these large groups of the gigantic
Raptores.

In the Bateleur Eagle (Helotarsus ecaudatus) there is a small median palatine (Plate
XXV. fig. 9, mspg) between the ethmo-palatine lamine.

In Neophron percnopterus I find a delicate knife-shaped vomer (Plate X XV. figs. 14,
15, v), the right. fork of which is evidently aborted by a small mesopterygoid (mspg);
and besides this, on the same side, there is another granule of bone, belonging to the
same region (fig. 16, mspg). At the end of vomer, thinned out, however, in front, there
is a small medio-septomaxillary (msmz).

In Gyps fulvus there is an unlooked-for amount of differentiation of the palatine arch ;
for even in old birds a sharp sutural line runs obliquely two thirds across, almost sepa-
rating the transpalatine (Plate X XV. figs. 17, 18, ¢pa) from the rest of the bar. Here also
I find a strong wedge-shaped mesopterygoid (mspg) permanently distinct.

For the skull of Cathartes aura the reader is referred to Prof. Huxley’s paper (op.
cit. p. 440, fig. 22); it is an instance of “indirect desmognathism,” the maxillo-pala-
tines not being developed more mesiad than in Plovers and Pigeons; but the bond
being formed of an ossified tract of the septum nasi and inturned alinasal fold, this

forms a bony bridge similar to that seen in the Cariama; but there it is the anterior

138 MR. W. K. PARKER ON THE STRUCTURE

part of the alinasal fold. These South-American types agree together in the very
limited ossification of the nasal sacs, so different from the Old-World Rapaces.

In the King Vulture (Sarcorhamphus papa) the palate agrees with that of its con-
gener Oathartes (Plate XXYV. fig. 19); but I find a small medio-palatine (mpa), and
also much larger and more functional basipterygoids (pg). In this bird the Eustachian
passages (ev) open out into a very patulous, triangular fossa; and all the basicranial
structures are very strongly soldered together. The basitemporals (6¢) have also a
large mammillate knob on each side for muscular attachment.

On the Structure of the Skull in the Owls (Strigidee).

The Owls are the most distinct group of the Rapaces, and in respect of their basis
cranii come very naturally after the Cathartide. They agree with the Cariama in their
digestive organs, and, what is also quite unaccountable, with the Pigeons, in having a
curious deficiency in the occipital arch, a hole, like an artificial pin-hole, being left unos-
sified, as it was formerly non-chondrified, over the great cranio-spinal opening. ‘The study
of the skull of the Owls in nestlings and young fledglings is of great interest, these ob-
servations being made before the skull has become one light, spongy, undivided mass.

In a fledgling of the Barn-Owl (Aluco flammeus) the details can be well made out
(Plate XXIV. fig. 5), and the likeness and unlikeness of the type to that of the highest,
the Falcon (fig. 4), and the lowest, the Cariama (fig. 3), is clearly seen.

The relation of the pterygoids to the basipterygoid processes (pg, dpg) is quite like what
is seen in the King Vulture; but the parasphenoid (pas) is very swollen and is early fused
with a very thick and spongy interorbital septum. Not only are the outworks of the
ear of larger extent, but the tympanic cavity has large “anterior recesses” and great
upper chambers, corresponding to the open gallery that connects the right and left
chamber in the Crocodiles. The Barn-Owl has a chain of small tympanic ossicles, six
on each side, running along the inferior surface of the junction of the “lower otic pro-
cess”’ of the quadrate and the wing of the prootic, to which it is articulated.

This lower otic process is as large relatively, and as divergent inwards from the
“upper otic process,” as in the Frog; it turns, however, more backwards than in the
Batrachian.

The orbital process, or blunt-pointed free pedicle of the mandibular suspensorium (qua-
drate), is far severed from the trabecular base, and turns forward as well as inwards. ‘The
rest of the mandibular arch is feeble; its united halves form an acute angle in Aluco
flammeus, and a much wider angle in Asio and Striv. Soitis with the palate, this part not
being nearly so outspread in Aluco flammeus as in the other types. Still in this species
the pterygoids (pg) diverge very much, and articulate with basipterygoid processes
(bpg) very far apart, on the massive parasphenoid (pas). The pterygoids are slender,
sigmoid, and cultrate; they are toothed in front, and have a small epipterygoid crest (epg)
behind. In this species the palatines (pa) are almost parallel, and they are very contrary
to most of the bones, being thin and lathy. In this species the palatines are as simple
as in the Common Fowl, the transpalatine process being suppressed, and the muscular

AND DEVELOPMENT OF THE BIRD’S SKULL. 139

grooves almost invisible. In older birds a small interpalatine spur can be seen, but
in the young it is not evident. The ethmo-palatines (fig. 7, epa) are very short, and
backwardly placed; they are high, looking obliquely forwards and upwards, and become
spongy; in front they send forwards a flat half-coiled horn of bone; this is lost as a
separate process in the adult, for it coalesces with the much enlarged vomer, each
process adding to the depth of its shelving sides, and leaving a large fossa above, which
opens into most exquisite air-cells.

The palatines are kept apart by a triple wedge, the vomer (v) below, and the early-
coalesced mesopterygoids (mspg) above. This structure can only be seen in very young
birds before they leave the nest. Now the vomer is a smallish spindle; afterwards it
is a large dagger-shaped bone, with a deeply sulcate upper edge.

The thin fibrous prepalatine bars (fig. 5, prpa) form rests for the huge, ear-shaped,
swollen maxillo-palatines (map); their tip articulates with the imside of the splintery
palatine process of the premaxillary (ppx). The dentary part of that bone (dpz) is not
strong, and the beak part is hollowed below. ‘The fore end of the ossified septum nasi
coalesces with the premaxillary, and thus an extensive bridge is formed over the anterior
palatine space and under the external nostrils, the same as, but much more extensive
than, that seen in the Cariama (fig. 3, al), where the ossification is arrested.

- The hollow part of the beak in front of the nostrils has a bony septum between its
air-cavities, formed by the premaxillaries themselves (px), and lying in front of and
below the true septum nasi.

A considerable chink, 35 of an inch wide, separates the maxillo-palatines (map) ;
so that, notwithstanding their great size, these birds are only “ indirectly desmognathous.”’
The septum nasi is thoroughly ossified, but not the outer ale. It is of great antero-
posterior extent.

The orbital septum is a thick cushion of soft bone, very much bevelled away below,
but running forwards at its upper part; this arises from the direction taken by the great
cranio-facial cleft, which runs forward as well as upward.

The weak diverging jugal bars are composed of three elements, as in Accipiter (mz,
J; 9j); both the premaxillary and the maxillary have an angular process.

But these three elements are not all that is to be seen in the jugal bar; for that addi-
tional bone which I have called the postmaxiilary is here present (Plate XXIV. fig. 8,
ptmx); it is a smallish oval grain of bone, and can be seen in young birds not long
escaped from the nest. In this bird also can be seen an additional bone behind the orbit,
the counterpart of the ichthyic postfrontal or ‘“ sphenotic.” The lacrymal (partly
shown in fig. 8, 7) is very large and spongy, and does not send out a superorbital process
as in the “ Diurne.”

In the Long-horned Owl (Aso ofus) the palatines are supplied with a large vomerine
keystone (Plate X XV. fig. 10, v). Here the inner part of the palatine arch may have had
a single medio-palatine, or a pair of mesopterygoids; but they are not distinct, although
the specimen was a bird of the first summer; these sutures are early lost in the Barn-
Owl; and the medio-palatine of the adult is often formed of two mesopterygoids in the
young. I suspect it is so in this case.

SECOND SERIES.—ZOOLOGY, VOL. I. U

140 ‘MR. W. K. PARKER ON THE STRUCTURE

The vomer (v) wedges in between the small ethmo-palatine laminze (epa); and above it
is a similar bone, the median septo-maxillary (msm): both the bones are similar in
shape; but the vomer lies behind the bone above it. In the large Ketwpa ceylonensis
there are two similar bones (Plate XXV. fig. 11, v, mpa); but they are much smaller,
and the vomer is in front of the other, which is a medio-palatine.

In Athene noctua these parts are all coalesced together. The palatine keystone, made
carinate by asmall vomer, just shows traces of its sutural connexion with the ethmo-pala-
tines. This bird has large superorbital processes to its frontals, not to its lacrymals.

In the Hooting-Owl (Strix stridula) these parts remain separate (Plate XXV. figs. 20,
21). There is a small vomer (v), and over it a similar bone, as in Asio otus; both of these
arrested bones are in front of the short ethmo-palatines (epa); but behind these there
is a veritable medio-palatine (fig. 20, mspg, fig. 21, mpa)*; and the antero-superior
ossicle (msma) has the same position as the small median bone which I have called
the “median septo-maxillary ””’ in the Harrier (Circus cyaneus).

This is the bone which may possibly be a rudiment of the vomer of the other side
(right or left); for in the “ Picidze,” certainly, I have seen submesial bones join the mid
line; and the stronger push the weaker out of place.

In none of the Rapaces have I seen any remains of the paired “ septo-maxillaries ’’ so
common in the Passerinz, where they are small (although so large in Ophidia and Lacer-
tilia). Asa correlate of this state of things, the labial cartilages appear to be wholly
suppressed in these birds.

In Stri2 stridula (Plate XXYV. fig. 20) the transpalatine region (tpa) is as well developed
as in the Diurnal types; altogether it has a most instructive skull, and one quite unri-
valled for its swollen, spongy lightness of osseous tissue.

We have, even in this short survey of so small a tract of their structure, seen that the
Rapacious birds have much in common, and that the least specialized, as the Cariama,
marvellously wedges itself in among the roots of the most specialized of all, even the
Falcons and the Owls.

On the Structure of the Skull in the Laughing- Gull (Gavia ridibunda).

Several years ago a friend supplied me with a goodly collection of the young of this
species, and I took an early opportunity of working them out.

One of the first things which struck me was the Pluvialine character of these pulli, and
this at once struck me as being completely in harmony with resemblances I had noted
in these birds to the common Lapwing (Vanellus cristatus).

I am not sure which side of the boundary between the Plovers and Gulls one elegant
type—the Pratincole (Glareola)—should be placed; nor whether any man knows the
exact proportions of the Plover and of the Tern it contains in its organization. The fact
that Gulls are neither “ Altrices,” proper, nor true “ Preecoces ”’ makes their relation to
the Pluvialine types the more instructive; for the amount of super-pluvialine metamor-
phosis is not great, although real and measurable.

* This bone has been lettered in two ways, by mistake ; both mpa and mspg are correct, as a sengle mesopterygoid
becomes a medio-palatine.

AND DEVELOPMENT OF THE BIRD’S SKULL. 141

My views as to these relationships were fairly pledged several years since (see “ Gal-
linaceous Birds and 'Tinamous,” Trans. Zool. Soc. vol. v. p. 150); and I was soon led to
see how many large Families of Birds were constructed according to the pattern of the
Plover*.

But this was only one among several nascent generalizations germinating in my head
at the time ; so that when Prof. Huxley’s new ideas were given out, they fell into a recep-
tive soil, so far as I was concerned.

In this, as in other branches of zoology and morphology, there has been for many
years a remarkable harmony in our views; and nothing more struck and delighted
me, in that coincidence, than my friend’s commendation of one of the large groups
that seemed to me to gather towards one point: I refer to the double “ Gallo-Anserine”
family.

That the embryology of both the Goose and Fowl] was marvellously similar, I had then
seen; but I was not aware how close the Palamedeas on one hand come to the
Curassows and mound-making Megapods on the other.

But that which concerns us most at present is the huge number of birds with open
palates (‘‘Schizognathe”’) that have either a filial relationship to the Plover, or are
cousins-german to it.

It thus comes to pass that when once the structure of a Plover’s skull is thoroughly
understood, it serves as a norma to which the skulls of whole groups of birds can be
brought for comparison.

This is true, not merely of the related Schizognathous birds, but also of those Grallee
which are desmognathous; for the likeness and the difference between the split skull
of a fowl and the downd skull of a Palamedea is very similar to the likeness and the
unlikeness of the skull of the Crane on the one hand, and of the Heron and Stork on
the other.

There is one thing which is apt to mislead the zoologist in the consideration of the
great group of “ Carinate” birds; they, with the “‘ Ratitz,” are unconsciously compared
with the whole of the Mammalia. At first view it might be thought that the Ratitze
equalled, as a group, the “non-placental”” mammals, and the Carinatz the placental ;
but the groups are not properly comparable; the Carinate birds, notwithstanding their
number and variety, could not be equalled with more than two or three at most of the
Mammalian Orders, and not with the whole herd of the Placentalia.

Those morphological modifications which lie at the root of the diverse specializations
of the Carinate birds take place, as we pass from type to type, in the gentlest manner; all
the types of skull may be seen to spring from a fundamental form, rich in Rhyncho-
saurian, and even in Batrachian characters. Such a skull the Ratitee all possess.

After analyzing a skull of this kind, and finding how it is built up, as it were, accord-
ing to old styles of skull-architecture, familiar to him who knows the lower or cold-
blooded types, the morphologist is brought to a stand; for here the lines of life diverge.

* This. was handled excellently by Prof. Huxley in his “ Classification of Birds” (Proc. Zool. Soc. 1867, pp. 426—

431 and 456-458).
v2

142 MR. W. K. PARKER ON THE STRUCTURE |

Standing in the ways to see and to judge which is the right path for each Carinate
type, the worker finds it no “ far call” from the order-joining Tinamous to the Plovers on
one hand, and the Fowls on the other. j

The sucker which grows into the Passerine tree is not so clearly discernible in the
Tinamine stock—in such Tinamous as now exist; but just above where the fork is, the
Hemipods are unmistakable in their relation to the harsh-voiced Passerines of the southern
hemisphere.

Above, when the Tinamine branch forks amongst types of no great elevation, and in
which the Dromzeognathous skull has passed into the Schizognathous, we see the nobler
forms (“ Aves aérize”’) just superimposed upon and, as it were, growing from the lower
forms—the “‘ Preecoces.”

From above the Sand-Plover arose the Pigeon, in its older and now extinct forms;
and from above old types of the Gallinaceze the fundamental or stock-form of the Rapa-

cious bird sprung. The genus Dicholophus is but little changed from such a primitive
plunderer.

Skull of Gavia ridibunda. 1st stage.

There were five stages of the young of the Laughing-Gull amongst those examined by
me; to these are added the adult; the largest young were just ready for flight, the
smallest only about a week old.

After these had been worked out, I obtained half-ripe young of the Silvery Gull
(Larus argentatus) from the “ Lesser Orme’s Head,” near Llandudno ; but these, although
studied in various ways, have not been wholly worked out by me; their condition at that
early stage differs but little from embryos of the Common Fowl (see Phil. Trans. 1869,
plate Ixxxii., 3rd stage).

In the first stage the primordial skull (chondrocranium) is thoroughly formed; it has
also most of the osseous centres commenced, and some of them ankylosed to their neigh-
bours. The occipital region is very instructive ; the basiocciptal (Plate XX VI. fig. 2, bo)
is a long oval piece, enclosing the starved notochord, and separated by large cartilaginous ~
tracts from the lateral pieces, the exoccipitals (eo). These bones are very irregular
pentagons; they are creeping over the hinder part of the auditory capsule, the canals of
which are seen shining through the clear cartilage (figs. 1-3, hse, psc). The ex-
occipitals already half enclose the foramen magnum (fm) ; but a good space exists between
them and the double keystone piece, the superoccipital (so). This latter bone is com-
posed of two halves, which are only half-soldered together ; above they are quite distinct.
They now form a very elegant bony plate with many sides, some having a scooped and
others a rounded outline.

The archway of the foramen magnum is finished by the concave margin of the
coalesced part; above, behind, and below the parietals (p), the margin is a bifoil.
Between these margins the bone has an indrawn waist; on each side, at this part, there
is a non-cartilaginous space—a “fenestra” or “ fontanelle,” the lateral occipital fon-
tanelle so familiar to osteologists in Pluvialine and Anserine types (see in Vanellus
cristatus, Trans. Zool. Soc. vol. v. plate xxxvii. figs. 1, 2, & 4, J. 0. f).

AND DEVELOPMENT OF THE BIRD’S SKULL. 143

Outside these spaces the bone is ear-shaped, the substance of it spreading towards
the ear-capsule.

These capsules are very large, and being impacted into or, at least, grafted upon the
large occipital arch and wall, appear very considerably in the back of the head.

Ossification has appeared in them, and the largest bony piece, the prootic (pro), is seen
behind the quadrate (q) in the lateral view (fig. 1); the other centres will be described in
more advanced stages, but the opisthotic is already well developed, although much
smaller than the prootic; the epiotic has not yet appeared.

The common sphenoidal region is at this stage a part full of interest to the mor- |
phologist. The three divisions of the great winged parasphenoid, although ankylosed, are
very evident. They are the rostral portion in front (figs. 1 & 2, pas), and the basi-
temporal plates (d¢) behind. These three bones are setting up ossification in the
overlying chondrocranium, although developed at first as parostoses and quite inde-
pendent of it.

As in the Ichthyopsida, these primordial splints are very large in relation to the rest

of the skull. That this should be so, shows how little the Bird is related to the ordinary
Lacertian type of Reptile, where the only remnant (not always present) of the para-
sphenoid is the anterior part. On each side of the scooped (Eustachian) part of the
median bone there is a cartilaginous bud; this is the basipterygoid process (pg), a
typically Pluvialine structure.
_ Behind these knobs the bone spreads into large wings that enclose the “ anterior
tympanic recesses.” Beneath these imperfect trumpets lie the basitemporal bones (6¢) ;
they have nearly melted into each other at the mid line, and above they are being
soldered to the tympanic wings of the parasphenoid.

But the overlying cartilage around the pituitary body and beneath the optic nerve (2)
has become osseous directly from the bone beneath; this is the true basisphenoid (ds) ;
the square inturned large alisphenoids (qa/s) are also ossified.

A large bony leaf is seen in front of the orbital septum ; it is trifoliate behind, and does
not nearly reach the membranous window between the eyes (iof).

In the figure (fig. 1) the septal portion of the nasal labyrinth has been cut away (see
this part from below in fig. 7); this is at the cranio-facial “ notch,” and the part in front
is only supplied by the orbito-nasal branch of the fifth nerve, and not by the olfactory.

This bony plate is the perpendicular or mesethmoid (pe) ; its huge size has relation to
the abortive development of the anterior sphenoid, and its bony matter will not cease to
grow until it has reached that of the posterior sphenoid (4s) beneath the optic nerve (2).

Nothing but development could possibly throw the least light upon this anomaly ;
there are no orbito-sphenoidal alee, and the presphenoid is merely that bar of cartilage,
afterwards bone, which is seen bounding the posterior half of the interorbital fenestra
(tof) behind; it reaches neither the top nor the bottom of the interorbital chondro-
cranium.

The top of the chondrocranium finishes behind in a median projection, behind and
between the upper turbinal scrolls (aliethmoids); this is a rudiment of that cartilaginous

144 MR. W. K. PARKER’ ON THE STRUCTURE

cloth which is thrown over the brain-sac in the Salmon and Polypterus (see my memoir
on the “Salmon’s Skull,’ Phil. Trans. 1873, plate v. ¢.cv), the tegmen craniv.

The reason for the membranous space between the eyes is not primarily teleological.
Nature is not husbanding her cartilage as being poor in that substance; it is a secondary
cleft, tending to divide the overlying ethmoidal structures from the crest developed on
the coalesced trabecule.

The roofing bones are seen to be well developed, the frontals, parietals, and squamosals
(fp, sq); the latter are elegant subfalcate bones, running far along the postorbital part
of the frontals, and somewhat trilobate below, where they clamp the ear-capsule. The
columella (s¢) is ossified in its medio-stapedial region, and the quadrate also, not its pedicle
or orbital process; its otic facets and its double lower condyles are all soft as yet.

Tn this stage I have only brought into view the hinder facial parts ; the pterygoid (pg)
has now a long splint-like process which reaches to the corresponding crus of the vomer,
not here shown, but seen in fig. 12, v*.

Afterwards the absorbents fret a suture across the base of this spur, and convert it into
a separate mesopterygoid, which, however, is only distinct for a few weeks, for it soon
coalesces with the postero-superior edge of the palatine.

The relation of the jugal to the quadrato-jugal, and of the latter to the quadrate, ito
which its hooked end is inserted, is here shown (fig. 1, /, ¢/, q).

The long dentary, which has coalesced with its fellow at the “mentum,” has been
removed, but the proximal part of the free mandible is shown (fig. 4, outside; fig. 5,
inside).

Meckel’s cartilage (mz) is still large, and runs far forwards; its articular end, with its
short Pluvialine posterior and internal angular processes (pag, iag), is ossifying ; there is
an endosteal patch and a long ectosteal plate (ar).

I see no separate coronoid, a bone often indistinct in birds; but the splenial, angular,
and surangular (sp, ag, sag) are large and still free.

In these long-faced birds the glossal portions of the hyoid are more than their hypo-
hyals; they deserve the name here of cerato-hyals (fig. 6, chy); they are unossified and
confluent.

There is a common cartilaginous basibranchial element (40r), pointed and delicate
behind, and only the proximal part of the first branchial arch (thyro-hyals, brs) is
ossified ; the dorsal element is long, slender, and soft.

-The nasal cartilages have been cut away and turned on their dorsal face to show their
laminar outgrowths (fig. 7); the section was made close behind the hinge or notch (see
fig. 1), and a line in front of the meso-ethmoidal bony centre (pe).

Unlike what we see in Mammals, the turbinal that figures most largely here is that
belonging to the ale nasi. This alinasal turbinal (at)) is here a simple foliaceous
outgrowth (see it in section, fig. 8). Behind this, and rising above it, is the inferior tur-
binal (itd). This is a simple plate at its commencement; further back it is coiled upon
itself. In most birds this scroll has two coils.

* This is a primordial relation of the pterygoid and the vomer; it is found in the lowest Urodela, to say nothing
of the Rhynchosauria. See Huxley, Proc. Zool. Soc. 1874, pl. xxix., and Ginther, Phil. Trans. 1867, pl. i. fig. 2.

AND DEVELOPMENT OF THE BIRD’S SKULL. 145

Behind the inferior turbinal, and also above it, is the coil of the upper turbinal (wd),
which is merely the aliethmoidal region of the nasal roof turned in upon itself.

This is represented as cut through the middle in the inverted figure (fig. 7, wtb); this
and the front wall of the pars plana (see fig. 15), which has no definite middle turbinal on
it, form the proper olfactory territory.

Skull of Gavia ridibunda. 2nd stage.

A somewhat more advanced stage (Plate X XVI. fig. 11) is shown in section as far as
to the cranio-facial isthmus (cfc); this figure supplies the deficiency of the last in not
showing a sectional view ; and it also shows some advance in the osseous deposits.

This section has been madea little to the near side of the mid line, and shows the para-
sphenoid and basipterygoid and the whole interorbital septum uninjured. The back-
tilted large auditory capsule is seen to lie both on and im the occipital arch; for the bony
matter creeping over the junction of the anterior and posterior semicircular canals is not,
in the bird, a large separate epictic bone as in reptiles; a rudiment appears afterwards
in this bird as in some others; but at present the super- and exoccipital is largely walling
in the otic mass.

The main periotic centre, the prootic ( pro), is now a large oblong bone, and separated
by the unossified auditory crest from the alisphenoid (als), and by synchondrosis below
from the basisphenoid (0s) and the basioccipital (do). The auditory nerve, or portio
mollis, pierces it, as also the portio dura, or facial nerve. Outside these foramina the
bone is scooped for the superincumbent brain, and becomes a mere crest; the edge
itself is still soft.

A clear wedge of bone is seen between the prootic and the exocciptal (eo) ; this is the
opisthotic (op); in the specimen it was becoming ankylosed to the prootic, but I have
diagrammatically given it a sharp suture from observation of the first stage.

The large passage (8) between the exoccipital and the opisthotic is for the glosso-
pharyngeal and vagus nerves ; the anterior and posterior condyloid foramina (9) are seen
on the edge of the exoccipital.

A large unossified space still exists on the crown of the arch of the anterior semi-
circular canal, and also in the recess for the cerebellar flosculus below it. If the bony
matter from the superoccipital had been somewhat less, then this space would have cor-
responded exactly to the region occupied by the epiotic bone in the Reptiles; its rudi-
ment has not yet appeared in the Gull.

The lozenge-shaped basioccipital is nearing the side piece (40, eo); but there is a large
spheno-occipital synchondrosis, underfloored by the basitemporal plate (07).

From the great submucous bones, the parasphenoid proper, and the basitemporals
(pas, bt), an upgrowth of bone has taken place, metamorphosing the overlying chondro-
cranium.

This bone, the basisphenoid (4s), underprops the optic nerves at their exit, and
embraces the dipping pituitary body, which passes down, as in osseous Fishes, to lie on
the submucous bony tract ; in the chondrocranium the pituitary space is open.

Into this pouched space the internal carotid (éc) enters; behind this space the cerebral

146 MR. W. K. PARKER ON THE STRUCTURE

mass lies on a shelving wall, the postpituitary wall, a scooped and slanting tract of bone,
split at the mid line, from which the notochord has retreated. This fissure is the
remains of the posterior basicranial fontanelle of the Reptile.

The prepituitary region has the anterior clinoid wall reaching the bony alisphenoid
(als), and a pinched front portion, which forms the postero-inferior angle of the inter-
orbital plate, and which afterwards coalesces with the huge perpendicular ethmoid below
the presphenoid.

This great mesethmoidal plate has increased in size, and is growing into the frontal
region. The frontals themselves at present fail to wallin the space caused by the
abortion of the orbito-sphenoids in the chondrocranium; there is, therefore, a post-
orbital fontanelle (po).

The subfalcate squamosal helps the alisphenoid (sq, als) to finish the side walls; it

clamps the prootic, overlies the alisphenoid, reaches the frontal, and supplements the
parietal (p) *.

Skull of Gavia ridibunda. 3rd stage.

This stage, in pulli a few days older than the last, has been worked out principally to
display the exquisite architecture of the great skull-floor, including the occipital, tem-
poral, and posterior sphenoidal regions.

At this stage we see best this most curious piece of morphological mosaic, not quite
unlike the wall-work of the test of an Hehinus, but of a more complex type by far, and
yet having fewer parts as far as mere number goes.

Indeed two very distinct strata are condensed and calcified to produce these results ;
and this difference is indicated in the figures by the colouring of the inner stratum,
formed by ossification of the chondrocranium, and the outer, which is the inner stratum
of the submucous connective web, is left uncoloured.

The first thing that strikes the eye in the under view is the long bony dagger and the
deltoid bone which lies beneath and behind it. These divisions of the ichthyic para-
sphenoid are the rostrum and the basitemporals (Plate XX VII. figs. 1-3, pas, bt); on each
side the rostrum widens between the basipterygoid cartilages (dpg), which are lessening ;
at the mid line it is scooped correlative to the meeting together of the right and left
Eustachian tubes (ew), and behind it simulates the basitemporal wings, widening and
spreading.

_ These hinder wings form the “ anterior tympanic recess ”’ (atr, ppg) : they are long,
faleate, notched, and grooved plates of bone, and with the help of the underlying
deltoid plate, the coalesced basitemporals (02), they form a pair of very elegant bony
trumpets leading into the drum of the ear, and are a specialization of the first visceral

* One of the older antomists (Owen), not seeing the squamosal in the inside of the struthious skull, took one of
his remarkable intellectual leaps, and supposed that it was always absent from that part, a hard thing to find in any
of the ‘ Carinatee.” ,

This great theorist wanted the bone on the outside and further down for transcendental uses; it, however, refuses,

to leave its own supratemporal region, for it must be ready to graft itself on the auditory mass in all the hot-blooded
Vertebrata.

AND DEVELOPMENT OF THE BIRD’S SKULL. 147

cleft. The hinder angles of the basitemporal plate are greatly produced, forming horns
that underlie the exoccipitals, whilst the posterior concave edge has lying on it the fore
end of the basioccipital (40).

This latter bone has on each side, between it and the paired exoccipitals, a reentering
angle of soft cartilage in front; but behind the three meet in the very substance of
the transversely oval occipital condyle; here, in meeting, they form a suture like a
leech-bite.

For the most part the occipital arch is well ossified (Plate XXVII. figs. 1-4); but the
whole of the outer edge of this transversely crested structure is still unossified, the car-
tilage not merely existing on the tympanic wings of the exoccipitals, but also on the
lower half of the superoccipital ridge (¢eo, so). The original suture between the two
superoccipitals is still visible above; but the “ lateral occipital fontanelles ”’ are nearly
filled in by bone on each side the narrow waist of the median part of the keystone.

We note here already the partial obliteration of the two very important Pluvialine
characters in these Sea-Mew chicks, namely, the basipterygoids and these occipital
windows. Here specialization has taken place in two ways :—first, by arrest and absorp-
tion; and secondly, by the secondary covering in of a region not filled in by the
chondrocranium.

The next important point to be noticed in these chicks of the third stage is the recol-
lection, as it were, of the epiotics left out in the first growth of bony territories; they
are, however, very small, and broken up into two or even three subcentres (Plate XX VII.
fies. 2, 5, & 6, ep).

This is seen many a time in ornithic morphology ; the bones which get the first start
in growth are large, whilst those which lose it are small, feeble, and overshadowed.

The upper view of the skull-base (fig. 2) shows well the round, deep, pituitary cup,
pierced below by the internal carotids (ic); also the slit below the hinder pituitary
wall, which is the old gap caused by the retirement of the notochord. The elegant basi-
occipital lozenge somewhat wedges itself into the end of the large, long, compound
basisphenoid.

But perhaps the most notable bones of all are the prootics; immense are they if com-
pared with those of the mammal. They are scooped laterally for the brain, and have
a lunate notch in front, which by a similar notch in the alisphenoid (a/s) becomes the
foramen ovale (5).

The prootics tilt themselves back to such a degree that the great anterior semicircular
canal is thrown, at its junction with the posterior canal, into the fore edge of the super-
occipital (figs. 4 & 5, asc, so).

If this figure (2) be compared with the rest, it will be seen how neat is the carpentry
by which the great ethmo-trabecular plate rests in the grooved upper surface of the
parasphenoidal rostrum (pas).

There it stands as a fixture; but this plate, the great meso-ethmoid, is almost sawn
through from below in front of the rostrum, and its under beam is the light and loosely

braced yomer.

SECOND SERIES.—ZOOLOGY, VOL. I. x

148 MR. W. K. PARKER ON THE STRUCTURE

Skull of Gavia ridibunda. 4th stage.

A little older than the last, the young of this stage yielded me the perfect longitu-
dinally vertical section (Plate X XVI. fig. 12) now to be described: it runs close after
and further illustrates the third stage.

The first sight of this object, and indeed of its figure, suggests feathery lightness ; and
the whole sum of its specialization, as compared with what is seen in the lower classes of
Vertebrata, may be taken as typical of this group (the Carinatze), in which the Reptile
so marvellously culminates.

The actual brain-cavity only occupies one third of the entire length of the head; the
nasal region proper, where the olfactory nerves are distributed, is exactly in the middle;
the ear-capsules run into the hinder arch; and the eyes are fairly halfway between the
other sense-capsules.

The great mesethmoid (pe) has now reached the interorbital fenestra (tof) above,
and behind it the cartilage ends in a retral spike, and in a groove below this lies the
olfactory crus.

Then in front of that bony plate we see how the cranio-facial axis has been gnawed
away, and a main and a lesser notch are seen leaving still a thick cartilaginous isthmus
connecting the small septum nasi (sz) with the main ethmoid.

In front of the former plate there is a large open gap before we reach the short, solid,
hooked end of the premaxillary (pz).

The feeble maxillary (ma) is left im sitw, and its maxillo-palatine plate (map) is seen
rising obliquely and leaf-like, on each side of the cranio-facial notch; inside of it is the
forked and notched double vomer (v); the rest of the face has been removed.

Behind, we see what has been already described in the second stage (compare figs. 11
& 12), but with this difference, namely, that the bony territories are not hedged in with
cartilaginous balks, but have their margins edge to edge.

The prootic is still distinct; but the epiotic fragments are now lost in the coadapted
faces of the prootic and superoccipital ; also the opisthotic has coalesced with the pro-
otic and exoccipital.

The base of the cranial cavity is an ascending floor, concave below and behind, and
convex as it begins to close in in front: the brain sits on the rounded upper edge of the
presphenoid in front (Plate XXVI. figs. 16 & 17, ps).

Here the floor is the original membrano-cranium, the cartilage being stunted in this
part ; and any orbito-sphenoidal bones are merely such osseous centres as appear in the
fontanelle. Inthe Fowl there are two on each side, supplemented by the orbital plate of
the frontal. I see no evidence of any distinct centre here in this species; in the old
bird the bony matter merely runs a little way into the fontanelle from the presphenoid.

Skull of Gavia ridibunda. 5th stage.

In these ripe chicks the sutures are becoming obliterated fast, especially those of the
ossified chondrocranium; but there are still many clear divisional lines in the investing
or outer part of the skull.

AND DEVELOPMENT OF THE BIRD’S SKULL. 149

The occipito-otic region is now one large shell of bone (Plate XX VII. figs. 7-10); in
the hind region (fig. 10) the “ lateral occipital fontanelles ” (dof) are still open.

The basitemporal plate (fig. 9, df) has now obtained its perfect form; it is subtriangular,
but the sides are convex, and the base is emarginate, and it has ear-like processes at
its angles; these help to floor in the tympanic cavity. From the parasphenoidal rostrum
(pas) the basipterygoids have disappeared, so that the Pluvialine stage has been passed
in this part of the skull.

All the elements of the face are very slender and elastic, both those of the outer and
also those of the inner series. The great perpendicular ethmoid (pe) has reached the
“notch” in front and above, and hs nearly met the basisphenoid behind and below.
There are still large postorbital and a large interorbital fenestra ( pof, tof).

The nasal cartilages (the fore part of which had been removed in the specimen figured)
are all cartilaginous, with the exception of the back wall or pars plana (pp), which
is made bony by a centre of its own.

Outside this there is an anvil-shaped lacrymal, Pluvialine in form, but answering to
that of the larger types; its lower crus rests upon the outstretched antorbital plate
(Plate XXIV. fig. 7, 7, pp). The upper turbinal coil (aliethmoid) divides the groove for
the olfactory nerve from that for the trabecular or naso-orbital.

The nasal processes of the premaxillaries were laid upon ‘and wedged in between the
upper plate of the nasals (xpx,n). The upper plate of the nasals lies upon and is wedged
in between the frontals (7, f). The frontals are sinuously applied, behind, to the fore
margin of the parietals (f, p); this harmony-suture is the “coronal” of human anatomy.
A “sagittal suture” runs along the whole skull and face, from the fore end of the nasal
fossa, where the premaxillaries are soldered together, up to the notch in the upper surface
of the superoccipital (fig. 10, so), the remains of the divisional line of its two primary
halves.

The falcate squamosal (sq) elegantly binds the whole cranial cavity together on each
side; it runs upwards and downwards obliquely, behind, lying like a tile over the slant-
ing occipital edge.

The nasals, as is the wont of the Pluvialine types, are sharply split into two crura
(figs. 7 & 8, 7). These run forwards, tying down the upper and lower branches of the
premaxillaries (7, npx, dpa).

The pterygoids (pg) are long, slender, and inbent. They are pedate behind, having
an epipterygoid process, with an oval hollow above; and the proper end of the bone has
an acetabular facet ; both these embrace similar convexities on the fore margin of the
quadrate (q).

In front the pterygoid, having lost its mesopterygoid spike, which is now part of the
palatine, fits, by a short tooth, into a shallow socket in the end of the palatine (pa). In
front, the palatine is a long, slender needle of bone, which is now tied to, and afterwards
anchyloses with, similar needles, namely, the fore part of the maxillary (ma) and the
palatal process of the premaxillary (ppz).

The outer edge of the two-keeled hinder part of the palatine has no more outgrowth
as a proper transpalatine portion than the Plovers. (See my “ Gallinaceous Birds and

x 2

150 MR. W. K. PARKER ON THE STRUCTURE

Tinamous,” Trans. Zool. Soc. vol. v. plate xxxvii. figs. 1-5, for views of the skull of
Vanellus cristatus, useful for the whole comparison between the Larine and Pluvialine
skulls.)

The solid part of the premaxillaries is short. The maxillary widens at its middle, to
form the elegant spoon-shaped maxillo-palatine process (map), which is obliquely placed,
and has its “bowl” outside. The rest of that bone and the two pairs of jugals (7, ¢/)
are very slender spicules of bone.

The vomer (Plate XXVILI. figs. 9 & 9a, v) has the form so common in Pluvialine, Gruine,
Ralline, Alcine, and Ardeine types; it is formed of two lanceolate centres that unite for
their anterior two thirds by a sharpish angle. They become carinate below, and the
carina is, in the young bird (fig. 9a), divided into two tracts. These afterwards are joined
by fresh bone into one keel, which forks where the bone forks to form the crura that
unite with the under surface of that part of the palatine which was the distinct meso-
pterygoid bone.

The very pluvialine mandible (fig. 7, d, ar) has still a suture where the anterior and
posterior parts unite ; there is also an oval fenestra in front of the hinge.

Skull of Gavia ridibunda. 6th stage.

In the skull of the oldest specimen I have examined of this species the occipital
fenestree and the basipterygoids are entirely gone. The bone is light and thin, but
rather dense; scarcely a trace of suture is to be seen—for instance, only above the
cranio-facial hinge. The only bones free are the vomer, the quadrates, and the mandible.
The latter bone has its sutures between the dentary and the hinder part persistent.

The nasal capsules are only ossified in the olfactory region (aliethmoid and pars plana) ;
the vestibular cartilages remain soft, viz. the aliseptal with its inferior turbinal, and the
alinasal with its turbinal.

But there is a structure of intense interest attached to the outturned end of the pars
plana, and not differentiated from it in the cartilaginous state ; this is the “os uncinatum”’
(Plate XX VII. fig. 11, ow). This is a small triangular wedge of bone which rests upon
the zygoma. It is well shown in the Albatross (Diomedea). I have lately described it in
Dicholophus ; traces of it occur in Alcea torda and Uria troile; but its highest development
is found in certain arboreal birds—Plantain-eaters, Parrots, &c.

{his bone is the ethmo-palatine, or joining-piece between the trabecula and palatine
in front, and belongs to the same category as the basipterygoid.

On the roof of the skull large and elegant fosse exist, on which lie the long, tongue-
shaped supraorbital or nasal glands. In old birds the two long ceratohyals are only
soft at their apex or ventral extremity ; they ossify to a great extent, ankylose in the fore
part, then are somewhat bowed out apart, and then come close together in front of
the basihyal. This latter bone sends osseous matter into more than the front half of
the urohyal. The lower thyrohyals are ossified, The upper and more slender pieces
are half soft below and at the upper ends.

But little difference would be found between this bird in its development and the

AND DEVELOPMENT OF THE BIRD’S SKULL. 151

growing young of a species of Podiceps, Colymbus, Alca, or Uria; they all specialize
by passing just a stage or so beyond the simple Pluvialine pattern, but in different
degrees. The Petrels are close congeners of the Gulls; many of the smaller kinds
retain their basipterygoids. A large number of water-birds keep their “lateral occi-
pital fontanelles” open throughout life. (See, on the Pluvialine types, Huxley, Proc.
Zool. Soc. 1867, pp. 426-431.)

EO ot B® 09

DESCRIPTION OF THE PLATES.

PLATE XX.

. 1. Vertical section through head of embryo Fowl (Gallus domesticus), about the beginning of the

sixth day of incubation. x 7 diameters.

. The same, taken further back through the eyeballs and nasal sacs. x 7 diameters.

. The same, behind the nasal walls. x 7 diameters.

Part of same section. x 27 diameters.

. A section further back, through the eyes, pituitary space, and lower jaw. x 7 diameters.

. Section through ear-sacs of the same embryo. x 7 diameters.

. A longitudinally vertical section of the hind face of an embryo of the House-Martin (Chelidon
urbica). x 15 diameters.

ig. 8. Inner view of auditory structure and mandible in a fledgling Carrion-Crow (Corvus corone).

x 3 diameters.

. 9. Os quadratum and stapes of adult Jackdaw (Corvus monedula). x 6 diameters.
. 10. Stapes of Pipmg-Crow (Gymnorhina tibicen), inner view. x 10 diameters.

. 11. The same, edge view. x 10 diameters.

g. 12. Palatine view of skull of Starling (Sturnus vulgaris). x 4 diameters.

g. 13. A species of Cardinal (Cardinalis, ?sp.). x 54 diameters.

Prats X XI.

.1. Palate of embryo of Brown Linnet (Linota cannabina) of the sixth day of incubation.

x 15 diameters.

.2. Part of same. x 150 diameters.
. 3. Part of embryo of same bird, seventh day of incubation. » 20 diameters.
Fig.

4, Vomer and part of palatines of a fledgling Sparrow (Passer domesticus), seen from above.
x 44 diameters.

152 MR. W. K. PARKER ON THE STRUCTURE

Fig. 5. Palate of common Wren (Troglodytes vulgaris). x 51 diameters.
Fig. 6. Part of same. x 11 diameters.

Fig. 7. Side view of face of the Wren. x 54 diameters.

Fig. 8. Palate of adult Goatsucker (Caprimulgus europeus). x 24 diameters.
Fig. 9. Part of same, seen from above. x 44 diameters.

Fig. 10. Part of face of same bird, upper view. x 24 diameters.
Fig. 11. Part of palate of young Goatsucker, seen from above. x 4 diameters.

Puate XXII.
Fig. 1. Palate of a still younger Goatsucker (Caprimulgus europeus). x 4 diameters.
Fig. 2. Palate of Humming-bird (Patagona gigas). x 34 diameters.
Fig. 3. Part of same view. x 6% diameters.
Fig. 4. The same, seen obliquely. x 62 diameters.
Fig. 5. Cranium of same bird, seen from above. x 34 diameters.
Fig. 6. Part of same, side view. x 34 diameters.
Fig. 7. Hyoid bones of same bird. x 63 diameters.
Fig. 8. Palate of a nestling Humming-bird (? Lampornis) from Barbadoes. x 63 diameters.
Fig. 9. Part of same. x 10 diameters.

Fig. 10. Part of same, seen from above. x 10 diameters.

Prats XXIII.

Fig.
Fig.
Fig. 10. Stapes of same, outer view. x 74 diameters.

. Palate of same, with lower plate removed, seen from below. x 24 diameters.

Fig. 1. Palate of Scythrops nove-hollandie. x 24 diameters.
Fig. 2. Section of palatine bones of the same bird. x 21 diameters.
Fig. 3. Side view of face of same bird. x 21 diameters.
Fig. 4. Palate of Megalema asiatica. x 3 diameters.
Fig. 5. Side view of face of same. x 3 diameters.
Fig. 6. Palate of Podargus. x 1z diameter.
Fig. 7. Side view of skull of same. x 17 diameter.
8
9

. Side view of palate of same. x 21 diameters.

Fig. 11. The same, inner view. x 7% diameters.

PuatTe XXIV.

Fig. 1. Skull of Gyps fulvus, from above. + nat. size.

Fig. 2. Skull of Dicholophus cristatus, from above. + nat. size.
Fig. 2a. Part of side view of the same skull. x 2 diameters.
Fig. 3. Palate of same. x 14 diameter.

Fig. 8a. Vomer of same. x 3 diameters.

Fig. 4. Palate of Falco tinnunculus (nestling). x 2 diameters.
Fig. 5. Palate of Aluco flammeus (fledgling). x 2 diameters.
Fig. 6. Part of same, from below. x 4 diameters.

. Part of same skull, side view. x 4 diameters.
. Skull of Buteo vulgaris (recently fledged), part of side view. x 13 diameter.

Fig.

4,
5
6
Fig. 7. The same, from above. x 4 diameters.
8
Fig. 9

AND DEVELOPMENT OF THE BIRD’S SKULL. 153

Piatt XXV.
Fig. 1. Upper beak and nasal opening of Falco tinnunculus (nestling). x 8 diameters.
Fig. 2. The same in section, seen from the inside, and half the inferior turbinal showing. x 3 diameters.
Fig. 3. Nasal capsule of same, with bones removed, from below. x 3 diameters.
Fig. 4. The same in longitudinal section. x 3 diameters.
Fig. 5. Transverse section of same, through the beginning of the inferior turbinal. x 6 diameters.
Fig. 6. Transverse section of same, through alz nasi. x 6 diameters.
Fig. 7. Palate of Accipiter nisus (fledgling). x 4 diameters.
Fig. 8. Part of same, side view. x 4 diameters.
Fig. 9. Part of palate of Helotarsus ecaudatus. Nat. size.

Fig. 10. Part of palate of Asio otus, lower view. x 3 diameters.

Fig. 11. Similar figure of Ketupa ceylonensis. x 2 diameters.

Fig. 12. Vomer of Falco peregrinus (?), side view. x 3 diameters.
Fig. 13. The same, under view. x 38 diameters.

Fig. 14. Vomer of Neophron percnopterus, from below. x 38 diameters.
Fig. 15. The same, side view. x 3 diameters.

Fig. 16. Pterygoid of same bird. x 38 diameters.

Fig. 17. Part of palatine of Gyps fulvus, upper view. Nat. size.

Fig. 18. The same, from below. Nat. size.

Fig. 19. Palate (part) of Sarcorhamphus papa, from below. Nat. size.
Fig: 20. The same in Sfria stridula. x 2 diameters.

Fig. 21. Part of same, oblique view. x 4 diameters.

Note.—The letters mspg in fig. 20 and mpa in fig. 21 refer to the same bone; fig. 21 is inverted (by mistake).

Prate X XVI.

Fig. 1. Skull of Gavia ridibunda (first nestling), side view, partial. x 3 diameters.
Fig. 2. Same, from below. x 3 diameters.

Fig. 3. End view of same. x 3 diameters.

Fig. 4. Hinder part of mandible of same, outer view. x 3 diameters.

Fig. 5. Same, from within. x 3 diameters.

Fig. 6. Hyoid of same specimen. x 3 diameters.

Fig. 7. Nasal labyrimth of same. x 5 diameters.

Fig. 8. Transverse section of nasal vestibule of same. x 20 diameters.

Fig. 9. A like section, further back. x 20 diameters.

Fig. 10. Another section still further back. x 20 diameters.

Fig. 11. A longitudinal view of the inside of the skull of second nestling (partial). x 3 diameters.
Fig. 12. A similar (perfect) section of the skull of fourth nestling.

Fig. 13. Section of alee nasi of fourth nestling. x 20 diameters.

Fig. 14. Section through ethmoid of fifth nestling. x 5 diameters.

Fig. 15. Section near “ notch” of ethmoid of fourth nestling. x 5 diameters.

Fig. 16. Another section of same, through interorbital septum. x 5 diameters.

Fig. 17. A like section, behind the fenestra. x 5 diameters.

154 MR. W. K. PARKER ON THE STRUCTURE OF THE BIRD’S SKULL.

Puate XXVILI.

Fig. 1. Basis cranii of third nestling, from below. x 3 diameters.
Fig. la. Alisphenoid of third nestling. x 3 diameters.
Fig. 2. The same basis cranii, from above. x 3 diameters.
Fig. 8. Side view of same. x 3 diameters.
Fig. 4. End view of same. x 8 diameters.
Fig. 5. Inner view of auditory region of the same. x 6 diameters.
Fig. 6. Inner view of the other side. x 6 diameters.
7. Side view of skull of fifth nestlmg. x 1}? diameter.
8

. Upper view of same. x 14 diameter.

Fig.
Vig.
Fig. 9. Lower view of same. x 1? diameter.

Fig. 9a. Vomer of same, side view. x 4 diameters.

Fig. 10. End view of same skull. x 12 diameter.

Fig. 11. Part of side view of skull of the adult bird. x 6 diameters.

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THE
TRANSACTIONS

OF

THE LINNEAN SOCIETY

OF

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART THE FOURTH.

tO Ne DeOrN:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET :
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER- ROW.

- M.DCCC.LXXVII.

wh ibaa . iy ah
FO i) CUS ; i Bg Mot OE ry
; aie RAUCH GN MRD Oe
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tO gi VO a wey sh ;
Ae aN i , SASS mL A AORTA ato Ny mee: mea: (Ck ws yoy are Hes
, an Re Res AN tae oN Ps sib <n 3 su ha) shane: ua

ROOD.

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epee cnc aoe 9 :

00%, Mi

CONTENTS.

PART IV.—Janvuary, 1877.

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups. By
JRO WESEWOODS MOAN, Pens aw SMe 5) 2 om ee pare dlns

IX. On a New Genus of Trematoda, and some new or little-known Parasitic Hirudinei.
By Joan Dents Macponatp, W.D., F.R.S., Deputy Inspector-General R.N.,
Professor of Naval Hygiene Army Medical School. (Communicated by G. EH.
DOBSON UMCA. SIE Beet Sa caustiod Tron Wa, Pees series hen, 209

X. On the Osteology and Pterylosis of the Spoon-billed Sandpiper (Eurynorhynchus
pygmeus, Linn.). By Jonn ANDERSON, ID., F.R.S.H., FLS., P.ZS., §¢.,

Curator of Indian Museum, and Prof. Comp. Anatomy at Medical College,
(Cian, sh ccm “ese We cpa: gh ble Rete Reis iyi ghey tala decal ci ae NT

PP etnat eo

\

Tain! a Ho See

NT I meee cc Cam

ah BAMo'VvO Waits Anion, Saw pinden’) wavy ayoradqobiqars oh Xp sage
ach aseg DRONE 2) 50! cq 1c po Ae I A Rs gs Joona

: F \ i ‘
| ianibnit toners winorkattil 0 osm amos baw hone Yami awl
* GAB Any d -eolosaeeh ysge 2 AL ML ,ctenoagell ava caOk Wi,
A yh GelnaioonmeteO ) ~ Joos Londo gare, QnSi YEE Ioan, Yo wweadhor a
} GOk, ms ¥ Pier ag. i fe) se a A ae aie 5 te (Gh i ohooh weoenokh,

- ado tomrstik) setabane holid-coog’ oY Ap eine Id huey yooleaied say
Lah BSN 2AM AT AG woonseck aah yh” CW atrotarg yay
AYN Ihibskl Io ymoinak .qudy Apvk Gan see svibal Wyo solo
Mec ae on) ne i erage acs, Se

[ 155 ]

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups.
By J. O. Westwoop, JZA., P.LS., Sc.

(Plates XXVIII.-XXXIIL.)

Read June 17th, 1875.

NOTWITHSTANDING the vast additions which have been made during the last
quarter of a century to the lists of species of Lepidopterous insects, especially by the
entomologists of our own country, the observation of Latreille, in his most valuable and
classical work the ‘Genera Crustaceorum’ &c., vol. iv. p. 186 ( Lepidopterorum ordo
entomologorum scopulus: horum insectorum etenim instrumenta cibaria simplicia; an-
tennee pro sexu diverse; metamorphoses permultorum nobis ignotz; idcirco nepotes
nostri methodum optimam soli conficient”’), is almost as true now as it was when
written, seventy years ago. In fact, the circumstance of such great numbers of species,
for the most part of exotic origin, being added to the already numerous tribes of Butter-
flies and Moths, without, in the greater number of cases, any actual knowledge of their
transformations or special structure being furnished by the captors or describers, has only
tended to render that “still darker which was dark enough before.” If, at the same
time, we turn to the generic distribution of these insects, we are bewildered by the
infinite number of new groups which have, as it appears to me, most unnecessarily been
introduced into the science, and the never-ending change in the employment of old
generic names. ‘This has in a great degree resulted from an insufficient appreciation of
the relative value of the characters which constitute a species and the illogical raising
of such characters to a generic rank. Thus no sooner had I published a monograph of
the Australian species of sac-bearing Moths, for which I adopted the generic name of
Oiketicus, proposed by the Rev. Lansdown Guilding, in the ‘Transactions of the Lin-
nean Society,’ in which paper I described the varying structure of the antennz, wing-
veins, and cases formed by the larve, than my specific details were seized upon, and each
species erected into a genus, characterized from my figures. Again, we have seen in a
recent American publication some of the Swallow-tailed Butterflies, which had uni-
versally been considered a species of the old genus Papilio, separated off from the
rest, and formed into different genera; whilst the old generic name Papilio has been
entirely removed from the family, and applied to one of the genera, or rather to one of
the species of the genus Vanessa—each of the other species of the latter group being also
raised to generic rank, whilst almost every species of North-American Hesperiide has
been formed into a separate genus. If, again, we look at the general classification of the
Order in the hope of supplying the desideratum of a ‘“ methodus optima,’ to which
Latreille especially alluded in the paragraph quoted above, I fear we are but little further
advaniced towards the attainment of that end than our forefathers. By the employment
SECOND SERIES. —ZOOLOGY, VOL, I. : ¥

156 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

of the transformations, and especially by the structure of the veins of the wings, feet,
antenne, palpi, tongue, and other characters of the imago, we have arrived at certain
results as to the characters of certain groups which had formerly been comparatively only
partially examined ; yet the want of a general revision of all the groups of the Order in
the same philosophical manner, has still left us in doubt as to the proper clue to the clas-
sification and relations of the families and still higher primary divisions of the Order.
I do not hesitate to say that the attempts to characterize and systematize the families
and subfamilies of Nocturnal Lepidoptera hitherto made are total failures.

Another difficulty, unknown to the older writers upon this order of insects, has arisen
from the exploration of numerous adjacent localities, which, whilst it has added greatly
to our knowledge of new and quite distinct species, has also shown that the wide geo-
eraphical range of a species is often attended with the development of slightly modified
races, which have by some writers been indifferently regarded as distinct species, or have
been sunk to the rank of local varieties. Thus, of the gigantic types of the diurnal
Lepidoptera which, from their size, have been well named Ornithopterus, and which are
natives of the East, we find the single species, Papilio Priamus, in Mr. Kirby’s Cata-
logue, made to comprise not fewer than seventeen of these local forms. These have been
specifically named, and regarded by others as distinct species, namely :—1. P. Pri-
amus, Linn.; 2. Panthous, Linn.; 3. Richmondia, G. R. Gray; 4. Cassandra, Scott;
5. Euphorion, G. R. Gray ; 6. Pronomus, G. R. Gray; 7. Poseidon, Doubleday ; 8. Cronius,
Felder; 9. Boisdwalii, Montrouzier; 10. Oceanus, Felder; 11. Arruana, Felder ;
12. Urvilliana, Guérin; 13. Triton, Felder; 14. Pegasus, Felder; 15. Archideus, Felder ;
16. Lydias, Felder; 17. Craesus, Wallace.

Another series of Hastern Butterflies, of which Papilio Paris is the type, distinguished
by having their wings powdered with golden-green atoms, exhibits a number of permanent
variations of no higher specific rank than those of the Priamus group in their respective
localities ; yet we find each of them, in Mr. Kirby’s Catalogue, given as a distinct
species.

The solution of this difficulty can only be hoped for by the very careful investigation
of such Protean species in their native haunts, with reference to their appearance in the
adjacent and intervening localities, so as, if possible, to determine how far the specific
type becomes modified (at least, so far as the markings of the wings are concerned) by
change of situation and other local circumstances. And here it is to be observed that
possibly too much weight has been eiven to the mere markings of the wings, without
sufficient attention to the question whether they are accompanied by structural differ-
ences. It was, for instance, with considerable doubt that I at first regarded several of
the Heliconiiform species of Oastnia described below as specifically distinct; the inves-
tigation of their wing-veins, however, satisfactorily proved that they were structurally
different from each other. But paradoxical as it may at first appear, the publication
of descriptions of Butterflies and Moths (unless belonging to well-defined and previously
characterized genera) has probably become the greatest obstacle in the way to our know-
ledge of the real “ methodus optima” desiderated by Latreille. Ido not hesitate to
affirm that the publication of the hundreds, nay, even thousands of careless and insuffi-

GENUS CASTNIA AND SOME ALLIED GROUPS. 157

cient descriptions, especially of Moths, in this country during the last twenty years, will
embarrass and disgust the student to such an extent as to retard the inquiry into the
general distribution of the Order, as well as the relations of the various families inter se,
for years to come. Without going to the extreme length which Mr. Scudder has done
in extending the characters of every genus over four or five closely printed pages (em-
bodying every point of the structure of the type in every stage of its existence, thereby
converting specific characters into generic ones), I yet hold that the description of a
Lepidopterous insect, without a thorough examination of its structural characters, is not
only detrimental to the science, but disgraceful to the author*.

In like manner the entire alteration in the old and well-established systems of celassi-
fication, both of the Rhopalocerous and Heterocerous Lepidoptera, and the substitution in
their stead of other systems which have only the charm of novelty and the love of change
to support them, is, as it appears to me, detrimental to the real advance of science. I
see, for instance, no reason or even advantage in removing the six-legged Papilionidee
from the head of the order, and substituting in their stead the Nymphalide, with their
imperfect fore feet, advocated by the German writers, and servilely adopted by their
English followers ; neither can I approve of the displacement of the gigantic Bombycide
(Saturnia, &c.) from the head of the Nocturnal Lepidoptera.

We thus perceive that, with respect to the discrimination of many species, or the limita-
tions of genera and families, or the general distribution of the primary groups of the
Order, we are scarcely further advanced than we were in the days of Latreille. The
genus Castnia of Fabricius affords an illustration of the various difficulties to which I
have above alluded. By Linnzeus and all the writers of the last century the species of
this interesting genus with which they were acquainted were regarded as Butterflies,
being arranged under the genus Papilio, on account of the clubbed structure of the
antennze. On the dismemberment of that great group, at the commencement of the
present century, by Fabricius and Latreille, the genus Castnia was established; but it
was still retained by the former as the 7th genus of the Butterflies, between the genera
Cethosia and Huplea, characterized only by the structure of the palpi and antenne + ;
whilst Latreille, in all his works, placed it (with Agarista &c.) at the head of the
Crepuscularia, between the Hesperiidee (with which the Uranie were united) and the
Sphingidee.

The connexion of these insects with the Diurnal Lepidoptera rests only on the structure
of their antenne, and on their evidently diurnal habits evinced by the brilliancy of their
colours. But when the antenne are carefully examined they do not bear out this con-

* It would. in this place, indeed be unjust not to refer to the admirable manner in which Mr. Stainton has worked
out the economy and structure of the small and difficult groups of the Linnean genus Tinea, in his different works on
the Microlepidoptera, whereby he has laid down the groundwork of a classification of these beautiful tribes, which
had previously been enveloped in doubt and difficulty. His figures also of the veining of the wings of these small
and difficult insects, as well as those given by the late Dr. Herrich-Schiffer in his ‘Systematische Bearbeitung der
Sehmetterlinge von Europa,’ will greatly assist the student in his investigation of the relations of these insects.

t System. Glossat. (Hd. Lliger, Mag. Ins. yi. 277, 1822). In Mr. Children’s abstract of this system, given in Taylor’s
Phil, Mag. for 1830, the generic name Castnia is misprinied Castina.

Y 2

158 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

nexion, differing materially from the real Rhopalocerous structure; whilst their con-
nexion with the Hesperiide is more remote than with any other group of the Diurna,
in consequence of the simple arrangement of the veins of the wings in the Hesperiidz.
The complicated arrangement of the wing-veins in Castnia, and the metallic gloss of the
very large scales (which are much larger than in any other Lepidopterous genus) with
which most of them are clothed, and the rich coloration of nearly all the species, in like
manner separate them from all the other groups of Heterocerous Lepidoptera.

The large and robust structure of the Castnia, together with their very ample wings,
separate them from all the groups to which they are assimilated by their transforma-
tion, except the gigantic Hepiali and Cossi, from which they are at once distinguished
by the veining of the wings and their clavate antenne.

In the complicated arrangement of the wing-veins, and especially in the existence of
the minute supplemental postcostal cell, we find an approximation to the Geometride ;
the details of so many species of which are illustrated by Herrich-Schaffer. But the weak
structure of these moths and their geometric larvee, destitute of several pairs of the
ventral pro-legs, will not allow any approximation between the Castnie and Geometre.

Another curious circumstance connected with the group, and which is not without its
significance in respect to their general relation, is the analogical resemblance exhibited
by some of the species to that strange family of Butterflies, the Heliconiide ; to these latter
we find similar resemblances exhibited by species, not only of other Heterocera, as
among the Arctiidee, but also amongst the most typical of all the butterflies forming the
family Papilionidee, as well as amongst the Pierides and Hrycinides.

Hence with regard to the actual relations of the Castnie, in the uncertainty which
still exists as to the preponderance to be given to the characters derived either from the
immature or perfect state of the articulated animals, we can arrive at no other con-
clusion than that which Dr. Boisduval has given in his last work, ‘ Hist. Nat. des Ins.
Lepidopt. hétérocéres :’'—‘‘ La famille des Castnides appelées Hesperi-sphinx par lillustre
Latreille, quoiqu’elles n’aient pas plus de rapports avec les Hesperies qu’avec les
Sphinx. Ce sont des insectes d’une création 4 part, qui sous leurs premiers états, ont a
la fois les habitudes des Sésiides et des Zeuzérides, mais auxquels il est difficile d’assigner
une place” (Pref. p. iv). .

The veining of the wings in this genus offers several modifications of much interest with
reference to the typical arrangement of these organs in the order Lepidoptera. In their
simplest form in the fore wings they may be thus described :—Ist. A strong simple branch-
less vein, running parallel with and close to the front margin of the fore wing (the costal
vein); 2nd. A vein arising close behind the base of the costal vein, but dividing into six
branches before reaching the extremity of the wing (the subcostal or postcostal vein and
its branches); 3rd. The strong median vein, separating into four branches; 4th. The
subanal vein, simple and branchless, but sometimes accompanied by one or two rudimental
longitudinal vein-like folds. In most instances, in order to give support to the middle of
the fore wing, there is a transverse veinlet connecting the 6th branch of the postcostal
(6 5*) and the 4th branch (¢ 8*) of the median veins, forming a closed cell (the discoidal
cell). But in all the families of Butterflies (except the Papilionide) the median vein has

GENUS CASTNIA AND SOME ALLIED GROUPS. 159

apparently only three branches, and the postcostal vein only five branches; the two
intermediate branches, namely the 6th branch of the postcostal (6 5*) and the 4th
branch (¢ 3*) of the median veins, have been assumed to be distinct from those veins,
and to constitute the remains of a vein intermediate between the postcostal and the
median, of which the basal portion has been aborted or seen only in a fold running
towards the base of the wing between those two veins. To these two detached branches
Mr. Doubleday applied the names of the 1st or upper discoidal nervule, and the 2nd
or lower discoidal nervule; the little transverse veinlet connecting the 5th branch of
the postcostal vein with the upper discoidal nervule he termed the upper disco-cel-
lular nervule; that connecting the upper and lower discoidal nervules together he
termed the middle disco-cellular nervule; and that connecting the fourth median (or
lower discoidal) nervule with the third branch of the median vein he termed the lower
disco-cellular nervule (these three disco-cellular nervules closing the discoidal cell); the
longitudinal fold or obsolete vein between the bases of the postcostal and median veins
within the discoidal cell, he termed the discoidal nervule.

From the very great number of specimens of Lepidopterous insects of all families,
both diurnal and nocturnal, which I have examined, I prefer to treat the first or upper
discoidal nervule as part of the system of the postcostal vein and the second or lower
discoidal nervule as portion of the median system. But in order to mark the distinction
between these two aberrant nervules or branches and the true branches, or veinlets, or
nervules of the postcostal and median veins, I have, in the Plates of details of structure
illustrating this memoir, distinguished the upper or first discoidal nervule with the mark
65*, and the lower or second discoidal nervule as ¢ 3*.

In thus differing from the system of venation proposed by the late Mr. Edward
Doubleday in his paper on Argynnis, published in the ‘ Linnean Transactions,’ and in
the introduction to the ‘ Genera of Diurnal Lepidoptera,’ I cannot but admit the propriety
of the adoption of the stems of the veins as of primary importance—contrasted with the
plan of Herrich-Schaffer and other German writers, who seem to regard the branches as
of primary value, counting them (quite independently of their origin from the stems)
numerically upwards from the anal angle, from 1 to 12:—No. 1 being the anal vein
(my d); Nos. 2,3, 4 the three branches of the median vein; 5 and 6 the two disco-
cellular veins (or my ¢3* and 0 5*); 7, 8, 9,10, 11 the five branches of the postcostal
vein, and 12 the costal vein+.

Many years ago I had a long controversy with the late M. Alex. Lefebvre on this
subject, in consequence of his adoption of the principle that the fold indicating the vein
in the discoidal cell of the fore wing, and running thence towards the outer margin,
formed the true le of demarcation between the branches of the postcostal and median —
veins. The fold in question is, of course, the representative of the discoidal vein; and an
examination of the figures accompanying this memoir, or those of Herrich-Schiaffer’s

+ Inhis elementary plate, H.-Schiffer omitted one of the branches of the postcostal vein, and so gave only 11 ribs
(as he terms the branches). Assmann (Schmett. Schles. tab. A. f. 1) counts the “‘ Adern” of the fore wing from 1 to 14,
adding the two supplemental anal veins, which are sometimes, although very rarely, to be observed.

160 MR. J. 0. WESTWOOD ON THE LEPIDOPTEROUS

work, will show the complete irregularity in the position of this discoidal vein and its
-divisions.

The large moths of the genus Gastropacha are excellent instances in which the
arrangement which I have endeavoured to establish is shown. In G. repanda (H.-Schaffer,
Syst. Bearb. Schm. Eur. pl. xviii. fig. 3), the median vein is seen to have four branches,
the upper one of which (¢ 3*) arises from the third branch (¢ 3), at the lower extremity
of the discoidal cell, whilst the other, upper, accessory branch, my 0 5*, arises from the
preceding, 0 5, far beyond the discoidal cell, forming as clearly a portion of the post-
costal system as c¢8* does of the median, there being no trace of a discoidal vein within
the discoidal cell.

Aglaope infausta (H.-Schaffer, Syst. Bearb. Schm. Eur. pl. xiii. fig. 32) presents us with
an interesting example, in which the arrangement of Mr. H. Doubleday is excellently
shown. Here the last branch of the postcostal vein (65) extends to the tip of the fore
wings, and the discoidal cell is divided into two nearly equal parts by the discoidal vein,
which, extending nearly from the base of the wing, branches into two equal branches
(my 6 5* and ¢ 3*), leaving the three branches of the median vein at equal distances
apart.

In Leucophasia sinapis (see H.-Sch. op. cit. pl. v. fig. 11) the arrangement of the
branches is very instructive. Here, in consequence of the very small size of the discoidal ~
cell, all the branches of the postcostal vein arise quite regularly on the anterior side of the
vein beyond the cell, the fifth (6 5) extending to the tip of the wing; but the two accessory
and the three branches of the median vein are all equally independent at their origin at
the end of the short oval discoidal cell, and appear to be parts of one system of veins
distinct from the postcostal.

In many of the Geometridee, as may be seen especially in the 9th plate of Herrich-
Schaffer’s work on European Lepidoptera, vol. vi., not only is the small postdiscoidal
cell, as I have called it (cellula accessoria of H.-Sch.), found in the Castnie present,
but it is even divided into two cellulets by a more or less oblique cross vein.

In the Hesperiide, on the other hand, the discoidal cell of the fore wing is elongate-
ovate, and all the branches of the postcostal and median veins, as well as the two
accessory branches (0 5* and ¢ 8*), arise from the extremity of the discoidal cell at
nearly equal distances apart, without any trace of the discoidal vein within the cell; so
that it is impossible, except from analogy, to determine whither each of the several
systems of veins extends.

The same arrangement is also seen amongst the He erbeeroes Lepidoptera, as in the
fore wing of Scardia mediella (H1.-Sch. op. cit. vi. pl. Microl. ii. fig. 1), and in Nemotois
scabriosella (ib. pl. iv. f. 32).

In some species of Hesperiidee, however, as in Hudamus (Megathymus) yucce, the
postcostal and median systems are easily distinguished, although their two outer
branches, 0 5* and ¢ 3*, are united by an extremely slender and angulated veinlet
imperfectly closing the discoidal cell.

From the numerous figures given in Pls. XXVIII. and XXIX., it will be seen that
the branches both of the postcostal and median veins exhibit much irregularity in their

GENUS CASTNI4 AND SOME ALLIED GROUPS. 161

arrangement as compared with the simple system seen in so many other groups of
Lepidoptera. The number of branches is normally present; but the 2nd, 3rd, 4th,
and 5th branches of the postcostal (d 2, 63, 64, and 65) are generally arranged so that
they form a small oblong premedian or postcostal cell. Looking at Pl. XXVIII. fig. 6
(C. cacica), we see that this is effected by the branch 63 throwing off a very short
transverse branchlet connecting it with 6 4; or, rather, we may say that 6 4: commences
transversely, and then suddenly bends at a right angle, and runs longitudinally, so that
the little transverse branchlet must be regarded as its basal portion. In 45 we have
the anomaly that the 5th branch arises at a short distance beyond the 1st branch, and
forms the hinder margin of the postcostal cell, throwing off a little oblique branchlet,
by which itis united to the 4th branch, and which in fact forms part of the extremity
of the postcostal cell. In like manner the median vein (c) has its third branch (c 3)
strongly angulated at a right angle at its base, whilst the branch ¢ 3* (lower discoidal
nervule of Doubleday) is carried backwards by a fold (z), the discoidal nervure of
Doubleday, and almost to the base of the wing f. Moreover the veinlet connecting 6 5*
and ¢ 3* is angulated below its middle, throwing off backwards a fold which unites
with 3, and which indicates the normal position of the discoidal nervure of Doubleday.
The object of this complicated arrangement, which has not hitherto been described,
appears to be to form a strong network of corneous veins for strengthening the wing,
so as to enable it to perform the strong flight which these muscles doubtless enable
the insect to perform.

Modifications of the arrangement described above exist in many of the species of
Castnia. :

In the splendid new species C. veraguana (fore wing, Pl. XXVIII. fig. 9) the distribution
is nearly similar to that of C. cacica; but there is no little transverse branchlet connecting
63 and 04, and consequently there is no closed postcostal cell, and the branch ¢ 3* arises
from ¢ 3 in the ordinary manner at a little distance in front of the transverse base of
e 3, making an acute angle instead of being pushed backwards a little distance, and
making an obtuse angle as in C. cacica. The curious species C. cochrus (Pl. XXVIII.
fig. 11) agrees with C. veraguana in this respect.

In C. Botsduvallii and other closely allied species (Pl. XX VIII. fig. 8) the small narrow
posteostal cell exists; but it only emits one branch at its extremity, which branches off
into 63, 64, and 05, and the angulated base of 05* arises from the posterior margin
of the postcostal cell near its extremity.

In C. icarus (Pl. XXVIII. fig. 13) the small postcostal cell exists, emitting 6 2 from its
front margin a little beyond the middle, whilst from its extremity one branch is emitted
which forks into 6 3 and 6 4, whilst 0 5 arises from the extremity of the postcostal cell,
close to the base of preceding branch, and 0 5* from 0 5 at a little distance from its base.

In C. Hubneri (Pl. XXVITI. fig. 14) there is no closed small postecostal cell, the little
transverse veinlet between 03 and 04 being wanting. In Gray’s figure, pl. 18. f. 2,
the vein is misrepresented as closing the cell.

In C. Latreiilii (Pl. XXVITI. fig. 15) the small postcostal cell exists, terminating in an

+ C. linus scarcely differs from C. cacica in the arrangement of the veins of its fore wings (Pl. XXIX. fig. 9).

162 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

acute point from which a single branch is emitted, which forks into two branches, 63
and 04, 65and 0 5* arising together from the hind margin of the cell at three fourths of
its length from the base. C.mygdon (Pl. XXVIII. fig.7) and C. inca agree with C. La-
treillu in this respect.

Among the Helicoinoid species with elongated fore wings, forming the genus Gazera,
Boisduval, C. cononia (Pl. XXIX. fig. 8) has a small postcostal cell, the 2nd branch of
the postcostal vein arising shortly before its extremity ; the 3rd and 4th branches arise
from its extremity a little distance apart, the short transverse branchlet connecting
65 with 6 5* springs close to the emission of 65 from the postcostal cell ; the 5th branch
arises from the postcostal cell nearly opposite the base of the 2nd branch, and appears
to extend backwards to the origin of the 1st branch of the postcostal. The branches
b 5* and ¢ 3* extend backwards, forming an acute angle within the discoidal cell, and
emit a branch-like fold (discoidal nervure) towards the base of the wing; ¢3 likewise
is extended backwards, uniting with this discoidal nervure, forming with it a rudimental
(postdiscoidal) cell.

In C. equadoria (Pl. XXVIII. fig. 10) the arrangement of the postcostal branches
agrees with C. cononia, except that the little transverse vein uniting branches 65 and
b 5* is rather nearer the base of the wing.

In ©. mimica (Pl. XXIX. fig. 11) and C. linus (Pl. XXIX. fig. 9) the postdiscoidal cell
is also present ; but it only emits one branch at its acute extremity, which forks at some
little distance beyond the cell into the two branches 03 and 0 4.

In C. truvilla (Pl. XXVIII. fig. 12), on the other hand, the postdiscoidal cell is not
completed, there being no connexion between the branches 63 and 64; the relative
position of the postcostal branches is otherwise identical with that of C. linus.

In C. Marcel Serresi (Pl. XXIX. fig. 2), forming the type of Boisduval’s genus
Ceretes, the small postdiscoidal cell is present, emitting the branches 0 2, 63, 64, and
65 from its extremity at smail distances apart, and the short transverse veinlet con-
necting 0 5 and 65* (and closing the discoidal cell dc) from its hinder margin beyond
the middle. The discoidal cell is traversed by a longitudital vein-like fold, the hinder
portion of the cell (d c*) forming a distinct cell by the addition of a veinlet extending
backward from the basal junction of the median branches ¢3* and ¢ 3—thus leaving
a narrow tract between it and the basal portion of the median vein c, which may be
called the premedian cell, p.m. c.

In C. chremes (P1. XXIX. fig. 4) the arrangement is nearly similar; but the vein-like
fold in the discoidal cell is less strongly defined, and the little transverse vein which
closes the discoidal cell does not arise from the hind margin of the postcostal cell, but
is pushed forwards and arises from the 5th postcostal branch close to its base.

On comparing the figures of the wings of C. cacica (Pl. XXVIII. figs. 6 and 6 a) with
those of C. Marcel Serresi (Pl. XXIX. figs. 2 and 3), it will be seen that the general
arrangement both of the veins and cells is identical.

The arrangement of the veins of the hind wings has been comparatively neglected in
the Lepidoptera, although Dr. Boisduval has partially, but not satisfactorily, used this
character in the generic distribution of his Castniaires. In the vast majority of the

GENUS CASTNIA AND SOME ALLIED GROUPS. 163

species of Lepidoptera the number of branches of the veins in the hind wings is smaller
than in the fore wings; but it will suffice to examine the wings of Hepialus humuli
(Pl. XXIX. figs. 27 & 28) to be convinced that the same elements normally exist in
the hind as in the fore wings. It will be seen from the lettering of the branches, that
the costal, subcostal with its six branches, median with its four branches, and anal veins
are present; and even the discoidal cell is cut up into the three discoidal, subdiscoidal,
and premedian divisions. This will enable us to determine satisfactorily the normal
condition of the veins and branches when some of the latter are not present in the hind
wings.

In C. cacica (Pl. XXVIII. fig. 6a) the subcostal vein has only two branches, the
second of which is united with the base of ¢ 3* by a very thin curved fold, whilst the
veinlet connecting the base of ¢3 with that of ¢3* extends backwards, and is continued
into a discoidal nervure extending nearly to the base of the wing.

In C. juturna (Pl. XXIX. fig. 1) the same arrangement exists, except that the thin
fold arising a little beyond the base of 6 2 and ¢ 3* is strongly angulated, and a very
slight longitudinal fold extends from the angle towards the base.

In C. cochrus (Pl. XXVIII. fig. 11a) the fold connecting the extreme base of ¢ 2
and ¢38* is almost obsolete, but is seen to spring from the discoidal vein at some distance
before the origin of ¢ 3*, thus leaving the discoidal cell nearly entirely open.

In C. Hubneri (Pl. XXVIII. fig. 14a) the discoidal is fairly divided into three portions,
namely, the discoidal, subdiscoidal, and premedian portions.

In C. paradoxa the vein or fold connecting the branches 62 and ¢ 3* is entirely
wanting, and the true discoidal cell is wanting; but its premedian portion is present,
in consequence of the branch c 3* being extended backwards, nearly to the base of the
wing. i

C. acreoides (Pl. XXIX. fig. 14) agrees with C. paradoxa, except that ¢2 and c3 are
united together at the base beyond the cell.

In C. linus (Pl. XXIX. fig. 10) the almost indistinct fold connecting the base of the
subcostal vein with the discoidal nervure is pushed still further back, and thus differs
materially in its relation to the branches of the median and subcostal veins.

In C. mimica the arrangement is the same as in C. linus, except that the transverse
fold closing the cell is obsolete, being only indicated by two minute tubercles at the
place of its origin (Pl. X XIX. fig. 12).

In both sexes of C. Marcel Serresi (Pl. X XIX. fig. 3) the postcostal vein and its two
branches are quite free; but c3* extends backwards beyond the base of ¢3, forking
into two branches, which curve backwards to the base of the wing, thus forming a sepa-
rate subdiscoidal cell, the anterior or discoidal part being open, and a very narrow pre-
median cell.

In the female of C. chremes (Pl. XXIX. fig. 5) the same arrangement takes place, but
the discoidal cell itself is partially closed by a thin fold; and in the male the branches
e3 and ¢ 3* arise close together at the extremity of the premedian cell.

Lhave thought it advisable to trace these veins and their branches to a normal state
in order, if possible, to ascertain their significance, either as sexual, specific, subgeneric,

SECOND SERIES.—ZOOLOGY, VOL. I. a

164 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

or generic characters. It appears to me that we have now sufficient materials before us
to assume that they possess no higher than specific importance.

The palpi in C. dedalus are closely applied to the face, extending upwards to three
fourths of the height of the eyes, with their inner margins close together; the apical
joint very minute, but distinct and pointed, but not at all projecting. In C. atymnus
they do not reach higher than half the height of the eyes, and the terminal joint is less
prominent.

In C. eudesmia the palpi are longer, and extend as much as one fourth of their length
above the top of the eyes; they are rather obliquely porrected, standing off from the
face, and are more slender. In C. decussata they are parallel, and reach to the top of
the eyes. In C. Hubneri they only reach to half the height of the eyes, and are much
more rugosely squamose. In C. Marcel Serresi they are small, not reaching so high as
half the height of the eyes, slender, oblique, pointed, and finely scaled. In C. cochrus
they are small, not reaching higher than the epistome, and are thickly squamose, but not
rugose. Such is also their structure in the Heliconiiform species, as well as in C. linus
and acreoides.

The geographical range of the whole of the species of Castnia and its immediate allies
extends only to Southern and Central America.

The transformations of the species of Castnia are of great interest, in connexion with
the natural relations of the group and its position with regard to the Rhopalocerous
division of the Order in which it was placed by the early systematists.

Madame Merian, in the 36th plate of her work on the insects of Surinam, gave a set
of figures as representing the transformations of Castnia licas. The caterpillar is eyi-
dently that of one of the larger Morphideous Butterflies; and the pupa is a chrysalis
fastened by its tail to the plant, with its head downwards, resting on the leaf, but with-
out any indication of a central girth of silk round the body. Dr. Klug was fortunate,
however, to obtain a pupa of C. therapow from the bulb of a Catasetwm, sent from
Central America to the Royal Gardens at Potsdam. ‘This pupa, figured in the Appendix
to his Memoir on the genus Synemon, agrees with those of Sesia, Cossus, Zeuzera, and
Hepialus, in its elongated general form and in the transverse rows of reflexed spines
with which the segments of the abdomen are furnished. It is, however, to Dr. Philippi,
of Santiago, Chile, that we are indebted for a knowledge of the larva of the genus Castnia.
In the ‘Stettiner entomologische Zeitung’ for 1863, p. 337, and plate iil., he has de-
seribed and figured the transformations of C. ewdesmia, the larva of which he found in
the stem of Pourretia coarctata, one of the Bromeliacez. It is 44 inches long, and
closely resembles the larva of a Prionus or Cossus, with a large prothoracic and short
meso- and metathoracic segments, with six thoracic, eight ventral, and two anal feet. It
is white and fleshy, with a fulvous-brown-coloured head and extremity of the body.
When full-grown it forms a large cocoon of bits of leaves, twigs, and other vegetable
matter, fastened together with silken threads. The pupa is dark chestnut-coloured, with
the abdominal segments paler.

In the Hopeian Collection at Oxford is preserved one of these cocoons (Pl. XXVIII.
fig. 4) from which I extracted a pupa of C. eudesmia, nearly arrived at the imago state,

GENUS CASTNIA AND SOME ALLIED GROUPS. 165

so that the scaly covering of the fore wings could be easily removed or raised, showing
the markings of the wings of the future moth of a diminished size (Pl. XXVIII. fig. 3).

The classification of the species of the genus Castnia has not hitherto been successful,
owing to the imperfect examination of the minute specific details of structure of many
of the species; nor can I hope to have been more fortunate than my predecessors.
Dalman, who described seventeen species of the genus in his memoir in the Swedish
Academy’s Transactions, remarked in his introduction that C. linus differed from the rest
in its more elongated and partially hyaline wings and abdomen, longer than the wings,
adding,—“ An proprium etsi propinquum genus he discrepantiz indicant?” In his
list of the species at the end of his paper he places C. cronus in a separate second section,
and C. linus in a third, without any characters; and his description of the former of
these two species is confined to its colour and markings.

Mr. G. R. Gray, m his memoir in the Entomological Society’s ‘Transactions,’ proposed
to divide the species into five sections :—

1. Those with the palpi very long, closely applied to the head, and covered with short
scales. The fore wings are subtriangular, thickly covered with scales, with the exterior
margin straight or slightly rounded ; the lower wings beneath with one guide (or bristle
working in the socket at the base of the fore wing). C. cyparissias, licus, evalthe,
Fonscolombet, &e. (pl. xiii. fig. 1).

2. Those with the fore wings elongate, triangular. The hind wings are expanded,
with the posterior margin somewhat truncated: C. ardalus, palatinus.

3. Those with the palpi short, reaching halfway towards the base of the antenn, and
covered with long scales; the fore wings covered with small scales, with the exterior
margin rounded. The outer margin of all prominently fringed with long scales: C.
Hiibneri (pl. xiii. fig. 2).

4, Those with the wings covered with minute scales, partly diaphanous; the fore pair
with the exterior margin rounded ; similar also in the hind pair, with three wing-guides :
C. cochrus, linus, and acreoides.

5. Those with the antennz similar to the others, but the hook at the apex is formed
of large scales. ‘The head of the male is very broad, and the eyes are much larger than
in the female. All the wings have the outer margin rounded, with a narrow fringe of
small scales, and with four wing-guides. ‘The sexes differ much in colour: C. nicon
(pl. xii. fig. 8), thats.

The plate accompanying the memoir of Gray gives the veining of the upper and lower
wings, the clava of the antennze, and the palpi of the 1st, 3rd, and 5th of these groups.

Mr. F. Walker, in his Catalogue of the Lepidoptera Heterocera, Part I., neglected the
various excellent characters pointed out by Mr. G. R. Gray, and divided the species
simply in accordance with the form of the wings, thus :—

Div. 1. Body thick.

Subdiy. 1. Fore wings slightly angular at the tips; hind wings rounded.
Sect. 1. Wings very broad. C. dedalus, Latreillii, atymnius, cochrus, &c. (16 species).
Sect. 2. Wings rather narrower; fore wings longer. C. satrapes, therapon, ardalus (Brecourtii) ,
invaria (.carus, var.), inca, palatinus, subvaria (phalaris).

Z 2

166 MR. J.O. WESTWOOD ON THE LEPIDOPTEROUS

Subdiv. 2. Fore wings more distinctly angular at the tips: hind wings slightly angular. 5 species—
C. Dalmanni, hegemon, orestes, Boisduvalii, and amycus.
Div. 2. Body more slender.
Subdiv. 1. Hind wings slightly angular. C. mygdon and phalaris.
Subdiv. 2. Hind wings quite rounded.
Sect 1. Fore wings rounded at the tips, moderately long. C. actinophous (acreoides), decussata,
cronis, strigata (Godarti), Hubnerz.
Sect. 2. Fore wings slightly truncated at the tips. C. chremes, thais.
Sect. 3. Fore wings rounded at the tips, very long; resembling the Heliconii in structure. C. linus

and acreoides.
Div. ?. C. Kirstenit.

Dr. Boisduval, in his ‘Species général des Lépidoptéres hétérocéres,’ published in
February 1875 (although dated 1874), has divided the Castnie (Tribu des Castniaires)
into the following genera :—

1. Castnra (including 53 species, without any attempt at subdivision). Characterized by
the “Palpes divergents de trois articles écailleux, montant 4 peine au niveau du
front, le dernier article petit et conique” (as shown in Gray’s figures 10, 1¢, 1d).
The “antennes en massue, cylindriques, terminées par une trés-petite houppe
soyeuse’”’ (as in Gray’s figure 1 a), the ‘‘ pattes—avec les jambes munies de deux
ergots.”’ The other characters cannot be regarded as generic.

2. CERETES (n. g., including C. chremes, thais, and Fabric (Marcel Serresi) = Gray’s
5th group and Walker’s Div. 2, Subdiv. 2, Sect. 2, characterized by the “ téte petite”
(overlooking the difference of the size of the head in the two sexes). Antennze with
a “longue massue fusiforme se terminant en pointe”’ (contrary to Gray’s descrip-
tion), “ palpes trés-velus appliqués fortement sur le front, non divergents, & articles
indistincts.” Fore wings with the “sommet un peu pointu,” and hind wings with
the “deux nervures médianes trés-rapprochées et coudées en sens inverse, mais ne
se joignant pas pour former une cellule fermée.” This last character is evidently
taken from Gray’s fig. 3, and answers to the hind wing of C. Marcel Serresi, but
not to that of C. chremes, in which the cellule is pro tanto ‘ fermée” by the fold, as
shown in my figure. The fact that Dr. Boisduval excludes C. diva, Butler (tricolor,
Felder) from this genus, and unites it (p. 531) with Castnia, notwithstanding its
extremely close affinity with C. chremes, is a sufficient proof of the unsatisfactory
character of the genus.

3. ORTHIA, n.g., having the head small, the antenne “en massue fusiforme, terminée par
une petite pointe, palpes tres-velus, trés-courts, a articles indistincts. Ailes allongées,
assez étroites, les inférieures ayant prés de la base une petite cellule elliptique
fermée.”

This genus is divided into two uncharacterized sections :—

+1. C. therapon, Koll. (paradoxa, Bdv.).
2. C. acreoides, Bdy.

GENUS CASTNIA AND SOME ALLIED GROUPS. 167

t13. C. pelasgus, Fab.
A. C. unifasciata, Felder (Voy. Novara).
5. O. augias, Bdy. (Herr.-Sch.).
6. O. nexa, nu. sp., regarded as the probable female of O. augias.

The genus Orthia, as constructed by Dr. Boisduval, is quite untenable. C. therapon
and acreoides have long fore wings, with a small front costal cell, the discoidal cell
closed as above described; whilst the hind wings have the discoidal cell open; that is,
there is no junction between the branches 6 2 and ¢3*, the small cell near the base of
the wing noticed by Boisduval being the cell which I have termed premedian. This’
cell, indeed, exists in C. pelasgus, the first of Boisduval’s second section of Orthia; but
the fore wings have no postcostal cell, the second and third postcostal branches arising
from a single base; and the 4th, 5th, and 6th branches also arising from a single basal
portion of the vein; the discoidal cell itself is quite open, there being no connexion
between the branches .65* and ¢3*. The very broadly triangular wings of C. pelasgus
will also remove it from the two preceding, and indeed from O. augias, which possesses
a small postcostal cell, a closed discoidal cell, and a closed discoidal cell in the hind
wings. I do not hesitate therefore to reject the genus Orthia.

4, GAZERA, n.&., characterized by the “‘ Antennes en massue allongée, terminées par une
5) oO” £
petite houppe soyeuse a peine visible. Palpes courts arrivant & peine au milieu du
front, de trois articles, le troisiéme trés-distinct, nu et pointu. Téte assez petite.
> P

Ailes allongées Héliconiformes, a écailles moins grossiéres que dans les Castnia pro-
prement dites; les inférieures ayant la cellule discoidale étroite fermée en angle
aigu. Abdomen un peu plus long que les ailes inférieures.”” Containing

1. C: limus, F. 4. C. columbina, Bdv.
2. C. heliconioides, Bdv. 5. C. zagrea, Feld.
3. C. personata, Web. 6. C. simulans, Bdv.

The elongate ovate form of the fore wings of the species of this group (to which, how-
ever, C. therapon and acre@oides must be added) is scarcely sufficient to separate them
generically (seeing the variable form of the wings in the other species) from the rest.
The other characters proposed by Boisduval are too variable in the different species to
possess generic value.

Finding it impracticable to carry out a distribution of the species of Castnia founded on
the form of the wings, as partially proposed by Mr. Walker, and rejecting Boisduval’s genera
for the reasons given above, I prefer to regard the species as constituting a single genus,
and shall content myself by dividing them into two sections :—1st. Those with the fore
wings more or less broad and triangular. 2nd. Those with the fore wings more or less
elongate-ovate, the latter nearly corresponding with Boisduval’s genus Gazera.

Genus I. CastTni..
Sect. I. Species with the fore wings more or less broad and triangular.

1, CASTNIA D&aDALUS. C. alis fuscis metallico nitentibus, anticis fascia media angusta,
obliqua lateribus parallelis fere ad angulum internum extensa alteraque subapicali

168 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

magis irregulari e lunulis connexis, albis; alis posticis fuscis extus nigricantibus,
fasciis duabus macularum parvarum albidarum (margine externo parallelis), interna
interdum minus distincta. Long. corp. unc. 23. Expans. alar. ant. unc. 6-73.

Papilio dedalus, Cram. i. pl. i. f.1, A, B; Dalm. Monogr. Castn. p. 7, 1 (Casin. ded.) ; Act. Holm.
1824, p. 898; Walker, List Lep. Het. B.M.i. p.16; Boisduval, H. n. Lép. Hét. p. 499.

P. cyparissias, Fabricius, Gen. Ins. Mant. 257; Ent. Syst. i. 1. p. 80; Herbst, pl. 118. f. 1,2; Shaw’s
Nat. Miscell. pl. 574, p. 142 (Castnia, c.) ; Godart, Enc. Méth. ix. 797, Ins. pl. 12. f. 1 (Castnia
cyp.); Gray, Trans. Hnt. Soc. u.. 142.

Hab. Cayenne, Amazonia. In Mus. Hopeiano Oxonie, et Boisduval.

2. CASTNIA GERON. C. alis fusco-griseis, certo situ violaceo viridique nitentibus, supe-
rioribus supra fasciis duabus albis, subtus medio pilis longis barbatis; inferioribus
supra basi discoque barbatis, margine postico flavido maculatis, subtus dense squa-
mosis, duplici serie e maculis albis obsoletis. Exp. alar. antic. unc. 5 lin. 4
(14 centim.).

Castnia geron, Kollar, Lep. Bras. Ann. Wien, i. 217, pl. 13. f. 1; Walker, List Lep. Het. B. M. i. p. 20;
Boisduv. H. n. Lép. Hét. p. 496.

Hab. Brasilia. In Mus. Vindobon.

3. CASTNIA VERAGUANA (Pl. XXX. fig. 1). OC. thorace alisque anticis castaneo-brunneis
his basi czeruleo viridique nitentibus, in medio fascia lata valde angulata, antice
albida, postice brunnescente, punctisque nonnullis ovalibus versus apicem albidis ;
alis posticis chalybeo-nigris, fascia pone medium (e maculis ovalibus composita)
margineque postico aurantiaco-rufis, abdomine nigro. Long. corp. unc. 24. Exp.
alar. antic. unc. 6 lin. 7.

Hab. Veragua (Salvin). In Mus. Hopeiano Oxoniz (olim Saunders).

This very fine new species is allied to C. cacica, Bdv., H.-Sch., but has the fore wings
rather less triangular, and they do not possess the small oblong cell seen in other species,
formed by the connexion of the second and third branches of the subcostal vein by a
small transverse veinlet (see Pl. XXVIII. fig. 9).

4. CASTNIA SCHREIBERSII. C. alis anticis olivaceo-fuscis, medio multo pallidioribus et
luteo-albidis, fascia obliqua intus attenuata e costa, ante medium, ad medium disci
-extensa, ubi attingit maculam magnam rotundam brunneam parum distinctam et
ad apicem ale extensam, angulo apicali interno in maculam alteram maximam
intus dilatatam brunneam; alis posticis nigris, vitta longitudinali pone medium
disci antice dilatata, guttisque nonnullis (8-5) submarginalibus pallide ceruleis.
Long. corp. une. 2-24. Expans. alar. antic. unc. 54 (14°15 centim.).

Castnia Schreibersui, Mikan, Delect. Flore et Faun. Brasiliensis (1820) ; Boisduval, H. n. Lép. Hétér.
p. 500.

C. Latreillti, Godart, Enc. Méth. ix. 797; G. R. Gray, Trans. Ent. Soc. L. ii. p. 142; Walker, List

Lep. Het. B. M. pt. i. p. 17; Perty, Del. An. Art. Brasil. pl. 30. f. 7; Schaufuss, ‘Nunq.
Otiosus,’ p. 8.

GENUS CASTNIA AND SOME ALLIED GROUPS. 169

C. actor, Dalman, Monogr. p. 8, pl. 1. f.1; Act. Holm. 1824, p. 398, pl. 1. fig. 1.
Eupalamnides ctesiphon, Hubner, Exot. Schm., ii.

Obs. Variat presertim mas vitta, vel guttis ceruleis alarum posticarum interdum
obsoletis, ut in figura Dalmanni. ‘Talia vidi in Mus. Parisiensi cum nomine C. Godarti
inscripta (nec C. Godartii, Ménétriés= C. strigata).

Hab. Brasilia, Rio Janeiro, Minas, Pernambuco. In Mus. Britann., Hopeiano
Oxonize, et Boisduval.

5. Castnra cactca. C. alis supra nigro-fuscis, ciliis albis; anticis striga tenui albida
fere recta margini postico fere parallela, punctoque parvo in medio disci interdum
obsoleto: alis posticis fascia denticulata paullo pone medium, ad angulum analem
dilatata, maculisque quinque rotundis submarginalibus aurantiacis (3) vel rufis (2).
Variat interdum guttis duabus parvis pallidis inter fasciam et marginem externum
alarum anticarum. Long. corp. unc. 2. Expans. alar. ant. unc. 53.

Castnia cacica (Boisduval, MS.), H.-Schaffer, Aussereur. Schm., i. f. 143; Boisduval, H. n. Lép. Hétér.
p- 502; Walker, Cat. Lep. Het. B. M. p. 1580.

Mas? C. procera, Boisduval.

Obs. An excellent figure of this fine species is contained in Jones’s unpublished
‘Icones,’ Dandi, pl. 63, from the collection of Mr. Francillon, but without indication of
locality or name.

Hab. Ad ripas fluv. Magdalenze, Colombia, Polochic Valley (Salvin), Bogota; Nica-
ragua (Belt). In Mus. Hopeiano Oxonie, et Boisduval.

6. CasTNIA PROCERA. ‘“ D’un noir brun 4a reflet verdatre. Les ailes supérieures, dont
la frange est blanche, sont traversées, sur leurs tiers postérieur, par une raie blanche
droite, réguliére, ne touchant pas tout-a-fait la céte. Ailes inférieures offrent un
peu au-dela du milieu une bande transverse, assez étroite, rouge, un peu crénelée, et
dilatée 4 langle anal; entre cette bande et l’extrémité il y a, en outre, une série de
quatre gros points de la méme couleur. Différe de la cacica par ses ailes supérieures
un peu plus pointues au sommet, par l’absence de la petite tache blanche dis-
coidale, par ses ailes inférieures dont la bande transversale est moitié plus étroite, et
enfin par le dessous des ailes, qui est entiérement d’un brun uniforme, ainsi que le
corps. Le méme port et la méme taille que la cacica.” (Exp. 13 centim.)

Castnia procera, Boisduval, H. n. Lép. Hétér. p. 5038.

Hab. Guatemala. In Mus. Boisduval.

7. CASTNIA PYLADES. C- alis nigro-fuscis metallico nitentibus, anticis fascia lata fusco-
lutea media oblique ad angulum internum apicalem extensa, marginibus irregulari-
bus fuscoque irrorata alteraque abbreviata ejusdem coloris subapicali irregular,
strigaque nigra undulata apicali; alis posticis fuscis, fascia subapicali utrinque
dentata rufo-fulva, nigro maculata maculis versus angulum analem plus minusve
confluentibus. Femina differt alis anticis inter basin et medium fascia indistincta
subalbida. Long. corp. unc. 12-2. Exp. alar. antic. unc. 43-53-64.

170 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Papilio pylades, Cram. pl. 397. f. A,B; Herbst, Icon. pl. 185. f. 1,2; Shaw, Nat. Miscell. pl. 895 5
Godart, Enc. Méth. ix. 797 (Castnia p.); Dalm. Ins. pl. 12. f. 2; Monogr. p.9; Act. Holm. 1824,
p. 899; Gray, Trans. Ent. Soc. L. ii. p. 142 ; Walker, List Lep. Het. B. M.i. p.17; Boisduval, H.
n. Lép. Hétér. p. 501.
Hab. Cayenne, Para. In Mus. Britann., Hopeiano Oxonice, et Boisduval.

8. CASTNIA ZERYNTHIA. C. alis anticis olivaceo-fuscis medio pallidioribus lutescentibus
basi et apice magis nigricantibus, macula magna cuneata obliqua nigricante e costa
ad medium ale extensa albo marginata vittaque nigricante e medio fere ad basin
marginis interni, maculaque magna trigona costali inter medium et apicem albo bi-
euttata; alis posticis, basi fuscis, medio cum abdomine albis, limbo lato nigro, fulvo
guttato. Long. corp. unc. 24. Expans. alar. ant. unc. 5$ (14 centim.).

Casinia zerynthia, G. R. Gray, Trans. Ent. Soc. Lond. i. p. 144; Walker, List Lep. Het. B.M. 1. p. 18;
Boisduval, H. n. Lép. Hét. p. 506.

C. Langsdorfii, Ménétriés, Enum. Corp. An. Mus. Petrop. pl. xi. f. 1.

Hab. Brasilia, Bolivia. In Mus. Britann., Hopeiano Oxonie, Boisduval.

9. CasTNIA IcARuUS. C. alis anticis fuscis margine interno late griseo, fasciis duabus
parallelis albis obliquis e costa fere ad angulum internum extensis, punctisque
nonnullis interdum fasciam parvam simulantibus versus apicem; alis posticis
leete rubris basi griseis, strigis duabus undulatis pone medium margineque externo
nigris maculisque nonnullis prope medium costz albis et nigris. Long. corp. maris
une. 14, foem. une. 1g. Expans. alar. antic. maris unc. 34, foem. une. 43-5
(10-12 centim.).

Papilio icarus, Cram. i. pl. 18. f. A, B; Shaw’s Mise. pl. 672; Godart. Enc. Méth. ix. p.798 ; Ins. pl. 12.
f. 3 (Castnia ic.) ; Dalman, Monogr. p. 10; Act. Holm. 1824, p. 899; Gray, Trans. Ent. Soe. L. ii.
p- 144; Boisduval, Consp. Lep. Guatemala, p. 75; H.n. Lép. Hétéroc. p. 503; Walker, List Lep.
Het. B. M. i. p. 19; Hiibner, Samml. exot. Schmett. 1. (Urbanus celebris ic.).

Hab. Cayenne, Brasilia, Guatemala, Maranham. In Mus. Britann., Hopeiano Oxoniz,
et Boisduval.

Var. (C. invaria) alis posticis maculis discoidalibus fere obliteratis. “ Fusca, corpus
subtus et abdomen testacea: ale cyaneo viridique nitentes; antice fasciis tribus
hyalinis [albis]; posticee rufse basi albidze, fascia antica [sc. versus costam] maculari
hyalina [alba], fascia breviore margineque postico nigris, hoc maculis 2 vel 3 rufis.”
Expans. alar. ant. unc. 47.

Castnia invaria, Walker, List Lep. Het. B. M. i. p. 23; Boisduval, H. n. Lép. Hét. p. 530.

Hab. Brasilia. In Mus. Britann.

10. CastNIA PAPAGAYA (Pl. XXX. fig. 6). Corpore crasso fulvo-fuscescente, abdominis
lateribus magis rufis; alis anticis ovali-trigonis, in medio saturate fulvescentibus,
basi et margine apicali obscurioribus maculaque media costali fusca bipartita,
medio pallidiore, fasciaque obliqua irregulari fusca inter medium ale et apicem,
ad costam et marginem internum dilatata, in medio angustata et ibi subobliterata,
maculam parvam albam prope costam includente; alis postic’s rotundatis chalybeo-

GENUS CASTNIA AND SOME ALLIED GROUPS. 171

nigris, fasciis tribus transversis e maculis ovalibus compositis, rubris, fimbriaque lute-
scente; antennis rufescentibus; alis subtus rufis, anticis nigro variis. Long. corp.
une. 13. Expans. alar. antic. unc, 33.

Hab. Papagaya (Rogers). In Mus. Hopeiano Oxonie (olim Saunders).

This species is remarkable for its large robust body and short strong wings.

11. Castyra Datmannit (Pl. XXX. fig. 5). C. alis ferrugineis, maris fasciis duabus
obsoletis, foemine fascia alba angusta et punctis hyalinis duobus; posticis rubris,
basi strigis maculisque juxta marginem nigris, maris disco flavo et foeminz albo.
Expans. alar. antic. maris unc. 34, foem. une. 4.

Castnia Dalmannii, Gray, Trans. Ent. Soc. L. ii. 145; Walker, List Lep. Het..B. M. 1. p. 26; nec
Boisduval, H. n. Lép. Hét. p. 512, which = C. hegemon.

Fom. C. Grayi, Boisduval, H. n. Lép. Hét. p. 498.

Hab. Brasilia. In Mus. Britann., Boisduval.

Dr. Boisduval’s description of the wings of the insect to which he has applied the name
of C. Grayi, will be seen from my figure (Tab. XXX. f. 5) and Gray’s description to agree
entirely with that of the female of C. Dalmannii, whilst the description of his C. Dal-
mannit as completely agrees with C. hegemon. His description of C. Grayi is as follows :—
* Ailes supérieures d'un brun un peu ferrugineux, A reflet verdatre; elles ont entre le
milieu et le sommet une raie oblique d’un blanc pur, allant de la céte au bord interne,
précédée en dehors d’un petit point de sa couleur: ailes inférieures d’un brun olivatre
a la base, marquées au milieu d’une large tache blanche transversale, atteignant le bord
abdominal; ensuite le contour est rouge, divisé en arriére par un cordon de taches
noires assez grosses et bien marquées. Un peu plus grande que |’ Japyz.”

Hab. Brasilia. In Mus. Boisduval.

12. Castnta Paunasta. C.- alis anticis olivaceo-fuscis, fascia valde sinuata et irregulari,
grisea, e costa (ultra medium), tune dilatata subbifida et inde versus basin marginis
interni extensa, maculaque magna fere rotundata ejusdem coloris versus angulum
posticum internum, liturisque duabus arcuatis subapicalibus albis; alis posticis
nigris fascia lata fere media alba seu flavescente, striga rubra maculari pone medium
(in limbo late nigro) guttisque submarginalibus albidis. Long. corp. maris unc. 1%;
foeem.une. 13. Expans. alar. antic. maris une. 31, foem. une. 4 (9-12 centim.).

Var. a alis anticis magis nigricantibus, posticis fascia alba venis nigris gracilibus divisa.

Castnia Pallasia, Eschscholtz, in Kotzebue, Reise, iii. p. 217, pl. 6. f. 27; Thon, Archiv, ii. pl. 1. f. 5;
Boisduval, H. n. Lép. Hét. p. 507. :

C. ardalus, Dalman, Monogr. p. 17; Act. Holm. 1824, p. 403; Gray, Trans. Ent. Soc. L. ii. p. 147 ;
Walker, List Lep. Het. B. M. i. p. 24.
C. Brecourti, Godart, Ene. Méth. ix. p. 798; Gray, Trans. Ent. Soc. L. u. 147.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie (Burchell, 29 Dec. 1825, Catombi,
Brazil), Boisduval.

Var. 8. Mas e Papagaya fasciis variis alarum minus dilatatis.

SECOND SERIES.—ZOOLOGY, VOL. I. 2 A

172 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

13. CASTNIA GALINTHIAS. C. alis anticis acuminatis, fuscis, nitidis, lineolis longi-
tudinalibus albis et nigris alternis (in interstitiis yvenarum); alis posticis nigris,
fascia media (venis divisa) seriebus duabus macularum plus minusve distinctarum
albis, margine postico obscure fulvo. Long. corp. fem. une. 13. Expans. alar.
antic. unc. 33.

Castnia galinthias, Hopfter, Neue Schmett. tab. iv. f. 4; Boisduval, H. n. Lép. Het. p. 412.

Hab. Brasilia, Novo Friborgo. In Mus. Hopeiano Oxonie et Berol.

14. CasTNIA HARMODIUS. C. alis fusco-nigricantibus metallico tinctis, anticis fimbria
alba, fascia media obliqua postice attenuata e costa ad angulum internum extensa,
altera sinuata, subapicali minore e maculis quinque formata; alis posticis fascia
intermarginali maculari rufa, ad angulum analem dilatata et striga obliqua nigra ~
divisa. Long. corp. unc. 13. EHxpans. alar. antic. une. 3g—-44.

Obs. Striga parva obliqua fasciam rubram submarginalem alz posticee in duas partes
dividens interdum obliterata est, inde fascia integra evadit.

Papilio harmodius, Cramer, ii. pl. 223. f. C, D; Dalman, Monogr. Act. Holm. 1824, p. 399 (Castnia h.) ;

Gray, Trans. Ent. Soc. L. ii. p. 143; Walker, Cat. Lep. Het. B. M.i. p. 21; Boisduval, H. n. Lép.
Hét. p. 525.

P. syphax, Fabricius, Syst. Ent. p.48; Ent. Syst. ii. 1,p.45; Herbst, Ins. vi. p. 73, pl. 134. f. 1, 2; Jones,
Tcones, 11. pl. 74. f. 1.

P. pirrha, Oliv. Enc. Méth. Ins. pl. 16. f. 2.

Castnia licas, var., Godart, Hnc. Méth. x. p. 797.

Hab. Cayenne, Maranham, Amazonia, Surinam, Para, Demerara. In Mus. Britann.,
Hopeiano Oxonie, Boisduval.

15. CASTNIA ATYMNIUS. OC. alis anticis nigro-fuscis metallico nitidis, fascia media albida
obliqua fere recta ad angulum analem extensa alteraque subapicali valde sinuata e
maculis 5 vel 6 formata (interdum obliteratis) ; alis posticis fuscis extus plus minusve
brunneis, fascia magna alba obliqua media versus costam attenuata, et ad angulum
analem valde dilatata, fimbria partim alba; antennis apice fulvis. Long. corp. une.
13-13. Expans. alar. antic. unc. 33-33.

Var. a maculis subapicalibus albis in alis anticis valde distinctis. (Colombia, Nicaragua,
Bogota.)

Var. 8 fascia media alarum anticarum alba distincta, fascia maculari subapicali fere
obliterata squamis nonnullis pallidioribus tantum distinguenda, vel omnino obliterata
(ut in figura Pertii), alis posticis brunneis albo fasciatis. (Colombia, St. Hstavan,
Venezuela. )

Var. y fascia media et fascia subapicali alarum anticarum fere obsoletis. (Nicaragua,
Santa Martha, Guatemala, Mexico.) .

Castnia atymnius, Dalman, Monogr. p. 12; Act. Holm. 1824, p. 400; Gray, Trans. Ent. Sec. L. un.

142; Walker, List Lep. Het. B. M.i. p. 17 (C. atymnus) ; Boisduval, Cons. Lepid. p. 75; H. n. Lép.
Hét. p. 528.
C. spixii, Perty, Del. An. Art. Bras. pl. 31. f. 3.

GENUS CASTNIA AND SOME ALLIED GROUPS. 173

Obs. C. futilis, Walker (List Lep. Brit. M. pt. vii. 1856, p. 1581), from Nicaragua, is
described from two old faded and rubbed specimens of this species in the British Museum.

Hab. Brasilia, Guatemala, Nicaragua, Honduras, Venezuela, Columbia, Bogota. In
Mus. Britann., Hopeiano Oxoniz, Boisduval.

16. Castnia Licus. C-.alis fusco-nigris metallico nitidis, anticis fascia media recta postice
attenuata ad angulum internum extensa, alteraque subapicali smuata maculari;
alis posticis fascia magna alba obliqua, versus costam attenuata, ad angulum
analem valde dilatata, serieque marginali macularum rubrarum. Long. corp. une.
14-14. Expans. alar. unc. 33-4 (7-11 centim.).

Var. a fascia maculari subapicali alarum anticarum fere obliterata.

Var. 6 alis anticis lutescentibus, basi, apice, marginibus fascixe medie nigris, maculis
alarum posticarum pallide fulvis.

Papilio licus, Fabricius, Ent. Syst. ii. 1, p. 47; Herbst, Ins. pl. 134. f. 38, 4; Drury, Ins. i. pl. 16. £.1, 2;
Merian, Ins. Surimam. pl. 86; Oliv. Enc. Méth. Ins. pl. 12. f. 4; Jones, Icones, iu. pl. 73. f£. 3, 4;
Godart, Enc. Méth. ix. p. 797 (Castnia licus) ; Cramer, iii. pl. 223. f. A, B; Dalman, Monogr.
p. 13; Act. Holm. 1824, (Castnia licus); Gray, Trans. Ent. Soc. L. 1. p. 143; Walker, List
Lep. Het. B. M.i. p. 18; Cuvier, Régne An. ed. Crochard, Ins. pl. 145. f. 1, 2; Boisduval, H. n.
Lép. Hét. p. 526; Chenu, Ency. Pap. 235. f. 402.

Obs. I am inclined to believe that this supposed species is formed of individuals of
C. atymnius, the chief difference consisting in the submarginal rows of red spots on the
hind wings, which, indeed, vary both in number and size in the specimens in the Oxford
Museum. Several faded specimens in the Hopeian Collection, marked C. futilis by Mr.
Walker, only differ from similar faded specimens of C. atymnius (also labelled C. futilis
by Mr. Walker) in having a few slight orange marks near the angle of the hind wings.

Var. a alis posticis fascia brevi alba e disco ad angulum internum extensa margine
aurantiaco (Walker, op. cit. vii. p. 1582).

Hab. America meridionalis, Ecuador, Trinidad, Cayenne, Surinam, Demerara, Para,
Brazil, Ega (Amazonia); mouth of river Una, near Para, 19 Jan. 1830 (Burchell);
Upper Orinoco (larva in Orchidearwm radicibus). In Mus. Britann., Hopeiano Oxonie,
Boisduval.

17. Castyta Botsptvattr. Mas C. alis anticis purpureo-fuscis metallico nitidis, basi,
macula trigona media costali, altera magna trigona inter medium cost et apicem
(guttas tres pellucidas gerente), his maculis intus cum altera magna trigona (e
medio marginis interni ad angulum externum extensa) connexis; alis posticis basi
nigris, in medio macula magna aurantiaco-rufa marginis antici, in strigam tenuem
ad marginem analem extensa, limbo late nigro, rufo plus minusve maculato, maculis
duplici serie congestis. Long. corp. unc. 14. Exp. alar. antic. unc. 23—4.

Foemina differt abdomine albo, macula postica alarum anticarum extus griseo-suffusa,
alisque posticis fascia media lata ad marginem analem extensa maculisque mar-
ginalibus omnibus flavidis.

Castnia Boisduvalii, Becker, MS.; Walker, List Lep. Het. B. M. i. p. 27; Schaufuss, Nung. Otiosus,
p- 9; Boisduval, H. n. Lép. Hét. p. 510.

2A 2

174 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

3. C. Besckei, Ménétriés, Enum. Anim. Mus. Petrop. u. pl. xi. f. 3.
9. C. Herrickii, Boisduval, MS.; Herr.-Schaffer, Samml. aussereur. Schmett. fig. 144; Walker, List Lep.
Het. B. M. p. 1581.
Hab. Brasilia, Bahia, Pernambuco, Papagaya, Parana, Bolivia. In Mus. Britann.,
Hopeiano Oxoniz, Boisduval.

18. CastN1a oRESTES. C. alis anticis plus minusve pallide luteo-fuscis, basi, fascia e
medio coste fere ad angulum internum extensa (ramum curvatum intus pone
medium disci emittente), macula magna irregulari trigona subapicali (guttas tres
indistinctas hyalinas gerente) margineque apicali obscurioribus ; alis posticis basi
eriseis, medio macula magna triangulari, e margine anali fere ad costam extensa,
alba, limbo lato nigro fasciis duabus irregularibus e maculis aurantiacis plus minusve
distinctis compositis notato. Long. corp. unc. 1-1}. Expans. alar. 25-32.

Castnia orestes, Boisd. H. n. Lép. Hét. p. 509.

C. umbrata, Ménétriés, Enum, An. Mus. Petrop. pl. xi. f. 2; Walker, List Lep. Het. B. M.1. p. 26.

Hab. Brasilia, Rio Janeiro, Venezuela. In Mus. Britann., Hopeiano Oxonie,

Boisduval.

19. CastNIA IncA. C. alis anticis fusco, luteo et brunneo variegatis, maculis duabus
pallidioribus costalibus, una media subtrigona, altera semirotundata inter primam et
euttas tres hyalinas subapicales, hujus macule margo posticus niger obliquus, et cum
striga tenui obliqua prope basin marginis interni retro extensa continuus; alis
posticis basi fuscis; medio fulvo-rufis, ultra medium nigris, fascia maculari albida
submarginali. Long. corp. unc. 14. Expans. alar. antic. unc. 383-43 (9-10 centim.).

Var. a alis posticis serie macularum parvarum rotundarum parum distincta, inter fasciam
fulvam et maculas submarginales.

Castnia inca (Boisduval, MS.), Herr.-Schaffer, Samml. aussereur. Schm. f. 488, 489; Walker, List
Lep. Het. B.M. i. p. 24; Hopffer, Neue Schmett. Heft ii. pl. 4. £2; Boisduval, H. n. Léep. Hét.
p- 524.

Hab. Mexico prope Oajaca, Honduras, Venezuela. In Mus. Britann., Hopeiano

Oxoniz, Boisduval.

20, CASTNIA HECATE. (. alis fuscis olivaceo nitidis, fascia ocellari (margini postico fere
parallela) postice dilatata strigaque obliqua alba e medio coste ad medium disci
extensa, ubi fasciz ocellari se conjungit ; alis posticis rufo-fulvis, fascia pone medium
limboque tenui marginali nigris. Long. corp. unc. 17-13. Expans. alar. antic.
unc. 3¢-4¢ (8 centim. ).

Castnia hecate (Boisduval, MS.), Herrich-Schiffer, Samml. aussereur. Schmett. f. 439; Boisduval, H.
n, Lép. Hét. p.531; Walker, List Lep. Het. B. M. p. 1581.

Hab. America meridionalis, insula Haiti. In Mus. Hopeiano Oxonize et Boisduval.

9]. Castnta srricata. C.alis fuscis, metallico nitentibus, albo variis, scil. strigis duabus
longitudinalibus basalibus, fascia submedia, valde angulata et in medio interrupta,
dimidio antico integro, postico e maculis 4 subovalibus formato; inter hanc fasciam

GENUS CASTNIA AND SOME ALLIED GROUPS. 175

et apicem striga obliquae maculis 3-4 formata serieque subapicali macularum, in
medio interrupta et versus apicem interdum fere obliterata; alis posticis nebula
media costali fulvescente, fascia maculari pone medium alteraque submarginali ¢
maculis minoribus formata albis. Long. corp. unc. 1. Expans. al. ant. unc. 23
(6-63 centim.).

Casinia strigata, Walker, List Lep. Het. B. M. i. p. 30; Boisduval, H. n. Lép. Hét. p. 519.

C. Godartui, Ménétriés, Enum, Corp. Anim. Mus. Petrop. pl. 11. fig. 4, p. 130.

Hab. Brasilia, Pernambuco. In Mus. Britann., Hopeiano Oxoni, et Boisduval.

22. CASTNIA DECUSSATA. C. alis fuscis viridi nitentibus, fascia angulata (literam magnam
V supinam simulante) alteraque valde sinuata e maculis 5 composita subapicali,
maculis alteris duabus versus angulum internum positis albis; alis posticis fascia
irregulari pone medium, e maculis 5-7 oblongis composita, strigisque nonnullis fere
obsoletis marginalibus prope angulum analem. Long. corp.unc. 1%. Expans. alar.
ant. unc. 2-3 (6-6 centim.).

Castnia decussata, Godart, Enc. Méth. ix. 799; Gray, Trans. Ent. Soc. Lond. ii. p. 146; Walker, List
Lep. Het. B. M.i.p. 29; Hiibner, Zutrage, iii. f. 689, 640; Boisduval, H. n. Lép. Hét. p. 519.

A figure of this species is contained in Jones’s ‘ Icones,’ pl. 89. f. 1, without any
reference or locality.

Var. a alis magis maculatis, = C. melaleuca, Boisduval, MS. in Mus. Parisiensi.
Hab. Brasilia. In Mus. Britann., Hopeiano Oxoniz, et Boisduval.

23. CASTNIA PAPILIONARIS. (Tab. XX XI. fig. 3.) Alis anticis obscure fuscis metallico
nitidis, puncto parvo ad apicem cellule discoidalis, fasciaque tenui recta inter
medium et apicem ale pallide flavis, fimbria alba; alis posticis nigris, fascia pone
medium, versus angulum ani attenuata, cerulescente. Long. corp. unc. 13. Exp.
alar. antic. unc. 4.

Castnia papilionaris, Walker, List Lep. Het. B. M. part xxxi. p. 42; Boisduval, H. n. Lép. Hét. p. 500.

Obs. The ochraceous colour of the body described by Mr. Walker is simply the result
of the rubbed condition of the specimen in the British Museum.
Hab. Bogota. In Mus. Britann., et Hopeiano Oxonie.

24. CASTNIA SATRAPES. (Tab. XXXI. fig. 4.) Alis anticis viridi-fuscis, macula magna
marginis interni (guttam fuscam arcuatam includente) fasciaque lata subapicali
albidis fusco irroratis, squamisque nonnullis subapicalibus albis; alis posticis
aurantiacis, venis radiantibus limboque nigris, hoc albido maculato. Long. corp.
une. 1 lin. 5. Exp. alar. antic. une. 4 lin. 2 (74 centim.).

Var. a maculis pallidis alarum anticarum intus connexis, ut in figura Kollari.

Var. 8 maculis pallidis alarum anticarum separatis, alis posticis aurantiacis nigro
limbatis.

Var.-y maculis pallidis alarum anticarum separatis, alis posticis flavidis nigro limbutis,
limbo serie macularum rubrarum intus notato. (Tab. XX XI. fig. 5.)

176 MR. J. 0. WESTWOOD ON THE LEPIDOPTEROUS

Casima satrapes, Kollar, Lep. Bras. Ann. Wien. Mus. ii. 216, pl.12. £.3; Walker, List Lep. Het. B.
M.1i. p. 33; Boisduval, H. n. Lép. Hét. p. 5388.

Hab. Brasilia. In Mus. Hopeiano Oxoniz et Boisduval.

25. CASTNIA PALATINA. C. alis anticis rufo-brunneis medio pallidioribus, macula magna
subovali e medio coste versus medium disci oblique extensa, gutta parva hyalina
subapicali alis posticis basi fuscis, medio flavidis versus costam aurantiacis, limbo
nigro striga submarginali flavo-maculari. Long. corp. unc. 13. Expans. alar. ant.
une. 33 (8 centim.).

Papilio palatinus, Cramer, ii. pl. 159. f. B,C ; Godart, Ene. Méth. ix. p. 799 (Castnia palatina) ; Dalman,
Monog. p. 18; Act. Holm. 1824, p. 403; Gray, Trans. Ent. Soc. Lond. ii. p. 147; Walker,
List. Lep. Het. B. M.i. p. 25; Boisduval, H. n. Lép. Hét. p. 513.

Hab. Cayenne, Surinam, Demerara, Para. In Mus. Britann., Hopeiano Oxoniz,

Boisduval.

26. CASTNIATHERAPON. Alis elongatis, anticis supra fulvis, striga tenui nigricante obli-
quae basi marginis postici fere ad apicem ducta, macula media disci fusca versus
basin vitta nigricante marginata, margine interno apiceque fuscescentibus, hoc
obscure biguttato ; alis posticis rufo-aurantiacis, limbo apicali nigro, hoc guttis septem
albidis notato. Long. corp. lin. 18. Expans. alar. ant. unc. 3 lin. 4 (74 centim.).

Castnia therapon, Kollar, Lep. Bras. Ann. Wien. Mus. ii. 218, pl. 13. f. 3; Walker, List Lep. Het. B. M.

i. 23; Boisduval, H. n. Lép. Hét. p. 540 (Orthia th.).
C. paradoxa (Boisduval, MS.), Herr.-Schaffer, Samml. aussereur. Schmett. f. 16, 17.

Hab. Brasilia. In Mus. Hopeiano Oxonie et Boisduval.

27. Casrnta cuirarcHa. (Tab. XX XI. fig.1, 2.) Foem. alis anticis magnis obovalibus
apice subacutis, stramineo-albis, brunneo-fusco variegatis, maculis in medio e
costa literam h magnam reversam valde obliquam rude simulantibus, striolaque
tenui denticulata fere ex apice versus basin marginis interni extensa, apiceque
ipso fuscis, hoc punctis tribus albis (intermedio multo majore) notato, limbo etiam
fusco; alis posticis leete fulvis basi magis rufis, fascia rufescente inter medium et
apicem alze seriem macularum nigrarum includente limboque denticulato nigro ;
corpore toto fulvo thorace saturatiore, subtus pallidiore.

Mas multo minor, alis anticis saturate fulvis notulis foeminze multo minus distinctis ; alis
posticis basi paullo obscurioribus, aliter ut in foemina notatis. Long. corp. maris
fere unc. 13, fem. unc. 13. Expans. alar. antic. maris unc. 33, foem. une. 5.

This species is closely allied to C. inca, the figures of which hitherto published appear
to represent female specimens. The female of the present species has the fore wings
much more strongly variegated ; and the broad black border of the hind wings in C-. inca,
with its row of pale straw-coloured spots, is replaced by a richly coloured and marked
margin in C. elitarcha. The cell and branches of the postcostal vein of the fore wing
are arranged as in V. Latreillii, the third and fourth branches separating at a consider-
able distance beyond the cell.

Hab. Panama (Salvin), Nicaragua (Belt). In Mus. Hopeiano Oxonie.

GENUS CASTNIA AND SOME ALLIED GROUPS. 177

28. CASTNIA PHALARIS (Fab. nec Godart). (Tab. XXX. fig. 3.) (C.alis anticis fuscis basi
et margine externo obscurioribus, disco nonnihil pallidiore, fascia lata obliqua nigri-
cante e medio costz ultra medium disci extensa, inde in plagam magnam rotundam
angulum posticum apicalem occupantem (medio pallidiorem) dilatata, maculaque
magna trigona subapicali (guttas tres subhyalinas includente) obscura; alis posticis
nigris limbo lato aurantiaco-fulvo, intus irregulariter undulato, striga submarginali
fusca margineque ipso interdum nigro dentato. Long. corp. lin. 10-15. Expans.
alar. antic. une. 24-84.

Papilio phalaris, Fabricius, Ent. Syst. ii. 1, p. 45; Jones, Icones, iti. pl. 75. f. 3, 4; Donovan,
Naturalist’s Repos. ii. pl. 47. f.1, 2 (nee Godart, nec Boisduval, =mygdon, Dalm.) ; Dalman, Monogr.
Castnia, p. 22; Act. Holm. 1824, p. 405; Gray, Trans. Ent. Soc. Lond. ii. 145; Walker, List Lep-
Het. 1. p. 19.

Castnia subvaria, Walker, op. cit. p. 25.

C. musarum, Boisduval, MS. in Mus. Parisiensi.

Var. a striga submarginali fusca alarum posticarum fere obliterata, margine ipso fulvo.

C. dionea, Hopfier, Neue Schm. pl. 5. f. 3; Boisduval, H. n. Lép. Hét. p. 523.

Hab. Minas Geraes Brasiliec. In Mus. Britann., Hopeiano Oxoniz, Berolini,
Boisduval.

C. phalaris was described by Fabricius from the cabinet and Icones of Mr. Jones,
whose figures were published by Donovan in his ‘ Naturalist’s Repository.’ The Fabri-
cian description is simply “alee supra fusce, anticee immaculate, postice fasciis duabus
fere marginalibus e maculis exalbidis.” Godart applied the description of Fabricius to
the insects subsequently described and figured by Dalman under the name of P. mygdon,
with which, indeed, the two rows of spots on the outer margin of the hind wings
described by Fabricius agree both in form and colour.

29. CASTNIA EUDESMIA. C. alis anticis glauco-fuscis, costa tenui margineque interno
late albo pulverosis, fasciisque duabus postice acuminatis e costa ultra medium
disci extensis; alis posticis nigris, fasciis duabus macularibus inter medium et mar-
ginem posticum sanguineis albo irroratis, margine ipso anguloque anali griseis, hoc
fulvo squamoso. Long. corp. une.13. Hxpans. alar. ant. unc. 34 (9-11 centim.).

Castnia eudesmia, Gray, Trans. Ent. Soc. L. ii. p. 145 ; Herrich-Schiffer, Samml. aussereur. Schm. f. 140,

141; Gay, Historia fisica de Chile, Lep. pl. 5. f. 8, p. 47; Walker, List Lep. Het. B. M.i. p. 19;
Boisduval, H. n. Lép. Hét. p. 517.

Hab. Chili; Brasilia? In Mus. Britann., Hopeiano Oxoniz, Boisduval.

30. CasTNra cocHRuUS. C.alis nigris; anticis in medio subpallidioribus, venis nisris,
striga obliqua tenui nigra pone medium disci versus apicem alee extensa, vittaque
subalbida postica adjecta; alis posticis nigris macula media alba transverse ovata
in sex partes divisa, venis guttisque nonnullis plus minusve obsoletis versus angulum
externum albis; abdominis segmentis 5 apicalibus rubris, nigro et albo marginatis.
Long. corp. unc. 13. Expans. alar. antic. unc. 4-42.

Papilio cochrus, Fabricius, Ent. Syst. 11. 1, p. 42; Donovan, Nat. Repos. pl. 156; Jones, Icones, iii.

pl. 77. f.1; Godart, Enc. Méth. ix. p. 798 (Castnia c.) ; Gray, Trans. Ent. Soc. Lond. ii. p. 147;
Walker, List Lep. Het. B. M.i. p. 20; Boisduval, H. n. Lép. Hét. p. 517.

178 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Castma maris, Dalm. Monogr. p. 16; Act Holm. 1824, p. 401, pl. 1. f. 4.
Prometheus casmilus, Hiibner, Samm]. exot. Schmett. ii. f. 1, 2.
Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie, Boisduval, &c.

31. CasTNIA RUTILA. C. alis anticis testaceo-fulvis, macula obscuriore in medio cost,
intus bifida, ramis curvatis, ad venam mediam extensis, serieque irregulari punc-
torum obscuriorum inter medium et marginem apicalem, scil. 2 versus apicem, supe-
riore majore nigricante guttam albam includente, 3 in medio majoribus, et. 2 versus
angulum internum multo minoribus, serieque marginali macularum cuneiformium
obscuriorum; alis posticis testaceo-rufis, fascia submarginali e maculis 8 nigris

formata, margineque ipso tenui nigro intus undulato. Exp. alar. antic. cire. une. 5.
Castnia rutila, Felder, Reise Novara, pl. 79. f.1; Boisduy. H. n. Lép. Hét. p. 496.
Hab. Amazonia. in Mus. Felder.

32. CasTnta Fasrici. C-.alis anticis luteo-fulvis (“ ferrugineis ”’), nebulis obscurioribus
fuscescentibus, scil. macula seu circulo juxta medium coste, fasciaque lata inter
medium et marginem apicalem, cum margine parallela, versus costam et marginem
internum dilatata, hujus fasciz marginibus undulatis et presertim postice obscu-
rioribus; alis posticis rufis (‘‘ pallide testaceis’’), fasciis tribus nigris, scil. prima ante
medium abbreviata, secunda mediana, tertia pone medium, his duabus crenatis,
serieque submarginali macularum sex nigrarum ovalium notatis. Expans. alar.
une. 4¢ (54 Gray, an recte P).

Castmia Fabricii, Swainson, Zool. Ill. iii. pl. 149; Thon, Ent. Archiv, 1829, f. 1236; Gray, Trans. Ent.
Soc. L. ui. p. 144; Walker, List Lep. Het. B. M.i. 19 (nec C. Fabricii, Godart, Ene. Méth. ix. 800,
=C. Marcel Serresi feem., nec C. Fabricii, Boisduval, H. n. Lép. Hét. p.539,=C. chremes.

Hab. Brasilia (“ diamond district ’’).

33. CASTNIA JaAPYX. C. alis rufo-brunneis, fascia obliqua pone medium obscuriore, in
mare macula parva pallidiore ad apicem cellule discoidalis punctoque hyalino inter
cellulam et marginem externum, in feemina maculis 5 rotundis inter se distantibus
discoidalibus flavescentibus ; alis posticis fulvis extus magis testaceis, basi fusces-
centibus, in foemina fasciis duabus submarginalibus e maculis parvis nigricantibus
formatis, his maculis in maribus fere obliteratis (nonnullis versus angulum ex-
ternum tantum apparentibus). Long. corp. unc. 14-1. Expans. alar. ¢ une. 33
(10 centim.), @ une. 44 (114).

Athis japyx, Hubner, Samml. exot. Schmetterl. ii. 362, f. 1, 2, ¢ ; Boisduval, H. n. Lép. Hét. p. 497.

Castnia Kirstenti, Thon, Archiv, ii. p. 7, f. 6-10, g¢ ; Walker, List Lep. Het. B. M. p. 33.

C. Fonscolombei, Godart, Enc. Méth. ix. p. 799; Gray, Synops. Trans. Ent. Soc. ii. p. 144; Walker, List
Lep. Het. B. M. i. p. 20.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie, Boisduval, &c.

34, CASTNIA HEGEMON. C. alis anticis rufo-badiis, macula media costali fasciaque
obliqua pone medium ale (ad medium marginis interni et prope apicem dilatata
ef male determinata) magis brunneis; alis posticis basi nigricantibus, disco flaves-

GENUS CASTNIA AND SOME ALLIED GROUPS. 179

centi, margine anali albo, limbo lato rubro, strigis duabus undulatis nigris versus
angulum analem latioribus et ibi coalitis. Fcemina (sec. sententiam Boisduvalii)
differt fasciis alarum anticarum magis distinctis, postica seepe interrupta ; alis posticis
disco niveo, strigis nigris marginis postici magis determinatis. Long. corp. unc. 14.
Expans. alar. ant. unc. 33.
Castnia Hegemon, Kollar, Lep. Brasil. Ann. Wien, 1. 217, pl. 13. f. 2; Walker, List Lep. Het. B. M. i.
p- 26; Boisduval, H. n. Lép. Hét. p. 514.
C. Dalmannii, Boisduval, H. n. Lép. Hét. p. 512 (nec Gray =Grayi, Bdv. p. 498).
Dr. Boisduval has mistaken this very distinct species for C. Dalmannii of Gray, and
has applied the name of Grayi to the latter insect.
Hab. Brasilia, Rio Janeiro. In Mus. Britann., et Hopeiano Oxoniz.

35. CasTNIA creLA. “Ailes supérieures d’un gris-blanchatre chatoyant en vert ou en
violet, avec la base, le bord terminal, une grosse tache sous-costale et une bande trans-
versale de couleur brune; la bande transversale située sur le tiers postérieur est
sinuée; elle se dilate sur la céte et sur le bord interne, de sorte qu’elle est un peu
étranglée dans son milieu. Ailes inférieures d’une belle couleur orange, avec un arc
discoidal noir; leur tiers postérieur est marqué d’une rangée de taches un peu
oblongues, un peu plus pales que le fond, séparées par des nervures noires et ren-
fermées entre deux lignes ondulées d’un noir foncé.” Expans. alar. antic. 8 centim.

Casinia ciela (Boisduval, MS.), Herr.-Schaffer, Samml. aussereur. Schmett. figs. 486, 487; Boisduval,
H. un. Lép. Hét. p. 532; Walker, List Lep. Het. B. M. pt. xxxi. p. 43.

Hab. Bahia, Penambuco. In Mus. Boisduval.

36. CastNIA areus. “Ailes supérieures pointues au sommet, d’une teinte brune
obscure, qui oblitére en partie des bandes d’une couleur plus foncée, disposées comme
dans les espéces précédentes (Phalaris et Mygdon) mais beaucoup moins distinctes ;
cependant on voit toujours dans la fissure de la bande bifide trois petites taches
blanches plus ou moins marquées, un peu confluentes. Ailes inférieures noires, avec
trois rangées paralleles de taches mal alignées, dont celles des deux premiéres séries
sont blanches, plus ou moins cerclées de roux vif et celles du bord marginal, qui
sont presque pontiformes, d’un fauve orangé ainsi que la frange de langle anal:
chez la femelle, on voit sur les secondes ailes une tache fauve entre le dise et la
base.” Expans. alar. antic. 12 centim.

Castnia argus, Boisduval, H.n. Lép. Hét. p. 522.

_ This species, which I examined in Dr. Boisduval’s collection, appears to me to be
scarcely different from C. mygdon, being only of a larger size, and with a few more spots
towards the base of the hind wings.

Hab. Brasilia. In Mus. Boisduval.

37. CASTNIA MYGDON. C. alis anticis olivaceo-fuscis viridi nitentibus ; basi, faseia irre-
gulari obliqua e medio costz ad angulum internum extensa (ramum internum paullo
‘pone medium emittente ad medium marginis interni currentem), maculaque magna

SECOND SERIES.—ZOOLOGY, VOL. I. | 2B

180 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

costali subapicali (guttas tres subhyalinas includente) margineque apicali obscurio-
ribus ; alis posticis nigricantibus, inter medium et marginem posticum fasciis duabus
irregularibus e maculis subovalibus albidis aut flavidis formatis maculisque nonnullis
plus minusve obscurioribus versus medium disci positis. Long. corp. unc. 14. Exp.
alar. antic. unc. 25-3 (8-12 centim.).

Castnia mygdon, Dalm. Mon. p. 20, pl. 1. f. 2; Act. Holm. 1824. p. 403; Gray, Trans. Ent. Soc. ii.
p- 145; Walker, List Lep. Het. B. M. 1. p. 28.

C. phalaris, Godart, Enc. Méth. ix. 799 (nec Fabricius, &c.); Dalman, Monogr. Castn. p.22; Act, Holm.
1824, p.405; Gray, Trans. Ent. Soc. Lond. 11. p. 145; Boisduval, H. n. Lép, Hét. p. 521.

Sympalamides mimon, Hiibner, Samml. exot. Schmett. figs. 1, 2.

Hab. Brasilia, Parana, Papagaya. In Mus. Britann., Hopeiano Oxoniz, Boisduval.

38. CASTNIA CHELONE, “ C. alis anticis maris cinereis, foeminze fusco-cinereis, fasciis
duabus, maris murinis obsoletis, foeminze albis ; posticis aterrimis, cyaneo-micantibus,
fascia sinuato-angulata, fusca albido-marginata.”’ Exp. alar. antic. ¢ une. 2 lin. 9
(7 centim.), $ 3” 5” (9 centim.).

Castnia chelone, Hopffer, Neue Schmett. Heft ii. tab. iv. figs. 1, 2; Boisduval, H.n. Lép. Hét. p. 530.

Hab. Mexico. In Mus. Berol.

39. CastniA MENneErrigst. “Ailes supérieures pointues au sommet, d’un brun noir assez
obscur, avec deux bandes transverses d’un noir plus foncé, dont l’antérieure bifide
et la postérieure sinuée ; outre cela, ces deux bandes paraissent réunies par un rameau
latéral ; le long de la céte on voit aussi deux taches plus pales que la teinte générale.
Ailes inférieures noires, marquées vers leur milieu d'une grande tache jaune trans-
versale, sinuée en avant, suivie en arriére, du cété de ’angle externe, de quatre 4
cing points orangés, alignés et d’un autre point semblable rejeté en dehors. La
goutticre abdominale est blanche.”

“A peu pres de la taille de la femelle de C. Boisduvalii, dont elle a le port.”

Castnia Menetriesi, Boisduval, H. n. Lép. Heét. p. 511.

Hab. Brasilia. In Mus. Boisduval.

49, Castnia Humpoupti. “ Ailes supérieures d’un brun noirdtre, avec un reflet olivatre
tres-prononcé; elles sont traversées du milieu de la céte a langle anal, par une
bande blanche, atténuée inférieurement; elles ont, en outre, du coté du sommet,
une bande courte, flexueuse, maculaire, bien nette, composée de cing taches un peu
oblongues. Ailes inférieures d’un brun roux & leur extrémité, traversées par une
bande d’un blanc pur, sinué en dedans et s’élargissant insensiblement, pour arriver
4 Vangle anal; leur frange est entiérement blanche. Un peu plus grande que
V Atymnius, avec les ailes plus pointues au sommet.”

Castnia Humboldti, Boisduyal, H. n. Lép. Hét. p. 528.

Obs. Fascia 5-macularis subapicalis alarum anticarum, in individuo Boisduyalliano
guttas duas subcostales albas supra gerit; heec fascia in individuis Muszei Gallici e ma-
culis fere obsoletis constat.

Hab. Nova Granada. In Mus. Parisiensi et Boisduval.

GENUS CdASTNIA AND SOME ALLIED GROUPS. 181

41. Castnra sazasta. “Ailes supérieures d’un brun noirdtre, 4 reflet verdAtre: elles
sont marquées d’une bande linéaire trés-étroite, ou plutét d’une raie enfumée, allant
du milieu de la cdte a angle interne; ou voit, en outre, vers le sommet, mais trés-
faiblement, l’empreinte de la bande sinuée du dessous. Ailes inférieures noires vers
la base, avec un reflet vert, brunes 4 l’extrémité a reflet rougeatre, elles sont tra-
versées, comme dans les espéces du méme groupe, par une bande d’un blane pur,
mais & reflet violatre, légérement sinuée en dedans, et s’élargissant insensiblement
pour gagner l’angle anal. Frange des quatre ailes entiérement noire.”

De la taille de ? Atymnius avec les ailes plus pointues au sommet.

Castnia salasia, Boisduval, H.n. Lép. Hét. p. 529.

Hab. Mexico. In Mus. Boisduval.

42. CASTNIA EVALTHE. - C. alis latis, nigro-fuscis, anticis fascia media obliqua ad angu-

Tum analem extensa (ibique attenuata) alteraque subapicali tenuiore (interdum obso-
leta) albis aut flavis; posticis fascia media maculari vel macula magna discoidali
flava (interdum externe rufo late marginato) fasciaque maculari submarginali
anguloque anali late rufis. Expans. alar. ant. unc. 24-22 (7-10 centim.).

Var. a alis anticis bifasciatis, alis posticis fascia media submaculari flava, angulo anali
et maculis 5 vel 6 submarginalibus rufis. (Cramer, pl. 17. f. E, F.) Surinam;
Keuador. In Mus. Hopeiano.

Var. 8 maculis discoidalibus in alis posticis in maculam integram majorem coalitis.
(C. euphrosyne, Perty, Delect. pl. 31. f.1; Walker, List Lep. Het. B. M. i. 15.)
Brasilia et Demerara. In Mus. Hopeiano.

Var. y differt e var. 8 macula discoidali flava in alis posticis extus rufo late marginata,
colore rufo cum maculis submarginalibus plus minusve coalito. (Dalman, tab. i.
f. 3.)

Var. 6 similis var. a, at fascia subapicali in alis anticis obliterata maculisque fascize
medize posticarum angulo externo proximis fere obsoletis.. (C. Viryi, Boisduval,
p. 515, pl. 11. fig. 8.) Mexico et Nicaragua. In Mus. Hopeiano. Sore

Papilio evalthe, Fabricius, Syst. Ent. p. 480 ; Ent. Syst.iii.1, p.45; Donovan, N. Hist. Ins. India, Lep. pl.1;
Herbst, Icon. pl. 137. f. 1, 2; Boisduval, H. n. Lép. Hét. p. 514 (Castnia e.) ; Dalman, Monogr.
p- 14, tab. i. f. 3; Act. Holm. 1824, p. 401; Godart, Enc. Méth. ix. p. 798; Gray, Trans. Ent.

Soc. L. u. p. 143; Walker, List Lep. Het. B. M. i. p. 21.
P. dardanus, Cramer, i. pl. 17. f. E, F.

C. euphrosyne, Perty ut supra; Gray, op. cit. p. 144.
E. Viryi, Boisduval ut supra.
Hab. Brasilia, Para, Cayenne, Mexico, Ecuador. In Mus. Britann., Hopeiano Oxonize
(individua 8), Boisduval. .

43. CASTNIA AMycuS. C. alis anticis nigro-fuscis pallidius variegatis, squamis magnis
griseo-albis indutis, puncto seu fasciola subobsoleta ad apicem cellule discoidalis
fasciaque plus minusve obsoleta alba vel albida pone medium cost ad angulum
analem extensa margine interno dentato, fasciolaque subobsoleta abbreviata sub-
apicali, margine postico ante medium fasciculo elongato squamarum instructo ; alis

2B2

182 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

posticis nigris, medio disci ferrugineo fasciaque pone medium e maculis parvis plus
minusve obsoletis maculisque nonnullis submarginalibus rufis. Long. corp. lin.
91-14. Exp. alar. antic. unc. 13-3.

- Variat disco alarum posticarum omnino nigro, maculisque rufis multo minoribus fere

obsoletis.

Papilio amycus, Cram. iii. pl. 227, D, E.; Dalman, Monogr. p. 19 (Castnia a.); Act. Holm. 1824, p. 408 ;
G. R. Gray, Trans. Ent. Soc. L. ii. p. 145; Godart, Enc. Méth. ix. p. 798; Walker, List Lep.
Het. B. M.i. p. 27; Boisduval, H.n. Lép. Hét. p. 504.

Obs. As this species differs in the greater or less amount of red markings on the hind
wings, [am much inclined to consider the C. meditrina of Hopfier to be an extreme
variety in which the red has entirely disappeared. The less acute apex of the fore
wings relied upon by Boisduval appears to me to be the sexual (female) character.

Hab. Surinam, Brazil, Papagaya. In Mus. Britann., Hopeiano Oxonie (7 individua),
Boisduval.

44. CASTNIA MEDITRINA. “ C. alis anticis virescenti-nigris, fascia undulata obliqua gut-
tisque ad costam tribus albis ; posticis aterrimis cyaneo micantibus.” Expans. alar.
ant. une. 2 lin. 5.

The hind margin of the fore wings has the dilated patch of elongated scales noticed
in the description of the preceding, and to which neither the description of Hopffer
nor that of Boisduval allude.

Castnia meditrina, Hopfier, Neue Schmett. Heft i. tab. v. f. 4; Boisduval, H. n. Lép. Hét. p. 505.

An var. C. amycus ?
Heb. Brasilia. in Mus. Berolini.

45. Castnta HvBNERI. Calis anticis fulvo-fuscescentibus, fascia maculari plus minusve
continua e medio coste ad angulum internum extensa alteraque parallela e ma-
culis sex magis distinctis inter medium et apicem albis; alis posticis basi mar-
gineque lato abdominali fuscis, dimidio costali rufo-fulvo; dimidio externo nigro,
maculis 7 cuneiformibus pone medium irregulariter dispositis, serieque submarginali
macularum rufo-fulvarum; fimbria lata pallide fusca.

Casinia Hiibneri (Boisduval, MS.), Latreille in Cuvier, R. An. 2d ed. iii. pl. 20. f. 2, p. 489; Gray,
Trans. Ent. Soc. Lond. i. p. 146; Walker, List Lep. Het. B. M.i. p. 30; Boisduval, H. n. Lép.
Hét. p. 516.

Castnia Sternbergii, Kollar, Lep. Bras. Ann. Wien, 1. 216, pl. 12. f. 4.

Synemon Sternbergii, Mikan, Delect. Fl. et Fr. Brasil. ; Schaufuss, Nunquam Otiosus, p. 11.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxonie, Boisduval.

46. CASTNIA JUTURNA. C. alis anticis fuscis, basi et margine interno griseo-squamosis,
fasciis 4 albidis, duabus anticis obliquis, e costa, marginem posticum et externum
haud attingentibus, duabus posticis brevioribus magis transversis, 3ta e maculis
obcuneatis composita, 4ta venis rufis in maculas divisa; alis posticis basi fuscis,
raacula magna nigra sanguineo cincta, medio fascia alba valde sinuata sanguinco

GENUS C4STNIA AND SOME ALLIED GROUPS. 183

extus marginata; dimidio externo alarum nigro, fascia maculari subapicali san-
guinea, maculis versus angulum externum albo irroratis. Long. corp. unc. 13. Ex-
pans. alar. antic. unc. 3/4.

Castnia juturna, Hopffer, Neue Schmett. pl. iv. f.3; Boisduval, H. n. Lép. ee. p. 508.

Castnia lorze, Boisduval, MS. olim.

Hab. Brasilia (Minas). In Mus. Hopeiano Oxoniz, Boisduval.

47, CASTNIA CRONIS. C. alis anticis nigris, costze basi maculisque thoracis rufis vel
albis, plaga magna discoidali postice dilatata, alteraque subcuneata in medio coste,
fascia abbreviata obliqua e maculis tribus composita pone medium, altera e maculis 9
ovalibus submarginalibus albis, imargine ipso albo guttato ; alis posticis abdomineque
albidis, margine apicali tenui nigro, intus crenulato. Expans. alar. antic. unc. 3-34.

Papilio cronis, Cramer, i. pl. 60. f. C. ??, pl. 178. f. A; Oliv. Enc. Méth. Ins. pl. 12. f.7; Dalman, Mo-

nogr. Castn. p. 23 (Castnia cr.) ; Act. Holm. 1824, p. 504; Godart, Enc. Méth. ix. 801; Gray,
Trans. Ent. Soe. L. ii. p. 146; Walker, List Lep. Het. B. M.i. p. 29. n. 34; Boisduval, H. n. Lép.

Hét. p. 534.
Obs. M. Boisduval, H. n. Lép. Hét. p. 534, considers that the figure given by Cramer,
i. pl. 60. f. C, as the male of C. cronis (Pieris cronissa, Godart, Enc. M. ix. Suppl. p. 814;
Pieris cronis, Boisduval H. n. Lép. i. p. 548) is manufactured out of the head, abdomen,
and hind wings of a Pieris attached to a mutilated thorax and fore wings of a Q. cronis.

Hab. Surinam. In Mus. Britann., Loddige et Herrich-Schiffer (valde mutilata).

48. CasTNIA cronipDA. “ Ailes supérieures noires, avec des taches blanches disposées
a peu prés comme dans la cronis, savoir, une trés-grande longitudinale sur le milieu,
une médiocrement grande sur la cdte et les autres plus petites 4 l’extrémité;
ces derniéres sont alignées au nombre de quatre ou cing seulement, et sont pré-
cédées vers le sommet de trois autres taches blanches allongées, il y a, en outre, sur
les nervures, une série de petits points blancs, trés-peu indiqués; Torigine de la
céte est rouge comme dans la cronis. Ailes inférieures d’un blane un peu jaunatre,
avec une large bordure noire, divisée par un cordon de gros points d’un blanc un
peu jaunatre. Le corselet est noir, marqué de deux points rouges, avec le collier
blane interrompu; l’abdomen est d’un blanc jaundtre avec l’extrémité d’un jaune
d’ocre.”’

“La femelle ne differe du male que par sa taille plus grande et par la bordure des ‘ailes
inférieures plus large avec des points plus gros.”

- Expans. alar. antic. g 8}, 9 103 centim.

Castnia cronida, Herr.-Schafler, Samml. aussereur. Schm. fig. 142; Boisduval, H. n. Lép. Hét. p- 535 2
Walker, List Lep. Het. p. 1581.

Except in the less complete development of the subapical row of spots on the fore
wings, and the wider black margin of the hind wings, this supposed species is evidently
identical with C. cronis, which appears, from Gray’s description of Mr. Loddige’s spe-
cimen, to be subject to a certain amount of variation.

Hab. Guyane francaise. In Mus. Stuttgard et Boisduval.

184 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

49. CASTNIA LicoipEs. “Elle a tout-a-fait le port et la taille de lespdce précédente
(Zicus) dont elle est trés-voisin, mais la massue de ses antennes est jaune. Ses ailes
supérieures sont de méme d’un brun-noir chatoyant en vert, avec une bande linéaire
blanche dentée en dehors, allant du milieu de la céte 4 l’angle anal. Il n’y a de bande
sinuée & l’extrémité; la frange est blanche. Les ailes inférieures sont d’un noir-brun,
traversées dans leur milieu par une bande blanche a reflet un peu violet, dentée en
dehors, s’élargissant insensiblement pour arriver 4 langle anal; le bord terminal est
divisé par un cordon de sept taches rouges, dont l’anale est double et plus petite.”

Castnia licoides, Boisduval, H. n. Lép. Hét. p. 527.
Urbanus licus, Hiibner, Samml. exot. Schmett. v. i. Celeb. a, fig. 1. 2.

Hab. Brasilia, Santa Catherina. In Mus. Boisduval.

50. CASTNIA INORNATA. “ Ferrugineo-fusca, subtus cervina, abdominis dorso nigricante
maculis luteis ornato, alis anticis fasciis duabus fuscis obliquis indeterminatis, ma-
cula reniformi obscure cinerea fusco marginata.”’

«Female ferruginous-brown, fawn-colour beneath. Frontal tuft porrect lanceolate,
extending rather beyond the head. Antenne subclavate; tips pale cinereous. Abdomen
blackish above, shorter than the interior border of the hind wings, each segment with a
transverse luteous spot. Fore wings with two irregular oblique brown bands, which are
diffuse on the outer side, and have between them a reniform dark cinereous mark, which
is bordered with brown; underside with two irregular dark brown bands beyond the
middle, the first composed of patches, the second much abbreviated hindwards, marginal
space fawn-colour. Hind wings with a fringe of very long hairs, which extends across the
disk from near the interior border. Hind wings beneath with an irregular abbreviated
ferruginous line across the disk, including at its fore end a whitish dot.” Length of the
body 10 lines, of the wings 31 lines.

Hab. ? In Coll. T. Chapman, Glasgow.

Castnia inornata, Walker, Cat. Lep. Heteroc. p. 106.

This species belongs to the group of which Chremes is the type.

51. CastNIA PENELOPE. ‘ Dem Icarus in Form und Zeichnung ganz ahnlich, die Ober-
seite aber in allem blasser, die Zcichnung auf den Hinterfliigeln braun statt schwarz,
die fussere Querbinde der Oberfliigel sowie weissen Tropfen der Unterfitigel fast:
verloschen : auf der Unterseite der Oberfliigel alle drei Binden bestimmt ausgepragt,
im Uebrigen opalisirend wie Zcarus und veilleicht nur Varietét.” Lat. 78 mm.

Hab. Brasilien.
Castnia penelope, Schaufuss, Nunq. Otios. p. 9.

52. CASTNIA ALBOFASCIATA. ‘“ Grosser als mimon, oben ebenso gezeichnet, die Oberfliigel
jedoch heiderseits der braunen Querbinden mit weisser, schmaler Kante eingefasst
und mit verwischtem Weiss ausgefillt. Unterfliigel mit drei weissen Flecklagen,
dieselben mit Schwarz und Zimmetbraun eingefasst, ahnlich wie bei Phalaris ; doch

GENUS CASTNIA AND SOME ALLIED GROUPS. 185

ist das Weiss uberwiegend ausgebreitet; unten dem letateren sehr ahnlich (vedlleicht
nur dessen 9 ?).” Lat. 88 mm.
Hab. Brasilia. In Mus. Kaden olim.

Castnia albofasciata, Schaufuss, Nunq. Otios. p. 10.

| Obs. The three following species form the genus CHRETES of Boisduval. See
supra, pp. 166, 167.]

53. CASTNIA CHREMES. C. alis anticis apice acutis, fuscis aut brunneo-fuscis, basi parum
pallidioribus, fascia obliqua pallidiore e medio costze ad medium disci extensa inde
dilatata, parte basali versus medium marginis postici extensa, parte externa ad
angulum posticum extensa, fascia altera abbreviata subrecta pallida inter priorem et
apicem, strigaque valde indistincta undulata prope marginem posticum: alis pos-
ticis maris nigris purpureo lete nitidis, punctis duobus (vel 1) costalibus, margine
ad angulum externum punctisque nonnullis submarginalibus late aurantiaco-rufis ;
foeminze rufo-aurantiacis, fasciis duabus maculisque marginalibus nigris: alis anticis
subtus pone medium fascia obliqua recta nigra in utroque sexu. Expans. alar.
antic. ¢ unc. 23-84, 2 unc. 3-34.

Papilio chremes, Fabricius, Ent. Syst. 111.1. 47 9 ; Jones, Icones, iii. pl. 72. f. 2; Walker, List Lep. Het.
B. M.1. p. 31 (Castnia chremes) ; Boisduval, H.n. Lép. Hét. p. 536 (Ceretes chremes) .

Corybantes nicon, Hiibner, Samml. exot. Schmett. ii. Versic. 1. f. 1, 4; Gray, Trans, Ent. Soc. L. ii,
p. 148.

Castnia thalaira, Godart, Enc. Méth. ix. 800 (2).

Castnia Fabricii, Swainson, teste Bdv. op. cit. p. 539.

Papilio thats, Drury, Il. ii. pl. 16. f. 4 (aot of Walker).

Foeem. var. major (exp. alar. antic. unc. 33). “‘ Ferrugineo-fusca, aurato viridique micans ;
pectus et abdomen aurato-ochracea; ale antic fasciis tribus abbreviatis fuscis ;
postice late ochraceze, subpurpurascentes, postice atre fasciis duabus arcuatis
ochraceis, 1ma submaculari, 2da maculari.”

Castnia morphoides, Wik. List Lep. Heter. B. M. pl. 31. f. 48, = C.nicon, Hib. Exot. Schmett.

Length of wings 44 lines. “ Nearly allied to C. icarus; but the markings of the fore
wings are different.”

Brazil. In Mus. Brit.

Foem. var. minor. (exp. alar. antic. unc. 2-24).

Castnia thats, Boisduval, op. cit. p. 537.

Hab. Brasilia, Minas Geraes, Rio Janeiro, between Sapezal and Conceicao (Burchell,
30 October, 1827). ‘‘ Umbroso sylvz dense, volatu et coloribus Phalene,” Burchell,
MS. In Mus. Britann., Hopeiano Oxonie, Boisduval.

From the great diversity of the sexes of this and the following species, and the variable-
ness in the intensity of their markings, there has been great confusion in their nomen-
clature.

The description of C. chremes (female) given by Fabricius is scarcely sufficient to
distinguish it from the smaller species, although his words “ fasciis duabus” would imply
simple and not irregular stripes, and his description of the fore wings, “ subtus fere con-

186 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

b)

colores et dilutiores,” apply to it better than to the female of C. Marcelserresii. Mr.
Walker, from an inspection of the original drawings of Jones’s ‘ Icones.’ Danai, pl. 72,
referred the Fabrician insect to the C. nicon of Hubner. A copy of the figure of Jones
was made by Donovan, and is preserved in the Hopeian collection at Oxford. By the
kindness of Mr. Drewitt, the relative of Mr. Jones, and the present possessor of the
volumes of his ‘ Icones,’ I have been enabled to verify the figure and insect, the fascia
beyond the middle of the fore wings being quite straight, and extending to the anal
angle; the apical margin of the fore wings is but slightly rounded; and the black edge
of the hind wing does not extend to the outer angle. iS

The figure of Drury of his Pap. thais (iii. pl. 16. f. 4) exhibits several characters
hitherto unnoticed, which clearly identify it with Hitbner’s figure of the female of
C. nicon. These are the subtruncate (instead of strongly rounded) apical margin of the
fore wings, the curved (instead of quite straight and transverse) inner edge to the
central pale fascia of the fore wings, and the fulvous outer angle of the hind wings, towards
which the second black fascia emits two or three short strigze or small oval spots. In
the females of C. Marcelserresii these strigee or spots are united to the marginal fascia,
which is extended to the outer angle.

The males of the two species are at once distinguished by the undersides of their fore
wings being broadly black in the male of C. chremes and pale brownish grey in that of
C. Marcelserresi.

ITecannot but think the unique female described by Boisduval as agreeing with
“la figure donnée par Drury trés-exacte,” is a very small female of ¢hais, measuring
only 5 centimetres in the expanse of the fore wings, whereas the female which he
describes of thais (his C. chremes) measures 9 centimetres.

The Oastnia morphoides of Walker is certainly only a large female of C. thats, agreeing
closely with Hiibner’s figure (wicon 2).

The following is the description of an insect from Georgia, in North America, which
Herr Schaufuss considers may be a variety of this species.

“ (. thais, var.? Schaufuss {[Nunq. Otios. p.10]. Viel kleiner als Thais, Dr. Die Binden
oben fast verloschen oder schmaler, unten orangegelb: Oberfligel mit nur einer
aus dreieckigen Flecken zusammengesetzen Binde, die in die Randflecken, welche
oben nur angebraunt erscheinen, verlauft.” i

Hab. Georgia, America.

54. CASTNIA MARCELSERRESII. C. alis anticis ovatis (margine apicali versus apicem
rotundato), livido-fuscis aut fusco-fulvescentibus, fascia recta obscura e medio costz
ad medium marginis interni extensa, macula magna obliqua obscura intus subito
dilatata et interdum cum fascia irregulari e costa versus angulum analem extensa
connexa, apice obscuriore et parum angulato: alis posticis maris fuscis, serie
subapicali lunularum pallidiorum vix distinctarum (singulis punctum nigrum in-
cludentibus) : alis posticis foeminz rufo-fulvis, fasciis tribus undulatis nigris (ultima
marginali intus dentata): alis anticis subtus in utroque sexu pone medium macula
magna bipartita costali fasciaque subapicali plus minusve distincta nigris. Long.
corp. une. 1. Expans. alar. antic. unc. 23-3.

GENUS CASTNIA AND SOME ALLIED GROUPS. 187

Mas. Castnia Marel-serresii, Godart, Enc. M, ix. 800; Geyer, in Hiibner, Zutrige exot. Schm. iii. f. 711,
712; Gray, Trans. Ent. Soe. ii. p. 148 (excl. syn. P. thats, Drury).
Fem, C. Fabricii, Godart, Enc. M. ix. 800; Boisduval, H. n. Lép. Hét. p. 538 (nec Swainson).
C. thats, Walker, List Lep. Het. B. M. i. 31 (not of Drury).
Hab. Brasilia. In Mus. Britann., Hopeiano Oxoniz (5 mares, 6 foem.) &c.

The male of this species was described by Godart under the name adopted above, and
the female under that of Fabricii, with the observation that “ce Lépidoptére parait étre
Pun des sexes de celui que M. Swainson a figuré sous le nom de C. Fabricii.”” This sup-
position being erroneous, I have not thought proper to follow M. Boisduval in retaining
the name of C. Fabricvi for this species, but have adopted the usual plan of preferring
the name given to the male insect, C. Marcel-serresit. Gray, followed by Walker, erro-
neously considered the P. thats of Drury to be identical with the female of this species.

55, CasTNIA Diva. C. alis anticis apice acutis, supra nigro-fuscis, fascia latissima albida
ferrugineo-squamosa, e medio coste ad angulum analem extensa, pone medium sub-
interrupta, macula parva discoidali, alterisque parvis ovalibus pone medium albis
aut albidis brunneo circumdatis squamisque nonnullis subapicalibus ferrugineis :
alis posticis basi leete purpureis, fascia lata pone medium nigra, margineque postico
versus angulum externum aurantiaco intus undato. Expans. alar. antic. unc. 23-44.

Castnia diva, Butler, Lepid. Exot. pl. 17. f. 1, 2.

C. tricolor, Felder, Reise Novara, Zool. Bd. ii. Lep. lxxix. fig. 3 (sine descriptione) ; Boisduval, H. n. Lép.
Hét. p. 531.

Hab. Nicaragua, Columbia. In Mus. Britann., Hopeiano Oxonie, Felder, Boisduval.

Scetion Il, Species with the fore wings elongate-ovate. (See supra, p. 167, and
Plate XX XT.)

= Genus GAZERA, Boisduval, in H.-Schaffer, Samml. aussereur. Schm. ; Hist. nat. Lép. Hét. p. 546.

56. (1) Casrnia srmunans. “ Ailes entiéres, les supérieures d’une teinte roussdtre
enfumée depuis la base jusqu’au-dela du milieu, avec une tache oblique d’un brun
noiratre & lextrémité de la cellule discoidale: la partie postérieure de ces mémes
ailes est d’un brun noir, coupée par une bande oblique, droite, partant de la céte et
n’arrivant pas jusqu’au bord externe, outre celail y a, & partir de angle interne, une
rangée de trois ou quatre points jaunatres qui ne remonte pas jusqu’au milieu du
bord extérieur; le bord interne, la nervure médiane et la cdte sont plus ou moins
brunatres. Ailes inférieures d’un fauve roussitre, traversées au-delA du milieu par
une bande noire maculaire, elles ont en outre a l'extrémité une espéce de bordure
noiratre formée des taches cunéiformes ; cette méme bordure offre vers l’angle anal
deux points jaunatres et un point semblable vers l’angle externe. Taille de
Lycorea ceres.”

Gazera simulans, Boisduval, H. n. Lép. Hét, p. 547, pl. 11. fig. 4.

This species is the representative of Zycorea ceres, one of the Heliconian butterflies.
It is closely allied to, if not a variety of C. personata, Wk.

Hab. Columbia. In Mus. Boisduval.
SECOND SERIES.—ZOOLOGY, VOL. I, 2¢

185 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

57. (2) CAsTNIA PERSONATA. “ Fusca, aureo viridi micans, subtus flava; antennze apice
flavescentes ; abdomen subauratum vittis duabus ochraceis : ale cervine, sat angustz,
lituris nigro-fuscis, fascia marginali flavescente maculata; anticee vitta discali,
strigas quatuor emittente, postice vittis duabus parallelis extus abbreviatis. Nearly
allied to O. linus. Length of wings 36 lines.”— Walker. ~

The above description is unintelligible, as may be seen by M. Boisduval’s attempt to
translate it into French, and by his inability to perceive its apparent identity with his

Gazera simulans. The general colour of the four wings is obscure luteous, with black

markings; these consist of an oblique patch, arising on the costa before the middle,

dilated towards the middle of the wing into an elongated patch; beyond the middle of
the costa is a broad oblique band reaching to the middle, followed by four more distinct
luteous longitudinal spots preceding the broadly black apex ; the inner margin is dusky :
the hind wings are rather lighter luteous in the middle of the disk; the costa, a central
bar, and the apical margin black, the latter with four white dots on its inner edge near

the outer angle. Expans. alar. antic. lin. 36.

Castnia personata, Walker, List Lep. Het. B. M. xxxi. p. 48; Boisduval, H. n. Lép. Hét. p. 546 (Gazera

personata).

Hab. Guyaquil. In Mus. Britann.

58. (3) CasTNIA cononiA. (Plate XXXII. fig. 5.) Alis anticis elongato-ovalibus apice
rotundatis, dimidio basali obscure luteo-fulvo, striga postcostali plagaque magna
prope marginem internum nigris, hac striolam pallidiorem includente; macula
media (prope costam) postice angulata ejusdem coloris fasciaque obliqua (e maculis 5
ovalibus constante) inter medium et apicem alz albida; alis posticis saturate fulvis,
fascia postcostali maculis quinque ovatis pone medium, ad marginem analem majori-
bus et connexis, limboque apicali nigris ; capite albo punctato, collo postice fulvo ;
fascia transversa media thoracis flavida; abdominis fusci lateribus anguste fulvis,
segmentis ventralibus basalibus et apicalibus fulvis, intermediis nigris. Long. corp.
unc. 14. Exp. alar. antic. une. 33.

Hab. Ecuador. In Mus. Hopeiano Oxonie (olim Saunders).

This species is nearest allied to C. ecuadoria, with which the arrangement of the cell
and branches of the postcostal vein of the fore wings nearly agrees; but the transverse
vein closing this cell is longer, in consequence of which the branches 3 and 4 arise
wider apart.

59. (4) Castnza cratiInA. (Plate XXXII. fig. 4.) Alis anticis oblongo-ovatis apice
subrotundatis fulvis; striga costali prope basin, altera subcostali tertiaque multo
majore prope marginem internum, striga undulata obliqua discoidali, macula rotunda
media cum margine antico connexa, plaga magna obliqua inter medium et apicem,
limboque apicali dentato nigris ; alis posticis aurantiaco-rufis, striga subcostali, fascia
lata media e margine anali fere ad angulum externum extensa limboque dentato
nigris ; capite albo maculato, collo lato fulvo fascia transversa tenui flavida ; abdomine ©

GENUS CASTNIA AND SOME ALLIED GROUPS. 189

nigricante lateribus segmentisque 5 apicalibus subtus fulvis; thorace subtus albido
vario; antennis albidis. Long. corp. unc. 13. Expans. alar. antic. unc. 43.
Hab. Amazonia. In Mus. Hopeiano Oxoniz (olim Saunders).
This species is closely allied to C. cycna, but has the fore wings less rounded; the
arrangement of the cell and branches of the postcostal vein agrees with that of
C. cycna.

60. (5) Castnra EcuaDoRIA. (Plate XXXII. fig. 6.) Alis anticis oblongo-ovatis apice

- rotundatis; e basi ultra medium obscure lutescentibus, apice lato fusco; cellula
basali fusco marginata, macula media rotunda subcostali, striga obscura media,
alteraque majore marginis interni fuscis ; alis posticis obscure fulvis nebula magna
ex angulo anali versus medium ale extensa fimbriaque fuscis ; corpore fusco ; collari,
fascia media transversa thoracis, lateribusque abdominis lutescenti-albidis ; antennis
lutescentibus: alis subtus magis rufescenti-fulvis, apice fusco, posticis stria tenui
transversa subbasali fusca. Long. corp. unc. 14. Expauns. alar. antic. unc. 34.

Hab. Ecuador. In Mus. Hopeiano Oxonie (olim Sawnders).

This species so closely resembles C. ¢ruailla in general form and appearance that it
might easily be considered to be a variety of that species; but the arrangement of the
veins proves them to be totally different, there being a very distinct elongate-ovate cell
in C. ecuadoria, formed at the base of the branches of the postcostal vein, the 2nd, 3rd,
4th, 5th, and 5*th being emitted at nearly equal distances apart at the extremity of the
cell (Plate XXVIII. fig. 10).

61. (6) Casrnta zAGRmA. “ Ailes supérieures noires avec une bande longitudinale fauve,
sinuée, un peu irréguliére, coupée par les nervures partant du milieu de la base et
se prolongeant vers le bord postérieur ; outre cela, il y a sur le milieu de la céte un
petit trait, et au-dessous de celui-ci une tache un peu triangulaire de la méme
couleur, divisée par les nervures, l’extrémité postérieure offre deux rangées paralléles
de taches blanches lavées de jaune; mais la premiére rangée ne descend pas au-dela
du milieu de Vaile.” Expans. alar. antic. une. 44.

Gazera zagrea, Felder, Reise d. Novara, Zool. Bd. ii. Abth. 2, pl. 79. f. 2 (sine descr.) ; Boisduval,
H. n. Lép. Hét. p. 547.

Hab. Panama. In Mus. Felder.
This is the Heterocerous counterpart of Papilio zagreus, which isitself the counterpart.
of one of the Heliconian butterflies.

. Nearly allied to C. cratina, Westw., but with the fulvous markings of the fore wings

and the black markings of the hind wings much less developed in size. Tore wings

black, with a longitudinal fulvous vitta of irregular width, extending from the base of

the wing along its middle, rather deflexed and narrowed at its extremity; considerably

beyond the middle of the wing an oblique cuneated patch of the same colour arises at

the middle of the costa, followed by a row of six suboval spots, the lowest of which ranges

with the extremity of the central bar; beyond these is a submarginal row of nine white
2c2

190 | MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

spots, the middle ones being the smallest ; hind wings fulvous, costa narrowly black,
with a row of semiovate black spots beyond the middle, not extending to the outer
angle; apical margin black and strongly dentate within, the teeth near the anal angle
uniting with the spots of the inner row. Described from the figure of Dr. Felder.

62. (7) CASTNIA TRUXILLA. (Plate XXXII. fig. 3.) Alis fuscis, anticis oblongo-ovatis
apice rotundatis macula magna media lutescente e costa oblique versus angulum
internum extensa, guttam parvam obscuram fuscescentem versus costam ineludente ;
eutta parva lutea prope angulum internum nebulisque duabus oblongis pallidioribus
prope marginem internum; alis posticis ovalibus fuscis, basi ipsa ad marginem ana-
lem squamis fulyis notata; corpore fusco, capite albo-maculato, patagiis fulvo-
marginatis ; abdomine fusco, lateribus anguste fulvis; abdomine subtus basi et apice
fulvis, segmentis intermediis nigris, griseo marginatis ; antennis fuscis, apice fulvis ;
alis subtus pallidioribus, costa posticarum fulva. Long. corp. une. 14. Expans.
alar. antic. unc. 33.

Hab. Columbia. In Mus. Hopeiano Oxoniz (olim Saunders).

This obscurely coloured species is remarkable for its elongated fore wings (which are
rounded at the apex) and its oval hind wings. In the fore wings there is no cell formed
by the basal portion of the branches of the postcostal vem; nor is there any transverse
connexion between the 3rd and 4th branches of the same vein. (See Plate XXVIII.
fig. 12.)

68. (8) CAsTNIA coLUMBINA. “ Ailes supérieures d’un noir foncé; elles ont ala base, deux
bandes longitudinales, d’un roux un peu enfumé, dont V’inférieure se termine par deux
rayons blanchatres: outre cela, il y a sur le milieu de la céte un petit trait blane et
au-dessous de celui-ci, prés de la principale nervure, une tache blanche arrondie;
plus bas et plus en dehors deux taches blanches cunéiformes, ’extrémité postérieure
offre deux rangées paralléles de taches blanches inégales, les unes rondes, les autres
allongées: ailes inférieures d’un fauve vif, avee une bordure noire crénelée, et une
rangée de points blancs; outre cela elles offrent au-dela du milieu deux ou trois
points noiratres disposés transversalement.”’ Hxpans. alar. antic. 12 centim.

Gazera columbina, Boisduyal, H. n. Lép. Hét. p. 546.

Hab. Nova Granada. In Mus. Boisduval.
Obs. This species is nearly allied to C. salvina, but differs in the markings of the hind
wings.

64. (9) CasTNIA SALVINA. (Plate XXXII. fig. 1.) Alis anticis oblongo-ovatis nigrican-
tibus, basin versus vittis duabus longitudinalibus obscure rufescentibus, maculisque
novem, magnitudine variis, discoidalibus, serieque submarginali macularum novem
albidarum (maculis apicem appropinquantibus longioribus) ; alis posticis fulvis, costa
limboque lato integro nigris, hoc maculas sex submarginales parvas albas includente :
capite et thorace nigris, illo albo maculato, patagiis albo marginatis : antennis fulvis,

GENUS CASTZNIA AND SOME ALLIED GROUPS. 191

basi obscurioribus: alis subtus similiter ac supra coloratis, posticarum costa et
striga parallela subcostali nigris: abdomine fulvo, apice obscuriore; subtus medio
sterni abdomineque albidis. Long. corp. unc. 13. Exp. alar. antic. unc. 43.

Hab. Veragua (Salvin). In Mus. Hopeiano Oxoniz (olim Saunders).

The arrangement of the branches of the postcostal vein in this species is peculiar, there
being no basal cell formed by the transverse junction of the middle branches; the third
branch springs from the second at a considerable distance from the base of the latter;
and the 4th and 5th branches arise at a considerable distance from the transverse base of
the 6th branch (i. e. 65*), which forms the termination of the discoidal cell.

65. (10) Castnia cyona. (Plate XXXII. fig. 2.) Alis anticis subovatis fusco-nigris,
basi, costa, striga media (e basi ad medium alz extensa) margineque postico obscure
fulvis, striga media subcostali obliqua, macula ovali discoidali, fascia obliqua inter
medium et apicem maculisque 9 magnitudine variis albidis; alis posticis lete au-
rantiacis, extus in flavum pallescentibus, limbo intus acute dentato nigro: capite
albo-punctato; antennis albidis basi nigricantibus: thorace fascia antica transversa
fulva maculisque duabus posticis albidis: abdomine fusco, lateribus ventreque
fulvis. Long. corp. unc. 1}. Expans. alar. antic. une. 4.

Var. minor (exp. al. ant. unc. 34) fascia basali alee antics et striga marginis postici
obsoletis, coloribus etiam magis vividis. ~ ;

Hab. Columbia. In Mus. Hopeiano Oxoniz (olim Saunders).

The broadly ovate fore wings and the richly coloured hind wings, with their strongly
dentated black border, distinguish this species, which agrees with O. ecwadoria in the
arrangement of the cells and branches of the subcostal vein of the fore wings.

66. (11) Castyia mimica. (Plate XXXII. fig. 7.) Alis fusco-nigricantibus, certo situ
parum metallicis, anticis elongatis basi obscurioribus, squamis nonnullis rufis juxta _
tegulas; posticis macula magna transverso-oyali inter medium et angulum analem
flavescente: capite nigro, punctis duobus ante antennas, linea tenui postica pone
oculos, thoracis macula minuta antica albis; tegulis rufis; abdominis segmentis
intermediis lateribus rufo angulariter maculatis, squamisque albis strigas breves
laterales ad basin segmentorum formantibus: antennis nigris, clava subtus fulva:
coxis anticis et 2 posticis geniculisque pedum 4 anticorum rufis: abdomine subtus
utrinque albo maculato, segmentis intermediis macula parva rufa in medio notatis.
Long. corp. unc.1j. Expans. alar. antic. une. 3 lin.

Hab. Amazonia. In Mus. Hopeiano Oxonie (olim Saunders).

C. mimica, Felder, Reise d. Novara, Ins. Lep. pl. 79. f. 4: (sine descript.).

This remarkable species is, as to its colours, unique in the group; its wings are also
of peculiar form, giving it an aspect quite alien to its allies; it possesses the elon-
gated cell at the base of the branches of the subcostal vein; but the erd and 4th
branches arise at some distance beyond the cell. (Plate XXIX. figs. 11, 12.)

192 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

67. (12) Casrnra tinus. C. alis nigris hyalino-albido variis, venis nigris; anticis macula
magna subtrigona, venis divisa, fascia media e costa ad angulum internum api-
calem extensa in duas partes interrupta, macula magna ovali subapicali in sex
partes venis divisa, serieque submarginali punctorum alborum: alis posticis macula
maxima discoidali transverso-ovali, serieque externa macularum conoidearum magni-
tudine decrescentium versus angulum analem, ubi in puncta alba evadunt. Long.
corp. une. 14-14. Expans. alar. antic. unc. 83-4.

Variat venis in disco pallido alarum posticarum concoloribus nec nigris, maculis prope
angulum externum minoribus punctisque versus angulum analem fere obliteratis.

Papilio linus, Cramer, iii. pl. 257. fig. A; Herbst, Ins. pl. 78. f. 4.; Dalman, Monogr. p. 24; Act. Holm.
1824, p. 405 (Castnia linus) ; Gray, Trans. Ent. Soe. L. ii. p. 147; Walker, Cat. Lep. B. M. p. 32;
Boisduval, H.n. Lép. Hét. p. 544, f. 15.

Var. C. heliconioides, Herrich-Schaffer, Samml. exot. Schmett. pl. 4; Walker, op. cit. p. 1582; Bois-
duval, H. n. Lép. Hét. p. 545 (Gazera hel.).

Hab. Brasilia, Surinam, Cayenne, Guatemala. Mus. Britann., Hopeiano Oxoniz, ©
Boisduval.

Dr. Boisduval has separated two specimens in his collection from Cayenne from the
more ordinary Brazilian ones, considering the former the true representatives of
Cramer’s figure of C. linus from Surinam, and giving to the Brazilian insects a new
name, G. heliconides (recte Heliconioides, as previously published by Herrich-Schaffer).
I cannot, however, regard these insects otherwise than as belonging to one species, dif-
fering both individually and geographically, and even sexually: variations of the latter
character are exhibited in the form of the wings, which modifies to a certain extent
the form of the markings. Thus a female specimen in the Hopeian Collection from
Cayenne has the fore wings broad and almost ovate; and a male from the Amazons
(also with the veins of the disk of the hind wings concolorous) has the fore wings re-
markably long, as will be seen from the admeasurement of one of the fore wings of
these individuals as compared with those of two Brazilian specimens with black veins.

Veins of disk of hind Veins of disk of
wings concolorous. hind wings black.
Cayenne 9. Amazon. Brazilian 9. Brazilian 3.
Lengthofforewing .... .. . 48centim. 50 AT AS
\WVACRAN OVE Chg), Poe GBA Vat Ia AAD. bo 23 223 22
Length of outer margin from extreme
Apex tOnmMnerane |e. aael Ne eran siekime ome, 30 28 26
Length from base toinnerangle . . . 30 ,, dl 29 30

It will thus be seen that instead of the Brazilian individuals being “un peu plus grands
avec les ailes plus larges” than those of the Cayenne ones, as stated by Boisduval, the latter
are the largest in every respect, and more especially in the width of the wing.

The Cayenne female before me has the large spot of the hind wings more ovate,
the black anal margin extending further towards the base than in the Brazilian spe-

‘cimens; the veins in this large spot, in this female and in three other specimens in the

GENUS CASTNIA AND SOME ALLIED GROUPS. 193

Hopeian Collection, are concolorous, and not clothed with black scales as in the Brazilian
specimens; and the marginal spots near the outer angle of the hind wings are much
shorter than in the latter, whilst the small dots near the anal angle are reduced to a very
few scales. In the four Hopeian specimens, with pale veins in the disk of the hind wings,
the pale markings are less strongly yellow than in the Brazilian ones; and in the Ama-
zonian male above noticed the two apical spots of the terminal macular fascia are much
larger than the three following, whilst the three next are reduced to minute dots, the
one close to the inner angle being as large as the 8rd to 5th. Thus neither in size,
shape, nor markings can we specifically distinguish between the specimens of this

species.

68. (18) CasTNIA ACRMOIDES. C-alis anticis fuscis basi magis lutescentibus, medio fulvo
tinctis; venis omnibus dilute nigris, strigis longitudinalibus obscurioribus inter
venas fasciaque obliqua albido-ochracea e costa ultra medium, versus medium
marginis apicalis extensa; alis posticis aurantiaco-rufis, venis limboque apicali intus
dentato nigris, hoc serie macularum parvarum (subtus multo majorum) flavarum
notato. Long. corp. unc. 13. Expans. alar. ant. unc. 3 lin. 1.

Casinia acreoides, Boisduval, Sp. Gén. Lép.i. pl. 14. f.1; H. n. Lép. Hét. p. 540; Guérin, Icones R.
An. Ins. pl. 83. f. 4; Griffith, An. K. Ins. pl. 53. f.4; Gray, Trans. Ent. Soc, L. 1. p.148; Walker,
List Lep. Het. B. M. i. p. 32. ‘

Castnia actinophorus, Kollar, Lep. Brasil., Ann. Wien. i. p. 215, pl. 12. f. 2; Walker, List Lep. B. M.
i. p. 29; Boisduval, H. n. Lép. Hét. p. 520; “Gray (mn Ann. Kingd.) as C. acreoides,” Hrichs.
Berichte, 1839.

Hab. Brasilia. In Mus. Britann., Hopeiano Oxoniz, Boisduval.

This species bears so close a resemblance to Acrea thalia as often to be mistaken for
it. Boisduval states that its larva feeds on certain species of Bromeliacez.

It is surprising that Mr. Walker should not have perceived from Kollar’s excellent
figure that C. actinophorus (placed by Walker in his section 1 of subdiv. 2) is identical
with C. acreoides of Boisduval, which he places in his section 8. Deceived by this error,
Dr. Boisduval, who, like Walker, simply repeats or translates Kollar’s character of
actinophorus *, has also given the new supposed species as distinct, placing C. actino-
phorus in the genus Castnia, and C. acreoides in that of Orthia.

Genus IJ. Orruta, Boisduval (pars).

Ale magne, late ; antice subtrigone, cellula discoidali aperta, loco ven disco-cellularis tantum crassitie
minuta nodiformi ramorum 05* et c3* indicato, ramo supero discoidali (65*) e vena postcostali
pone cellulam emisso, cellula nulla accessoria postcostali: alee postice cellula angusta dimidiata vena
mediana connexa, cellula ordinaria aperta: abdomen alas imferiores haud postice superans.
(Plate XXIX. figs. 16, 17 +.)

* «¢ Alis anticis nigris radiis plurimis viridi-fuscis, basi fasciaque abbreviata dilute flavis: posticis luride rubris,
radiis limboque nigris, hoc flayo maculato. Long. corp. lin. 9-12. Exp. alar. ant. lin. 27-33.”

+ In figure 16 the small lettering of the two branches 6 5* and ¢3* is placed too high—the former branch, which
looks like-the terminal branch of the postcostal vein, and the latter branch, which looks like a fourth branch of the

194 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Having restored C. therapon and acreoides (forming Boisduval’s first section of
Orthia) to Castnia, and O. augias and nexa belonging to the Agaristides, I have been
compelled to regard C. pelasgus, Fabr., and wnifasciata, Felder (which appears to me to
be identical with pelasgus), as a separate genus, for which I have, unwillingly, retained
the name of Orthia.

OrtHIA PELASGUS. Alis anticis nigris bronzeo-viridi tinctis, fascia lata lutescente aut
alba e medio costee ad angulum internum extensa, medio ejus paullo latiore apiceque
infero paullo incurvo ; alis posticis CL aoa, margine externo parum cerules-
cente. Expans. alar. ant. unc. 275-23.

Papilio pelasgus, Fabricius, Sp. Ins. ii. p. 54; Ent. Syst. i. pt. 1. p. 46 ; Herbst, Ins. pl. 136. f. 6; Oliv.

Ene. M. Ins. pl. 12. f. 5; Cramer, iii. pl. 202. fig. D; Dalman, Monogr. p. 21; Act. Holm, 1824,
p- 404 (Castnia p.) ; Godart, Enc. Méth. ix. p. 801; Gray, Trans. Ent. Soc. L. 1. p. 146; Walker,
List Lep. Het. B. M.i. p. 22; Boisduval, H. n. Lép. Hét. p. 54 (Orthia p.).

Castnia unifasciata, Felder, Reise d. Novara, Zool. Band ii. Abth. 2, pl. 79. f. 5 (sine deser.) ; Bois-
duval, H. n. Lép. Hét. p. 542 (Orthia unif.).

H.-Sch., Samml. exot. Sch. pp. 56 and 78, gives O. augias, H.-S., as = pelasgus, F.

Except in the white colour of the fascia of the fore wings, I can see nothing to separate

Dr. Felder’s figure from C. pelasgus of Cramer.

Hab. Brasilia, Guiana, Surinam, Ega, Bolivia. In Mus. Britann., Hopeiano Oxonize,

Boisduval.

Genus III. Synemon, E. Doubleday. (Plate XX XIII.)
In Lord’s Discov. in Australia; Klug, Abh. k. Akad. Berlin; Walker, List Lep. Heter. Brit. Mus. part i.;
Boisduval, H.n. Lép. Hét. 2
1. Synemon Sopuia. Nigro-fusca, alis anticis maris fuscis griseo-brunneoque nebulosis,
nebula obscura baseos alteraque submedia puncto albo adjecto; venis griseis, famine
nigricantibus squamis plumbeis irroratis, fasciam subapicalem maculis nigris divisam
formantibus, macula parva submedia fasciaque abbreviata costali maculari pone
medium : alis posticis maris basi nigris macula ovali ante medium, fascia maculari
pone medium maculisque submarginalibus flavis; famine nigris, macula magna
aurantiaca baseos, fasciaque lata abbreviata transversa marginis analis, macula
magna versus angulum externum, alterisque 3 vel 4 submarginalibus fulvis: alis
subtus luteis, fasciis ramosis nigris. Expans. alar. antic. lin. 18-22.
Hesperia? Sophia, White, in Grey’s Exped. Australia, App. p. 474, fig. 7; Doubleday, in Lord’s Discov.
Australia, App. i. p. 516, pl. 3. £. 5 (Synemon Sophia) ; Walker, List Lep. Het. B. M.1. p. 34, vii.
p. 1583; ma Abhandl. kon. Akad. Berlin, 1850, p. 248, pl. annex. f. 1,2; Boisduval, H. n. Lép.
Hét. p. 550.
Hab. Australia, King George’s Sound. In Mus. Britann., Hopeiano Oxonie,
Boisduval.

median vein, being separated from each other by the slenderly indicated fold running into the discoidal cell (the letters
c 3* in the plate being improperly placed opposite to its extremity), the closing of which by the transverse disco-
cellular yein is indicated by two small swellings, omitted by the engraver, opposite to each other in the middle of the
wing.

GENUS CASTNIA AND SOME ALLIED GROUPS. 195

2. SYNEMON THERESA. Griseo-fusca, disco subalbido, marginibus omnibus fuscis, venis
nigricanti-fuscis, in medio et ad marginem apicalem dilatatis, macula obliqua media
fusca albo extus marginata fasciaque subapicali e maculis ovalibus 4 et 3 fuscescenti-
bus inter venas obscuras formata; alis posticis fulvis basi fuscis, macula parva ante
medium fulva, margine externo tenui serieque macularum submarginalium fuscis.
Expans. alar. antic. lin. 18.

Synemon theresa, Doubleday, in Lord’s Discov. Australia, Append. i. p. 517, pl. 3. f.6; Walker, List Lep.
Het. B. M.i.p. 35, vi. p.84; Klug, Abhand. kon. Akad. Berlin, 1850, p. 249, pl. 3. £.5,6; Boisduval,
H.n. Lép. Hét. p. 552.

Hab. Nova Hollandia. In Mus. Britann., Hopeiano Oxoniz, Boisduval.

3. SyneMoN Lava. Nigro-fusca, subtus albida; antennis albo annulatis: alis anticis
fuscis, albido conspersis, maculis duabus irregularibus (una majore ante medium,
altera minore pone medium subcostali) albido separatis fasciaque apicali maculari
nigro-fuscis ; subtus aurantiacis, fusco maculatis : posticis aurantiacis, macula basali
fasciisque duabus nigris, prima arcuata, mediana, irregulari, secunda lata marginali,
serie guttarum fulvarum ornata; subtus albo guttatis. Expans. alar. antic. lin. 20.

crete leta, Walker, List Lep. Het. B.M.i. p. 36; Boisduval, H.n. Lép. Hét. p.553.

Hab. Nova Hollandia. In Mus. Britann.

4, SYNEMON SELENE. S. corpore fusco, subtus albido, abdomine supra griseo; alis
anticis fusco-griseis, medio late fuscis, ante apicem fusco maculatis, fascia maculari
transversa interrupta ante medium lunulaque media albis; posticis fuscis luteo tri-
fasciatis ; omnibus subtus albido-testaceis fusco maculatis: ¢ 2. Expans. alar. ant.
lin, 203-21.

Synemon selene, Klug, Abhandl. kén. Akad. Berlin, 1850, p. 249, pl. annex. f. 3,4; Boisduval, H.n. Lép.

Hét. p.553; Walker, List Lep. Het. B. M. vii. p. 1584.

Hab. Nova Hollandia. In Mus. Berlin.

5. Synemon ais. JS. corpore griseo, subtus pallido; alis anticis supra fusco-griseis,
fascia tranversa sinuata ante medium maculaque media griseis: posticis luteis fascia
transversa medio profunde sinuata baseos, maculis ad marginem margineque fuscis,
omnibus subtus luteis fusco-maculatis 9. Expans. alar. antic. 12+ lin.

Synemon nais, Klug, Abhandl. kén. Akad. Berlin, 1850, p. 250, pl. annex. f. 7; Walker, List Lep. Het.
B. M. vii. p. 1584; Boisduval, H.n. Lép. Hét. p. 553.

Hob. Nova Hollandia. In Mus. Berlin.

6. SYNEMON caTocaLoipEes. Nigyo-cinerea, subtus alba; antennis albo annulatis ; alis
anticis squamis plurimis pallide cinereis, basi maculis tribus alteraque versus
medium marginis interni nigris, macula magna pone medium nigro-ferruginea,
fascia arcuata guttarum albarum strigisque quatuor brevibus subapicalibus nigris,
margine apicali fusco intus crenulato; alis posticis lzete rufis, fascia parva media

SECOND SERIES.—ZOOLOGY, VOL. I. 2D

196 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

transversa versus marginem analem, costa Jate et irregulariter nigra, limboque
apicali nigro guttis nonnullis subapicalibus rufis. Expans. alar. antic. lin. 22.
The above description is made from the type specimen in the British Museum.
Synemon catocaloides, Walker, List Lep. Het. B. M. xxxi. p. 44; Boisduval, H. n. Lép. Hét. p. 551.
Hab. Swan River. In Mus. Britann.

7. SYNEMON PARTHENOIDES. Alis anticis brunneis obscurius nebulosis, fascia lata sub-
maculari e medio coste fere ad angulum internum extensa, venis divisa, altera
breviore bifida literam v simulante, inter precedentem et apicem, albis; alis posticis
nigris, macula rotunda aurantiaca prope basin, seriebus duabus macularum ejusdem
coloris inter medium et apicem, interiore in medio interrupta, altera versus angulum
externum abbreviata. Hxpans. alar. antic. fere une. 2.

Synemon parthenoides, Felder, Reise d. Novara, Zool. Bd. ii. Abth. 2, tab. 79. f. 7,8; Boisduval, H. n.
Lép. Hét. p. 550, pl. 11. fig. 5.

Hab. Noy. Hollandia. In Mus. Boisduyal.

8. SYNEMON mopsa. Nigro-fusca; alis subtus fulvescentibus, maculis testaceis, anticis
supra pallide fuscis vittis duabus albis basalibus, gutta alba ad apicem areole
discoidalis, pone medium fascia transversa maculari alba seriem macularum fus-
carum includente, margine apicali subgriseo; alis posticis pallide castaneo-fuscis,
nebulis nonnullis submarginalibus fuscescentibus. Hxpans. alar. antic. 15 lin.

Synemon mopsa, Doubleday, in Lord’s Discov. Australia, App. i. p.518, pl. 3. f. 7; Walker, List Lep. Het.
B. M.i. p. 35; Boisduval, H. n. Lép. Hét. p. 552.

S. theresa, var., Klug, Abh. kon. Akad. Berl. 1850, p. 249.

Hab. South Australia. In Mus. Britann., Boisduval.

9. SYNEMON ICARIA. Alis anticis griseo-albido nebulosis, basi obscuriore, fascia e medio
costee ad marginem internum (ibique dilatata) fusca, fasciis duabus inter medium
et apicem sinuatis obliquis; interna versus angulum internum seriem lunularum
formante, externa (subapicali) abbreviata, lunulis submarginalibus fuscis: alis pos-
ticis flavo-aurantiacis, macula magna basali marginis abdominalis cum striga un-
data submedia e margine costali fere connexa nigris, limbo apicali angusto nigro,
aurantiaco guttato. Expans. alar. antic. unc. 14.

Synemon icaria, Welder, Reise d. Novara, zool. Th. Bd. ii. Abth. 2, pl. 79. f. 6; Boisduval, H. n. Lép. Hét.
p. 554.

Hab. Nova Hollandia, Swan River. In Mus. Boisduval.

10. SyNEMON PLANA. (Plate XXXIII. fig. 8.) Fusca, abdomine pallido depresso: alis
anticis fusco-badiis, macula ovali inter basin et medium albido cincta, strigaque
tenui conica postice adjecta, macula indistincta pallida ad apicem cellule discoidalis
maculisque duabus majoribus inter medium et apicem nigricantibus, lineis albidis
marginatis; margine apicali tenui nigro, intus serie punctorum albidorum notato:

’

GENUS CASTNIA AND SOME ALLIED GROUPS. 197

alis posticis leete fulvis, guttis 3 et 1 pone medium margineque apicali tenui nigris :
alis subtus albis; margine tenui apicali omnium fusco, squamis luteis irrorato ;
anticis prope apicem punctis duobus, posticis cum 4 pone medium nigris. Expans.
alar. antic. unc. 13.

Synemon plana, Walker, List Lep. Het. B. M. i. 37, Boisduval, H.n. Lép. Hét. p. 554.
Hab. Nova Hollandia, Hunter River. In Mus. Britan., et Hopeiano Oxonie.

11. Synemon pirecta. (Plate XX XIII. fig.6.) Fusca, alis anticis concoloribus, fascia

recta alba, e medio costz fere ad angulum internum extensa, lunula abbreviata inter
medium et apicem minus distincta, spatio inter fascias plagam ovalem fusco-nigram
formante; alis posticis nigris, macula curvata inter basin et medium, fascia macu-
lari pone medium valde sinuata guttisque nonnullis ovalibus submarginalibus leete
aurantiacis ; alis subtus rufo-fulvis nigro maculatis. Hxpans. alar. une. 2.

Hab. Australasia. In Mus. Hopeiano Oxonie.

12. Synemon NupTA. (Plate XXXIII. fig.7.) Plumbeo-cinerea, macula oblique trans-

versa ad medium cellule discoidalis extus squamis albis marginata, fascia media
obliqua pone medium e costa ad medium disci extensa, altera abbreviata costali inter
hance fasciam et apicem strigaque subapicali e maculis conoideis nigris formata ; alis
posticis nigris extus cyaneo nitidis, plaga magna basali trigona, maculis duabus
pone medium prope marginem internum alteraque prope angulum externum san-
guineis: alis anticis subtus nigris, fascia abbreviata media costali alteraque sub-
apicali rufis ; posticis nigris guttis tribus albis discoidalibus in triangulum dispositis,
alteraque parva rufa inter duas posticas. Expans. alar. unc. 13.

Hab. Australasia occidentalis. In Mus. Hopeiano Oxonie.

13. SyneMon NorHA. (Plate XX XIII. fig.10.) Fusca, alis anticis cinereis, nigro variis,

macula magna indistincta inter medium disci et marginem posticum alteraque
subcostali, inter medium et apicem, in strigam punctorum nigrorum postice ad
marginem internum extensam terminante, strigaque maculari submarginali, margi-
neque ipso tenni nigricantibus ; alis posticis lete aurantiacis, basi fascia media (ad
costam paullo dilatata) limboque tenui marginali nigris, hoc serie guttarum auran-
tiacarum notato: alis subtus rufo-fulvis, anticis striga interna basali fasciaque brevi
(pone medium costze) extus guttis albidis notata; alis posticis macula ante medium,
fascia media limboque nigris, omnibus guttis albidis postice notatis. Expans. alar.
antic. une. 14.

Hab. Australasia, Albany (Brewer). In Mus. Hopeiano Oxonie.

14. SYNEMON OBSCURELLA. (Plate XXXIII. fig.9.) Fusco-nigricans, alis anticis striga

tenui abbreviata obliqua media alba, lunulisque duabus inter medium et apicem
pailide luteis (externa obscuriore); alis posticis nigricantibus, gutta parva lutea

inter basin et medium, dimidio apicali fulvo-luteo, striga undata subapicali, in medio.

interrupta, fuscescente: alis anticis subtus basi nigris, dimidio apicali fulvo, macula
2D 2

198 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

rotunda versus costam nigra; alis posticis nigris, ultra medium fulvis striga sub-
undata submarginali fusca. Hxpans. alar. antic. unc. 1}.
Hab. Australasia. In Mus. Hopeiano Oxonie.

15. SYNEMON HESPERIOIDES. (Plate XX XITI. fig. 11.) Fusca, alis anticis versus basin
squamis albidis variis, macula parva conoidea ad apicem cellule discoidalis, maculis
duabus (lineis tenuibus marginatis) strigaque apicali lunularum parvarum albidis;
alis posticis brunneis, pone medium striga fere indistincta macularum nigrarum,
margine apicali fusco: alis subtus brunneis, anticis in medio et ad apicem, posticis
in medio, serie irregulari circularum margineque apicali squamis albido-sericeis
irroratis. Expans. alar. antic. une. 15.

Synemon hesperioides, Felder, Reise d. Novara, zool. Th. Bd. uu. Abth. 2, pl. 82. fig. 12; Boisduval, H.
n. Lép. Hét. p. 555 (regarded as one of the Hesperiide).

Hab. Nova Hollandia. In Mus. Hopeiano Oxoniz.

16. SynemMon vacans. (Plate XXXIII. fig. 12.) Alis anticis fusco-cinereis, fascia
transversa nigra ante medium e costa, postice acuminata, marginem posticum vix
attingente, fascia altera abbreviata subcostali venis divisa nigra pone medium, extus
pallidius margivata, guttis tribus ovalibus nigris subapicalibus alterisque quinque
submarginalibus margineque ipso tenui nigris; alis posticis nigris, fasciis duabus
arcuatis aurantiacis: alis subtus fulvo-luteis, anticis macula irregulari media alte-
raque pone medium subcostalibus nigris, apice albido nigroque vario; alis posticis
magis fulvescentibus, medio circulo fere integro nigro notato, limbo tenui nigro,
fulvo guttato. Hxpans. alar. antic. lin. 19.

Hab. Nova Hollandia (Strange). In Mus. Hopeiano Oxonie.
Obs. Heec species cum sequente a reliquis differunt alis anticis angustioribus, margine
antico in medio parum concayo-sinuato nee convexo.

17. Synemon GRratiosa. (Plate XX XIII. fig.13.) Alis anticis fuscis, dimidio apicali
pallidiore, macula irregulari transverse obliqua submedia, intus squamis pallidi-
oribus irrorata, macula altera subcostali conoidea pone medium nigra, serieque
submarginali punctorum nigrorum (in medio interrupta), margine ipso tenui ob-
scuro; alis posticis aurantiacis, macula parva nigra versus costam et pone medium
posita limboque tenui nigro, in medio aurantiaco guttato: alis anticis subtus obscure
fulvis apice albidis, maculis duabus mediis duabusque inter medium et apicem fuscis,
margine fusco, albido guttato; alis posticis fulvis, maculis tribus mediis, externa
majore et ad costam extensa, limbo nigro, extus albido guttato. Hxpans. alar. antic.
mmbae, dees

Hab. Nova Hollandia. In Mus. Hopeiano Oxoniz.

Genus IV. Tascina. (Plate XX XIII. fig. 5.)

Insectum orientale Castnias Americe meridionalis mundo antiquo representans, cum quibus antennis
necnon numero venarum alarum congruit, his autem aliter dispositis ; palpis exiguis, lingua spirali

GENUS CASTNIA AND SOME ALLIED GROUPS. 199

obsoleta, alisque anticis apice acuminatis e Castniis facile distinguitur. Caput magnum ; oculi maximi,
laterales, antice approximati, spatio angusto inter se separati; palpi tenuissimi, breves, erecti, ad
medium frontis vix attingentes, articulo penultimo longo, ultimo minuto acuto. Vestigium nullum
lingue spiralis detexi. Antenne circiter 50-articulate, clavatz, articulis primis 32 filiformibus ; apice
acuto, parum incurvato, fasciculo parvo setarum terminato. Thorax robustus. Abdomen mediocre.
Tibie antic breves, in medio calcari armate. Tarsi antici elongati, pulvillo mediocri, unguibus
acutis, curvatis, simplicibus, setisque duabus elongatis apicalibus instructi. Ale sat magne, antice
trigonz, margine apicali fere recto, apice acuminato producto ; postice ovali-trigone, tenaculo (seta
crassa curvata prope basin costz) anticis ligatz: antic vena postcostali paullo ante medium ale
bifurcata, ramo antico ramulos tres emittente, quorum tertius (6 3) apicem ale attingit, ramo
postico etiam tres emittente ; vena mediana ramulos quatuor emittente, quorum superior (seu disco-
cellularis, c 3*) cum inferiore (65*) ven postcostalis venula nulla transversa conjungitur : ale
posticse vena postcostali bifida, ramulis hberis et cum venulis mediane haud conjunctis, vena mediana
e basi bipartita, singula parte etiam bifida, inde rami 4 mediani; vena anali et subanali simplicibus.

TASCINA ORIENTALIS. (Plate XX XIII. fig. 5.) Alis anticis saturate fuscis, fascia fere
recta obliqua pone medium costz fere ad angulum internum extensa albida, alis
posticis obscure aurantiaco-rufis, imbo lato fusco; corpore nigricante, abdominis
apice rufescente. Long. corp. unc. 14. Expans. alar. unc. 34.

Hab. Singapore (Wallace). In Mus. Hopeiano Oxonie.

Genus V. Hecatresta. (Plate XX XIII. figs. 1—4.)
Hecatesia, Boisduval, Monogr. Zygen. p.11; Walker, List Lep. Het. B. M. i. p. 54.

Corpus sat crassum, abdomine in maribus barbato. Caput parvum, ocellis duobus instructum. Antenne
elongatz, cylindricze, pone medium in maribus fusiformes vel subfusiformes, in foeminis minus
incrassate. Palpi ultra medium faciei assurgentes, dense hirsuti, articulis tribus, primo crassiore
curvato, tertio dimidium secundi paullo superante. - Lingua elongata spiralis. Thorax valde hirtus.
Ale, insecto quiescente, tectiformes, breves; anticee in maribus spatio magno reniformi vitreo
transverse striolato, prope costam, instructz, hac in medio in tuberculum corneum elongatum dila-
tata, ramos venz postcostalis dislocante, scil. ramo primo curvato marginem posticum arez vitre
formante, ramis 2, 3 et 4 subapicalibus ad basin inter se zque distantibus, vena quinta prope basin
arez vitreze emissa, vena cellulam discoidalem claudente fere indistincta at curvata; vena mediana
4-ramosa, ramis tribus ultimis ad basin valde approximatis: ale anticee in feemina area vitrea
earentes; cellula magna discoidali, apice fere recta transversa, cellulaque parva angusta subcostali
distincta, ad apicem ejus ramos tres emittente, medio bifido: ale posticz cellula discoidali ad
trientem alee extensa, ibique transverse et indistincte desinens, vena 3 ¢* e medio ejus emissa. Pedes
satis graciles, tibiis barbatis ; tibiee antice: calcari forti medio instructee.

This curious Australian genus was first described by Boisduval, and placed by him in
the family Zygzenidze, but removed to the Agaristides in his recent memoir on that tribe
published in the ‘ Revue de Zoologie,’ 1874, in which position, between Husemia and Aigo-
cera, it was placed by Walker. It appears to me, however, to be more nearly allied to
Castnia by means of the genus Synemon, agreeing with Castnia in the possession of the
small narrow subcostal cell formed by the branches of the postcostal vein, the fourth
of which arises from the third at some distance beyond the extremity of this small cell,
in the normally formed wing of the female. Until, however, we become acquainted with

200 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

the transformations of Hecatesia, and have learned whether the larva is xylophagous,
radicivorous, or phyllophagous, and the pupa armed with rows of recurved spines or
simple, we cannot decide on the true relations of the group.

1. HecatTesiaA FENESTRATA. (Plate XXXIIT. fig. 1.) Hem. thorace nigro-fusco, albo
trivittato; alis anticis nigro-fuscis, fasciis duabus transversis parum macularibus,
prima ante, secunda pone medium alarum; alis posticis luteo-fulyis, limbo lato
punctoque medio subcostali nigris; abdomine fulvo-luteo, maris fusco-barbato, serie
dorsali punctorum nigrorum. Expans. alar. antic. lin. 14-18.

Obs. The female, here figured for the first time, differs from the male only in wanting
the vitreous spot on the fore wings. |

Hecatesia fenestrata, Boisduval, Monogr. Zygen. p. 11, pl. 1. f. 2; Voy. de lAstrolabe, Ent. p. 171;
Spec. Gén. Lépid.i. pl. 10 B. fig. 7; Mon. Agaristides, Rev. Zool. 1874, and sep. issue, p. 23;
Chenu, Encycl. Hist. Nat. Pap. 236, f. 403 ; White, in Grey’s Journ. Exped. Austral. Append. p. 475,
pl. 8. f. 2; Walker, List Hep. Het. B. M. 1. 54.

Hab. Nov. Hollandia. In Mus. Britann., Hopeiano, &e.

2. HECATESIA THYRIDION. horace fusco-nigro, albo trivittato; alis anticis fusco-nigris,
fasciis tribus transversis macularibus guttisque nonnullis subapicalibus albis; area
vitrea in maribus angustiore quam in precedente specie; alis posticis aurantiacis,
limbo lato albo guttato punctoque subcostali mediano nigris ; abdomine supra auran-
tiaco, maculis nigris dorsalibus et lateralibus nigris ; subtus albo; palpis et pedibus
nigris, albo variis; antennis in utroque sexu pergracilibus. Expans. alar. antic.
bin, IP

Hecatesia thyridion, Feisthamel, in Voy. de la Favorite, v. Suppl. p. 19, pl. 5. fig. 1; White, Append. Capt.
Grey’s Journ. Exped. Austral. p. 475, pl. 8. f. 1 (m. andf.) ; Boisduval, Mon. Agaristides, p. 24;
Walker, List Lep. Het. B. M. i. p. 55.

Hab. Nova Hollandia, Albany (Brewer), Swan River (Curtis), West Australia

(Damell). In Mus. Britann., Hopeiano Oxonie.

3. HECATESIA EXULTANS. (Plate XXXIII. figs. 2m, 3/f.) Atra: capite et thorace flavo
suttatis; abdomine serie dorsali punctorum fulvorum, lateribus fulvo nigroque
variis : alis anticis nigris, maris margine antico in medio dilatato, pone medium pro-
funde emarginato, feeminze forma ordinaria, in utroque sexu guttis numerosis
flavis et griseis, fascias septem irregulares maculares alternatim flavas et griseas for-
mantibus ; fimbriis omnibus griseis nigro maculatis: alis posticis aurantiacis, limbo
lato maculaque trigona subcostali mediana nigris: capite subtus orbitu oculorum
et basi palporum flavo hirtis: pedibus nigris, tibiis fulvo hirtis, apicibus et tarsorum
artic. 1 et 2 apicibus albis; calcaribus longis nigris: abdomine pone medium ma-
culis sex transversis albis (per paria dispositis) : alis anticis subtus absque squamis
griseis; posticis guttis nonnullis albis in limbo nigro. Expans. alar. antic. cire.
une. 1,

Hecatesia exultans, Walker, List Lep. Het. B. M.1. p.58; Boisduval, Mon. Agarist. p. 24.

GENUS CASTNIA AND SOME ALLIED GROUPS. 201

Obs. The typical specimen of this species, described by Mr. Walker as a male, is a
female, wanting the vitreous spot of the fore wings, which exists in the true male, the
wings being of a remarkable form in the latter sex.

Hab. Swan River, Nov. Holl. (Diggles). In Mus. Britann., Hopeiano Oxonie.

The insects described in the remaining portion of this memoir have been regarded by
various authors as belonging to the group of which Castnia is the type—from which,
however, they are entirely distinct, belonging to other Heterocerous groups.

Genus OTHria, Westw.

Dr. Boisduval’s species of Orthia, O. augias and O. nexa, must be referred to the
Agaristides, the veining of their wings agreeing exactly with that of Agariste. I propose
to place them therefore as a separate genus of Agaristides, under the name of Othria.
From Zgocera they are distinguished by their geographical distribution being con-
fined to South America.

In O. augias (Plate X XIX. figs. 18, 19) the fore wings have a small postcostal cell,
the branches 62 and 43 arising far beyond the cell, as does also 05, whilst 05* springs
from a little branch from the middle of the hinder margin of the postcostal cell, and the
discoidal cell of the hind wings is imperfectly closed by a very slender arched disco-
cellular vein or, rather, fold.

1. OruRia avetas. “ Ailes noires sans aucun reflet; les ailes supérieures, dont les
nervures forment des stries grisatres longitudinales sont traversées, du milieu de
la céte a l’angle interne, par une bande d’un jaune @ocre, trés-droite. Ailes infé-
rieures sans taches, avec les nervures un peu plus claires.” Exp. alar. ant. une. 22.

Orthia augias (Boisduval, MS.), Herr.-Schaffer, Aussereur. Schmett. f. 18; Boisduval, H. n. Lép. Hét.
p- 542 (incorrectly referred in his text, but with doubt, to Orthia pelasgus).

Hab. Brasilia (Rio Janeiro), Minas Geraes (Rogers), Brasilia occidentalis (Rogers).

In Mus. Hopeiano Oxoniz, Boisduval.

2. OTHRIA NEXA. “ Hlle a le port et la taille de O. augias; ses quatre ailes sont d’un

noir terne, sans aucun reflet, avec l’extrémité des nervures grisatre comme chez
augias. Lesailes supérieures sont traversées, du milieu de la céte a l’angle interne,
par une bande d’un jaune d’ocre trés-pale un peu recourbée a son extrémité comme
dans pelasgus ; les ailes inférieures sont sans taches. Cette espéce pourrait bien étre
une variété femelle d’augias. Hlle ne différe que par la bande des ailes supérieures
notablement plus pale, formant un tant soit peu le crochet a son extrémité inférieure,
et par le dessous des ailes supérieures dont la céte est piquée de gris.”

Hab. ——? In Mus. Boisduval.

Orthia neva, Boisduval, H.n. Lép. Hét. p. 543.

202 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

3. OTHRIA AMAZONICA, Westw. Alis brevioribus, latioribus, anticis subtrigonis; atra,
alis fascia fulva e medio costee fere ad angulum analem extensa, postice nonnihil
incurva, alis subtus omnino concoloribus. Expans. alar. ant. une. 1, lin. 11.

Hab. Amazonia. In Mus. Hopeiano Oxonie.

Had not Dr. Boisduval described an insect as the probable female of O. augias, with
erey veins at the extremity of the fore wings as in the male, I should have been induced
to regard this Amazonian insect as the female of O. augias. The veins of its wings are
arranged exactly as in O. augias.

4. Orarta Linpier. Atra, collo, abdominis lateribus, alis anticis, macula magna basali
trigona alteraque transversa ovali pone medium alisque posticis (limbo lato nigro
excepto) fulvis, subtus magis stramineis. Expans. alar. 273 une.

Agarista Lindigii, Felder, Lep. Voy. Novara, pl. 107. f. 6.

Hab. Villa Nova Amazonize. In Mus. Hopeiano Oxonie.
Obs. Individua e Nova Granada et Ecuadoria allata paullo minora (exp. unc. 1g) macu-
lisque fulvis minoribus.
Obs. Individuum majus (exp. unc. 24%;) maculis albidis in alis anticis, alisque posticis
pallidioribus, limbo nigro angustiore.
In this species the second branch of the postcostal vein arises near the upper end of
the small accessory postcostal cell. In the hind wings the veins are arranged as
in O. augias.

5. OraRIA AMALTHEA. Atra, annuio colli, lateribus abdominis, puncto trigono in medio
fasciaque tenui abbreviata pone medium alarum anticarum, alis posticis (limbo
lato nigro excepto) fulvis. Expans. alar. antic. unc. 144.

gocera amalthea, Dalman, Analecta Entom. 49. 24; Walker, Cat. Lep. Het. B. M. i. p. 58.

Hab. Brasilia, Peru, Bogota? In Mus. Hopeiano Oxoniz.
“In this species the fore wings are more elongated, and the second branch of the post-
costal vein of the fore wings arises exactly at the tip of the small accessory post-
costal cell.

6. OTHRIA coLUMBINA. Tota atra, fascia obliqua parum flexuosa e medio costx fere
ad angulum analem extensa, fulva, postice parum latiore. Expans. alar. antic.
une. 2-(5-

Hab. Columbia (Chesterton), In Mus. Hopeiano Oxonie.
The wing-veins are arranged as in O. amalthea.

7. OTHRIA ECUADORINA. Corpore cum collo nigris, alis latioribus atris; anticis fascia
tenui recta obliqua pone medium nec marginem costalem nec angulum analem
attingente, maculaque magna ovali discoidali alarum posticarum aurantiacis, margine
anali nigro. Expans. alar. antic. unc. 2.

Hab. Eouador. In Mus. Hopeiano Oxoniz.

GENUS CdASTNIA AND SOME ALLIED GROUPS. 203

This species agrees. with the preceding in its black and orange colours; but it wants the
small accessory postcostal cell, and the branch 6 2 arises at half the distance between the
discoidal cell and the tip of the wing, followed at a short distance by 63, whilst 55 arises
from the hind edge of the main stem, slightly in advance of the origin of 62. (See Plate
XXIX. fig. 23.) In other respects the veins agree with O. augias.

Several other species, evidently congeneric with the above, are figured by Dr. Felder
in the 137th plate of his work on the Lepidoptera of the ‘Novara’ voyage, under the
names of Agarista Doleschallii, A. Semperi, A. Moorei, A. alethe, and A. Batesii.

The following insects must also be removed from their proximity to Castnia and
referred to the Agaristides.

Genus Damas, Boisduval, Voy. Astrol. pt. i. p. 259.

The insects of this genus are black and orange like the Othrie, but are natives of the
Kastern Archipelago.

1. Dantas ELEGANS, Boisduval, Voy. Astrol. pt. i. Lép. 259. 1.
Hab. New Guinea.

2. DAMIAS MELAXANTHA, Boisduval, Voy. Astrol. pt. i. Lép. 260. 2.
Cleis posticalis, Guérin, Voy. Duperrey, Atlas Ins. pl. 18. f. 5.
Hab. Port Praslin, New Ireland.

3. Damas DicHROA, Boisduval, Voy. Astrol. pt. i. Lép. 260. 3.
Hab. Offack and Bourou.

4, Damas varia, Walker, List Lep. Het. Brit. Mus. i. p. 15.

Hab. Richmond River, Australia.

I cannot distinguish this insect specifically from an insect collected in Batchian by
Mr. Wallace, now in the Hopeian Museum at Oxford, described by Mr. Walker under
the name of Burgena transducta.

Genus BureGEna, Walker.

BuURGENA TRANSDUCTA, Walker. (See Plate XXIX. fig. 21.)

The fore wings are narrow; but in the veining of its wings it agrees with Agaristu,
the small accessory postcostal cell emitting only one branch at its extremity, the branch
6 2 arising at a considerable distance beyond the cell, whilst 65 arises from the hinder
edge of the stem close beyond the accessory cell.

Genus Hrspagarista, Walker.

HESPAGARISTA INTERJECTA, Walker, List Lep. Het. B. M.i. p. 14.

Brought by Mr. Angus from Caffraria, and figured by him in his fine work on the
Kaffirs, pl. 30.f.10. Agrees with the Othrig in general form and antenne ; but the wings
are much more maculated, the extremity of the body is tufted, and the veining of the

SECOND SERIES.—ZOOLOGY, VOL. I. 25

204 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

fore wings differs with respect to the small postcostal cell, which emits one branch, 05, at
its extremity, whilst the main stem of the postcostal vein arises from its anterior edge
near the extremity, b 2 arising at a little distance beyond the cell, followed at a very
short distance by 03, 04 extending to the extremity of the wing. (See Plate XXIX.
fig. 22.)

Genus RorHta.

Another group of species, also closely allied to the preceding, are natives of Mada-
gascar, and have the small postcostal cell very narrow (thus differing from the typical
Eusemie, in which the small cell is wanting, Pl. XXIX. f. 20). They also have the
branch 6 2 arising from its upper extremity, and running free nearly to the tip of the
wing, 03 arising halfway between the cell and the tip of the wing, whilst 0 5 arises
close beyond the extremity of the postcostal cell; such at least is their arrangement
in C. pales and simyra. They may advantageously be formed into a separate genus,
to which the name of Rothia may be applied. Here belong

1. RovHiA PALES.

Agarista pales, Boisduval, in Guérin, Icon. R. An. pl. 83. f. 1; Faun. Ent. Madag. pl. 10. f. 1, 2.
Eusemia pales, Boisduval, Mon. Agarist. p. 75. no. 85; Walker, List Lep. Het. B. M. 1. p. 54.

Hab. Vicinity of Tananariva, Madagascar. In Mus. Hopeiano Oxoniz.

2. ROTHIA ERIOPIS.
Agarista eriopis, Boisduval, in Herrich-Schaffer, Auss. Schm. f. 31; Mon. Agaristides, p. 71 (Eusemia er.).

3. ROTHIA AGRIUS.
Agarista agrius, Boisduval, in Herrich-Schiffer, op. cit. f. 83; Mon. Agaristides, p. 72 (Husemia agr.).

4, ROTHIA PEDASUS.

Agarista pedasus, Boisduval, in Herrich-Schiffer, op. cit. f. 32 (nec 34 ut in Bdv.); Mon. Agaristides,
p- 73 (Eusemia ped.).

5. Rorura stmyra. Atra, capite et collo flavo-guttatis: alis anticis elongatis nigris,
cutta minima humerali fasciaque transversa parum sinuata (postice paullo latiore) e
medio costee fere ad angulum analem extensa flavescentibus; alis posticis atris,
fimbria ad angulum externum alba; corpore infra, pedibus, et abdominis lateribus
lutescentibus ; alis posticis subtus dimidio basali rufo. Expans. alar. une. 23.

Hab. Madagascar. In Mus. Hopeiano Oxonie.
The arrangement of the wing-veins of this species was noticed above.

Genus AIGIALE, Felder. (Wien. ent. Mon. iv. 110, 1860.)
Type Agiale Kollari (Castnia hesperiaris, Walker, List Lep. Het. B. M. pt. vii. p. 1588).

This Mexican insect is no other than the large Hesperian butterfly first mentioned by
me (in the ‘ Genera of Diurnal Lepidoptera,’ p. 506) as remarkable for wanting spurs to

GENUS CASTNIA AND SOME ALLIED GROUPS. 205

the hind tibie*, and which was subsequently formed into a separate genus by Dr.

Felder, under the name of dgiale Kollari. The arrangement of its wing-veins (Plate

XXIX. fig. 25), and, indeed, the whole structure of the insect, are entirely Hesperideous.
Specimens are in the Hopeian collection at Oxford, presented to me by Mr. Coffin.

Genus MEGATHYMUS.
Megathymus, Scudder, Syst. Revis. Amer. Butterflies, p.12 (Salem, 1872).

The type of this genus is the large butterfly figured by Boisduval and Leconte (Iconogr.
Lépid. Amér. Septentr. pl. 70), under the name of Hudamus? yucce, by whom also the
larva and pupa were represented, but in an unsatisfactory manner.

Walker, List Lep. Het. B. M. pt. 7, p. 1583, gavea short and very insufficient description
of this insect under the name of Oastnia yucce, which, he says, “ connects the Castnie
with the Hesperie. Had he been aware of the figures of the preparatory states given in
Leconte and Boisduval’s work, he would have seen that in all respects (and most especially
in the larva form) this is a true Hesperian. In its wing-veins it agrees with Agiale
Kollari, except that the angulated vein closing the discoidal cell is very slender.

Since the present memoir was presented to the Linnean Society, an admirable memoir
on this insect, with figures of its various characters in the preparatory and perfect states,
has been published by Dr. Charles V. Riley, in the ‘ Transactions of the Academy of
_ Sciences of St. Louis,’ vol. 11. January, 1876, and subsequently introduced into the
‘ Highth Annual Report on the Noxious, Beneficial, and other Insects of the State of
Missouri, by the same author, who has arrived at the opinion that the imsect is a
large-bodied Hesperian, and that it has no real relation with Castnia.

DESCRIPTION OF THE PLATES.
Puare XXVIII. |

Details of Castnia.

Fig. 1. Larva of C. eudesmia.
Fig. 2. Pupa of C. eudesmia, dorsal view.
Fig. 3. Pupa of C. eudesmia, with one of the pterothece raised up, showing the fore wing and its marks.

I
2
3
Fig. 4. Cocoon of C. eudesmia.
5. Venation of fore and hind wings of C. papilionaris.
6. Venation of fore wing of C. cacica.
Fig. 6 a. Venation of hind wing of .C. cacica.
Fig. 7. Characteristic portion of the veins of fore wing of C. mygdon.

Fig. 8. 7 i i of C. Boisduvallii.

* In ‘H. paniscus the two spines in the middle of the two hind tibie are wanting (H.-Schifter, Syst. Bearb.
pl. vi. f. 17).
2E2Z

206 MR. J. O. WESTWOOD ON THE LEPIDOPTEROUS

Fig. 9. Fore wing of C. veraguana.
Fig. 10. Characteristic portion of the veins of fore wing of C. ecwadoria.

Fig. 11. +3 » of fore wing of C. cochrus
Fig. lla. e », of hind wing of C. cochrus,
Fig. 12. : » of fore wing of C. truvilla.
Fig. 13. 33 5, Of fore wing of C. tcarus.
Fig. 14. 5 »,. of fore wing of C. Hiibneri.
Fig. 14a. ss » of hind wing of C. Hiibneri.
Fig. 15. & » of fore wing of C. Latreillei.

Note.—The same numbers and small letters are used in the above figures of the venation of the wings, as
well as in the following Plates, to express the corresponding parts in the different species.
a, costal vein; 6, postcostal vein with its five branches, 61, 62,63,64, and 65, together with
6 5*, the upper disco-cellular vein, which is here regarded as portion of the postcostal system ;
c, median vein and its branches, c 1, ¢ 2, and ¢ 8, together with c3*, the lower disco-
cellular vem, which is here regarded as portion of the median system; y and z represent the
bases of 6 5* and c3* within the discoidal cell; d, the postmedian vein; pc.c, postcostal cell ;
de, discoidal cell ; de*, hinder division of discoidal cell ; pmc, premedian cell.

PLATE XXIX.

Fig. 1. Characteristic part of the hind wing of Castnia juturna.

Fig. 2 a » fore wing of C. Marcel-Serresii male.

Fig. 3 i » hind wing of ditto ¢, and annexed partial diagram of 2.
Fig. 4 3 » fore wing C. chremes, male.

Fig. 5. A » | hind wing of C. chremes, male and female.
Fig. 6 Ss » fore wing of C. diva.

Mig. 7 Bs », hind wing of ditto.

Fig. 8. Fore wing of C. cononia.

Fig. 9. Characteristic part of the fore wing of C. linus.

Fig. 10. » hind wing of ditto.

Fig. 11. a » fore wing of C. mimica.

Fig. 12. 4 » hind wing of ditto.

Fig. 13. Fore wing of C. acreoides.
Fig. 14. Characteristic part of hind wing of ditto.
Fig. 15. 55 » hind wing of C. therapon.
Fig. 16. Fore wing of Orthia pelasgus.
Fig. 17. Hind wing of ditto.
Fig. 18. Fore wing of Othria augias.
Fig. 19. Hind wing of ditto.
Fig. 20. Fore wing of Husemia.
Fig. 21. Characteristic part of fore wing of Burgena transducta.
Fig. 22. Fore wing of Hespagarista interjecta.
Fig. 23. Characteristic part of the fore wing of Othria ecuadorina.
Fig. 24. % eS fore wing of Agarista Lindigii.
Fig. 25. Fore wing of Agiale Kollart.
Fig. 26. Fore wing of Huschemon Rafflesii.
27. Fore wing of Hepialus humili, fem.
Fig. 28. Hind wing of ditto.

Fig.

Fig.

oo

GENUS CASTNIA AND SOME ALLIED GROUPS.

. Castnia veraguana.
. C. ardalus, var.
. C. phalaris.

Castnia clitarcha, ¢ .
. C. chtarcha, @.
. C. papihonaris.

cycna.

PLate XXX.
Fig. 4.
5.
6.
PuatTe XXXII.

Fig. 4.
5.

Puate XXXII.

Fig

Puate XXXII.

207

Castnia amycus, var.
C. Dalmanni, 2 from Brit. Mus.

C. papagaya.

Castnia satrapes, var.
C. satrapes, var.

. 5. C. (G.) cononia.
6. C. (G.) ecuadoria.
7. C. (G.) mimica.

1. Hecatesia fenestrata, 9 ; 1a. Head and antenna of male; 1%. Palpus; 1c. Palpus denuded;
1d. Fore leg; 1e. Fore wing of male.
2. H. exultans 3.

3. H. exultans 2.

4. Wings of H. thyridion.
5. Tascina orientalis; 5a. Head, seen in front; 5 6. Ditto, sideways; 5 ¢. Hxtremity of antenna;
5 d. Fore legs; 5e. Extremity of fore tarsus; 5/f. Hind leg; 59. Venation of fore wing;
5h. Ditto hind wing.

Fig. 6. Synemon directa.

ihe

S. nupta.

8. S. plana.
9. S. obscurella.

Fig. 10. Synemon notha.
ll. S. hesperioides.
12. S. vagans; 12 a. Venation of wing of same.
13. S. gratiosa.

14a. Fore wing of Synemon Sophia; 146. Hind wing of ditto; 14c. Head and base of antenna;
14d. Palpus of S. Sophia; 14 e. Palpus, denuded, of ditto; 14,f. Fore leg of ditto; 14.9. Middle
leg of ditto; 14. Hind leg of ditto.

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JOWestwood del. Trans. Linn Soc.Ser.2.Zoou Vou 1.Tas.28

CLaiwetes

Fi y. 15. O Papizonarts.

et Hanhart imp.
a VENATION & DEVELOPEMENT OF CASTNIA.

JO Westwood! del. Trans Linn Soc. SER. 2.Zooi Vou. 1 Tas 29.

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Burov Transdicta.

a= sll Agurisia Limiigu. ] = ese
Euschemon Fattlesiv. 2 Aigiale Kollar:
CBerjeau lith. Hamhart imp
VENATION, SPECIES OF CASTNIA &c.

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JOWestwood. del TRANS. LINN Soc SER 2 Zoo. Vou. Tas 30

CBerjean hth. Hanhart imp

NEW SPECIES OF CASTNIA.

JOWestwood del. TRANS. Linn. Soc.SER.2.Zoon Vou.1 Tas 31.

CBerjeau lith Hamhart imp.

NEW SPECIES OF CASTNIA

JOWestwood del. Trans Livw Soc Ser.2.Zoon Vou.1 Tas. 32

OG) ruxilla

ClGleratina. —~

Cl (oz. Creer

CBerjeau. lith : : Hanhart imp.
NEW SPECIES OF (GAZERA.) CASTNIA.

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ER.C. Z00L 1. Tas. 33.

MS GLLELIDOD Sophia.

Hanhart mp.

Pred

[ 209 ]

IX. On a New Genus of Trematoda, and some new or little-known Parasitic Hirudinei.
By Joun Denis Macponatp, U.D., F.R.S., Deputy Inspector-General R.N.,
Professor of Naval Hygiene Army Medical School. (Communicated by G. E.
Dosson, M.A., M_B., F.LS.)

(Plate XXXIV.)
Read April 6, 1876.

MANY striking points of resemblance are traceable between the Trematoda and the
Hirudinei; but this is merely indicative of a representative relationship, or one of
analogy. The numerous errors into which our forefathers fell in grouping together
really incongruous things may be referred to the deceptiveness of primd facie resem-
blance, and the failing to recognize the intrinsic difference between the kind of relation-
ship here alluded to and that of a genuine affinity. This, I am quite sure, is one of the
greatest difficulties with which modern evolutional biology will have to contend. An
analogous property of gemmation in certain animals, in common with plants, being the
innocent cause, so to speak, of the dendritic or tree-like form, so deceived the earlier
observers that even their very word Zoophyte is accepted by us as a merely descriptive
term, in a sense which is precisely the converse of that which its etymological construc-
tion would indicate. The balance was so far in favour of the plants, that the doubt
gave rise to the additional complication of confounding this group with the Corallines.
Now, we know that there are also other representative relationships between the two
great organic kingdoms, such as conjugation, fission, contractility, and motion, in con-
nexion with which errors of judgment are continually creeping into the dicta of science.
I might further say that the light available to Linnzus and his immediate followers with
regard to these matters was scarcely less than that which we have at present to guide us
in the more difficult study of the primary modes of evolution observable in the ovum,
with the view of establishing the affinity or antipathy of the resultants, however different
or identical in type these latter may seem to be. Indeed, even up to the present, facts
are not wanting to show that very similar beings originate in a very dissimilar way, and
vice versd. Nevertheless we look forward with pleasure to the results that appear to be
promised to the labourers in this line of research.

- If we look upon the Trematoda as representing the Hirudinei, coupling also the
Oligocheta with the Polycheta, and the Rotifera with the Articulata, Prof. Huxley’s
interesting classification, embodying a large proportion of the results just alluded to,
would seem to sanction the idea. Thus the corresponding part of his table may be
arranged, without material change, in the following manner :—

1. ARCH ZOSTOMATA. 2. DEUTEROSTOMATA.
Trematoda. Annelida.
Oligocheta. Polycheta.

Rotifera. Arthropoda.

210 DR. J. W. MACDONALD ON A NEW GENUS OF TREMATODA,

There is surely something more than coincidence in this. There may be misgivings
as to the position of the Oligocheta; but perhaps something will soon turn up to recon-
cile it.

One or two additional points of analogy between the Trematoda and the Hirudiner
will be seen in the description of the forms which I have now to present to your notice.

The little Trematode represented in the first figure of the plate was found by me
creeping about in the respiratory siphon of a large Melo, or melon-shell, in Shark Bay,
Western Australia. It was between an eighth and a tenth of an inch in length, tallowy
white (to use an American expression), and so much resembled a leech as to deceive me
at first sight; for I had already seen white leeches, both parasitic and free, and I did not
expect to find a Trematode walking abroad. The fore part of the body was slender, taper-
ing, and depressed ; the oral sucker was subterminal, and of the usual cup-like form ; and
after a slight constriction, a bulbous pharynx was distinctly seen, from which the ceso-
phagus passed backwards for some length, to what would appear to be its bifurcation ;
and the appearance of the czeca could be traced to within a short distance of the posterior
extremity. But the ventral disk constituted the most striking feature in this little animal,
on account of its great size and complex structure. It occupied more than the two
posterior thirds of the inferior surface of the body, which was laterally compressed
instead of depressed in this locality. The disk, or virtually the foot, might therefore be
said to be narrow and elongated in form, and not unlike that of Aspidogaster, though
very different in structure. Thus, it presented four rows of alveoli, or deep sucking-pits
(fig. 3), the marginal series being the largest, with their long diameter running trans-
versely, while that of the intermediate ones was longitudinal. All the contiguous
borders of these little pits were so connected by straight bands of muscular fibres as to
divide the whole of the intervening area into triangular spaces, the centre of each of
which was ocupied by a curious tentaculiform cirrus, capable of complete retraction from
the apex, as in the common instance of the finger of a glove (fig. 4). Six rows of these
cirri, with thirty in each row, would make up 180, and four rows of alveoli, thirty in
each row, 120; so that the disk altogether presented a very remarkable appearance.

The water-vascular system, as in the Zrematoda generally, opened posteriorly. The
aperture, however, was subterminal and dorsal, or just above and within the hinder
border of the disk. The tortuous vessels of this system were seen running forwards on
either side of the body as far as the space between the pharyngeal bulb and the oral
disk, whence they turned backwards upon themselves, as shown in fig. 5. The genital
opening, as in most other cases, was situated in front of the ventral disk (fig. 1, 6).

About the same time I found a large black leech on a species of Myliobatis, presenting
at least two characters which were quite new to me—namely, a row of branchial leaflets
on each side of the body (figs. 8 & 10), and a caudal sucker alveolated (figs. 11 & 12
very much after the manner of the Tremotode already described.

The head of this creature was depressed, spatulate, and pointed in front (fig.8). The
anterior third of the body was without appendages, constituting a kind of neck, but the
rest was furnished on either side with a row of heart-shaped leaflets on short pedicles.
Over the base of these, on every fourth segment, was situate a pulsating projection,

AND SOME NEW OR LITTLE-KNOWN PARASITIC HIRUDINEI. 211

connected, no doubt, with the water-vascular system, as in Pontobdella. The rich black
body, moreover, was ornamented with six rows of white spots (fig. 10).

The alveoli of the caudal disk were disposed in rays rather than in circles, with
radiating muscular bundles between them ; they were circular and cuplike, with a central
retractile portion, reminding one of the mechanism of the suckers of the Cephalopoda.

Attached to the body in a very irregular manner, but chiefly at its fore part, were
several of the double tubular spermatophora shown in fig. 9. These curious bodies
I have also found on other marine Hirudinei, but always with some characteristic dif-
ference. Fig. 6, for example, represents a small black leech with white tubercles, refer-
able, apparently, to the genus Pontobdella, found on Rhinobatis in the same seas ;
and fig. 7 is its double-barrelled spermatophore, which is quite different from fig. 9,
though obviously of the same nature. Very little is positively known of the generative
processes of the marine leeches ; but the facts here mentioned may one day meet with a
satisfactory explanation. i

Fig. 13 is a small white leech, with two eyes, a retractile proboscis, and, in short, the
general anatomy of Glossiphonia, but characterized by possessing seven pairs of ramose
lateral branchize, taking up about the middle third of the body. It was found on the
conjunctiva of a green turtle at Huon’s Island, on the reef to the northward of New
Caledonia. Fig. 14 is another specimen of the same species, but much larger, taken in
Shark Bay; both figures are about double the natural size. Fig. 15 shows one of the
branchize as seen with a power of a quarter of an inch.

In systems of classification both the Oligocheta and the Hirudinei come under the
head of Abranchiata. But this is evidently a mistake; for, in the first place, it is difficult
to deny the branchial function of the ciliated finger-like organs surrounding the vent
in Oken’s genus Prato, of which I have found good examples in the Taro beds of the
Fiji Islands ; and, from what has been above advanced, there can be no doubt of the
existence of external branchize in the Hirudinei.

I have now only to say, by way of apology, that I have written this paper without
having the opportunity of consulting any recent researches on the subject; so that, if I
have mentioned things already known to science, they may at least be taken as corro-
borative matter.

DESCRIPTION OF PLATE XXXIV.

Fig. 1. A little Trematode, with large and complex ventral disk, found in the respiratory siphon of a
species of Melo, Shark Bay: a, oral sucker; 6, genital opening; c, ventral disk; d, external
opening of water-vascular system. x 20 diameters.

2. Posterior extremity of the disk, with cirriexserted. x 150.
3. Central portion of the disk, x 300: a, alveoli of the border; 6, intermediate alveoli; c, openings
of the cirri.
4, Small portion of the disk, x 350, to show the cirri in different conditions : a, completely retracted ;
b, partially exserted, with the apex still inverted ; c, completely protruded.
SECOND SERIES.—ZOOLOGY, VOL. I. 25

oOo CON DS Ot

ON SOME NEW OR LITTLE-KNOWN PARASITIC HIRUDINEI.

. Tortuous course, loop, and return of water-vascular canal.

. Small black Pontobdella, parasitic on Rhinobatis.

. Spermatophore of the same.

. Black leech, with branchial leaflets and alveolated caudal disk, found on Myliobatis.
. Spermatophora of the same.

10.
I).
12.
13.
14.
15.

Portion of the body of the same, slightly magnified.

Inferior surface of the caudal disk, magnified to the same extent.

Marginal portion of the disk, more highly magnified.

White leech, with lateral branchie, on conjunctiva of Green Turtle, New Caledonia.
Larger specimen of the same species, Shark Bay, W. Australia.

One of the branchiz. x 300 diameters.

Trans. Linn. Soc. Ser. Z.Zoon Vou I. Tas. 34

JD. Macdonald del.

=
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Hanhart imp

TREMATODA & HIRUDINEI

CBerjean lith

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Bullen

X. On the Osteology and Pterylosis of the Spoon-billed Sandpiper (Burynorhynchus
pygmeeus, Linn.). By JOHN ANDERSON, I.D., .R.S.E., F.L.S., F.ZS., §¢.. Curator
of Indian Museum, and Prof. Comp. Anatomy at Medical College, Calcutia*.

(Plate XX XV.)

Read June 15, 1876.

AS Mr. Harting has already fully stated + all that has been ascertained regarding the
bibliography and life-history of this remarkable little bird, the following observations
will be confined to a description of the particulars indicated in the title of this paper.
These are founded on a specimen procured many years ago at Chittagong in Hastern
Bengal, on the mud flats at the mouth of the Karnafulie river }.

The Skull.—The distinguishing feature of the skull, as is well known, is the expansion
of the distal end of the beak into a quadrangular-shaped spoon, which nearly equais in
breadth the greatest transverse diameter of the cranium. In the specimen on which these
remarks are based all the sutures of the skull have disappeared, the palatines having also
united at their postero-internal angles. The orbit is very large, as in Tringa, its
longitudinal diameter equalling the interval between its posterior margin and the oc-
ciput. The skull has the giobular and smooth character of the pluvialine group of birds.
There is a feeble lambdoidal ridge, large occipital imperfections of ossification, and a
triangular foramen magnum, with the apex of the orifice directed backwards and slightly
upwards, as in Tringa minuta. The occipital and basicranial regions are somewhat
tumid, which is also a character of the Tringa skull; and the palatine and interorbital
regions also conform to it in their general characters. The latter region, however, is
nearly flat, and is more depressed than in any species of Tringa I have examined ; and
the groove of the frontal suture is not prolonged on to it, although the interlacrymal
depression is the same as in Tringa.

The occipital area of the skull is directed upwards and backwards, and, as defined by
the lambdoidal ridge, is of very limited extent compared with the parieto-frontal capacity
of the cranium. The two supraoccipital imperfections of ossification form its most
_ posteriorly prominent part, as is the case with the generality of skulls which have these
fontanelles—so much so, that they are continuous from side to side when the skull is
viewed in profile. They are symmetrical, and are narrower above than below; and
from their lower margins a groove is continued forwards along the sides of the foramen
magnum, defined externally by the tumidity of the exoccipitals. Inside the skull this

* T am indebted to the Trustees of the Indian Museum for their permission to illustrate this paper from the
materials in their possession.
+ ‘The Ibis,’ 1869, new ser. vol. v. p. 427.
+ This is one of the “three specimens in spirits” mentioned by Mr, Harting in the paper referred to (p. 433) as
haying been sent by Mr. J. E. Bruce from Chittagong in 1856.
2F2

214 DR. J. ANDERSON ON THE OSTEOLOGY

venous groove is seen to run along the upper surface of the inward projection of the
periotic; and at its highest part it is partially bridged by an outwardly and upwardly
projecting spicule from the epiotic. In Zringa minuta there are no occipital fontanelles ;
but the groove occurs along the lower half of the side of the foramen magnum, and opens
into the skull by a small foramen ; whilst in 7. salina the fontanelles and groove are as in
Eurynorhynchus. The squamous region is slightly concave, and the postorbital margin
is the same as in Tringa, there being a stylet-like postorbital process projecting down-
wards and forwards, separated by a very shallow notch from the descending process of the
squamosal, which is deeply bifurcated and more strongly developed than in Tringa—
although, as in that genus, the temporal fossa is very feebly marked. Immediately before
the foramen magnum there are from three to four minute foramina for the cranial
nerves ; and slightly anterior to these a larger one, separated from the paroccipital con-
tinuation of the previously mentioned groove. ‘The paroccipital process in this, as in the
Limicoline skull generally, is thrown much forwards onto the base of the skull, and is
directed forwards and downwards. Its inner side is perforated by a small foramen,
which, however, does not occur in Tringa minuta or T. salina; and the apex of the
process nearly touches the lateral expansion of the basisphenoid; the upper wall of the |
imperfect ring so formed is partially divided into two separate channels. Although the
orbit has the same general characters as in Tringa, the septum is more complete, having
a greater expansion superiorly. The ethmoidal ale are also more strongly developed
than in that genus, their superior and inferior external angles being more marked and more
outwardly projecting. The lacrymal is also more pronounced and formed somewhat
differently from that of Tringa, in which the downwardly prolonged portion that abuts
against the ethmoidal alee is a very fine osseous thread; while in Hurynorhynchus the
curve is less and the bone is stouter, and a sharp, but short, process is given off from the
outer margin, where it turns into the ethmoidal plates. The superior or orbital angle
of the laerymal is also stronger and more outwardly projecting than in Tvringa. The
nasals have a small anteriorly projecting lamina developed from their external margins
in a line with the posterior border of the nostril, and which does not occur in 7. salina
and 7. minuta. The quadrate bone, although proportionally larger, has the same form
asin Tringa; but it differs in having the articular surface for the mandible more deeply
concave, and, in the inner: side of the same surface having a much shallower notch.
The air-cells open behind, and between the proximal facets, by a round orifice, which
appears to be occasionally supplemented by a smaller one, and on the inside of the bone
by two or three such openings. The pterygoid has the general form and relations of the
Schizognathous. skull, but is much stronger than in Tringa. The basipterygoid process
is more produced than in that, genus; and there is a small process posterior and internal
to it, which, does. not occur, or is only feebly developed, in Tringa.

The palate is essentially Schizognathous; but, as already mentioned, the palatines have
ankylosed in the mesial line, and their vomerine processes have amalgamated with the
vomer, which is short and bifurcate at its tip, and does not extend forwards beyond the
origin of the nasal process of the maxilla. The palatines have much the same form as
in Charadrius pluvialis. The external lamina is at a slightly lower level than. the

AND PTERYLOSIS OF EURYNORHYNCHUS PYGMAUS. 215

internal, although the under surfaces of the palatines are but slightly concave. A notch

occurs behind the middle of their internal margins; and from it posteriorly a small de-
pendent lamina is prolonged backwards, gradually diminishing in size towards their
postero-internal angles. he concavity of the palatines is continuous with the notch.
In Tringa the notch is more strongly marked, and there is a contraction of the external
margin of the palatine in its neighbourhood, the dependent plates also in that genus
being more developed than in Hurynorhynchus; and the portion before the notch is like-
wise more concave. An anteriorly directed foramen occurs near the inner border a
short way before the notch. The maxillo-palatines in no way differ from those of Tringa.
The internasal septum has a large imperfection of ossification in its centre; and its
inferior border is jagged, terminating under the middle of the aforementioned lateral
lamina of the nasals. It has much the same capacity as in Tringa, in which, however,
its lower border slopes upwards in an unbroken line. The jugals and maxille call for
no remark.

As already stated, the distinctive feature of the skull is the great development of the
‘distal extremities of the preemaxille into a broad four-sided spoon—two anterior and
two lateral borders, the former meeting at an obtuse angle with
each other, and at an acute angle with the latter. It is
marked by a longitudinal mesial groove on its under surface,
and by a corresponding low ridge on its dorsal aspect. At
the point of divergence, where the nasal and palatine pro-
cesses are given off, there is an orifice, which, penetrating
between the plates of the premaxille, divides into a great
number of canals for the transmission of the nerves and blood-
vessels. These canals appear on the surface as so many fine
ridges, radiating forwards and outwards from the point of
entrance, and open along the anterior and lateral margins of fig. 1. Bill of Eurynorhynchus
the spoon, more especially the former. On the under surface ipo gpocers ta puoi,
there is one well-marked regular row, with a few somewhat Fig. 2. Upper view, head of
scattered foramina behind it; and on the same surface, on a ee sve een ePite
slight eminence, external and separated from the mesial .
furrow by a slight concavity, there is, on each side, a row of from 6 to 8. foramina.
A similar structure prevails in the mandible; but the positions of the ridges. and orifices
are reversed. A similar but less-intensified structure also prevails in Tringa..

Vertebral Column.—There are 15 cervical vertebree, as in Tringa. The atlas is a
narrow ring, with the odontoid ligament strongly ossified, and a pair of. obscure processes
on the under surface of the segment. In Tringa minuta the ligament is unossified, and
the two hypapophyses are much more developed. The axis has a thick nodose spinous
process, with a falcate backwardly projecting under process, and with two nodose processes,

Fig. 1.

* Through the kindness of Mr. Harting and the Editor of ‘ The Ibis’ the use of the above woodcuts has been
obtained. In the paper in ‘ The Ibis,’ 1869, Mr. Harting mentions them as being from the pencil of Mr. Swinhoe.
They are here introduced that the head with feathering of this singular bird may conveniently be compared with.
the bones of the same, given in Plate KXXXY.

216 DR. J. ANDERSON ON THE OSTEOLOGY

one on either side of its base. In the same species of Tringa the two spines have much
the same form as in Hurynorhynchus, but the nodose processes are absent in the inferior
one. All the remaining vertebrae of Hurynorhynchus, in their number, even to the
minute details of their structure, conform to the same parts in Zringa. The ribs also
call for no remark.

Shoulder- and Wing-bones.—The scapula is rather more dilated in its posterior half
than in Tringa minuta; and the suprascapular border is bevelled off to a point from its
lower margin, whereas in the latter species it is less pointed. The coracoid has much
the same proportion to the scapula and to the total length of the sternum as in Tringa ;
the head of the bone has the same form as in 7. minuta; similarity also prevails in the
shafts and distal ends; and both birds have hooked epicoracoid processes of one type.
The clavicles are the same as in the species of Sandpiper compared.

The sternum only differs from the sternum of Zringa in the anterior border of the
keel being less forwardly curved. The notches are the same in both forms; and a
similar imperfection of ossification exists at the anterior superior border of the keel.

The humerus is proportionally longer to the antibrachium than in 7. minuta. From
the humerus of the latter it is also distinguished by the feeble ridge occurring on the
preaxial side of the proximal extremity of the humerus of that bird being developed in
Eurynorhynchus into a strong triangular anteriorly curved process. The antibrachium
and manus have the same relative proportions as in Trimga.

Pelvis and Hind Limb.—The chief difference that exists between the sacrum of the
Spoon-billed Sandpiper and that of 7. minuta is the slightly greater dimensions of the
ilio-sciatic foramen. In Hurynorhynchus there is narrowing of area immediately below
the foramen, and the long free portion of the pubis is given off nearer to the acetabulum,
whilst it is more divergent from the ischium. The femur is very little more than
one half as long as the tibia, being proportionally shorter than in 7. minuta, the femur
of which it otherwise closely resembles, although somewhat stouter. The cnemial
process of the tibia is the same in both birds; and the perfect fibula of Hurynorhyn-
chus is more than one half as long as the tibio-tarsus. In all other respects the hind
limbs of the two genera are alike.

The Hyoid Arch.—The entoglossal cartilage is considerably longer and stouter than in
Tringa salina; and the entoglossal bone is shorter and broader, with a much larger
imperfection of ossification than in that species; but im other respects the remaining
elements are alike, only larger in Hurynorhynchus, owing to its greater size.

The Viscera.—in Tringa salina the intestine measures 7°75 inches, and in Huryno-
rhynchus pygmeus 8°75 inches. In the former the large intestine is 0°78 inch, and its
cecal appendages 0°62 inch long, while in the latter the great intestine is 0°88 and its
excum 0:70 long. The stomach and proventriculus are the same in these two forms.
The liver of Hurynorhynchus appears to be somewhat smaller, and differs from that
of Lringa salina in having the right lobe flattened externally, and much more pointed at
its lower end, which terminates in a sharp projection, this being somewhat the character
of T. minuta. Much importance, however, cannot be attached to such slight hepatic
differences, because these are liable to occur in individuals of the same species. The

AND PTERYLOSIS OF EURYNORHYNCHUS PYGMAUS. 217

tongue is longitudinally grooved, and fringed at its extremity in the same way as in
Tringa; but it is nearly thrice as broad as in that genus. It rapidly diminishes in
breadth towards its extremity, and contracts to a sharp point.

Pterylosis.—There are 12 rectrices and 24 remiges, of which the first two are the
longest and of equal length, the pinion adhering to the Tringa type in having ten
remiges. The wing, however, is more deeply bifid than in Tringa minuta or T. salina;
this is owing to the greater length of the posterior secondaries, and to the slightly shorter
character of the feathers intervening between them and the first primaries, the 23rd and
the 24th feathers of the wing being of nearly equal length.

The spinal tract follows the Limicoline type. The interhumeral clefts are much more
strongly marked than in 7. salina; and the posterior part is feeble at its commencement,
and its two halves meet at the tail. The spinal space is thus long and lanceolate. In
the anterior portions of the posterior clefts the feathers are arranged obliquely from
before backwards and outwards, in rows of from 3 to 4 feathers, which become more
and more longitudinal in their course, till at last, in the undivided extremity of the
spinal tract the feathers are in transverse rows of generally 5 feathers—the rows
being slightly backwardly convex, and the feathers diminishing in numbers as the tract
is traced backwards.

The oil-gland is pyriform and bare, with the exception of a circlet of feathers around
its orifice. The ventral tract is divided by the ventral space immediately before the
sternal keel; but in Tringa salina the division occurs slightly anterior to that. Its
branch, or pterylosis lateralis, is broad and strong, as in 7. salina, and is prolonged.
aleng nearly two thirds of the chest. The truncal branches are rather feeble at first,
but are stronger and broader on the abdomen, as in 7. salina. The humeral tract is.
strong, and at its origin unites with the last mentioned. The femoral or lumbar tract
originates from the posterior extremity of the spinal tract, as in Tringa, and has the
same general character. The lateral neck-space extends two thirds up the neck; and the
other spaces are the same as in Vringa and its allies.

Concluding Remark.—The foregoing examination of the osteological and other cha-
racters of Hurynorhynchus reveals only one important feature wherein this bird struc-
turally differs from the genus Tringa, namely the singular expansion of the bill, the
structural equivalent of a similar modification in Platalea leucorodia. Indeed this
species, as pointed out by Mr. Harting (/. c.), was originally placed in the genus Platalea
by Linnzeus *.

* Mus. Ad. Frid. i. Prodr. p. 26 (1764).

218

Fig. 1.

OSTEOLOGY AND PTERYLOSIS. OF EURYNORHYNCHUS.

DESCRIPTION OF PLATE XXXV.

Skeleton of the Spoon-billed Sandpiper (Eurynorhynchus pygmeus),in profile. Of natural dimen-
sions; and all the other illustrations are also drawn to the same scale.

. The skull, seen from above, and displaying the expansion of the premaxillaries and their

foramina.

. The palatal or lower base of the same cranium.

. An upper view, or oral surface of the (lower) mandible, its terminal width corresponding in

outline to the premaxillaries.

. A side view of the hyoid bone.

. The bones of the right wing, on their outer or dorsal surface.

. The right humerus, its imner aspect.

. Femur of right side, seen from in front.

. Tibia and fibula of same leg, also their anterior aspect.

. Tarso-metatarse of right limb, front surface.

. Pelvis and caudal vertebre, viewed from above. ‘The posterior rib of the right side remains.

. An inferior view of the sternum, furcula, coracoids, and costo-vertebral parts of the chest of

Burynorhynchus pygmeus.

1

S ANLETSO?, TRANS. LINN. Soc. SER. 2 Zoou. Vou I, Ph 35.

OSTHOLOGY OF THE
a, Lith THE SPOON-BILLED SANDPIPER, Banks & Co, B
HURYNORHYNCHUS PYGMABUS.

THE
TRANSACTIONS

OF

THE LINNEAN SOCIETY

OF

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART THE FIFTH.

LONDON:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:
SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXVII.

TOUS 28 Lire noose

oP a a ae et

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ROU) CLG IU RRRRRI be te a em ase

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LEE Oe Ue

CONTENTS.

PART V.—Ocrosun, 1877.

/XI. On Steere’s Sponge, a new Genus of the Hexactinellid Group of the Spongida.
By James Movriz, WD., FLS., F.GS., Fe. (Plates XXXVI. & XXXVII.)
page 219

/XII. Notes upon the Oxystomatous Crustacea. By EpDwarp J. Mirrs, 7.L.S., F.Z.S.,
Assistant in the Zoological Department, British Museum. (Plates XX XVIII.

IN MY ee JRE Es OE TG cute CRO UM Ie baenng halt 1.935

vXIII. On the Domestic Pig of Prehistoric Times in Britain, and on the mutual Rela-
tions of this variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andaman-
ensis, and Sus barbatus. By Grorce Rouixeston, UD. PRS, F.L.S.,
Linacre Professor of Anatomy and Physiology, Ozford. (Plates XLI.-
XLII.) MR iit Rg Nee id tee ag Sol me e OONL

vXIV. On two new Forms of Deep-sea Ascidians, obtained during the Voyage of H.MU.S.
‘ Challenger. By H. N. Mosstzy, IA., F.R.S., Fellow of Exeter College,
Oaford. (Communicated by Dr. J. Murtsz, F.LZ.8.) (Plate XLIV.) . . 287

/XV. On new Forms of Actiniaria dredged in the Deep Sea; with a Description of
certain Pelagic Surface-swimming species. By H.N. Mosetey, V.A., F.R.S.,
Fellow of Exeter College, Oxford, late Naturalist on board H.M.S. ‘ Chal-
lenger. (Communicated by Dr. J. Muniz, F.L.S.) (Plate XLV.) . . 295

teh) RE Ay bee

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| hy Sor ery IBS aa i vi ty urs pig eh if KR: Lay L hae 1M radish sat
; Dre lena Teer rhc oenteng s if. aon Dewey weed: % cotai hh tii Rag
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Al oo) Woes LA EATS LL DROME era M A ee? ail ie ih OH 43

Fe 8

XI. On Steere’s Sponge, a new Genus of the Hexactinellid Group of the Spongida.
By James Muniz, ICD., F.LS., F.GS., Se.

(Plates XXXVI. & XXXVI.)

Read January 20, 1876.

THE Order of Sponges now designated Hexactinellida has only of late years been
prominently brought forward to the notice of naturalists. As early as 1833 by Quoy and
Gaimard, and again more definitely in 1841 by Stutchbury, at least two or three forms
of the group had been registered in zoological literature ; and later on a few new members
were occasionally added. From about 1860, or even a few years later, up to the present
time a number of new and most curious genera and species have been minutely described.
'The issue of the monographs of Bowerbank, Oscar Schmidt, and Hackel, among others,
have greatly helped to advance and give prominence to the history of the Spongida; but
the study more especially of the Siliceous Sponges, however, received an unexpected
impetus when the first results of the deep-sea explorations became known. Besides
Dr. Gray, ever active in many fields, the labours of Carter, Wyville Thomson, Saville
Kent, and Perceval Wright, among our own countrymen, have borne good fruit so far as
the group in question is concerned. Not only has the deep-sea dredging yielded a partial
resolution of some of the difficulties of structural organization of certain of the more
singular kinds, but another item of considerable importance has become evident, as fresh
faunal ground has accrued: I allude to distribution. The geographical dispersion, then,
of the Siliceous or, as some prefer to call them, Vitreous Sponges, where unmistakable
closely allied kinds are widely severed, suggests many possibilities, geological and other-
wise, though no explanation is capable of easy proof from our as yet limited data. Every
new genus or species entered into the catalogue thus has a cumulative significance,
bearing ratio, moreover, to its peculiarities of build, where these either exhibit novelty,
intermediate characteristics, or set at rest moot points. On grounds such as these the
noyel specimen I herewith call attention to I believe possesses unusual interest.

History or Spectmen.—Professor Joseph B. Steere, M.A., LL.B., of the University of
Michigan, U.S., a few years ago resolved on enriching the Museum collections of his
alma mater. Young, enthusiastic, and fond of adventure, after an exploration and
somewhat lengthened stay in the regions of the Amazon, sending home valuable material,
he traversed the continent, crossing the Andes. Thence by the Pacific Ocean towards
China, he remained awhile in Formosa. Afterwards making his way to the Philippines,
he resided for longer or shorter periods in several localities of the group. In devious.
voyages, visiting the more notable islands of the Malay archipelago, he gathered a vast
array of objects of Natural History. Time speeding, he betook himself homewards, and
en route made a hasty stay in London, during the beginning of the month of October 1875.

Circumstances did not permit of his boxes and cases of material being unpacked here,
SECOND SERIES.—ZOOLOGY, VOL. I. 26

220 DR. J. MURIE ON STEERE’S SPONGE,

a few loose packages only being opened to gratify the curiosity of friends. Mere accident
in conversation led me to a sight of a sponge the subject of the present communication.

Its appearance and history in a trice convinced me of its rarity, and I advised the
owner to take advantage of the extensive collection at the British Museum, and after _
comparison &e. to describe the specimen. If ultimately deposited in the distant Michigan
Museum, the chances were its value and uniqueness might temporarily be lost sight of
among the plethora of his collections, and a long while elapse ere opportunity offered to
bring it before the scientific public. Neither time nor indeed inclination to a special
study of the group rendered Professor Steere desirous thereupon to enter on the task;
but still, knowing the interest attached to the object, he requested my drawing out a
notice of it for publication. This could have been little else than a mere superficial
survey had he not given me possession of the tempting specimen for a brief period, to
facilitate closer inspection and to procure a photograph of it. He still more generously
gave me a portion, quite enough for a thorough examination of its microscopic structure,
as the accompanying illustrations bear witness.

The following is Professor Steere’s verbal account to me of how and where the sponge
was obtained. When in the Philippines, 1875, his attention one day was called to what
at first sight he deemed to be a branching coral, hanging in the sun near one of the
native dwellings. On purchasing it, he learned from the fisherman, its first possessor,
that it had accidentally been dragged up by his hooks* from a depth of what he supposed
to be 400 to 600 feet, 7. e. roughly, then, between 70 and 100 fathoms. The locality where
originally obtained is the narrow deep channel between the strip-like and parallel islands
of Negros and Zebu. This channel is said to be some 100 miles long, and varies from
2 to 3 miles in breadth. The water is comparatively still, or with only a slightly percep-
tible current, never, generally speaking, being boisterous. This condition in the main
results from the construction of the channel, which to some extent is land-locked, as
reference to a chart of the district shows.

As regards the nature of the sea-bottom, Professor Steere believes it to be sandy. As
to the geological aspect of the surrounding country, according to him the island of Zebu
is chiefly characterized by a limestone formation ; but the larger island of Negros bears
apparently marks of volcanic origin.

He further informed me that the Huplectella and other peculiar sponges hitherto

* Dr. Meyer (Ann. & Mag. Nat. Hist. (4) 1874, vol. xiii. p. 67) has drawn attention to the mode of fishing for
sponges adopted by the natives of the Philippines. A number of small hooks are fixed to a light framework, and this,
sunk by a stone, is dragged on the bottom in depths quite 50 fathoms or more, and the hooks catch the objects as the
boat drives with the current. A confirmation of this has been published by one of the staff of the ‘ Challenger’
(the ‘ Times,’ 30th April, 1875, and quoted by Bowerbank, P. Z. 8. 1875, p. 506). But see also Mr. T. J. Moore’s
letter to the late Dr. J. EK. Gray on the mode of dragging for the “‘ Regadera ” (Huplectella) with a bamboo and nails
at Talisay, in the island of Cebu, illustrated (Ann. & Mag. Nat. Hist. 1869, vol. i. p. 196). Itagrees substantially with
what Professor Steere relates of how the fisherman obtained his specimen of sponge. In many of the Euplectellas sent to
this country, ragged holes or other slight injuries to the spicular framework indicate where jt has been caught by
the fisherman’s hooks. Since this paper was read, I have myself seen in Edinburgh one of the fishing implements in
question, brought home by the officers of the ‘Challenger’ Mxpedition; and I may state, moreover, that a capital

drawing representing a similar native instrument was exhibited at one of the meetings of the Linnean Society by
Capt. Chimmo, F.L.S.

A NEW GENUS OF THE HEXACTINELLID SPONGID. 22M

brought from the Philippine group are obtained by the natives in plenty on the opposite,
eastern, side of the island of Zebu, but that no sponges are said hitherto to have been
got from the sound or channel above mentioned.

One day, while in the forest, Professor Steere was suddenly apprised of the arrival of
the ‘Challenger’ Expedition, by a note from Professor Wyville Thomson, who had heard
of him and his whereabouts. He hastened on board, passing, as he assures me, a most
agreeable day in the company of our countrymen. He doubts if the English naturalists
obtained any other specimens of this sponge (Dendrospongia), though he told them of his
one, and where and how acquired*.

I may only further add to the history of the specimen that its transport has been an
adventurous one. Its great size and fragile nature prevented packing up in the boxes
with the other objects collected. Safe carriage then depended on its being carried about
by hand. To this end a native grass cord was fastened to the stronger portion of the
root, the more delicate branches were partly wrapped in cotton-wool, and the object
inserted in a paper bag, with a loop of the cord out. Thus slimly protected it afterwards
was borne for hundreds of miles on foot and on horseback; afterwards on shipboard it
was slung to the cabin roof, and there freely dangled during the subsequent long voyage
homewards. The wonder is such a delicate brittle organism escaped injury and smash, or
was preserved tolerably intact through such a journey.

EXTERIOR AND Sections.—This example of Steere’s sponge may well be compared in
outward figure to a shrub, or even with greater propriety, so far as superficial appearance
is concerned, to some of the branching corals. The above remark of course only applies
to the siliceous skeleton, the sarcode and dermal membrane being quite unknown to me.
The lower basal end is laterally expanded and to a certain degree flattish, the thickness
being less by one fourth than the breadth. The inferior left end (see fig. 1, Pl. XXXVI.)
has a depending, slightly curved, broken stump or kind of resemblance to a root, which,
however, may have been part of a horizontal branch; in this case the whole sponge might
have had a greater tilt or inclination to the right than the drawing exhibits. To the
inferior right corner a smaller broken branch projects upwards and slightly outwards.
Above this the basal portion mounts as a gently sinuous stem, giving off three irregular,
short, stoutish main branches about equidistant above each other. The middle one has
a sinistral obliquity, but only a short and considerably worn part of it remains. The
right main branch is given off lowest, and has received rather rough usage and consi-
derable abrasion ; about 2 inches from its commencement a knob only shows evidence
of a branch now absent. As much again higher up is a trifurcation, and each of these
secondary branches bifurcates, terminally producing the long upright but curvilinear
whorled surface branches, one, if not two, of these exhibiting short forked ends. The
left main branch also divides into three secondary branches expanding outwards; each
of these again bifurcating and trifurcating, and some terminally being forked like those
on the right. When in the living state doubtless this sponge has a somewhat more

* Since reading this communication, Sir Wyville Thomson’s return has enabled me to show him the accompanying
Plates ; the species he does not recognize as being among their collections, although it is possible a less perfect spe-
cimen or fragmentary portions may be found among the stores of material as yet imperfectly worked out.

262

222 DR. J. MURIE ON STEERE’S SPONGE,

arborescent outline than that now represented (its dendritic characters suggesting the
generic term), but I have preferred abiding by the photograph rather than adding branches
where fractured portions indicate their former presence.

Dimensions of sponges are so exceedingly variable that this character is only of very
subsidiary importance. Withal in the original description of a type full mastery of detail
is needful, for one never knows on what trivial peculiarities important issues may after-
wards be raised. Admeasurements in the present instance can be but rough approxi-
mations and not mathematical verities. On comparing the great Askonema of the
Portugal coast with its relatively diminutive representative Crateromorpha of the
Philippines, Carter*, in his excellent commentary on the group, says :—“ In short, like
most things in the west, if the Hexactinellidee do not surpass in beauty, they do in size,
for the most part, those of the east.” Steere’s sponge, however, has an area equalling
the large vase-like Askonema, its branched character, though, differing widely.

Its greatest vertical height is 3linches; from tip to tip of the most distant upper
branches is a trifle over 28 inches. Some 8 or 9 inches may be allowed for the length of
the broad lower, trifid stem portion; and the several longer branches vary, three measured
being respectively 16, 20, and 23 inches in their entire length. Taken by compasses, the
breadth quite at the flattish base, or owhat for convenience’ sake may be termed the root,
is fully 54 inches, but the same in the opposite diameter or thickness is barely more than
12 inch. Higher up and just below the broken middle main branch the stem lessens in
breadth to 3 inches across. The girth taken round each of the three so-called short main
branches, where about to divide into their numerous long upper branches, is from 44 to
5 inches. The long whorled branches have a circumference, at what further on will be
explained as the frills, of from 3 to 34 inches, that is, a diameter of about an inch; the
erooved portion intervening between the whorls, however, very little exceeds half that
diameter.

In allusion to the superficial aspect as it looks to the eye and under a hand-lens, the
way in which the specimen has been tossed and carried about should not be forgotten.
Whilst many of the top twigs and branches here and there have wonderfully escaped
injury, and the surface can be made out in detail with tolerable accuracy, other portions
are greatly abraded, and particularly so the stem part. he latter seems to have been
covered by a dermal membrane, now almost entirely rubbed off, at spots only a kind of
fluffy residue proving its former presence. But the abrasion notwithstanding eliminates
other features, especially the marked nature of the pores and oscula as connected with
the internal tubular and canal system. The inferior cracked and fractured pieces also
clearly point out that the said root- and stem-spicules are to a certain extent stronger and
more solid than those higher up.

The colour of the entire siliceous skeleton save lower parts is frosty white, not quite so
lustrous and pearly asis the “glass rope” of Hyalonema, nor so dull as Dactylocalys ; but
yet the marginal spicules are translucent, and in spots, or under certain lights, some parts
are more brilliant than others. The lower stem and so-called root is as a whole to the
eye of a browner tint, its spicular meshwork with a pale yellowish hue. But this coloration

* Ann. & Mag. Nat. Hist. (4) 1878, vol. xii. p. 369.

A NEW GENUS OF THE HEXACTINELLID SPONGID. 223

is evidently due to the presence of a greater amount of dried-up sarcode mingled with
interstitial foreign substances. This darker shade, moreover, brings out in relief specks
and lines of pure solid white, viz. sparsely scattered minute shells of Serpula. These
partially are superficial and partially are deeper seated in the sponge network. In one
hole I observed a small Crustacean, seemingly an Amphipod.

The exposed surface of the stem has been rubbed down moderately even, but never-
theless is far from smooth, being drilled by innumerable holes—in fact resembles shaven
cancellous texture, as is its real nature. The orifices are irregular in size and pattern,
though the most prominent are either circular or ovoid. The larger have a diameter of
nearly 3/5 inch, and are more widely scattered than the greatly more numerous and much
smaller intervening ones. Here and there also the former are so obliquely cut across or
worn as to form short shallow grooves or canaliculi; whereas the finer more angular holes
spring from a delicate lacework, of which more hereafter. 'The said perforations evidently
correspond to the oscula and pores of other sponges. The general surface, moreover,
exhibits several broad but very shallow impressions or concavities which traverse the stem
obliquely. These slightly scooped areas are seemingly traces of the deep hollows so
well marked between the whorls of the upper branches presently to be spoken of.

The distinguishing feature par excellence of the branches is a series of tufts or rosettes,
so continuous and interwoven in the main as to present a whorl running successively
round from base to apex. Hach branch in this particular may be likened to a thick
stranded coil of rope, or, still better, resembles the spiral twist of the horn of the Indian
Antelope (Antilope cervicapra). In some instances where two forks or branchlets arise
close together from a branch, before they tend to divaricate widely apart, their spiral fringes
in passing each other so interlock as to form bridges of union (see fig. 2, Pl. XXXVI.).
To the naked eye, but still more manifest with a pocket-lens, the spiral tufty elevation is
seen to be composed of a bunch of long parallelly placed spicules, which issue from the
axis of the branch at an oblique angle, and slightly spread out at their free ends. The
radiation, however, tends upwards or chiefly towards the direction of the spire; and thus
in each turn the upper border of the fringe or frill gently overarches the intervening
depression, its lower border proportionally less so. The apex of each branch is closed,
ending with a semitwist of the spicular frill. The said spicules are like so many glass
bristles; but their true character is considerably masked by their being bound together
throughout by short transverse and oblique secondary spicula, which produce a network
further to be described under “ Microscopic Structure.’’ What is more plainly visible in
this outward examination is a superficial frosty-like network of fine spicules, these, where
best preserved, causing a woolly appearance; their precise nature will again be discussed.
The spiral hollow between the frill differs from the latter rather in degree than in kind,
the relative absence of long protuberant spicula being its characteristic. The depth
varies, but averages a quarter of an inch. Its bareness brings out two features never-
theless, which are subdued in the frills. One is multitudinous oscula and pores, already.
sufficiently alluded to in the stem; the second, even more deeply interesting, is the presence
of a most lovely, delicate gossamer lacework, composed of minute spicules forming a
rectangular chequer. This is not everywhere intact, but sufficient traces of it exist as to.

224: DR. J. MURIE ON STEERE’S SPONGE,

render it likely that it extended over a great part of the branch, probably excepting the
summits of the whorled frills. Whether it extended over the stem it would be hazardous
affirmatively to assert, though the chances are it may have had a partial covering, basing
judgment upon what we know of other forms possessing a veil. Dr. Millar first drew my
attention to this structural point, and then we thought it only stretched over the large
orifices of the stem, a view we afterwards found erroneous. The significance of this
surface layer, both in a morphological and physiological point of view, I am inclined to
think is of considerable importance, but, as will be seen, it is not confined alone to this
form of hexactinellid sponge.

Having run over the most noteworthy points of the external features of our specimen,
I come to the interior. In studying this, pieces were cut across and in the long direction
of the axis. The transverse sections of both stem and branches disclose not a hollow
cylinder as in Aphrocallistes and Aulodictyon, a funnel-shaped or narrowed fusiform
cloacal cavity as in Rossella and Meyerina, nor yet quite such close areole as in Dactylo-
calyx, but rather one may say a kind of compromise between the last two. While tubular,
these openings are unequal, the larger corresponding to the exterior oscular apertures,
the lesser to the pores aforementioned. The latter, angular and all sorts of shapes, are
bounded by the small cross and tangental spicules; the former, rounder or ovoid, are the
open passages between the larger-sized bundles of vertical and obliquely radiating spicules.
In the centre of the body of the stem the open tubular character is best marked towards
the periphery; radii here are more visible, between which the short secondary spicula
intrude, while quite at the edge the frostwork of dermal sexradiate spicula encircles.

The longitudinal median sections still further explain the preceding, and morever, in |
some respects, agree with Meyerina, though in the general appearances and relative
thickness of the spicula an approach is made possibly nearer to Dactylocalyx. Longi-
tudinal fibrous-looking spicula appear to run up the entire length of the axis, but are in
fact a continuous repetition of stout sexradiate spicules, the cross bars being the’ short
arms. These median vertical pillars, moreover, most obviously branch off curvilinearly
towards the margins; and it is evidently the production of these which gives rise to the
exterior whorls. Thus there is a certain plumular arrangement, the obliquely directed
spicules corresponding to the barbs, the smaller annectent spicules to the barbules, and
the dermal appanage of terminal sexradiates to the so-called extradivisional fluff of owl-
feathering.

. Mrcroscoric Srructure.—Before adverting to this it behoves me to mention the in-
valuable assistance derived from my friends Dr. John Millar and Mr. H. J. Carter. To
the former I am indebted for making my best illustrative microscopical specimens, and
also with liberal hand placing others of different genera at my disposal for comparison,
besides discussing points connected with the interpretation of structure. Mr. Carter’s
labours and knowledge of Sponges, and especially of the group in question, are well
known and justly appreciated. When I add, then, that he has carefully examined portions
of the sponge, compared these with the figures in my Plates, and, moreover, measured the
spicula, drawing out a descriptive analysis of same for my use, it will be admitted I have
been amply helped and stimulated in investigating Professor Steere’s specimen. In my

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 225

own examinations of the structural peculiarities I have used object-lenses of both low
and high defining powers, according as delineations of the coarser parts or of the minute
spicula required.

The different constituents of the skeleton whereof Dendrospongia is built up I enume-
rate as under-mentioned; the measurements of spicules are after Mr. Carter.

1. The Vitreous or Glassy Fibre (figs. 4 to 18 inclusive)—This composes the basal
structure of the sponge, forming its tolerably compact root, stem, and cylindrical branches.
In the living state it supports the sareode, which is charged with other minute spicula,
to be described in the sequel. The glassy fibre is thickest in the central or oldest portions
of the root, stem, and branches, and thinnest towards the circumferential parts of same.
Admeasurements of that of the centre yield a thickness of +4, inch, whilst the smaller
fibre is about z 55 of an inch in diameter. -Hvery intermediate thickness between these
is met with, and, as will be understood, the earliest condition of the growing spicula may
even be less than the lowest dimension above mentioned. As has previously been adver-
ted to in remarks on the sectional naked-eye view, the fibre presents main longitudinal
and innumerable intercrossing branches—that is, extending upwards and outwards round
the axis, leaving room for the canals of the excretory system, which take the same
direction. These complicated anastomosing reticulations, however, are products of hex-
actinellid spicula, the various arms of which may either retain their straight character,
or, bending to a lesser or greater extent (figs. 12 and 22), give rise to sinuosities as the
case may be—their overlapping union and growth, with here and there ultimate coales-
cence of adjoining arms, producing all manner of configuration, and, indeed, the appear-
ance of fibre throughout. The whorls, to some extent already explained, are due to the
extension towards the circumference of the long, pointed, terminal filaments of, so to say,
certain strands of the spicular fibre. Thus the protruding pitchfork character of the
outermost hexactinellid fibres (over which the dermal network, fig. 9, is attached) results
from the central and horizontal arms of the spiculum being elongated and turned
outwards (fig. 12) towards the free surface of the sponge-branch, while the sixth or
inner arm is in continuation with the rest of the spicular glassy fibre internally. The
fibre itself is less or more roughened, that is, covered with short spines, very pronounced
and prominent in the younger spicular fibre.

With regard to these spiny productions on the fibre, Mr. Carter has called my attention
to a point concerning them, the importance of which I
was not then aware of. Under a high magnifyine-
power the said elevations are found not to be simple,
conical, and pointed, but, instead, “‘mucronate,”’ that is,
with the summit divided into two, four, or more expan-
ded points (see woodcut, fig. 1). He further observes
that the only other instance he knows of in which this
form of spine is present is in Parrea densa (‘ Annals,’
1873, vol. xii. p. 463, pl. xvii. figs.5 and 6). The accom- Diagram of spinous processes on the
panying woodcut is a reproduction of his sketch, kindly vitreous fibre of Dendrospongia, on the
forwarded in a note to me.

Fig. 1.

scale of 4, to =, inch. After Carter

226 DR. J. MURIE ON STEERE’S SPONGE,

2. Acerate spined Spicula (figs. 17, 18, and 19)—These are of two sizes, large and
small. Both sorts are long, acerate, fusiform, inequilateral, one half being thicker than
the other. The thicker moiety is rather more abruptly pointed than is the attenuated
one. The larger-sized spicula average yo inch long by gop inch at their thickest part.
The smaller acerate spicula in most respects agree with the preceding, but are not more
than one fourth or one fifth the size of the others. More frequently I have met with
them as represented (fig. 19), in bundles, whereas the larger spicula are usually single and
apart when obtained. The spines covering the large spicula (very manifest in fig. 18)
point away from the thick end, and they all slope in the same direction, closely applied
and almost parallel with the shaft. Hach spine is simple, smooth, and conical, supported
on a projecting basal portion of the shaft. These acerate spined spicula, and particularly
the big kind, may be regarded as subskeletal, being situate throughout the structure of
the sponge, though chiefly towards the surface. They appear to be a very common form
among the Hexactinellid sponges.

According to Mr. Carter, when elongated they apparently pass into the anchoring
spined spicules of the bearded species of the Hexactinellida. Moreover such spicula he
regards as “subskeletal,” because he avers they are often drawn into the vitreous fibre,
with which they become indistinguishably incorporated durimg life; he even holds that
the larger scopuline spicules, presently to be described, are occasionally similarly amal-
gamated.

3. Scopuline Spicula (figs. 13 to 16).—Two forms of these obtain, likewise differing
in dimensions. (a) The larger of the two spicula (fig. 13) consists of a straight shaft and
terminal arms from two to four in number. The shaft is microspined, inflated and pointed
at the free extremity, and the opposite end terminates in a palmate, bifid, or quadrangular
inflation, springing therefrom two to four and occasionally five arms. These latter are
opposite, extended fork-like almost in a line with the shaft itself, and are microspined
and indistinctly capitate, or with convex head. The shaft of these large scopuline spicules
is 2; inch long, and the arms yygq inch long. (4) The smaller scopuline spicula
(fig. 15) have also a straight shaft and from two to four terminal arms. They differ from
(a), besides size, in the arms being opposite and expanded laterally or flower-like. Both
shaft and arms are microspined, the arms very indistinctly capitate. In dimensions these
second kind of scopuline spicules have a shaft z3e inch long, the arms yy'95 inch long.
Besides their terminal arms, both kinds of the spicula under consideration are often more
or less provided with shorter armlets, which spring from the body of the shaft at right
angles, and vary in number as well as in length (figs. 14 & 16). While they are often
hardly traceable in the large scopuline form (a), they on the other hand are occasionally
profuse in the smaller kind (2), numbering in the latter frequently as many as twelve,
which usually are longest towards the middle of the shaft. These different figured sco-
puline spicula pass by almost imperceptible gradations into each other, though in most
slides under the microscope the two sorts are readily distinguished even when lying in
heaps, as is shown in fig. 11.

Mr. Carter remarks that the said scopuline spicula belong to the sarcode inside the
dermal meshwork, where their original arrangement has been destroyed by the contracting

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 227

of this sarcode in drying and subsequent partial destruction of it by mucedinous fungi.
They thus now appear in little detached masses without any definite arrangement,
although, here and there, in the dry sarcode which holds them together, a circular hole
may be seen indicative of their position and office in supporting the internal porous sar-
codic expansions during the lifetime of the sponge.
Indeed the sarcode generally has been destroyed
throughout this sponge in this way, and now only
hangs in small fragments here and there in the
otherwise naked fibre (consult portions figs. 11 &
12). Such, he observes, is the fate of most sponges
under these circumstances, especially if they are
exposed to dampness owing to the salt-water not
having been entirely soaked out of them at the
commencement.

A. Spicular Rosettes (figs. 24.&25).—Flesh-spicula
of the rosette type obtain, and, as in the preceding,
are likewise of two forms, a small and a large kind. Diagrammmatieal representation! Ofehe pros
(a) In this the small rosette or staple form (fig. 24) able mode of arrangement of the scopuline
the figure is relatively globular, and is g{¢ inch in spicula of Steere’s sponge, after a pen-and-ink
diameter. It consists of six straight, smooth arms ‘etch by Mr. Carter. Sarcode membrane,
parting at right angles from the centre, each arm Beret ihy, pore mea ye sede
terminating in a little discoid inflation from which
spring four to eight rays spreading in what Carter
terms a “ flewr-de-lis” (“* Hexactinellidee and Lithistide,” Ann. & Mag. Nat. Hist. 1873,
vol. xii. p. 355; the spinulo-multifurcate hexradiate stellate spiculum of Bowerbank,
Monog. Brit. Spong. vol. i. p. 259, pl. vill. fig. 192). Hach ray terminates in an inflation
which is expanded into a circular convex head bordered by four spines opposite and
recurved. These rosettes (a) are chiefly congregated about the angle of the dermal
meshwork and relatively numerous.

(6) The large rosettes (fig. 25), contrariwise, are very rare, and still more globular in
appearance, whilst they range from ;g to gz inch in diameter. Their centre or, so to
say, nucleus appears to consist of six short, stout arms, each arm terminating in a qua-
drangular inflation, from which radiate several long straight rays. The rays themselves
(very rarely met with bifurcate) end in a quadrangular or more often pentangular cap or
head (fig. 26), with free convex surface, but provided with recurved spines at the angles
of the opposite surface. This kind of rosette resembles the so called “ multiradiate
spicula of the sarcode” of Askonema setubalense, Kent (Monthly Microsc. Journ. 1870,
p- 246, pl. lxiv. fig 9).

5. Sexradiate or Dermal Hexactinellid Spicula (figs. 20 and 23, and in position figs. 3
and 9).—These forming by the coalescence of the adjoining spicula the delicate veil
latticework or rectangular dermal meshwork, have already had passing allusion called
to them when speaking of the exterior characteristics. The six arms part at right angles

from the centre, the horizontal ones being 735 inch long; the vertical are shorter.
. SECOND SERIES.—ZOOLOGY, VOL. I. 24H

tween the spicular bars and quadrangular
spaces, that is, when the sponge was alive.

228 DR. J. MURIE ON STEERE’S SPONGE,

Towards the fixed end (or centre) the arms are thickest, and throughout are minutely
tuberculate or subspined; the arm extremities are smoothish and pointed. It is by the
overlapping of the horizontal arms (fig. 23, a), and this to a considerable extent, that the
squares are produced, whilst the horizontal arms respectively project outwards and inwards.
The outermost tip frequently bears the small rosette (fig. 23, 6) already described.

Besides the foregoing spicula (save the large rosette), all present in abundance, I met
with, by chance, at least in one of the slides of objects examined, another form of minute
but not measured spiculum (fig. 27). Whether this had accidentally got amongst the
débris shaken out of the Dendrospongia, or otherwise forms a constant part of the latter’s
skeleton, I cannot say positively. Possibly this spiculum in question forms but a seg-
ment of a “rosette with short arms and long, straight, pointed rays,” similar to what Carter
figures from Huplectella aspergillum (Ann. & Mag. Nat. Hist. ser. 4, vol. xii, pl. xii. fig. 3);
only one or two examples were observed by me. I have figured it rather to arrest future
attention than to insist on its being an integral part of the sponge under consideration.

REMARKS ON THE FOREGOING.—From what precedes it will readily be admitted’ that
Steere’s sponge is one among the many already singular forms of the Vitreous Sponges
of considerable interest. In size, shape, and structural detail it differs sufficiently from
others of the known Hexactinellida, as I presume, to warrant generic and specific dis-
tinction, while it no less combines certain peculiarities assimilative and explanatory, to
some extent, of moot points in the construction and history of this altogether remarkable
group. Its dendroid or shrub-like branching aspect has suggested the generic appellation
I submit; and the collector’s name, to whom I am so much indebted, I convert into its
specific title. Its habitat but adds another to the many strange and delicate marine
organisms hitherto obtained in the Philippine seas; and doubtless when hereafter studied
in its live conditions some curious traits in its history may be unfolded.

DENDROSPONGIA STEERII, gen. et sp. nov.

Hexactinellid sponge characterized by its dendritic or shrubby contour ; occasionally
attaining a height of 3 feet or possibly more. Branches forking or dichotomous, with
continuous whorled series of spicular tufts from base to apices. Skeleton only known;
basework composed of relatively stoutish glassy fibres of coalesced, sexradiate and spino-
mucronate spicula, disposed in tolerably compact trabecule. Main direction of fibre
longitudinal to axis, in parallel straightish or slightly bent lines where continued into
exterior whorls; intercrossing fibres more irregular, as are the very numerous excretory
canals. Oscula and pores of moderate size distributed all over the free surfaces. Flesh-
spicula abundant, and of scopuline, acerate, and rosette shapes. A dermal veil of
‘slender, interwoven hexactinellid spicula probably clothes the major portion or possibly
the entire sponge.

Hab. Philippines, Channel between Negros and Zebu, from 70 to 100 fathoms ?

In discussing and assigning the affinities of this choice, indeed elegant, sponge, one
naturally first looks to the kindred stock from the Eastern Archipelago. To the Huplec-
tela group, including the genus Habrodictyon (Corbitella and Heterotella, Gray), their

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 229

open tubular unbranched and widely netted structure, besides basal cushion or root of
great long anchoring spicula, sufficiently distinguish and separate them from our specimen
in question, apart from the shape of the various spicula themselves. In Deyerina the
more approximated condition of the glassy fibres composing the stem, the somewhat
spiral and therefore to a certain extent whorled nature .of outer, pronounced elevations
of the same, and marked veil or latticework suggest closer ties with Dendrospongia; but
Meyerina is unbranched, and besides has a birotulate form of flesh-spicule entirely absent
in that compared. But, moreover, the Huplectellide, spite of their looser build of vitreous
fibre, have frills bearing some relation to the said whorls, and an exceedingly delicate
veil, though their true spicular skeleton, as well as that of Meyerina, has not such rough-
ened nor spino-mucronate anastomotic fibre. In Aphrocallistes its branching character
makes a closer approach, so far as mere outward configuration is concerned; but in the
Malaccan as well as Atlantic species the widish tubular branches, relatively thin walls,
absence of exterior whorled elevations, and other differences of framework stamp sepa-
ration from Dendrospongia. The former genus notwithstanding has scopuline and flesh-
‘spicula, and, at least in A. beatriz, such echinate closely reticulate main fibre as to show
relationship with the latter, though not of the closest tie. The massive subglobose
species of Dactylocalyx from Malacca, and even the branching forms of this genus from
other regions, though possessing an anastomotic network of glassy fibre bearing a certain
resemblance to some parts of Dendrospongia, nevertheless in other particulars strikingly
depart from the latter. Such genera as the goblet-shaped Crateromorpha and Labaria,
and the “sculptor’s mallet”’-like Hyalonema, in most aspects show recedence of type
from our Zebu representative.

Again, other genera of the Atlantic and American seas, the purse-figured Holtenia,
Rossella, and nut-like Sympagella, the bird’s-nest Pheronema,and the felt or blanket sponge
Askonema,'to wit, present relationships rather with one another than in any specially
marked degree to that under consideration. To Awlodictyon and Furrea, however, there
undoubtedly seems to be a nearer affinity, if much weight is alone to be placed on the
flesh-spicular identity. Examples of neither of those two genera, so far as I am aware,
have ever been obtained in a perfect and complete form, save the diminutive 7. infundi-
buliformis, Carter ; but still sufficient is known to yield a useful comparison. Both are
‘to some extent diminutive branching Hexactinellid sponges, but even in this respect widely
different from the great, handsome, dendritic Steere’s sponge. They are, besides, hollow

-‘and less or more tubular, with very thin walls and no trace of whorled tufted branches.
In one species of Farrea the closely reticulate vitreous fibre is thickly spined (and mucro-
nate according to Carter); there also obtains scopuline, acerate?, and rosette-shaped
spicules, bearing considerable likeness to those of Dendrospongia.

With all this similarity to species of Farrea in spicular form, considerably departed from
“in other features, it seems to me that Steere’s sponge in reality presents intermediate skeletal
‘characters between Meyerina, Aphrocallistes, and Furrea, while yet so characteristic
‘from all in several respects, that it is hard even to admit positive close affiliation to one or
“other.- I expect the future may reveal some form from the eastern seas that may modify
‘the difficulties of their relationships.

Dig

230 DR. J. MURIE ON STEERE’S SPONGE,

With regard to the homology of the so-called root, body, beard, and dermal spicules of
at least the major section of the sexradiate sponges, I venture to offer a few mere
passing remarks. I apprehend that the presence of a veil or thin dermal layer of lattice-
work is far more constant among the Hexactinellida than is generally suspected, its deli-
cacy in many instances, I fear, causing its destruction in whole or part ere examination
of specimens is made. In Meyerina it is a very beautiful structure, readily apparent to
the eye even at a distance. It exists like a spider’s web on the surface of the Huplectel-
lide. A fine sheet lies upon the Askonema’s felt. Labaria and Hyalonema both possess
such latticework. Holtenia and Rossella characteristically are veiled. Even Dendro-
spongia I am inclined to believe has such a covering spread all over its whorled branches
during life. In other genera the evidence borne by fragments leads to the conclusion of
its being a structural characteristic common to all. It seems to be for the support of the
dermal sarcode in which the pores are situated.

The beard and whiskers, so termed by some authors, of such forms as Pheronema,
Rossella, Holtenia, &c., evidently are the homologues of the spicula, shorter, it is true,
composing the elegant sinuous frills of the Huplectella series. These latter frills, however
much differing in aspect, moreover correspond essentially with the whorled tufts of Den-
drospongia, and with the elevated obliquely ranged spicular vent-ridges in Meyerma. In
these and doubtless other forms they appear to be more or less modified spicules in direct
relation with the true skeletal vitreous fibre, and by or to which the finer dermal lattice-
work is attached.

The long knitting-needle-like bundle of the glass rope in Hyalonema, the anchoring
spicules so well known in Holtenia &c., and the hairy-like bunch of lengthened spicula
at the root of Huplectelia, have all strictly a homological relationship. Indeed nearly all
the Hexactinellida that have been procured in any thing like a perfect condition offer
some remnants, representing spicula specially adapted to retaining a hold of foreign
bodies to which they are rooted, or the so-termed roots are sunk in the sand or mud.

Thus, again, however unlike may he the pattern of the vitreous fibre composing the
body or basis of such forms as Dyliusia and Iphiteon, as contradistinguished from that
of Orateromorpha, Habrodictyon, Euplectella, Dendrospongia, and Hyalonema, whether
the sponge be shallow, cup-shaped, tubular, branched, or more solid, this, so to speak,
basal or body-fibre of simple, fused, or compactly built sexradiate spicula has a common
homological significance.

However much modified, then, I am inclined to think the whole of the Hexactinellid
group, besides their bond of unity in six-armed spicula, are each provided with well-deve-
loped or traces of veil and root, superadded to such glassy fibre as more properly constitutes
their basal or body skeleton.

Postscript.—It was not until some time after I had laid my paper before the Society
that two contributions on the hexactinellid sponges by Dr. William Marshall, of Weimar,
in the Zeitsch. fiir wissensch. Zool.*, came under my notice. The first published of these

* “Untersuchungen iiber Hexactinelliden,” Z. f. w. Z. 11875, vol. xxv. (Supplementheft 2), pp. 142-243, pls.
xi-xvii.; and “Ideen iiber die Verwandtschaftsverhiiltnisse der Hexactinelliden,” Z. f. w. Z. 1876, vol. xxvii. pp.
113-136,

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 231

bore a special interest to me, as containing a photograph of a new genus and species of
sponge—Sclerothamnus Clausii, which I perceived had close affinity with Steere’s sponge.
Their identity at the first glance I certainly did not appreciate, possibly from his evi-
dently being a much smaller imperfect specimen, and also doubtless owing to the blurred
ill-defined photographic representation of it. A further careful study of his descriptions
and close comparison of figures has, however, convinced me that in the said Sclerotham-
nus and Dendrospongia, without a doubt, we have the same generic type, and as regards
specific distinction between them I am really at a loss to point out differences. 4 for-
tiori, I regard them as identical; and therefore Dr. Marshall’s name to this remarkable
form has the claim of priority, and my own will hereafter necessarily be sunk to that of a
synonym. Hence Sclerothamnus Clausii, Marshall,=Dendrospongia Steerii, Murie. The
examples on which Marshall founds his description and diagnosis are two odd pieces
got in an old cabinet of the Leiden Museum, locality and history unknown. My data,
therefore, clears habitat, and in many other respects supplies points wanting in his com-
munication. Dr. Marshal], notwithstanding, has observed and figured the ‘“‘ mucronate”
character of the glassy fibres, on which Mr. Carter has laid some stress, and he moreover
calls attention to certain phenomena connected with the growth and coalescence of the
_ glassy spicular fibre, besides what he regards as monstrous spicula.

In his ideas upon the relations of the Hexactinellida he constitutes a division SYNAv-
Loip#=— Das Lumen der Réhren der verschiedenen Nadeln hangt, wie diese selbst,
continuirlich miteinander zusammen, so dass das ganze Zittergewebe des Schwammes von
einen gleichfalls zusammenhingenden Rohrsystem durchzogen ist ”—in which the only
generic type is Sclerothamnus, whose characters are :—“ Zu den zusammenhaingenden
Zittern gesellen sich als frei bleibende Nadelformen Besengabeln.” ‘'The single species
as yet known, Sel. Clausii, according to him, has the following diagnosis :—“ Polyzoisch,
buschférmig, mit dichotomisch verzweigten, nicht in einer Ebene gelegenen Aesten. Zwei
Formen von Besengabeln. Zitterwork mit Hockerchen besetzt.”

Into his second division AsyNAULOID# and its subdivisions, and with his general con-
clusions and remarks on the relationships &c. of the Hexactinellida, neither space nor
other circumstances permit me herein to enter.

Apprnpix. By H. J. Carrer, F.RB.S.

With Dr. Murie’s description of Dendrospongia Steerii I should hesitate to interfere, even if I were not
well acquainted with the general and microscopic structure of the more durable parts of the sponge itself,
such is the descriptive power and truthfulness of the author. :

Professor Steere’s specimen is certainly the largest and noblest mstance of the Hexactinellida on record,
and from its branched form contributes to show that the Hexactinellida are subject to the same influences
in this respect as those which determine the variation in form and size of all other sponges, which appears,
to a certain extent, to be almost unlimited.

It is not a little remarkable, too, that a similar specimen, only not half so large (16 inches high), should
have been deseribed and figured by Dr. William Marshall, of Wiemar (Zeitschrift f. wiss. Zool. xxv.
Suppl., 20th November, 1875), just two months before Dr. Murie brought Prof. Steere’s specimen to the
notice of the Linnean Society. Still more remarkable is it that just nineteen years before this, and before
the same society (viz. 17th February, 1857), large fragments of this species should have been uncon-

232 DR. J. MURIE ON STEERE’S SPONGE,

sciously brought, and subsequently figured among the root-detritus of Prof. Owen’s Euplectella cucumer,
still where it was then, viz. in the possession of Dr. Arthur Farre (Trans. Linn. Soc. vol. xxu. pl. xxi.
fig. 1), with a microscopic portion, highly magnified im fig. 8, which had been “discovered by Dr. Farre”
himself, and thus had been thought worth figuring independently of the Huplectella—showing, as the
sequel will prove, that the peculiarities of the structure had not passed unnoticed, although the sponge
itself had not been discovered.

Lastly, it is worthy of remark that Dendrospongia appears to exist in the neighbourhood of the Euplec-
tellas both at the Seychelles and among the Philippine Islands.

Besides the fragments of Dendrospongia Dr. Farre also discovered another structure, which Prof, Owen
likened to a “harrow” (l.c. figs. 9,9 a). After this Dr. Bowerbank obtained permission of Dr. Farre to
examine the enéire specimen of Euplectella cucumer, and (among other fragments) brought away with him
specimens respectively of those discovered by Dr. Farre, to which I have above alluded. These Dr.
Bowerbank examined, described, and figured respectively, under the general name of Furrea occa (Proc.
Zool. Soc. 1869, pl. xxiv. figs. 1 and 7), conceiving the “ harrow-like tissue” to have belonged to “the
dermis,” and the other structure to the body of this sponge.

We know, however, now, from enéire specimens of the sponges represented by these two structures,
that they belong to two distinct species, and that the ‘“harrow-like tissue” is the dody-structure of
Farrea occa, and not identical with the dermal layer of the Hexactinellida, probably existing (according
to Dr. J. Millar, who first directed especial attention to it) throughout all the species, although often,
from its delicate structure, more or less rubbed off.

In 1873, just after having, by the kind assistauce of Dr. Farre, examined his justly celebrated specimen
of Huplectella cucumer myself, I also brought away specimens of the structures figured both by Professor
Owen and Dr. Bowerbank (J. c.), and, having examined them microscopically, came to the conclusion that
the “ harrow-like tissue” should remain as Farrea occa, and the other structure be designated Farrea
densa, as belonging undoubtedly to a different sponge (Ann. & Mag. Nat. Hist. 1873, vol. xii. p. 454,
pl. xv. fig. 5). Hence a fragment of the latter with its peculiar spines was figured (J. c. fig. 6), mention-
ing also (p. 463) that among the spicules boiled off from this fibre was a scopuline one like that .of
Aphrocallistes beatriz (which is nearly identical with that of Dendrospongia) “in great abundance.”

As soon, therefore, as Dr. Murie, through Dr. J. Millar, kindly put me in possession of an actualfragment
of Dendrospongia, I saw that the body-fibre was identical with that of Warrea densa, and that the scopulme
spicules were also identical, but I had observed no rosette. However, on turning to a mounted specimen
which had not been boiled in nitric acid, I found not only the rosette but the peculiar head of one of the
scopuline spicules also present with the body-fibre. It may be here remarked, respecting the rosette, that
Dr. Marshall states (Zeitschrift, /. c. p. 185) “ bei Sclerothamnus und Aphroeallistes findet sich keine
analoge Nadelform.” Thus we have traced Dendrospongia back to the Linnean Transactions of 1857.

Whether Farrea densa, Sclerothamnus Clausii, or Dendrospongia Steerii is to take precedence in nomen-
clature I care not. Dr. Murie’s description and illustrations take precedence with me as to accuracy and
fullness of detail. This lasts for ever ; but a name may be disputed to the end of time.

Dendrospongia Steerii evidently belongs to that group of sponges which possesses for one of its pecu-
‘liarities the scopuline spicule, which must not be confounded with the rosette that, so far as my obser-
vation extends, always accompanies it. The scopuline spicule is often taken into the skeleton-fibre, and
therefore may be regarded as a subskeléton spicule ; while the rosette, bemg essentially a flesh-spicule

(that is, confined to the sarcode) is seldom or never included in the vitreous fibre. Hence, if the skeleton-
fibre alone remains, the rosette (as was probably the casein Dr. Marshall’s specrmen) may have fallen
through it on the decomposition of the sareode, since, as I have before stated, when this takes place the
rosette passes out through the skeleton almost as easily as small pebbles through the meshes of a fishing-
net. In such instances, therefore, the absence of the rosette may not indicate that’ there never was any.

The rosette, again, often presents itself under a variety of forms in each species of the Hexactinellida,
as described and figured by Dr. Murie in Dendrospongia Steerit.

Fig. 9.

A NEW GENUS OF THE HEXACTINELLID SPONGIDA. 233

DESCRIPTION OF THE PLATES.
Prata XXXVI.

. Steere’s Sponge (Dendrospongia Steerii). Drawn on stone from a photograph, aided by a piece

of the specimen itself, and reduced ? nat. size. In accordance with the text the lettering
used signifies:—ro, root; s, stem; 7, right, m, middle, and /, left main branches; 0, secondary
branch; f, fork.

. Portion of a terminally forking branch, of the natural dimension. w, whorl or spiral tuft; or,

interlocking branch of same; g, intervening groove or spiral depression between the tufts.

. Asmall segment of a branch of Dendrospongia at one of the spiral grooves. Enlarged about

twice nat. size. This shows the appearance of the veil or delicate latticework of square
meshed spicules, spread over the surface and partially extending to the raised whorls; openings
of the oscula or vents are seen beneath the veil.

. Fragment of the interior of a branch, illustrating mode of skeletal arrangement of the large glassy

fibres. Enlarged about twice nat. size.

PuaTe XXXVII.

. Sketch of the foreshortened summit of a branch, exhibiting termination of spines &c. Of

nat. size.

. A transverse section (somewhat enlarged) of the stem at the base of one of the branches, and where

the ruff or whorled spicula have been considerably rubbed away. The central framework has
many irregularly contoured canals and of different sizes; towards and at the peripheral margin
the glassy fibres appear to radiate, the most marked of which constitute what remains of
the abraded whorls.

. Another cross-section made higher up, and where branching occurs; hence its more compressed

oval form, also in part due to the exterior spicula being worn off. Nat. size.

. A median longitudinal section of part of arubbed branch of D. Steeriz, and where only indications

of the spicular whorls are present. Enlarged about once and a half nat. size. This illustrates
the exact disposition of the internal main body of the glassy fibres or basal structure of the sponge..
The long, large, strong, and vertically directed bundles of spicula incline outwards or with
slightly sinuous curve follow a certain spiral arrangement, the marginal elevations cor-
responding to the whorls and the depressions to the grooves. A network of short cross or
oblique spicula bind together the axial and longitudinally directed stouter sorts. Canals and
oscula permeate freely.

A greatly magnified representation of a portion of the framework or glassy fibre as seen on its
superficial aspect. It exhibits in position the so-called veil-layer or lattice-lke squares of the
dermal hexactinellid spicula (see fig. 28, a, 6), also here and there deep-seated hazy patches of |
the flesh-spicula &c.

Fig. 10. Another portion of the glassy fibre, from the interior, still further enlarged, the dotting showing

its spmy roughening. At a one of the larger rectangular interspaces between the reticulated

vitreous spined fibre is partly shown; 4 is within a similar small circular area; ¢ indicates a

thin spined spicule, so bent and united at tips as to form a complete circle; springing from its

middle is a still more delicate needle-shaped spicule (d), which lies across ¢; below is a mass
“ of the common fibre variously agglutinated.

ON A NEW GENUS OF HEXACTINELLID SPONGIDA.

. Another piece of the vitreous fibre, wherein a, b, two main longitudinal branches, are seen, by

degrees, to approach, and further on, at c, to unite, and ultimately, by coalescence, to con-
stitute but a single thickened fibre. The areas between the glassy fibre are charged with loosely
arranged scopuline and other flesh-spicula. Specimen also very considerably enlarged.

. A couple of what may be compared to sprays of the terminal filaments of the stout glassy fibre,

greatly magnified. These, though compound branched, are characteristically pitchfork in
appearance, the free tips towards the right (corresponding in part to what is shown in fig. 9)
being those that, so to say, protrude through the veil latticework, and hold it in place even in
the dried condition of the skeleton. In some of the terminal single fibres the sexradiate con-
dition is marked, but the axial fibre, nevertheless, ends in a tapering point. Minute dermal
and flesh-spicula abound in this specimen.

. One of the longer scopuline flesh-spicules whose shaft is microspined.
Fig. 14.

Another spicule of the same kind, with secondary or spinal arms, set at right angles to the long
axis of the shaft.

. Scopuline spicule, with microspined shaft and head, bearing expanded arms.

. Variety of the last, with secondary or spinal arms.

. One of the large (subskelatal) acerate spimed spicula.

. Portion of same, still further enlarged, to demonstrate arrangement of the spines on its shaft.
. A group of the smaller-sized acerate flesh-spicula.

. A hexactinellid spiculum, with short axial arms, all the arms being slightly roughened or

microspined.

. Sketch of the pomt of junction of a sexradiate spicule, the four arms of which persist, but only

a rudiment of one of the axial arms 1s apparent in a central knob.

. A couple of the circumferential younger sexvadiate glassy fibre, the arms being unequal in

length, the longer ones tending to curvature.

. A few of the dermal hexactinellid spicula, 7. e. those composing the lattice-lke veil. At a the

mode of union of the squares is displayed, and at 6 a single sexradiate spicule is shown, with
a rosette in position tipping the free extremity of the axial arm.

4. A highly magnified representation in perspective of one of the small rosettes of the fleshy,

dermal, rectangular meshwork. (See fig. 236 for its position with respect to the free axial
arm of the sexradiate spicule.)

. The larger kind of multiradiate rosette, sparsely met with among the débris of the sponge.
. A foreshortened view, or from above, of the cap or head of one of the rays of the rosette, fig. 25.
. A spicule of a minute kind, of which only one or two were observed among the loose substance

shaken off the Dendrospongia.

All the figures of the spicula, from 18 to 27 inclusive, are enormously enlarged ; and the relative propor-
tions as to size of the different sorts have not been strictly adhered to, though in the main approximated.
For their accurate measurements &c. consult description in body of text.

Berjeau & Iterson lith,

D

ENDROSPONGIA STI

T

levine! lit

Trans Linn. Soc:Si

ER.Z.Z00L Vou.l Tas 36.

M& N.Hanhart imp.

Trans.Linn. Soc. Ser.2.Zoou Vou lTas 37

Berjeau lith. Hanhart imp

SKELETAL DETAILS OF DENDROSPONGIA.

Ab

[ 235 J

XII. Notes upon the Oyxstomatous Crustacea. By Epwarp J. Mizrs, 7.L.8., £.Z.8.,
Assistant in the Zoological Department, British Museum.

(Plates XX XVITI.—XL.)
Read June 15th, 1876.
I. Family LEUCOSIID A.
Remarks on the Literature of the Subject.

SINCE 1855, when Mr. Bell published, in the 21st volume of the Society’s ‘ Trans-
actions,’ his excellent monograph of the Leucosiide, several important memoirs have
appeared, adding considerably to our knowledge of the genera and species of this
family, which includes some of the most striking and beautiful forms in the whole range
of the Brachyura. In the ‘ Proceedings of the Academy of Natural Sciences of
Philadelphia’ for 1858 Mr. W. Stimpson published diagnoses of several new genera
and species of Zeucosiide, collected during the United States’ Expedition to the
N. Pacific, chiefly in the Chinese seas. In the ‘ Journal of the Boston Society of
Natural History,’ vi. 1857, the ‘ Annals of the Lyceum of Natural History of New York’
for 1860 and 1871, and in the ‘ Bulletin of the Museum of Comparative Zoology,’ ii.
1870, the above-mentioned author has described a considerable number of new genera
and species from different parts of the North-American coast. M. Alph. Milne-Edwards,
in the ‘ Annales de la Société Entomologique de France,’ v. 1865, the ‘ Nouvelles Archives
du Muséum d’Histoire Naturelle,’ x. 1874, and the ‘ Journal des Museum Godeffroy,’
part iv. 1873, has described and beautifully figured many new and interesting genera
and species from different parts of the world. Species of this family have also been
described or noticed by Gibbes, Herbst, Hess, Kinahan, and others.

In the present essay the species of this family in the collection of the British Museum
which do not appear to have been hitherto recorded, are described; and remarks are
added upon a few other species in the national collection.

Descriptions of New Species of the Family of Leucosiidee.

Leucosta, Fabricius.

The species of this, the typical genus of the family, are remarkable in many instances
for the beauty of the coloration and markings of the polished carapace. ‘They are found
throughout the tropical and warmer temperate zones of the oriental region, from the Red
Sea to the islands of the Pacific.

Twenty species are enumerated by Bell in his monograph of the family. Stimpson, in
1858, added three species from the Chinese seas to the list. A species from the
Australian coast was characterized in 1865 by Hess; M. Alph. Milne-Edwards in 1874
described and figured three species from New Caledonia; and in 1875 I described one

_ SECOND SERIES.—ZOOLOGY, VOL. I. 21

236 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

from the Samoa Islands. There are in the national collection eight species of this
genus, apparently distinct from any that have been hitherto recorded.

1. LevcosIA FUSCOMACULATA, sp, n. (Pl. XX XVIII. figs. 1-3.) Allied to ZL. wrania,
Herbst, but less convex, with a more distinct, marginal beaded line; the carapace
distinctly punctated on the sides; anterior part of the thoracic sinus much
narrower, and its cavity, above the point of insertion of the thoracic anterior legs,
filled by a confluent series of but three or four closely placed, flattened tubercles,
whereas in L. urania the tubercles are at least twice as numerous, and remote from
one another. Abdomen of male smooth, 4-jointed. Colour light, yellowish brown,
with very indistinct spots and patches of dusky grey on the carapace. Length % in.;
breadth 2 in.

Hab. Wed Sea, Gulf of Suez.

2. LevucosIA PULCHERRIMA, sp. n. (Pl. XXXVIII. figs. 4-6.) Carapace smooth,
convex, sparsely punctate towards the front and anterolateral margins, which are
defined by a beaded line which terminates above the point of insertion of the first
pair of ambulatory legs. Hepatic regions concave in their front portion, and
slightly convex behind. Arm with a line of large tubercles on its anterior and
posterior margins, upper surface with a group of small tubercles at base, followed
by four larger; lower surface with many closely placed tubercles at base; wrist .
and hand smooth, fingers not quite meeting at base when closed. Thoracic sinus
deep, narrow, and well-defined in front, with a beaded margin; its cavity, above
the base of the anterior legs, filled with a series of about half-a-dozen small
tubercles. Colour stone-grey, inclining to brown on the back and white on the
ambulatory legs, a circle of rose-pink upon the back of each branchial region, a
trilobate pattern formed by three confluent semicircles of the same colour upon each
side of the anterior half of the carapace (these are confluent in the middle line, and
together somewhat resemble in outline a butterfly with the wings extended); there
are three white spots in the centre of each trilobate pattern; the basal half of the
fingers is orange; and there are rings of the same colour surrounding the joints of the
ambulatory legs and tubercles of the arms, also small spots of orange near the
lateral margins and upon the inferior surface of the carapace. Length 1), in.;
breadth +4 in.

Hab. Lizard Island. —

In all structural characteristics this species is nearly allied to ZL. neocaledonica,
A. M.-Edw.; but it is preeminently distinguished from this and all other species of the
genus by the strikingly beautiful coloration and markings of the carapace. The thoracic
sinus is also deeper, narrower, and better-defined in front, with a beaded margin. Only
a single specimen (2) in the collection.

3. LEUCOSIA ORNATA, sp. n. (Plate XXXVIII. figs. 7-9.) This species, in all struc-
tural characteristics, very closely resembles L. pulcherrima; but it is more strongly

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 237

punctated, the concavity and convexity of the hepatic region is more decided, and
there are but four clearly distinct tubercles in the cavity of the thoracic sinus. The
colour is dark brown; there are three white spots upon each side of the gastric
region, and a small red circle on the back of each branchial region, The bases
of the fingers, tubercles of the arms, and joints of the ambulatory legs are banded
with orange, as in many species of the genus. Length 1 in.; breadth % in.
Hab. Ceylon,
This species was presented to the Trustees of the British Museum by H. W. H. Holds-
worth, Esq., with a valuable series of Crustacea from Ceylon. The specimen is a female.

4, LEUCOSIA BRUNNEA, sp.n. (Plate XX XVIII. figs.10-12.) Carapace convex, smooth
and polished, but very slightly punctated towards the anterolateral margins, and
with a finely beaded line continued to the middle of the posterolateral margins.
Front very slightly projecting beyond the eyes; behind it there is a short longi-
tudinal median ridge between the concave hepatic regions, Thoracic sinus wide, its
anterior margin straight, not beaded, and forming nearly a right angle with the
anterolateral margin of the carapace. Arm with numerous, small, crowded tubercles
upon its infero-anterior surface, upper surface with a group of very small tubercles at
base, beyond them four tubercles placed in a quadrangle; anterior and posterior
margins with a line of tubercles; wrist smooth; inner margin of hand with
a granulated line; fingers straight. Colour brown, inclining to red on the back of
the carapace. Length 13 in.; breadth 1 in.

Hab. Singapore.
This species is distinguished from its congeners by the form of the thoracic sinus &c.

5. LeUCOsIA RETICULATA, sp. n. (Pl. XX XVIII. figs. 18-15.) Carapace subglobose,
smooth ; sides with a beaded line extending from the front of the hepatic regions
along the anterolateral and about halfway along the posterolateral margins. Front
flat above, and not produced beyond the eyes; anterior margin straight. Thoracic
sinus short; anterior margin smooth, not beaded, and not defined at either ex-
tremity. The cavity of the thoracic sinus, above the insertion of the anterior legs,
is filled by a series of about half-a-dozen flattened tubercles. Anterior legs strong;
arm tubercular on the proximal half of its upper surface, the distal half smooth and
polished; on the anterior and posterior margins the tubercles extend nearly to the
extremity of the arm; on the anterior surface, and below at base, the tubercles are
numerous and crowded; wrist smooth, hand compressed, outer margin smooth,
inner with an obscurely beaded line. Abdomen of male smooth, without any
tubercle on the penultimate joint; terminal joint very small, the others of about
equal width throughout. Colour greyish green, yellowish on the sides and back of
the carapace, which is marked with reticulating lines ofa darker green. Ambulatory
legs yellowish brown. On each side of the carapace, above the point of insertion of
the anterior legs, there isa group of from six to a dozen small red granules. Length
11 in.; breadth 1 in. i

212

238 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

Hab. Western Australia, Shark’s Bay.
This species is at once distinguished by its colour and the group of red granules on
each side of the carapace. Specimens of both sexes are in the collection.

6. Lrvcosta WHITMEE! (Plate XX XVIII. figs. 16-18.)
Leucosia whitmeei, Miers, Ann. & Mag. Nat. Hist. xvi. p. 342 (1875).

This species was described by me in 1875 from a specimen from Samoa, presented by
the Rey. 8. J. Whitmee to the Trustees of the British Museum. Specimens are in the
national collection, from Ovalau in the Fiji Islands, which probably belong to this
species, with which they agree in structural characteristics ; but all traces of the original
coloration and markings of the carapace have disappeared.

7. Leucosia PERRY, sp.n. (Plate XX XVIII. figs. 19-21.) Carapace rather depressed,
with a continuous beaded line defining the lateral and posterior margins, and with
a slight longitudinal median ridge between the hepatic regions. Front tridentate,
not concave above. Thoracic sinus entirely absent. Anterior legs rather slender ;
arm closely tuberculated above and below at and towards the base; wrist and hand
smooth, hand rather compressed, but not keeled or beaded; fingers, when closed,
meeting only at tips. Abdomen of male narrow, triangular, with the sides straight,
and witha slight tubercular prominence on the penultimate segment. Colour of
carapace deep brown, with two or three small, irregular, paler spots in front of the
branchial regions ; arms greyish; hands and ambulatory legs paler brown. Length
3 in.; breadth about 4 in.

Hab. Western Australia, Shark’s Bay.

This species is distinguished from all others of the genus having a tridentate front by
the entire absence of the thoracic sinus. In this respect it resembles L. orbicularis, Bell ;
but this species has the front much less produced forward, with the anterior margin
concave.. I have named it in honour of its discoverer, W. Wykeham Perry, Esq., by
whom it was presented to the Trustees of the British Museum, with three others of the
new species described in this paper *.

8. LEUCOSIA PUBESCENS, sp. n. (Plate XXXVIII. figs. 22-24.) Carapace convex,
smooth and polished; anterolateral margins with a finely beaded line, which does
not extend beyond the commencement of the posterolateral margin. Hepatic
regions concave. Front tridentate, flat above. Thoracic sinus with its anterior

* The following is a list of the species of Crustacea (besides several interesting specimens too young for deter-
mination) that have been presented by Mr. Perry to the collection of the British Museum :—

Lambrus nodosus. Geograpsus crunipes. Matuta obtusifrons, sp. 0.
Neptunus sanguinolentus. Sesarma obesum. Calappa hepatiea.
Caphyra rotundifrons (young). S. quadratum. Renuiipes pacificus,
Gelasimus annulrpes, var. Leucosia reticulata, sp. D. Squilla armata,

G, gaimardi. L, perryi, sp. .

Grapsus pictus. Matuta crebrepunctata, sp. D.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 239

margin smooth, not beaded, and not defined at either extremity, its cavity above
the base of the anterior legs filled with a short close pubescence. Upper surface of
arm with a group of small tubercles at base, followed by two larger, and with a
patch of dense short pubescence. Anterior and posterior margins with a line of
tubercles; lower surface densely tuberculated; wrist and hand smooth, fingers
straight. Colour greenish grey, with a series of eight marginal spots of darker
green, two spots of the same colour on the gastric region, and four on the cardiac
region. There are also two white spots on each side of the gastric region. Ab-
domen of male smooth, with all the joints except the last coalescent. Length nearly
1 in.; breadth 3 in.

Hab. Western Australia, Shark’s Bay.

This species is most nearly allied to Z. pallida, Bell, but differs from it in the form of
the thoracic sinus, which is wider in front; the arm of L. pallida has not the patch of
pubescence on its upper surface at base that always exists in specimens of L. pubescens ;
the hand is slenderer, with a granulated line on its inner margin.

Myra, Leach.

Myra mMAmMitiARts, Bell. (Plate XXXVIII. figs. 25-27.) Some specimens from
Australia, in the British-Museum collection, differ in several particulars from the
adult specimens of IZ. mammillaris, but are, I think, the young of that species. In
these examples the carapace is scarcely longer than broad, and there is a faintly
marked longitudinal median carina. The hepatic regions are more strongly convex,
the pterygostomian regions more decidedly angulated ; the tubercles on the posterior
margin of the carapace are more conical (though not acute), the anterior legs are
much shorter, and the tubercle on the penultimate segment of the abdomen of the
male much more prominent than in adult specimens of WM. mammillaris. Length -
inch; breadth 2 inch.

It is probable, if this determination be correct, that both If. carinata and W. elegans,
Bell, are founded on specimens that have not attained quite their full size, and that the
median carina, characteristic of these species, will be found to disappear entirely in fully
matured animals. Should, however, the examination of a larger series of specimens
prove the examples above described to be specifically distinct from IL mammillaris, they
may be designated by the name of WZ. angulata.

Noursta, Leach.

1. NuRsIA stNuATA, sp.n. Carapace moderately convex, broader than long, with three
tubercles placed on the cardiac region in the form of a triangle, a longitudinal
median ridge extending from the front to the posterior margin, and a transverse
curved ridge that extends from the cardiac region to the second tooth of the pos-
terolateral margin. There is also a slight elevation in front of each anterior tubercle
of the cardiac region. There are well-marked depressions between the ridges
towards the margins. Posterolateral and posterior margins deeply sinuated, with

240 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

seven broad, triangular, nearly equal teeth. Anterior legs slender; arm granu-
lous and rounded on its anterior and inferior and the proximal half of its posterior
margins ; wrist and hand smooth; hand slightly compressed, but scarcely carinated.
Hab. Australia, Moreton Bay.
In both NV. plicata and N. abbreviata the anterior legs are smooth, the arm trigonous,
and the hand angulated and carinated. In N. plicata the teeth of the posterolateral
margins are unequal; in IV. abbreviata they are nearly ol)solete.

2. Nursta pLicaTa? (Plate XXXVIII. fig. 28.)

? Cancer plicatus, Herbst, Naturg. Krabben u. Krebse, iii. pl. lix. fig. 2 (1803).

Nursia hardwickii, Leach, Zool. Miscell. i. p. 20 (1817); M.-Edw. Hist. Nat. Crust. ii. p. 137 (1887).
Nursia plicata, Bell, Trans. Linn. Soc. xxi. p. 307 (1855), no figure,

Hab. Indian Ocean.

Mr. Bell observes of this species, that the only specimens with which he was acquainted
(those in the British-Museum collection) were brought from India by General Hard-
wicke. He evidently was unaware of the true habitat of the specimens of NV. plicata,
and confounded them with NV. hardwickii. These specimens were in the collection
in 1852, three years prior to the date of the publication of Mr. Bell’s essay; and one of
them is undoubtedly figured by him as W. plicatus (pl. xxxiv. fig. 4). In the British-
Museum copy of Herbst’s work the description and plate containing the figure of his
Cancer plicatus are unfortunately wanting; and therefore I cannot tell to which of the
undoubtedly distinct species, NV. sinuata and N. hardwickii, it is to be referred. JV.
hardwickii of all ages is distinguished from the WN. sinwata by the anterior legs, which
are flat, smooth on the upper surface, with the anterior and posterior margins of the
arms thin and sharp-edged, not rounded, as in NV. sinuata,

The specimen I have figured is the typical example of NV. hardwickii.

ARcANTA, Leach.

ARCANIA GRANULOSA, sp.n. (Plate XX XVIII. fig. 29.) Carapace shining, subglobose,
uniformly and distinctly granulated, with ten short, acute, equidistant, smooth
marginal spines—and one on the intestinal region, above the two posterior marginal
spines. The frontal spines short, projecting but slightly over the eyes. Anterior
legs slender, about twice as long as the carapace; arm distinctly, wrist and hand
very finely granulated. Length nearly 4 in.; breadth + inch.

Hab. Australia, Moreton Bay.
This species is distinguished by the granulated carapace (which is not at all tubercu-
late or spinulose on its upper surface) from the other species of the genus.

CRYPTOCNEMUS, Stimpson.

This genus was established by Stimpson in 1858 for a Crustacean discovered in Japan,
and remarkable for the extension of the carapace laterally over the ambulatory legs, to
which he applied the specific name of C. pentagonus. A second species, C. grandidierii,
was described by M. A. Milne-Edwards in 1865, from Zanzibar.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 241

CRYPTOCNEMUS HOLDSWORTHI, sp.n. (Plate XX XVIII. figs. 30-32.) Carapace nearly
transversely oblong, with the frontal region projecting; the lateral margins nearly
as long as the anterior margins, and forming nearly a right angle with the straight,
entire, posterior margin; smooth, without granules or tubercles, convex in the
centre, and slightly concave towards the margins. There is a longitudinal median
ridge between the front and cardiac region, and a less distinct, oblique ridge between
the cardiac region and posterolateral angles of the carapace. Anterior legs (in
female) smooth, with the arm trigonous, hand slightly compressed and cristate
above, the margins straight, not sinuated; fingers nearly as long as palm, meeting
throughout when closed. Ambulatory legs small, and entirely concealed beneath
the carapace. Abdomen of female broad, punctulated, 4-jomted; the terminal
joints coalescent. Length } in.; breadth 3 in.

Hab. Ceylon.

This species is named after its discoverer, H. W. H. Holdsworth, Esq. In C. gran-
didieri, A. M.-Edw., from Zanzibar, the front is broad and truncate, the elevated regions
of the carapace marked with granulated lines, and its posterior border emarginate. In
C. pentagonus, Stim., from Japan, the carapace is differently proportioned, the entire
lower surface of the body smooth, and the fingers only one third the length of the hand,
which has greatly expanded horizontal crests and deeply sinuated margins.

If. Family MATUTID A.

Historical Summary.—Although the remarkable crustaceans of the Oxystomatous
genus Matuta are among the commonest of the species of Brachyura inhabiting the
Oriental region, the differences in the armature and coloration of the carapace and anterior
legs are so slight and so numerous that their determination is a matter of great difficulty ;
and most of the later naturalists have been content to admit one, or at most but two
species, distinguished mainly by the coloration of the carapace. I have, after careful
examination of the large series of specimens in the British Museum, arrived at the con-
clusion that there are at least nine well-marked species in the national collection.

In the following condensed summary of the history of the genus, the principal authors
are mentioned who have contributed to our knowledge of the classification of the species.

Fabricius, in the Supplement to his ‘ Entomologia Systematica’ (1798), established the
genus Matuta for his previously described Cancer victor, and added another species,
M. planipes, which he characterizes simply as “thorace postice striato.’ It is impos-
sible, from this brief diagnosis, to identify the species with any of those described below
with any degree of certainty.

In 1817, Dr. Leach, in the third volume of his ‘ Zoological Miscellany,’ p. 12, pub-
lished an article on the ‘‘ Characters of the Genus Matuta, with Descriptions of the
Species.” In this essay four species are described, and two figured ; the Matuta vicior
of Fabricius is referred to, but not described; no attempt is made by Dr. Leach to
identify with his own the species of Herbst, Desmarest, Latreille, and earlier authors.
The characters employed in the definition of the species are unfortunately such as vary

249 MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

with the age and sex of the specimen ; and (in the case of I. peronii and WZ. lunaris)
examples of widely separated species are brought together, and exist in the nationa
collection under the same specific name. In the case of these species I have therefore
considered the specimen figured by Dr, Leach the type of the species. Of IZ. banksia
no labelled specimens are now in the collection; and I have been guided by the descrip-
tion alone.

In 1825, M. Latreille, in the tenth volume of the ‘Encyclopédie Méthodique d’ His-
toire Naturelle,’ published an excellent article upon the genus Matuta, in which he indi-
cates, for the first time, what I believe to be the natural subdivisions of the genus, based
upon the sculpture of the external surface of the hand. He describes a new species,
M. doryphora, and in a footnote points out distinctive sexual characters. His essay is
not noticed by M. Milne-Edwards in ‘ L’Histoire naturelle des Crustacés.’ This natu-
ralist, in his second volume of the work in question (1837), admits but two species of
this genus, WZ. victor and WV. lunaris, Herbst, characterized respectively by the carapace
being marked with numerous scattered dots or reticulating lines.

De Haan, in the fifth fasciculus of his elaborate work on the Crustacea of the ‘ Fauna
Japonica’ of Von Siebold (1841), reunited under one name, Matuta victor, all the species
of the genus, but distinguished six varieties, to which no original names are ‘applied —
although he quotes as synonyma the names of several of the species of the earlier
authors, as referring to them, whether rightly I am not in every case able to determine.
Dana, in 1852, in the first part of the ‘Crustacea of the United-States Exploring Expe-
dition,’ xiii. p. 395-6, notes the sexual differences in the striation of the ridge on the
mobile finger in the specimens collected by the Expedition, but adds nothing to our
knowledge of the species.

Hess (1865), in the ‘Archiv f. Naturgeschichte,’ xxxi. p. 158, pl. vi. f. 18, described
and figured a new species, MZ. picta, from Sydney. I. Hilgendorf, in the ‘ Reisen in
Ost-Afrika’ of Baron van der Decken (1869), figures details of a species he refers, I
believe, correctly to JZ. victor (op. cit. Crustaceen, p. 93, pl. iii. fig. 2), and adds remarks
upon the striated areas upon the inner surface of the hand (which, he thinks, serve the
purpose of producing sound), and upon the specific distinctness of I. victor and I.
lunaris.

Lastly, in 1874, M. Alphonse Milne-Edwards, in the ‘Nouvelles Archives du Mu-
séum,’ x. p. 54, reunites, under the name of I. victor, all the species of earlier authors,
remarking that it is impossible to admit the numerous species proposed by different
authors, every intermediate degree being found between forms which at first might
appear distinct.

I have endeavoured to retain the names of the early authors for the species hereafter
described, wherever possible; but their identification is often a matter of considerable
difficulty, owing to the insufficiency of the descriptions and the inaccuracy of the figures.
The species of the genus Watuta are distributed throughout the whole Oriental region ;
but I am not aware that any have been recorded from any other of the great geogra-
phical areas.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 243

A Revision of the Species of the Genus Matuta.

Marotta, Fabricius.

In the species of this genus the carapace is usually slightly broader than long, broadest
and with a strong, conical, acute, lateral spine at the junction of the anterolateral and
posterolateral margins. The front is about equal to the orbit in width, slightly con-
cave above, with a projecting median lobe, which in some species is notched, in others
entire. The anterolateral margins are rather convex, with about six small tubercles
behind the external orbital angle, and following them usually three granulated, larger
tubercles. The posterolateral margin is straight, with a tubercle, that is sometimes
obsolete, placed at some distance behind the lateral line. In the central region of the -
carapace are six tubercles, of which some, or all, may be obsolete; three of these are
placed in a median transverse series, one anterior on each side of the gastric, and one
posterior in the centre of the cardiac region. The anterior legs are robust, and may be
closely applied to the body; the arm is smooth, the wrist slightly tuberculated upon its
external surface; the upper margin of the hand has three prominent, acute lobes or
spines; on the upper part of the external surface are three transverse ridges, or two
parallel series of three tubercles; below these, on the outer surface, a ridge, or series of
tubercles and spines, the direction and form of which varies in the different species of
the genus; on the inner surface two small striated areas. The external surface of the
mobile finger in the male is usually transversely striated or beaded. The four following.
pairs of legs have the two terminal joints more or less expanded and compressed; the
expansion of the terminal joint is greatest in the first and fourth, and least in the third
pair of legs; and this joint is acute at the extremity in the first three, and rounded in
the last pair of legs.

(A) Hand of male with an interrupted ridge or series of spines and tubercles crossing its outer surface

obliquely, and continued along the outer surface of the lower (immobile) finger. Upper (mobile) finger
with a strongly beaded ridge extending throughout its whole length.

(a) Carapace covered with numerous dark-red spots.

1. Marura vicrrix. (Plate XX XIX. figs. 1-3.) site a Berk]

Cancer victor, Fabr. Spec. Ins. 1. Append. p. 502 (1781) ; Ent. Syst. ii. p. 449 (1798)?
Matuta victor, Faby. Ent. Syst. Suppl. p. 369 (1798) ; ? M.-Edw. Hist. Nat. Crust. ii. p. 115, pl. xx.
figs. 3-6 (1837) ; Régne Animal de Cuvier, Atlas, Crustacés, pl. vii. fig. 1.
Matuta peronii, Leach, Zool. Miscell. m1. p. 18, pl. 127. figs. 1, 2 (1817) @.
Matuta lesueurii, Leach, Zool. Miscell. mi. p. 14 (1817) 3. he
Carapace finely but distinctly granulated; two anterior tubercles of the carapace obso-
lete, the remainder, as well as that upon each posterolateral margin, faintly marked ;
lateral marginal spine long, and directed slightly forward. Front slightly emar-
ginate, with the lobes rounded. Hand not granulous upon its external surface, tie
‘tubercles on the upper half sharply defined; a small tubercle, followed by a very
strong, conical spine at the base of the oblique ridge, which is interrupted at base,
SECOND SERIES.—ZOOLOGY, VOL. I. 2K

24,4) MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

and obscurely defined. Ridge upon the external surface of the mobile finger strongly
beaded. Spots upon the carapace numerous, minute, simple, scattered, and without
any tendency to form reticulating lines. Length 1} in.; breadth nearly 2 in.

Hab. Red Sea; Zanzibar; Indian Ocean; Borneo; Australia.

This is the commonest and most widely distributed species of the genus. In the
females the ridge upon the external surface of the palm is replaced by a series of two
acute spines of unequal length, alternating with three small tubercles. The ridge upon
the external surface of the mobile finger is very obscure or obsolete.

I adopt the name of IZ victrix for this species, as it is certainly figured by M. Milne-
Edwards as Matuta victor in the Atlas of the ‘ Histoire Naturelle des Crustacés;’ and
it is also the commonest species of the genus. Perhaps the Matuta doryphora of La-
treille is also to be referred to this species. The description hardly suffices for its deter-
mination with certainty.

1a. MATUTA VICTRIX, var. CREBREPUNCTATA. (Plate XX XIX. fig. 4.) __
Cancer lunaris, Herbst, Naturg. der Krabben und Krebse, i. p. 140, pl. vi. fig. 44 (1790) ?

This differs from the preceding in having the carapace more coarsely granulated, the
spots very numerous, crowded, and forming small compound spots, each having the form
(under a lens) of a small curved line or circle with paler centre. In the females the
form of the hand is similar to that of IZ. victria.

Hab. Japan; Fiji Islands; New Hebrides.

This variety, on account of its crowded, dark-red spots, presents a very different ex-
ternal appearance from the preceding; and I am in some doubt whether it ought not to

be separated as a distinct species.

(b) Carapace with numerous looped reticulated or anastomosing lines.

2. MATUTA RUBRO-LINEATA, sp.n. (Plate XX XIX. figs.5 & 6.) Carapace finely but

eu distinctly granulated. Two anterior tubercles of the carapace obsolete, the others,

Ne as well as the tubercle upon each posterolateral margin, faintly marked. Lateral

| marginal spine long and acute. Front distinctly emarginate, the lobes more acute
than in MW. victriv. Hand of male nearly as in that species, but with the spine at
base of the oblique ridge shorter. The lines upon the carapace light red, often
interrupted and more crowded anteriorly, rarely forming complete loops and circles.
Length 14 in.; breadth 14%, in.

Hab. Indian and Pacific Oceans; Cheefoo.

A specimen presented by the Smithsonian Institution is named Jf. lunaris.

Tn the females the oblique ridge upon the palm is replaced by a series of five promi-
nences, the second of which is an acute spine, the others small and tuberculiform. The
ridge upon the mobile finger is obsolete. In coloration this species resembles Maiwia
lunaris, but differs in the oblique ridge upon the external surface of the hand, and the
beaded ridge upon the external surface of the mobile finger.

MR. E. J. MIERS ON THE OXYSTOMATQUS CRUSTACEA. 245

3. MATUTA LINEIFERA, sp.n. (Plate XX XIX. fig. 7.) Nearly allied to the preceding,
but with the carapace more finely granulated, the tubercles more strongly marked,
the two anterior distinct, as are also those of the posterolateral margins. Lines
upon the carapace dark red, very fine, thread-like, usually uninterrupted, and
forming complete but irregular loops and circles. Length 13 in.; breadth 14 in.

Hab. N.W. Australia, Nicol Bay.
Only a single male adult specimen is in the collection. A larger series of specimens
might show that it is but a variety of the foregoing. |

(B) Hand of male with an interrupted ridge, or series of tubercles and spines, crossing its external surface
parallel to the lower margin, and terminating at the base of the upper margin of the lower (immobile)
jinger. The ridge upon the outer surface of the mobile finger obsolete, or smooth, or very finely striated at
base, and beaded only towards the distal extremity.

(a2) Carapace marked with spots or blotches.

A, MATUTA GRANULOSA*, sp.n. (Plate XX XIX. figs. 8&9.) Carapace rather coarsely
granulated, especially towards the lateral marginal spines. ‘Two anterior dorsal
tubercles obsolete, the others and those on the posterolateral margins very indistinct.
Lateral marginal spines short, conical, and rather obtuse; front slightly but
distinctly emarginate, with the lobes rounded. Hand of male with the upper and
lower part granulous between the tubercles; the ridge upon the outer surface in
the male divided into five acute tubercles, of which the second is the largest. Ridge
upon the external surface of the mobile finger thin, smooth, not beaded. Spots
upon the carapace brownish red, of the same shape as in J. victrix, var. crebre-
punctata, but less numerous. A brownish-red patch upon the basal half of each
lateral spine. Length and breadth nearly 13 in.

Hab. Hastern seas.

In the female of this species the tubercles of the ridge upon the external surface of
the palm are disposed as in the males. The ridge upon the external surface of the
mobile finger is obsolete.

5. Matura BANKsII. (Plate XL. figs. 1 & 2.)
Matuta banks, Leach, Zool. Miscell. ii. p. 14 (1817) ?

Carapace very coarsely granulated, especially in the centre and towards the lateral spines ;
the tubercles, as well as those of the posterolateral margins, very strong and distinct.
Lateral marginal spines of moderate length, acute; front obtuse and rounded, or

* Since the description of this species was drawn up (from three specimens, of which the exact locality is unknown,
in the collection of the British Museum) I have had the opportunity of examining a series of seventeen specimens of
both sexes and different ages in a collection made by Mr. Mac Farlane on the islands of the Torres Straits. These
examples show that M. granulosa is never to be confounded with its nearest allies, M. banksi and M. maculata,
from which it is always distinguishable by its coloration, the absence of the anterior pair of dorsal tubercles, dc.
The characters generally are subject to little variation: the only points I have observed are, that in some specimens
the tubercles are rather more clearly defined, and the ridge on the outer surface of the mobile finger faintly striated
at its distal extremity, as in other species of this section of the genus.

2K 2

IAG MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

very obscurely emarginate. Hand slightly granulous in its lower half, the tubercles
_very strongly marked, ridge in the male divided into three acute tubercles alter-
nating with two spines. Ridge upon the outer surface of the mobile finger very
finely striated in the greater part of its length, and obscurely beaded towards
its distal extremity. Spots upon the carapace pink, more numerous and crowded
upon the anterior portion, but not disposed in reticulated lines. Length and
breadth about 1¢ in.
Hab. Indian Ocean; China Seas; Philippine Islands.
A specimen from the Philippines has the spots more scattered, and a large pink blotch
upon the penultimate and last joints of the second and fifth pair of legs.
In the female of this species there are two or three conical, acute tubercles and spines
upon the external surface of the hand. Ridge of mobile finger obsolete.

6. MATUTA MACULATA, sp.n. (Plate XL. figs. 3 & 4.) Carapace distinctly granulated,
the granules coarser upon the raised parts of the carapace, but the tubercles, both
of carapace and posterolateral margin, nearly obsolete. Lateral marginal spine
very long and acute; front obtuse, or but very faintly emarginate. Hand granulous
upon its inferior surface, the tubercles upon its upper part distinct, the ridge in the
male with five prominences, of which the first, third, and fifth are indistinct tuber-
cles, the second and fourth larger, acute. Ridge upon the mobile finger as in the
preceding species. Spots upon the carapace small and crowded anteriorly, pos-
teriorly larger, scattered, and each in the form of small curved lines and circles.
Length and breadth about 1,4); in.

Hab. Kastern and Chinese Seas; Panagatan Shoal.
In the female the tubercles of the ridge upon the hand are nearly asinthe male. The
ridge of the mobile finger is obsolete.

7. Marura prcta. (Plate XL. figs. 5—7.)

Cancer lunaris, Leach, Zool. Miscell. iii. p. 18, pl. 127. figs. 8-5 (1817), nec Herbst.

Matuta planipes, Desm. Consid. Crust. p. 102 (1825), nec Fabricius.

M. picta, Hess, Archiv f. Naturg. xxxi. p. 158, pl. vi. fig. 13 (1865).

Carapace distinctly granulated. Tubercles of the carapace and posterolateral margins
stronglymarked. Lateral marginal spine of moderate length, directed slightly forward..
Front obtuse and rounded or slightly emarginate. Hand not granulous upon its
external surface; tubercles upon the upper part rounded and obtuse, ridge on
external surface divided into four or five lobes, of which the second is an acute
spine, the remainder small, obtuse, or truncate. Ridge upon the mobile finger
distinctly striated in the greater part of its length, and beaded at its distal extremity.
Spots upon the carapace very numerous and minute, often disposed in reticulated
lines. Length 13 in.; breadth 144 in.

Hab. Ceylon; Indian Ocean; Zanzibar; Mauritius; Seychelles ; Australian Seas.
In the single adult female of this species in the collection of the British Museum, the
tubercles upon the lower part of the hand are stronger and interspersed with granules.

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA. 24,7

The second and fourth of the series of prominences on the external surface of the hand
are acute, spiniform. The ridge upon the outer surface of the mobile finger is obsolete.
The example figured by Hess under the name of Jf. picta is almost certainly the young
of this species ; and I have therefore adopted his specific name. It is easily distinguished
by the tendency of the spots upon the carapace towards a disposition in reticulating
lines.

The Matuta distinguenda, Hoffmann, Recherches sur la Faune de Madagascar, &c.,
Crust. p. 27, pl. vi. figs. 49-52, vii. 538-57 (1874), may, I think, be this species, although
in the figure, which is evidently much exaggerated, the front is represented as entire,
the tubercles much more prominent than in any species of the genus known to me,
and the immobile finger of the hand of the female much abbreviated.

(6) Carapace marked with looped, reticulating, or anastomosing lines.

8. MATUTA OBTUSIFRONS, sp. n. (Plate XL. figs.8 & 9.) Carapace rather coarsely
granulated, especially in the centre and towards the lateral spines; tubercles all
strongly marked and distinct. Lateral marginal spines of moderate length, acute.
Front small, obtuse, rounded, and entire. Hand very faintly granulous upon its
external surface, tubercles upon the upper part not strongly marked; ridge in the
male nearly as in the preceding species. Ridge upon the mobile finger very finely
striated for the greater part of its length, and obscurely beaded towards the distal
extremity. Lines upon the carapace dark red, forming irregular loops and circles,
which are smaller and more crowded anteriorly, and interspersed with spots. Length
and breadth 1} in.

Hab. Fiji Islands; New Hebrides.
In the female the same differences in the sculpture of the hand are observed as in the

preceding species.

9. Matura LunaRis. (Plate XL. figs. 10 & 11.).
Cancer lunaris, Herbst, Krabben u. Krebse, iii. p. 43, pl. xlviii. fig. 6 (1799). Jvo!
Carapace very finely granulated; tubercles of the carapace and posterolateral margins
obsolete (except in quite young specimens). Lateral marginal spines acute, of
moderate length. Front divided by a triangular emargination into two sharply
defined acute lobes. Hand not granulous, or very faintly granulous below; tuber-
cles upon the upper part strongly marked; ridge upon the external surface in the
male divided into five lobes, of which the second is acute, spiniform, and the others
small, tuberculiform. Ridge upon the mobile finger obsolete. Lines upon the
' carapace pale red, more interrupted than in the preceding species. Length and
breadth about 13 in.
Hab. Yndian Ocean, Ceylon.
The form of the hand is almost exactly the same in the maie and female of this
species.

248

MR. BE. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

DESCRIPTION OF THE PLATES.

Puate XXXVITL.

Illustrations of Species of the Family Leucosiide.

Fig. 1. Leucosia fuscomaculata, sp. n., male: natural size.

Fig. 2. The same: lateral view of its thoracic smus, enlarged.

Fig. 8. The same: outline of the abdomen, natural size.

Fig. 4. L. pulcherrima, sp. u., female: natural size.

Fig. 5. The same: lateral view of its thoracic sinus, enlarged.

Fig. 6. The same: outline of the abdomen, natural size.

Fig. 7. L. ornata, sp. u., female: natural size.

Fig. 8. The same: a lateral view of its thoracic sinus, enlarged.
Fig. 9. The same: outline of the abdomen, natural size.

Fig. 10. L. brunnea, sp. n., female: natural size.

Fig. 11. The same: inferior view, showing angular sinus, enlarged.
Fig. 12. The same: outline of the abdomen, natural size.

Fig. 18. ZL. reticulata, sp. u., male: natural size. ;
Fig. 14. The same: lateral view of its thoracic sinus, enlarged.
Fig. 15. The same: outline of the abdomen, natural size.

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. 16. L. whitmeei, Miers, male: natural size.

g. 17. The same: lateral view of its thoracic sinus, enlarged.

g. 18. The same: outline of the abdomen, natural size.

.19. L. perryi, sp. u., male: natural size.

. 20. The same: lateral view, showing absence of thoracic sinus, enlarged.

21. The same: outline of the abdomen, natural size.

. 22. L. pubescens, male: natural size.

23. The same: lateral view of thoracic sinus, enlarged.

. 24. The same: outline of the abdomen, natural size.

. 25, Myra mammillaris, Bell, young male: natural size.
. 26. The same: lateral view, enlarged.

. 27. The same: outline of the abdomen, natural size.

. 28. Nursia plicata, Herbst, male: natural size.

. 29. Arcania granulosa, male: enlarged.

. 80. Cryptocnemus holdsworthi, female: enlarged.

Fig.

S:

31. The same: outline of its abdomen, enlarged.
32. The same: arm of female specimen, greatly enlarged.

Puate XXXIX.

Illustrations of Species of the Genus Matuta.

. 1. Matuta victrix, Fabr., male: natural size.
. 2. The same, hand of maie: natural size.

. 3. The same, hand of female: natural size.

© Cc “EO Or be CO

MR. E. J. MIERS ON THE OXYSTOMATOUS CRUSTACEA.

(© COND OF SB

. M. victrix, var. crebrepunctata, sp. n.: natural size.
. M. rubrolineata, sp. n.: natural size.

. The same: hand of male, natural size.

. M. lineifera, sp. u.: natural size.

M. granulosa, sp. n.: natural size.

. The same: hand of male, natural size.

Prats XL.

Illustrations of Species of the Genus Matuta.

. Matuta banksii, Leach: male, natural size.
. The same: hand of male, natural size.

. M. maculata, sp. n.: male, natural size.

, The same: hand of male, natural size.

. M, picta, Hess: male, natural size.

. The same: hand of male, natural size.

. The same: hand of female, natural size.

. M. obtusifrons, sp. n.: natural size.

. The same: hand of male.

. 10. M. lunaris, Herbst: female, natural size.
. 11. The same: hand of female, natural size.

249

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SPECIES OF LI

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[ 251 ]

XIII. On the Domestic Pig of Prehistoric Times in Britain, and on the mutual Relations
of this variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andamanensis, and
Sus barbatus. By Grorce Roxiestoy, W.D., F.R.S., F.L.S., Linacre Professor
of Anatomy and Physiology, Oxford.

(Plates XLI.—XLIII.)
Read June 15th, 1876.

PORTIONS of two skeletons of domestic pigs having been put into my hands by the
Rev. William Greenwell, F.S.A., from an interment of the so-called late Celtic period, 2. e.
of the ultimate or penultimate century before the Roman conquest of this country,
I determined to compare them with such other specimens of Swide as might by any
possibility be genetically connected with them. Among these other specimens I may
mention, first, several specimens of the Wild Boar, Sus scrofa, var. ferus, from the alluvial
deposits of this neighbourhood, and now in the Geological Series of the Oxford Museum,
under the charge of Professor Prestwich, F.R.S.; secondly, five specimens of the Indian
Wild Hog, Sus cristatus, kindly lent me by Sir Walter Elliot, K.C.S.1., F.L.S.; thirdly,
two skulls of Sws andamanensis, presented to the Oxford University Museum by my friend
Prof. J. Wood-Mason, of the Indian Museum, Caleutta; and fourthly, four skulls
of Sus barbatus from Borneo. The extensive series of skulls of Suidze contained in the
British Museum, those in the Royal College of Surgeons of London, and the specimens
of wild- and domestic-swine skulls contained in our own collection were also used for
this comparison.

It may be well at the outset to specify the several points of wide and general interest
upon which such an inquiry as the ensuing may be brought to bear. First among these
I would mention its bearings upon the now so commonly discussed questions relating to
the early migrations of our own species. The pig was one of the earliest, possibly the
very earliest, of animals which man domesticated; and the question of the source or
sources whence it was derived has consequently an “ethnographisch-archzeologische Bedeu-
tung” (to use the words of Fischer *, in his analogous investigation as to the sources
whence the jade and nephrite of early European times were procured) of the first import-
ance. Gibbon + has remarked that “man is the only animal which can live and multiply
in every country from the equator to the poles ;” and he has proceeded to aver that “ the
hog seems to approach nearest to our species in that privilege.’ Asa matter of fact,
Gibbon here, as so often elsewhere, was very nearly though not quite exact: the north-
ward limit of the range of the Wild Boar may perhaps be taken as somewhere between

* H. Fischer, ‘ Nephrit und Jadeit nach ihren mineralogischen Eigenschaften.’? Stuttgart, 1875. For similar
investigations as to the sources of the cultivated plants and the weeds of prehistoric times, see Keller’s ‘ Lake-
Dwellings,’ translated by Lee, pp. 303 and 343.

+ ‘ Decline and Fall of the Roman Empire,’ chap. ix. note 9, p. 352. Smith’s edition.

SECOND SERIES.—7ZOOLOGY, VOL. I. 21

252 PROF. ROLLESTON ON THE DOMESTIC PIG

55° * and 60° N., and that of the tame Pig as 64° N., whilst that of the Common
Fowl + extends probably a little, and that of the Dog certainly much, further northward.
The mention of the Australian Dingo eee another amendment of the somewhat
cynical remark of our great historian.

Secondly, a very useful light is thrown, or may be thrown, upon the question of the
extent to which the influences of civilization act upon our own species, by the analogous
inquiry into the effects which domestication has been able to produce upon an animal
linked so closely with ourselves in a many-sided commensalism. The least pleasant
aspect of that commensalism, that, namely, which is presented to us by the facts of our
solidarity with swine in the maintenance of the alternation of the forms of life of Tenia
and Trichina, must, it may be remarked, upon any view of the origination of the four
species concerned, force upon our attention considerations of the very greatest gravity.
To take somewhat lower ground, from such a study as that of the variations of the Pig
under domestication, we may obtain safe criteria for estimating the relative effects of
food, whether scanty or abundant, of early or late exercise of the sexual functions, and
of intercrossing, upon the formation of facial characteristics. The importance of not
overlooking the influences of sex and of age is nowhere more forcibly pressed upon our
attention than in an examination of a series of skulls of Suide. It has often been over-
looked in disquisitions on the skulls of Hominide.

Thirdly, whether the question at issue as regards man between the Polygenists and the
Monogenists will, as has been predicted (Darwin, ‘ Descent of Man,’ ed. 2nd, p. 180,
1874), “die a silent and unobserved death” or not, there can be no doubt that illus-
trations of the argumentations whereby that question has been, or ought to have been,
dealt with, can be furnished nowhere more fitly and fully than in an inquiry into the
distinctness or non-distinctness of the various races of swine.

Three distinct views have been advocated as to the relationship of the domestic to the
wild swine, Sus scrofa, var. ferus. We may take as the first of these that advocated by
Professor Steenstrup t, and stated by him in the following plain words :—‘ Il n’y a pas
de transition 4 observer entre les sangliers et les plus anciens cochons domestiques.”

Tn this view Professor Steenstrup will find Mr. Samuel Sidney coincide in the opening
sentences of his work ‘On the Pig;’ and a view very closely similar to it was put
tentatively forward in the year 1821 by a savant who combined the functions of a Pro-
fessor of Materia Medica with those of Director of the Botanical Garden at Berlin,
Professor Link, in the following words, to be found in his work ‘ Die Urwelt und das
Alterthum,’ i. p. 192.

« Das zahme Schwein stammt nach allen Naturforschern von den wilden Schweinen
ab, und auch die Alten waren schon dieser Meinung §; doch scheint mir die Sache

* For the northward range of the Wild Boar and the tame Pig, see Brandt und Ratzeburg, Med. Zool. p. 89;
Fitzinger, Sitzungsberichte d. Acad. d. Wiss. Wien, 1864, p. 887; Radde, Reise im Siiden von Ostsibirien, i. 236;
Zimmermann, Geographische Geschichte, i. 189, 1778. Middendorff, in his ‘Sibirische Reise,’ p. 1062, 1867, gives
56° N. lat. as the extreme actual northward limit of the Wild Boar. (Since I wrote as above, Mr. H. N. Moseley has
procured for me a skull of a tame pig from Stene i B6, Lofoten Islands, 2° above the arctic circle.)

+ For the northward range of the domestic Fowl, Gallus domesticus, see Brandt and Ratzeburg, J. c. p. 150.

+ Bulletin du Congrés International d’Archéologie préhistorique 4 Copenhague en 1869, p. 163.

§ Varro, ‘ De Re rustica,’ 1. ii. c. 1.

OF PREHISTORIC TIMES IN BRITAIN. 253

keinesweges entschieden. Die Starke, Grosse und Farbe des wilden Schweines wiirden
keinen Unterschied machen, da die wilden Thiere starker, grésser, und dunkler gefarbt
sind, als die zahmen, aber die grossen Hauer des wilden Ebers scheinen doch nicht bloss
Vergrosserung zu sein. Die Feltdecke des zahmen Schweines findet sie niemals auch
bei dem fettesten wilden Schweine, die gestreifte Farbe des Frischlings ist sogar merk-
wurdig, die Stirn der wilden Schweine ist mehr gewélbt, die Ohren sind kiirzer, mehr
zugerundet, der Riissel langer, andere Verhaltnisse an den innern Theilen zu geschweigen.
Hs fehlt ganz an Beispielen, dass die Zihmung solche Veranderungen hervorgebracht.
Vielleicht stammt das zahme Schwein von einer orientalischen Art ab, welche gross aber
unschadlich sein soll, und hin und wieder in einigen Reisebeschreibungen erwahnt wird *.
Doch erfordert die Sache noch eine genauere Untersuchung. Das siamische Schwein aus
dem ostlichen Asien abstammend, ist ohne Zweifel eine besondere Art.”

Precisely the opposite view was held by Mr. Youatt, who at p. 35 of his work on “ The
Pig,” London, 1847, says, when speaking of the Wild Boar, “ no one can for a moment
doubt that itis the parent stock from which the domesticated breeds of swine originally
sprang.” Blasius (Saugethiere, 1857, p. 509), Fr. Cuvier (Hist. des Mamm., 1824), and
Giebel (Saugethiere, p. 225, 1859), would I apprehend, agree with the extreme view
enunciated by Mr. Youatt. An intermediate view is put forward by Riitimeyer in his
inestimable work, ‘Die Fauna der Pfahlbauten,’ pp. 186-190: according to him, what,
owing to the slow spread in Germany of improved breeds, may still be called das gemeine
Hausschwein, has originated from Sus scrofa, var. ferus ; whilst the Berkshire breeds, he
thinks, may owe their origin to Sus celebensis; and his Sus scrofa, var. palustris, ‘das
Torfschwein,” a domestic pig known as “ das Biindtner Schwein,” and Sus indicus may
represent a distinct stock, if not species. Nathusius, in his ‘Schweineschadel,’ p. 175,
agrees with Riitimeyer as to the origin of the large-eared race common in Central Europe,
but suggests Sus vittatus, of the islands of Java, Borneo, Amboyna, and Batchian, as the
parent stock of the widely spread domestic breed known as Sus indicus. In this latter point
he agrees with S. Miller. Fitzinger (/. c.) differs from Nathusius in supposing, without,
perhaps, adequate reasons, Sws leucomystax to be the parent stock of the Chinese, Cape,
Portuguese, and Cleveland breeds, whilst Sus cristatus, the Wild Pig of Hindostan, he
suggests as the parent stock of the Siamese and Sardinian races. Temminck (‘ Fauna
Japonica,’ p. 57, pl. xx., cit. Nathusius, /. ¢. p. 167) suggests that Sus leucomystax may
be the parent of the domestic Sws indicus; and though Nathusius, p- 167, demurs to
the enunciation of this statement as being definitely proved, his objection amounts to
little more than saying that Sus lewcomystax is probably not specifically distinct from
Sus vittatus, which Nathusius himself holds to be the parent stock of Sus indicus.

My own views, as based upon the data available to me, are to the following effect :—-
The prehistoric domestic swine which have come into my hands appear to me to be more
nearly affined to Sus scrofa than to any of the Asiatic wild swine with which I am
acquainted ; secondly, without wishing to affirm absolutely that too much weight has
been laid by Nathusius upon the shortness of the lacrymal for differentiating Sus indicus,
as we now see it, from Sws scrofa and its progeny, I am inclined to think that sub-

* Otter, Voyage en Persie, t. i. p. 207. D. Maillet, Description de l’Egypte, t. ii. p. 176.
22

254 PROF. ROLLESTON ON THE DOMESTIC PIG

equality, if not actual equality, between the malar and the orbital borders of the
lacrymal bone may be found in prehistoric skulls of the Sws scrofa, var. ferus, and
especially in the female skulls of that variety of Sus which, in other points, such as the
slenderness of the snout, differ from the “‘ Torfschwein,” the representative in those
times of Sus indicus, according to Riitimeyer ; and, thirdly, I think it is possible to show
that, whilst Sus cristatus, Sus leucomystax, Sus vittatus, and Sus timorensis form a closely
connected group of Swide non verrucosi, with which, again, Sus andamanensis and Sus
papuensis are to be allied, all these subspecies differ in points of considerable if not of
specific value from Sus verrucosus of Java, from Sus celebensis, and, finally, from Sus scrofa
of the Palearctic region as well as from the non-verrucose Sus barbatus of Borneo,

It may be well to begin with this latter point first, and to show that the group of
Eastern Pigs, of which the Wild Pig of India, the Malay peninsula, and the Lancay
Islands, Sus cristatus, may be taken as a type, is always distinguishable from Sus scrofa,
var. ferus, of Europe, and Asia north of the Himalayas. ‘This view is not equivalent to
one which should lay it down as certain that they are specifically distinct, a question
which it is not proposed to raise here. I shouldagree, however, with Mr. Jerdon, whose
book on the Mammals of India, 1874, came into my hands subsequently to the formation
of my opinion, in holding that the Indian Wild Hog was “ as worthy of specific distine-
tion as many other recognized species”’ (/.¢. p. 241), though this is not to say much.
I should not, however, entirely accept his statement to the effect that the head of the
Indian Wild Hog was longer and more pointed than that of the Kuropean Wild Boar,
though this has been laid down by Colonel Sykes (Proc. Zool. Soc. 1831, p. 30) as being the
state of the case; for in measuring the relative lengths of the nasal and fronto-parietal
regions of the Indian Wild Hog, I have come to think that precisely the reverse of this
statement is usually, though not invariably, the case, the Indian Hog having the nasal
bones shorter relatively to the rest of the roof-bones of the skull than Sus scrofa, var.
ferus. Neither do I agree with Colonel Sykes in holding that the straightness of the plane
of the forehead will differentiate the Indian at least from modern European Wild Boars:
some concavity is produced in the mesial contour-line of large Prehistoric Wild Boars,
not by any angulation at the junction of the facial with the cranial bones, as in modern
tame swine of highly cultivated breeds, but by the upgrowth of the back portion of the
skull roof-bones and the occipital transverse crest ; modern European Wild Boars, how-
ever, which are much inferior in size to their prehistoric and, indeed, to their medizeval
predecessors, have the fronto-parietal and nasal lines forming one unbroken straight line.

Colonel Sykes’s words, “ Tail never curled or spirally twisted,” appear to me (J. ¢. p. 11)
to be said of the tame variety of the Indian Hog; but though Fitzinger (Sitzungsbericht.
Akad. Wiss. Wien, 1864) specifies the form in which the caudal vertebree are carried as
one of the specific marks in each of his descriptions of the Swid@, and though Linnzeus
uses the words (Hd. xiii. Syst. Nat. p. 65) “Cauda sinistrorsum recurvata” for differen-
tiating Canis familiaris from Canis lupus, s. Canis cauda imcwrvata, | am inclined to
think too much weight may be laid upon this point.

Thoroughly trustworthy figures of the European Wild Boar, such as that given by
Schreber (Saugethiere, taf. cccxx.), or that in Buffon’s Hist. Nat. v. pl. xiv., and Fréd.

OF PREHISTORIC TIMES IN BRITAIN. 255

Cuvier, Hist. Mamm., represent it as having the root and tip of its tail lying evenly
between two points; though the Vienna zoologist just referred to says of this appendage
in this animal, “‘Schwanz geringelt, kurz, nicht ganz bis zum Fersengelenke reichend.”’
And as an indication of the trifling value of such a point as this, it may be remarked that
of two female specimens of the very well-marked species, Sus barbatus, one young, the
other old, figured by S. Miller, Verhand. i. taf. 30, one has the tail curled, and the other,
the elder one, has it straight.

Mr. Blyth (cit. Jerdon, 1. ¢.) holds 8. cristatus to be only a variety of the Wild Boar of
Europe, but still to be a well-marked race. De Blainville (Ostéog. Sus, p. 129) sees no
differences of morphological importance between any of the Asiatic Swine and the Euro-
pean Wild Boar, and says, “‘ La premiére espéce que le squelette nous permette de distin-
guer par des caractéres susceptibles d’étre lus et exposés est celle quise trouve dans toute
l Afrique au dela de |’ Atlas et jusqu’a son extrémité la plus méridionale et méme au dela
dans la grande ile de Madagascar, et qui est connue sous le nom de Sus larvatus”’ (Pota-
mocherus africanus, Porcus madagascariensis, nobis). Giebel follows De Blainville in

this. Dr. Gray, in Proc. Zool. Soc. 1852, p. 180, said that, after examining ten skulls of the

European Wild Boar and its offspring from this country, from the Gambia, and from the
Cape, as also twelve skulls of the Wild Hog from Continental India, he could not discover
any constant easily described character by which the European and the Indian kinds could
be distinguished. And, he adds, “ this is the case in the many other genera allied to
the Pigs.” It is true, no.doubt, that many animals, such as “ the lion and the tiger,
the fox and the jackal, the ass and the zebra, are far more strikingly differentiated by
their pelage than by their skulls,’ as Professor Huxley (‘ Prehistoric Remains of Caith-
ness, p. 132) has taught us; still it yet remains to be proved that differences which,
though only skin-deep, are constant and permanent, will not ultimately be found to be
correlated with more or fewer differences of the deeper-lying parts, either of a purely
qualitative or of a quantitative kind.

In a disquisition the ultimate object of which is the attainment of clearer views as to
the origin of our tame pigs, the question meets us at the outset whether there exists
any marked difference between the wild stocks under comparison as regards their suscep-
tibility of domestication. Upon this point I have to say that I find, in opposition to
what Mr. Samuel Sidney has laid down in the first chapter of his work on the Pig,
that Sus scrofa, var. ferus, is credited by most trustworthy authors with as great a
capacity for domestication as any wild animal, including its wild Asiatic congeners, upon
which observations are recorded as to this particular susceptibility. Pallas’s words in his
* Zoographia,’ i. p. 269, as to the Wild Boar of the Palearctic Region, called by him, not
inconveniently, Sus ewropeus, in contradistinction to the China or Siam Pig, called by
him Sus indicus, says plainly and emphatically “ Porcelli cicurari assuescunt facile et cum
domesticis generant.” Radde’s utterances (Reisen im Siiden von Ost-Sibirien, Bd. i.
p- 286) are even more to the point, as they affirm the like of wild swine of greater age.
They run thus “So muss ich gestehen dass sie sehr friedlicher Natur sind, und es mir
mehrmals passirte mittelalte Wildschweine sich mir bis auf vier Faden Weite nahen zu
sehen.”

256 PROF. ROLLESTON ON THE DOMESTIC PIG

The Asiatic pigs, secondly, of the group represented by Sws cristatus, though not, within
my knowledge, those known as Sus verrucosus, nor those known as Sus barbatus, have very
_similar and very numerous testimonies borne to their educability and capacity for attach-
ment to man. Fitzinger, indeed, says of the domesticated Chinese Pig, his “ Sus leuco-
mystax sinensis,” that it resembles the domesticated European Pig generally in its habits
and character, but that it shows much more attachment than the European farm-pig to
the persons who take care of it, and will even follow them about, although it is other-
wise troublesome and obstinate. (Sitzungsberichte d. Akad. Wiss. Wien, 1858, Bd. xxx.
p. 235). In Formosa, when the Dutch first became acquainted with it, in the beginning
of the 17th century, every native woman, we are informed, on the authority of Ogilby
(Atlas Chinensis, ii. p. 8, ci¢. by Swinhoe Proc. Zool. 1870, p. 643), had “a great pig
running after her, as we use to have a dog.” A closer intimacy than this has been
observed to exist between Homo sapiens asiaticus, Linn., and Sus sinensis, Linn., by Pro-
fessor Huxley (cit. Galton, Trans. Ethnol. Soc. vol. iti. 1865, p. 127), who has seen
sucking-pigs nursed at the breasts of women, apparently as pets, in islands of the New-
Guinea group. As regards the wild races, Sir Walter Eliiot tells me, in a letter of date
May 15, 1876,—“ I have seen the young of Sws cristatus, which had been captured by
some of the Indian nomad communities, and reared by them, running about among the
domestic stock ; so that it would be hard to say where the line should be drawn”’*. ‘The
wild Sws papuensis has been found by Europeans, as we are informed by M. R. P. Lesson
(Voyage de la Coquille, 1826, vol. i. p. 176), to be more thoroughly domesticable than the
half-wild state in which some of the natives are content to leave it would have led us to
expect, and to set up relations of mutual amity, not only with the human, but also
with canine companions.

_ As regards the Aithiopian region, the Wild Hog is reported to us by Dr. Barth + as
consorting on terms of perfect amity, and, indeed, intimacy, with other domesticated
animals than the dog, and also with the natives. These are his words relating to a
district in Central Africa :—‘*‘ Naked young lads were splashing and playing about in
the water together with wild hogs in the greatest harmony; never in any part of
Neegroland have I seen this animal in such numbers as here about the Shari. Calves
and goats were pasturing in the fields with wild hogs in the midst of them.”

It is impossible to be perfectly certain what wild hog this may have been; still it can
scarcely have been any other than Sus sennaariensis, which has been supposed (see Darwin,
‘Domestication,’ i. p. 71, 2nd ed.) by J. W. Schiitz to have been the parent stock of
Sus scrofa, var. palustris, of Rittimeyer, but underneath the entry of which, in the
British-Museum Catalogue of 1869, p. 338, I find the following note:—‘ Dr. Murie
says he has often seen and eaten the true Wild Boar of the genus Sus in Africa, as
well as the Potamocherus, on the west coast. I have never seen any, or the skull
of one.” Like Dr. Gray, I have had no quite satisfactory opportunities for forming an

* Tt would appear that this difficulty has been felt by others to be a very real one. Colonel Walter Campbell
tells us, at p. 325 of his ‘Indian Journal,’ 1864, that he fears “the young gentlemen of the present day have taken
to spearing village pigs instead of wild boars,” and that he has “seen the thing done before now.”

+ Referred to by Nathusius, p. 147, /.c.; or see his ‘ Travels in Central Africa’, vol. iii. p. 311, 1857.

OF PREHISTORIC TIMES IN BRITAIN. 257

opinion, such as inspection of authentic skulls gives, as to the relations of Sus sennaariensis
to Sus scrofa; but whilst going over the series of skulls of Suidz in the British
Museum, I came upon one which, though entered under the head of Potamocherus
africanus, ought, I make no doubt, to be entered under the head of Sus sennaa-
riensis, or, aS I should prefer to call it, Sus scrofa, var. africanus. This skull is
numbered 715 a; and of it we have the following history from Dr. Gray (Proc. Zool. Soc.
1868, p. 35; Brit. Mus. Catalogue, 1869, p. 342) :—* A skull without its lower jaw
(715 a) was brought home by Captain Alexander from his Expedition to Damara, and
presented to the British Museum. It is recorded in Mr. Gerrard’s ‘Catalogue of the
Bones in the British Museum’ as Sus capensis (p. 277). It is the skull of an adult
animal, with the crown of the grinders much worn. It is probably the skull of a
female, as it agrees with all the characters of Potamocherus; but it has only a well-
marked ridge across the upper part of the base of the sheath of the upper canine, and
the upper margin of the nose is not dilated nor swollen.”” When I took this skull into
my hands I was uncharitable enough to suggest to Mr. Gerrard that it was a skull of
Sus cristatus wrongfully assigned to Damara as its habitat. Leaving this low ground, I
came to think that it might have come from some descendant of pigs of the Sus indicus
breed which had run wild at the Cape and reverted to the Sus cristatus form. But I
have now no doubt that this isa skull of such a Wild Boar as those of which Dr. Murie
speaks; and if this be so, the Sus of Africa is not readily to be distinguished from Sus
eristatus, at least by cranial characters. Dr. Gray, under the heading Sus sennaariensis
(Proc. Zool. Soc. ¢. c. p. 32, and British Mus. Catalogue, 1869, p. 338), suggests that the
skull figured by De Blainville (Ostéographie, tab. v.) may have belonged to a Sus sen-
naariensis. It is described as “ Sus scrofa egypti” by De Blainville; but, as far as I
can judge from the drawing, it has all the characters, to be hereinafter detailed, which
distinguish Sus scrofa, var. ferus, from Sus cristatus, and consequently from such a
skull as that labelled 715 a, and brought home by Captain Alexander. Of course there
is no @ priori difficulty in the way of our supposing that the Wild Boar either of
Palestine or of Algiers, both well-known animals, may have extended into Hgypt, a
country which has so much both of the Palearctic and of the Aithiopian fauna in oceu-
pation of its territory. Anyhow Dr. Murie and Dr. Barth are sufficient witnesses to the
fact that a true Sus is found in Africa south of the Atlas and Sahara. I cannot, therefore,
accept Mr. Wallace’s statement (‘Geograph. Distrib. of Animals,’ vol. i. pp. 253, 286-322)
to the effect that a true Sus is not to be found in the Athiopian region. The mistake
made, and handsomely acknowledged, by Fitzinger (Sitz. Akad. Wiss. Wien, 1864, Bd. 49,
i. p. 389), in naming certain young specimens of Sus sennaariensis by the name which
Fr. Cuvier had given to the Masked Boar, supposing them to be the young of that species,
would not have occurred with adult specimens. What Fr. Cuvier called Sus larvatus, we
know as Potamocherus africanus ; to save further confusion of names and errors of fact,
it would be well to drop the name Sus larvatus altogether. Potamocherus, which I
observed to resemble Sus cristatus in having a large lacrymo-frontal ridge, is, of course,
specifically distinct from it. Sus sennaariensis, if,as I think is most likely, closely
allied to Sus cristatus, is another instance of the wide distribution of ‘“‘ Pachyderms,” a

258 PROF. ROLLESTON ON THE DOMESTIC PIG

point on which Riitimeyer insists, in a different tone (Herkunft unserer Thierwelt,
p. 34) from that adopted by Gibbon, but not less categorically, nor, indeed, less strikingly,
remarking, as he does, that at the present day the Hyrax and the Hippopotamus are the
only genera of ‘‘ Pachyderms” confined to one quarter of the globe.

The Athiopian region therefore must be held to possess a true Sus; and as to domestica-
bility, the Palearctic, the Oriental, and the Mthiopian Suidze have possibly equal claims*.

There would be little weight to be laid upon mere differences in size, even if much
ereater differences did exist between Sus scrofa, var. ferus, and Sus cristatus than a
comparison of the measurements given in the British-Museum ‘Catalogue of Carni-
vorous Pachydermatous and Edentate Mammalia,’ 1869, pp. 334, 337, 338, or a compari-
son of the skulls themselves shows to be the case; for Pallas’s+ words and Nathusius’s ¢
are both clear to the effect that differences of as much as 50 per cent. do exist between
individuals of the Sws scrofa, var. ferus; and a comparison of what I believe to be the
skull of an old Wild Sow, from the alluvium of the Thames, near Oxford (Pl. XLI. fig. 2),
with that of a modern Wild Boar (Pl. XLII. fig. 5), will put the matter beyond all
question.

According to Nathusius }, the differences in size between the largest and the smallest
wild swine are so considerable as to have caused them to be distinguished by such names

* Pigs fulfil excellently well the six conditions enumerated by Mr. Francis Galton, 1. c., as necessary for domesti-
cation, viz.:—1. That the animal should be hardy; 2. That it should have an inborn liking for man; 3. That it
should be comfort-loving; 4. That it should be found useful; 5. That it should breed freely; 6. That it should be
gregarious. Adsop, Ailian, and Lactantius (cit. by Bochart, Hierozoicon, i. 698) have, in various ways, remarked on
the peculiarity of the pig as contrasted with other domestic animals, in that it is usetul only when dead, giving
neither milk, as does the cow, nor wool as does the sheep. With this peculiarity is connected the fact, useful for the
often difficult task of deciding whether a particular skull came from a wild or a domestic breed, that domestic pigs
are usually made useful while young. Riitimeyer, indeed (J. c. p. 52), gives it as one of his reasons for supposing Sus
scrofa, var. palustris, to be represented by a wild as well as by a tame stock, that its remains are usually those either of
yery old or quite young individuals. A pig will father while quite young; and whilst gaining nothing in its capacity
of manufacturer of food in its own body, it loses in its capacity of a breeding animal with increase of age. This is
not the case with the cow; and the discovery, therefore, of remains of very old individuals of this species only jns-
tifies us in inferring that the cow was a scarce and valuable animal in the period and place to which it belonged.
See Riitimeyer, ‘ Fauna der Pfahlbauten, p. 10.’

The special value of the pig as a domesticated animal is commonly expressed in an estimate that “twice the weight
of tood may be obtained from hogs than can be obtained from the same cost of food by means of any other animals ”
(Richardson, 7. c. p.42). In a little more detail, it is to be remarked that the pig, as a meat-producer, stands at an
advantage (to the consumer), first, in the smaller relative weight of its ‘‘ offal” as compared with the entire weight
of its body, but secondly, and chiefly, in the large proportion of fat, the kind of food which is eminently the hardest
for a savage or for the poor to procure, which it will store up upon almost any dietary. For this, see Lawes and
Gilbert’s invaluable Paper in the Royal Society’s Transactions for 1859, ‘‘ On the Composition of some of the Animals
fed and slaughtered as Human Food,” page 565 for relative proportion of offal, pages 513 and 5438 for storing up of
fat. It may be here remarked that De Blainville (Ostéographie, G. Sus, Introduction, p. 107) may very likely be
right in suggesting that the pig may have furnished animal food to the earliest races of man before either cow or
sheep, but that he could not have had our knowledge of the very various kinds of animals which, even in these days,
furnish lower races of men with animal food, when he supposes that animal food must have been supplied by one or
other of the three animals named. And his reason for such truth as his conclusions contain is, it is to be feared, but
a poor one; it runs thus, “le Cochon étant Vanimal qui sympathise le moins par ses qualités affectives avec l’espéce
humaine adi étre celui qu’elle aura le moins répugné 4 tuer de sangfroid pour s’en nourrir ; ce qui aura eu lieu plus
tard et avec bien plus de répugnance pour le Mouton et pour le Beeuf.”

+ Zoographia, 1831, p. 267. + Schweineschidel, 1864, p. 65.

OF PREHISTORIC TIMES IN BRITAIN. 259

as “ Hauptschweine”’ and “ Kiimmerer”’ respectively ; the latter of which terms has an
equivalent in the word ‘“‘ Wreckling,” applied in some parts of England to the super-
numerary pig in a litter, 7. e. to the one which makes the litter exceed the number of ayail-
able teats, and fares accordingly. Two or three of the commonly reported facts * as to the
pairing and period of reproduction of the wild swine account very sufficiently for these
great differences in their size. Though the males are monogamous, severe battles never-
theless take place between them for the possession, it is said, of the largest females; the
smallest females consequently are left for the vanquished, which will usually be the
smallest males. Hence a great difference in the two sets of offspring would be reasonably
expected. But, further, it is known that the wild, like the tame + swine, will breed long
before the period of maturity ; and that the offspring of such unions, whether both of the
parents or only one be immature, are likely to be smaller in size as well as fewer in
number, needs no argument.

Whilst no & priori probability can be gathered from any greater domesticability in
favour of the claims of either Huropean, Asiatic, or African Sus to be the exclusive source
of our domestic pigs, and whilst mere size equally fails to differentiate these races,
the point of the relation between the length and the height respectively of the lacrymal
bone on which Nathusius has laid such weight (Schweineschidel, passim et pp. 9, 10, 838,
91, 92, 175), and to which Mr. Darwin has assigned so much importance (‘ Animals and
Plants under Domestication,’ vol. i. p. 70, ed. 1875), though in the immense majority of
cases enabling us at once to differentiate the skulls of Sus cristatus, as indeed of the other
Asiatic pigs without facial warts, from those of Sus scrofa, var. ferus, also sometimes fails
us. Having measured a very considerable number of skulls of Sus cristatus from very
various parts of India, and having invariably found them to have the orbital border of
the lacrymal shorter than, or at most only just equal in length tothe malar, and bearing
in mind the constant reference made by Nathusius to the morphological and classificatory
value of this proportional difference in the tame variety, the so-called Sws indicus, I was
entirely unable to understand how that author could say (/. ¢. p. 185) that two skulls of
Sus cristatus furnished to him by Mr. E. Gerrard differed from skulls of Sus scrofa, var.
ferus, only in being smaller altogether. But after measuring the skulls from Sir Walter
Elliot’s collection and those in the British Museum, with the result of feeling certain
that, from the contour, proportions, and, in adult males, the texture and sculpturing,
together with the lacrymal of Sus cristatus, it was always possible to distinguish such
skulls from those of our Wild Boar, I came, in the Royal College of Surgeons, upon a
skull which, whilst possessing certain other peculiarities (to be hereafter detailed) as
distinctive, more or less, of Sus cristatus, did combine with them the long lacrymal of
Sus scrofa, var. ferus. This skull is numbered 3251 a, and was pointed out to me by
Professor Flower, with his often experienced kindness, as being a skull of Sus crzstatus,

* See Richardson, ‘ Domestic Pigs,’ pp. 18, 19 (Warne, London); Samuel Sidney, ‘The Pig,’ p. 4 (Routledge,
London) ; Blasius, ‘ Siiugethiere,’ p. 509, 1857 ; Wagner, ‘Saugethiere, p. 426, 1835; Brandt & Ratzeburg, ‘ Medi-
cinische Zoologie,’ p. 88, 1829.

+ For period at which tame sows will breed, see Sidney U. c. p.61; Low, ‘On the Domesticated Animals of the
British Islands,’ p. 415.

SECOND SERIES.— ZOOLOGY, VOL. I. 2M

260 PROF. ROLLESTON ON THE DOMESTIC PIG

which, together with two others, with the same appearance as to textural condition,
had been brought out of the stores of the Museum into its series. Both these other
skulls possessed the short lacrymal so characteristic, as the subjoined tables of measure-
ments will show, of Sus cristatus and its allies, together with the other points usually
observed in that animal’s skull. The principal points, besides those of general facies
and proportions, which appeared to me to justify this assignment of skull 32514 (Royal
College of Surgeons) were, in the absence of the shortness of the lacrymal:—first, the
great prominence of what may be called the lacrymo-frontal ridge, that part of the
frontal bone, to wit, which les between the channel for the supraorbital nerve mesially
and the upper border of the lacrymal bone outwardly ; secondly, the great relative deve-
lopment of the part of the third molar which is posterior to the two anterior bicus-
pidate lobes of that tooth; thirdly, the absence of convexity backwards in the naso-
frontal suture. These three points are usually present in Sus cristatus, as seen in
figures 4 and 6, appended to this paper, and taken from two Indian specimens; and they
are usually found to be accompanied by the fourth peculiarity of a short lacrymal. One
or other of these characters may be absent; but in an undoubted specimen of an adult
male Sus cristatws | have never seen more than one of these missing; whilst it is rare
for the second, and very rare for either the first or the third, to be found in undoubted
specimens of the Palearctic Wild Boar. Ina skull figured by Mr. Richardson (/. ¢. p. 50),
from “an excavation in an island on Loch Gur, a lake in the neighbourhood of Limerick,”
and ‘“‘ found in company with skulls of oxen, goats, sheep, red deer, reindeer, and our
extinct gigantic deer, sometimes erroneously styled the Irish Elk,” but considered by
Nathusius (p. 150) to have belonged to a domestic animal, it is true that the lacrymo-
frontal ridge is represented as of great size ; but we must set against the assigning of much
importance to this fact the considerations that the drawing is taken from a reconstructed
skull, that it is obviously inaccurate in some points, as, for example, like a drawing in
S. Miller’s Verhandel. (Taf. 28 bis, fig. 3), im having an extra tooth posteriorly to its
canines, and that it may consequently be supposed to he likely to be inaccurate in other
particulars also. The fronto-lacrymal ridge is, of course, in the adult underlain by a pro-
longation of the frontal sinuses; it is, however, visible enough in very young specimens
of domestic pigs, which show other points of affinity to the Sws indicus long before the
frontal sinuses are fully developed; and I am ixclined to think it may sometimes,
though certainly not always, be detected in very young specimens of Sus scrofa, var.
ferus, such as the one figured by Nathusius, J. c. (Taf. 1. fig. 1, Taf. 1. fig. 13). Though
the fulness of this region is due in the adult partly to its beimg underlain by frontal
sinuses, which are relatively small in the early stages of the animal’s life, there
is still some justification for regarding this structure as an instance of the retention by
the adult of an early structural arrangement; for it is easy to understand that the
contour described by the external tables in early youth may be carried out conform-
ably by the bloodvessels of the scalp as the animal grows older. A parallel to
such a process is furnished to us very frequently, though by no means so nearly
universally, by the retention in the adult Sus eristatus of that fulness and convexity of
the vertical aspect of the fronto-parietal region which is characteristic of Sws scrofa

OF PREHISTORIC TIMES IN BRITAIN. , 261

both wild and domestic, as well as of Sus cristatus and Sus indicus, at birth*. Lieu-
tenants W. E. Baker and H. V. Durand, in their paper on “ Subhimalayan Fossil
Remains of the Daduptr Collection,” in the Journal of the Asiatic Society of Bengal,
vol. v. 1886, p. 664, observe of the two fossil skulls which they describe, that in both
“there is in the frontal plane a total absence of convexity. As this plane ascends
there is a tendency to concavity, in consequence of the parietal crests being more
strongly marked than in the existing species, and thus producing the appearance of
a gentle hollow where, in the common Wild Hog, there would be a gentle swell.”
Further on, in the same paper, the authors remark :—“ From the form of the cranium,
the shape of the canines and incisors, and the other points in which the fossil differs
from the existing species of the country, a specific difference may be inferred; for the
dissimilarity, although less than that which occurs between the Babyrussa, the 8. larvatus
and the Sus scrofa, or Common Hog, is too remarkable, particularly in the shape of the
canines of the lower jaw, to admit of the fossil bemg considered as a mere variety of
the Sus scrofa.” Sir Walter Elliot, however, to whom I owe this reference as well as
other things, writes me to the effect that the skulls sent by him to me “do not seem to
differ much from the Subhimalayan fossil specimens figured and described,’ as above
specified. And it is worthy of being put upon record that of five skulls of modern Wild
Indian Hogs thus sent by Sir Walter Elliot, three show the upgrowth of parietal crests,
which Lieutenants Baker and Durand had supposed to be characteristic of the fossil
animal, and to contribute towards justifying its claim to be considered specifically dis-
tinct. These three skulls have the following labels and histories appended to them by
Sir Walter Elliot, which, when coupled with the localities assigned below to the British-
Museum specimens (specimens not, so far as I can see, different in any essential point
from Sir W. Elliot's), bear importantly on the question of the unity of Sus cristatus :—
No. 71. Large Boar, killed near Rajkote, in Kattywar, June 4, 1832. He was with
a large sounder, and ripped two horses severely. Rajkote is in the extensive
open plains of the Kattywar peninsula.
No. 330. Niigherry Hills, 1840.
No. 428. Jaggiapettah, 1851. On the east side of the Madras Presidency, in the
Masulipatam district, on the high road from Masulipatam to Hyderabad.

These three skulls agree in having their third molars considerably worn and their
canines large, their muscular insertion-surfaces marked with polygonal reticulations + in
some places, and with arborescent markings in others, and, thirdly, in the spar-like hard-
ness and density of the bones generally ; and they must be supposed consequently to have
belonged to old and powerful male animals. In all of these points they differ more or
less from the other two skulls, also of male but of younger and less powerful animals.
But such differences as these are far from being of specific value, either in comparison
of modern races with fossil ones, or in comparison of modern races inter se. All the five
skulls, however, lent to me by Sir Walter Elliot possess the lacrymo-frontal ridge
developed into a very considerable prominence; and though every now and then I have

* See Nathusius, pl. i. figs. 1 & 3, pl. ii. fig. 13, and pp.8 & 18. Compare the mesial fulness in the frontals of

S. papuensis. T For similar reticulation in Bos primigenius see Rutimacyer, Fauna der Pfahlbauten, Taf. iii. fig. 3.
2m 2

962 PROF. ROLLESTON ON THE DOMESTIC PIG

had occasion, in going over the extensive series of skulls of Indian Wild Hogs which
have been available to me, to note an almost, I have never noted an entirely complete
disappearance of it. The fronto-lacrymal ridge is not, even in the aged hog, a mere
expansion or dilatation of the frontal bone: it is underlain, it is true, by an arm of the
frontal sinuses; but it has thick and independent walls of its own. Though not quite
exactly homologous with the supraciliary ridges of human anatomy, it is nevertheless
very closely comparable with them. In some varieties of the human species the supra-
ciliary ridges, whether underlain by sinuses or not, are very constantly developed into great
prominences. In a classification, indeed, of human crania proposed by Dr. Williamson *,
“skulls with the supraciliary ridge so prominent as to overshadow the face”’ formed one out
of four classes into which all crania were divided. It is not necessary to go so far as this
for acknowledging that this character may become, in the Suidee at least, a race charactery.

The second in order of the three points which, taken together, as they usually exist
together, with the fourth, that relating to the lacrymal, help us to identify Sus cristatus, is
complexity of its third molar, at least in male specimens, and specially the complexity of
that part of the tooth which lies posteriorly to the two primary bicuspidate lobes.

A reference to my description of fig. 8 (where the third lower-jaw molar of Sws
andamanensis represents this often complex tooth reduced to great simplicity) will enable
the reader to understand this second point of distinction between Sus cristatus and Sus
scrofa more readily than any verbal description, however long, without such reference.
It may be sufficient to say here that the division of the third molar which lies posteriorly
to the two anterior lobes, is, in the male Sus cristatus, equal sometimes in size to and
sometimes even greater than both those other lobes taken together. This is not often,
though it is occasionally, the case in Sus scrofa, var. ferus, of modern times. Nor is it
the case in the females of Sus cristatus; so that the greater development observed in the
males may perhaps be due to the working of the law of sexual battle. The large size of
the canines postulates a large determination of blood to the jaw; and the large size of the
third molar, a tooth evolved at the period of sexual maturity, when the animal “ venerem
et preelia tentat,” may be, to use a word suggested long ago by myself }, “ tautogeneous ”
with it. Still the fact that the third molars are small both in Sus barbatus and Sus bar-
birussa shows that this smallness cannot always be explained on physiological grounds.

The third point, that of the straightness of the naso-frontal suture, appears to have
some classificatory value, whether we look at it with the light furnished by its
importance and validity in the classification or identification of other animals, or in that
furnished by the facts of its own developmental history.

The nasal and the frontal bones together form a roof over the ethmoid and the turbinal
bones ; and as there is no apparent physiological reason why they should contribute in

* Observations on Human Crania in Museum, Fort Pitt, Chatham, 1857.

7 It is observable in the pigmy Nepal pig, Porcula salviania, as also in the African Potamocherus. It does not
appear to me that male Suidz possess it more markedly than females; and herein it differs from the supraciliary-
ridge development as seen in our own species, as also from the race-mark to be next mentioned. An analogous
eminence, rudimentary in S. papuensis, occupies the middle frontal line over an area homologous with the human

glabella in each of four pig’s skulls brought from the Admiralty Islands by H.M.S. ‘ Challenger.’
~ Nat. Hist. Rey. Oct. 1861, p. 486.

OF PREHISTORIC TIMES IN BRITAIN. 263

different proportions towards the securing of this end, the fact that they do so is of so
much the greater morphological value. In Sus cristatus the naso-frontal suture very
ordinarily ruus, as may be seen in fig. 4(and in Sus andamanensis, in fig. 3), straight across
the roof of the ethmoid, at right angles to the long axis of the skull; or the frontals may
intrude themselves mesially between the nasals, making thus the contour of the suture to
be convex forwards. Precisely the reverse is the case in the adult European Wild Boar.
Some weight has been laid * upon a similar conformation in the skull of the Tiger, as
being of service in differentiating it from the skull of the Lion; and though it is not
pretended that an equally great distinctness can be supposed to exist between the two
animals now under comparison, still the structural differences in the two sets of cases
are analogous.

But when we come to look at the skulls of developing Pigs, we see that real value
attaches, from this point of view also, to the relatively greater or less extension back-
wards of the nasal bones, and the contour described consequently by the naso-frontal
suture. A tape crossed at right angles to the long axis of the skull from one infra-
orbital foramen to the other, passes very closely in front of, and often parallel with the
naso-frontal suture in the very young pig ; the suture gets removed further and further
away from it as the pig increases in age. Nathusius (Taf. iii. figs. 11 and 13) has
figured the skulls of a young Wild Boar and of an adult Wild Boar upon the same plate ;
and the straight line of the suture of the former contrasts most instructively with the
backwardly arching contour of the latter. The straightness, therefore, of the naso-frontal
suture may be supposed to illustrate the principle that climatic or other conditions may
cause structural arrangements to be permanently retained in certain races whilst they
are obliterated in others. The retention of the prolongation of the sagittal suture over
the frontal region is believed, with much reason, to be hereditarily transmitted in our own
species; and I incline to think that the persistence of the frontal tubera with very
much of their infantile eminence, which we see not rarely in adult men, may be taken as
furnishing another parallel to the retention in some degree, by the adult Sus cristatus, of
the characters of the young animal’s naso-frontal relations.

In the fourth place, as regards the value of the relative shortness of the lacrymal
bone as a means for differentiating the skulls of Sws cristatus and its allies from
those of Sus scrofa, var. ferus, I was for a long time of opinion that the same might
be said of this all but invariably observed peculiarity of Sus cristatus, which I have
already said, however shortly, of the straightness of its naso-frontal suture; and
that all this may, mutatis mutandis, be repeated as to the lacrymal’s peculiarities
in the two subspecies there can be no doubt. Anybody who will examine the figures
given in Spix’s ‘ Cephalogenesis, or the reproduction of them in Erdl’s ‘Tafeln der
vergleichenden Anatomie des Schidels, 1841, can convince himself of the fact that
the malar border of the lacrymal is very short as compared with its orbital border in the
young Pig +; and if he extend his observation to the various adult Suide, he will find

* Ost. Catalog. Royal College of Surgeons of England, 4506. p. 706.

+ Though I do not suppose thatit would be possible to say in 1876 what Dr. Gray said (Proc. Zool. Soe. 1868, p. 19)
m 1868, to the effect that Nathusius’s works were not to be had either in the Library of the British Museum, or in

264) PROF. ROLLESTON ON THE DOMESTIC PIG

that this side of the bone goes on growing from a condition of permanent inferiority in
Sus andamanensis till it reaches subequality or entire equality with the orbital length in
some Suidee—such as Sus cristatus, Sus vittatus, Sus leucomystar, Sus taiwanus, Sus
timoriensis, Sus papuensis, aud Sus barbatus, all Suidee without facial warts—and finally
exceeds the orbital border in length considerably in Sus scrofa, var. ferus, and. ordinarily
in Sus verrucosus (and Sus celebensis ?), and disproportionately, it may be added, in the
African Wart-Hog, Phacocherus.

There is no doubt that the younger the pig the greater is the distance separating its
lacrymal’s length from equality with or superiority to its lacrymal’s height; but the
subjoined tables and the description of my figures show that there is no constant relation
to be observed between the growth of the entire facial skeleton out of the short proportion
of early days into the elongated muzzle of the adult, and the longitudinal evolution of the
lacrymal factor of that snout. Thelongest snout, such as that of the adult Sus barbatus,
may, as Nathusius has remarked with surprise (p. 167), show a lacrymal with the
proportions of the domestic Sus indicus, or the immature Sus scrofa, var. ferus; and
I venture to suggest that the elongation of the lacrymal, which Professor Owen taught
us to call a “ mucodermal bone,’ may be correlated with the evolution of the facial
warts which are found in all the Suidee with such lacrymals. If these two structures are
thus correlated, we come to be able to explain how it is that the length of the lacrymals
is, though not very variable, still as variable as we have found it to be; for the facial
warts themselves are a variable structure, as we should from several analogies expect them
to be. If they were not so, it would be difficult to explain how it is that in many
zoological descriptions * of our European Wild Boar, no mention is made of the presence
of warts (small ones, it is true) immediately below the eyes; and inasmuch as they are
so variable, and probably more liable + to disappear in the female sex, we can, on the
hypothesis of the evolution of the Jacrymal in length being correlated with their

the library of either the Zoological, Royal, or Linnean Society, it may nevertheless be convenient to give here the
results of his measurements of the borders of the lacrymal bones at different periods of the life of Sus serofa. They
stand thus at p. 10 of his work of 1864 :—

Newly born. 2 months old. 6 months old. Adult.

Lacrymal bone. millim. millim. millim. millim.
Ieleiohitiy ena eet ai 13 18 21
Frontal border ...... 11 20 42 60
Malar border........ 3 6 25 35
Ratio of height...... 1 Eg 2-6 3
Ratio of frontal border i 18 3:8 5:5
Ratio of malar border 1 2 8:3 11:6

* De Fatio, in his ‘ Faune des Veridbrés de la Suisse,’ 1869, p. 354, goes, if I understand him rightly, further
than this, by using the words “ Pas de saillies sur la face en dessous des yeux,” in his definition of “Le Sanglier ordi-
naire, das wilde Schwein.”

+ Dr. Gray, in his description of Potamocherus africanus (P.Z.8. 1868, p. 34), says of the male animal’s face,
that it is “ swollen and often warty on the sides in front,” and of the females, that “ the side of the nose is simple.”
Fitzinger, however, whose descriptions of the external characters appear to be carefully done, does not say that any
such sexual difference exists in this species or in any other of the Suide with warts. There are other reasons,
however, for the suggestions in the text, of which the sexual limitations of the facial callosities in the Orangs may be
taken as an example.

OF PREHISTORIC TIMES IN BRITAIN, 265

presence, understand how the lacrymal is sometimes found to be shorter than we should
have expected it to be in skulls such as those of the wild sows from prehistoric deposits
in this country, measurements of which are given below.

As the interpretation I suggest of the proportions observed to exist between the length
and height of the lacrymal bone depends upon the method of gradations, as applied not
only to several varieties, but indeed to several species of Suidee, as well as upon a history
of the development of the bone, it will be well now to give a short list of those species—
pointing out the particulars which justify their claim to that rank, and contrasting them
with the aggregate of characters which do enable us, as a matter of fact, to differentiate
such forms as Sus cristatus from Sus scrofa, possibly without justifying us in considering
them distinct species.

And first of Sws barbatus. I am in no way inclined to give too much weight to dif-
ferences in colouring or in character of hair or bristles; still a glance at the drawing of
this animal in 8. Miler’s Verhandl. Taf. 30, showing its half black, half tawny, wavily,
not crisply, curling beard, its ochraceous dorsal stripe, and its tail ending in a consider-
able brush, impresses one with the idea that it is impossible that the bony substructures
should not make some approach at least to a similar diversity from other forms of Suide.
This anticipation was fully borne out by an examination of four skulls, two of which are
in the British, and one in the Oxford Museum—and of Schlegel’s figure of the skull
described by him, and 8. Miiller, Verhandl. pp. 173, 179-181, and Taf. 31. figs. 4 &5.

The skull of Sus barbatus is absolutely longer than the skulls either of Sus scrofa, var.
Jerus, or Sus cristatus; and relatively to the body its length is considerably greater,
being, as given by Fitzinger, no less than # of the length of the trunk. The contour
described by its sagittal suture is, as my figures show, strikingly different from that
-deseribed by the corresponding suture in most other Suidze; its highest point is some
way in front of the plane of the occipital squama, and occupies a level far above the plane
occupied by the anterior half of the frontal or by the nasal bones; and these latter bones
make up more than one half of the entire length of the skull, resembling herein the
typical Suws scrofa, whilst the naso-frontal suture resembles that of Sus cristatus. Its
maximum interzygomatic width is in the middle of that arch, not at its posterior
border as in mest other Suide.

In addition to these larger points, the following may be mentioned as having a great
morphological importance, though relating to smaller structural peculiarities. The third
molar consists of three lobes; but, large as is the jaw and the canine armature of this Sus,
the most posteriorly placed of the three lobes is more simple than even the very simple
posterior lobe of the pigmy Sws andamanensis, having only one cusp prominently marked
in the upper, and four in the lower jaw, whilst the entire posterior lobe is little, if at all,
greater in antero-posterior extent, and much smaller in transverse, than either of the
anterior, differing thus altogether from Sus eristatus.

A second point, relating to a small structure as measured by the callipers, which is a
very large one, however, to the morphologist, is the permanent retention by Sus barbatus
of the mesopterygoid of Parker (Phil. Trans. 1874, p. 324, pl. xxxvi. fig. iv. ms.pg) as a
distinct bone. ‘This peculiarity was observed in all the four skulls of Sus barbatus

266 PROF. ROLLESTON ON THE DOMESTIC PIG

examined by me; and its obvious general significance is increased by the fact that it is
in the area of the mesopterygoid that the great basicranial cavities are excavated in the
Babirussa and Phacocherus, and are represented rudimentarily in Sus vitiatus.

Knowing what we do* of the affinity of the fauna of the subregion of Ceylon and
South India to that of Malaya, there is no @ priori improbability in a view which
should accept the Sus ceylonensis of Blyth + (Journal of the Asiatic Society of Bengal, xx.
p- 178) as identical with the Sus barbatus of Borneo. Mr. Blyth’s words, J. c., are as follows:
—“ Sus zeylanensis. Skull longer than that of the Indian Boar, nearly straight in profile,
very much contracted at the vertex. Palate contracting posteriorly to less than 1” from
the magnitude of the last molar, which is considerably larger in both jaws than in the
Wild Boar of India, the upper measuring 13” long by +8” broad anteriorly. Vertex nar-
rowing to 1” only in breadth. Total length of skull from vertex to tips of nasals 162”.
Altogether the skull approximates closely in contour to the figures of the skulls of Sus
barbatus by Dr. 8S. Miller and M. Temminck.’’ Dr. Gray appears to have had access,
which I have not had, to a photograph of this Sws zeylanensis, and says that, judging
from it, ‘the skull is much shorter and thicker than the skull of 8. barbatus. The pho-
tograph is much more like that of Sus verrucosus.” For my own part, I cannot think
Mr. Blyth would ever have spoken of a real male Sus barbatus as having a skull with a
straight profile ; Miller and Schlegel figured a female skull, whereas Mr. Blyth’s, I am
sure, was a male’s. As regards the contraction of the vertex, which shows Mr. Blyth’s
skull to have been a male’s, this is sometimes exceeded by the old and strongly muscular
Indian Wild Hog, such as Sir Walter Elliot’s No. 71 (for history of which see p. 261),
where it is only 8”, whilst in the skull No. 72 it is 1:8”, showing an oscillation, owing to
the varying action of the temporal muscles which entirely deprives it of any morphological
value; and the measurements of the molars, finally, assure me that there is no need to
add the words “ und wahrscheinlich auch Ceylon” to the word “ Borneo” as the “ Vater-
land” of Sus barbatus, as Fitzinger has done, J. c. p. 393. Miller and Schlegel, p. 179,
give Borneo, if I rightly read their words, as the “ habitat’ of this well-marked species.

Secondly, of Sus verrucosus. The soft and perishable parts of Sus verrucosus are even
more interesting and of greater importance than the bones; for they show us that wild pigs
do have appendages—warts, to wit, covered with long bristles, and attached to the corners
of the lower jaw, like those of the Irish Greyhound Pig, once so plentiful in Galway ; and
they thus do away with one of the objections to Mr. Darwin’s views, stated fairly (by him-
self) in the work on Animals and Plants under Domestication, i. p. 79, ed. 1875. If it
is a profitable thing to lay Midler and Schlegel’s figure of Sus verrucosus (tab. 28. J. c.)
alongside of Richardson’s figure of the “ Old Irish Greyhound Pig” (/. c. p. 49; Darwin,
l.c. p.79), it is profitable also to read the Dutch letterpress | of the two former authors,

* Tennant, Nat. Hist. of Ceylon, pp. 61-68, 186; Wallace, Geographical Distribution, i. p. 328.

7 See Gray, Proc. Zool. Soc. 1868, p. 24; Brit. Mus. Cat. 1869, p. 331.

+ The words of the two Dutch naturalists are, Verhandel. p. 177. ‘‘ De deer jonge voorwerpen dezer sort zijn niet
gestreept, en onderschieden zich daardoor van de jongen van Sus vitéatus en yan de meeste, ja, misschien van alle overige

soorten.” IJtis much to be regretted that Dr. Gray did not enter a note of this most important fact in his papers
above referred to, in which the ‘ Verhandelingen’ are often cited.

OF PREHISTORIC TIMES IN BRITAIN. 267

from which we learn, when helped by the translation given by Fitzinger, that the young of
Sus verrucosus are never striped, and hereby have it suggested to us how it may be that
so many of our domestic breeds never have their young striped—none, in fact, except the
Westphalian and the Turkish. Rittimeyer (J. c. p. 187) has suggested that the so-called
“« Berkshire breed” of domestic pigs may have its parent stock in Sus celebensis, as
figured by Schlegel and S. Miller, which resembles it in the great transverse develop-
ment of its skull, the verticality of its forehead, of its occiput, and its temporal fosse,
in the height of its zygomatic arch, and specially in the remarkable height and massive-
ness of its lower jaw. There is no doubt that Sus celebensis is very closely allied to
Sus verrucosus, having not only the same general facies but the same peculiarities,
though on a smaller scale, as those whence the Java pig has taken its name. The chief
point of difference which Riitimeyer points out between the figured Sus celebensis and the
Berkshire breed depends upon the artist, 1 apprehend—being, as it is, a greater length of
the molar series, which, however, consists of 8 teeth in the figure of Sus celebensis (Ver-
handelingen, pl. 28 dis, fig. 3); and it is possible enough that the young of Sus celebensis
(the contrary not having been definitely recorded) may be like the young of Sus verru-
cosus in not being striped, and that they may thus have resembled the young of the
Irish Greyhound Pig, as when old they actually do resemble it in the development
of the mandibular warts.

As regards the skulls of Sws-verrucosus, figured tab. 32 by Miller and Schlegel, the
lacrymal in fig. 4 appears to have the favourable relation of length to height which is
characteristic usually of the other Suide with facial warts; whilst in fig. 1, which was
taken from a very old individual, the height and length would appear to be subequal, as in
the skull (fig. 6) of Sus vittatus, on the same plate. I should have set less store by this
variation in the two drawings than I do, if Nathusius had not distinctly recorded (p. 179
l. c.) that this latter is the relation in the Sws-verrucosus skull in his possession. On the
other hand, all the undoubted Sws-verrucosus skulls which I have been able to measure
have presented the elongated form of this bone; and the closely allied species Sws cele-
bensis, as figured in Miiller and Schlegel’s fig. 8, tab. 28 b¢s, may or may not have the
long lacrymal usual in that pig. But, on the whole, I should certainly accept the position
that the elongation of the lacrymal was not constant in Sus verrucosus, though it has
been so in the skulls [ have seen; and with this position I should hesitate to assign to
this peculiarity all the value which Nathusius’s insistence might lead one to assign to it.

Taking, in the second place, this variability in the proportions of the lacrymal bone,
and coupling with it, first, the absence of striping in the young, and secondly the
presence of mandibular warts, such as one of our domestic breeds, viz. the Galway
Greyhound Pig, still retains, or did till quite lately retain, we may be tempted to think
that some form allied to Sus verrucosus or Sus celebensis may have been the single
parent stock of all our domestic breeds, except, of course, such as the Westphalian and
Turkish, which are striped in their youth, and would be referred to the indigenous Sus
scrofa, var. ferus, as their parent stock. In favour of sucha view, we have, of course, the
general principle “ Entia non sunt multiplicanda preeter necessitatem.”

But the wide range* of Sus vitiatus, over Java, side by side with Sus verrucosus,

* Tt is interesting to compare with these statements, as to the geographical distribution of the Asiatic Suide, the

SECOND SERIES.—ZOOLOGY, VOL. I. 2N

268 PROF. ROLLESTON ON THE DOMESTIC PIG

over Borneo with Sus barbatus, and, as stated, also over Amboyna, Macassar, Banka, and
Sumatra, gives it special claims to attention. This Pig appears to me to be very closely
allied indeed to Sus cristatus, and to be similarly and readily distinguishable from our
European Wild Boar. In this latter point I differ from Riitimeyer (J. c. p. 187), whilst I
should agree with him in considering it all but identical with Sus leucomystax. The
claims of Sus vittatus and Sus lewcomystax, and Sus taivanus of Formosa (occurring, as
they do, in an area comprising Japan as well as Java), to have given origin to Sus
indicus, the domestic Chinese pig, in days sufficiently far off to have allowed the tendency
to striping of the young to become eliminated, are very strong, and can scarcely be
considered antagonistic.

Closely allied as Sus cristatus is to these races, its severe struggle for existence entailed
by its habitat on a continental area tenanted by the tiger (Felis tigris), as well as other
Carnivora unfriendly to Suidee, appears to me to have specialized it, especially as regards
its dental armature and the bones which carry it, into divergence from the probable line
of parentage of the inconveniently so-called Sus indicus. Sus andamanensis, on the
other hand, and, I am inclined to think, one or two other of the Asiatic Swide, show, from
a precisely opposite cause, that of restriction to a very confined area, the same divergence
from what I have imagined the present stock of the Chinese breed may have been.
But against any such speculations about what we do not see in the darkness of past
ages we have to set what we can see by travel in the broad daylight of the present, viz.
that almost all Swid@ are readily domesticable by savages in almost every quarter of the
globe; and what savages do now they may very well have done formerly.

I have not been able to institute any satisfactory examination into the relations of the
Athiopic tothe Asiatic Suide; and I should welcome an opportunity of examining skulls
and skins of the true Sws seen by Dr. Murie and Dr. Barth in Central Africa. It would
be additionally satisfactory if investigations could be set on foot as to the existence or
non-existence in this Sws of the Cystic form of Tenia soliwm, which certainly exists in
Sus cristatus. Dr. Cobbold informs me that Tenia mediocannellata is the common tape-
worm of Indian as of other patients ; but I apprehend that, as it has been so very definitely
laid down by others + that the Pig is at least one source whence inhabitants and sojourners
in India become infested with tapeworm, it would be premature to conclude the reverse
even from his authoritative statements. It must be very difficult to prove a negative here.

The facts of most direct importance, however, in investigations as to the relationships

following words from M. Gabriel de Mortillet’s memoir in the ‘ Revue d’Anthropologie,’ iv. 4, 1875, p. 653, as to
the origin of bronze:—* Reste le groupe de V’extréme Orient Asiatique. C’est la évidemment ou il faut chercher
Yorigine du bronze. Les principaux gisements sont dans la presqu’ile de Malacca et surtout dans Vile de Banca,
mais ils s’étendent dans d’autres iles de la Sonde et remontent jusque dans l’empire Birman ou I’étain est encore
exploité actuellement dans le district de Merguy. Ce minéral, dans tous ces gisements se recueille de la maniére la
plus simple et la plus facile dans les alluvions. Ce sont bien certainement les alluyions les plus riches du monde en
étain et celles qui occupent la plus grande étendue. Il est donc tout naturel que ce soit celles qui les premiéres
aient attiré Vattention de Vhomme. Le cuivre se rencontre dans les mémes régions. Tout le monde connait les
gisements de cuivre des iles de la Sonde, Timor, Macassar, Borneo. Ja Birmanie anglaise présente des mines de
cuivre 4 cdté des ses exploitations d’étain. Le pays se trouve donc dans les meilleures conditions pour avoir yu
naitre Vindustrie dw bronze.” Gallus bankiva, the parent stock of our Common Fowl, is found over the same area.
tT Notably by Dr. Charles A. Gordon, Medical Times and Gazette, May 2, 1857, p.429; and by Dr. T. R. Lewis,
Appendix B. to Highth Annual Report of the Sanitary Commissioner with the Government of India, 1871.

OF PREHISTORIC TIMES IN BRITAIN. 269

of Prehistoric to modern races are those of the great variations observed—first, in the
entire size of the individual animals, and, secondly, in the proportions of particular bones,
and notably of the lacrymal bones, in specimens from the same species or subspecies of
Suide. In view on the one side of this twofold variability which is explicable upon
acknowledged physiological principles, and affects the wild races both of Europe and
Asia, and, on the other, of the aptness for domestication possessed by Sus scrofa, var.
ferus, it appears to me to be unsafe to postulate for Prehistoric British Swine any other
parent stock than the one just named. On the other hand, such is the diffusibility and
transportability of Saws that it is not impossible, nor inconceivable, that the domestic
Buropean Pig, even of the Stone Age, may have had an Asiatic or African origin. As
regards the Bronze Age, indeed, if its tin and copper did really come from the East, such
a view cannot be said to be even improbable. But the acceptance of it does not seem to

me to be necessitated by the facts.

Weasurements of the Orbital and Malar Borders of the Lacrymal Bone*.
I. Sus eristatus and its allies.

Skull of Indian Wild Boar, Sus cristatus, no. 72. Sir Walter Elliot’s collection. Killed about 1830,
at Haugul, S.W. corner of Dharwar province, in South Mahratta country.

Height of lacrymal 1in.; malar border 1 in.

Skull of Indian Wild Boar, Sus cristatus, no. 46. Sir Walter Elliot’s collection, shot at Dendelly
12th April, 1831, in the great forest tract between Dharwar and Goa.

Height of lacrymal 1in.; malar border 1 in.

Skull of large Indian Boar, Sus cristatus, killed near Rajkote, in Kattywar, June 4, 1832. He was
with a large “sounder,” and ripped two horses severely. No. 71 in Sir Walter Elhot’s collection.

Height of lacrymal 1-0 in.; malar border 0°8 in.

Skull of Indian Wild Boar, Sus cristatus, no. 428 in Sir Walter Elliot’s collection. From Jaggia-
pettah, on the east side of the Madras Presidency, in the Masulipatam district, on the high road from
Masulipatam to Hyderabad. An old male with some obliteration of sutures.

Height of lacrymal 0°95 in.; malar border 0°95 in.

* To this paper I append a number of measurements, taken from various Sude, of the length and height of the
lacrymal bone, this point haying had great weight laid upon it by Nathusius, and holding good within very con-
siderable limits. I have not given any measurements relating to the widening of the palate (a second point insisted
upon by Nathusius), because, as regards the wild races, I have found that this widening of the interpremolar, as
opposed to the intermolar transyerse diameter of the palate, is sometimes found in specimens which undoubtedly
belong to wild European boars, whilst, on the other hand, the two measurements are usually subequal in the other
wild races, such as Sus cristatus, which I have measured. In taking the length of the lacrymal along its malar
border, some little ambiguity is caused by the fact that in many specimens the lip of the orbit is a little everted, so
as to resemble to some extent the spout of a mortar rather than the rim of a cup at the line of j unction of the malar
with the orbital border of the lacrymal. Where the differences between the ‘heights and lengths are measured by
tenths or twentieths of an inch or by millimetres, this structural arrangement may make the measurements vary
importantly. There is, however, always a line separating the part of the lacrymal which is to be considered as
belonging to the inner aspect of the orbit from that which is strictly facial ; and from this line the measurements have
been taken. The frontal border of the lacrymal, again, often bends downwards just before reaching its orbital edge,
just posteriorly to the plane of the lacrymal canals, thereby curtailing the height of the bone for the distance corre-
sponding with this deflection. My measurements have been taken in the plane occupied by the lacrymal canals, so

as to ayoid this source of fallacy.
(>)
2N 2

270 PROF. ROLLESTON ON THE DOMESTIC PIG

Skull of young female, Sus cristatus. British Museum, 716 m. “Terai, Nepal; Dr. Oldham.”

Height of lacrymal 1-1 in. ; malar border 0°75 in.

Skull of adult female, Sus cristatus,716n. British Museum. “India; Professor Oldham.”

Height of lacrymal 0:1 in.; malar border 0:7 in.

Skull of young female, Sus cristatus, about 9 months old. British Museum, 716g. “ Nepal; B. H.
Hodgson.”

Height of lacrymal 0:9 in.; malar border 0°75 in.

Indian pig, Sus cristatus, British Museum, 716g, under 5 months of age, very convex frontals, slightly
tumid lacrymo-frontal ridges.

Height of lacrymals 0°5 in. ; malar border 0:25 in.

Indian pig, Sus cristatus, British Museum, 716 v. “Nearly adult female, Nilgherris.”

Height of lacrymal 0-95 in. ; malar border 0°6 in.

Young male of Sus indicus; im collection of British Museum, not catalogued; with 3rd molar just
coming into use—that is to say, about 18 months old.

Height of lacrymal 0:9 in.; malar border 0°9 in.

Skull of Indian Wild Hog, Sus cristatus, no. 3251 a, Royal College of Surgeons of England. ‘This is
the skull of an adult male probably, the complexity of the posterior lobe of the 3rd molars being very
great. The naso-frontal suture resembles that of Sus scrofa rather than that of Sus cristatus. ‘The
lacrymo-frontal ridge is very largely developed.

Height of lacrymal 0°] in. ; malar border 1:25 in.

Skull of Indian Wild Hog, Sus cristatus, no. 3251 6, R. C. 8S. England.

Skull of Indian Wild Hog, Sus cristatus, no. 3251 c. R, C. 8. England.

Height of lacrymal 0:1 in., malar border 0:95 in., in both.

These two skulls were probably from the same district as no. 3251 a; but they differ from it, and agree
with all the other specimens of Sus cristatus,in having the height and the malar length of the lacrymal
subequal.

Skull of Wild Sow from Andamaa Islands, Sus andamanensis, no. 1514.0, Oxford University Museum.

Height of lacrymal 17 mm.; malar border 8 mm.: adult.

Skull of Sus vittatus from Amboyna. Alfred Russel Wallace, Esq. British Museum, “1362 ec.”
An adult male with a nearly straight naso-frontal suture, and a convex fronto-parietal region.

Height of lacrymal 0°55 in.; malar border 0°8 in.

Sus vittatus. British Museum, 1362 f. “Skull of animal developing the hinder molar.” The
lacrymo-frontal ridges are convex, the frontal region is flat, the naso-frontal suture, however, is straight,
as in S. cristatus.

Height of lacrymal 1:05 in.; malar border 0°6 im.

II. Sus scrofa, var. ferus, and its allies.

“« Sus libycus.” British Museum, 713 a. ‘Hab. Asia Minor, Xanthus (Sir Charles Fellowes). ‘The
skull is very distinct from all the skulls of the Wild Boars from Germany in the British Museum.” The
animal was just about 18 months old: its interectorbital region is still convex; but it has no very marked
lacrymo-frontal ridge; its naso-frontal suture is convex backwards, as in Sus scrofa, var. ferus ; its nasals
are 8 in. long. as against 142 of total skull-length; and the height of the lacrymal is less than its
inferior length. Here it resembles Sus scrofa rather than Sus cristatus.

Height of lacrymal 1in.; malar border 0:15 in,

OF PREHISTORIC TIMES IN BRITAIN. 271

Wild Boar from Germany, Sus scrofa, var. ferus, 3253a: Royal Coll. Surg. England, adult.

Height of lacrymal 0:9 in.; malar border 1:5 in.

Wild Boar from Germany, Sus scrofa, var. ferus. Oxford University Museum, 1513 a. All teeth in
place.

Height of lacrymal 1 in.; malar border 1:5 in.

Wild Boar from Germany, Sus scrofa, var. ferus. Oxford University Museum, 1518. Presented by
Prof. Max Miiller ; with all teeth, except 3rd molar, in use; 16 months old.
Height of lacrymal 0°75 in.; malar border 1:05 in.

Young Wild Boar, Sus scrofa, var. ferus. Royal College of Surgeons of England. With first molar
only through, therefore between five and six months old, with frontals still very convex in vertical aspect,
but with flat lacrymo-frontal ridge.

Height of lacrymal 0°7 in. ; malar border 0-7 in.; frontal border 1-5 in.

Skull of old Wild Sow, Sus scrofa, var. ferus, from alluvium near Oxford. Figured Plate XLI. fig. 2,
from the Geological Collection, Oxford University Museum.

Height of lacrymal 17 millims.; malar border 18 millims.

Skull, from peat, of Wild Sow, Sus scrofa, var ferus. British Museum. With 3rd molar through.

Height of lacrymal 0°8 in.; malar border 1-05 in.

Skull from silt of old bed of Lea river, shown to me in British-Museum Geological Collection by
Mr. Davis. Under 18 months.

Height of lacrymal 0-6in.; malar border 0°5 in.
* Second skull from same locality, and in British Museum, with all the teeth in place, and the third
molar with considerable complexity in its posterior lobe.

Height of lacrymal 0:6 in.; malar border 0°8 in.

Skull of Sus scrofa, var. ferus? British Museum, 7132. Skull of a young animal from the Zoological
Society’s Museum. The animal was about 16 months old, all the teeth being in place except the
third molars above and below. ‘There is still some convexity across the frontals, though flattening is
beginning on either side towards the ectorbital processes. The lacrmyo-frontal ridge is not markedly
convex; the nasals are convex backwards in the middle line.

Height of lacrymal 0-9 in.; malar border 0°75 in.

Skull of Sus scrofa, var. ferus? British Museum, 713%. Skull of young animal about 6 months old,
from the Zoological Ssciety’s Museum. This skull was entered in the Catalogue of the Zoological Society’s
Museum under the label A. 763; but no locality was recorded under this heading. The frontals are
convex from the level of the supraorbital foramina backwards ; the fronto-nasal suture is straight, nearly
as it is represented in the young German Wild Boar (Nathusius, /. c. Taf. iti. 12). Lacrymo-frontal
ridge is slightly convex.

Height of lacrymal 0°75 in.; malar border 0°5 in.

IIL. Sus verrucosus, var. ceranica.

Brit. Mus. 712d. ‘Skull, adult. A Wild Boar from Ceram. Collected by Mr. Wallace.” Of this
skull [have noted that its third molar is comparatively simple, whereas the other specimens of Sus ver-
rucosus in the British Museum have it complex. All the other specimens of Sus verrucosus catalogued
p. 330, Catalogue 1869, as Sus verrucosus, with the exception of no. 1862 d, which was received from
Mr. Wallace as Sus vittatus, have been noted by me as having the lacrymal long in relation to its height,
and as having complex molars. As, however, Nathusins, /. c. p. 179, has noted the lacrymal to be short in
a specimen obtained by him from Java, and as, of two skulls figured by Solomon Miiller (Verhandel.
tab. 32. figs. 2 and 4), one has the lacrymal short and the other long, it is clear that this species varies

272 PROF. ROLLESTON ON THE DOMESTIC PIG

in this matter. The closely allied species Sus celebensis cannot be said quite certainly, from Miller’s
pl. 28 dis, fig. 3, to have, or not to have, the long lacrymal usually found in the pigs with facial warts.
Height of lacrymal 1-0 in.; malar border 0-75 in.

IV. Sus barbatus, BoRNEO.

Skull of Sus barbatus. 1519 d, Oxford University Museum. Figured Plate XLIII. fig, 7.
Height of lacrymal 1:2 in.; malar border 1-2 in.

Skull of Sus barbatus. Procured in Borneo by the Rey. C. Spencer Bubb.
Height of lacrymal 1:2 in.; malar border 1:2.

Skull of Sus barbatus. Brit. Mus. 712a; called Euhys barbatus, Hand-list, p.58, 1873.
Height of lacrymal 1:15 in. ; malar border 1 in.

In Sus barbatus the lacrymo-frontal ridge is not very prominent, though recognizable ; in the shortness
of the lacrymal, which is 4 constant character in this species, so far as recorded in 8. Miller’s figures
and elsewhere, and in the characters of the naso-frontal suture, it resembles Sus cristatus and its allies
rather than Sus scrofa.

BIBLIOGRAPHY.

Baker and Durand, “Subhimalayan Fossil Remains,” Journal As. Soc.} Bengal, vol. v. 1836, p. 664.

Bartlett, Proc. Zool. Soc. 1861, p. 264.

Blainville, ‘Ostéographie,’ Sus.

Blasius, ‘ Naturgeschichte der Saugethiere Deutschlands, 1857.

Bochart, ‘ Hierozoicon,’ 1663, i. 696, 978.

Brandt und Ratzeburg, ‘ Medizin. Zoologie,’ 1829, pp. 80-150.

Buffon, ‘ Hist. Nat.’ tom. v. pp. 181-137, pl. xxiv. fig. 2, pl. xv.

Campbell, Colonel Walter, ‘ My Indian Journal,’ 1864, p. 325.

Crawford, J., Trans. Ethn. Soc. i. p. 439, 1863.

Cuvier, F'. et Geoffroy, ‘ Hist. des Mammiféres.’

Cuvier, G., “Oss. Foss.’ 11. p. 119.

Daubenton, Buffon, /. c. p. 189.

Erdl, ‘ Tafeln der vergleich. Anatomie des Schiadels,’ 1841.

Hyton, Proc. Zool. Soc. 1837. :

Fitzinger, Sitzungsberichte d. Akad. Wiss. Wien, Bd. 29, 1858, p. 861; 50, 1864, p. 383.

Forrest, ‘ Voyage to New Guinea,’ 2nd ed. 1780, p. 97.

Galton, Francis, Ethnological Society’s Trans. i. p. 127, 1865.

Giebel, C. G. ‘ Siugethiere,’ p. 225, 1859.

Gordon, C. A., Esq., M.D., Medical Times and Gazette, May 2, 1857, p. 429.
' Gray, Proc. Zool. Soc. 1852, p. 180, 1868, Jan. 9; Brit. Mus. Cat. Mammals, 1869, p. 325; Hand-
list, 1873, p. 57.

Hodgson, B., Journal Asiatic Society Bengal, x. p. 911

Jerdon, ‘Mammals of India,’ 1874, p. 241.

Link, ‘ Urwelt,’ i. p. 192, 1821.

Linneus, ‘Systema Nature,’ ed. xin. p. 217.

Low, ‘ Domesticated Animals of the British Islands.’

Miller, Solomon, ‘ Verhandlingen,’ taf. 28-32.

Nathusius, ‘Die Racen des Schweines,’ 1860; ‘Die Schweineschidel,’ 1864.

Pallas, ‘ Zoographia Rosso-Asiatica,’ p. 267, 1831.

Parker, W. K., Phil. Trans. 1874, p. 324.

OF PREHISTORIC TIMES IN BRITAIN. 273

Pictet, ‘Origines Indo-Européennes,’ 1859, p. 369.

Radde, ‘ Reisen im Sitiden yon Ost-Sibirien,’ i. p. 236.
Richardson, ‘ Domestic Pigs,’ London, Warne.

Riitimeyer, ‘ Fauna der Pfahlbauten,’ 1861.

Schreber, J. C. D. von, ‘ Die Saugethiere,’ 1835, Taf. 320 &e.
Sclater, Proc. Zool. Soc. 1861, p. 63; 1863, p. 122.

Sidney, S., ‘The Pig,’ London, Routledge.

Spix, ‘ Cephalogenesis.’

Swinhoe, Proe. Zool. Soc. 1862, p. 361; 1864, p. 382; 1870, p. 369.
Sykes, Proce. Zool. Soc. 1831, p. 104...

Temminck, ‘ Fauna Japonica,’ Taf. 20.

Wagener, A., Gétting. Gel. Anzeigen, Ix. p. 535, 1839.
Youatt, ‘The Pig, 1847.

Zimmermann, Geogr. ‘ Geschichte,’ i. p. 189, 1778.

Postscriet, March 22, 1877.

Srnce the publication of an abstract of the foregoing paper in ‘Nature’ for July 20.
1876, I have been enabled to compare the undermentioned skulls and figures of Swide,
and to consult the undermentioned volumes and memoirs which had not previously been
accessible to me. I have also been favoured with valuable information by the letters
of a number of scientific correspondents named below. The information thus gained, in
addition to that upon which my paper was based, has been in the main confirmatory of
the views I had come to independently of it; the very important fact, however, that the
young of Sus celebensis are striped, as communicated to me by Dr. A. B. Meyer, of the
Royal Zoological Museum at Dresden, is directly contradictory of the suggestion which
TI threw out at p. 267, as is also the fact published by him as to the striping of the young
of the nearly allied §. verrucosus directly contradictory of the statement of Miller and
Schlegel given above, p. 266.

List of Sources of Information on Suidee made available to me since July 1876.

I. A skull of S. celebensis °, kindly lent me by Dr. A. B. Meyer, Director of the
Royal Zoological Museum at Dresden.

II. A skull of a Sus, sp.?, 2, from Ternate ; also lent me by Dr. A. B. Meyer.

III. Two skulls of the Domestic Pig of Bengal (S.cristatus, var. domesticus), pre-
sented to the Oxford University Museum by E. Lockwood, Esq., Monghyr,
Bengal.

IV. Two skulls of 8. scrofa, var. ferus, presented by Dr. Fiedler, of Dresden.

V. ‘A skull of S. scrofa, var. ferus, presented by M. le Marquis de l’Aigle, Chateau
de Compiégne, Pise, France.

VI. Askull of §. scrofa, var. domesticus, from Stene i Bé, Lofoten Islands, 2°above Arctic
Circle; presented by H. N. Moseley, Esq., Fellow of Exeter College, Oxford.

VII. Two bronze statuettes of S. scrofa, var. ferus, from the Gallo-Roman period in

' France; presented by John Evans, Hsq., F.RB.S.

274. PROF. ROLLESTON ON THE DOMESTIC PIG

VIII. A Memoir by Prof. L. Riitimeyer, “Neue Beitrage zur Kenntniss des Torfschweins,”
in the ‘ Verhandlungen der naturforschenden Gesellschaft in Basel,’ iv. 1, 1864,
pp- 182-186.
IX. The Inaugural Dissertation “ Zur Kenntniss des Torfschweins,” by Johann Wil-
helm Schutz, Berlin, May 4, 1868, referred to by Mr. Darwin in ‘ Animals and
Plants under Domestication,’ vol. i. p. 71, ed. 2nd, 1875.
X. A memoir by Prof. R. Hartmann on the ‘“ Verbreitung der in nordéstlichen Afrika
wild lebenden Saugethiere,” in the ‘ Zeitschrift fir Hrdkunde zur Berlin,’ Bd. iii.
1868, in which (at pp. 349-352) there is an account of §. sennaariensis, which
the Professor and Dr. Schutz consider to be the parent stock of S. scrofa,
var. palustris.
XI. Letter from Dr. A. B. Meyer, of the Royal Zoological Museum, Dresden.
XII. Letter from Professor R. Hartmann, of Berlin.
XIII. Letter from Professor Fitzinger, of Vienna.
XIV. Letter and drawing from Professor Busk, of London.
XV. Letter from the Rey. C. Spencer Bubb, late Missionary in Borneo.
XVI. Letter from G. Dobson, Esq., B.A., M.B., F.LS.

A comparison of the skull of the old male of S. celebensis lent me by Dr. A. B. Meyer,
and the measurements of it given below, with the skulls of S. andamanensis and with the
descriptions of the skull of S. verrucosus given by Nathusius, Riitimeyer, and Miller and
Schlegel, will show that while agreeing with these skulls in general contour and propor-
tions it is intermediate between them in actual size. The highest point in the sagittal
contour is, asin the Swide above named, a little way anterior to the upper edge of the
occipital squama ; the transverse arc from one orbital process of the frontal to the other is
even more convex than in any other Asiatic Pig; the zygomatic arch is short, deep, and
roughly sculptured; the lower canine’s alveolus reaches back to a prominent external pro-
tuberance in the plane of the anterior lobe of the third molar, a point further backwards
than it usually attains to. The posterior lobes of the third molars, though consisting
of 4-5 cusps, have not the great antero-posterior length relatively to the rest of the tooth
which we find in specimens of the same sex of S. cristatus.

As in S. verrucosus, the bony palate is nearer to the basicranial bones than in most
other Swide.

_ Its measurements (being those adopted by Professor Riutimeyer, ‘ Verhandlungen
Gesellschaft Basel,’ 1864, p. 163, from Nathusius, ‘ Abbildungen von Schweineschadeln,’
1864, p. 6) are as follows :—

Measurement of Skull of Sus celebensis.
millim. inches.

Length from foramen magnum to tip of intermavxillaries ......... 281 = 103
Length of occipital crest to tip of intermaxillaries ............... 325 = 12.8
Length from occipital crest to tip of masals ....................6.-- 320 = 12°55
Meneth cof masal sMiiiancs. sues snaps seceiseonoy os se aRReRe eee elise is ih == Gl
Anterior part of frontal and sagittal sutures ..............000:000e i = Be

OF PREHISTORIC TIMES IN BRITAIN. 275

millim. inches.
Posterior part of frontal and sagittal suture .....................05 y= Bs
From occipital foramen to VOMET ..............-.ceseeeenec eee ee eee Ae — nel,
Brom occipital foramen to palate 272.20... -. 2... eee erence we 2 iO
Dentizerousypart Or maradllaeseseeseeeesese ce eseee sacar tee sea 20 — ee,
THRONES FOIE CHF JOBINS ons egonsecnsoopsossooousbecsacss0ass090R0EFogoRR: Cl ee:
Greatest interzygomatic width. ..................-ss20ceeeeerec gene eee 1440 = 55
Greatest trondalllavaditeeeaee peeves sata sses eee stsctls sates -ctace er 3 = Bh
Shanalligaty texerell WIGAN, 5. asosnosoesqcsboboosenansoeaneesonaussHeceopenseces ol = ie
Intermaxillary width ................... SNe SESERS OSA ODER EOROROEDCAATNG 37 = 145
Ibigast iaeisalll wally 645 45d nodesasdsesoosanaseeteseudes souesogecesenouse so8Ds Ales
Itatkemn@lere Tmalidiy 1. sous asongoHansdesbaeseoseoossuscusovasseucdscosbosens 43 = 17
InntieryomermnGllaie WrGHHd, 225000 secacogonosacosodacocsooncasnoapanesendn 800 an ss INS
Qreqijoriivall NEVE, ascoocroradepsonsesoosuedscosccksbnvasbosscergnnagoeRBODS 1 ee
millim 6 3
Lacrymal orbital border ............ 22 = -9 compare 1 in Sus verrucosus, Riitim.
Lacrymal malar border............... aa Pe rh ee lalicss
Lacrymal upper border ............ 31 = 14 » 184
millim te
Length of S. celebensis, sec. Dr. A: B. Meyer............... 1160 = 45°67
is. ee Miller & Schlegel ............ 950 = 37-40
Length of tail of S. celebensis, sec. Dr. A. B. Meyer...... 250 = 984
5 3 Mill. & Schleg........ 150 = 59
Height of S. celebensis, sec. Dr. A. B. Meyer ............ 550-570 = 21-22
33 53 Miller & Schlegel ......... 560 =

Dr. A. B. Meyer’s specimen was more brown than Miiller’s and Schlegel’s figure,
pl. 280is; the tuft at the angle of the mouth red-brown, and the hair on the nape
longer. As to his measurements of S. celebensis, Dr. A. B. Meyer observes that they are
taken from a full-grown animal, and, being larger than those of Miller and Schlegel,
bring S. celebensis more nearly on to a level with S. verrucosus. Its smaller size suggests
the dwarfing action of a confined insular habitat undergone in some of the upheavals or
subsidences of Celebes.

Dr. A. B. Meyer informs me that the young of S. celebensis are striped, in contradic-
tion to the suggestion which I (see p. 267, supra), being impressed with the closeness
of the affinity between this Sws and S. verrucosus (which Miiller and Schlegel had de-
elared not to be striped when young), had thrown out. He informs me also that he
has the authority of General von Schierbrant, who has lived thirty years in Java and is
a first-rate sportsman, for saying that Miiller and Schlegel (/.c. p. 177) are not correct in
what they say is the case with S. verrucosus. If this be so, the claim which S. verrucosus
would have had to be considered the parent stock of our improved breed of pigs falls to
the ground. On the other hand, I cannot, knowing the great modificatory power which
domesticating influences of one kind or other have been proved to possess over the highly
plastic porcine organism, and bearing in mind the similarity between the Irish Grey-
hound, Pig, and §. verrucosus, agree with Professor Riitimeyer (p. 184, 7. c. 1864) im ex-
cluding this Svs from consideration when we are speculating as to the parentage of
our domestic Pigs.

SECOND SERIES.—ZOOLOGY, VOL. I. 20

276 PROF. ROLLESTON ON THE DOMESTIC PIG

The skull of the Sws from Ternate lent me by Dr. A. B. Meyer belonged to a young
male, the third molars of which were just coming into place, and which may be supposed
therefore, according to the analogy of S. scrofa, var. domesticus, if that may be taken as
any guide, to have been about eighteen months old (see Nathusius,/.c. p. 21). It differs
from skulls of S. cristatus in the flatness of its lacrymo-frontal ridge, in the convexity
backwards of its naso-frontal suture, and in being somewhat smaller than male skulls of
that variety of Sus usually are. The posterior lobes of its third molars are not quite so
large as they usually are in S. eristatus ; but to this point I am not inclined to assign as-
much importance as some other writers do (see Riitimeyer, ‘Fauna der Pfahlbauten,’ pp. 33
and 188, and, per contra, Nathusius, J. c.pp.49 and 103). In the straightness of its vertical
contour it contrasts with WS. celebensis, S. andamanensis, S. vittatus, as figured by Muller
and Schlegel, /.c. plate 32. fig. 6, and with S. barbatus as figured here, Plate XLIII.
fig. 7; and resembles S. cristatws, as it does also in its general facies. Dr. A. B. Meyer
thinks that this Saws may possibly deserve to be considered a new species, S. ternatensis ;
but having had abundant experience of the facility with which the wild pigs of that
region cross considerable arms of the sea, he would, I gather, express himself with much
caution as to its relations to S. papwensis and the -Suidze of the neighbouring islands, vol-
canic and other, tenanted by swine. ‘The colouring,. however, of the head of this Sus
differs from that of any other Sws seen by Dr. A. B. Meyer, or figured by Schlegel—the
head being covered all over with long black hair, except in the region occupied by a broad
yellowish brown streak beginning between the eyes and descending to the snout, where
it broadens. .

Measurements of skull of Sus ternatensis.
millims.’ inches,

Length from foramen magnum to tip of intermaxillaries ......... 290 = 114
occipital squama 5 Sail Mihalis 355 = 141
a n THD OL MASS s5o00scesdoonsoa00 B15 = ss
Inen’e tivo feniasalls eee eee Vo ats SEES AR Ee pet ee cman eons
Anterior part of line of frontal and sagittal sutures ............... 8 = 32
TROANSTAONE JORNAG Oil SEVENG INE) .oooocona0eag0sedadandaaoanangacaocagnecs0aed 9 = 37
Mio lant Merial. ates ae tgeduoiaetaroeoslen ene see semer eee berien das bepasepretaiaseek 1443 = 5:65
Incisive sbomevlenetinii. veer yes teenie reli = saa ei Ae Le emanate i —
Greatest interzysomatic width ...............csenseeeeeee eee scese snes 3a osoo
Greatesttaciall awaditlssenomencesmretece: var aa or yseee oer ceee teen cas % = 88
east sacial-wid thes ss. de shen sels naecni se eeee ea pm eae an EE ET 86 = 1:45
Juaneracnarailehar (Gli dhl eas dedusnaanusasdudandecadsosaauavscudaedsdauaaseus 40 = 16
Weds tanasalawadthy ex gccee saree ee a Acerca sani ean erat Rae eanyaa Oa —
Ibayrsrsaavolere Aili eas asda osadebe Me naneetronee aon ordradé stiadadsersasdse¢ A) == hy
Tbniterprem olla swa dit nes main a seh eres eee eee aa eee rep eee aoe 44. = 1°75
(Ojoroyamhiel levered ohh aneamae ancoouatadainansocendaseddesicare asians bate coecooees Wl = 4r4
Ibeereyamen!l Guelomitaall WORCIE 5. sovo0vesconosenoescononbasoecopcdno or vscoce 2 = i
Pe HIT DIATMOOTOSL lent ayes: es oncck oa REET Oa 8
55 LUPPCEMDOLUER crap wnacastere anise aaiaccen cate a Cece MPR Eee 34 = 13

A comparison of the two skulls of the domestic Pig of Monghyr, Bengal, which I owe to
the kindness of H. Lockwood, Hsq., and which, he informs me, are kept in large numbers

OF PREHISTORIC TIMES IN BRITAIN. 277

by the pariah castes of his district, féed on human ordure, and are sold out of the district
to the Chinamen at Calcutta, with the skulls of the wild Pig of Hindostan (S. eristatus)
shows that they differ from them in little else than their smaller size. They may be
spoken of, therefore, as S. cristatus, var. domesticus; and they are as distinguishable
from any variety of S. scrofa, var. domesticus, or indeed of S. indicus, as is the wild S.
eristatus from other wild Suide.

It is noteworthy that one of the skulls belonged toa very old sow. In these, as in the
days when Juvenal spoke (Sat. vi. 160), of another eastern country as a place where

“ Vetus indulget senibus clementia porcis,”
it is a rare thing for a domestic Pig to be allowed among western nations to live long
enough to wear down its third molars. Mr. H. N. Moseley, however, informs me that
domestic Pigs are kept in the Chinese Buddhist monasteries till they die of old age and
infirmities ; and of India, Meiners, in his ‘Allgemeine kritische Geschichte der Re-
ligionen,’ 1806, Bd. i. p. 193, says, ‘‘In Asien war von jeher Hindostan, wie in Afrika,
Heypten, der Thron des Thierdienstes.”’

It is noteworthy, secondly, that with the worn-down molars of this aged domesticated
Pig were correlated abscesses on both sides of the lower jaw, much as might have been
the case (see J. R. Mummery, Esq., Trans. Odont. Soe. ii. 2, 1869, p. 72) in ill-fed human
beings with similarly worn-down teeth.

Mr. Lockwood informs me that the young of the domestic Pigs of his district are
striped like the young of wild Pigs.

The skull of a wild boar, from France, presented by the Marquis de l’Aigle, shows that
the posterior lobe of the third molar in the lower jaw may attain the same proportions
in S. scrofa, var. ferus, that it does in S. cristatus.

The skull of §. scrofa, var. domesticus, procured for me by H. N. Moseley, Esq., from
the Lofoten Islands, two degrees within the Arctic circle, has the long, low, lacrymal
characteristic of S. scrofa, var. ferus, a fact of particular interest when coupled with the
information, also procured for me by him, to the effect that the young domestic pigs of
that region are occasionally born with stripes, and with his observation that the old pigs
have a very wild-boar-like appearance. These facts should be borne in mind as telling
against the views propounded by M. André Sanson, in his memoir “Sur la prétendue
transformation du Sanglier en Cochon domestique,” in the ‘ Journal de l’Anatomie et de
la Physiologie,’ tom. iv. 1867, p. 38.

In Professor Hartmann’s memoir above referred to, it is stated (p. 850) that certain
Negro tribes, who disobey Mahommedan precepts by eating as well as domesticating S.
sennaariensis, excuse themselves by saying it was formerly the custom to do so, a fact which
goes some little way to disprove the view that this true Sws can be merely a feral variety
of S. scrofa, imported by Huropeans. Professor Hartmann, in a letter of date Sept. 28,
1876, says, “ Sus sennaariensis ist ein kleines dem europaischen Torfschwein (der Pfahl-
bauten) ahnliches Schwein, echtes Sus, welches wild durch einen grossen Theil von
Mittel-Afrika vorzukommen scheint . . . Sus scrofa ferus in der Sahara und in Algypten
nicht selten, findet sich angeblich ebenfalls in Sennaar, indess weiss ich hieritber nichts

vollig Sicheres.”
202

278: PROF. ROLLESTON ON THE DOMESTIC PIG

Professor Busk, in a letter to me of Dec. 17, 1876, informs me that in the Etruscan
Museum at Florence, amongst numerous little bronze articles extracted from the ancient
Etruscan tombs, there were many figures of animals, one in particular being a very well
made statuette of a pig, which to his eye very closely resembles the Berkshire breed, the
only point in which it differed being the comparatively large eye, whilst the rest of the
contour was quite what we might expect to see at an ordinary cattle-show. The animal
was represented apparently as having a close curled tail. With it were a good many
statuettes of stags, the horns of which were of the type of the phocene Cervus ctenoceros.
In the very perfectly restored Htruscan tomb erected in the Museum, with all its original
contents and frescoes, Professor Busk noticed, amongst other figures of animals, one very
well drawn of a monkey climbing up a tree or pole.

I should suggest that the monkey and the pig, both alike, are representations of
animals from the same quarter of the globe as that whence the kinsmen of the Htrus-
caus in the time of Solomon brought, every three years, into Mediterranean regions, the
ivory, apes, and peacocks, the Sanskrit names of which still remain to speak to their
habitat. See Max Miller, ‘ Lectures on the Science of Language,’ 1861, ser.i. p. 190, and
ser. li. p. 234, for the source whence these animals and copper came to Europe.

Professor Busk sent me by the same post odontograms of the teeth of S. eristatus,
S. scrofa, var. ferus, and S. scrofa, var. domesticus, which show that in S. crisiatus the
antero-posterior length of the third molars is much greater in relation to that of the rest
of the molar series than it is in either of the other Suide named. I have above, p. 262,
noted that the sexual differences in this matter are very considerable in S. cristatus ;
and those produced by domestication are also not insignificant—points which somewhat
impair the value of these statements of relative proportions, though no less an authority
than Riitimeyer avers, ‘ Fauna der Pfahlbauten,’ p. 188, that “mit viel grésserer Zihigkett
das Gebiss den Species-Typus ausseren Linflussen gegeniiber aufrecht halt als die Schddel-
bildung’’ (cf. however, Nathusius, /. c. pp. 49, 103, and Studer, Mittheil. Ant. Gesell.
Zurich, xix. 3, 1876, p. 67). Professor Busk also observes that in S. crista/us nearly all
the teeth, except molar 8, are wider in both jaws in proportion to their length than they _
are in the other two, and that from this we may suppose that the Indian pig is more
exclusively herbivorous than the tame or wild animals with which he has compared it.
Jerdon speaks of S. cristatus, 1. c. p. 248, as being in general almost entirely “ vegetable
feeders.” Captain Baldwin’s views (‘The Game of India,’ pp. 154-5, 1876) are to the
same effect.

Professor Busk’s drawing and description of this Etruscan pig, coupled with a similar
figure of a pig found in the ruins of Herculaneum (see Nathusius, /. c. p. 142, fig. 38,
and Darwin, /. c. p. 71), furnishes a good illustration of Riitimeyer’s saying (Fauna der
Pfahlbauten, p. 190), that though the modern breed of domestic pigs is recent enough
to have been introduced by steamboats, it nevertheless had been represented con-
tinuously from former ages by the Biindtner-Schwein in the valleys of the Grisons.
A figure of a sow suckling three young ones may be seen on an Umbrian medal of
probably the third century 8.c. at latest, figured by M. Sambon, in his ‘ Monnaies
Antiques de l’'Italie,’ 1870, pl. v..5, with the short snout bent on the chanfrin, the

OF PREHISTORIC TIMES IN BRITAIN. 279

pendent ears, and the mane limited to the nuchal region, which justify us in considering
it to have been intended to represent a tame variety. A pig, figured ibidem, pl. iv. 4,
from Etruria has a snout of such slenderness as to correspond very closely with the
description given of §. scrofa, var. palustris, whilst it contrasts in other points very
strikingly with the wild boars represented from Apuleia and Lucania, on the other side
of Italy (¢didem, plates xiv. 1, xv. 17 and 37, xix. 4), which have as close a zoological as
legendary connexion with the wild boar of Calydon, on the other side of the Adriatic
(see ‘ Thesaurus,’ Brandenburg, i. pp. 318 & 464, 1696, and ‘Thesaurus Numismaticus,’
1. p. 400, and tab. xl.).

Of the two bronze statueties given me by Mr. John Evans, one has the long slender
snout, and the mane reaching the whole length of the convex back, from the prominent
ears to the curled tail, which may justify us in considering it as intended for a wild boar;
the other combines the sturdy straddles and the long and large erect mane, beginning on
the forehead and in front of the erect ears, characteristic of a wild boar, with a snout as
disproportionately short, and tusks as reduced as we ever see them in the highest-bred
modern Chinese pig. The characters of the wild and tame varieties, however, being thus
inaccurately and inartistically combined in these statuettes, causes them to contrast dis-
advantageously with the Italian works of art just mentioned; but they furnish us with
a conclusive answer to the weak reasoning of De Blainville (‘ Ostéographie,’ Sus, p. 170),
expressed in the following words :— Du temps de César, il parait cependant qu’elle (la
culture du cochon) n’était pas encore parvenue dans les Gaules, car il n’est nullement
question de cet animal dans ses Commentaires ; elle s’y est donc propagée depuis la con-
quéte, d’ou elle a passé en Angleterre, qui ne possédait pas méme de sanglier dans ses
foréts!” If further answer were required to this astounding statement, a reference to
Mr. Evans’s work on British Coinage would furnish it—figures of the boar, some of
which are exceedingly characteristic, being given there on pls. vi., vill., xi., Xil., and XiiL.,
from those ancient coins. But all well-informed antiquaries are aware that the wild
boar is one of the earliest animals figured in Celtic works of art (see ‘Hore Ferales,’
p- 185, pl. xiv.; Montellier, ‘Mémoires sur les Bronzes Antiques,’ Paris, 1865; and
Stephens, ‘ Literature of Kymry,’ p. 250).

Professor Riitimeyer’s paper in the ‘ Verhandlungen der naturforschenden Gesellschaft
in Basel,’ iv. 1, 1864, is, I apprehend, referred to by Mr. Darwin when he says (‘ Animals
and Plants under Domestication,’ 1. p. 71, 2nd ed. 1875) that “‘ Riitimeyer himself seems
now to feel some doubt” as to whether the “ 'Torfsechwein”’ existed as a wild animal during
the first of the stone period. No reference is given /.c. to any memoir of Professor Riiti-
meyer’s; and I became, after writing the foregoing paper, acquainted with the one in
question from a mention of it made by Herr Edmund Naumann in ‘ Archiv fir Anthropo-
logie,’ Bd. viii. 1, 1875, p. 19, in a discussion on “‘ Die Fauna der Pfahlbauten im Starnber-
ger See.” From a perusal of this paper of Professor Ritimeyer, I am inclined to think that
he would regard the skull which I have figured (Plate XLI. fig. 2) asa skull of S. serofa,
var. palustris. I have spoken of it as the skull of a wild sow, considering, as said above,
p- 259, that early breeding may, in a species admitting of such a wide range of structural
oscillation, and notably in the matter of mere size, account for a very great distance be-

280 PROF. ROLLESTON ON THE DOMESTIC PIG

tween its male and female representatives (see for similar view as regards our own species
‘Journal of Anthropological Institute,’ vol. ii. 1875, p. 122). Iam not inclined to with-
draw from this view even after reading Prof. Riitimeyer’s and Herr J.W. Schutz’s memoirs.
For in the former of these I find (p. 151) that the jive skulls used for description are
acknowledged to be skulls of sows, and four of them to have been skulls of o/d sows, and
the measurements given at p. 163 have been taken exclusively from skulls confessedly
female (see p. 161). And I learn from Herr Schutz’s essay (p. 44) that Steenstrup (cit,
Wiegmann’s Archiv, xxvii. n. 112) had distinctly stated that the “Torfschwein ” was (as
I had hinted without any knowledge of his views, see p. 254, supra) simply the female repre-
sentative of 8. serofa, var. ferus. Professor Riitimeyer had not at the time of writing his
‘Fauna der Pfahlbauten,’ 1861 (see p. 33), a single perfect skull of the “ Torfschwein”’
available to his comparison ; the almost perfect skulls treated of in his memoir of 1864 (see
p- 150) have caused him to modify the view put forward at p. 190 of the earlier work,
and in the later one he allows (pp. 158-160) that, both in the matter of the length of the
lacrymal and in the absence of widening of the palate anteriorly, the 8. scrofa, var.
palustris, was more nearly affined to 8. scrofa, var. ferus, than to S. indicus. To show this
was the reason for giving a great number of my measurements, supra, pp. 269-272. I take
this opportunity of saying that the very small breed of the Scottish highlands and islands,
with suberect ears, usually of a dusky brown colour, with an arched back and coarse
bristles along the neck and spine, spoken of by Professor Low in his ‘ Domesticated
Animals of the British Islands,’ p. 429, may perhaps be such a breed as Professor Riiti- -
meyer suggests, in his later paper (p. 168, see also p. 148), should be looked for in
Eastern Europe or Western Asia or in fossil forms*.

Mr. G. EH. Dobson, F.L.S., ina letter to me of date Jan. 15,1877, informs me that the
young of the Pig of the Andaman and of the Nicobar Islands are striped.

The Rev. C. Spencer Bubb informs me that the young Borneo domestic Pig is some-
times striped and sometimes not, whilst the young Chinese Pig is never striped; and he
adds that there are certainly two domestic breeds in Borneo.

* T suspect that latitude has more to do with the production of such varieties than longitude, and still operating
causes more than geological. One of these causes is suggested by the words of Varro, ii. 4. 13, cited by Dureau de la
Malle, ‘ Economie politique des Romains,’ ii. p. 149: “Porcique nati hieme, fiunt exiles propter frigora.”

OF PREHISTORIC TIMES IN BRITAIN. 281

DESCRIPTION OF THE PLATES.
Puate XLI.

Fig. 1. Side view of partially reconstructed skull of Sus scrofa, var. domesticus, wanting nasals and inter-

maxillaries, from late Celtic interment at Arras, Hast Riding of Yorkshire. Oxford Museum.
It is, as Nathusius has well pointed out, /. c. p. 147, by no means always easy to be absolutely
certain as to the question whether a particular pig’s skull belonged to a wild or to a domesticated
individual. The difficulty is increased when the animal is young, as in this case, the last molar
having only just come into use, and the animal consequently being only about 18 months old,
and when as the small size generally, and especially thesmall size of the third molar and the
canine, may be taken, I believe, to indicate that it is of the female sex. It is true that the
paucity of cusps in the molars has been taken as indicating the wild state; a comparison,
however, of the male and female molar series in Sus cristatus has suggested to me that the
greater size of the molars really depends upon a greater supply of blood, such as the male
molars would get, by virtue of sharing the greater supply lavished on the canines, and such as
a well-fed domesticated animal’s molars would get im common with all its other structures and
organs. The comparatively vertical occipital squama is one main anatomical point in favour
of this skull having belonged to a domesticated specimen; the pterygoid, on the other hand,
has much of the obliquity characteristic of the wild Sus scrofa, var. ferus. When we consider,
however, that this skull was found in an interment containing a human body, together with
portions of another skull of a pig of the same age, the probability that it belonged to a tame
individual appears to be very great. The subjoining measurements show that the lacrymal bone,
though not so long relatively to its height as is often the case in the non-domesticated Sus
scrofa, is yet longer than it is im Sus indicus, or in any of the Eastern pigs from which Sus
indicus can with any probability be supposed to have descended.
inches.

Length from anterior external angle (apex) of frontal to middle of occipital ridge 5:9

eneth from apex of frontal toyapex ofmaxilla) 592 9 5 6 5 5 4 ns 4
Length from anterior border of orbit to temporal ridge. . . ..... . 38
henethifromyanterior border of orbit to‘posterior 9. . 5). ae «oy
Greavestpwidubgotocemput “esc jo es o, ie || | eptctud b tbs cen neug iy acdsee
hen et hyor molar Series) viene ypaceesen be hah Gace. eat ice Brekeiiiecteh ehidoie
ers HG OCCiPUti rae Aare tes ae Ren Cee Brit ce Coc tea de 378
Evers nigomlaenyanale ia ener oe eee ee a OR ed eens ise OTT.
Tengthvor lacrymo=malar sninre ey nue Welland yi tly yi OI

The difference here noted between the height of the lacrymal and the length of its lower
border amounts but to a tenth of an inch ; but in the female of Sus indicus, with the same stage
of dentition, Pl. XLII. fig. 4, the height of the lacrymal, instead of being one tenth of an inch
less, is 0°35 inch greater in length than its lacrymo-malar border.

Fig. 2. Skull of Sus scrofa, var. ferus, old 2, from alluvium of Thames valley, obtained for the Oxford
University Museum from the cutting for the drainage works near Iffley, 1876, by Professor
Prestwich, F.R.S.

This skull combines the general contour and the slender snout of Sus scrofa, var. ferus, with
a lacrymal bone differing little in its proportions from the lacrymals so characteristic of the
Asiatic pigs, less Sus verrucosus (and celebensis). ‘The fronto-parictal region does not form one

282

PROF. ROLLESTON ON THE DOMESTIC PIG

continuous slope in the same plane as is the case in the Wild Boar of Germany (Pl. XLII. fig. 5);
but this difference may be observed in skulls certainly of Wild Boars from the Thames-valley
deposits ; whilst the great wear of the teeth (see p. 258, note * above) and the slenderness and
length of the naso-facial region are much in favour of considering this specimen to have
belonged to the wild race. I have placed side by side with the measurements of this skull the
measurements given by Riitimeyer, /. c. pp. 45 and 183, of his “ Torfschwein,” Sus scrofa, var.
palustris. Unhappily, Rtitimeyer has never been able to procure (see pp. 43 and 45, note 1) a
skull of this variety of Sus with the facial bones in connexion with the braim-case, nor has
Nathusius (see p. 149, 7. c.) ever been able to see an uninjured lacrymal bone from the same
animal. These facts, whilst making the value of this skull (the opportunity of figuring which ~
I owe to the kindness of Professor Prestwich) greater, make the value of the comparison of its
measurements less.

The instructive observations of Nathusius (J. c. pp. 99-101), to the effect that ill-nourished
pigs have the entire length of their skulls greater as measured from the occipital crest to the
apex of the snout, whilst the portion of that length made up by the frontal and parietal is
somewhat shorter, and the nasal portion proportionally longer, when coupled with the fact of
the great wear of the teeth in this specimen, enable us to explain the one great point of
inferiority, that of the length of the fronto-parictal region, which this skull’s measurements
show us, compared with those of the “Torfschwein.” It may be added, that the true explana-
tion of Dr. Gray’s statement (Brit. Mus. Catal. 1869, p. 329), that the nasal bones of the skull
elongate as Suid increase in age, ‘and especially as they reach adult and old age,” is probably
that he had im his mind’s eye skulls of old and ill-fed wild pigs, such as this specimen., On
the other hand, when we are comparing such skulls as this with the “Torfschwem” of
Riitimeyer, we must recollect that he represents this latter variety of pig, and what he supposes
to be its still surviving representative, the “ Biindtner-Schwein,” as having a short snout (see
pp. 42, 45, 181-185). The classificatory value, however, of such a peculiarity is much reduced
by the results of such experiments as those of Nathusius just referred to.

Measurements of Iffley Skull and of Riitineyer’s “ Torfschwein,” pp. 45, 183, 1. c.

Iffley Skull. Torfschwein.

inches. inches.

Greatest frontal width between ectorbital processes . . . . . 32 37
Lease WHO OR TOMER 6 s 6 5 6 6 6 2 8 2 ob wo op) OS 0-9
Greatest interzygomatic width . . . . . ot eevee, TAG 4 7—4e9
Height of occipital from inferior border of fame magnum . . 35 38-45
Length of vertex from level of supraorbital foramen to ridge of

OUSIDE 6 56 6 : : ORS An
Horizontal distance com anterior Bonde of oie to oueeenee of

temporal fossa. . . : ‘ is lade Se Me aol le ae eo 31
Length of intermaxillary aoe Resin hordes pr acorns anh ao coed 1:9-2°4.
Maximum length of skull from apex of intermaxillaries . . . . 10°9 —
Length of nasal hones (approximately). . . ..°. . =. . . O55 —
Length from apex of intermaxillaries to inferior border of foramen

magnum . . 5 ; - 103 —
Length from anterior een Pls ened 0 of fr intel to eae of

occipital ridge. . . . 5 : 5:9 =:

Length from middle of ponte suture to male of sicelinttl
TCE Fat ata me eas cull ict ates No (MES aictg io ae, QoaMi RU uN Oa mae od iS _

OF PREHISTORIC TIMES IN BRITAIN. 283

Tffey Skull. Torfschwein.

inches. inches.

Length from anterior border of orbit to posterior. . . . . . 4I4 —
Greatest width of occiput . . . + hire Ppeie —
Breadth of nasal at commencement of eeeteontal Peck which

is point of maximum width ofnasals . . . . ..... 4d _
Maximum width of intermaxillaries . ......2.. =. «18 —
Height of lacrymal along rim oforbit . .... . . . . 17 millims.
Length of lacrymal along malar suture. . . ... =... =. IJ8wmillims.

Fig. 3. Sus andamanensis. 15146, Oxford University Museum, adult 9.

The skull from which this drawing was taken was that of a Wild Sow from the Andaman
Islands, procured for me by my friend J. Wood-Mason, F.G.S., of the Calcutta Museum. It is
about the same size as the two prehistoric British skulls, figured figs. 1 and 2, and as the
“ Torfschwein ” of Riitimeyer—resembling this latter im the one important particular, that of
its long fronto-parietal region, in which it differs from the British skull (fig. 2). It differs
from all the pigs’ skulls here figured in the exaggeratedly disproportionate shortness of the
malar border of its lacrymal bone, which is, as in the newly born European pig, little more
than half the length of the orbital border. In its convex frontal region we have, again, a
character retained in the adult Asiatic which is transitorily represented in the European pig. This
convexity of the fronto-lacrymal ridge is well marked and, like the straight naso-frontal suture
and the two other peculiarities specified, serves to differentiate Sus andamanensis from specimens
of similar size from the Palearctic region, and to show that it is a member of the group repre-
sented by Sus cristatus. Again, this skull resembles Sus cristatus and differs from Sus scrofa
in bemg somewhat shorter, and having the paroccipitals, temporal bulle, and pterygoid processes
more crowded together. The length of the nasals is a little less than that of the fronto-
parietal region in the vertical aspect, as is often, though not always, the case in Sus cristatus,
though very rarely, if ever, in Sus scrofa.

Neasurements of Skull of Sus andamanensis, female.

inches

Greatest frontal width between ectorbital processes. . . . .... =.=. 33
Meas tawidthnoumvervex ssa st ro mater eee to er ee a sy) ee OS
Greatest interzygomatic width. . . . rach TUT cee sa eee mercer Ae OD,
Height of occiput from inferior border of adteh magnum .. . . 37
Length of vertex from level of supraorbital foramina to middle of eee Heth 4:5
Horizontal distance from anterior border of orbit to temporal ridge . . . . 3:05
Length of intermaxillary along alveolar border . . ....2.2.2.~. 20
Maximum length of skull from apex of itermaxillaries. . . . .... 10: 8
Length ofnasals . . . . é db SME RS
Length from apex of stir ee erticid to ee ae of anaes magnum . . 91
Length from apex of frontal to middle of occipital ridge . . . 6 Jo ra BS
Length from middle of fronto-nasal suture to middle of occipital eo sees om ORO)
Length from anterior border of orbit to posterior . ...:. .. =.=... «CLS
Greatest width of occiput . . . : : 2°5
Breadth of nasals at commencement of naso- ee sate aie is me peadeet

partofnasals. . . . BEB dy! CST heaters ochre maces end eet cocaine 60)
Maximum width of as tate) LEMON Pee Ree 2 ey eee et soe eee ve
Heizhtiotlacryndl-alonetrim! of orbit’. 9. 2): UR OS SOB 2? 17 enim

SECOND SERIES.—ZOOLOGY, VOL. i. 2P

284

Fig.

Fig.

PROF. ROLLESTON ON THE DOMESTIC PIG

Length of lacrymal along malar suture BR enn SNE gs Gling. Tease Neen peevadnli ns
[GRIN OE uOleGEE) 5 6 5 4 56 9 0 0 50 9 0 8 9 6 oe cs 4 6 ADCS,
Thensthvor premolar Series) ir itll ss i-inr a cE

Pratt XLII.

4, Skull of Wild Indian Sow, Sus cristatus, British Museum. See ‘Catalogue of Carnivorous,
Pachydermatous, and Edentate Mammalia,’ 1869, p. 334 (where this skull is catalogued as
“716 m. Skull of young female. Length 10}, height 74 inches. Terai, Nepal; Dr. Oldham”),
or Hand-List, published 1878, p. 64.

This skull has been figured to show the points in which it contrasts with the British pig’s
skull (figure 1) on the one hand, as to racial characters, and with the Indian boar’s skull (figure
6) as to sexual characters, agreeing, as it does, with the one as to sex, and with the other as
to age.

It is, as may be seen by constructing two triangles with the three first measurements given
of it and of that of Sus cristatus (fig. 6) respectively, a very much smaller skull than that ;
the principal points, however, enumerated in the description of that skull, as differentiating
it from Sus scrofa, can be recognized in its smaller contours. The third molar in the upper
jaw has the part posterior to its second principal lobe much smaller relatively and absolutely
than is the case in Boars of Sus cristatus; this posterior factor, however, effloresces into as
many as 7 cusps in the upper and 5 in the lower jaw of this specimen.

Measurements of Skull 716 m, British Museum Collection. Sus cristatus, from
Terai, Nepal.

inches. inches.
Vertes lemme 9 6 5 0 0 oo 6 Alle Length of lacrymal along malar suture 0°75
Rixtreme hercht.- ani EHiS Interpremolar transverse diameter of
Bee 5 6 oo eo 0 ep 6 oo NO (HAWS soe) oo Go 6 oo LG
Height of lacrymal along rim of orbit iil iumiemim@kie 6 6 56 5 6 6 5 « o lve

5. Skull of Wild Boar, Sus scrofa, var. ferus, from Germany. No. 1513 a, Oxford University Museum.
The entire dentition was in place; the portion of the third molar which is posterior to the four
primary cones is very small as compared either with the rest of the tooth or with its homologue
in the Indian Wild Boar (fig. 6). It is, however, considerably in front at a line drawn as a
perpendicular to the plane of the cutting-edges of the molars from the lacrymal canal. This
skull differs markedly from the skull just named in the greater length, relatively to the
height, of its lacrymal bone, in the greater length of its nasals relatively to the length of the
fronto-parictal region, in the absence of any great convexity in the part of the frontal abutting
upon. the lacrymal, and in the convexity backwards of its fronto-nasal suture.

Measurements of Skull of German Wild Boar, no. 1513 a, Oxford University Museum.

inches. inches.

Extreme length. . . ... . . 165 \Wichin GP meRAI), 5 6. 6 oo oo CS
Taine Inerlng 5 oc 0 6 6 9 9 a | Ol Maximum interzygomatic width . . 59
Base-line from anterior margin of fora- Maximum frontal (imterectorbital)

men magnum to apex of intermaxil- Wilda 6 6 2 o blo 6 0 o 5 ay

laricsmeyte Gee! ca ee LAO Height of lacrymal along rim of orbit 1:0
Length of nasals, approximatively . 80 Length of lacrymal along malar suture 1°2
Fronto-parietal length in vertical aspect 6°8

OF PREHISTORIC TIMES IN BRITAIN. 285

Prats XLITI.

Fig. 6. Skull of Indian Wild Boar, Sus cristatus, killed at Haugul, Dharwar Province, from the collection

of Sir Walter Elliot, K.C.S.I. No. 72.

In this animal the third molar is only just coming into place, and its very large multicuspi-
date posterior lobe is not so far forward relatively as in the European Wild Boar (fig. 5)
nor in other Indian hogs of greater age. It shows, however, very plainly the poiuts, several,
if not always all of which have, by their presence in every specimen of Sus cristatus which I
have examined, enabled me to distinguish it from Sus scrofa, var. ferus. It has the relatively
short lacrymal—a tape, stretched as an arc across the long axis of the skull, from the anterior
inferior angle of the bone on one side to the homologous point on the other, passing over the
frontals, and not over any part of the nasals; the naso-frontal suture, which lies entirely in
front of such an arc, is straight, and not convex backwards; the portion of the frontal which
is bounded internally by the supraorbital channel, and externally by the lacrymal bone, is
markedly convex. The nasals are broader, as is nearly always the case, and shorter also, which
is not by any means always the case, relatively to the fronto-parietal region of the vertex,
than in Sus serofa, var. ferus.

Measurements of Indian Wild Boar, no. 72, Sir Walter Elliot’s collection.

inches.

Extreme length , 16:9
Toa Inara 4G Gc ee ee el
Base-line from ant. margin of foramen

magnum to apex of intermaxillaries 13
Henethyofmosalsisne mylulal eur uae
Length of fronto-nasal suture to

middle of occipital ridge . 58
Width of nasals at apexof frontals . 2
Maximum interzygomatic width . 6
Maximum frontal (interectorbital)

Width: Uo \yaaiewecs ors His Ue et Al

Fig. 7. Skull of Sus barbatus, Borneo.

inches.
Minimum vertical width . . . . 41:8
Height of lacrymal along rim of orbit 1
Lengthof lacrymalalong malar suture 0:9
Interpremolar transverse diameter of

palate soe vunyn eat toon cee
Intermolar . tian yeti 2°05
Length of posterior upper molar . 1°35
IDFRENOMIN Seo a REOMPON Guu Beach Hub nor MES)
Length of third lower molar . . . 1:6
peccadt hy. eee, oe la eke §©6(OF8

1519 d, Oxford University Museum.

This skull, like the other three skulls of the same species examined by me (of which two are

in the British Museum), differs from those already described in large points as well as in small
ones; and there can be little reason for hesitating to accept it as specifically distinct from
them and, indeed, from all other Suidz.

The contour described by the middle line of its nasal and fronto-parietal regions superiorly,
the relations of the greatest width and greatest lengths both of the entire skull and of the nasal
bones, the position of the plane of its greatest interzygomatic width, not posteriorly, but in the
middle of the zygomatic arch, are points of large difference. The exceeding simplicity of its
third molars and the persistence of the mesopterygoid as a distinct bone, are points of small
difference, but yet of great morphological importance.
resembles the other Suid without facial warts.

Its naso-frontal suture and lacrymo-frontal ridge are more like those of these pigs than those
of Sus scrofa.

In the shortness of its lacrymal bone it

286 ON THE DOMESTIC PIG OF PREHISTORIC TIMES IN BRITAIN.

Measurement of Skull of Sus barbatus, 1519 d, Oxford University Museum.

inches. inches.
xtreme lensthy alc seni) ere LOW Maximum frontal (interectorbital)
DAIS eee a Bene circle. cua gh OF width . 45
Base-line utes Ala bark asic G37: Minimum vertical . 1°4
Length of nasals fio pan of postero- Lacrymo-malar line 1)
lateral tips . . ; AL, enrliOr Height of lacrymal 1:2
Fronto-parietal region in eel aspect Length of posterior upper molar 1:4
from same point as preceding mea- Breadth of anterior, which is much the
surement to ea ridge in straight widest lobe acy i 0:9
lime... Meld ice Menem nese ogi O30: Length of posterior lower molar 1:5
Width of batty Nias Meantitye oti (5 Breadth of anterior lobe . 0:8
Maximum (interzygomatic) widih 5 OS Intermolar space at narrowest point 11

Fig. 8. Left lower third molar of Sus andamanensis, female, 24 times the natural size.

This tooth shows the three divisions of the third molar of the true Suidz in great simplicity.
There are two bicuspidate lobes corresponding to the two principal lobes of molar 1 and molar 2,
and, like them, enclosing a single azygos lobe in the middle line between them. This azygos
lobe is developed from the second bicuspidate lobe. There is no ridge developed in Sus anda-
manensis on the anterior part of the tooth, é.e. on the face in contact with molar 2. Poste-
riorly to two bicuspidate lobes, which already show signs of wear, are seen five smaller cusps,
occupying in all a much smaller space than the rest of the tooth. In the males of Sus cristatus
these five smaller cusps would, as in well-fed domestic pigs, occupy a very much larger space
relatively and absolutely than they do here, or even in Sus scrofa, var. ferus. In Sus barbatus
the third lower molars are as simple as they are in Sus andamanensis, and, like this Sus, have the
third molars of the upper jaw simpler still than the lower. The five posterior cusps consist of
one placed mesially in the interspace between the primary cusps of the second pair, of three
placed like the dots in the sign -.-, and of a fifth, not constant, accessory cusp placed on the
inner side. The four latter are represented in the upper jaw of this pig by two, and in Sus
barbatus by one main cusp.

Trans. Linw. Soc. Ser.2.Zoon Vou.) Tas. 41

D.SCROFA wan FERUS. ?
Thames valley.

Caldo

Fig. 1
S.SCROFA va: DOMESTICUS. 2
Colite aierment, Yorkshire.

alba

Fig. 3.
S_ANDAMANENSIS. 2

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XIV. On two new Forms of Deep-sea <Ascidians, obtained during the Voyage of
HM.S. ‘ Challenger” By H.N. Mosstey, I.A., F.R.S., Fellow of Exeter College,
Oxford. (Communicated by Dr. J. Muniz, F.L.S.)

(Plate XLIV.)

Read December 21st, 1876.

THE two Ascidians herewith described and delineated are most unusual and aberrant
in several particulars and quite new to science.

The first form was obtained from the great depth of 2900 fathoms. I propose to
name it Hypobythius calycodes, in allusion to its occurrence at a vast depth and its cup-
like form. It was unfortunately imperfect when obtained. It is remarkable for
having a series of cartilaginous plates developed in its otherwise rather soft and
yielding test, which plates are arranged in symmetrical patterns, as will be seen by the
figures.

The other form, which I propose to name Octacnemus bythius, is most remarkable, on
account of its stellate eight-rayed form, and abnormal in the details of its structure. It
was obtained from 1070 fathoms. It appears to be entirely without near allies in the
group of Ascidians. The gill-sac is flattened out so as to be nearly horizontal, and
there is no gill-network. Peculiar developments of the muscular tunic are prolonged
into the eight curious conical protuberances of the test. The principal viscera are con-
tracted within a small rounded mass or nucleus, as in Salpa.

As only a single specimen of each of the forms was dredged by H.M.S. ‘Challenger,’
the specimens themselves are, I presume, unique; and since they are highly interesting
in many respects, are well worthy of the Linnean Society’s notice.

I. HyPoBYTHIUS CALYCODES, gen. et sp.nov. (Plate X LIV. figs. 1-6.)

On July 5th, 1875, the trawl was used in 2900 fathoms in the North Pacific Ocean,
lat. 37° 41’ N., long. 177° 04’ W. ‘The bottom consisted of a red clay with numerous
concretions of peroxide of manganese. A gigantic Hydroid, a Monocaulus* seven feet in
the length of its hydrocaulus, fragments of Hyalonema, a Holothurian, and a few other
animals were obtained, and with them the Ascidian now to be described in detail. The
specimen was considerably lacerated, but almost all its essential structures were pre-
served. The animal has the form of an inverted cone, compressed laterally so as to have
an oval transverse section. From the apex of the inverted cone is continued a cylindrical
stem, which is enlarged towards its inferior extremity (Pl. XLIV. fig. 1). That side
of the animal on which the ganglion lies will be denoted the dorsal. The animal is
viewed from the ventral surface in fig.1. The test is hyaline and extremely transparent.

* See a letter by G. J. Allman, F.R.S., in ‘ Nature,’ vol. xii. Oct. 28, 1875.
SECOND SERIES.—ZOOLOGY, VOL. I. 2 Q

288 MR. H. N. MOSELEY ON TWO NEW

Where it is simple it is thin and flexible, but in certain spots it is strengthened and
rendered stiff by the presence in it of rounded or plano-convex masses or plates of denser
tissue, which are tough and cartilaginous in consistence, and which are disposed over the
surface of the test in a nearly symmetrical manner. ‘These plates are extremely con-
spicuous when the test is held up to the light and viewed by transmitted light, because
they refract the light strongly; and the pattern formed by them on the test when
thus viewed has a very peculiar appearance. The disposition of the plates on the
ventral surface’ of the body will be seen from fig. 1, that on the dorsal surface from
fig. 2.

A series of globular lobes range on either lateral margin of the body, and give it here
considerable rigidity, and a ridge of slightly condensed tissue runs across the body at
the upper margin of its dorsal surface. A series of flattened plates is disposed over the
inferior region of the ventral aspect of the body, whilst the superior region of the same
aspect, covering the gill-sac and perforated by the inhalant aperture, is entirely devoid of
plates, very thin and flexible, and most perfectly transparent. On the dorsal aspect of
the body a very large plate occupies the middle line inferiorly, whilst immediately above
it two pairs of oblong plates (fig. 2,d) form a stiff shield for the principal viscera, which
lie upon them. ‘Two series of oval plates range on either side of the larger median ones,
and extend up as far as the exhalant aperture. The test-tunic is continued downwards
from the upper region of the body to form the outer wall of the cylindrical stem, thus
forming a tube. ‘The lower end of this tube is widened out into a funnel-shaped mouth,
and in the specimen had apparently been torn away from some object of attachment. -

The substance of the test is composed of transparent hyaline tissue, in which are
embedded the small bodies represented in fig. 4, the larger of which have a length of
from 007 to 014 millim. They are irregular in form, sometimes crystalline, or with
apparently crystalline contents. They are not sensibly altered in appearance by the
action of acetic acid, and no effervescence is produced in the test-tissue by that re-
agent. The bodies are present in the greatest abundance in the test-tissue at the base
of the stem. In the plates of denser tissue they are rather less abundant than else-
where. The test forms a simple sac continuous with the tubular cavity of the stem.

The exhalant orifice is an aperture in the test situate at the end of a short tube projecting
externally on the dorsal aspect just below the nerve-ganglion. Into it the ducts of the
generative glands and the rectum open. The inhalant aperture was entirely obliterated
in the only specimen obtained; it must have lain on the ventral aspect of the body,
since the dorsal wal! was intact. The arrangement of the muscular fibres and remnants
of attachment of the gill-sac seemed to indicate the position for it given in the figure,
where it is introduced conjecturally.

Closely attached to the inner surface of the test-wall is a delicate tunic (the mantle)
containing muscles. The muscles occur in the form of very fine bands, which have
a nearly parallel course. The series of bands springing from near the region occupied
by the heart follow the curved inner surface of the test-cavity towards its superior
margins. ‘The muscles are disposed most thickly in the lateral regions. The mesial
region of the dorsal surface is entirely devoid of them, but they extend over the whole

FORMS OF DEEP-SEA ASCIDIANS. 289

ventral wall. Only the lateral fibres are indicated in the figure; they are prolonged
superiorly in a horizontal direction along the upper margin of the dorsal wall of the
test-cavity. The tubular cavity of the stem is filled by a core composed of muscular
fibres embedded in gelatinous tissue, a prolongation of the mantle.

Of the gill-sac only a small portion remained intact in situ, but fragments here and
there attached indicated an arrangement as shown in the figure. The small portion
im situ lay over the nerve-ganglion. The fenestrations in the membrane are small,
simple, and irregular (fig. 3).

The mouth is situate nearly in the middle line (m, fig. 1). It leads by a short trans-
parent cesophagus to a stomach (s), which has opaque walls corrugated externally ; and
this viscus leads into a rectum which curves up to end at the exhalant aperture.

Beneath the stomach is a tubular heart with a wide vessel leading from it downwards
towards the stem.

In the loop formed by the cesophagus, stomach, and rectum is the large ovisac or
ovary, which is circular in outline. The elongate tubular testis running parallel to the
rectum in the middle line terminates posteriorly internally to the ovisac, its lower end
spreading out into a series of ramifications, which appear very like large nerve-fibres.
The testis-tube opens into the exhalant aperture close to the rectum, as does also the
oviduct, which lies to the dorsal side of the testis-tube and in the same line with it.

The ovisac was full of very large ova, measuring as much as 1°5 millim. in diameter
(fig. 5). These ova, on slight pressure being applied, were discharged from the oviduct.
They were found to consist of a very thick transparent test, with contents composed of
oily yelk-globules, without a germinal vesicle.

The testis was tumid, and full of an opaque white matter, which was discharged on
pressure in tenacious threads composed of spermatozoa.

The spermatozoa are very small. They were examined under a Hartnack No. 10
(immersion system). They consist of an elongate rod-like head, measuring ‘005 millim.
in length, and an excessively fine tail, the length of which could not be determined.
All the spermatozoa examined had a small transparent vesicle attached to one side of
their heads, as shown in the figures (fig. 6).

This Ascidian appears to be allied to Boltenia; but it is especially remarkable because
of the occurrence in its test of symmetrically arranged cartilaginous plates, which form
apattern. The name Hypobythius calycodes is proposed for it.

II. OcTACNEMUS BYTHIUS, gen. et spec. nov. ' (Plate XLIV. figs. 7-13.)

This stellate Ascidian was trawled March Ist, 1875, in 1070 fathoms, lat. 2° 33’S., long.
144° 04’ E., about 40 miles north of Rossy Island, Schouten Islands. From its peculiar
appearance, due to the presence of the eight long radiating conical processes of the test,
the animal was at first supposed to be a Medusa. The single specimen was considerably
injured, the muscular networks maintaining their attachments in only three of the conical
processes, but the test was entire.

The test of the animal is gelatinous and hyaline. On the under surface the body
presents a flat area of a nearly oval form (fig. 7). The border of this base is thickened

2Q2

290 MR. H. N. MOSELEY ON TWO NEW

into a slightly prominent, rounded ridge running round the periphery of the entire basal
area, and, further, is indented slightly opposite the interspaces between the long conical
processes, so as to have an undulating outline. Towards one end of the base (which
end of the animal will be termed anterior, since it is that in which the nerve-ganglion
lies), and in the middle line, is a prominence, also oval in outline (fig.7, pa). This pro-
minence is formed of a process of the basal part of the test. It terminates outwardly
in a tangled mass of rootlets, massed amongst which was found much sand and shell-
particles from the bottom. The Ascidian was evidently attached by this process or
pedicle.

Above the margin of the base the body of the animal is somewhat contracted, but its
walls then again spread outwards and extend into eight wide conical processes. The
processes terminate in abruptly narrowed tentacular-like tips (fig. 7, ¢, ¢), which are im-
perforate, and in which no sense-organ or any special structure could be discovered.

On the upper aspect of the body the eight conical processes are directly continuous
with the upper surface, which is somewhat hollowed or saucer-shaped.

Towards its anterior part in the middle line the upper surface is perforated by a trans-
verse slit, the inhalant aperture (fig. 8, 7a). The exhalant aperture is situated in front of
this, and at a lower level, proceeding from the wall of the body just above the base as
a short cylindrical tubular process (figs. 7 & 8, ea). The inhalant aperture is enclosed bya
pair of simple rounded lips, and is without tentacles.

The test forms a wide cavity, which extends freely into the capacious hollow conical
processes. The two apertures, inhalant and exhalant, form the only communication
between this cavity and the exterior.

A flat horizontal membrane is stretched across the test-cavity in such a manner as to
separate off an upper chamber communicating with the inhalant aperture from a lower
communicating with the exhalant. This membrane, in the central region of the body, is
thick and of an opaque white colour (fig. 9, 7). This thickened central portion runs out
peripherally into eight processes, directed to the intervals-between the long conical pro-
cesses of the test. Opposite these intervals the processes become attached, or give origin,
to bands of muscular fibres, which bands, after a short radial course, in which their
fibres remain parallel, split into two halves (fig. 7, 7m). The halves of the bands diverge
at an angle from one another, and proceed to the tips of the long conical processes, where
each is joined by the tip of a corresponding half-band from the next adjoining process.
The half-bands, as they run towards the tips of the conical processes, give off a series of
transverse muscular threads, which, passing from band to band, form a series of loops
one beyond the other, continued almost to the tip of the conical processes. These radial
muscles are apparently the homologues of the longitudinal muscles of ordinary Ascidians.
They would become longitudinal were the upper part of the discoid body of the animal
drawn upwards, so 4s to make the respiratory cavity tubular instead of saucer-shaped.
A second series of muscular threads lies beneath the radiating bands just described
about their points of bifurcation, and extending thence almost to their points of origin.
This second set of muscles takes a circular direction (fig. 7, em), and is continued round
the entire circuit of the animal, the several strands, of which there are about twelve,

.

FORMS OF DEEP-SEA ASCIDIANS. 291

appearing to be continuous throughout their length. These circular muscles lie beneath
the radial ones; and were the respiratory cavity elongated into a cylinder, the radial or
longitudinal muscles would thus be internal in position, the circular external. In order
to prevent confusion, and because of the difficulty of drawing them clearly, not nearly
the entire number of transverse and circular strands is inserted in the figures in the
accompanying Plate (figs. 7 & 8). Their arrangement is shown in detail in fig. 12.

Over the muscular meshwork thus formed, and extending from it to be continuous
in all directions with its thickened central portion, the horizontal membrane is continued
as a thin and transparent lamina. Opposite the indentations in the margin of the
thickened central portion of the membrane, 7. e. between the processes or thickened
folds attached to the muscular bands, this thin lamina is loose and hangs in bags or
depressions.

In the bottom of each of these depressions is a slight three-cornered elevation (fig. 8, a),
at the tip of which a perforation or aperture may possibly be present; but in the
specimen examined such could not be made out. At the tip of each small elevation were
seen only three minute closely apposed villous folds of the membrane, between which no
aperture could be detected. The central horizontal opaque white membrane is merely
a thickened part of the general membrane, which is spread over the muscular meshwork,
and reaches to the tips of the large conical processes. It is probably respiratory in
function, and represents a gill.

The membrane was observed to be attached to the inner surface of the test-wall at the
intervals between the conical processes ; but the specimen was too much injured to allow
of the investigation of the extent and manner of its attachment within the conical pro-
cesses. It appeared to be attached laterally on either side to the inner walls of these
processes, and is probably reflected so as to line their cavities. No normal perforated
gill could be discovered in any part of this membrane; but probably the central
thickened portion has a respiratory function, and possibly the eight small elevations may
prove to be openings. No reflection of the membrane over the inner surfaces of the upper
and lower walls of the test was observed.

The membrane was composed of an irregular mesh of fibrous tissue, with numerous
rounded gland-like cells, and numerous nerves proceeding to the radial muscular bands.

The mouth lies in the anterior part of the thickened portion of the membrane above
described, and at a short distance behind the inhalant aperture. Behind it is the endo-
style. The mouth isasimple oval aperture, encircled by a few sphincter muscular threads.
It leads directly into the digestive tract, which is embedded in a compact nucleiform mass,
which is seen conspicuously through the transparent test on viewing the animal from
beneath. The nucleus lies entirely beneath the horizontal membrane, which is attached
round the margin of themouth. ‘The exact arrangement of the viscera inside the nucleus
was not determined. A short tubular rectum projects from the anterior and inferior
extremity of the nucleus.

Posteriorly to this, on the inferior surface, is a well-developed ovary, and behind this,
again, the testis. The ovary is a racemose gland, whilst the testis is composed of short
cecal tubes. In minute structure these organs agree closely with the testes and ovaries
of other Ascidians.

292, MR. H. N. MOSELEY ON TWO NEW

Between the mouth and rectum, in the middle line, lies the heart-shaped nerve-

ganglion, immediately beneath which is a spherical body, which had the appearance of a
thin-walled sack full of a milky fluid. The wall of this sae is composed of a layer of
cells of rectangular outline, from the inner surface of which fine hair-like processes
depend (fig. 13). The hairs are not straight, but wavy, and appeared stiffer than
ordinary cilia. This sense-organ was not in sufliciently good preservation to allow of
more accurate histological investigation. A large nerve from the ganglion passed directly
over it in the middle line.
. A pair of strong muscular masses is developed at the posterior part of the nucleus,
one mass lying on either side. These muscles appear to take origin from the under-
side of the thickened horizontal membrane above, and to terminate on the sides of the
posterior part of the nucleus. The terminations of the muscular slips composing the
muscles are bifurcate (fig. 10, mm). Other narrow transverse muscular bands are present
between the mouth and rectum, embracing the fore part of the nucleus (fig. 10, m0).
The use or homologies of these muscles are not apparent.

The endostyle lies in the middle line, at the back of the nucleus, between the pair
of posterior muscles of the nucleus (fig. 10,2). It is very short. It showed the cha-
racteristic structure of the Ascidian endostyle—long, fine, granulated, spindle cells, packed
close side by side, with their long axes at right angles to the length of the organ.

The relation of the endostyle to the horizontal membrane was not observed, since the
organ was only discovered after the nucleus had become detached from the membrane.
The endostyle was then found tucked in between the paired posterior muscles of the
nucleus. It is placed in the position given to it in the figures, because this seems to be
the necessarily correct one. The organ, at all events, is on the ventral side of the
animal, or on the side of the mouth opposite to that on which the nerve-ganglion lies,
which is its normal seat.

The length of the base of the Ascidian was 5°5 centims., breadth 4°75 centims.; extreme
length between tips of the protuberances 7'5 centims.

On the whole this very perplexing animal appears to be an Ascidian in which the
respiratory sac is flattened out so as to become nearly horizontal, and in which no gill-
network is present. In Cystingia (Bronn, KI. und Ord. ii. p. 181) a gill-network cannot
be distinguished.

The radial muscles belong to the longitudinal set of other Ascidians and are internal ;
the circular are external in relative position. I can find no homologues of the muscles
of the nucleus. In having the viscera contracted into so small a nucleus, the animal
resembles Salpa. The nerve-ganglion is abnormal in position, in being situate on the
nucleus. It, nevertheless, is normal, in lying between mouth and anus, whilst the endo-
style is on the opposite side of the mouth as in other Ascidians.

The name Octacnemus bythius is proposed for this curious eight-rayed deep-sea form.

The above account is the result of an examination of the animal in the fresh condition.
A further investigation of the specimen as preserved in spirit is much to be desired.

FORMS OF DEEP-SEA ASCIDIANS. 293

DESCRIPTION OF PLATE XLIV.

Figs. 1-6 inclusive illustrate the structure of Hypobythius calycodes.

Fig. 1. The animal viewed from the ventral surface (i. e. the surface opposite to that on which the nerve-
ganglion lies). The viscera are seen through the transparent body-wall lying on the opposite
side of the cavity of the test. At the lateral margins the test swells into a series of lobes
composed of masses of compact gelatinous tissue, which lobes give a certain rigidity and
support to the test. Similar masses or plates of compact tissue are seen on the ventral
surface of the test in its lower region. In the single specimen obtained a longitudinal rent
divided the ventral wall of the test, and of the mantle and gill-saes, along the middle line down
to about the point indicated by the letter h.

This figure represents the animal of half the natural size, and the lettering applies as
follows :—

a, a. Plates of dense test-tisswe, similar to the other plates also seen, which are unlettered.
6. Muscular core of the stem of attachment of the animal. c. Prolongation of test forming the
outer tissue of the stem of attachment, here, at the end of the stem, widened out into a funnel-
shape. za. The inhalant aperture. No trace of this aperture remained in the specimen as
obtained ; its most probable position only is given. It may have lain somewhat more nearly
opposite the mouth on the ventral surface. ea. The exhalant aperture. ng. Nerve-ganglion
with sense-organ seen through the test and across its cavity. o. The ovary; od, the oviduct;
and ¢ the testis. gs. Margin of gill-sac attached by a series of fibres to the membrane lining
the interior of the test. Externally to the gill-sacs the directions of the lateral muscular bands
and their anterior prolongations along the dorsal margin are indicated. m. The mouth; s, the
stomach; and /, the heart. 6v. Large blood-vessel leading from the heart towards the stem.

Fig. 2. Sketch of the upper portion of the animal, much reduced in size, and as viewed from the dorsal
aspect, to show the symmetrical arrangement of the plates of thickened test-tissue.

ea. The exhalant aperture. d. Set of four plates covering the generative and digestive
organs. Beneath there is a large elongate oval plate.

Fig. 8. Portion of the gill-sac, to show its small and irregular fenestrations, which have a diameter of

about *10 millim.
. Bodies, partly crystalline or with crystallime contents, occurring in the tissue of the test.
. Ovum from the ovary; actual diameter 1°5 millim.

D> oO

. Spermatozoa, one of them viewed sideways, so as to show the attachment of the vesicle to the head.
Actual length of the heads of the spermatozoa 005 millim.

Figs. 7-18, inclusive, illustrate the structure of the Ascidian Octacnemus bythius,
trawled March 1, 1875, from 1070 fathoms.

Fig. 7. The animal viewed from below: natural size. The central part of the figure is occupied by the
flat irregularly oval base, with undulating outline and thickened margin (6). Beyond the
margin of the base the eight large conical processes are seen to radiate outwards, and terminate
in (¢) abruptly narrowed tentacle-like tips. The short pedicle of attachment (pa) projects
from the anterior part of the base ; the other parts (nucleus, respiratory membrane, and muscles)
are seen through the transparent wall of the test constituting the base. The nucleus is turned
over slightly to the left hand.

ea. Exhalant aperture. 6. Thickened border of the base. pa. Pedicle of attachment.
“7. Therectum. n. The nucleus. cm. Circular muscles. rm. Radiating muscles. g. Thickened
horizontal part of mantle-membrane.

294 MR. H. N. MOSELEY ON TWO NEW FORMS OF DEEP-SEA ASCIDIANS.

Fig. 8. The animal viewed from above ; also of the natural size. The long conical processes are seen as
in fig. 7, but their upper surfaces are flatter than their lower, and pass directly into the smooth,
somewhat hollowed, general upper surface of the animal.

This upper surface is perforated towards its anterior end by the transverse slit-like inhalant
aperture (¢a). Anteriorly part of the lateral wall and base are seen projecting beyond the out-
line formed by the union of the bases of the tentacles, and projecting from it the tubular exha-
lant aperture (ea). The mouth, endostyle, muscles, and membrane are seen through the
transparent upper part of the test. The rectum and nerve-ganglion are seen through the test
and horizontal membrane as well; they are somewhat directed to the left, being normally
in the median line. In both drawings the transverse muscular slips within the long. conical
processes are represented far too stout, and are not numerous enough. ‘These derivatives of
the radiating muscles are far less stout than the circular muscular slips.

za. Inhalant aperture. ea. The exhalant aperture. m. Themouth. e. Endostyle. r. Rec-
tum. ng. Nerve-ganglion. «a. Pyramidal eminences in the bottoms of the bag-like depressions
of the gill-membrane ; possibly apertures.

Fig. 9. Schematic, vertical, and longitudinal section, through the animal, along the middle line, z.e.
following the direction of the arrows in fig. 8.

ia. Inhalant aperture. ea. Exhalant aperture. m. Mouth. mn. Nucleus. mn. Muscles of
nucleus. 7. Rectum. g. Respiratory membrane. rm. Radiating muscles. ¢. Thickened
margin of the base. pd. Pedicle of attachment with its fibrous roots.

Fig. 10. The nucleus, enlarged. The horizontal membrane is attached round the mouth, which opens
above it, whilst the whole of the remainder of the nucleus lies below it. The cut edge of the
membrane (g*) is seen to pass between the nerve-ganglion and mouth, and (g) between the
posterior muscles of the nucleus and the endostyle.

m. Mouth with sphincter muscles. e. Endostyle, with membrane alongside. rm. Radiating
muscle (the median anterior band). g*. Edges of cut membrane. ng. Nerve-ganglion. so.
Sense-organ. m6. Transverse muscular bands on the anterior aspect of the nucleus. 7. Rec-
tum. mn. Posterior muscles of the nucleus. s. Stomach. ¢. Testis. 0. Ovary.

Fig. 11. The nerve-ganglion and sense-capsule, much enlarged.

ng. Nerve-ganglion. n,n. Nerves passing round the mouth, one on either side. m. Mouth.
so. Sense-capsule, across which runs a large nerve-stem, n*, proceeding from the inferior extre-
mity of the ganglion. The two filaments proceeding from the sense-capsule are of uncertain
nature.

Fig. 12. Muscular arrangement at one of the points of bifurcation of the radiating muscles, enlarged.

rm. Radiating muscles, appearing as a broad flat band towards their inner part, or part nearer
the centre of the discoid animal. The fibres composing the band commence at a, where their
pointed extremities are embedded in the thickened radial process of the thickened central part
of the horizontal membrane. Externally the band bifurcates, and the two halves of it, passing
into the long conical processes, give off thin transverse looping fibres on their outer side. cm.

The circular muscular fibres which lie behind the radiating ones. The inner ones curve inward,
the outer outward.

Fig. 13. Structure of the wall of the sense-capsule.

a. Part of the layer of cells composing the wall of the sense-capsule, seen in perspective.
b. Single cell, with its hairs or cilia.

TRANS. Lm. Soc Ser.Z. Zoot Vor.) Pi 44

HN Moseley. del?

OBerjean hth. Hanhart imp.
NEW DEEP-SEA ASCIDIANS

[ 295 ]

XV. On new Forms of Actiniaria dredged in the Deep Sea; with a Description of
certain Pelagic Surface-swimming Species. By H. N. Mosztny, WA., F.R.S.,
Fellow of Exeter College, Oxford, late Naturalist on board H.U.S.* Challenger,
(Communicated by Dr. J. Murine, £.L.8.)

(Plate XLV.)
Read February 15th, 1877.

THE deep-sea Actiniaria herewith to be described were obtained on various occa-
sions by the dredge or trawl during the voyage of H.M.S. ‘ Challenger.’ Actiniaria of a
variety of forms were obtained in deep water frequently during the Expedition and in
very various localities, and only a small part of these are accounted for in the present
paper. Several circumstances, such as rough weather, and the fact that, as frequently
was the case, the dredge or trawl was only got up to the surface at nightfall, combined
to prevent all the Actiniaria obtained being worked at in the fresh condition, and very
often, when other circumstances were favourable, some more important subject called
for immediate attention. A few of what appeared to be the most interesting forms were,
however, sketched and partly worked out whilst the animals were fresh, and the
results are here given. The pelagic Actinias ((/imyadine) were obtained either with
the surface-net or in the trawl.

Family ACTINTIAD &.
Subfamily Minyapinaz.

NAUTACTIS PURPUREUS, n. sp. (Plate XLV. figs. 1 & 2.)

A small floating Actinia was found amongst the contents of the towing-net on
August 26th, 1874, lat. 13° 39’ S., long. 150° 53’ H., between the New Hebrides and the
north-east coast of Australia. The animal was very small, not more then 7 millims. in
height, when measured with the parts about the mouth protruded. The body is cylin-
drical, with the aboral surface much flattened. The costz are slightly elevated rounded
ridges, 24.in number, nearly equal in size, and without tubercles. The tentacles are 12 in
number, disposed in two series, 6 primary larger ones lying nearer the mouth than the
6 somewhat smaller secondary ones; they are simple, very short indeed, reduced
in fact to rounded conical tubercles. Between the tentacles at the margin of the disk
are a series of small tubercles of triangular outline. The mouth is linear. The cosial
and aboral surfaces of the animal are of a chocolate colour with a tinge of lilac. Near
the margin of the disk is an irregular line of bright orange-yellow. The margin of the
mouth is of a dark burnt-sienna colour. From it radiate out streaks of chocolate
colouring, which fork to embrace the bases of the transparent white tentacles and the
intertentacular tubercles. These tubercles are of a glistening but opaque white, with
bright orange colouring on their lateral borders, and a dark spot in their centre.

SECOND SERIES.—ZOOLOGY, VOL. I. 2k

296 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

The animal on retracting the protruded parts of the mouth became of a simple cylin-
drical form with the disk concave.

The absence of tubercles on the costa, and their presence on the calicinal margin
only, and the extreme shortness of the tentacles, places this Minyad near the genus
Nautactis. It differs, however, from the Nawtactis olivacea*, as described by MM.
Milne-Edwards and Haime, in having 24 costs instead of 22, and in the nature of the
tentacles, which in J. olivacea are tuberculated, simple and very small towards the
centre, and multilobate towards the circumference of the disk.

Larva Minyap. (Plate XLV. fig. 3.)

On Nov. 2, 1874, off the Philippines, in lat. 12° 48’ N., long. 122° 10' B., a larval
Minyad was obtained with the towing-net from the surface.

This larva was about 1°5 millim. in height. The body was ciliated over its entire outer
surface. No tentacles were as yet formed in the larva, but the mouth was maintained
everted, and showed six distinctly marked lobes. Six primary mesenteries were well
developed, and six intermediate secondary just indicated. Upon the larva being
compressed, a single pair of mesenterial filaments was emitted from its mouth. The
filaments and the ectoderm were crowded with simple, elongate, ovoid nematocysts.

OcEANACTIS, gen. nov.

Characters.—Body transparent in the contracted condition, smooth and spherical in
shape; when expanded hemispherical, provided with rounded costal ridges and a single
row of costal tubercles. Tentacles simple, elongate, conical, and in two rows. Base
very small, entirely invisible in the contracted condition, with an aperture in the centre
communicating with the body-cavity.

- OCEANACTIS RHODACTYLUS, n. sp. (Plate XLV. fig. 4.)

On July 10th, 1874, the trawl used in 700 fathoms in lat. 37° 36’ S., long. 179° 24' E.,
about forty miles off New Zealand, came up full of a tenacious blue clay. In the clay
were found two Actinias in a contracted condition. When thus contracted they were
nearly spherical, with a tense and perfectly smooth transparent external tunic, through
which the mesenterial septa were plainly visible, as also the ovaries and the stomach-
wall. No trace of any base or basal aperture could be made out in this condition of the
animals. The oral orifice appeared as a small, slightly protuberant reddish cone. Hach
animal on the whole looked not unlike a small onion. Placed in fresh water for the
night the animals expanded to the form shown in the accompanying figure.

The body of the animal in the expanded condition is nearly hemispherical. The upper
part of the stomach is protruded as a semitransparent cone beyond the circle of ten-
tacles. There are twenty mesenterial septa, the spaces between which rise up as rounded
ridges or costee on the wall of the expanded body. ‘Ten of these bear, just above the
origin of the tentacles, rounded prominent tubercles.

* ¢ Histoire Naturelle des Coralliaires,’ Milne-Edwards et J. Haime, tome i. p. 230.

AND PELAGIC SURFACE-SWIMMING SPECIES. ; 297

The tentacles, 20 in number, are simple, elongate, conical in shape, and disposed in
two series. The tentacles of the outer series are the larger, and correspond to those
costze which bear tubercles. The inner are in connexion with the intermediate coste,
and arise from the edge of the disk just internally to the larger tentacles. At the aboral
pole of the animal is a small aperture, surrounded by a small circular area with tumid
folds representing the base. This aperture is in direct communication with the body-
cavity, for one of the ovaries became protruded through it without any violence having
been used. There is hence no separate chamber for the air-vesicle as in Minyas, but the
condition is like that in Peachia. The general transparent appearance of the animal,
however, and absence of base sufficient for attachment in an animal of this short hemi-
spherical shape, point to its being a free-floating and not a deep-sea form. Perhaps the
condition is here somewhat as in Nawtactis, where the base may be reversed, the air
extruded, and the body attached from time to time to floating bodies (M.-Hd. & H. /.c.
p- 230). The body of the animal is transparent and bluish coloured, with a slight
dusting of light red on the costee. The base is red. The costal tubercles are transparent
and marked with red, especially at their lines of origin from the costz. The tentacles
are of the same light red colour and somewhat transparent. The margin of the mouth,
which presents ten slight indentations, is also tinged with red. In the contracted con-
dition of the animal the ovaries show through the body-wall as yellow bodies near the
aboral pole. The stomach also is seen in the same manner as a transparent whitish
cylinder. The tentacles and tubercles are full of nematocysts of the common elongate
form, and about ‘04 to ‘05 millim. in length.

The animal apparently belongs to the group of the Minyadine (M.-Edwards & Hatate,
1. c. tome i. p. 227), but does not seem to come exactly within either of the three genera of
that subfamily, viz. Winyas, Plotactis, or Nautactis. Erom Nautactis it is distinguished
by the simplicity of its tentacles, from Plotactis by its single row of costal tubercles, and
from Minyas by not possessing a series of tubercles on each costa. It is further
distinguished from all three genera by the perforation of its base.

Subfamily ActTininaz.
ACTINIA ABYSSICOLA, sp.n. (Plate XLV. fie. 5.)

This animal, found adherent to the stem of a species of Mopsea, as depicted, is drawn
out and elongated in a direction transverse to the central axis of the body in relation
with the slender cylindrical form of its support.

The adherent surface of the base is extremely elongated, and of the same width
throughout, and is closely applied to the Mopsea-stem lapping round it, the region of
junction of the opposite edges being marked by a perfectly straight line. Along this
line of union the two edges are somewhat coalescent, but separable without the appli-
cation of much force. At either end of the elongated base the body swells out into
rounded masses, narrowed at the extremities, and evidently representing the fleshy
lobes present around the bases of normal Actinias, which are thrown into this form in
this instance by the extreme contraction and elongation of the attaching surface. Within

2R2

298 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

the rounded masses is a very stout and prominent muscular ring, pulled out into an
oval form, and somewhat contracted laterally; and within this is a smoother space
rounded from side to side, and with its inner margin bounding an oval opening,
immediately within which le the tentacles. This region of the body is coloured bright
pink, with a few radial streaks of slightly darker colour, and contrasts with the muscular
ridge and lobes external to it, which are of a pale pinkish yellow. The muscular ridge
is especially pale in colour. This ridge and the coloured space being immediately
internal to it, represent the vertical wall of the body of normal Actinias. The tentacles
are small, about 60 in number, disposed in two rows, and of a very pale pink colour.
The disk is dark pink in colour and of an oval form. The mouth very irregularly oval.

The greatest length of the largest specimen obtained was 34 millims. Another
specimen measured 23 millims.

Actinia abyssicola was obtained on two occasions from deep water, viz.:—On May 6th,
1873, in lat. 40° 17’ N., long. 66° 48’ W, from 1350 fathoms; and also on May 29th,
1878, to the south-west of Bermuda, from 1075 fathoms.

The two specimens procured on May 6th came up alive, but never expanded them-
selves more than as seen in the figure.

Very similar Actinias were dredged frequently from deep water attached to rod-like
bodies, such as Aleyonarian stems. I find in my notes, e.g., that one came up from
1400 fathoms, attached to a Hyalonema-spicule (lat. 12° 8’ S., long. 145° 10° H.), near
Raine Island, N.E. coast of Australia, August 29, 1874. I identified the species, how-
ever, only in the two above-cited instances.

ACTINIA GELATINOSA, n. sp. (Plate XLV. fig. 6.)

The body of the single specimen of the animal obtained was gelatinous and transparent.
The base was closed tightly around the stem of a dead Gorgonoid. The opposite
margins of the basal surface were approximated beneath this stem, and closely united
and coalesced along a linear suture. The basal surface thus formed a tightly fitting
sheath, in which the supporting Gorgonoid stem was held. The sheath and stem reached
only to a point opposite the mouth of the animal, the base having been drawn towards
the stem in forming its attachment, so that part of its margin here, instead of its central
point, lay beneath the mouth. Nearly one halfof the mass of the body thus lay in front
of the end of the stem of attachment. This free half was movable on the remainder of
the body, as if on a hinge (fig. 6,@); and as the specimen was turned about, it fell over
so as to shut over or enclasp the other firmly attached portion of the body, and thus
close the cavity of the animal as with a lid. The tentacles appeared to be numerous,
and disposed around the margin of the disk; but owing to their extreme softness were
too much injured to allow of their being counted. The body of the living animal was of
a transparent bluish colour. The tentacles were of a light brick-red colour, the mesen-
teries of a dark madder tint, being coloured with what I have elsewhere termed
“ polyperythrin’”’*.

* Consult H. N. Moseley, ‘‘ On the Colouring-matters of various Animals,” &c., Quart. Journ. of Microsc. Science
(new ser., January 1877), vol. xvii. p. 2. '

AND PELAGIC SURFACE-SWIMMING SPECIES. 299

It is possible that I may have been deceived as to the folding over of the body, for it
may be noted that the animal was injured by the dredge; but it appeared to me that the
hinge-like folding place was natural, and not artificially produced.

This animal was dredged between the Banda group and Amboyna, in lat. 4° 21’ S.,
long. 129° 07’ E., on Oct. 3, 1874.

EDWARDSIA CORIACEA, sp. n. (Plate XLV. figs. 11-14.)

The body is cylindrical, and when fully extended of nearly equal: diameter throughout,
and worm-like in appearance and movement. When the animal is at rest with the
tentacles fully expanded, as shown in fig. 11, the posterior portion of the body is dis-
tended and somewhat ovoidal. The entire median region of the body is covered with a
dense coriaceous test, thickly beset with grains of sand, shells of Foraminifera, &c., the
whole forming a tough brown tube, into which the anterior region of the body is
completely retractile. In the contracted condition the animal is pointed at the anterior
extremity, whilst the posterior thin-skinned region is distended and pushed out into
the form of a rounded disk (fig. 12). The animal thus in the act of retraction of its
anterior extremity must obtain firmer hold in the sand by distension of the posterior
part. The base is imperforate and of a pinkish vioiet colour. The general colouring
of the anterior retractile region of the body and disk is a pale pinkish white. The ten-
tacles are 16 in number, disposed in two series, of which the external are most brightly
coloured. They are marked by equidistant transverse bands of reddish brown, the outer-
most placed on the tips. On the disk are yellow streaks between the bases of the ten-
tacles, and on the light external tentacles reddish-brown streaks placed internally to
these on either side. The oral protuberance is uncoloured. The remainder of the
anterior retractile region of the body, for a short interval beneath the tentacles, is un-
coloured. Lower down it has short longitudinal streaks of reddish brown, corresponding
to the tentacles in position, the streaks corresponding to the external row of tentacles
being broader than those opposite the intermediate ones. The streaks are absent
opposite four of the tentacles of the external row. Thus, passing round the body, are
seen a colourless space, then three coloured streaks, then another colourless space, and
so on (fig. 14). The length of the specimen as obtained, when at rest and contracted
(fig. 12), was about 275 centims.; when fully extended, with the tentacles expanded,
about 4 centims. JZ. coriacea is allied to H. beautempsis (M.-Hd. & Haime, Hist. Nat.
des Coralliaires, tom. i. p. 285) in having only 16 tentacles, but differs from it in the
fact that the tentacles are disposed in two series, in the details of colouring, and in the
structure of the test.

This Actiniad was procured with the dredge from 600 fathoms off Cape St. Vincent, on
January 16th, 1873. It was closely contracted when found in the dredge, but expanded
on being placed in fresh sea-water, and lived several days.

CoRALLIMORPHUS, gen. nov.

Characters—Body rigid, smooth, gelatinous, not contractile, without pores, but with
an adherent base; disk circular and large; tentacles non-retractile, elongate, conical,

300 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

with a rounded terminal knob, of several sizes, disposed in regular series at the margin of
the disk and in two circlets on its surface.

CoRALLIMORPHUS PROFUNDUS, n. sp. (Plate XLV. figs. 7, 8, & 8a, 80.)

The animal is in form a short cylinder, with flat upper surface or disk, and is rounded
below, where the base takes the form of an excavation, its margins being turned inwards.
Attached to the centre of the disk, but occupying not nearly its whole area, was, in the
single specimen obtained, a small nodule of peroxide of manganese, of which many were
brought up by the trawl. The wall of the body of the Actinia is firm and rigid, and the
animal is unable to contract so as to shut in the disk and tentacles, 60 tentacles are
present, of which 48 are attached to the margin of the disk, and 12 spring from its
surface. The marginal tentacles are of three sizes—24 being very small, 12 much larger,
and 12 intermediate in dimensions. The disk, which has a diameter of about 4 centims.,
shows a series of radially disposed rounded ridges, corresponding to the summits of the
intermesenterial chambers. Of those corresponding to the 12 larger tentacles, 6 are
broader than the others, and terminate towards the mouth in broader ends; these may
be considered as primary. A pair of them coincide in direction with the long axis of the
elongate mouth. These primary rays of the disk bear each a tentacle at a distance from |
the margin of the disk of about one third their length. The 6 secondary rays bear
smalier tentacles, which are placed much nearer to the margin of the disk. The tentacles
are elongate conical, with a small knob at their extremities. The mouth is of an elongate
oval form, and is sunk in a slight eminence in the centre of the disk. The animal is
covered with a hyaline transparent integument, through which the ochre-yellow coloured
inner tunics and viscera show themselves. The ovaries are especially conspicuous,
being of a dark ochre-yellow, of which colour also is the mouth. ‘The base is of a dark
brown.

The animal has three kinds of nematocysts or thread-cells. One of these is exactly of
the same structure as figured by Mébius as occurring in Caryophyllia Smithii*, being
extremely large and of an elongate cylindrical form, with narrowed ends. These cells in
the present species are remarkable for their great size, being apparently the largest
nematocysts known. They measure about ‘18 millim. in length, whereas the cells of
Caryophyllia Smithit measure only -102 to ‘071 millim. They are thus specially favour-
able objects for examination, and I clearly saw in them the peculiar triple invagination of
the wide first part of the thread figured by Mobius; and I also saw the various stages of
development of the cells exactly as figured by him.

The second kind of nematocysts is of nearly the same form as that just described, but
a little shorter. It contains a thread which is of uniform dimension, and beset through-
out its length with short spines: the thread is simply coiled in a somewhat irregular spiral
within the cell. These two kinds of thread-cells occur together in the tentacles of the
animal. The third form of nematocysts is shown (figs. 8 a, 8 0) as seen in the expanded and

* Mobius, “ Ueber den Bau, den Mechanismus und die Entwicklung der Nesselkapseln,” Abh. nat. Ver. Hamburg,
1866. (Cited by Allman, figures copied by Dana.)

AND PELAGIC SURFACE-SWIMMING SPECIES. 301

unexpanded conditions. These cells are smaller than the others, being of about one fourth
the length of the larger ones. They are of an ovoid form, but slightly bent. They have
a thread which, when protruded, has a very wide, fusiform commencement, round which
is wound a spiral line, bearing spines, the succeeding fine portion of the thread-being
unarmed. In the contracted condition a triflmg invagination takes place in the first
broad part of the thread, as in the largest cells; and the spines are of course reversed in
inclination, being turned backwards towards the base of the cell. These cells seem to
form an interesting transition towards the peculiar three-spined thread-cells of Hydroids ;
they occur on the base of the animal.

A single specimen of this interesting form was dredged in the South Pacific Ocean in
lat. 39° 4 S., long. 105° 5’ W., on Noy. Ist, 1875, from 2025 fathoms. A closely similar
specimen was likewise obtained which differed only in having darker, madder-coloured,
radial streaks on the disk, and in being much larger, measuring 10°5 centim. in diameter
of the disk. It was abnormally irregular in having 52 instead of 48 marginal tentacles,
caused apparently by the intercalation of an additional half-system of 4 tentacles; but it
may evidently be referred to the same species. It was dredged, together with a large
number of other Actininze, in lat. 33° 42’ S., long. 78° 18’ W., on Dee. 17th, 1875, from
1375 fathoms.

It seems necessary to refer the above-described form to a new genus. It resembles
the genus Discosoma* in being rigid and unable to contract the disk and cover the
tentacles, but differs from it in that the tentacles are here not very numerous, and that
they are provided with a knob at the tip, as in Corynactis. Similar or kindred forms
were often dredged in deep water by the ‘Challenger.’ They seem closely allied to the
simple disk-shaped corals, such as Stephanophyllia, in which the tentacles are also
knobbed and disposed at the margin, and on the surface of the disk, in a closely similar
manner. They lave further similar thread-cells to these solitary corals, and, indeed, ap-
pear to differ from them only in having no calcareous corallum developed. Indeed, in one
species of Stephanophyllia, obtained in deep water, the calcareous skeleton was in a most
rudimentary condition, being a mere delicate network of calcareous trabecule. So like
in appearance were these forms of Actininze to the living corals, that it was only by
feeling them as they were found in the dredge that I became convinced that they had no
calcareous skeleton, and I macerated several in caustic potash in the hopes of finding
traces of a corallum present in them. They seem to approach these disk-like corals very
closely, and possibly to represent forms from which the corals were developed.

CoRALLIMORPHUS RIGIDUS, n. sp. (Plate XLV. figs. 9 & 10.)

A form allied to that just described, and evidently to be referred to the same genus,
was obtained amongst the Moluccas in 1425 fathoms. The body of the animal is some-
what cartilaginous, tough and firm, maintaining its shape, and not contractile. Its form
is that of a truncate cone, the base being of less than half the diameter of the oral disk.
The lateral walls of the body present smooth, slightly projecting, rounded ridges or costee,

* Vide MM. Edwards & Haime, J. c. tome i. p. 255.

302 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

corresponding in position to the intervals between the attachments of the mesenteries.
The base is hollowed out into a cavity (fig. 10), which cavity had probably been occupied,
as in C. profundus, by some object to which the animal had been attached. The cavity
is surrounded laterally by a thin but firm wall, formed of the obliquely projecting wall
of the body, the margins of the cavity being thus somewhat undercut, so as to afford
increased holding power. The bottom of the cavity, or actual base, is smooth and
imperforate. The oral disk is elevated and rounded towards the centre. The tentacles
are 72 in number: of these, 12 larger or primary are disposed at equal intervals along
the margin of the disk, with 12 secondary and 24 still smaller tertiary ones placed
symmetrically between them. On the surface of the disk itself, at about half the distance
between the margin of the mouth and that of the disk, is a circle of 12 tentacles, placed
opposite the primaries, and intermediate in size between these latter and the secondaries.
Halfway, again, between the line of origin of this inner circle of tentacles and the disk
margin is a further circle of 12 smaller tentacles, placed opposite the secondaries. The
tentacles are elongate-conical in form, tipped by very distinct knobs. The bases of the
tentacles are very firm and cartilaginous. The general colour of the animal is a very
pale pinkish yellow. The disk has a dark madder colouring * in parts, and rings of the
same colouring girdled the tentacles just beneath their knobs. The single specimen
obtained was much rubbed, and hence the exact distribution of the pigment was
uncertain.

The specimen was dredged between the Banda group and Amboyna, in lat. 6 21’ S.,
long. 129° 7’ H., from 1425 fathoms, on Oct. 3rd, 1874.

Family CERIANTHID&.

CERIANTHUS BATHYMETRICUS, sp. nov. (Plate XLV. figs. 15-20.)

This species is contained within a tube formed of the felted threads of its large
nematocysts, as is the case with the other species of the genus. The tube is 11 centims.
long, and about 14 centim. broad in the broadest part. It is represented of the natural
size in the figure (fig. 15). ‘The opening at the upper extremity is in the form of a longi-
tudinal slit. The tube is beset with foraminiferous shells. Embedded in its substance
were found spicules of a Hexactinellid sponge, and also amongst its fibres were nume-
rous small nematoid worms (fig. 20). The animal contained within the tube is about
91 centims. long. There is an opening at the apex of the conical posterior extremity of
the body as in other species of the genus. The tentacles are disposed in two series. ‘Those
of the outer series are 5 millims. in length, and are, in the one perfect specimen available
for examination, 28 in number. Those of the inner series are about 1 millim. in length
and 14 in number. The upper part of the digestive cavity corresponds exactly to that
figured in Bronn’s ‘ Klassen und Ordnungen des Theirreichs,’ 2te Bd. Taf. vii. fig. 4,
after Haime, from Cerianthus membranaceus, and termed ‘“‘ Magensack;’’ but the digestive

* The madder colouring on both species of Corallimorphus here described was due to the substance 1 have termed
polyperythrin (Quart. Journ. Micros. Science, J. ¢.).

AND PELAGIC SURFACE-SWIMMING SPECIES. 303

cavity in C. bathymetricus does not terminate, as in C. membranaceus, above the line of
commencement of the mesenteries, but is continued downwards for some distance, its
walls consisting of a thin and delicate membrane, supported by the mesenteries and
having an irregular lower border. This lower sac is evidently in this case the true
stomach, since it was filled in the specimen examined with food, consisting of Globigerina-
shells together with small Crustacea. The main part of the mesenteries are not con-
tinued to the apex of the body, but only extend down about 3 millims., and from these
mesenteries are prolonged the ovaries, beyond the terminations of which is a clear space
of about 7 millims. in length. There is no canal formed of the opposed mesenteries and
leading from the mouth to the posterior part of the body-cavity ; but this is represented by
three or four stout muscular threads derived from the mesenteries, which are attached
to the inner surface of the body-wall near the posterior opening. These muscular
threads are branched superiorly, and thus each is derived from two or more mesenteries.
The animal was not examined until it had been put with its tube in spirit, under the
belief that it was an annelid, and its colouring, if it had any, was therefore destroyed.
Only two kinds of nematocysts were found in it, EAE portions of the epidermis,
the tentacles, and the test were examined.

The large, characteristic Cerianthus thread-cells, which form the main bulk of the
animal’s tube, resemble those of C. membranaceus in all respects except that the thread
is wound in an even spiral around the periphery of the internal cavity of the cell, the
axis of the spiral being at right angles to the longer diameter of the cell. In C. mem-
branaceus the thread is, as figured in Bronn (J. c. Taf. 1. fig. 7), irregularly doubled up
inside the cell. These cells are in C. bathymetricus ‘06 millim. long by ‘03 millim. broad.

The other kind of thread-cell observed was of the common elongated ovoidal form,
with the thread wound in a spiral about the longer axis of the cell (fig. 5). The
largest of these cells are of the same length as those forming the bulk of the test, but
they are very much more slender. They are not very abundant in the test, but occur
in large numbers on the tentacles. Thread-cells of all sizes and stages of development
are to be found in various regions of the body, but all of them are referable to these two
forms.

C. bathymetricus differs from the hitherto described species in the number of tentacles
arrangement of the digestive cavity, absence of a canal formed by the mesenteries,
the form of the thread-cells, and in its very small size. Two specimens were obtained
on June 19th 1873, in 2750 fathoms, in lat. 35° 29' N., long. 50° 53’ W., between the
Bermudas and the Azores.

GENERAL REMARKS.

The occurrence in the deep sea of representatives of well-known shallow-water forms
of Actiniaria is of considerable interest. The Hdwardsia here described seems to have
undergone no modification of any moment in a descent to 800 fathoms. The Ceri-
anthus, which is from the great depth of 2750 fathoms, is much dwarfed, and differs
in some details of its anatomy from the shallow-water forms; but it is astonishing
that the same genus should be able to adapt itself to shallow tropical seas and the ice-

SECOND SERIES.—ZOOLOGY, VOL. I. 28

304 MR. H. N. MOSELEY ON DEEP-SEA ACTINIARIA

cold water of the deep sea. A Cerianthus occurs at the Island of Mactan, opposite Zebu,
in the Philippines, which has a tube 1 foot 4 inches long, the animal itself being
6 inches in length. I found this Cerianthus abundant in water only 1 foot in depth at low
tide about the reefs, with its tentacles expanded, in the full glare of the tropical sun.
Yet a species of the same genus can exist at a depth of three miles, where no solar
light exists, and in water at a temperature close to freezing-point. .

The occurrence of a Nematoid worm in abundance in the substance of the tube of the
Cerianthus at this great depth is a very striking fact. The fact that some of the deep-
sea Actinias still retain a vivid colouring in the dark depths is also of great interest and
importance. I have dwelt on such facts and on the colouring-matters of deep-sea
animals generally elsewhere *. The usual absence of rocks or other objects offering a
wide surface for attachment seems to have led to the development of the peculiar elon-
gated base in Actinia abyssicola and A. gelatinosa, adapted to adhere to dead Alcyo-
narian stems, and to the hollowing out of the base of Corallimorphus for attachment to
small bodies, such as manganese nodules.

The genus Corallimorphus is of especial value, as appearing to be the nearest ally
amongst Actiniaria of many simple discoid corals, and for the excessively large size of
one form of its thread-cells or nematocysts ; these nematocysts seeming to be the largest
hitherto observed in any animal. There can be little doubt that Oceanactis is a pelagic,
and not a deep-sea form. Pelagic animals, such as Pyrosoma, Meduse, &c., were con-
stantly found in the trawl, caught by it on its way up or down, and mixed up with the
deep-sea animals from the botom. Pelagonemertes was obtained in this way.

DESCRIPTION OF PLATE XLV.

Fig. 1. Nautactis purpureus: side view, magnified. Actual height of the animal 7 millims.

Fig. 2. Enlarged view of portion of the disk of the same, to show arrangement of the colouring.

Fig. 3. Larval Minyad from the surface-net at the Philippines. Enlarged 15 diameters. Actual height
1:5 millim.

Fig. 4. Oceanactis rhododactylus. Twice natural size.

Fig. 5. Actinia abyssicola, attached to a Mopsia stem. Twice natural size.

Fig. 6. Actinia gelatinosa. The animal is shown of half the natural size. The line of continuation of
the rod to which the Actinia was attached within its sheath is shown by dotted lines: a, spot at
which the body doubled upon itself.

Fig. 7. Corallimorphus profundus. Side view, showing the attachment of the base to a small manganese
nodule.

Fig. 8. The same. Direct view from above of the disk. Figs. 7 and 8 both of natural size.

Fig. 8a. Nematocyst of same in quiescent, and 84 in active condition. x 510. Actual length of
cell 044 millim.

Fig. 9. Corallimorphus rigidus. Diagram of the disk, to show the arrangement of the tentacles and
their proportionate sizes.

* «On the Colouring-matters of various Animals, and especially of Deep-sea forms dredged by H.M.S. ‘Chal-
lenger.’” By H. N. Moseley. (Quart. Journ. of Microsc. Science, vol. xvii. p. 1, 1877.)

Fig.
Fig.
Fig.

AND PELAGIC SURFACE-SWIMMING SPECIES. 305

. 10. The same, seen in vertical section, to show the form of the oral disk, the mode of origin of the

tentacles, and peculiar cavity in the base. Figs. 9 and 10 both half natural size.

g. 11. Edwardsia coriacea, at rest, with the tentacles expanded. Twice the natural size.

. 12. The same, when completely contracted.

. 13. Portion of the disk, with three tentacles enlarged, to show the colouring.

. 14, Arrangement of the coloured streaks on the body-wall of the same, immediately beneath the

edge of the disk.

. 15. Cerianthus bathymetricus. Tube of the animal viewed intact, of natural size.
. 16. The same, tube laid open, to show the animal én situ.
. 17. The same, animal laid open and dissected. T, Outer tentacles; ¢, ner tentacles; or, mouth

and plaited upper region of the digestive tube; s, stomach, or lower portion of the digestive
tube, in which were found the remains of a crustacean ; m, mesenteries, from which proceed in
one region three muscular threads, which are prolonged to the apex of the body near the
posterior opening, where they are attached; g, generative glands.

18. Large nematocyst of same, and of the kind of which the tube or test is mainly composed.

19. Second form of nematocyst of same.

20. Small Nematoid parasite occurring in the wall of the tube.

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OF

THE LINNEAN SOCIETY

OF

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART THE SIXTH.

LONDON:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXVII.

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CONTENTS.

PART VI.—NovemBeER, 1877.

J v1. On the Birds collected by Professor J. B. Steere in the Philippine Archipelago.
By R. Bowpter Swarrt, F.L.S., 7.Z.8., &c., Senior Assistant, Zoological Depart-
ment, British Museum. (Plates XLVI-LIV.).... .. . . page 307

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[ 307 ]

XVI.—On the Birds collected by Professor J. B. Steere in the Philippine Archipelago.
By R. Bowpier Sarre, £.L.8., F.Z.S., §c., Senior Assistant, Zoological Depart-
ment, British Musewn.

(Plates XLVI.-LIV.)

Read November 16th, 1876.

ON his rettirn from the Philippines, Prof. Steere passed through London in the autumn
of 1875, and delighted me with an account of his travels in the Hast. At that time his
collections were packed up, and it was not possible to examine them ; but this year (1876)
he returned to England, and was so good as to place his birds in my hands for description.
This would have been more difficult to accomplish had it not been for the very elaborate
memoir published in the spring of last year by the Marquis of Tweeddale (then Viscount
Walden). Previously to this the ornithology of the Philippines was not concisely
described; and I must record my great obligations to his memoir on the birds of the
Philippine Archipelago. I must further acknowledge Lord Tweeddale’s kindness
in allowing me free access to his fine collection of Philippine birds, thus enabling me
to make comparisons of many rare and difficult species. He has seen all the specimens
obtained by Dr. Steere, and agrees with me as to the correctness of my determinations.
Knowing that Dr. Steere had visited several islands hitherto untrodden by the
naturalist, I was prepared to see several new species; but I was indeed surprised at the
large number of novelties which he has brought home with him. Forty new species
were discovered by him; and there is no doubt that even this number would have been
inereased but for the severe attacks of a fever which he contracted in Balabaec. As it is,
however, the results are most considerable, and Dr. Steere will receive the hearty
applause of all naturalists for the vigorous way in which he combated the difficulties of
climate and personal danger in pursuit of science, and reaped such an abundant harvest
in face of these trials. The danger from pirates alone may be estimated from the fact
that he collected many of his specimens in the company of native hunters and their
dogs—the latter being indispensable companions on an expedition, to give warnings
of the approach of piratical Malays. Again, in Basilan it was impossible even to
bathe under the shelter of the fort without keeping firearms at hand for personal

defence.
The following sketch of his expedition is extracted from an article communi-
cated by me to ‘Nature’ of the 3rd of August, 1876 :—‘ Leaving Hongkong for
Manila, in May 1874, Dr. Steere crossed the island of Luzon by way of Mauban and
Lucban to the Pacific, passmg some time on the mountain of Ma-hay-hay, near the
Laguna de Bay. In July he went by steamer to the colony of Puerto Princesa, on the
east side of the island of Palawan, where he stayed a month. Thence he crossed to
the island of Balabac and remained a month, afterwards visiting the south-east corner
of the island of Mindanao and resting for a month and a half at Zamboanga and the
SECOND SERIES, ZOOLOGY, VOL. I. 27

308 MR. R. B. SHARPE ON BIRDS COLLECTED BY

Indian village of Dumalon in the same province. The island of Basilan, lying between
Mindanao and the Sooloo group, was next visited; and here he stayed two weeks, after
which he returned to Zamboanga, and thence to Manila. In the month of December he
again went south, stopping at Tlo Ilo, on the island of Panay, and visiting the mountains
in the interior. After a short stay at the neighbouring island of Guimaras he crossed
over to Negros, journeying on horseback round the north end of the island; thence in a
native boat he traversed the sea to Zebu, which he crossed, till he arrived at the town of
the same name, where he took horse again and rode southward, crossing the island once
more and passing over the strait to the town of Dumaguete, on the island of Negros.
Dr. Steere then went back to Zebu and crossed to the island of Bohol ; after passing
round part of this island he returned to Zebu and afterwards to Manila, where he
visited the Negritos on the north side of the Bay of Manila, leaving finally in April for
Singapore.”

In the preparation of this paper I have followed, as far as possible, Lord Tweeddale’s
memoir, in order to make it supplementary to the latter; and, before commencing the
detailed account of the species, it may be as well to compare the results obtained by
Dr. Steere with those recorded by Lord Tweeddale in 1875.

The number of birds included by the last-named author was 219. From this total
I am only aware of one species that should be deducted, viz. Circus wruginosus. For
this error I myself am responsible, as I recorded in the ‘Catalogue of Birds’ a young
Marsh-Harrier, which I have since come to the conclusion is only an immature Circus
spilonotus (cf. Sharpe, ‘ Ibis,’ 1876, p. 30). Dr. Briiggemann has added a Pitta from
Luzon, in the Darmstadt Museum. :

The result of Dr. Steere’s expedition has been to add more than sixty species to the
avifauna of the Philippines; and by restoring to its position as a species Dicewm papuense
from Mindanao, and by adding the few birds known from the Sulu Islands, we have now
atotal of 286 Philippine species. The following remarks give some idea of the actual
alterations made in our knowledge of the different islands.

PHILIPPINE ISLANDS.

In the above-mentioned paper of Lord Tweeddale’s no less than 57 species are cata-
logued as being of Philippine origin, without any exact locality being known for any of
them beyond the above vague habitat. I am able to reduce this number by eight, as
follows :—

. Circus eruginosus is C. spilonotus, juv.

. Brachyurus erythrogaster is from Luzon (Mus. J. Gould).
. Zeocephus rufus is from Luzon (Mus. Brit.).

. Padda oryzivora is from Mindanao (Steere).

. Sturnia violacea is from Negros (Séeere).

. Calenas nicobarica is from Mindanao (Steere).

. Ortygometra cinerea is from Zebu (Steere).

. Gorsachius melanolophus is from Mindanao (Steere).

OTe oP CO DS

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 309

Many of the remaining 49 species are doubtfully Philippine, and I believe that a con-
siderable reduction will be ultimately made.

Inzon.—Dr. Steere remained so short a time in this island, and then only collected in
the neighbourhood of Manila, a locality already well worked, that he made no additions
to our knowledge of the ornithology of Luzon, the few birds obtained by him being
all well-known residents. Lord Tweeddale notices 133 species from Luzon. From this
total Totanus calidris must be deducted as being inserted in error, as it already appears
in the “ Philippine” column ; but, one species being omitted in counting up the total,
there remain still 133 species known from Luzon. Cataguan is treated by Lord Tweed-
dale as a separate locality; but it should really be considered as part of Luzon, as it is
situated on one of the irregular peninsulas of the south-eastern corner of the island.
This will enable us to add 8 species collected by Cuming at Cataguan, and now in the
British Museum, viz. Penelopides panini, Zeocephus rufus, and Puffinus leucomelas.
In addition to this, Mr. Gould’s collection contains the Red-breasted Pitta (Brachyurus
erythrogaster), from Manila, which can therefore be removed from the “ Philippine”
column to the Luzon list. The Pitta recently described by Dr. Briggemann as P. kochi
(Abhandl. Bremen, v. p. 65, pl. ii. fig. 6) makes the present number of species known
from Luzon 138.

Panay.—The number of species included by Lord Tweeddale was 9; and to these
Dr. Steere has added 22, making a present total of 31 species known from this island.
Of the acquisitions fowr are new, viz. Craniorrhinus Waldeni, Humyias panayensis,
Xantholestes panayensis, and Dicewm dorsale. The others are chiefly well-known
Philippine forms occurring to the notice of naturalists for the first time in Panay.

Guimaras.—To the 27 species collected by Mr. L. C. Layard and Dr. Meyer in this
island Dr. Steere has only added 4, viz. Philentoma albiventris, Dendrophila enochlamys,
Diceum hematostictum, and Cinnyris speratus. Of these, all but the last are new to
science. The total number of species now recorded from Guimaras is therefore 31.

Negros.—To the 38 species of birds recorded from this island by Lord Tweeddale,
Dr. Steere adds 21, the following being new species :—Oriolus Steerit, dithopyga mag-
nifica, Anthreptes chlorigaster, Phabotreron nigrorwm, the latter being the P. leucotis
of Lord Tweeddale’s list. The total number of species noticed from Negros is now 59.

Zebu.—Highteen species are enumerated by Lord Tweeddale; and: Dr. Steere only adds
one, Ortygometra cinerea, a well-known bird from the Malayan islands. It was stated
to occur in the Philippine archipelago; but hitherto no particular island had been
assigned as its exact habitat.

Camiguin.—This island was visited by Mr. Cuming; but I cannot say whether his
collections of birds made here were large, as, unfortunately, the bulk of his ornithological
consignments were dispersed without a complete record of their contents having been
made. I have discovered, however, two species from the present island in the Museum,
viz. Craniorrhinus leucocephalus and Brachyurus sordidus.

Mindanao.—Dr. Steere obtained 28 species in this island, in addition to the 19
included by Lord Tweeddale; to the latter should also be added Philentoma cyaniceps,
which is in the British Museum from Mindanao. The new species obtained by Dr. Steere

27 2

310 . MR. R. B. SHARPE ON BIRDS COLLECTED BY

were Brachywrus propinguus and B. Steer. This large Pitta, discovered near Zamboanga,
is one of the most interesting of Dr. Steere’s discoveries.

Basilan.—This island lies to the south of Zamboanga, and has apparently never before
been visited by a naturalist. 23 species were obtained by Dr. Steere; and of these
the following were new:—Chrysocolaptes maculiceps, Halcyon Winchell, Hypothymis
superciliaris, Setaria ruficauda, Zeocephus cinnamomeus, Macronus striaticeps, Dendro-
biastes basilanica, Irena melanochlamys, Hypsipetes rufigularis, Orthotomus cinereiceps,
O.frontalis, Diceum hypoleucum, Kudrepanis pulcherrima, Sarcophanops Steer. Con-
sidering that all the collecting done on this island was accomplished within a short
distance of the Spanish fort and settlement, it may be confidently hoped that a good deal
still remains to be discovered in Basilan.

Malamauwi.—Dr. Steere informs me that this little island is really nothing but a con-
tinuation of Basilan, from which it is separated only by a narrow channel about 100
yards across. This strait, however, is very deep, allowing a large ship to pass in; and it
is a favourite halting-place for the Spanish gun-boats. The distance between these two
islands is so small that doubtless the same species of birds are common to the two.
Malamaui is a low flat island, but very heavily wooded. Of the 8 species obtained by
Dr. Steere, only one, Swrniculus velutinus, did not occur in any other of the Philippine
Islands ; but three of the new species discovered on Basilan were also found on Malamaui.

Cujo.—Dr. Steere did not collect any birds here; so that at present we know only of
the 5 species recorded by Lord Tweeddale.

Palawan.—On this island Dr. Steere broke new ground; and of the 31 species obtained
by him the following were new :—Chrysocolaptes erythrocephalus, Zeocephus cyanescens,
Nixornis Woodi, Ptilocichla falcata, Phyllornis palawanensis, Irena Tweeddalii, Criniger
frater, Cittocincla nigra, Arachnothera dilutior, Aithopyga Shelleyi. Of the remaining
21 species recorded, 13 at least are well-known Bornean and Malayan forms, showing
the strong Bornean element in the ornithology of the island, as might almost have
been expected from its proximity to Borneo. At the same time the presence of
such birds as Tanygnathus luzoniensis, Leucocerca nigritorquis, and Brachywrus sordidus
(not B. Muelleri), shows great affinity to the true Philippine avifauna. We must wait
therefore for thorough exploration of the island before we can determine for certain its
zoological affinities ; but there can be little doubt that it holds an intermediate position
between Borneo and the other islands of the Philippine archipelago. !

- Balabac.—Dr. Steere was so prostrated with fever during his residence here that he
was unable to do as much work as he would have done had his health been good. He
met with only 9 species, two of which, Dryococcyx Harringtoni and Parus amabilis, were
new, and not found elsewhere. Most of the remaining ones are known from Borneo or
from the other islands of the Philippine group.

Sulu Islands—Myr. Hugh Low has been so good as to give me some birds which he
had obtained from Si Butu ; and although they consisted of only two species, they clearly
show the nature of the Sulu avifauna, and its close relations to that of the Philippines.
One of them is an apparently new species of Sarcops; and the other is a new Oriole
(Oriolus suluensis, Sharpe). As far as I am aware, only two other species, Pelargopsis

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 311

gigantea and Carpophaga Pickeringi, have been recorded from the Sulus; and the blockade
maintained by the Spaniards renders exploration in this archipelago difficult.

At p. 128 Lord Tweeddale alludes to a number of Indo-Malayan genera which are
wanting in the Philippines. The researches of Dr. Steere enable us to supply the
following—Anthreptes, Aithopyga, Arachnothera, Prionochilus, Mixornis, Macronus,
Phyllornis, Tora, Criniger, Buchanga, Hulabes, Tiga, Dendrophila. Besides these the
Eurylemidz are represented by a new genus, the Timeliidze by two, and the Cuculide
by one. All of these new genera are very closely related to well-known Indo-Malayan
forms. It should be noted, moreover, that Palawan, the Bornean character of whose
avifauna has already been alluded to, supplies us with most of the missing genera.

In the present paper 45 species are added, which are presumably peculiar to the
Philippines ; so that out of 285 species, 151 are peculiar (¢f Walden, J. ¢. p. 128).
Following Lord Tweeddale’s example, I give a table “‘ showing by families the proportion
of species peculiar to the Philippine Islands.”

sélses sélsek 38328
Families. S 2 3 By g Families. ee o8e Families. BE\ 8 3 "3
A | 4 oto AA oe ABA oS
Psittacides .......... 9 9 Mundi dsoenneteere 2 Gourideaeneeeeeeene 4
Falconide :......... 14 4 Rittidss) Sa ssnee tse 5 5 Phasianide ........ 1
Sirieidecsts Meee tte cao. 4 3 || Timeliide .......... 4 4 || Tetraonide.......... 2 1
Bicidee cities cy tas's0 9 7 || Crateropide ........ 2 INTIRACHD. co 50005000 1 1
Trogonide .......... 1 il Trenides oss ake 3 3 || Rollulide .......... 1 1
Meropide .......... 2 1 Brachypodide ...... 8 6 || Megapodide ........| 2 1
Coraciide .......... 1 Saxicolidease eee 6 3 || Charadride ........| 8
Alcedinide.......... | 15 G) ||| Siyllvattebsis Gon6occc5n 10 6 || Glareolide.......... 1
Capitonide.......... 2 Motacillide ........ 5 1 Gallinulide ........ 4 1
Cypselide .......... 2 er RaridasnesraSerecy ta ae 4 3 || Rallide ............ 8 2
Caprimulgide........ 3 3 || Meliphagide ........ 1 Dens || eParridse espe a seu 1
Cuculide .......... 12) 11 Nectariniide ........ 16/ 13 Scolopacide ........ 9
Bucerotide.......... 5 5 Certhiide .......... 1 1 Ciconidee .......... 1
amit dea ys 22 heer er cncrste 3 1 Conyideeanrreeie see il 1 ISRUISGED 5 5 50 Ganac0 oc 11 1
Artamide .......... il SUIICED oooce cose 7 3 PSTERONGEDS 3.5 oe Cac oee 4 2
Campephagide ...... 8 4 || Kurylemide ........ 1 1 Procellariide ........ 1
Dicruride .......... 3 Py }| VsaiarelGE}e5 Sooecaes 1 IGAGES Gade apoolee 5
Muscicapide ........| 14 | 11 IRMETCED.6 oadcooccas 6 | odicipidear ye ene 1 il
Hirundinide ........ 3 Treronidz mere 13 9 Pelecanide.......... 6
OME sosccoscadut 4 4 || Columbide.......... 4 3) | THERES. 5 cong edso ox 1

I now proceed to a detailed catalogue of Dr. Steere’s collection; and in this I follow
almost entirely the arrangement adopted by Lord Tweeddale, though in some few
instances not agreeing with the order of the different families: this, however, is a small
matter, my object being to render the presen paper as far as possible a supplement to
the memoir above mentioned.

312 MR. R. B. SHARPE ON BIRDS COLLECTED BY

PSITTACT.
PLYCTOLOPHID.
Cacatua, Vieill.

1. CaAcATUA HH{MATUROPYGIA (Mill.): Walden, Tr. Z. S. ix. p. 132.
a. 2. San Bernardino, Negros, Jan. 1875. Eyes pink.
[I saw a small species of Cockatoo in Balabac apparently of the same kind as the _

Negros bird; but, owing to a severe attack of fever, the three I shot were not skinned.
In Palawan these birds are also plentiful.—J. B. 8. |

PSITTACID A.
Prioniturus, Wagler.

2. PRIONITURUS DISCURUS (Vieill.): Walden, ¢. ¢. p. 132.

a. & ad. Isabella de Basilan. Beak nearly white; iris grey.
b. 2 juv. San Bernardino, Negros, Jan. 1875. Eyes dark.
c. juv. Balabac, Sept. 1874.

[In little flocks near San Bernardino, among the wild plantains. Owing to their
similarity of colouring, they were not easy to see among the foliage.—J. B. 8S. ]

T have carefully examined the above specimens, and have compared them with Lord
Tweeddale’s series. I can recognize but one species, although great difference exists in
the form of the middle tail-feathers. The one from Basilan is an adult, with a long
shaft and broad racket, whereas the other two have these plumes only slightly elongated,
and the spatulee a mere widening of the attenuated apex of the feather. The species was
not before known from any of the above-mentioned islands.

TANYGNATHUS, Wagler.
3. TANYGNATHUS LUZONIENSIS (L.): Walden, ¢. ¢. p. 133.

a. Puerto Princesa, Palawan.

6. 2. Dumalon, Mindanao, Oct. 1874. Eyes nearly white.

c. 9. San Bernardino, Negros, Jan. 1875. yes light yellow.
d. Negros.

[In flocks near Dumalon, coming into the fruit-trees in the town. The natives eat
them.—J. B. 8.]

Negros was known as a habitat for the species, it having been obtained by Mr. L. C.
Layard; but Palawan and Mindanao are both new localities for it.

Cyctorsitra, Hombr. & Jacquinot.
4, CYCLOPSITTA LUNULATA (Scop.): Walden, é. c. p. 133.
a. Manila.
Only one specimen collected, and that without the scale-like collar.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 313

Loricuxts, Blyth.

5. LoricuLus curysonotvs, Scl.: Walden, ¢.¢. p. 137.
a. Zebu.
Agrees with Zebu (typical) examples in Lord Tweeddale’s collection.

6. LoRIcULUS REGULUS, Souancé: Walden, z. ¢. p. 135.
a. Panay.

[The native names for all these little Parrakeets are Cust and Culacissi, both being
derived from the curious whistle of the birds. They seem to reside amongst the cocoa-
nut palms, and are frequently captured and kept by the natives.—J. B. 8.].

Lord Tweeddale and I carefully compared Dr. Steere’s specimen, and came to the
conclusion that it was an individual of L. regulus not adult. Panay is therefore a new
habitat for the species.

ACCIPITRES.
FALCONIDA.
ACCIPITRIN &.

ACCIPITER, Briss.

7. ACCIPITER ViRGATUS (Temm.): Sharpe, Cat. B. i. p.151. Teraspiza virgata, Walden,
t.c. p. 141.
a. ad. Negros.

Tn rich adult plumage with red breast. It is in all likelihood a male, as it has the fol-
lowing measurements :—Total length 11:8 inches, culmen 0°85, wings 6:9, tail 5-5, tarsus
21. The above dimensions, however, do not tally with those of either sex given in my
Catalogue; and it is probable that a further revision may prove that more than one race
is included under the title of A. virgatus. Negros is a new habitat for this Sparrow-
Hawk.

AQUILIN A.
SPILORNIS, G. R. Gray.

8. SPILORNIS HOLOSPILUS (Vigors): Walden, 7. c. p. 142.
a. ad. Mount Ma-hay-hay, near Manila.

Burastur, Hodgs.

9. BurastuR iInpicus (Gm.): Walden, ¢. ¢. p. 143.
a. 3 juv. Zamboanga, Mindanao. LHyes greenish yellow.
6. ad. Panay.
This species, a migrant from the China coast, visiting the Moluccas in winter, is now
first recorded from the above two islands.

314 MR. R. B. SHARPE ON BIRDS COLLECTED BY

PICARLA.

PICIDA.
THRIPONAX, Cabanis.
10. THRIPONAX JAVENSIS (Horsf.): Walden, 7. c¢. p. 146.
a. g. Zamboanga. LHyes yellow.
6. 9. San Bernardino, Negros, Jan. 1875. Eyes yellow.
c. 2. Negros.
d. 9. Puerto Princesa, Palawan.
[Climbing about the dead trees, generally in pairs.—J. B.8.]
I agree with Lord Tweeddale that we cannot separate the Philippine Black Wood-
pecker from the Javan bird. It was not known before from Palawan and Negros, but
had been recorded from Mindanao by Wagler.

CHRYSOCOLAPTES, Blyth.
11. CHRYSOCOLAPTES XANTHOCEPHALUS, Walden & Layard: Walden, ¢.c. p. 147.
a. 6. San Bernardino, Negros, Jan. 1875. yes carmine.

[I saw more of this Woodpecker, which was not rare; but I had the greatest difficulty
in getting about the forests in Negros, owing to the obstruction caused by the trees
blown down in a recent typhoon.— J. B.S. ]

This species was originally described from the island of Negros; but only the female
was known. Dr. Steere has now procured the male, which shows that the species is an
excellent one. The description of the male is as follows :—

Adult male-—General colour above ruddy crimson, all the feathers olive-brown at the
base, the upper tail-coverts entirely of the latter colour, with a crimson mark on their
margins; scapulars and wing-coverts exactly resembling the back; the bastard wing,
primary-coverts, and quills brown, externally yellowish olive, the secondaries brown,
inclining to olive on the outer webs, which are externally crimson like the back, all the
quills spotted with white on the inner web, much more largely on the secondaries; tail-
feathers dark brown, with a slight wash of olive on the outer webs; crown of head bright
crimson, with yellow bases to the feathers ; lores, eyebrow, and entire sides of face bright
yellow, as also the throat, which is bordered on each side with a narrow black mousta-
chial line, and has also another down the centre of the throat; sides of neck yellow, with
three lines of black running down from behind the ear-coverts; neck all round, and
fore neck, scaly in appearance, the feathers being yellowish buff fringed with black ;
remainder of under surface yellowish buff, the sides of the body somewhat streaked with
olive-brown; under wing-coverts dull fulvous, with olive-brown margins to the feathers,
which are also slightly washed with red ; bill stone-brown (in skin) ; legs yellowish, claws
black; “iris carmine’’ (S¢eere). Total length 10°5 inches, culmen 1°6, wing 5:6, tail
3°8, tarsus 1:05,

12. CHRYSOCOLAPTES MACULICEPS, sp. n. (Plate XLVI. fig. 2.)
Ad. supra coccineus, plumis ad basin olivascentibus et celatim albo maculatis; tectri-

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 315

cibus alarum dorso concoloribus nec maculatis; tectricibus primariorum nigrican-
tibus extus olivascenti-aureis; remigibus nigricantibus interne albo ovatim macu-
latis, extus olivascenti-aureis gradatim coccineis; supracaudalibus nigris, versus
apicem olivaceis et fulvo maculatis; cauda nigricanti-brunnea olivaceo lavata, rec-
tricibus externis basaliter fulvo maculatis ; pileo summo olivascenti-brunneo, fulvido
transversim maculato, coccineo adumbrato, maculis fulvis hoc colore nitentibus;
loris et facie laterali fulvescenti-flavis, regione parotica postica genisque anguste
nigro striatis; gula etiam fulva, vitta malari distincta nigra; collo undique marmo-
rato, plumis lete fulvis nigro marginatis, torquem collarem squamosam forman-
tibus; corpore reliquo subtus ochrascenti-fulvo, plumis pectoralibus nigro margi-
natis, reliquis nigro irregulariter transfasciatis; subalaribus pectori concoloribus ;
rostro corneo, mandibula ad basin flavicante; iride coccinea. Long. tot. 10, culm.
1:55, alee 5°55, caudee 3°55, tarsi 1:05.
a. Isabella de Basilan, Oct. 1874. Eyes carmine.

The nearest ally to this species is C. hematribon; but the Basilan bird differs in its
paler red and more spotted head, buff-coloured ear-coverts, spotted mantle and upper
tail-coverts. Dr. Steere marked on his label that one bird was “full-grown,” not being
young, as it might have been supposed to be.

13. CHRYSOCOLAPTES ERYTHROCEPHALUS, sp. n. (Plate XLVI. fig. 1.)

Ad. supra auratus vix coccineo lavatus; remigibus brunneis extus auratis, secundariis et
tectricibus alarum interscapulio concoloribus; pileo cristato undique coccineo;
gutture paullo pallidiore, vitta mediana angusta nigra, altera utrinque malari an-
gusta nigra; preepectore et torque collari ochrascenti-fulvo, plumis nigro anguste
circumdatis ; corpore reliquo subtus fulvescenti-albo, nigro variegato, plumis an-
guste nigro marginatis, hypochondriis et subcaudalibus irregulariter nigro transfasci-
atis; axillaribus albidis nigro maculatis; subalaribus nigris albo ovatim maculatis,
margine alari aureo lavato; supracaudalibus nigricantibus, olivaceo terminaliter
lavatis; cauda nigra; rostro virescenti-corneo; pedibus saturate plumbeis; iride
coccinea. Long. tot. 10-0, culm. 1°7, alee 5°65, caudee 3°4, tarsi 1:15.

a. 6 ad. Puerto Princesa, Palawan.
This species appears to be distinguished from all the other members of the genus by its
red throat and head.
Trea, Kaup.
14. Ties JAVANENSIS (Ljungh): Salvad. Ucc. Born. p. 55.
a. §- Puerto Princesa, Palawan. Lyes black.
fam unwilling on the strength of one specimen only to institute a new species; but
certainly this three-toed Woodpecker differs from all the others with which I have com-
pared it in the uniform sides of the fore neck, which are brown, like the rest of the
under surface, the latter being barred across. It appears to agree best with a Bali bird
obtained by Mr. Wallace, and is probably the same in worn plumage.
SECOND SERTES.—ZOOOLGY, VOL. T. 2uU

316 MR. R. B. SHARPE ON BIRDS COLLECTED BY

TROGONID A.

HARPACTES, Swains.

15. HARPACTES ARDENS (Temm.): Walden, ¢.c. p. 149.
a. 2. Isabella de Basilan.
[I found plenty of Trogons in the forests some little distance inland, but I only suc-
ceeded in preserving one female bird.—J. B. 8.]
The single bird obtained being a female renders it difficult to identify the species ; but
as 7. ardens is known from Mindanao, it is possible that it extends also into Basilan.

MEROPIDA.
Meroes, Linnzeus.

16. Mrrors pHinippinus, L.: Walden, ¢. ¢. p. 149.

a,b. Negros.
c. Panay.

d. Dumalon, Mindanao, Oct. 1874. yes red.

e. Zamboanga, Mindanao.

[At Zamboanga I found them flying over a wet rice-field taking dragon-flies on the
wing, and returning to stakes to- devour them. In the other islands I saw them in the
open.—J. B. 8.]

Panay and Mindanao are new habitats for this species.

17. Mzrors Bicotor, Bodd.: Walden, ¢. c. p. 150, pl. xxvi. fig. 1.

a, b. Panay.
c. 2. Dumalon, Mindanao, Oct. 1874. Eyes carmine.

[Found in swamps, perching in the low trees at Dumalon. In Panay it was shot
more in the open, settling on weeds and sticks a little way from the ground.—J. B. S.]
Not known before from either of the above-mentioned islands.

CORACIIDA.

Eurystomvs, Vieillot.

18. EURYSTOMUS ORTENTALIS (L.): Walden, @. c. p. 152.
| a, b, c. Dumalon, Mindanao.
d, e. . San Bernardino, Negros, Jan 1875. Eyes hazel.
{Found in abundance in the Indian village of Dumalon, in the island of Mindanao, as
well as in Negros. In the former place they were perched on limbs of dead trees in the
edgings of clearings; and in Negros they were seen along the sides of the streams on

limbs overhanging the water.—J. B. 8.]
Although recorded from Negros, Mindanao is a new locality for the species.

PROF, STEERE IN THE PHILIPPINE ARCHIPELAGO. 317

ALCEDINID A.
ALCEDININE.
Atcrpo, Linneeus.

19. ALCEDO BENGALENSIS, Gm.: Walden, ¢. c. p. 152.
a. juv. San Bernardino, Negros, Jan. 1875. Eyes dark.
6. $. Dumalon, Mindanao. Eyes hazel.
[Very generally distributed throughout the Philippines, and found on nearly every

stream.—J. B.S. ]
_ Previously to Dr. Steere’s visit, we only knew of this species from Luzon.

PELARGOPSIS, Gloger.
20. PELARGOPSIS LEUCOCEPHALA (Gm.): Sharpe, Monogr. Alced. pl. 31.
a. 6. Puerto Princesa, Palawan, July 1874.
_ [J found this species in mangrove swamps in the island of Palawan.—J. B. 8.]
The above specimen belongs to P. lewcocephala of Borneo, not P. Gouldi, which is
probably confined to Luzon.

21. PELARGOPSIS GIGANTEA, Walden, Ann. N. H. (4) xiv. p. 123 (1874): Meyer, ¢. ec.
pp. 401, 475.
a. Dumalon, Mindanao.

[Frequently seen flying along the sea-beach, fishing over the open sea, and perching
upon the low trees on the sandbars.—J. B. S.]

Lord Tweeddale described this distinct species from Sulu in the Sulu archipelago, where
Dr. A. B. Meyer obtained it; and now Dr. Steere has procured it in Mindanao. Itisa
representative of P. melanorhyncha of Celebes, but has a reddish bill, like P. leucoce-
phala &c. I compared the Mindanao bird with Lord Tweeddale’s type specimen.

Cryx, Lacépéde.

22. CEYX MELANURA, Kaup: Walden, ¢. ¢. p. 153.
a. 6. Dumalon, Mindanao. LHyes hazel. |

[I only saw the one specimen of this pretty Kingfisher, in deep thick jungle, perching
low. Its food consisted of small crabs.—J. B. 8. |

The single bird collected by Dr. Steere differs from every other Philippine specimen
which I have examined in having the wing-coverts black with rufous edgings, whereas
all the others which have fallen under my notice have had them black spotted with
blue; and Lord Tweeddale has a nestling resembling the adult in this respect. At the
same time I do not like to found a new species on a single example.

HALcyon, Swainson.

23. Hancyon @uLARIS (Kuhl): Sharpe, Monogr. Alced, pl. 70. Entomobia gularis

(Kuhl), Walden, ¢. ¢. p. 154.
2U 2

3818 MR. R. B. SHARPE ON BIRDS COLLECTED BY

a. 6. Maasin, Panay, Jan. 1875. yes dark.
b,c. 3; d. 2. San Bernardino, Negros, 1875. Eyes dark.

[There is scarcely any difference in the sexes; but the female seems a little lighter
below, and to have rather a more greenish tinge on the scapulars, back, and tail.

I noticed this species along the streams, sometimes fishing, but as often feeding on
the ground, where its diet seemed to consist of earth-worms. I saw this, or a closely
allied species, near Zamboanga, in Mindanac, but failed to secure it.—J. B. S. |

Mr. Layard obtained this Kingfisher in Negros; but Panay is a new locality for it.

24. Hatcyon HomBRONI, Bp.: Sharpe, Monog. pl. 115. Actenoides hombroni, Walden,
t.c. p. 155.
a. 3. Dumalon, Mindanao. Eyes hazel.

[Only met with on one occasion, in low thick jungle which was covered every day by
the sea.—J. B. 8.]

This Kingfisher is only known from the original type specimen in the Paris Museum,
which was obtained in Mindanao by Messrs. Hombron and Jacquinot. I have never
seen a skin before; and the plate in my ‘Monograph’ was drawn from a coloured
sketch sent over to me from Paris. I therefore give a full description of the adult
male in Dr. Steere’s collection :—

Adult male.—Head and nape bright blue, more brilliant on the sides of the head,
over the eye, and on the nape; round the latter a narrow line of deep black; ear-
coverts tawny chestnut; along the lower line of the lores a streak of black feathers,
reaching below the eye, and widening behind the latter, being here shaded with blue;
cheeks bright blue, forming a broad band; sides of neck and hinder part of the latter
deep tawny, varied with narrow black edgings to the feathers; mantle blackish, mottled
with tawny spots, these being subterminal with a narrow black frmge; middle of back
and scapulars and wing-coverts green with a slight shade of verditer, each feather
having a distinct subterminal spot of ochraceous buff; quills blackish, externally washed
with greenish, the primaries edged with ochraceous, the secondaries with the same sub-
terminal spot of ochre as on the wing-coverts; lower back, rump, and upper tail-coverts
bright silvery cobalt, the sides of the back and the lateral coverts blackish washed with
blue; tail-feathers deep blue, with black shafts; throat white, slightly washed with
tawny; rest of the under surface deep tawny, whiter on the centre of the abdomen; the
breast-feathers with narrow nearly obsolete blackish margins; thighs externally black-
ish, internally deep tawny; feathers at side of vent, adjoining sides of lower back, deep
blue, the outer web more or less ochraceous; under wing-coverts and axillaries deep
tawny ; the quills blackish below, edged with pale tawny-buff along the inner web; bill
coral-red, the culmen black (in skin); iris hazel. Total length 11°83 inches, culmen 2-0,
wing 4°95, tail 4°15, tarsus 0°75.

25. Haucyon WINCHELLI, sp.n. (Plate XLVII.)

9 ad. supra nigra, purpureo lavata; dorso postico, uropygio et supracaudalibus argenteo-

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 319

cyaneis, his purpurascentibus;. loris et torque collari postico castaneo-fulvis ;
pileo nigro, lateraliter cyaneo, supercilium latum ab oculo summo usque ad
nucham productum exhibente; facie laterali cum regione parotica et colli lateribus
nigris purpureo vix lavatis; alis nigris, vix extus purpurascenti lavatis; cauda
purpurascenti-nigra ; genis et corpore subtus leete cervino-fulvis, abdomine albicante ;
subalaribus et remigibus intus cervinis; rostro nigro, ad basin mandibulze carnoso ;
iride brunnea. Long. tot. 10-0, culm. 2°05, alee 4:1, caudee 3:2, tarsi 0°55.

a. ©. Basilan (Isabella). Eyes hazel.
[This Kingfisher inhabits the forest; and its food consists of beetles, dragon-flies, Ke.
I met with it at a long distance from water.—J. B. S.]
This species is named, by Dr. Steere’s request, after his friend and old tutor, Mr.
Winchell.

26, Hatcyon cHoris (Bodd.): Sharpe, Monogr. pl. 87. Sauropatis chloris, Walden,
t. c. p. 155.
a.ad. Negros.
6. 2 juv. Dumalon, Mindanao, Oct. 1874. Eyes dark.
ec. g ad. Isabella de Basilan.

{General in the islands of Mindanao, Panay, Negros, and Zebu, perching on low
trees, and searching for worms on the ground; they also fish occasionally. They
usually frequent cane-fields and open plains, perching on low timber.—J. B. S.]

Not before recorded from Mindanao, nor from Basilan.

CYPSELID.
Macropreryx, Swainson.
27. MAcropTERYx comATus (‘Temm.): Walden, ¢. c. p. 158.
a, 6. Negros.
c,d,e. 6. Dumalon. Eyes dark.

[In great abundance along the sea-beach at Dumalon, sitting towards evening in
clusters on the limb of a tree. In Negros I shot the pair right in the woods.—
eal bears

The birds from Mindanao belong to the small race, or true WZ. comatus of Tem-
minck, and agree in their measurements with Bornean examples. The specimens from
Negros, on the contrary, are much larger, as Lord Walden found to be the case with
the Luzon birds. The following measurements exhibit these differences in size :—

Long. tot. ale. caude. rect. ext. tarsi.
a. 6 ad. Negros (Steere) . . . . 6:6 5:45 2:0 315 0.3
boRads 5 x Speer en 6:4 5°35 2:0 315 0:3
ce. g ad. Dumalon ,, <i 5:8 5-1 2-0 2°95 0°25
dG ads)" ,, 35 = TORS 5:8 4:8 1:8 3°0 0°25
e. ¢ ad. 5 5 ffs) cee 6:0 4°95 18 29 0°25
ip & ath Ibelonern (hoy) 6 Give ss 6:0 4-9 16 30 0°25

qed re a 61 4:95 1:85 31 0°25

320 MR. R. B. SHARPE ON BIRDS COLLECTED BY

Contocata, G. R. Gray.
28. CoLLOCALIA TROGLODYTES, Gray: Walden, ¢. c. p. 158.
a. Panay.

[Shot at some distance from the sea, in the hills.—J. B. 8.

Compared with the typical specimens, the Panay bird is more blue above, with a
purplish gloss on the wings. Underneath the throat and breast are washed with silvery
white. The types are much browner, with very little purple gloss above, and scarcely
any of the silvery white below. They are also a little larger (wing 3°65-3:75 inches).
These differences, however, are only due to age, Dr. Steere’s Specimens being older;
the measurements are as follows—total length 3:7 inches, culmen 0:15, wing 3°65,
tail 1-65, tarsus 0°35. Y

The two specimens in the Museum have no exact locality to them; for they are
registered as from “ Malacca,” at the end of a Philippine collection, both received from
the late Mr. Cuming. It is probable that they are really from the Philippines; but
there is no proof of this.

CUCULID®.
CUCULIN &.
CAcoMANTIS, S. Miller.

29. CACOMANTIS MERULINUS (Scop.): Walden, ¢.c. p. 160.

a. 3. Malamaui, Oct. 1874. Eyes and inside of mouth cherry-red, legs yellow.
b. 2. Maasin, Panay, Jan. 1875. Eyes grey.

Originally described by Sonnerat from Panay, and recorded by Prof. Schlegel from
Luzon (Govers). It also occurs in the southern Philippines, as Dr. Steere shot it in
Malamaui; and it is probably found throughout the archipelago.

Curysococcyx, Boie.

30. CHRysococcyx BASALIS (Horsf.): Salvad. Ucc. Born. p. 62.

a, Negros.

Agrees with a Flores specimen collected by Mr. Wallace. Total length 5-7 inches,
culmen 0°55, wing 3°7, tail 2°6, tarsus 0°6.
Eupynamis, Vigors and Horsfield.

31. HUDYNAMIS MINDANENSIS (L.): Walden, J. ¢. p. 162.

a. 2. San Bernardino, Negros, Jan. 1875. Eyes carmine.

[I found this in thickets of low trees about the sugar-cane fields and along a little
river near San Bernardino.—J. B. 8. ]
The specimen collected is in the rufous and black plumage.

SuRNIcULUS, Lesson.
32. SURNICULUS VELUTINUS, Sp. 0.
3 ad. supra velutino-nigra vix purpureo lavata; scapularibus alisque metallice viri-

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 321

descenti-nigris vix chalybeo nitentibus ; uropygio, supracaudalibus caudaque metal-
lice virescenti-nigris, illis chalybeo nitentibus; facie laterali et corpore subtus
velutino-nigris haud metallicis; tibiis niveis, plumis longis obtectentibus externe
albis, interne nigris; remigibus subtus nigris, albo transfasciatis, pogonio interno
albo ovatim maculato fasciam unicam formante; rostro et pedibus nigris, his subtus
flavis; iride saturate brunnea. Long. tot. 8°3, culm. 0°8, ale 4°55, caude 4:1,
tarsi 0°65.
a. 2. Malamaui, October 1874. Eyes dark.

This pretty little Cuckoo is closely allied to S. dugubris (Horsf.), but is at once to be
distinguished by the velvety black colour of the upper and under surface, and by the
absence of metallic gloss on these parts, what little colour there is being of a purplish
shade. The size is also somewhat smaller.

PHENICOPH HINA.
DASYLOPHUS, Swainson.

33. DASYLOPHUS SUPERCILIOSUS (Drap.): Walden, ¢. c. p. 162.
a. 3g. Mount Ma-hay-hay, near Manila.

[Only found in Luzon, and never seen by me in any of the other islands. It was
plentiful in the neighbourhood at the foot of Ma-hay-hay in the thickets about the
villages—J. B. 8. ]

Dryococcyx, gen. n.

Closely resembling Rhinococcyx, Sharpe (P. Z. 8. 1873, p. 604), but distinguished at
a glance by the shape of the nostril, which is a small circular hole situated in a deep
perpendicular groove which separates the lores from the upper mandible.

Fig. 1.

Profile of Dryococcya Harringtoni.

34. Dryococcyx HARRINGTONI, sp. n.

¢ ad. supra metallice viridis vix olivascens; alis dorso concoloribus, primariis extus
chalybeis ; cauda viridi, terminaliter 4 castanea, rectricibus externis 3 castaneis;
capite summo olivascenti-brunneo; collo postico castaneo lavato, torquem collarem
vix formante; fronte et genis anticis mentoque cinereo lavatis; facie laterali et
corpore subtus toto lete rufis, abdomine et subcaudalibus castaneis; regione oculari
nuda coccinea; rostro pallide viridi; iride coccinea. Long. tot. 18, culm. 1°65, ale
6°35, caudee 9°3, tarsi 1°55.

a. 3. Balabac, September 1874. Eyes currant-red, as well as the excrescences on the face.

322 MR. R. B. SHARPE ON BIRDS COLLECTED BY

[I saw others of this species when I reached the jungle on the hill-sides during an
excursion I made into the mountains.—J. B. 8. |

This new form of Coucal is most nearly allied to Rhinococcyx curvirostris, from which
it differs not only in the structural peculiarities of the nostrils but also in colouring.
It has an olive-brown head and partial rufous collar round the neck, and has none of the
greenish black colour on the abdomen and thighs, these being rufous like the breast.
By Prof. Steere’s desire I have named this species after Mr. Harrington, Professor of
Botany in the University of Michigan, and an old schoolfellow and college friend of the
discoverer’s.

CENTROPODINS.
CEnTROCOCCYX, Cabanis and Heine.

35. CENTROCOCCYX VIRIDIS (Scop.): Walden, ¢. c. p. 168.

a. Zamboanga, Mindanao.

PyrRHocentorR, Cabanis and Heine.
36. PyYRRHOCENTOR MELANOPS (Less.): Walden, ¢.c. p. 164.
a. 3. Malamawi, October 1874. yes cherry-red.

BUCEROTID.
Buceros, Linnzeus.
37. BucrRos HyDROcORAX, L.: Walden, t. c. p. 164.
a. juv. Head only. Mount Ma-hay-hay, Manila.
(This Hornbill was plentiful on the above-named mountains.—J. B. S.]

CRANIORRHINUS, Cabanis and Heine.
38. CRANIORRHINUS LEUCOCEPHALUS (V.): Walden, ¢. ¢. p. 165, pl. xxvii. figs. 1, 2.

a. Zamboanga, Mindanao.

39. CRANIORRHINUS WALDENI, Sharpe, Journ. Linn. Soe. Zool. vol. xiii. p. 155; Elliot,
Monogr. Bucerot. part 1.

Ad. supra niger, ubique metallice olivascenti-viridi nitens; alis dorso concoloribus ;
cauda fulva, rectricibus quatuor intermediis pallide castaneis, omnibus late nigro
terminatis et dimidio basali nigro; pileo et collo undique saturate castaneis, plumis
supraocularibus et regione parotica antica fulvescentibus ; regione periophthalmica,
genis et gutture toto nudis; corpore reliquo subtus nigro viridi nitente; rostro
rubro, mandibula et crista rostri maxime sulcatis. Long. tot. 28, alee 14°6, caudze
10°8, tarsi 2:1.

a. 3. Panay.

[I shot this Hornbill on the highest ridge of the mountains west of Ilo-Ilo. This is
the only place where any of the virgin forest is left; and here alone I saw these birds.
They were not very rare; but I could only get a single specimen, as they flew so high in
the trees that ry gun could scarcely reach them.—J. B. 8.]

PROF, STEERE IN THE PHILIPPINE ARCHIPELAGO, 328

PENELOPIDES, Reichenbach.
40, PENELOPIDES PANINI (Bodd.): Walden, ¢.c¢. p. 166, pl. xxviii. figs. 1, 2.

a. Negros.

PASSERES.
LANIIDA.

Lantus, Linneeus.
41. Lantus tuctonensis, L.: Wald. ¢. ¢. p. 171, pl. xxix. fig. 1,

a. Adult. Panay.
6. ¢ imm. Basilan (Isabella). Eyes hazel.

[In the cultivated country of Panay, in the lower country. In Basilan they were
found about the clearing in the vicinity of the fort at Isabella.—J. B. S.]

Dr. Steere adds two more islands to the known Philippine habitats of this species ;
but, as this Shrike is a migrant from China, it probably visits every one of the group.
Mr. Low and Mr. Everett have lately discovered it in North-western Borneo.

ARTAMIDA.
ARTAMUS, Vieillot.

42. ARTAMUS LEUCORHYNUSs (L.) : Walden, ¢.c. p. 174,
a. Dumalon, Mindanao, October 1874.
[Very plentiful at Dumalon and also in Negros, flying much like Swallows over open
ground, and lighting upon stumps.—J. B. 8.]
This Swallow-shrike was not before known from Mindanao.

CAMPEPHAGIDA.
GRAUCALUS, Cuvier,

43, GRAUCALUS SUMATRENSIS, Mill. ; Salvad. Uce. Born. p, 150.
a, 6. Puerto Princesa, Palawan.
[Found in similar situations to G. striatus.—J. B. 8.]
This is a well-known Bornean and Sumatran bird; and its occurrence in Palawan is
one of the evidences of a Bornean element in that island.

44, GRAUCALUS STRIATUS (Bodd.): Walden, ¢. ¢. p. 175, pl. xxx. fig. 1.

a. 3. San Bernardino, Negros, Jan. 1875. yes reddish.
6. 3. Malamaui. Eyes light yellow, nearly white.
c. Zamboanga, Mindanao.
[The difference in the colour of the eyes in specimens a, 0 is curious, as they are nearly
identical in plumage. ‘They were found in low trees in oper country.—J. B. 8.]
A very full account of this species is given by Lord Tweeddale (/. ¢.), describing all
the changes of plumage. Malamaui is a new locality.
SECOND SERIES.—ZOOLOGY, VOL. I. 2X

o24 MR. R. B. SHARPE ON BIRDS COLLECTED BY

LALAG#, Boie.

A5, LALAGE poMINIcA (Miuill.): Walden, 7. c. p. 178.

a. Panay.
b. ¢. Dumalon, Mindanao. Hyes dark.
c. 9. Malamaui, Oct. 1874. Eyes dark.
This species was added to the Philippine avifauna by Dr. Meyer, who met with it in
Zebu and Guimaras. All the islands mentioned above are new localities for it.

Prricrocotes, Boie.
46. PpricrocoTus 1GNEUS, Blyth: Sharpe, Str. F. 1876, p. 209.

a,b,c. ¢ ad. Puerto Princesa, Palawan. Eyes black.
d,e. 2 juv. Puerto Princesa.

- [The first specimen I shot in a mangrove tree; and the other four I killed from the door
of my house as they were eating fruit on a small clearing. The iris in all was black.—
J. B.S.)

This species of Pericrocotus is known from Malacca and Borneo, being very common
in the north-western part of the latter island. The island of Palawan is therefore an
entirely new locality for it. Prof..Steere’s series comprise a pair of fully adult males,
a third in full plumage, excepting a few yellow feathers on the throat, and two young
males. One of the latter is in the dress of adult P. flagrans, showing that at first the
young male resembles the old females; for there can be no doubt that P. flagrans is only
the hen of P. zgneus.

DICRURIDA,

Dicrurvs, Vieillot.

47. DickURUS MIRABILIS, Walden: id. ¢. ¢. p. 181.

a. 6. San Bernardino, Negros, Jan. 1875.
6, Negros.

[Always in low thickets, and feeding on fruit. Rather hard to shoot, owing to their shy
habits.—J. B.8.] geen: .
BucHanea, Hodgson.
48, BUCHANGA CINERACEA (Horsf.): Sharpe, Cat. of Birds, iil. p. 250.
a. 2. Puerto Princesa, Palawan. yes hazel.

[Procured in the deep woods a good way from the sea.—J. B. 8. ]

Compared with (typical) Javan examples, the present specimen shows complete iden-
tity. It isnot quite in full plumage, but is assuming the grey coloration of the adult, the
young feathers being downy in texture and not of such a blue-grey colour as in the old
bird, the shade of the feathers being of a dull ashy. The measurements are as follows
—total length 9 inches, culmen 0:9, wing 4°7, tail 3°75, to tip of outer feather 4°85,
tarsus 0°7,

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. By)

MUSCICAPIDA.
PuitEntomA, Hyton.
49, PHILENTOMA ALBIVENTRIS, sp. n.

Ad. schistaceo-cyanescens, plumis frontalibus medialiter argentescenti-albido striolatis ;
dorso scapularibusque leete castaneis, his summis vix schistaceo lavatis; tectricibus
alarum schistaceo-cyanescentibus, majoribus intimis extus castaneis; remigibus
nigris, primariis vix schistaceo limbatis, secundariis extus castaneis; rectricibus
duabus centralibus nigris, duabus proximis extus castaneis, intus nigris, reliquis
omnino castaneis; facie laterali sordide cyanescente ; gutture et pectore toto schis-
taceo-cyanescentibus, plumis anguste argentescenti-albo medialiter striolatis, abdo-
mine toto albo; crisso et subcaudalibus pallide castaneis, hypochondriis imis fulvo
lavatis; subalaribus cinereis, majoribus albicantibus. Long. tot. 6:4, culm. 0°65,
alze 3°05, caude 3°45. tarsi 0-7. |

a. ad. Guimaras.

[Found in the thick woods on low trees.—J. B. S.]

This species closely resembles P. cyaniceps, which is in the Museum, from Mindanao,
but is distinguished at once by its white belly. It is not improbable that it may be a
sexual plumage of P. cyaniceps.

Ruipipura, Vigors & Horsfield.
50. RHIPIDURA NIGRITORQUIS, Vigors. Leucocerca nigritorquis, Walden, t. c. p. 182.
a. &. Puerto Princesa, Palawan. yes black.
b. 9. Zamboanga, Mindanao. : Eyes dark.
[Found by me springing about over the knees of thé mangrove-trees, feeding appa-
rently on insects.—J. B. 8.]
I cannot see the slightest difference in the sexes of this bird, which seems also to be
the same specifically from the different islands of the Philippines.

Cyornis, Blyth.
51. CYORNIS PHILIPPINENSIS, sp. n. Cyornis banyumas, Walden, ¢. c. p. 152.
C. similis C. banyumati, sed abdomine et subcaudalibus albis pspaepead Long. tot.
5°8, culm. 0°45, alse 2:95, caude 2°6, tarsi 0-7.
a-c. Panay.

This is the C. banyumas of Lord Tweeddale’s list ; but he does not omit to notice certain
differences. On comparing Dr. Steere’s specimens with a large series from Labuan and
other localities, I notice that the belly and under tail-coverts are white in the Philippine
birds, while they are always rufous in those from Borneo. I think, therefore, that the
former must be kept distinct.

Hypotuymis, Boie.
52. HypoTHyMIs AZUREA (Bodd.): Walden, 7. c. p.182.
“a, 6. 2. Basilan (Isabella). Eyes dark brown,
[Pretty abundant in the island of Basilan.—J. B. S. |

326 ; MR. R. B. SHARPE ON BIRDS COLLECTED BY

This wide-spread species appears to be found in most of the islands of the archipelago,
having been noted from Guimaras, Negros, and Luzon.

53. HypoTHYMIS SUPERCILIARIS, sp. n.

¢ ad. cyanescenti-cinereus, pileo saturate ceruleo; fronte et linea angusta super-
ciliari argentescenti-cyaneis; loris et plumis periophthalmicis cinereo-nigris ; facie
laterali, gutture et pectore summo cyanescentibus ; corpore reliquo subtus albicante,
cyanescenti lavato ; tibiis cyanescenti-cinereis ; tectricibus alarum dorso concoloribus,
plumis intus nigricantibus; remigibus nigricantibus, extus leete cyaneis, secundartis
intimis omnino cyaneis; rectricibus intermediis cyaneis, reliquis nigricantibus extus
cyaneis; subalaribus pallide cyanescenti-cinereis ; rostro nigro; pedibus brunneis;
iride saturate brunnea. Long. tot. 6:2, culm. 0-6, ale 3:1, caudie 3:0, tarsi 0°65.

a, 6. Isabella de Basilan, October 1874. Eyes dark brown.

[A true Flycatcher in habits, flying from limb to limb, and catching flies on the wing.
—J. B.S8.] .

The species nearest to this new one seems to be ZZ. indigo; but the latter is a much
smaller bird. The general tint of the upper surface is of a much greyer blue in
H. superciliaris, while the bright cobalt wings and tail also distinguish it in addition to
the silvery blue frontal line and eyebrow. The under surface is greyish blue, not greenish,
the colour extending over the throat and chest, leaving the rest of the under surface
bluish white. In H. indigo the breast is also greenish, leaving only the abdomen
white. .

Eumytias, Cabanis.
54. EUMYIAS PANAYENSIS, sp. n.

Ad. sordide cyanescens, pileo vix leetiore; tectricibus alarum minimis dorso concolori-
bus, majoribus cum remigibus rectricibusque nigricantibus dorsi colore marginatis ;
facie laterali et corpore subtus vix leetius cyanescentibus, abdomine medio albicante
cyaneo lavato; crisso et subcaudalibus subalaribusque albis. Long. tot. 6-0, culm.
0°55, alee 2°95, caudee 2°45, tarsi 0°7.

a. ad. Panay. .

Closely allied to #. sordida, Walden (£. ceylonensis, Gray, Handl. B.i. p. 326. no. 489,
deser. nulla), but distinguishable by its broader bill and by its brighter cyaneous under
surface. In #. sordida the breast is grey, with a brighter blue shade on the chin, and
the lores and feathers in front of the eye and at the base of the lower mandible are
black. In #. panayensis the breast is rather bright greyish cyaneous, a little more
silvery on the throat, lores, and base of mandible, the latter parts not showing any
black.

Burauis, Boie.

55, BUTALIS MANILLENSIS, Bp.: Walden, ¢. c. p. 183.

a. Panay.

The single specimen collected by Dr. Steere proves that 6. manillensis of Bonaparte

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. B27

is the species more familiarly known as B. griseosticta of Swinhoe; and of course
Bonaparte’s name will take precedence over the latter title. This identification was to a
great extent predicted by Lord Tweeddale (J. ¢.).

XANTHOPYGIA, Blyth.
56. XANTHOPYGIA NARCISSINA (Temm.): Swinh. P. Z.8. 1871, p. 380.
a. &. Dumalon, Mindanao. Eyes hazel.

A lovely specimen in full plumage. This is the first time that the species has been
known from the Philippines, where it is doubtless only a winter visitant.

XANTHOLESTES, gen. n.*

57. XANTHOLESTES PANAYENSIS, sp. 0.

é. supra olivascenti-flavus, pileo saturatiore; uropygio leete flavo; tectricibus alarum,
remigibus et rectricibus brunneis flavido marginatis; loris flavis; subtus leetissime
flavus, gutture vix obscuriore. Long. tot. 4°7, culm. 0°45, alee 2°2, caude 2:2,
tarsi 0:5.

a. ad. Panay.
Closely allied to XY. helianthea, Wall. (Walden, Tr. Z. 8. viil. p. 66, pl. vii. fig. 1), of
which it is a representative species, differing in its smaller size and olive-greenish head
and face, the latter being yellow in X. helianthea.

Srrarra, Blyth.
58. SETARIA RUFICAUDA, Sp. D.
$ ad. supra fulvescenti-brunneus, pileo vix obscuriore; supracaudalibus caudaque
castaneis, rectricibus vix apicaliter brunnescentibus; tectricibus alarum dorso
concoloribus, majoribus saturate brunneis extus dorsi colore lavatis; tectricibus
primariorum nigricantibus extus angustissime olivaceo lavatis; remigibus saturate
brunneis extus fulvescenti-brunneo lavatis, secundariis intimis dorso concoloribus ;
loris cum genis et facie laterali cinereis, regione parotica fulvescenti lavata ; cula et
pectore abdomineque toto albis, pectore summo olivaceo lavato; hypochondriis
olivaceis; tibiis cinerascenti-brunneis; subalaribus subcaudalibusque albis, illis
extus olivaceis; rostro nigro; pedibus pallide brunneis; iride saturate brunnea.
Long. tot. 5°4, culm. 0°6, alee 2°85, caudz 2°35, tarsi 0°7.
a, b. 8. Isabella de Basilan, October 1874. Eyes hazel.
| Has the habits of a Flycatcher; found in rather low deep woods, and only met with
in Basilan.—J. B. 8. |
This new species is similar to S. pectoralis, Salvad., which it strongly resembles below ;
but it differs in its grey cheeks and bright-chestnut upper tail-coverts and tail. In the
latter character it comes very close to S. affinis, which also has the upper tail-coverts and
tail red; but then the latter bird has the lores and sides of face white, not grey.

* Mzyiolestes, in which genus these birds have been placed, is a synonym of Hemipus; cf. Sharpe, Cat. B. iii. p. 305.

328 MR. R. B. SHARPE ON BIRDS COLLECTED BY

ZEOCEPHUS, Bonaparte.
59. ZEOCEPHUS CINNAMOMEUS, sp. n. (Plate XLVIIT. fig. 1.)

3 ad. omnino cinnamomeus, subtus paullo letior, abdomine albicante; remigibus nigri-
canti-brunneis, extus et intus cinnamomeis; cauda saturate castanea, rectricibus
quatuor medianis versus apicem brunnescentibus. Long. tot. 7°5, culm. 0°75, ale 3°4,
caudee 3°35, tarsi 0°7.

a. &. Isabella de Basilan. Eyes dark.

Although not possessing such a strongly graduated tail as Z. rufus*, this bird is
apparently congeneric, and may be distinguished by its brighter and clearer cinnamon-
red colour, and especially by its white belly. Like Z. rufus it has a wattled ring round
the eye.

[Of this new bird I only saw one specimen, which I distinguished by its bright red
colour among the green leaves. It was shot in a thick forest in the interior of the island
of Basilan, at some distance from the sea.—J. B.8.]

60. ZEOCEPHUS CYANESCENS, sp.n. (Plate XLVIII. fig. 2.)

9 ad. schistaceo-cyaneus; loris et margine angusto frontali, mento summo et macula
ad basin mandibule posita, nigris; remigibus nigricantibus dorsi colore extus
lavatis; cauda dorso concolori, sed saturatiore; rostro ¢yaneo; palpebra carun-
culata cyanea; iride nigra. Long. tot. 85, culm. 0°85, ale 3°85, caudze 4:25,
tarsi 0°75.

a. &. Puerta Princesa, Palawan, July 24. Eyes black ; inside of mouth and tongue leaf-green ;

eyelid and bill light blue.

[I saw a few more of this species in the island; but it did not appear to be very
abundant.—J. B. 8.]
This species is totally distinct from the other two rufous-coloured ones.

HIRUNDINID A.
Hirunpo, Linneeus.
61. Hirunpo rustica, L.: Dresser, B. Hur. part XXX.
a, 6. & juv. Dumalon, Mindanao. LHyes dark.
[Found in some numbers; not seen far from the beach.—J. B. S.]

These birds seem to be our Common Swallow and not H. javanica. The specimens
procured were both young birds; but I do not doubt the identification.

ORIOLIDA,
G2. OnroLus CHINENSIS, L.: Sharpe, Cat. B. iii. p. 208. Broderipus acrorhynchus
(Vig.), Walden, ¢. ¢. p. 185.
a. 2 ad. San Bernardino, Negros, Jan. 1875. Baill flesh-colour.

b. g juv. Dumalon, Mmdanao. Hyes dark.
c, ¢ ad. Balabac, Sept. 1874. Eyes “ currant-”red.

* Tord Tweeddale states (¢. c. p. 183) that the precise locality of Z. rufusis not known. We have two specimens in
the Museum collected by Cuming at Cataguan, in Luzon.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 329

[Pretty generally distributed through all the islands of the Philippine group, but
rather shy. At Bernardino it seemed to be common, feeding on the fruits of the
wild bananas.—J. B. 8. ]

Between the adult male and female there is scarcely any perceptible difference beyond
a slightly more greenish tinge on the back and scapulars of the latter.

The young bird hasa blackish bill; and the yellow of the back has a very strong wash of
greenish; the wings and tail are browner, the centre tail-feathers being olive; the horse-shoe
mark is blackish washed with yellow; and on the breast are several fine streaks of black,

63. ORIOLUS SULUENSIS, Sharpe, Cat. B. ili. p. 205.
a, 6, c, d. Si Butu, Sulu archipelago.

Mr. Low very kindly presented to the Museum four specimens of an Oriole which
at first appeared to me to resemble Dr. Cabanis’s description of his Oriolus formosus ;
but, thanks to the kindness of Dr. Briggemann, who procured me the loan of some
specimens of the latter bird, I find that the species is really very distinct, as may be
seen by consulting Mr. Dawson Rowley’s plate (Orn. Misc. pl. lvi.).

64. OrtoLus SteERt, Sharpe, Cat. B. ii. p. 218, pl. x.

g ad. supra sordide aureo-flavus, regione parotica et colli lateribus dorso concolori-
bus; loris, genis, gutture toto et preepectore cinereis; corpore reliquo subtus albo,
plumis medialiter nigro late striolatis; subcaudalibus lete aureis; tectricibus
alarum minimis et medianis dorso concoloribus, majoribus remigibusque nigricanti-
bus extus flavis, secundariis intimis dorso concoloribus; tectricibus primariorum
et primariis nigris extus cinereo limbatis; subalaribus cinereis, externis flavo extus
limbatis; rectricibus nigris aureo apicatis versus basin pogonii interni flavicantibus,
pennis duabus centralibus flavicantibus, apicaliter nigricantibus; rostro rubro ;
iride coccinea. Long. tot. 7-5, culm. 0°9, alee 4°45, caude 3°1, tarsi 0°9.

a. &. Basilan (Isabella). Eyes cherry-red.
6. Negros.

[In Negros I found these Orioles feeding on the wild bananas, everywhere rather shy,
—J. B.8.] ;

The Negros example is rather larger and darker in colour, with broader black ovate
streaks on the belly; the yellow spot on the outer tail-feathers too is rather larger.
Total length 8°3 inches, culmen 1:05, wing 4°7, tail 3:45, tarsus 0°9. Although this
indicates a larger race, I do not propose to give it a specific name. .

This new Oriole resembles at first sight O. wanthonotus, which it equals in size and in
the broad black streaks on the belly; but it is quite distinct from this and from every
other Oriole by reason of its grey throat and fore neck and yellow upper surface. In form
it is a true Oriolus, approaching in colour some of the species of Mimeta.

PITTIDA.
Bracuyvrts, Thunberg,
65. BRACHYURUS STEERII, sp.n. (Plate XLIX.) Sharpe, ‘Nature,’ August 1876.

2 ad. supra olivascenti-viridis; uropygio leetissime cyaneo; supracaudalibus et cauda

330, MR. R. B. SHARPE ON BIRDS COLLECTED BY

nigris; pileo summo et capitis lateribus nigerrimis; tectricibus alarum letissime
cyaneis, majoribus nigris cyaneo terminatis; remigibus nigris, secundariis intimis
dorso concoloribus, externis extus viridibus, primariis 4°, 5°, 6° extus prope basin
macula alba ornatis; gula alba; corpore reliquo subtus lete cyaneo; pectore et
abdomine medio nigerrimis; crisso et subcaudalibus scarlatinis; rostro nigro;
iride saturate brunnea. Long. tot. 7°5, culm. 1:1, alee 4°75, caude 1°7, tarsi 1°7.

a. 8 ad. Dumalon, Mindanao.

b. 8 juv. Dumalon, Mindanao.

[This bird was probably not very rare in the neighbourhood of the village of Dumalon,
as I had one or two more specimens of it, which unfortunately were destroyed.—
dig 18s tS

There is no species with which this beautiful Pitta can be directly compared, as it
differs so thoroughly from all the described forms. Its bright blue breast and white
throat distinguish it from all the black-headed members of the genus.

A second specimen is apparently a younger male from the same locality. It differs
only in the colouring of the belly, which is pale scarlet all the way up, becoming blacker
in certain parts, and showing that the whole is becoming black as in the full-plumaged
bird described.

66. BRACHYURUS PROPINQUUS, Sp. 0.

B. affinis B. erythrogastro, sed tergo cyanescente, plaga transversali dorsali tantum
viridi; pileo castaneo-rufo; collo postico lete cyaneo; gutture toto nigricante,
mento tantum rufo maculato; preepectore nigerrimo ; plaga pectorali lete cyanea,
pectoris lateribus viridescentibus ; corpore reliquo subtus latissime scarlatino ; sub-
alaribus brunneis sordide cyanescenti lavatis, alis ut in B. erythrogastro coloratis.
Long. tot. 6°5, culm. 0°75, ale 3°6, caude 1°5, tarsi 1°35.

a. & ad. Balabac, Sept. 1874. Iris grey.
6. g juv. Dumalon, Mindanao. Iris hazel.

As might be expected, this bird is very closely alliefl to B. erythrogaster, which is
apparently a Manila species. We have compared the Balabac skin with four examples
of the true B. erythrogaster, and find that it cannot be united to that species. The
differences may be summarized as follows :—

B. erythrogaster. B. propinquus.

Back entirely green, including the scapulars, the Back entirely cobalt, meluding the scapulars ;
lower ones only of the latter washed with blue at only the middle of the back’ green washed with
the tip; rump and upper tail-coverts and tail blue, forming a band across the back.
blue.

Throat brown, gradually shading into black on Throat entirely brownish black, becoming jet-
the fore neck. black on the fore neck. ;

Chest green, with a slight wash of blue on the Chest bright blue, the sides only greenish.
central portion.

Under wing-coverts greyish brown. Under wing-coverts dull blue.

This species is intermediate between B. erythrogaster and B. cyanonotus of Ternate,

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 6881

differing from both in its blackish throat and blue under wing-coverts. The other
characters are duly enumerated above. The young bird from Mindanao seems to belong
to this species; but being immature, it is rather difficult to say.

67. BracHyurus sorpipvs (Miull.). Melanopitta sordida, Walden, J. c. p. 187.
a,b. 3 Q. Puerto Princesa, Palawan.
c. g. Basilan.

[Common in both the above-mentioned islands. They alight on the low stumps and
logs in the woods, and emit a curious shrill note at intervals of about a minute. On
being disturbed they fly off rapidly and disappear in the jungle, but very soon return
to the same spot, and commence whistling again directly the danger appears to have
passed. They are said by the natives to raise a large heap of leaves and nest on the top
of the mound.—J. B. S.]

Considering the number of Bornean forms shown by Dr. Steere to inhabit Palawan, it
is interesting to note that the Black-headed Pitta of that island is the true Philippine
species, and not the Bornean one, B. Muelleri. These two birds are closely related ;
but the Bornean species almost entirely wants the black patch on the belly, and has the
thighs blackish instead of ochraceous brown.

TIMELIIDA.
Macronvts, Jardine and Selby.
68. MacRONUS STRIATICEPS, sp. n.
¢ ad. supra fulvescenti-brunneus, scapis pallide fulvescentibus ; uropygii plumis longis,
laxis, scapis basaliter albis ; tectricibus alarum dorso concoloribus et eodem modo
striatis, majoribus autem fulvescenti-brunneis concoloribus; remigibus brunneis
extus castaneis ; cauda brunnea, rectricibus extus castaneo lavatis; pileo nuchaque
nigris, clare medialiter albo striatis; facie laterali alba, plumis utrinque nigro
marginatis, quasi striolatis; gula pure alba; corpore reliquo subtus albido, plumis
pectoralibus olivaceo striatim marginatis, corporis lateribus, tibiis subcaudalibusque
olivascenti-brunneis albido striolatis; subalaribus olivascenti-brunneis ; remigibus
subtus cineraceis, intus rufescentibus; rostro nigricante; pedibus brunneis; iride
brunnea. Long. tot. 5:2, culm. 0°6, ale 2°15, caudze 2°15, tarsi 1°8.
a. 2. Isabella de Basilan. Eyes hazel.
6b. g. Malamaui, October 1874. Eyes hazel.
[The sexes are alike. Shot in exactly the same localities as Mixornis Woodi.—
J. B.S.]
5 Mrxornis, Hodgson.
69. Mrxornis Woopl, sp. n.
Ad. similis WY. pileate, sed facie laterali cinerea, mento albo et dorso olivascenti-
cineraceo distinguendus. Long. tot. 5:2, culm. 0°6, ale 2°35, caude 2°25, tarsi 0°8.
a, 6. S. Puerto Princesa, Palawan.
c. 6. Puerto Princesa. Eyes flesh-colour.
‘d. &. Puerto Princesa. Eyes hazel.
[Only found in the deep brush.—J. B. 8. ]
SECOND SERIES.—ZOOLOGY, VOL. I. Pas

332 _ MR. R. B. SHARPE ON BIRDS COLLECTED BY

PTILOCICHLA, gen. n.

Allied to Turdinus, but distinguished at a glance by the long drooping sickle-shaped
plumes on the sides ofthe lower back. The type is :—

70. PYILOCICHLA FALCATA, sp.n. (Plate L. fig. 3.) .

g ad. pileo summo nuchaque rufescenti-brunneis; fronte laterali et striga postoculari
cervino-rufis ; loris anguste albis; macula anteoculari et regione parotica nigris;
genis gulaque albis, vitta malari angusta nigricante; dorso toto nigricanti-brunneo,
plamis late medialiter fulvis, dorsi postici plumis longis, laxis, falcatis, dependenti-
bus, brunneis vel nigricantibus late albo striatis; alis rufescenti-brunneis, remigibus
intus nigricantibus ; cauda omnino rufescenti-brunnea; corpore subtus nigricante,
late albo medialiter striato; tibiis, subalaribus et subcaudalibus brunneis, fulvo
striolatis ; rostro flavicanti-corneo, mandibula flavicante; pedibus saturate brunneis.
Long. tot. 6°8, culm. 0:95, alee 3:2, caudee 2°4, tarsi 1:3.

a. &. Puerto Princesa, Palawan.

[When first I saw this curious bird he reminded me at once of the Ant-birds
(Formicariide) which I had seen in South America. The little fellow was in the
thickest wood, hopping about on the ground, and puffing himself out like a ball of
feathers, presenting a most remarkable appearance.-—J. B. 8.]

DENDROBIASTES, gen. 0.

Closely allied to Alcippe, but distinguished by its long wings, which reach to the end of
the tail. The type is :—

71. DENDROBIASTES BASILANICA, sp. n. (Plate LITT. fig. 1.)

2 ad. supra rufescenti-brunnea, uropygio paullo fulvescenti lavato; supracaudalibus
castaneis; pileo magis fuscescenti-brunneo; loris, supercilio indistincto et facie
laterali fulvescentibus ; annulo periophthalmico letius fulvo; regione parotica ful-
vescenti-brunnea, auguste albido striata; corpore subtus albicante, gutture vix
fulvido lavato; pectoris lateribus, torque pectorali, hypochondriisque fulvescenti-
brunneis ; tibiis et subalaribus rufescentibus, margine alari rufescenti-brunneo; alis
caudaque supra saturate brunneis, tectricibus dorsi colore extus lavatis, remigibus
et rectricibus extus castaneis; rostro nigricanti-brunneo; pedibus flavicantibus ;
iride brunnea. Long. tot. 4°2, culm. 0°55, ale 2°4, caudz 1°6, tarsi 0°8.

a. 9. Isabella de Basilan. yes hazel.

[Shot in open ground near the fort at Isabella, inhabiting the low bushes in this
neighbourhood.—J. B. 8.]
This seems to be an entirely new form of Timeliine bird, remarkable for its long wings.

Wing of Dendrobiastes basilanica: nat. size.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 300

PHYLLORNITHID”.

PuyLiornis, Boie.

72. PHYLLORNIS PALAWANENSIS, sp.u. (Plate L. figs. 1, 2.)

s ad. supra lete viridis, pileo antico magis flavicante; tectricibus alarum dorso con-
coloribus basaliter cyaneis, minimis late purpureis plagam distinctam exhibentibus,
extimis et tectricibus primariorum purpurascenti-cyaneis; remigibus nigris, extus
cyaneis, secundariis viridibus versus basin cyaneis; cauda sordide cyanea, viridi plus
minusve lavata; loris et plumis ad basin mandibule positis viridi-cyaneis; facie
laterali aureo-flava, regione parotica postica virescente; oculo plumis purpureis
circumdato ; genis albis, fasciam mystacalem formantibus; gutture toto late aureo-
flavo ; corpore reliquo subtus viridi; subalaribus cineraceis, viridi lavatis, margine
alari cyanea; rectricibus intus cinerascentibus, pogonio interno versus basin albi-
cantibus; rostro nigro ; iride saturate brunnea. Long. tot. 7:0, culm. 0°85, ale 3°5,
caudee 2°8, tarsi 0:8.

a. 6; 6,¢. 2. Puerto Princesa, Palawan. July 1874. Eyes dark.
[Shot in low bushes and trees in some numbers, eating fruit about the borders of the

clearings. They were generally in small flocks.—J. B. S.]

The sexes seem to be alike in every respect excepting as regards the moustache, which
is blue instead of white in the male.
The present species bears some resemblance to the female of P. Sonnerati, but is

' distinguished by the blue eye-ring and the blue on the wings and tail. It seems to be

quite a distinct species from any yet known.

Tora, Horsfield.
73. [oRA scAPULARIS, Horsf.: Salvad. Ucc. Born. p. 190.

a. g. Puerto Princesa, Palawan. Eyes black.
b,c. g;d. 2. Puerto Princesa. Eyes grey.
[Shot in the same locality as the Jrena, and found always in pairs.—J. B. 8.]
This is an interesting addition to the birds of the Philippines; but, like so many of
the Palawan birds, it is a well-known Bornean species.

IRENID.
Trena, Horsfield.
74. IRENA TWEEDDALII, Sharpe, Cat. B. iii. p. 268. (Plate LI. fig. 1.)

I. similis F. puelle, sed colore cyaneo vivide argentescenti-turcoso minime purpurascente
distinguenda. Long. tot. 8-0, culm. 1:05, alee 4°85, caudze 3°85, tarsi 0°7.
a. ad. Puerto Princesa, Palawan.
[This bird was eating fruit in low bushes near Puerto Princesa. A number of convicts
were hard at work making a road; and two or three of these birds were feeding in
close proximity, quite undisturbed by the noise of the men at work.—J. B. 8.]

This undoubtedly new Irena has the short tail-coverts of J. pwella, and agrees exactly
2x2

Bes MR. R. B. SHARPE ON BIRDS COLLECTED BY

with that species in the distribution of the black and blue colours ; the latter, however,
instead of being purplish, is of the most brilliant silvery turquoise-blue. I have named
this species, which bears the same relation to J. puella that J. turcosa does to I. cyanea,
after Lord Tweeddale, who has so carefully worked out the species of this genus, and
to whom we are indebted for his most excellent paper on the birds of the Philippines.

75. IRENA MELANOCHLAMYS, sp. n. (Plate LI. fig. 2.)

3. ad. similis I. cyanogastri, sed interscapulio scapularibusque nigris haud purpureo
lavatis distinguenda. Long. tot. 10-0, culm. 1:05, alee 4°8, caudze 41, tarsi 0°7.
a. &. Isabella de Basilan. Eyes carmine.

[Only seen on one or two occasions. It is apparently rather rare in the part of the
island which I visited.—J. B. 8.]

BRACHYPODIDA.
POLIOLOPHUS, gen. n.
76. PoLroLoruus vrosricrus.—Ivus ? wrostictus, Walden, ¢. c. p. 191, pl. xxxii. fig. 2.
a. 3. Negros. yes hazel; a yellow ring round the eye.

The combination of an eye-wattle and a crest seem to me to indicate that this bird
should be placed in a distinct genus from Pycnonotus or Otocompsa; and I therefore
propose the name of Poliolophus. The colouring of the bird is very badly rendered

in the plate accompanying Lord Walden’s paper—the greyish crest not being defined,
and the eye-watile entirely omitted.

CriINIGER, Temin.
77. CRINIGER FRATER, Sp. 0.

g ad. affinis C. gutturali, sed pectore ochrascenti-flavo et subcaudalibus ochrascentibus
distinguendus. Long. tot. 8°8, culm. 0-95, ale 4:0, caudee 3-7, tarsi 0°85.
a. 3. Puerto Princesa, Palawan, July 1874.

This new species is very closely allied to two others, viz. OC. gutturalis and C. gularis,
all of them having brown heads and olive-green backs. It differs from C. gularis in
being larger, with a longer beak, and in having the under surface from the throat
downwards ochraceous yellow,, varying towards pale fawn. Other characters are the
brown tail with olive-green edges, and the greyish white cheeks with silvery white
shaft-streaks. In (. gularis the ear-coverts are uniform ashy brown, the tail is dull
chestnut-red with pale ochraceous brown edgings; the under surface, with the excep-
tion of the throat and fore neck, is bright yellow, including the under tail-coverts.

To C. gutturalis the Palawan bird is very similar, but it differs in the colouring of the
under surface. C. guttwralis has the chest ashy brown washed with olive, the under
surface pale yellowish with tawny under tail-coverts; C. frater has the throat white,
and the entire rest of the under surface ochraceous yellow, including the under tail-
coverts. The flanks in both birds are olivaceous, and they are both alike in the grey
facial markings and the white-streaked ear-coverts.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 335

Hypsiprtes, Vigors.
78. HYPSIPETES PHILIPPENSIS (Gm.): Walden, #. c. p. 191.
a. 3. San Bernardino, Negros, Jan. 1875. Hyes reddish hazel.
[Feeding on the wild bananas at the foot of Malaspina volcano.—J. B. S.]

79. HYPsIPETES RUFIGULARIS, sp. 0.
3 ad. H. philippensi persimilis, sed genis brunneis haud striatis et gutture toto rufo con-
colori distinguendus. Long. tot. 9°8, culm. 1:1, alee 4°45, caudze 4°25, tarsi 0°85.
a. 3. Malamaui, Oct. 1874. yes hazel.
b. Sg. Isabella de Basilan. Eyes cherry-red.

[Feeding on fruit, and resembling Thrushes in their actions. Very abundant both in
Basilan and Malamaui, which is a flat island, only separated from Basilan by a narrow
deep channel about 100 yards broad.—J. B. S.]

This species differs from H. philippensis in its uniform rufous throat and white breast
without the conspicuous white shaft-streaks. It is also a decidedly larger bird.

TURDIDA.
Monricona, Boie.
80. Monricona sonirartia (Miill.): Walden, ¢. c. p. 192.
a. Panay.
[I got this Rock-Thrush on the top of the ranges of mountains in Panay. They were
scattered about on the mountain-sides, but were extremely shy, hopping about the fallen
timber, and disappearing on the first alarm of danger—J. B. 8. ]

Pratincona, Koch.
81. Pratincona cApRrata (L.): Walden, ¢. c. p. 198.
a. &. Panay.

[Shot in a field about halfway up the mountains in Panay, as it was flying about from
one bush to another.—J. B. 8.|

The discovery of this species in the Philippines is very interesting, as since Brisson’s
time it had not been met with there, and Lord Tweeddale appeared almost inclined to
doubt its real occurrence there.

Crrrocincia, Gould.

82. CITTOCINCLA NIGRA, sp.n. (Plate LII. figs. 1 & 2.)

3 ad. nigra, indigotico nitens; alis nigris, tectricibus alarum omnino dorso concolori-
bus; cauda nigra, quadrata, rectricibus quatuor intermediis nigris, reliquis albis,
basaliter nigris; facie laterali et corpore subtus omnino indigotico-nigris; corporis
lateribus et hypochondriis albo maculatis; tibiis nigris; subalaribus nigris; sub-
caudalibus albis; rostro nigro; pedibus nigricanti-brunneis; iride nigra. Long.
tot. 9°0, culm. 0°7, ale 3°4, caudee 4°6, tarsi 1:05.

336 MR. R. B. SHARPE ON BIRDS COLLECTED BY

g. mari similis, sed minor et abdomine albo, hypochondriis rufescentibus distinguenda.
Long. tot. 7-2, culm. 0°6, alee 0-15. caudze 3°6, tarsi 1:0.
a, 6. 3. Puerto Princesa, Palawan, July 1874. Eyes black.
c. 2. Puerto Princesa, July 1874.

The chief difference between the genera Cittocincla and Copsychus seems to lie in the
long tail of the former, this being much longer than the wing in Citfocinela but about
equal to it in Copsychus. 'Therefore the present bird is a new species of the greatest
interest, as being a true Cit/ocincla in structure with the plumage of a Copsychus; and of
course its black breast distinguishes it from C. swavis, C. macrura, &c. The difference
in the sexes is very extraordinary, and quite unlike any thing else known to occur in
this genus.

SYLVIID A.

Puyiioscorvs, Boie.

83. PHYLLOSCOPUS LUGUBRIS, Blyth: Seebohm, Ibis, 1877, p. 78.
> a. 8. Dumalon, Mindanao, Oct. 1874. yes hazel.

IT am able to state, with pleasure, that my friend Mr. Henry Seebohm, who has so
thoroughly worked out the Phylloscopi, has identified the above specimen for me as
P. lugubris. He considers that it is a young male, a bird of the year, not haying
quite completed its moult.

CIsTICOLA, Kaup.
84. CISTICOLA, sp. ?

a. San Bernardino, Negros, Jan. 1875. Eyes light hazel.

[Shot in grass in fields of sugar-cane near 8. Bernardino.—J. B.8.]

OrtHoromus, Horsfield.

85. ORTHOTOMUS FRONTALIS, Sharpe, Ibis, 1877, p. 112, pl. u. fig. 1.

g ad. supra olivascenti-viridis, uropygio paullo flavicante; tectricibus alarum dorso
concoloribus; remigibus saturate brunneis, extus dorsi colore layatis; rectricibus
olivascenti-brunneis, pallidius terminatis et macula saturatiore brunnea subter-
minali notatis et anguste flavido limbatis; fronte et supercilio angusto castaneis ;
pileo summo cum nucha et colli lateribus cinereis; facie laterali albida, regione
parotica cinereo lavata; genis et gutture toto sericeo-albis; corpore reliquo subtus
fulvescenti-albo, hypochondriis et subcaudalibus flavicanti-viridibus; tibiis pallide
castaneis; subalaribus albis flavicanti lavatis, margine alari lete flavo; rostro
saturate brunneo, mandibula flavicante; pedibus flavicanti-brunneis; iride pallide
brunnea. Long. tot. 3°7, culm. 0°55, ale 1°6, caudee 1:5, tarsi 0°75.

a. 3. ad. Zamboanga, Mindanao. yes light hazel.
b,c. 2 juw. Basilan (Isabella). Eyes grey. |
[This small Tailorbird was very abundant in the grass near Isabella de Basilan.—
J.B.8.]

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 307

The two young specimens seem to me to belong to the same species as the adult
male: but this is by no means certain; for in one specimen I can detect a slight rufous
shade on the sides of the crown, the adult O. frontalis of course not having this
colouring.

86. ORTHOTOMUS CINEREICEPS, Sharpe, Ibis, 1877, p. 113, pl. ii. fig. 2.

é ad. pileo toto cum nucha et colli lateribus saturate cinereis; loris fulvescentibus ;
facie laterali et vitta angusta malari schistaceo-nigris; regione parotica sericea
alba; dorso toto et tectricibus alarum leete olivaceo-viridibus, his majoribus letius
flavo marginatis; remigibus saturate brunneis, extus late dorsi colore lavatis;
rectricibus saturate brunneis, anguste olivascenti-flavo limbatis et terminatis, ma-
cula subterminali nulla; mento et gule lateribus albis; gula media et jugulo
nigris, plumis paucis vix albido marginatis; pectoris lateribus et hypochondriis
subcaudalibusque olivascenti-flavis; corpore reliquo subtus medialiter sericeo albo;
tibiis olivascentibus; subalaribus flavidis, margine alari lete flavo; rostro corneo-
nigricante, mandibula flavicante; pedibus pallide brunneis ; iride pallide brunnea.
Long. tot. 5, culm. 0°65, alee 1°85, caude 1:9, tarsus 0°9.

a. 6 ad. Basilan (Isabella). Eyes light hazel.

[Found in the grass near the fort at Isabella de Basilan.—J. B. 8.]

87. ORTHOTOMUS CASTANEICEPS, Wald. Zz. c. p. 195.
a. ad. Panay.
[Shot in the long grass on the mountains at a height of 2000 to 3000 feet.—J. B. 8.]
This is a perfectly good species. I have compared Dr. Steere’s skin with the type in
Lord Tweeddale’s collection from Guimaras, and find them identical.

88. ORTHOTOMUS RUFICEPS (Less.): Salvad. Ucc. Born. p. 248.
a. ad, Puerto Princesa, Palawan.

I have compared this specimen with Bornean and Sumatran examples, and can find no
difference in plumage. Total length 4°7 inches, culmen 0°65, wing 1:95, tail 1:85, tarsus
1:85. It is the first occurrence of the species in the Philippines.

MOTACILLIDA.
CorYDALLA, Vigors.

89. CoRYDALLA LUGUBRIS, Walden, ¢. c. p. 198.
a. Negros.
[Found in the open grassy plains.—J. B.S. |
Lord Tweeddale having kindly lent me the typical specimen from Guimaras, I have
been able to compare Dr. Steere’s Negros skin ; and I find them quite the same.

300 MR. R. B. SHARPE ON BIRDS COLLECTED BY ©

Antuus, Bechstein.

90. Anruus eusTavi; Corydalla gustavi, Swinhoe, P. Z. 8. 1871, p. 366.
a. Isabella de Basilan.

Having carefully compared the Basilan skin, I came to the conclusion that it was
the same as the Chinese species, which is, therefore, recorded from the Philippines for
the first time.

PARID A.
Parus, Linn.

91. PARUS AMABILIS, sp. n. (Plate LIII fig. 2.)

? ad. pileo undique cum regione jugulari nitide indigotico-nigris; collo postico et inter-
seapulio leete sulfureis; dorso reliquo pulchre cinereo; supracaudalibus indigotico-
nigris; cauda nigra, rectricibus omnibus albo terminatis, reliquis etiam extus
omnino albis; alis indigotico-nigris, tectricibus alarum late albo terminatis, fascias
alares formantibus ; remigibus albo apicatis, primariis extus cinereo lavatis, secun-
dariis intimis etiam versus apicem pogonii interni albis; corpore reliquo subtus
leete flavo; subalaribus albis, macula nigra prope marginem tectricum inferiorem
posita; rostro nigro. Long. tot. 4°5, culm. 0°45, ale 2°85, caudee 1°75, tarsi 0°75.

a. ad.; b. juv. Balabae.

The young bird is very different from the adult. It is brown above, with an olive-
greenish tinge, the head and face browner. Below it is yellow, duller than in the
adult, the throat olive; wings and tail browner than in the adult, with the same white
spotting, but not so pronounced.

92. PaRuUS ELEGANS; Machlolophus elegans (Less.), Walden, ¢. c. p. 199.

a,b. Palawan; ¢. Guimaras.

SITTID A.

93. DENDROPHILA FRONTALIS, Horsf.: Salvad. ¢. ec. p. 161.
a,b. 6. Puerto Princesa, Palawan. Eyes light chrome-yellow.

[A genuine Nuthach in its habits. I shot my two specimens during a little sojourn
that I made among the herdsmen on the opposite sides of the bay from Puerto Princesa.
They were running about on the timber logs exactly as our own Nuthach would do at
home.—J. B. 8.]

Not to be distinguished from Bornean examples.

94. DENDROPHILA @NOCHLAMYS, sp. nu. (Plate LIII. fig. 3).

2 ad. similis D. frontali, sed ubique letius colorata; collo laterali et postico cum inter-
scapulio colore pectoris vinaceo adumbratis; sed preecipue vitta angusta lorali alba
et macula parva alba ad basin mandibule posita distinguenda. Long. tot. 5-0,
culm. 0°65, alee 3:0, caudee 1°7, tarsi 0°7.

a. ad. Guimaras.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 339

{Found running up and down the trees in the manner of Nuthatches. I saw a pair,
but only managed to secure one.—J. B. 8. |

This beautiful new species resembles D. frontalis, but is larger and altogether more
richly coloured; the vinaceous colour of the underparts overspreads the sides of the
neck, and the hinder neck and mantle, the rest of the back being bluish. The chin
alone is white on the under surface; and the species is further distinguished by a narrow
white line running along the base of the lores to the eye from the beak, and by a second
small white spot at the base of the lower mandible.

NECTARINIIDA.

DIC HINA.
Diczum, Cuvier.

95. DicmuM HaMAtTOosTIoTUM, Sharpe, ‘ Nature,’ Aug. 1876, p. 297.

3 ad. supra indigotico-niger, dorso postico magis schistaceo; supracaudalibus nigris ;
tectricibus alarum dorso concoloribus, majoribus cum remigibus et rectricibus
nigricanti-brunneis, extus virescenti limbatis; facie laterali nigra, genis et corpore
subtus albis, hypochondriis cinereo lavatis; fascia preepectorali transversali schis-
taceo-nigra ; pectore et abdomine medialiter lete coccineis, illo utrinque schistaceo-
nigro maculato; subcaudalibus et subalaribus albis, remigibus intus albo margi-
natis; rostro et pedibus nigris. Long. tot. 3°7, culm. 0°45, ale 11, caude 1°2,
tarsi 0°55.

a, b. ad. Guimaras.
c, d, e. ad. Negros.
fg. &- Negros. Hyes hazel.

[Found in all cases in the cocoa-nut trees, flying about and perching on the bamboo
receptacles which are hung under the blossom-stems to catch the juice. The natives
said they were feeding on the latter; but whether this was the case, or whether they
were eating the insects which were attracted by the juice, I was unable to determine.—
J. B.S. |

Two young birds differ from the adult in being slaty grey above, the wings and tail
blacker, with greyish margins to the feathers; under surface of body ashy whitish,
washed with yellowish on the abdomen, and with greyish on the sides of the body. In
one specimen the red colour on the under surface is appearing, and the blackish feathers
on the upper surface: in both the lower mandible is yellowish at the base.

Guimaras and Negros specimens are identical; but the slaty black on the fore neck
and breast varies in extent. It is, however, well developed in every specimen, and
shows clearly that the species is not Diceum papuense of Mindanao, which is the bird
Mr. Gould has figured as the female of his D. retrocinetwm.

96. Dicatm HYPoLEUCUM, Sharpe, ‘ Nature,’ Aug. 1876, p. 298.
3 ad, supra niger, pileo vix viridi nitente; facie laterali collique lateribus nigris; genis,

et corpore subtus toto cum subalaribus subcaudalibusque albis; als caudaque
SECOND SERIES.—ZOOLOGY, VOL. I. - 2 2

340 MR. R. B. SHARPE ON BIRDS COLLECTED BY

nigris ; rostro et pedibus nigris; iride saturate brunnea. Long. tot. 8-6, culm. 0:5,
alee 2°1, caudee 1:05, tarsi 0°55.
is Be MValanaau October 1874. Eyes hazel.
b. $. Isabella de Basilan. Eyes hazel.
[Found in the mangroves in the same localities as the Nectariniide.—J. B. 8.]
This species is distinguished at a glance by its uniform white under surface.

97. DiczuM DORSALE, Sharpe, ‘ Nature,’ Aug. 1876, p. 298.

3 ad. supra lete schistaceo-cinereus, pileo clariore; plaga interscapulari aurantiaco-
miniata; tectricibus alarum dorso concoloribus, extimis anguste olivaceo lavatis;
remigibus nigricantibus, extus dorsi colore marginatis, secundariis paucis olivaceo
limbatis, intimis dorso concoloribus; supracaudalibus schistaceo-cinereis; cauda
nigra, rectricibus schistaceo limbatis; loris nigris; facie laterali saturate schistacea :
subtus leetissime aurantiacus, gula et abdomine imo flavis; subalaribus albis, axilla-
ribus pallide flavis; tibiis intus cinereis, extus flavis; rostro nigricante, mandibula
versus basin pallida; pedibus saturate brunneis. Long. tot. 3°5, culm. 0°5, ale 1:9,
caudee 1:05, tarsi 55.

? a mari diversa: supra olivascenti-viridis, uropygio magis flavicante, facie laterali
pileo concolori; subtus leete flavicans, pectore et subalaribus leetioribus. Long. tot.
3°7, culm. 0°5, alee 1:9, caudee 0°9, tarsi 0°55.

a.g. Panay.
[Shot in the highest part of the island, in the remains of the virgin forest on the highest

range of the mountains west of Ilo Tlo.—J. B. 8.]

The bird which I consider to be the female of this species has unfortunately lost its
label. The only Dicewm with which this new species can be confounded is D. trigono-
stigma; but it is at once distinguished by its yellow throat and blue lower back and

rump.

98. DicmuM TRIGONOsTIGMA (Scop.): Salvad. ¢. c. p. 166.
a. 6 ad. Negros.

The appearance of this species in Negros is somewhat remarkable. As the specimen
entirely agrees with others from Borneo, it might have been expected to occur in
Palawan.
PRIONOCHILUS, Strickl.

99. PRIONOCHILUS, sp.
a. 2 juv. Puerto Princesa, Palawan. Hyes dark.

The only specimen obtained by Dr. Steere is a young bird which I'am unable to refer
with certainty to any species. It is evidently the young of a bird belonging to the same
eroup of Prionochilus as P. percussus and P. santhopygius; and, from its yellow spot
on the rump, it must be closely allied to, if not identical with, the latter. The occurrence
of the genus is interesting; but the exact species can only be determined on the exami-

‘nation of the adult.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 3841

ARACHNOTHERA, Temm.

100. ARACHNOTHERA DiLuTIOR, Sharpe, ‘ Nature,’ August 1876, p. 298; Shelley,

Monogr. Cinnyr. part 3.

ad. supra sordide olivascenti-flavus, uropygio paullo cinerascente,
brunneo, collo postico vix olivascenti lavatus; alis cinerascenti-brunneis, tec-
tricibus vix olivaceo limbatis, remigibus extus olivaceo lavatis, secundartis latius ;
cauda saturate brunnea, rectricibus olivascenti-flavo marginatis, extimis versus
apicem cinerascentibus, tribus externis pogonio interno albo apicatis; facie laterali
saturate brunnea, loris nigricantibus, plumis ad basin mandibule positis et mento
albicantibus; gutture albido, cinerascenti lavato; corpore reliquo subtus flavi-
canti-albido, subcaudalibus albis; fasciis pectoralibus leete aurantiacis ; subalaribus
albis vix flavido lavatis; remigibus subtus cinerascentibus, intus albo marginatis ;
rostro nigricante, mandibula basaliter albida; pedibus saturate brunneis; iride
saturate brunnea. Long. tot. 5-7, culm. 1:5, alee 2°7, caude 1:9, tarsi 0°75.

Eyelids yellow.

pileo saturate

a. 3. Puerto Princesa, Palawan, J uly 1874. Eyes dark. Breeding.
[Shot in the mangroves, but not very abundant.—J. B. 8. ]

Cinnyris, Cuvier.
101. Cryyyris sppratus: Nectarophila sperata (L.), Walden, ¢. c. p. 200.
a,b. § 2. Negros.
c-t. g ad.; k. § juv. Guimaras.
[The pair obtained in Negros were shot together flitting about a dead tree about
20 feet in height, and were apparently nesting. In Guimaras it was very common, hover-
ing about the bamboo cups which are hung up to catch the sap in the cocoa-nut

blossoms.—J. B. 8.]

102. CrynyRis sUGULARIS (L.): Arachnechthra jugularis, Walden, ¢. c. p. 200.

a,b,c. g. Negros.
d,e,f.3 2. Panay.
g. 2. Zamboanga, Mindanao. Eyes hazel.

EUDREPANIS, Sharpe.
Allied to Hthopyga,- but at once distinguished by its rounded tail, which has not the
elongated central feathers of the above-named genus. The type is

103. EUDREPANIS PULCHERRIMA, Sharpe, in Shelley’s Monogr. Cinnyr. part 3.—ditho-
pyga pulcherrima, Sharpe, ‘ Nature,’ 1876, p. 297.

g ad. supra saturate olivascenti-viridescens, plumis basaliter griseis quasi marmoratis ;
plaga frontali letissime metallice violacea; capite laterali nigricaate, linea supra-
parotica metallice violacea; dorso postico et uropygio lete sulfureis ; supracauda-

libus metallice chalybeo-viridibus, rectricibus nigris hoc colore layatis: subtus

lete sulfureo-flayus, macula magna preepectorali miniata; subalaribus albis ; sca-
222

BA2 MR. R. B. SHARPE ON BIRDS COLLECTED BY

pularibus dorso concoloribus; tectricibus alarum metallice viridibus; remigibus
nigricanti-brunneis, extus olivaceo lavatis: rostro nigro; iride coccinea. Long.
tot. 3°5, culm. 0°75, alee 1:8, caudze 1-05, tarsi 0°55.

a. $. Basilan, Isabella. Eyes carmine.

Shot in the mangroves near the town of Isabella.

104. dAitHopyeA MAGNIFICA, Sharpe, ‘ Nature,’ August 1876, p. 297; Shelley, Monogr.
Cinnyr. part 3.
44, similis 4. flavostriate, sed major, rostro fortiore, abdomine et subalaribus nigris
distinguenda. Long. tot. 5:0, culm. 0°95, alee 2°3, caude 2:0, tarsi 0°6.
a-e. Negros.

This is a large representative of Athopyga flavostriata from Celebes, but is distin-
guished at once by its large size and blackish under wing-coverts and belly.

105. AirHopyeA SHELLEYI, Sharpe, ‘Nature,’ August 1876, p. 297; Shelley, Monogr.
Cinnyr. part 3.

g ad. pileo circumscripte chalybeo; loris nigris; facie laterali tota cum colli laterali
et postico et interscapulio saturate scarlatinis; dorso olivascenti-viridi, tectricibus
alarum minimis dorso concoloribus; tectricibus majoribus remigibusque brunneis
extus olivaceo marginatis; plaga uropygiali lete sulfurea; supracaudalibus et
rectricibus duabus centralibus lete chalybeis, reliquis nigris extus chalybeo mar-
ginatis: subtus lete sulfureus, vitta malari chalybea; gula quoque flava, utrinque
searlatino marginata; pectore scarlatino striolata; subalaribus albis. Long. tot.
4-1, culm. 0°65, ale 1°85, caudz 1°85, tarsi 0°55.

a, b, c, d. Puerto Princesa, Palawan, July 1874. yes black.
e. Balabac.

[I met with this species on both the above-mentioned islands. The first one I shot
was in the town of Puerto Princesa, in the mangroves, fluttering about among the
blossoms. I afterwards fell in with it a few miles inland in the forest. In .Balabac I
shot it also in the mangrove.—J. B. S.]

106. ANTHREPTES MALACCENSIS (Scop.): Salvad. ¢.c. p. 178.
a,b,c. 8; d,e. 9. Puerto Princesa, Palawan. Eyes carmine or cherry-red.

~ The specimens collected agree with Bornean examples.

107. ANTHREPTES CHLORIGASTER, Sp. n.

A. similis A. malaccensi, sed major, capite colloque lete metallice viridibus nee purpu-
rascentibus, dorso reliquo et plaga magna humerali leetissime violaceis, tectricibus
alarum majoribus clare castaneo marginatis; remigibus et gutture ut in 4. malac-
censi picturatis; corpore reliquo subtus viridiscenti-flavo ; cauda nigricante, metallice
violaceo late lavata. Long. tot. 5-2, culm. 0°75, alee 2°75, caudee 1:95, tarsi 0°7.

a,b. 3. Negros.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 343

[Not found in mangroves, like 4. malaccenis, but feeding on the blossoms of the
cocoa-nut trees.—J. B. 8.]

These two specimens, collected by Dr. Steere, are precisely similar the one to the
other; and as they differ from a series of at least twenty specimens of 4. malaccensis,
I have no doubt that the green breast is a constant character, and that the species is

therefore distinct.
CORVIDA.

Corone, Kaup.
108. CoRONE PHILIPPINA (Bp.): Sharpe, Cat. B. iii. p. 42.—Corvus philippina, Bp.:
Walden, ¢. ¢. p. 201.
a. Negros.

[Crows were found in most of the islands; but I only preserved one specimen. The
native name is ‘ Quawk,’ from the note. I saw birds apparently of this species in Negros,
Zebu, Panay, Mindanao. In Negros they were very plentiful about the house where I
was staying, and very impudent, coming down into the yard and stealing the jerked
beef which was being cured at the time.—J. B. 8.]

Dr. Steere’s specimen measures—total length 18°5 inches, culmen 2°45, wing 11:9,
tail 8:0, tarsus 2°05.
STURNIDA.

STURNIN A.
StTuRNIA, Lesson.
109. STURNIA VIoLACEA (Bodd.): Walden, ¢. c. p. 203.

a. Negros.
6. 6. Dumalon, Mindanao.

[In large numbers in flocks in both the above localities.—J. B. S.]
This Starling was not before known from either of the above islands.

Catornis, G. R. Gray.
110. CALORNIS PANAYENSIS (Scop.); Walden, ¢. c. p. 205.

a. Panay.
b,c. 6 9. Maasin, Panay, Jan. 1875. yes light carmine.

[In large flocks in the open fields, feeding in the cultivated grounds. They were tearing
to pieces the flowers of a leguminous tree.—J. B.S.]

111. Catornis cHALYBEUS (Horsf.): Salvad. Uce. Born. p. 271.
a. °.Puerto Princesa, Palawan. Hyes carmine.
b. $ juv. Puerto Princesa. LHyes grey.

The only adult specimen procured by Dr. Steere is a female in rather dull plumage.
It seems to me, however, to agree sufficiently well with a hen bird sent by Mr. Low
from Labuan; but it appears to be a little larger in its measurements than the Bornean
birds, the dimensions of Dr. Steere’s specimen being as follows—total length 7°6 inches,
culmen 0°85, wing 3°95, tail 2°65, tarsus 0°9.

B44 .. MR. R. B. SHARPE. ON BIRDS COLLECTED BY: :

EULABETINA.
Sarcops, Walden.

112, SARCoPS CALVUS (L.): Walden, ¢.c. p. 205.

a, b, c. Negros.
d. 8. San Bernardino, Negros, Jan. 1875. Eyes reddish hazel ; bare parts flesh-colour.
[ I met with this bird in great abundance in the upper part of the island of Negros,
frequenting the groves of wild bananas, and feeding on the fruit of these trees.—J. B. 8.]

118. Sarcops Lowit, sp. n.

S. similis S. calvo, sed interscapulio argentescenti-griseo nec nigricante, et corpore
subtus argentescenti-griseo distinguendus. Long. tot. 12°5, culm: 1:15, ale 5:1,
caude 5:0, tarsi 1:3. |

a. Si Butu, Sulu archipelago.

This species appears to be distinguished from the ordinary Sarcops calvus by its grey
mantle, which resembles the rest of the back, and by the colour of the under surface,
which. is silvery grey with a narrow blackish line down the centre of the body,
whereas in S. calvus the mantle is black, and the under surface also, ouly the flanks

being grey.
Gracuna, Linneus.

114, GRACULA JAVANENSIS (Osbeck): Salvad. ¢. c. p. 274.

a, Puerto Princesa. ;
6, c. 8 ¢. Puerto Princesa, Palawan. yes black, bill red, wattles bright orange-yellow.,

[These Minas were not common in Palawan, but were found in the high trees in the
jungle, making discordant noises. The natives catch them and cut their tongues, in
order to make them 'talk.—-J. B. 8.]

Identical with Bornean specimens.

EURYLAMIDA.
‘SARCOPHANOPS, gen. 0.
Exactly resembling Serilophus in general character, but differing at once in the large
wattle round the eye. The type is

115. SARCOPHANOPS STEERI, sp.n. (Plate LIV.)—Hurylemus Steerii, Sharpe, ‘ Nature,’
August 1876, p. 297. |

é ad. supra saturate cinereus ; uropygio et supracaudalibus nitide purpurascenti-vinaceis ;
cauda leete castanea ; capite summo nitide et saturate purpurascenti-brunneo ; torque
collari et colli lateribus albis; loris albis capitis colore apicatis; facie laterali et
gula tota nigris; corpore reliquo subtus lilacino, hypochondriis vix vinaceis, ab-
domine imo albido; subcaudalibus fulvescentibus; tibiis cinereis; axillaribus albis;
subalaribus nigricantibus; alis nigris, secundariis intimis basaliter albis, extimis
extus aurantiaco-flavis, fasciam alarem transversalem exhibentibus; remige primo

PROF. STEERE IN THE. PHILIPPINE ARCHIPELAGO. o45-

extus versus basin auguste albo; caruncula opthalmica pallide indigotica; rostro
pedibusque pallide indigoticis. Long. tot. 6:8, culm. 0:9, ale 3°45, caudse 2°75,
tarsi 0°85.
? ad. mari similis, sed pectore albo nec lilacino distinguenda.
a,b. 3; c. 9. Isabella de Basilan. Bill, wattle about eyes, and feet light indigo-blue. yes
colour of gold-stone.
- [I only fell in.with this species on one single occasion in the island of Basilan, when I
found a little flock of about seven individuals in the thick deep jungle towards the
centre of the island. They were feeding on fruit, and were very tame, having probably
never been disturbed before by a gun, and did not take flight until I had secured three
of theirnumber. Of all the birds I ever saw, this one has the most beautiful eye. I can
only describe it as being of the colour of “ gold-stone;” that is to say, it was like a piece
of clear crystal crowded with specks of gold. All three specimens had the iris exactly

the same.—J. B. 8. ]
This is the gem of Dr. Steere’s discoveries. The eye-wattle, the peculiar coloration,

the difference in the sexes, render the genus entirely distinct from all the other members
of the Hurylemide.
PLOCEIDA.
Pappa, Reichenbach.
116. Pappa oryztvora, L.: Walden, ¢. ec. p. 207.
a. Zamboanga, Mindanao.

[I saw plenty of these in little flocks of from five to seven, perching on the roofs and
nesting in the latter.—J. B. 8.]

Identical with Javan examples.

Munta, Hodgson.

117. Munta arricaPrLua (V.): Salvad. Ucc. Born. p. 265.
a, 6. Zamboanga, Mindanao. LHyes reddish hazel.

Dr. Steere’s example is not referable to If, Jagori, Cab., according to the diagnoses
given by Lord Tweeddale (¢. ¢. p. 207), as it has a black head. The black on the belly
extends right up to the throat in a continuous patch, this perhaps being a character of
the male only. After examining a large series of these chestnut-and-black Finches in
the British Museum, I have come to the conclusion that Lord Tweeddale’s identifications
are correct; but I believe that with increased material the number of species will have to
bereduced ; at present, however, as Dr. Steere’s specimen is clearly not referable to
true W. Jagori, it must be kept as M. atricapilla, with Labuan examples of which it
agrees best, having the same black head.

118. MuniA LevcocAsTRA (Blyth) : Salvad. Uce. Born. p. 267.
a. %. Puerto Princesa, Palawan. Hyes dark red.

[In large flocks in the rice-fields, where they made great havoc.—J. B. 8.]
I have compared the above specimen with Malaccan individuals, and have found them

identical.

346 MR. R. B. SHARPE ON BIRDS COLLECTED BY

COLUMBA.
TRERONIDAN.

TRERON, Vieill.
119. TrERON Nastca, Schl.; Salvad. Ucc. Born. p. 288.
a. 9. Puerto Princesa, Palawan. Eyes orange.

I have compared this specimen with a Sumatran pair collected by Mr. Wallace, and
find them identical.

OsmoTRERON, Bonaparte.
120. OSMOTRERON VERNANS (L.): Walden, ¢. c. p. 210.
a, b. Maasin, Panay.
[Shot in cane-fields and open thickets.—J. B. 8. ]
Panay is a new locality for this species.

121. OsMOTRERON AXILLARIS (Gray): Walden, é.c¢. p. 211.
a, Zamboanga, Mindanao.
b. Negros.

[Very abundant in both the above places, and called by the Spaniards “ Llorosa,” or
the “ weeper,” from their mournful note. ‘They were flying about in flocks and feeding
on fruit.—J. B. 8. |

This Fruit-Pigeon was not before known from Mindanao.

PHABOTRERON, Bonaparte.
122. PHABOTRERON NIGRORUM, sp. 0.

P. similis P. leucoti, sed gula et plaga faciali pallide cervina distinguenda. Long.
tot. 9:2, culm. 0°7, alee 4°95, caude 49, tarsi 08°5.
a. &. San Bernardino, Negros, Jan. 1875. yes with a narrow rig of carmine;, crop full of
Chil peppers.

[A Ground-Pigeon, by no means uncommon, but difficult to preserve on account of
the looseness of the plumage. It was found in the corners of the cane-fields, where the
red peppers had grown up wild.—J. B. 8.]

On comparing Dr. Steere’s adult male bird with those from Luzon in Lord Tweeddale’s
collection, it appears to differ in having the throat pale fawn-colour instead of rufous,
and the cheek-stripe is fawn-colour instead of white. As other Negros birds in Lord
Tweeddale’s cabinet exhibit the same differences, we may fairly suppose them to be con-
stant; and in that case the species requires a name.

CarpPoPHaGa, Selby.
123. CARPOPHAGA NEA (L.): Walden, ¢. c. p. 215.

a. Negros.
b. 9. Sau Bernardino, Negros, Jan. 1875. Eyes carmine.
c. Puerto Princesa, Palawan.

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 347

[Pretty generally distributed, as I saw it in most of the islands, Mindanao, Palawan
and Balabac, as well as in Negros. It was feeding in most instances on the fruit of the
wild fig. —J. B. S.]

This wide-spread species probably occurs on all the islands.

Myristictvora, Reichenbach.
124. MyRIsTiIcIvora BICOLOR (Scop.): Walden, ¢. ¢. p. 217.
a. 6. Balabac. yes bright red.

[This species was plentiful in Palawan and Balabac. In both islands they were found
at certain times of the day coming down to the mangroves along the sea-shore in large
flocks. What they were doing there I could not find out, unless they found something
to feed upon along the beach. Off Balabac and Palawan were several sandy islands
where the larger Pigeons nested in large numbers; and during my stay at Balabac I saw
the natives returning from an expedition to one of these with a number of young birds
and eggs, probably of this and the preceding species.—J. B. 8.

Hemrenaca, Bonaparte.

125. HEMIPHAGA POLIOCEPHALA (Gray): Walden, ¢.¢. p. 217.
a. San Bernardino, Negros.
[Found in high woodlands in pairs or two or three together and not in flocks.—
J. B.S.]

COLUMBIDA.

Macropyeia, Swainson.
126. MAcROPYGIA TENUIROSYRIS, Gray, Walden, ¢.¢. p. 218.
a. &. Isabella de Basilan. Eyes hazel.
6. Negros.

[Shot near streams. They seem to frequent rocky ground in the neighbourhood

of water.—J. B.8.]
I have compared the above two specimens with others in Lord Tweeddale’s collection
from Luzon and Negros, and could not find any difference in the Basilan example.

'

TurtuR, Selby.

127. TurTUR DUSsUMIERI (Temm.): Walden, ¢. ¢. p. 218.
a. 2. Maasin, Panay, Jan. 1875. Eyes brick-red.
[This species is found in the open fields and roads in pairs upon the ground.—J. B. 8.]
Panay is a new locality for the species.

128. Turtur HUMILIs (Temm.): Walden, ¢. c. p. 219.

a. Mount May-hay-hay, Luzon.
SECOND SERIES.—ZOOLOGY, VOL. I. 3A

348 ' MR. R. B. SHARPE ON BIRDS COLLECTED BY

GOURIDA.
PHLoceNas, Reichenbach.
129. PHLOGaNAS LUZONICA (Scop.): Walden, 7. c. p. 221.
a, b, c. Manila.
[The natives snare these birds in some numbers; and all my specimens were procured

from a friendly priest, who gave them to me out of his aviary. I saw the same or an
allied species in numbers at the north end of the island of Negros.—J. B. 8.]

CHALCOPHAPS, Gould.
130. CHatcopHars tnpIca (L.): Walden, ¢. c. p. 221.
a. 9. Maasin, Panay, Jan.1875. yes dark.
b. g. Puerto Princesa, Palawan. THyes dark.

[This Dove seemed to me to be a common Pigeon in most of the islands.—J. B. 8.]

GALLIN &.
PHASIANIDA.

Gaus, Linnzeus.
131. GaLLus BANKIvA, Temm.: Walden, 7. c. p. 223.
a § ad. Panay.
[I also saw Jungle-fowl in Mindanao, where they were plentiful, and also in Palawan
and Balabac. They are difficult to shoot, as they are very wary; and I have hunted for
hours with birds crowing all about me without getting a shot. The Indians of the

forest-villages snare them, and often keep them alive, crossing them with the domestic
fowls.—J. B. 8.]

MEGAPODIIDA.
MeEcApopius, Quoy et Gaimard.

132. Mrcapopius Ltowtt, Sharpe, P. Z.8. 1875, p. 111.
a. @. Dumalon, Mindanao, Oct. 1874. Eyes hazel.
b,c. pull. Balabac.

[In Balabac I found large numbers of these birds, and procured both eggs and young.
They were nesting on the sea-beach on the edges of the brush, the nests being
about 12 feet across and about 8 or 4 feet in height, and made of sand and bits of
shells and sticks. The natives. say that sometimes 50 or 60 eggs are found in one
of them. In Mindanao, in the mountainous country, some way inland, I found
heaps of earth piled up against fallen logs, which the natives said were Megapodes’
nests; I dug into them myself, but did not find eggs. On one occasion the nest had
been burrowed by a huge land-lizard.—J. B. S.]

The Megapode collected by Dr. Steere is not the true MZ. Cumingi of Manila, but
appears to be identical with I. Lowit of Labuan. It is a very dark-coloured bird,

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. 349

leaden grey underneath, and measures as follows—total length 12 inches, culmen 1°15,
wing 9°38, tail 3°6, tarsus 2°55. The dimensions do not agree with those given by me
(P. Z. 8. 1875, p. 111); but the dark leaden-grey under surface agrees exactly with
Labuan birds.
GRALLA.
CHARADRIIDA.
CHARADRIUS, Linneeus.
133. CHARADRIUS FULVUS, Gm.: Walden, ¢. c. p. 226.

a. Balabac, Sept. 1874.
RALLIDA.

OrTYGOMETRA, Linnzeus.
134. ORTYGOMETRA CINEREA (V.): Walden, ¢. ¢. p. 230.
a,b. g 9%. Naga, Zebu Island. Eyes reddish hazel; feet olive-yellow.

[These birds were found in a grassy wet marsh; they were’ actively running about
through the grass and water, flying short distances, and alighting on tussocks. They
were in great abundance.—J. B. 8. |

This is the first exact occurrence of the species in any of the Philippines; though
specimens were obtained by Mr. Cuming in the archipelago, no island was recorded as
the habitat.

CICONIID.
MELANOPELARGUS, Reichenbach.
135. MELANOPELARGUS EPIscoPUS (Bodd.) : Salvad. Uce. Born. p. 356.
a. Negros.

[This Stork was killed out of a flock of fifteen or twenty individuals as they flew low

over the mangroves towards the high lands in the interior, and apparently on their way

to roost.—J. B. 8.]
This is the first time that the species has been noticed in the Philippines.

ARDEIDA.
Busuucus, Pucheran.
136. BuUBULCUS CoROMANDUS (Bodd.): Walden, ¢.c¢. p. 237.

a. Puerto Princesa, Palawan.

[White Egrets were noticed in the rice-fields in nearly every island.—J. B. S.]

Buroriwes, Blyth.
137. Buroripes savanica, Blyth: Walden, ¢. c. p. 287.

a. Negros.
Gorsacuivs, Bonaparte.

188. GoRSACHIUS MELANOLOPHUS (Rafil.): Walden, ¢. ec. p. 288.

a. Dumalon.

oAQ

390 MR. R. B. SHARPE ON BIRDS COLLECTED BY

[Found at some distance from the water, running along a road in the forest, where
there were some shallow puddles of water.—J. B. S.]

LARIDA.
STERNINZ.
STerNA, Linn.

139. STERNA BERGII, Licht.: Saunders, P. Z.S. 1876, p. 657.
a, b. Balabac, Sept. 1874.

[In abundance, flying round the coast of Balabac in flocks.—J. B. 8.]

The specimens were seen by Mr. Howard Saunders; and he pronounced them to be
of this species.

The following Table is drawn up on the model of Lord Walden’s, and embraces all the
species known to inhabit the Philippines. The new species discovered by Dr. Steere are
here printed in small capitals; and those which he has added for the first time to the
Philippine avifauna are printed in italics.

Philippines.

| Polynesian Region.

| Malay peninsula,
| Australian Region.
|| Papuan Region.

| Panay.

| Zebu.

| Camiguin.
| Mindanao.
| Basilan.

| Malamaui.
| Cujo.

| Palawan.
| Balabae.

| Sulu archip.
| Celebes.

| Borneo.

| Jaya,

| Sumatra.
| Timor.

| Tndia.

1. Cacatua hematuropygia...............- a
2. Prioniturus discurus.................-. 5
3. Tanygnathus luzoniensis ..............-+ no
Cyclopsitta lunulata..................6 30
Loriculus philippensis.................- ae

9 TEAWUS)s os oo cGo0dD000000000G0 Pe Bloc

3 ORO, oo00000900000000000 oA[o allo olfoo

99 WATT 6 cong o0g00n0n0000008 sallsailorolloallae

i chrysonotus).))..)ece le - ee o.dlloa|lolle alloo
Wal cousevertisn was sececnaisctoetee tater: aol een lteol cles
. Microhierax erythrogenys is
> ZNSE THMIVAUSFNTS) 5 6505G0000000000000bnN0 #
pm. SOWIE. cosccccansca0go00005000 %
5 ANCE WARM 3 2g00550000000000000 .
. Spizaetus philippensis.................-; As
. Spilornis holospilus
p IBMT THON 55 4a56509000Gn00b00900
. Haliaétus leucogaster
. Haliastur intermedius
. Klanus hypoleucus
21. Baza magnirostris
22, Circus melanoleucus :
23. ,, spilonotus
24, Bubo philippensis...................00-
25. Scops megalotis
26. Ninox philippensis
27. Scelostrix candida
28. Thriponax javensis
29. Mulleripicus funebris
30. Chrysocolaptes heematribon o
31. ” xanthocephalus.......... 30
32. oy lucidus

x x
5 ee
Eom

#

Lok * | Guimaras.
: & *% * | Negros.

> & & & | Luzon.

*
*
*
bo
kh

To
SSRN STR ONE SOON OU
LR Re eT

#

%
eR
Ho RO

#

*

2%

34, ERYTHROCEPHALUS ......|.. dolla alle allo.alla allo olla allo ollo offo- off 3
BD. Tig@ Javanensts ..... 0... cece cee neeenac cles
36. Jungipicus maculatus ..
Oipmulannactesiard cuore rence niet rerit te
38. Merops philippinus
32. 4, _ bicolor

ae =

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO. abl

Philippines. a & é| <
A A 2 a a %
a EWEN lel VAI SIEW ei ay Wel 8.2/5
25/2) 2) s\2)5| 2) 2) .|2lS| 2/218) [Eel slel ele):
5/5 /2|2/3/8/2| Zl eislalelals leis lais|alzlelsie
ARS) aisl/Olalalalolalalalolals jaja lala |4| ala
. Eurystomus orientalis ..........4..--. wel xe x |e le ele | oe * wlelaelalelelele
. Alcedo bengalensis........ siotehaied Bs sper: walle n]e [ae ]e feo] x as weleelelalalelel se |
. Aleyone cyanipectus ........... dsio.o ce %
3. Pelargopsis gouldi ............+....... ~-|%
> gigantea ......... Uboonouda cafeefe fe c|e ete te [ae fe cle ele ole ole ol xe
on leucocephala......seveeeeees calls alla c[oalle-ollo allsolfo clle alla allo alles |
pCeyxc rE, 58o55cqub0duub 00000000 calks ile olocleolloalloalies |
oy TEORCGUD, cogannosdon Usa oonagee Halen cts cect alle ate os cle lente ele fool «lle | ae |x |. «tae
» philippinensis ...... vooosodsgbee - «| x
. Halcyon gularis .......... bon oodocd ot ela fale
ULE coanacooUudo Arevads cored’ *
» coromanda ...... Eiainrlahe renee *
59 chloris ....... Boisetitancte ge tieh ony eeloe leet lala}. ele |e
. SWANCHELIL | cial sveyerctals stele a siete wefecfe ete efecto ete ol= «| se
Ps ombroninemm mer iecireyastr colle aiiagiio dliaciloollac| |e
: is JOHAGRENT, oon eugcdncncaogeccus Bae |
. Santholema hemacephala ...........- sly vale] x
: a TOSCAM Eira rtene heleysloraloetobst ate cclleulercllo oles
. Macropteryx comatus.................. wef fee} ole * |e]. .|*] %
. Collocalia troglodytes ................ ole |e
. Lyncornis macrotis......... oobuooooeaD ole 1
. Caprimulgus manillensis .............. Beles “ #
$n ise Gegoqooou50s00d5 *
. Cacomantis merulinus ........ RO ores Oalltd Pa\les|lo allo alloallcallaallo alle»
. Chalcococcyx amethystinus ............|-- *
. Chrysococcyx basalts .......0e eevee eens |e n allo olloall es
. SURNICULUS VELUTINUS .............-|++ sallecliocle llealle alla-alioolles
. Hierococcyx strenuus .......-.......- aoe
b op [eee] Fasccoocogonge0o *%
. Eudynamis mindanensis .............. wale efaefae fae fe |e eda
. Desylophus superciliosus .............. os | xe
Lepidogrammus Cumingi..............- «le
. Centrococeyx viridis .................. welacle el} | elle ele
. Dryococcyx HARRINGTONI .......... oro slate oallo elloal lal ection lana cea eretle3
Pyrrhocentor melanops.............+.. Sallartic olla clic olla olla oll eslts oll es
. Buceros hydrocorax ..........00++005 calles j
. Craniorrhinus leucocephalus............ 6 lle allo ollo cllo ollo ol [Eales i
Ry INV/ATED ENT ae ater lal-eietelel oes colon |e
. Penelopides panini.................-5- ele lela x
} op MEMES 6 GooeGanddcocec wa feede
} ILENE REBT! Soocboo0a0g6000000C0 Gallo oljoallfalle olle alles |
9 schach . tie tle alten eee ee eeee nee #
», lucionensis ..... DO AOE TOO RCS tio Se ae ee ae ea (ae
. Artamus leucorhinus.................- colacde-lalade ete -lale-le> wxlelaelale-le dels %
. Graucalus striatus ......)..02.-ce eee es edlesloaic slesleeioelesio alle
i ue SUMAETENSIS occ e cece scenes Jalloollodieclloale alloca atloolle clio alles
. Wolvocivora cerulescens .............. alse jlo
. Lalage dominica eee wee ew tere ewe eee eof] HR] sede Le fe ef fee] ele ele fe ele ele ele Paes Kyle sfe ole Ve
. Pseudolalage melanoleuca.............. ole
. Hyloterpe philippinensis .............. --|%
. Pericrocotus cimereus.............20005 Alls alloallactio alle ole clla allo elle allaolle allo clo aloo eatlaniea les
é + UGNEUS: tet AA areh ah sPayecchaveeecn seat: delle aloollecleollocleclscloelaciocl fala cloniocllestaicila cles
. Dicrurus balicassius ..........0.0+000- Boies
Bs mirabilis ...... araedarshisrats aeelocatacs dollo olloclfe allie
« Buchanga cineraced 1... eevee veeneeeee salle cl[ealla clio clloollectlaa|is alle clo alles
. Philentoma cyaniceps ..............-- sallestoollocie ooo cles
b PP ABINAORABS coacoasconodoo Ballo alls oles
- Leucocerca nigritorquis..............-.. sllfsllodleale olleslleel|eello ello allo oll es
. Cyornis philippinensis ................ wef alate e]e | ae
. Hypothymis azurea .................. Salts |loolles lee lle clloclle alles|locilacilaclioollo alloallealtes||ealESilo all £2
% SUPERCILIARIS............ pollocicollo cic clac|laelloclies
. EUMYIAS PANAYENSIS .......0.000.002 solle sllee
. Butalis manillensig..................6% Bela
. AanthopyGid NArctssind . 1... .veeesaeees Jellocle ile clocliaallocl|e3
. XANTHOLESTES PANAYENSIS .......... Be) loo
A SE PARTAWR UTC AUDA) se) </dsteleyo1 4 siete crosses BallNollbollecoiio allaallen| tac lee
b ADENINE HUNKS 5 GogopeoguuHueoudouoc le
'

MR. R. B. SHARPE ON BIRDS COLLECTED BY

Philippines.
z a/8|.|3 Ae
Jel 3/3) 18/8] 8l 2 slslzleis| |é
aE: SBA HEBEAEOE
AJAIS(A\8|S5/4/S ls l6lalalalslals|a
107. ZEoCEPHUS CINNAMOMEUS ...........-- calleelico|lp ilo dlloallerg|loolle
108. op CYANESCENS .........-.--- selloHoolloclle sto Wo elloolle olla allo alles
WO9. Berundo rustica . 00.5. c en cence sec csee dol latallaiollacollpalloval enlle™ ;
110. ” BMV), 6 cog ood oocnd05n0e. Solloa| alo oliodipclnelaoloclieciioaissllsallaallesi|=s|lasiies
111. Cecropis daurica...........-. soo0eccoe «lig Seale
112. Oriolus chinensis........ goaderegabende Solles|le allsl celica hall ealle allo allo ollo cll =
1B 5) SMIIMEDTs cong congeaagdoxcce00c Melleclloaleellerlecleellectiealloellecte cls nes
114, oj PUMAIDIGRL oS Hacamgodooecoadeee oellb olla oleic lo alle pllo oles
1B, og, olay Sep oo obconax0¢00K0 %
116! Durdus obsemrus). shes... eden eee snl Re
117. > chrysolaus).-. 2... Cee Gece *
118. Brachyurus erythrogaster ..,........... soll es
119. 9) PROPINQUUS ...+.ssseeeee- pollee|io clic clo clic cla cll Sailo olla allo cic oltex
120. n SOWGWGWS ooaoaondooggenoace salaeiia als alae twstae te cl als ale ll ae
121. 5 STEER ...... Aeharic Chunagsts arllaRerateraltteltexa em itae
122. MACRONUS STRIATICEPS ..........+... 00) o6|o alo alio alle ole el[o of ell ee
TOR WooxorENAs| NWWOOWE ooocosgapac eo cascade 2 ele le PP ka
TOM, [Parra Oyonerenc/A TVUE(OYNIWN 5 oc coon oD OCaeOO aaliaaliociiaceeliccleelcallo alls allaciles
125. DENDROBIASTES BASILANICA ......-... Pelle ollacheciecllec|iacloolles
126. Megalurus palustris ...........-..5-.. > all
127. Crateropus caudatus ....... ocononroseS %
198 ImEwAUDWEEDDAGIE . $0... cece h ee rle celle gical celica colle elloalloollas|lo oll =
IPE, 5 OVRIMOERISIMES goo accuocens0no0KG ~ ole |e
130, ,, MBLANOCHLAMYS ...........- Salamon) ele dio dices oloolle3
131. PHYLLORNIS PALAWANENSIS .....0----|--\..}e-|..fe-Jeelecfee[e ele ede |
132. Tora scapularis ...... He OOITOE Ge Btothe oe calla alioctla cle alle lle elle alle ollo olfo ols
133. Ixus golavier ........ non0D0CCUoONGDCS sll
B45 0) SWeRSIS Beale gnoonace o900000n)oa||
135. Poliolophus urostictus ......0..0- odo o}]-a\lo ollo all =
136. CRINIGER FRATER........ aocacooce Daelloollo alice allo allo cle cia lf olla ollo olla oll ex
137. Hypsipetes philippinensis.............. sof ae le el ae fae | ae
138. RUFIGULARIS ..... Re cpt ne laa a neallee | xe
139. Monticola solitaria SA ede CaREE OG oe oc wl el oe Sflecllec|[antto olfeclloalfe oll alle of elle
140. Pratincola caprata aes aren ates cra adeeotel stars bis Ps ele. Se PS gel aal aera a ae iy males
141. Cittocincla luzoniensis ................ ciel
142. 9 NL GURVAG ole fey artustey ets aane tenc eecuetele oollacliogl ouallbelleeleoalGrallor alle ells alte
143. Copsychus mindanensis.............. oalecle elle *
144, Calliope camtschatkensis .............. *
145. Gerygone simplex .............. on0000 > oll es
146. Phylloscopus magnirostris.............. allo lol's alle clic lic cloalic elle olla cle cilaalle ole nlic ol ca lloolle
147. 9) UMHS osoonacoovevedos Prllnelleetie alinelie cler| (2
148. Acrocephalus orientalis................ 3.0) e olleallo oll 22
149. Cisticola semirufa ........ gaootorceace ues %
150. Orthotomus derbianus ..............-. *
151. 9) CHUANG IS Gooocenoceeoce wf ed ae | ae
152. 9 TUMGTES. Geos ouedecoessane echo clecte cle ele cleelioolie clic allooll es
153. s CINEREICEPS.............. B.a[s clo o}\s allo oleeli= ale ollcs
154. a INZONAY NTI Alc bGanosconcoat a ollc clio olla allele clic ell ail ee
155. Budytes viridis .......... La nee Heya ce Neve RecA TTR Te 1 Ra get ag ad
156. Calobates melanops .................. ool ss * eles
157. Motacilla luzonensis ...........0++.0. *
158. Corydalla lugubris...............00005 celecllachealles
WSO Anthus Gustav. v0. secs cue neces ve ches Soon Sole le aleeheioelles
AGO;sBarnselepamsy s 2) e)siaevatec seman delehels %
IGN, gy ARUBA co san one ELS net LE A Bs OR Te Ve eat ll aca etal
162. Dendrophila frontalis................0% pee ltimetion o
163. H GNOCHLAMYS ............ palle alles i
164, Zosterops meyeni ....... Sere ece ee tera welled slates
165. Myzanthe pygmea.................... wl gelln of ge
166. Diceeum retrocinctum ........ dgrooece Selle eels alles
167. D9 jAYMUEWES ooocoupanneonocaauge a ea ee a eg *
168. y H@MATOSTICIUM ............ ello ole alleles
169. A | UrarOysmOCWENe Cagodor padoueouD aA tea bee %|
170. yy WON 5 soneo0 ag 0c aga0accas oolle olla ;
171. UGCROLUTIO oaocaccaccovccoo PASS ME
172. Prionochilus, Ro soaodvegaadatbodsHeces|(on Pee Maa eel el Salle cllsctlecalles
173. ARACHNOTHERA DILUTION ............|.. ARR be Pasa lec i s|loulls

| Malay peninsula.

| Australian Region.
| Polynesian Region.
| Papuan Region.

| Timor.
| India.

PROF. STEERE IN THE

PHILIPPINE ARCHIPELAGO.

Philippines. a
5
: | 3S 3 & A
| lelel (eidlel@l lelellalel (2/2).
3/812] £)2| 2\4|4|21s/8|2|2|2| 2 <#lZ/414
4/8 /s [A/S 6/5 |4/4]6/8)4/2/6]/8|3|a) 4/6) 4
174. ANTHOPYGA SHELLEYI ................).0).5 calla * |
175. = MAGNIFICA. 22.0. .ce eee wee lee lee ole
176. EUDREPANIS PULCHERRIMA............).-J.-].. bleed *
7c. Cinnyris speratus) (os. «Jacl > | - x|--| ale
178. 5 FURIES cose oosocogussasonalles pale sleallealleslloniles
179. Anthreptes malaccensts .........00.+000)--|e-].- pallesilocticailon ls
180. es CHLORIGASTER) 2... 500..5.]s-|-s]. 0]. - re
181. Rhabdornis mystacalis ................].. %
182. Corone philippima ...............-0..0/.- Pa ee ee | x
183. Acridotheres cristatellus ..............).. *
14> Sturmia: violaceay A nlss 5 cider. pss aicin’ cin oelaelioe Slo siies|lodllooilesiie |x
185. Calornis panayensis ..................|-- xl e|--|ele
186. s GiHh|ETS 55-s580aceresoses cog: allo {baile oltosal le Aes
TES /, SPSS GENIE co Suton nesocococude se. sla] ele]
188 3 OweHtide sesotoouuaocen seu O65 Iealealle 35 atoll a ode |g
189. Eulabes javanensis 2.0... ce eee veces eee eee salle mal oO ioral 2
190. SARCOPHANOPS STEERT ..............].. alle Bille x
HOls Passer*montanus.%..% 6.0. cde we eee oe z|--|. BeGolleallad thine lagi ore #
NODS Pade ay OryZiVOEM) wc las aleiclalsicieiem <loereselae fie -|. salle oll silo lelelalels
GB, Wikies Feo Soo soe goonsbonUsoNee soso x]. a
TDs by elmer, coo cose uo ooooet aH ae aesllas calle ol|ee
195. ,, TMNT) 5 Sar od boeine OUT SeNS SO so *
GS ee LCUCOR AS LIeum ielteteyovereitereleleleleled-lo!| ee jae -|x
WOVE OxyCerca Jago. . hase. oes sows ce eeen| a> *
IGS, IraWe TEOSIe a Gen decocsooouDosacdao ae slodlo ale 5 eee se *
199. Osmotreron vermaDS ..........0.0+-00- elelad--|.. as 50 zlelele|..|=
200. ” cpa Peat heels Seases EAS ace acto ais Ea od ie oe “|x
OOIS Weucotreron pironiert. <.).. sje ~ 2 ee oe = xl..lx
202. Rhamphiculus occipitalis..............).. %
203. Phabotreron amethystina ..............|.- %
204. 0 IGIGONG Co ocanondoseceDcusos &
205. or NEG HORMONE | Pyeleseyorcies ster ekeeee| Seni Jee |e
206. Carpophaga cenea .........0..0 eee eeeles PA slo allele ol ger lets es « glelxl.-|x
pA | Bike ee SIAL Slecbgde ciel eqectanie ai
208. Ptilocolpa griseipectus ................|.. *
209. Myristicivora bicolor..................|-- Ne slersl esl clxlael. ele] a] e|--lel-
210. Hemiphaga poliocephala ..............].. Pao eee
211. Jantheenas eriseigularis................|.. #{--|e-]
212. Macropygia tenuirostris................/.. x|--|--[e|- we
AIG, Aime GiechiniOul oe po stub bouoc ue bbl lan xlael--l
D4.) 5, InpiemiiGy SoGoodeencaopouerougs os }e ede
215. Foleo raezonr cays 154.2 pce stancetots wet eile x
216. Chaleophaps indica ..............2..5[-: xlel--|xl- % a ae
Bis Calisstdseotiarieal ots ects sle 1c RL PMI HN Lk aa S.
218. Geopelia striata .... 0... .ceeese sees ees #|--|..]..1. =|
POG allusybanlkdivene sey etcrcvcnetajcce lop: sivebax siete ever ee xlels|- --|*
DI0s Arharophita tsps pee tala: let-teiclafercieiata see % 2
221. Excalfactoria chinensis................ *% wlale|aele-|s
OVP eurnixocell atari tose teleice cee % -
22E. IOI, A, oo SereooadocHodaduccdood %
294, Mecapodius Cumingi...............00. x
225. 5 It dork oS a=ppo bose sede coldalla 4 as sles
2G, Chimchisuwlias) joouaddocusaaeacdc0sllse Salsllesy (6 Aa fee fae]. fae] ae | se |e
327, Sam EnOMINSH OAD bop. caneusgeavn lodliecltclk let & al sae
298, Aigialitis Geoffroyi............2.ee00-s = salle Re dleelles|les
OMIM dubia; 0050 as theca atom ome: y alld
230 or WOME, coooocoapoocecaccc Palas ne wefaelafee|- |. fae
231. Esacus magnirostris ..............000- False : Rolls ¥ #| 3 Dalle
232. Himentopus autumnalis ..............]. aly ce sale Ba lelleal incl eslealles
233. Himantopus leucocephalus ............|..]..].. Pas «le. x lel «|e % Q
OpuL, Cibmenik, Quemalis! saogoouooconeaeueuKc F %* all salleatleallecllocleslle
235. Porphyrio pulverulentus .............. x RNG
236. Gallinula chloropus .............+....|.. * Me salle aio alles
Bevin (Colliers GME, “GogcooncnuegonooendG x Ae sollodljo alte:
238. Hrythreea phoenicura .............+...)-- te aal{sol| es x |e]
939. Ortygometra cinerea ...........2-++---|--]--|-- eNealloc salle:
240; -Porzana pygMea .....-.-.cceeeeesaee x. 2c :

353

:|¢
5.2
81) 3
21 2|8
alle
i
2).2\8
s| 2
= a
zg
E|als
Ellie:
4) |
dl
a |
|
|
= * |
vole el ae}

MR. R. B. SHARPE ON BIRDS COLLECTED BY

* * *

1
Philippines. g
: A
|| |ele]./8 Sl el alc alle
és) alE/s/Slel2(zlelEla eiclelelzia
oa Q Ala] slolsa io ie | s
E lelsl2is|é|alalSe Ale G lelaelz|alé
PML, Vetoes WEEE 5g oooueobODeoHNDOODDDO # | oes eae
242. Rallina fasciata ........... baddoncce | ox || | ae
243, Amaurornis olivacea ............+005 +] x
244, Typotenidia torquata .............. *
245 os philippensis............ Foor xlecfeferfe-
246, a HAPEND soooncacgo00da0e. <- || x «|e | |e] Rae |e
247, Hydrophasianus chirurgus........ pooa)ooll . sefeetae fects
248, Numenius pheeopus ................ wef ee +| % % | ele]
249. Rhyacophilus glaveola ........++..--/++| % a Paella
250. Tringoides hypoleucus .............. Boller x lee | ele].
251. Totanus calidris ................ pocolt3l] oc x fae laelec|ael-
252. n CamescensS .........06- Rikieie Tats Boll 6% lal a&lelecde-
28. Dring rutCOllisysys ese) iclo+ slelelels ss e\e\e)° | x sfoe | ae | ae |e
254. Gallinago scolopacina............+ on ol[ool| & safeafe de
259. megala ...... SodOdOOHUGG 4 ++] & *
256. Rhynchiea capensis ...........- oagalesliiios
201. Ardea PULPULED . oe sence ocicie ewan ot |
258. Ardetta flavicollis .................. #I oo.
259, 4, += clmmamomea..............-- +]
260. on... HIGHS Sha bono Rd 0b OaCK0 000 ey 36 s+. 0 in
261. Bubuleus coromandus ..........+.+- -o| x +l ae]. * |: #
262. Herodias garzetta ....... poodooeca0s | x Sol aig .
263. 3 Intermedia .......... i nreiene Paleo . Pal -
264. Butorides javanica ................ ef] # etfs | ae
265. Nycticorax manillensis ...... seeeeee eles] oe
266. 8 BRMGINS oconos000000000000 o| sla le] e |
267. Gorsachius melanolophus ............ ..[ e- sof ete] a [el alls
268. Melanopelargus episcopus ......+. Gooooo! & +|
269. Querquedula multicolor.............. ++] x
270. Anas luzonica........ badooodccodedg)ool 6
271. Dendrocygna vagans ............+... +] % “hee
| 272. Nettapus coromandelianus ..........|-+| ae ap
273. Puffinus leucomelas ................ | x
| 274, Larus, sp..............-% sonoo0d00GS | #
275. Hydrochelidon leucopareia .......... | x H/e |e
276. 90 Ss o0a00000 seseeceesailnsl|
277. Sterna Bergit ............ SHaoKodey S safe fret ete ete eta ate cle e[e ale cleo] a
| 278. Onychoprion anesthetus ............ wef we fate cpa ete ef cde fees ele ete ele o]e olae Lae |e oe] ae].
| 279, Anous stolidus...... apooacumoAGBACOD %
280. Podiceps philippensis....... pae000000 #
281. Pelecanus roseus........ g0000000000 sfeo]
282. = philippinensis ............ ef ae Jee fe efecto e |e ete ete cds cfeefe efecto cle ate el ae dee] ae |e «
283. Dysporus piscator .................. * safe ete cfe efecto ete cle ete fae ]e ete -]e
284. | BWBs.ccoopssocac0s0sdesqan (O° + el ele |e | *
| 285. Phalacrocorax, sp........+--s+..0es+ *
286. 9 oes OSU eo eE | #
287. Plotus melanogaster ................]-- ge fe cfe [ae fe cde ete cfe cde cle ele efe alee] ae | ae | ae [ae fe ete
Motallyrrrteeie 49) 187 |31)31)59|19) 2 |48,23) 8) 5 |31) 9 | 4

* To the above add Brachyurus Kochi from Luzon, making therefore 138 from that island.

| Australian Region.
| Polynesian Region.
| Papuan Region.

| India.

*

PROF. STEERE IN THE PHILIPPINE ARCHIPELAGO.

DESCRIPTION OF THE PLATES.

Puate XLVI.

Fig. 1. Chrysocolaptes erythrocephalus, n. sp.
Fig. 2. C. maculiceps, n. sp.

PiatE XLVII.
Halcyon Winchellii, n. sp.

Pirate XLVI.

Fig. 1. Zeocephus cinnamomeus, n. sp.
Fig. 2. Z. cyanescens, nu. sp.

PLATE X LIX.

Brachyurus Steerii, adult and young, n. sp.

Puate L.

1. Phyllornis palawanensis, male, n. sp.
Fig. 2. The same, female.
3. Pitilocichla falcata, u. sp.

Puarte LI.
Fig. 1. Irena Tweeddalii, n. sp.

rx

a
oe

cw)

. I. melanochlamys, u..sp.

Puate LIT.

Fig. 1. Cittocinela nigra, male, un. sp.
Fig. 2. The same, female.

Puatre LILI.

Fig. 1. Dendrobiastes basilanica, n. sp.
Fig. 2. Parus amabilis, n. sp.
Fig. 3. Dendrophila enochlamys, u. sp.

Prats LIV.

Sarcophanops Steerii, male and female, n. sp.

SECOND SERIES.—ZOOLOGY, VOL. I.

oB

ARTI. ant ,

sit

| a Dba sic ne Bae
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Lye bay cate a alisha Neat ‘Sites a ae .
é i ae iran Seen a Aue risen car

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Wa te aN em hu reins ihe haa) 3 of Sisiion <—
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sai Be it MOREE a s git inte
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an cu snag find shyiba, sola oe
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ie al i al ‘pbelpamnhnry sonar a ee
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Raney

| j Tee
‘ nae ; a. oth ¢ states tire ne ei

i . }
| . . i

AE lO FADE OE—

RB Sharpe. Trans.Linn.Soc.s

1. CHRYSOCOLAPTES ERYTHROCEPHALUS.
GF calemans hth Z.C MACULICEPS.

TRANS Linn S¢

oy

fs ad

IER Z

ZOOL VOL

HALCYON WINCHI

ea

M&NHanhart imp

RB.Sharpe.

JG Keulemans hth.

EPHUS CINNAMOM!
2.4.CYANESCENS.

a

ay

Trans.Linn.Soc.SeR.Z.Zoou Vou. |. Pu 48.

Use

M&N Hanhari up.

walpy TIAA LS SQOUNAHOVE

nayfnip

6) Td] 10 1007 7 YAS 00C'NNIT SNVE, ‘eduey gay

-
=

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Sharpe

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Z. PHY LMORNIES
3. PTILOCGICHILA

PALAWANE
LCATA

NSIS

1S. LINN SAG.

¥
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a ; Pate) |
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1M ‘
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ne ‘ M4
;
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al
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RENA TWEEDDALII
MELANOCHLAMYS.

M&N Hanhart imp

i}
4

.B.Sharpe. Trans. Linn.Soc. 5eR.2.Zoon Vou lPu 52

J.GKeuylernans lith. CITTOCINCLA NIGRA, é&? Mé&NHanhart imp

RBSharpe.

JG@Keulemans hth.

Trans. Linn.Soc.SER 2. Zoo. |

Z.PARUS AMAI

DENDROBIASTI
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HS BASILANICA ; M&N Hanhart amp
DENDROPHILA NOCHLAMYS

RB Sharpe. Trans Linn.Soc.Ser.2.Zoon Vou 1.Pu.54

J.&Keulemans hth SARCOPHANOPS STEERII. ¢ &2. M&N Hanhart imp.

THE
TRANSACTIONS

OF

THE LINNEAN SOCLETY

OF

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.

PART THE SEVENTH.

LONDON:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

“ M.DCCC.LX XVIII.

Pet APT 3B)

Pa AS ie TS Ste TS

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Oe, CANE ASE ETE: Oe PRY eae ly

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ARP TAC

ve ¥

CONTENTS.

PART VII.—Aveust, 1878.

XVII. On the Male Genital Armature in the European Rhopalocera. By F. Bu-
CHANAN WuitE, U.D., F.L.S., ¥c. (Plates LV.-LVII.) . . . page 357

XVIII. Morphology of the Mammalian Ossicula auditis. By AtBan H. G. Doran,
F.R.C.S. (Communicated by Professor Fiower, F.RB.S., F.L.S., Fe. (Plates
TEAVALLI IN 3 DD. St NY) |e eek SS haere Cree uae aS ora Rl) Sr /

XIX. On the Annelida obtained during the Cruise of H.U.S. ‘ Valorous’ to Davis
Strait in 1875. By W.C.M‘Intosn, WD., LL.D., PRSS. L. & £., FL.S.,
Cj; (elate IXeVve and. 2 pWOodcuts)iu ys en a) 2 ge 4) ees 2 499

XX. On the Sacral Plexus and Sacral Vertebre of Lizards and other Vertebrata.
By St. Grorce Mivarr, Sec. L.S., and the Rev. Roperr CiarKke, F.L.S.
(lanes XeVily Cee Vvalieeand OkWoodeuts\Ma seas 42 2 =. = ols

| vs ol OS ee A Oe

‘ k Jae h Jen 17 5 ii ; f j ELEN
a
x A F u A
«th Wo MT ane ell piety Uh Sah, et REE hae Dai lta aay

\ bi tps ; ' ‘ , ro x
i AES ONE.” PET eatethe cr SAA i eee OANA :

‘ : Me
: Qh 8 Pe ree ar SOARES a. Sovak ea Barth) ,

fj
HaAy MLCRO UE Re OS eR ORR Via ies AS ‘a iNhaae a
eas hy AEA AAS CO choke Oe ae
ae : Cah aeimanty % bine 4 ae ee el og
Ge
r eS t ay NE ANCE ’ Y Sea Ny 4 3 tes
TO Ue RR MTT ACR CCS A ME eT 6
ne ty svohiao te. Hh Mine 14 aS V2) ae ;
~

[ 357 ]

XVII. On the Male Genital Armature in the European Rhopalocera. By
F. Bucuanan Waite, I.D., F.LS., Sc.

(Plates LV.—LVII.)

Read December 21st, 1876.

IN several orders of the Insecta the structure of the organs upon which the perpetua-
tion of the species depends has afforded in very many cases the best and surest cha-
racters for the discrimination of species. But in the Lepidoptera, one of the largest
of the orders, and that on which perhaps the greatest amount of attention (scientific or
otherwise) has been bestowed, the structure of these parts has not only not received
the consideration it deserves, but has been almost entirely ignored *. Moreover those
lepidopterologists who have not altogether neglected to examine this structure have been
of opinion that it affords generic rather than specific distinctions.

- With this opinion, however, my observations prevent me agreeing; for I think that
not only generic, but, in many (if not in every) species, good specific characters are to be
found; and to show that this is the case is my object in laying before the Society the
following notes.

In them I have confined myself to the European Rhopalocera. Whether every family
of the Lepidoptera will show equally good characters my researches have not yet been
extensive enough to enable me to decide; but from what I have seen I am inclined to
think they will. The distinctive characters do not always lie in the same parts, nor are
they equally perspicuous in every group; but in one part or other, if not in all, some
distinctive character is to be found, and, allowing so much for individual variation, the
difference of structure is constant and appreciable.

An examination of the hind body of a male Trichopteron will show that it is furnished
with certain corneous appendages, protruding from the interior of the apical segment,
which appendages have been termed the ‘ appendices anales.” Of these there are
normally three pairs—the appendices superiores, inferiores, and intermedie.

If the hind body of a male Lepidopteron is denuded of the scales and hairs with which
it is clothed, somewhat similar appendages (more or less covered externally with scales
and hairs) will be seen protruding, to a greater or less extent, from the orifice of the
seventh or apical segment (PI. LV. figs. 2 & 3). In many cases the appendages protrude
far enough to permit of a good view of their form; but frequently it is necessary to
remove part of the apparent apical (7th), and occasionally of the penultimate (6th) or
even of the antepenultimate (5th) segment, before they can be properly seen.

On making the necessary dissection the appendages will be found to be three in

* A notable exception is afforded by Mr. S. H. Scudder’s various Memoirs on the Hesperide of North America
(* Proceedings’ and ‘ Memoirs’ of the Boston Soc. of Nat. Hist. &c.).

SECOND SERIES.—ZOOLOGY, VOL. I. 30

358 DR. F. B. WHITE ON THE MALE GENITAL

number—one superior, and two lateral. The lateral are the equivalents of the appendices
inferiores in the Trichoptera, and in this paper will be termed the “' harpagones”
(harpdgo -onis, a grappling-hook). The upper appendage may be the equivalent of the
pair known as the appendices superiores in Trichoptera; but though sometimes deeply
bifurcate, it yet consists of only one piece, which I will call the “ tegumen”’ (¢égimen,
a cover).

On dissecting the parts more deeply, it will be seen that the tegumen, though most
largely developed in the dorsal are (or tergite) of the segment, is continued as a chitinous
ring more or less round the ventral arc (or sternite) (Pl. LV. figs. 4a, 10 ¢, 11 ¢, 12¢, and
18c). The harpagones arise either from within the ventral portion of the tegumen, or
are superimposed on that part (PI. LV. figs. 4d,10d, 11d, 12d, 13d).

I will not at present say any thing further about the deeper anatomy, or concerning
the development of these structures, which probably represent the eighth abdominal
segment.

More or less immediately below the tegumen is situated the anal opening of the
alimentary canal (Pl. LV. figs. 10 a, 11 a, and 12a); and below that, again, is the intro-
mittent organ with its sheaths (Pl. LV. figs. 4c, 106, 116, and 126). In the latter good
characters will probably be found to exist; but as very frequently a good view thereof
cannot be obtained in a dry specimen, and as the soft parts are more or less altered by
desiccation, further allusion thereto will not be made in this paper. Part of the structure
is incidentally figured in Pl. LVI. fig. 26a and Pl. LVI. fig. 18 p.

In the female Lepidopteron the armature is much less developed, and the parts suffer
so much in desiccation that an examination of fresh specimens is absolutely necessary.
In some cases, however, the structure of the apical and penultimate abdominal segments
present interesting structural characters.

Before proceeding to give some details of each part of the armature, I may state that
I have examined many species in addition to those mentioned and figured.

The drawings have been made by the camera lucida, except the diagrams Pl. LY.
figs. 10, 11, 12, & 18,

They have been made from dry specimens, which are better suited for the purpose than
fresh ones. In the latter the necessary operation of removing the hairs and scales is
apt to result in causing the parts to assume unnatural positions, while the parts are
sufficiently hard not to be altered in form by desiccation.

_In giving side views I have only figured the parts next the eye, though frequently the
corresponding parts on the opposite side are more or less visible; nor have I figured
the sheaths or other parts of the intromittent organ that were visible in some cases.

The drawings (and descriptions) only attempt to give an idea of what is visible without
an alteration of the position of the object under the microscope. The same part viewed
in a very slightly different position will often present quite a different form; so that to
arrive at a correct idea of the structure the part itself must be examined. As much care
as possible, however, has been taken to place in identical position the parts figured in the
different species, in order that a just comparison may be made.

ARMATURE IN THE EUROPEAN RHOPALOCERA. 359

1. Structure of the Apical Segment.

In discussing this and other points of the genital armature, it will be convenient to
take one species as the type or standard with which to compare all others; and in
selecting Hpinephele Hyperantus, L., as the type or standard species, I wish to remark
that I do not claim for it any special position amongst the butterflies.

The typical structure, then, is that in which (looking at it from the side) the hind
margin of the segment is perpendicular, or nearly so, for about half the distance (less or
more) from the upper margin to the lower margin, and then slopes obliquely forward
(te. towards the head of the insect)-till, at the lower margin, it nearly meets the hind
margin of the penultimate segment (Pl. LY. figs. 1 & 2). This form I shall ferm the
typical or angular. The oblique portion may be either straight, or slightly concave, or
convex. The upper margin, or dorsal surface, is slightly sloped backwards, so that the
height of the orifice is rather less than that of the penultimate segment. The lateral
surfaces are gently rounded towards the longitudinal furrow (pleurite) which marks on
each segment the division between the dorsal and ventral ares. or this reason the orifice
viewed from behind presents a long oval outline, slightly constricted at about one third
the distance from the lower margin. Viewed from above, the hind margin of the segment
presents a flatly rounded outline (PI. LV. fig. 3).

2. Modifications of the Apical Segment in the various Families.

I must first premise that for this and other parts of the structure the appearance is
described as seen in profile from the side, unless when stated otherwise.

Paprtionip#. Structure various. In Papilio Machaon (P\. LVI. fig. 40 4) only the
upper half is visible, and is triangular in form, assuming the character of the tegumen,
for which I mistook it. In P. Alexanor there is a similar modification. In P. Poda-
lirius (Pl. LVI. fig. 394) the hind margin is perpendicular. In Thais Polyxena
(Pl. LVII. figs. 1,4, B) and 7. Medisicaste (Pl. LVII. figs. 2, a, B) the hind margin is
concave, and the segment assumes to a certain degree the character of the tegumen. In
Parnassius Apollo (Pl. LVI. figs. 3, A, B, c) and P. Delius there is a considerable departure
from the type, the segment being long oval in outline, excavated for half the distance above,
and nearly closed below. In P. Mnemosyne the departure from the type is not so great.

Prerip#. Hind margin more frequently oblique throughout than angular (Pl. LVI.
figs. 27 a, 28 A, 29, 30.4, 31 A,B, 33.4, 37 A,B, and 38,8). In Colias there is an oblong
prolongation in the direction of the dorsal mesial line (Pl. LVI. fig. 36 a from side, 8B
from above). In Leucophasia the structure is very aberrant. From the side the
outline is oblong, rounded at the apex; from above oblong, rounded, and deeply emar-
ginate at apex; from below it appears widely split to the base, with a triangular piece
partly filling the aperture (Pl. LVI. figs. 34.4 side, B above, c below).

Lyca/nip#. Angular (Pl. LVII. figs. 4 a, 12 a, 13 a), or perpendicular (Pl. LVILI.
figs. 17 4, 18 a, 194), more rarely concave (Pl. LVII. fig. 144).

Eryctntip#. Angular, approaching oblique (Pl. LVI. fig. 26 a).

LisyTtHEID#. In the single European species, Libythea celtis, the structure is aberrant.

3c 2

360 DR. F. B. WHITE ON THE MALE GENITAL

The dorsal surface of the segment is prolonged into two stout terete horns (Pl. LVI.
figs. 25 a side, B above), which, seen from the side, are slightly curved downwards ;
there is a tooth on the margin of the segment below the horn; and the remainder of the
hind margin is concave.

ApatuRiIp#. Angular, approaching concave (Pl. LVI. figs. 24, 34, 4 a).

NymMpHaLip#. Frequently concave and the orifice contracted (Pl. LV. figs. 264,
29 4), or concave and the orifice not markedly contracted (Pl. LV. figs. 14 a, 17 a), or
angula (Pl. LV. figs. 22a, 24a, 38). In some species of Argynnis the structure is
peculiar. In 4. Miobe (Pl. LV. figs. 184 side, Babove) the apex is deeply emarginate in
the dorsal mesial line. In A. Adippe (Pl. LV. figs. 19 a side, B above) the same part is
toothed and is not emarginate. Some doubts have been expressed as to the distinctness
of these two species. Thus, in the ‘Entomologist’s Monthly Magazine,’ vol. viii. p. 166,
Mr. A. G. Butler gives it as his opinion that they are only forms of one species; and at
p- 206 he reiterates this opinion, and cites Freyer in support of it. I think, however,
that the structural difference now pointed out is sufficient to show that the views of
those who consider both to be good species are correct.

DanatpmH. I have not sufficient material to enable me to give the structure in this
family.

Satyrip#. Frequently angular or oblique. In several species aberrant structure
occurs in the form of stout spines, arising from the hind margin. In their simplest
form these are hairs, not much stouter or persistent than the other hairs which clothe
the segment, and may be seen in Satyrus Fidia (Pl. LVI. fig.11). In Satyrus Tolaus
(Pl. LVI. fig. 10) they are more spiny, and each is emarginate at the tip. In Satyrus
Hermione (P1. LVI. figs. 12 « side, B above) they are considerably stouter, are reduced
to two or three in number on each side, and arise from a special process of the segment.
In Epinephele Janira (Pl. LVI. fig. 14.4) there is on each side a single spine, which
appears to consist of a mass of smaller ones, slightly unequal in length, agglutinated
together. In several other species there is a trace of these appendages.

HesPerip#. Angular (Pl, LVII. fig. 28 a, &c.), or nearly perpendicular (Pl. LVII.
figs. 25 a, 314, &c.).

3. The Tegumen.

‘Viewed from above, the tegumen is oval-acuminate in outline, but truncate at the
base ; the basal half is ovately spherical; and the acuminate apex is somewhat terete,
and curved slightly downwards. Nearly halfway between the base and apex a slender,
curved, spine-like lobe (which will hereafter be termed the “‘side lobe’’) is given off on
each side, and curves downwards, inwards, and backwards, so that it is not very well
seen from above (Pl. LV. fig. 3, undissected, and fig. 8, dissected out).

Viewed from the side, the outline is somewhat triangular, with the hind margin concave,
and the upper margin convex ; from about the middle of the hind margin the side lobe
arises and curves backwards alongside and below the apical half of the tegumen (Pl. LV.
fig. 4b, dissected).

Viewed from below, the tegumen is seen to be concave, with the edges slightly

ARMATURE IN THE EUROPEAN RHOPALOCERA. 361

incurved, and gradually approaching backwards till they meet and join in the apical half
(Pl. LV. fig. 9, dissected).

The standard form of the tegumen is that in which, looking from above, the organ is
much longer than broad, and tapers from a moderately broad base to a moderately sharp
apex, and, looking from the side, when the apical half gently curves downwards and
the side lobes are well but not inordinately developed. The standard length is that in
which the tegumen slightly exceeds the harpagones, when viewed from the side and the
parts are in their normal position. As will be seen, there is often a very considerable
departure from this typical form. ,

4, Modifications of the Tegumen im the various Families.

PAPILIONID#. Structure very abnormal. In Papilio it is not visible without dissec-
tion, and is shortly triangular and deeply emarginate—in fact, almost bilobed (P1. LVI.
fig. 39 B). As pointed out above, the upper margin of the last segment assumes the
character of the tegumen, and the tegumen proper appears in the character of the side
lobes (Pl. LVI. fig. 404 & B). In Thais the construction is somewhat on the same plan as
in Papilio, but the last segment is less tegumen-like (LVII. figs. 1&2). In both these
genera I at first was deceived by the form assumed by the apical segment. In Parnassius
the tegumen is larger, but various in form. In P. Apollo (Pl. LVII. figs. 3 B & c) and
P. Delius the apex is short and slightly developed, and the side lobes (or what appears
to be them) are exceedingly developed, and on the same plane as the apical lobe. In
P. Mnemosyne the structure is rather different: the apex is bilobed, resembling the
side lobes in the other species; and the side lobes proper are smaller and situated
below.

Prerip#. Structure various. Side lobes obsolete or very slightly developed. Tegumen
rather short, acuminate-triangular, with entire apex (Pl. LVI. figs. 273, 308, & 373),
or somewhat bifurcate (Pl. LVI. figs. 31c, 345, & 38 B).

Lycznipm. Structure still further modified, the development of the part being rather
in a lateral than a longitudinal direction. In Thecla (Pl. LVII. figs. 4-8), Awrotis (Pl.
LVII. fig. 9), and Thestor (Pl. LVII. fig. 10), the tegzumen may be regarded as decidedly
transverse and broadly developed laterally; and in these genera it is relatively much
larger than the other parts. In Zycena (Pl. LVII. figs. 11, 12, & 13) the development
is still decidedly but more narrowly lateral. In Polyommatus (Pl. LVII. figs. 14-24),
while in some species the development is transverse, in the majority it is more longi-
tudinal, the sides, however, and not the centre, being produced. P. Alsus (Pl. LVII.
fig. 24) is an exception, as in this species the greatest development is in the direction of
the mesial dorsal line, as we find in the type.

In this family the side lobes are frequently present, but, instead of arising from each
side of a central prolongation, spring from the under and inner side of the lateral prolon-
gation (Pl. LVII. figs. 7B, 104,128 & c, 19 B, &.). In P. Alsus (Pl. LVII. fig. 24 4)
their situation is more normal.

Erycinip#. Short and broadly triangular. Side lobes developed, but nearly hidden
(Pl. LVI. figs. 26 B & c).

362 DR. F. B. WHITE ON THE MALE GENITAL

Lisytnerpa2. Somewhat typical, but short, and side lobes either not developed, or
hidden (Pl. LVI. fig. 258).

Apaturip®. In Apatura (Pl. LVI. figs. 2 & 3) somewhat typical, but shorter, and
no side lobes. In Charaxes (Pl. LVI. fig. 4B), structure very abnormal. Viewed from
above and dissected, the tegumen is transversely oblong, four times as broad as long,
and the outer angles developed into short acute tubercles. No side lobes visible.

NympHatips#. In the majority approaching the standard form, but rather short, and
base rather broad. Side lobes usually obsolete or not apparent, but in some cases well
developed. Apex of tegumen usually entire, but occasionally bifurcate (Pl. LV. figs. 16 c,
21 8B, 23.8), or abnormally formed (PI. LV. figs. 18 c, 19 c, 26c, 318).

Satyrip#. Structure typical, and side lobes well developed (Pl. LVI. figs. 5-24).
In Canonympha (Pl. LVI. figs. 16 & 17) there is a tendency in the apical prolongation
to become a little wider before the tip, and again narrowed. In Melanargia (Pl. LVI.
fig. 5 a) the apex of the side lobe is abruptly bent downwards.

Hesperip#. Structure more or less aberrant. Viewed from the side, the tegumen
is often more or less abruptly bent downwards, frequently very abruptly (Pl. LVII. figs.
28 B, 30, 338A).

Viewed from above, it is often longly triangular, the apex slender and produced
(Pl. LVIL. figs. 25 B, 268, 278), or more or less longly bifurcate (Pl. LVII. figs. 28 c,
31 B, 320,338). In Hesperia linea (Pl. LVII. figs. 268 & c), and H. lineola (Pl. LVII.
figs. 27 4 & B) there is a process underneath the apical half resembling the side lobes
coalesced. In dineola this process forms a groove in which the apical half lies, the process
being prolonged beyond the apex. ‘The side lobes are not otherwise apparent.

5. Structure of the Harpagones.

Viewed from the side, the harpago appears as a somewhat triangular convex plate
(more convex on the lower than on the upper margin), extending beyond the hind
margin of the last segment. The apex is prolonged, rather incrassated, and toothed and
curved upwards and inwards towards the tegumen (Pl. LV. fig. 2). On dissection, the
shape of the harpago is seen to be a long oval, with the free end prolonged and curved
(Pl. LV. fig. 4). Viewed from below, the shape is long oval, with a convex surface,
and the apex prolonged and curved inwards towards the other harpago (PI. LV. fic. 5).
Viewed from the inside, the harpago is seen to be deeply concave, with the lower margin
~ much incurved, and joined to the upper margin near the apex, to make the long tubular
tip (Pl. LV. fig. 6), which is toothed on the upper margin (Pl. LV. fig. 7, more highly
magnified).

This is perhaps one of the simplest forms that the harpago assumes; but, as we shall
see, it is often, in other species, provided with lobes or processes, generally arising from
the upper margin, which is also, especially towards the apex, frequently more or less
toothed. The interior surface is also sometimes furnished with spines or processes ;
but, except where these are visible when the harpago is viewed from the outer side,
frequent reference will not be made to them on the present occasion.

ARMATURE IN THE EUROPEAN RHOPALOCERA. 363

6. Modifications of the Harpagones in the various Families.

Papiiionip#. Usually well extruded, large and broad, without many or large pro-
cesses (Pl. LVI. figs. 39a, 404; Pl. LVII. figs. las, 2438). Parnassius forms an
exception, the harpagones not being visible without dissection (Pl. LVIL. figs. 3 4,D) in
Apollo and Delius, and only partly visible in Mnemosyne.

Prerm#. Usually well extruded, large and broad, and without conspicuous processes
or spines (Pl. LVI. figs. 27-33, and 36-38). Lewcophasia is (as in other respects) very
exceptional. The harpagones require dissection to render them visible to any extent,
are very small, and situated rather inferiorly than laterally. In Z. sinapis (Pl. LVI. figs.
34D, F) they seem to be coalesced ; in Z. lathyri (Pl. LVI. fig. 35) they appear to be less
closely united.

Lycamyip#. Usually requiring more or less dissection before a good view can be
obtained; often not larger than the tegumen, and frequently much smaller; shape
various ; with or without processes or lobes. In Thecla (Pl. LVILI. figs. 4-8), Aurotis
(Pl. LVII. fig. 9), and Thestor (Pl. LVITI. fig. 10) usually small, not very conspicuous,
and without processes. In Zycena (Pl. LVII. figs. 11-13) rather larger, and sometimes
furnished with a spine. In Polyommatus (Pl. LVI. figs. 14-24) as large or larger than
the tegumen, and frequently with the apical half bilobed or subbilobed. P. Alsus (Pl.
LVII. fig. 24.4) has a long narrow curved process arising from near the base of the
upper margin.

Erycinip#. Requiring dissection; short and broad, and without processes (Pl. LVI.
fig. 268).

Lisyrunip#. Well extruded; large and somewhat typical in form; without pro-
cesses (Pl. LVI. fig. 25).

Apvaturip&. Well extruded; large and broad, without processes (Pl. LVI. figs. 2—4).

Nympnatip#. Variously extruded; large, and often furnished with large processes.
In some genera there is an approach to the typical form, and no processes (PI. LV. figs.
36-38); in others the harpago is broader, and also without processes (Pl. LV. figs. 32
& 33); in others there is a large process arising from the upper margin at the apex,
near the middle, or near or at the base (PI. LV. figs. 14-31 & 35, 36). In many dissec-
tion is necessary for a proper view of the parts.

Satyrip#. Usually well extruded; large but not broad, and often with prolonged
apex; without- processes like those so frequent in the preceding family, but often with
a large tooth or tooth-like lobe about the middle of the upper margin (Pl. LVI. figs. 6,
8, 9, 11, 12). The apex and upper margin (especially towards the apex) is often toothed
(Pl. LVI. figs. 5, 20, 22-24).

Hespermp#. Moderately well extruded; rather large, without large processes. Apex
often somewhat bilobed (Pl. LVII. figs. 25-29), upper margin with a large tooth
(Pl. LVII. fig. 30).

The following Table shows the characteristic variations of the structure of the different
parts in the various families :—

DR. F. B. WHITE ON THE MALE GENITAL

364

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ARMATURE IN THE EUROPEAN RHOPALOCERA. 365

7. Value of all the Armature in showing Affinities and Differences between Families,
Genera, and Species.

The arrangement of the families given in this paper is that adopted by Dr. Staudinger
in his ‘Catalog der Lepidopteren des Europzischen Faunengebiets,’ not as necessarily the
best classification, but as that of the most recent list of the European Lepidoptera. My
observations have been confined to the species inhabiting Europe proper.

PaPinionipz. Affinity with Lyczenidz and Hesperide, and with Pieride. Generic
differences great ; specific differences sufficiently well marked *.

Prerip#. Affinity with Papilionide and Erycinide, and with Nymphalide (Nym-
phalides). The genera rather closely allied, especially Aporia and Pieris, and Antho-
charis and Zegris, which latter two have scarcely generic differences. Colias is well
separated by structure of apical segment. Lewcophasia is very aberrant in all respects.
The specific differences are strongest in the structure of the tegumen in Pieris, Antho-
charis, and Gonepteryx, and of the harpagones in Leucophasia and Colias.

LycznibD#. Affinities with Papilionide and Hesperide, and perhaps Charawes in
Apaturide.

Thecla, Aurotis, and Thestor form one group, and have scarcely generic differences.
Lycena is well differentiated from them, and approaches Polyommatus, which connects
the family with Hesperide. In Polyommatus there are at least four groups of species :
—Ist, Beticus and its allies; 2nd, Arion and Argiolus (perhaps forming each the type
of a group); 8rd, Corydon, Adonis, Icarus, Astrarche, &c., which are rather closely
allied; and, 4th, minimus (Alsus), which diverges considerably from the type of the
family. (I have not yet succeeded in detecting any difference between Astrarche and
Artaxerxes, which, indeed, are usually united.)

Erycinipa. Affinity with Pieridee and Nymphalide (Nymphalides). ,

Lisytaeip#. Affinity with Nymphalide (Nymphalides), Apaturide, and Satyride.

* In his “Systematic Revision of some of the American Butterflies” (Rep. Peabody Ac. Sc. for 1871, pp. 24-82),
Mr. 8. H. Scudder has revived many old (Hiibnerian, &c.), and founded several new genera. Without expressing
any opinion as to the validity of these genera (which seem to have been somewhat unnecessarily multiplied), or of
Mr. Scudder’s changes in nomenclature, it may not be amiss to point out how the structure of the genital armature
(the characters derived from which are only once or twice used in the work in question) supports, in some instances,
Mr. Scudder’s views.

Papilio Podalirius, L., and P. Machaon, L., are given as the types of the genera Iphiclides, Hb., and Amaryssus,
Dalm., respectively—a generic separation which the structure, especially of the apical segment, rather confirms.

Of the genus Vanessa, polychloros, Antiopa, and urtice are referred to three different genera. The structure of
the armature supports the separation of wrticew (genus Aglais, Dalm.), but not of the other two.

The genus Argynnis is split up into several, two of which include the species we have had océasion to treat of.
Aglaia and other of the large species belong to the restricted genus Argynnis; the smaller species are contained in
the genus Brenthis, Hb. The character on which those and some other genera are founded by Mr. Scudder (it need
hardly be remarked that the genera, as Hiibnerian, are of no yalue) haye been shown by Mr. T. L. Meade (Canadian
Entomologist, viii. 232, Dec. 1876) to be for the most part so inconsistent that their value is very questionable.
Attention may therefore be called to the difference of structure of the tegumen in at least some of the European
species of Argynms and Brenthis, as giving, at any rate, subgeneric characters. In the Hesperide the generic separa-
tion of Thaumas, lineola, &c., from sylvanus &c., seems to be warranted.

SECOND SERIES.—ZOOLOGY, VOL. I. 3D

366 DR. F. B. WHITE ON, THE MALE GENITAL

Probably an examination of more than the single European species belonging to this
and the preceding family would reveal greater affinities.

Apaturip&. Affinity with Nymphalide and Libytheide. The two genera are very
different in the structure of the tegumen; and the species of Apatura are sufficiently
well separated by the form of the harpagones.

NyMPHALIDm. Related to the Apaturide and Satyridz by the Nymphalides, which,
again, are connected with the Vanessides by Araschnia and Pyrameis.

LTimenitis, Neptis, and Nymphalis have scarcely generic differences.

Araschnia and Pyrameis are generally different from Vanessa; but Grapta is scarcely
so. Vanessa is nearer to Argynnis than to Melitea.

The specific differences are often great. In the larger genera the species arrange
themselves in groups. Thus in Vanessa, wrtice and Jo form one group, and polychloros,
Aanthomelas, L-album, and Antiopa another. In Argynnis the large and small species:
form each a group, containing several subgroups.

Satyrip#. Affinities with Nymphalide (Nymphalides) and Libytheide. Generic
differences not strong, except in Melanargia and perhaps Gineis. Pararge, Epimephele,
and Satyrus scarcely separated. Species sufficiently well differentiated.

Hzsperrp#. Close affinity to Lyceenide, less close to Papilionid, and more remote
to Melitea, in Nymphalide. Genera and species fairly well separated *.

I regret very much that, from want of material, I have been unable to examine,
amongst the Rhopalocera, any but European species. It is much to be desired that
some one who has at his command a large collection of the butterflies of all regions
should investigate more extensively than I have been able to do, the structure of the
genital armature.

DESCRIPTION OF THE PLATES.

Note.—Where the outline of the hind margin of the last segment is ragged, part has been cut away
(e.g. Pl. LV. f. 168).

Puate LY.

Fig. 1. Apex of hind body of Epinephele Hyperantus, before denudation.

. The same denuded : side view.

. The same: viewed from above.

. A side view of the same, with the integuments of the apical segment removed. a, the tegumen ;
b, side lobes; c, intromittent organ; d, harpago.

5. The same: viewed from below ; the harpagones only.

6. View of the inside of the harpago.

7. Apex of harpago, more highly magnified.

8. Tegumen from above, with the side lobes.

9. Tegumen from below, with the side lobes.

EB Oo 0

* Cf. Scudder and Burgess, ‘On Asymmetry in the Appendages of hexapod Insects, especially as illustrated by

the Lepidopterous genus Wisoniades’ (Proc. Boston Soc. Nat. Hist. xii. pp. 282-306) and other works of Mr. Scudder
on the Hesperide.

Fig. 10.

Fig.

ARMATURE IN THE EUROPEAN RHOPALOCERA: . 367

Diagrammatic plan of parts, from behind, in Epinephele Hyperantus. a, anal opening; 6, intro-
mittent organ; c, tegumen; d, harpagones. The same lettering applies to figs. 11, 12.

. The same in Argynnis Aglaia.
. The same in Pieris brassice.
. Side view (the apical half of harpago is cut away).

NyYMPHALIDz.

. Argynnis Lathonia, L. a, from the side; 3, from above.
. Argynnis Paphia, li. a, from the side; 3, from above.
. Argynnis Euphrosyne, L. a, from the side; 3B, the same dissected ; c, tegumen, from above,

dissected.

. Argynnis Aglaia, Lu. 4, from the side; 8, from above.

. Argynnis Niobe, L. a, from the side ; B, from above ; c, from the side; D, from above, dissected.
. Argynnis Adippe, Lu. a, from the side ; B, from above; c, from the side, dissected.

. Argynnis Laodice, Pall. a, from the side; 8B, tegumen, from above: both dissected.

. Argynnis Amathusia, Hsp. a, from the side; B, tegumen, from above.

. Melitea Phebe, Knoch. a, from the side; 8, from above.

. Mehitea Athalia, Rott. .«, from the side; s, from above: both dissected.

. Melitea Britomartis, Assm. A, from the side; 8, the same, dissected ; c, from above, dissected.
. Melitea Didyma, O. 4, from the side; B, from above.

. Vanessa urtice, L. a, from the side; 8, the same, dissected ; c, from above, dissected.

. Vanessa polychloros, u. From the side, dissected.

. Vanessa L-album. From the the side, dissected.

. Vanessa xanthomelas, Esp. A, from the side; 8, the same, dissected.

. Vanessa Antiopa. a, from the side; 8, from above: both dissected.

. Vanessa Jo. a, from the side; 8, from above: both dissected.

. Pyrameis Atalanta, L. a, from the side, dissected; 8, appendage on inner side of harpago;

Cc, apex of tegumen.

. Pyrameis card, Lu. a, from the side, dissected ; 8, appendage on inner side of harpago;

Cc, apex of tegumen.

. Grapta Egea, Cr. A, from the side; 8, from above.

. Grapta C-album, L. a, from the side; 3, from above.

. Araschnia prorsa, L. (=Levana, L., var. gen. 1.). a, from the side; 3, from above.
. Neptis Lucilla, ¥. From the side.

. Limenitis Sibylla, L. From the side.

Prats LVI.

NympPxHaip# (continued).

. Nymphalis populi, L. From the side.

APATURIDE.

. Apatura Iris, l. a, from the side; 3s, from above.
. Apatura Ilia, Schiff. a, from the side ; 3, from above.
. Charaxes Jasius, L. a, from the side; 8, tegumen, from above, dissected.

SATYRIDZ.

. Melanargia Larissa, H., var. Herta. a, from the side; 8, tegumen, from above.
. Pararge 4igeria, L. 4, from the side; 8, tegumen, from above.

3D 2

Fig.

DR. F. B. WHITE ON THE MALE GENITAL

7. Pararge Rowellana. a, from the side; 8, tegumen, from above.
8. Satyrus Arethusa, Esp. 4, from the side; B, tegumen, from above.
9. Satyrus Statihnus, Hufn. From the side.
10. Satyrus Iolaus, Bon. From the side.
11. Satyrus Fidia, L. From the side.
12. Satyrus Hermione, L. a, from the side; 8. apex of last abdominal segment, from above.
13. Epinephele Lycaon, Rott. From the side.
. Epinephele Janira, . 4, from the side; 8, tegumen, from above.
15. Canonympha Iphis, Schiff. From the side.
16. Canonympha Typhon, Rott. Tegumen, from above.
17. Cenonympha Pamphilus, L. Tegumen, from above.
18. Cenonympha Hero, L. Tegumen, from above.
19. Gineis Aello, Hb. From the side.
20. Erebia Stygne, O. (= Pirene, Hb.). a, from the side; 8, tegumen, from above.
21. Hrebia Goante, Esp. From the side:
22. Erebia Tyndarus, Esp. From the side.
23. Erebia Melampus, Fuessl. 4, from the side; B, inner side of harpago.
24. Erebia Epiphron, Ku. 4, from the side, parts protruded more than naturally ; 8, mner side of
harpago.

I
o

LisyTHEID2.
25. Libythea Celtis, Esp. a, from the side; 8B, from above.

ERYCINID A.

26. Nemeobius Lucina, L. a, from the side; s, the same, dissected; c, tegumen, from above,
dissected.
Pirrip#.

27. Pieris napi, L. A, from the side; 8, tegumen, from above.

28. Pieris Daplidice, L. a, from the side; 8, from above.

29. Pieris brassice, LL. A, from the side; 8, tegumen, from above.

30. Aporia crategi, L. a, from the side; B, tegumen, from above.

31. Anthocharis cardamines, lu. a, from the side; B, from above; c, tegumen, from above.

32. Anthocharis Damone, Feisth. Tegumen, from above.

33. Zegris Eupheme, Esp. a, from the side; 8, tegumen, from ahove.

34. Leucophasia sinapis, . A, from the side; 8, from above; c, from below; pv, from the side;
dissected ; n, tegumen, from above, dissected; F, harpagones, from below, dissected.

35. Leucophasia Duponcheli, Stgr. (= Lathyri, Dup.). Harpagones, from below.

36. Colias Edusa, F. a, from the side; 8, from above; c, tegumen, from above, dissected.

37. Gonepteryx rhamni, L. a, from the side; B, from above.

38. Gonepteryx Cleopatra, L. a, from the side; B, from above.

PApiLiIonIpz.
39. Papilio Podalirius, L. a, from the side; 8, tegumen, from above, dissected.
40. Papilio Machaon, L. a, from the side; 8, from above, dissected.

Pruate LVI.

1. Thais Polyxena, Schiff. a, from the side; 8, from above; c, tegumen.
2. Thais Rumina, L., var. Medisicaste, I. a, from the side; 8B, from above.

“EO o1

ARMATURE IN THE EUROPEAN RHOPALOCERA. 369

. Parnassius Apollo, L. a, from the side; s, from above; c, from below; p, from the side,

dissected.
Lycanip2.

. Thecla ilicis, Esp. a, from the side; 8, the same, dissected; c, tegumen, from above, dissected ;

D, harpagones, from below, dissected.

. Thecla quercus, Li. a, from the side; 8, tegumen, from above: both dissected.
. Thecla pruni, Lu. a, from the side; 8, tegumen, from above : both dissected.
. Thecia betule, L. a, from the side; c, tegumen from above; B, harpagones, from below: all

dissected.

. Thecla rubi, i. a, from the side; B. tegumen, from above; c, harpagones, from below: all

dissected.

. Aurotis roboris, Esp. 4, from the side; B, tegumen, from above: both dissected.
. Thestor Ballus, F. a, from the side; 8, tegumen, from above: both dissected.
. Lycena Alciphron, Rott., var. Gordius, Sulz. a, from the side; c, tegumen, from above; ., har-

pagones, from below : all dissected.

. Lycena Eurydice, Rott. a, from the side; B, from the side, dissected; c, tegumen, from above,

dissected.

. Lycena virgauree, LL. A, from the side; B, from the side, dissected; c, tegumen, from above,

dissected ; p, sheath of intromittent organ, from above.

. Polyommatus Beticus, L. 4, from the side; 8, from above, dissected.

. Polyommatus Telicanus, Lang. a, from the side; B, the same, dissected.

. Polyommatus Arion, L. a, from the side; 8, from above : both dissected.

. Polyommatus Corydon, Poda. 4, from the side; 8, tegumen, from above.

. Polyommatus Adonis,Hb. 4, from the side; 8, tegumen, from above.

. Polyommatus Icarus, Rott. a, from the side; 8, tegumen, from above.

. Polyommatus Optilete, Knoch. a, from the side; 8, tegumen, from above: both dissected.

. Polyommatus Astrarche, Bgstr., var. Artaxerxes, L. a, from the side; 8, tegumen, from above:

both dissected.

. Polyommatus Afgon, Schn. a, from the side; B, tegumen, from above: both dissected.
. Polommatus Argiolus, L. a, from the side ; 8, tegumen: from above, both dissected.
. Polyommatus minimus, Fuessl. (= Alsus, F.). a, from the side; B, tegumen, from above: both

dissected.
HEsPERIpz.

. Nisoniades Tages, Lu. a, from the side; 8, tegumen, from above, dissected.
. Hesperia Thaumas, Hufn. (=Linea, F.). a, from the side; c, the same, cissected; s, tegumen,

from above, dissected.

. Hesperia lineola, O. A, from the side ; 8, tegumen, from above, dissected.
. Hesperia Sylvanus, Esp. a, from the side; 8, the same, dissected; c, tezumen, from above,

dissected.

. Syrichtus serratule, Rbr. 4, from the side; 8, tegumen, from above, both dissected.

. Spilothyrus alcee, Esp. (= Malvarum, Hffg.). a, from the side, dissected.

. Syrichtus malve, L. (= Alveolus, Hb.). a, from the side; B, tegumen, from above.

. Carterocephalus Palemon. Pall. (=Paniscus, ¥.). , from the side ; B, the same, dissected ; c, tegu-

men, from above, dissected.

. Cyclopides Morpheus, Pall. (=Steropes, Schiff.). a, from the side; B, tegumen, from above : both

dissected.

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F B. White, del. Trans. LINN. Soc. SER. 2 Zoou.Vou.l. Pu.55.

GENITAL ARMATURE EUROPEAN RHOPALOCERA

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GENITAL ARMATURE EUROPEAN RHOPALOCERA

(ff 74

XVIII. Morphology of the Mammalian Ossicula audits. By AuBan H. G. Doran,
F.R.C.S. (Communicated by Professor Fiowsr, F.B.S., F.L.S., §c.)

(Plates LVITI.—LXIV.)
Read December 21st, 1876.

AMONG John Hunter’s original collection are some preparations in spirit, repre-
senting the tympanic region of mammals, with the small ear-bones i situ, but only a
very few imperfect dry specimens of the internal ear-bones themselves, though those of
man are better represented. Little was added to this department at the Royal College
of Surgeons’ Museum until 1866, when Mr. Toynbee’s valuable pathological auditory col-
lection showed the want of a corresponding series in the physiological department. For-
tunately, about the above date, the nucleus of such a collection offered in that of Dr. Max
Hibrich, of Munich, which the College authorities purchased. The Conservator, Prof.
Flower, wisely availed himself of opportunities to increase this; and at the present date
a very complete series of most of the genera of Mammalia is the result.

In drawing out a catalogue of these, I was led to study minutely and compare the
various groups one with the other. I soon found that, excepting the labours of Prof.
Hyrtl, of Vienna, on the osteology of the entire auditory apparatus in the Mammalia
(‘ Vergleich. Anat., Untersuch. iiber das innere Gehororgan des Menschen und der
Saugethiere,’ Prague, 1845), little or nothing in a connected form had been done to
elucidate the characters of the small ear-bones of Mammalia. Occasional desultory
remarks are to be found scattered in papers on human and comparative anatomy; but
these in most cases are entirely lost sight of. This suggested to me, in preparing and
arranging the Museum series, to take notes thereon and the critical comparison of
the objects themselves and collation of the literature on the subject, have resulted in the
following descriptive and comparative account of the ossicula auditis in the class
Mammalia.

In such small objects as the internal ear-bones it is at first difficult to apprehend the
value that may be attached to them as regards physiological influence on audition, and
still less as to their morphological importance; but if, for instance, those of man be
compared with those of marsupials, the wide difference is at once apparent. Again, in
such groups as the Carnivora or Cetacea, the family characters derived from these
diminutive ossicula are in the main very distinctive.

As the small ear-bones of the human being, for various apparent reasons, have neces-
sarily had most attention called to them, and moreover as man, in other respects, is con-
sidered by some a standard of comparison, I have consequently taken the human ossicula
- first into consideration. Moreover I have described their peculiarities in such a way
that the homologies of the parts of each ossicle in the lower mammals may be more
clearly understood.

372 MR. A. H. G. DORAN ON THE MORPHOLOGY

The drawings forming the accompanying Plates are from nature, and in all cases
enlargements. By their representation side by side in series a much better idea of the
family and generic distinctions are obtained than by mere verbal descriptions. Indeed
the latter have grown inordinately in my hands; and but for the circumstance that the
full data whereon my conclusions are founded are absolutely necessary, pending the fact
of this series of ossicula being unique, or certainly the most extensive now extant,
I should gladly have dispensed with this portion of the labour, and curtailed my text
accordingly.

In the Journal of this Society (Zool. No. 68, vol. xiii. p. 185) and in the Proc. Roy.
Soc. (No. 172, vol. xxv. p. 101) I have already indicated briefly a few of my results, and
in the latter referred to those gentlemen to whom I have been indebted for objects,
advice, and assistance.

Tue AUDITORY OSSICLES IN MAN.

The Malleus.—In the human malleus (Pl. LVIII. figs. 1 & 2, taken from a fully
developed foetus) the portion of the head above the articular surface is prominent,
smooth, and convex, projects more on the outer than the inner side, and is broadest
in its lateral measurement. Posteriorly it bears the articular surface for the incus,
which has very elevated borders and is about three times as broad as it is deep vertically.
This surface lies very obliquely compared with the mallei of most mammals, so that
its external extremity is much higher than its internal. It is generally described
as “ saddle-shaped,” though, from a functional point of view, Helmholtz + has objected
to this expression. In most works on human anatomy it is regarded as if it were one
single facet; but on comparing it with the same surface on the malleus of a Cat
(Pl. LVIII. figs. 30 & 31), it will be seen that it should be considered as made up of two
facets, much less distinct, it is true, than in Felis and most other mammals, but far
better-marked than in certain monkeys. A very faint groove, more marked in some
human mallei than in others, and not reaching either extremity, divides them, running
in the very oblique long axis of the whole surface. The more internal and upper facet
(Pl. LVIII. fig. 30, a@ * a), or portion of the articular surface above the groove, repre-
sents that facet which is almost completely superior in position in most lower mammals
(fig. 31, a* a), the more external (d* 0) below this groove corresponding to the lower
facet in many other mammals. Both rise into a high vertical convexity about the
middle of the whole surface, which is there much contracted ; their planes slope down-
wards towards the groove, so that a concavity is formed, plainly visible when the arti-
cular surface is viewed sideways. Now it may be remarked that of all the portions of
the three ossicula there is none that requires so minute and careful an investigation as
the articular surface of the malleus; and although the results of such labour are often
puzzling, and not invariably satisfactory, still, as a whole, many distinctive points of
value will be found in this region among the different orders of mammals. It will be
shown, when speaking of the ruminants, that the characters of this surface undergo

+ See his remarks on the malleo-incudal articulation in his work, ‘The Mechanism of the Ossicles and the
Membrana Tympani,’ translated by J. Hinton, New Sydenham Society, 1874.

OF THE MAMMALIAN OSSICULA AUDITUS. 373

conspicuous changes during growth in some genera, whilst in others no such alterations
take place. Hence the advisability of procuring young as well as adult ossicula, before
coming to conclusions with regard to peculiarities in any group,.is manifest.

Between the head and the manubrium is the short, constricted “ neck.” This is
rather flattened laterally, so as to be narrower in its vertical than in its horizontal measure-
ment. On its external aspect is a sharp sigmoid ridge *, convex forwards at first, where
it is near the anterior border of the articular surface, then concave forwards, where its
anterior end loses itself on the root of the manubrium (PI. LVIII. fig. 30). I call
particular attention to this ridge, as it is almost constant in those animals where the
malleus has a distinct neck, and is very plain in the mallei of the fissiped Carnivora and
most Ungulates, appearing as the sharply curved compact neck itself, the remainder
being a thin lamella of bone joining the processus gracilis. In the Common Badger
(Meles taxus) it will be seen to curve so sharply as to form a distinct projection.

Observe that this sigmoid portion—evidently the essential part of the neck, if we give
that name to the bony isthmus which seems made to unite the head to the manubrium—
runs away from the processus gracilis, but is lost on the base of the manubrium. This is
easily accounted for when we remember that the neck is originally developed from the
ineurved dorsal end of the mandibular arch, the apex of which forms the manubrium itself,
Professor Parker has recently shown 7.

On the anterior aspect of the neck, close to the root of the manubrium, is the root or
place of origin of the processus gracilis. Now, after examining numerous mallei from
human foetal and adult skulls, I have hardly ever found a trace of the wide lamina of
thin bone filling up the angle that process forms with the neck as in the fissiped
Carnivora. But as I believe the sigmoid posterior part of the neck in Homo to
represent the similar curved isthmus which constitutes the whole neck (independently
of the lamina) in these animals, I consider it correct to say that the stout compact
portion of the ‘neck’ of human anatomists in front of that sigmoid ridge is really
the same ossification as the upper part of the lamina, when such exists—the lower
being absent in any form, whether compact or papery. In good specimens from a
fetus at full term, it will be seen that a little fringe of friable bone extends as far
upwards as the front of the head, which latter, in these cases, sends forward a slight
projection, identical with a far more developed process seen in some lower animals,
especially in many of the Rodentia with laminated mallei, and notably m the Musquash
(Fiber zibethicus); in these extreme cases that projection may readily be taken for
the processus gracilis, if the anatomist overlook the fact that its root is not in the right
position for the origin of the latter.

The form of the human processus gracilis is well known. It cannot be seen entire
unless taken from a feetal or very young human skull, not only on account of its fra-
gility, but also because it actually atrophies to a mere stump before adult life ¢; so that
after removing a malleus from the temporal bone of a full-grown subject, the anatomist,

* Henle describes this as “a sharp border” (“ Kante’’), but he does not mention its sigmoid form.
+ “ On the Structure and Development of the Skull in the Pig,” Phil. Trans. 1874.
+ See Helmholtz, op. cit.

SECOND SERIES.—ZOOLOGY, VOL. I. 3E

374 MR. A. H. G. DORAN ON THE MORPHOLOGY

unaware of this fact, has frequently searched in vain along the Glaserian fissure for this
process supposed to have been broken off. It forms with the neck an angle of 125°, and
runs almost horizontally forwards to the fissure. It is generally bowed regularly, and
is concave forwards; sometimes it is somewhat wavy; but in all cases the degree of cur-
vature is slight. When well preserved, the processus gracilis at birth is longer than the
manubrium.

Below and internal to the processus gracilis the insertion of the tensor tympani muscle
is sometimes marked by a faint elevation, close to or upon the manubrium representing
the “ processus muscularis” of Hyrtl, which attains a great development in some lower
animals. In the human malleus, however, an elevation is occasionally seen as marked
as its homologue in the Bears, presently to be described.

That important process the manubrium, or handle, now claims our attention. In
Homo it is below the average length, far shorter than in ruminants or terrestrial Car-
nivora, though longer than in some Seals or Whales. It forms with the neck an angle of
about 140°; Hyrtl gives 150°; but he must have taken the axis, not of the upper and greater
part of the manubrium, but of the recurved portion at the tip, from strongly curved
examples. It is of a stout make, and broader at the base than in the Simiide; it is
much flattened laterally, the sides looking respectively antero-superiorly or postero-
inferiorly ; they are slightly convex, notwithstanding their compression. ‘The extremity
is always slightly recurved, though more in some specimens than in others, and per-
ceptibly spatulate and smooth towards the tympanic membrane. The edges of the
manubrium are both thick and blunt; and the outer edge, though of a certain breadth,
and giving attachment to fibres along its entire length, is rarely separated from each side
by sharply defined borders, so as to constitute a true and distinct outer surface, as seen
in many lower animals, and particularly in the Canide *. This edge ends below in the
spatulate dilatation, above in a very distinct sharp projection, well curved outwards.
This is the “ processus brevis:” its good development is a prominent feature in the
human malleus; and it will be seen to be better marked in the higher than in the lower
Quadrumana, though it occasionally reappears in the latter. In most mammals it is
merely an angle at the outer aspect of the base of the manubrium ; in many it does not
exist even in that form, especially where the neck of the malleus is very short.

Such is the human malleus. It may be seen as a good central type of the bone in
mammals, a type which very frequently reappears as we go downwards in the scale—a
malleus characterized by a well-developed,head, a moderately long, constricted neck, and
a broad-based, stout manubrium. It holds an intermediate position between the almost
or quite neckless type seen in the Cebide, in Tupaia, and in the true Squirrels, and the
broadly laminated form so well seen in the terrestrial Carnivora and the artiodactylate
Ungulata. It will be seen to be repeated, with modifications, in the higher monkeys, re-
appearing among Carnivora in the Ichneumons, and certainly approached (with some
important distinctions) in the Seals. It is also imitated in some of the Insectivora,

* Tn a recent contribution to the ‘ Archiy fiir Ohrenheilkunde,’ Trautmann describes and figures this condition as
if constant in Homo. It is only in a very few specimens of the human malleus, however, that I haye distinguished
two distinct borders running completely from the processus brevis to the tip.

OF THE MAMMALIAN OSSICULA AUDITUS. 375

Rodentia, &c. As for its comparison with the malleus of the Anthropomorpha, such a
consideration may be more conveniently deferred until the latter have been described
individually. The same remark applies to the other ossicula.

The Incus.—In the embryo this little bone holds a very prominent position, as the
proximal end of the hyoidean arch. In comparative anatomy it has hardly as yet
received the attention it deserves, on account of its having been supposed to vary less
in different mammals than do the other ossicles. My own observations lead me to con-
sider that in reality the chief objection to the incus as a distinctive feature in the skeleton
is the frequency with which it varies in form in different individuals of the same species.
Still it possesses important distinctive features in many orders of mammals. The incus
of Homo is of even more central type than the malleus.

The “‘ body” is well developed both in length and in height, and is considerably com-
pressed laterally. Its posterior part, or that which lies over the processus brevis, is more
developed than the anterior. The articular surface looks upwards and forwards, and is
shaped to fit that of the malleus; its more external limit is on the posterior segment of
the body, which is higher and more external than the anterior, in accordance with the
high position of the external extremity of the opposing surface on the malleus, already
described. On the other hand, that boundary of the surface on the incus opposing the
inner, which is the much lower extremity of the corresponding area on the malleus, lies
upon the lower and more internally placed anterior segment of the body. Into the
deep groove between the two extremities fit the convexities of the two facets on the
malleus. The very oblique position of the incus renders any purely verbal explanation
of the relations of this surface somewhat confusing; and the difficulty is increased in
this case chiefly because, in describing the malleus, I have acted in conformity with its
condition in the lower animals, where, as I have already remarked, that which is almost
the upper end of the articular surface in Man is in them rather the outer extremity, the
boundary which lies almost quite superiorly in most mammals being almost internal in
our species ; hence the homologue of their upper facet in Homo has an interno-superior,
and that of their lower facet an externo-inferior position.

Around the inner margin of this articular surface of the incus is a distinct groove, like
that which is often so strongly developed in the Seals (especially in Cystophora and
Phoca); and there is a distinct concavity on the inner surface of the body, at the bottom
of which is a minute nutrient foramen, often, however, filled up, even in the incus of a
new-born child.

The “processus brevis”’ of a human incus is, as a rule, not much shorter than the so-
called “processus longus.” It may, with more reason, be called the “ posterior crus.”
In Man, however, it lies as much superior to the rest of the ossicle as posterior; and in
some animals (the Dugong, &c.) it is almost absolutely superior; but the term given
above is less clumsy than “ postero-superior crus.” This posterior crus is broad at the
base and blunt at the point, where, on the inner aspect, is a minute pit or concavity,
generally admitted to be a true articular surface. It is moderately divergent from the
body. The processus longus, or stapedial crus, is slender and slightly convex inwards at
first, then forms a gentle curve, suddenly turning inwards at the extremity, which bears

3E 2

376 MR. A. H. G. DORAN ON THE MORPHOLOGY

a flat elliptical disk on a narrow pedicle. This is the “ os orbiculare,” better termed the
“Sylvian apophysis’ *. Among anatomists, Sappey gives reasons for considering it a
separate bone; but as it is now recognized as being formed on the proximal end of the
hyoidean arch, and, on the other hand, is separate from the long crus in an early stage of
development, it would appear correct, on such premises, to consider it an epiphysis of
the incus.

The Stapes—Whilst the ossicles already described take, in Homo, a rather central
type, this little bone assumes in the Bimana an extreme and highly developed form.
The beautiful bicrurate stapes of Man differs strikingly from the simple columelliform
representative of that ossicle in the Ornithorhynchus; and the study of the types inter-
mediate between those two extreme forms is highly instructive.

The “head” is broad and shailow; its free aspect is elliptical and a little concave, to
articulate with the os orbiculare. The crura are long and widely divergent, so as to
leave a large aperture between them and the base. The anterior crus is the more
slender and the straighter of the two; it is widely grooved towards the aperture in its
upper half; but the channel generally narrows close to the base. The posterior crus has
a curve or shoulder, often very marked, not far from the head; it is deeply channelled
towards the intercrural aperture by a groove almost as deep near the base as near the
head. ‘This sulcus joins, under the head, that which exists on the opposite crus.

Sometimes the anterior crus is more curved than usual, but seldom so sharply as the
opposite limb of the stapes. In such cases it is still the more slender, and the groove is
less marked near the base than it is on the posterior crus. I have examined several
specimens of short, stumpy stapedes, from both infants and adults, where it was found
very hard to distinguish the anterior from the posterior crus by their own characters
alone +.

The base, or foot-plate, is reniform, its upper border being a wide arch, its lower
slightly concave in the middle. Its posterior extremity is always well rounded off; and
its anterior is usually much sharper; but this distinction is often the least marked where
the crura are most alike.

The base is slightly convex towards the vestibule. I have never seen a true umbo
like that which is found on the stapes of many monkeys. On the tympanic aspect it is
formed into a shallow tray by the prolongation along its margin of the high edges of the
channel within the crura. This condition is very frequent in animals where that ossicle
is well developed and lightly made. A bony ridge}, traversing the long diameter of
the tympanic aspect of the foot-plate, is sometimes seen, but is not constant. The aper-
ture between the crura of the human stapes is larger, when measured vertically as well

* A beautiful description of this structure and the incudo-stapedial joint is to be found in Kysell’s “Beitrige zur
Anatomie des Steigbiigels,” Arch. f. Ohrenheilkunde, vol. v.

+ But the straightness of the anterior crus is generally much more apparent in the stapes of an infant than in that
of an adult. A specimen from an adult Bushwoman in the College collection is of a very marked infantile type
in the form of its crura. ‘This agrees with the characters of the rest of her skeleton, described in Flower and Murie’s
“ Account of the Dissection of a Bushwoman,” Journal of Anatomy and Physiology, vol. 1.

+ The “ crista stapedis.” See Hysell, loc. cit.

OF THE MAMMALIAN OSSICULA AUDITUS. 377

as horizontally, than in the lower animals, even including the Elephant, as Hyrtl justly
observes; but in the Golden Mole (Chrysochloris) and a few other fossorial animals
the width of the aperture at the base is proportionally greater, though the head is much
nearer the foot-plate.

The presence of a minute tubercle on the inner side of the head of the stapes does not
seem constant in our species. The occasional ossification of a part of the stapedius tendon
in Homo is well known; it is a constant feature in many animals.

For the most recent minute observations on the anatomy of the human ossicula, the
works of Magnus *, Helmholtz +, Eysell }, Rudinger §, Trautmann ||, Urbantschitsch {,
and others should be consulted.

THE OSSICULA OF THE SIMIIDA.

In a set of ossicles from a large mature fetus of a Gorilla (Pl. LVIITI. fig. 3) the incus
much resembles that of Homo, the stapes is not so large as in the human fcetus at birth,
and the crura are a little more slender and straighter. The tympanic ring from the
same foetal Gorilla has a considerable part of the floor of the external meatus already
developed, which is not the case in Homo at birth. Both rings are preserved in the
College of Surgeons’ collection.

I have examined a single malleus of the adult Troglodytes gorilla. It is above
the average size of the same bone in Man. The free upper portion of its head is
more developed than in Homo or Simia, and is somewhat flattened vertically, at the
same time protruding considerably forwards, as in the Chimpanzee. The articular sur-
face remains oblique, so that, as in our species, its outer extremity, or the homologue of
that extremity in other animals, lies very much higher than does the inner. The
outer and upper segment of the articular surface is far larger than the inner **—another
point in which this genus differs from Man and the Orang. The upper and lower facets
are saddle-shaped; the vertical convex ridge is very near the inner margin of the whole
surface; and hence the external, concave, sloping portions of both facets are very wide.
A very faint groove divides them, indeed, as in Man; it is only relatively that they can
be spoken of as two separate facets. This is the case with the remaining Quadrumana.

The neck is rather short and very narrow; the size of the head and the length of the
handle make it look less developed than it really is. The sigmoid ridge described in
Man is well marked. In the mallei from a fetal Gorilla the processus gracilis is very
slender, especially at the root.

The manubrium is very long, resembling in that respect the Orang’s, and differing
from the rather shorter manubrium of the Chimpanzee and Man. It is somewhat narrow
at the root, and well recurved at the tip, which is more thoroughly spatulate in Homo and

* « Beitrage zur Anatomie des mittleren Ohres,” Virchow’s Archiv, vol. xx. + Op. cit. = Op. eit.
§ ‘Atlas des menschlichen Gehérorgans,’ Munich, 1875.

|| “Der gelbe Fleck am Ende des Hammergriffs,” Archiv f. Ohrenheilk. vol. xi.

@ “Zur Anatomie der Gehérknéchelchen des Menschen,” bzd. vol. xi.

** This condition is not nearly so marked in specimens from new-born Gorillas.

378 MR. A. H. G. DORAN ON THE MORPHOLOGY

the other anthropoid apes; but the processus brevis is quite as well developed as in our
species.

In the Chimpanzee (Pl. LVIII. fig. 4) the malleus is much more human than in the
Gorilla or Orang-utan, and is as big as most of the numerous mallei removed by me
from adult male skulls in the dissecting-room of St. Bartholomew’s Hospital. In the
shape of its head, which projects markedly forwards, and in the nature of its articular
surface, of which the outer segment is much the widest, it approaches 7. gorilla more
than Homo or Simia; but in neck and manubrium it is very human, the only differ-
ence being that the latter, in this ape, is rather narrower at the base, and more curved
than in Man, and its well-developed processus brevis is directed upwards, and hardly
outwards. In length the handle does not exceed that of our species—another prominent
distinction from the other two apes.

The body of the incus resembles that of Homo; the processus brevis is more slender,
and ends in a sharp point, with no trace of any depression on it. The processus longus
is rather stouter and shorter than in Man; it forms, with the posterior crus, a right
angle.

The stapes of an adult Chimpanzee is smaller than the smallest specimen of that
bone from Homo in the College collection. The crura are almost equally curved; they
are shorter and more slender than in Man, but wider apart at their insertion. They
are well grooved towards the aperture, which is very wide. The base resembles that
of Homo, though less distinctly reniform, and equally rounded off at both extremities.

In the Orang-utan (Pl. LVIII. fig. 5) the head of the malleus is rather less developed,
and less regularly globular than in Man, and does not project conspicuously forwards
as in Lroglodytes. The whole bone, however, is of about the average size of its human
representative. The articular surface is shaped as in Man, but is less extensive; it is
hardly wider at its external or externo-superior extremity than internally. The neck is
as narrow as in the last genus, but rather longer; and it seems as if the sigmoid ridge
had become almost effaced in the process of elongation, being hardly perceptible, though
so marked in Troglodytes.

The manubrium is as long and slender as in the Gorilla; but the processus brevis is
very ill developed, though distinctly bent outwards. The extremity of the manubrium
is less dilated than even in Homo. .

In the incudes of Simia, in the College collection, removed from a young male Orang,
the body is much more developed *, particularly behind, than in Man, though not as
wide horizontally. Hyrtl asserts that the body is markedly smaller. The posterior
crus is as small and pointed as in Troglodytes; but the processus longus is short and
slender, and more incurved than in Homo; the crura too are more divergent.

The stapes has a very slightly developed head, and much more slender crura than in
Troglodytes. ‘The anterior is quite straight, the posterior slightly curved. The base
is distinctly reniform, and equally rounded at each extremity.

* Less of the body is taken up by the articular area than in the incus of Homo, as that surface is smaller abso-
lutely and proportionally in this ape.

OF THE MAMMALIAN OSSICULA AUDITUS. 379

In Aylobates (Pl. LVIII. fig. 6) the malleus (which is rather small even in a large
Gibbon’s skull) resembles in many respects that of the Chimpanzee. The upper part
of the head projects forwards; its articular surface is very wide externally, after the
Troglodytes type. From the fore part of the head projects a thin fringe of bone, which
does not join the root of the processus gracilis as in the Cynomorpha, and is repre-
sented, as in this genus, in some mallei from new-born children. The neck is short
and constricted ; the manubrium is as short as in Man, and much curved; it has a very
prominent, outwardly directed processus brevis, and a distinctly spatulate dilatation at
the tips; the whole angle forms, with the neck, an angle about as wide as in Homo, not
being so obtuse as in the Cercopithecide. Moreover the neck, though short and con-
stricted, is longer than in Semnopithecus, which has the most distinctly necked malleus
of all the next group. Altogether the affinities of the malleus of a Gibbon tend more
towards Homo than to the tailed monkeys.

In H. syndactyla and H. leuciscus (and probably in other Gibbons, only not con-
stantly) a distinct “processus muscularis” for the attachment of the tensor tympani
exists. In the latter it is seen as a distinct projection from the inner edge of the manu-
brium, at some distance from its root. In H. syndactyla it is reduced to a very small
tubercle on the root, almost on the site of the attachment of the tendon in Homo.

The incus in the Siamang (H. syndactyla) has a body of the same form as in Homo ;
in a specimen of that ossicle from H. leuciscus the body is higher and narrower, whilst
in an incus from another Hylobates the body is as narrow as in some of the Cebide.

The stapes of Hylobates has always curved, divergent crura, and a wide aperture *.
This is particularly marked in H. syndactyla. In this ossicle the Gibbons are allied to
Homo, Simia, and Troglodytes, in a manner not nearly so apparent in the malleus, and
still less in the incus. The very straight and not very divergent crura of the stapes in
all the tailed monkeys, even Semnopithecus, distinguish that group of the Quadrumana
not less from Hylobates than from Homo+. With regard to the ossicula of the Simiide,
in general, it may be briefly stated, in conclusion :—

I. That the ossicula of Troglodytes niger are, taken as a whole, most like those of
Homo. T. gorilla closely resembles Homo in its incus and stapes, but less in its malleus ;
Simia more resembles Homo in the head and articular surface of the malleus.

IJ. Though the incus of some species of Hylobates exhibits a tendency to low type, in
the malleus, and particularly in the stapes, this genus is quite anthropoid.

III. In their ossicula, but most markedly in the stapes, these apes are much more allies
to Homo than to the lower monkeys.

* In the Hylobates of uncertain species just referred to, the stapes bore on its foot-plate a distinct umbo, or central
‘convexity on the vestibular aspect.

7 From want of a sufficient number of good specimens when I contributed the communication already quoted to
the Royal Society, I was under the impression that the Gibbon’s stapes was of a low type; but I am now, after
more fayourable opportunities, compelled to modify that assertion.

380 ' MR. A. H. G. DORAN ON THE MORPHOLOGY

Tur AUDITORY OSSICLES OF THE CYNOMORPHA, OR TAILED OLD-woRLD MONKEYS.

The study of the ear-bones of the Old-world monkeys yields interesting results; for
they all possess certain types in common; but in comparing them with the Anthropo-
morpha we find definite grades of distinction, particularly between Semnopithecus and
Macacus.

In examining the malleus of an HEntellus (Semnopithecus entellus, Pl. LVIII. fig. 7),
the free upper portion of its head will be seen to be well developed and globular, dis-
tinctly projecting forwards. In the majority of the mallei of this species and genus a
distinct fringe of bone, of papery consistence, is found on the front border of the head,
joining the root of the processus gracilis (which is stout, stumpy, and very brittle in
the Cynomorpha). This (which is the exception in Man, and when existent the frmge
never joins the root of our processus gracilis, and is as little indicated in the anthro-
poid apes, excepting, perhaps, Hylobates) is a clear approach to the laminated type of
malleus seen in the Carnivora, &c. The articular surface is shaped much as in Man
and Simia, and is as little divided into the two facets of the typical mammalian malleus
as is the case in Homo. The neck is very short, but, being much constricted, is quite
distinct. The manubrium forms with the axis of the neck and head an angle of at
least 155°, distinctly wider than in any of the last family. This process is of moderate
length, though the shortness of the neck makes it look rather longer than it actually
is. It is almost of equal breadth laterally, till near the tip, where it turns up rather
_ sharply, and is there discoidal rather than spatulate.

Immediately above this dilatation is a distinct convex projection, giving a wavy look
to the manubrium seen in many lower monkeys. There is always a well-developed
and everted processus brevis in this genus; and on the inner edge of the handle, above
the projection just described, on the opposite or outer margin, is a distinct ‘ processus _
muscularis ” (as Hyrtl styles it, in describing the carnivorous malleus, and as it will be
henceforward designated in this paper), to which, in the recent skull, the tensor tympani
muscle will be found attached, as I have verified by dissection. Its position in this
and in neighbouring genera, well down the manubrium, reminds the observer of the
same process in Herpestes and in many Rodents.

In its very distinct though very short neck, and well-developed processus brevis, the
malleus in the genus Semnopithecns resembles the Simiidze more than any of the fol-
lowing genera except Colobus, where we find similar characters; but there the form
of the head is different. But the wide angle between the body and manubrium, the
peculiar form of that process, with a tendency to a discoidal termination, and the
presence of a distinct processus muscularis, as well as the trace of a lamina between
the head and processus gracilis, are points that ally this genus far more closely to other
Old-world monkeys.

The incus, on the other hand, in all the skulls of this genus which I have examined,
more resembles that of the anthropoid apes than in the succeeding genera, being broad
in the body, measured horizontally, and not very deep, with a considerable distance
between the origin of each crus, which diverges as much as in Man.

OF THE MAMMALIAN OSSICULA AUDITUS. 381

But the stapes of Semnopithecus assumes to perfection the type almost constant in the
lower monkeys. The head is deep rather than wide, the crura practically quite straight,
the base rather broad and almost plane, and the aperture much smaller than in the
Simiide. Thus, when we take into consideration all three ossicula in this genus, we
* see at once that the majority of their salient features ally them, as a whole, to lower
genera. The nature of the incus, and the few points given above regarding the malleus,
show a decidedly high,tendency in Semnopithecus, compared at least with Macacus.

In the malleus from a large skull of Colobus satanas (Pl. LVIII. fig. 8), propor-
tionally smaller than the same ossicle in a Semnopithecus of the same size, the head is
rather ill-developed and flattened laterally—a condition seen, but never to so marked a
degree, in Macacus and Cynocephalus. But there is a distinct, short, constricted neck,
very like that of the last genus, with a well-developed processus brevis ; only the processus
muscularis is abortive, and the tip only slightly spatulate. The incus is distinctly higher
and narrower than in the genus Semnopithecus ; and its crura arise nearer to each other.
Hence, as to these bones, Colobus must take a lower place than Semnopithecus, the ill-
developed head of the malleus being apparently a generic peculiarity, and the incus
allying it decidedly to succeeding genera. But the well-developed processus brevis, and
the still distinct neck, are marked affinities to the genera already considered.

Cercopithecus (P|. LVIII. fig. 9) is central among these Old-world monkeys in its
ossicula. The head of the malleus is fairly developed; the articular surface, though
shaped as in Homo, assumes the position usually observed in lower animals, the outer
extremity lying but little higher than the inner. A trace of lamination, as described in
Semnopithecus, exists; the root of the processus gracilis is very thin and fragile. The
neck is always extremely short (in C. cynosurus, however, I have found it nearly as long
as in S. entellus); and the manubrium runs almost in its axis. This is similar in
form to that of the last two genera, and bears a distinct processus brevis, never so well
- developed as in any higher monkeys, excepting Simia. In C. diana it is more prominent
than in C. cynosurus or in C. lalandit. The extremity is always spatulate, often dis-
coidal (as in a Diana-monkey, of which the ossicula are in the College collection). But
there does not seem to be any convexity above the tip, as described with Semnopithecus.
A more or less distinct trace of a processus muscularis is always seen upon the inner
border of the manubrium*. The incus is very high and narrow in C. cynosurus, but in
the other species resembles that of Colobus; the processus longus is very straight and
slender. The stapes is as described with Semnopithecus; in one from a fine C.diana the
base is so broad that the whole bone forms almost an equilateral triangle.

In Wacacus (Pl. LVITI. fig. 10) the malleus assumes an extreme condition in this
group. The head presents a surface flattened laterally, and projecting considerably for-
wards. This is best seen in WU. nemestrinus and WZ. erythreus; in I. cynomolgus it is
nearly as small as in Colobus. There is a trace of lamination and a stout processus

* This is represented in a sketch of the malleus of C. sabeus in Owen's ‘Anatomy of Vertebrates, vol. iii.
fig. 177 c.
SECOND SERIES.—ZOOLOGY, VOL. I. 3F

382 MR. A. H. G. DORAN ON THE MORPHOLOGY

gracilis, of which a great part is persistent in adult Macaques. Although the neck does
exist, it is extremely short, especially its anterior wall, the manubrium running almost
in its axis. The latter seems to spring from immediately below its articular surface ; its
extremity is often quite discoidal; the processus brevis may be completely obsolete, but
it is fairly marked in some species; the processus muscularis is in the same position as in ~
Semnopithecus, and is generally well developed.

The body of the incus is always either square or higher than broad; the anterior
segment is nearly as high as the posterior. Hence it approaches (in the first pecu-
liarity) the form seen in some specimens from Hylobates, and (in both the above points)
many of the Cebide. Both processes are rather long and slender; the long crus is
very straight till close to the tip. The stapes has straight crura and a broad base, as.
in all the past genera.

In Cynopithecus (Pl. LVIII. fig. 11) the head of the malleus is as ill-developed as in
Colobus, the neck and manubrium closely resemble the same in the Macaques. The
incus has an extremely stout square body; the anterior segment is very well developed.
The crura are very much stouter and shorter than in any Macacus of the same size,
and shorter, though of the same thickness as in Cynocephalus. 'This incus, with its
square body and short stout crura, is as different from the anthropoid type, and as
strikingly reminds the observer of the shape of the bone in Cebus and Ateles, as the high
narrow incus of Hylobates and Macacus diverges from the type of Homo in a direction
towards the form prevailing in Mycetes and in other less typical Cebide.

The stapes, too, though very similar to the prevailing form in this group see Semno-
pithecus), has a narrower base; nor are the crura so far apart as usual.

In Cynocephalus (Pl. LVIILI. fig. 12) the characters of the malleus are like those of
Macacus, but rather more marked. The head is not so developed in proportion to the
size of the bone, is extremely flattened laterally, and projects very sharply forwards. The
neck is also extremely shortened, especially in its anterior wall; the manubrium runs —
almost in its axis. This process has a very well developed processus brevis, and a sharp,
extremely prominent muscular process almost halfway down the inner margin of the
handle.

The incus has a rather shallow body, and very stout crura; the anterior is rather long.
It is clearly of a higher type than in Macacus.

The stapes of the Baboons does not differ from the common type in this group, and is
in outline almost an equilateral triangle.

In conclusion, it may be said of the Old-world tailed-monkeys :— —

I. That they all depart from the apes and Man, and resemble lower monkeys and
most other mammals in the straight and little-divergent crura of the stapes. A pro-
cessus muscularis on the malleus is almost constant.

II. In the distinctly-necked malleus, with a well-formed head, and in the incus,
broad between the crura, Semnopithecus approaches the Simiide, especially Hylobates ;
but the stapes is not at all anthropoid.

III. In Cercopithecus the malleus is nearly as high in type, but the incus is either
square-bodied or high and narrow, as in lower monkeys.

OF THE MAMMALIAN OSSICULA AUDITUS. 383

TV. In Colobus (and Cercocebus) the head of the malleus is almost as ill-developed
and sometimes more ftattened than in Macacus, and the incus is of the same form.

V. In Macacus, Cynopithecus, and Oynocephalus the malleus has a very short and
constricted neck; the manubrium, which bears a processus brevis and muscularis, forms
a very wide angle with the rest of the bone; the incus and stapes exhibit constantly
the tendency to low types, already seen occasionally in higher Quadrumana.

THE OssICULA OF THE CEBIDA.

The ossicula of the Spider-monkeys aman present some singular peculiarities not
observed in neighbouring genera.

The head of the malleus (Pl. LVIII. fig. 18) is extremely flattened vertically as well as
laterally ; seen from above it appears as a smooth, convex, almost quadrilateral surface.
The articular area is very wide, far more extensive than in any higher monkey; the
upper segment or facet is not so wide as the lower, and they meet each other at a very
open angle without a trace of agroove. The upper lies almost vertically above the lower
segment, so that the whole surface resembles the condition seen in the lower Mammalia,
and is unlike the type of Homo. Anteriorly a little friable layer of bone fills up a
part of the angle between the head and the processus gracilis, the stump of which is
stout. This homologue of the lamina seen in some mammals is often broken off and
absorbed in the mallei of old Spider-monkeys. The head never bulges prominently
forwards.

So far the malleus bears characters common to the other Cebide; but it differs from
them all in possessing a singular neck, quite unlike that of the Anthropomorpha, though
almost as long. Through the kindness of Prof. Garrod I have been enabled to examine
numerous skulls of Ateles melanochir, A. hybridus, and A. ater, from specimens once
living in the Zoological Gardens; and I find these characters best marked in A. mela-
nochir. Immediately below the articular surface the neck is very stout, and it becomes
narrower close to the handle, though even there it is much less constricted than in the
apes and Old-world monkeys. The very short and curious manubrium forms with the
neck an angle often not more than 120°, though occasionally much wider. Constricted .
near the base and tip, it bulges in a conspicuous manner in the middle, and terminates
in a wide discoidal enlargement. The processus brevis is represented by a prominent
rounded knob. There is no processus muscularis; and the tendon of the tensor tympani
muscle is inserted into the root of the manubrium close to the neck. This malleus is
certainly the most specialized among all the Quadrumana.

The incus has avery well-developed and very square body; in a specimen from 4. hybridus
I find it slightly longer horizontally than vertically. The anterior segment is almost as
high as the posterior; and the articular surface, looking rather outwards, appears unusally
deep from the outer surface of the body, and unusually shallow from the inner aspect.
The incus, on this account, will be seen to be most like that of Cebus, except in the last
feature, not showing that tendency to great height and narrowness so frequent amongst
other American monkeys.

The stapes has a well-developed head, and Ise slender crura, not very divergent, and

3F 2

384 MR. A. H. G: DORAN ON THE MORPHOLOGY

both inserted some distance from the extremities of the base, which is slightly convex
towards the vestibule.

The cruraare longer than in most Cebide, where the stapes closely resembles that of the
Old-world monkeys, only differing from them, as a rule, in the greater length of their
limbs; hence this distinctive feature of the entire family is best marked in Afeles.

In the Howlers (Mycetes) the head of the malleus is of the same form as in A/eles,
but still more flattened vertically and still less developed, the height of its convexity

Yhardly rising above the level of the upper border of the articular surface (Pl. LVIII.
fig. 14). This latter is extremely wide in all directions and very shallow; both facets
are very saddle-shaped. There is absolutely no neck; but immediately from below the
articular aspect springs the manubrium, which is tolerably long, forms an even, rather
wide curve, and becomes gradually narrower towards the extremity, which ends in a
perfect disk. There is not a trace of a processus brevis; and the processus muscularis is
almost obsolete. In IZ. seniculus the manubrium runs almost in the axis of the body;
in 1. laniger and WH. ursinus it forms a distinct angle.

The malleus of Wycetes, so different from that of Ateles, where there is a thick neck
and a manubrium bulbous in the middle and possessing a tubercular processus brevis,
may be considered a good type of this bone among the Cebide, and to a certain extent
among the less-typical Lemurs. It differs strikingly from the same bone in Man, Apes,
Old-world monkeys, and the genus Lemur.

The incus is very deep vertically, and narrow horizontally, with thick and short crura
arising close to one another. The articular surface has the peculiarity noticed in Afeles.
This. bone, too, is very typical among the Cebide.

The stapes is of the ordinary monkey type, the base concave laterally, with a trace of
an umbo ina specimen of I. laniger in the College collection, but not in one of JZ. wrsinus,
where the crura are almost fused; a minute aperture which exists between them is, I have
reason to believe, accidental *.

In Cebus (Pl. LVITI. fig. 15) the malleus differs little from that of Mycetes; the head
is almost identical in form. The manubrium runs still more closely to the axis of the

_ body; and no rudiments of a processus brevis or muscularis can be traced. The handle
does not form the free and regular curve seen in most Howling-monkeys (Mycetes), but
is slightly bent near its root, and straighter towards its extremity, which is a little turned
up and almost a perfect disk. é
. The incus departs from the type of Mycetes, Pithecia, Nyctipithecus and the Marmosets,
the body being generally longest horizontally, or quite square. The posterior crus is
rather thick at the base; it diverges, as in most Cebide, very little from the body. The
processus longus is often long and thin, and always divergent.

The stapes is of the usual monkey form; but in most specimens the posterior crus is
always slightly bowed and thicker than the anterior.

In the Douroucoulis (Wyctipithecus, Pl. LVIII. fig. 16) I do not find any distinction

* « Bei den Briillaffen ist die Basis konkayv, aber in der Mitte als Umbo gegen das Vestibulum vorgetrieben. Bei

einem Sapajou dessen Species unbestimmt ist, finde ich in beiden Ohren die Crura des Stapes verwachsen.”—Hrrrt,
op.cit. But I have noticed an Umbo in an ape (see Hylobates).

OF THE MAMMALIAN OSSICULA AUDITUS. 385

of importance in the malleus from the same bone in Cebus, except that a rudimentary
processus muscularis is to be found halfway down the inner border of the manubrium in
Nyctipithecus. The incus, on the other hand, is always high and narrow; the posterior
erus hardly diverges from the body, the slender processus longus diverging considerably.
The stapes has long slender crura and a comparatively narrow base.

In the Squirrel Monkey (Saimaris sciurea, Pl. LVIII. fig. 17) the malleus has the
general characters described with Cebus; but the articular surface is still shallower, and
the manubrium runs much less nearly in the axis of the body, often forming an angle
of not more than 150°. The incus is higher than broad, resembling that of Myctipi-
thecus in its crura, as does the stapes.

In a single specimen of the malleus and incus of the Black Saki (Pithecia satanas,
Pl. LVIII. fig. 18) the former ossicle has a manubrium rather thicker at the base, and
more distinctly flattened laterally (as in most mammals) than in the above-described
genera. ‘This process, too, forms the same wide curve seen in Mycetes; and the incus
has the same extremely high narrow body as in the Howlers; the vertical measurement
of the posterior segment is even greater than in those monkeys.

After careful observation, I consider that the malleus in the Hapanip# (Marmosets)
is intermediate between the same in the Cebide and in the Nycticeboid lemurs. The
description given of the malleus in the genus Cebus will apply to Hapale with two important
distinctions—namely, that the articular surface is always narrower and deeper in the
Marmoset, and in that animal the whole body is bent more inwards on the manubrium,
a character much more prominent in the Nycticebide, but in Hapale the head of the
malleus is longer in its vertical measurement than in Nycticebus. The body of the
incus is quite square in H. jacchus; but in a specimen of H. melanura in the College
collection it is as high and narrow as in the Douroucoulis (NMyctipithecus felinus and
N. trivirgatus). The stapes is of a different form from the type prevalent in the last
family. The head is very long; or, more correctly speaking, the crura are fused half-
way to their insertion, where they suddenly diverge towards the base. This is very
- like the condition seen in many of those marsupials where the crura are not fused

throughout. Hence in the genus Hapale may be found the lowest type of ossicle seen
in all the Primates (Pl. LVIII. fig. 19).

In Midas (Pl. LVIII. fig. 20) the malleus is of much the same form as in Hapale, the
articular surface also is rather deeper and narrower than in the Cebide; but the ten-
dency of the body to be bent inwards on the manubrium is not so plainly seen, at least
in YN. edipus, rufimanus, and geoffroii, as in Hapale. In all these species the incus is
high and narrow. The stapes in WZ. edipus has a small head with free crura and a wide
aperture; but one from UW. geoffroyii in the College collection has the peculiarities of
that ossicle in Hapale.

The absence of a neck to the malleus of most of the New-world monkeys seems not to
be due to an absolute suppression of that part, but rather to its being bent down below
the articular surface, and lying actually in the axis of the manubrium. In many mallei

from the genus Cebus a slight rounded projection may be detected high up on the outer
edge of the handle. In such cases, if the small segment of the manubrium above that

386 MR. A. H. G. DORAN ON THE MORPHOLOGY

prominence were raised to the exact axial line of the head (for even in these cases the
manubrium is never literally in that axis, but always slightly bent on the head), the rest
of the handle retaining its angle, we should have an ossicle almost exactly like the
malleus of Ateles. The processus muscularis, placed so low down as the inner edge of
the manubrium, would then be brought close to the neck, its normal or, at least, most
usual position.

THE OSSICULA OF THE LEMURID.

In the genus Lemur (Pl. LVIII. fig. 21) the malleus has a well-developed head, rising
considerably above the level of the articular surface, which is tolerably deep, and much
like the same in the Old-world monkeys. A faint trace of a neck exists ; the manubrium
forms an angle of 150° with the rest of the ossicle, and ends in a broadly spatulate or
discoidal termination. In the specimens of ZL. varius and L. albifrons in the College
collection, the manubrium is much flattened near the base, as in Homo; and in ZL, albi-
frons a distinct processus brevis exists; but in L. catta, L. nigrifrons, and L. xanthomystax
the base of the handle is narrow, and hardly perceptibly flattened; but probably in any
species of this genus a processus brevis may be occasionally developed. A small area of
lamellar bone is generally to be seen between the head and the stump of the processus
gracilis. The malleus of Lemur bears more resemblance to the same in the Old-world
monkeys than to the Ceboid type—not so, probably, from any nearer relation of the Lemurs
to the Macaques &c., which is most unlikely, but rather from the fact that the malleus
of the American monkeys is highly specialized, while it happens that the same in Lemur
and the tailed Old-world monkeys is of a far more generalized type in all cases.

The incus decidedly varies in form in different species of Lemur. In the College
specimens of that bone in Z. varius the body is wide horizontally and shallow vertically,
with stout widely divergent crura. In ZL. nigrifrons and L. xanthomystax the body is
both shallow and narrow, the crura stout and but little divergent, so that the ossicle has
avery tooth-like appearance. In Z. albifrons and L. catta the body is square, and the
processus longus is slender and very divergent. cH

The stapes has always long and extremely slender crura, very slightly bowed outwards,
with a wide aperture. The base is invariably convex towards the vestibule, and rather
narrow vertically. The space between the crura is in this genus always occupied by a
bony canal, which runs between two projections of the petrous bone above and below
the fenestra ovalis, and transmits a small blood-vessel—a frequent condition among some
Insectivora and Rodents. The same condition is seen in a skull of Hapalemur in the
Museum of the Royal College of Surgeons.

Among the Indrisinee, Propithecus (Pl. LVIIL. fig. 24)* has a malleus more like
that of Cebus than Lemur. The head is of much the same shape, and the articular
surface of the same depth, as in the latter; but there is the same absence of all trace of
neck, so frequent in the American monkeys. The manubrium is sharply curved in the
middle, and terminates in a very large disk. The incus is wide horizontally; the pos-

* Sketched from a young specimen. The presence of the processus gracilis, with some friable tissue between it and
the neck, alters its aspect. :

OF THE MAMMALIAN OSSICULA AUDITUS. 387

terior segment is much deeper than the anterior; and the crura are thick, and both
divergent. The stapes resembles that of Lemur; but the crura are stouter. In both
the skulls of Propithecus in the College Museum a bony canal passes between the
crura of the stapes, as is the case in a skeleton of Indris (Lichanotis) brevicaudatus. In
a skull of the Avahi (Jndris laniger) I have found the trace of a bare vessel, not supported
by bone, instead of an ossified intercrural canal.

The malleus of Galago (Pl. LVIII. fig. 22) strikingly resembles that of the Hapalide ;
but the upper part of the head is better developed, the articular surface is still narrower
and deeper, and the head is much shorter vertically than in Hapale or Midas. It is
quite neckless, and bent very much inwards on the manubrium—an exaggeration of a
condition frequent in the mallei of Primates where the neck is suppressed. The manu-
brium is slender throughout, even at the base, and ends in a very distinct disk; there
is a processus muscularis halfway down the inner margin. The incus is very high and
narrow, as in Wyctipithecus, Pithecia, &c.; and the anterior segment is actually as high
as the posterior, and projects forwards conspicuously. The processus brevis runs in the
long axis of the posterior part of the body, with hardly a trace of divergence, the pro-
cessus longus diverges extremely. This arrangement of the crura recalls Nyctipithecus,
and is usual in the Lorises. The crura of the stapes are slender, and perfectly straight ;
the intercrural space is moderate; and no bony canal appears to occupy it in the
recent skull.

In a specimen of the mallei (Pl. LVIIT. fig. 23) and incudes of-Cheirogaleus cookii,
the malleus, although resembling Galago in its head and articular surface, has a
manubrium which springs from immediately below the articular area of the head, yet
bears a distinct projection corresponding to the processus brevis of Man. This pecu-
liarity is seen in some squirrels, a family where gradations between the necked and
the neckless malleus are as numerous as in the Primates. The handle, too, in Chei-
rogaleus is rather flattened naturally. This ossicle altogether is related not distantly
to the same in the genus Lemur; but the incus assumes those conspicuous features seen
in Galago; the divergence of the anterior crus is extreme. There appears to be no
bony intercrural canal supporting the stapes in this genus.

In studying the Nycticebide or Lorisine, I find in the malleus of Wycticebus
(Pl. LVIII. fig. 26) that the characters of the neckless type of that ossicle are all
prominent. The head is very short in its vertical measurement, very much bent
inwards on the manubrium, and bears a deeply cut but not very wide articular surface.
All these features are seen in the Galagos, although the last two are approached in the
Marmosets. The singular manubrium is long and very straight; its outer border is
somewhat wavy; and there is a distinct projection and processus muscularis for the
tendon of the tensor tympani halfway down the inner margin. The handle terminates
in a large disk. The strong resemblance of the malleus of Nycticebus and of some other
Primates to the same ossicle in the Tupaiidee and Squirrels is interesting, especially as
they are all small thickly furred arboreal animals.

The incus has a shallow body, quite unlike that of Galago; but the anterior segment is
nearly as high as the posterior. Its articular surface is completely on the top of the

388 MR. A. H. G. DORAN ON THE MORPHOLOGY

body (strictly expressed, on the antero-superior aspect), and does not lean outwards, as
described in Ateles, and as found more or less in all the Cebidze and Hapalide. More-
over this surface is in Nycticebus very saddle-shaped, indeed almost convex. The pos-
terior crus is stout, short, and slightly divergent; the stapedial crus is stout, straight,
long, and very divergent, supporting an “os orbiculare”’ or Sylvian apophysis, which is
small as in all Lemurs.

The stapes has slender crura, not very divergent ; the base is rather thick and almost
plane. I have never found any intercrural bony canal in any skull of Nycticebus, Arcto-
cebus, Loris, and Perodicticus.

In a malleus of V. javanicus the articular surface is deeper cut than in WV. tardigradus.
The characters of the ossicula of Loris gracilis are almost identical with those of
Nycticebus ; only the head of the malleus is even still shorter vertically, and the anterior
part of the body of the incus projects forwards somewhat as in Galago. In Arctocebus
calabarensis (Pl. LVIII. fig. 27) the extreme characters of the neckless malleus are even
better-marked than in Zoris; the head of the malleus is shorter than in any of
the Platyrrhine monkeys, and the articular surface is far deeper cut than even in
Hapale. :

Perodicticus potto (Pl. LVIII. fig. 25) resembles Nycticebus in its malleus and
stapes; but the incus is higher than broad, its articular surface being, however, similar
in position to the same in the tardigrade Loris.

In Tarsius spectrum (Pl. LVIII. fig. 28 *) the malleus, which is quite neckless, has a
head of square outline, not nearly so compressed vertically as in the last family, and
projecting considerably forwards. The articular surface is deep, and rather wide; the
manubrium is slender, ending in a spatulate extremity; there is a trace of a processus
brevis ; and the tensor tympani is inserted into the inner border of the manubrium midway
between the tip and the root. The incus has a square body; both crura diverge con-
siderably. The stapes is small, even in proportion to the size of the animal; its erura
are very thick, and its base almost plane.

- Whilst the characters of the ossicula of the aberrant Tarsiws are somewhat negative,
the sole representative of the Chiromyide presents very salient features in its ear-
bones. :

All the ossicula of the Aye-aye (Chiromys madagascariensis, Pl. LVIII. fig. 29) are
much larger in proportion to the size of the animal than inany tailed monkey. The mallei
in the College collection are larger than any from A‘eles, and as large as that ossicle in
Cynocephalus. The head is much flattened vertically ; its upper free surface is broad
and concave, far more developed than in the other Lemures. The articular surface is
deeply cut and wide vertically; the two facets are well-marked: the upper is very wide,
and the lower is very small. There is a distinct short neck, twisted, together with the
head, somewhat inwards; the manubrium is long, rather narrow laterally at the base,
and forms a similar curve and angle to what is seen in the anthropoid apes. There is a
small angle corresponding to the processus brevis, and a distinct processus muscularis

* Taken from a new-born Tarsius. The remark made in the foot-note to the description of Propitheeus applies
to this as well.

OF THE MAMMALIAN OSSICULA AUDITUS. 389

on the inner edge of the manubrium nearer the root than the tip. The outer aspect
of the manubrium facing the tympanic membrane, though very narrow, is distinctly
bordered off from both sides by sharp edges, till it dilates very gradually into the narrow
spatulate, recurved extremity. This distinct outer surface to the manubrium is seen
in the Carnivora, Ungulata, and many Rodents. As it is in the Rodentia that the
processus muscularis is so frequently placed completely on the manubrium, the extensive
lamina being generally absent in that order, it follows that the malleus of Chiromys is
more allied to the Glires than to the flesh-eating and ungulate Mammalia.

The incus is remarkable for the sharp projection backwards of its posterior segments ;
the erura are stout, short, but neither are strongly divergent; the Sylvian apophysis
is extremely elliptical, or rather key-hole-shaped in outline, being broad horizontally
and very thin vertically. The crura of the stapes are stout, the posterior much bent,
as in Homo; the intercrural space is wide, and not occupied by a bony canal in the
recent skull.

The above researches into the ossicula of the Platyrrhini and the Lemuride will now
be briefly summarized.

I. In Afeles the malleus is of peculiar shape. It has a short neck and a rounded
prominence corresponding to the processus brevis. The articular surface is shallow, as
in all the Cebide. The incus has not a very high body; the crura of the stapes are
longer than in other monkeys.

IT. In Mycetes and Pithecia the neck of the malleus is quite suppressed (together
with the processus brevis), and the manubrium at its root runs in the long axis of the
head. The incus has a very high narrow body. The stapes, in these and the genera of
the Cebidze yet to be mentioned, ‘has not so long crura as in Afeles, but is of the
same form as in the Old-world monkeys.

III. In Cebus, Nyciipithecus, and Saimaris the malleus much resembles that of
Mycetes; but the head is bent on the root of the manubrium. In Cebus the body of
the incus is not higher than in Afeles; but in the other genera it is almost as high as in
Mycetes or Pithecia.

IV. In the Hapalidz the malleus has a rather deeper articular surface than in the
Cebidze; it is quite neckless; and the head is bent on the manubrium. ‘The incus is
generally high in the body. In the Marmoset the crura of the stapes are often fused for
some distance below the head, as in mammals of a very low grade.

V. Among the Lemuridx, the Nycticebide, Galaginide, and Propithecus (at least in
young specimens of P. diadema) much resemble the Hapalidze and Cebus in the malleus ;
but the articular surface is deep, deeper than even in Hapale; the head is very
short, and strongly bent on the manubrium. The incus has a very high body in the
Galagos, but not in the Nycticebide. Tarsius and Perodicticus resemble Nycticebus.

VI. But in Lemur the malleus loses the specialized form common in the other
Lemuride and the New-world monkeys. There isa constricted neck, and a more or less
developed processus brevis; the manubrium forms a distinct angle with the neck,
instead of the head being bent on it as in the Galagos &c. The incus is not high and

SECOND SERIES.—ZOQOLOGY, VOL. T. 36

390 MR. A. H.G. DORAN ON THE MORPHOLOGY ~

narrow. ‘There is always a bony canal between the crura of the stapes. This is seen in
the Indrisinz as well, but does not appear to exist in Galago.

VIL. In Chiromys the ossicula are large, and their affinities indistinct. The malleus
hasa much flattened head and a trace of processus brevis; in most respects it is of the
rodent type. The incus is of a somewhat peculiar shape; the stapes has curved crura,
a character seen in the higher monkeys, but also in many of the Sciuromorpha.

THE OSSICULA OF THE CARNIVORA.

The Fissipedia are almost as remarkable for the very trifling variations in the ear-
bones of the different families, genera, and species as the Pinnipedia are for the boldly
defined individual characters of all three ossicula in closely allied genera. The study of
these bones in the latter suborder is rendered all the lighter on account of their great size.
Yet while the Pinnipedia maintain, in the malleus at least, a certain adherence to the
common and most typical form of ossicle, the Fissipedia possess a type of malleus
unusual except among certain Ungulates and Rodents.

The Fissipedia must first be considered; and the study of their ossicula shows that
distinctive features of any clear value in classification are chiefly confined to the malleus,
the variations in the form of the ineus being more limited, although of interest in some
families.

The divisions of the suborder used here are those given by Flower* (P. Z. 8.
1869, p. 4).

The typical malleus of a fissiped carnivorous mammal may thus. be described :—It has
a small head, with a deeply cut articular surface, of which the facets are divided by a deep
well-marked groove; the neck is thin, long, cylindrical, and more or less sharply curved,
bearing on its inner side a muscular process for the tendon of the tensor tympani. The
manubrium forms with the neck an angle seldom over 100°; it has an angular pro-
jection at the outer side of its base, which may be dignified by the name of processus
brevis, although never developed to the extent seen in Man. It is narrow laterally;
but the side towards the membrana tympani is constantly separated from the two other
sides by sharp borders, and is often very broad,

But the great and conspicuous feature is the broad lamellar expansion between the
head and neck, on the one hand, and the very slender processus gracilis, which often
appears as merely a thickened lower border of that bony lamina, undoubtedly a minor
modification of the condition existing to an extreme degree in the Hedgehogs, Shrews
and Bats. This is observed to a corresponding extent in the Ruminants and larger
Muride.

The incus in the Fissipedia is always small in proportion to the malleus, directly con-
trary to what obtains with the Seals. Excepting extreme shortness of the processus
brevis in some families, there are few distinctions of importance between the incudes of
the various members of this group. The same applies to the stapes; and such slight
differences as exist will be noted in the description of individual genera or species.

* Tn his paper “ On the Value of the Characters of the Base of the Cranium in the Classification of the Order
Carnivora, and on the Systematic Position of Bassaris and other disputed Forms,”

OF THE MAMMALIAN OSSICULA AUDITUS. 391

In the Frnip#& the malleus (Pl. LVIII. fig. 32), a bone of light construction, exhibits
the general characters already described. The head is moderately developed; the arti-
cular surface is narrow vertically, but very deeply cut; the upper facet is nearly flat,
the lower quite convex; they are separated by a groove, which is very sharply marked
towards the inner limit of the articulation, but fades completely before reaching the
outer border. In the characters of this surface the smaller Cats may be known from the
Civets. The manubrium is not only narrow at the base, but its outer aspect, though
sharply bordered anteriorly and posteriorly, is extremely narrow throughout; and the
whole process, though well recurved at the tip, is not strongly bowed, with its concavity
forwards. The last two points distinguish the feline malleus from that of the Dogs,
the Hyzna, and Proteles. The extremity is only slightly spatulate, as in most Carnivora.
From the inner side of the thin, curved neck proceeds a long sickle-shaped processus
muscularis, as in the Canidze and the Weasels, much more slender than in Profeles and
Hyena, and better developed than in Cryptoprocta, the Viverride, Procyonide, and
Ailuride ; in the Bears it is absent. The lamina is well developed, though never as
wide as in the smaller families; even in the largest Lions and Tigers it remains papery
and semitransparent, not so compact as in Hyena.

The body of the incus (Pl. LVIII. fig. 42) is deep; its posterior crus is always well-
developed; the Sylvian apophysis is small.

The stapes (Pl. LVIII. fig. 56) has straight crura, with a wide aperture between them ;
and the thin but broad base is more or less convex towards the vestibule.

All the above characters are best seen in the Lion, where the lower facet of the arti-
cular surface of the malleus is unusually concave. In all the animals this facet lies
rather external to the upper, the outer extremity of the articular surface is higher
than the inner, but never to the extent seen in Man and in many monkeys. In the
Tiger, the manubrium is rather more slender than in the Lion. In the Cat (Pl. LIX.
fig. 1) the angle at the outer aspect of the base of the manubrium projects so much as
to form a true “processus brevis.” In the Leopard, the groove between the facets
of the malleus almost reaches the outer border of the articular surface, and there is a
distinct concavity of both facets external to the point where the groove ceases, an
approach to the Viverrine type. In the Cheetah (felis jubata) the manubrium is some-
what shorter than in the above cats, and more distinctly spatulate at its termination.
The Serval has a stapes with long slender crura, much less divergent than in most
Carnivora. In F. lynx and F. caracal the malleus has a stouter processus gracilis, and
is altogether more solid than in other cats of the same size. This is not the case with
F. caligata, which has a very broad lamina to the malleus, as in the European Lynx.
These smaller Felidz retain the characters of the articular surface of that bone as seen
in the Lion.

In the Hymyip# (Pl. LVIII. fig. 34) the malleus is of stouter build than in any
other family of the Fissipedia, excepting, perhaps, certain large Canidee; the head is
greater than in the Cats, yet not large in proportion to the development of the rest
of the bone. The articular surface is wider than in Felis; and the groove, well marked
internally, fades in the middle of the articulation; the facets it divides are flatter.

3G 2

392 MR. A. H. G. DORAN ON THE MORPHOLOGY

The neck is of the character seen in most Carnivora, but proportionally rather’ thicker.
The manubrium is much thicker than in Felis, though not broad at the base laterally.
Tts outer surface is moderately broad above, but very broad towards the point; thus,
lying exactly intermediate between Fels and Proteles, the whole process is rather’
more curved, with the concavity forwards, and less recurved at the tip than in the
Cats. The great feature in this and the next family is the processus muscularis, which
is very stout at the base, blunt at the tip, and almost straight. The lamina is denser
than in the Cats and Dogs. The incus (PI. LVIII. fig. 48) resembles that of a large Felis.
The stapes (Pl. LVIIT. fig. 58) is rather larger than in the other Carnivora; its crura
are slender, as in the greater Felide, not thick, as in Wolves.

In the Aard-Wolf (Proteles, Pl. LVIII. fig. 35) the head of the malleus is smaller and
less convex than in Felis or Hyena; its articular surface approximates to the Viverrine
type: both the facets are convex internally; but they run into one another, forming a marked
concavity external to the point where the groove ceases, the latter only running half across
the surface, as in Hyena. The manubrium has its outer surface broad throughout, as
in the Dogs; this character distinguishes it from Hyena, which it resembles in having
an almost equally straight and solid processus muscularis. The body of the incus is
shallow, as in Viverra and Canis; both crura are very stout and well developed.

Among the Viverride two distinct varieties of malleus are to be found. One, unlike
any other among the Fissipedia, is seen in Herpestes and Suricata, and in all probability
in allied genera; the other, which will be described first, is found in the genera Viverra,
Paradoxurus, Nandinia, Arctictis, Cynogale, and Hupleres.

The malleus of the Civet and its allies possesses very negative characters, or may be con-
sidered to present, together with those of the Cryptoproctidz, Procyonidz, and Ailuride,
a central type, from which those of the Cats, Dogs, and Bears are much specialized, the
above-named families exhibiting in this bone very slight tendencies towards the modi-
fications observed in those more important groups, which they respectively approach in
their general anatomy. Taking the most prominent point of distinction between the
mallei of terrestrial Carnivora, namely the form and development of the processus
muscularis, we find that process long in the Cats, and absent in the Bears. Now,
as it is of moderate length in the Civets and Racoons, from a quite natural moditication
of the characters of the bone in the two former families, it comes to pass that the
latter animals, very different from each other in the whole skeleton, come close to each
other as far as the malleus is concerned. F

The head of this bone in Viverra (Pl. LVILI. fig. 37) is small, its free surface pro-
jecting but little above the upper margin of the articular area. The articular surface
is narrow vertically and deeply cut; both facets are convex; the groove between them
is very deep, and, as in the Cats, it runs from the inner border to the middle of the
surface, where, however, it does not fade, as in Felis, but bifurcates; the two branches
run to the outer surface, and enclose a triangular, somewhat concave space, which may
be considered a third facet, and is clearly a further modification of the condition de-
scribed in Proteles, where the two branch grooves do not exist, but the facets run,
externally, into a concavity. In many faintly marked Viverrine mallei the bifurcation is

OF THE MAMMALIAN OSSICULA AUDITUS. 393

effaced; then the articular surface appears precisely as in Proteles. The neck is always
thin, and much curved towards the manubrium, which latter process is very long and
slender in all directions, even near the root; the angle corresponding to the processus
brevis is blunt; the outer surface, though well bordered, is extremely narrow throughout,
the terminal dilatation being almost imperceptible; and the point is little recurved.
The whole manubrium is very straight, as in the Cats, never forming the curve seen in
the Dogs and, to a less degree, in Hyena and Proteles. The processus muscularis is a
small mammillary projection, never quite absent. The processus gracilis forms a wide
angle with the neck, so that the lamina is extensive.

The incus (Pl. LVIII. fig. 45) is a small bone with a shallow body, bearing a pro-
portionally deep articular surface. Both crura are well developed; the posterior diverges
little from the body.

The stapes (Pl. LVIII. fig. 60) has always very slender crura, with a large space between
them, and a thin, broad base ; the former are inserted close to each extremity of the latter.

The Viverrine malleus strikingly resembles that of certain ruminants ; but the articular
region in those ungulates is of a different character, and the other two ossicles are
unlike those of the Carnivora.

Viverra cwetta exhibits all these characters. The processus muscularis of the malleus
is well developed, and, in a small way, approaches the straight conical type seen in Hyena
and in Proteles. In V. malaccensis that process is very short, and the manubrium is
very slender and almost styliform at the tip. In Genetta vulgaris the processus muscu-
laris is short, but still prominent; otherwise the malleus is as in Viverra; but the crura
of the stapes, though very thin, are not so extremely slender in this animal and in the
remaining Viverride as in the true Civet Cat. In Nandinia binotata and Paguma
grayit, as well as in Cynogale bennettii (Pl. LIX. fig. 13 a), the processus muscularis is
still short; but in Paradoxurus bondar and musanga it is reduced to a conical tubercle.
In Hupleres goudotiu (Pl. LIX. fig. 10) the head of the malleus is proportionally larger
than in the Civet; but the processus muscularis is as long, and the neck as well deve-
loped, as in that animal. The incus has a slender and well-developed processus brevis,
as in most of the Cats, Dogs, and Viverridze. The stapes is larger in proportion to the
incus than even in Herpestes. In Arctictis binturong (Pl. LIX. fig. 2) the malleus is
much more distinct. The neck makes a very short curve before joining the root of the
manubrium, so that the whole bone appears “hog-backed.” The manubrium is much
broader at the base and more recurved at the tip than in any of the preceding genera
—a distinct approach to Herpestes; but the processus muscularis, a mere tubercle, as in
Paradoxurus, still arises from the neck, and the lamina is well developed*.

One cannot fail to be struck by the great difference between the malleus of Herpestes
(Pl. LVITII. fig. 88) and that of the preceding Viverridee. The Ichneumons are the only
Carnivora where that auditory ossicle approaches the somewhat central type seen in
Man. The head is large, with a free orbicular surface; it bears an articular surface
sinilar to that of the Civets. The neck is short and stout; the manubrium is much
broader at the base, and more recurved at the extremity, than in any of the past genera

* The figure is from a specimen where the lamina was mutilated.

394 MR. A. H. G. DORAN ON THE MORPHOLOGY

except Arctictis, and forms a very obtuse angle with the neck; the processus brevis is
generally well marked. The short, stout, conical processus muscularis does not spring
from the neck, but from the inner edge of the manubrium, a perceptible distance beyond
its root, approaching a characteristic of some rodents. ‘The lamina is very little deve-
loped, indeed is not perceptible until close examination.

The incus (Pl. LVIII. fig. 47) is rather larger than in the Civets; the stapes is large
(Pl. LVITII. fig. 61), and resembles that of Genetta and Arctictis.

The species do not differ much in their ossicula. The head of the malleus in Herpestes
ichneumon and in H. paludosus is particularly large. In the manubrium of the latter, Prof.
Flower has detected a minute foramen which perforates it close to the root of the pro-
cessus muscularis. In the above species and in H. nepalensis, where the head is smaller,
the processus brevis is well marked; but it is much blunter in H. griseus and H. pulve-
rulentus. In the latter (Pl. LVIII. fig. 38) the processus muscularis is a more con-
siderable distance down the inner edge of the manubrium than in the malleus of any
other ichneumon ; its incus is proportionally larger.

In Suricata zenik (Pl. LIX. fig. 3) the ichneumon type of malleus is maintained. The
head is large, the neck extremely short. ‘The manubrium leaves the neck at a wider
angle than in Herpestes, and is distinctly shorter, broader at the base, and more spatulate
at the tip. The processus muscularis is situated on the inner side of the manubrium, a
perceptible distance from the neck; and the lamina is very little developed.

In the Cryptoprocta the malleus closely resembles that of Viverra; the head is a little
larger, the articular surface similar; the manubrium is even more slender; the processus
muscularis is almost as short as in Paradoxurus. The incus (Pl. LVIII. fig. 44) is a
larger bone than in the Civets; and the stapes (Pl. LVIII. fig. 59) has thicker crura than
in Viverra.

Thus Cryptoprocta ferox is incontestably more allied, as far as its ossicula are con-
cerned, to the Viverride than to any other family; and the latter are clearly more
related to the Cats by the type of their ear-bones, as Hyena and Proteles are related
to the Dogs.

In the Canidz the ossicula have far more marked distinctive features than they bear
in the Viverridee.

The malleus (Pl. LVIII. fig. 33, and Pl. LIX. fig. 5) often rivals that of the Hyzna
in stoutness of build. The head cannot be said to be remarkably developed ; the articular
surface is narrow, and deeply cut; both facets are concave; the groove dividing them
never bifurcates, as in well-marked viverrine mallei; but the facets join in a concave area
external to the point where the groove fades, as in Proteles. The neck is only mode-
rately curved; the manubrium, the most distinctive feature in the canine malleus, is
long, rather thin at the root, and very strongly curved, with the concavity looking
forwards. The outer aspect is a distinct surface, not only sharply bordered from the
sides, as in all Carnivora, but likewise very broad throughout, though rather broader
towards the extremity. Thus the Hyznide, as has been already remarked, resemble the
Dogs in their manubrium ; but Proteles is still more canine. On the other hand, the
Canidee have a long sickle-shaped processus muscularis projecting from the inner side of

OF THE MAMMALIAN OSSICULA AUDITUS. 395

the neck, as in the Felide; it is never straight and conical asin Hyena. The lamina
is broad, and bounded below by a well-marked splinter-like processus gracilis. The body
of the incus (Pl. LVIILI. fig. 43) is shallow; and the crura, which are both of nearly equal
length, are conspicuously far apart, even at their origin. The stapes resembles that of
the Cats. Hyrtl considers that the aperture between the crura is much smaller in the
Dogs; but it is more correct to say that in the large Canide the crura are more massive
than in the Lion or Leopard, and encroach more on that space, which, in small dogs, is
often as wide as in the domestic cat. The anterior crus is often curved.

In large specimens of the Dog the outer surface of the manubrium is so expanded at
its borders that the rest of that process appears as a sharp ridge springing from the inner
side of that external aspect. In the Wolf the same surface of the manubrium is even
broader than in the malleus of a Newfoundland or a Mastiff (although I have prepared
a malleus from a small dog where it is nearly as broad); and the whole process is
extremely curved; the extremity, however, is not much turned up, as in the straighter
manubria of many other mammals. The crura of the Wolf’s incus are extremely far
apart; and the anterior crus of the stapes is always well curved, and thicker than the
posterior. In the Dog the crura are far more similar in straightness and bulk. (See
Pl. LVIII. figs. 33, 48, 57, & Pl. LIX. fig. 5.)

The other species in this genus have ill-marked specific characters. In the European
Fox the neck of the malleus is shorter than in the Dog or Wolf; the outer aspect of
the manubrium is nearly as broad as in Canis lupus; but in C. vulpes (Pl. LIX. fig. 6) it
is nearly of equal breadth throughout. The crura of the incus diverge as much as in
the Dog, and hence less than in the Wolf; the anterior crus of the stapes is slightly
curved, but not much thicker than the posterior. In a specimen of C. fulvus I found
the manubrium rather longer than in the Fox. The crura of the stapes were both
straight, and the opening between them very small. In the Arctic Fox (C. lagopus) the
malleus has a much longer neck than in the two foxes above named, and a longer pro-
cessus muscularis than in any of the preceding Canide ; the incus is wolf-like. In the
Cape Hunting Dog (Lycaon pictus, Pl. LVIII. fig. 41) the processus muscularis is stouter
and straighter, and proportionally larger than in any other of the Canide; it closely
resembles that of Proteles; but the outer surface of the manubrium is quite canine, being
very wide, and almost of equal breadth throughout ; the incus is most like a dog’s in aspect.
In the Long-eared Fox ( Ofocyon) the neck and manubrium mallei are both rather short ;
and the processus muscularis is as long as in Canis (Pl. LIX. fig. 4).

In Canis lupus and familiaris the lamina of the malleus is wider than in the Foxes,
Iycaon or Otocyon.

The MustELIp# possess auditory ossicles which are not so remarkable for any single
distinctive feature or prominently developed process as for their very characteristic generai
appearance. ,

The malleus of the Glutton (Gwlo, Pl. LVIII. fig. 39) has a rather elongated head; the
free globular portion, fairly developed, projects considerably upwards. The articular sur-
face is always wide, contrary to other land-Carnivora ; but the facets are of the same pattern
asin Viverra. The neck is long, and only slightly curved, except in the Badger ; the manu-

396 MR. A. H. G. DORAN ON THE MORPHOLOGY

brium is always short, broad at the base, much recurved at the tip, though not curved
as a whole, with the concavity forwards, as in Dogs; and it forms an extremely obtuse
angle with the neck. There is always a rudimentary processus brevis. The muscular
process is always stout at the base, and tolerably long. The lamina. is very long and
narrow, owing to the very acute angle formed between the neck and the somewhat stout
processus gracilis—a condition diametrically opposite to that seen in the Procyonide,
and a great modification from most other Carnivora.

Careful comparisons seem to lead to the conclusion that the musteline malleus is
intermediate between the typical carnivorous form and that seen in Herpestes and
Suricata. By slightly enlarging the head in this family, and fancying the lamina ab-
sorbed, and shifting the processus muscularis to the root of the manubrium, we have
an ichneumon’s malleus without altering the manubrium or the angle it forms with
the neck. But by widening the lamina, lengthening and thinning the handle, and
placing it at nearly a right angle to the neck, the weasel’s manubrium may be brought
to the type prevailing in the other Carnivora already described. The thick curved
processus muscularis reminds the observer of Canis, and completely separates it from
the Procyonide and Urside. In some foxes there is a marked tendency to shortening of
the manubrium, and to a wide angle between that process and the neck; but it curves
more, as a whole, with the convexity backwards, and less at the tip; then, on the other
hand, the weasel’s manubrium is never so broad at its outer surface as in these vulpine
Canidee.

It may seem at first remarkable that the malleus in the Mustelide is unlike that of
the Bears, except perhaps in the head and articular surface; but after all, there are
considerable differences in the tympanic bullae beneath them. (See Flower, P. Z. S.
1869, p. 11, figs. 2 & 4.)

The incus (Pl. LVIII. fig. 55) is unusually distinctive in the weasels, having a very
short, slightly curved processus brevis, as pointed out by Hyrtl, at least in Mustela and
Lutra. The tendency to shortness of this process is as marked in most bears as in many
weasels, and nearly as great in the Mluride and Procyonide, except Nasua. The pro-
cessus longus stretches well forwards, and bears a rather large pedunculated Sylvian
apophysis.

The stapes (Pl. LVIII. fig. 62) has straight crura and a stout base; the former are
inserted each close to the extremity of the latter. The base is very bullate towards the
vestibule in small weasels, as it is in Hyrax, Geomys, and Phalangista cookie.

In the Pine Marten (Wartes abietum, Pl. LIX. fig. 7) all the above characters are well
marked in all the ossicles. The outer surface of the manubrium is broad at the base and
tip, which is well recurved and spatulate. The processus longus of the incus is shorter than
in other weasels; the stapes is larger, with a well-formed bulla. In Mustela vulgaris,
M. putorius, and WM. erminea the neck of the malleus is generally rather shorter than in
Martes and Gulo. The base of the stapes of the Mink (Mustela vison) is very bullate.
In Jetonyx zorilla the malleus has a very stout deflected muscular process; the base of
the stapes is slightly inflated (Pl. LIX. fig. 8). In the Glutton (Guwlo luscus) the large
malleus has a very elongated head and neck, a wide articular surface, and a short diver-

OF THE MAMMALIAN OSSICULA AUDITUS. 397

gent manubrium. The incus has a very short processus brevis, but a long, far-stretching
processus longus ; the whole bone is very like a Bear’s. The stapes has thick crura; and
the base is absolutely concave in the horizontal direction.

In the Badger (Meles tazus) (Pl. LVIII. fig. 40) the malleus is remarkable. The
head is as long as in Gwlo; the articular surface is even wider; and the facets are almost
flat, whilst in the above-described genera of weasels they are both quite convex*. The
most distinctive feature is a conical projection, formed by the neck making a sharp
angular bend, instead of a curve, close above the manubrium. It appears at first sight
to be the processus brevis, and thus makes the handle look longer than it really is. But
on close examination, the true representative of the processus brevis may be seen, pro-
jecting outwards, a perceptible distance below the conical projection. The incus resem-
bles that of Gulo. A very slight representative of the above-described projection exists
on the neck of the malleus of the American Badger (Taxidea americana). As regards MW.
taxus, this process (Pl. LVIIT. fig. 40) is equally prominent and well developed in five pairs
of mallei from skulls in the College collection. Its evident relation to the orbicular promi-
nence on the malleus of the Shrew (Sorex) and other mammals will be discussed later on.

The malleus of Helictis nipalensis resembles that of Meles in general aspect; but there
’ is no projection from the neck; the muscular process is not as long as in Mustela.

In the Otter (Zutra) (Pl. LIX. fig. 9) the malleus resembles the Weasel’s; the neck
is much constricted close to the manubrium. The incus has an extremely small pro-
cessus brevis ; the processus longus is stout and divergent. This character is better seen
in the Sea-Otter (Hnhydra), where that crus supports a large flask-shaped Sylvian
apophysis. -Aonyx (Pl. LIX. fig. 18) closely resembles Lutra in its ossicles.

In the Procyonip# the malleus is devoid of good distinctive features, and is of
slender make, with a broad lamina, as in the Civets. The head is not much developed ;
the articular surface is always narrow, and shaped very much like the same in the
Viverride ; the manubrium is generally long and slender, but the outer aspect, though
narrow, is broader than in that family. The muscular process is always present and
always short. The processus gracilis leaves the neck at a very wide angle, so that the
lamina suddenly becomes broad to an extent seldom imitated in the Civets. In a well-
preserved malleus of a member of this family this feature is very prominent; but the
lamina is often mutilated.

The incus has generally an ill-developed processus brevis, never so small as in
Mustela; the crura of the stapes are straight, and, together with the base, are thicker —
than in Cats, Dogs, and Civets. :

In the Raccoon (Procyon lotor, Pl. LIX. fig. 11) these characters are well marked : the
processus muscularis of the malleus is as large as in Viverra ; the end of the manubrium is
distinctly spatulate ; and the whole process is long and slender. The lamina, too, widens
suddenly from the great divergence of the processus gracilis. The processus brevis of the
incus is very much more slender than the stapedial crus, which is very broad, and turns
up sharply at the tip. The base of the stapes is very convex. In the Coati (Naswa nasica,

* Tn this and many other Mustelide the lamina is sometimes perforated, probably from atrophy or ill develop-
ment. Owen describes such a condition in Gulo and Lutra (Anatomy of Vertebrates, vol. iii.).

SECOND SERIES.—ZOOLOGY, VOL. I. 3H

398 MR. A. H. G. DORAN ON THE MORPHOLOGY

Pl. LIX. fig. 12) all these features in the malleus and incus are very feebly marked: the
neck of the malleus is rather short; the processus muscularis is a mere tubercle; the ma-
nubrium is very slender, and the lamina wide but not wider than in a typical Civet’s
ossicula. The processus brevis of the incus is more developed than inthe Raccoon. The
stapes is of stouter make than in Procyon, or in any Felis or Viverra; and its thick base
is very convex. In the Kinkajou(Cercoleptes caudivoloulus, Pl. LIX. fig. 14) the malleus
is much more distinct in type: the head is more developed upwards than in the other Pro-
cyonidee, and resembles that in many bears and weasels; the articular surface is deeply cut
and very narrow ; the neck is long, the manubrium is rather short, broad at the base, and
well recurved at the tips, as in the Mustelide ; but it does not form nearly so wide an angle |
with the neck, and the lamina has the characters of the Raccoon’s. The incus (Pl. LVIII.
fig. 53) has avery small sharp-pointed processus brevis, and a stout processus longus, as in
Procyon. It may be here noticed that, while the tendency to great shortness of the former
process exists in the Mustelidee, Procyonide, Aluridee, and Ursidé alike, in the first two
families the processus longus is stout and short; in _dlurus and the Bears it stretches out
far,asin Man. The base of the stapes in Cercoleptes (Pl. LVIII. fig. 63) is almost flat.
In the Cacomix! (Bassaris astuta) the malleus (Pl. LIX. fig. 15) more resembles that of
Viwerra and Procyon; but its head projects more than in either of those animals, and
the processus gracilis is extremely divergent, so that the procyonine type of a wide
lamina is best marked in Bassaris. This rare animal has a peculiarity already described
and better marked in Meles. The neck of the malleus forms a sharp conical projection
just above the manubrium (Pl. LIX. fig. 15, a, »), which being readily mistaken for the
angular upper extremity of the base of the latter, representing the processus brevis,
makes the whole manubrium look unusually long. In reality the handle is as short as
in Cercoleptes, and nearly as broad at the base; the extremity is distinctly recurved and
only slightly spatulate. The incus (Pl. LVIII. fig. 52) has a very small processus
brevis ; the stapedial crus is stout, and rather longer than in Procyon ; it supports, as in
the rest of the family, a sessile elliptical Sylvian apophysis.
In the Panda (#lurus fulgens) the malleus (Pl. LIX. fig. 16), asa whole, somewhat re-
sembles that of Procyon. 'The head forms a narrow projecting knob above its articular
surface, which is broader than in the last family. The neck is rather short, and almost as
sharply curved as in Viverra. The lamina has the same depth as in the Procyonide,
owing to the divergence of the processus gracilis. The manubrium is rather short, not
very broad at the base, and but little recurved at the tip; it does not form a very broad
angle with the neck : thus it is intermediate in characters between those of the Weasels
and the typical Procyonide, and resembles its representatives in most Bears. The
processus brevis of the incus (Pl. LVIII. fig. 51) is very short and sharp-pointed; the
processus longus is rather thin, and stretches forwards considerably, as in the Ursidze ;
it bears at its extremity a distinctly pedunculated and discoidal Sylvian apophysis.
‘The Ursip# are the only terrestrial Carnivora where a large malleus is to be seen with
an obsolete processus muscularis (Pl. LVIII. fig. 36). The head of that ossicle is well
developed, often projecting considerably upwards; the articular surface is rather wide,
as in the Mustelide; but the groove between the facets is generally faintly marked.

OF THE MAMMALIAN OSSICULA AUDITUS. 399

The neck is somewhat stout and boldly curved. The manubrium is not unlike the
same in Felis, but generally shorter and broader near the base; it turns up at the tip,
which is slightly spatulate; but the outer aspect is rather narrow. It never forms the
bold curve seen in the Canide, Hyzenide, and Protelide. The processus muscularis is
a mere tubercle or almost imperceptible elevation close to the root of the manubrium.
The processus gracilis always diverges considerably from the neck, so that the lamina
is very broad, as in Procyon, whereas the same is narrow in Distela.

The incus (Pl. LVIII. fig. 49) has a very small processus brevis, whilst the stapedial
crus stretches far forwards, and supports a Sylvian apophysis in the form of a disk on a
very narrow peduncle. The stapes (Pl. LVIII. fig. 64) has a well-formed head and
rather slender crura.

In the Polar Bear ( Ursus maritimus, Pl. LVIII. figs. 36, 49, & 64) all these characters
are well marked. The neck forms a sharp curve close to the manubrium, rather ap-
proaching the condition described in Cercoleptes; but it is placed lower, and does not give
an appearance of false length to the manubrium. This process is stout, sharply angled
at the base, and well recurved at the tip. The processus muscularis is represented by
a tabercle nearly as large as in Paradoxurus. The shortness of the processus brevis
of the incus, and the length and divergence of the other crus, are very marked; the
latter bends, forming a prominent knuckle, before contracting to give attachment to the
peduncle of the Sylvian apophysis. There isa rather wide aperture between the crura of
the stapes; the base is broad and plane.

In the Spectacled Bear ( Ursus ornatus, P]. LIX. fig. 17) hardly a trace of the processus
muscularis can be found on the malleus ; the manubrium is rather long and slender, little
recurved at the tip, and not sharply angled at the base. In an incus in the College col-
lection there is a slight tubercular projection close to the extremity of the processus longus,
just above the Sylvian apophysis. The crura of the stapes are rather thick, with a
small aperture between them.

The malleus of the Himalayan Bear (U. tibetanus) is more like that of U. maritimus,
especially in the manubrium, which is not very long, and is broad at the root, sharply
angular at the base and well recurved at the tip; but there is hardly a trace of processus
muscularis. The anterior crus of the stapes is distinctly curved.

In the Sloth Bear (Welursus labiatus) the head of the malleus is less developed than in
an Ursus of the same size. The manubrium is less stout and recurved than in the large
species of the typical genus, but is as angular at the base. There is no trace of any
processus muscularis; the lamina is very wide, and bounded in front by a prolongation
ofthe head, which joins the processus gracilis, a condition seen less plainly in other Car-
nivora, but constant in some other mammals.

THE OssICULA OF THE PINNIPEDIA.

The ossicula of the majority of the Seals and their allies are not only very large but
also-well-marked and readily distinguishable in the different genera or even species.
The ossicles of the Otariidz differ from those of the other families in their absolutely

3H 2

4.00 MR. A. H. G. DORAN ON THE MORPHOLOGY

and proportionally small dimensions. The ear-bones of an adult Otaria jubata are very
much smaller than those of any small Seal, such as Phoca hispida.

The malleus from the skull of a Sea-lion (Otaria stellert) (Pl. LIX. fig. 20) that
measured 0°305 metre from the premaxilla to the front of the foramen magnum, is itself
only 0-008 metre in length from the summit of the head to the tip of the manubrium ;
but this and the other ossicles, though small in these eared-seals, are always of stout
make. ‘The head of the malleus, instead of projecting well above the articular surface,
as in Man, or forming a smooth convexity in front of it, as in most of the Fissipedia, is
hollowed out anterior to that surface, and forms a shallow pit as in all other Pinnipedia.
The articular surface is very wide laterally and rather narrower vertically ; each facet has
so high a vertical convex ridge meeting its fellow that the whole area is quite convex,
forming a prominent angle in a horizontal view, though, viewed perpendicularly, the
planes of the facets do slope towards each other, and form a distinct concavity. The
groove dividing these prominent facets is but faintly marked. The neck is thin,
much constricted towards the manubrium, and never long. The manubrium is longer
than in the true Seals, and shorter than in the Fissipedia; it is broad, and flattened
laterally at the base, narrowing laterally, and turning well up at the point, which is only
slightly spatulate; it does not curve as in the Dog and Hyzna. There is a sharp angle
at its base; and its outer aspect forms a true well-bordered surface, very narrow, except
at the base and at the slight dilatation near the extremity There is no trace of a pro-
cessus muscularis, nor of any lamina, the processus gracilis running close under the
head and neck, to which it is fused by a medium of very compact, not lamellar, bone.
All the characters of the Otarian malleus are thoroughly phocine, putting aside its
small dimensions. Yet Hyrtl says,—“ Bei Otaria ist er (der Hammer) nicht grosser als
bei der Fischotter, und hat den gemeinsamen Typus aller Fleischfresser.’ Had that
distinguished anatomist said ‘“ common type of all aquatic Carnivora,” he would have
been correct; but he means the reverse, as the context shows.

But the incus does not approach any of the singular forms observed in the other
Fissipedia. It assumes very much the form seen in Man, though smaller in proportion
to the malleus, having high compressed sides, a rather short, but stout, blunt-pointed
and only slightly divergent processus brevis, and a long, slender, far-stretching stapedial
erus. ‘This is a modification of the forms seen in the arctoid Carnivora (compare figs. 49
& 50, Pl. LVIII.), where the posterior crus is short and not divergent. The extended or
widely divergent character of the processus longus is, it wili be remembered, very
marked in Ursus maritimus, in Alurus, and to a less extent in Bassaris. The little
stapes has stout crura, often, but not invariably, fused; they are not inserted precisely
into the extremities of the base, which is very thick, and quite concave horizontally.

In Otaria jubata the ossicula are largest : the incus is as large as in Homo, and similar
in form; the posterior part is much higher than the front of the body; the sides are
almost flat. The whole bone is wonderfully light in make compared with the incus in some
true Seals. ;

In Otaria stelleri (Pl. LIX. fig. 20) the concavity on the front of the head of the
malleus is very shallow, the neck and manubrium are moderately long ; the posterior part

OF THE MAMMALIAN OSSICULA AUDITUS. 401

of the body of the incus is not much higher than the anterior, the latter being well’
developed ; the anterior or long crus stretches well forwards, and supports, as in the
other Eared Seals, a Sylvian apophysis in the form of a perfectly circular sessile disk.
The crura of the stapes are quite fused, and the base is very thick; the bone strongly
resembles the same in the Dolphins. In Otaria falklandica (Pl. LIX. figs. 18, 19)
the depression in front of the head of the malleus is deep and well defined, the neck is
very short and constricted, the manubrium shorter than in Steller’s Sea-lion, though,
from the ill-development of the neck, it looks as long. The body of the incus is not
quite so broad as in the last two species, the posterior part is much higher than the
front, the processus longus does not reach so far forwards as in O. stelleri and jubata.
The two crura of the stapes are very thick and straight, with a distinct space between
them, and a base like that in O. stelleri. It is more regular in form than in any true
Seal; but a stapes of Phoca grenlandica in the College collection makes a near approach
to it, excepting that the ossicle in the Greenland Seal is much larger.

In the Walrus (7richechus) (Pl. LIX. figs. 21, 22) the malleus has a large broad head,
of a rhomboidal form, its anterior and posterior aspects being very much flattened, and
very wide in the middle, tapering at their upper and lower extremities. The former
surface bears a deep concavity, as in Otaria and the Phocide; but, unlike either, there is
asmooth rounded surface, of small extent, above the level of the articulation, repre-
senting the globular part of the head seen in other mammals. Posteriorly lies the
articular surface, as broad horizontally and as narrow vertically as in the Seal, but
concave in both directions; the facets are only slightly saddle-shaped, and divided by a
very distinct but shallow groove. The neck is short, not broad near the head, and
extremely constricted close to the manubrium; it is also flattened antero-posteriorly.
It bears no vestige of a muscular process; and the root of the processus gracilis is very
blunt. The manubrium, which forms little more than a right angle with the neck,
is extremely short; it is proportionally broad at the base and at its outer aspect; there
is a distinct everted processus brevis at its upper extremity. The whole bone is not so
large in proportion to the whole skeleton as in the true Seals, but is larger than in any
Otaria. It is plainly a mere modification of the Seal-type of malleus, the articular
surface being the most distinct feature ; the shortness of the malleus is only a slight
exaggeration of the condition seen in Stenorhynchus, where, on the other hand, the neck
is shorter than in Trichechus. It far less resembles the Otarian malleus.

The incus is a large bone, with the posterior part of the body well developed and
bulging externally ; the processus brevis is long, thin, and rather divergent. The pro-
cessus longus is stout, stretches far forwards, and is broadest in the middle, without a
trace of a groove on its broad inner aspect. It forms a gentle sigmoid curve, and bears
on its extremity a thick, sessile, discoidal Sylvian apophysis. This bone, too, is very
seal-like in the thinness of the processus brevis. The long, far-reaching processus
longus allies it to O¢aria; but I have examined one incus of Trichechus where that crus
was rather short; and moreover divergence and length is seen in the same process in
the Cystophorine Seals, but masked by the huge over-developed body of the incus.

The stapes is a high narrow bone, with a small head, thick crura, completely fused,

402 MR. A. H. G. DORAN ON THE MORPHOLOGY

without a trace of an aperture between them in any specimen I have seen, and a narrow,
rather thin hase, concave horizontally. The whole is shaped like a bottle with a short
neck, and very unsymmetrical on account of the outline of the anterior crus being by
far the most convex. This stapes strongly resembles that of MJacrorhinus, while Otaria
closely assumes in its stapes the type of the Dolphin. These two animals approach,
though less nearly, the Sirenian form of that bone.

The Cystophorina, the first subfamily of the Paocipm, are remarkable for the great
difference between the auditory ossicles of two well-known species otherwise closely
allied, and often included in the same genus, viz. the Hooded Seal ( Cystophora cristata)
and the Elephant Seal (Macrorhinus proboscideus). The latter most resembling Tri-
chechus in its ear-bones, it will be described first.

The head of the malleus of Macrorhinus proboscideus (Pl. LIX. figs. 23, 24) is flat-
tened laterally, as in the Walrus; it is still larger in proportion to the rest of the hone,
but is rather triangular than lozenge-shaped, the upper aspect being rounded off and
very broad, not produced to a blunt point. Anteriorly the head is marked by a deeply
concave pit, quite triangular, its basal boundary being uppermost—that is, corresponding
to the broadest or upper part of the whole head. The articular surface is very small,
not extending to the periphery or upper part of the broad posterior aspect of the malleus.
It exhibits the extreme of vertical narrowness and horizontal breadth so general among
the Phocidz. Both facets have a very prominent vertical convex ridge; so that the
whole surface is horizontally quite convex, though laterally slightly but distinctly con-
cave; the groove dividing them is extremely faint. The neck is very short; and the
processus gracilis springs from it as a prominent splinter of bone, connected with the
head by a thin and very brittle layer of compact, not lamellar bone. There is no trace
of a muscular process. The manubrium is very short, though not quite as short as in
Trichechus and Stenorhynchus ; it forms almost a perfect right angle with the neck; the
base is very broad; it is much recurved at the tip, which is spatulate; the outer aspect
is very wide at the base, corresponding to the processus brevis; it widens again near the
termination.

In general configuration the malleus of IZ. proboscideus closely resembles that of
Stenorhynchus ; indeed the only differences I can find are in the wider articular surface
and the still shorter manubrium, forming a wide obtuse angle with the neck in the latter
Seal. In Zrichechus the articular surface is different from that of Macrorhinus, and the
manubrium is still shorter. In Cystophora and Phoca the neck is long, and the manu-
brium forms a very wide angle with the rest of the bone.

The incus is of unique form. It is more than double the bulk of its own malleus, and
exceeds in size its representative in Balena and in most specimens of Manatus. The
oreat feature is the immense development of the posterior part of the body, which leans
over to the outer side of the crura to an extraordinary degree. The anterior part
projects forwards like a lip, and is also inclined somewhat outwards at its uppermost
part. The articular aspect is very small, occupying, to an equal extent, both the pro-
jecting posterior and the more level anterior part of the body; internal to it is a deep
pit, divided by a groove, which runs in a line with the ridge dividing the facets. This

OF THE MAMMALIAN OSSICULA AUDITUS. 403

latter peculiarity, also seen in Stenorhynchus, distinguishes the incus of Macrorhinus
from that of Phoca and Cystophora. The processus brevis is a short process, very stout
at the base and blunt at the tip; it does not diverge from the body; the long crus is
short compared with the dimensions of the whole incus, rather thin, quite cylindrical, and
stretches well forwards from the body, with a wide curve, as in the Bears, the Sea-lions,
and the Walrus. This bone is only a caricature, so to speak, of the same condition seen
in Phoca (compare fig. 23, with fig. 27, Pl. LIX.); but whilst the posterior part of
the incus of Macrorhinus assumes and exaggerates the Phocine type, the long crus is,
as just noticed, rather like the same in the last families, Otariide and Trichechide ;
so that the incus of Macrorhinus is a truly central form.

The stapes strongly resembles that of Trichechus, only itis of much stouter build; the
head is even smaller; the crura widen close below it, and then slowly approximate to the
base; they are entirely fused ; but on the front aspect of the bone is a deep concavity,
representing the normal aperture. The base is a thick bordered facet, and is quite saddle-
shaped, being very convex vertically, though nearly plane horizontally.

In Cystophora cristata (Pl. LIX. fig. 25) the head of the malleus is flattened antero-
posteriorly, and somewhat oval, being broadest horizontally; there is a concavity in
front and a facet behind, convex laterally and concave vertically, as in Phoca and
Wacrorhinus, &c.; but it occupies nearly the whole of the posterior aspect, being far
wider than in the latter, while the whole head is smaller. The neck is extremely long,
more so than in any other Seal, and almost perfectly cylindrical, being but little con-
stricted near the handle; it is, moreover, very straight. The manubrium is short,
rather narrow at the base, well recurved and dilated at the extremity; it forms a very
obtuse angle with the neck, and bears a prominent blunt processus brevis. Altogether
this bone resembles the malleus of Phoca, from which it may at once be known by the
length of its neck.

The posterior part of the body of the incus is well developed, and bulges internally,
but not nearly to the extent seen either in Stenorhynchus or Macrorhinus. There isa
shallow groove round the inner side of its facets, as seen ina more developed form in
Phoca. The processus brevis is short and stumpy; the longer crus is shorter and
stouter than in Macrorhinus; still it stretches well forward, and in that process
alone the incus of this animal resembles that of the last-described species rather than

Phoca.
_ The stapes has an ill-developed head and very thick crnra, both croofed. but the
anterior more so than the posterior. There is a considerable round aperture between
them; the base is broader than in the last family, and slightly concave.

This stapes is truly intermediate between that of the Pinnipedia already noticed and
those to follow. It differs from the former in having an intercrural space and a wider
base, and from the latter in that the space between the crura is very much more limited,
and, the crura being almost parallel, the base is therefore narrower.

Tn the SrenorHyNCHINA (see Pl. LIX. fig. 26, Lobodon carcinophaga*), the head of the

* Syn. Stenorhynchus serridens, Owen, P.Z.8. 1843, p. 131, & Cat. Ost. Mus. Coll. Surg u. p. 641.

404 MR. A. H. G. DORAN ON THE MORPHOLOGY

malleus closely resembles that of Macrorhinus. The excavation in front of its large, flat,
triangular head is deeper, the articular surface is similar but wider, the neck and the
manubrium are even shorter; the latter differs from that of the Elephant-seal (Macro-
rhinus) in being more recurved and spatulate at the tips; its outer aspect is very
broad.

There is uo trace of a processus muscularis, or of the special articular process seen
in Phoca vitulina and hispida; but I find the tendon of the tensor tympani is inserted
in a depression on the exact site of the latter process in a malleus of L. carcinophaga.
This would seem to indicate that the extra articulating process of some Seals is a
modified processus muscularis, though nearer to the head than that process is usually
placed; but, on the other hand, it will be seen that in Phoca barbata, where no trace
of the second articulating process can be seen, there exists a sharp, tubercular processus
muscularis in the exact position of the same in the Felide and Canide.

In the incus the posterior part of the body is more developed than in any Seal ex-
cepting Macrorhinus. Internal to the facets is a deep pit, with a deep groove, which
runs in a line with the groove on the articular surface ; this peculiarity is better marked
in Lobodon and Stenorhynchus* than in the Elephant-seal. The posterior crus is very
short and stout; the processus longus is short, thick, and sharply curved; it is almost
plane, not channelled, on its inner aspect, and the Sylvian apophysis is placed a little
to the posterior side of its extremity.

Hence this bone resembles that of Macrorhinus in its peculiar depression internal to
the articular surface, but Phoca in its processus longus, which, however, is not grooved
as in that animal.

The stapes is rather aberrant. In the Crab-eating Seal (L. carcinophaga) it has long,
very divergent, and much flattened crura, with or without an aperture. 'The base is wide,
thick, and concave horizontally. Inthe Sea Leopard (8. leptonyx) the flattening of the
crura is very marked ; both limbs have rather crooked outlines.

Among the true PHoctna, the ossicula are remarkable for marked differences in closely
allied species. These will be enumerated after describing the ear-bones of the common
Seal (Phoca vitulina) (Pl. LIX. figs. 27, 28, & Pl. LX. fig. 1).

The head of the malleus is flattened, as in the Seals already described; it is rather
small, and nearly circular in outline. Its upper border, the site of the globular part
of the head in most mammals, is quite thin and sharp. The anterior aspect is com-
pletely occupied by a deep concavity (Pl. LIX. fig. 27.¢). The articular surface occupies
more than half the posterior aspect; so that it is more extensive than in Macrorhinus,
but not quite so large as in the other Seals already described. As in most of them, it
is narrow vertically, and very broad horizontally; and from the sharp, perpendicular
ridge on the facets, it is convex in the latter sense, though vertically it is distinctly con-
cave; even the ridges slope downwards to meet each other, and are divided (or, rather,

* Prof. Flower informs me that Prof. Peters (Monatsb. Akad. Berlin, 1875, p. 393), in a footnote, has notified that
the term Stenorhynchus applied by F. Cuvier (1824) to the Small-nailed Seal had been preoccupied by Lamarck (1819)
for a genus of Crabs; hence Prof. Peters proposes to substitute the term Ogmorhinus for the Seal-genus in question.
T haye preferred, however, to use Stenorhynchus as that at present best known to naturalists.

\

OF THE MAMMALIAN OSSICULA AUDITUS. 405

merely indented) by a faint groove. The neck is thin and subcylindrical, rather long,
but never so regularly cylindrical or so prolonged in this species and genus as in Cysto-
phora cristata. Close to and a little below the articular surface there springs from the
neck a little projection in the form of a much-truncated cone, directed inwards and
slightly downwards and forwards; its extremity is quite plane and round, and articulates
with a similar projection from the incus (Pl. LIX. fig. 28, & LX. fig. 1, eas). This
peculiarity, as well as the concavity in the head, is already well known to anatomists.
The processus muscularis is absent. The manubrium closely resembles that of Cysto-
phora, but is slightly longer ; besides, it has a distinct processus brevis, which is curved
upwards towards the neck in well-preserved specimens.

The incus closely resembles that of Cystophora; the articular surface is surrounded
posteriorly and internally by a deep ear-shaped depression; the posterior crus is very
short, and generally thick; the stapedial crus is short, rather thin, boldly curved
inwards, and deeply grooved on its internal aspect, bearing on the outer and interior
aspect of its base a projection as well developed and similar in form to that on the back
of the neck of the malleus, with which it articulates (Pl. LIX. fig. 28). When these
ossicles are articulated, a small foramen exists between the ordinary and the peculiar
phocine articulations (Pl. LX. fig. 1). The extremity of the processus longus, which is
rather thin, bears a Sylvian apophysis, happily compared in its form by Hyrtl to a house-
fly’s proboscis.

The stapes has very stout, rather short and crooked crura; the anterior is most bent;
they diverge considerably towards the base, and have an almost circular aperture between
them ; the base is also thick, and quite concave in its long diameter.

Such are the ear-bones of Phoca vitulina; and no essential variations have been found
in numerous specimens examined by me at the Royal College of Surgeons. In a
specimen of the left ossicula of the Harp-seal (Phoca grenlandica) (Pl. LIX. fig. 30)
the neck of the malleus does not bear a trace of the singular extra articular process ;
but Hyrtl has observed such a projection in this species. There is a faint protuberance
at the back of the neck, close to the manubrium, which may resemble the processus
muscularis seen in P. barbata. The neck is nearly as long (though not nearly so regu-
larly cylindrical), and the manubrium as short, as in Cystophora. The incus (Pl. LX.
fig. 3) wants the extra articular projection, as might be expected ; its crura are otherwise
exactly similar to P. vitulina; the limbs of the stapes, though rather straighter, are quite
as divergent. .

In the Bearded Seal (P. barbata, Pl. LIX. fig. 29) the extra articular process of the
malleus is quite absent; but on the back of the neck, near the root of the manubrium, Is
a distinct processus muscularis (absent in the def¢ malleus of one specimen in the College
collection, though present on the right side). In describing the Stenorhynchina, I dis-
cussed the relationship of the true processus muscularis to the extra articulating process
and the deep depression found on its site in that Seal. The fact that in P. barbata there
is a true processus muscularis much lower down the neck than the articulating process
is placed in P. vitulina, is against the probability that the latter is a modification of the
former; but the insertion of the tendon of the tensor tympani into a deep pit situated in

SECCND SERIES.—ZOOLOGY, VOL. I. 31

406 MR. A. H. G. DORAN ON THE MORPHOLOGY

the malleus of Stenorhynchus and Lobodon, on the exact site of that articulating process,
is in favour of that theory, which cannot be disproved till a malleus is found with both
the processes coexisting. |

The incus of P. barbata (Pl. LX. fig. 2) most resembles that of P. grenlandica, as
both necessarily want the extra articulation; but the groove around the inner side of the
facets is faint. The anterior crus of the stapes is very crooked.

In a complete set of the ear-bones of the Ringed Seal (P. hispida, Pl. LIX. fig. 31)
the malleus and incus closely resemble those of P. vitwlina, having the extra articular
process on both bones; there is no trace of a processus muscularis. The stapes has a
very large space between the crura, which are very divergent; and the anterior is much
bent; the base is very broad; and, in fact, this bone in this little Seal has all the
distinctive features of this subfamily well pronounced.

I have only procured a single incus of the Grey Seal (Halicherus grypus, Pl. LX.
fig. 5); it resembles that of P. vitulina, except that the extra articulating process is
much less developed. .

The foregoing observations on the auditory ossicles of the Carnivora may be con-
veniently abridged in the following manner :—

I. The ossicula of the Fissipedia differ from those of the Pinnipedia very strongly, in
type as well as in consistence.

Il. The laminated form of the malleus prevails throughout the Fissipedia, excepting
in Herpestes and allied genera, where the lamina is nearly obsolete.

III. The Felidz possess a malleus which bears a long, curved, and slender processus
muscularis, and a long manubrium, with a narrow, but distinct and sharply bordered
outer aspect. The processus brevis of the incus is slender and rather long. The crura
of the stapes, as in all these Carnivora, are straight and moderately divergent.

IV. In the Viverride the malleus (except in Herpestes and Swricata, whereof the
peculiarity is mentioned above; moreover the processus muscularis is ox the manu-
brium, away from the root, in these genera) has a very ill-developed processus muscularis,
and the outer surface of the manubrium is even narrower than in the Cats, yet still
distinctly bordered. The incus and stapes are very similar to those of Felis.

V. The extreme narrowness of the outer aspect of the manubrium and the very
small processus muscularis of the tolerably large malleus, of Cryptoprocta ally it more
closely to Viverra than to Felis.

VI. In Hyena the processus muscularis of the malleus is very stout and straight.
The outer surface of the manubr’ wm is very broad near the tip.

VII. In Proteles the processus muscularis is as stout and as straight as in Hyena;
but the outer surface of the manubrium is very broad throughout, as in Canis.

VIII. In the Canidz the processus muscularis of the malleus is as long, curved, and
slender as in the Cats, but the outer surface of the manubrium is very broad throughout.

IX. In the Mustelide the handle of the malleus is much shorter, and forms a wider
angle with the neck than in any of the above Fissipedia. The processus brevis of the
incus is very short, as in the Bears.

OF THE MAMMALIAN OSSICULA AUDITUS. 4.07

X. In the Procyonide the manubrium of the malleus is long, its outer aspect as
narrow as in the Viverride, and the processus muscularis as ill developed as in that
family. The processus brevis of the incus is shorter (except in Masa) than in the already
named groups, excepting the Weasels.

XI. In Bassaris and Aluwrus the malleus has the same characters asin Procyon. The
lamina is very wide, as in most Bears; but the processus muscularis appears never to
be absent. The processus brevis of the incus is very short.

XII. In the Urside the manubrium is long, with a very narrow outer surface; the
processus muscularis is very short, or quite obsolete, the lamina wider than in the Cats
and Dogs. The processus brevis of the incus is very short, and the stapedial process
long and divergent from the body. In the latter ossicle it is allied to the Weasels, Pro-
cyonide, Bassaris, Hlurus, and to the Kared Seals.

XIII. In all the Seals the ossicles are of very dense consistence; and in all but the
Otariide they are very large, absolutely and proportionally.

XIV. In the Otariidz the ossicles are small; the incus is of the ursine type, but the
malleus and stapes decidedly phocine.

XV. In TPrichechus the malieus resembles that of Phoca, excepting that the manu-
brium is extremely short. The incus is very like that of the Phocina, but the stapes
like that of Macrorhinus.

XVI. In the Cystophorina, the ossicula of Cystophora are very similar to those of
Phoca; but in Macrorhinus the malleus and ineus more resemble those of the Steno-
rhynchina, whilst the stapes is of the Walrus type.

XVII. In the Stenorhynchina the malleus has a very large head, and a manubrium
almost as short as in Trichechus; the body of the incus is very bulky. The stapes is of
slender make compared with the same in the Walrus and Macrorhinus. Lobodon and
Stenorhynchus are very similar in their ear-bones.

XVIII. In the Phocina the head of the malleus is excavated, and the articular sur-
face of a distinctive form, as in all the Fissipedia. The neck is not as long as in Cysto-
phora cristata, but longer than in the other Seals; the manubrium is of moderate
length. The incus has often a special process, articulating with a corresponding deve-
lopment from the malleus. The stapes has rather stout crura; an aperture is constant,
being never effaced as in Zrichechus and Macrorhinus. Neither is that ossicle nearly
so bulky as in those genera; but it is a little stouter than in the Stenorhynchina.

Tue OssICULA OF THE RODENTIA.

This order is remarkable for extreme variety in the form of the auditory ossicles in
different families: nearly every type of malleus may be observed among Rodents; a
peculiar kind of ankylosis between that ossicle and the incus is exclusively found in
certain groups of this order; and there is likewise great variety in the form of the
stapes.

To study the peculiarities of the ossicula of the ScruRIDA, a set should be selected from
a skull of one of the fine large foreign. species, such as the Malabar Squirrel (Sciwrus

312

4.08 MR. A. H. G. DORAN ON THE MORPHOLOGY

maximus, Pl. LX. fig. 6). The head of the malleus of that animal is large, rising well
above the articular surface, but much flattened laterally. The articular area is deeply
cut; each facet has a vertical ridge running down the middle, meeting its fellow at the
groove of separation. Of the two segments thus formed in each facet the outer are most
concave, the inner almost flat. There is absolutely no neck; and the manubrium runs
almost in the axis of the body, and springs from close under the lower border of the arti-
cular surface. This process in S. maximus is narrow throughout, but flattened laterally,
and terminates in a well-developed disk. Halfway down its inner border is a slight
projection ; this is the processus muscularis (Pl. LX. fig. 6, pm),—very frequently in that
position among Rodents. The processus gracilis, which I have found entire ina large
adult skull, is very long, thin, and straight; and a small area of friable opaque bone
extends from the head to its root. The incus has a moderately developed body, a short,
pointed, and non-divergent posterior crus; and the processus longus is broad at the root
vertically, long, very straight, and widely divergent; it turns up very sharply at the
extremity, which is tipped by a minute elliptical Sylvian apophysis. The stapes is
large, with extremely slender crura; the anterior is the thinnest ; both are slightly curved
and very divergent. There was no bony canal between the crura in the skull whence
the stapes was extracted for the College collection ; but an intercrural canal was found in
S. vulgaris, S. bicolor, and S. carolinensis. The base is thin and slightly bullate towards
the vestibule; its extremities, especially the anterior, extend considerably beyond the
attachment of the crura.

In the Jelerang Squirrel (S. bicolor) the ossicula are very similar, but the termination
of the manubrium is spatulate rather than discoidal. This process, moreover, is broader
at the root; and the processus muscularis, on the inner border, is very prominent. In
the Common Squirrel (S. cwlgaris) the manubrium mallei is even broader, but the
termination is an almost perfect disk. The processus muscularis is in the same position
as in S. mazimus. In a young British Squirrel’s skull I found the processus gracilis
of the malleus nearly five millimetres in length, running free for two fifths of its length
from the ossicle, beyond which it becomes a little dilated, and is fused by its inner border
to the neighbouring bone; its extremity is a little rounded.

In Tamias striatus (Pl. LX. fig. 9) the malleus resembles closely that of S. vulgaris in
every respect, especially in the manubrium; but at the point where that process leaves
the body of the ossicle there is a blunt angular projection, a rudimentary processus brevis
The body of the incus is very shallow. The stapes has a bony canal between its
crura *.

In the European Souslik (Spermophilus citillus, Pl. LX. fig. 10) the malleus, in general
respects like that ossicle in Sciwrus vulgaris, has a still more distinct trace of a processus
brevis (pb) than has Tamias. The incus is of the Zamias form; and there is a bony
canal between the crura of the stapes.

The malleus of Anomalurus fraseri (Pl. LX. fig. 8) is of about the same size as that of
Sciurus bicolpr; the manubrium is broader and more flattened laterally than in the true

* In the Bristly Ground Squirrel (Xerus setosus) the ossicula are very similar.

OF THE MAMMALIAN OSSICULA AUDITUS. 409

Squirrels, and ends in a very narrow, spatulate, and not discoidal termination. There is
no trace of a processus brevis. The body of the incus is well developed, the anterior
part as much as the posterior. The stapes differs from that of the true Squirrels ; its
crura are short, stout, and almost straight, not very divergent. There is no intercrural
canal in the recent skull. The base is rather thick, quite flat towards the vestibule, and
projects more than a millimetre beyond the insertion of both crura. In the general
characters of its ossicula, Anomalurus is as nearly a true Squirrel as it has recently been
shown to be, in its general anatomy, by Mr. H. R. Alston *.

In the Flying Squirrels (Pteromys, Pl. LX. fig. 7) the malleus, in general resembling
the British Squirrel’s, has a more distinct trace of the processus brevis on the manubrium
even than Spermophilus. The termination of the handle is spatulate. The body of the
incus is as shallow as in Tamias, Spermophilus, and many of the smaller species of Sciwrus.
The stapes is of the same form as in the above-named genera, and unlike that of Anoma-
lurus, having slender, widely divergent crura, occupied in lifetime by a bony canal; the
anterior crus is inserted into the base some distance from its extremity, the posterior
joining the base almost precisely at the opposite end.

The above descriptions show that Sciwrus is the most specialized of the genera already
examined, particularly in the absence of any processus brevis to the malleus, which is
not as broad as in most Rodents. The distinctions from Sciwrus in the other genera
already discussed will be seen to be truly transitional when allied families are
considered.

In the Marmots (Arctomys, Pl. LX. fig. 11) the sides of the malleus are flatter than in
Sciwrus, and the head rises into a conspicuous blunt point high above the articular surface,
which is extremely deep and narrow. All this gives a peculiar aspect to the ossicle.
The manubrium is much broader at the base than in any true Squirrel, though other-
wise of the same type; the extremity is very faintly spatulate, the inner border very
angular. The neck is quite wanting; and there is hardly any trace of a processus
brevis. ‘The incus has a very wide shallow body; the crura are very similar to those
in Sciwrus. The stapes is a proportionally very large bone, with slender, bowed, and
widely divergent crura. There is a dense bony canal between them during lifetime; and
they are inserted almost exactly into the corresponding extremities of the base, which is
very narrow vertically, and slightly convex towards the vestibule. The characters of the
malleus and stapes separate Arctomys from the Squirrels: in the former bone this genus
approaches the Beaver, where, however, there is a neck and a very prominent processus
brevis. The Marmot’s malleus, with its broad, flattened manubrium, is of the type most
prevalent in this order.

In the Dormice, Myoxip# (Pl. LX. fig. 18, Myoaus glis), the head of the malleus is as
compressed laterally and as flattened at the top, above the articular surface, as in Sciurus,
and the articular surface is as deep and narrow vertically. So far it differs from
Arctomys, where the head rises high and the articular surface is wide and shallow. How-
ever, the malleus of Myoxus resembles families yet to be described in having a faint

“On Anomalurus, its Structure and Position,” Proc. Zool. Soc. 1875.

410 MR. A. H. G. DORAN ON THE MORPHOLOGY

trace of a neck and a characteristic manubrium, very thin, very broad at the base, and
still broader at the middle, where an angular projection from the inner border constitutes
the processus muscularis. There is a distinct angle corresponding to the processus brevis ;
and the outer aspect of the manubrium, though very narrow, is distinctly bordered off
from the sides, as in the Carnivora; the extremity is distinctly spatulate. The malleus of
both Myoxus glis and M. muscardinus much resembles the same among certain Muridee,
where the lamina, conspicuous in Jus, is nearly suppressed.

The incus in the Dormice has a longer and more slender processus longus than in
other families of Rodents; the posterior crus is short and very thick.

The stapes has very slender and straight crura, neither of which is inserted into the
extremity of the base, which is quite plane. No bony canal exists between the crura in
the two species above mentioned.

In the Beavers, Castorip# (Pl. LX. fig. 12), the head of the malleus is still large in
proportion to the size of the whole ossicle, as in all the above families; it is also rather
flattened laterally: the articular surface is very wide vertically, as in the Marmots; but
the head does not rise high above it. There is a narrow lamina between the head and
the processus gracilis, and a distinct, much constricted neck. The manubrium is very
broad laterally ; at the outer aspect of the base is a very prominent angle (pd); it is
recurved and spatulate at the tip; but the processus muscularis (pm) is placed close
to its root, so that there is no angular projection on the inner border. Such is the case
in many Rats and Hares; but in the form of the head of the malleus the Beaver more
resembles the Squirrels.

The processus brevis of the incus is very short and pointed, it diverges distinctly, as in
most of the Muride. The body is square and well developed; the processus longus is
rather long, and supports a well-developed Sylvian apophysis. The stapes of the Beavers
is peculiar, it is large, with very slender, widely divergent, and rather short crura, both
inserted some distance from the corresponding extremity of the base, which is very large,
being long horizontally and wide vertically. There is no bony intercrural canal.

In the Murnipa, Rats and Mice, the malleus retains the singular form seen, with modi-
fications, in Arctomys, Castor, &c., in its less typical genera; but in the genus Mus and
allied genera the broad lamina seen in Sorex, the Carnivora, and the Ungulata reappears.
Even in more aberrant genera the lamina is always developed, though to a less degree.

In Mus decumanus (Pl. LX. fig. 14) the head is little developed and deeply excavated
by the articular surface; the neck is long, thin, and much curved, and, together with
the head, is united to the processus gracilis by a broad area of lamellar bone (/); this
lamina is bordered above by a linear prolongation of compact bone from the front
of the head (phm), the processus gracilis bordering it anteriorly and below. From
the outer aspect of the neck projects an orbicular apophysis (oa), a prominent
spherical knob of bone, as seen in the Shrews. It displaces the manubrium considerably
outwards; and, as in Sorea, it is probably an extreme development of the boss sent
outwards from the shoulder of the incurved capitular portion of the first postoral
bar *. The tensor tympani is not inserted into it, but into a minute tubercle close to

* «On the Structure and Development of the Skull of the Pig,” by Prof. W. K, Parker, F.R.8., Phil. Trans.
1874, p. 301,

OF THE MAMMALIAN OSSICULA AUDITUS. ALL

the root of the neck, and just below it. The relations of the processus muscularis to
the orbicular apophysis will be considered when the ossicula of the Shrews are described,
as certain interesting homologues are seen among the Insectivora, throwing much light
upon the subject.

The manubrium is much flattened laterally, very broad at the base; and, as in Castor,
the processus muscularis is at its root, not halfway down its inner margin. The pro-
cessus brevis is an angular projection concealed by the orbicular apophysis, which lies
above and internal to it.

The incus has an ill-developed, slightly divergent, short crus. The processus longus is
stout, and projects far forwards.

The stapes, as in other Muride, has slender, long crura, not very divergent towards
the base, which is thin and narrow; there is no intercrural canal.

us ratius closely resembles the above in its ossicles; in IZ. musculus the orbi-
cular apophysis is very well developed, and was first described in that animal by
Hagenbach.

In the Golden-bellied Beaver-Rat (Hydromys chrysogaster) the malleus, being larger,
shows the murine characters very well (Pl. LX. fig. 16); the spherical knob of bone is
proportionally less developed than in the smaller mice. Below this orbicular apophysis
a tubercular projection can be detected; this is the real processus muscularis; I have
seen the tendon of the tensor tympani still attached to it. The processus brevis of
the incus is very short.

In Hapalotis mitchelii (Pl. LX. fig. 15) the orbicular apophysis is ill iewclepen and
the tubercular processus muscularis, close to the root of the manubrium, is plainly
visible with the aid of a pocket-lens. The manubrium itself is rather shorter than in
Mus. The anterior crus of the incus is far stouter than the posterior. The stapes is of
the same form as in the past genera of the Muride; and, as in them, there is no bony
canal between the crura.

In the Water Voles (Arvicola, Pl. LX. fig. 17) the lamina of the malleus is less deve-
loped, the orbicular apophysis is quite absent, and the manubrium is rather short and
very broad at the base. The processus muscularis is tubercular, and is placed on the
inner side of the neck, close to the manubrium, as in Mus. In A. cmphibius I have
found a bony canal between the crura of the stapes, but not in 4. arvalis.

In the Musquash (Fiber zibethicus, Pl. LX. fig. 18) the head and lamina of the malleus
resemble those of Arvicola; but from the front of the head a very stout process of bone
(ib. p hm) runs to join the upper border of the lamina: as the true processus gracilis is
very slender, the former might, on hurried observation, be mistaken for it. This process
from the head exists in the mallei of most mammals, but rarely to so highly developed
an extent as in Fiber. Its homologue will be seen to form a conspicuous feature in the
Cetacean malleus. The manubrium is extremely broad and flattened, even for a Rodent ;
the processus muscularis is at: the root of the manubrium, asin Mus. The incus has an
ill-developed processus brevis, and a long, divergent processus longus.

The stapes has the same form as in ws and the other Muride, the crura being long,

412 MR. A. H. G. DORAN ON THE MORPHOLOGY

slender, and not very divergent near the base. There is a bony canal between them in
the recent skull. — .

In the Hamster (Cricetus frumentarius) the malleus has nearly as well-developed a
lamina as in Jus; there is no orbicular apophysis; and the manubrium is not so broad
at the base as in the other Muride. The processus brevis of the incus is very ill-
developed, a character more pronounced in Castor and Fiber than in Mus. There isa
bony canal between the crura of the stapes.

Asan example of the Siphneinze, the malleus of Hilobius talpinus is somewhat rat-like
in form, having a small head and a distinct lamina; the manubrium bears a processus
brevis close to its root. It is sometimes partially ankylosed to the incus. The incus
itself has a very ill-developed body with well-developed crura, a feature seen in the
fossorial rodent families distinct from the Muride, and about to be considered. ‘The pro-
cessus longus is much longer than the posterior crus. The stapes has very slender crura,
between which runs a very dense bony canal in the recent skeleton.

Coming to the SpaLaciD#, considerable variety will be found in the ossicula of pro-
minent genera. In Spalax typhius (Pl. LX. fig. 19) the head of the malleus is ill-
developed and laterally flattened, with a very shallow articular surface, hardly any trace
of alamina, and a very constricted neck. The manubrium runs almost in the axis of the
neck; it is short, and is very broad at the base, much laterally compressed; there is a
prominent angle representing the processus brevis. In its malleus Spalax bears the
same relation to Geomys that Castor does to Arctomys, the mallei in Geomys and
Arctomys being quite neckless ; otherwise, in the flattened heads, with wide and shallow
articular surfaces, they all bear a certain relation to one another; and all seem inter-
mediate between Sciurus and Mus.

The incus of Spalax much resembles that of Hllobiws; but the body and articular
surface are still shallower *; the crura are very similar in form and proportion. The
stapes is of an extraordinary and quite unique type. It is very large, measuring from
head to base two millimetres, the base of the skull measuring from the preemaxilla to the
occipital foramen not quite five centimetres, whilst in a skull of Capromys, measuring
over eight centimetres, the stapes is only a millimetre and a quarter in height. The
head is a small narrow rod; the anterior crus is very straight, and runs forward at a great
angle to the posterior, which is much bowed, and inserted into the corresponding
extremity of the base almost vertically under its origin. The base is well developed,
convex, both vertically and horizontally, and very reniform, its lower border being con-
cave. There is no bony intercrural canal in the recent skull.

In the Bay Bamboo Rat (Rhizomys badius, Pl. LX. fig. 20) the head of the malleus is
well developed and not much compressed laterally : the articular surface is much deeper
than in Spalax or Hllobius; and the neck is long and constricted. There is a trace of
a lamina; and the head is prolonged into a point forwards as in Fiber; near the root
of the manubrium is a small tubercular processus muscularis. The manubrium is longer
than in Spalax, and forms a sharper angle with the neck; it is of similar shape, and
very broad laterally, The malleus of Rhizomys bears a strong resemblance to that of

* The posterior part of the body is much deeper than the anterior.

‘OF THE MAMMALIAN OSSICULA AUDITUS. 413

some of the less-typical Muride, especially Fiber. The ossicle is decidedly not so
specialized as that of Spalax. The incus has a very shallow body; the posterior crus is
less developed than in the other genera, though the processus longus is stout and long
—a murine character. The stapes has straight crura, more divergent than in the Muride,
and a very broad base; there is no canal between the crura in lifetime.

In the subfamily Bathyergine the genus Bathyergus exhibits in its ossicles a feature as
distinctly allying it to the Cavies and Porcupines as the past fossorial rodents are allied
by other peculiarities to the Muride, and more distantly to the Sciuride.

The head of the malleus (Pl. LX. fig. 22) is much elongated anteriorly, but not so
much as in the Hystricide. The neck is thin and constricted; and there is a distinct
laminar expansion beneath it. The manubrium is rather short ; it is tolerably wide near
the base, but gets very narrow towards the tip, and has a distinct processus brevis, and a
small stout tubercle on its inner border, representing the processus muscularis, as in many
Squirrels. The head of the malleus is completely fused to the body of the incus by their
articular surfaces, the contour of which is deeply marked all round the seat of ankylosis ;
the bony union involves the neck and a part of the inner border of the manubrium,
from the root to close above the tubercle for the tendon of the tensor tympani, the manu-
brium being fused to the processus longus of the incus nearly down to the point where
it turns inwards towards the head of the stapes. This ankylosis was observed by Hyrtl.
Whilst the malleus differs from the hystricine malleus in the head being less produced
forwards, the incus is distinguished from that of the Porcupines in having the very
shallow body and proportionally large crura seen in the rest of these fossorial animals.
The posterior crus is particularly long. The processus brevis is as well developed as in
Spalax or Ellobius, and hence different from the same in Khizomys.

The stapes (Pl. LX. fig. 22) is a large bone, though not proportionally as great as in
Spalax. Both crura are straight, not very long, and widely divergent. The base is very
broad, and extremely convex towards the vestibule ; the crura are inserted some distance
from both extremities. There does not appear to be any bony canal between the crura ;
at least such does not exist in the skulls of Bathyergus in the College collection.

Among the Saccomy1p®, in the Pouched Gopher (Geomys bursarius, Pl. LX. fig. 21)
the malleus is much flattened laterally, and the articular surface is extremely shallow,
long yertically and narrow in horizontal measurement ; there isno lamina, no trace of a
neck ; and the manubrium, which runs almost in the long axis of the head, is little com-
pressed laterally, and has no processus brevis. It more resembles the malleus of a
Beaver or Marmot than the same in Sciwrus or Mus.

The incus is of singular form; its body is extremely shallow; so is the articular
surface; the processus brevis is very stout and blunt, and disproportionately large,
whilst the anterior crus is very slender throughout, bearing a well-developed pedun-
culated Sylvian apophysis at its extremity. The stapes has rather stout well-diverging
crura; the aperture is occupied, in the recent skull, by a very delicate ossified canal for an
artery. On its base there projects towards the vestibule a very large bulla, better
developed than in Mustela or Hyraz, and rivalling the same condition already described
by Hyrtl as existing in Phalangista cookii.

SECOND SERIES.—ZOOLOGY, VOL, I. BE

414 MR. A. H. GZ. DORAN ON THE MORPHOLOGY

It is evident that Geomys is more nearly allied to the families already described than
to the Cavies, and is not in any way so related to the latter, in the characters of its
otosteals, as is Bathyergus.

In the Jerboas, Drpoprp#, very interesting transitional characters may be observed.

In one adult skull of Dipus egyptius I have found the malleus and incus unanky-
losed (see Pl. LX. fig. 23) on both sides; in other crania of this Jerboa these ossicles were
firmly united. The head of the malleus is extremely small, and deeply excavated by its
articular surface. The neck is slender and constricted; there is a broad and distinet
lamina, along the front of which a portion of the head is prolonged as in Fiber (phm).
The manubrium is very thin and fragile; itis compressed laterally and broad at the base,
which has a curved upper border, ending externally in a true processus brevis (p 6).
The outer aspect of the handle dilates into a very broad, spatulate, slightly recurved
extremity ; its inner border is stout, and has a sharp angle not far from the tip; but the
tensor tympani muscle is not inserted into that projection, being attached to a little
tubercular processus muscularis where that border joins the neck (pm). The incus
of Dipus has a moderately developed body, a stout posterior crus, and a slender very
divergent processus longus. The stapes has long, exceedingly slender crura, not widely
divergent ; the base is narrow horizontally. I have found a canal between the crura in
some skulls of Dipus, but not constantly.

Altogether the malleus of Dipws approaches to that of the Muride, having a small
head and wide lamina, as in ws, and avery broad manubrium, as in Fiber. In its
frequent fusion with the incus it allies itself to the Porcupines and Cavies, where, how-
ever, the head is always well-developed at the expense of the lamina.

The malleus of Pedetes caffer (syn. Helamys capensis) is more Hystricine, since the
head is much developed at the expense of the lamina, and the bone is firmly fused to the
incus. The stapes, too, has well-divergent crura.

Among the Porcupines, Hystricip#, the fusion of the two outer ossicles is extreme,
and the line of separation is barely perceptible externally, in Hystrix; but in none of
the rodents to be described does this bony union extend beyond the normal articular
area as in Bathyergus.

The malleus of Hystrix cristata (Pl. LX. fig. 24) has a very well-developed head,
which projects very prominently forwards, taking the shape of a Phrygian cap, the
convex border being above, the concave below; the blunt apex points forwards and
downwards. In a perfect specimen the processus gracilis (pg) may be seen running close
to the lower border of the head, a very limited area of lamellar bone (/) connecting
them. ‘The manubrium, much flattened laterally and very fragile, is trapezoidal rather
than triangular, on account of the sharp angle on its inner border, which is here a true
processus muscularis, the tendon of the tensor tympani being actually inserted into it.
The broad upper border of the base, well-developed here, as in Fiber or Dipus, forms
a very obtuse angle with the outer border, so that the site of the processus brevis
(pb) is not so prominent as the angular processus muscularis which is readily
mistaken for it in a detached bone. The outer border dilates very gradually into a
narrow, spatulate, slightly recurved extremity. The incus, which is firmly ankylosed

_OF THE MAMMALIAN OSSICULA AUDITUS. : A415

to the malleus in quite young skulls, has a well-developed body and stout crura, the
posterior diverging as much as the processus longus. The crura of the stapes are very
straight, rather short, and very divergent; the posterior is inserted into the base at a
marked distance from its extremity. There does not ever appear to be any bony canal
between the crura, either in Hystrix or Cercolabes. The base is very broad vertically as
well as horizontally, and rather convex towards the vestibule.

In Hystriz javanica the angular projection on the inner border of the manubrium
mallei is even better-marked than in H. cristata. In the Guiana Tree-Porcupine (Cerco-
labes insidiosus, Pl. LX. fig. 25) the manubrium is like that of Hystriz; but the head is
more massive than in the common Porcupine’s, and still more than in the elongated
mallei of Cavia, Chinchilla, &c. The stapes has a rather narrow but very stout base,
convex towards the vestibule; and the crura, as in M/us and Hydrocherus, are not widely
divergent. There is no bony intercrural canal.

Among those Rodents arranged by Mr. Alston (/. c.) under the family Ocropontipa,
in the Ground-Rat (Awlacodus swindernianus, Pl. LX. fig. 27) the head of the malleus is
still more produced forwards than in Hystriz, and there is as little trace of a lamina; but
in the manubrium it differs from the Porcupine’s in having no projection on the inner
border, the processus muscularis being a prominent tubercle close to the root of that border.
The outer aspect ends in a very broad spatulate dilatation. The fusion of the malleus and
incus is complete; the line of junction is very faint. The processus brevis of the incus
does not diverge so much as in Hystrix. The crura of the stapes are slender and very
divergent ; both are inserted into the base a distinct distance from each extremity; and
there is no bony intercrural canal in the recent skeleton. The base is very broad
vertically and horizontally.

In Fournier’s Capromys (Capromys pilorides, P|. LX. fig. 26) the head of the malleus
is as much produced forwards as in Aulacodus ; the manubrium, however, resembles that
of Hystriz, as the processus muscularis (pm) is placed on the middle of the inner border*.
There is the same obtuseness of the angle corresponding to the processus brevis (p 6) as in
all these Hystricomorpha. The fusion of the malleus and incus is so complete that the line
of demarcation is often quite effaced, especially posteriorly. The stapes is very small in
proportion to the other ossicles, as is seen in Dasyprocta; the crura are thick, short, and
not widely divergent ; the base is very broad ; and an intercrural bony canal exists in some,
but not all, of the skeletons of this rodent in the Museum of the Royal College of Surgeons.

Hyrtl states that in the Coypu (Myopotamus) the malleus and incus are not anky-
losed ; no ossicula of this animal have yet been prepared for the College collection.

In a skull of an Octodon cumingii, six weeks old, the author found the malleus and
ineus quite unankylosed (Pl. LX. fig. 28). The former ossicle is rather different from
the same in the above animals. The upper part of its head is ill-developed, and hardly
projects above the articular surface; the latter is narrow and deeply cut; the neck is
very short; the manubrium is long, and not so broad at the base as in the Porcupines
and Rats; it bears a trace of a processus brevis (pd); and the processus muscularis
(pm) is halfway down the inner border of the handle. There is, as usual in this

* The letters pm in the lithograph are placed too near the processus longus of the incus.

3K 2

416 - MR. A. H: G. DORAN, ON THE MORPHOLOGY

section of the Rodentia, a narrow rudimentary lamina (/). The ineus of Octodon has
‘a well-formed body and slender crura. The crura of the stapes are very thin; its base is
wide; there is a dense bony intercrural canal in the recent skull*.

In the CHINcHILLIDA, the malleus of Chinchilla lanigera (Pl. LX. fig. 29) has a head
extremely produced, so as to look like a slightly flexed index finger, a very charac-
teristic feature. The manubrium is very broad, resembling that of Aulacodus rather
than that of Hystrizx, the processus muscularis being on the inner border close to its
root and not further down. ‘This ossicle isvery completely fused by its articular surface
to the ineus, of which the body is well developed; and both crura are stout and short.
The crura of the stapes are very slender; and there is a distinct bony canal between them
in the recent skeleton.

Among the Dasyproctip#, the Spotted Cavy (Celogenys paca) has the malleus and
incus completely fused. I have found, in a mutilated specimen prepared for the
College collection, that the line of demarcation between the ossicula was best-marked
above and internally; Hyrtl describes it as best-marked below and only partially indi-
cated above. The head of the malleus is not so prominent forwards as in Hystria.
There is no canal between the crura of the stapes in the recent skull.

In the typical genus Dasyprocta (Pl. LX. fig. 30) the fused malleus and incus (anky-
losed even in very young skulls) closely resemble the same in Hystrix. 'The head of the
malleus, however, is less produced, and the two otosteals are more flattened than in the
Porcupine. The stapes is small in proportion to the other ossicles ; the crura are short,
straight, and not widely divergent, so that the aperture is small. The base is very broad,
projecting far beyond the insertions of each crus. In one out of three skulls of D. aguti
I have found a bony intercrural canal; none existed in a cranium of D. antillensis.

In the Patagonian Cavy (Dolichotis patachonica, Pl. LX. fig. 31) the malleus and
incus are completely ankylosed. The head of the malleus is stouter and straighter, and
at the same time more produced than in Dasyprocta. The boundaries of the articular
surface are clearly marked on the outer, but very faint on the inner aspect of the fused
bones. The manubrium of this representative of the Caviip# is trapezoidal as in Hystria.
The angle at the outer aspect of the base is very obtuse, but still it represents the
processus brevis (p06); the well-developed and tubercular muscular process (pm) is
seated on a sharp angle on the inner border, which is very thick above and quite sharp
below that process. The outer border broadens regularly from the processus brevis to
the tip, which is slightly everted and spatulate, its edges spreading out widely from the
thin compressed sides of the manubrium. The body of the incus is both broad and deep ;
and its crura are of almost equal thickness and length. The stapes is proportionally
large, the crura very straight, rather long, and divergent; they are deeply grooved
within; and in the recent skull a bony canal runs between them. The base is broad,
extending considerably beyond the insertions of the crura; it is convex towards the
vestibule ; the convexity is separated from the periphery by a deep channel, so as to
form a large umbo. |

* Tn the figure of the malleus of Octodon the letters pm haye been placed where pg should have been written ;
and J is marked too close to the head of the malleus.

OF THE MAMMALIAN OSSICULA AUDITUS. 417

In the Guinea-pig (Cavia caprera, Pl. LX. fig. 33) the malleus and incus, which are
ankylosed in new-born specimens, resemble strongly those of Dolichotis; but the head of
the malleus is more produced and more slender. The manubrium has the same general
characters; only its thin flattened sides and spatulate extremity are much broader than
in the Patagonian Cavy; there is a distinct trace of a processus brevis. The stapes has
all the characters seen in Dolichotis; there is sometimes a bony canal between its
rura.

In Hydrocherus capybara (Pl. LX. fig. 32) the malleus and incus are ankylosed in the
adult, but often free in young skulls. The head of the malleus is stout and less produced
than in Cavia and Dolichotis; and the manubrium mostly resembles the same in the
latter. The crura of the stapes are long, rather crooked, and slightly divergent; the
base is somewhat narrow, stout, and extends beyond the insertions of the crura. No
bony intercrural canal exists, or at least it is not constant.

Among the Glires Duplicidentati, the LEporiD& possess a malleus which does not
assume the extreme types seen in different Rodents, such as the Sciuride and the
Hystricomorpha. It assumes the less-specialized characteristics of the Marmots, Bears,
and some of the Muride.

In the English Hare (Lepus ewropeus, Pallas) the head of the malleus is ill developed,
rising hardly at all above the articular surface; however, it sends a narrow projection
towards the trace of the lamina. The articular surface is rather deeply cut, and rather
narrow laterally; the facets have a vertical convex ridge across their middle as in many
animals in other orders. Between the body and the processus gracilis is a very fragile
and rather narrow bony lamina, generally damaged in the removal of the bone, but plainly
visible if the malleus of a new-born hare or rabbit be prepared ém st¢w with the tympanic
bone. I have examined it in its natural position in the adult hare and rabbit,
and certainly have found the lamina partially absent. That it should atrophy during
- adolescence is highly probable, seeing that the processus gracilis itself is partially absorbed
in Man and in many lower animals. —

The neck of the Hare’s malleus is very distinct, beimg much constricted and. rather
short. The manubrium is of the sword-shaped form seen in the Beaver, being very thin
and flattened laterally, broad at the base, and well recurved. ‘There is a trace of pro-
cessus brevis (Pl. LX. fig. 34, Z. cuniculus, pb); the muscular process (pm) is situated
on the inner border very close to the neck. The outer edge is very narrow, though
distinctly bordered off from the sides and hardly dilated at the tip. This broad manu-
brium, without an angle on its inner margin, may be considered a central type among
Rodents, differing, on the one hand, from the narrower manubrium of the typical
Squirrels, and, on the other, from the trapezoidal handle of the malleus in the Hystri-
comorpha.

The incus has an ill-developed posterior crus and a very divergent stout processus
longus, bearing a well-developed elliptical and pedunculated Sylvian apophysis. Of
course the articular area is deeply cut; otherwise, particularly in the crura, this ossicle
resembles the same in most Rats and Squirrels.

A418 MR. A. H. G. DORAN ON THE MORPHOLOGY

The stapes has perfectly straight and well-diverging crura; the base is broad, and
projects beyond their insertions. There is no bony intercrural canal in the recent skull.

In the Rabbit (Lepus cuniculus, Pl. LX. fig. 34) the ossicles are identical in form with
those of the Hare. In both these animals the processus muscularis may be situated
absolutely on the neck of the malleus, and not at all on the inner edge of the manu-
brium as is usually the case.

The conclusions arrived at, after the study in detail of the ossicula of the Rodentia or
Glires, is as follows :—

I. In the Sciuromorpha there is a tendency in the malleus to suppression of the neck
and processus brevis. This is complete in Sciurus and Anomalurus; but the neck and
the process are gradually developed in Xerus, Tamias, Spermophilus, Pteromys, Myoxus*,
and Castor, the process being perceptible as a slight projection in the first four genera,
whilst it is well developed and there is a distinct neck in the Dormouse and Beaver.
In Arctomys there is no neck, and the processus brevis is as ill developed as in the
Ground-squirrels and Pteromys; the malleus is extremely flattened, and projects above
the articular surface. As in all other rodents where the incus is free f, it has a rather
square solid body, with the posterior crus but little divergent. The stapes is large, with
wide, slender, divergent crura (least so in Anomalurus) ; a bony canal runs between the
crura in lifetime.

The great breadth of the manubrium, so distinctive of Rodents, is well marked in
Castor and Arctomys, least in Sciwrus. The processus muscularis is situated on its
inner border.

II. In the Myomorpha the laminated malleus reappears in its best-developed form in
Mus, Hydromys, and allied species; in some of these Muride there is an orbicular
apophysis, so frequent in the Chiroptera and the Shrews. In the less-typical genera the
lamina is narrow, the manubrium extremely broad, with a prominent processus mus-
cularis on its inner border. The stapes has long slender crura, little curved or diver-
gent. A bony intercrural canal is rare. ,

III. In the fossorial Rodents remarkable varieties are seen in all the ossicula, especially
in the stapes, which is bullate in Geomys and of unique type in Spalax. Hillobius, Spalaz,
Rhizomys, and Geomys} show distinct affinities to the Muride in the form of the
malleus, whilst in Bathyergus we find that the malleus and incus are fused even to a
srveater extent than in the Hystricidee.

TV. In the Hystricomorpha the malleus is almost invariably fused to the incus in the
adults, but often freein young subjects. In Dipus § the malleus is more like those of
some of the Muride than is that ossicle in the rest of the group. The head of the
malleus is produced forwards moderately in Hystria and its allies, also in Cavia,

* Myowus is rather Sciuromorphic than Myomorphic in its ossicula.

+ Excepting Spalax and its allies.

+ The absence of a neck to the malleus in Geomys allies it to the Squirrels.

§ As the malleo-incudal ankylosis is frequent in Dipus, and probably constant in Pedetes, these genera are not
placed here among the Myomorpha.

OF THE MAMMALIAN OSSICULA AUDITUS. 419

Dasyprocta, and Hydrocherus, but to an extreme degree in Awulacodus, Capromys, and
especially in Chinchilla. The stapes is small, with well-developed crura, not very diver-
gent. A bony canal runs between the crura in some specimens from several species, but
is not constant. t

V. In the Duplicidentati the characters of the ossicles are rather central. The malleus
is of much the same form as in the less-typical Muride; but the head is less developed.
There is never any fusion of the malleus and incus in Lepus.

THE OssICULA OF THE UNGULATA.

The different families of Perissodactyla offer more sharply defined characters in their
auditory ossicles than do the even-toed Ungulates.

Suborder PERISSODACTYLA.

Among the RurNocerorips, the ossicula are all very small in proportion to the
size of the animal. In Rhinoceros bicornis (Pl. LXI. fig. 1) the malleus, from the top
of the head to the top of its long manubrium, measures one centimetre, the incus is
two millimetres high between the crura, and its body is barely four millimetres broad,
all much inferior measurements to the same in a Horse, and hardly above the pro-
portions in a Tapir.

The malleus has an extremely deep and wide articular surface, reducing the head to a
mere cup-shaped process, with none of the free globular portion seen more or less in
most mammals. The upper facet (very prominent from the ill-development of the head)
is not as large as the lower; both are slightly saddle-shaped, and run into each other at
the bottom of the deep excavation without any sharp dividing ridge. The neck is cylin-
drical, curved, and rather narrow, bordered anteriorly by the lamina. The manubrium
is very long and slender: it is bent somewhat inwards as it leaves the neck, curving
boldly outwards near the extremity, which is distinctly spatulate. It forms nearly a
right angle near the neck; a well-rounded tuberosity occupies the site of the processus
brevis; there is a small tubercular processus muscularis on the inner aspect of its root.
The sides of the manubrium are flattened and narrow throughout, the inner border is
rounded, not sharp; the outer angle is still bordered, though extremely narrow. In
front of the neck is a laminar expansion (Pl. LXTI. fig. 1, 2), which is continued (rather
than merely blended below, as in Carnivora and Ruminants) into a very large lamelliform
processus gracilis (pg), shaped like a streamer or pennant, and measuring, in a speci-
men in the College, twelve millimetres, the manubrium being only nine millimetres long.
A process from the head (pm) bounds the anterior and upper border of the
lamina.

Comparing this bone with the corresponding ossicle in other Perissodactyla and
Artiodactyla, the presence of a lamina might lead the observer to the assumption that
the malleus of Rhinoceros is more like that of the Pigs and Ruminants than that of the
Equide; the same remark applies to the very similar malleus of Tapirus. Buta very
little consideration must convince him that the lamina in the Rhinoceros is of a different
type from that of the even-toed Ungulates. It is not, as in them, a broad lamella of bone

420 MR. A. H. G. DORAN ON THE MORPHOLOGY

stretched from the root and adjacent part of the processus gracilis to the neck and head,
but a narrow strip of lamellar bone running with that process far in front of the head
into the Glaserian fissure. Indeed the thin sickle-shaped processus gracilis of Equus
is probably a modification of it. The Rhinoceros’s malleus approaches a type very
frequent in the Insectivora, the Bats, the Marsupials, and some Edentata, as Priodon,
where the processus gracilis is elongated in a very similar manner. But more will be
said on the subject when the ossicles of the latter groups are discussed.

The incus of Rhinoceros is of a very peculiar form, the body being very broad and
shallow, and the crura both widely divergent ; the posterior is long, stout, and blunt-
pointed; the processus longus is rather short, boldly curved, deeply grooved on its inner
aspect, and bears on its extremity a greatly developed Sylvian apophysis, pedunculated
with a slightly concave oval surface towards the stapes. It is absolutely and, as I
believe, proportionally the largest Sylvian apophysis existing, greater than the same in
Galeopithecus, to which Hyrtl gives the palm; but he does not describe the ossicula of
the Rhinoceros.

The stapes has a very large head, slightly convex, to fit the apophysis; the posterior
crus is quite straight, the anterior curved forward just before its insertion. In the spe-
cimen in the College collection there is a thin lamina of bone between the upper halves
of the crura. The base is broad and convex.

The ossicula of R. sondaicus and R. sumatrensis are almost identical in character.

Among the Tarrrips, the malleus of Tapirus bairdi (Pl. LXI. fig. 2) resembles con-
siderably that of the Rhinoceros, particularly in the large excavation for the incus, and
the very slight development of the upper part of the head; but the neck is shorter, more
curved, and not so closely defined in front from the lamina; the manubrium is not so
much bent inwards, and is very little curved outwards at the tip, which is less broadly
spatulate. Besides the processus muscularis there is a small tubercle on the inner side of.
the neck, not on the root of the manubrium as in Rhinoceros. The lamina is broad, and
runs into a similar processus gracilis, which is slender and fragile.

The incus has a very broad body; its interior segment is much less developed than the
posterior, by which it differs from Rhinoceros and approaches Hquus.

In the Equip, the typical Equus caballus (Pl. UXT. fig. 3) has a malleus with a well
developed globular head, exceeding, in that respect, other Ungulates, and a wide, deeply
cut articular surface, whose edges are raised above the level of the rest of the head, upon
which it encroaches very little. The facets are almost plane, but slightly saddle-shaped ;
the groove between them is not well marked. The neck is rather long, somewhat
flattened, and twisted a little outwards near the root of the manubrium, which leaves
it at a right angle, is long, but broader at the base than in the Tapir and Rhinoceros,
The manubrium is only slightly spatulate and recurved at the tip, and has a narrow but
well-bordered outer surface. ‘The processus brevis is represented by a blunt tubercle,
much bent forwards; the muscular process is a small tubercle on the inner side of the
neck, close to the root of the manubrium. The processus gracilis is long and sickle-
shaped, not nearly so broad as in Rhinoceros; and there is little trace of a lamina:
hence the equine malleus is a less specialized bone than that of the other odd-toed

OF THE MAMMALIAN OSSICULA AUDITUS. 421

Ungulates, and far more removed from the Artiodactyle form. The incus is very much
larger than that of the Rhinoceros, both absolutely and proportionally. Its body is rather
broad, and cannot be distinguished posteriorly from the stout, short, and little-divergent
processus brevis; the anterior part is very little developed ; the processus longus is more
slender than the processus brevis, though still rather broad at the base; unlike the pos-
terior crus, it diverges considerably from the body. The stapes is quite triangular, has
a well-developed head, with extremely straight crura, not very divergent towards the
base, and inserted a perceptible distance from each extremity. The base itself is broad
and almost plane.

In Hquus asinus, and still more in #. zebra, the processus muscularis of the malleus
is more developed than in #. caballus, appearing as a slightly curved blunt-pointed pro-
cess. The incus in both the former has a slender and more divergent processus brevis ;
in the Zebra the whole body is more developed. The Sylvian apophysis is a flat sessile
disk; in adults it fuses more or less completely with the processus longus, as in Man.
The stapes of #. zebra is very horse-like, the head being very broad and stout, the crura
straight, and the aperture triangular, with sharp angles; the base is thick, with an
umbo towards the vestibule. The stapes in £. asinus has a smaller head; the anterior
crus is distinctly curved ; and the intercrural aperture is round rather than triangular.
A tubercle, with an ossified piece of the stapedius tendon attached, is often seen in
the stapedes of the Equide and also in many Artiodactyles.

SUBFAMILY ARTIODACTYLA.

This group is interesting for the frequent recurrence of the broadly laminate form of
malleus which prevails among the fissipede Carnivora. ‘The incus, as will be shown, is a
bone of great interest and distinctive value in these animals.

In the Hippopotamus (Pl. LXI. figs. 4 & 5) the malleus has a large head, with a well-
developed spherical upper aspect, and a very broad articular surface, of which the upper
facet is of great breadth, almost plane, but has a vertical convex ridge near the outer
border ; while the lower facet is very narrow, and vertically much elongated, so as to be
tongue-shaped and quite convex in both directions. ‘They blend together undivided by
_ any sharp groove; and the entire surface is somewhat shallow. The neck is short, very
broad towards the neck, and constricted close to the manubrium; it is considerably
flattened laterally; and from its inner aspect, a considerable distance from the root of
the handle, springs a long, straight processus muscularis, of equal thickness throughout,
and hence blunt-pointed. The manubrium is long, it leaves the neck at an obtuse angle,
and is flattened laterally, but not very broad at the base. The processus brevis is a
sharp angular projection bent rather backwards ; the outer aspect of the handle is sharply
bordered from the sides, and spatulate at its extremity, which is but a little recurved ;
the whole process is almost straight. The processus gracilis is very slender and united
for some distance to the neck and head by spongy, not lamellar bone.

This bone is very like the same in the pig, the chief distinction being in the cha-
racter of the articular surface; the spongy bone between the neck and processus

SECOND SERIES.—ZOOLOGY, VOL. I. 3L

422 MR. A. H. G. DORAN ON THE MORPHOLOGY

eracilis represents the lamina so marked in the other Artiodactyla; it exists in the
same modified form in the still larger malleus of the Elephant.

The incus has a very high and rather narrow body; the upper part of the articular
surface is very broad, the lower part forms a deep, narrow groove, to fit the lower facet
of the malleus. Both crura are very short and divergent; there is a small tubercle
over the outer aspect of the processus longus, which crus is concave on its inner side.
This high-bodied incus is thoroughly porcine. The stapes has a comparatively small
head, with a flat tubercle on its posterior side for the stapedius tendon. The crura
are both straight, the posterior is much thicker than the anterior, and they diverge very
considerably, leaving a large space between them. The base is thin, broad, and rather
convex; the whole bone forms an equilateral triangle. Hyrtl considers the aperture
between the crura to be small; but it is very wide in three stapedes of Hippopotamus
in the College collection.

In the Wart-Hog (Phacocherus ethiopicus) the malleus resembles that of Sus, about to
bedescribed ; but the following distinctions exist : 1, the upper part of the head is narrower,
and projects more prominently forwards; 2, the articular surface is wider and shallower,
the lower facet is nearly as large as the upper, both are only slightly and evenly
saddle-shaped, and the groove between them is very faint instead of being sharply cut.
This is a distinct approach to the Hippopotamus. The manubrium is almost straight
towards the extremity. The incus is like that of Sus, except that the processus ‘ brevis”
is absolutely longer than the stapedial crus, and is slender and blunt-pointed. ‘The
stapes is pig-like; its crura are extremely slender.

In the Surpm (Pl. LXI. fig. 6) the malleus is of the broadly laminate type. The head
is well developed, though not so large proportionally as in the Hippopotamus, and
projects forwards rather than upwards; the articular surface is narrow and deep. The
groove dividing the facets is very sharply defined ; the upper is larger than the lower; and
both are saddle-shaped, a vertical convex ridge running near their outer borders, leaving
to each a very narrow outer segment,—a very different condition from that which exists
the Carnivora, where the malleus is of a very similar form. The neck is very narrow,
and boldly curved; a stout lamina connects it with the thin processus gracilis. From
the inner aspect of the neck, a slight distance from the root of the handle, springs a long,
stout, and straight processus muscularis, similar to that of the Hippopotamus. The
manubrium leaves the neck almost at a right angle; it is not very broad laterally at the
base, and has a sharp processus brevis pointing vertically upwards. Its outer aspect
has well-defined borders, but is very narrow; the extremity turns up slightly, but dis-
tinctly, and is but little dilated.

The incus has a well-developed exceedingly square body, with short processes, as in
Phachocherus and Hippopotamus, and contrasts strongly with the incus of the Ox and
Deer, and still more with that of the Perissodactyla. Its articular surface is deeply cut,
to fit that of the malleus; the lower facet is narrow and shallow, a natural and necessary
variation from the same in the Hippopotamus, corresponding to a similar difference in
the malleus. The stapedial crus is scarcely longer than the processus brevis, and sup-
ports a small, discoid, pedunculated Sylvian apophysis. ‘The stapes tends to the quadri-

OF THE MAMMALIAN OSSICULA AUDITUS. 4.23

lateral type frequent in the Ruminants, the head being very broad ; the crura, wide apart
at their origin, are slender, and diverge little towards the base; the aperture is large,
and almost circular. The base is stout, with a very convex umbo towards the vestibule,
as Hyrtl has previously observed.

In the genus Sus all the above characters are strongly marked; the lamina, however,
is much encroached upon even ina feetal pig. In Babirussa alfurus the articular surface
of the malleus has but a faint line between the facets, the lower of which is considerably
narrower than the upper. This approaches the type of Phacocherus. The processus
muscularis is very long and stout; the end of the manubrium is slightly but distinctly
recurved.

In the Peccary (Dicotyles tajacu, Pl. LXI. fig. 7) the malleus has a longer and a
thinner neck than in Sws, much less abruptly curved, and a wide, perfectly papery lamina
between it and the processus longus; the manubrium, which forms a more obtuse angle
with the neck than in the other pigs, has a rather wider outer surface, and a distinctly
broader and more spatulate extremity.

The more compact and less laminar expanse of bone between the neck and the pro-
cessus gracilis in the Pig, as compared with the Peccary, might be explained by its
greater size; and, indeed, in the larger animal in any Order there is often a tendency
to such thickening of that area; still, in the large malleus of the Lion this lamina is
as thin, in comparison with the rest of the bone, as the corresponding structure in the
Cat and Genet; on the other hand, it is denser in the Hyzena, which is intermediate
in size between them. Hence the degrees of density of the lamina constitute fair
generic distinctions.

The incus of Dicotyles is even higher and narrower than in Sus; the processus longus
is rather longer. The stapes has a broad head, very straight crura, and a very thick
base, with an umbo convex towards the vestibule, as in Sus.

In the Chevrotains, Tracunip# (Pl. LXI. fig. 8), the head of the malleus is small;
and the articular surface is deeply cut, with a sharp groove between the facets, which are
both saddle-shaped, a sharp convex ridge running vertically down their surfaces, near the
outer margin, as in the Pigs, whilst in most of the Bovidee the facets are nearly plane.
The lower facet is almost as wide asthe upper. The neck is shorter than in Sws, and con-
sequently much shorter than in most Ruminants; but the malleus resembles that of the
latter group of Artiodactyla in the bluntness of the angle at the outer side of the base of
the manubrium, which, moreover, forms a very obtuse angle with the neck; the processus
muscularis is short, but stout. The incus invariably retains the solid square form seen
in the Pigs; and the crura are as short. The stapes has a small head and more divergent
crura than in Sus; the base is very thick and quite plane; the crura are both inserted a
perceptible distance from its extremities. Hence this bone, with its divergent limbs,
has the form of an almost equilateral triangle, as seen in some Antelopes, particularly in
Gazella, differmg from the almost quadrilateral stapedes of both Bos and Sus. The
corresponding ossicle in the Cervide, having a narrow base, with straight, equal crura,
is intermediate in form.

In Tragulus javanicus the manubrium does not form quite so obtuse an angle with

oL 2

ADA MR. A. H. G. DORAN ON THE MORPHOLOGY

the neck as in Tragulus napw*. On the inner side of the neck of the malleus of 7. hanchil
is a small tubercle-like processus muscularis. The Sylvian apophysis on the end of the
long process of the incus is a thick disk, wider in diameter than the end of the processus
longus, and pedunculated.

In the CamreLipm or Tyxopopa (Pl. LXI. figs. 10 & 11, Camelus dromedarius) the
upper part of the head of the malleus above the articular surface is not much developed ;
and, as in the Pigs, it projects forwards, but does not rise above the level of the articular
area in an upward direction. This surface is very wide vertically ; its edges are much
raised above the level of the rest of the head. The facets are almost plane; the lower is
as long as, but narrower than, the upper; they run into each other without any trace of a
groove. The neck is very short, rather broad towards the head, and very constricted
near the manubrium ; there is only a very small fragile lamina stretched between it and
the processus gracilis. The manubrium is extremely long, rather slender even at the
base, and does not form a very obtuse angle with the body; a bold rectangular projection
corresponds to the processus brevis; the outer aspect is narrow, but sharply bordered,
the extremity distinctly spatulate and only a little recurved.

The incus is very singular and distinctive in form (Pl. LXI. fig. 9, Lama peruana).
It is extremely high and narrow; and the anterior part of the body is nearly as deep as
the posterior—a rare condition in any Order}. The articular surface is equally distinctive ;
it is very wide, corresponding to that on the malleus, and absolutely convex from above
downwards; its margins are very prominently raised above the level of the body; its
lower and anterior extremity stops very far short of the root of the processus longus.
Hence the incus in the Camelide is quite distinct from that of other even-toed Ungu-
lates, even from those where it is high and narrow. The stapes has a small narrow
head, long and rather slender crura, not very divergent; and both are inserted some
distance from the extremities of the base, which is very thin, particularly in proportion
to the same in other Ungulates; it bears a base of an umbo.

The malleus of the Camel and Llama are very similar, in the former the head projects
more forwards than in its American representative. In Camelus and Lama pacos there
is a small, tubercular processus muscularis on the inner side of the neck of the malleus,
hardly perceptible in LZ. peruana.

In the Prong-buck, or American Antelope (Antilocapra americana, Pl. LX1I. fig. 12),
the head of the malleus is very little developed, the upper portion projecting neither
upwards nor forwards; the articular surface is very wide and rather deep, its borders
are much raised; both facets are almost plane; the upper is very broad; the lower is
narrowed, and is sometimes prolonged into a short tail-like extremity, running onto
the neck; but this feature is not constant. I have noticed the same peculiarity
in individual mallei of Goats and Antelopes. The groove between the facets is
distinct. The neck is very long, cylindrical, and only gently curved, it is joined to the
slender processus gracilis by a well-marked lamina. From the inner aspect of the neck
projects a very stout, straight, blunt-pointed processus muscularis. The slender manu-

* The processus muscularis is sometimes obsolete in these species.
+ In Camelus the body of the incus is not so high and narrow as in Lama.

OF THE MAMMALIAN OSSICULA AUDITUS. 425

brium has a well-bordered outer aspect of almost equal breadth throughout; there is a
sharp angular projection at its base; and it is but slightly recurved and very little
dilated at the tip.

The incus is high and narrow; its articular surface is deeply cut ; and when there is a
tailed prolongation of the lower facet on the malleus, a corresponding peculiarity is seen
on that facet on this bone. Both crura are very short; the lower is very divergent and
boldly curved. The stapes is almost quadrilateral; the head is very broad, with a pro-
minent process for the stapedius tendon; the crura are stout and almost parallel; the
anterior, which is the thinnest, is inserted into the extremity of the base, which is rather
thin and almost plane.

In general characters the ossicula of Antilocapra closely resemble those of the Bovide.
The body of the incus is almost as stout on square as in the Pigs and Tragulus; but
this feature is seen in Gazella.

THE Bovyip2.

In this family it is hard to find distinctive features in the auditory ossicles of different
species, while, to add to the difficulty, in many instances these bones undergo considerable
_ changes during the growth of the animal; so that little reliance can be placed on any
conclusions based on the observation of one specimen alone from any species. It is
necessary to examine duplicates as well as examples from different species of the
same genus. Through the kindness of Sir Victor Brooke, Bart., the author has been
enabled to collect a large series of the ear-bones of this order from the skulls in that
gentleman’s fine collection of skeletons of Ruminants.

Beginning with the Common Sheep (Ovis aries), it is highly important to select
complete series of the ossicula from individuals of all ages, for reasons immediately to
be explained.

Taking a set from the skull of an adult sheep (PL LXI. fig. 16), the malleus will
be seen to have the upper and anterior part of its head ill developed, as in Antilocapra,
and unlike Sws and its allies. The articular surface (a most important region to observe
closely) is wide vertically as well as laterally, and also deeply cut; each facet is dis-
tinetly saddle-shaped, with a sharp vertical ridge down the middle. The neck is cylin-
drical and sharply curved, as in most mallei of this type; the lamina is of moderate
breadth. The processus muscularis is curved and prominent, but not very long. The
manubrium is long and slender, compressed laterally, but not broad even at the base,
on the outer extremity of which is a rectangular projection, hardly to be called a pro-
eessus brevis. The outer aspect is sharply bordered and extremely narrow, hardly dilating
at the extremity, which is but little recurved.

From repeated observations, I find that the malleus of an adult domestic Ovis closely
resembles that of a foetal Bos tawrus of about the sixth week (Pl. LXI. fig. 15); but at
birth the characteristic, extremely shallow articular surface, and strongly curved manu-
brium, are always found in the ox; the outer aspect of that process, too, is very broad
in foetal calves, though never in adult sheep. ‘The same articular surface is very deeply
cut in the mallei of lambs.

4.26 MR. A. H. G. DORAN ON THE MORPHOLOGY

The incus in an adult sheep has a well-developed body, which is by no means shallow,
but never as square and solid as in the Pigs and their allies. ‘The great development of
the crura in Ovis makes the body look smaller than it really is. The processus “ brevis ”
is very long and straight, and almost cylindrical in form, though a little compressed
vertically. The processus “longus,”’ well divergent from the other crus, is absolutely
shorter; it bends very abruptly above its termination, which bears a very well-formed
pedunculated and perfectly elliptical Sylvian apophysis. In the incus of a new-born lamb
(Pl. LXI. fig. 17) the body is much shallower, as in many less typical Bovide; that part
of the same ossicle in the adult sheep, on the other hand, clearly approaches the high
well-developed type seen in adult oxen.

The stapes in a large adult skeleton is quite trapezoidal; the head is large and square,
bearing a slight trace of a process for the insertion of the stapedius muscle on its inner
aspect. The crura are thick and deeply channelled towards the aperture ; they are wide
apart at their origin and almost parallel; the anterior is the thinner and longer; so that
base is further from the head in front than behind. The base is thick, its upper border
is strongly arched, and both extremities project very perceptibly beyond the insertion of
the crura.

But in the new-born lamb the stapes is generally quite triangular, the head being
much less developed, and the crura close together at their insertion. It will be seen to
retain features observable among many adult Antelopes—that ossicle in the Sheep, when
full-grown, approaching, as does the incus, the type seen in Bos.

The Ox will now be considered ; and it will be demonstrated afterwards how every grade
of transitional features in all the ossicula, between Ovis and Bos compared, are seen in
every kind of combination in Goats and Antelopes.

In a malleus taken from a large Ox’s skull (Pl. LXI. fig. 14) the head appears no more
developed above the facets than in Ovis; but the articular surface, wide horizontally, is
very wide vertically, the facets gaping far apart. The upper is almost plane, though
still slightly convex; the lower is more perceptibly elevated, and is divided from the
upper by a very faint groove. The neck is short and very strongly curved; the lamina
is moderately wide; the processus muscularis projects from the inner side of the neck
some distance from the manubrium, and is rather short, almost straight, and blunt-
pointed. The manubrium is boldly curved, with the concavity forwards; its outer
aspect is rather broad and sharply bordered, dilating but little and very gradually
towards the tip. The base projects upwards, outwards, and a little backwards, and may
be said to bear a true processus brevis. .

The malleus of a foetal calf at the sixth week (Pl. LXI. fig. 15), on the other hand,
is much more like that of an adult Sheep, and is almost as large. The articular surface
is wide and deep vertically, as in Ovis, but not so deep as in the adult Bos; and the
facets are more perceptibly convéx. The manubrium too is almost straight, and slightly
recurved at the tip, instead of being well bowed forward. In the mature foetus the
malleus has assumed the adult Ox’s character.

The incus in the full-grown Bos has a very well-developed body, its outer and inner
aspects both showing a wide, convex, bony surface; the postero-superior border over

OF THE MAMMALIAN OSSICULA AUDITUS. 427

the processus brevis is twice as long as the antero-internal; and the articular surface,
well to the front, is of necessity wide. The processus brevis is as long and cylindrical
as in Ovis, and much curved outwards; the processus “longus” is not so long, but it
stretches out further than in the Sheep, and does not turn up so sharply near the Sylvian
apophysis, which is similar in form. In the fcetus of a calf at the sixth week the body
is about as well developed, proportionally, as in an adult domesticated Sheep.

Taken as a whole, the Ox has the best-developed incus of all the Artiodactyla, inas-
much as the body is in all directions larger than even in Sws, where the crura are charac-
teristically short, whilst those processes in Bos are even longer than in Sheep and
Antelopes (where the body is shallower), and as long as in the Deer (where it is generally
very shallow and broad).

The stapes of the adult Ox is absolutely oblong. The head is very broad, with a pro-
minent tubercle for the stapedius tendon, which latter is partly ossified. The crura are
quite parallel, or even nearer together at the base than the head in very large spe-
cimens; they are well channelled towards the aperture; the anterior is very thin and
straight, and a little longer than the posterior, which is thicker; the base is very stout,
and projects well beyond the crura. The whole bone is only an extreme modification of
the form seen in Ovis; indeed in the feetal calf that bone is still trapezoidal, or almost
triangular, the head being less broad, and hence the crura nearer together at their
origin.

In the adult Zebu (Bos indicus) the ossicula are more slender than in the Ox; the
manubrium of the malleus is not so boldly curved; the stapes is even more oblong than
in B. taurus; its crura are very long and close together.

The ossicula of the Burrhel (Ovis nahoor) are more like those of a new-born lamb than
those of a full-grown domestic sheep. The body of the incus is shallower, though the
crura are as long; nor is the stapes so quadrilateral as in O. aries.

In the adult Goat (Capra) the malleus has an articular surface as wide as in Ovis, and
hence not so wide or so plane as in Bos. The manubrium is distinctly curved, almost as
much as in the Ox in C. bez, less so in C. hircus, C. megaceros, and C. egagrus, judging
from specimens in the College collection. The body of the incus is sometimes shallow in
this genus; but in the skulls of large goats it becomes as high as in the adult Sheep.
The posterior crus is not very long; the anterior is sharply bent, as in Ovis. But even in
large adults the stapes remains triangular, though the head is always thick, and the crura
a distinct distance apart at their origin; they diverge considerably towards the base,
which is very wide and thick, projecting beyond their insertions, and is quite plane
towards the vestibule. The crura are almost of equal thickness; but the anterior is
rather the longer.

In the Chamois (Rupicapra tragus, Pl. LX1. figs. 22-34) the articular surface of the
malleus is wider than in Ovis or Capra, and almost as wide and as level as in Bos; but
the manubrium is slender and almost straight, as in the Antelopes. The incus resembles
the full-grown Sheep’s in the development both of its body and crura; and the stapes is
of the same trapezoidal form.

428 MR. A. H. G. DORAN ON THE MORPHOLOGY

In the malleus of Nemorhedus (Capricornis) swmatrensis (Pl. LX1. fig. 29) the articular
surface is as in Ovis; the manubrium is slightly curved; and the lamina is extremely
wide. The stapes is, on the other hand, positively triangular, as in most Antelopes, the
head being rather small, the crura close together at their origin, and very divergent ; they
are rather thick; the base projects but little beyond them, and is not very wide. Ina
single malleus of Oreotragus saltatrix, from an adult skull in Sir V. Brooke’s collection,
the articular surface is narrower vertically than in Ovis; the manubrium is long, straight,
and very slender. The malleus in Saiga tartarica (Pl. LXI. fig. 23) is very sheep-like
in the nature of its articular area and in the form of its manubrium; the incus (PI.
LXI. fig. 35) has a very shallow body, with widely separating and diverging crura; the
stapes is quite triangular. .

In Gazella (Pl. LXI. fig. 18) the malleus retains most of the characters of Ovis; but
the articular surface is often wider, the processus muscularis is almost always shorter
and smaller, the angle at the base of the handle is so prominent as to form a true pro-
cessus brevis, and the extremity of the manubrium is well recurved and spatulate, the
whole outer aspect of that process being rather wide. The body of the incus is always
well developed in both directions, so as to take from the length of the crura (which, on
actual measurement, are fairly long) and to make the whole bone resemble the same in
the Pigs and Tragulinee, where the processes are very short, especially the processus
brevis. ‘The stapes is a large bone, of very delicate make, and almost an equilateral
triangle in form; the head is small, the crura very slender and divergent, the base very
wide and plane, extending well beyond the insertion of the crura. The ossicula of
G. dorcas, G. levipes, and G. bennettii are very similar.

The malleus of Antilope cervicapra is of the Gazelle type; but the stapes has stouter
and less-divergent crura. The malleus of Procapra picticaudata (Pl. LXI. fig. 27) is
similar; the incus (fig. 38) has as large a body as in any Gazella. The same remarks
apply to Nanotragus pygmaeus and N. montanus (fig. 13), though even in adults of the
latter species the body of the incus remains shallow; at least such is the case in the
College series. The stapes approaches the quadrilateral form of that of Ovis, the head
being larger and the crura more nearly parallel than in Gazella.

In Cephalophus (fig. 28) the articular surface is almost as wide on the malleus as on
that ossicle in Bos, the processus muscularis is very short, and the manubrium is of the
long, straight, and slender form frequent in Antelopes yet to be noticed. The incus
(fig. 39) in C. maawellii is very like a Sheep’s; the stapes is triangular, though the crura
are very divergent. The malleus of Nesotragus moschatus is of the Cephalophus type ;
in the incus from an adult Tetraceros quadricornis (fig. 37) the body is as shallow as ina
lamb; the malleus (fig. 25) and stapes resemble those of Cephalophus ; the same applies
to the malleus of Nemorhedus goral (fig. 30), where, however, the processus muscularis is
longer than in C. mergens or C. maxwellii; the incus has a shallow body and very long
divergent crura. In the adult Kobus ellipsiprymnus (fig. 24) the articular surface of the
malleus is quite as wide as in Bos, but the manubrium is well recurved and not very
slender, altogether more like that of Gazelia than of the other Antelopes. The incus
has a very high body, quite as high as in the Ox; thecrura are rather short. This ossicle

ON THE MAMMALIAN OSSICULA AUDITUS. 429

exceeds the incus of the Pigs in height as much as the incus of some other Antelopes
exceeds that of Sws in breadth.

In the Harnessed Antelope (Tragelaphus scriptus) the malleus has a wide articular
surface and a very long, slender manubrium, which is almost straight. In Damalis luna-
tus, D. pygarga, and Alcelaphus bubalis (Pl. LX1. fig. 19), the articular surface is always
wide, asin Bos, but the manubrium is rather broad at the base and more curved than in
Tragelaphus. In all these genera the incus has a very shallow body, with very large,
thick, and long crura, somewhat as in the Cervide. The only stapes in the College col-
lection (from .4. bubalis) is triangular.

Catoblepas gnw and OC. gorgon are remarkable for the great length and slenderness of the
manubrium mallei, which is slightly turned up at the tip (fig. 26). The head is very
small, the articular surface is wide and smooth, and the processus muscularis is mode-
rately developed. The stapes of C. gorgon is triangular, of C. gnu more oblong from the
crura being more parallel. In both the anterior crus is the thinner.

In the Musk Ox (Ovibos moschatus) the malleus (Pl. LXI. fig. 31) resembles that of
Bos, but the manubrium is straighter, the body of the incus (fig. 40) is not so high, and
the processus longus is as sharply curved as in the Sheep. As the crura of the stapes
diverge a little, that ossicle takes the form of an isosceles triangle with a narrow base.

The left ossicula from an adult Anoa (figs. 32, 41) all resemble those of Bos in the
broad articular surface and well-curved manubrium of the malleus, the high body and
long erura of the incus, and the slender anterior crus of the stapes, which bone, how-
ever, is triangular from the divergence of the crura. The head bears a large tubercle for
the stapedius.

In Camelopardalis giraffa the auditory ossicles (fig. 21) are remarkably like those of
the Ox, and undergo similar changes in the growth of the young animal.

The articular surface of the malleus in an adult and a young male is still wider and
more plane than the Ox; the manubrium is strongly curved. The incus is all but iden-
tical in form with the Ox’s; the body is very well-developed; the stapes is almost rhom-
boidal; the anterior crus is much longer than the posterior, and more slender (fig. 21).

In the ossicula of a new-born or very young Giraffe, the articular surface of the
malleus is deeper, and not so wide vertically nor so plane as in the adult, the manu-
brium is much straighter, the incus has a far shallower body, and the stapes is almost
triangular. All these characters remind the observer of the changes which occur in the
development of Bos.

The ossicula of the Deer are very small and light, a set from an adult Cervus (Pl. LXI.
fig. 20) of large size being all smaller than a series from Gazella dorcas. The foetal
type of Bos prevails more strongly and persistently among adult Deer than even in
Ovis. The head of the malleus in the Red-deer is very little developed : and the articular
surface is deep, very saddle-shaped, and narrow vertically; the manubrium is long and
slender, only slightly bowed and recurved at the tip; the processus muscularis is a short
stout tubercle; the lamina is very broad. The incus has a very shallow, ill-developed
body, with a long, stout, diverging posterior crus, and a short processus “ longus,” sharply

SECOND SERIES.—ZOOLOGY, VOL. I. 3M

430 MR. A. H. G. DORAN ON THE MORPHOLOGY

curved, and bearing a distinct, pedunculated, elliptical Sylvian apophysis. The stapes
is quite triangular, with a small head and slender crura, which are almost equal and not
very divergent. There is generally a trace of a tubercle on the head of the stapes for
- the stapedius tendon.

There is no distinctive feature between any of the ossicula in Cervus elaphus, C. aais,
C. mexicanus, O. pudu, Dama vulgaris or Capreolus caprea. In Cervulus muntjac the
lamina of the malleus is very wide, and, contrary to Hyrtl’s assertion, the stapes (in the
College collection) is by no means thoroughly triangular ; nor can any trace of the pear-
shaped tuberosity described by that author be seen in the specimens examined by me.

In Hydropotes inermis the ossicula are identical in form with those of Cervus; the
same applies to two sets of the ear-bones of Moschus moschiferus (figs. 33, 42) from two
different skulls in the College collection. The shallow-bodied incus, with its long, stout,
divergent processus brevis, is strikingly distinct from the square-bodied, short-limbed
incus of Tragulus (Pl. LXI. fig. 8).

THE OssICULA OF THE PROBOSCIDEA.

The malleus of Elephas (Pl. LX. figs. 38, 39) is the largest seen in any terrestrial
animal, measuring nearly an inch in length from the head to the tip of the manubrium.
The portion of the head above the articular surface is not much developed, either
superiorly or anteriorly ; it is smooth and globular, and widest from behind forwards.
The articular area is shallow and wide vertically; its facets, which are only slightly
convex, run into each other even more insensibly than in Man, not a trace of a groove
dividing them. The neck is of a very peculiar form. It is very broad and flattened
laterally, and bent strongly outwards on itself. The segment above the bend is formed
of rather spongy bone, with a very irregular surface, generally marked with more than
- one nutrient foramen; posteriorly is a sharp ridge of the same sigmoid form as that seen
in Homo; and anteriorly a thin splinter-like processus gracilis is given off without a
lamina. The lower half of the neck is of very compact bone, smooth on the surface;
it gives off the manubrium at a little more than a right angle. The handle is of the
same stoutness as the lower portion of the neck; it is rather short, broad at the base
which bears a prominent everted processus brevis, flattened laterally, and moderately
recurved at the tip, There is a distinct sharply bordered outer surface to the manu-
brium ; but it is narrow, and very little dilated towards the extremity.

This bone is the most distinctive by far of the three ossicula in the Elephant. The wide,
shallow articular surface is seen among the larger Ruminants; but the breadth and short-
ness of the manubrium distinguish it from that of any Ungulate. There is no lamina;
but the anterior part of the less dense upper segment of the neck probably represents it *,
especially as the sigmoid ridge behind that part represents the entire neck of a laminated
malleus. The manubrium recalls a form not rare in the Rodents; but in this great
animal there is no trace of any processus muscularis, the tensor tympani being
attached to a tubercular elevation on the angle formed by the bend of the neck. TI had
the advantage of dissecting the internal ear of the old female Indian Elephant that died

* See remarks on the malleus of Sus, Phacocherus, and Hippopotamus.

OF THE MAMMALIAN OSSICULA AUDITUS. 431

in the Zoological Gardens in the summer of 1875. The tendon of the tensor tympani
was on both sides attached to the elevation referred to above; yet asimilar tubercle may
be seen in other mammals where a processus muscularis also exists. The great size and
peculiar development of this ossicle seems to mask all features which might relate it to
any other Order in particular.

The malleus is as large in Z. africanus as in H. indicus; but in the former it is of
somewhat lighter make, and the neck is not so broad nor so abruptly bent, while the
manubrium is longer.

The incus of the Elephant has a well-developed body, much deeper posteriorly than in
front, where it is excavated for the articular surface. The crura are both short; the
processus brevis diverges but little from the body, and is rather slender for the bulk of
the whole bone; the processus longus is more divergent, and turns up very slightly at
the tip, which bears a small, round, flat and sessile Sylvian apophysis. This bone,
taking even its large development into consideration, cannot be referred to any type
prevailing among the Ungulates. An approach to it is seen in many Rodents, espe-
cially as regards the much greater divergence of the processus longus compared with
the posterior crus.

The stapes in Hlephas indicus (Pl. LX. fig. 36) is quite triangular; the head is
well developed, and bears a tubercle on its posterior aspect for the stapedius tendon; the
posterior crus is quite straight the anterior is slightly curved; the base is broad, and
convex towards the vestibule; it projects considerably in most specimens beyond the
insertion of the posterior crus.

Tn a single specimen of the stapes of E. africanus, in the College collection, the head is
broad, but shallow, and both crura are quite straight; the bone is triangular, but trun-
cated at the apex (fig. 37).

The peculiarities of the Ungulate ossicula may thus be summarized :—

I. In this Order the central* laminated type of malleus is the rule; the processus
muscularis is generally present, but is not often of any length; the incus is rather large,
and variable in form; the stapes has a broad head, and is sometimes quadrilateral.

II. In the Equide the malleus is most generalized; the head is well developed, and
the lamina almost obsolete. Inthe Rhinoceros and Tapir the malleus is of a very low
type, almost recalling that of the higher Marsupials. The stapes in all Perissodactyla is
triangular, with a wide head.

III. In the Tylopoda the malleus possesses characters seen both in the Artiodactyla
and Perissodactyla, the incus is of peculiar form, the stapes is triangular.

IV. Among the Artiodactyla the Suide are distinguished by the production forwards
of the head of the malleus and its deep articular surface, even in the larger species. The
incus has a very square body, with short crura; the stapes is quadrilateral. Hippo-
potamus resembles the Pigs, excepting that the stapes is perhaps more triangular.

V. In Tragulus the incus precisely resembles that of Sus, but the malleus and stapes
are like the same in the smaller Antelopes.

* My reasons for using this term when speaking of the laminated malleus will be found in the description of the
mallei of the Marsupials.

3M 2

432 MR. A. H. G. DORAN ON THE MORPHOLOGY

VI. In the Bovidee the head of the malleus is not produced forwards, as in the Pigs ;
the articular surface is wide and shallow in the adults of the large species. The crura
of the incus are always long; the body is solid, and very high in the Ox, shallower in the
Sheep: the stapes is quadrilateral in the larger members of this family. The feetal
ossicula of the great Bovidze much resemble the ear-bones of the full-grown smaller kinds.
The gradations of type between Ovis and Bos, and the intricate distinctions between
the ossicula of different genera of Antelopes, have already been discussed.

VII. The ossicula of the Giraffe are very similar to the same in the adult Ox.

VIII. In the Cervide the malleus is of the type seen in the Calf, or in the adults of
smaller Bovidee. The processus muscularis is stout, but short in the genus Cervus; the
incus has a very shallow body, with very large crura, even in Moschus, which may thus
' at once be distinguished from Zragulus. The stapes is always triangular.

Tur OssicULA OF THE HYRACES.

In Ayrax (Pl. LX. fig. 35) the upper part of the head of the malleus is a wide,
smooth, spheroidal surface; the articular area is extremely shallow, wide in both direc-
tions; and there is not a trace of any groove dividing the facets. The neck is narrow and
rather long; there is nolamina. The manubrium is well recurved at the tip, which is
slightly spatulate; close to its root a small tubercular processus muscularis is generally
perceptible. The outer aspect of the whole manubrium is sharply bordered and narrow ;
there is a rudimentary processus brevis.

The incus is remarkable for a peculiarity already noted by Hyrtl: the body is ex-
tremely small, whilst the crura are both long and divergent. That anatomist describes
those processes as thin; but in both Hyraa dorsalis and H. capensis they are stout
and well-developed.

The stapes is perfectly triangular, with a small head and very straight crura. The base
is thick and bullate towards the vestibule *.

The ossicula of the Hyraces recalls rather those of the Equide than their repre-
sentatives in any Artiodactyle or Rodent. The malleus is not unlike that of Hgwus,
where also the processes take considerably from the size of the body of the incus; the
triangular form of stapes prevails, too, among the Perissodactyla.

THE OSsiIcULA OF THE INSECTIVORA.

In the different families that compose this Order we meet with great variations in the
form of the ossicula, which are sometimes of almost as low a type as their representatives
in the Marsupials; in other instances they assume characters seen even in the higher
Primates. Tt is in the Insectivora that we so often find a bony canal between the crura
of the stapes, giving support to a small vessel, the “ Steigbiigelarterie” of Hyrtl. This
canal has already been described as existing in the genus Lemur and in some Rodents.
It reaches, however, the highest degree of development in the Order now under con-

* According to Prof. Owen, “ the base of the stapes is rarely ossified beyond the circumference.” In the speci-
mens in the College collection a bulla exists.

OF THE MAMMALIAN OSSICULA AUDITUS. 433

sideration ; Hyrtl devotes a long and interesting chapter (J.c.) to the vessels of the
mammalian tympanum, chiefly dwelling on the stapedial artery and its. branches.
According to that anatomist, this vessel seems, in Hrinaceus and other Insectivora, to
end ultimately in branches distributed to the orbit, whilst the homologue of the internal
carotid is given off before the stapedial artery enters the canal between the crura of the
stapes; but the distribution of these vessels, and the extent to which they are sur-
rounded by bony canals, differ in certain families. Still that is a subject. which does
not pertain to the distinctions of the auditory ossicles in the same animals.

The ossicula of the Shrews (Sorex) are best studied in the crania of some of the large
American species (Pl. LXITI. figs. 1, 2).

The malleus is readily detached from the skull by maceration, together with the
tympanic bone, which is almost identical in its ring-like undeveloped character to its
homologue in the Marsupials. By the aid of a fine needle the ossicle may be separated
from the tympanic without fracturing the processus gracilis.

The head is small, as in most laminated mallei. It bears a rather deeply cut articular
surface, with two facets, of which the uppermost is the larger, and the lower the more
convex. The neck is long and thin, and midway between the head and manubrium it is
bent on itself, forming a sharp angular projection. At the point where it joins the
manubrium is attached a prominent globular process of bone (Pl. LXII. figs. 1, 2, oa),
which projects downwards and backwards in the natural position, and seems, as it were,
to push aside the handle outwards and forwards. The manubrium itself is as thin and
slender as in most Marsupials. A distinct angle, corresponding to the processus brevis,
exists; it may be seen at the base of the manubrium, projecting anteriorly to the
globular process. The extremity of the handle is almost styliform, although, with the
aid of a strong lens, I have detected not only a slight spatulate dilatation, but also an
outer surface to the manubrium, eeincateliae narrow, yet sharply bordered from the very
thin and compressed sides *.

The lamina is as wide as in a typical terrestrial Carnivore, and runs forward with the
processus gracilis without receiving a thin process from the front of the head, such as is
seen to join the processus gracilis in most laminated mallei. That process is flattened,
narrow near the root, but wide in the middle, tapering to a point at the Seer: ; it is
closely applied to the tympanic bone.

A very small tubercular elevation on the inner side 6f the neck, close to where it joins
the manubrium, represents the processus muscularis. To study the relation between
this process and the orbicular projection from the neck, a malleus of a Sorex should be
placed side by side with one from a Gymmnura, where both exist well developed + (PI.
LXII. figs. 2 oa & 14).

The exact morphological signification of the orbicular process is a difficult point to
decide. It exists in many Insectivora andin many of the Muride. It clearly cannot be

* For a fuller explanation of this expression, “ sharply bordered from the sides,” see descriptions of the malleus in
the terrestrial Carnivora.

+ This elevation can only properly be seen by placing a very clean malleus of Sorex on a piece of black paper, and
viewing it under the microscope with a 1-inch object-glass,

434 MR. A. H. G. DORAN ON THE MORPHOLOGY

the processus brevis of Man, as the sharp angle at the base of the manubrium, which
represents it in the Shrew, may be plainly distinguished (with the aid of a lens), and
seen to be pushed outwards by this very orbicular apophysis under discussion. Nor
is it the processus muscularis; for such a process exists in Sorex. In the Bulau
(Gymnura rafflesii), as in the Shrew, a large, curved, and conspicuous processus
muscularis exists, as well as a large, well-formed orbicular apophysis. The tolerably
large malleus of this animal (fig. 14) is very favourable for examination to eluci-
date the question. I consider it highly probable that this orbicular process is the
homologue of the sharp projection seen in the neck of the malleus of the common
Badger, and already described in this paper. It is true that a sharp projection is
also to be found on the neck of the malleus of Sorex; but in that little animal the
projection lies midway between the head and the handle, whereas in Meles the sharp
projection is placed close to the manubrium, in the exact position where the orbicular
process is to be found both in Sores and in all other animals where it exists.

Hence it is not the homologue of any named portion of the malleus in any of the
Mammalia; nor is it represented, except as an orbicular process, in any member of that
class, putting aside the angular projection on the Badger’s malleus*. In all probability
embryology will be able to show that it represents some part better developed in the
homologue of the mammalian malleus among the Sauropsida and Ichthyopsida.

The incus of Sorex has a fairly developed body ; the processus brevis is very short and
non-divergent ; the stapedial crus is rather long, and diverges considerably from the
body; it supports a Sylvian apophysis, very minute (even proportionally), and mounted
ona slender pedicle. Altogether this ossicle is of a type very frequent among the Mar-
supials; yet it will be remembered that extreme shortness of the processus brevis
and marked divergence of the processus longus are seen in Mammals as high as Ursus
and Mustela.

The stapes of Sorex has a minute vessel running between its crura in the recent skull,
but unsupported by any bony canal; so that it drops out and is readily lost in maceration.
The head is ill-developed, broad, and shallow ; the crura are much curved, especially the
posterior ; and the aperture is very wide, as in Homo. The hase is narrow horizontally,
and very narrow vertically. The stapes of Sorex is evidently of a much higher type
than are the two outer ossicles.

The malleus of the Daesman (I/yogale moschata) must hold an important share of the

-anatomist’s attention when he undertakes the study of the orbicular apophysis, so
frequent in this Order (see Pl. LXII. figs. 3, 4).

The upper portion of the head is even less developed than in Sorex, projecting but
little above the upper edge of its articular surface, which is narrow in its horizontal, but
extremely wide in its vertical measurement. The incus in the recent skeleton lies, I
have observed, as much posteriorly as in the higher Primates. Both facets are of about
the same size; hence the lower is proportionally larger than in most animals; they are
nearly plane, and not divided by a sharp groove. The neck is short and curved. Ata
first glance the outline of the lower facet of the articular surface looks like a portion of

* And in the malleus of Bassaris and a few other animals.

OF THE MAMMALIAN OSSICULA AUDITUS. 435

the neck; and a sharp angle is seen where the latter commences, very like the projection
which actually exists in the middle of the neck of the malleus of Sorer. But, by
examining these ossicles in the Shrew and in Wyogale by means of a lens, it will
be seen that in the former the articular surface is not deep vertically, but the neck is
long and sharply angled, whilst in the Daesman the articular surface is long, and the
neck short. The lamina is wider than in Sorex; the processus gracilis is slender and
partially ankylosed to the tympanic bone in the adult. Close to the root of the manu-
brium there springs from the inner side of the neck a round-headed process (Pl. LXII.
fig. 3, ow) mounted on a long pedicel, to which I find that the tendon of the tensor
tympani is attached in the recent skull.

The manubrium is long, and forms a very obtuse angle with the neck; it is much
flattened laterally, and rather broad at the base, so as to be of the sabre-like outline
frequent among Rodents. The outer angle of the base is blunt and slightly bent towards
the neck ; the same aspect of the rest of the manubrium is sharply bordered from the
sides, and distinctly dilated at the extremity.

It is instructive to compare the malleus of Myogale with the corresponding ossicle in
Gymnura, which possesses an undoubted orbicular apophysis homologous to that of
Sorex, and an equally distinct processus muscularis. The latter process, moreover,
actually coexists with the former in many Shrews, or at least is represented by a
tubercle.

The attachment of the tensor tympani tendon to the single process in Myogale makes
that projection a functional analogue of the processus muscularis; but since it projects
backwards, as the orbicular apophysis does in Gymnura, where the muscular process
points inwards and forwards, it seems to me that this solitary process of the Daesman
must be considered as a homologue of the orbicular apophysis of the Shrew and Gymunura,
serving functionally as a muscular process, the true representative of the latter being
suppressed in this animal.

The incus much resembles that of Sorex; but the body is better developed, especially
posteriorly. The processus brevis is very short and non-divergent; the stapedial crus is
long, straight, and ends in a rather large discoidal Sylvian apophysis.

The stapes has a very wide aperture, occupied in the recent skeleton by a large bony
canal. The head is broad and shallow; there is a small tubercle for the insertion of the
stapedius tendon. Both crura are very slender and divergent ; the anterior is consider-
ably the more curved. The base is of necessity wide horizontally ; it is distinctly concave
towards the vestibule, without any bulla or umbo.

The ossicula of If. pyrenaica are very similar to those of the Russian Myogale. In
this animal and in the skulls of the latter and larger species I have always found a very
stout bony canal between the crura of the stapes. Hyrtl (op. cit.) observes :—

“In Sorex and Myogale the vessels run exactly as in Tulpa, without, however, being
enclosed within the tympanum in bony canals.” In describing the vessels of the tym-
panum in the Mole, he had previously remarked that “the bony canal does not fill the
three-cornered aperture of the stapes; on that account the stapes can be moved up and
down on the canal like a ring on the finger.” If this be “exactly” the case in spe-

436 _ MR. A. H. G. DORAN ON THE MORPHOLOGY

cimens of Myogale examined by Hyrtl, or if the stapedial intercrural artery was not, in
his examples, enclosed within the tympanum in a bony canal (whereas in the College
specimens that osseous tube nearly blocks up the whole aperture), it follows that there is
great variety among individual specimens in this respect, as I have actually BOSE eL in
the crania of Squirrels and Cavies.

In the Mole (Zalpa europea, Pl. LXII. fig. 5) the head of the malleus is rather large
for an ossicle of the laminated type; it forms a distinct globular prominence above the
level of the articular surface, also projecting forwards. The area for articulation with
the incus is wide laterally on account of the head being hardly flattened at all in that
direction ; but it is not long vertically; the facets form a sharp angle with one another,
and the uppermost is the larger. The neck is short and curved, the lamina wide
and rather opaque for so small a malleus; the processus gracilis is thin and very fragile.
here is no orbicular process; and the processus muscularis is reduced to a scarcely
perceptible tubercle.

The manubrium is short, and forms an angle of about 120° with the neck. The outer
angle of the base, corresponding to the processus brevis, is considerably bent forwards ;
the whole manubrium is rather broad, laterally, at the base, but narrows rapidly, so as
to be almost styliform at the extremity.

The incus of the Mole has an ill-developed body ; and the processus brevis is almost
obsolete. The processus longus is most characteristic, and has been long familiar to
comparative anatomists. It is deeply channelled out on its front and inner aspect, and
very long. It terminates in a proportionally small, pedunculated Sylvian apophysis.

The stapes is similarin form to that of Myogale; the crura are very slender and well
arched. ‘The intercrural canal is very small, and justifies Hyrtl’s remark quoted in the
above description of the Daesman’s stapes.

The extreme difference between all the ossicles of this animal and those of the Golden
Mole (Chrysochloris) is most instructive and significant; the great distinctions between
the malleus of Zalpa and that of Condylura,; where the incus is very similar, are also
interesting, but less to be expected.

The malleus of Scalops has a large head, as in the last genus; its lamina too is of the
same form. ‘There is a small tubercular processus muscularis close to the root of the
manubrium, which latter fragile process is very readily broken in examining the attach-
ment of the tensor muscle of the tympanum in the recent skull.

‘The incus (Pl. LXII. fig. 6) is more generalized than in either Talpa or Condylura, as
the processus brevis is not so rudimentary ; and though the stapedial crus is long, it is

not so remarkably broad as in those animals. As in them, however, it is deeply
channelled. P

The malleus of Condylura (Pl. LXII. figs. 7, 8) is doubly interesting :—first, because
it is so different from that of Zalpa; secondly, because it possesses, like Myogale, a pro-
cessus muscularis remarkably like the orbicalar apophysis seen in the Shrew.

The head of the malleus in the Star-nosed Mole is very small; nor is the articular
surface deep vertically ; its upper facet is wider than the lower. The neck is very long
and narrow, and is bent into a very sharply projecting angle midway between the head

OF THE MAMMALIAN OSSICULA . AUDITUS. 437

and manubrium ; the lamina is very wide and thin. Here it may be observed that in all
the above characters this ossicle resembles the same in Sorex. It has not either the
well-formed head of alpa or the vertically wide articular surface and short neck of
Myogale. The processus gracilis is broader at the root than in the Shrew; and in form,
being of equal breadth for the greater part of its length, it resembles the same in the
Phalangers and other Marsupials *.

The manubrium is rather short, it is broad at the base, with a very prominent ex-
ternal angle, not produced into a processus brevis. Its outer aspect, though extremely
narrow, is nevertheless separated from the much-flattened sides by distinct borders; the
extremity is very faintly dilated, so as to appear styliform+. The whole manubrium is
separated from the neck by a very deep constriction, so as to be readily broken off.
From this constriction, on the inner aspect, arises a very prominent globular process,
attached by a very narrow peduncle.

It is as difficult to decide whether in this animal the above is the analogue of the true
globular apophysis of the Shrew and Gymnura, or whether it is the processus muscu-
laris, as is the case in the malleus of Myogale. In Condylura the process certainly
points backwards, as in Myogale, like the globular apophysis of Gymnura; but (likewise
as in Myogale) the tendon of the tensor tympani is attached to the root of the process
in the Star-nosed Mole, which may hence be either considered the homologue of the
processus muscularis, or else as the homologue of the globular apophysis made use of, so
to speak, by the tensor-tympani tendon as a point of attachment.

The incus in Condylura is almost identical in character with that of Talpa in the
smallness of its body, the extremely slight development of its processus brevis, and the
large deeply channelled processus longus.

Chrysochloris possesses a malleus of a perfectly unique pattern (Pl. LXIT. figs. 9, 10).
Otto and Rudolphi first appear to have observed the extraordinary development of the
head of the malleus; but the former repeated the error of the latter, who took the
enormously enlarged head for a fourth ossicle. Hyrtl discovered their mistake, and
settled the true nature of the singular pear-shaped prominence from the malleus of the
Cape Golden Mole. The error of the first two anatomists above named must have arisen
from their having attempted to remove the ossicle by pulling at its manubrium with
forceps without enlarging the aperture of the meatus; in doing so the bone is readily
broken. If the malleus be extracted by chipping away some of the bone forming the
prominent convexity in the temporal fossa behind which the enlarged head is lodged,
then there is no fear of its breaking at the neck ; but it cannot be drawn safely through
the meatus. When removed entire, it may be examined separately with little fear of
damage, as it is by no means fragile.

The head of the malleus of Chrysochloris is extremely elaine so as to be over a
quarter of an inch in length. MHyrtl describes it as pyriform, although, being rather
thick and cylindrical at its root, convex behind, and slightly concave in front, it

* This process is united, partly by bone, to the tympanic ring in the adult Condylura.

+ [have never found the extremity of the manubrium to be absolutely styliform in any mammal except in its
modified homologue in the Dolphins.

SECOND SERIES.—ZOOLOGY, VOL. I. 3N

438 MR. A. H. G. DORAN ON THE MORPHOLOGY

may be better compared in shape to a grenadier’s bearskin. It is formed of dense and
brittle bone, and lies in a concavity which bulges into the temporal fossa. There can
be no doubt of its nature as a simple though extreme elongation of the ordinary
caput mallei. On its root posteriorly is seated, very obliquely, the equally modified
articular surface, which is long and narrow, slightly concave, without any indication
of division into facets, and prominently raised above the level of the contiguous parts
of the ossicle.

The neck is long, stout, and cylindrical; and a small depression on it extends towards
the root of the head, and probably represents the site of the lamina, the ridge of bone
below it being the root of the processus gracilis. I have found the ossicle slightly
adherent to the tympanic bone at that ridge, and have reason to believe that part of the
processus gracilis is modified and enlarged, and assists in forming a portion of the pro-
cess chiefly developed from the head. The manubrium (mm) is very short, being hardly
the length of the neck; it bears a very prominent processus brevis, and is scarcely
dilated at the extremity. Hyrtl does not appear to recognize either neck or processus
brevis in the Golden Mole.

T have detected a small tubercular processus muscularis in the neck of the manubrium
of C. trevelyani, where the head does not form so great a prominence in the temporal
fossa.

The incus of Chrysochloris is as modified in form as is its malleus. The body is of the
long and narrow shape of the articular surface of the outer ossicle, and is much con-
stricted before giving off its crura; its long axis intersects a line drawn from the tip of
the long to that of the short process, instead of being parallel to the latter as in other
mammals. Both crura are long, and stout, and rounded; the stapedial crus ends in a
very minute Sylvian apophysis.

The stapes has a small head, and short, very divergent crura; a stout bony canal runs
between them in the recent skeleton. The base is long horizontally, and very broad
vertically in the middle; it is convex towards the vestibule without being bullate. In
many respects this stapes resembles that of Spalaz.

In the Hedgehog (Hrinaceus europeus, Pl. LXII. fig. 13) and its allies the malleus
assumes a very characteristic form. The head of that ossicle is very small, and bears
a perfectly saddle-shaped articular surface, the two convex facets insensibly merging
into each other, without any groove to separate them. The neck is extremely long, and
forms a wide arch without any angular prominence as in Sorex. From its inner side, a
perceptible distance from the root of the manubrium, springs a straight, narrow, and
prominent processus muscularis; and nearer to the handle there projects from the neck a
considerable prominence, not so large as in the Shrew, or even as in Gymnuwra, but
clearly the homologue of the orbicular apophysis in those and in other Insectivora.
The manubrium is slender, and but slightly dilated, and recurved at the tip; the angle
corresponding to the processus brevis is very blunt, and pushed outwards and forwards
by the rudimentary orbicular apophysis.

The lamina is wider than in any other animal; the processus gracilis is reinforced
by a distinct process from the front of the head; and they together form an extremely

OF THE MAMMALIAN OSSICULA AUDITUS. 439

broad process, ankylosed to the tympanic ring in the adult. This process is broadest
towards its extremity, which is well rounded; it is much flattened laterally; and as
it stands at right angles to the plane of the lamina, it presents a very characteristic
appearance. Close to its root, at about the point of junction of the processus gracilis
with the process from the head, it is perforated for the passage of the chorda tympani
nerve.

The malleus of #. dealbatus is very similar to that of the common British species.

The prolongation of its processus gracilis is even wider and longer; but the lamina is not
quite as broad.
_ That the rod-like projection from the inner side of the neck of the Hedgehog’s mal-
leus is the true processus muscularis there can be no doubt, as the tendon of the
tensor tympani muscle will be found attached to it m the recent skull; nor, from the
nature and position of the prominence nearer to the malleus, can it very well be other-
wise than the homologue of the orbicular apophysis of Sorex, which is not utilized in
this animal for the attachment of the muscle, a distinct processus muscularis existing
as well. It seems extraordinary that the malleus of the common Hedgehog, broken in
half, should ever have been taken for two ossicles; yet Hyrtl has corrected that erro-
neous opinion enunciated by a previous observer. Such a mistake could scarcely have
been made if the homologies of the fragments of that or any other ossicle had been care-
fully made out.

The incus of the Hedgehog lies almost vertically above the malleus in the recent
skeleton. Its body is wide and shallow; the processus brevis is very short, but stout,
and pointed. The stapedial crus is long, stout, rounded, and very divergent from the
body; near its root, to the inner side, it bears a small tubercle, and at its extremity a
well-developed Sylvian apophysis on a narrow pedicle. In their general characters the
ineudes of the Hedgehog and of most of the musteline and arctoid Carnivora agree; but —
the remaining ossicles are totally dissimilar.

The stapes of the Hedgehog has a very: small head, with a small tubercle for the
stapedius-tendon on its posterior aspect. Both crura are very slender, grooved towards
the aperture, and much curved, so as to be furthest apart midway between the head and
the base. The posterior is often the straightest. A large vessel passes through the wide
aperture in the recent skull, not enclosed in a bony canai. The base is narrow and
almost plane towards the vestibule.

In the Bulau (Gymnura Rafflesii, Pl. UXII. fig. 14) the head of the malleus is better
developed than in Hrinaceus, but still small; the articular surface is of similar character .
The neck is much shorter and more sharply curved. From its inner aspect, near the
handle, there projects a very conspicuous slightly curved processus muscularis ; and
closer behind the manubrium is a prominent, but sessile, convex process, eviden tly the
homologue of the orbicular process in the Shrew. The presence of both these projections,
equally well developed in this animal, is of great value in considering, by comparison,
the homologues of the single process in Myogale and in other Insectivora. The manu-
brium is much laterally compressed, but not very broad at the root; the site of the pro-

cessus brevis is rounded off, and not prominent. The lamina is much narrower than in
3N 2

440 MR. A. H. G. DORAN ON THE MORPHOLOGY

the Hedgehog ; and the processus gracilis is produced into a much more slender appen-
dage, which bears, however, a foramen for the passage of the chorda tympani.

In the specimen in the College series of ossicula, the incus of Gymnura is extremely
like that of Hrimaceus, but the processus brevis is longer.

The stapes in Gymnura has rather straighter crura and a broader base than in the
Hedgehog. I have not found any bony canal between the crura in the skulls examined
by me.

In the Tenrec (Centetes ecaudatus) the head of the malleus (Pl. LXII. fig. 11) re-
sembles that of the Hedgehog; but the neck is shorter even than in Gymnura, and is
bent on itself into a sharp angle, as in Sorex. It bears a short processus muscularis,
and a very large, convex, but sessile, orbicular apophysis. The manubrium is quite
hedgehog-like, the lamina still narrower than in Raffles’s Gymnura; the processus
gracilis is perforated near the root of its free part beyond the lamina by a foramen
for the chorda tympani nerve. This ossicle in the Tenrec decidedly shows affinities to
Sorex.

But the incus, otherwise like that of Hrinaceus, has a well-developed and divergent
processus brevis. This is not the case in the Shrews and Moles, though it is observed in
Ericulus and, to a certain extent, in Gymnura. In the Macroscelide and Tupaiidee this
process is more developed than in the Hedgehog; but the body is higher than in that
animal and in Centetes.

The stapes is of the Hrinaceus type; the crura are slender and bowed, and not sepa-
rated by any bony canal in the recent skeleton; the base is moderately broad.

In Hriculus nigrescens (Pl. LXII. fig. 12) the malleus is more like that of Centetes
than that of Hrinaceus, the neck being short and sharply curved, and the lamina narrow.

The orbicular apophysis is not well developed; and the processus muscularis is reduced
' to a tubercle. The processus gracilis is perforated by a very distinct foramen for the
chorda tympani.

The incus, as in Centetes, has a body and processus longus similar to that of the
common Hedgehog; but, as in the former animal, the processus brevis is rather long,
thin, and divergent.

The stapes has a very broad base compared with that of Hrinaceus ; the head and crura
are very Similar in character.

The malleus of Solenodon cubanum is much like that of Centetes; it has a prominent
orbicular apophysis and a much more developed processus muscularis than has Hriculus.
The free part of the processus gracilis is unfortunately broken off in the College
specimen.

In the Macroscelide the malleus assumes a characteristic form widely differing from
the types of Hrinaceus and Sores.

In the Elephant Shrew (Wacroscelides alexandri, Pl. UXII. fig. 15) the caput mallei
is ili-developed and much taken up by the articular surface, which is composed of two
very distinct facets separated by a sharp groove. The lower is nearly the size of the
upper and more anterior; both have a high vertical convexity running across them.
The neck is rather short and curved; it is extremely constricted near the manubrium ;

OF THE MAMMALIAN OSSICULA AUDITUS. 441

the constriction appears all the greater on account of the narrowness of the lamina.
Close to the narrowest part, on its inner aspect, the neck bears a distinct, curved pro-
cessus muscularis. I can detect no trace of any orbicular apophysis.

The manubrium is long and rather narrow at the base, which bears externally a trace
of a processus brevis. Its outer aspect is not only sharply bordered from the sides, but
remarkably and almost uniformly broad; the extremity is but slightly dilated and
recurved. ;

The lamina is extremely narrow, owing to the processus gracilis joining rapidly a
process from the head, and thus leaving little room for any area of lamellar bone. The
processus gracilis thus reinforced forms a thin straight bar of bone, which runs half
across the front of the tympanic cavity, to fit into the Glaserian fissure, quite different
from what prevails in the preceding families of the Insectivora, where that process runs
downwards at once, and is curved so as to fit accurately to the tympanic ring.

The incus lies almost vertically above the malleus in the recent skull; it has a
well-developed square body; the part above the processus longus overhangs that pro-
cess. The processus brevis is stout, blunt-pointed, and does not diverge from the body.
The stapedial crus is straight and freely divergent; it supports a very small Sylvian
apophysis.

The stapes has an ill-developed body, and long, slender, extremely straight crura. The
aperture is wide and triangular, though not as wide as in the Shrews and Hedgehogs.
A very stout bony canal fills it during lifetime. The base is rather broad, and convex
towards the vestibule.

The malleus of Rhynchocyon cirnet (Pl. LXII. fig. 16) resembles that of Macroscelides
in general characters, particularly in the narrow neck and singular processus gracilis ;
but the head is better-developed, and the processus muscularis is still stouter and more
curved. The incus is very like that of the Elephant-shrew ; its stapedial crus bears a
fairly developed Sylvian apophysis. The crura of the stapes are very slender.

Itis remarkable that in Petrodromus (fig. 19) (the ossicula of which I have been able
to examine through the kindness of Dr. Giinther) the malleus is much more Shrew-like
than in the Hlephant-shrew. The lamina is wider; and there is a trace of an’ orbicular
apophysis. The processus muscularis and the manubrium are of the same form as in
Macroscelides. As for the incus of Petrodromus, it is not only quite unlike that of Sorea
or Hrinaceus, but in its high and narrow body it diverges also from Muacroscelides and
Ehynchocyon, and tends towards Tupaia. The stapes has straight crura, with a bony
canal between them in the recent skeleton.

In Tupaia the malleus differs from that of any other insectivorous mammal (Pl. LXII.
fig.17). The head of the malleus is broad antero-posteriorly, but flattened laterally. It
supports a very shallow articular surface, saddle-shaped, and only indistinctly divided
into two facets. There is not a trace of any neck or lamina; the tensor-tympani tendon
is inserted into the root of the manubrium. ‘This latter process arises from the head
directly, and has no angle at the outer side of its base corresponding to the processus
brevis. It is not very broad at the base, and is very little dilated at the tip; its outer
aspect is distinctly bordered from the sides.

442 MR. A. H. G. DORAN ON THE MORPHOLOGY

In being neckless and devoid of lamina, the malleus of Twpaia much resembles that
of some of the lower Primates, especially IJidas or Hapale, or certain Lemurs, the
shallow articular facets being a Ceboid feature. It cannot be said to resemble the mallei
of the Sciuridze so closely; for though the latter have a similar head, and are neckless, the
manubrium is of a different type.

The incus has a high and narrow body; the processus brevis is rather long, thin, and
pointed, but hardly diverges from the body; the stapedial crus is scarcely longer than
the short process; it is curved and divergent, and bears a very small Sylvian apophysis.
In general characters this bone is very like the same in many Monkeys and Lemurs. The
author has never seen the long process so divergent as to form (with the body and
short process as the top-piece) a letter T out of the incus, as Hyrtl states of Tupaia
javanica.

The stapes has a very small head and extremely slender, straight crura; the base is
rathernarrow. ‘The aperture is filled by athick bony canal in the recent skeleton, whence
this frail ossicle can with difficulty be removed.

In the Colugo (Galeopithecus volans, P|. LXII. fig. 18) the head of the malleus is much
flattened antero-posteriorly, and is produced a little forwards and upwards above the
articular area. The surface for articulation with the incus is of the type of Homo and
the higher Primates, being narrow vertically and wide laterally ; the outer extremity is
placed much higher than the inner ; and the two facets are faintly indicated, the upper
lying rather internal to the lower; both have a vertical convex ridge internal to the
middle line of the whole surface. The neck is very short and-constricted; and a very
narrow lamina (which seems to me to be absorbed in the adult skull, as the processus
gracilis is in Man) connects it to the slender processus gracilis. It bears a small tuber-
cular processus muscularis on its inner aspect.

The manubrium forms little more than aright angle with the neck, and is long, broad,
flattened laterally, and recurved ; it possesses a distinct trace of a processus brevis. Its
outer aspect is distinctly bordered from the sides, but very narrow, and prominently
spatulate at the recurved extremity.

The affinities of the malleus of Galeopithecus are puzzling. At first, judging from its
articular surface, narrow neck, and large flattened manubrium, with a trace of a pro-
cessus brevis, it seems allied to the highest Primates. But, on the other hand, in the
Manis the articular surface is shaped much as in Homo; and in Galeopithecus the head is
flattened, and the narrow lamina and slender processus gracilis connected with it also
resemble the corresponding parts in the scaly Edentata; and the manubrium still more
nearly resembles that of the Manis than that of Man, having an outer aspect separated
by broad borders from the sides. It approaches the form of the malleus in Lemur more
than that of any other of the Lemuridz ; but its head is not so well developed, nor of the
form seen in any Lemur or Monkey, and its neck is more constricted and distinct than in
any of these animals except the Apes and highest Cercopithecide. It differs widely from
the malleus of the Zarsius, the Nycticebide, and Galagos ; but it is more like that of
Chiromys, where the head is better-developed, though the articular surface, neck, and
manubrium are very similar.

OF THE MAMMALIAN OSSICULA AUDITUS. 443

Tt differs, comparing it with the other Insectivora, very completely from the Shrews’
and Hedgehogs’ in the smallness of the lamina and general form, and as widely from
that of the Tupaiide in possessing a constricted neck and in other respects; but it cer-
tainly has points of resemblance to the malleus of Rhynchocyon and Macroscelides, the
main distinctions being the form of the articular surface and the character of the lamina
and processus gracilis, though the latter feature is not very different in this animal from
that of the Elephant-shrew, except that the processus gracilis is more slender.

The incus is very high and narrow, especially over the short crus; in this respect it
exceeds the Zupaia incus and approaches that of Mycetes and Galago, though it does
not project over the processus longus as in the latter. The processus brevis is very short,
and curved backwards. The stapedial crus is stout, not very long, and bears a well-
developed very elliptical Sylvian apophysis on a narrow pedicle ; it does not diverge from
the body so much as in Tupaia; but the incus altogether resembles that of the latter
animal more than that of any other insectivorous mammal.

The stapes of a young Galeopithecus has an ill-developed head, with slender bowed
erura, and a large aperture not occupied by a bony canal in the recent cranium. The
base is broad and very convex; it projects far beyond both crura. In a pair from an
adult skull in the collection the aperture is filled by a thin lamina.

In most respects both incus and stapes of this animal more resemble those of the
Macroscelidze and Tupaias than of the Shrews, Moles, and Hedgehogs. They are not
strongly allied to the same ossicles in any definite group of the Primates, though strik-
ingly like them in one or two characters above given. Hyrtl considers that the incus of
Galeopithecus bears a larger Sylvian apophysis than in any other animal; but I find it
larger in Myogale, Rhinoceros, Ovibos, and several other mammals.

The above researches on the ossicula of the Insectivora lead to the following con-
clusions :—

I. These bones all offer great variations in the different families of this Order ; and
there is no constant positive character to be foundin any of the three ossicula. The
stapes is never Sauropsidan or columelliform, as in Manis and the Marsupials; but the
very frequent wideness of its intercrural aperture is perhaps not so much a sign of high
type, as an accidental feature in relation to a bony canal passing through it, or, at
least, on account of a large vessel, unsupported by such a canal, running between
the crura *.

Il. Chrysochloris has the most specialized ossicula in this Order. The extraordinary
malleus appears as if rather modified from the highest or anthropoid type than from the
more central laminated form of the ossicle. The almost equally peculiar incus, in the
development of its body and short crus, must be considered as of rather high type than
otherwise ; and the stapes is quite the reverse of Sauropsidan in all its characters.

IIT. The ossicula of Galeopithecus are very generalized in character. The malleus
has several characters seen in the highest Quadrumana and in some Lemurs, most of
which, however, are reproduced in certain mammals as low as the Hdentata. The incus

* More will be said, however, on this topic in the description of the stapes in the insectivorous Bats.

AAA MR. A. H. G. DORAN ON THE MORPHOLOGY

is of a type hardly seen except in certain Cebide and Lemuride; the stapes has no par-
ticularly high affinities. In the malleus Galeopithecus is decidedly allied to the Macro-
scelidee among Insectivora, and to no other family in this Order; but in the incus it more
approaches Tupaia.

IV. In the Tupaiidee the malleus assumes to an extreme degree the neckless and non-
laminated type common in most Cebide and some Lemurs, as well as in Sciwrus; but the
manubrium is rather of the form prevailing in the above-named Primates than of that
seen in the Squirrels; and so in every respect is the incus. All the ossicula differ from
the varied forms in other families of the Insectivora, except that the incus somewhat
approaches in type that of Galeopithecus. Taken as a whole, the ear-bones of Tupaia are
higher in type than in any other family belonging to this Order.

V. The Macroscelide have a more specialized malleus than can be seen in any other
insectivore except Chrysochloris; the extremely constricted neck and the narrow lamina
and processus gracilis running straight forwards to the Glaserian fissure are highly
characteristic. The processus muscularis is well-developed. The head is better-deve-
loped in Rhynchocyon than in Macroscelides. In Petrodromus the malleus is far more
like that of Sorex than in the two former genera; but the incus is not in any sense
Shrew-like.

VI. In the Shrews, the Myogalidze, Talpidee, Centetidee, and Hedgehogs proper one
common feature exists—a malleus with a wide lamina and a processus gracilis united to
the tympanic ring after the fashion of the Marsupials—in short, a low type of malleus.
The processus brevis of the incus is always ill-developed except in the Centetidze and
Gymnura. The intercrural aperture of the stapes is always wide.

VII. In the Soricide the malleus has a moderately wide lamina, an orbicular apo-
physis, and a more or less distinct processus muscularis, to which, and not to the
spherical protuberance, the tendon of the tensor tympani is inserted. Asa whole the
malleus is very much like that of a Marsupial. The stapedial crus of the incus is not
very broad.

VIII. In the Myogalide the malleus differs in several respects from that of the Shrews
proper: the manubrium is of stouter make ; and an orbicular process exists, which pro-
bably is the homologue of that in the malleus of Sorez, but. certainly functionally
replaces the processus muscularis, absent in Myogale. The incus resembles that
of Sorex.

IX. In the Talpide the malleus of Condylura most resembles that of Sorex, but has an
orbicular process serving as a processus muscularis, as in Myogale; its incus resembles
that of Talpa. In Scalops the incus has not the chief distinguishing feature seen in
Talpa and Condylura; but its malleus is of the form seen in the common Mole, where
the head is better-developed than in the Shrews and Hedgehogs, and the lamina not so
broad. The incus of Talpa has a very broad and channelled processus longus.

X. The Erinaceide and Centetidee have a more specialized malleus than the Shrews,
Daesmans, and Moles. The great feature is the very broad processus gracilis perforated
to transmit the chorda tympani nerve; the processus muscularis and an ill-developed
orbicular process coexist ; the lamina is very broad. In Centetes and Hriculus the pro-

OF THE MAMMALIAN OSSICULA AUDITUS. 445

cessus brevis of the incus is well developed; that crus is smalier in Gymnura and
abortive in Hrinaceus.

XI. The orbicular apophysis of the Shrews, Daesmans, Myogale, and Hedgehogs allies
them to the Muridz, where also the malleus is laminated.

THE OssICULA OF THE CHIROPTERA.

The malleus of Péeropus (Pl. LXII. figs. 26, 27) is attached to the ring-like tympanic
bone much in the same manner as among the Insectivora and Marsupials. The anterior
or upper extremity of that bone is fissured, and the processus gracilis fits into the cleft,
beyond which is a flat surface, to which the extremity of the process is ankylosed. The
portion of the tympanic external to the cleft is very thick, and it runs over the outer
surface of the lamina and neck of the malleus, a distinctive feature not observed in the
non-placental mammals or in the Insectivora, the tympanic ring reaching, in those
animals, not further backwards than to the root of the processus gracilis.

The head of the malleus is small, and projects very little beyond the articular surface,
which is placed as much posteriorly as superiorly, and is shallow, but deep vertically. It
is distinctly divided into two facets, although they are not separated by a sharp hori-
zontal groove; the upper is larger, and lies a little more internally than the lower, and
both are faintly convex. The neck is narrow and well curved, as in most laminated
mallei; the lamina is rather narrow, and formed of opaque and brittle, not semitrans-
parent, papery bone: the processus gracilis is broad and thin at its root, the lower
border being, however, well defined, and the lamina is prolonged upon it; this process
ends in a point as in most Insectivora, not in a foliaceous dilatation as in many
Marsupials.

T have found, in dissecting the ear of a fresh Bat (Pteropus), that the tendon of the
tensor tympani is inserted into the point of junction of the neck and manubrium, at the
most usual site, in fact, of the processus muscularis, which does not exist; but there is
a distinct prominence close to the manubrium at the place where a large orbicular
apophysis is seen in the Shrew (Sorex), Mouse (Mus), &c. The relations of that process
to the processus muscularis have been already fully discussed in describing the ossicula
of the Shrew and other Insectivora. Suffice it to say here, that in Péeropus the rudiment
of the orbicular apophysis does not replace functionally the processus muscularis as in
Myogale and Condylura. ‘This rudiment is better marked in the College specimen from
P. hypomelanus than in that from P. edulis in the same collection.

The manubrium is slender and rather long, broad at the base and flattened laterally ;
in fact, sword-shaped. Its outer and upper angle (not to be confounded with the rudi-
mentary orbicular apophysis which pushes it aside) is somewhat blunt, so that no pro-
cessus brevis exists; the outer aspect is very narrow, but bordered sharply from the
sides, and ends in a distinct spatulate dilatation.

It is evident that a verdict of “low type” must be passed on the malleus of
Pteropus. It has no anthropoid, ceboid, or lemuroid characters. It is not the typical
laminated malleus of the Carnivora and the Ruminants. It is decidedly lower than the

SECOND SERIES.—ZOOLOGY, VOL. I. 30 .

446 MR. A. H. G. DORAN ON THE MORPHOLOGY

malleus of the Muride, where the lamina is broad and semitransparent as in the
Carnivora. Its general resemblance to the malleus of the Marsupialia is very evident.

The incus of Pteropus has a well-formed body, not deeply excavated. The processus
brevis is distinct, but very short; the stapedial crus is rather long, and ends in a pedun-
culated almost discoid Sylvian apophysis. In relation to other animals its characters
are negative.

The stapes has a small head with slender crura; the posterior quite straight, the
anterior generally a little curved *. They do not diverge so much as in Man; but there
isa free aperture not occupied in the recent condition by any intercrural canal. The
base is rather narrow horizontally, and almost plane towards the vestibule; it projects
but little beyond the insertion of each crus.

The stapes of Pteropus must be considered of rather central type, for it shows no sign
of any tendency towards the columelliform stapes of most Marsupials; but its crura do
not diverge as in the anthropoid Apes and Man, nor even to the extent seen in Ruminants
and Carnivora. It most resembles the stapes of those Insectivora where the crura are
not widely bowed to admit the passage of a bony canal, or the same ossicle in the Mice.
This variety of stapes, with its straight crura, diverging but little, and its consequently
narrow base and narrow but distinct aperture, lies between the carnivorous type, where
the aperture is larger, but the crura as straight, and the form seen in Didelphys, where
the crura are free throughout, yet leave a very narrow aperture between them.

I have experienced great obstacles in the study of the ossicula of the insectivorous
Bats, as in museum specimens the bones of the auditory region are frequently lost in
macerating the skull; still more generally are the incudes and stapedes wanting, and
the delicate malleus is frequently mutilated. Nevertheless I have succeeded in procuring
specimens of at least the latter ossicle from Bats belonging to all the chief divisions of the
group, nor, judging from duplicates, does there seem to be great variety among them.

In studying the malleus of Asellia tridens, as an example of the RHINOLOPHIDA, a
very strong general resemblance to the same bone in the Shrew (Sorex) may be observed
ataglance. It is absolutely necessary, in studying the ossicula of the smaller Chiroptera,
to place thoroughly clean white specimens upon a piece of black paper; they may then
be viewed by direct illumination under the microscope. A 1-inch object-glass is suffi-
ciently strong for this purpose.

The head of the malleus in Asellia tridens + (Pl. LXII. fig. 20), though small com-
pared with the area of the entire ossicle, as in all laminated mallei, is well formed
and convex. ‘The articular surface is rather deeper cut than in Pferopus; but the facets
are of similar character, the upper and anterior being the larger. The neck is narrow,
longer even than in Sorex, and resembles its homologue in the Shrew in being sharply
bent midway between the head and the manubrium. At the apex of this angle I
find, projecting internally, a curved process (pm), remarkably like a second process
(acpm).

* Hyrtl (op. cit.) states that the crura “in Pieropus edulis and P. vulgaris [sic] are convergent before their insertion

into the base.” This is not the case in the College specimens.
¢ In Mr. Dobson’s ‘ Monograph of the Asiatic Chiroptera,’ Asellia is reduced to a subgenus of Phyllorhina.

OF THE MAMMALIAN OSSICULA AUDITUS. 447

At first sight the anatomist may be inclined to consider that pm (fig. 20) is accessory,
and acpm the homologue of the processus muscularis; for acpm lies in the usual
position occupied by that process in laminated mallei; and though the muscular process
is often found lower down, as in the Cavies, where it is seated on the manubrium, it is,
on the other hand, rarely seen higher up than acpm. But whilst I have never
found both processes coexisting in the malleus of any other Bat*, the upper of the two
observed in Asellia (pm, fig. 20) is constantly observed in the remaining insectivorous
Chiroptera (Pl. LXIJI. figs. 22, 24, 25, pm). As, then, the lower process, acpm, in
Asellia, is the peculiarity of this (and probably of a few allied Bats), and the upper is the
most frequent in others, it is reasonable to consider the latter, pm, as the homologue of
the processus muscularis, so common in other orders, the lower acpm being an acces-
sory process. From the base of acpm a sort of splint or buttress runs for a short
distance across the lamina; it must not be mistaken for the thin processus gracilis below
i, though it is most probably ossified as a part of that element.

Besides the structures just described, a large orbicular apophysis exists, identical in
position to the same in Sorex and Mus, and as large in proportion to the entire bone as
in those animals, but less conspicuous, because it is sessile, not pedunculated. It is far
better developed than in Pteropus.

The processus gracilis is slender, and appears as the thickened free border of the
lamina; it is joined by a process from the head, the homologue of what has been
already very frequently described in this memoir; the two form a rather broad curved
process, which fits into a cleft in the ring-like tympanic bone. The lamina is very
broad, and formed entirely of semitransparent bone; a buttress from the lower processus
muscularis runs across it, but is lost before the processus gracilis joins the process from
the head.

The manubrium is slender and long, well compressed laterally, and narrow, in that
sense, even at the root. It is not triangular, as is the case in most laminated mallei, but
truly quadrilateral ; for the inner aspect as well as the outer forms a narrow surface
sharply bordered from the sides almost to the point. A tendency to this condition may
be observed in the malleus of the Shrew and in some Carnivora, but never to so complete
an extent ; for in Asellia this inner surface is as broad as the outer. In the great majority
of animals the inner aspect of the manubrium is a simple border, sharp or blunt as the
case may be. The outer border or, rather, surface is very narrow, but ends below ina
distinct spatulate dilatationt. Above it forms a rather sharp angle, slightly incurved
towards the neck, in fact a rudimentary processus brevis.

In the well-formed lamina and orbicular and muscular process the malleus of Asellia
tridens resembles that of certain Insectivora and Rodents, and differs from the less
specialized type observed in Péeropus. It must be borne in mind that the characters of
the malleus in Asellia and other insectivorous Bats almost coincide with the distinguish-
ing features of the same ossicle in two tribes of mammals belonging to different Orders, but

* Excepting Phyllorhina fulva and P. speoris, where the malleus in all other points precisely resembles that of
Asellia tridens. + This dilatation is shown in fig. 21, the malleus of Plecotus.

30 2

448 MR. A. H. G. DORAN ON THE MORPHOLOGY

alike remarkable for the small size of the species, namely the Muridz and Soricide.
There may be some functional explanation of this similitude.

Among the VESPERTILIONIDA, the malleus of Plecotus awritus (Pl. LXITI. fig. 21) closely
resembles that of Asellia ; but the lower process (fig. 20, ac pm) is absent. The extremity
of the manubrium is rather widely spatulate.

The stapes has long crura, both almost straight; unlike most Bats, they diverge but
little; the aperture is on that account long and narrow; itis not occupied, in the recent
skull, by any bony canal.

In Vesperugo serotinus (Pl. LXII. fig. 25) the malleus also much resembles that of
Asellia; the upper or true processus muscularis alone exists, so that we miss the lower,
which in Asellia sends a conspicuous buttress from its base across the lamina. The
orbicular apophysis is smaller and more pointed in Vesperugo.

The incus of Vesperugo has a well-formed body, deeply excavated by the articular
surface; the processus brevis is short, but stout; the processus longus is very long and
divergent; it is slightly grooved on its upper and anterior aspect, and ends in an almost
discoid Sylvian apophysis, mounted on a very slender peduncle. In general characters
this incus is of a low type; the same form is frequent among Marsupials. In all the
insectivorous Bats the body of the incus is deeper and better developed over the pro-
cessus longus than it is over the shorter crus. The minute stapes of Vesperugo has a
small broad head, with a tubercle on its posterior aspect. The crura are slender and
very divergent; the posterior is quite straight, the anterior a littlecurved. The aperture
is very wide; the base has a rather thick border, so as to be slightly concave towards the
vestibule. It is much narrower anteriorly than behind—a well-marked feature here,
seen to a less extent in the stapes of Homo, and still less distinctly in most mammals.

The completely bicrurate stapes of these small Chiroptera, with a moderate or wide
aperture, is interesting when we consider the low type of the other two ossicula; but the
wideness of the aperture may not so much indicate high type, but may be considered as
merely due to the passage of an artery * between the crura; the same remark, as I have
already observed, applies to many Insectivora where the stapes has a wide aperture.
Whether a vessel passes through the opening free or unsupported by a bony canal is of
small moment ; still some may consider the wideness of the aperture as a sign of high
type, since in Lemur, where a bony canal exists, the crura do not diverge widely ; so that
in the Bats and Shrews the wideness may not be entirely for the benefit of the canal or
artery, especially as these structures do not always completely block up the aperture.
In many Rodents I have found the blocking complete without the crura being much
bowed or very divergent, as in some skulls of Oapromys.

The malleus of Vespertilio nattereri (Pl. LXII. fig. 22) is very similar to that of Ves-
perugo. The processus muscularis, too, in this Bat, seems in position to be the homo-
logue of the upper process in Asellia, especially as, in the place of the lower distinct
process in the latter, I observe in Vespertilio a faint eminence which sends a pro-

* Seo Hyrtl, op. cit. pl. ii. fig. 13. This artery is given off from the carotid together with the occipital.

OF THE MAMMALIAN OSSICULA AUDITUS. 449

longation across the lamina. The processus longus of the incus is very long. The stapes
is identical in form with that of Vesperugo*.

Among the EMBALLonuripz WMolossus obscurus shows in its malleus (Pl. LXII.
fig. 24) all the typical appearances above described in Asellia; but the processus mus-
cularis in Yolossus, as in most Bats, represents the upper process described as existing
in Asellia. The orbicular apophysis is not so large as in the last-mentioned genus. The
quadrilateral condition of the manubrium is very wel! marked.

Among the Puyiuostomip# I fail to find any distinctive characters. In the malleus
of Chilonycteris osberni the processus muscularis is very stout and long, and I have
preserved the tensor tympani muscle actually inserted into it in a specimen in the
College collection; but the process is the same in position as that in Molossus, hence
corresponding to the upper process in Asellia. The reader is referred to the description
of the malleus in the latter genus for arguments on the homologies of the two processes.
The mere fact of the upper process being found the most constantly throughout the
Chiroptera is, as already remarked, in favour of its being the true processus muscularis:
the lower being so very rarely found.

The incus of Chilonycteris has an extremely long processus longus, but the processus
brevis is much less developed than in the Bats already described; this in itself is
decidedly a sign of low type.

In Mormops blainvillii the malleus has a similar character to that of Chilonycteris; the
processus muscularis is in the same position (Pl. LXII. fig. 28).

I regret I have been unable to procure ossicula from the crania of Bats belonging to
other genera. Sufficient material, however, has been examined to warrant certain definite
conclusions on the characters of the ear-bones of the Chiroptera.

I. In all the Chiroptera the ossicula much resemble those of the Soricide and allied
Insectivora, particularly in the malleus.

Il. In the frugivorous Bats the malleus does not resemble that of Sorex so strongly
as does the same ossicle in the remaining section of this Order; indeed it approaches the
marsupial type im general characters, but bears a strongly marked rudiment of the
orbicular apophysis seen in the Shrew.

III. In all the insectivorous Bats the malleus bears a long, curved processus mus-
cularis, and in some genera (at least in Asellia) a process apparently accessory to it.
A well-formed orbicular apophysis is constant, the lamina is always wide, and the
manubrium is quadrilateral, the outer and inner aspects being sharply bordered off from
both the sides.

IV. The incus in all the Chiroptera has a very short processus brevis and a very long
divergent processus longus, as in: Sorex and the Marsupials. In this point the Shrews
and Bats differ from the Muridze, where the malleus is similar in character to the same
ossicle in the former animals.

V. The stapes is never columelliform, or even approaching that low type, but, on the
contrary, the aperture is generally wide; this is partly necessary, at least, for the

* Prof. Owen describes in Vespertilio noctula ‘‘a retention of the columelliform confluence of stapes and incus *
(Anat. of Verteb. vol. iti.).

450 MR. A. H. G. DORAN. ON. THE MORPHOLOGY

transmission of an artery unsupported by any bony canal, but may be considered as
well (for reasons given above at length) to indicate an approach to the Primates.

VI. Judging from their ossicles it may be said that the Chiroptera are very closely
allied to the Insectivora far more than to any other order of mammals.

THE OSSICULA OF THE CETACEA.

The highly specialized tympanic and petrous bones of the Whales and Dolphins are
now familiar, not only to the anatomist, who confines his studies to recent animals, but
also to the student of paleontology, on account of their great density and consequent
durability. The already known peculiarities of the soft parts in the auditory apparatus,
as they exist in this Order, may be conveniently studied in the fine preparations made by
Hunter, and now preserved in the Museum of the Royal College of Surgeons*. His
observations, quoted in the Catalogue +, are full and satisfactory as far as concerns the
common species described therein, but preparations of the tympanum and labyrinth of
rarer types remain unrepresented in that series. Further researches in the anatomy of
the cetacean tympanum and membranous labyrinth are much to be desired.

But, in accordance with the scope of this paper, the morphological characters of the
ossicula must alone occupy our attention. I will describe them relatively, as touching
the distinction of each ossicle in the different genera, and comparatively as regards the
homologies of the several parts of the highly modified cetacean ear-bones to correspond-
ing portions in the more familiar types of terrestrial or partially terrestrial mammalia.

In the College collection there is one fine specimen of the malleus of Balena mys-
ticetus (Pl. LXII. fig. 28), and another, precisely similar in character, attached to the
tympanic bone (No. 2488, Osteol. Ser., Pl. LXIII. fig. 13), probably from B. australis. At
the first glance, no part of this bone, as it is seen in other animals, can be distinguished
in these specimens, excepting the articular surface; after a little observation, the pro-
cessus gracilis will be recognized; and by further careful study of the specimen, the
observer will find that the modifications are not quite so great as they at first appear.

The peculiarities of this ossicle may be rendered more intelligible by first examining
it i situ, as in Pl. LXIII. fig. 18, which includes part of the (left) tympanic and petrous
bones in B. australis and is viewed from in front. The malleus is seen to be firmly
ankylosed to the tympanic (tb) by its processus gracilis (pg) and by certain appendages
(7., phm), presently to be explained. That ossicle is, in Balena, anterior to the membrana
tympani; and the processus gracilis runs, not forwards, but outwards, to a special pro-
jection of the margin of the tympanic bone to which it is attached. This projection
probahly represents the anterior extremity of the annulus tympanicus of other mammals,
the part in front being an extension of that ossification to surround the beginning of the
‘Eustachian canal. The petrous bone (pet) lies much above the tympanic; hence the
unusual position of the articular surface of the malleus, as Cuvier has observed. It
follows that the homologous parts of the malleus of Balena and the same bone in an
ordinary mammal will not be found in the same respective aspects of each bone.

® Nos. 1582-1598 8, Physiological Series. + Physiology, vol. iii. p. 115.

OF THE MAMMALIAN OSSICULA AUDITUS. 451

The malleus of B. mysticetus (Pl. LXII. fig. 28) measures about half an inch or a little
more in length. The head is very large; its intero-inferior aspect is smooth and quite
convex; externally and a little anteriorly it is very deeply excavated, the upper border
of the cavity extensively overhanging it, especially towards the front, whilst behind there
is a deep notch in that border. Hyrtl justly compares the excavation to that seen in the
anterior aspect of the malleus of the Seal (Phoca); and from the extremely overhanging
border, the bone assumes a form seen in certain shells, as Van Beneden has observed ;
indeed, putting aside the articular region and manubrium, the head much resembles the
tympanic bone to which it is attached. On its upper aspect is situated the articular
surface, which looks a little inwards and is deeply concave. It is made up of two
facets, both of a semicircular form; the upper and external is almost plane, and by far
the most extensive; it is separated from the lower, which is convex, by a sharp groove.
In front of the head, on the outer aspect, is a deep channel, distinct superiorly, but
absent below. This cuts off a portion of the malleus, which, close to the channel, is
broad and flattened, bearing a distinct tubercle (Pl. LXII. fig. 28, & Pl. LXIIL. fig. 18,
pm), which projects upwards and inwards; but beyond the tubercle it gradually nar-
rows as it runs forwards, ending in a hook-like point (mz), which curves downwards and .
outwards, and is longer and narrower in B. australis than in B. mysticetus. The
tubercle (pm) is the processus muscularis, as may be seen in preparation No. 1598 B in
the Physiol. Ser. Coll. Surg., where the tensor tympani muscle remains still attached to
it. The hook-like process (mz) is without doubt the true manubrium ; into it is inserted
the apex of the curious prolongation of the membrana tympani characteristic in this
Order. :

The homologous parts of the malleus in Balena (Pl. LXIII. fig. 13) may be com-
pared with the same in the more familiar malleus of Felés (Pl. LXIII. fig. 14).

Cuvier states, in his ‘ Lecons d’Anatomie Comparée,’ that in the Dolphin there is no
manubrium, and that the malleus of the Whale “resembles it in every respect, but is
double its size.’ Hyrtl (J. c.) correctly denies this assertion, and remarks of the Ceta-
cean manubrium :—“ It is only very short, and reduced to a rounded bony tubercle,
separated by a deep incision from the head. . . . The processus minor [sic] is represented
by two small pointed appendages, to which a triangular process of the membrana tym-
pani attaches itself.” But Van Beneden remarks* of Balena mysticetus :—“< The
malleus possesses neither any distinct apophyses nora manubrium.” Lastly, Huxley +
asserts, in describing the anatomy of the Porpoise:—‘“‘ The tympanic membrane is
concave externally; and, as is usual in the CuTacra, is connected by a ligament
with the handle of the malleus.” I cannot see how the curved process of the malleus
of Balena can represent any thing otherwise than the inflected splinter of bone on the
blunt extremity of the malleus of the Dolphin {, which is the true manubrium ofall other
mammals; and, judging both by analogy and by the preparation above referred to, the

* ¢ Ostéographie des Cétacés,’ Yan Beneden and Gervais.
+ ‘ Manual of the Anatomy of Vertebrated Animals,’ 1871.
+ Pl. LXIII. fig. 15, mn, from a Globiocephalus.

452 MR. A. H. G. DORAN ON THE MORPHOLOGY

tubercle can hardly be fairly denied to be the homologue of the processus muscularis.
Taking these assertions for granted, and recognizing henceforth in this treatise the
curved process as the handle of the malleus, it will be shown that the manubrium is far
better developed in Balena than in the Dolphins or Sperm-whales, and still more than
in the Ziphioid Physeteride.

From the malleus to the tympanic bone runs, almost directly outwards, a process over
half an inch in length. It is made up of two stout splints or columns, one forming its
posterior, the other its anterior border, and a broad flooring of thin bone connects
them by their inferior borders ; this latter is continuous with the floor of the excavation
in the head of the malleus, the posterior raised and everted margin of that cavity
running to join the posterior splint, the anterior coming distinctly from the corresponding
margin of the depression in the malleus of B. australis; in that of B. mysticetus the
anterior splint is given off in front of the groove, at least so I find it in the spe-
cimens I have examined; but this appears to be a trifling point, probably subject to
variety.

The whole of this process, highly characteristic of the Cetacea, has for years been
‘recognized by most authorities as the processus gracilis. One anatomist, Buchanan, who
has carefully studied the anatomy of the ear in this Order, and written a most lucid
description of the Whale’s membrana tympani, represents this bony connecting medium
in a Plate *, and there calls it “the long handle of the malleus ;”’ whilst the process mn
in Pl. LXIT. fig. 28, & Pl. LXIIT. fig. 18, at the end of this paper,’is represented in
Buchanan’s illustration as very much longer than I can find it in any specimen in the
College Museum, and styled “the short [sic], slender, or gracilis processus of the mal-
leus.” Here is a great confusion of terms, as other anatomists make the “long” and
“‘eracilis’? process synonymous; then, too, the expression “long handle” is very vague.
Hunter (Phil. Trans. 1787) merely says that the process from the membrana tympani is
inserted “into that bone” (@. e. the malleus), and speaks of the other process, that is the
bony attachment of the malleus, without naming the homologies of those parts of the
ossicle in question. Home (Phil. Trans. 1812) figures the ossicles very correctly, but calls
the manubrium the “ short handle,” and the processus gracilis the “long handle.”

But when we come to compare once more the process connecting the malleus to the
tympanic bone in Balena with the more typical malleus of Helis (Pl. LXIII. fig. 14), we
- shall find that, more strictly speaking, the anterior splint (fig. 13, pg) is the homologue
of the processus gracilis, whilst the posterior (phim) is the prolongation towards it from
the head of the malleus seen so frequently in terrestrial mammals; then the floor of
thin bone between them (2) represents the lamina (fig. 14, 7) of the Carnivora, Ruminants,
&c. The whole of the above connecting process between the malleus and tympanic bone
is rather longer in Balena, and perhaps in Balenoptera, than in the Dolphins and
Ziphioid Whales.

Tn its large excavated head and short curved manubrium, the malleus of Balena bears
a distant resemblance to the same in Macrorhinus and Stenorhynchus among the Seals.

* «Physiological Tlustrations of the Organ of Hearing.’ London, 1828.

OF THE MAMMALIAN OSSICULA AUDITUS. 453

The incus of B. mysticetus (Pl. LXII. fig. 29) and of B. australis, which exactly
resembles it, is more generalized than in any other cetacean (excepting, perhaps, Wega-
ptera and Neobalena). From the position of the articular surface of the malleus, the
incus lies above and internal to it, with the corresponding surface downwards. The body
is very solid and square. The posterior crus is short; stout, and recurved at its extremity.
The processus longus, not so oyer-developed as in other Cetacea, is still very stout and
broad to its extremity; it forms a bold sigmoid projection. A faint groove cuts off the
Sylvian apophysis from the rest of the processus longus; it does not expand or project in
any way from the crus itself. The articular surface on the body is very prominent; its
upper facet is almost plane and larger than the lower, as in the malleus; a sharp ste:
which separates them, fits into the groove between the facets of the malleus.

The stapes (Pl. LXII. fig. 29) is nearly half an inch long and very narrow. The head
is well developed, and bears a rough surface posteriorly for the insertion of the stapedius
tendon. The crura are long, straight, and slender; they diverge but little, and are con-
nected by a lamella, which has only a very small aperture in it; this lamella is evidently
an ossification of the thin membrane thrown across the crura in the recent stapes of a
typical mammal. ‘The base is generally immovably soldered to the fenestra ovalis *.
The stapes of Balena bears a certain resemblance to that of Stenorhynchus, and differs
from all other Cetacea in the length and slenderness of its crura. )

Tn describing the process which unites the malleus to the tympanic bone, the anterior
column or border of that process (Pl. LXIT. fig. 28, Pl. LXIII. fig. 18) was assumed to
represent the true processus gracilis, and the posterior (phm) as homologous to the
prolongation from the head of the malleus, which in other mammals generally joins it.
This posterior column runs as an elevated ridge two or more inches over the outer
surface of the scroll-like tympanic bone, ending in a broad lamina, which is closely
applied to the upper part of the deep oblique groove which separates the anterior from
the posterior portion of that bone, which would seem to indicate that the posterior
column was the processus gracilis. But the anterior, which seems, from its position
between the head and manubrium, to be the true processus gracilis, is soon lost on the
immer side of the tympanic bone, where it makes a sharp ridge in some whales (Phy-
seter &c.), as if the root of Meckel’s cartilage had become embedded in that bone during
ossification. Altogether, then, arguments are in favour of the anterior column being the
homologue of the processus gracilis. This can alone be verified by the examination of
very small foetal Cetacea.

In the Pike Whale (Balenoptera rostrata) the auditory ossicles assume much more
thoroughly the type most frequent in the Cetacea, excepting Balena.

Indeed the malleus (Pl. LXIT. fig. 30) is as extreme in its modifications as that of
Globiocephalus, Phocena, and Delphinus, being only exceeded in specialization by the
Ziphioids. It is less like the malleus of Balena than is the same ossicle in Physeter or
even Orca.

* In Balena mysticetus; but I have since found this not to be invariable, and in an adult petrous bone of
B. australis in the College Museum the stapes has fallen out.
SECOND SERIES.—ZOOLOGY, VOL. I. 3P

454 MR. A. H. G. DORAN ON THE MORPHOLOGY

The head is less developed than in Balena, especially below and internally *; it has a
deep depression externally, of which the borders are elevated, but not nearly so over-
hanging as in the Greenland Whale. The articular surface is very similar, and the
upper facet is the larger. In front of the head is a very prominent well-rounded tubercle,
on the under and inner surface of which is a sharp splinter of bone (mm), with its point
directed backwards; it lies flat against the malleus.

Hyrtl, in denying Cuvier’s assertion that the manubrium is absent in Dolphins and
Whales, continues :—‘“ It is only very short, and reduced to a rounded bony tubercle,
separated by a deep incision from the head.” But in describing the tympanum of Mo-
nodon he remarks :—‘‘ The fleshy process of the membrana tympani is half an inch long,
and attached to a ledge [ Leiste| on the outer border of the malleus.” Now the malleus
of Balenoptera is very similar to that of Monodon, and still more to that of Delphinus
and Phocena. By carefully dissecting the tympanum of a Monodon, and examining the
preparations of that cavity from all the remaining above-mentioned genera as seen in
the College Museum (Nos. 1590-3 and 1596), I find that the tensor tympani tendon is
almost invariably inserted into the most prominent and anterior part of the rounded
tubercle of the malleus, whilst the process from the membrana tympani + is inserted, as
Hyrtl says, into the splinter of bone described above; it is, however, on the under, not
the outer surface of the malleus. Hence it must be correct to consider the bony splinter,
Hyrtl’s “ Zeiste,” as the manubrium, whilst the bony tubercle } should be considered
the homologue of the processus muscularis of other mammals. The short curved process
in the Greenland Whale, well divergent from the body, is evidently an intermediate
condition between the long manubrium of terrestrial quadrupeds and the flattened
splinter of Balenoptera and the Dolphins.

The process joining the malleus to the tympanic bone is very similar to that in
Balena, but shorter. The anterior pillar, or true processus gracilis, is very stout, and
the lamina is rather denser than in Balena. i

The incus (Pl. LXII. fig. 31) assumes, like the malleus, the form prevailing in genera
presently to be referred to. The body (which bears an articular surface similar to that
of Balena) is very ill developed, and the posterior crus is almost atrophied ; it measures
about a line in length, though tolerably stout. The processus longus is enormously
developed, and incurved so abruptly as to present anteriorly a knee-like projection. On
its broad extremity it bears a smaller and narrower, thick and elliptical Sylvian apo-
physis, of about the size and shape of the base of a Squirrel’s stapes. Hence the incus
of Balenoptera differs from that of Balena more than the latter is different from that
of most terrestrial Mammalia.

The stapes (Pl. LXII. fig. 31) is not so long as in the Greenland Whale, though its
crura are longer than in the Dolphins. The head is well developed, and bears a rough

* Tt is much flattened on its outer and inner aspects.

+ See Pl. LXIII. fig. 16, where the tendon of the tensor tympani and the process from the membrana tympani
(in Monodon) are both indicated.

+ The exact site of the insertion of this tendon is marked by a rough surface in Balenoptera, Orca, and most
other Cetaceans.

OF THE MAMMALIAN OSSICULA AUDITUS. 455

tubercle for the insertion of the stapedius tendon. The crura are very stout, straight,
and more divergent than in Balena. The base was not soldered to the fenestra ovalis
in the stapes removed from a skull of an adult male in the College collection, but could
be displaced uninjured without any force being required to detach it; the same was the
ease with the stapes of an adult Balenoptera in the British Museum. ‘The base is
quite concave towards the vestibule. The aperture between the crura is small and
circular. :

The malleus of the small New-Zealand Fin-Whale (Balenoptera huttonii, Gray) in the
British Museum almost precisely resembles that of B. rostrata; the tubercle is propor-
tionately less developed, and the splint-like manubrium of the northern species is, in
B. huttonii, reduced to a mere rough surface, as in many Dolphins.

A stapes from the gigantic Steypireythr (Balenoptera sibbaldii), kindly presented to the
College by Professor Turner, has longer crura than that of B. rostrata, but is only half
as large again *.

The malleus of Megaptera nove-zelandi@ (Pl. LXII. fig. 32) has its head as flattened
as in the last described genus; but in front, instead of a short round tubercle, is a stout,
long, blunt-pointed process curved downwards. On the inner aspect of the latter is a
sharp spine (mn), directed backwards, and representing, as in Balenoptera, the manu-
brium. The blunt tip of the process (pm) has a rough surface, evidently for the insertion
of the tensor tympani; and posteriorly, close to the head, is a prominent tubercle, only
faintly indicated in Balena and Balenoptera, the analogies of which are not evident,
though it decidedly is not a muscular process in the last-named genera, as preparations
in spirit, already quoted, clearly show.

I have not had the fortune of examining the incus of a Megaptera, and cannot
find a detailed description of it in any work on the Ear or on Cetacea. Van Beneden
states that it is small, and that one of its articular surfaces is ill developed (in I. boops) ;
in the illustrations to his work, that ossicle is represented with a body almost as well
developed as in Balena, and the “short crus” is depicted rather stout, long, and pointed,
the “long” process not very thick.

The stapes of Megaptera nove-zelandia, and also that of MW. longimana, Gray, has
as long crura and as wide an aperture as in Balena; in a specimen of the former
it was found ankylosed to the petrous bone; in the latter species (judging from an adult
specimen in the British Museum) such ankylosis is at least not invariable.

In the greater length of the process bearing the homologue of the manubrium and the
insertion of the tensor tympani, in the form of the stapes and probably of the incus,
Megaptera tends more towards Balena than to Balenoptera; but, on the other hand,
the flatness of the caput mallei relates it very nearly to the latter; and the whole process
from the head is produced and shaped not so much in-the manner seen in Balena, but
rather as in Platanista.

The malleus of Neobalena marginata + (Pl. LXII. fig. 33) is highly interesting, since,

* See Appendix to this section.
+ Dr. Hector (Trans. N. Z. Inst. 1872) considers that this species, known to Dr. Gray under that name through .
oP 2

456 MR. A. H. G. DORAN ON THE MORPHOLOGY

like that of Megaptera, it exhibits intermediate characters between Balena and Baleno-
ptera. It is larger than the malleus of an adult Balena; its head is less flattened than
in Balenoptera. The upper facet of the articular surface is extremely wide and almost
plane; the lower is small, and as convex as in most Cetaceans; it must be remembered
that in the Greenland Whale and Southern Whale the former facet is rather larger than
in Balenoptera. From the front of the head a process projects, which bears on its inner
side a sharp spine (mn), directed more downwards and less compressed against the body
of the malleus than in the last-mentioned whale, but neither so long nor directed so much
downwards as the stout curved manubrium of Balena, which it represents (Pl. LXII.
fig. 28, Pl. LXIII. fig. 18, mn).

In the plates illustrating Camper’s ‘Observations Anatomiques sur la Structure et le
Squelette des Cétacés’ the ossicula of Physeter are very correctly figured. He speaks .
of “la manche, ou lapophyse de Rau”’ as if those terms were synonymous ; the site of
the insertion of the tensor tympani is not mentioned or indicated.

The malleus in the Cachalot and in the Ziphioid Whales possesses highly distinctive
characters.

This ossicle in Physeter macrocephalus (Pl. LXII. fig. 34) is very small in proportion
to the bulk of the animal. In general form it may be compared to a half-split pea,
which it hardly exceeds in size. The head is small and convex; it does not bulge or rise
high over the upper facet. The depression on its outer aspect exists, but is not so
extensive as in any of the Baleenoidea. The articular surface is very wide, considering
the ill-development of most of the elements of the bone. Its upper facet is very broad
and almost plane, the lower is much smaller, convex, and everted, like the mouth of a
pitcher.

The process connecting the malleus to the tympanic bone is extremely short, but the
same elements can be distinguished in it as in Balena.

But the most distinctive feature in the malleus of Physeter is the suppression of the
projecting tubercle which in other Cetacea bears the homologue of the manubrium and
the insertion of the tensor tympani. On the anterior aspect of the head is a sharp, pointed
tubercle (pm) looking directly upwards; directly below it is a very small blunt projec-
tion, pointing, in some specimens, a little forwards; in most examples it is hardly per-
ceptible. This (mz) is the representative of the manubrium, and is much better deve-
loped in some of the Ziphioid Whales shortly to be described. In specimen No. 1584,
Physiological Series, in the Museum of the Royal College of Surgeons, showing the
auditory apparatus of Hyperoodon, where the malleus is very similar in form to the same
ossicle in Physeter, the insertion of the membrana tympani to the tubercle (mn,
Pl. LXI1. fig. 35), and of the tensor tympani muscle into the upper process (pm), is well
displayed. Hence mn in figs. 34 & 35 is the representative of the manubrium in Phy-
seter and Hyperoodon, whilst pm is the homologue of the processus muscularis.

piece of whalebone only, is identical with the Caperea antipodurum, described in the B. M. catalogue from a solitary
tympanic bone. Gervais and Van Beneden consider Caperea antipodarum to be the same as their Balena antipo-
darum, which they distinguish from B. australis, and rightly, so far as the mallei are concerned.

OF THE MAMMALIAN OSSICULA AUDITUS. 457

The incus of Physeter has a much smaller body than that of Balena; but this
portion of the ossicle is, on the other hand, not so reduced in size as in Balenoptera and
the Dolphins. The articular surface is very wide; the upper facet is the larger and
planer, the lower is very concave, to fit the corresponding parts on the malleus. The
ridge dividing them is most distinct anteriorly. The processus brevis is slender, but
rather long; the processus longus is very stout, but not so big as in the Delphinoidea
and in Balenoptera; it bears at its extremity a small, sessile, elliptical Sylvian
apophysis.

The stapes has extremely stout crura, not so divergent as in the Dolphins ; the aperture
is sometimes obliterated. The footplate has a very thick border, and is a little con-
cave towards the vestibule. I have never found this ossicle ankylosed to the petrous
bone.

In the Ziphioid Whales the ossicula much resemble those of Physeter.

In Hyperoodon rostratus (Pl. LXII. fig. 35) the malleus has a larger head than in the
Cachalot ; that portion of the ossicle is also more convex, and rises a little above the
upper border of the articular surface.

There is hardly any trace of a depression on the outer aspect of the head; it seems
rather as if the bone above had grown into it, and reduced it to a horizontal slit. The
process holding the malleus to the petrous bone is extremely short, but its homologies
can be easily made out. The articular surface is, proportionately to the rest of the
ossicle, considerably smaller than in Physeter, nor is the upper so much larger than the
lower facet.

The processus muscularis (Pl. LXII. fig. 35, pm) is even less developed than in
Physeter; the manubrium (mz) is a still more insignificant tubercle.

The incus also in general characters resembles the same in the Cachalot, but possesses
a character which it shares with the two other Ziphioid Whales whose ossicula the author
has examined, and in which it differs from Physeter. This feature is the long, flattened,
and very divergent processus brevis, which is narrowest in the middle. The articular
surface has its facets divided by a very sharp ridge, equally defined in its whole course ;
the upper facet is plane, the lower concave, as in Physeter, but consistently with the
characters of the articular area in the malleus of Hyperoodon the upper facet is, in its
incus, not so much larger than the lower, as is the case in the Cachalot. The body is
developed to the same extent as in Physeter; the processus longus is as stout but
shorter than its representative in that Whale.

The stapes in Hyperoodon (Pl. LXII. fig. 35) has very stout crura, with hardly a
vestige of an aperture. The base is narrower than in Physeter, and not nearly so thick at
its margin ; towards the vestibule it is quite concave.

The stapes, but not the other ossicula, of Ziphius cavirostris is figured in a plate
illustrating Prof. Turner’s paper on that Cetacean*. It resembles, apparently, that of
Hyperoodon.

The malleus from the fine adult specimen of Berardius arnuaii in the College Museum
(Pl. LXIII. figs. 1 & 18) bears, to a well-marked degree, all the chief peculiarities seen

* Trans, Royal Soc. Edinb. 1872.

458 MR. A. H. G. DORAN ON THE MORPHOLOGY

in Physeter and Hyperoodon, The head is better developed than in either of those
Whales. It is half as long again horizontally as vertically, whilst in Hyperoodon
this portion of the ossicle, though not more prominently convex than in Berardius, is
almost a hemisphere. It rises very prominently above the facets. The depression on its
outer aspect is small and slit-like; the process holding the ossicle to the tympanic bone
is as short as in the Physeteride already described, and made up of the same elements
(compare Pl. LXIII. fig. 1 with figs. 13 & 14). The specialized modifications in the
processus muscularis and manubrium are very marked in the malleus of Berardius. The
processus muscularis (fig. 1, pm) is nearly as long as in Physeter, so that it is better
developed than in Hyperoodon. Here it may be observed that it is only in the Phy-
seteridee that the processus muscularis can be found as an actual process among the
Cetacea, sincein the Whalebone Whales and Dolphins its homologue is either a rough sur-
face or a depression on the tubercle, from which also springs the manubrium; but itis this
very tubercle which is wanting in the Cachalots and Ziphioid Whales, where the processus
muscularis is distinct.

In Berardius the manubrium is much better developed than in Hyperoodon or
Physeter. It is seen as a small, sharp spine *, arising close to the root of the processus
muscularis, and running directly downwards, whilst the other process runs straight
upwards. The two processes together lying close against the head, resemble the con-
trivance seen on board ship for belaying cable. The articular surface is extensive even
in proportion to the rest of the malleus; its upper facet is of the same form as in
Hyperoodon, plane, and very elevated at its margin; the lower is smaller and convex,
everted as in the Ziphioid Whale just mentioned.

Henee, in its malleus, Berardius is more perfectly specialized than either Pleat
or Hyperoodon. This ossicule is certainly, among these three Whales, largest and
bulkiest in Berardius, smallest, proportionately, in Physeter, owing to the ill-develop-
ment of the portion of the head external to the facets in that animal.

The incus (Pl. LXIIT. fig. 1) resembles that of Hyperoodon rather than Physeter; the
processus “‘ brevis” is still longer, more divergent, and flattened and expanded at its
extremity. Close to its root the body (as is the case in Hyperoodon) is not so reduced
and shallow as in Physeler; the groove dividing the facets is not very sharp in
Berardius. The processus longus is quite similar in character to that of Hyperoodon.

- The stapes is longer than in the other Physeteridee here described. In the College
specimens there is a prominent tubercle on the inner aspect of the head for the tendon
of the stapedius. The cruraare long, stout, and straight, a slit-like but pervious aperture
divides them. The base is narrow, its margin is rather thick, and it is quite concave
towards the vestibule.

I have examined the malleus and incus from a skull of Mesoplodon hectori in the
British Museum. These bones are much smaller than their representatives in the above-
described Physeteride. The malleus has a head as small as in Physeter, moderately
convex, and produced backwards to a prominent blunt point. The head projects slightly
above the facets, as in Hyperoodon; the articular facets are proportionately still smaller

* Vl. LXIII, fig. 1, where this spine is lettered im in place of mn, its true homologue.

OF THE MAMMALIAN OSSICULA AUDITUS. 459

than in that Whale, nor is the upper much larger than the lower. The processus mus-
cularis is a short blunt point. I found a few shreds of the tendon of the tensor tympani
adherent toit. The manubrium is represented by the faintest trace of a tubercle; there
is a ridge above it, into which a part of the process from the membrana tympani is pro-
bably inserted as in Platanista.

The processus gracilis and its accessories eonnectiae the malleus to the tympanic bone
is extremely short, and the depression in the head above it almost effaced. See Pl. LXIII.
figs. 19 & 19 a, where this bone in the closely allied Wesoplodon grayii is shown.

The incus is of the same character as in Hyneroodon and Berardius; it is rather

smaller in proportion to the malleus. The ridge separating its facets is almost as sharp
as in Hyperoodon.

Taken as a whole, and judging from the incus as well as the malleus, Mesoplodon
resembles Hyperoodon rather than Berardius. In the ill-development of the head of the
malleus we have a character which should place this Whale between Physeter and
Hyperoodon. In the smallness of the articular surface of the malleus it differs alike from
the Cachalot and Berardius, and approaches Hyperoodon.

The stapes has no aperture, and in form coincides with the same DSelele | in Hyperoodon,
confirming the remarks in the last paragraph.

In the Delphinidee the ossicles may be said to be central in character among the
Cetacea, the Baleenoidea and Platanistidee departing from them towards other mammals,
the Physeteride diverging, on the other hand, from the Dolphins in the highly specialized
characters of the malleus and incus. It is remarkable that among the Whales proper
the genus Balenoptera should so closely resemble the Dolphins in its ossicula.

In Delphinus the malleus (Pl. LXIII. figs. 2 & 3) in general form reminds the observer
strongly of the same in Balenoptera*. The outer aspect of the head is less flattened,
the articular facets are similar, the upper being the larger and planer, the lower the most
convex; but instead of the large rounded tubercle seen in the Whale just mentioned, a
small, blunt, slightly excurved process projects from the head in Delphinus. On its
inner surface may be observed a small spine (mm), directed backwards, hardly perceptible
to the eye, yet readily detected by rubbing the finger forward over the internal aspect of
the malleus. This represents the very perceptible homologue of the manubrium
described as existing in the Pike-Whale, and equally or more developed in other genera
of Dolphins rt.

The body of the incus is very feebly developed, being, as it were, absorbed by the
stapedial crus. Its articular surface has facets divided by a sharp ridge, as in the
Baleenoidea, and unlike Physeter and Berardius. The processus brevis is very slender
and much shorter than in the Ziphioids; the processus longus is very stout, and shaped
like its representative in Balenoptera; the Sylvian apophysis is of the same nature.

The stapes is about a line in length, with fairly divergent crura and a very thick base,
quite concave towards the vestibule. ‘The aperture is closed, but always represented in

* The ossicula of a fetal Dolphin, four inches in length, are all of precisely the same form as in the adult.
+ The homologous parts of the malleus in the Dolphins are indicated in Pl. LXIII. fig. 15, representing that
ossicle in Globiocephalus (see also figs. 3 to 12).

460 MR. A. H. G. DORAN ON THE MORPHOLOGY

this genus by a minute pit; and in no specimen of any Dolphin of any age in the Col-
lege Museum was it found ankylosed to the petrous bone, nor in any other Cetacean except
Balena mysticetus, and then not invariably. Hyrtl, who has described the ossicula of
the Cetacea (op. cit.) far more extensively than any anatomist before him, states that all
trace of the aperture is effaced “in Delphinus longirostris, braziliensis, tursio and
rissow ;” but he asserts that it is also absent in Monodon, though a distinct trace is seen
in the two College specimens from different skulls; and although he finds that the
aperture “ reappears in the Porpoise,” it does not seem more distinctly indicated in the
examples in the College collection than in Delphinus, and is decidedly not pervious.
The distinguished anatomist continues :—“ Possibly this depends on the age of the
individual specimen ; a conjecture, however, not justified by the persistence in all other
animals of the form of the stapes in different degrees of age.” I have not found the
aperture more blocked up or less clearly indicated by a depression in adult than in
young Delphinoid crania; and, on the other hand, a specimen of the stapes of Bradypus
tridactylus in the College collection, from an adult Sloth, has a much smaller aperture
than another stapes from a cranium of a younger animal of the same species.

In Globiocephalus (Pl. LXIITI. figs. 4 & 15) the head of the malleus is rather larger
than in Delphinus, and the projecting tubercle rather smaller; but it is marked at its
apex by a distinct rough surface for the insertion of the tensor tympani tendon (pm),
and the manubrium (mm) is represented by a very sharp spine pointing backwards and
flattened against the inner side of the ossicle. The bony process connecting the malleus
to the tympanic bone is rather long, and the same elements as described in Balena
may be observed in it ; the processus gracilis (pg) or the anterior column is thick. The
incus has the same characters as in Delphinus ; the stapes, too, is very similar, but some
specimens have a pervious aperture.

In a Lagenorhynchus in the British Museum the ossicula are of the same form as in
the last genus, and the manubrium is as well developed (fig. 11). In Phocena the
malleus (fig. 5) most resembles that of Globiocephalus; the incus is very similar, and
the stapes has generally a very minute but pervious intercrural aperture. The malleus
is connected by bone to the tympanic, though by a somewhat thin processus gracilis,
contrary ts Hunter’s experience. .

In Delphinapterus albicans (Pl. LXIII. fig. 8), the Beluga or White Whale, the
malleus much more resembles that of Delphinus than the same ossicle in G'lobiocephalus,
the process in front of the head being rather long and curved outwards. The recurved
splinter representing the manubrium in other Dolphins is quite absent, and represented
by a rough ridge (mn), hardly visible except by aid of alens*. ‘There is no excavation
on the outer aspect of the head of the Dolphins as in Balena; it seems as if the edges
so prominent in the Greenland Whale had rolled in and filled up a cavity, and to a less
extent in Physeter.

The incus of Delphinapterus is of the form prevailing among the Delphinide. The
stapedes in the College collection have neither so much as a trace of an aperture.

* As the malleus of this Dolphin is larger than that of most Globiocephali, the absence of the spine-like manu-
brium (as in the smallest Delphint) is not due to diminished proportions or development of the entire ossicle.

OF THE MAMMALIAN OSSICULA AUDITUS. 461

The malleus of a large Orca from Tasmania (Pl. LXIII. fig. 7) in general aspect
appears like a magnified malleus of Delphinus; but some of its elements are propor-
tionally more highly developed than in the typical genus. The head is large and convex ;
the depression on its outer aspect is very deep, and almost recalls Balena. The size of
the processus gracilis and the elements associated with it, connecting the ossicle to the
tympanic bone, contrasts strikingly with the reduced proportions of the homologue of that
process in the malleus of the Physeteride. The articular surface is as small as in Delphi-
nus, and the facets are of similar type; but the tubercle from the head of the malleus is
far larger than in any of the Dolphins of which the ossicula have been already described.
it is much more prominent than in Balenoptera. At its upper extremity, looking
forwards, is a conspicuous rough depressed surface for the insertion of the tensor-
tympani tendon, Below is a sharp splinter-like process representing the manubrium.
This is much more developed than in Delphinus, or even Globiocephalus; and instead of
being directed backwards, and pressed, as it were, against the head, it is directed con-
siderably in a downward direction. ‘The whole tubercle assimilates to its homologue in
Balena, as the same in Megaptera recalls Platanista in general features.

The incus of Orca is quite Delphinoid; its short crus is thin and rather long, the
stapedial crus very stout. The stapes is similar to that of Delphinus; it has a minute
aperture.

In a smaller Grampus (Pseudorca meridionalis, Flower) all the ossicula much resemble
those of Orca. The malleus has the tubercle bearing the manubrium a little less
developed (fig. 6), so as to approach Globiocephalus.

The homologies of the splinter of bone inflected from the process projecting from the
bead of the malleus in most Cetacea, and of the rough surface or actual tubercle seen on
the point of that process, have been discussed in the description of the ossicula of
Balenoptera; and specimens in the College Museum illustrate the relations of the soft
parts in Dolphins, Whales, and Hyperoodon. I have dissected several ears of Dolphins,
Porpoises, and of a Narwhal, and the malleus from the latter dissection (Pl. LXIII.
fig. 16) is mounted in the special collection of ear-bones on which this treatise is
founded. The tendon of the tensor tympani is left attached to the apex of the process
from the head; and the whole membrana tympani is also retained, dried and varnished ;
its broad, flat, and pointed process may be seen attached by the point to the splinter-like
manubrium, Hyrtl’s “Leiste.” All the above preparations conform, on the whole, to
the descriptions given by that distinguished anatomist, and by Buchanan, concerning
the anatomy of the Cetacean tympanum. ‘The dissection above described was suggested.
by a figure of the malleus of Monodon in Hyrtl’s work.

The malleus of the Narwhal (Monodon, Pl. LXIII. figs. 9 & 16), though essentially
the same in type as that of Globsocephalus, is much elongated antero-posteriorly, and the
head merges into the broad process in front without any constriction or trace of sepa-
ration. The articular surfaces are of equal size(as Hyrtl observes), the upper being not
larger than the lower, as in all other Cetacea. The processus gracilis is of the same
type as in Balena and the rest of the Order; it is rather long. The incus has an even

SECOND SERIES.—ZOOLOGY, VOL. I. 3Q

462 MR. A. H. G. DORAN ON THE MORPHOLOGY

thicker stapedial crus than in other Delphinoidea*; the posterior is short and small; and
the facets are, of necessity, of equal size; they are divided by a sharp and prominent ridge.
The stapes is of the ordinary Dolphin type, with a very thick base, generally (but not
always) very concave towards the vestibule. It was found unankylosed to the fenestra
ovalis in a large adult skull in the College Museum. ‘The ossicle in this case had a
minute almost pervious depression between the crura.

In the ossicula of the PLATANISTID# we find interesting modifications. In the Gangetic
Dolphin (Platanista gangetica) the head of the malleus (Pl. LXIII. figs. 12 & 17) is
of the same form as in Delphinus, and the upper facet is larger than the lower; but the
process in front is extremely elongated, so as to be longer than the head itself, from
which it is hardly more distinct than in Monodon, except that a faint groove, generally
observed in the true Dolphins, divides them above and internally. Instead of a rough
and only slightly concave surface at the very extremity of the process, as is found in
most Cetacea to mark the insertion of the tensor tympani, there is a very deep pit on
the external aspect of the projection, actually nearer its root than its pot; this is
highly characteristic (fig. 17). The point itself is sometimes slightly hooked; and
from it to close under the articular surface, all along the inner aspect of the process,
runs a very distinct groove, ending posteriorly in a faintly marked tubercle representing
the manubrium (fig. 12, mn) It seems as if the groove were homologous with the outer
surface of that process in other Orders of the Mammalia, the hooked or blunt point
corresponding to the processus brevis. Many of the fibres of the fleshy prolongation of
the membrana tympani to the malleus are attached to the bottom of that groove, and
are often seen still adherent in imperfectly macerated specimens. In Monodon and the
Dolphins proper some of the fibres are in the same manner inserted along the back of the —
anterior projection from the head of the malleus, as well as to the splinter-like more
definite homologue of the manubrium +.

The processus longus of the incus (fig. 20) isas thick as in Monodon, and the articular
surface for the head of the stapes isas small; the posterior crus is even less developed
than in Delphinus. The stapes is of the same form as in the smaller Dolphins, without a
trace of any intercrural aperture; there is sometimes a large tubercle for the stapedius-
tendon on the inner aspect of the head.

By the kind permission of Dr. Giinther I have been enabled to examine the ossicula
from the skull of Inia geoffroyit belonging to the British Museum. ‘The general shape
ofthe malleus (Pl. LXIII. fig. 10) reminds the observer more of Delphinus than Pla-
tanista. The head and facets resemble the same in the Gangetic Dolphin ; but the process
in front is not nearly so produced as in Platanista. The tensor tympani is inserted into

-a pit (at pm) as in the Gangetic genus, which, however, is seated in Inia quite at the
tip of the process from the malleus, on the site of the same insertion in Delphinus. 'The
manubrium is a very distinct reflected spine (mm); and there is a groove running

* Hyrtl (op. cit.) directs attention to the small size of the articular surface for the head of the stapes on the end of
this process. :

+ In making a fresh dissection of the tympanum of Plataiista, I found a strong band of fibres inserted into the
groove, and the tensor tympani was inserted into the depression above described (Pl. LXIII. fig. 17).

~OF THE MAMMALIAN OSSICULA AUDITUS. 463

forward from it. Hence the malleus of Jnia allies it, on the whole, more to Delphinus
than Platanista. The articular surface for the stapes on the stout processus longus
is as wide as in Delphinus; the processus brevis is rather long, but slender. The
stapes is of the type prevailing among all the Dolphins ; it has an intercrural aperture.
Tn a full-sized adult Inia these ossicula are less than half as large as their representatives
in a Platanista of the same age and proportions.

In taking a general concluding view of the auditory ossicles of the Cetacea, it may be
remarked :—

I. The ossicula of these animals differ from those of other Mammalia rather in their
solidity than in any great size in proportion to the whole skeleton, and more in extreme
modifications of detail than in solidity of composition; for in size they are exceeded by
the Seals and Sirenia, in density by the latter, but in specialization of form by no other
mammals.

II. The general diagnostic features are :—in the malleus, constant, firm bony ankylosis
to the tympanic bone through the medium of the processus gracilis, with ill-development
or complete suppression of the manubrium; in the incus, great development of the
stapedial crus ; in the stapes, thickness of the crura contracting or obliterating the aperture.
This latter bone always fits firmly into the fenestra ovalis, to which, however, it is in
no species constantly ankylosed.

III. The genus Balena possesses undoubtedly the most generalized ossicula among
all the Cetacea, particularly in the case of the incus. The malleus has the least-modified
form of manubrium, the whole bone distantly resembling that of certain large Seals
(Macrorhinus, Stenorhynchus), as does also the stapes. |

IV. The malleus of Neobalena is the next least specialized. That of Megaptera is
much more modified in regard to the aborted condition of the manubrium.

V. Both in the malleus and incus Balenoptera is far more modified than any of the
above genera*. ‘The incus is quite Delphinoid, unlike either the Balena or Physeter
type. The malleus is even more modified than in the Dolphins ; its manubrium is of more
tubercular form, and the whole bone is more flattened.

VI. In the Physeteride the malleus is extremely modified. The tubercle projecting
from the head in other Cetacea is here obsolete; the manubrium and processus muscu-
laris are reduced to two small spines. That representing the manubrium is almost sup-
pressed in Physeter, Hyperoodon, and Mesoplodon, but is well developed in Berardius.
The head is largest in Berardius; in Physeter the articular surface is very extensive.
The incus is intermediate in type between the same in Balena and Delphinus. In the
the Ziphioid Whales the short crus is long and of peculiar shape. The stapes differs a
little in character from that of the true Dolphins. ,

VII. In the Dolphins the malleus is never so modified as in the Ziphioid Whales: the
tubercle projecting from its head is pointed, not rounded as in Balenoptera; and the
seat of insertion of the tensor tympani muscle is indicated by a roughened, depressed
surface. The manubrium may be quite absent (Delphinapterus) or reduced to a small

* See Appendix to this scction.
3Q 2

464 MR. A. H. G. DORAN ON THE MORPHOLOGY

spine, depressed against the side of the ossicle (Globiocephalus, Lagenorhynchus, Pho-
cena); or the spine may be well developed and directed downwards, away from the rest
of the malleus (Orca, Pseudorca). Part of the process from the membrana tympani is
inserted along a narrow groove on the tubercle. Monodon differs from other Dolphins
in the characters of its articular surface, and the length of the head and tubercle of the
malleus.

VIII. All the Dolphins agree in the characters of the incus, which has an ill-developed
body and processus brevis, but a very large stapedial crus. The stapes, too, is very
similar in character in all these Cetacea.

IX. Platanista differs remarkably from the true Dolphins in the great length of the
process from the head of the malleus, as well as in minor particulars. Such distinctions
are hardly perceptible in Inia.

Appendix to Remarks on Cetacean Ossicula—The malleus of a foetal Balenoptera
sibbaldii in the Museum of the University of Edinburgh bears a long process in front of
the head, with the same homologues, apparently, as are seen in Megaptera. Whether
or not this process is arrested, whilst the head increases during the growth of B. sib-
baldii is a question only to be solved by the examination of the malleus in an adult of
this species. If the process remains long, then it is truly remarkable, since it allies this
Balenoptera to the genus Megaptera, and separates it from B. rostrata and B. huttoni.
The incus from the same footus is of the Balena type, as the body is very well deve-
loped, the long crus by no means so large as in Balenoptera rostrata. For this observa-
tion I am indebted to the kindness of Prof. Turner. I may further add that the malleus, in
the skeleton of Macleayius (Hubalena) australis at the British Museum, is of the Balena

type.
THE OssICULA OF THE SIRENIA.

On account of their large size and conspicuous peculiarities, the ossicula of the Dugong
and Manatee have already become familiar to most comparative anatomists; and,
through their great durability, even the same bones in Rhytina and Halitherium have
been described respectively by Claudius * and Krauss +; but the most prominent cha-
racters which have rendered them interesting to zoologists are due to their structure and
consistence, the homologies of their various segments and processes being, on the whole,
much easier to distinguish than is the case with the ear-bones of the Cetacea.

he malleus of the Dugong (Halicore australis, Pl. LXIII. figs. 26, 27) is a heavy
pone, weighing on an average 40 grains, and measuring about an inch from the root of
the processus gracilis to the tip of the manubrium; the largest specimen never attains
the size and weight of the malleus of species of either Manatus or Rhytina, though, from
Krauss’s account, it appears to be longer than in Halitherium schinzi.

It is important to bear constantly in mind the position of this bone in the Sirenian tympa-
num. In ordinary mammals the head of the malleus lies far back in the tympanic cavity,

* «Tas Gehérorgan von Rhytina stelleri,’ Mém. Acad. Imp. des Sciences de St. Pétersbourg, 1867.
+ Leonhard u. Bronn’s N. Jahrb. 1862, p. 385.

OF THE MAMMALIAN OSSICULA AUDITUS. 465

so as to be often invisible from the external meatus; the processus gracilis stretches well
forwards from the rest of the malleus ; and the articular surface is placed almost wholly pos-
terior. In the Cetacea, on the other hand, the head of the malleus is much less posterior in
position, the processus gracilis runs from it directly outwards to join the tympanic bone,
and the facets look inwards. In the Dugong and Manatee the malleus holds an inter-
mediate position, as may be seen by examining a skull of Halicore in the College
Museum (No. 2633), where all the left ossicula are in situ. The head lies quite anterior
to the posterior horn of the dense annulus tympanicus, its manubrium projecting directly
outwards from it; the processus gracilis runs as completely forwards as in terrestrial
mammalia; but, again, the articular surface is as internal as in the Whales and
Dolphins.

The massive head of the malleus of Halicore australis is of irregular shape, about as
long antero-posteriorly as vertically, and slightly flattened from within outwards. Its
uppermost portion is wide, slightly convex, and does not project above the upper facet
as in most Mammals. The articular region lies entirely on the very broad inner aspect
of the ossicle. It is made up of an upper facet and two lower, which latter represent
the single inferior facet in other mammals, the extero-inferior segment of the articular
surface of the human malleus; they form almost a right angle with each other, and an
obtuse angle with the upper facet, a very distinct groove dividing them. The superior
facet is reniform, with its long axis vertical, and its concavity forward; its surface is
distinctly convex from above downwards. The smaller inferior facets* are oval; and
their edges generally coalesce along the high ridge which divides them, and whence
they slope. On the outer aspect of the malleus is a very deep groove running forwards
and slightly downwards; its upper border is thick and prominent; its lower is formed by
the root of the manubrium. This projecting upper border is homologous with the sigmoid
ridge on the neck of the human malleus, as may be seen by laying an ossicle of Homo
and of Halicore side by side, with the manubrium forwards. Hence it must represent
the solid curved neck of the laminar type of malleus in the Carnivora, Ruminants, and
other Mammals+. Completely in front of the malleus of the Dugong is the processus
gracilis (Pl. LXIIT. figs. 26, 27), which is broad at its root, but narrows rapidly.
Owing to the whole ossicle being placed unusually forwards this process is far shorter
than in terrestrial Mammals. Its upper surface is smooth and convex, being con-
tinued from the upper surface of the head; its lower is flat and very short, separated
by a deep horizontal groove from a prominent tubercle on the front of the head below
it. The processus muscularis is a blunt tubercle, looking forwards from the inner
aspect of the malleus, and is continued backwards and upwards as a very promi-
nent convex ridge, which is lost on the head of the malleus behind the inferior facets,
near the ridge on the anterior aspect already described. Strictly speaking, the whole
must be considered as the processus muscularis, having its root on the true neck (the

* These evidently represent the inferior facet of other animals, the convex ridge being here much exaggerated,
even to the point of dividing that facet into two.
tT See the account of the human malleus, antea.

466 MR. A. H. G. DORAN ON THE MORPHOLOGY

outer and anterior projection), as in many other animals. The manubrium stands out
from the head completely external; and its point is directed rather backwards: it is
separated from the rest of the malleus above by the groove on the outer aspect of the
ossicle; below it is blended with the base of the ridge continued from the processus
muscularis. It is wide at the root, and triangular in form; its upper border is very
broad, particularly towards the groove, and sharply bordered from the sides ; externally
it runs into the outer aspect by forming a prominent bold convexity, instead of a pro-
cessus brevis; the outer aspect is also sharply defined, its borders being continued from
above, and towards the extremity it is spatulate: in some specimens this dilatation is
very marked, in others hardly perceptible. The sides of the manubrium are flattened,
and somewhat concave near the root; the inner border is rather sharp, and quite concave ;
it blends with the head as aforesaid below the processus muscularis. It is wide at the
root, and triangular in form: its upper border is very broad, particularly towards the
groove, and sharply bordered from the sides; externally it runs into the outer aspect by
forming a prominent bold convexity instead of a processus brevis; the outer aspect is
also sharply defined, its borders being continued from above, and towards the extremity
it is spatulate; in some specimens this dilatation is very marked, in others hardly per-
ceptible. The sides of the manubrium are flattened and somewhat concave near the —
root; the inner border is rather sharp, and quite concave, it bends with the head, as
aforesaid, below the processus muscularis.

The incus of Halicore (Pl. LXIII. figs. 28, 29) is singularly placed, its “ processus
brevis” being absolutely superior in position to the rest of the bone, instead of being
postero-superior, whilst the stapedial crus is equally inferior; the facets look directly
outwards on account of the internal position of those on the malleus. The body is twice
as wide as it is deep; its inner aspect, which is wide, smooth, and convex, fits into a
depression in the petrous bone; its articular surface consists of a concave reniform facet,
corresponding to the upper one on the malleus, separated by a sharp projecting angle
from two oval lower facets, which look towards each other, and are divided by a deep
eroove, into which the projecting angle between the lower facets on the malleus is locked
in the recent skeleton. The posterior of the two lower incudal facets is borne on a very
prominent tubercle projecting from the outer and more posterior surface of the body of
the incus—a most characteristic feature, by which this ossicle in Halicore may be at once
distinguished from the same in Manatus, and apparently in Rhytina and Halicore.

The processus brevis is very short, and lies above and somewhat anterior to the rest of
the ossicle; it is firmly ankylosed to the petrous bone; but being thin and brittle, the
whole incus is generally detached if the skull is dropped on the ground, or struck
violently after maceration. The processus longus arises far from the outer crus on
account of the great width of the body, to which it lies inferiorly and slightly posteriorly.
It is short, slightly curved, flattened extero-internally, and of almost equal thickness
throughout ; the oval disk at the extremity is generally separated from the rest of the
crus by a faint groove, so as really to represent the os orbiculare, the existence of which
in Halicore seems to be denied; as a rule, the larger the mammal the less developed is
its Sylvian apophysis,

OF THE MAMMALIAN OSSICULA AUDITUS. 467

The stapes of the Dugong (fig. 30) measures about half an inch in length, and is of
very solid consistence; as the crura diverge but little, the base is narrow; the anterior
extremity of the latter and of the fenestra ovalis is placed much lower than the posterior,
although, after examining five skulls of Halicore in the College Museum, I do not find
the long axis of the base of the stapes or of the fenestra vertical, as has been asserted ; the
same remark applies to Manatus. 'The head is well developed, and the articular surface
is concave; the site of insertion of the stapedius is generally indicated by a slight depres-
sion. The anterior crus is half the thickness of the posterior, and both are rather
crooked; the aperture is circular, generally just large enough to admit a small sewing-
needle, but in some specimens it is of the calibre of a No. 1 English catheter. The base
is convex, most so posteriorly, and is narrowest towards the front ; it never appears to be
ankylosed to the fenestra ovalis, or even tightly fitted imto it, as in the Cetacea. The
broad well-formed head and comparatively thin anterior crus somewhat remind the
observer of the stapes of Bos; but in the other ossicula of Halicore and the other Sirenia
few, if any, distinct analogies to the same bones in any particular order of Mammalia can
be made out.

Since the extinction of the Rhytina the American Manatee (W/. americanus) has borne
the largest and heaviest malleus known in any animal, as Hyrtl has already stated.
Indeed, to judge from Claudius’s account of that ossicle in Rhytina, of which the head,
he remarks, is of about the size of an ordinary hazel-nut, it hardly can have exceeded,
as a rule, the malleus of a large Manatee, though I have never seen the processus
gracilis of the latter “at least 13!" in diameter,” which is the case in the specimen from
the extinct Sirenian he describes. The specimen in the collection of ossicula in the
College Museum (Pl. LXIII. figs. 21, 22), from a skull of WZ. americanus, in the Com-
parative-Anatomy series, measures over an inch in length, and weighs over sixty-one
erains; the head is quite as large as a middle-sized hazel-nut. This bone, as well as the
other ossicula, lies, in the recent skull, in the same position as has already been described
in Halicore; but it offers many points of difference in its structure from the malleus of
the latter animal.

The head is much longer antero-posteriorly than vertically. All the facets are larger ;
and the upper, though reniform, has its concavity backwards, and its surface is distinctly
concave above. In the College specimen there is a prominent peg-like process (Pl. LXIII.
fig. 21) projecting immediately below the inferior border of this facet, bearing supe-.,
riorly another small facet; not a trace of this can be seen in any specimen from -
M. senegalensis in the same series. The lower facets in Manatus americanus are of
much the same form as in the Dugong; but at the top of the prominent angle their
borders do not blend, but are nearly a line apart. The outer aspect of the malleus is
very wide and quite convex; the groove is very shallow anteriorly, and quite effaced
behind, in which direction, however, is a very sharp and prominent ridge analogous to a
similar projection in Halicore, the homologies of which have already been discussed.
The processus muscularis is a stout incurved process, continued backwards as a con-
spicuous elevation towards the former ridge, precisely as in the Dugong.

468 MR. A. H. G. DORAN ON THE MORPHOLOGY

The manubrium differs in form from that of Halicore. The upper border is very sharp
and narrow; it forms a semicircle, and projects against the membrana tympani in the
live subject; the outer border is short, and its margins, united above, diverge but little
towards the extremity ; its surface is concave vertically, and slightly convex horizontally.
The inner border is much blunter than the outer in Manatus.

The processus gracilis is not relatively so well developed as in Halicore, and projects,
as a fragile splinter of bone, from the massive head; it is firmly ankylosed to the
tympanic ring; and, from its very nature, the whole malleus is easily broken off from
the ring.

The incus (figs. 23, 24) is placed in the tympanic cavity precisely as in the Dugong,
the processus longus lying above the body, and the other crus running downwards from
it. The body is even shallower and wider between the crura than in Halicore ; its inner
surface is less extensive and less convex, and much overhung by the upper facet of the
articular area, which facet is very wide, reniform, and faintly concave; the two lower
facets look towards each other, and are separated by a deep groove, into which the cor-
responding prominence between the opposing surfaces on the malleus are fitted during
lifetime. There is not nearly so marked a prominence behind the upper and more
posterior of these facets in the Manatee, like that which projects so conspicuously on the
posterior side of the Dugong’s incus. The processus longus is rather long, slightly
curved, and flattened extero-internally; it widens out towards the point, where it is
ankylosed to the petrous bone. The processus brevis is very short, stout, and curved;
the articular surface for the stapes is very wide, almost circular, and nearly plane; a
faint trace of a rim can be seen above it, separating it from the crus, as the homologue
of the os orbiculare.

No stapes of this Manatee exists in the College collection; but that of I, senegalensis,
of which there are several examples, will be shortly described. The malleus from an
adult of the latter species is rather smaller than in the American kind ; the upper facet
is not so wide, and there is no peg-like projection internal to it. The processus muscu-
laris is extremely stout and prominent; but the manubrium is not so broad as in
M. americanus.

The incus of Manatus senegalensis very closely resembles that ossicle in the American
Manatee; the processus longus is of similar form and length; the articular surface is
slightly different, in accordance with the characters of that surface on the malleus of the
African species; hence the upper facet is smaller, and does not overhang the inner side of
the body as much, and the two lower facets gape wider than in the American species.
In none of the existing Sirenia does the incus reach the size of that of the Seal
(Macrorhinus proboscideus), where it is larger than in any other animal.

But as is the case with the malleus, so is the stapes of Manatus (fig. 25) largest among
mammalian stapedes. It is over an inch in length, and weighs about one drachm; it is
of the dense consistence of the other ossicula in this Order. In shape it differs from the
stapes of Halicore, being over twice as long as it is wide. The head is well developed,
and bears an almost cireular articular surface. Both crura are long and hardly

ON THE MAMMALIAN OSSICULA AUDITUS. 469

divergent; both are crooked; but the anterior is the thinner and straighter; the
posterior is very broad inferiorly, often bulging prominently just above the base. The
aperture appears as an oblique slit, a quarter of an inch long from above; but on the
lower aspect it is more circular: in a large specimen it is of the calibre of a darning-
needle. The sides of the crura bounding it are slightly concave, but not absolutely
grooved. The base is rather narrower than are the crura a little above it, but it is
wider than the head; it is convex towards the vestibule; and its margin is generally
rather concave above, in accordance with the form of the fenestra ovalis in this animal.

According to Claudius *, the malleus and incus of Rhytina closely resemble those of
Manatus. In the work already referred to, these ossicles are figured and described ; and
that anatomist does not name any point in which the extinct differs from the above-
named existing Sirenian as far as these bonesare concerned. ‘The inner border of the
manubrium, he says, is shorter, narrower, and sharper than the outer (that is, the upper
and outer surface of our description); but he does not state whether the latter is broad,
with sharply defined margins, as in the Dugong, or sharp and narrow, as in the Manatee.
The incus is placed, according to his account, precisely as in the existing species; and
the short crus is ankylosed to the petrous bone; the processus longus is described as
long and strong, though in the plate it does not appear to be longer than in Manatus.

Dr. Krauss + has had the good fortune of procuring and examining all three ossicula
of Halitherium schinzi, which he figures both separate and articulated. From the
letterpress and the plates it appears that the malleus is intermediate in form between
the Manatee’s and Dugong’s. The body is figured as short antero-posteriorly, asin Hali
core ; but the manubrium is represented more of the Manatws-type; the articular surface_
though not distinctly described or figured, must, judging from the incus, be as in the
latter animal; the processus gracilis is rather long, thin, and brittle. As for the incus,
it is both described and figured as much more like that of Manatus than Halicore; the
articular surface is represented as large as in the Manatee, and similar in its facets; more-
over there is no prominent tubercle on the outer aspect of the body, backing up one of
the facets, as I have observed and described in Halicore. 'The stapes is described as
very similar to that of Halcore, “‘ whilst that of the Manatus is much larger, thicker,
- and rounder.” The illustration confirms this statement, but represents the crura as
rather longer than in the Dugong; the anterior crus is much the thinner ; the head is
broad; and the whole bone decidedly, as far as one can judge from a plate, approaches
the quadrilateral type of the larger Ruminants.

In conclusion, it may be said of the ossicula of the Sirenia :—

I. That their dense structure and clumsy form, consistent with the characters of the
whole skeleton, distinguish them at a glance from their homologues in all other Orders.

IT. That their modifications lie much more in their peculiar general conformation than

\

* Mém. Acad. Imp. des Sciences de St. Pétersbourg, 1867.
t As already quoted in the Jahrb. 1862.

470 MR. A. H. G. DORAN ON THE MORPHOLOGY

in the suppression or peculiar development of any of their processes, which are so remark-
able in the Cetacea.

III. Not only do they differ clearly from the Cetacean ossicula, but their modified
general structure may be said to mask or obliterate any homologies to the ossicula in other
mammals, which otherwise might be observable—the only exception being the approach
to a quadrilateral type of stapes in Halicore, apparently more marked in Halitheriwn,
which, whatever be its value, cannot fail to remind the anatomist of the bovine form of
that ossicle.

IV. Halicore differs distinctly from Manatus in the antero-posterior shortness of the
body of its malleus, in the smallness of its articular facets, in the form of its manubrium,
in the tubercle on the outer and posterior aspect of the body of the incus supporting the
articular surface, and in the shorter incus with more divergent crura. —

V. In Rhytina the malleus and incus appear very like those of Manatus. But in
Halitheriwm the characters of the ossicula are extremely intermediate—the incus being
most like that of Manatus, the stapes like the Dugong’s, and the malleus possessing
features seen in both.

VI. The distinguishing feature by which WM. americanus may be known, by its ossi-
cula, from IM. senegalensis, is a slight difference in the characters of the malleo-incudal
articulation.

Tab OSSICULA OF THE EDENTATA.

As is the case with the rest of the skeleton, the ear-bones in this Order exhibit con-
siderable variety of type; the characters of the malleus and incus are very general;
whilst the stapes assumes the Sauropsidan form in one group, and approaches it in
several others.

In the two recent genera of the PayropHaGa the malleus offers points of difference too
marked to be overlooked. ‘This ossicle in the Ai, or Three-toed Sloth (Bradypus tridac-
tylus, Pl. LXTV. fig. 1) has a rounded, well-developed head, projecting a little upwards, but
more forwards, and rather flattened laterally. The articular surface is deeply cut, the facets
lying almost at right angles to one another ; they are narrow horizontally; the upper is
almost plane, the lower perceptibly convex ; the neck is long, slender, and almost straight.
By laying the malleus of Homo by the side of one from this Hdentate, the identity of the
sigmoid ridge of the former with the much less curved neck, just described, in the latter
will at once become very evident. ‘The processus gracilis forms a very acute angle with
the neck, leaving a narrow space between them, homologous with the lamina, but filled
up with thick and friable rather than lamellar bone. There is no trace of any processus
muscularis, either on the neck or the handle. The manubrium forms an angle of over
150° with the rest of the malleus; it is of medium length, very thin, and compressed
laterally, and broad at the base, where, externally, is seen a sharp angle, representing the
processus brevis. The external aspect is bordered from the sides by sharp edges, as in
the Carnivora, &c., but very narrow; the extremity is slightly spatulate, and perceptibly
recurved.

OF THE MAMMALIAN OSSICULA AUDITUS. ATL

Thus the malleus of Bradypus has very general characters. It cannot be considered
a low type, like that of any marsupial; but its affinities are rather in the direction of
Mammals undeniably higher than the Edentata, such as the higher Primates, the
Ichneumons, and the Horses, not from any positive characters, but rather from the
absence of salient modifications so prominent in animals much nearer to them in other
respects.

The incus (Pl. LXIV. fig. 1) has a rather shallow body, with proportionally large
crura. The processus brevis is very stout and rather long, sharply pointed and very
divergent from the body; the processus longus is well developed and broad. I have
not observed any bifurcation on its inner surface in three incudes from different
individuals in the College Museum, although Hyrtl names that peculiarity in describing
the incus of Bradypus. The Sylvian apophysis is small, but very distinct, being attached
to the crus by an extremely narrow pedicle. The affinities of this ossicle are very
uncertain ; the stout divergent processus brevis recalls the type of medium-sized Bovide ;
but in the Sloth the incus is smaller in proportion to the malleus than is the case with
the Ungulata.

The stapes in Bradypus tridactylus (fig. 1) is a very interesting bone, remarkable for
having occasional (not constant) features which relate it very decidedly to the Didelphia
and Sauropsida. In one pair belonging to the College series the head is moderately
developed, the crura are stout, rather long, and but little divergent, leaving between
them a narrow but very distinct aperture. In another pair in the same collection
the right stapes has no aperture, whilst the left has a minute foramen between its
erura. In all, the base is very broad, recalling its form and proportions in the Marsupial
stapes.

Yet Hyrt! remarks (op. cit.), “It is most remarkable that Bradypus tridactylus has
only one crus, which is broad; Bradypus didactylus (sic), on the other hand, has two
very thick crura, which, on that account are separated by a space hardly the width of a
hair. The simple crus of the stapes of B. tridactylus is very thin and transparent, whilst
the stapes in B. didactylus is massive and conical.” He then remarks that he has only
examined the stapes cf B. tridactylus in an embryo.

From Hyrtl’s observation, and from the distinct and complete fusion of the crura in
the stapes on one side only in the specimen already referred to, we see in the Ai a
singular wavering between the higher and lower type of this ossicle in the Vertebrata
that possess it. The intercrural aperture in none of the College specimens is as large as
it is in the columella of the Golden Hagle (Aquila chrysaetos), so that the presence of that
space must not be considered unreservedly as a tendency to higher type.

The ossicula of the Unau (Cholepus didactylus) differ from tlie above in some respects
(see Pl. LXIV. fig. 2). The head of the malleus is even more developed, and is some-
what produced forwards, as in the Hystricide. The articular surface is similar; the neck
is rather longer, but more curved, assuming the sigmoid form of the ridge on the human
malleus. The lamina is similarly represented by a narrow tract of rather thick bone.
Indeed the malleus of both the Sloths is distinctly intermediate between the non-lami-

3R 2

472, MR. A. H. G. DORAN ON THE MORPHOLOGY

nated form of Homo and the broadly laminated type of the Carnivora and Artiodactyla*.
The manubrium of Cholepus is longer than in Bradypus, and forms little more than a
right angle with the neck; this alone gives the whole malleus a different aspect from
that of the Ai. The processus muscularis is represented by a distinct tubercle on the
inner side of the root of the manubrium; the latter is even broader than in Bradypus.

The incus of Cholepus does not differ from that of the Ai; the Sylvian apophysis is
of the same character.

The stapes of the Unau has even less divergent crura than that of Bradypus; ‘the
aperture is either very small or quite obliterated: these two conditions may be seen in
different examples in the College collection.

Turning to the ENromopHaGa, the ossicula in the genus Dasypus are larger than in
the Sloths. In the College collection all three in Dasypus villosus are absolutely
larger than in Bradypus.

In the fine malleus of the 6-banded Armadillo (Dasypus sexcinetus, P|. LXIV. fig. 4)
the head is well formed and globular, without being produced forward or in any par-
ticular direction. The articular surface is deeply cut, the facets lying at right angles to
each other; the upper is very wide and nearly plane, the lower is barely half its area
and more convex. The neck is short and bent; the processus gracilis is arched towards
the body of the ossicle, and then curved downwards and forwards; the space thus formed
is narrow, and occupied by a true lamina of really lamellar bone. The processus mus-
cularis is a bony tubercle on the inner side of the neck, close to the root of the handle.

The manubrium is long and straight; it forms an angle of about 140° with the neck ;
it is compressed laterally, but not very broad at the base. A true processus brevis
exists—the angular projection at the external aspect of the base being considerably
everted, and hence is not merely the simple apex of an angle formed by the upper edge
of the base and the outer side of the manubrium. This outer aspect, as in Bradypus, is
very narrow, yet distinctly bordered from the sides; the extremity is very slightly
spatulate and but little recurved. In the well-formed head, decurved processus gracilis,
narrow lamina, and long manubrium this malleus recalls that of the Horse (Hquwus).

The body of the incus (Pl. LXIV. fig. 4) is shallow and broad; the processus brevis
(which lies much more superiorly than posteriorly) is short and sharp-pointed, it
diverges very little. ‘The processus longus is thin, long, and very divergent; the pro-
cessus Sylvii is absolutely sessile, and hardly separated in the adult by any trace of a
groove; but a distinct channel is to be seen in incudes from new-born Armadillos. The
characters of the crura very plainly and completely divide Dasypus from the Sloths.

The stapes is a large bone of very light and elegant form (fig. 4). The head is long
and flattened, with a small tubercle for the insertion of the stapedius. The crura are
long, straight, and conspicuously slender ; they are well grooved towards the aperture,
which is very wide. The base is broad horizontally, but not very deep vertically ; it is
distinctly convex towards the vestibule.

* Refer to the remarks on the human malleus (wnted), particularly the description of the sigmoid ridge and its
homologies.

OF THE MAMMALIAN OSSICULA AUDITUS. 473

In every respect this stapes approaches the type of the higher Mammalia, as does that
ossicle in the Ant-eaters and Orycteropus, contrasting strongly with the Sauropsidan
stapes of Manis and the tendencies in that direction already described in the Sloths.

In Dasypus villosus and D. minutus the ossicula are almost identical with the same in
D. sexcinctus. The stapes in the Hairy Armadillo is even more slender than in the six-
banded species ; the processus muscularis is more prominent in the malleus of D. mimutus
than in the larger species.

In the Peba Armadillo (Zatusia peda, Pl. LXIV. fig. 3), the ossicula resemble greatly
those of Dasypus. The processus muscularis mallei is obsolete in the College specimens ;
the crura of the incus are of similar character ; the stapes is of slender form with a wide
aperture. I have not found the head of this ossicle to be unossified in any young or
adult Zatusia, as Hyrtl asserts of T. hybrida; but that portion is readily broken
off from the fragile crura. The malleus in the new-born Zatusia (fig. 7) has an ill-
developed head and a similar manubrium to that of Priodon (fig. 5). It is remarkable
that whllst the malleus of the adult Zatusia or Dasypus resembles that of Hquwus, the
same ossicle in the young of those genera or in the adult Priodon reminds the observer of
Rhinoceros. ‘This is at least remarkable when we remember the relationship of the
ossicula of the adults in some of the Ungulata to those of the young in others.

In Priodon gigas (Pl. LXIV. fig. 5) the malleus has a much less developed and less
globular head than in the above-described Armadillos: the articular surface is of similar
form; the neck is thinner, longer, and straighter. The lamina is rather wide, and pro-
longed a little on the strongly decurved processus gracilis, which is joined by a styliform
process from the front of the head, which has its homologue in the Carnivora, Rodentia*,
Ungulata, Cetacea t+, and most other Orders. The processus muscularis is wanting.
The manubrium forms little more than a right angle with the neck; it is much more
slender than in Dasypus, and bent outwards on the rest of the malleus; there is no pro-
cessus brevis, but, on the contrary, a very blunt projection at its site. The ill-developed
head, peculiar lamina, and the form of the manubrium give it a general resemblance to
the malleus of Rhinoceros.

The incus of Priodou (fig. 5) has a shorter processus brevis than in Dasypus; the long
crus is much stouter, and has a notch or fork on its upper margin. Hyrtl has observed
a similar appearance in the Sloths. The processus longus bears a distinct, sessile, and
very elliptical Sylvian apophysis.

The stapes (fig. 5) has much stouter crura than in Dasypus or Tatusia, with a small
intercrural aperture. The base is rather narrow; and in no wise does this ossicle bear
any resemblance to that of Bradypus or Manis.

In the Three-banded Armadillo (Volypeutes tricinctus, fig. 6), the malleus of the
adult much resembles that of Priodon; but the lamina is very wide, so that the ossicle
is modified from a low type in the same direction as in the Carnivora, Artiodactyla, and
Muride. The neck remains straight, and the outer angle of the base of the manubrium

* See remarks on the malleus of the Musquash (Fiber).
+ Description of the malleus of Balena, anted.

AGA, MR. A. H. G. DORAN ON THE MORPHOLOGY

blunt, as in the Giant Armadillo; but there is a distinct tubercular processus mus-
cularis in the same position as in Dasypus.

The incus of Tolypeutes resembles that of Dasypus and Tatusia much more than
Priodon, the processus longus being long, slender, straight, and very divergent.

But the stapes from the adult specimen in the College Museum is of more Sauropsidan
type than in any other Edentate except Manis. ‘The head is long and narrow; and the
slender crura keep together till close to the foot-plate, where they diverge, leaving a small
aperture betweenthem. The base is broad vertically as well as horizontally. It is the
manner of divergence just described, and not merely the smallness of the aperture, which
relates this stapes of Tolypeutes to obviously lower forms. As Manis in its stapes with
absolutely fused crura allies itself to Dasypus, Phalangista, Perameles, the Ornitho-
delphia, and most of the Sauropsida, so does Tolypeutes tend, in the character of the
aperture in that ossicle, to Macrapus, Didelphys, and some Birds.

In the squamate Edentata all three ossicles present positive points of peculiarity.

The malleus of Manis dalmanii (Pl. LXIV. fig. 8) has a large head, much flattened
antero-posteriorly ; the front is but slightly convex, and it does not project beyond the
upper margin of the articular surface. This latter has a very prominent border, and in
form is not unlike the same articular surface in Man and the Anthropoid Apes. It is
placed very obliquely, so that its outer extremity * lies much higher than the inner
(fig. 10); it is also divided into two facets, one superior and internal, the lower
being the more external. Hach presents a vertical convexity; and they slope towards
each other, so that the whole articular surface appears concave when viewed late-
rally. But this surface is far wider in proportion to the entire head than it is in
Homo, nor does the head in Manis project prominently above it as in Man; moreover
the division into facets is even less distinct +. The neck is very short and curved; the
~ lamina is of medium width, and formed of not very thin or transparent bone; there is no
processus muscularis. The manubrium forms an angle of about 130° with the neck, and
resembles that of Dasypus; a distinct processus brevis exists; the manubrium is rather
broad at the base, and much flattened laterally ; the extremity is distinctly recurved.
The character of the articular surface and the form of the head distinguish the malleus
of Manis alike from the same in the Sloths and the Dasypodide.

The incus (fig. 8) lies in the skeleton with its articular surface looking directly
forwards; it is rather broader than deep in the adult Manis daimanii. The postero-
superior part of its body (which is really absolutely superior in Janis) is well developed
and bulging, but not so markedly deeper than the other part of the body over the pro-
cessus longus as is the case in some species of Manis. The crura are very short, and
diverge but little ; the stapedial crus bears an ill-developed elliptical Sylvian apophysis,
which is pedunculated.

The stapes of the Pangolin (Manis) is most remarkable, being absolutely Sauropsidan ;

% This is actually the wpper extremity in Manis; but being the homologue of the outer in most mammals, that
term is retained, as in describing a similar condition in the human malleus.

+ On the subject of the facets, I must again refer to my observations on the homologies of the articular surface in
the human malleus.

ry

OF THE MAMMALIAN OSSICULA AUDITUS. 475

the head is very ill developed; the column does not exhibit the least trace of any division
into crura ; the foot-plate is almost circular, and in diameter almost equals the length of
the column. <A tendency to this form of the base has already been described in Bradypus ;
and the gradations from the completely bicrurate stapes of Dasypus, through the form
seen in Tolypeutes, to that in Manis are most instructive to observe.

In Wanis tricuspis, M. longicauda, and WZ. pentadactyla, the malleus is identical in form
with that of I. dalmanii; but in the incus (Pl. LXIV. fig. 9) the portion of the body over
the processus brevis is extremely high, so that the whole ossicle assumes a form seen in
Mycetes and Pithecia; the processus brevis is thinner and slightly more divergent than
in YW. dalmanii, though both processes are as short. The stapes has a shorter column,
without any trace of division into crura, and it is absolutely shorter than the diameter
of the foot-plate, which is not so circular as in the species first described.

In the Great Ant-Eater (Myrmecophaga, P|. LXTV. fig. 11) the malleus has a large head,
very much flattened laterally and produced forwards; its upper portion, very wide antero-
posteriorly on account of the lateral flattening, does not project at all above the level of
the articular surface, which is rather deeply cut, and much wider vertically than from
without inwards. The lower facet is the most concave, and is divided from the upper
by a very faint groove. The neck is long and almost straight; there is a very narrow
lamina; the manubrium forms an angle of about 150° with the neck. The handle is
long and slender; there is no processus brevis, but a very blunt angle in its. place; the
extremity is very slightly spatulate and scarcely recurved. The sides are flattened, and
not broad near the base ; there is no trace of any processus muscularis. This malleus of
the Great Ant-eater bears a certain resemblance to that of Priodon, from which it may
at once be distinguished by the form of the head.

The body of the ineus of Myrmecophaga (fig. 11) is almost square, although rather
broader than deep. The processus brevis is moderately long, divergent, and blunt-
pointed; the processus longus is well developed without being very stout, and bears a
distinct elliptical Sylvian apophysis on a broad pedicle. The whole bone is large in
proportion to the malleus.

The stapes (fig. 11) has an ill-developed head and long very slender crura, with a
wide aperture between them. The base is thin, but very broad vertically as well as
horizontally. Thus this bone in the Great Ant-eater approaches the type of Dasypus
rather than Bradypus, Tolypeutes, or Manis. The vertical breadth of the foot-plate
(that is to say, the tendency to roundness in that portion of the ossicle) is the oniy
approach towards the Sauropsidan columella, the crura tending towards much higher
types.

In Tamandua tetradactyla (Pl. LXIV. fig. 12) the head of the malleus, though flat-
tened laterally, projects much more above the upper margin of the articular surface
than in Myrmecophaga, so as to be very convex above. The neck is a little more
curved, and the lamina wider; the manubrium is very similar in form. The incus has
an even squarer body ; its processus longus bears a similar Sylvian apophysis.

In Cyclothurus didactylus (P\. LXIV. fig. 18) the head of the malleus is as prominent
_ above asin Tamandua; the neck is shorter, and the manubrium is much broader at the

A76 MR. A. H. G. DORAN ON THE MORPHOLOGY

base, than in the two larger Ant-eaters. The whole bone somewhat resembles the same:
ossicle in Bradypus. The incus has a shallow body ; and the end of its stapedial crus bears
a very elliptical and sessile Sylvian apophysis, so narrow vertically as to be almost
rod-like. The stapes in a young specimen in the Collection has no trace of any division
between the crura, and in general resembles that ossicle in the Sloths.

In the Ant Bear (Orycteropus ethiopicus, Pl. LXIV. fig. 14) the head of the malleus
is ill-developed, the articular surface is shallow and wide vertically, both facets are but
faintly convex. The neck is long and curved ; it bears internally and nearly one line
distance from the root of the handle a prominent tubercular processus muscularis. There
is a wide and transparent lamina. The manubrium is very long, and forms a right angle
with the body; it is rather broad at the base, and presents an angle corresponding to the
processus brevis ; the extremity is very little dilated or recurved. In the characters of the
head the malleus of the Aard-vaark (O. capensis) resembles Priodon ; but the greater
length of the manubrium, the broad lamina, and the very distinct processus muscularis
distinguish it readily from the latter.

The body of the incus is shallow ; both crura are long and divergent; the Sylvian apo-
physis is almost obsolete, being only separated from the long crus by an indistinet
groove.

The stapes has an ill-developed head, and two long slender crura bowed outwards; a
thin bony lamina is spread between them below, closing the lower part of the aperture,
which otherwise would be wide. The foot-plate is not very wide horizontally, and is narrow
vertically. Hence the stapes of Orycteropus, on account of the latter feature, is of a
still higher type than that of Myrmecophaga.

Reviewing, then, the ossicula of the Edentata as a whole, it may be asserted :—

I. That in the Bradypodide the characters of the malleus and incus are fairly gene-
ralized, whilst the stapes assumes to a certain extent, though never completely, Sau-
ropsidan characters. Cholepus differs from Bradypus in the form of the malleus.

II. That in the Loricata the genera Dasypus and Tatusia present much higher cha-
racters in their ossicula than are found in Priodon and Tolypeutes. In the first two
genera the stapes is of a high type, both as to its crura and its base; in Priodon that
ossicle is more Sloth-like; in Yolypeutes it resembles that of the Kangaroos and other
Marsupials, or even some Birds, but not the complete columelliform type of Manis and
most Aves.

III. Manis has the most positive characters of interest to be found among all Hdentata :
the malleus is more specialized than in the other groups; the incus has peculiar though
less unusual characters; and the stapes is more absolutely Sauropsidan in every respect
than in any other placental mammal.

IV. Inthe Ant-eaters the malleus may be known from that of other Edentata by the
form of its head. The stapes is of as high type, as to the free separation of its crura, as
that of Dasypus; but the base has a tendency to assume the circular rather than oval
form seen in the columella of Birds. Cyclothurus, in its malleus, tends more towards

OF THE MAMMALIAN OSSICULA AUDITUS. ATT

Bradypus and Dasypus than do Myrmecophaga and Tamandua; and its stapes, In one
young specimen at least, approaches much more to the Sloth type.

V. In Orycteropus the malleus, quite unlike that of any Ant-eater, Sloth, or Manis,
approaches to a certain extent that of Priodon, from which it has, on the other hand,
very distinctive features. The stapes is of as high a type as in Dasypus.

VI. That many instances are observed in this Order of the ossicula of the adults of
some genera resembling those of the young of others; especially does the malleus of
Priodon or Tolypeutes, however old and well developed the specimen it be removed
from, remind the observer of the malleus of a new-born or fcetal Dasypus or Tatusia.
The highest degree of development is not necessarily seen in the largest species, as is
at once evident on comparing the malleus of Priodon gigas with that of Dasypus
mmutus.

THE OSSICULA OF THE DIDELPHIA.

The study of the ear-bones of the Marsupials is of necessity peculiarly interesting, as
it is on them that we must depend to a great extent for elucidation concerning the
obscure homologies of the proximal elements of the mandibular and hyoidean arches in
the Mammalia and Sauropsida. Considering how undoubtedly inferior these animals
are to placental mammals, it must be clearly understood that their auditory ossicula
are, on the whole, of proportionally high type; for though these little bones in adults of
this subclass resemble, as a rule, their representatives in the foetus of many higher
animals, still hardly one persistent embryonic feature observable in any Order of the
Didelphia is not also to be found in many highly organized Monodelphia. Indeed, in
large Orders, as the Ruminants, or smaller ones with families very specialized and
different in important features from each other, as the Hdentata, certain genera, as
already shown, retain in a most remarkable and instructive manner characters which
disappear during advance to maturity in genera closely allied to them, where the species
may be either larger or of distinctly higher conformation.

Those families will first be described where the stapes has divergent crura, as this
character is manifestly a high one. But it must here be remarked that the divergence
is slight, and only commences a little above the base-plate; never has any Marsupial
been found to possess a stapes with a wide aperture extending freely in the direction of
the head as in Man, most Insectivora, and many Rodents; nor can examples be found
where, as in the Cetacea, the smallness of the aperture is due rather to thickness and
over-development than to non-divergence of the crura. On the other hand, the inter-
mediate condition seen in this section of the Marsupials exists already in many mono-
delphous mammals, as in Priodon, Tolypeutes, and to a certain extent in Rhinoceros and
the Marmosets, as the author has already indicated. The families where this type of
stapes is found are the Macropodide and the Didelphyide.

Secondly the ossicula of those families will be described where the stapes is absolutely
columelliform, without any aperture or any indication of divergence of crura. Even this
still lower type is represented in higher mammals, namely in Manis. The stapedes of

SECOND SERIES.—ZOOLOGY, VOL, I. 38

A478 MR. A. H. G. DORAN ON THE MORPHOLOGY

many Cetaceans and Seals are in no way homologous with that of this section, for a reason
siven in the last paragraph. This group will include the remaining families of the
Didelphia—to wit, the Peramelide, Phalangistide, Dasyuride, and Phascolomyide.
Among the larger species in these groups a small aperture occasionally exists.

The malleus of an adult Kangaroo (Macropus major, Pl. LXIV. fig. 31) is of light con-
struction, though in no wise so thin and papery as in the Bandicoots. The head is small,
and only moderately developed, as in the majority of laminated mallei in the Monodelphia ;
though much flattened laterally, it is not broad in that sense, nor does it project in any
direction beyond the articular surface. This latter is placed more superiorly than behind,
as in most Hdentates, owing to the position of the incus. It has a raised and prominent
border; owing to the lateral flattening of the head, it is rather narrow, but deeply cut.
Its facets are of the character seen in most of the higher Mammalia, and hence differ
from the type of Man, Monkeys, and Manis. The upper or, rather, the anterior is wide
and almost flat; it is twisted a little more inwards than the lower, which is narrow and
convex; the groove between them is well marked. The neck is rather short and slightly
curved; the lamina is very narrow. The long and slender manubrium forms a right
angle (or even a few degrees less than 90°) with the neck ; the site of the processus brevis
is very obtuse ; and the sides are compressed, though not broad even near the root; there
is a distinctly bordered outer surface, as in Carnivora, Ruminants, and others, very
narrow, and but slightly spatulate near the tip, which is hardly perceptibly recurved. A
very slight trace of a tubercle at the junction of the neck with the root of the manubrium
represents the processus muscularis.

From the front projects the remarkable processus gracilis, already well known to ana-
tomists*. It is much longer than the manubrium, and curved downwards and back-
wards, and also somewhat outwards. It is attached by membrane (or, in some individuals,
actually ankylosed) to the tympanic ring, and is flattened and papery in character, rather
broader below than at the root, but ending in a point; near the body of the malleus it is
separated from the lamina by a thin ridge; and the narrow lamina itself is prolonged on
the anterior edge of the processus gracilis. The anterior part of the head of the bone is
not prolonged on to the front of the lamina, as in most laminated mallei; on the con-
trary, it is separated from the lamina by its very high border 7.

Such is the malleus of Macropus. Let us review its peculiarities, and compare them
with those of the same ossicle in higher mammals; for the more the latter possess cha-
racters in which the Macropodian malleus differs from that of the Primates, the lower,
in that respect, must such Monodelphia be considered.

In the lateral flattening and general form of the head the mallei of Carnivora and
Ruminants, and other Monodelphia where the lamina is well developed, all resemble the
malleus of Macropus; so does that of Myrmecophaga and Tamandua; only in the Ant-

* See Hyrtl, op. cit.; Peters, ‘Berichte der Berliner Academie,’ 1867; his conclusions in his paper for that year on

this subject were admitted to be incorrect by himself in an article in the same publication, 1868. See also, on the
latter, Huxley, Proc. Zool. Soc. 1869.

+ This high border is most marked on the external aspect of the malleus. It is very distinct in all the Kangaroos
and Wombats. (See Pl. LXIV. fig. 31.)

OF THE MAMMALIAN OSSICULA AUDITUS. 479

eaters just named the flattening is greater and the head itself larger. In the narrow
curved neck the same animals resemble the Kangaroos rather than Man and the higher
Monkeys, and differ in having a distinct neck from the Cebide. But as to the lamina,
which is absent in Homo and the Quadrumana and Lemures, it is well developed in the
small-headed malleus of a Carnivore or Ruminant, but ill developed, though existent, in
Macropus. A similar degree of partial development has already been noted in some
Edentata, as in Cholepus or Myrmecophaga; and it is to be remembered that the same
condition may be said to exist in Priodon, whilst Tolypeutes possesses a lamina as wide
as in many Carnivora; so that the latter genus differs from Priodon in one way, as
Dasypus does in a much higher direction, the head of the malleus being almost
anthropoid in development. On the other hand the lamina is again suppressed in Wanis,
and the articular surface is singularly like that of Homo, though the head is ill deve-
loped, and the stapes of a lower type than in Macropus, much lower than in Dasypus.

From the above facts, it appears to me to be justifiable to consider that the narrow-
necked and narrow-laminated type of malleus, as seen in Macropus, is the lowest form,
which has become developed, in higher mammals, into two higher forms :—one where
the lamina is more developed, as in the Carnivora; the other where the neck is stouter
or even merged into the head, as in Homo and the Quadrumana. The latter form
must be considered the highest, not only because it exists in Man, but also because
the former, or laminated type, most resembles the Kangaroo’s malleus in the form
of the head, and in the neck being equally long, narrow, and slightly curved; indeed,
in the very Orders where the laminated type prevails, the occasional partial or complete
suppression of the characteristic lamina is often associated with an increased develop-
ment of the head, and a greater stoutness or shortness of the neck, as I have already
observed in Herpestes and Suricata.

On the other hand, a form of malleus exists in other Monodelphia where not only are
the head, neck, and even the manubrium strikingly like the same in Macropus, and the
lamina of like small development, but the latter is prolonged upon a well-developed pro-
cessus gracilis in a similar manner. This is the case in Rhinoceros and Priodon.

Hence uot only is the WMacropus type of malleus the lowest of the three, but the lami-
nated type is clearly the second or the intermediate form, the large-headed and thick-
necked form being, therefore, unquestionably the highest; indeed it is almost as much
above the laminated as it is above the Macropoid variety. These observations suggest
an easy test of the grade of any mammal when judged by its malleus; and, as a rule,
the other ossicula agree with it.

In a young Halmaturus, measuring two inches without the tail, I have found the
malleus of similar form; but the head is much larger in proportion to the body than in
Macropus*; thé lamina is the same in character and similarly prolonged on the pro-
cessus gracilis.

Supposing, then, that both head and lamina, as existent in the young Halmaturus,
remain in a comparatively arrested state of development, the adult must necessarily

* The same, it is worth recording, is the case in the malleus of the adult Phalanger (Phalangista).

38 2

480 MR. A. H. G. DORAN ON THE MORPHOLOGY

have a malleus of the type of Macropus, Priodon, or Rhinoceros; let the head remain
undeveloped, and the lamina take an extra growth, then the laminated type will be seen
in the full-grown animal; but if the head and neck grow, they must necessarily encroach
on the lamina, and the adult will have a malleus like that of Man.

The foetal malleus of Dasypus or Tatusia, very Macropoid in character, undergoes the
latter change, whilst in Tolypeutes it grows into a laminated hammer, and in Priodon
remains unchanged in the adult. Almost similarly the mallei of the adult Rhinoceros
and Hquus differ, whilst that of the foetal Horse is remarkably like the adult Rhinoceros’s.
Again, the mallei of foetal Ruminants ultimately overshoot the latter animal in type,
but do not ultimately reach the high form of Hquus.

The incus of Macropus (Pl. LXIV. fig. 27) lies in a very similar position to that of
many Hdentates, the processus brevis being almost superior; and the stapedial crus is
directed rather backwards as well as inwards. The body is very small in proportion to
the crura, and deeply excavated by the articular surface. The processus brevis is stout
and blunt-pointed, rather long, and little divergent from the body. The processus longus
is long, and rather slender, it is not much curved, and supports a very distinct almost
circular Sylvian apophysis on a narrow pedicle ; altogether it more resembles the singular
ineus of Perameles than that of any Monodelphian mammal.

The stapes (figs. 27 & 42) has a small head and two slender crura united for halfway
towards the base, where they suddenly diverge, leaving a distinct triangular aperture ;
the outline of the crura is slightly concave. There is frequently a spur-like process for
the insertion of the stapedius-tendon, rather on the inner crus than on the head. The
base, which projects considerably beyond the insertion of the crura, is much more nearly
circular than in most placental mammals; the upper border particularly is very. much
curved.

All these characters are seen almost literally imitated in Tolypeuies. I deem it hardly
necessary to waste time in showing at length how this type is intermediate between the
bicrurate stapedes, with large apertures, as in Homo, and the columelliform type of the
Ornithodelphia; still less is there any occasion to demonstrate which is the lowest, and
which the intermediate form, that being seen at a glance if a Man’s, a Kangaroo’s, and
an Echidna’s stapedes be put in a row. The Macropoid type of stapes is approached, it
must be remembered, in animals as high as the Marmosets.

All the ossicula of other species of Macropus, as well as those of Hypsiprymnus and
Lagorchestes, resemble the same in M. major. The crura of the stapes divide in a
similar manner, and the base is broad and flat; nor has it the least tendency to become
bullate in the smaller Kangaroos, such as Hypsiprymnus murinus (Pl. LXIV. figs. 27,
28, 29, & 30).

The DipELPHYID# will now be described, as they resemble the Kangaroos in that the
stapes is not completely columelliform.

The malleus of the Opossum of the United States (Didelphys virginiana, P|. UX1V.fig.15)
is a little smaller than the same ossicle ina small Hypsiprymnus ; in its characters it closely
resembles that of Wacropus. The head is ill developed, and the articular surface deeply
cut, with facets of the same character as in the family just described. The neck is of the

OF THE MAMMALIAN OSSICULA AUDITUS. 481

same slender build; it is very straight until close to the manubrium, where it curves
suddenly. The lamina is decidedly wider than in any Kangaroo, and prolonged upon
the processus gracilis, forming a curved process applied, as in other marsupials, to the
tympanic ring very closely, though readily detached from the latter even in the oldest
male specimens, without the aid either of a lens or of any great skill in dissection. This
process is of more regular form in Didelphys than in Macropus, being of nearly equal
breadth throughout except at the extremity, which is slightly dilated. No foramen can
be detected in it, as in Hrinaceus, which possesses a somewhat similar but still more
uniformly broad processus gracilis.

The manubrium is even more slender and straight than in Macropus; but at the outer
angle of its base, the site of the processus brevis in Homo, it is peculiarly broad and flat-
tened, projecting towards the neck on the outer aspect. ‘The extremity is almost styli-
form; still there is a distinctly bordered though extremely narrow outer surface*, which
may be seen by a magnifying glass to be slightly dilated towards the tip. This malleus
may be said to be a trifle higher in type than that of the Kangaroo, exhibiting certain
affinities to some Insectivora.

The incus of the Virginian Opossum has a deeper body than in the Kangaroo, with the
articular surface as deeply cut ; the processus brevis is thin, long, and rather divergent
from the body, lying almost vertically above the latter in the recent skeleton. The pro-
cessus longus is very long, and bears at its extremity a circular Sylvian apophysis on a
very long pedunele, which is broad, and fiattened where it joins the extremity of the
apophysis +. A high development of this process is frequent among the Insectivora; but
it is also seen in other families of the Marsupialia, where the peduncle is of similar form.

The stapes has a small head, sometimes bearing a tubercle for the stapedius-tendon.
The slender crura diverge rather higher than in Macropus, so that the aperture is more
distinct and more triangular. The base, which is thin and flat, projects far beyond the
erura, is more oval, and hence not so deep vertically as in the Kangaroo (Pl. LXIV.
figs. 15 & 41).

The stapes of Didelphys is certainly of a higher type than that of Macropus or any
other marsupial, both in the greater divergence of the crura and in the characters of the
base. It much resembles that of some small American Monkeys and of certain Rodents
and Insectivora. Classifying the different types of stapes on the same principle upon
which the mammalian malleus was classified after describing that ossicle in Macropus,
it may be asserted that we find two extreme types:—on the one hand the stapes of
Homo, the Simiidee, and some Rodents (as Arctomys) and Insectivora {, where the crura
are freely divergent and curved with a large aperture; and on the other hand the
completely columelliform stapes of the Ornithodelphia, Phalangistide, and Manidee.

* See the description of the ossicula of the Carnivora, anted.

+ This apophysis is, of necessity, very easily broken off in preparing the ossicula for anatomical observation.

+ The stapes in these latter and clearly inferior animals has a much less-developed head than in Homo; and the
wideness of the aperture is necessary for the transmission of a bony canal. It may be said to constitute another
and a rather high type.

482 MR. A. H. G. DORAN ON THE MORPHOLOGY

Between these extremes will be found :—first, the stapes with straight crura and a
moderate aperture, as in the Carnivora and Muride, including the Cetacean form,
where the opening is filled more or less by the thickness of the crura; then the stapes,
with thin straight crura and a very small aperture, as in Bradypus; and lastly the
variety seen in Midas, Didelphys, Tolypeutes, and Macropus, where the crura are fused
for a considerable way from the head, diverging near the base, the separation beginning
much earlier in the first two than in the last two genera just named, so that the former
possess a subvariety of higher type; from this last we find examples, in several Orders,
of insensible gradations towards the still higher straight-limbed type, with a free aper-
ture, as in the Carnivora.

In Didelphys opossum all the ossicula are of very similar character; the lamina of the
malleus is even wider; and there is a little tubercle on the inner side of the neck,
representing the processus muscularis, very feebly indicated in D. virginiana. In both
species the processus gracilis is held in a chink between two forks of the tympanic ring
during lifetime, as in most of the Marsupialia. |

In the remaining families of marsupials the divergence of the crurais either completely
absent in the stapes or exceptional and inconstant.

In the ossicula of the Tasmanian Wolf (Thylacinus cynocephalus) (P1. LXIV. fig. 17),
so like a Canis in its external appearance, there is hardly one point to be found in which
they resemble the ear-bones of a dog.

The head is very ill-developed, though not much flattened laterally, and distinctly
convex upwards. ‘The articular surface is deeply cut, with prominent margins; the lower
facet is very narrow. The neck is very thin, long, and narrow, forming a very prominent
humplike bend midway between the head and the handle. The lamina is very broad and
prolonged upon the processus gracilis, to form with it an extremely long process, which
is very narrow, except near the club-shaped, curved, and flattened extremity. The pro-
cessus muscularis is indicated by a very faint elevation on the inner side of the neck.
The manubrium is slender, and very similar in character to that of Macropus*.

Altogether, even in the character of the lamina, the malleus of the Tasmanian Wolf
more resembles that of certain Insectivora than that of any Carnivorous mammal.

The incus has an ill-developed body with a very short processus brevis ; the stapedial
crus is long and very divergent; it bears a small elliptical Sylvian apophysis on a rather
long pedicle.

-The stapes (figs. 17 & 43) is absolutely columelliform. The head is very small, and
the column long, and rather broader near the base; there is not a vestige of an aperture.
The base is wide horizontally, but shallow vertically, with the upper border well curved,
contrasting with the almost circular base of the Ornithodelphia and Sauropsida, and
constituting a higher type of the columelliform stapes, in that the footplate assumes the
same form as is observed in the higher mammals, where that ossicle is bicrurate.

* The front and lower part of the head ends in a point, ‘and is not prolonged on to the lamina joining the pro-
cessus gracilis in the Carnivora &c, With regard to the manubrium in Dasywrus, consult my observations on Myr-
mecobius further on.

OF THE MAMMALIAN OSSICULA AUDITUS. 483

In the smaller species of Dasyures, as, for instance, Dasyurus maugei (Pl. LXIV.
fig. 16), the malleus is of very similar character to that of Thylacinus, though the neck
is not so boldly curved, and the lamina is narrower. The ineus and stapes are identical
with the corresponding ossicles in the Tasmanian Wolf. Not a trace of any aperture
exists in the stapes.

In Dasyurus ursimus the malleus has a larger and rounder head, a much narrower

lamina, and, in particular, a much stouter and stronger manubrium, than in the above-
~ described species; its sides are of considerable breadth near the root. The processus
longus of the incus bears a small Sylvian apophysis on an extremely thin and fragile
pedicle. The stapes is absolutely columelliform. ‘The two inner ossicles are smaller in
an adult D. ursinus than in a full-grown Thylacinus; but the malleus of the former is of
nearly the same size and of more solid make than in the Tasmanian Wolf.

The malleus from a small Phascogale (Pl. LXIV. fig. 18) in the collection resembles
that of Dasywrus in every respect; the lamina is wide. The processus brevis of the
ineus is very short; the stapedial crus bears a similar but proportionally larger Sylvian
apophysis compared with the same in Thylacinus. The little stapes is columelliform
without an aperture; the stapedius-muscle attached to it is very large, as in most of these
animals.

In Uyrmecobius the malleus is of the same light build as in the Thylacine. The head
projects considerably above and in front of the lamina. The neck is very long, but
not niore curved than in Dasywrus. The lamina is as wide and smooth as. in the Tas-
manian Wolf, and hence more extensive than in the Dasyure and Phascogale ; it joms the
processus gracilis to form the usual long curved process applied to the tympanic ring.
The manubrium is of the same form as in the other Dasyures ; but its outer angle (which
in this family is generally sharper than in the Kangaroos) is conspicuously bent out-
wards so as to form a genuine processus brevis. The incus has an extremely short pro-
cessus brevis, a deeply cut articular surface as in other Dasyures, and a distinct Sylvian
apophysis on a long pedicle. The stapes is perfectly columelliform.

Among the PHaLANGIsTIDz the malleus of Phalangista vulpina (Pl. LXIV. fig. 20)
may be selected as a typical example. The head, though small, is better developed than
in the Dasyures; and though it does not rise high above and in front of the facets, its
upper surface is wide antero-posteriorly, projecting backwards over the articular surface.
This latter is rather deeper cut than in the Dasyures; and the upper facet is very wide,
especially vertically. The neck is rather short, but curved, the lamina rather narrow ;
and the processus gracilis is of equal or almost equal breadth throughout, so as to look
like a curved piece of steel ribbon. This is much more like the form of that process
in the Didelphyidz than in the Dasyures. The manubrium is almost straight, and is
slender and little recurved at the tip, as in the Kangaroos; the angle corresponding to
the processus brevis is rather obtuse.

A notable feature in this family is the large size of the incus. In Phalangista vulpina
it is decidedly slightly larger than in Macropus major, and larger than in any Dasyure,
nearly twice the size, in fact, of the same ossicle in Thylacinus; yet the malleus of au
adult Vulpine Phalanger is hardly larger than that of Dasyurus maug@i, where the incus

ASA MR. A. H. G. DORAN ON THE MORPHOLOGY

is hardly ever one third the size. The body of the incus of P. vulpina is well developed
and not much encroached upon by the articular surface, which has well-defined edges.
The processus brevis is stout and blunt-pointed ; it diverges little from the body. The
processus longus is long, rather straight and divergent; it bears a small pedunculated
Sylvian apophysis. |

The stapes in large well-formed skulls of this species has a small aperture as in
Macropus; its base is much longer horizontally than vertically ; and there is a small umbo
on the centre of the base.

In Phalangista cookii the distinctive characters above given are still better marked.
The upper part of the head is well developed in the malleus, projecting well backwards
over the articular facets. The lamina is very narrow, but produced on to the processus
gracilis; they together form a curved ribbon-like process of very even breadth through-
out. Theincus is as large and well-developed a bone as in P. vulpina. The stapes
(Pl. LXIV. fig. 21) is most remarkable for a character already noted by Hyrtl(/.c.): the
crura are united throughout without a trace of any aperture, forming a stout pillar, much
shorter than in any other Marsupial ; and the base is inflated into a large bulla projecting
prominently into the vestibule. I have not found this peculiar feature in any other
Phalanger; and since Hyrtl represents it precisely as it appears in a specimen recently
procured from a Cook’s Phalanger preserved in spirit in the store-room of the College
of Surgeons, it may be concluded that the bulla is a constant feature in this species.
Its morphological value does not appear to be great, as a bullate stapes occurs spo-
radically, so to speak, among higher mammals—for instance, in animals not closely allied
to each other, such as Mustela, Geomys, and Hyraz.

In Petawrus (Pl. LXIV. fig. 22) the malleus has even still more pronounced features
than in Phalangista cookii. The head is markedly of the form already described in
Phalangista vulpina; the neck is shorter, and the lamina rather wider; but in the
Flying Phalanger the manubrium is of stouter make in the above-described species, and
in the larger species (as in the fine and typical malleus of the Taguan, Petawrus
taguanoides, Pl. LXIV. fig. 23) it ends in a very marked spatulate dilatation *.

The incus of Petaurus is a large bone with similar characters to that of Phalangista.
The stapes often has a distinct perforation, though otherwise it is columelliform; the
base is much wider horizontally than vertically.

In the characters of the malleus and of the large incus, the Flying Phalangers show
certain affinities to the Sciuromorphic Rodents.

In the Koala (Phascolarctos fuscus, Pl. LXIV. figs. 24, 34) the malleus does not
possess the distinctive features so well marked in other Phalangers.

The head is very small and ill-developed, projecting little either upwards or backwards ;
it is not much flattened laterally ; the articular-surface is rather wide; and the upper
facet is planer but much larger than the lower. The neck is long, thin, and curved, the
lamina rather narrow ; and the processus gracilis forms with it a similar process to that
in Phalangista, but more prolonged. The manubrium presents a great peculiarity

* This dilatation is marked in Phalangista cookii, though the end of the manubrium is almost styliform in

P. vulpina.

OF THE MAMMALIAN OSSICULA AUDITUS. 485

(fig. 34): it is rather broad at the base and flattened laterally, very slightly curved,
and dilated at the tip; but above, at the site of the processus brevis, it is not only
broad, but presents a distinct concavity; the raised margins (of which the outer appears
continued from the neck), very wide apart above, almost coalesce below, but keep slightly
apart towards the extremity, forming a very narrow outer surface to the manubrium *.
The incus of the Koala is smaller in proportion to the malleus than in the Phalangista;
and the processus brevis is very short and ill-developed ; the stapedial crus bears a pedun-
culated Sylvian apophysis.

The stapes is absolutely columelliform, without any trace of an aperture; it bears a
small bony process for the attachment of the stapedial tendon.

Hence in all three ossicula Phascolarctos exhibits characters in which it differs from
Phalangista, and resembles rather more the majority of Marsupials.

In the Wombat (Phascolomys vombatus, Pl. LXIV. tigs. 32, 33) the most interesting
feature is the enormous development of the processus gracilis of the malleus.

The head of the malleus is even less developed and more compressed laterally than in
the Dasyuridz ; it is much encroached upon by the articular surface, which is both
deeply cut and wide; the lower facet, though more convex, is not much narrower than
the upper, from which it is divided by a well-marked groove. The neck is thin and
curved, as in the Kangaroos and Dasyures; and the lamina is narrow. The manubrium,
as in most Marsupials, is slender, and forms little more than a right angle with the
neck. The outer aspect of the base is very broad, though not so modified as in the
Koala. The extremity of the handle is narrow but distinctly spatulate. The processus
gracilis, joined by the narrow lamina, forms a curious process three times as long as
the manubrium, which is closely applied to the tympanic bone, and can readily be
detached from it even in the adult. At first it is narrow and almost of as even breadth -
as in the Phalangers; but its distal two thirds form a broad sabre-like or leaf-like
process, sharp-edged, much flattened, and ending in a tapering point. The whole pro-
cessus gracilis is only an extremely developed form of the type seen in the Rhinoceros
and in many of the Insectivora and Marsupials. The foliaceous extremity is readily
broken off if the specimen be not carefully prepared.

In the incus the body is fairly developed, especially above the processus brevis, though
the whole bone is not so large in proportion to the malleus as in Phalangista. The pro-
cessus brevis is rather short, but stout at the base; it diverges but little from the body.
The stapedial crus is long and thin, and supports a large elliptical Sylvian apophysis on
a slender pedicle. The stapes is quite columelliform ; the head forms a narrow elliptical
capital to the fused crura; just beneath it is a spur-like process, to which the tendon of
‘the stapedius may be found attached in the recent or even in the imperfectly macerated
‘skull. The base is not of the Sauropsidan character, being as narrow vertically as
in Homo.

The ossicula of P. latifrons are very similar in character. They are all rather larger in
the Wombats than in the largest Dasyures (such as D. wrsimus) or Thylacinus.

* The blunt outer aspect of the base of the manubrium in the mallei of large. Dasyures and other Marsupials
exhibits an approach to the above-described peculiarity in the Koala. ~

SECOND SERIES.—ZOOLOGY, VOL. I. 3T

486 MR. A. H. G. DORAN ON THE MORPHOLOGY

In the PrramMeLip# the malleus is of light and fragile construction, although wholly
osseous and not very different in character from the same ossicle in many older Mar-
supials (Pl. LXIV. fig. 25).

The head is very small, with a deeply cut articular surface, which has two facets, the
upper and anterior being the widest and plainest of the two, The neck is long and
thin; it forms, by suddenly bending on itself midway between the head and manubrium,
asharp projecting angle, as in the equally fragile malleus of the Soricide. The slender
manubrium is rather broad and flattened at the base, and is blunt externally at the point
corresponding to the processus brevis. It ends in an almost styliform extremity. The
lamina is narrow, and joins the processus gracilis to form a curved process slightly broad
near the tip, as in the Dasyuride ; this is fitted into the tympanic bone as in other
Marsupials.

The incus of Perameles has a rather high and narrow body. ‘The processus brevis is
rather long and thin, and not divergent from the body. ‘The processus longus is thin,
long, and very divergent ; it ends in a very conspicuous and perfectly elliptical Sylvian
apophysis, supported on a pedicle which is extremely thin at its attachment to the
stapedia] crus, but becomes broad and flattened before it ends in the extremity of the
apophysis. This is but an extreme development of the condition so frequent in the
Didelphia.

The stapes is completely columelliform ; and the column isas short as in the Passerine
birds; the base, too, is rather wide vertically. This ossicle is generally imperfectly
ossified even in the adult Bandicoot.

In Cheropus all the ossicula are of the type seen in Perameles (Pl. LXIV. fig. 26).

All the ear-bones of this family must be considered as of a very low type, the fragile
. malleus of the adult recalling that ossicle in the foetus of higher mammals. The ill-
development of the processus brevis incudis and the persistent distinct character of the
Sylvian apophysis, as well as the perfectly columelliform stapes, combine to force upon
the observer the conclusion that, in respect of the ossicula, the Peramelide are the
lowest of the Didelphia; for in no other family are the little bones to be found all
three so thoroughly of low development in every detail.

In concluding this review of the Ossicula of the Marsupialia, it may be said, by way of
a summary :—

I. That the ear-bones in this subclass are always of a low type, yet not as a rule so
inferior to those of the Monodelphia as might be inferred, judging from the rest of
their skeletons.

II. That no point showing a low grade of development in any ossicle in any Mar-
supial is not occasionally met with in higher mammals, excepting that in the latter the
ear-bones are never seen to retain to so high a degree the foetal consistence as may be
observed in the adult Perameles.

III. That the Bandicoots or Peramelidz are, in respect of their ossicula, the lowest
family in the whole subclass, both in the type of all three bones, and in their frail
papery consistence compared with the rest of the cranium.

OF THE MAMMALIAN OSSICULA AUDI'TUS. 487

IV. That in the same respect the American Opossums or Didelphyide must be con-
sidered the highest of the Marsupials. The malleus is, in type and consistence, similar
to that of many Insectivora; the incus has the body and short crus as well as the pro-
cessus longus well developed; and the stapes is more perfectly bicrurate than in any other
allied family.

V. The Macropodidee are very central among Marsupials in their ossicula. The
malleus is of low type, but tolerably firm osseous consistence; it bears the peculiar
highly-developed foliaceous processus gracilis aiso seen in the Wombats; the incus is
fairly developed; and the stapes is always partially bicrurate.

VI. The Phalangistidee come next in order. They are decidedly inferior to the
Kangaroos in that the stapes is generally (though not invariably) columelliform; more-
over the malleus is of the form seen in a new-born Macropus; but in the size and
good development of all parts of the incus they are as high or even higher than the
American Opossums. Phascolarctos is lower in all respects than Phalangista; its
malleus is of a very distinctive character.

VII. The Wombats are slightly lower than the last family, the stapes being always
columelliform, and the ineus having a rather short processus brevis, but a stapedial
crus rather of the Perameles type. The malleus is not so much of high type as an
extreme form of the specialized corresponding ossicle in Macropus, the processus gracilis
being very large, curved, and flattened.

VIII. The Dasyures have the stapes invariably columelliform, and the incus of low
type; but all their ossicles are of more solid consistence than in the Bandicoots
(Peramelidee).

THe OssIcuLA OF THE MoNOTREMATA.

Next to the ear-bone of Man, those of Hchidna and Ornithorhynchus are of all other
mammalian ossicula the best known to the anatomist. From the conformation of the
base of the skull in the Monotremata, the malleus cannot fail to obtrude itself on the
glance of the student cursorily examining the cranium. ‘Then these little bones are of
profound interest to the morphologist when studying this subclass, which is so un-
-doubtedly allied in many respects to the Sauropsida, where two out of three ossicula
are suppressed. Good general descriptions of the ear-bones of the Monotremata, with
clear figures, are to be found in the works of Huxley *, Hyrtl +, Peters ¢, and others.

I leave such important questions as What bones are the homologues of the mam-
malian malleus and incus in the Sauropsida? to distinguished authorities in the united
branches of morphology and embryology. Next, I need hardly remind the anatomist
that, as far as the ossicles are concerned, the Monotremata wear a perfectly mammalian
uniform, having malleus, incus, and stapes. If any theorist consider this Order more
Sauropsidan than mammalian, on account of the most marked feature in the first two

* Proc. Zool. Soc. 1869; see woodcut, p. 404. t+ Op. cit.
+ “ Ueber das os tympanicum und die Gehérknochelchen der Schnabelthiere &c.” Monatsberichte der Akad. der
Wissensch. Berlin, 1867.
3T 2

488 MR. A. H. G. DORAN ON THE MORPHOLOGY

bones, he must, for the same reason, place the Porcupines and Cavies nearly as low in
the scale, and decidedly in an inferior position to the Marsupials; and as for the stapes, it
is of a type precisely imitated in the placental Manis.

It remains for me to describe the ear-bones of this Order as I have detailed the same
in higher milk-secreting vertebrata, investigating carefully their exact homologues in
the former.

In Echidna hystrix the two outer ossicula are proportionally much larger, better
developed, and more firmly osseous than in Ornithorhynchus. It is on that account
advisable to commence with the description of the ear-bones of the so-called Porcupine
Ant-eater.

Let the united malleus and incus of an Hchidna be removed from the skull. In so
doing observe that the processus brevis of the incus lies anterior as well as superior to
the stapedial crus in the natural condition, an exaggeration of what is seen in the
Sirenia, Marsupials, and other mammals. Even in Homo the short crus is almost as
superior as it is posterior to the rest of the incus. Observe also, if the skuil be im-
perfectly macerated, how large the tensor tympani muscle is, as Professor Huxley *
has already remarked. Were it attached to the inner edge of the manubrium, as in
the Rodents, it would require still greater care and preparation, to remove the malleus
entire, than even is necessary in those animals. Now let the united malleus and incus
be examined from the inner aspect (Pl. LXIV. fig. 36), and the complete outline of the
incus may plainly be seen. The inner aspect of its body is free, square, and faintly.
concave, whilst antero-inferiorly it is limited by a convex border fitting into a concavity
in the malleus. The whole, at this region, has precisely the same appearance as may be
seen in articulating the incus and maileus in a Marsupial, or in any mammal where the
latter ossicle has a small ill-developed head.

But view the bones from without (fig. 35), and it will then be observed that the
malleus encroaches on the whole of the outer surface of the body of the incus, so that
only the processus brevis (pd) and the tip of the processus longus (pl) can be seen. As
it covers the greater part of the long process, on turning to the inner aspect of the
ossicles the singular appearance of a lamina of bone, reflected from the outer border of
the stapedial crus towards the manubrium, is at once accounted for, that lamina being
simply an integral part of the malleus appearing from behind at the point where, from
this aspect, the incus ceases to cover it.

‘Hence the incus is articulated to the malleus by its whole inner surface, including the
greater part of its long crus, and also by its convex inferior border. It is firmly anky-
losed to the malleus; and even in a young skull the ossicles cannot be separated unin-
jured without difficulty.

Bearing in mind the above remarks, the description of the malleus by itself will now
be more comprehensible. The head is narrow and concave; externally it is prolonged
upwards into a thin scale (Pl. LXIV. fig. 35, shm), of which the wide convex free
border reaches to the base of the short crus, and conceals still more of the long process

* Proc. Zool. Soc. 1869, “« On the Representative of the Malleus and Incus of the Mammalia in the Sawropsida.”

OF THE MAMMALIAN OSSICULA AUDITUS. 489

of the ineus. This scale hides the concave excavation of the head, which, on the other
hand, is plainly visible from the inner aspect of the united bones (fig. 36, hm).

From the inner aspect, too, it may be seen that the head sends forward a long, pro-
minent, and very straight process to join the processus gracilis *. The analogue of this is
already familiar to us in most laminated and in other mallei. The neck is rather long,
and not strongly curved; its free border appears from in front of the incus when the
' bones are viewed from within, as just described; in the opposite direction it blends
insensibly with that part of the bone representing the lamina in higher mammals.
- From the outer aspect the neck is more clearly defined. Close to the root of the
manubrium a small tubercle rises from the inner side of the neck, to which the tensor
tympani muscle will be found attached in imperfectly macerated specimens. Hence
this tubercle is the processus muscularis.

The manubrium of Lchidna is of a type very frequent among the lower mammalia.
It is of moderate length, compressed laterally, but not so thin and broad at the root as
in some Rodents. The outer angle of its base (the site of the processus brevis, which is
absent) is very blunt, broad, and well rounded; the outer aspect is very narrow, though
sharply bordered from the sides, and ending in a narrow but distinct spatulate dilata-
tion. The entire process, in slenderness, bluntness of its outer angle, &c., recalls the
same in Rhinoceros, Priodon, and the Marsupials. It cannot be said to bear any clese
resemblance to any process of the os quadratum or any neighbouring bone in the
Sauropsida.

The processus gracilis is readily distinguished, as it starts off from the malleus, forms
a deep curve, and joins the process from the head, to form the long straight splinter of
bone which articulates with the tympanic. This condition is well known; but anatomists
hitherto have described the united part as if it were solely and entirely the processus
gracilis, ignoring the process from the head, which I have frequently described in
animals where it is well marked, notably in Fiber. Whilst this latter process, as above _
stated, is traced along the united portion from the inner aspect, the continuation of the
processus folianus proper must be looked for in viewing the malleus from without; then
it can be seen to terminate in a broad lamellar expansion, on which the process from the
head loses itself posteriorly.

Between the neck and the free curved portion of the processus gracilis (and hence best
seen from the outer aspect of the ossicle) the analogue of the lamina so frequent in the
higher mammals may be distinguished : it is neitWer broad nor thin, and runs on towards
the united processes, with which it blends insensibly. This, it may be remembered, is the
same as in the Marsupials.

The body of the incus has already been referred to, and the free inner aspect described,
as well as the manner in which it is wholly covered externally by the malleus+. The
processus brevis is short and flattened extero-internally; it projects from the outer

* See Pl. LXIV. fig. 36. The proccss from the head (phm) is here distinguished from the processus

gracilis (pq).
+ Excepting its anterior edge, which is just left uncovered.

490 MR. A. H. G. DORAN ON THE MORPHOLOGY

aspect above the scale-like process from the head of the malleus, which covers the
body. The processus longus is not much longer than the short crus; it is straight and
flattened, and wholly concealed from without by the malleus, except its extremity, which
is bent inwards, and bears a very small but distinct Sylvian apophysis.

It will be seen at a glance that the kind of ankylosis between the two outer ossi-
cula in Echidna is totally different from their mode of union in Porcupines and Cavies,
where the malleus and incus are joined on the same level externally as internally ;
and, besides, there is always in the latter animals a peculiar development of the head
of the malleus forwards, away from the incus, instead of the scale-like prolongation of
the head on to the outer side of the latter ossicle. .

The incus of Echidna is, after all, not so very much smaller in proportion to the
malleus than in many Marsupials; only it is more flattened; flatness is also seen in the
same ossicle in those Rodents where it is fused with the malleus. I have never found
it to be so loosely attached to the malleus as in young skulls of the Duck-bill.

The stapes is absolutely columelliform. Its head is very small, with no trace of any
tubercle for the stapedius-tendon, which does not appear to exist in this animal*. The
column bears not a trace of any subdivision into crura; it is rather long, and the foot-
plate is almost perfectly circular.

What relation does the stapes of this avowedly low mammal bear to the columelliform
representative of the same in higher animals on one hand and in the Sauropsida on the
other? Is the column of the stapes of Hehidna formed of the same element or elements
as in Aves ?

By referring to the description of the stapes in the Marsupialia, it will be seen that
this bone is of the same perfectly columelliform type in the Dasyures, Wombats, Bandi-
coots, and most Phalangers, as in the Echidna; and among the Edentata, Manis has a
similar type of stapes. In some of the above animals the base of the ossicle in question
- is less perfectly circular than in Echidna, and hence more like the foot-plate of the
bicrurate stapes of Homo. Then the column is shorter in some of these animals than in
the Porcupine Ant-eater.

But it happens that all the above characters rather strengthen the alliance between
Marsupials, the Manide, and the Monotremata, than show that those animals are parti-
eularly ornithic or reptilian.

In some birds where the column is almost as short as in these mammals, it divides
distinctly into two crura, as distinctly as in Macropus and Tolypeutes. This is the
case with the Nightingale (Pl. LXIV. fig. 45) and Redstart; or at least it is so in the
specimens in the College collection. In others, where the column is very long, it may
also divide into two crura near the foot-plate, as is very clear in the Hagle (Aquila,
Pl. LXIV. fig. 46). Besides, the foot-plate in many Finches and other birds, and in
Crocodiles and other reptiles, is not of the circular outline seen in the Echidna, but as
long horizontally, and narrow vertically, as in the stapes of the highest mammals.

Then, without plunging into unsettled questions of development, does the column of

* Huxley. I have never seen a vestige of any representative of that muscle.

OF THE MAMMALIAN OSSICULA AUDITUS. 491

the stapes of these inferior mammals represent the same or only a part of the elements
making up the column of a bird’s columella ?

In all these mammals we have an incus with a processus longus and Sylvian apo-
physis articulating with the stapes, which is columelliform; but the column is clearly
identical with the two crura of the higher Mammalia, as the transitional condition
seen in Macropus &c. clearly proves.

In birds the column of their stapes or “columella” is certainly the same, whether
short (as in the Finches, Pl. LXIV. fig. 44) or very long (as in the Hagle, Albatross, &c.,
figs. 46, 47); for the same “ extrastapedial” and “suprastapedial” elements are found
attached to its outer extremity in the same manner in both varieties. But at least part
of the column is recognized by the latest investigators into embryology * as the “ medio-
stapedial’’ element which represents the long crus of the incus in mammals, and
begins as the proximal incurved extremity of the hyoid arch in both the Mammalia
and the Sauropsida.

Now, this long crus of the incus always existing in mammals, it follows, if the most
advanced embryologists have not erred, that the column of the Marsupial and Mono-
trematan stapes (as well as that of Manis) represents only a part of the columella in
Aves. It is evidently formed entirely from the mass of cells which grow from the
periotic capsule to meet the proximal end of the hyoidean arch (Parker, J. c.) ; for since
that end becomes the processus longus of the incus and its Sylvian apophysis in these
mammals (identical with the mediostapedial element in birds), no segment of that
arch can take any share in the formation of their stapes, as if is seen to do in the
Sauropsida.

In examining the ear-bones of Ornithorhynchus paradoxus, the first thing that strikes
the observer is the relatively small size of the malleus and incus compared with those bones
in Hehidna, and.their imperfectly ossified semimembranous texture, in which they exceed
those of any other mammal. The stapes in the Duck-bill is nearly as large as in the
Porcupine Ant-eater (Pl. LXIV. fig. 37).

If the united malleus and incus of Ornithorhynchus be more closely investigated, it
will be found that in ossicles from young skulls they may be readily disarticulated ;
but in specimens from the adult this is not invariably the case. Professor Peters +
asserts that these bones remain free throughout life in this animal; but in a set which
I prepared for the College collection, from a full-grown skull, the malleus and incus
remained ankylosed on the right side. ee

The manner of union between these two ossicula is precisely the same as in Hehidna.
The incus is seen from the inner aspect (Pl. LXIV. fig. 38), but almost completely
concealed in viewing the malleus from without (fig. 37) {.

The portion of the malleus representing the head in higher mammals (fm) is ex-

* Consult summary at the end of Professor Parker’s memoir “ On the Structure and Development of the Skull of
the Pig,” Phil. Trans. 1873.

+ Loe. cit.

+ In Pl. LXIV. figs. 37 & 38, the same abbreviations are used as in the figures of the corresponding ossicles

in Echidna.

492, MR. A. H. G. DORAN ON THE MORPHOLOGY

tremely small; and hence the greater narrowness of the malleo-incudal articulation, as
seen from within, compared with Echidna. But it sends up from without a scale-like
process (sim), as in that animal, which conceals the body of the incus and the root of
its long process; so that on viewing the bones from within, the same appearance of a
lamina of bone, reflected from that process to the malleus, may be noted. When the
incus can be detached, a smooth slightly concave space on the process from the head
comes into view. This has been already figured in Peters’s memoir already quoted.

From the head a very stout and straight process is sent forwards, blending with the
very thin papery processus gracilis. The neck is longer, and much better defined than
in Lchidna, and forms a very wide curve. The true processus folianus is merely the
slightly thickened free border of the narrow but distinct lamina; it joins the well-deve-
loped process from the head. It is only by a careful comparison of these elements in
Ornithorhynchus with the same in laminated mallei of placental mammals that the error
can be avoided of taking the process from the head for the processus gracilis *.

The processus muscularis is of a unique form. It is a wide angular projection from
the inner side of the neck above the manubrium. The processus gracilis (and its adjunct
just referred to) is united with the tympanic bone in the same manner as in Hehidna.
This bone is here less developed and more sharply curved than in that animal. Hence,
united with the malleus, it bears a very characteristic distinctive appearance.

The manubrium is short, ill developed, and so imperfectly ossified, as to shrink when the
malleus is dried. Its outer angle is blunt, very broad, and slightly concave, somewhat
as in Phascolarctos. Its shank is very slender, and slightly dilated at the extremity ;
the outer aspect is excessively narrow, but distinctly bordered from the sides.

The incus is small and flattened, as in Hehidna. Its inner surface is more concave, and
narrower where it joins the malleus. The crura are short, stout, and straight, and almost
of equal length; the stapedial crus bears a very faint indication of a Sylvian apophysis.

The stapes is absolutely columelliform, and not so firmly osseous as in Hehidna ; the
column is rather longer, the head broader, and the almost circular footplate is not
quite as large. As a processus longus of the incus exists, the column must be the
exact homologue of the same in Echidna, and of the two crura of the stapes of the
higher mammals, and, for the same reason, represents only part of the column of the
Sauropsidan columella. In fact the presence of the processus longus of the incus throws
the stapes of the Monotremata both towards other mammals, and away from the birds.

The result of the above considerations shows :—

1. That, in their ossicula, the Monotremata are as low as in the rest of their skeleton,
yet, as might be expected, thoroughly mammalian.

2. That their distinguishing features are a peculiar form of articulation between the
malleus and incus by means of a scale-like development from the head of the former,
and the presence of an absolutely columelliform or unicrurate stapes. Otherwise the
three ossicula are not much modified from their representatives in the lower Marsupials.

* Compare phm and pg in Pl. LXIV. figs. 36 & 38, with the same in Pl. LXIII. fig. 14.

OF THE MAMMALIAN OSSICULA AUDITUS. 493

3. That a definite incus has long been admited to exist, articulated, as above noticed,
with the malleus, and ankylosed in the Hehidna, and generally in the adult Ornitho-
rhynchus. The method of ankylosis is quite different from that seen in certain Rodents.

4. That as a processus longus of the incus exists, and hence in all probability is the
proximal end of the second postoral arch, it follows that the column of the stapes of
these animals (and of all other mammals where it is of similar shape) represents the two
crura of the stapes of higher mammalia, but only a part of the column of the Sauropsidan
columella, which is generally admitted to be formed partly from the ‘‘ mediostapedial ”’
element of the hyoid arch. But a further careful study of the embryo of these animals
is necessary to settle this point otherwise than by analogy.

5. That the malleus and ineus of Ornithorhynchus differ from those of Echidna in
matters of detail, but in no essential features which might ally either animal more or
less than the other to higher mammals or to the Sauropsida, excepting, some might say,
the semimembranous consistence of the malleus of the Duck-bill.

DESCRIPTION OF THE PLATES.
Pirate LVIII.

Throughout the Plates the deft malleus and incus are represented as viewed from the inner side, that
position being the most convenient for displaying the homologous elements. The “ natural position,”
z.e. that in which the manubrium of the malleus is turned towards the observer, is objectionable, because
foreshortening of the ossicles interferes with a ready comparison of their various processes and relative
dimensions as seen in different animals. The ossicula are enlarged 8 times natural size, unless otherwise
specified. The letters used im the various figures bear the following signification :—as, articular surface
of malleus; , neck ; pb, processus brevis ; pg, processus gracilis; a, a*, upper facet, and 6, b*, lower
facet of malleus; sr, sigmoid ridge of malleus ; emn, external surface of manubrium; pl, processus
longus ; ¢, concavity head of malleus ; eas, extra articulating surface on malleus and incus ; gr, groove
on incus; an, angular projection on neck of malleus; oa, orbicular apophysis ; mall, malleus; Im, fig. 22,
Pl. LX., in error for pm; pm, processus muscularis; hm, head of malleus (fig. 1, Pl. LXIII., lettered
wrongly for mn, see footnote p. 458) ; pg, processus gracilis; J, lamina; phm, process head of malleus ;
acpm, accessory processus muscularis ; mn, manubrium; pet, petrous bone; 7b, tympanic bone; shim,
scale head of malleus ; ms, medio-stapedial element.

Figs. 1-17. Left ossicula: 1,2, Homo*; 8, Troglodytes gorilla; 4, Troglodytes niger ; 5, Simia satyrus ;
6, Hylobates syndactylus ; 7, Semnopithecus entellus ; 8, Colobus satanas; 9, Cercopithecus cephus ;
10, Macacus erythreus ; 11, Cynopithecus niger ; 12, Cynocephalus anubis ; 18, Ateles melanochir ;
14, Mycetes ursinus ; 15, Cebus; 16, Nyctipithecus trivirgatus; 17, Saimaris sciurea.

Fig. 18. Left malleus and incus: Pithecia satanas.

Figs. 19-22. Left ossicula: 19, Hapale jacchus; 20, Midas edipus ; 21, Lemur albifrons; 22, Galago
crassicaudata.

* The remaining figures of the left ossicula show the malleus placed as in fig. 2, and the incus as in fig. 1. I had
intended that the human malleus and incus should be placed together in the desired position in the same group—
namely, fig. 2; but by an oversight the object was not reversed on being drawn on stone.

SECOND SERIES.—ZOOLOGY, VOL. T. 3U

494 MR. A. H.G. DORAN ON THE MORPHOLOGY

Fig. 23. Left malleus and incus: Cheirogaleus cookii.

Figs. 24-26. Left ossicula: 24, Propithecus diadema (young) ; 25, Perodicticus potto; 26, Nycticebus
tardigradus.

Fig. 27. Left malleus: Arctocebus calabarensis.

Figs. 28, 29. Left ossicula: 28, Tarsius spectrum (young); 29, Chiromys madagascariensis.

Figs. 80, 31. Articular surface of right malleus: 30, Homo; 31, Felis, much enlarged ; a, a*, upper facet;
6b, b*, lower facet.

Figs. 32-41. Right mallei of various Carnivora, arranged so as to show the principal distinguishing
features seen in the different families, more especially the processus muscularis and the relative
breadth of the outer aspect of the manubrium: as, articular surface of malleus; n, neck ;
1, lamina; pg, processus gracilis; pm, processus muscularis; pd, homologue of processus
brevis; emn, external surface of manubrium:—82, Felis leo; 33, Canis lupus; 34, Hyena
maculata; 35, Proteles cristatus; 36, Ursus maritimus; 37. Viverra civetta; 38, Herpestes
ichneumon ; 39, Gulo luscus; 40, Meles taxus, showing angular projection on neck; 41. Lycaon
pictus.

Figs. 42-55. Left imcus: 42, Felis domestica; 43, Canis familiaris; 44, Cryptoprocta ferox ; 45, Viverra
cwetta; 46, Paradoxurus musanga; 47, Herpestes pulverulentus ; 48, Hyena; 49, Ursus mari-
timus ; 50, Otaria falklandica; 51, Alurus fulgens; 52, Bassaris astuta; 53, Cercoleptes caudi-
volvulus ; 54, Lutra vulgaris ; 55, Mustela erminea.

Figs. 56-64. Left stapes: 56, Kelis domestica; 57, Canis lupus: 58, Hyena; 59, Cryptoprocta feroz ;
60, Viverra civetta; 61, Herpestes pulverulentus; 62, Mustela erminea ; 63, Cercoleptes caudi-
volvulus; 64, Ursus maritimus.

Puate LIX.

Figs. 1-3. Left ossicula: 1, Felis domestica; 2, Arctictis binturong ; 3, Suricata zenik.

Fig. 4. Left malleus and incus: Octocyon lalandii.

Figs. 5-12. Left ossicula: 5, Canis familiaris; 6, Canis vulpes; 7, Martes abietum; 8, Ictonyx zorilla ;
9, Lutra vulgaris **; 10, Hupleres goudotit ; 11, Procyon lotor ; 12, Nasua nasica.

Fig. 13. Left malleus and incus: Aonya indigitata.

Figs. 13a-16. Left malleus : 13a, Cynogale bennettii ; 14, Cercoleptes caudivolyulus ; 15, Bassaris astuta ;
an, angular projection on neck; 16, Hlurus fulgens.

Figs. 17, 18. Left ossicula: 17, Ursus ornatus; 18, Otaria falklandica (c in this figure and in the mallei
of others of the Pinnipedia, represents the concavity on the anterior surface of the head of the
malleus). j

Fig. 19. Right ossicula: Otaria falklandica.

Fig. 20. Left ossicula: Otaria stelleri.

Vig. 21. Right ossicula: Trichechus rosmarus.

Fig. 22. Left ossicula: Trichechus rosmarus.

Fig. 23. Right ossicula: Macrorhinus proboscideus; c, concavity of malleus.

Fig. 24. Left ossicula: Macrorhinus proboscideus.

Fig. 25. Right ossicula: Cystophora cristata; c, concavity of malleus.

Fig. 26. Left ossicula: Lobodon carcinophaga.

Fig. 27. Right ossicula: Phoca vitulina; c, concavity of malleus.

Fig. 28. Left ossicula: Phoca vitulina; eas, extra articulating surface on the malletis and incus;
gr, groove on the incus.

Figs. 29-31. Left malleus: 29, Phoca barbata; 30, Phoca greniandica; 31, Phoca hispida.

* The processus muscularis was broken off in the specimen from which this figure was taken.

OF THE MAMMALIAN OSSICULA AUDITUS. 495

Puate LX.

Fig. 1. Left malleus and incus: Phoca vitulina (articulated).

Figs. 2-5. Left incus: 2, Phoca barbata; 3, Phoca grenlandica; 4, Phoca hispida; 5, Halicherus grypus.

Figs. 6-35. Left ossicula: 6, Sciwrus maximus; 7, Pteromys nitidus; 8, Anomalurus fraseri; 9, Tamias
striatus ; 10, Spermophilus citullus; 11, Arctomys marmotta; 12, Castor fiber; 13, Myoxus
glis; 14, Mus decumanus (oa, orbicular apophysis) ; 15, Hapalotis mitchellii; 16, Hydromys
chrysogaster ; 17, Arvicola amphibius ; 18, Fiber zibethicus (p.h.m, process from head of malleus) ;
19, Spalax typhlus ; 20, Rhizomys badius ; 21, Geomys bursarius ; 22, Bathyergus maritimus ; .
23, Dipus egyptius ; 24, Hystrix cristata; 25, Cercolabes insidiosus ; 26, Capromys pilorides ;
27, Aulacodus swindernianus ; 28, Octodon cumingii; 29, Chinchilla lanigera ; 30, Dasyprocta
agutt; 31, Dolichotis patachonica; 32, Hydrochwrus capybara; 33, Cavia caprera; 34, Lepus
cuniculus ; 35, Hyrax capensis.

Figs. 36, 37. Stapes: 36, Elephas indicus (adult) ; 37, Elephas africanus.

Fig. 38. Right ossicula: Hlephas indicus.

Fig. 39. Left ossicula: Hlephas indicus.

Puate LXI.

Fig. 1. Left ossicula: Rhinoceros bicornis.
ig. 2. Left malleus and incus: Tapirus bairdi.

Figs. 3, 4. Left ossicula: 3, Equus caballus ; 4, Hippopotamus amphibius.

Fig. 5. Right ossicula: Hippopotamus.

Figs. 6-9. Left ossicula: 6, Sus scrofa: 7, Dicotyles tajacu ; 8, Tragulus javanicus ; 9, Lama peruana.

Fig. 10. Right malleus: Camelus dromedarius.

Fig. 11. Left malleus : Camelus dromedarius.

Figs. 12-21. Left ossicula: 12, Antilocapra americana ; 13, Nanotragus montanus ; 14, Bos taurus (adult) ;
15, Bos taurus (foetus 6th week) ; 16, Ovis aries (adult); 17, Ovis aries (foetus, at birth) .
18, Gazella dorcas (adult) ; 19, Alcelaphus bubalus ; 20, Cervus elaphus (adult) ; 21, Camelo-
pardalis giraffa (adult) .

Figs. 22-33. Left mallei of various Ruminants* (young) : 22, Rupicapra tragus; 23, Saiga tartarica ;
24, Kobus ellipsiprymnus ; 25, Tetraceros quadricornis; 26, Catoblepas gnu; 27, Procapra
picticaudata ; 28, Cephalophus maxwellit ; 29, Nemorhedus sumatrensis ; 30, Nemorhedus goral ;
31, Ovibos moschatus ; 32, Anoa depressicornis ; 33, Moschus moschiferus.

Figs. 34-42. Incudes of various Ruminants (left): 34, Rupicapra tragus; 35, Saiga tartarica; 36;
Kobus ellipsiprymnus ; 37, Tetraceros quadricornis; 38, Procapra picticaudata; 39, Cephalo-
phus maxwellii ; 40, Ovibos moschatus; 41, Anoa depressicornis ; 42, Moschus moschiferus.

eI

Puate LXII.

As many of the ossicles are here represented much magnified, a smaller figure is, m all such cases,
placed to the left of the greater, the smaller being drawn on the same scale as in all the other plates, 2. e.
three times the natural size.

Fig. 1. Right ossicula of a large American Sorex: oa, orbicular apophysis; pb, processus brevis.

Fig. 2. Left malleus from the same Sorer. Fig. 6. Left meus: Scalops aquaticus.

dig. 3. Left malleus: Myogale moschata, Fig. 7. Right malleus; Condylura cristata.
Fig. 4. Right ossicula: Myogale moschata. Fig. 8. Left malleus and incus: Condylura.
Fig. 5. Left ossicula: Talpa europea. Fig. 9. Right malleus: Chrysochloris capensis,

* The laminz were more or less mutilated in the originals from which several of the figures in this group were

taken.

3U 2

496

Figs

Fig.
Tig.
Hig.
Fig.
Fig.
Wig.
Fig.
Fig.
Fig.
Fig.

Fig.

MR. A. H. G. DORAN ON THE MORPHOLOGY

. 10-18. Left ossicula: 10, Chrysochloris ; 11, Centetes ecaudatus ; 12, Ericulus nigrescens ; 13, Eri-
naceus europeus; 14, Gymnura rafflesit ; 15, Macroscelides alexandri; 16, Rhynchocyon cernet ;
17, Tupaia tana; 18, Galeopithecus.

19. Left malleus: Petrodromus.

20. Right malleus: Asellia (Phyllorhina) tridens ; acpm, accessory processus muscularis.

21. Right malleus: Plecotus auritus (outer aspect).

22. Left ossicula: Vespertilio nattereri.

23. Right malleus: Mormops blainvillic.

24. Left malleus: Molossus obscurus.
25. Left ossicula: Vesperugo serotinus.
26. Right ossicula: Pteropus edulis.
27. Left ossicula: Pteropus edulis.

28. Left malleus: Balena mysticetus.

29. Right incus and stapes: Balena mysticetus.
30. Right malleus: Balenoptera rostrata.
31. Right incus and stapes: Balenoptera rostrata,

. 82. Left malleus: Megaptera nove-zelaendiz.
. 33. Right malleus: Neobalena marginata.
. 84. Left malleus, right incus and stapes: Physeter macrocephalus.

35. Left ossicula: Hyperoodon rostratus.

Pratre LXIL.

.1. Left malleus and incus: Berardius arnuxit.
. 2. Left ossicula : Delphinus.
gs. 8-12. Right malleus: 3, Delphinus; 4, Globiocephalus ; 5, Phocena; 6, Pseudorca meridionalis ; 7,

Orca; 8, Delphinapiterus albicans ; 9, Monodon; 10, Inia geoffroyit; 11, Lagenorhynchus ; 12,
Platanista.

, 13. Left malleus: Balena australis, in situ (pet, petrous bone ; tb, tympanic bone).
. 14. Left malleus: Felis, for comparison of the homologous parts in figs. 13, 15, 16,17, 18, &e.

Figs. 15-17. Left malleus: 15, Globiocephalus ; 16, Monodon (with tendon of tensor tympani and process

Spe ste ete see eS
Set Ge Se Se (So Se Sa So So
3 oe of Ue Fe Ge Je de de ihc}

=
pas
Q

from membrana tympani attached) ; 17, Platanista.

. 18, 19. Left malleus: 18, Berardius arnuxii; 19, Mesoplodon (Oulodon) grayii.
. 19a. Right malleus: Mesoplodon grayii (inner aspect, showing the facets, the groove for the process

from the tensor tympani, and the pointed posterior extremity of the head).

. 20. Right incus: Platanista,

91. Left malleus: Manatus americanus.

.22. Right malleus: ditto.

23. Right incus: ditto.
24. Left incus. ditto.

. 25. Stapes: Manatus senegalensis.

26. Left malleus: Halicore australia.
27. Right malleus: ditto.
28. Right incus: ditto.

¢. 29. Left incus: ditto.
g. 30. Stapes ditto.

OF THE MAMMALIAN OSSICULA AUDITUS. 497

Prats LXIV.

Figs. 1-6. Left ossicula: 1, Bradypus tridactylus ; 2, Cholepus didactylus ; 3, Tatusia peba ; 4, Dasypus
sexcinctus ; 5, Priodon gigas; 6, Tolypeutes tricinctus.

Fig. 7. Left malleus: Tatusia hybrida (new-born).

Fig. 8. Left ossicula: Manis dalmanit.

Fig. 9. Left meus: Manis tricuspis.

Fig. 10. Left malleus: Manis dalmanii. Head, much enlarged, to show articular surface.

Fig. 11. Left ossicula, Myrmecophaga jubata.

Fig. 12. Left malleus and incus: Tamandua tetradactyla.

Figs. 13-22. Left ossicula: 13, Cyclothurus didactylus ; 14, Orycteropus capensis; 15, Didelphys virgi-
nana ; 16, Dasyurus maugei; 17, Thylacinus cynocephalus; 18, Phascogale ; 19, Myrmecobius
fasciatus ; 20, Phalangista vulpina; 21, Phalangista cookii; 22, Petaurus flaviventer.

Fig. 23. Left malleus: Petaurus taguanoides.

Figs. 24-27. Left ossicula: 24, Phascolarctos fuscus; 25, Perameles; 26, Cheropus castanotis ; 27,
Macropus laniger.

Fig. 28. Left malleus: Halmaturus Bennettii (in situ).

Fig. 29. Left malleus and incus: Lagorchestes.

Fig. 30. Left ossicula: AHypsiprymnus.

Fig. 31. Right malleus: Macropus major.

Fig. 32. Left ossicula: Phascolomys vombatus.

Hig. 33. Right malleus: Phascolemys vombatus.

Hig. 34. Right malleus: Phascolarctos, showing great width; upper portion of the manubrium.

Fig. 35. Right malleus and incus: Echidna (Tachyglossus) hystrix, from without; shm, scale-like
process from head of malleus, concealing body of incus; pd, pl, processus brevis and longus
of incus.

Hig. 36. Left malleus and incus: Echidna, from within ; hm, head of malleus; pg, processus gracilis ;
phim, process from head of malleus, blending with processus gracilis.

Fig. 37. Right malleus and stapes: Ornithorhynchus paradozus.

Fig. 38. Left malleus: Ornithorhynchus.

Figs. 39-48. A series of stapedes of mammals, “ columellz ” of Sauropsida, displaying the gradations of
type in the Mammalia of the occasional reappearance of the bicrurate stapes in Aves: ms,
medio-stapedial element in columella of Sauropsidan, equivalent, according to Professor Parker,
to the processus longus of the mammalian incus.

Figs. 39-43. Srapes: 89, of Man; 40, White Bear; 41, Virginian Opossum ; 42, Great Kangaroo ; 43,
Tasmanian “ Wolf” (Thylacinus).

Figs. 44-48. Columella: 44, of House-Sparrow; 45, Nightingale; 46, Golden Eagle; 47, Wandering
Albatross (Diomedea exulans) ; 48, Nilotic Crocodile (adult).

N.B. The stapes of Homo, 39, is drawn on the scale maintained as the rule throughout these plates ;

so likewise are 46,47, and 48. The remaining stapedes are represented of the same size as fig. 39, for a

better comparison of their general forms.

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BIMANA, QUADRUMANA LEMURIDAE & CARNIVORA.

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PINNIPEDIA, RODENTIA, HYRACES & PROBOSCIDIA.

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r 499 |

XIX.—On the Annelida obtained during the Cruise of H.M.S. ‘ Valorous’ to Davis
Strait in 1875. By W. C. M°Invoss, ICD., LL.D., FRSSL.S #., PLS, §e.

(Plate LXV.)
Read June 21st, 1877.

FOR the opportunity of examining the collection made during the voyage of the
‘ Valorous,’ I am indebted, as on many similar occasions, to Dr. Gwyn Jeffreys, the
scientific chief of the expedition. The specimens were not quite in such good pre-
servation as those collected, for instance, in the dredging expeditions of H.M.S. ‘ Por-
cupine, and Dr. Jeffreys’s yacht ‘Osprey,’ since specially prepared bottles and spirit
were sent on these occasions. No group of marine Invertebrates is more difficult to
preserve in a satisfactory manner than the Annelida; and it may not be inopportune
again to draw attention to the necessity of pursuing, as far as possible, the method
indicated below *.

From the days of Otho Fabricius onward the Annelids of the Greenlandic seas have,
in comparison with those of other parts, received unusual attention. Thus, besides the
mention of between 40 and 50 marine Polycheeta by the foregoing author in 1780,
A.S. Cirsted published, in 1843, his ‘ Gronlands Annulata Dorsibranchiata,’ a work
illustrated by eight good lithographic plates. He described about three dozen species,
and calculated that at least fifty Greenlandic dorsibranchs were known. Moreover, in the
year 1818, it was in the same region (Baflin’s Bay) that Sir John Ross discovered the great
depth at which marine animals lived, by dragging up from 1000 fathoms a fine Astro-
phyton and mud containing “* Lumbricus tubicola” (? Hyalinecia tubicola, O. F. Miller).
The laborious papers of Dr. Malmgren of Helsingfors, which include the Greenlandic
collections made by O. Torell and Amondsen, give us, with the exception of the remarks
of Mobius (which I have not seen) the most recent enumeration of the Annelidan

* On removing the Annelids from the dredge or collecting-jar they should be immersed for some hours in large
flat vessels containing the strongest methylated spirit, or, if this is inconvenient, placed sparingly in capacious bottles
-of the same fluid. After the lapse of two or three hours they should be taken out of the large flat vessels and put
in separate bottles (few in each) of the same spirit. In the case of those immersed in the latter at first, the liquid
should be poured off and replaced by fresh spirit. It is during the first twenty-four hours that the greatest care is
necessary ; for if by inattention the specimens are allowed to soften, their subsequent preservation is always
unsatisfactory. The bottles should be systematically surveyed again within a day or two; and, as a rule, in any
case in which the colour of the spirit is deep or its transparency clouded, the fluid should be removed and fresh spirit
or ahsolute alcohol again added. In some collections the annelids are only represented by their cuticular coats
enclosing, as in a bag, a mass of pulp: much time is lost in specific determination, while for thorough investigation
they are worthless. Further, there is generally no difficulty in preserving the majority of the hardy forms alive for
many months in deep bottles half filled with sea-water, and, in the case of those which bore in sand, a layer of the
latter on the bottom. One or two specimens only should be placed in each jar; and all should be supervised daily
in hot weather, for the removal of the injured and dead and the renewal of the seawater.

500 DR. W. C. MSINTOSH ON THE ANNELIDA

fauna of the region. From the works of Dr. Malmgren the list published in the ‘ Arctic
Manual’ for 1875 was drawn up by Dr. Liitken, of the University Museum, Copenhagen.
The list comprises 101 species of marine Polychzta. The remarks on the Annelids of
Novaia Zemlia by Dr. Ehlers *, and those of the Siberian Seas by Prof. Grube +, show
that not a few forms have a wide range in the arctic waters {.

Though an accident which happened to the ‘Valorous’§ prevented the extensive
dredging operations contemplated by Dr. Jeffreys, and limited the species of marine
Polycheta to about 68, yet of this comparatively small number 13 are new to the
Greenlandic area, and, in addition, about 9 are new to science. Finally, so far as our
present knowledge of the Annelids goes, there is no doubt that Dr. Jeffreys’s opinion
that the Greenlandic Invertebrates are more European than American, is substantiated.

EUPHROSYNE BOREALIS, Cirst. Station 1, 70° 30’ N., 54° 41’ W., off Hare Island,
Disco, 175 fathoms, on a bottom of sandy mud. The specimens measured about an
inch in spirit, with 25 body-segments, besides the first and last. The branchize in the
rows agreed with the description of Prof. M. Sars (viz. 7 or 8 in each row), though
sometimes one or two were deficient. Some are simple, the majority bifid, and that
nearest the median line generally trifid. The bristles are much more prominent organs
than in #7. foliosa, Aud. & Kd.; and the spur on the smooth kind is much shorter. The
bifid bristles, again, have bolder transverse serrations. TZ. foliosa occurs abundantly
between tide-marks in Herm, whereas the arctic species frequents deep water, and
ranges to Spitzbergen and Finmark. No example has yet been found in British waters.
Amongst the bristles are many specimens of a small red mite.

The form of the branchize seems to be very characteristic in the British forms, viz.
EL. foliosa, Aud. & Kd., and HL. armadillo, Sars.

Nycuia crrrosa, Pallas. Fine specimens from Holsteinborg Harbour, from 7 to 35
fathoms. This form has a wide range, from British waters to Scandinavia, Iceland,
Spitzbergen, and the Gulf of St. Lawrence.

Nycu1a AmonpsEeNI, Malmgren. Station 3, 69° 31’ N., 56° 1’ W., sandy mud, 100
fathoms, and Godhavn Harbour, Disco, in 5-20 fathoms. A common species in the
Gulf of St. Lawrence, Canada.

Eunoa Girstepi1, Malmgren. Holsteinborg Harbour, in 35 fathoms. The specimens
measure an inch and three quarters long, and are larger than those from Canada. It
ranges to Iceland, Spitzbergen, and Finmark.

HarmMornok impricata, L. Station 4, 67° 50’ N., 55° 27’ W., 20 fathoms on shell-
sand; station 5, 66° 59’ N., 54° 27' W., 60 fathoms on sand and shells; Holsteinborg
Harbour, 7-35 fathoms; outside Godhayn Harbour, in 80 fathoms; station No. I.
H.MLS. ‘ Alert,’ 65° 00’ N., 53° 00’ W., 30 fathoms, on a rocky and gravelly bottom. This
ubiquitous form is abundant; and there is nothing peculiar in size or coloration; indeed

* Ann. Nat. Hist. 4th ser. vol. xi. p. 464.

+ Mém. de Savants étrangers (St. Petersburg), t. viii. 1858.

+ The forthcoming ‘ Report on the Annelida of the Austrian North-Polar Expedition, by Dr. Emil von Maren
Faller of Vienna, will afford further valuable information on the subject.

§ See “ Reports on the ‘ Valorous’ Expedition,” Proceed. R. 8. vol. xxv. No. 1738, p. 182. <A preliminary
notice of the Annelids also appears in the latter.

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 501

much larger examples occur in my collections from Exeter Bay, Greenland. Some
varieties approach Hvarne, but there is no difficulty in discriminating them.

EVARNE IMPAR, Johnston. Station 4, 20 fathoms; Station 6, 64° 5’ N., 56° 47' W.,
410 fathoms, in sandy mud; Holsteinborg Harbour, in 10 fathoms; Godhavn Harbour,
5-20 fathoms.

EUPoLyNoé occipENTALIS, M°I. Station 1, 175 fathoms, in sandy mud. The frag-
mentary specimen has scales, which were absent in the Canadian example*. In spirit
these have a ferruginous-brown hue, and the surface is almost smooth, the minute
spines being insignificant. On the outer border they have rather numerous, long, slender
cilia (PJ. LXV. fig. 1), the tips of which are very slightly dilated. Parasitic Pedicelline
abound on the bristles. The Lepidonotus sublevis of Prof. Verrill + may be this or an
allied form, but his description is too lax for determination.

EUS©HENELAIS ABYSSICOLA, n. sp. Station 16, in the middle of the North Atlantic,
55° 10’ N., 25° 58’ W., in 1785 fathoms, on a bottom of Globigerina-ooze. This is repre-
sented by a minute and fragmentary example, which at first sight was considered to be
a Leanira. In Sthenelais and Stgalion the ventral bristles are more or less boldly
bifid ; in Husthenelais, while the bristles lean to the former types in regard to terminal
segmentation and outline, yet many have perfectly tapered tips without a trace of the
secondary process. In the present form every one of the inferior bristles (Pl. LXV.
figs. 2 & 2a) has a filiform tip, but the upper group (fig. 2) more nearly approach the
characteristic terminal portions in Leanira; it needs but the shortening of the tips of
the processes and the better definition of their transverse markings to convert them into
those of an ordinary Leanira, for the same gradation between the superior and inferior
series of the ventral bristles is maintained in that group. Moreover, the steps
between Sigalion and Sthenelais on the one hand, and Leanira on the other, are
further indicated by the diminution of the serrations on the distal end of the shaft of the
bristle, such being distinct in Husthenelais hibernica}, very rudimentary in the present
form, and altogether absent in Leanira.

The injured fragment is devoid of scales and is eyeless. The palpi resemble those of
Eusthenelais hibernica; the tentacular cirri (and probably the tentacle also) are short.
_A few lobate papillee occur on the feet, as in the last-mentioned species, and the ventral
cirrus does not seem to reach the tip of the setigerous division.

PxHoto# minutTA, Fabr. 7-10 fathoms in Holsteinborg Harbour; 5-20 fathoms in
Godhavn Harbour. The specimens are large.

NEPHTHYS cHcA, Fabr. Station 4, 20 fathoms, on shell-sand; 10 fathoms in Hol-
steinborg Harbour; 5-20 fathoms in Godhavn Harbour. Many Infusoria are attached
to the bristles.

NEPHTHYS CILIATA, O. F. Miller. Station 1, 175 fathoms, off Hare Island, Disco, in sand
and mud; Station 3, 100 fathoms, in sandy mud; 10 fathoms in Holsteinborg Harbour,
5 to 20 fathoms in Godhavn Harbour, Disco. Swarms of a commensalistic form, re-
sembling at first sight a little Leech, occur between the feet, and quite differ from similar

-* Ann. & Mag. Nat. Hist. 1874, vol. xiii. p. 264. + Invert. Anim. of Vineyard Sound, p. 581.
+ Trans. Zool. Soc. vol. ix. pt. 7, p. 407, pl. 73.
SECOND SERIES.—ZOOLOGY, VOL. I. 3X

502 DR. W. C. MSINTOSH ON THE ANNELIDA

forms (Zoxosome) occurring in the British species in having a shorter pedicle, with a
larger basal portion for attachment, while the tip of the body is bluntly rounded and
presents only an aperture on one side.

PHYLLODOCE GR@NLANDICA, Cirst. Station 4, in 20 fathoms, on shell-sand; Godhavn
Harbour, 5-20 fathoms. The large specimens exceed in size those from British waters.

Eurawia viripis, O. F. Miller. 7-12 fathoms, Holsteinborg Harbour.

EULALIA PROBLEMA, Malmgren. Station 5, 66° 59’ N., 55° 27’ W., 60 fathoms, on
sand and shells. In spirit the specimen is dusky brown dorsally, and the lamelle are of

_the same hue; ventrally it is paler. The tips of the bristles are comparatively short,
and the enlarged distal end of the shaft finely spinulose. Dr. Malmgren received only
a single injured example from the same (western) shores of Greenland.

Hreone arctica, Malmgren. Station 5, 60 fathoms, on sand and shells. A minute
species, in which the proboscis (in the present examples) forms a clavate process, smooth
throughout the greater part under a lens, but toward the terminal dorsal region of the
partly extruded organ a distinctly papillose area occurs. In some there is a division of
the proboscis into a firm basal region and a more membranous distal portion. It seems
to come close to #. pusilla, the chief differences between the species, according to Dr.
Malmgren, being that in the latter the body tapers equally anteriorly and posteriorly,
and the head is longer than broad; whereas in E. arctica the body tapers less, and the
antero-posterior and transverse diameters of the head nearly correspond. The superior
lamella of the foot touches the next division in the anterior and central regions of the
body in #. pusilla, but is separate in #. arctica. ‘The anal cirri are elongate (linear-
fusiform) in the former, broadly ovate in the latter. The condition of the proboscis in
FE. pusilla is not noticed. In the present form the inferior lamella is not longer than
the bristle-papilla. It was formerly procured in Spitzbergen and Finmark.

ETEoNE FLAVA, Fabr. 5-20 fathoms, Godhavn Harbour, Disco.

ANCISTROSYLLIS GR@NLANDICA*, n. sp. A remarkable form, apparently allied to
the Syllide, was dredged at Station 6, 64° 5’ N., 56° 47’ W., in 410 fathoms, on
a bottom of sandy mud. ‘The specimen is about 3 of an inch long. The head
forms a short blunt cone, divided by a somewhat shallow fissure into two frontal
lobes (Pl. LXV. fig. 20), each of which has a short tentacle at the base, while
toward the posterior border another short conical tentacle occurs in the middle line.
Immediately below the great lateral eye on each side are two somewhat larger
tentacular processes, which resemble those on the adjoining segments of the body.
The latter is slightly tapered anteriorly, and more distinctly so posteriorly. The
segments are deeply cut, especially behind the anterior third and for some distance
along the wide part of the body. They are 57in number. Each foot, in the anterior
third, has a short dorsal and a smaller ventral cirrus (Pl. LXV. fig. 3), a single large
and characteristic hook, which somewhat resembles those on the first body-segment
of Paramphimome pulchella, Sars, placed above the former, and a slender longitudinally
striated bristle (generally enclosed in the tissues). The inferior division bears, in a promi-
nent lobe, a spine and a series of simple bristles, with a slight wing on one side—best

* dyxiorpov, hamus.

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 503

developed in the lower group, which also have a claw at the tip. The latter process is
indistinct in the delicately tapered superior series. The cuticle of the foot has many
cylindrical papillee; and similar organs occur on the dorsal and ventral cirri, those
on the latter, however, being less marked. The relationships of this form are not easily
made out; for while agreeing in some respects with the form of the head in such as
Microsyllis, it wholly differs in other points, so that it is only provisionally ranged
amongst the Syllide.

NEREIS ZONATA, Malmgren. Station 1, 175 fathoms, in sandy mud, off Hare Island,
Disco ; Station 8, 100 fathoms, on a similar bottom. In a large male example a peculiar
change occurs in the bristles; for besides the ordinary kind, toward the anterior third
of the body, a dense tuft of translucent bristles exists in each division of the foot. In
shape these resemble the tips of the Heteronereis forms, though they are only partially
developed, the tips alone projecting through the skin, while the shafts form a dense
mass in the neighbourhood of the spine. The distal division of these bristles (fig. 4)
resembles the blade of a sword, with a finely serrate edge, and tapering to an acute point,
which is slightly prolonged in a filiform manner. Moreover, a tendency to the foliaceous
condition of the feet accompanies the appearance of the Heteronereis-bristles, processes
appearing at the base of the dorsal cirri, a foliaceous growth at the inferior bristle-
papilla, and a process on each side of the ventral cirrus. Such, as usual, are best
marked posteriorly. The specimens are tinted with fine reddish-brown bars.

NEREIS PELAGICA, L. Station 4, 20 fathoms, shell-sand; Station 5, 60 fathoms, on sand
and shells; Station 6, 410 fathoms, in sandy mud; Godhavn Harbour, 5-20 fathoms;
H.M.S. ‘ Alert,’ Station 1, 65° 00’ N., 58° 00’ W., 30 fathoms, on a rocky and gravelly
bottom.

LUMBRICONEREIS FRAGILIS, O. F. Miller. Station 1, 175 fathoms, off Hare Island,
Disco; Stations 3 and 6, wt anted; Station 7, 63° 9 N., 56° 43’ W., 1100 fathoms, on a
bottom of mud; Station 16, in 1785 fathoms, in Globigerina-ooze; outside Godhavn
Harbour, 5-20 fathoms. This species, common in Britain, has hardly been differentiated
in the Catalogue of the. British Museum from the abundant southern type Notocirrus
tricolor, Mont.

NoTHRIA CONCHYLEGA, Sars. Stations 1, 3, 5, wt anted. The specimens are large and
in very firm tubes of small pebbles, very similar to those from the Gulf of St. Law-
rence. The British examples have tubes composed of flat bivalves, fragments of Echino-
derm-tests, &c.

GLYCERA CAPITATA, Cirst. Stations 4 and 7; Station 9, 59° 10'N., 50° 25’ W., in 1750
fathoms, and on a muddy bottom.

GLYCERA SETOSA, (rst. On the surface of the sea at Ritenbenk Kulbrud, Waigat
Strait; a fragment from Station 4. This form is readily distinguished by the great
length of the feet and bristles and by the structure of both. The nerve-trunks are
rounded and more distinctly separated than usual in the group. The longitudinal
ventral muscles do not seem quite to cover the trunks in the soft preparation, though
a thin layer of tissue does.

3x 2

504 DR. W. C. MSINTOSH ON THE ANNELIDA

ARICIA GRENLANDICA, n. sp. (Pl. LXV. figs. 5-9). A fragmentary specimen (anterior
region) was dredged at Station 4, in 20 fathoms. Many of the examples of the genus
procured elsewhere by the dredge are in the same condition (7. e. represented by the
anterior region), while those procured by digging are not uncommonly devoid of the
same part. The general form of the head and body corresponds to that of its allies,
and especially to Aricia Kupfferi, Ehlers *, a form described from the collection of the
‘ Porcupine.’ It is at once distinguished, however, from the latter, 4. Cuvieri, and
the abundant Scotch 4. Latreillic by the absence of the pectinate rows of papillee on the
ventral surface. The snout is elongated and pointed, and a little pigment exists at the
base of the region dorsally, but there is no evidence of distinct eyes in the preparation.
The lateral folds of the mouth turn smoothly round to form the posterior border of
the first bristle-bearing segment. The succeeding sixteen bristle-bearing feet (2nd to
17th) constitute the anterior region, characterized by the pectinate rows of papille
along the lateral border of the foot and the numerous strong bristles. The papille are
few in number, generally about six, and the upper is considerably longer than the others :
ail are simple, conical, hypodermic papille. The stout bristles of the inferior division of
the foot are somewhat like those of M. Claparéde’s 4. feetida, consisting of thick shafts
with peculiarly curved tips (Pl. LXV. fig. 5a), the striated brownish region ending in a
fine point, beyond which the translucent terminal wings proceed to the tip. The edges of
the latter (fig. 5, seen from behind) are finely serrated. Besides the foregoing, there are
numerous short tapering bristles of the same type as the dorsal fascicle, and a few large
hastate brown spines (fig. 6), which posteriorly sometimes project in a conspicuous
manner. The dorsal fascicle consists of a series of stout tapered bristles, having the
characteristic transverse markings so common in the group, and apparently due to
minute rows of spines transversely arranged. The tips of the bristles are much attenu-
ated, and the distance between the transverse rows (which become mere lateral serrations)
increases with the diminution of the bristle.

After the form of the foot alters, the bristles of the dorsal fascicle are much more
slender and the distal spinous rows are further separated. Amongst these are some shorter
forms (also with spinous shafts) which terminate in a fork, one of the prongs being
longer than the other and both slightly thickened at the tip (fig. 7). The web of the
fork has fine chitinous hair-like processes. A distinct curve of the shaft occurs below
the fork. The ventral bristles are much larger, with evident serrations and finely
tapered tips, the peculiar narrowing which takes place being less easily noticed than n
A. Latreillvi and others.

The branchize commence on the 5th bristled segment (fig. 9).

SCOLOPLOS ARMIGER, O. F. Miller. Stations 4 & 7, ut antea; Holsteinborg Harbour,
30 fathoms.

NAIDONEREIS QUADRICUSPIDA, (rst. A fragment from Station 7, 1100 fathoms, on a
muddy bottom.

AMMOTRYPANE AULOGASTER, H. Rathke. 5-20 fathoms, in Godhayn Harbour.

* Zeitsch. f. w. Zool. Bd. xxv. 1, p. 57.

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 505

OPHELIA Limactna, H. R. Station 4, 20 fathoms, in shell-sand. Of good size, but
not larger than at St. Andrews.

TACHYTRYPANE JEFFREYSII, n. sp. (Pl. LXV. fig. 10). Station 9, 59° 10'N., 50° 25’ W.,
1750 fathoms, on a bottom of mud. The body of this species is elongate, being upwards
of 2 inches long in spirit and about +5 of an inch in breadth, rounded in front, but marked
by the usual ventral ridges throughout the rest of its extent. There are about thirty
segments, besides the first and last, the anterior being short, those in the middle very
long. The head forms a short cone, with a minute filiform process at the tip. No
bristles are visible under a lens, the body being smooth and iridescent, like that of Lino-
trypane apogon, for the fine transverse lines are not conspicuous; very minute simple
bristles are, however, present. There are no cirri. The last caudal segment is separated
by a well-marked furrow from the rest of the body, and terminates in a slender recurved
process bent downward and forward. In the structure of the body-wall Tauchytrypane
Jeffreysti forms an intermediate link between Linotrypane and the other forms, con-
necting both with Ammotrypane and Ophelia. The cuticle (fig.11) is greatly thickened
(as in Limotrypane), and a special process passes inward from the median line on the
ventral surface toward the nerve-cords; it is further marked by fine cross strie. From
each side of the nerve-trunks the oblique muscle proceeds to the body-wall, and cuts off a
segment of the longitudinal muscular layer. The intestinal canal is filled with GJodi-
gerina-mud. .

AMMOTRYPANELLA ARCTICA, n. sp. Station 16, 1785 fathoms, on Globigerina-ooze.
A somewhat small form, measuring about ? of an inch long, and more slender than a
young Ammotrypane aulogaster of the same length, while the contour is different. The
cephalic lobe (Pl. LXV. fig. 12) especially diverges, being bluntly rounded, and furnished,
on each side of the base dorsally, with a considerable pigment-speck. The succeeding
segments are somewhat narrow, as in allied forms (e. g. Ammotrypane), and each has a
tuft of simple silky bristles, which, in a dorsal view (as in fig. 12), project prominently
outward and backward in the first seven or eight segments. The following twelve or
thirteen segments are larger, but their bristle-tufts are minute, and being sunk in the
groove opposite the attachment of the oblique muscle, do not show laterally in ordinary
views. ‘Toward the posterior end the bristles again become somewhat more distinct, and
slender cirri still remain on several feet. The caudal segments are short, and the last is
funnel-shaped and terminates ina smooth rim, The ventral sulcus commenees anteriorly
at the buccal orifice, and is well marked in front of the caudal process. The cuticle is
dense. The condition of the specimen is unfavourable for minute work. The total
number of segments seems to be about thirty.

A still more minute example comes from the same station. It presents no cirri;
but the state of the specimen (which appears to have been dried) is unfavourable for
the preservation of such organs. A papilla, considerably larger than in 4. aulogaster,
exists at the tip of the snout. The bristles are long in front of the caudal process,
which is cylindrical and appears to have an even rim. This may be an injured young
example of 4. aulogaster, though the snout and contour differ.

506 DR. W. C. M‘INTOSH ON THE ANNELIDA

Like Tachytrypane, the two foregoing had the intestinal canal filled with fine Globi-
gerina-rud.

TRAVISIA GLANDULOSA, n. sp. (Plate LXV. figs. 15, 16).

From the same depth (1785 fathoms, Station 16) is a Travisia, which, while agreeing
in form, presents certain differences from the ordinary 7. Forbesit, Johnst. The example
has been partially deprived of cuticle. The greater part of the basement tissue of the
body-wall is marked by prominent rows of punctures or dots, resembling those in the
Sipunculide. In the central region a larger row runs transversely across each seg-
ment, towards the posterior border. In comparing these with similar specimens of
T. Forbesii, the markings are much more uniform in the latter; and though here and
there a large papilla occurs, there is none of the uniformity characteristic of the new
form.. The dotted appearance is caused by the projection of the papille inward, as
observed in transverse section (Pl. LXV. fig. 16); and each has a tubular connexion
with the outer surface, like that noticed in the ventral papille of Aphrodita and Her-
mione. The cirri near the tail are much shorter and less distinct than in 7. Forbesi.
The specimen is so imperfect that some hesitation is felt in naming it; but the structure
of its body-wall is very different when contrasted with 7. Forbesii.

ScALIBREGMA INFLATUM, H. R. A fragmentary (small) example in 80 fathoms, out-
side Godhavn Harbour.

TropHonta PLUMOSA, O. F. Miller. A fragmentary specimen, in 175 fathoms, off
Hare Island, Disco (Station 1). The inferior hooks have their points somewhat less
curved than in the ordinary British forms, and the mud-and pebble-coated warts are a
little coarser.

FLABELLIGERA AFFINIS, Sars. Holsteinborg Harbour, 30 fathoms ; Godhavn Harbour,
5-20 fathoms. The specimens are considerably larger than those usually met with in the
British seas.

ScOLECOLEPIS CIRRATA, Sars. Station 1, 175 fathoms; Station 6, 410 fathoms. The
fragment is large.

From the first-named station is also the anterior region of a form which resembles
the foregoing in general outline. The head, however, instead of presenting the pair of
large, brownish, and somewhat faleate eyes in front of the median (occipital) tentacle,
exhibits only two small brown specks, placed close together. The foliaceous dorsal
pinne are also larger; and no trace of branchie occurs on the fragment, which con-
sists of about twelve segments. The bristles present no feature of interest.

ScoLECOLEPIS (?) JEFFREYSII, n.sp. Station 4, 20 fathoms, on shell-sand. This is repre-
sented by a fragment (about } of an inch in length) of the anterior end, and is closely
allied to Scolecolepis, but distinguished by the form of the snout, the arrangement of
the branchize, and other particulars. The head (Pl. LXV. fig. 13) is bluntly pointed
and divided into three regions by two antero-posterior curved lines, which are marked
with dark pigment posteriorly, and therefore very distinct. he central region is also
tinted with dark pigment posteriorly ; and in front of this is the short and somewhat
clavate tentacle, which has a constricted base, the whole rising from an elevated portion
of the snout. The body is flattened from above downward, and narrowed posteriorly

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 507

in the fragment. The first segment bears only the dorsal and ventral bristle-tufts ;
the second has, in addition, a dorsal cirrus; while the fourth possesses a well-marked
branchial process. The branchize remain of considerable size (¢. e. nearly meet in the
middle of the dorsum) for about eight segments, and then diminish to the fifteenth, in which
they are very minute. The bristles are simple striated structures, with a slight wing, as in
S. cirrata; and they increase in length so as to project outward and backward at the pos-
terior part of the fragment. In transverse section (Pl. LXV. fig. 14) the body-wall presents
(at the end of the fragment) an ovoid outline. The cuticle is well marked, and the hypo-
derm thick, especially along the entire ventral surface. The dorsal longitudinal muscles
are considerably less than the ventral, and a large gap occurs in the median raphe.
The ventral longitudinal muscles are separated by a central interval, at the bottom of
which the nerves lie. The oblique muscles pass over to the raphe at the summit of the
latter. The circular muscular coat is not much developed, but the lateral vertical bands
supplement it. The alimentary canal is rounded, has a thick circular muscular coat,
and the glandular inner surface is richly ciliated.

PrionosPio STEENSTRUPI, Malmgren. Fragmentary examples from Stations 3 & 16.
The stout anterior bristles show peculiar dotted markings, which are also present in the
short and curved inferior bristles of the median segments. The hooks have more than
one tooth above the great process.

CIRRATULUS crRRaATUS, O. F. Miller. Holsteinborg Harbour, 30 fathoms.

CAPITELLA CAPITATA, Fabr. Station 3, 20 fathoms. An injured fragment of a small
example.

MaLDANE Biceps, Sars. A single injured fragment in 175 fathoms, off Hare
Island, Disco (Station 1).

Ma.pane Sarst, Malmgren. Holsteinborg Harbour, 30 fathoms. Fragment.

Roprneé Lovent, Malmgren. Holsteinborg Harbour, 30 fathoms. A fragment in a
tube sparsely coated with clear quartzose sand-particles and rather brittle.

NIcOMACHE LUMBRICALIS, Fabr. Station 1, 175 fathoms, off Hare Island, Disco. The
size, like that of the Canadian examples, is large. The tubes are composed of sand or
sandy mud. on

AXIOTHEA CATENATA, Malmgren. Godhavn Harbour, 5-20 fathoms; and a fragment
from Station 1. Tubes composed of sandy mud. .

PRAXILLA ARcTICA, Malmgren. Station 7, 1100 fathoms, in mud. The species is
smaller than P. pretermissa, and the hooks agree with those figured from a Zetlandic
example in Trans. Roy. Soc. Edinb. xxv. pl. xvi. fig. 18. The tubes are composed of
very fine sand, and have a hyaline inner coat.

OwENIA FILIFoRMIS, D. Ch. Station 5, 60 fathoms; Station 16, 1785 fathoms; out-
side Godhavn Harbour, 80 fathoms. The tubes at the latter station closely resemble
the stalks of Alcyonidium, growing in proximity, while a fragment from 1785 fathoms
has a tube composed of Globigerine united by the tough secretion.

CISTENIDES GRANULATA, L. Holsteinborg Harbour, 10 fathoms; Godhavn Harbour,
5-20 fathoms. The tubes offer a noteworthy character, since they are in most cases con-

508 DR. W. C. MSINTOSH ON THE ANNELIDA

structed of transparent quartzose sand-particles set in the opaque secretion, so that, at
first sight, they seem pitted all over.

CISTENIDES HYPERBOREA, Malmgren. Station 1, 175 fathoms, off Hare Island, Disco.
The tubes of this species generally present a darker hue and less regular masonry. The
cement in both is more abundant than in Pectinaria belgica or Amphictene auricoma, so
that the architecture in the latter is more finished.

AMAGE AURIcULA, Malmgren. Station 4. A small fragment seemed to approach
Samytha sexcirrata, Sars, but its condition negatived satisfactory results.

AMPHITRITE CIRRATA, O. F. Miiller. 5-20 fathoms, Godhavn Harbour, Disco.

NicoLEA ARcTICA, Malmgren. 5-20 fathoms, Godhavn Harbour. The specimens are
larger than the following, and the pedicles of their branchiz are longer.

NICOLEA ZOSTERICOLA, Cirst. & Gr. Holsteinborg Harbour, 30 fathoms.

LEZNA ABRANCHIATA, Malmgren. Stations 5 & 6. A fragment also from the latter
presents certain affinities with Lanassa Nordenskidldi, Malmgren; but its condition is
such that a degree of doubt remains. The body is grooved along the ventral aspect
behind the ventral scutes. The bristles resemble those shown by Malmgren in Laphania
Boecki, that is, they have wings at the tip, and the latter in some is short and broad.
The hooks, again, have elevated crowns, with about 5 teeth above the large process, as
in Lanassa.

THELEPUS CIRCINNATUS, Fabr. A small example from Station 5; fragment from Hol-
steinborg Harbour, 10 fathoms; large specimens from Station 1 (30 fathoms), 65° 00' N.,
53° 00' W., H.M.S. ‘ Alert.’ It is well to remember that the posterior hooks show two
teeth above the large fang.

TRICHOBRANCHUS GLACIALIS, Mgrn. Station 12, 1450 fathoms, on a bottom of Glo-
bigerina-ooze and stones. ‘The tubes are chiefly composed of Globigerine.

TEREBELLIDES StrRemiI, Sars. Station 6, 410 fathoms; Station 7, 1100 fathoms;
Godhavn Harbour, 5—20 fathoms; and outside the latter in 80 fathoms.

SABELLA CRASSICORNIS, Sars. Fragments from Godhavn Harbour appear to belong
to this species. The occurrence of the branchiz in the tube showed that they did not
pertain to S. pavonia, as first supposed. The present species and S. spetsbergensis, Mgrn.,
seem to be closely allied, but the condition of the specimen was such that absolute accu-
racy was not attainable. The pigment on the branchie resembles that found in the
latter form. The tube is thin and tough.

EUCHONE ANALIS, Kroyer. Station 4, 20 fathoms on shell-sand ; Holsteinborg Harbour,
10 fathoms. One was in a fine hyaline tube coated with coarse quartzose sand and
minute fragments of shells.

CHONE INFUNDIBULIFORMIS, Kroyer. Station 4; 5-20 fathoms in Godhavn Harbour.
The hooks (Pl. LXV. fig. 17) differ from Dr. Malmgren’s figure. One was in a tough
brownish tube. .

Another Chone, evidently belonging to a different species, was dredged in the latter
locality, but it had apparently been dried before being placed in spirit. The collar is
large and of a similar form to that in C. infundibuliformis. The branchie (which are

OBTAINED DURING THE CRUISE OF H.M.S. ‘ VALOROUS.’ 509

about twenty in number on each side) are fragmentary, and appear in process of develop-
ment, as they scarcely exceed in length the diameter of the body. The long anterior
crotchets (Pl. LXV. fig. 19) slightly differ (in curves) from those of C. infundibuliformis,
while the posterior hooks (fig. 18) still more diverge, a comparison with one from the
same region in the former being made by referring to fig. 17. Whether this is the
Chone Duneri, Mgrn., or another species, the brief remarks and somewhat unsatisfactory
figures given by the author prevent determination. The posterior hooks especially
disagree.

The remarkable differences in the structure of the muscular system which distinguish
Euchone and Chone, on the one hand, from Sabella and Dasychone on the other, are like-
wise associated with corresponding differences in regard to the arrangement of the nerve-
cords. The fasciculi of each dorsal and ventral longitudinal muscle in Chone infundi-
buliformis form a complex double spiral in transverse section; while in Huchone the
dorsal also form a double spiral, less elaborate than in the foregoing, and the ventrals
each form asimple spiral. The nerve-trunks in both are much more closely approximated
than in Sabella and Dasychone.

SrrpuLta? A fragmentary tube of a minute, shining, quinquecostate species, with four
transverse ridges anteriorly, and irregularly twisted; from Station 11, 1860 fathoms.

DirRYPA GRENLANDICA, n.sp. (Proc. Roy. Soc. vol. xxv. no. 178, p. 219, woodcuts 1 & 2.)
Station 12, 1450 fathoms, on G‘lobigerina-ooze and stones.

A. Tube of Ditrypa grenlandica, enlarged. 3. The aperture, still more highly magnified.

The tube is about half an inch in length, not much thicker than a thread, and curved
from end to end like a bow (see woodcut, fig. 1,4). It tapers very gradually from the
anterior to the posterior end, contrasting in this respect with the more decided diminu-
tion in D.arietina. The oral aperture (fig. 1, B) forms a smooth and slightly constricted
rim, which is narrower than the tube almost by the depth of the ridges. The latter are
eight in number, and run from the anterior to the posterior end of the tube, though, it °
must be added, none of the specimens are quite perfect. It differs from any other
Ditrypa known to me in its slender form and the well-marked longitudinal ridges. In
the most perfect specimen the latter coalesce to some extent posteriorly, so that the tip
of the tube is somewhat quadrangular.

SPIRORBIS BOREALIS, Daud. Godhavn Harbour, 5-20 fathoms. Tube only.

SPIRORBIS VERRUCA, Fabr. Station 5, 60 fathoms. Tube only.

SPIRORBIS SPIRILLUM, L. Station 5.

SECOND SERIES.—ZOOLOGY, VOL. I. 3Y

510 DR. W. C. MSIINTOSH ON THE ANNELIDA

A remarkable tube (woodcut, fig. 2) occurred at Station 16, 1785 fathoms, amongst
Globigerina-ooze. It consists of a minute
tube (about the thickness of a slender silk
thread) of fine greyish mud, and having an
enlargement at one end, the blunt tip being
partly formed by a Globigerima-test. The
enlarged portion is tufted with what at first
sight (under a lens) appear to be minute
and finely ramose alge; but the micro-
scope shows that these delicately branched
processes are composed of the same elements This (ann Sinton 106), bey ee ahs
as the tube.

The Nemerteans are represented by a species from Stations 4 & 5, Holsteinborg Harbour
and Godhavn Harbour. The colour in spirit is brownish purple on the dorsum, whitish
beneath. The short body and large proboscis distinguish it from Nemertes Neesii; but it
may be related to the Amphiporus grenlandicus, Girst.—the more so, as the pigmentary
portion of the skin is apt to fall off in spirit-specimens. The longitudinal muscular coat
is cut into numerous and somewhat regular fasciculi by the radiating fibres. The base-
ment-tissue outside the circular coat is largely developed. The proboscis conforms to
the typical structure. If Mirsted’s NV. Neest was like this form, the British species widely
diverges. Another species, probably a Tetrastemma, comes from Station 5, but its con-
dition prevents accurate determination.

DESCRIPTION OF PLATE LXV.

Fig. 1. Portion of the outer edge of an anterior scale of Eupolynoé occidentalis. x 210 diam.

Fig. 2. One of the upper series of the ventral bristles of Husthenelais abyssicola. 2a, another of the
same series more closely resembling those in Sthenelais. x 350 diam.

Fig. 8. Foot from the anterior third of Ancistrosyllis grenlandica. x 210 diam.

Fig. 4. Heteronereis-bristle from a posterior foot of Nereis zonata, Mern., g. It is not completely
developed, and only the tips project beyond the fleshy part of the foot. x 210 diam.

Fig. 5. Anterior strong bristles (from the lateral pectinate row) of Aricia grenlandica, seen from behind.
5 a, another from the same part, viewed laterally. x 350 diam.

' Fig. 6. Large bristle from the anterior papillose feet (inferior division) of the same form. x 210 diam.

Fic. 7. Two of the bifid bristles from the region behind the pectinate feet (in the same animal). The
curve of the shaft is indicated in that on the right. x 350 diam.

Fig. 8. Tip of the inferior group of bristles after the change of type. The peculiar narrowing of the
organ towards the tip is apt to escape notice. x 350 diam.

Fig. 9. The 20th foot of the same species (not well preserved). @, branchial process. x about 20 diam.

Fig. 10. Tachytrypane Jeffreysti, nu. sp. Enlarged under a lens.

Fig. 11. Transverse (vertical) section of the body-wall of the same, about the anterior third. The speci-

men was not well preserved. x 20 diam.

OBTAINED DURING THE CRUISE OF H.M.S. VALOROUS. 511

. Ammotrypanella arctica, nu. sp. Enlarged under a lens.
. Scolecolepis Jeffreysii, n.sp. Enlarged under a lens.
. Transverse section of the anterior region (shortly behind the branchiz) of the same. The right

side is somewhat distorted from imperfect preservation: em, vertical muscle; om, oblique
muscle; dm, dorsal longitudinal muscle; ac, alimentary canal ; v, blood-vessel ; n, nerve-cords.

x 90 diam.

. Papille on the inner surface of the cuticular tissues of Travisia glandulosa. x 90 diam.
. Large papille of one of the transverse rows, as seen in transverse section of the body-wall of the

same. x 350 diam.

. Hook from posterior third of Chone infundibuliformis, Kroyer, from Godhayn. x 350 diam.
. Hook from the posterior third of the other Chone. x 350 diam.

. Crotchet from the anterior region of the latter species. x 350 diam.

. Anterior end of Ancistrosyllis grenlandica. Enlarged under a lens.

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XX. On the Sacral Plexus and Sacral Vertebre of Lizards and other Vertebrata.
By St. Gtorce Mivarrt, Sec. L.S8., and the Rev. RoBert CuaRKE, /.L.S.

(Plates LX VI., LX VII.)
Read May 3rd, 1877.

INTRODUCTORY REMARKS.

It has of late been recognized that in any attempt to answer the question, which
vertebra of any lower animal answers to the first sacral vertebra of Man, the nervous no
less than the osteological relations of the parts should be carefully investigated; and it
has been considered that the nervous rather than the osteological relations should be
deemed the more important—in fact it has been sometimes assumed that the nerves
must be taken as the fixed points, and that the bones must rather have their homology
decided by the nerves than vice versd. Should it be possible to show that in any group
of reptiles both the nervous and the osteological relations of any vertebra constantly
agree with the nervous and osteological relation of Man’s. first sacral vertebra, the
homology between two such parts may well be taken as thereby established; but if
either of these sets of relations exhibit discrepancy, then of course such homology
cannot be considered satisfactorily determined. Nor can we justly set aside osteo-
logical in favour of nervous resemblances if it should turn out that the nerves
themselves exhibit notable variations of condition as we pass from one allied form to
another—a@ fortiori if there should be variations in this respect even within the limits
of a species. It might surely be anticipated that more or less variation would be found
to exist in nervous as well as in skeletal structures; and in the event of such
anticipations being justified, the determination of sacral homology must depend upon a
comparison of the values of the conflicting claims of differing degrees of resemblance in
both the osseous and nervous systems—unless we prefer to consider the osteological
sacrum and the nervous sacrum as distinct structures, which may or may not com-
pletely coincide, and may or may not widely diverge. .

Professor Gegenbaur, in his “ Beitrage zur Kenntniss des Beckens der Vogel”, has
come to the conclusion? that there is such a complete agreement between the anterior
of those two Saurian vertebree the expanded and elongated transverse processes of which
abut against the ilium and the first sacral vertebra of Man. In both such transverse
process abuts against the ilium, and in both (as he represents it) such vertebra is that
one which is directly postaxial to the issuing forth of the most preaxial of those nerves
the fibres of which go to the sciatic plexus and not at all to the lumbar plexus.

With respect to the second vertebra abutting against the ilium, however, he represents

1 See the ‘ Jenaische Zeitschrift, Band vi. Heft 2.
* As shown by his diagram fig. 2, at page 201.

514 MESSRS. MIVART AND CLARKE ON THE SACRAL

an osteological but not a nervous agreement with Man’s second sacral vertebra. The
divergence consists in this, that whereas in Man the nerve issuing preaxially to every
sacral vertebra, except the last, goes entirely to the sacral plexus, Professor Gegenbaur
represents the nerve in Saurians which issues preaxiad of the second ilium-joining
vertebra as only contributing but a very small part of its fibres to that plexus. The
arrangement thus generally indicated by him is supported by descrip-

tions of the nervous conditions which he found to exist in a certain Fig. 1.
number ef different species.

Professor Hoffman! has also given brief notices and rough diagrams
of the arrangement of the lumbar and sacral plexuses in a variety of
Batrachians and Reptiles.

These two authors, however, do not agree in their account, nor
have we been fortunate enough to entirely agree with either of them ;
and, indeed, the discrepancies manifested by the species we have
examined have seemed to us somewhat noteworthy, and such as to
render it desirable that we should go somewhat further into detail as
to nerve-distribution than has been done in the memoirs referred to.

Amongst the Lacertilia, Professor Gegenbaur has investigated the Veyves of Lizard (from
composition of the lumbar and sacral plexuses in Chameleo vulgaris, Gegenbaur).
Lacerta viridis, Lacerta agilis, Uromastix spinipes, and Grammato- er, crural; 1s, ischi-
phora barbata. Professor Hoffman has examined it in Chameleo, atic nerves; “other
Monitor, Gecko, Lacerta viridis, Iguana tuberculata, Urothropus, and i.
Polychrus marmoratus. We have made use of, for examination, Cha-
meleo vulgaris, Monitor arenarius (two specimens), Lacerta viridis, Teius tequexin, Iguana
tuberculata (two specimens), Agama colonorum, and Grammatophora barbata.

In our descriptions we shall make use of the following nomenclature :—The nerve
coming out in Lizards between the elongated transverse processes which abut against
the ilium we shall speak of as the intersacral nerve ; the nerve issuing out in front of
the more preaxial of these elongated transverse processes we shall call the first presacral
nerve; the nerve still more preaxial we shall name the second presacral, and so on.
The nerve issuing out postaxial to the more postsacral of the two elongated transverse

processes we shall call the jirst postsacral nerve; the nerve still more postaxial we shall
call the second postsacral, and so on.

NERVES OF THE LUMBAR, SACRAL, AND PRECAUDAL REGION OF LACERTILIA.
Chameleo vulgaris. (Plate LXVL. fig. 1.)

According to Professor Gegenbaur?, the sciatic nerve in this animal appears to be
formed by three roots, one intersacral and two presacral, the first presacral being the
largest. The intersacral nerve gives off branches to the postsciatic nerves, while the second
presacral nerve bifurcates, giving one branch to join the third presacral, so forming the
crural nerve, which thus has but two roots.

' See ‘ Niederlindisches Archiy fiir Zoologie,’ Band iii. Heft 2, p. 143 (1876). * L.c. pp. 199, 200.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 515

Professor Hoffman! finds the sciatic nerve of the Chameleon to be formed by four roots,
having, in addition to those given by Gegenbaur, a postsacral root also, formed by a
branch from the first postsacral nerve. He also represents the nerve formed by branches
from the second and third presacral nerves (and thus having but two roots) as dividing
into two important branches, one the obturator nerve, the other the crural.

In our specimen of Chameleo vulgaris we found that the sciatic nerve might be said
to have either three or four roots, according as the branch (41a) connecting together
the first and second postsacral nerves should be considered a branch going from the
former to the latter, or from the second postsacral nerve to the first one. But our
specimen differed importantly both from the descriptions of Gegenbaur and Hoffman in
that, in the first place, the crural (15) and obturator (5) nerves were formed by three
roots instead of two, and, in the second place, that the position of all the roots of the
sciatic and crural nerves was different.

According to both Gegenbaur and Hoffman the sciatic nerve in this animal is formed
by the intersacral and the two presacral nerves, with or without the addition of fibres
from the first postsacral nerve, while of these roots the first presacral is the largest.
We, however, found the sciatic nerve (33) to be formed by the intersacral (29), first
presacral (12), and first postsacral (80) nerves; while of these roots the intersacral was
the largest. Again, both Gegenbaur and Hoffman found the first presacral nerve to be
undivided and to go wholly to the sciatic plexus, while we found the first presacral nerve
to bifurcate, one branch (12) descending to the sciatic, the other (11) ascending to the
lumbar plexus.

According to both Gegenbaur and Hoffman this lumbar plexus is formed from the
second and third presacral nerves, while we found it to be made up from the firsts
second, and third of these nerves (2, 9, and 11). Having united into a single trunk, we
found that this trunk almost immediately divided (as represented by Hoffman) into the
crural (15) and (5) obturator nerves, whereof the latter traversed the small pubic
foramen, called obturator foramen by Professor Hoffman?.

Lacerta viridis. (Plate LXVI. fig. 2.)

Professor Gegenbaur represents the sacral plexus of Lacerta as formed of three roots,
the middle one of which (that formed by the first presacral nerve) is the largest, and
sends no branch to the lumbar plexus; while the second presacral nerve bifurcates,
sending one branch to the last-mentioned plexus and one to the sacral plexus. The
lumbar plexus is formed by but two roots, one being part of the second presacral nerve,
and the other part of the third presacral nerve.

Professor Hoffman® quite agrees as to Lacerta viridis with Professor Gegenbaur, save
that he represents.no nervous connexion whatever between the intersacral nerve and the
first postsacral.

In one specimen of this species we found four roots to the sciatic nerve, there being a
distinct contribution to it from the first postsacral. Moreover we found as many as
three roots to the lumbar plexus, the fourth presacral nerve (2) concurring in its

1 Z.¢. pp. 174, 175, fig. 6. * L.c. tab. xi. figs. 9 and 10, fo. $ L.c. pp. 175, 176, fig. 9.

516 MESSRS. MIVART AND CLARKE ON THE SACRAL

formation with the whole of the third (9) and the preaxial bifurcation (11) of the
second presacral nerve. ‘The crural (15) nerve (that passing over the brim of the
pelvis) we found to be entirely formed by the preaxial part of the bifurcation of the
second. presacral, giving off, moreover, a filament to join the obturator nerve (5), which
has thus three roots :—1, the filament just mentioned; 2, the third presacral nerve; and
3, the fourth presacral nerve. The disagreement between the conditions found by us
and those described by the authors referred to was thus less striking in Lacerta than in
Chameleo. The sacral plexus was formed from the lumbo-sacral nerve (12), the first
presacral (21), the intersacral (29), and the first postsacral (30).

Teius tequexin. (Plate LXVI. fig. 3.)

A specimen of Teius teguexin was (with other specimens, as will be hereinafter
mentioned) most liberally placed at our disposal by Professor Flower, the Curator of the
Museum of the Royal College of Surgeons, from the stores of that Institution.
Unfortunately, in spite of its magnificent external appearance, it proved to be, internally,
in a state very unfavourable for dissection, while it was the first of all save one
examined by us. We cannot therefore speak with confidence as to the number of nerves
which contributed to form the lumbar plexus; but we did not observe that more than
two did so, the third (9) and part of the second (11) presacral; but we are certain that
three nerves contributed to the sacral plexus in addition to the lumbo-sacral nerve (12)
or postaxial bifurcation of the second presacral. The first postsacral (80) certainly
contributed to it, as well as the first presacral (21), thus differing from the conditions
described and figured by Professor Gegenbaur; while of the sacral roots the ¢ntersacral
(29), and not a presacral root, was the largest.

Grammatophora barbata. (Plate LXVI. fig. 4.)

This species is described by Gegenbaur! as only differing from his other Lizards in the
equality of the two branches of the bifurcating second presacral nerve. Its sacral nerve
therefore, according to him, is not reinforced by the first postsacral nerve.

In our specimens we did find it so reinforced, the sciatic nerve having thus four roots,
of which the first presacral (21) is (as Gegenbaur says) the largest. The lumbar plexus
we found to be formed, not by two but by three nerves—the fourth (2), third (9), and
second (11) presacral, the last bifurcating, and so giving off (12) the lumbo-sacral nerve
(or most preaxial root of the sciatic) as well as (11) the most postaxial root of the crural ;
but the two last-mentioned nerves we did not find of equal size, and the lumbo-sacral was
the more slender. On the right side we found the three roots of the lumbar plexus all
unite together, and then give off two branches, viz. the obturator (5) and crural (15)
nerves. On the left side, however, the fourth and third presacral nerves united and then:
gave off two branches; while the preaxial bifurcation of the second presacral also
bifureated, its bifurcations joining the two branches just mentioned, and then running
on as the obturator (5) and crural (15) nerves.

1 L.c. p. 200.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 517

Agama colonorum. (Plate LXVI. fig. 5.)

This animal was the only one found by us to present that character figured’ by
Professor Gegenbaur, and described by him as existing in all Lizards”, namely, the non-
junction with the sacral plexus of the first postsacral nerve (80). The most postaxial of
the sciatic roots was the intersacral nerve (29), which gave off one or two branches (411),
going to the cloaca and tail respectively. The first presacral nerve (21) formed the
stoutest root; but besides a rather stout lumbo-sacral nerve (12) (forming the postaxial
bifurcation of the second presacral) the sacral plexus also received a slender branch (46)
from the third presacral, which bifurcates at its origin like the second presacral. In
no other Lizard whatever have we found the sacral plexus receive a branch from the
third presacral. Professor Gegenbaur mentions no such condition, while Professor
Hoffman has only found it? in one specimen of Polychrus marmoratus. This was not
the case in another specimen‘ of his, thus showing (what our own dissections have
convinced us of) that two individuals of the same species may present considerable
nerve-differences. The sciatic nerve (38) gave off, as usual, a branch high up to the
obturator internus muscle (28).

In Agama the lumbar plexus was formed by three roots. The most preaxial was the
fourth presacral nerve (2); the second root consisted: of the greater part of the third
presacral nerve (9), the rest of that nerve (46) going to the sacral plexus. The most
postaxial root consisted of a slender branch of the second presacral nerve (11). This
slender branch passes over the before-mentioned outer slender branch (46) so as to forma
marked cross on each side of the body. The obturator nerve (5) was principally formed
by the fourth presacral (2), but received an accession from the third presacral; it gave off
a branch (7) to the iliacus. The crural nerve was formed mainly from the third presacral
nerve (9), but received a branch from the fourth presacral and also the slender branch
before mentioned (11) from the second presacral.

Iguana tuberculata. (Plate LXVII. fig. 1.)

Professor Gegenbaur does not appear to have examined the sacral nerves of this
species; but Professor Hoffman? describes the sciatic as arising from three roots, namely,
1, a bifurcation of the second presacral nerve (i. e. the ilio-lumbar nerve); 2, the whole
of the first presacral nerve; and 3, a branch from the intersacral nerve. No branch is
described or figured as going to it from the first postsacral nerve. The lumbar plexus
is described as formed by four roots:—1, the fourth presacral nerve; 2, the preaxial
bifurcation of the third presacral; 3, the postaxial bifurcation of the same third pre-
sacral; and 4, by the preaxial bifurcation of the second presacral. The first and second
roots unite together to form the obturator nerve, and the third and fourth roots unite
together to form the crural nerve.

We have examined two individuals of Iguana tuberculata, for one specimen of

1 L.¢. p. 201.
2 « Vor allem ist die Betheiligung postsacraler Nerven am Plexus ischiadicus ginzlich aufegehohen.”.
3 L.c. p. 178, fig. 13. 4 Luc. fig. 12.

Ҥ L.c. pp. 176, 177, fig. 11. His figure does not agree with his description as regards the obturator nerve.
SECOND SERIES.—ZOOLOGY, VOL. I. 3Z

518 MESSRS. MIVART AND CLARKE ON THE SACRAL

which we were indebted to the great kindness of Professor Flower, and to Professor
Parker for the other. We found both sides to agree as to the arrangement of the roots
of the lumbar and sacral plexus, and at the same time to differ from the description
above taken from Hoffman, as also from that given by Gegenbaur as existing in Lizards
generally. We found the sacral plexus and sciatic nerves to be formed by four roots,
the most postaxial of which comes from the first postsacral nerve. We found the lumbar
plexus to be formed by three roots, the most preaxial of which is the fourth presacral
nerve. Fibres from all the roots of the lumbar plexus appear to go to form the obtu-
rator nerve; but the crural nerve seems to be formed from the second and third presacral
nerves only.

The main difference between our specimens and the figure and descriptions of the
authors referred to, however, consists in the concurrence of the first postsacral nerve in
the formation of the sacral plexus.

In order the better to determine the essential nature of the nerve-roots, we thought it
desirable to follow out their branches to the muscles which are supplied by them; being
favoured by the large size and fine condition of the specimens examined.

The fifth presacral nerve (1) we found to have no connexion with the lumbar plexus,
but to send its branches to the pyramidalis! and other abdominal muscles.

The fourth presacral nerve (2) bifurcated, the more preaxial branch (3) going to the
internal oblique? and perhaps other abdominal muscles. The postaxial bifurcation (4)
went to form the obturator nerve, but sometimes, before doing so, gave off a branch (7) to
the iliacus?. The muscle is, however, also supplied by branches given off (7) from the
obturator nerve itself, as on one side of the specimen figured.

The third presacral nerve (9) joins a branch (11) from the second presacral nerve, and
it may communicate with the last-mentioned nerve by an additional short neryous
connexion (14).

The second presacral nerve (11+12) bifurcates, the preaxial bifurcation (11) joining
the third presacral nerve, with or without the aid of the short connecting branch (14).
Hither at this junction (13), or from the second presacral nerve before the junction, the
crural nerve (15) is given off. The postaxial bifurcation joins the sacral plexus. The
second and third presacral nerves having thus jgined together, they are continued on
by a single branch (10) which joins the postaxial bifurcation (4) of the fourth sacral
nerve, before described, to form the obturator nerve (5), which, after giving off a branch
(6) to the iliacus, perforates the pubic foramen and. then divides into branches, which
respectively go to the pectineus, obturator externus, tibial adductor, adductor magnus, and
gracilis muscles. or

The crural nerve (starting, as has been said, either directly from the preaxial bifur-
cation of the second presacral nerve, or else from the junction of this bifurcation with the
third presacral) passes over the brim of the pelvis, and, giving off two or three branches

‘Tn naming the muscles, we have followed the nomenclature adopted in the paper “ On the Myology of Iguana
tuberculata,” published in the ‘ Proceedings of the Zoological Society’ for 1867, p. 766. The pyramidalis is there
described at p. 772.

? This muscle is described at p. 771 of the paper referred to. * For zliacus see U. ¢. p. 787.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 519

(16) to the iliacus, then continues on to be distributed to the crwreus (20) and rectus
femoris (19), giving off in its course branches to the tsascalted) gluteus medius (17) and
maximus: (18).

The first presacral nerve (21), which is the largest of all, goes entirely to join the
postaxially extending bifurcation (12) of the second presacral nerve already mentioned ;
and these then, having effected a junction, are continued on by a single trunk (22) to
join the intersacral nerve (29).

The intersacral nerve (29) gives off, in the male, very close to its root, a very small
branch (81) to the penis, and then receives accessions, in the shape of two slender
branches (32 and 32), from the first postsacral nerve (30). The intersacral nerve, before
joining the conjoined trunk formed by the two more preaxial nerves, may give off a
branch (86 a) to the femoro-caudal muscle’, or a branch to this muscle may be given
off (86) from the great sciatic nerve itself.

The three roots of the sacral plexus (the intersacral and two presacrals) having thus
joined into one whole mass, the said mass divides into three unequal branches. The
smallest of these branches bifurcates and goes to the obtwrator internus®? muscle (23).
A second branch, which is rather slender, curves round the tendon of the femoro-caudal,
and gives off branches to the gracilis (24), the semimembranosus (25), the semitendinosus
(27), and the pyriformis (28) and biceps femoris muscles.

The main branch from the sacral plexus, the great sciatic nerve (33), gives off three
small branches to the gluteus medius (34) and maximus, to the ilio-peroneal (35), and to
the femoro-caudal (36) muscles. It then continues on down the thigh, just where the
deep edges of the gluteus maximus and ilio-peroneal are adjacent to each other, and,
distally, close to the delicate tendon of the femoro-caudal muscle*. When about two
thirds down the thigh, the sciatic nerve divides into the internal (37) and external (38)
popliteal; and later the latter again divides into two branches (89 and 40).

The first postsacral nerve (30), much smaller than the three preceding nerves, very
soon gives off a very long branch (44), which descends into the femoro-caudal muscle
and passes down the tail. It then bifurcates, but before or after bifurcating supplies
the slender branches (32 and 32) before mentioned which go to the intersacral nerve.
Its preaxial bifurcation ends by small branches (45), which supply the skin, transversus
perinei, and external, lateral, and caudal muscles. Its postaxial bifurcation (41) passes
under the tendon® from the caudal muscles which is attached to the tuberosity of the
ischium. It becomes connected by a very short branch (41 a) with the second postsacral
nerve, and then ends in delicate branches, which go to the external skin, transversus
perinet’ and sphincter cloace muscles.

The second postsacral nerve (42), more delicate than the first, gives off a branch (43)

1 This muscle is described at p. 791 of the paper referred to. Therein doubt was expressed as to correct naming

of the muscle, the name of which was chosen from the resemblance of the muscle in question to the gluteus maximus
of the Echidna. Its neryous supply coming from the crural is a strong argument against its being really the gluteus

maximus. 2 For this muscle see p- 794 of the paper before referred to.
* See Lc. p. 793. * For this tendon see J. c. p. 794, figs. 17 & 18 y.
° See lc. p. 774. ® See 7. ¢. p. 773.

382Z 2

520 MESSRS. MIVART AND CLARKE ON THE SACRAL

postaxial to the cloaca, and, after becoming connected, by the short branch (41 a) already
mentioned, with the first postsacral, it ends in delicate fibres whieh supply the sphincter
muscle and mucous membrane of the cloaca.

Monitor arenarius. (Plate LXVILI. fig. 2.)

The genus Monitor does not appear to have been examined by Professor Gegenhaur ;
but Professor Hoffman has described and figured! the arrangement of its lumbar and
sacral nerves. He represents its sacral plexus as formed from four roots, the most
postaxial of which is the first postsacral nerve, while the most preaxial is the lumbo-
sacral nerve coming off postaxiad from the second presacral nerve. The root consisting
of the first presacral nerve is the thickest, and does not bifureate, the whole of it going
to the sacral plexus. The second presacral nerve is shown dividing at once into three
branches, one of which is the lumbo-sacral nerve, another joins a branch of the third
presacral to form the crural nerve, while the remaining branch continues on as the
obturator nerve.

We have been fortunate enough to obtain two fine and large specimens of Monitor
arenarius, for one of which we are again indebted to the great kindness of Professor
Flower. In both these specimens we found on both sides of the body the same arrange-
ment of the nerve-trunks, which differed considerably from the conditions described by
Professor Hoffman. We found the sacral plexus to be formed of but three roots, the most
preaxial root consisting of the postaxial bifurcation of the first presacral nerve, while the
intersacral nerve was slightly the largest, and the first postsacral much the slenderest.
The first presacral nerve bifurcated, one branch going to the sacral plexus as already
mentioned, the other continuing on, as the crurai nerve, over the brim of the pelvis, but
being connected on each side with the second presacral nerve by a pair of delicate
nervous cords. The second presacral nerve having received (or given off) these nervous
cords, received an accession from the postaxial branch of the third presacral nerve, and
is then continued on as the obturator nerve.

We have thought it well to investigate the distribution of the nerves in this species,
being a species sufficiently remote from Jguana. We have, however, found a very close
similarity in this respect between these two animals.

The fourth presacral nerve (1) is like the fifth of Zguwana; it goes to the pyramidalis
and other abdominal muscles, and takes no part in the formation of the lumbar plexus.
The third presacral nerve (2) is like the similarly numbered fourth presacral of Jgwana,
except that we did not note that its postaxial branch (4) gave off any nerve to the
iliacus, although on one side of our specimen it gave off a branch which joined another
(10 a) from the second presacral, and thus continued on to the iliacus. The second
presacral nerve (9) is like the third of Jguana, except that it is less intimately fused
with the nerve postaxiad to it, with which it is only connected by commissural fibres
(14, 14, and 14.0). Having joined the postaxial branch (4) of the nerve before it, it is
continued on as the obturator nerve (5), giving off branches to the iliacus (6) before it

1 ZL. c. p. 175, fig. 7.

PLEXUS AND SACRAL VERTEBRZ OF LIZARDS. 521

passes through the pubic foramen. Having passed through the foramen, it divides (8),
and is distributed to the same muscles as in Zywana. On one side we found that minute
branches (16) went to the last-named muscle from the outermost of the commissural
fibres above referred to (14a). The first presacral nerve is like the second presacral of
Iguana, bifureating similarly, and sending one branch to the lumbar and one to the
sacral plexus. The former branch (11) is continued on (18) to become the crural nerve
(15), receiving on its way the commissural fibres before mentioned (14, 14 a, and 14d)
from the second presacral nerve. Having passed over the brim of the pelvis it divides
and supplies the very same muscles as in Jywana. The postaxial bifurcation (12) joins
the sacral plexus; but before doing so gives off a branch (23), going to the obturator
internus, the gracilis (24), and the semimembranosus (25) muscles. As in Zguana, the
branch to the obturator imternus (23) is isolated, while the fibres going to the gracilis
and semimembranosus (24 and 25) come off from the branch which also supplies the
semitendinosus (27) and the pyriformis (28). This difference is explicable by the fact
that the connexion between the sciatic and first postsacral nerves is less in Wonitor than
in Iguana. Therefore if the obturator internus, part of the semimembranosus, and the
gracilis are supplied by fibres coming from the first presacral, while the semitendinosus
and pyriformis ave supplied with fibres from the first postsacral, the whole set might be
supplied by the same main branch in J/guana (on account of the greater fusion above
mentioned), while they could not be so in Monitor. The intersacral nerve (29) is like
the intersacral nerve of Jgwana, save that it is connected with the first postsacral (30) by
only one connecting branch (82) instead of two. It thus appears that in Monitor there
is ne nerve answering to the first postsacral of /gwana, which seems entirely suppressed.
The sciatic nerve (33) was as in Jgwana, save that it gave no special branch to the femoro-
caudal muscle and no branch to the gracilis, semimembranosus, semitendinosus, and
pyriformis muscles. The first postsacral nerve (80) is like that of Jgwana, save that it is
(as has been said) less united with the intersacral nerve and the sciatic plexus, and
that it gives off the branch going to the semitendinosus (27) and pyriformis (28) muscles
as well as to part of the semimembranosus (26), which latter comes off from the sciatic
nerve in Jguana. This branch, like the branch supplying most of the same muscle in
Iguana, carves round the tendon of the femoro-caudal muscle. The same branch which
supplies the last-mentioned muscles also sends nerves (36) to the femoro-caudal itself.
As in Iguana, it gives off a branch (41) which passes under the tendon going from the
caudal muscles to the tuberosity of the ischium, and supplies the transversus perinei and
sphincter cloace muscle. It is also connected by a commissural branch (41 a) with the
second postsacral nerve; but in Monitor it is the commissural branch which is continued
(44) into the tail. The second postsacral nerve answers to the second postsaeral of Jgwana,
and gives off branches (42) to the mucous membrane of the cloaca and skin; but the
second branch to the cloaca which exists (43) in Jguana was not found in Monitor.

Thus the differences between these two very divergent Lacertilian forms may be thus
stated :—In Jguana four nerves contribute to form the sacral plexus; in Monitor only
three. In Jgwana the second presacral nerve bifurcates and goes to the two plexuses:
in Wonitor it is the first presacral that does so. In Zyuana the second, third, and fourth

522 MESSRS. MIVART AND CLARKE ON THE SACRAL

presacral nerves unite to form the lumbar plexus; in Monitor it is the first, second, and
third which doso. In Jguwana there is a branch to the iliacus (7) which was not detected in
Monitor. In Iguana the semimembranosus is not supplied from the first postsacral nerve ;
in Monitor it is so supplied. In Jguana the intersacral and first postsacral nerves unite
by two commissures; in Monitor by only one. In Jguana the femoro-caudal is supplied
by the sciatic nerve; in Monitor by the first postsacral. In Jguana more of the first
postsacral nerve joins the sciatic plexus than in Monitor, and the semitendinosus and
pyriformis are, in Monitor, supplied by the first postsacral instead of, as in Jgwana, from
the sciatic. In Iguana the obturator internus has a branch to itself, which comes from
the root of the sciatic. In Monitor it is supplied, in common with the gracilis and
semimembranosus, by a branch which is given off by the most preaxial root (12) of the
sacral plexus.

GENERAL CONDITION OF THE NERVOUS AND OssEOUS STRUCTURES OF THE SACRAL
REGION IN LIZARDS.

It appears, then, that in Lizards generally (judging from the specimens we have
examined and from the observations of Professors Gegenbaur and Hoffman), the
lumbar plexus may be formed by from two to three roots, and that the most preaxial of
them is never in advance of the fourth presacral nerve, while the most

postaxial root is never more postaxial than the first presacral nerve. Fig. 2.
Only in Monitor and Chameleo have we found this plexus to have
a root as postaxiad as the first presacral nerve, and neither of the jam

authors referred to represents it with so postaxiad a root.

The sacral plexus may have from three to four roots, but we have
generally found four. The most preaxiad of these is the lumbo-sacral
nerve (12), formed from the third presacral nerve (as sometimes in
Polychrus), or from the second presacral (as in Grammatophora, Lacerta,
Teius, Agama, and Iguana), or from the first presacral (as in Monitor).
The most postaxiad root is from the second postsacral, as in our spe-
cimens of Chameleo.

marmoratus 1m two

Generally, according to our examinations, the roots are the first jnaiviguals (from
(or first and second) presacral, the intersacral, and the first postsacral. Hoffman).
According to the authors referred to, it is generally the first and second
presacral and the intersacral, and never any postsacral. It cannot be — ‘schiatic nerve; o,
doubted, however, but that the roots preaxiad to the first vertebra aves f ie
with enlarged transverse processes may vary in number from one to Bie Be hy,
three, and that there may be two, one, or none behind the posterior postsacral vertebra;
vertebra with enlarged transverse processes. In all the Lizards re- s,s, vertebre with
ferred to there are two such vertebree with enlarged transverse pro- _—“"/arged transverse
cesses abutting against the ilimm. Are both these vertebrae sacral or 9 OS™
neither ; or if only one, which ?

The relation of the nervous roots to these transverse processes varies from species to

Nerves of Polychrus

crural nerve; 2,

is)
S

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 523

species, and even in different individuals (apparently, from Gegenbaur’s and Hoffman’s
works) of the same species. Shall we, then, consider that the nerves are the fixed points ?
If so, it is evident that we must extend our survey, and consider the nervous conditions
of the sacral region in other orders and classes of animals.

Crocodiles present no marked differences from Lizards, according to Gegenbaur,
who describes the sciatic nerve as formed from three roots—one intersacral, one pre-
sacral, and one postsacral, the intersacral one being much the largest. Professor Hoff-
man + agrees in this.

In Tortoises, according to Gegenbaur?, there are four roots to the sacral plexus—1,
one postsacral, as large as the next preaxiad, ¢.e. 2, the intersacral, and 3 and 4, two
presacrals, one of which is the largest of all. According to Hoffman? also there are four
roots; but he represents the postsacral root as by far the most slender, and the inter-
sacral root as larger than either of the two presacral nerves.

Fig. 4.

Nerves of Alligator (from Hoffman). Nerves of Tortoise (from Hoffman).
¢, crural nerve ; 2, ischiatic; 0, obturator; s, s, €, crural nerve; 2, ischiatic nerve; 0, obturator
vertebrae with expanded transverse pro- nerve; s,s, vertebrae with expanded trans-
cesses; 1 pr—3 pr, presacral vertebra ; verse processes; 1, 2, 3 and 4, presacral
1 po, postsacral vertebra. vertebra.

In all the Reptilia enumerated, the transverse processes which abut against the ilium
are wholly or in part parapophysial, and are in serial relation to (serial homologues of) the
capitular processes (or the capitular parts of the whole transverse processes) of the more
preaxial vertebra. The junction of the sacral vertebra with the ilium is much post-
acetabular in Saurians, but in Crocodiles and Tortoises (some at the least) it is about
opposite the acetabula.

THe Sacral REGION oF BATRACHIANS.

Here we have almost always but a single vertebra with enlarged transverse pro-
cesses abutting against the ilium. .

In the tailed Amphibia we have in Cryptobranchus japonicus, according to Professor
Humphry 4, a sciatic nerve formed from three roots, two of which are preaxial of the

_ | L.c. p. 187, fig. 14, * L.c. p. 200. ° L.c. p. 167, fig. 5.
* Journal of Anat. and Physiol. 1872. The muscles and nerves of C. japonicus, p. 51, woodcut.

524 MESSRS. MIVART AND CLARKE ON THE SACRAL

enlarged transverse process, while the third is postaxial to it, the obturator and crural
nerves being formed by two roots, one preaxiad to, the other

coincident with, the most preaxial root of the sacral plexus. Fig. 5.
The third spinal nerve, preaxial to the ilium (fig. 5, a), bifur- ary
cates, one branch forming the obturator nerve (fig. 5, 00), the
other contributing to form the crural (a.cr). The second spinal ,..,°
nerve, preaxial to the ilium (6), also bifurcates, one branch
joining the postaxial bifurcation of the preceding nerve to form Neves of Cryptobranchus (from

the crural, the other joining the spinal nerves, pre- and post- ae

axial to the ilium (¢ & d), to form the sciatic (sc). Hoffman merely cites! Professor
Humphry, and he gives a wrong reference.

In Salamandra maculosa and Triton cristatus, according to Hoffman?, the sacral nerve
is formed by one root preaxiad and two postaxiad of the transverse processes joining the
ilium. Dr. J. G. de Man? agrees with this representation, save that he places all the
roots one vertebra more preaxiad. We found, in Zriton cristatus (Pl. LXVI. fig. 6),
the same conditions as those described by Dr. de Man; but there was an interesting
difference between the two sides of the individual examined, inasmuch as there was a
lumbo-sacral nerve (12) only on one side, so that the sciatic had two roots (21 and 30)
on one side and three upon the other.

According to Professor Gegenbaur*, Salamandra maculosa
has three roots to its sacral plexus—(1) a delicate branch from
the nerve coming out preaxially to the large transverse pro-
cess; (2) one large root formed by the whole of the nerve,
i.e. coming out postaxially to the large transverse process ;
and (3), a delicate branch from the nerve next postaxiad.
Thus Gegenbaur agrees with Hoffman and differs from
Dr. de Man.

We found that a specimen (woodcut, fig. 6) of this species,
dissected by us, agreed as to its sacral roots with the obser-
vations recorded by Dr. de Man and figured by him in his nerco pvith tose of eaeceie
Table vii. fig. 4; but the branch connecting the second post- _ the Plates and as described in
sacral (9) nerve with the next postaxial nerve (12) appeared _ the text.
to us to be rather formed of fibres going preaxiad to the
crural nerve (15) than of fibres going thence postaxiad to the sacral (33). If we were
right in this view, the sacral nerve had but two roots, one issuing preaxiad (12) and the
other postaxiad (30) to the ilium-joining vertebra. Moreover, in our specimen, a branch
from the third preaxial nerve (2) joined, but very slightly so, the lumbar plexus on one
side, while on the other side it approximated to, but did not join, that plexus, and then
went on its way.

In the tailless Batrachia, the sciatic nerve appears, according to Hoffman, to be

a.cr

Fig. 6.

Lumbo-sacro-caudal vertebree and
nerves of Salamandra maculosa.
The lettering and numbers

1 ZL. ¢. p. 149, fig. 1. ? L.¢. p. 149, fig. 2.
* « Myologie comparative de extrémité postérieure chez les Amphibies,” Niederlindisches Archiv, ii. p. 58.
‘ L.c. p. 200. 5 L. ¢. pp. 156, 157, figs. 3, 4.

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 525

formed of three roots, one preaxiad and another postaxiad to the ilium-supporting ver-
tebra, while the third root is still more postaxiad. In Batrachians the transverse pro-

Fig. 7. Fig. 8.

Nerves of Dactylethra, fig. 7, and of Rana and Hyla, fig. 8 (from Hoffman).
¢, crural nerve; 2, ischiatic nerve; 0, obturator nerve; 1 pr—3 pr, presacral vertebrae; s. vertebra with enlarged
transverse processes.

cesses abutting against the ilium are parapophysial, not diapophysial, in nature, thus
agreeing with those of Reptiles.

THE SACRAL REGION OF MAMMALS.

In Man the great sciatic nerve has six roots, the most preaxial of which (the fourth
lumbar nerve of anthropotomy) may be called the second presacral of the nomenclature
here adopted, since the next postaxiad (the fifth lumbar nerve) comes out preaxial
to the first transverse process abutting against the ilium, and would thus be our first
presacral. The next postaxiad (the first sacral nerve of anthropotomy) may be reckoned
as equivalent to what we have termed intersacral in Lizards, while the remaining three
may be reckoned as equivalent to our first, second, and third postsacral nerves.

Now if we take the nervous relations as our exclusive test, then the vertebra postaxial
_ to the first sacral root of Lizards must be the homologue of the first sacral vertebra of
Man; and in that case this vertebra would, in Monitor and Chameleo, be the first with
expanded processes abutting against the lium. In Iguana, Teius, Lacerta, and Gramma-
tophora, however, it would be the vertebra preaxiad of such first ilium-joining vertebra,
while in Agama and, at least sometimes, in Polychrus', it would be the second vertebra
in front of such ilium-joining vertebra.

In an inferior mammal, such as the Cat (Pl. LXVI. fig. 7), we found the sacral plexus
to be formed by four roots, two very much larger than the others. The most preaxial (12)
root (the largest but one) is formed by part of the nerve coming out between the pen-
ultimate and ultimate lumbar vertebre; the next (the largest) comes out (21) in front

' See ante, woodcut, fig. 2, p. 522.
SECOND SERIES.—ZOOLOGY, VOL. I. AA

526 MESSRS. MIVART AND CLARKE ON THE SACRAL

of the first sacral vertebra; the third (the smallest but one) comes out (29) from the
most preaxiad sacral foramen; the last (the smallest) comes forth from (29 a) the second
sacral foramen. Thus here the sciatic nerve is formed from nerves issuing out of the
sacrum in a less proportion than in Man, and more by nerves issuing preaxiad to the
sacrum—not that the nwmber of the latter is greater than in Man, but that their relative ~
size, compared with those from out the sacrum, is greater. Compared with Lizards, it would
seem, with respect to nerves, that the first and second sacral vertebree of the Cat answer
very well to the two vertebree with enlarged transverse processes of Lizards, while osteo-
logically they, of course, also answer very well to them. There can be little doubt, how-
ever, but that the two first sacral vertebree of the Cat are to be considered homologous
with the anterior human sacral vertebrae; and therefore it would seem that the two
iliam-joining vertebree of Lizards should be considered homologous with the anterior
human sacral vertebree.

In Man, the Cat, and also in other mammals, down to the Hchidna, the transverse
processes abutting against the ilium are parapophysial, like those of the other classes
before described—viz. Reptiles and Batrachians. In all mammals which we have ex-
amined, however, the junction of the sacral transverse processes with the ilia is pre-
acetabular, although that junction is much less preacetabular in position in Man than it
is in most mammals.

Altogether, from the osseous and nervous conditions considered jointly in the groups
hitherto referred to, we would venture to propose the following definition of a “sacral
vertebra” in Mammals, Reptiles, and Batrachians:—“A vertebra with parapophysial
transverse processes which abut against the ilium preaxiad, or postaxiad, or opposite to the
acetabulum, and having a root of the sciatic plexus coming forth either immediately pre-
axiad or postaxiad of it.” This definition will exclude from the sacrum of Man, as not
abutting against the ilium, the more posterior vertebree called “sacral” in anthropotomy.
But in the lower mammals (even already in Apes) the number of so-called “sacral”
vertebree augments more or less with age by the ankylosis of the sacral vertebrz, so as
to render the extent of the ‘‘sacrum” very variable. It would surely be well, then, to
distinguish the human sacral vertebre (like the ribs) into true and false, those being the
true sacral vertebree which abut against the ilium.

THE SACRAL REGION IN BrrRDs.

The determination of the homological relations of the different parts of the postdorsal
part of the spinal column of Birds is a matter of much difficulty. It might be antici-
pated, however, from the close relations and probable genetic affinity between Birds and
Reptiles, that the true sacral vertebra of the former class might be readily determined
by the aid of reptilian characters. But however close may be the morphological afti-
nities, they are so strangely disguised by great physiological differences, that the results
of comparison are much too unsatisfactory to justify the anticipation above referred to.
The enormous forward prolongation of the iliac bones and the great power and activity
of the erect and vigorous legs of birds, compared with the backwardly extending ilia and
the sprawling and feebly pushing legs of Saurians, combine to produce in Birds a redun-

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 527

dancy of both osseous and nervous structures compared with those of the existing Rep-
tilia. Could we dissect the different forms of Pterosawria and Dinosauria, these diffi-
culties and obscurities would no doubt disappear; but the hiatus is too great between
the most Sauroid of existing Birds and any form of Lizard for the homological rela-
tions, in this respect, to be readily and easily solved.

In Birds generally, we find that a greater or less number of the vertebre which are
postaxial to those bearing elongate ribs, develop short parapophyses which abut against,
and ankylose with, the preaxiaily extending, preacetabular part of the ilium. Postaxial
to these come vertebree (which have been called1+ lumbo-sacral) which develop no para-
pophyses at all, or only minute rudiments of such, a fossa (the acetabular or renal
fossa) resulting from their absence.

Behind these, again, there are generally vertebree with elongated parapophyses abutting
against the ilium; and the first one, two, or three of these have often a more or less
decidedly different aspect from those which postaxially succeed. These vertebrae, which
we will provisionally call, for distinction, “posterior parapoyhysial,”’ abut against the
postacetabular part of the ilium in the Struthionide and in most other birds. In the
Merganser they are placed very much postaxiad to the acetabulum. In Otis they
are opposite the middle of the acetabulum, and they are even more preaxiad in Tetrao
tetric and Gallus, while in the Peacock (Pavo) they abut against the preaxial part of
the acetabulum. Thus in birds these posterior parapophysial vertebre (whatever may be
their true nature) sometimes have a position relative to the acetabulum nearly approach-
ing that which is found in the sacral vertebre of some mammals, though generally their
relation to it is that of the sacral vertebree of Saurians.

Ina few birds none of the vertebre postaxiad to the acetabulum develop parapophyses.
This is the case, é. g., in the Hornbill (Buceros), in which the most postaxiad vertebra, with
parapophyses abutting against the ilium, abuts against the front margin of the aceta-
bulum. In the Woodpecker (Pica) the position of the most postaxiad of these parapo-
physes is still more removed preaxially from the acetabulum.

In some Parrots the most postaxial of these vertebree with parapophyses is even more
preaxiad to the acetabulum than in Pica; but in other Parrot-forms there are also one or
two vertebree with parapophyses opposite the middle of the acetabulum, and separated
by an interval from the preacetabular vertebrae with parapophyses.

Judging, then, from the skeleton alone, we should be inclined to consider these posterior
parapophysial vertebre abutting, in most Birds, against the postacetabular part of the
ilium as the homologues of the lacertian sacral vertebre which also abut against the
postacetabular part of the ilium. And just also as we should consider the sacral vertebree
of Alligators and Tortoises, in spite of their more preaxial position with relation to the
acetabulum, as really homologous with the sacral vertebre of Lizards, so we should con-
sider the more preaxially situated posterior parapophysial vertebre of Otis, Tetrao,
Gallus, and Pavo as homologous with the postacetabular posterior parapophysial vertebrze
of the Struthionide and of most birds. What, then, is to be said as to the vertebrz of

1 Trans. Zool. Soc. vol. x. p. 9.

4a 2

528 MESSRS. MIVART AND CLARKE ON THE SACRAL

this region in such forms as Buceros and Pica and the Parrots? Osteologically they
are indeterminable, but they do not stand alone; the similar vertebrae of the Diver
(Colymbus) are also osteologically indeterminable from their ankylosis and lateral com-
pression, and the corresponding vertebrae of the Penguins can hardly be discriminated.

Quite osteologically indeterminable, again, is such a sacrum as that of the Frigate Bird
(Fregata), in which none of the vertebree of this region are devoid of parapophyses, and.
in which there is therefore no differentiation into anterior and posterior parapophysial
vertebrae within the pelvic part of the vertebral column.

We must turn, then, for aid to the nervous conditions of the parts.

Now in birds (fig. 9) the sciatic plexus and nerve has (as repre-
sented by Professor Gegenbaur) as many as six roots!; and if
we are to take nervous conditions as our test, and if we are to
consider the most preaxial parts of the sciatic nervous mass
in Birds as corresponding with the most preaxial parts of the
sciatic mass in Lizards, then we must consider that some or ae,
other of the avian vertebree without transverse processes (an-
terior to Gegenbaur’s vertebrae a and 0) must answer to those
vertebrae of Lizards which have enlarged transverse processes
to abut against the ilium.

But we have seen that the amount of variation in Lizards
is such that, if nerves be taken as our test, we must deny that
the ilium-joining vertebree of one species answers to the ilium-
joining vertebrae of other species—nay, even that, in indi-
viduals of the same species, we ought (according to this view)
to call the ilium-joining vertebree “sacral” in one and
“lumbar” in another. So to do, however, would surely be Nerves of Birds (from
ings alse b i Ho Be longated

If Professor Gegenbaur is, as he doubtless is, right as to Datimece ot aa
the sciatic nerve-roots in Birds, then, as there are six such against the ilium, opposite
roots, and the most preaxial of them comes forth in front of the acetabulum; er, crural
the most postaxial of those vertebrae with parapophyses which — °tve 00¢, obturator nerve;
are preaxiad to the vertebrae without parapophyses, it follows |” wo
that such vertebra would be included within that definition peeve

: 1'-5', vertebree postaxiad to
of “sacral”? which we have above proposed as applicable to a5.
Mammals, Reptiles, and Batrachians. But, at the same time, :
at least the first of the posterior parapophysial vertebre (7. e. the first vertebra postaxiad
to those without parapophyses) would also enter into that definition; and thus in most
birds we should have sacral vertebre separated from each other by interposed vertebrze
not sacral, which would be a very forced and unnatural conception.

But it might, perhaps, be suggested that as the ilia are so antero-posteriorly extensive
in Birds, the preacetabular and most preaxial of the vertebree giving out sacral roots in

transverse processes abutting

© iby Oj AVIn

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 529

Birds answer to the sacral vertebree of mammals, which are preacetabular ; while the post-
acetabular and most postaxial of the vertebree giving out sacral roots in Birds answer to
the sacral vertebra of Reptiles, which are postacetabular in position. . But, as has been
said, in some Birds the posterior parapophysial vertebre abut beside the acetabulum ;
and there is so little difference in this region between Man and the Peacock that it
would seem to be very unreasonable not to consider the more preaxial of the posterior
parapophysial vertebree of that bird as being the homologues of the anterior sacral ver-
tebre of Man—and this the more so, seeing that the difference is bridged over by the
Tortoises and Crocodiles, which have their sacral vertebree beside the acetabulum. May
we not, then, take the nervous conditions as our guide, and say that every vertebra which
has a sacral root coming forth, either pre- or postaxiad of it, is “sacral’’? But then the
last two lumbar vertebree of at least many mammals would be sacral; while if we deter-
mine that only those vertebree shall be sacral which give forth from their postaxial end
a root going entirely to the sacral plexus, then the last lumbar of Man and the Cat would
be “sacral,” as also the first presacral vertebra of Teiws and Agama, while the second
sacral vertebra of no Lizard we have examined would be “ sacral’”’ at all !

If we were to add to our definition of a sacral vertebra, as proposed for Mammals, Rep-
tiles, and Batrachians, the words “ together with any or every vertebra without parapo-
physes and interposed between vertebree abutting against the ilium, and having a root
of the sciatic plexus coming forth either immediately preaxiad or postaxiad of it,’’ we
should have a definition which would include all the vertebree which might be considered
“sacral” in most Birds; but it would leave out in an arbitrary and unnatural manner the
vertebree in such forms as Buceros and Pica, which vertebree are plainly homologous with
those vertebree which in most Birds are without parapophyses, but are interposed between
vertebrae which have parapophyses. Neither, moreover, would it apply to Fregata.

According to Professor Gegenbaur, the most preaxiad root of the sacral plexus is, in
some species at least, the fifth spinal nerve in front of the posterior parapophysial ver-
tebree; and if he has correctly determined his vertebree! a and J in Pica and Buceros,
then the vertebra which he considers the fourth presacral will alone enter into even our
enlarged definition of a sacral vertebra as above given, and which will therefore be a
manifestly incomplete one for birds. The same would be the case with those Parrots
which have no parapophysial vertebree in this region, save greatly preaxial to the aceta-
bulum ; while in those which have such vertebre, five or six vertebree would be included
under it.

On the whole, then, seeing, on the one hand, the manifest homology between the
sacral vertebre of Man and Lizards (by the help of those of Crocodiles and Tortoises), and,
on the other hand, the manifest homology between the sacral vertebra of Lizards and
the posterior parapophysial vertebrae of most Birds, we think it better to consider the
latter vertebree in Birds as alone truly sacral, and to regard such forms as Buceros, Pica,
and certain Parrots as differing from the rule of the class by the suppression of their
parapophysial processes, and Fregata as differing from the same rule by the development
of parapophyses in all the vertebre of this region.

> L. c. plate vii. figs, xxix. and xxx.

530 MESSRS. MIVART AND CLARKE ON THE SACRAL

The sacral vertebree in Birds may be defined, then, as “vertebre having one of the
more postaxial roots of the sciatic plexus coming forth either immediately preaxiad or
postaxiad, having parapophysial transverse processes abutting against the tlium, and placed
mmediately postaxiad to vertebre which are devoid of such parapophyses, or else being
the homologues of vertebre so conditioned in most other Birds.” By the combination of
these two definitions, the former one for Mammals, Reptiles, and Batrachians, and this
latter one for Birds, it seems that the sacral vertebre may be defined in all Vertebrata,
above Fishes, which have pelvic limbs.

What, then, are we to consider those vertebree of Birds to be which are preaxial to the
sacrum—those, namely, which are generally more or less entirely devoid of transverse
processes? It appears to us that in Birds this region of the spine is really augmented in
extent and in the number of its vertebree in unison with the augmentation in the number
of the sciatic nervous roots. Such vertebre may, of course, be considered as corre-
sponding with the human lumbar vertebre; but inasmuch as they are so constantly
developed in Birds, and are so peculiar in that class, it may be as well to distinguish
them by a distinct appellation, and to call them, as one of us has elsewhere* before sug-
gested, ‘“lumbo-sacral vertebree.”

At first sight it might be well supposed that these vertebra answer to the sacral ver-
tebree of Man; but it is evident which of the vertebrze of Reptiles answer to the human
sacral vertebree ; and we agree with Professor Gegenbaur in thinking that the sacral ver-
tebree of Birds may be determined by the sacral vertebree of Reptiles. Upon this view the
normal type must be taken as existing in Reptiles and Batrachians, compared with which
we have in Man a multiplication of vertebre, postaxial to the true sacral vertebra, giving
their nerves to the sacral plexus, while in Birds we have a similar multiplication of the
vertebree which are immediately preaxial to the sacrum. The sciatic plexus in Man is
mainly furnished from the true sacral vertebree and the posterior vertebree which have
coalesced with them: it is furnished mainly from the presacral (lumbo-sacral) vertebree
in the Bird. That there should be this difference is, however, very little surprising when
we consider that even in the Cat it is the presacral and not the sacral vertebrae which give
forth the greatest part of the fibres of the sciatic plexus. Yet no one would dream of
calling the last lumbar vertebra of the Cat “sacral” on account of such increase in
volume of the presacral part of its sacral plexus. As to augmentation in the number
of roots, the variations we have found to exist in Lizards afford abundant ground for
declining to change the designation of a vertebra from “lumbar” to “sacral,” or vice
versd, on this account alone.

1 Trans. Zool. Soc. vol. x. p. 9.

Gt o

PLEXUS AND SACRAL VERTEBRA OF LIZARDS. 5381

DESCRIPTION OF THE PLATES.

Prats LXVI. fig. 1. Lumbo-sacral plexus of Chameleo vulgaris.
2. Ditto, Lacerta viridis.

3. Ditto, Teius teguexin.

4. Ditto, Grammatophora barbata.
5. Ditto, Agama colonorum.
6. Ditto, Triton cristatus.
7. Ditto, Cat (Felis catus).

1. Lumbo-sacral plexus and nerves of Iguana tuberculata.
2. Ditto of Monitor arenarius.

Priare LXVII. fig.

The following lettering applies consecutively to the different figures.

NERVES.

. Spinal nerve, third preaxial to the one giving off the lumbo-sacral nerve (. e. fifth presacral nerve

in Lguana, fourth in Monitor).

. Spinal nerve, second preaxial to the one giving off the lumbo-sacral nerve (fourth or third pre-

sacral).

. Branch to abdominal muscles.

. Branch continued on into the obturator nerve.

. Obturator nerve.

. Branch or branches to éhacus.

. Other branch or branches to ilacus.

. Obturator nerve dividing to supply the pectineus, obturator externus, tibial adductor, adductor

magnus, and gracilis muscles.

. Spinal nerve, next preaxial to that which gives off the lumbo-sacral nerve (third or second pre-

sacral nerve).

. [ts connexion with the obturator nerve.
. Preaxial bifurcation of the nerve, the postaxial bifurcation of which is the lumbo-sacral nerve

(second or first presacral nerve).

. Lumbo-sacral nerve (generally the postaxial bifurcation of either the second or the first presacral

nerve, but in Salamandra the entire first presacral) .

. Continuation of nerve 11 towards the obturator nerve.
. Connexion between nerves 11 and 9.

. Crural nerve.

. Branches to iiacus.

. Branch to gluteus medius.

. Branch to gluteus maximus.

. Branch to rectus femoris.

. Branch to crureus.

. Spinal nerve intermediate between the spinal nerve giving off the lumbo-sacral nerve and

22.

23

the more preaxial of the vertebree with enlarged transverse processes (either Ist presacral
nerve or absent).

Trunk formed by 12 and 21, and going to sciatic nerve.

Branch to obturator internus.

532

24.
25.
26.
27.
28.
29.

ON THE SACRAL PLEXUS AND SACRAL VERTEBR# OF LIZARDS.

Branch to gracilis.

Branch to semimembranosus.
Another branch to semimembranosus.
Branch to semitendinosus.

Branch to pyriformis.

Intersacral nerve. _

29 a. Second intersacral (in Cat).

30.
3l.
32.
33.
34.
35.
36.

First postsacral nerve.

Branch to penis.

Commissure or commissures between 29 and 30.
Sciatic nerve.

Branches to gluteus maximus and medius.
Branch to tlio-peroneal.

Branches to femoro-caudal.

36 a. Extra branch to femoro-caudal.

37.
38.

Internal popliteal nerve.
External popliteal nerve.

39 and 40. Divisions of external popliteal nerve.

Al.

Branches to ¢ransversus perinei and sphincter cloace.

41 a. Branch connecting first and second postsacral nerves.

42.
43.
44,
45.
46.

Second postsacral nerve, ending in branches to mucous membrane of cloaca and to skin.

Branch to cloaca.

Branch to femoro-caudal, and extending down to the tail.

Branches to skin and lateral caudal muscles.

Branch in Agama (i.e. in fig. 5, Plate LXVI.), extending postaxiad from the third presacral
nerve to the lumbo-sacral nerve.

VERTEBRA.

A. Most preaxial vertebra articulating with ilium.

B. Second (or more postaxial) vertebra articulating with ilum.

I. Vertebra preaxiad to A.

II, III, IV, and V. Vertebrz successively more and more preaxial.

Ik,

Vertebra postaxial to the more postaxial vertebra articulating with ilium, or to the only vertebra
which so articulates.

TI*, ILI*, and IV*. Vertebrz successively more and more postaxial.

Trans Linn.Soc.5zr.2.Zoon Vou. Pu 66

Agama colonerunv Triton cristatus.

Chameleo vulgaris

Granmatophora barhata

TZ. re Pe = ae
; Tews tegwexi.
Lacerta viridis. 2

CBerjean bth. SACRAL PLEXUS & VERTEBR& OF LIZARDS &c. Hanhart imp

-
‘
ws
eat ,
Ny
ee
fags tt
‘
ase
.

aa

j
“
We
" rah |
‘

fg. l

Trans. LINN.SOC. 51

ER 2.Z00L You.l.Pu.67

CBerjeau hth

3S?

Monitor

PL!

al

RT

24

BRA OF LIZARDS.

EXUS & VI

Hanhart imp

oe

THE

TRANSACTIONS

THE LINNEAN SOCIETY

LONDON.

SECOND SERIES.—ZOOLOGY.
VOLUME I.
PART THE EIGHTH.

EON DON:
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET:
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON-HOUSE;

AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

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XXI.

XXII.

XXIII.

XXIV.

CONTENTS.

PART VIII.—June 1879.

The Butterflies of Malacca. By ARTHUR GARDINER Borurr, F.L.S., F.Z.S.,
Senior Assistant in the Zoological Department, British Museum. (Plates
MODAN EINES. TXT re PE RET CRESS ar CUB SRS OS ORNS spageld533

On certain Organs of the Cidaride. By Cuaries Stewart, Esq., F.L.S.,
M.R.C.S., Lecturer on Comp. Anat. St. Thomas’s Hospital. (Plate LXX.) 569

On Hypsiprymnodon, Ramsay, a Genus indicative of a distinct Family (Pleopo-
dide) in the Diprotodont Section of the Marsupialia. By Prof. OwEn, C.B.,
F.RS., L.L.S., §c.. Superintendent of the Natural-History Departments in the
British Monscums (2 lates) IAXeXele SIXOXCLL, Pa 8.) ER oS

Descriptions of some minute Hymenopterous Insects. By J.O. Wxstwoop, W/.4.,
ee Sirece: (Plater XOMUUP er ee i on Sy ee ee eur oeS

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[ 583 ]

XXI. The Butierflies of Malacca*. By Anvuur Garpiver Burimr, F.L.S., ZS,
Senior Assistant in the Zoological Department, British Museum.

(Plates LXVIII., LXIX.)

Read December 21st, 1876.

THE generosity of Captain Stackhouse Pinwill, who has presented to the British
Museum the whole of the Lepidoptera collected by himself during some years’ residence
in Malacca and Penang, has enabled me to study (with, I trust, some degree of profit to
others as well as to myself) the Lepidopterous fauna of the Malayan peninsula. As I
was obliged to go over the whole of the species hitherto recorded from Malaysia, and as
Captain Pinwill’s collection is the most complete one hitherto received from the penin-
sula, I determined not to lose the present opportunity of making a complete list of the
Butterflies as yet known to occur at Malacca, comparing them at the same time, by means
of a table, with the species hitherto received from India, China, Siam, and the islands
to the south of the peninsula.

Of the 258 species now registered from Malacca, 36 appear to be endemic; of the
remainder rather more than a fourth occur either at Assam or Nepal, more than a
seventh at Moulmein, less than a seventh at Ceylon, nearly two fifths (apparently) in_
the island of Penang, about two elevenths at Singapore, about three sevenths in Borneo,
about three sixteenths in Sumatra, more than a third in Java, about two thirteenths in
Siam, rather more than a tenth in China, two species in the New Hebrides, and six in
Australia. We see therefore that, with the exception of the last-mentioned eight
species, the Butterflies of Malacca are limited to the Indian Region ; there are, however,
several forms occurring in the Australian Region which some Lepidopterists would not
regard as specifically distinct, such as the various allied forms of Danuais, Melanitis,
ycalesis, Doleschallia, Neptis, Diadema, Cynthia, Junonia, Lampides, Amblypodia,
Catopsilia, and Papilio. All of these are, nevertheless, characterized by slight but con-
stant differences, and consequently have a right to be regarded as distinct species.

It appears to me to be important to those who desire conscientiously to study the
geographical distribution of animals, to discriminate between even the most nearly allied
species. I believe that few things have more retarded zoological geography than the
reckless association together of so-called “local varieties” under the same specific name.
I will cite an instance in illustration of what I mean. Some Lepidopterists still assert
that Diadema bolina ranges from Northern India to New Zealand, whereas that species
does not occur outside the Indian Region. Allied forms, indeed, are common throughout
the Moluccas, Australia, and the South Pacific; and one of them (D. nerina), strange

* Two of the new species mentioned in the present paper, and an abstract of the whole, have already appeared in
the Society’s Journal (Zool. vol. xiii. p. 115 and p. 196 respectively).
SECOND SERIES.—ZOOLOGY, VOL. I. AB

534 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

enough, is found both in Java and the Australian Region, whilst a dwarfed form of it is
taken in the Philippines; yet D. nerina, although more widely distributed than D. bolina,
exhibits even less individual variation.

The fact of two species of Terias, discovered in the New Hebrides, turning up in
Malacca, is somewhat startling; but it is quite possible, in a genus where only a prac-
tised eye can at once detect the specific differences, that the same form of imago may
have been simultaneously developed in both localities ; moreover, breeding might (as it
often does) reveal constant and well-marked distinctive characters whereby the earlier
stages could at once be separated.

The following Note, which appears to me to be worthy of the consideration of natu-
ralists, has been communicated by my friend Mr. W. L. Distant, who was for some years
resident at Province Wellesley :—

** All catalogues of the Penang fauna must be accepted with some qualifications. The
dependencies of Penang and Province Wellesley are usually made more or less into one
zoological province, especially as regards the insects. This I found to be the case when
collecting there a few years ago, all collections being then made indiscriminately from
both localities.

*“The island of Penang is about 20 miles long by 9 miles broad, comprising an area
of about 107 square miles. <A group of granite hills occupies about two thirds of its
extent, running through its centre from north to south, bounded by a plain on their
eastern and western sides. These hills are densely covered with a beautiful forest on all
sides ; and their highest point, West Hill, is about 2600 feet above the sea.

“Province Wellesley is situated on the Malayan peninsula, and is separated from
Penang by a strait from 2 to 10 miles broad; it is about 35 miles in length, averaging
8 miles in width. Although it possesses many hills of a considerable elevation (yet not
equalling those of the island of Penang), its general character is more of a plain, in which
jungle tracts and cultivated lands of rice and sugar-cane are its characteristics. Man-
grove swamps, often of great extent, exist in the neighbourhoods of its many creeks and
rivers.

“There is little doubt that, when the floras and faunas of these two districts are
worked out separately and distinctly, a common character will be found to pervade them
both, but that many species will be found locally distinct and constant. Species may be
expected to exist in the lofty wooded regions of Penang that are absent in the plains and
less elevated hills of Province Wellesley. At present, however, when mixed collections
are made from both regions, the habitat ‘ Penang’ must be accepted as not representing
the insular fauna alone, but comprising also that of a portion of the Malayan
peninsula.”

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 539d

List of Species collected by Captain Pimwill.
RHOPALOCERA.
Family NYMPHALIDA.
Subfamily Danan, Bates.
Genus Evpia@a, Fabricius.

1. Evpr@a PHa@sus, Butler, Proc. Zool. Soc. p. 270. n. 3 (1866). od ?, Malacca.
This is clearly the species described by Felder as HZ. Castelnaut.

2. EupL@A OcHSENHEIMERI, Lucas, Rev. et Mag. Zool. p. 315 (1853). ¢ 2, Malacca.

3. Eupt@a Groret, Felder, Reise der Nov., Lep. i. p. 339. n. 470, pl. 41. fig. 7
(1865). & 2, Malacca.

4, EuPL@A MARGARITA, Butler, Proc. Zool. Soc. p. 279. n. 84 (1866). ¢ ?, Malacca;
2, Penang.

5. EUPL@A CHLO#, Guérin, in Deless. Souv- Voy. d. l’Inde, ii. p. 71 (1843). ¢ ?, Malacca.

6. HurpL@a Bremert, Felder, Wien. ent. Monatschr. iv. p. 398. n. 16 (1860). 3, Ma-
lacea. Mr. W. L. Distant took this species at Province Wellesley; he says that all
the species of Huplea which he captured were flying in the months of August and
September.

7. Eupt@a Ménérriést, Felder, Wien. ent. Monatschr. iv. p. 398. n. 15 (1860).
Malacca.

8. Eupt@a Prnwitti, n. sp. (PI. LXIX. fig. 9.) Primaries brownish piceous,
purple-shot, slightly paler along the external border: a long sericeous interno-median
streak : secondaries paler brown, deepest at base; two marginal series of whitish spots,
clear at anal angle, obsolescent and decreasing towards apex ; costal area greyish. Pri-
maries below paler than above, with a subcostal spot and an elliptical interno-median
spot pinky white; a spot in the cell and two beyond it bluish white, two or three white
dots at external angle; secondaries with the submarginal spots clear whitish; a spot in
the cell and seven in an arched series beyond it lilacine whitish; base and pectus
white-spotted. Expanse of wings 4 inches. ¢ 2, Malacca. ZH. Pinwilli is allied to
LE. Meénétriési, but larger, darker, and shot with purple.

9. Evpn@a MipAMus, (Papilio midamus) Linneeus, Mus. Lud. Ulr. p. 251 (1764).
$ 2, Malacca; ?, Penang. ‘ This species is very common in gardens ” (WV. L. Distant).

Genus CALLipL@a, Butler.
1. CALLIPL@A DIOCLETIANUS, (Papilio diocletianus) Fabricius, Ent. Syst. iii. 1, p. 40.
n. 118 (1793). ¢ ¢%, Malacca. ‘Common in gardens” (W. L. Distant).

2. CALLIPL@A VESTIGIATA, (Luplea vestigiata) Butler, Proc. Zool. Soc. p. 288. n. 58,

fig. 1 (1866). 3 ¢, Malacca.
4B 2

536 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

3. CALLIPL@A LEDERERI, (Huplea Ledereri) Felder, Wien. ent. Monatschr. iv. p. 397.
n. 14 (1860); Reise der Nov., Lep. ii. p. 317. n. 481, pl. 40. figs. 5, 6 (1865).
3 ?, Malacca.

Genus SALPINX.

1. SALPINX LEUCOGONIS, n. sp. Nearly allied to E. vestigiata, but smaller, the costa
of primaries not so strongly arched, the outer margin slightly inarched in the centre,
the submarginal spot on second median spot wanting, the subcostal spot above the
end of the cell much smaller, the spot on interno-median area widened into a notched
blotch, all the spots lilac; secondaries paler, only three lilacine whitish spots placed
obliquely near apex, no submarginal spots, anal angle white. Primaries below with
only two or three submarginal .and three or four marginal white dots; secondaries with
no white blotches beyond the cell. Expanse of wings 3 inches 2 lines, ¢, Malacca.

Genus Danats, Latreille.

1. Danats PLEXIPPUS, (Papilio plexippus) Linneeus, Mus. Lud. Ulr. p. 262 (1764).
3 ¢, Malacca; 3, Penang. “Common in August and September” (W. L. Distant).

2. DANAIS MELANIPPUS, (Papilio melanippus) Cramer, Pap. Exot. ii. pl. 127. figs. a, B
(1779). 3 2, Malacca; 3, Penang.

8. DANAIS VULGARIS, Butler, Ent. Mo. Mag. p. 164 (1874). 3 %, Malacca.

4. DANAIS SEPTENTRIONIS, Butler, Ent. Mo. Mag. p. 163 (1874). 3, Malacca ;
2, Penang.

5. DANAIS GRAMMICA, Boisduval, Sp. Gén. Lép. i. pl. 11. fig. 10 (1886). 3 ?, Malacca.

6. DANAIS MELANEUS, (Papilio melaneus) Cramer, Pap. Exot. i. pl. 30. fig. p (1775).
?, Malacca.

7. Danats crocHa, Butler, Proc. Zool. Soc. p. 57. n. 53, pl. 4. figs. 5, 6 (1866).
3 %, Malacca. “ Occurs at Penang in August and September” (W. L. Distant).

Genus IpzEopsis, Horsfield.

1. Ipropsts Daos, (Idea daos) Boisduval, Sp. Gén. Lép. i. pl. 24. fig. 3 (1836).
3 2, Malacca; ?, Penang.

Genus Hest1a, Hibner.

1. Hestta LyNncEvs, (Papilio lynceus) Drury, Ill. Ex. Ent. ii. pl. 7. fig. 1 (1778).
3, Malacca. ‘ The species of Hestia go by the local name of ‘the widow’ in Province
Wellesley; the genus was very abundant near the top of Penang Hill” (W. L. Distant).

2. HESTIA LINTEATA, n. sp. (Pl. LXIX. fig. 6.) Nearly allied to H. belia, but much
larger, the veins less broadly black-bordered; spots larger, excepting towards the costa
of primaries at apex; discoidal spot of secondaries very large; clavate markings ter-
minating the veins much longer, more slender in the middle. Hxpanse of wings 5 inches
6 lines to 6 inches 8 lines. o ?, Malacca (six examples).

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 537
Subfamily Satyrina#, Bates.

Genus Mexaniris, Fabricius.

1. MELANITIS LEDA, (Papilio leda) Linneeus, Syst. Nat. i. 2, p. 773. n. 150 (1766).
$ ?, Malacca, Penang. “Common in jungle-grass” (W. L. Distant).

Genus Letae, Hiibner. .

1. Levis evRopA, (Papilio europa) Fabricius, Syst. Ent. p. 500 (1775). 3,2, Malacca;

6, Penang. “In grassy places” (W. L. Distant).
Genus Mycaxzsts, Hiibner.

1. MycanEsis MINEUS, (Papilio mineus) Linneeus, Syst. Nat. i. 2, p. 768. n. 126 (1766).
d %, Malacea; 2, Penang.

2. MYCALESIS POLYDECTA, (Papilio polydecta) Cramer, Pap. Exot. ii. pl. 144. figs. H, F
(1779). 3, Malacca, Penang.

3. Mycatusis JANARDANA, F. Moore, Cat. Lep. E.I. C. i. p. 284 (1857). 3, Malacca.

4. MycaLEsis HESIONE, (Papilio hesione) Cramer, Pap. Exot. i. pl. 11. figs. C, D (1779).
3d ¢, Malacca, Penang.

5. Mycazsis Fusca, (Dasyomma fuscum) Felder, Wien. ent. Monatschr. iv. p. 401.
n. 27 (1860). Malacca.

YrrHima, Westwood.
1. Ypruima PHILOMELA, Hiibner, Zutr. ex. Schmett. figs. 83, 84 (1806). 3, 2, Malacca;
?, Penang.
2. YPHTHIMA (sic) METHORA, Hewitson, Trans. Ent. Soc. 3rd ser. ii. p. 291, ( Ypthima)
pl. 18. figs, 20, 21 (1865). ¢ %, Malacca; 2, Penang.

3. YPTHIMA CORTICARIA, n. sp. Wings above greyish brown, an inner submarginal
streak and an outer submarginal line blackish diffused; primaries with a large subapical
oval black ocellus, with two plumbaginous pupils and a testaceous iris surrounded by a
dusky line; a similar smaller unipupillated rounded ocellus near anal angle of second-
aries. Wings below white, densely reticulated with brown, two ill-defined, interrupted,
central, subparallel lmes; a submarginal streak and line as above: primaries with the
ocellus brighter and better defined than above; outer border pale brown, bounded by
the inner submarginal streak: secondaries with three ocelli, one at apex (below which
is sometimes another extremely small ocellus) and two placed obliquely at anal angle,
the lower one small and irregular. Expanse of wings Linch 8 lines. Allied to Y. nareda.
3 ¢, Malacca.

Subfamily ExyMnriin#, Kirby.

Genus Enymnias, Hiibner.

1. HiyMnras NIGRESCENS, Butler, P. Z. S. p. 520, pl. 42. fig. 1 (1871). 3 2, Malacca,
Penang. A long series of this species, exhibiting no variation.

538 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

2. ELYMNIAS MEHIDA, (Melanitis mehida) Hewitson, Exot. Butt. iii. Wel. pl. 1. figs. 2,3
(1863). Malacca.

Subfamily Morruin#, Butler.

Genus AMATHUSIA, Fabricius.

1. AMATHUSIA PHIDIPPUS, (Papilio phidippus) Linneeus, Syst. Nat. i. 2, p. 752 (1767).
3 ?, Malacca.

Two forms (probably seasonal varieties) occur in Malacca, the one with dark bands
below, the other with pale bands; in the female of the dark form the subapical yellowish
patch on the upper surface of primaries is more distinct. I have seen three examples in
Mr. Distant’s collection from Penang. It occurs in August and September.

Genus ZEvuxrip1A, Hibner.

1. ZEUXIDIA AMETHYSTUS, Butler, P.Z.S. p. 485. n. 5 (1865). 3 2, Malacca. The
female differs from the male much in the same way as in the sexes of Z. doubledayi.

Genus DiscopHora, Boisduval.

1. DiscopHorA MENETHO, (Papilio menetho) Fabricius, Ent. Syst. iii. 1, p. 83. n. 260
(1793). &, Malacca.

2. DIscOPHORA TULLIA, (Papilio tullia) Cramer, Pap. Exot. i. pl. 81. figs, A, B (1779).
& ?, Malacca. The typical male differs from that of the Indian form in the unspotted
character of the upper surface.

Genus XANTHOTZNIA, Westwood.

1. XANTHOTANIA BUSIRIS, (Clerome busiris) Westwood, Trans. Ent. Soe. ser. ii. vol. iv.
p- 187 (1858). 2, Malacca.

Genus THAUMANTIS, Htibner.

1. THAUMANTIS NOUREDDIN, Westwood, Gen. Diurn. Lep. p. 337. n. 6, note (1851) ;
Trans. Ent. Soe. ser. i. vol. iv. p. 175. pl. 20 (1858). ¢, Malacca. ‘‘ Occurs at Penang
in August and September” (WV. L. Distant).

2. THAUMANTIS PSEUDALIRIS. (Pl. LXVIII. fig. 1.) Zhauwmantis aliris 8, Westwood,
Trans. Ent. Soc. n.s. vol. iii. p. 176 (1856-58); Butler, Journ. Linn. Soc. xiii. p. 115.
This species differs from the male of Z. aliris from Borneo in having the band of
primaries above half the width, not notched, and yellower in tint; the basal area of all
the wings ferruginous. Below, the area beyond the band is not striated, and is of the
same rusty red colour as the broad outer border, the latter is also paler at the edge, and
is cut much shorter by the obliquity of the transverse band ; the basal spots are rusty
red instead of red-brown and black; the irregular band of secondaries is paler and much
more constricted in the centre; the external and discal areas are altogether paler and
more uniform in tint; the ocelli are considerably smaller and paler; the spots between

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 539

the ocelli and the yellow submarginal spots are obsolete. Expanse of wings 4 inches

7 lines. &, Malacca.
As Professor Westwood figures the Bornean female, I am obliged to rename the

Malacca male.

Subfamily NympHatin», Bates.

Genus DoLESCHALLIA, Felder.
1. DoLESCHALLIA PRATIPA, Felder, Wien. ent. Monatschr. iv. p. 399 (1860). ¢ 2,
Malacca. This species has hitherto been represented in the Museum collection by only
one female; it differs considerably from D. disaltide in both sexes.

Genus CHARAXES, Ochsenheimer.

1. CHARAXES JaLysus, Felder, Reise der Nov., Lep. iii. p. 438, pl. 59. fig. 5 (1867).
$, Malacca. Mr. Distant gave me an example of this species taken by: him at Province
Wellesley.

2. CHARAXES HEBE, Butler, P. Z. 8. p. 634, pl. 37. fig. 3 (1865). ¢ %, Malacca. The
male does not differ at all in pattern from the female.

3. CHARAXES SCHREIBERI, (Mymphalis Schreiber) Godart, Enc. Méth. ix. Suppl. p. 825
(1823). ¢, Malacca.

Genus Apottas, Boisduval.
1. ApoutAs aponta, (Papilio adonia) Cramer, Pap. Exot. iii. pl. 255, figs. C, D (1782).
3 3, Malacca.
2. ADOLIAS GARUDA, Moore, Cat. Lep. E.I. C. i. p. 186. n. 374(1857). &, Malacca.

3. ADOLIAS JAMA, Felder, Reise der Nov., Lep. ili. p. 431 (1867). ¢ 2, Malacca. This
species appears to be common.

4. 2 ADOLIAS LAVERNA, Butler, Cist. Hnt. i. p. 29 (1870). ¢ 2, Malacca. Now
that both sexes of this species have come from the same locality, I find that the male
from Borneo figured in my ‘ Lepidoptera exotica’ is a distinct species, the secondaries
of typical A. laverna being totally unrelieved by bright colouring, and much resembling
those of A. garuda.

5. ADOLTAS ASOKA, Felder, Reise der Nov., Lep. iil. p. 433, pl. 58. fig. 1 (1867). 3 ¢,
Malacca. ‘This species was not previously in the collection of the British Museum.

6. Apottas MoNINA, (¢ Papilio monina) Fabricius, Mant. Ins. ii. p. 51 (1787). 3 2,
Malacca. The female, although much like 4. puseda, is rather different. The following
is probably only a variation of 4. monina.

7. ADOLIAS LUDEKINGII, Vollenhoven, Tijd. Ent. v. p. 189, pl. 10. fig. 3 (1860).
3 2, Malacca.

540 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus SyMPHzADRA, Hiibner.

1. SYMPHZDRA DIRTEA , (Papilio dirtea) Fabricius, Ent. Syst. iii. 1, p. 59. n. 184
(1793). 32%, Malacca, 3, Penang.

“ Slices of cut pineapple placed along a road that ran by this jungle, were generally
sure in a short time, at the proper season, to be visited by a good supply of both males
and females.” “Old and fallen fruits of most kinds were attractive; but sliced pine-
apple was mostly used as bait” (W. LZ. Distant Hunt. Mo. Mag. vol. xii. p. 207).

Genus Tanatcra, Butler.

1. Tawaiticta arnuNA, (Adolias aruna) Felder, Wien. ent. Monatschr. iv. p. 400. n. 24
(1860). ¢, Malacca. Not previously in the Museum collection.

2. TANACIA PULASARA, (Adolias pulasara) Moore, Cat. Lep. E.I. C. i. p. 190 (1857).
3 %, Malacca. The male is very closely allied to 7. varwna of Vollenhoyen, but has
the apical area of secondaries white, and the band between the zigzag lines of primaries
broader.

Genus AtHyMA, Westwood.

1. ATHYMA LARYMNA, Westwood & Hewitson, Gen. Diurn. Lep. pl. 35. fig. 1 (1850).
3, Malacca.

2. ATHYMA LEUCOTHOH, (Papilio leucothoé) Linnzeus, Mus. Lud. Ulr. p. 292 (1764).
36, Penang.

3. ATHYMA KRESNA, Moore, P. Z.S. p. 12. n. 6, pl. 50. fig. 4 (1858). 3 2, Malacca.
4, ATHYMA SABRATA, Moore, P. Z.S. p. 18. n. 10, pl. 51. fig. 1 (1858). ¢ 2, Malacca.

5. ATHYMA NIVIFERA, n. sp. (Pl. LXIX. fig. 4.) Athyma nefte, Moore (ex parte),
P. Z. S.1858, p. 18. This species may be at once distinguished from A. nefte of Cramer
by the much narrower central band, which in the male is much more distinctly blue at
the edges; by the longer and narrower trifid subapical band on the male, and the deeper
colour of the underside. Expanse of wings 2 inches 5-9 lines. ¢ ?, Malacca. It is
also common in Borneo, and occurs at Assam.

6. Arayma amHarA, Druce, P. Z.8. p. 344, pl. xxii. fig. 2 (1873). ¢, Malacca. In
the examples from Malacca the discoidal white streak of primaries is reduced to one or
two white dots, connected by a pale brown streak.

7, ATHYMA CLERICA, n. sp. (Pl. LXIX. fig. 5.) Nearly allied to A. abiasa, but larger,
the spots of primaries more oblique and larger, the subbasal transverse white band of
secondaries narrower; the discal series of spots much larger, more inarched above anal
angle, forming a waved band, divided by the nervures; a well-marked greenish-grey
submarginal streak. Below much paler, with the differences of the upper surface; sub-
marginal series of lituree replaced by the submarginal streak, which is rosy greyish (not
eveenish, as above). Expanse of wings 2 inches 7 lines. 6, Malacca.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 541

8. ArHyma rp1Ta, Moore, P. Z.8. p. 16. n. 16, pl. 51. fig. 8 (1858). ¢ 2, Malacca.

9. ATHYMA PRAVARA, Moore, Cat. Lep. EI. C. i. p. 173, pl. 5a. fig. 4 (1857).
3, Malacca.
Genus Eurrpeus, Westwood.
1. Euripus EUpL@omES, Felder, Reise der Nov. Lep. iii. p. 415. n. 688 (1867).
3 %, Malacca. This species is a mimic of Calliplea diocletianus.

Genus EvLacura, Butler.

1. EULACURA OSTERIA, (Apatura osteria) Westwood, Gen. Diurn. Lep. p. 305, note
(1850). 3 ?, Malacca.
Genus LEBADEA, Felder.

1. LEBADEA ALANKARA, (Aconthea alankara) Horsfield, Cat. Lep. E.I. C. pl. 5. fig. 6
(1829). 3, Malacca. This appears to be the male of the succeeding species.
2. LEBADEA MARTHA, (Papilio martha) Fabricius, Mant. Ins. ii. p. 56. n. 555 (1787).
?, Malacca.
Genus Limenitts, Fabricius.
1. Limeniris PRocris, (Papilio procris) Cramer, ii. pl. 106. figs. E, F (1779).
$2, Malacca; ¢, Penang. ‘‘ Very common at Province Wellesley” (W. L. Distant).

Genus Neptis, Fabricius.

1. NEPTIS MAMAJA, n. sp. (Pl. LXIX. fig. 3.) Nearly allied to WV. ewrynome, but
always to be distinguished by the narrower externo-discal band of white spots in secon-
daries, colouring below rather brighter than in NV. ewrynome. Expanse of wings 1 inch
11 lines to 2 inches 4 lines. ¢ 2, Malacca and Penang. This appears to be the com-
monest Neptis in Malacca.

2. NEPTIS LEUCONATA, n. sp. (Pl. LXIX. fig. 1.) Wings above olive-brown, banded
and spotted with cream-colour; arrangements of markings as in WV. nata, but the bands
of secondaries wider and nearer together; pale brown intermediate streaks better defined.
Wings below slightly redder; bands and spots as above; intermediate streaks white.
Expanse of wings 2 inches. 6, Malacca.

3. NEPTIS GONONATA, n. sp. (PI. UXIX. fig. 2.) Wings of the same shape and with
the same spots and bands as NV. soma; but all these markings pure white instead of
sordid yellowish white. Wings below chocolate-brown; bands below straighter; sub-
marginal line of secondaries below white. Expanse of wings 2 inches 1 line. 3, Ma-
lacca. Intermediate in character between NV. nata and N. soma.

4. Neptis Nata, Moore, Cat. Lep. E.I. C. i. p. 168, pl. 4a. fig. 6 (1857).. 32,
Malacca.
Vikast Group.
5. Nepris viKasi, Horsfield, Cat. Lep. E.I. C. p. 168, pl. 5. figs. 2, 2a (1829).
3, Malacca.
SECOND SERIES.—ZOOLOGY, VOL. I. 4¢

5A2 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Columella Group.
6. Nupris CoLUMELLA, (Papilio colwmelia) Cramer, Pap, Hxot. iv. pl. 296. figs. A, B
(1782). & 2%, Malacca.

Tiga Group.

7. NEPTIS DORELIA, n. sp. (Pl. LXVIII. fig. 3.) Papilio hetiodore, Fabricius (nec
Cramer nee Moore), Mant. Ins. ii. p.52. n. 516 (1787). 5 %, Malacca.

The type of the Fabrician species differs from the pretargles of WV. tiga in the Horsfield
Cabinet from Java in having a single instead of a double central arched line across the
under surface of the secondaries; this line is, in some examples, slightly thickened.

8. Nepris T1r¢A, Moore, Proc. Zool. Soc. p. 4. n. 3 (1858). 3%, Malacca. This
species agrees with an example in the Banksian cabinet, but not with the Fabrician
type; there are two central brown curved lines across the underside of the secondaries,
partially or wholly filled in with paler brown.

9. Nupris PARAKA, n. sp. (Pl. LXVIII. fig. 2.) Almost exactly like W. hordonia
above, but brighter in colour, and with the tawny bands rather broader ; the submarginal
tawny streak of primaries deeply bisinuate so as to form two wide arches, and intersected
by a black line. ‘Wings below ochre-yellow, the black portions of the upper surface .
represented by brownish stains, and partially blotched with grey: the basal area of
secondaries crossed by two black oblique liturze; the centre band bordered by two inter-
rupted black lines; the outer border intersected (above the indication of the submarginal
tawny line of the upper surface) by a slightly waved black line. Expanse of wings 1 inch
7 to10 lines. & 2, Malacca. The typical N. heliodora very nearly resembles this
species on the upper surface ; but below it is very different.

10. NEPTIS DINDINGA, n. sp. (Pl. LXVIIL. fig. 6.) Larger than W. heliodora, the
bands broader, particularly the central band of secondaries, which is twice as broad and
extends nearly to the base; subapical patch of primaries twice as wide, and deeply
indented in the centre of its inner border; submarginal tawny streak replaced by pale
brown (but intersected by the black line) in the primaries, tawny but extremely slender
in the secondaries. Differences below much as. above; discal band of secondaries broad
and black, and intersected by a whity-brown streak. Expanse of wings 2 inches,

?, Malacca.
We have a specimen in the British Museum from Moulmein, which measures 2 inches

2 lines in expanse. It is a well-marked species,
11. Nepris norDoNIA, (3, deck hordonia) Stoll, Suppl. Cram. pL 33. figs. 4, 4 D
(1790). ¢, Malacca,
Genus PANnpITA, Moore.

1. Panprra stnore, Moore, Cat. Lep. E.I. C. i. p. 182, pl. 6a. fig. 3 (1857). ¢, Ma-

lacca; Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 543.

Genus HypoLimnas, Hiibner.

1. HYPOLIMNAS BOLINA, (Papilio bolina) Linneeus, Mus. Lud. Ulr. p. 295 (1764).
3 ?, Penang.

2, HyPoLIMNAs INCOMMODA, n. sp. ¢& very similar to the male of ZH. bolina, but with
the subapical band of primaries straighter on the underside, and the pale brown sub-
marginal spots narrower and darker: expanse of wings 3 inches 1 line. ? differs from
the female of H. bolina in having a broad oblique subapical white band on the upperside
of primaries, a large diffused sordid white patch just beyond the cell of secondaries, and
the submarginal spots all separated, small, and pale brown: expanse of wings 3 inches
ll lines. 3& 2, Malacca.

This form seems to replace H. bolina in Malacca; in some respects it approaches the
Javan species H. nerina.

Genus CetTHostA, Fabricius.

1. CETHOSIA METHYPSEA, n. sp. Allied to OC. hypsina and C. penthesilea: from the
former it differs in its smaller expanse, the regularity of the subapical oblique whitish
band of primaries, in the presence of an apical submarginal series of four whitish spots
connecting the lower extremity of this band with the costa, and in the much less pro-
minent angulation of the outer edge of the red band on the under surface of secondaries ;
from the latter it differs in its smaller expanse, much brighter coloration, the yellower
tint of the subapical band of primaries, the more irregular and much more broadly
white-bordered red band below; the primaries are alse more produced. Expanse of wings
2 inches 9 lines to 3 inches 3 lines. ¢, Penang. The larger measurement is taken
from examples previously in the collection. We have a female from Assam.

2. CETHOSIA HYPSINA, Felder, Reise der Nov., Lep. iii. p. 885 (1867). 3 2, Malacca.
Also in the Museum collection from Nepal and Assam.

Genus CrrrocHroa, Doubleday.

1. CrRRocHROA oRISSA, Felder, Wien. ent. Monatschr. iv. p. 899 (1863); Reise der
Nov., Lep. iii. pl. 49. figs. 7. 8 (1867). 3°, Malacca; Penang.

2. CIRROCHROA JOHANNES, Butler, P. Z. 8. p. 221, pl. 17. fig. 10 (1868). 3 2, Ma-
lacca.

3. CIRROCHROA BAJADETA, Moore, Cat. Lep. E.I. C. i. p. 150, pl. 3a. fig. 3 (1857).
?, Malacca. This seems to be Felder’s C. malaya.

4, CIRROCHROA ROTUNDATA, n. sp. ¢. Nearly allied to C. mithila of Moore, the
wings more rounded, primaries less produced; markings of primaries above almost
obsolete; outer undulated line of secondaries much more distinct; below all the bands
and spots tawny instead of ochre yellow: expanse of wings 2 inches 11 lines. Malacca.
I have no doubt that this species is distinct from C. mithila; the coloration of the bands
below is strikingly different.

Ac 2

544 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus TrERINOoS, Boisduyal.
1. TERINOS TEUTHRAS, Hewitson, P. Z. 8. p. 89 (1862). 2, Malacca.

2. TERINOS ROBERTSIA, Butler, Ann. & Mag. Nat. Hist. ser. iii. vol. xx. p. 400, pl. 8.
figs. 2-4 (1867). 3%, Malacca. This species appears to be tolerably common. Mr.
Distant took both the above butterflies at Province Wellesley.

Genus Messaras, Doubleday.

1. MESSARAS ERYMANTHIS, (Papilio erymanthis) Drury, Il. Ex. Ent. i. pl. 15. figs. 3, 4
(1773). 3%, Malacca; Penang.

Genus ATELLA, Doubleday.

1. ATELLA PHALANTA, (Papilio phalanta) Drury, Ill. Ex. Ent. i. pl. 21. figs. 1, 2 (1778).
3 ?, Malacca; Penang.

2. ATELLA sInHA, (Terinos sinha) Kollar, Hiigel’s Kaschm. iv. 2, p. 438 (1848).
3, Malacca.
Genus Cyntuta, Fabricius.

1. CYNTHIA EROTELLA, n. sp. Allied to C. erota, but much smaller, brighter in colour;
the primaries generally comparatively shorter, the costal margin more arched, the ocelli
smaller in the male, and the discal lunulated line much less strongly defined; secon-
daries of the female with four ocelli, with broad bright orange irides. Wings below
slightly more reddish in colouring, with scarcely a trace of pearly grey clouding on the
external area. Expanse of wings, ¢ 3 inches 2-8 lines, ? 3 inches 7 lines. 3 ?, Ma-
lacca.

I have examined a good series of examples, and am therefore confident of the dis-
tinctness of this species. In general size and the indistinctness of the lunulated discal
line it agrees with C. deione; but it is altogether brighter in colour, with longer tails
to secondaries, and a different arrangement of the lines on the under surface of the
wings. Mr. Distant took it at Province Wellesley.

Genus PARTHENOS, Hiibner.

PARTHENOS LILACINUS, n. sp. (Minetra gambrisius, Hewitson, Gen. Diurn. Lep. pl. li.
fig. 2). Differs from P. gambrisius in the lilac internal area of primaries and basal area
of secondaries above; in the blue-green basal areas of secondaries below, and the much
better defined series of black liturze beyond the hyaline band of primaries. Expanse of
wings 4 inches 2 lines. 3o ?, Malacea ; Penang.

I have seen long series of both the Indian and Malayan forms, and am therefore
satisfied that the differences are constant. The Fabrician type agrees with our examples
from Silhet.

Genus Crrust1s, Boisduval.

1. CyrestTIs RAHRIA, Moore, Cat. Lep. H.I. C.i. pl. 8a. fig. 2 (1857). 3 2, Malacca;
?, Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 545

2. CYRESTIS NIVEA, (Amathusia nivea) Zinken-Sommer, Nova Acta. Acad. Cur. xvi.

p. 138, pl. 14. fig. 1 (1831). 3, Malacca. ‘“ Not scarce at Province Wellesley ” (W. L.
Distant).

Genus Ereoris, Boisduval.
1. ERGoLIs ARTADNE, (Papilio ariadne) Linnzeus, Syst. Nat. i. 2, p. 778. n. 170 (1767).
3 ?, Malacca; ¢, Penang.
2. ERGOLIS MERIONE, (Papilio merione) Cramer, Pap. Exot. ii. pl. 144. figs. G, H (1779).
$, Malacca; Penang.
Genus JunoniA, Hiibner.
1. Junonta rpHita, (Papilio iphita) Cramer, Pap. Exot. iii. pl. 209. figs. C, D (1782).
$ ?, Malacca; ¢, Penang.

2. JUNONIA LAOMEDIA, (Papilio laomedia) Linneus, Syst. Nat. i. 2, p. 772. n. 145
(1767). 3%, Malacca. ‘Common at Province Wellesley ” (W. L. Distant).

3. JUNONIA LEMONIAS, (Papilio lemonias) Linnzeus, Mus. Lud. Ulr. p. 277 (1764).
36, Malacca; ¢? 3, Penang.

4, JUNONIA oORITHYA, (Papilio orithya) Linneeus, Mus. Lud. Ulr. p. 278 (1764).
3 2, Malacca. ‘‘Common at Province Wellesley” (W.L. Distant).

5. JUNONIA ASTERIE, (Papilio asterie) Linnzeus, Syst. Nat. i. 2, p. 769. n. 183 (1767).
3 2, Malacca; Penang.
Genus RHINOPALPA, Felder.

1. RurNopatpa FULYA, Felder, Wien. ent. Monatschr. iv. p. 399. n. 21 (1860).
3, Malacca.

Family ERYCINID A.
Subfamily NemeEosi1inz, Bates.
Genus ZmmeERos, Boisduval.

1. ZemEros EMESOIDES, Felder, Wien. ent. Monatschr. iv. p. 396. n. 10 (1860); Reise
der Nov., Lep. ii. p. 289, pl. 36. figs. 9-11 (“1865”). 32, Malacca. This species is
evidently very common.

2. ZeMEROS ALBIPUNCTATA (PI. LXIX. fig. 10), Butler, Cist. Ent. i. p. 236 (1874).
3 2, Malacca. We have the female of this species from Borneo. It seems to be a
common species.

Genus ABIsARA, Felder.

1. ABISARA sAvirRI, Felder, Wien. ent. Mon. iv. p. 897. n.12 (1860). 39, Ma-
lacea. This is quite distinct from the Indian species A. susa of Hewitson.

2. ABISARA KAUSAMBI, Felder, Wien. ent. Mon. iv. p. 397, n. 11 (1860). 392, Ma-
lacca. A common species, the male of which is very different from 4. echerius.

546 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

3. ABISARA HAQUINUS (Papilio haquinus), Fabricius, Ent. Syst. iii. 1, p. 55. n. 169
ie @, Malacca.
Subgenus Laxrra.

4. ABISARA TANITA, (Taxila tanita) Hewitson, Ex. Butt. ii. Tax. pl. i.(1861). ¢, Ma-
lacea. The rounded wings of this and the allied species distinguish them from typical
Abiscra; but Iam doubtful of its generic distinction. TZ. orphna, placed by Kirby as
type of Taxila, must, as I have shown elsewhere, give place to 7. egeon or T. fylla,
T. orphna having been indicated in the ‘ Genera of Diurnal Lepidoptera’ as not the type
of the genus.

Family LYCAINID A.
Subfamily Lycanina, Butler.
Genus Porrrra, Moore.

1. PortTIA SUMATRA, (Pseudodipsas sumatre) Welder, Reise der Nov., Lep. ii. p. 259,
pl. 86. figs. 24-26 (1865). ¢, Malacca; ¢, Penang.

Genus Grrypus, Boisduval.
1. Gerypus Horsrrerpi, (Miletus Horsfieldi) Moore, Cat. Lep. E.I. C. i. p. 19, pl. 1a.
fig. 21857). ¢ 2, Malacca; ¢, Penang.

2. GERYDUS NIVALIS ?, (Miletus nivalis) Druce, P. Z. 8. p. 348. n. 4 (1873). Malacca.
I cannot identify this species with certainty; our example measures only 10 lines in
expanse of wing.

3. GERYDUS SYMETHUS, (Papilio symethus) Cramer, Pap. Hxot. i. pl. 149. figs. B, C
(1779). 3, %, Malacca; ?, Penang.

Genus Anops, Boisduval.

1. Anops maLayica, Felder, Reise der Nov., Lep. ii. p. 221, pl. 28. fig. 18 (1865).
36 ¢, Malacca.

2. ANOPS SPERTHIS, (Anops sperthis) Felder, Reise der Nov., Lep. ii. p. 222 (1865).
3, Malacca; ?, Penang.

Genus Lipuyra, Westwood.
1. LipHYRA BRASSOLIS, Westwood, Proc. Ent. Soc. p. 31 (1864). ¢, Malacca.

Genus Lycmnopsts, Felder.

1. LycaNopsis HARALDUS, (Papilio haraldus) Fabricius, Mant. Ins. i. p. 82. n. 744
(1787). , Malacca. This is L. ananga of Felder.

Genus CasraLius, Hubner.

1. Casratius Rostmon, (Papilio rosimon) Fabricius, Syst. Ent. p. 523. n. 341 (1775).
3 2, Malacca; Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 547

2. CASTALIUS ETHION, (d Lycena ethion) Westwood & Hewitson, Gen. Diurn. Lep.
pl. 76. fig. 8 (1852). 2, Above very like C. roxus, but with the white band extending
nearly to the costa of primaries, internally excavated and externally broadly truncate-
clayate at its upper extremity; below white, spotted with dark brown, as in the male.
Expanse of wings 11 lines. ¢?, Malacca.

3. CASTALIUS ROXUS, (Polyommatus roxus) Godart, Enc, Méth. ix, p. 659. n. 14.2 (1829),
Horsfield, Cat. pl. 2. figs. 4, 4.a (1828). & ?, Malacca,

Genus Lamprpes, Hiibner,

1. Lampipes Patan, (Lycena patala) Kollar, Hiigel’s Kaschmir, iv. 2, p. 419 (1848).
3, Malacca.

2. LAMPIDES MACROPHTHALMA, (Lycena macrophthalma) Felder, Verh. zool.-bot. Ges.
xii. p. 483. n. 115 (1862). 3, Malacca,

3. LAMPIDES BEROH, (Lycena beroé) Felder, Reise der Nov., Lep. ii. p. 275, pl. 34,
fig. 36 (1865). 3 2, Malacca.

4. Lamprpes atura, (Cupido aluta) Druce, P. Z. 8. p. 349. n. 16, pl. xxxii. fig. 8 (1878).
32, Malacca. Nearly allied to Z. nora of Felder, from Amboina, but smaller, with
less acuminate primaries.

5. LAMPIDES ALMOoRA?, (Cupido almora) Druce, P. Z.8. p, 349. n. 14, pl. xxxii. fig. 7
(1873). 3, Malacca.

6. Lamprpes BzxTIcUS, (Papilio beticus) Linneeus, Syst. Nat. i. 2, p. 789 (1767).
3 ?, Penang.

7. LAMPIDES CHRULEA, (Cupido cerulea) Druce, P. Z.8, p. 349. n. 18, pl. xxxii. fig. 6
(1873). 3, Malacca.

8. Lampipes AGNATA, (Cupido agnata) Druce, P. Z.8. p. 106, pl. xvi. figs. 2-4 (1874).
3 2, Malacca.

9. LAMPIDES PSEUDELPIs, n. sp. (Pl. LX VIII, figs. 7,8), Nearly allied to Z. elpis, which
it resembles above; below, however, with the white transverse bands narrower, irre-
cular, and broken up, the ground-colour more distinctly bluish opaline, and the large
submarginal black spot of secondaries less broadly encircled by ochreous. Expanse of
wings 1 inch 6 lines. 32, Malacca,

10. LAMPIDES KANKENA ?, (Lycena kankena) Felder, Verh. zool.-bot. Ges. xii. p. 481
(1862). 2, Malacca,

Genus CATAPACILMA, n, gen,

Allied to Lampides and Miletus, but differs from both in having three tails to the
secondaries; the antenne are long, slender, and acuminate; the wing-cells and neura-
tion generally are very like that of Lampides; the wings below are spangled with
silver, much as in Wiletus (Hypochrysops, part, Felder).

548 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

1. CATAPHCILMA ELEGANS, (Hypochrysops elegans) Druce, P. Z. 8S. p. 350, pl. xxxii.
fig. 12 (1873). 3, Malacca; Penang. The type was described from Borneo. The
figure, which is extremely rough, represents the species with three to four tails to
secondaries; but either the fourth has been broken off in all the examples which I
have examined, or does not really exist.

Genus Lycawna, Fabricius.
1. Lycmya cacaya, Felder, Reise der Nov., Lep. ii. p. 278, pl. 34. figs. 11-18 (1865).
2, Malacca.
2. LyCmNA LyYSIZONE, Snellen, Tijd. voor Ent. pl. 7. fig. 2 (1876). 3, Malacca;
Penang. Rather larger than L. karsandra; paler below.

3. LycmNA KARSANDRA, Moore, P. Z. 8. p. 505, pl. 31. fig.7 (1865). ?, Penang.

4. Lycm@NA SANGRA, (Polyommatus sangra) Moore, P. Z.8. p. 772, pl. xli. fig. 8 (1865).
3, Malacca; ?, Penang.

Subfamily Tueciinz, Butler.

Genus AmpBiyropiaA, Horsfield.

1. AMBLYPODIA CENTAURUS, (Papilio centaurus) Fabricius, Syst. Ent. p. 520. n. 329
(1775). 3, Malacca; ?, Penang.

2. AMBLYPODIA EUMOLPHUS, (Papilio ewmolphus) Cramer, Pap. Exot. iv. pl. 299. figs.
G, H (1872). 32, Malacca.

3. AMBLYPODIA ANTHELUS, Westwood, Gen. Diurn. Lep. pl. 74. fig. 6 (1852). 3, Ma-
lacca. |

4, AMBLYPODIA ADATHA, Hewitson, Cat. Lyc. B. M. pl. 4. figs. 29-31 (1862). ¢ 2, Ma-
lacca. ‘This is quite distinct from A. micale.

5. AMBLYPODIA ATosIA, Hewitson, Ill. Diurn. Lep. p. 9, pl. 2. figs. 8, 9 (1868).
36 2, Malacca.

6. AMBLYPODIA AMPHIMUTA, Felder, Wien. ent. Mon. iv. p. 396. n. 6 (1860).
3 ¢, Malacca.

7. AMBLYPODIA EPrmuTA, Moore, Cat. Lep. Mus. E.I. C. i. p. 42. n. 62 (1857).
3 2, Malacca.

8. AMBLYPODIA ANTIMUTA, (Arhopala antimuta) Felder, Reise der Nov., Lep. ii. p. 233
(1867). ¢, Malacca.

9, AMBLYPoDIA AROA, Hewitson, Il]. Diurn. Lep. p. 18, pl. 2. fig. 12 (1863). 3, Ma-
lacea. A species from India, labelled by Mr. Hewitson as 4. aroa, is more nearly allied
to A. epimuta.

10. AMBLYPODIA METAMUTA, $, Hewitson, Il]. Diurn. Lep. p. 18, pl. 2. figs. 154, 155

(1863). 3%, Malacca. The whole upper surface of the male is coloured deep purple
as in the primaries of the female, but without the black border.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 549

11. AmpLypop1a HYPoMUTA, Hewitson, Cat. Lyc. Brit. Mus. i. p. 11, pl. 6. figs. 63, 64
(1862). 32, Malacca; ¢, Penang, Quite distinct from 4. amphimuta.

12. AMBLYPODIA APIDANUS, (Papilio apidanus) Cramer, Pap. Exot. ii. pl. 187. figs. F, G
(1779). 3, Malacca.

13. Amspiypop1 ? D1arpi, Hewitson, Cat. Lyc. B. M. p. 9, pl. 5. figs. 41, 42 (1862).
$, Malacca. This species would, I think, be better placed in Uéica.

Genus Hypotycana, Felder.
1. Hypotyca#Nna ERYLUS, (Polyommatus erylus) Godart, Enc. Méth. ix. p. 638. n. 60
(1823). 32, Malacca; 2, Penang.
2. HypotycmNa? erotus, (Papilio etolus) Fabricius, Mant. Ins. ii. p. 66 (1787).
$, Malacca. This species appears to me to agree better with Wyrina.

Genus ApHnzvs, Hiibner.

1. APHN@US LOHITA, (Amblypodia lohita) Horsfield, Cat. Lep. Mus. E.I. C. p. 106.
n. 88 (1829). 3&2, Malacca; Penang.

Genus Drvporrx, Hewitson.
1. DEvDORIx PETOSIRIS, Hewitson, Ill. Diurn. Lep. p. 22, pl. 9. figs. 30, 31 (1863).
2, Malacca.

2. DEUDORIX PHERETIMA, Hewitson, Ill. Diurn. Lep. p. 21, pl. 9. figs. 27-29 (1863).
3, Malacca.

3. DEUDORIX JARBAS, (Papilio jarbas) Fabricius, Mant. Ins. ii. p. 68. n. 648 (1787).
3, Malacca. In D. jarbas the veins of the primaries are blackened, which readily distin-
guishes it from D. melampus.

4. DEuDoRIx pomitia, Hewitson, Ill. Diurn. Lep. p. 19, pl. 6. figs. 6, 7 (1863),
3, Malacca.

Genus Ionaus, Htibner.

1. Ionaus Lonernus, (Hesperia longinus) Fabricius, Ent. Syst. Suppl. p. 4380 (1798).
?, Penang.

Genus Myrina, Fabricius.

1. Myrina mecistrA (?), Hewitson, Ill. Diurn. Lep. Suppl. p. 5, pl. 3. figs. 77, 78 (1869).
Malacca.

2. Myrina TRAVANA, Hewitson, Ill. Diurn. Lep. p. 38, pl. 17. figs. 59, 60 (1865).
3 2, Malacca. The male of this is like a Dewdorix on the upper surface, the female
like an Amblypodia.

3. Myrina cuirra, (Zhecla chitra) Horsfield, Cat. Lep. E.I. C. p. 97, pl. 1. fig. 5
(1829). ?, Malacca.

4, MyrInA MARCIANA, Hewitson, Ill. Diurn. Lep. p. 34; pl. 12. figs. 12, 13 (1868).
3, Malacca.

5. Myriva Lapiruis, Moore, Cat. Lep. Mus. E.I. C. i. p. 48. n. 79 (1857). 3, Malacca.
SECOND SERIES.—ZOOLOGY, VOL. I. 4D

550 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

6. Myrina THARIS, (Oxylides tharis) Hubner, Zutr. ex. Schmett. figs. 883, 884 (1837).
3 2, Malacca.

7. Myrina amrita, Felder, Wien. ent. Mon. iv. p. 395 (1860). ¢ ?, Malacca.
8. Myrina FREJA, (Hesperia freja) Fabricius, Ent. Syst. iii. 1, p. 263 (1798). 2, Penang.

Genus Loxura, Horsfield.

1. Loxura atymnus, (Papilio atymnus) Cramer, Pap. Exot. iv. pl. 331. figs. D, EH
(1784). 32, Malacca; 3, Penang.

Family PAPILIONIDA.

Subfamily Prerina, Bates.

Genus DrxtiAs, Hubner.

1. DELIAS METARETE, n. sp. Nearly allied to D. hyparete, but longer in the wing, the
greyish apical area of primaries more uniform, in the female with a series of five or six
whitish (not yellowish) decreasing streaks between the veins; external border of second-
aries pale grey in the male, broadly blackish in the female, the red markings distinctly
visible. Apical spots of primaries below bounded within by a slender angulated greyish
line: internal area of secondaries of a much brighter yellow colour; scarlet submarginal
spots about twice as large at anal angle, not extending above the second subcostal
branch, bounded inwardly by a slender interrupted black streak. Expanse of wings
3 inches 2 lines. 3 ?, Malacca; Penang. ‘This species occurs also in Borneo.

2. Dewias pionE, (3, Papilio dione) Drury, Ill. Ex. Ent. ii. pl. 8. figs. 3, 4 (1778).
?, Malacca. The female resembles that of D. egialea, excepting that the bands are white
instead of orange.

Genus Trertias, Swainson.
1. TeRtas Formosa, (Hurema formosa) Hubner, Zutr. ex. Schmett. figs. 979, 980
(1837). 3 ?, Malacca.

2. TmrtAs sari, Horsfield, Cat. Lep. Mus. E.I. C. p. 186 (1829). 3%, Malacca.
“Occurs at Province Wellesley ” (W. L. Distant).

8. TeRIAs HECABEOIDES, Ménétriés, Cat. Mus. Petr., Lep. i. p. 85, pl. 2. fig. 2 (1855).
3%, Malacca; 3, Penang.
4, TERIASINANATA, Butler, P. Z. S. p. 617. n. 35 (1875). %, Malacca (3 examples).
5. TeRtAs pumIvaRts, Butler, P. Z. 8. p. 617. n. 36, pl. lxvii. fig. 7 (1875). 3, Malacca.

I cannot find any distinguishing characters to separate the two preceding Malayan forms
from their representatives in the New Hebrides.

Genus CatopsritA, Hiibner.

1. Caropsri1a crocate, (Papilio crocale) Cramer, Pap. Exot. i. pl. 55. figs. C, D (1779).
3%, Malacca; ¢, Penang.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 5d51

2. CATOPSILIA CATILLA, (Papilio catilla) Cramer, Pap. Exot. iii. pl. 229. figs, D, E
(1782). 3%, Malacca; 2, Penang.

3. CATOPSILIA CHRYSEIS, (Papilio chryseis) Drury, Ill. Ex. Ent. i. pl. 12. figs. 3, 4
(1773). 3&2, Malacca; Penang.

4, CATOPSILIA SCYLA, (Papilio scylla) Linneeus, Mus. Lud. Ulr. p. 242 (1764). 2, Ma-
lacca; 3, Penang. The specimens are all of the C. etesia type, having the secondaries
bright golden orange; one example nearly approaches the brighter specimens of typical
C. scylla.

Genus Hrpomora, Hiibner.

1. Hezomora GLAUCTIPPE, (Papilio glaucippe) Linneeus, Mus. Lud. Ulr. p. 240 (1764).
3, Matacca.

Genus Appras, Hiibner.

1. APPIAS PLANA, n. sp. 6d. Constantly differing from the Javan A. leptis of
Felder in the absence of the black border of secondaries, a trace only of which exists at
the apex of these wings; also larger, the primaries more produced; the apex of primaries
below and the secondaries of a clearer cream-colour: expanse of wings 2 inches 8 lines.
6, Malacca. This species is also common in Borneo.

2. APPIAS CARDENA, (Pieris cardena) Hewitson, Ex. Butt. ii. Pier. pl. 3. figs. 17, 18
(1861). ¢, Malacca. Also common in Borneo.

3. APPIAS ELEONORA, (Pieris eleonora) Boisduval, Sp. Gén. Lép. i. p. 481. n. 64 (1836),
3%, Malacca; 2, Penang. The female nearly resembles that sex of A. enarete.

4, APPIAS NATHALIA, (Pieris nathalia) Felder, Wien. ent. Monatschr. vi. p. 285 (1862).
$ %, Malacca. Occurs also at Singapore. The type described from the Philippines.

5. Appras PANDA, (Pieris panda) Godart, Enc. Méth. ix. p. 147, n. 102 (1819).
$, Malacca. This appears to be only the worn male of 4. nathalia; but the black border
seems narrower.

6. APPIAS NERO, (Papilio nero) Fabricius, Ent. Syst. rig) Ls 708 sayy ag Zl (Gl),
8 %,Malacca; ¢, Penang. Occurs also in Java and Siam.

7. ApprAas FIGULINA, (2 Pieris figulina) Butler, Ann. & Mag. Nat. Hist. ser. 3, vol. xx.
p- 399, pl. 8. fig. 1 (1867). 3%, Malacca. The male is brighter in colour than
A. nero &, and golden yellow, with a greyish streak connecting the first and second
branches of primaries, and usually a greyish indistinct nebula across the disk of second-
aries. The'type of A. figulina was from Singapore; the species occurs also in Borneo.

Genus BELENOIS, Hithbner.
1. Brnenors oynts, (Pieris cynis) Hewitson, Ex. Butt. iii. Pier. pl. 8. fig. 54 (1866).
3%, Malacca. The form figured by myself in the ‘Transactions of the Entomological

Society’ may, I think, turn out to be distinct.
4D 2

552 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Subfamily Papitioninz, Bates,

Genus ORNITHOPTERA, Boisduyal.

1. ORNITHOPTERA RUFICOLLIS, nu. sp. 6. Allied to O. flavicollis: wings smaller, com-
paratively narrower; outer margin of primaries more distinctly inarched; collar carmine.
Expanse of wings 5 inches 4 lines to 6 inches 1line. ¢ Malacca.

Unfortunately no female examples have come; the male seems to be not uncommon.
Mr. Distant has an example of the female from Province Wellesley. The primaries are
of a rather more rounded form and duller black colour than those of the male; the streaks
are more grey in colouring; the secondaries have a broad deeply indented marginal
border, six large discal spots, the veins and the base black. It was taken on the 21st of
February 1869, on the lower part of “ Batu Kawan.”

Genus PAPILIO, Linneus.
1. PAPILIo ANTIPHATES, Cramer, Pap. Exot. i. pl. 72. figs. A, B(1779). ¢ 2, Malacca.

2. PAPILIO SARPEDON, Linnzus, Mus. Lud. Ulr. p. 196 (1764). ¢ ¢, Malacca; Penang.
3. PaprLio EvEMON, Boisduval, Sp. Gén. Lép. i. p. 234 (1836). 3 2, Malacca.

4, PAPILIO AXION, Felder, Verh. zool.-bot. Ges. xiv. p. 305. n. 224, p. 350. n. 128
(1864). 3¢, Malacca. A vwariety of this species occurs in which the bands and spots
are cream-coloured.

5. Paprito BATHYCLES, Zinken-Sommer, Nova Acta Acad. Nat.-Cur. xv. p. 157, pl. 14.
figs. 6, 7 (1881). ¢, Malacca. ‘‘ P. bathycles is common round puddles in wet
weather ” (W. L. Distant).

6. Paprnio ARYCLES, Boisduval, Sp. Gén. Lép. i. p. 231. n. 51 (1836). o, Malacca.
One example was taken at Province Wellesley by Mr. Distant.

7. PapinIo AGAMEMNON, Linneus, Mus. Lud. Ulr. p. 202 (1764). 39, Malacca;
?, Penang.

8. PAPILIO MALAYANUS, Wallace, Trans. Linn. Soe. xxv. p. 59 (1865). od ?, Malacca;
$, Penang. A distinct species, like P. sthenetus above, but more like P. erithonius
below. “‘ Excessively common in gardens, August and September” (W. L. Distant).

9. PAPILIO DEMOLION, Cramer, |Pap. Exot. i. pl. 89. figs. A, B (1779). 3 2, Malacca.

10. Paritio DreLEsseRrtTI, Guérin, Deless. Souv. Voy. d. lInde, ii. p. 68, pl. 17
(1843). 3, Malacca.

11. Papriio ctytta, Linnzeus, Mus. Lud. Uly. p. 296 (1764). 3, Penang.

12. Papriio potytes, Linnzeus, Mus. Lud. Ulr. p. 186 (1764). 3 ?, Malacca; ¢, Pe-
nang. Var. P. stichiws, Hibner, Samml. exot. Schmett. (1806-16). 3%, Malacca.
The male of the latter form has the yellow spots well defined on the underside of
secondaries, just as the female has a greater amount of red colouring on the under-

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 553

side than than typical P. polytes. I suspect this variation to be seasonal, and
not polymorphic.

18. PApiILio HELENUS, Linnzeus, Mus. Lud. Ulr. p. 185 (1764). ¢ 2, Malacca.

14, Papriio PREXASPES, Felder, Reise der Noy., Lep. i. p. 107, pl. 15. fig. d (1865).
$¢ alacca. Not previously in the Museum collection.

15. Paprnio iswaRa, White, Entom. i. p. 280 (1842). o, Malacca; ¢ ?, Penang.

16. Paprnio sATURNUS, Guérin, Deless. Souv. Voy. d. l’Inde, ii. p. 70, pl. 18 (1848).
6, Malacca.

17. Pariuio EsPert, n. sp. (Pl. LXVIILI. fig. 7.) 3, Papilio protenor (part), Esper
[nec Cramer], Ausl. Schmett. pl. 29. fig. 2 (1785-98). ¢. Primaries above grey, with a
diffused broad subapical white band, the base, veins, and broad streaks between them
black, a red patch at base of discoidal cell; secondaries precisely as in the male, but with
a red spot on anal angle; differences below as above: expanse of wings 5 inches 10 lines.
$ 2, Malacca; ?, Penang.

18. Paprito mEstor, (3, Papilio androgeos), Cramer, Pap. Exot. i. pl. 91. figs. A, B
(1779). Iliades mestor, Hiibuer, Verz. bek. Schmett. p. 89. n. 931 (1816). ¢. Differs
from the female of P. Esper in having the white patch of primaries transferred to the
external angle, and the wings longer; below, the differences are the same as in the
male. Expanse of wings § inches 5 lines. ¢ ?, Malacca; 3, Penang.

19. Paprnio acHaTES, (? Papilio achates) Cramer, Pap. Exot. ii. pl. 182. figs. A, B
(1779). 3%. Wings much more uniform in colour than in the two preceding species, a red
basal streak as in P. androgeos; the red submarginal spots below much more restricted,
as in the female: expanse of wings 5 inches 6 lines. ¢ ?, Malacca.

Most Lepidopterists have united the three preceding forms in their collections as
polymorphic varieties of P. agenor, Linnzeus, noticing only the differences of the
females. I am satisfied that all three (with other forms referred to P. memnon, agenor,
and protenor) are distinct species.

20. PaPInio VARUNA, White, Entom. i. p. 280 (1842). ¢2 Malacca; Penang. The
type was described from Penang, where it is evidently common.

21. PAPILIO DIPHILUS, Esper, Ausl. Schmett. pl. 40 B. fig. 1 (1785-98). ¢ ?, Malacca;
Penang.

Family HESPERITD A.
Genus Casyapa, Kirby.

J. Casyapa THRAX, (Papilio thraz) Linneeus, Syst. Nat. i. 2, p. 794 (1767). 3, Malacca.

2. CASYAPA IRAVA (Hesperia irava), Moore, Cat. Lep. E.I. C. i. p. 254 (1857).
3 ?,,Malacca.

554 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus Hesperia, Fabricius.

1. HESPERIA HARISA, (Jsmene harisa) Moore, Proc. Zool. Soc. p. 782 (1865). 2, Malacca.

2. Husreria virra, Butler, Trans. Ent. Soc. p. 498 (1870); Lep. Exot. pl. lix. fig. 9
(1874). o&, Malacca.

The examples from Malacca have two minute transparent dots placed obliquely beyond
the cell of primaries. I have no opportunity of comparing the type at present, and
therefore do not know if there is any trace of these dots in the Bornean form.

3. HESPERIA BADRA, (Goniloba badra) Moore, P. Z.8. p. 778 (1865). 3%, Malacca.

Genus Copanus, Hiibner.

1. Copatus ELIA, (Hesperia elia) Hewitson, Trans. Ent. Soc. ser. 3, vol. ii. p. 489. n.9
(1866). o%, Malacca. The type was described from Sumatra.

2. COBALUS CILIATUS, n. sp. Dark chocolate-brown; primaries with two small spots
placed obliquely upon the median interspaces, and three dots inan angular line beyond the
cell hyaline white; secondaries with a broad abbreviated transverse band from the ab-
dominal margin, and the fringe (excepting at apex) white; body above greyish; head and
prothorax shot with golden green; abdomen irregularly white at the sides. Primaries
below as above; secondaries crossed by a broad angular white band, its lower half twice
as broad as its upper, connected with the white fringe by a white spur upon the second
median interspace; body below white, abdomen laterally banded with black lituree, anus
brown. Expanse of wings 1 inch 7 lines. ¢, Malacca.

Genus PampuiLa, Fabricius.

Section Gegenes, Hibner.

1. Pampnia arta, (Lsmene aria) Moore, P. Z. 8. p. 784 (1865). 32, Malacca. The
type was from Bengal.

2. PAMPHILA JULIANUS, (Hesperia julianus) Latreille, Ene. Méth. ix. p. 763. n. 99
(1823). <3. Malacca. Originally discovered in Java.

3. PAMPHILA MATTHIAS, (Hesperia matthias) Fabricius, Hut. Syst. Suppl. p. 483 (1798).
3%, Malacca; 3, Penang.

Section Pamphila (typical).
4, PAMPHILA AUGIAS, Linnzeus, Syst. Nat. i. 2, p. 794. n. 257 (1767). &, Malacca;
Penang.

5. PAMPHILA MASOIDES, n. sp. Above very like P. mesa of Moore, deep chocolate-
brown, banded and spotted with tawny; below similarly marked to P. mesa, but suffused
with deep tawny throughout. Expanse of wings 1 inch 1 line. 3 ?, Malacca. This
species is at once distinguished from P. mesa by the tawny colour of its bands (in
P. mesa they are ochreous); the outer band of primaries is also somewhat less irre-
gular; and the ground-colour of the underside is tawny instead of yellow.

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 55d

6. PaMPHILA MARO, (Hesperia maro) Fabricius, Ent. Syst. Suppl. p. 482 (1798).
3, Malacca; 5 ?, Penang.

7, PAMPHILA NIGROLIMBATA, (Thymelicus nigrolimbatus) Snellen, Tijd. voor Ent.
p- 165, pl. 7. fig. 5 (1876). ¢ ‘Malacca.

Genus Piastinera, Butler.

1. PLASTINGIA CALLINEURA, (Hesperia callineura) Felder, Reise der Nov., Lep. iii.
p. 578, pl. 71. figs. 9, 10 (1867). 3, Malacca.

Genus CycnLoprpsEs, Hiibner.

1. CycLoPrpEs SALSALA, (Msoniades salsala) Moore, P. Z. 8.786 (1865). ¢ 2, Malacca.
Previously known from Bengal.

Genus Astictorrervs, Felder.
1. AstIcTOoPTERUS JAMA, Felder, Wien. ent. Mon. iv. p. 401. n. 29 (1860). 3, Malacca.

2. ASTICTOPTERUS XANITES. (Pl. LXIX. fig. 7.) %. Astictopterus xanites, Butler,
Trans. Ent. Soc. p. 510 (1870). 32, Malacca. The type was from Sarawak; the male
is rather smaller, and has the orange band of primaries abbreviated and of a deeper
colour.

3. ASTICTOPTERUS GEMMIFER, n. sp. Chocolate-brown; primaries with a broad post-
median bright orange band; end of cell and apical area of primaries and disk of
secondaries spotted, in certain lights, with shining amethyst-coloured spots. Hxpanse of
wings 1 inch 6 lines. ¢ ?, Malacca (5 examples).

4, ASTICTOPTERUS ARMATUS, Druce, P. Z.S. p. 359. n. 3, pl. xxxiii. fig. 7 (1878).
?, Malacca. The type was described from Borneo.

5. ASTICTOPTERUS DiocLES, (Nisoniades diocles) Moore, P.Z.S. p. 787 (1865).
g-%, Malacca. The type was-from Bengal.

. 6. ASTICTOPTERUS sinDU, Felder, Wien. ent. Mon. iv. p. 401. n. 30(1860). ¢, Malacca.
In this species the orange band is transverse, which at once distinguishes it from the
male of A. wanites. It is also smaller.

7. ASTICTOPTERUS STELLIFER, n. sp. Above much like a small 4. jama, deep choco-
late-brown. Below irrorated, particularly at apex of primaries, with bronzy golden
scales; internal area of primaries whity-brown; a small white spot at the end of the
cell in all the wings, and a small dull white spot close to the middle of the submedian
vein of secondaries; body below pale brown; palpi clothed with golden hair-scales ;
antennze black above, yellow below. Expanse of wingslinch. 6, Malacca (2 examples).
The under surface of this species reminds one of Cyclopides salsala of Moore.

556 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Genus PLESIONEURA, Felder.

1. PLESIONEURA FOLUS, (Papilio folus) Cramer, Pap. Exot. i. pl. 74. fig. F (1779).
3, Malacca; Penang.

2, PLESIONEURA ALYSOs, Moore, P. Z. 8. p. 789 (1865). 3 2, Malacca.

3. PLESIONEURA ASMARA (Moore in litt.). Similar to P. dan, but not tawny-tinted,
the three spots in the centre of primaries united, and hyaline white: expanse of wings
linch 8lines. ¢ 2%, Malacca (5 examples).

4, PLESIONEURA DAN, (Papilio dan) Fabricius, Mant. Ins. ii. p. 88. n. 798 (1787).
3, Malacca.

5. PLESIONEURA PINWILLI, n. sp. (Pl. LXVIIL. fig. 4.) Primaries black with a
bluish shot; a broad oblique shining pale-yellow subhyaline patch, separated by the median
nervure and its second and third branches into three spots; two small spots of the same
colour, placed obliquely below it, on the interno-median interspace : secondaries bright
orange, the base and the apical portion of external border chocolate-brown ; remainder
of outer border, a rounded spot at end of cell, a second near anal angle, and five, sub-
marginal, touching the outer border, black: head and thorax greenish grey, vertex of
head edged with sordid white ; abdomen orange banded with black. Primaries below as
above, excepting that there is a bifid whitish spot above the end of the cell, a whitish
spot at base of interno-median area, and that the inner margin is brown: secondaries
bright orange; the costal and outer borders irregularly purplish black ; fringe brown-
ish; a subcostal dash, a rounded spot at the end of the cell, and a reniform spot near
the anal angle black : body below and legs bright ochreous, palpi pale ochreous; neck
below white : antennz black above, testaceous below. Expanse of wings 2 inches 2 lines.
3, Malacca. Most nearly allied to P. tabrica of Hewitson from Darjeeling.

Genus TacrapEs, Hiibner.

1. TAGIADES RAVI, (Pterygospidea ravi) Moore, Proc. Zool. Soc. p. 779 (1865).
$2, Malacca. I have compared this with the type, with which it entirely agrees.
I have seen specimens of what is evidently a nearly allied species labelled as T. ravi; they
differ in having the hind borders of the secondaries above, and the abdominal half below,
white.

2. TAGIADES GANA, (Pterygospidea gana) Moore, P.Z.S8. p. 780 (1865). ¢ ?, Malacca.
Originally discovered at Bengal.

3. TAGIADES CALLIGANA, n.sp. (Pl. LXIX. fig. 11.) Closely allied to 7. atticus, but
the secondaries much narrower, and the two black spots towards anal angle wanting,
leaving only two submarginal black spots; submarginal spots below contiguous,
almost forming a border. Hxpanse of wings 1 inch 8 lines. ¢ ¢, Malacca
(2 examples).

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 557

A, TAGIADES LAVATA, n. sp. (Pl. LXIX. fig. 8.) Wings above dark smoky brown;
primaries with a sinuous series of five subapical hyaline white dots; secondaries with anal
half of external border snow-white, its inner edge sharply defined; body brown. Pri-
maries below paler; secondaries snow-white, with the costal area brown, its inner edge
diffused ; a small brown litura across the first median branch; body below whitish.
Hxpanse of wings 1 inch 6 lines. 6, Malacca. Allied to 7. pralaya.

HETEROCERA.

Family SPHINGIDA.
Subfamily Macroeiossin# (Grote), Butler.

Genus Macroeiossa, Ochsenheimer.

1. Macroeiossa PRoxIMA, Butler, P.Z.S.p. 4, pl. 1. fig. 1 (1875). ¢, Malacca. The
figure of this species in the ‘ Proceedings’ is not satisfactory, the yellow band being
left too broad, the diverging basal black streaks being also omitted.

Family ZYGAINID Ai.
Subfamily Zveaninz, Butler.

Genus Syntomis, Ochsenheimer.

1. SYNTOMIS BASIFERA, Walker, Journ. Linn. Soc. vi. p. 92. n. 37 (1862). %, Penang.
This species is intermediate in character between §. apicalis and S. detracta.

Family ARCTIID A.

Genus Spitosoma, Stephens.
1. Spriosoma,n. sp. A wholly white species. It is in very bad condition ; and there-

fore I prefer not to name it. ¢, Penang.
Family LITHOSIIDA.
Subfamily LirHostin a.

Genus ARrGiInA, Hubner.
1. ARGINA CRIBRARIA, (Phalena cribraria) Clerck, Icones, pl. 54. fig. 4. %, Penang.

Subfamily Hypsin.z, Butler.

~Genus Hypsa, Hubner.

1. Hypsa HELIconrIA, (Phalena-Noctua helicouia) Linneeus, Syst. Nat. i. 2, p. 839
767). 2, Malacca.

2. Hypsa susstminis, Walker, Lep. Het. Suppl. i. p. 212 (1864). 2%, Malacca;
3g, Penang.
SECOND SERIES.—ZOOLOGY, VOL. I. 4 is

558 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Subgenus Damatts, Hiibner.
3. Hypsa (DAMALIS) EGENS, Walker, Lep. Het. ii. p. 453. n. 12 (1854). ¢, Malacca.

Genus NrocuERA, Hiibner.

1. NEOCHERA MARMOREA, (Hypsa marmorea) Walker, Lep. Het. vii. p. 1674 (1856).
2, Penang.

Genus CaLLipuLA, Hiibner.

1. CALLIDULA ABISARA, n. sp. Nearly allied to C. sakwni from Java, but smaller,
duller in colour; the band of primaries narrower, more regular, and rounded at its
lower extremity ; wings below more densely reticulated with brown, and duller. Expanse
of wings 1 inch 2 lines. ¢, Malacca (1 example).

Apparently the sameas the Bornean form; but the type is notin good condition. Our
Bornean examples differ from C. sakuni in the colour of the band of primaries, which
is yellow with only the inner edge orange. This is probably the case with fresh
examples from Malacca.

Genus CiEostris, Horsfield.
1. CLEostRis caTamitA, (Tetragonus catamitus) Hitbner, Zutrige, figs. 653, 654 (1832).
3, Penang.

Family NYCTEMERIDA.

Genus Nyctemera, Hiibner *.
1. NYCTEMERA TRIPUNCTARIA, (Phalena tripunctaria) Linneus, Syst. Nat. i. 2, p. 864
n. 226 (1767). 3, Malacca.
2. NYCTEMERA CoLETA (Phalena coleta), Cramer, Pap. Exot. iv. p. 153, pl. 368. fig. H
(1782). o&, Malacca.
Genus Sucusio, Walker.

1. Secusio MuNDIPrIoTA, (Nyctemera mundipicta) Walker, Journ. Linn. Soc. iv. p. 184,
n. 7 (1860). ¢, %, Malacca; 2, Penang. The type was from Singapore. I have re-
ferred the above species, with NV. ¢rita (its nearest ally), WV. plagiata, N. annulata, and
NV. distincta, to the genus Secusio; they differ from the species of Nyctemera in the

Museum.
Family CHALCOSIID A.

Genus Mintonra, Walker.

1. Mrttonta Basatis, Walker, Lep. Het. Brit. Mus. ii. p. 365 (1854). ¢, Malacca
The bands, in the single example sent, are rather wider than in females from Java.

* Type NV. coleta, the first three species belong to Otroeda, the fifth to Amnemopsyche, the seventh to Secusio ;
the two remaining forms are congeneric..

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 559

Genus EvuscHEeMA, Hiibner.

1. EUSCHEMA SUBREPLETA, Walker, Lep. Het. ii. p. 406. n. 3 (1854). Hazis bello-
naria, Guénée, Uran. et Phal. ii. pl. 18. fig. 1 (1857). ¢ %, Malacca.

Genus AmegstA, Westwood.

1. AMESIA JUVENIS, n. sp. Most nearly allied to A. venusta: primaries rich purplish
chocolate; apical area, and a series of five marginal spots ultramarine blue; a series of
eight submarginal white dots; a sickle-shaped series of five discal white dots: secon-
daries red-brown, with two submarginal series of white spots, the inner or discal series
ill defined : body blue-black ; head and tegule spotted with blue. Wings below red-brown,
costal half of primaries purplish; base of costa metallic green ; two submarginal series of
white spots uniting into streaks at anal angle of secondaries, the inner series of prima-
ries deeply incurved, the three uppermost spots of the outer series edged internally
with blue; body below metallic green spotted with white. Expanse of wings 2 inches
9 lines. ¢&, Malacca. Hasily distinguished from its allies by its pale red-brown
secondaries.

2. AMESIA PEXIFASCIA, Butler, Journ. Linn. Soe. xiii. p.115. Primaries rich purplish
chocolate, external two fifths covered by a broad externally deeply digitate dentated
snow-white band, interrupted by the black nervures, the portion crossing the end of the
cell divided through the centre into two large spots; secondaries almost exactly as in
A. eupleoides of Herrich-Schaffer, but the anal angle dark greenish-grey; body deep
purplish chocolate, the three last abdominal segments blue-green ;, palpi, tegulee, and
thorax spotted with lilacine dots. Primaries below as above, but with a basal white
spot, two costal, two subcostal, two discoidal, and two interno-median blue-edged white
spots; secondaries below nearly as in A. ewpleoides, but the anal angle grey; body
below brown, with a lateral series of blue-edged white spots. Expanse of wings 3 inches
11 lines. ¢, Malacca. The most striking species in the genus.

Genus Latrion, Walker.

1. LAURION CORCULUM, n. sp. Sepia-brown; primaries crossed by an oblique narrow
sulphur-yellow band, beyond which the apical area is almost black. Wings below paler,
greyish towards the base; primaries with the costa as far as the band blue; secon-
daries with four apical marginal steel-blue spots; sides of pectus and anterior coxz
blue; venter banded with white. Expanse of wings 1inch1line. 4d, Malacca.

Genus CHaucosta, Hiibner.
1. CHALCOSIA COLIADOIDES (var. LATIFASCIATA). Chalcosia eoliadoides, Walker, Journ.
Linn. Soc. vi. p. 97 (1862). 2, Malacca. Differs from the Bornean form in the more
regular, non-lunated, broader, subapical, curved, cream-coloured band of primaries.

Genus Cyctosta, Hiibner.
1. CycLosIaA PANTHONA, (Phalena | Geometra| panthona) Cramer, Pap. Exot. iv. p. 68,
pl. 322. fig. C (1782). %, Malacca.
AE 2

560 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Family LIPARID A.
BIRNARA, 0. gen.

Allied to Pantana: wings rounded, semitransparent; costal nervure of primaries
terminating above the end of the cell; subcostal with four branches, the third and fourth
forming a fork to apex; radials emitted near together from the upper part of the cell;
discocellulars zigzag ; median nervure with four branches, the third and fourth emitted
near together : secondaries pyriform, costal nervure extending to apex ; subcostal emitting
two branches from a short footstalk; radial forming a fourth branch of the median;
discocellulars in an oblique slightly angulated line. Type B. nubdila, n. sp.

1. BrrnaRa NUBILA, n. sp. Primaries smoky grey, the veins white, base ochreous ;
secondaries greyish or sordid white; body ochreous, antenne brown. Wings below pale
grey, becoming smoky grey towards the outer margin; body ochreous. Expanse of wings
2 inches 1 line. 2, Malacca. Allied to Pantana bicolor from China, which, with Hloria
margimalis and Genusa discifera, may be referred to this genus.

Genus KETrELiA, n. gen.

Wings semihyaline, broad: primaries subtriangular; costal nervure extending to
beyond the end of the cell; subcostal emitting five branches, the first branch emitted
before the end of the cell, the second some distance beyond the end, immediately beyond
which the vein forks, the upper ray proceeding to near apex, and then forking again,
the lower ray simple; upper radial emitted from the upper part of the discocellular
line (which is zigzag); lower radial forming a fourth median branch: secondaries pyri-
form; costal nervure proceeding to apex; subcostal emitting two branches from the
superior extremity of the cell; radial emitted from the inferior extremity of the cell on
a level with the third median; upper discocellular acutely angular, its lower portion
twice as long as its upper, the lower extremity of the cell is consequently greatly in
advance of the upper: body slender; antenne rather short, broadly pectinated. Type
K. Lowi.

1. Kerrer1a Lowit, n. sp. Semitransparent white, the borders and veins opaque; a
pale brown straight line from the costa to the external angle of primaries, and from
the apex to the anal angle of secondaries; body white; palpi head, back of tegule,
base of primaries, sides of abdomen (more or less), and the greater portion of the legs
ochreous yellow; antennze pale brown. Expanse of wings2inches5 lines. 3, Malacca;
Sarawak (Low).

Our Malacca example being rubbed, I have taken as my type one of the three spe-
cimens kindly presented to the Collection by Mr. Low. The genus is allied to Penora.

Genus Oreyi1A, Ochsenheimer.
1. ORGYIA TURBATA, n.sp. Primaries ochreous, densely irrorated with ferruginous ;
the apex, a spot near external angle, and an ill-defined subbasal spot bright golden
yellow; subapical area red-brown; two transverse sinuated blackish lines, widely

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 561

diverging to costa; a black-edged reniform ferruginous spot at end of cell; a subbasal
transverse dusky line: secondaries reddish brown. Wings below sandy brownish;
primaries with a large dusky patch over the end of thecell. Expanse of wings 113 lines.
$, Malacca. I have seen a second example from Moulmein in Mr. Atkinson’s collection.

Genus Parasa, Moore.
1. ParasA BANDURA, Moore, Cat. Lep. E.I. C. ii. p. 417. pl. xia fig. 9 (1858-9).
3, Malacca.
Family SATURNIID A.

Genus Arracus, Hitbner.
1. Arracts atias, (Phalena-Bombyx Atiacus atlas) Linneeus, Syst. Nat. p. 808. n. 1.
3 ¢, Malacca.
Family HYBLAID A.

Genus Hyspuza, Fabricius.

1. HyBL#A CONSTELLATA, Guénée, Noct. ii. p. 391. n. 1261. ¢ ?, Malacca.

Family OPHIDERID A.
Genus OPHIDERES, Boisduval.

1. OPHIDERES FULLONIA,(<, fullonia) Clerck, Icones, tab. 42. figs. 3, 4 (1759-64) ;
Ophideres cajeta, Walker (nec Cramer), Lep. Het. xiii. p. 1223 (1857); ¢, Phalena-
Noctua pomona, Cramer, Pap. Exot. ii. p.122, pl. 77. fig. C(1779). 3 2, Malacca.

Clerck gives three figures on pl. 42, and names the central one O. fullonia; it may or
may not be the O. fullonia of Linneus. I have little doubt that the true O. cajeta
is identical with Walker’s O. bilineosa. According to the example in the Banksian
cabinet (which, however, is not the type), the O. dioscoree of Fabricius is a variety of
this species in which the silvery spot of primaries is developed into a streak be-
tween the median branches.

Family OMMATOPHORIDA.
Genus Nycrreao, Hiibner.

1. NycrIpao CREPUSCULARIS, (Phalena-Attacus crepuscularis) Linneeus, Mus. Lud.
Ul. p. 878; Clerck, Icones, pl. 53. figs. 3, 4 (mec 1, 2). Malacca.

Figures 1 and 2 of Clerck’s plate represent Guénée’s W. leucotenia. ‘* Comes into the
house at night; I have never known it to be taken in any other way” (W. L. Distant).

Family BENDID A.

Genus HuLopeEs, Guénée.
1. Hunopes caRANEA, (Phalena-Noctua caranea) Cramer, Pap. Exot. ii. p. 140,
pl. 269. figs. B, F (1782). Malacca.

562 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Family THERMESIID A.
Genus Coruza, Walker.

1. CoTUZA DREPANOIDES, Walker, Lep. Het. xv. p. 1552 (1858). o, Penang.

. Family URANIID A.
Genus NycTaALEMoN, Dalman.

1. NYcCTALEMON DOCILE, n.sp. Allied to WV. hector, from which it differs in its superior
size, deeper and redder coloration, narrower central white band, which is bordered
internally by a broad brown band, and externally by a narrow one; the costal border
not spotted with white; a broad oblique diffused discal brown band: secondaries with
the apical area much less streaked with black, the anal area much more streaked ; the
marginal interrupted black border narrower; the tails considerably longer and narrower
towards their extremities. Hxpanse of wings ¢ 5 inches 6 lines, ? 6 inches 4 lines.
3 ¢, Malacca (6 examples). This species is a well-marked local representative of
N. hector.

Family URAPTERIDA.

Genus Uraptpryx, Leach.

1. URAPTERYX MARGINIPENNIS, n. sp. Wings greyish brown, transversely streaked
with clay-colour, and reticulated throughout with transverse dark-brown hatchings:
primaries with the apex broadly, and the outer margin rather narrowly, nearly sulphur-
yellow; costa whitish; a yellow spot on the second median interspace: secondaries with
the outer margin rather narrowly sulphur-yellow; two decreasing submarginal black
dots above the angle of the wing: body greyish brown. Wings below grey, spot and
borders paler than above, reticulations obsolete; body below whitish. Expanse of wings
2 inches. 6, Malacca.

This curious species is not nearly allied to any species with which I am acquainted ;
it approaches most nearly to U. crocopterata; but the wings are very slightly angulated.

Family GHOMETRIDA, Guénée.
Genus AGATHIA, Guénée.

‘1. AGATHIA DISCRIMINATA, Walker, Lep. Het. xxii. p.591 (1861). 6, Penang.

Family ACIDALIID A.
Genus ACIDALIA.

1. ACIDALIA EXxcLUSA, ( Hemerophila? exclusa) Walker, Cat. Lep. Het. xxi. p. 320
(1860) ; Macaria obstataria, Walker, l.c. xxiii. p.928 (1861). do, Malacca. Walker
notes this species as Bremer’s Philobia cinerearia; but, as I have no Chinese examples
before me, I cannot certify the correctness of this determination. |

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 563

Genus ZANCLOPTERYX, Herrich-Schaffer.
1. ZANCLOPTERYX SAPONARIA, Guénée, Phal. ii. p. 16. n. 914 (1857). Malacca.

Family MICRONIID A.
Genus MicrontA, Guénée.

1. MicrontTA ASTHENTATA, Guénée, Phal. ii. p. 24. n. 925 (1857). Malacca.

Family ZERENIID A.
Genus Genusa, Walker.

1. GenusA BicuTTA, Walker, Cat. Lep. Het. iv. p. 818. n. 1(1855). 4, Malacca.
G. delineata and circumdata must be retained among the Liparide; they are not in any
way related to G. bigutta (which I consider the type of the genus).

Family MARGARODIDA.
Genus GLYPHODES, Guénée.

1. GLYPHODES AMETHYSTA, n. sp. Basal two thirds of wings hyaline white; external
third purplish pearly, bounded internally by a sinuated black line; primaries with a
tapering bronze band from costa to inner margin at basal third, a subquadrate spot of
the same colour at the end of the cell, costa brown-bordered, a bronze brown discal
diffused tapering streak; body probably brown, but too much rubbed for description.
Hxpanse of wings 1 inch 2 lines. 6, Malacca.

Family BOTIDIDA.
Genus Borys, Latreille.

1. Borys PrEROPHORALIS, Walker, Cat. Lep. Het. Suppl. iv. p. 1418 (1865). ¢, Ma-

lacca.
2. Botys, sp. —? Closely allied to B. contigualis. In bad condition. Malacca.

Therefore it appears that of the 280 species of determinable Lepidoptera collected by
Captain Pinwill, 41 are new or hitherto unnamed forms*.

* The following Table includes the whole of the species hitherto recorded from Malacca ; the species noted in this
paper from Penang only are necessarily omitted. The names of some of the genera formerly included under Huploa
haye been altered subsequent to the reading of this paper. Consult Journ. Linn. Soc. Zool. vol. xiv. p. 290 (1878),
“On the Butterflies in the Collection of the British Museum hitherto referred to the Genus Huplea of Fabricius.”

564 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

Tabular View of the Butterflies of Malacca.

EF 3
wi | 3 3 3
Species from Malacca. 3 x ; S| a
# 5 E FI hb a S$ ie UO is ‘3 Remarks.
al2lElélelalalelalelalé
Euploea phoebus, Butler....... eee * * * | *
>, Ochsenheimeri, Lucas .. | ele bok | | ae. ee
50 Grotei, Felder .......... 26/50 los l\oel}o0 loo ]] sof} co lpoo |} ao ff ob |] on | Wpao taro Cocarin.
>» Margarita, Butler........ callooloa|l 3 oolloo | oollool| Ss
a «GUE, CGF caccnc000d Fee lPestcan Weesacia | seal h cases brewed og
, . Bremeri, Felder ........ © \loo leo || &2 || € || 2 |] 2
» Meénétriési, Felder ...... eee Was, leer tod. Wee loo lee. [es
» Pinwilli, Butler.
mridernts! LANNE Pee aee Se IW Ges eel|s eee Sea lector repeal oe 9 enemy | ok
Calliploea diocletianus, Fabrictus ..| * |..|..|*% | * | %*|..| *]*
33 vestigiata, Butler ...... Reed lzesenl uate | tater incuced| aaaee|| Seen Se.
3 Ledereri, Felder ........
Salpinx leucogonis, Butler. *
Danais plexippus, Linn. ........ Elfen lfc [Rion litetee|ccte[tereg fo ( ea [ese
5, melanippus, Cramer ...... @ loo | ca |} &3 Joo lloo | ma ye I Ss
oy wallets, JW 5 20900000 * | *e].. fe. | & | ® *
», septentrionis, Butler ...... * |...) #1 *
» grammica, Boisduval...... oa | 43 |] ool} oo lool oo] oa] &
>, melaneus, Cramer........ reLibacdanel cs
crocea, Butler .......... Se cece ieezes Il Bet Ie *
Tdeopsis daos, Boisduval ........ * | * | x hs *
Hestia lynceus, Drury .......... * * | * | *
» Jlinteata, Butler.
Melanitis leda, Linn............- * | * | * | * * * | * | x
Lethe europa, Fabricius.......... * * * * *
Ceelites humilis, Butler.
Mycalesis mineus, Hewitson ...... ocd \Weserdl Reged lle eal neuen HOS
5 polydecta, Cramer...... eel iste: ere. seo arcade |
a janardana, Moore ...... Bred pecan liccea Ibevead ivcictel este (eevos | sss
on nautilus, Butler........ *
3 hesione, Cramer ......|..|+.|.-|*%]..|..]--|%
ie fusca, Felder .......... ited bvattecll vere. Meet eS aoa
Ypthima philomela, Mibner ...... Ae se oo | £3
of methora, Hewitson...... odhee lia ||. 3
corticaria, Butler.
Elymnias nigrescens, Butler ...... lela lliga las hea Iles
as Tutescens, Butler 000000 so ||oclloo| <3 <3 |] & |] &
mehida, Hewitson ...... weal gees lige aeoh| isaead a
Amathusia phidippus, Linn....... colfcallcol & loo | & loo | & | 3
Zeuxidia amethystus, Butler...... sca lhases) Ursseeelll essed hteceall BORE [RR
Discophora menetho, Hubricius.. ..| *
6 tullia, Cramer........ SE |e Aeon eestelll eae al ioe el | pesepll eek toda asks
Xanthotenia busiris, Westwood ..|..|..|..|..|*|*|*
Clerome gracilis, Butler.......... aie. IN ceo lpee aoa gterd |S)
» faunula, Westwood ...... syeclllageedllnes al Sa ao
Thaumantis noureddin, Westwood..|..|..|..|*|..|%*
sh pseudaliris, Buéler.
Doleschallia pratipa, Helder ...... BO ora |ieeo Il 3
Charaxes jalysus, Felder ........ Bee [deel Ree pon it. 3 *
55 hebe, Butler .......... a * | *
or Schreiberi, Godart...... oe * *
Prothoe caledonia, Hewitson...... Be kscot leceen || aren | eto lfc
Adolias adonia, Cramer.......... Br NeaSel hers |fearcah Incortad | beress cease fie
= gamle, WORe 2600500000 Ae (ea oS ee Tl ead eal aoe eee 3
», Jama, Melder............ ../| *

ee ee eee ee ee SS

MR. A. G. BUTLER ON THE

Species from Malacca.

Adolias layerna, Butler
ramada, Moore
asoka, Felder.
monina, Fabricius
ludekingii, Vollenhoven....
Sympheedra dirtea, Fabricius

i, ? emalea, Guérin.
Tanaécia aruna, Felder
a pulasara, Moore
Athyma larymna, Westwood
leucothoé, Linneus
kresna, Moore

> sabrata, Moore ....... 5
> wivifera, Butler ........
so eaaalngin?, 2 an oc cooc on
>» ¢lerica, Butler.
cy Mabie, WUE eeeene oadood
Peplavata,, Moore... 4. + ch.)
> urvasi, Felder.
Euripus eupleeoides, Felder.
EHulacura osteria, Westwood ......
Lebadea alankara, Horsfield ......
> Martha, Pabricwus........
Limenitis procris, Cramer........
Neptis mamaja, Butler ...... :

leuconata, Butler.
gononata, Butler.
nata, Moore .....
vikasi, Horsjield
columella, Cramer
dorelia, Butler
tiga, Moore ....
paraka, Butler.
dindinga, Butler
» hordonia, Stoll
Pandita sinope, Moore
Hypolimnas incommoda, Butler.
Cethosia hypsina, Felder
Cirrochroa orissa, Felder
johannes, Butler.
bajadeta, Moore
a rotundata, Butler.
Terinos teuthras, Hewitson
> robertsia, Butler
Messaras erymanthis, Drury
Atella phalanta, Drury
» sinha, Kollar
Cynthia erotella, Butler
Parthenos lilacinus, Butler ......
Cyrestis rahria, Moore
>» nivea, Zinken-Sommer
Ergolis ariadne, Linneus
>, merione, Cramer
Junonia iphita, Cramer
laomedia, Linneeus
lemonias, Linneus
orithya, Linneus
5, asterie, Jinneus
eudoxia, Guérin.

39

5S)

55 +tAOMCUlA, LANNEUS ......

| Assam & Nepal.

| Moulmein.

* * KF

* KK:

| Ceylon.

* KK:

SECOND SERIES.—ZOOLOGY, VOL. I.

* | Penang.

* KK KK KK KK KK K

* eK!

BUTTERFLIES OF MALACCA.

565

| Singapore.

| ‘Borneo.

KKK,

| Sumatra.

* *

| Jaya.

* * *K OK

* KK KK KOK KK

| iene

| Guna

* * *K *

| South Pacific.

| Australia.

Remarks.

Probably more than one spe-
cies.

Probably Felder’s C. nivalis.

AF

566 MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

F :
e g
i 1 ele g ales
Species from Malacca. 3B S)a| o/s} ¢ g a | moniaks:
elg/elel®/elalelalelelé
@D a) () =| ie) =| 3 |S | 9 5) 3
4/Sl/Ol/a la /MAlals/al/olal|<
Rhinopalpa fulva, Felder.
Zemeros emesoides, Felder ...... go Wiod-|| oo |I,oo Ilo) Es
», albipunctata, Butler...... .. | *¥
Albisara sayitri, Felder .......... *
Se kausambi, Felder........ a *|* | * | *
5 haquinus, Mabricius ....| * 23 || oo *
3) tamita, Hewson 3... . - * | *
» dumayjanti, Helder.
Poritia sumatre, Felder.......... * am || %
Gerydus Horsfieldi, Moore........ x |*|*|..|/*]..]..]..].. | Possibly more than one spe-
s nivalis, Druce .........- * * cies here.
3 symethus, Cramer ...... * *
Anops malayica, Felder.
» | Sperbthis, Welder .........- ao} oa|| ool]
Liphyra brassolis, Westwood.
Lycenopsis haraldus, Fabricius....)--|--]-+-]--|--|+-|*
Castalius rosimon, /% “Brien Soa eae x |x) *¥]xe]..] ee]. | Ee] &
45 ethion, Westwood ...... oo || £3] Selool £2 ico] cal] oo |] 3
sf elna, Hewitson ........ se ll art [ence] ten facie | tees) | rout a
3 roxus, Godart ........ sonllicn oo Woodlacd| call ool ts
Lampides patala, Kollar ........ jello TES foollosiias Wicolleeloolaolloo lao) thse strom IN, Ibagtisy,
p macrophthalma, Felder ..|..| * | * |..] *|*#]--]--|..]--]--]*
Bs peroé, elder .........- Soles lactic ll eolloolloc|laall gallon || oo fae || deyeeyiexman ina arith pyerines.
3 aluta, Druce ...,.....- eer) egoIL E.
565 elum@a, JOA sn 200006 da |loa|} oo *
Cranlen, IDFA sacoo50a\|c0 ||o0|| = | co]| 20 || &
oh agnata, Druce ........ oo lloolloollool| £3 | coil co} oa] 3
8 pseudelpis, Butler.
kankena, Felder........ solles eslicstico ll oolleci oot calico ao lloo || Jats chrom ng INigolnems.
Catapceci!ma elegans, Druce...... allan 23) e3lloo | <2 lee loo | sali oalloo || ao || AWE@ tite Somtinern Jimtie,
Lyceena cagaya, Felder .......... Balloo llco|loolloe|| <2 eo llesllae |colloo lea || damae wan tne Iai pmes.
», lysizone, Snellen ........ yo || oll eo || sa) oc ger Ge
GEMS, WOOO socodoosans se lloa|lood| £3 lloall oo) colds || delice oo |foo || Ayo tarorm Itemeill,
Amblypodia centaurus, Fubricius..|..| * | * | *|--|--| *|--| x
ae eumolphus, Cramer ..|..| *|--|..| ¥|--]--] *
3 anthelus, Westwood ..| ..| *
A adatha, Hewitson....|..|..|--|--| * | ¥
5 OSE, JADUSID d6o50\]50 lla | 00 |lc0 || oo || £3 |) 53
* amphimuta, Felder <.\\.. =. == |= || a || % | =
55 epimuta, Moore...... P \loollos|lroll eoilocllee|/oell as loo) co lloo | Atyqae som Ibnetin,
3 antimuta, Felder.
mj aroa, Hewitson ...... Pree (vee |ReOn ececaa eacallicoced leg
_ metamuta, Hewitson..|)..|..]--|-.-|--|--| *
an hypomuta, Hewitson ..|..|..|--|.-|.-] *
a3 apidanus, Cramer ....|..| ¥|--|--|.-| ¥]..] *
a IDreGhi, JEWS scoollcallosal|oo|)o0llaa]}o0 las |jo0 || &
a vihara, Felder.
inornata, Felder.
Hy polycena erylus, Godart ...... x) x] .. |e]... | &
etoluss LHAVRBUS 5550) |) £2 oo lao] do || $3 os] 8
Amphneus lohita, Horsf. ....:..... Caroll to Sale elles on letoneceliice [ae eulibe:
Deudorix petosiris, Hewson <2. .\\4.\| sa fe- |ae [es | aes | ina) oe te Also from Silhet.
a pheretima, Hewitson....)..|..|--]..| * | *
if jarbas, Mubricius ...... Pra | ere Wess orl cecal cts ec (stelle
aa doinitia, Hewitson...... ae MH) ape. W! Bee even RS Oe
Myrina megistia, Hewitson.
>, travana, Hewttson........ Oya Waren len ot] ecseenleo a) as, SmI
op Ga, ZOE on coanaco re | Pet call leo call eceead IS al owed ern ee
>» Marciana, Hewitson ..... 6 lNiceems| Need | ence nace cen ee genes

a

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA. 567

2 &
| é it)
. a |S aa Utyi Ieee || 1S, Sales
Species from Malacca. Bae) a ||P Es g/8 < FH "a Remarks.
slelele/P@lEleleisl/2/3ia
a Slol/Aalscoilsi/3]s8 lis |2)58
qi/al/ola/e2lalals |aijol3ass
Myrina lapithis, Moore .......... .. | ®]e- * | * *
ap Alitierch We Mdinae Benes 66 oc * . ..|% *
= enue, JUG ne canbuD O° *
>» anasuja, Felder.
Loxura atymnus, Cramer........ ..| |] *] * # * | * | *|..|--| Type from Coromandel.
Delias metarete, Butler.......... x Leelee) #1 &] &® *
PCO, DrUrg) ocr. «oto = acs | Sia] ec er Poy sea esr avert Pe
; ninus, Wallace.
Terias formosa, Hiuibner ........ 7 aes Hears | ese *
Pe SAITSPFLONS/(ELO, | ayes have =e 0 pip ficic || oe |) <3 * *
» hecabeoides, Ménétriés ....| * |--| * | *
PPINAnabaEw Eee... 4. =o Ee fea co Ihc ce, | ectese eto) ced] owe ol rel oal 3
Ba Puulaniswebutler 2242.2. Bon | Re | aia freer fet cM onl @ ll dew, fhdeee ear Hecke
senna, Felder.
Catopsilia erocale, Cramer ......| *| * | * | * *)*}*e |e] * *
= eatilla, Cramer........ x*|*|*/|* *|..| * * *
5 chryseis, Drury ...... *| * * 25 *
i scylla, Linneus ...... 3 * * *
Hebomoia glaucippe, Linneus ....| ¥ | * | *}-- é * * Possibly more than one
Appias plana, Builer............ oe [ee}eot--|--] ® species.
» cardena, Hewitson ....... J || Saalherel ly eve lretenl) 3
» leonora, Bowsd...........: ott Sem eer hess
» nathalia, Felder.......... .-]--|--]--| ®] ¥].-]..]-.]..]..]-- | Type from the Philippines.
~ jmmube Colin coccooue se Pea|| cio ool fee il clooeell bie Ihecs
Be ETICIOS -HULUIEGIUS) a1) < -tetterel™ PY bes teal lis ete I Sal fae PE 2d ed ae an Pe
figulina, UBULHery Siete totarar es fests eel ie onl es We Sele
Be lonce cynis, Hewitson ........ Pe ieescah feel (oe acetal aetna 3
Ornithoptera ruficollis, Butler ....|..|--|--| *
Papilio antiphates, Cramer ...... ce fee} ee] ee] ee | ® *
» Ssarpedon, Linn........... e | #]..) #]-. | *
» evemon, Boisd.....: ..../--]--]++| *]--| *
a) Gen, MWS. csoceoooees * | * || 3 *
» bathyeles, Zinken-Sommer .|..|--|--| * | *| * *
5, abycles, Bowsd. .......... * --| *¥| *| "=| *] *
» rama, Felder.
» agamemnon, Jinn. ...... *|*| *|* * *x | x
» malayanus, Wallace...... Seared ete || eatkcculine cates leer thse | ee \
» demolion, Cramer........ ..| ¥].-] *]-..] e]..] *| *{..]..]--| Am allied form came from
», Delessertii, Guérin ...... 5 lec | een Reveal fe S. India. }
3 polytes, Zinn. .........- x |.-| #1] *|.-| *]--| *| *]--|--|--]| & theseus type.
Pe belemusse Mem iaerte steers #].- |. * | * | ¥ *
3 prexaspes, Felder.
eis wara Voce meprte ere Peso | heb Me Sat epee 3
» saturnus, Guérin ........ ES | ere | ihe Ail echo cpodl ee 3 ie
A - Wispert Butler Vs.1).. oar we x
» mestor, Hiibner.......... SE ont! on] 3
pp 2eHabess Oi CmMe a 1-v-- lie aid (234 Weta ese Hols 4 Penal ted ll se etl 263
mm NERO Ae oo enone. eu ibare | calla
» diphilus, Hsper.....-.... *|--| *| * fa ||oo |! 2 || es|| £2
oO, CHIEU Goncacec 6 Peilleaoulhon *|*
erebus, Wallace ........ *
Casy apa thrax, Linneus ........ Peat NE | ean leone peel [ee * | 2. | celts. |e. | Also irom India,
3 TDANR MOONE | ect cen 3 od ..|-.| ¥] *#] *
Hesperia harisa, Moore.......... Fae cers [lecae: || eon” || Carentan |, petal neces | ee Type from Sikkim.
x malayana, Felder ...... Sol (earn fee An allied species comes
35 yan: /35/td 17s eee eEBrS lole Beet lead Mee (ey asa | from Jaya.
Bare Mare enero |e lee en ices ee eee |e. eal ee seal ecsypertrom. Bengal:
Cobalus elia, Hewttson ........-- Bea esc otal port went foc eal| © 3
es ciliatus, Butler.

4AEZ

568

MR. A. G. BUTLER ON THE BUTTERFLIES OF MALACCA.

: | :
a |g g a | 3
Species from Malacca. 2 =e so | $ £ | Remarks.
FlelSil al Slslsslalalele
#/S\e|s/e\ S12 | e\ 21/8) 2] 2
4/Sl/OlsA/e2\/Alalael/azl/o\;al<
Pamphila aria, Moore .......... 20 * ne Ne Type from Bengal.
“3 julianus, Latrelle...... 9 * |. * *
+ matthias, Fubricius ....| * * | * be * | & | x?
3 augias, Linneus ...... * * | * : Be ace al [ena *
r mesoides, Butler.
ws maro, Hubricius........ a * | * * |..|..|..]| Also from 8. India.
= nigrolimbata, Snellen ..|..|..|..|.-|--|--}-.| *
Plastingia callineura, Felder...... Soilleacoall odeetll | Reel Sea aa avog | ake
Cyclopides salsala, Moore ........ diced orca SeaHl saiecdl etedMlieeond leer: Type from Bengal.
Astictopterus jama, Helder ...... SSCA al as AO] coset ech all
= xanibes, Butler ....|..|:.)--|..|-- || *
3 gemmifer, Butler....)..|..|.-]..]--|--| *
3 armutus, Druce a All) ee *
5 diocles, Moore...... 4a * * Type from Bengal.
a sindu, Felder ...... ye *
ey stellifer, Butler.
Plesioneura folus, Cramer........ uae AM cry Ileana oie ltors * | x
a alysos, Moore........ -. | *¥ | #] |. | ® *
i asmara, Moore ...... yea ee * Also from the Khasia hills.
‘3 dan, Fubricius ....-. ue * Also from India.
ss Pinwilli, Butler.
Tagisdes ravi, Moore............ boll oe ll oa Nee ea lltoa || 3 * |..|..|.-. | Type from Bengal.
Heme, AWNOGRA ~sadsn00c00 BR eee meat Wee lleos |e a ils or |eael Male Type from Bengal.
a calligana, Butler.
3 lavata, Butler.
$5 trichoneura, Melder...... Doel cull aekee (ete chet all ieehod |e oe Also from Bengal & Silhet.
DHS HORNS oouaacaccc0g00 65 | 38 | 83 | 94 | 46 112) 41 | 87 | 39) 26) 2 | 6

. 1. Thaumantis pseudaliris (upper surface) 3. Fig. 1
2. Neptis peraka (both surfaces). 2
3. Neptis dorelia (both surfaces) . 3
4. Plesioneura Pinwilli (upper surface). 4,
5. Salpinw (Huplea) leucogoms (upper sur- 5

face) 9. 6
6. Neptis dindinga (both surfaces). 7
7. Papilio Esperi (upper surface) ? . 8
8. Lampides pseudelpis (upper surface) ¢. 9
2), v1 5 (both surfaces) 3. 10.

EXPLANATION OF PLATES.

Puate LXVIII.

11.

Puate LXIX.

. Neptis gononata (both surfaces).

. Neptis leuconata (both surfaces).

. Neptis mamaja (both surfaces).

. Athyma nivifera (both surfaces) g.

. Athyma clerica (both surfaces) ¢.

. Hestia linteata (upper surface).

. Astictopterus xanites (upper surface).
. Tagiades lavata (upper surface).

. Euplea Pinwillt (upper surface) 3.

Zemeros albipunctata (upper surface) 3.
Tagiades calligana (upper surface).

A.S Butler Trans. Linn. Soc. SER 2. Zoo. Vor 1.Tas.68.

Neptis peraka. Thaumantis pseudaliris.

Lampides pseudelpis . ?. Lampides pseudelpis. 3.

Papilio esperi.

© Berjeau. hith. BUTTERFLIES OF MALACCA. Mintern Bros imp.

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A.G. Butler

©. Berjeau lith.

Neptis leuconata.

Euplea poowilli.d.

BUTTERFLIES OF MALACCA.

Trans. Linn. Soc.SER.2.Zoo1 Vou 1.TAB-69.

[ 569 ]

XXII. On certain Organs of the Cidaride. By Cuarius Stewart, Esq., F.L.S.,
M.R.C.S., Lecturer on Comp. Anat. St. Thomas’s Hospital.

(Plate LXX.)

Read December 6th, 1877.

SOME years since, whilst examining a spirit-preserved specimen of Dorocidaris papil-
lata, I first noticed the organs which it is the object of this paper to describe. An
undetected leak in the cell in which they were preserved unfortunately allowed them to
dry before a drawing had been made. ‘To the kindness of the Rev. A. M. Norman I am
indebted for two specimens, which have enabied me to confirm my observation, and
make the necessary drawings.

Of the regular Echinoidea the families Diadematide, Echinometrade, and Hchinide
have long been known to possess external branchie in the form of five pairs of conical
hollow processes of the outer margin of the buccal membrane. The branchie project
outwards into the surrounding water, their interior being bathed by the fluid of the body-
cavity, of which chamber they are diverticula. Their under surface is amplified by
secondary diverticula, the whole organ having its walls strengthened by small calcareous
plates, and sometimes spicules. Large cilia clothing its exterior, keep a constant current
of water over it. The bases of the branchiz are attached to the edges of the ten notches
which are usually present at the buccal margin of the corona. As might be expected,
these notches are most marked where the branchize are largest, e. g. Toxopneustes,
Spherechinus, &c.

The existence of such branchize in the Cidaride was denied by Miller; but in the
magnificent ‘ Revision of the Hchini’ of Prof. A. Agassiz, p. 694, he says:—‘‘In the
Cidaridz these gills are found in cuts of the buccal membrane itself, close to the acti-
nostome (Miller denies their existence ; yet he has himself figured the cuts through
which the gills pass in a species of Cidaris).”

I have carefully removed the spines one by one from the buccal membrane of Doroci-
daris papillata preserved in spirit, and then examined it under the microscope without
finding a trace of cuts or branchize (Pl. LXX. fig. 1); but in dry specimens the actinostome
appears notched from the shrinking of the membrane, which here occupies a gap left
between the interambulacral plates; this membrane shows a few creases, but nothing of
a branchial character.

If the abactinal or dorsal half of the corona be now removed, five remarkable organs
will be seen (fig. 2), which I believe must functionally take the place of the absent
external branchiz. They are situated immediately beneath the outer forked extremities
of the five compasses, and are diverticula of the chamber that occupies the intervals
between the jaws and their muscles, and which is bounded by the reflection of the inner
lining (peritoneum) of the test over them.

570 MR. C. STEWART ON CERTAIN ORGANS

In a full-grown specimen of Dorocidaris these branchiz are about half an inch in
length, are gracefully curled upwards, and have on their under surface a double row of
compound secondary diverticula, which, largest near the base of the branchiz, are
gradually reduced in size towards their tips. The walls of these organs are exceedingly
delicate, and contain similar spicular plates to those found in the neighbouring peri-
toneum. On either side of the base of the branchize external to the oblique ligaments of
the compasses there is a small group of bulla-like elevations of the peritoneum similar to
those found on the sides of the gills.

The size of the jaw-chamber can clearly be increased by the raising of the compasses
when the transverse muscles contract, especially if the jaws be protruded. Such move-
ments of the parts are frequently going on; so that if the chamber communicate with the
surrounding water, as it probably does near the tips of the teeth, water would then pass
in and out of the space, and bathe the interior of the gills. (I have not yet had an
opportunity of examining Asthenosoma, but should think it most probable that similar
gills are present there to those found in the Cidaride.)

If this hypothesis of one of the functions of the compasses, their muscles, and ligaments
be correct, some light is thrown on what has long been a puzzle to me, viz. of what use
they are in the economy of the regular Echini. The compasses have been said to be
used as levers by which the transverse muscles separate the points of the teeth ; and the
oblique ligaments have been thought by some to be muscles approximating the points of
the teeth. The ligamentous nature of the latter, however, is quite evident both under
the microscope and to the naked eye; and as such they are described by Valentin and
Prof. Agassiz.

There are two reasons why I consider that the compasses have nothing to do with
movements of the teeth. First, there are, exclusive of them, powerful muscles by which
all necessary movements of the jaws can be accomplished, viz. five pairs of retractors
passing obliquely downwards and inwards from the auricles to the tips of the jaws (pyra-
mids), which can also separate the teeth, five pairs of protrusors arising between the
auricles and inserted at the bases of the jaws, the whole of the contiguous faces of the
jaws giving attachment to a muscle which, passing from one to the other, would approx-
imate the teeth. Secondly, the mechanical arrangement of the compasses is unfavour-
able to their acting as teeth-movers. It seems, then, possible that the function of the
compasses is to raise the peritoneum from the base of the jaws; at the same time this
would stretch the oblique ligaments, and raise this membrane from their sides. Another
function might also be to remove injurious strain from the madreporic canal when the
jaws are protruded.

Although the pedicellarize of the Cidaridze have been described by many observers,
I have not seen the following peculiarity of the armed or chambered pedicellarize men-
tioned. In this form, when the jaw ends in a terminal fang, the chamber has a second
opening in addition to the large triangular tooth-armed orifice near the tip, the fang
itself being hollow, and perforated external to and in front of the tip, a place which
corresponds exactly with the opening of the groove in the terminal fang of the gemmi-

OF THE CIDARIDA. 571

form pedicellarize of the Echinidz, and which in them probably serves as a channel for
the injection of a poison secreted by two sac-like glands appended to each jaw-like piece
of the pedicellarian head. It would appear from this that the armed pedicellarize of the
Cidaridee are the analogues of the gemmiform ones of the Echinide. The position of the
opening in the fang is admirably adapted for the injecting of poison into a wound;
the flesh, which is compressed on its inner or convex side, has a cavity torn in it in
front, into which the poison could readily pass, from the opening of the fang being
situated in its anterior border and not at its point. A similar contrivance for a like
purpose is found in the falces of spiders, poison-fangs of most venomous snakes, &c.

Mixed with the ordinary forms of pedicellariz on or near the apical system of
D. papillata, 1 have often found the large armed variety possessing four jaws; this is,
I believe, the second instance of such an exception to the rule of there being three only,
avery remarkable form having been previously described by Sir Wyville Thomson as
existing in Asthenosoma. It differs, however, in all other respects from those of the
latter genus, only varying in the number of its jaws from the ordinary form found in
Doricidaris.

The next point to which I wish to call attention is that in Cidaris tribuloides and
Phyllacanthus baculosa, in addition to the minute calcareous arches which bridge over
the first three or four of the inner ambulacral pores, counting from the inferior opening
of the corona, and which have already been described by J. Miiller under the name
of vertebral processes, there are between all the inner pores delicate solid spines, some-
times two in number, between each pair of pores; they project downwards, and tend to
arch over the ambulacral vessel. The vertebral processes, of which these spines are
but a modification, have already been shown, in the paper by J. Miller before
referred to, to be the homologues of the auricule of the Diadematide, Hchino-
metradz, and Echinide, and of the vertebral ossicles of the Asteroidea, which
in like manner arch over the ambulacral vessel and nerve. The continuation of the
representatives of these arches throughout the whole length of the ambulacra in some
of the Cidaridze becomes therefore interesting. These spines remind one of similar pro-
cesses scattered over the inner surface of the corona in the Clypeastride.

The genital gland of D. papillata shows a spicular framework consisting of calcareous
triradiate spicula, some of which are of large size, and have their angles more or less
filled by an irregular network ; occasionally also one of the radii is prolonged, and projects
as a spine from the surface of the gland. Sir Wyville Thomson, in his ‘ Voyage of the
Challenger,’ vol. ii. pp. 227, 228, describes an interesting new species of Cidaris under
the name of C. nutriz. He points out many features of resemblance presented by it
to D. papillata, and others in which it differs from that species, amongst which is men-
tioned the paucity of spicula in the walls of its genital glands, whereas D. papillata is
said to have them in great abundance. Having examined a considerable number of
the latter species, I find great variation in the number of the spicula, although their
general form is constant in similar specimens, those having comparatively small genital
tubules having the spicula most abundant, whereas where the tubules are of large size
(females ?) the spicula are often very scanty and small.

572 MR. C. STEWART ON THE CIDARIDZ.

DESCRIPTION OF PLATE LXX.

Fig. 1. Portion of buccal membrane of Dorocidaris papillata. The tips of the five teeth are seen in the
centre. The spines have been removed from one side to show the ambulacra and the absence of
actinal cuts and external branchie. x 14 diameters.

Fig. 2. Internal branchia of D. papillata. x 8 diam.

Fig. 8. D. papillata, having upper half removed to show jaws and internal branchiz : slightly enlarged.

Figs. 4 & 5. Internal and side views of two jaws of an armed pedicellaria, showing perforation of terminal
fang. x 80 diam.

Fig. 6. Portion of ambulacrum of Cidaris tribuloides from near buccal membrane, showing vertebral
processes. x 12 diam.

Fig. 7. Side view of portion of an ambulacrum from near buccal membrane. Phyllacanthus baculosa.
x 12 diam.

Fig. 8. Portion of genital gland of D. papillata. x 60 diam.

1U

Soc. See. 2. Zoor Vou], Pr

WN. DO

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Trans. L

CStewart del.

ALATA.
LLACANTHUS BACULORA.

PAPIL

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ORO

6.CIDARIS TRIBULOIDES

A

Hanhart oop,

Y

TPN

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[ 573 J

XXIII. On Hypsiprymnodon, Ramsay, a Genus indicative of a distinct Family (Pleopo-
didee) in the Diprotodont Section of the Marsupialia. By Prof. Ownn, C.B.,
FRS., FLS., &c., Superintendent of the Natural-History Departments in the
British Museum.

(Plates LXXI. & LXXII.)

Read January 17th, 1878.

‘THE modification of the hind foot in the Marsupial series is peculiar in the way in:
which the digits depart from the typical mammalian number, 5. The peculiarity is.
chiefly shown by an atrophy of the second and third digits of a kind like that. which
affects the third digit of the fore foot in the Aye-aye—namely, attenuation without loss
of length, and this in relation to a special use distinct and apart from. the foot as a loco-.
motive or prehensile organ.

Not only are the slender metapodials of the digits 11 and 111 (Pl. LX XII. fig. 10) wrapped.
up with the others in a common sheath of skin, but also the proximal and middle
phalanges, leaving only the distal ones with their claws free, but in close contact and of
equal length; and these claws have been uniformly observed in the Marsupials with
the so-modified hind foot to be used as scratchers and cleansers of the fur.

This condition of hind foot is found in carnivorous, frugivorous, and herbivorous Mar-
supials, and is associated with as many distinct formule of the dental system.

The modification of digits 11 and 111, moreover, may coexist with the normal develop-
ment of the digits Iv and vy, and with an abnormal one of digit 1, in the form of a
powerful opposing hallux, as exemplified in the grasping hind foot of the scansorial
phalangers (woodcut, fig. 1); or it may be associated with an abnormal development of
the digits Iv and v in both breadth and length, especially of Iv, with a reduction of
digit 1 to rudimental insignificance, as in the gradatorial Bandicoots (Perameles); or it
may be associated with a still greater development of digits rv and y, and especially of
Ty, with total loss of digit 1, as in the hind foot of the saltatorial Potoroos and Kan-
garoos (woodcut, fig. 2).

But in the subject of the present paper a completely developed hallux (woodcut, fig. 3;
Pl. LXXII. fig. 7,1) is combined with the fur-cleansing and locomotive toes of the
Poéphaga* ; and this singular osculant condition of foot is associated with the dentition of
the Hypsiprymnide. Thus the transitional character of the mammal under consideration
would seem to place the genus between the scansorial and saltatorial groups of the great
diprotodont division + of the Marsupial order. The analogous approach in digits mand
tt to the Kangaroo type of foot made by the Bandicoots (Peramelide) may be a mere
adaptive one in relation to their fur, and at any rate is an affair of one of the Polypro-

* «Outlines of a Classification of the Marsupialia,’ Trans. Zool. Soc. vol. ii. 1839, p. 332.
+ ‘Researches on the Fossil Remains of the Extinct Mammals of Australia,’ 4to, 1876, vol. 1. pp. xii, 107.

SECOND SERIES.—ZOOLOGY, VOL. I. 4G

574 FROF. OWEN ON HYPSIPRYMNODON.

todont suborder *, and must therefore be set aside in the consideration of the affinities
of Hypsiprymnodon, a genus which I regard as the type of a distinct group of the
Diprotodontia, for which group, from the typical completeness numerically of the toes,
the name Pleopodidee + is proposed.

Fig. 1. Fig. 3. Fig. 2.

Fig. 1. Sole, hind foot, of Phalangista.
Fig. 2. Sole, hind foot, of Kangaroo.

Fig. 3. Sole, hind foot, of Hypsiprymnodon.

This group is at present represented by a single species, Hypsiprymnodon moschatus,
Ramsay, the description of which, unaccompanied by illustration, excited much interest,
and a strong desire with the students of the Marsupial order to make a further acquaint-
ance with the new form {.

I have recently been gratified by the reception of skins of mature male and female
specimens kindly transmitted to me by the describer of the species, the present accom-
plished Curator of the Australian Museum, Sydney. The skull and bones of both fore
and hind feet were included in the dried skins; and these, being extracted, have afforded
the materials for the following notes, which, with the accompanying illustrations (Plates
LXXI. & LXXIT.), may be acceptable to the Society, although they add little to the
accurate description in the ‘ Proceedings of the Linnean Society’ of the capital of one of
our most remote and prosperous colonies.

* ¢ Researches on the Fossil Remains of the Extinct Mammals of Australia, 4to, 1876, vol. i. pp. xii, 105.

+ mdéos, full; cows, foot.

+ “ Description of a new Genus and Species of Rat-Kangaroo, allied to the Genus Hypsiprymnus, now proposed to
be called Hypsiprymnodon moschatus. By E. Pierson Ramsay, F.L.S., C.M.Z.S.,” Proceedings of the Linnean Society
of New South Wales, vol. i. pp. 33-35, 1876.

PROF. OWEN ON HYPSIPRYMNODON. 575

From the presence, direction, and relative size of the hallux, the Hypsiprymnodon
might be inferred to be a grasper, perhaps an occasional climber, though not so strictly
arboreal as the Dendrolagues and Phalangers. Its nearest affinities, as shown by the
skull and dentition, are to the Hypsiprymnide.

Both male and female of Hypsiprymnodon, as exemplified by the transmitted skins, are
of equal size; each measures 1 foot 833 inches from snout to end of tail, this being 5
inches 9 lines in the female, 5 inches 4 lines in the male, of which the tip of the tail
appears to be entire. The skins indicate a rather long and slender-bodied quadruped,
with a circumference at the haunches of about 8 inches, the trunk not tapering forward,
as in the Potoroos, in which family also the smaller size of the female is notable, though
the sexual difference of size is less in some of the Potoroos than in the Kangaroos.

If to a skull of 2 inches 7 lines (65 millims.) in length 10 millims. be added for a
snout shrunk in the dried skin, and 2 millims. for occipital integument, we get a length
of head of 8 inches. The limbs are less unequal in length than in the Potoroos placed
by Mr. Waterhouse in his subgenus Potorous, the species of which have “the hinder
legs shorter and the head more slender and pointed”’ than in the Hypsiprymni proper.

In the relative length of the head to the body, and especially to that of the hind limb,
Hypsiprymnodon recedes from the Poephagous group, especially as exemplified in the
Kangaroos and in such Potoroos as Hypsiprymnus campestris, Gd., and Hypsiprymnus
platyops, Gd. In this proportion Hypsiprymnus apicalis, Gd., comes nearest to Hypsi-
prymnodon.

The pelage of Hypsiprymnodon moschatus resembles that of the Potoroos, and differs
from that of Bandicoots, in which the hair is of two kinds, the outwardly visible
and longest being coarse and harsh to the touch, that beneath forming a soft, somewhat
scanty fur. In Hypsiprymnodon the fur is of moderate length, pretty closely applied,
having numerous rather longer interspersed fine hairs, the visible portion of which is
black or blackish, and pointed; these are relieved by the dark- and light-barred colour
of the visible part of the shorter hairs, all the hairs at their basal portions showing a
leaden greyish tint. The tips of many of the shorter hairs are of a brighter brown than
the rest, approaching to yellow. Mr. Ramsey describes the upper surface of the body of
the living or recent animal as “clothed with a close and rather stiff fur of rich golden
colour mixed with black; the head, face, and lower part of the legs dark-brownish grey ;
along the centre of the throat and chest a few patches of white” *. These, in the skin
of the male transmitted, are of a light yellowish tint; and the abdomen is a light brown.
“The young are of a more golden hue, and less white on the under parts” +. The
pelage is continued by shorter hairs from the trunk over the base of the tail for half
an inch or more, where it abruptly ceases. The rest of the tail is nude and covered
by reticularly disposed scales about 14 or 1 millim. in size to the gradually attenuated
extremity. These scales are almost black on the transversely convex upperside, and of
lighter leaden colour on the less convex underside. The diameter of the tail at the
beginning of the scaly part is 8 millims. A few minute very short hairs project from

* Tom. cit. p. 34. . t Ibid.
462

576 PROF. OWEN ON HYPSIPRYMNODON.

the interstices of the scales, almost requiring a magnifier for detection. The caudal cuticle
is in the form of lacertian scales in many Potoroos; but in these, as in Lagorchestes, the
scales are partially hidden by short decumbent hairs, varying in position in different
species ; the scaly tail is most sparingly clothed, and with short stiff hairs, in the sub-
genus Potorous, Wth. On the head there are a few scattered supralabial, malar, and
superorbital vibrisse, black, fine in texture, about an inch in length.

The muffle is nude, as in Bettongia and Potorous, the black naked skin being con-
tinued a short way back from the upper part of the nose. The nostrils are lateral,
opening close to the terminal nasal disk; they are curved, with the convexity forward ;
the disk is vertically grooved, and projects beyond the mouth. The lateral extent of the
oral cleft is 1 inch (26 millims.). From the nose-tip to the eye is 1 inch 5 lines (86
millims.). The palpebral cleft is 11 millims. From this to the meatus auditorius is
22 millims. ‘The irides are dark hazel”*. The ears are rounded; 20 millims. long by
22 millims. broad; the upper margin is slightly deflected. The basal half of the outer
surface is clothed with short fine yellowish hairs; the rest of the auricle is naked.

The limbs, fore and hind, are less unequal in length than in the Hypsiprymnide; the
forearm and paw measure 3 inches-2 lines (80 millims.), the leg and hind foot 4 inches
9 lines (122 millims.). The trunk does not taper forward, as in the Kangaroos, but
maintains a more equable thickness in agreement with the more equal length and strength
of the fore and hind limbs.

The first and fifth digits of the fore feet, (Pl. lxxii. fig. 5) though shorter.and smaller than
the rest, are by no means so reduced as in Perameles; they retain the general type of those
in Hypsiprymnus. The pollex, 1, is the shortest; its claw does not extend beyond the base
of the fore finger 11. The little finger, v, attains the distal end of the proximal phalanx
of the finger, 1v. The mid digit, 111, exceeds the adjoining ones by nearly the length of
its claw. The claws are small, subcompressed, subdecurved, not broadened above, as in
Macropus. The short hairs of the hand terminate at the base of the digits, the dorsal
tegument of which is smooth and scaly. On the palmar side of the hand the hairs cease
at the proximal border of the palm. This presents a pair of unequal longitudinally
oblong callosities, and, in advance of these, three smaller ones near the base of the digits
II, Iv, and v respectively. The longer hairs of the fore limb shorten, but less abruptly
than in the hind limb, at about a third of an inch from the wrist. The callous pad
beneath the hind foot extends from the base of the toes backward to the end of the
heel, reducing the coating of short and adpressed dark brown hairs to a part of the
upper surface.

The chief gradatorial toe, woodcut, fig. 3, Tv, is not so disproportionately long and strong,
nor is the fifth toe so weak, as in the Kangaroos and the more Kangaroo-like Potoroos.
In the Hypsiprymnide the proportions are most nearly approached by the species Potorous
murinus. The claw of digit tv has not the three-sided hoof-like character of that of the
Kangaroo; it is subcompressed, shorter and deeper than the cleansing-claws of 11 and 111.

The longer hairs of the leg terminate abruptly half an inch above the tarsus, like

* Tom. cit. p. 34.

PROF. OWEN ON HYPSIPRYMNODON. 577

the legs of a pair of trousers, beyond which the hairs become abruptly very short. They
cease behind, leaving the heel bare, but are continued over the front and sides of the
tarsus, along the upper surface of the metatarsus, and along that of the phalanges of
toe Iv as far as the claw. The dorsum of the second phalanx of the hallux and that
of the same part of toe v, are covered by scaly integument, like birds’ toes. The
same integument covers the dorsum of the common sheath of the cleansing-toes
11 and 116.

The skull of Hypsiprymnodon (Plate LX XII. figs. 1, 2, 3) is smaller, narrower, more
elongated and attenuated anteriorly than in any known Potoroo—consequently departs
further in general form from that of the Kangaroos. The alisphenoidal bulla is rela-
tively smaller than in the Potoroos. The frontals are relatively narrower in proportion
to their length than in most Potoroos, showing rather the form of those bones in some
of the smaller Kangaroos and in the Dendrolagues. But the ridges continued back-
ward from the postorbital processes are very faintly indicated in both sexes of Hypsi-
prymnodon; in Dendrolagus they bound the temporal fossee above by a well-marked
elevation: Hypsiprymnodon resembles Hypsiprymnus in this difference. The nasal
bones are larger and narrower than in any Potoroo. Their pointed ends project 4 millims.
in front of the naso-premaxillary suture. A line dropped from their tip would clear the
front border of the foremost incisor. The length of the nasals is 1 inch 3 lines (31
millims.); their basal breadth 4 lines (9 millims.). The fronto-nasal suture is on a
transverse parallel with the fore part of the orbits, not in advance thereof as in Potoroos;
its position is that which obtains in Dendrolagus; but it is transverse, not angular. The
lacrymal advances a little way upon the face; it is grooved externally, and conveys the
duct by a canal commencing in advance of the orbit, as in other Marsupials and in Birds.

The maxillary is pierced by two antorbital foramina, one small, 6 millims. in advance
of the orbit; the other, which is larger, and answers to the normal foramen in Kan-
garoos and Potoroos, is 8 millims. in advance. In Potoroos* and Kangaroos + the upper
premolar descends more or less in advance of the antorbital foramen ; in Hypsiprymnodon
the foramen is in advance of the premolar. The Potoroos differ from most Kangaroos in
the longer and narrower facial part of the skull anterior to the zygomata, which part
also tapers more anteriorly. Hypsiprymnodon carries further this difference, the cor-
responding part of the skull being relatively longer and more pointed than in Hypsi-
prymnus cuniculus e.g. The extent of the maxillary in advance of the premolar
exceeds that of the premaxillary; in the Potoroos cited below reverse proportions
prevail. The course of the maxillo-premaxillary suture resembles that in the Potoroos:
the crown of the small canine projects from the lower end of the facial part of that
suture, as in Potoroos. The palatal plate of the maxillary has a pair of minute fora-
mina on the transverse line a little in advance of the premolar. The chief vacuities of
the bony palate are the two hinder ones, each half an inch in length, together 4 lines
(13 millims.) in breadth; the pair of premaxillary fissures are relatively longer and

* Waterhouse, ‘Mammalia ;’ Hypsiprymnus penicillata, plate vi. fig. 3a; Hypsiprymnus minor, plate viii. fig. 3.
+ Owen, ‘Foss. Mamm. of Australia,’ plate Ixvi. fig. 1 (Osphranter rufus) ; plate Ixxx. fig. 1 (Phascolagus erubes-
eens); plate Ixxxiy. fig. 1 (Halmaturus ualabatus).

578 PROF. OWEN ON HYPSIPRYMNODON.

narrower than in Potoroos. The palatal vacuities are bound behind by slender trans-
verse bars of the palatines, and extend forward to the interspace betweeen the premolar
and first molar teeth. Mr. Ramsay rightly notes of the posterior palatal openings
the character of the anterior margins meeting the exterior lateral at an acute angle*.

The breadth of the skull outside the zygomata is 1 inch 3 lines (82 millims.); the
vertical diameter of the zygoma at the back part of the orbit is 5 millims. Jt does not
increase posterior to this part, as in Kangaroos, but rather loses in that dimension as it
extends to the tympanic bone.

3—3 1—1

The dental formula of Hypsiprymnodon is that of the Potoroos, viz. 7. = ne

Dp. a m. i = 30. The molar series in each jaw, according to characters of develop-
ment, include the teeth symbolized as p. 4, d.3, m.1, m.2, m.3+. In some Potoroos
(Hypsiprymnus Gilberti e. g.) the crown of the premolar deviates from the longitudinal line
of direction of the four following teeth by a slight turn outwards, and rather more so
in the mandible t. A similar outward bend of the premolar is more marked in Hypsi-
prymnodon §, in which the crown of the tooth, p. 3, presents the proportions shown in
H. Gilberti. In most Potoroos the fore-and-aft extent of the crown of the grooved and
trenchant premolar is relatively greater. The molars in Hypsiprymnodon repeat the
proportions most common in the Potoroos, the last (m. 3) being the smallest, whilst in
H. Gilberti the first (d. 4) is the smallest.

The length ofthe forearm and hand from tip of elbow to the nail, inclusive, is 3 inches
2 lines (80 millims.), of leg and foot, 4 inches 9 lines (122 millims.), of fore foot, in-
cluding carpus, 1 inch 8 lines (80 millims.), of hind foot, including tarsus, 2 inches 5 lines
(61 millims.), of face from tip of nose to base of ear 2 inches 5 lines (61 millims.).

The ulna and radius are in contact along their distal halves (Pl. LX XII. fig. 5). The
radius is more slender and of more uniform breadth than in Hypsiprymnus Gilberti or
Bettongia penicillata (Pl. LX XII. fig. 9). In these and other Potoroos the bone is pro-
duced ulnad at its distal lower two thirds asa sharp plate. The length of the radius is
1inch 9 lines (45 millims.) in Hypsiprymnodon moschatus, but is only 1 inch 6 lines
(40 millims.) in Bettongia penicillata, with twice the breadth, viz. 5 millims. ‘These
differences are notable for the exigencies of paleeontology in reference to detached bones
in caves, breccias, and drifts. The ulna of Hypsiprymnodon is 2 inches 2 lines
(55 millims.). The metacarpal and phalanges are longer and more slender in Hypsi-
prymnodon than in Hypsiprymnus, save the ungual phalanges of 11, 111, and iv, which are
both longer and stronger than in the Potoroos.

The tibia and fibula, after their proximal coarticulation, keep -apart for more than
half their length distad; in the rest of their course they are in close contact but not
ankylosed. In Bettongia penicillata the interosseous space occupies but one third of the
proximal portions of the shafts of the two leg-bones; in the rest of their extent the tibia
and fibula are ankylosed, with a slight linear indication of their primitive distinctness.

* Tom. cit. p. d4. t ‘ Odontography,’ 1844; Proceedings of the Zoological Society, 1877, p. 358. .
t Plate LXXII. fig. 8. § Ih. figs. 3 & 4.

PROF. OWEN ON HYPSIPRYMNODON. 579

The length of the tibia in Bettongia penicillata is 34 inches; in Hypsiprymnodon it
is 2 inches 8 lines.

The length of the foot-bones in Hypsiprymnus Gilberti (Plate LX XII. fig. 10) is
4inches 6 lines; in Bettongia penicillata it is 3 inches 10 lines; in Hypsiprymnodon
(ib. figs. 6 & 7) it is 2 inches 5 lines.

The entocuneiform bone (ib. figs. 6 & 7, e) is proportionally larger, especially broader,
than in the Potoroos in relation to the articulation of the metatarsal of the hallux, 1.
The rest of the structure of the tarsus adheres, save in the shorter proportion of the bones,
to the Poephagous type, exemplified in plate Ixxii. of my ‘Fossil Mammals of
Australia.’

Mr. Ramsay states that the habitat of Hypsiprymuodon moschatus is the Rockingham-
Bay district, and that it frequents the dense and damp positions of the scrubs which
fringe the rivers and clothe the sides of the coast-range. Here it is by no means rare ;
but, from its retiring habits and the density of the forest, is difficult to obtain. The
habits are chiefly diurnal, and its actions, when not disturbed, by no means ungraceful.
It procures its food by turning over the débris in the scrubs in search of insects, worms,
and tuberous roots, frequently eating the palm-berries (Piychosperma Alexandre), which
it holds in its fore paws, after the manner of the Phalangers, sitting up on its haunches,
or sometimes digging. Seldom more than one or two are found together, unless accom-
panied by the young. “In March 1874 I obtained from Mr. Broadbent a female with
two young in the pouch, very small. During the same month a half-grown young one
was shot in company with the adult male and female. They evidently breed during
the rainy season, which lasts from February to May. Both sexes havea strong, although
not disagreeable, odour of musk, which appears to be stronger in the female’’*.

In the skins transmitted to me the scrotum of the male was large, and in the usual
marsupial prepudendal position t. In the female the orifice of the pouch was in the
corresponding part of the abdomen ¢. Both the size and position of the hind thumb
(woodeut, fig. 3) indicate the foot to be a grasper ina much lower degree than that of
the Phalangers and Opossums. Mr. Ramsay does not, indeed, adduce any instance
from his own or his correspondent’s observations of the opposition, or the mode of oppo-
sition, of the hallux to the other digits in the act of seizing or of climbing. Although it —
might be inferred, if the fruit of the Ptychosperma Alexandre was plucked from the
branch and not seized after falling to the ground, that Hypsiprymnodon attained it
by climbing.

Even in that case the act would seem to be occasional or exceptional; and the sum of
the evidence tells for the ground-dwelling habits of this small and interesting Marsupial.
The hallux here is, indeed, of more moment to the biologist in its homological than in
its teleological relations. It points, as in the case of more rudimental and useless parts
in other species, to the ancestral type.

Hypothetically, the earliest Marsupials are assumed to be pentadactyle; and this
assumption is supported by the following facts. The majority of the extinct Marsupials

* Ramsay, tom. cit. p. 34. + Plate LXXI. fig. 1. + Ib. fig. 2.

580 . PROF. OWEN ON HYPSIPRYMNODON.

from British mesozoic formations are polyprotodont *; and most of the existing species
of that dental division of the Order are pentadactyle f.

Among the Diprotedonts { the scansorial Marsupials retain the pentadactyle foot
with a maximized hallux §. In the gradatorial Diprotodonts, development has wrought
on the two outer toes (Iv and v), and especially on the fourth, with ultimate loss of the
innermost or first toe. This modification, characteristic of the Potoroos, reaches its
extreme in Macropus and Cheropus. The instructiveness of the foot of Hypsiprymnodon
lies in its indications of an early stage of modification of the pentadactyle type in the
direction reached by the great Kangaroos. The fourth toe is, indeed, the largest
(woodcut, fig. 3, Pl. LX XIT. figs. 6 & 7, 1v), but not so much larger than the fifth (ib. vy).
The size of both, being less exaggerated, shows the cleansing-digits (ib. 11 & 111) on a less
dwarfed scale. Finally, the hallux (ib. 1) is not yet lost: it is small, indeed, and so
placed as hardly to be opposable to any other digit, unless to the metatarsal part of m1.

As the habit of progression on hard ground increased, the share taken by the fourth
toe, as being most on a line with or most directly continuing groundward the tibia
or main pillar of the leg, would determine a greater flow of blood to the moving
powers of that toe; and as the exercise of these powers would become more strenuous
with the increase of weight to be so moved, especially when the motion was accelerated
by leaps, so the fourth toe has come, at last, asin the great Boomer Kangaroo, to be almost
exclusively the instrument applied to the resisting earth in the saltatory mode of
locomotion.

The reduction of the toes in terrestrial birds follows an analogous course. I need
hardly premise that the fifth toe in the foot of the Lizard is not developed in any bird.
The first toe, or hallux, shows its opposable character and grasping function in the
Perchers. When retained in a ground-bird, Apéferyz e. g. (Pl. LX XII. fig. 11,1), it is, as
in Hypsiprymnodon, minute, and placed above the level of the other toes. In the Cassowary
(PL LXXIT. fig. 12) it is lost; the second toe (1) is reduced in length and breadth, but
is provided with an unusually long and slender claw, which the bird may be seen occa-
sionally to apply in preening the plumage; the third toe (11) is that which has acquired
the largest size and takes the greatest share in the act of support and progression ; the
outermost toe (Iv) is smaller, but not so reduced as the innermost (1, Pl. LX XII. fig. 12).
Here we have an analogous reduction to that shown in the Potoroo.

Finally, in the Ostrich, besides 1, the next toe (11) is lost; and the swift and powerful
course of the bird is allotted, as in the Kangaroo, to the two outer toes, and mainly to
one of these (111), the outermost toe (Iv) being much reduced, but still characterized by
its five phalanges (Pl. LX XII. fig. 18).

A thaumatogenist || may, indeed, contend that the species Struthio camelus was
created, at the beginning, by primary power, with but two toes to each foot, in size,

* © Researches on the Fossil Remains of the Extinct Mammals of Australia, with a Notice of the Extinct Mar-
supials of England,’ 4to, vol. i. pp. 12-75, plates i.-tii. (1877).

t Lom. cit. p. 105. + Ibid. p. 107.

§ Art. Marsupratra, ‘ Cyclopsedia of Anatomy,’ 8vo, 1841, p. 285, fig. 111.

|| ‘ Anatomy of Vertebrates,’ vol. ili, 1868, p. 814.

PROF. OWEN ON HYPSIPRYMNODON. 581

shape, and proportion best adapted to the need of this feathered denizen of the desert.
But the nomogenist *, in reply, asks for the significance, or an explanation, of the
retention of the metatarsal of the last toe (fig. 13, 11), the jointless pointed extremity of
which projects, like the splint-bone of the same.lost toe in the Horse, on the inner side
of the articular end of the huge metatarsal of the adjoining functional toe (111).

In speculating on the way of operation of the secondary law in the coming to be of
species and genera distinguished, among other characters, by the number and _pro-
portions of the toes, a nomogenist has hazarded the following supposition :—“ as the
surface of the earth consolidated, the larger and more produced mid hoof of the old three-
toed Pachyderms [ Paleotheriwm] took a greater share in sustaining the animal’s weight ;
and more blood being required to meet the greater demand of the more active middle toe,
it grew; whilst the side toes, losing their share of nourishment, and becoming more and
more withdrawn from use, shrank ; and so on, according to the hardening of the ground,
until only the hidden rudiments of metapodials [11 and Iv] remained, and one hoof became
maximized for all the work” +. The writer illustrates this notion by a subjoined
diagram of the structure of the foot in the ungulates of the Middle Hocene, the Miocene,
and the Pliocene periods {.

But since the publication of the volume above cited, the structure of the foot of an
older ungulate mammal, the Coryphodon of our plastic clay §, has been demonstrated by
the fossils of that genus discovered by Professor Marsh in the corresponding Tertiaries of
North America ||.

And here, as in the oldest known form of Marsupials, the primitive pentadactyle type
of foot is revealed, from which the successive perissodactyle hoofed forms have deviated
by concentration of size and force through strenuous use upon digit m1, and by
degeneracy through opposite physiological conditions affecting size and power, first of
digits 1 and v, which are lost in Palgotherium, next of digits m1 and tv, dwarfed in
Anchitherium and Hipparion, until finally these also disappear in Hquus.

An analogous modification under other functions of the hind foot is traceable in
certain extinct species of Bruta (Edentata, Cuwv.).

In Myloden digit 1 is obsolete; 1 is small, but retains the ungual phalanx;
development is concentrated on 111; the ungual phalanx is wanting and the others
stunted, in Iv and v J.

In Megatherium both digits 1 and 11 are gone; Iv and v are clawless, stunted, and
seem to have been developed in a common hoof-like callosity. The whole power of the.
foot as an uprooter is concentrated in digit 111**.

* ¢ Anatomy of Vertebrates,’ vol. iii. 1868, p. 814. t+ Tom. cit. p. 793. + Ibid. woodcut 614.

§ Owen, ‘ British Fossil Mammals,’ 8yo, 1846, p. 299, cuts 103, 10.

|| Marsh, “ Principal Characters of the Coryphodontide,” American Journal of Science and Arts, vol. xiv. July 1877,
p. 81, pl. iv. fig. 2. A more direct and continuous ancestry of existing Equines will shortly be described and
illustrated, in American Fossils, by the same author.

@ Owen, ‘ Description of Mylodon,’ 4to, 1842, p. 117, pl. xxii.

** Memoir on the Megatherium,’ 4to, 1860, p. 72, pls. xxv. andxxvi. The degree and course of mutilation and
of partial and exaggerated development of certain toes in the hind foot of other Bruta are detailed in pp. 74-77.

SECOND SERIES.—ZOOLOGY, VOL. I. 4H

582 PROF. OWEN ON HYPSIPRYMNODON.

In these ponderous slow-moving quadrupeds the exercise of the muscles and con-
comitant vigour of circulation and innervation in locomotion was inferior to those
affecting the main instrument in the swift course of the Horse and Kangaroo; the
claw-bearing toe mainly (if not exclusively) exercised in exposing the ramifications of the
root of the tree to be felled, was the subject of those influences which govern excessive
or exclusive development.

In conclusion, I may remark that the longest and strongest toe in the pentadactyle
hind foot of the Lizards is the fourth; and I deem it indicative of the nearness to
Saurians of the Implacentalia among mammals—an affinity so remarkably shown by
monotremes in other parts of the skeleton—that in the progressive reduction of the foot
in Marsupials the fourth toe, Iv, holds the preeminence.

In the march of Man the chief motive power is concentrated on the innermost toe
(1) as the main instrument, thence called the “ hallux,’”’ or “ great toe.” In this modi-

fication of the foot for bipedal station and locomotion the human kind stands alone in
the animal kingdom.

DESCRIPTION OF THE PLATES.
Puate LXXI.

Figs. 1 & 2. Hypsiprymnodon moschatus, Ramsay: nat. size. 1, male; 2, female.

Puate LXXTII.

Fig. 1. Side view of the skull of Hypsiprymnodon moschatus, Rms.
2. Upper view of the same.
3. Under view of the same. 1 1, 2, 3, incisors; c, canine; Pp 3, premolar; d4, milk-
molar; m 1, 2, 3, molars.
4.. Upper view of the mandible of the same. Letters as in fig. 8.
5. Bones of the forearm and paw of the same. 1, II, 111, Iv, V, digits.
6. Bones of the hind foot: £, entocuneiform ; 1 to v, digits.
7. The same, from the opposite (plantar) side.
8. Mandible of Hypsiprymnus Gilberti: c, canine ; p 8, premolar ; d 4, milk-molar; ™ 1, 2,3,
molars.
9. Bones of forearm and paw of the same. 1 to v, digits.
10. Bones of the hind foot, plantar aspect, of the same: 1 to v, digits. All the above
figures are of the natural size.
11. Bones of the foot of Apteryz.
12. The same bones of the Cassowary (Casuarius).
13. The same of the Ostrich (Struthio). The same letters are applicable to these three
figures, viz. e, epiphysis; 2, 3, 4, metatarsals; 1, 11, 111, 1v, digits.

ROwen. _ Trans. Linn. Soc. 5EeR.2.Zo00u-Vou.1.Pu72

; 17 AYP Ss TERY MN OD) On SRIOMENG Rio Even iaNiUiS7
CBerjean lith, TLAPTHRYX 1Z.CASUARIUS. 13.STRUTHIO Hanhart imp

[ 583 ]

XXIV. Descriptions of some minute Hymenopterous Insects.
By J. O. Wzstwoop, W/.A., F.L.S., Se.

(Plate LX XIII.)
Read April 4th, 1878.

THE fourth volume of the ‘Transactions’ of the Linnean Society (1798) contains the
figure and description of an almost invisible Hymenopterous insect, referred by the
author, Mr. Shaw, to the Linnean genus Jchnewmon, and described under the name of
Ichn. punctum, with a doubtful reference to the Ichn. atomos of Linnzeus. The specimen
is stated to have been taken on the surface of a window; and on various occasions I
have taken the same or a closely allied species creeping on windows, and of so minute
a size as to be only visible when seen against the light and in motion. The insect is
remarkable for its antennze terminating in an oval club (being a female), and the long
fringe of hairs with which the edges of the wings are furnished. It is at the present
day referred to the subfamily Mymarides among the Proctotrupide, and to the genus
Anaphes of Haliday (Entomol. Mag. i. 269).

The twenty-fourth volume of the ‘Transactions’ of the Linnean Society (1863) adds
materially to our knowledge of these minute insects by the publication of a memoir, by
Sir John Lubbock, ‘‘ On two Aquatic Hymenoptera, one of which uses its wings in swim-
ming.” In this remarkable memoir the author describes two different species—one
belonging to the Mymarides, and the other, as it seems to me, to an aberrant group of
the Eulophideous Chalcidide. Of the first of these two species as many as twenty-one
individuals were observed by Sir John Lubbock, each swimming in a basin of pond-water
with the assistance of its wings, and using its legs apparently only for walking, so that
its motion under water might almost be called a flight. This insect was at first regarded
as identical with Polynema fuscipes of Haliday, but was subsequently considered to be a
distinct species, and described under the name of Polynema natans, although that genus
possesses 13-jointed antennz in the males, whereas P. natans has only twelve joints in
its male antenne. As Sir J. Lubbock did not give any representation of the male
antenna, I have represented it in Plate LX XIII. fig. 1, from a specimen of the insect
communicated to me by Sir J. Lubbock. With regard to the genus to which this insect
belongs, it is to be observed that Dr. Arn. Foerster (Hymenopt. Studien, ii. Heft, pp. 118,
121) objects to the name Polynema, because there was a fish genus so named previously,
and to the synonymous name Hutriche of Nees von Esenbeck (Monogr. Ichn. Min.),
because Stephens had given the name of Hutricha to a Lepidopterous genus. He con-
sequently proposed for it the name of Cosmocoma. Considering that it is unnecessary
and inexpedient to alter such generic names because they happen to have been previously
employed in different classes of animals, I should have retained the generic name given

Au 2

584 PROF. WESTWOOD ON SOME

to the insect in Sir John Lubbock’s memoir, had it not been that the difference in the
number of joints in its antennz above noticed would not allow it to be arranged with
the other Polyneme, the number of joints of those organs in the Proctotrupide affording
the principal characters for generic classification. Hence I should prefer to arrange
Sir J. Lubbock’s insect in Haliday’s genus Anaphes, since it has 12-jointed male and
9-jointed female antennee, as in that genus, notwithstanding that Anaphes is placed in a
section having the abdomen sessile or subsessile, whilst it is pedunculated in Polynema;
but the figure of the female given by Sir J. Lubbock (pl. xxiii. fig. 2) shows so small
an amount of pedunculation as to throw doubt on this point; and it will be seen that
the wings of P. natans (figs. 4 & 5) agree exactly with those of Anaphes (figs. 8 & 9)*.

With regard to the natatorial habits of this curious insect, I may observe that the long
fringe of fine hairs along the margins of the wings is admirably adapted to retain a
bubble of air, needed for the respiration of the insect under water, which, as Sir John
Lubbock observes, takes place through spiracles in the usual manner. M. Victor
Audouin showed how this was effected in an insect belonging to the family Carabidee
and genus A/pus, which passed a considerable portion of its existence in salt water
beneath high-water mark, and which was clothed with long setee; and it is in like manner
owing to the sericeous covering of the Argyroneta that the diving-bell spider is able to
subsist under water.

It is a curious analogy that whilst these insects, the most minute of the Hymenoptera,
should be furnished with these beautifully fringed wings, the most minute species of the
Coleopterous order (viz. the Trichopterygia) should also have their wings similarly
fringed; no one, however, as far as I am aware, has ever observed a Trichopteryx under
water; and it may be suggested that the fringe of hairs in that genus is used as a supple-
ment to the narrow elongated wings in effecting the ordinary flight of the Beetle.

The generic name J/ymar has been retained for the singular little insect figured by
Curtis under the name of

Mymar PULCHELLUS (Plate LX XIII. fig. 3). Brit. Ent. pl. 411.

This insect has 18-jointed antennee in the male and 9-jointed in the female, with the fourt
joint remarkably elongated and slender ; the fore wings forming a long and filiform thread,
terminated by a dilated oval membrane (like a boatman’s paddle), which is deeply fringed
with hairs, whilst the hind wings are reduced to a short rigid bristle. The tarsi are
4-jointed; and the abdomen is fixed to the thorax by a long slender peduncle. As Mr..
Curtis figured the female of this insect, I have thought it advisable to give a figure of
the opposite sex (having taken specimens of both sexes by sweeping grass and low plants
on a hot bank near Richmond Park), as well as to show its difference from another
species of the genus, discovered in Ceylon by Mr. Staniforth Green, as well as from the
still more remarkable species collected in St. Helena by the late T. V. Wollaston.

* Tf this proposal as to P. natans being placed in the genus Anaphes be rejected, I beg to suggest that a new
generic name be given to it, and would carry out Sir J. Lubbock’s original intention, by giving to it the generic name
of Valkerella.

MINUTE HYMENOPTEROUS INSECTS. 585

MyMAR TAPROBANICUS (Plate LX XIII. fig. 4, ¢, and fig. 6, ?), Ward, in Ent. Mo. Mag.
Feb. 1875, p. 197.

In the work above referred to, the female only of this remarkable species is described.
I have now, however, the pleasure of describing both sexes, collected in Ceylon by
sweeping low herbage, by Mr. Staniforth Green, which are preserved as microscopical
objects in Canada-balsam slides; and the male is fortunately arranged so that its
structure can well be observed from above; but the female is fixed laterally. The head
is large, transverse, with a slight ridge in the male between the anterior part of the
eyes, the antennz being affixed on the fore edge of this ridge, considerably above the
mouth ; so that the geniculation of the antennz is quite of a different nature from that
of the Chalcididz. The antenne of the male are very long, 13-jointed, and quite filiform ;
the basal joint slightly curved, and the second joint short, the remaining eleven joints
nearly equal in length and slenderness; in the female the antenne are 9-jointed, the
first joint longer than in the male, the second joint rather short and thicker than the
apex of the preceding joint, the third joint slender and slightly longer than the second,
the fourth very long, slender, the fifth to the eighth small, and the ninth forming a large
oval mass. The thorax is elongate-ovate, the anterior part conical, and the hind part
truncated ; the abdomen is attached to the thorax in both sexes by a thickish peduncle,
slightly longer than the posterior coxe; the abdomen is very small in the male,
larger and pointed at the tip in the female, which is furnished with a short, sharply
pointed ovipositor, with two long lateral valves. The fore wings resemble those of IZ. pul-
chellus, except that they are not so strongly battle-door shaped at the tips; the disk of this
terminal part is furnished with two rows of short fine sete, andthe margins of this part
are furnished with strong bristles. The hind wings are about two thirds of the length of
the fore wings (thus differing from those of IZ. pulchellus), and are quite bristle-shaped ;
they have a few very short fine hairs on their margins and near the base; on the fore
margin they have three small sharp hooklets (fig. 7) for fastening them to the hind margin
of the fore wings. The legs are very long and slender, with 4-jointed tarsi and large
pulvilli. .

The whole insect is of an ochraceous colour, paler beneath, with the antennze and legs
concolorous; the apical half of the dilated portion of the fore wings is dusky, and the
tips of the antennze and tarsi are darker.

I also refer to the genus Mymar* avery singular species collected by the late lamented
T. Vernon Wollaston by sweeping low herbage in the Island of St. Helena, of which
fortunately, both sexes were procured, and which agree in having the fore wings of large
size, coriaceous, setose, convex, and, when shut, forming a semiglobular dome over the
abdomen, quite unlike any previously known Hymenopterous insect.

Mymar Wottastonit. (Plate LXXIII. fig. 8, d; fig. 9, 2.)
Niger, nitidissimus, glaberrimus, capite thorace paulo latiore, subquadrato-rotundato, convexo, postice
valde emarginato, thorace compresso, prothoracis lateribus subtus paullo dilatatis, pedunculo abdo-

* Tf it should be deemed necessary to separate this species from the genus Mymar, I beg leave to suggest that its
affinity therewith may be indicated by the employment of the generic or subgeneric name of Mymarilla.

586 PROF. WESTWOOD ON SOME

minis fere thoracis longitudine fulvo-testaceo ; abdomine oyali, nitido; antennis maris corpore toto
dimidio longioribus, nigris, articalis duobus basalibus fulvis ; pedibus elongatis, gracilibus, fulvis, apice
tarsorum obscuro ; tibiis posticis in utroque sexu setosis ; alis anticis ovalibus, convexis, semiglobosis,
coriaceis, nigris, valde nigro-setosis ; alis posticis fere linearibus, setosis. Long. corp. ¢ circ. ‘2 mill.,
9 -274 mill.; long. anten. maris °3 mill.

Habitat in gramimosis in Insula Sanctze Helene.

ALAPTUS Excisus. (Plate LX XIII. fig. 10, 3, and fig. 11, ?.)

I am indebted to Mr. Whitmarsh, of Wilton, near Salisbury, for an opportunity of
examining a very large number of glass slides, prepared for the microscope, containing
minute insects mounted in Canada balsam—an excellent plan for the examination of
such objects, so far as the observation of the general outline and of detached parts are
concerned, the gummy solution rendering the parts more or less transparent. Amongst
these specimens I found two insects belonging to the Mymarides, which I have no
hesitation in regarding as the male and female of the same species. Both specimens
had been reared from white blotches on oak leaves, evidently caused by the action of
the minute larva of one of the leaf-mining Vinee (Lithocolletes?). The blotches were
about 1} in. in diameter. The leaves were gathered on the 9th September, 1871; and
the little Mymars appeared on the 6th October; one of the moths appeared on the 16th
September, 1871, and two other kinds of parasitic flies on the 4th October following.

The action of the Canada balsam has destroyed the colours of the insects; so that the
following description is confined to structural characters; moreover the male insect
has unfortunately been fixed by the Canada balsam on its side, and the exceedingly
minute size of the creatures rendered any attempt at displaying them, by mee
the limbs in the usual manner, ineffectual.

The head in the male is of large size and of an oval form (seen laterally), transverse
in the female and widest behind; in this sex it appears to be furnished with two
large appendages, truncate at the tips, which may possibly be dilated palpi. The
antennee of the male are long and filiform, 10-jointed, the basal joint being the largest,
the remaining nine being nearly equal in size. The antenne of the female are 8-jointed,
the first joint large, the second smaller, the third considerably shorter and thinner than
the preceding, the fourth to the seventh gradually but slightly thickened, and the eighth
forming an elongated oval mass. ‘The details of the thoracic segments are not easily
determined, owing to the mode of preservation of the specimens; but the scutellum
seems to be of large size and semicircular. The abdomen is sessile, depressed, and
gradually pointed to the tip in the female, whilst it is more ovate in the other sex, with
the male ‘organ protruded. The wings are of equal size and shape in both sexes, the
posterior ones being as large as the anterior, which latter have a remarkable dilatation
near the base of the posterior margin, terminating in an acute notch; the remainder of
the margins of all the wings is fringed with long hairs; the legs are long, slender, and
terminated by 5-jointed tarsi with large pulvilli.

The 5-jointed tarsi, the number of joints in the antenne of the two sexes, the sessile
abdomen, and the very long narrow wings, agree with the characters of Haliday’s genus
Alaptus given by Walker in the‘ Annals of Natural History,’ vol. xviii. (1846) p. 50. Of

MINUTE HYMENOPTEROUS INSECTS. 587

this genus two or, more probably, only one species is recorded in this country, namely
A. minimus, “‘ferruginosus, antennis et pedibus pallidis ;” the supposed second species,
A. fusculus, “ Preecedente major colore obscurior antennis longioribus, vix revera species
distincta”’ (op. cit. p. 51). Another species, 4. pallidornis [? pallidicornis], is slightly de-
scribed by Foerster, found near Aix la Chapelle. It is scarcely half as long as 4. mini-
mus, with yellowish-white antenne (Hym. Stud. ii. p. 120). As these authors do not
mention the singular dilated and excised base of the fore wings, I consider the one before
us distinct, to which may be applied the name of ALAPTUS EXCISUS.

A, antennis maris corpore paullo longioribus, feminz corpori «qualibus ; alis anticis basi postice dilatatis

et subito excisis. Insecta minutissima. Long. cire. } mill.

The second insect described by Sir J. Lubbock, under the name of Prestwichia aquatica,
offers considerable difficulty as to its true position, in consequence of the loss of the
typical specimens (all six of which are described as having been females) and the different
descriptions given of the structure of the antenne. Thus, in the Latin description
(p. 140) supplied by Mr. F. Walker, we read “‘antennz graciles, subclavate, inarticulate ;
clava longi-fusiformis ;”’ and in his English description they are described as ‘‘10-jointed,
the first joint nearly as long as the flagellum; the second, third, fourth, fifth, and sixth
joints equal in thickness, successively decreasing in length; seventh fusiform; eighth,
ninth, and tenth forming an elongate-fusiform club.” In the specific description, by Sir
J. Lubbock, they are said to consist of six or seven segments, a conjectural second seg-
ment (shown by a trace of a division near the end of the basal joint) being short and
round, the third joint longer and increasing somewhat in size towards the apex, the
fourth similar in shape but smaller, the terminal part of the organ forming a club-like
mass, which appears to be composed of three segments, the last joint, however, being
very indistinctly marked. In his fig. 10, and in the upper antenna in fig. 11, the antenne
are represented as only 4-jointed, whilst in fig. 12 and in the lower antenna of fig. 11
they are figured as 5-jointed. From this diversity of description, however, we may conclude
that the first joint is considerably elongated, the antenna being geniculated at its extremity.
The real second joint (the supposed preceding joint being, I believe, an optical delu-
sion) is shown in fig. 12, followed by a shorter third joint, the remainder of the an-
tenna consisting of the clava, of which the first division alone is indicated in fig. 12,
whilst the second division, as described in the text, is more or less obsolete. As to
the intermediate third, fourth, fifth, and sixth joints mentioned in Mr. Walker’s English
description, they are as illusive as the “ antennz inarticulate” of his Latin description.
We are warranted, then, in concluding that the antennze were geniculate* at the end
of the long first joint, and that the terminal portion formed a more or less distinctly
8-articulated clava. Such characters, however, remove Prestwichia from the My-
marides, and associate it with smaller aberrant Hulophides. The structure of the
tarsi of Prestwichia also, in my opinion, removes them from the Mymarides, and
brings them, in like manner, to the Eulophides. It is true both Mr. Walker and Sir

* Tt is proper to observe that in most of the Chalcidide the antenne are inserted low down in the face, close to

the mouth, the basal joint extending upwards to the crown of the head, and the remainder of the antenna being bent
downwards at a sharp angle, and only reaching to the mouth when at rest.

588 PROF. WESTWOOD ON SOME

J. Lubbock distinctly describe the tarsi as 4-jointed, the latter adding that the fourth
segment (pl. xxii. fig. 15) is in all the legs very small. Referring to my genus Tricho-
gramma (as described in Mr. Taylor’s ‘ London and Edinb. Phil. Mag.’ 1838, p. 444), in
which the tarsi are described as 8-jointed,—a description which was adopted and
confirmed by Mr. Haliday, one of the clearest-sighted of entomologists,—Sir J. Lubbock
is “disposed to consider that the tarsus in Trichogramma is in reality 4-jointed, and
that Westwood and Haliday’s descriptions must be amended”’ (p. 141). Now, whoever
has studied these minute Hymenoptera is well aware that in many of them the
pulvillus of the tarsi is greatly enlarged, having the ungues affixed at the sides towards
the base, the tarsus itself being only 8-jointed (see, for example, the figures of the
tarsi of the Chacidideous genera published by Mr. Curtis in his ‘ British Entomology,’
especially those of Hulophus, pl. 183; Encyrtus, pl. 395; and Phagonia, pl. 427; and
T have not the least hesitation in affirming, from the inspection of Sir J. Lubbock’s
figures 10, 11, 14 & 15, that the tarsi of Prestwichia are only 3-jointed, with an enlarged
pulvillus, as more distinctly shown in fig.15. This character, therefore, also removes the
genus from the Mymarides—amongst which it is arranged by the Rev. 'T. A. Marshall in
his ‘Catalogue of British Hymenoptera Oxyura,’ between Cosmocoma (Polynema) and
Caraphractus.

The structure of the fore wing of Prestwichia is also indicative of a similar want of
relation with the Mymarides. As will be noticed by comparison of the figure of its wing
(fig. 18) with those of Polynema (fig. 4) and Anaphes (fig. 8), without dwelling on the shape
of the wing and its rounded extremity, it will be seen that the thick subcostal vein
extends half the length of the anterior margin of the wing, where, at its apical extremity,
“it turns inwards and ends abruptly ’’—a character never found in the Mymarides.
Sir John Lubbock, as above stated, has referred to my genus Trichogramma; and I
cannot but think that Prestwichia belongs to the little group of which the former genus
is the type, and which Foerster has formed into his family Trichogrammatoide (Hym.
Studien, Heft 2, p.87). In my original description of Trichogramma I described the
antenne thus :—“ breves, 6-articulate, articulo 1mo longo, 2do brevi, gracili, 3tio quam
2do majori, crassiori, 4to et 5to brevibus, 6toque maximo oblongo-ovato apice acuminato,”
accompanied by a figure both of the antenna and the 3-jointed tarsus. More recent
examination of the antenne leads me to think that the supposed second joint is only
the narrowed extremity of the long basal joint (as represented in the accompanying
Plate LX XIII. fig.12); nor am I clear that the large oval apical joint is not in reality
composed of two or three joints closely soldered together. The most remarkable cha-
racter of Lrichogramma consists of the curious mode in which the fine hairs of the
fore wings are arranged in lines, of which I am not aware of any similar instance
throughout the order Hymenoptera. Plate LX XIII. fig. 13, represents the fore wing of
Trichogramma evanescens, of which no figure has hitherto been published; and it will
be seen that Prestwichia differs from it in having the fine hairs scattered over the
wing irregularly.

In the Entomological Magazine (vol. i. p. 340, July 1833) Mr. Haliday published the
description of a new genus with trimerous tarsi, under the name of Calleptiles (giving

MINUTE HYMENOPTEROUS INSECTS. 589

as a synonym the name of Microma, printed without description in Curtis’s ‘ Guide’).
The type of this genus, C. latipennis, had been reared from subcutaneous larve in the
leaves of Aquilegia by Curtis, and is described as having “ alee hyaline radice obscuriores.”
To the description is appended an editorial note that the insect is probably identical with
my Trichogramma evanescens ; and in the ‘ Monographia Chalciditum,’ 1839, p. 18, the
last-named species reappears as belonging to my genus Péeroptrix, placed in a distinct
section (1), erroneously described as having 4-jointed tarsi (which is the distinguishing
character of Pteroptrix), with Calleptiles latipennis, Haliday, given as a synonym. In
the ‘Entomologist,’ 1840-1842, the insect again reappears, in the descriptions of Mr.
Haliday’s beautiful series of plates of Chalcididee, under the name of Trichogramma
evanescens mas ; but the figure of the insect (Entomol. Mag. pl. K. figs. 4, 4a—4d) differs
in several important respects from my insect and its details, and evidently represents
not only another species with a dark band across the fore wings, but probably a male
insect, whilst mine was most probably a female. In Mr. Haliday’s figures the number
of lines in which the hairs of the fore wings are arranged is considerably greater than in
my insect; and the antenne are evidently 7-jointed, and strongly setose, the first joint
of moderate size, the second about the length of the former and much thicker; the
third resembles the second, as does also the fourth, which, however, is larger than the
third, whilst the fifth, sixth, and seventh are evidently coalesced, together forming a
3-jointed oval mass.

A species of this group is described by Foerster in the ‘Verhandlungen’ of the
Rhineland Natural-Hist. Soc. vol. viii. p. 26, pl. 1. fig. 9, a, b,c, under the name of
Trichogramma Walkeri, which has the antennee 4-jointed or (if the last joint consists, as
the author supposes possible, of three joints soldered together) 6-jointed—the Ist of
moderate size, the 2nd and 8rd equal, subovate, and nearly as large as the basal joint,
and the terminal part of the antenneze forming an elongated oval mass, the whole
antenne being very slightly setose. In his ‘ Hymenopt. Studien,’ Heft 2, p. 89, Foerster
incorrectly alludes to this insect as having been described by him (loc. cit. supra) under
the name of Calleptiles Walkeri, and proposes, on account of the great length of the
ovipositor of the female, to form it into a separate genus Centrobia.

In Foerster’s figures of the antennz of Poropoea, those of the male are represented as
7-jointed, the first of moderate size, the 2nd small ovate, and the five following equal-
sized, twice as large as the 2nd, and shortly ovate. The female antenne are also 7-jointed,
the 3rd and 4th joints thicker than in the male, and the last three joints clearly uniting
to form a terminal oval mass. In Ratzeburg’s second species, Ophioneurus signatus, the
basal joint of the antennz is of moderate size, the 2nd as thick and nearly as large as
the first, the remainder (7 joints as figured) forming a large oval mass.

In the ‘Annals of Natural History,’ 2nd ser. vol. vii. pp. 211-213, Mr. Walker pub-
lished, from Mr. Haliday’s MSS., descriptions of several of these very minute insects
found in Ireland—namely, Myina annulipes and livens, Chetostricha (un. g.) dimidiata,
Trichogramma vitripennis, and Oligosita (n. g.) collina—accompanied by the following
note, also by Haliday.

SECOND SERIES.—ZOOLOGY, VOL. I. AT

590. PROF. WESTWOOD ON SOME

** Synopsis of the Trichogrammini.
«Tarsi trimeri: tibiz antice calcari apice inciso: antenne articulis 6, 3 extremis in clavam
coarctatis (an semper?) : abdomen subsessile: statura Apheline (Myine) fere et huic magis affines
videntur quam Hulophinis: Oligosita vero pedibus gracilibus, tarsis 2di paris elongatis, alis longe fim-

briatis Thysani speciem mentitur.
GENERUM CONSPECTUS.

* Ale antice seriatim pubescentes.

+ Vena costam sinu tantum attingens ad ortum radii. 1. Trichogramma, Westw.
tt Vena costam longius decurrens ante ortum radii. 2. Chetostricha, n. g.
** Ale vage pubescentes.

+ Ale: antice late, margine subtiliter ciliatee. 3. Brachista, nu. g.
tt Alse antice anguste longe fimbriate (plumatze). 4. Oligosita, n. g.”

Of the species of these minute insects having the hairs of the wings disposed in linear
series and 3-jointed tarsi, Foerster describes five genera, Viz. :—

1. Poropoea, Foerst. in Verhandl. nat. Vereins Rheinlande, 8th year, p. 28, pl. 1. fig. 10, de (1851)
= Ophioneurus, Ratzeburg, Ichn. der Forstinsecten, iii, p. 197 (1852), consisting of two species, which
are parasitic in the eges of Apoderus curculionoides and Rhynchites betule, which reside, in the larva state,

in rolled-up oak-leaves.
2. Trichogramma,Westw.
3. “ Chetosticha, Walker” [rectius “ Chetostricha, Haliday,” pace Foersteri].

4. Lathromeris, Foerster, and
5. Centrobia, Foerster.

But he adds to the family a section consisting of three other genera, in which the hairs
of the wings are not arranged in lines, viz. :—

6. Asynacta, Foerster ; 7. Brachista, Haliday ; and 8. Oligosita, Haliday.
And it is, I do not doubt, in this latter section that Prestwichia must be placed. ‘These
three genera are thus tabulated by Foerster :—

aa. Fligel nicht reihenweise behaart.

fe Filer siebengliederig. Asynacta.

jf. Fiibler sechsgliederig.

g- Vorderfliigel breit, am Rande mit zartem Haarsaum. Brachystica [rectius Brachista*, Hal.1.
gg- Vorderfliigel schmal, lang gewimpert. Oligosita, Hal.

The difficulty pointed out in a preceding part of this paper as to the precise character
of the antenne of Prestwichia prevents me from determining whether it does not in
reality enter into the genus Oligosita, Hal., of which the antennz are thus described :—
“ Subclavate, brown towards the tips, much more than half the length of the body, first
joint very long, second cup-shaped, third and following forming a spindle-shaped club.”

This description seems to indicate that the specimen described by Haliday was a male,
as these organs in the opposite sex would probably be differently formed; at all events,
in this uncertainty, I venture to describe and figure in this place a minute species, pro-
bably congeneric with Prestwichia (and of which I took several specimens on the 18th
of August, 1887, by sweeping grass in Richmond Park), under the name of

* Changed by Foerster as being too near to Brachistes, Wesmael.

MINUTE HYMENOPTEROUS INSECTS. 591

OLIGOSITA SUBFASCIATA. (Pl. LX XIII. figs. 14-19.)

Picea, capite et dorso thoracis fulvis, hujus parte postica picea, linea media longitudinali fulva, oculis
magnis, nigris ; antennis pedibusque fulvescentibus ; alis anticis nubila media pallide fusca notatis.
Long. corp. ? millim. LExpans. alar. 1} millim.

Habitat Richmond Park, comit. Surrey, mense Augusto capta.

The antennee (fig. 15) are about twice the length of the head, and consist of a long
basal joint, more slender in the middle, a second joint, not quite half the length of the
first, followed by a minute annular joint, the fourth joint being rather smaller than the
second, and three terminal joints forming an elongate oval setose mass.

The mandibles (fig. 16) are strong, horny, and terminated by four small obtuse teeth ;
the maxille (fig. 17) are large, and terminated by a large fleshy setose lobe, having a
very short cylindrical and apparently exarticulate palpus, with two terminal short sete ;
the lower lip (fig. 18) is nearly of equal size with the maxille, elongate-conic, rounded
at the extremity; the labial palpi very small, slender, cylindrical, and terminated by a
small seta. The tarsi (fig. 19) are 3-jointed, terminated by a large pulvillus, with two
small lateral ungues. The fore wings are dilated at the hinder extremity; they are well
clothed with fine setee, and their apical margin is fringed with very long hairs; the sub-
costal vein is free for a short distance from the base of the wing, where it unites with
the margin, and extends to the middle of the wing, terminating in a very short and
nearly triangular stigmatic branch extending into the disk. The hind wings are very
narrow, with a prominent hook before the middle of the costal edge, the margins with
_a fringe of long hairs, and a row of fine hairs on the disk.

Amongst the numerous minute Hymenoptera collected at or near Oolombo, in
Ceylon, by Mr. Staniforth Green, which he has mounted in Canada balsam and kindly
presented to me, are several Trichogramme and ailied species, of which I add descrip-
tions and figures as further illustrating the relations of Prestwichia,

Oxigosita StanrForTHi. (Pl. LX XIII. figs. 20, 21.)

Tota luteo-fulva, oculis fuscis; antennis perbrevibus, geniculatis, articulo primo magno, secundo semiovali,
tertio annuliformi, quarto parvo subrotundato, reliquis tribus clavam longe ovalem formantibus ; alis
anticis subangustis, angulo antico apicali rotundato-truncato, longe ciliatis, disco parum setigero,
ramo stigmatico distincto, apice angulato-truncato ; alis posticis angustis, utrinque longe ciliatis ;
abdomine elongato-conico ; oviductu longitudine abdominis, e basi hujus exeunte, apice serrato,
Long. corp. 0-017 une.

Habitat in graminosis in insula Taprobana (D. Staniforth Green).

The insect here described is a female, as shown by the exserted ovipositor (fig. 21),
which is finely serrated at the tip. The structure of its antennz and wings are very
clearly shown in the specimen, and lead me to infer that it cannot be generically separated
from Ol. subfasciata. The peculiar form of the fore wings, which have the hind
margin straight and produced, whilst the anterior apical angle is rounded off, gives the
wings a form quite unlike that of any other Hymenopterous insect.

At 2

592 PROF. WESTWOOD ON SOME

Oxicostta ? Nopicornis. (Pl. LXXIII. fig. 22.)

Tota fulvo-lutea, oculis obscuris, venis alarum pedibusque pallide luteo-albis ; capite lato; antennis
brevibus, geniculatis, articulo primo elongato, secundo brevi obconico, 4: proximis semiovalibus,
fere eequalibus, singulo secundo parum majore, setis longis antice subverticillatis, articulo apicali
(septimo) ovali, quam praecedens paulo minore; alis anticis magnitudine ordinaria, angulo antico apicali
rotundato-truncato, longe ciliatis, disco parum setoso, setis in lineas 5 irregulariter dispositis ; alis
posticis angustis utrinque longe setosis, angulo hamifero costali conico; genitalibus masculinis
porrectis quasi biarticulatis, parte basali subtus spma armata. An mas Ol. Staniforthi? long.
corp. 0014 une.

Habitat cum precedente in Taprobana (D. Staniforth Green).

Iam obliged to add a mark of interrogation to the generic name applied above to this
insect, from ignorance of the true structure of the male antenne of the type of Oligosita.
The specimen before me is a male, as shown both by the exserted genitalia and the
structure of the subverticillated antennee (fig. 23), with the terminal joints not forming a
triarticulated mass.

TRICHOGRAMMA (APROBOSCA) EROSICORNIS. (Pl. LXXIII. fig. 24.)

Corpus breve, contractum; abdomine sessili; capite magno ; antennis brevibus, geniculatis, articulo primo
longo, secundo paryo semiovali, tertio minimo annuliformi, ultimo (quarto) elongato, valde irre-
gulari, quasi eroso, subuodoso, nodulis verticillatis; mandibulis angustis, curvatis, apice bidentatis ;
maxillis cum labio massam ovalem apice breviter setosam formantibus palpisque brevibus exarticulatis
instructam; pedibus sat longis, tarsis 3-articulatis, pulvillo magno ; alis anticis latis, ciliis medioeriter
elongatis marginatis, ramo stigmatico elongato, apice crassso inflexo, setis disci in lineas circiter 10
dispositis ; alis posticis-angustis, fere linearibus, longius ciliatis, angulo hamifero costali prominente. |
Tota fulva, oculis obscuris, pedibus antennisque pallidioribus. Long. corp. 0:018 unc.

Habitat in insula Taprobana (D. Staniforth Green).

The form of the fore wings, which are relatively broad, and the linear arrangement of
the fine hairs on their disk, prove that this curious insect must be closely allied to our
English Trichogramma ; but the singular antennee (fig. 25), of which the terminal nodose
portion under my strongest power shows no trace of transverse articulation (although
the groups of verticillate hairs might be supposed to indicate so many joints which
have become ankylosed), render it necessary to form at least a separate subgenus for
its reception. In favour of the exarticulation of this terminal part, the genus Thysanus,
described by Haliday *, may be quoted, in which the terminal joint is exarticulated,
elongated, and cylindrical; it has been placed in the section Tetrastichoidee by Foerster.
The antennze appear, at first sight, to consist of only three joints, of which the last is
very long and linear, especially in the males; but between the second and third joints
there appear to be three very minute annuli (more distinctly shown in the figure of the
female than in the male antenne f). The tarsi are not described, except that the inter-
mediate pair are said to be “longissimi” (very long); the detached figure of one of the
tarsi represents it as five-jointed, with a scarcely visible pulvillus; but Foerster states

* Annals Nat. Hist. iy. 234, December 1839. This name has been unnecessarily (as it seems to me) altered by
Foerster to Triphasius (Hymen. Studien, p. 84).
+ ‘ Entomologist, No. 26, December 1842, plate K. fig. 3a male, fig. 36 female antenna; fig. 3c tarsus.

MINUTE HYMENOPTEROUS INSECTS. 593

that they consist only of four joints (J. c.), and hence the genus is placed by that author in
the section Tetrastichoide. The wings, however, are very strongly ciliated ; and notwith-
standing the difference in the number of joints of the tarsi, I should be inclined to place
the genus near Pteroptriz, Westw. (unnecessarily altered to Pterothrix by Foerster), and
not far removed from the Trichogrammatini.

EXPLANATION OF PLATE LXXIII.
The whole of these figures are highly magnified,

Fig. 1. Antenna of the male of Valkerella (Polynema) natans.
. Tarsus of the fore leg of the same species.
. Mymar pulchellus, male.

. Mymar taprobanicus, male.

1

2

3

4

5. Tibia and tarsus of the same insect.

6. Mymar taprobanicus, female.

7. Hooklets of wings of this species, necessarily represented much too coarse.

8. Mymar Wollastonii, male.

9. Mymar Wollastonii, female.

10. Alaptus excisus, male.

11. Alaptus excisus, female.

12. Antenna of Trichogramma evanescens.

13. Fore wing of the same species.

14. Oligosita subfasciata.

15 to 19. Parts of the same insect : fig. 15, antenna ; fig. 16, the mandible; fig. 17, maxilla;
fig. 18, labium ; and fig. 19, the tarsus.

20. Oligosita Staniforthii, female.

21. Extremity of ovipositor of the same insect.

22. Oligosita? nodicornis, male.

23. Antenna of O. nodicornis 3.

24. Trichogramma (Aprobosca) erosicornis.

25. Antenna of the same species.

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JOWestwood del. Trans. Linn Soc, See. 2 Vou.1Pu.73.

i

if

SPECIES OF MYMAR ALAPTUS, OLIGOSITA, & TRICHOGRAMMA.
Berjeau bth. With detacts all highly magnified. Hanhart imp.

7
a
.
—"
y
‘
hee
.

Abisara dumajanti, 566; haquinus,
546, 566; kausambi, 545, 566;
savitri, 545, 566; tanita, Hewit-
son, 546, 566.

Abranchiata, remarks on, 211.

Acanthozoon, n. g., Coll., 86.

armatum, el. (Pl. XVIIL.),
87, 95; papilio, 87, 95.

Accipiter nisus, skull of, 135.

virgatus, 313, 350.

Accipitres of Philippines, 313.

Accipitrine, 313.

Acidalia exclusa, 562.

Acidaliidee, 562.

Acrea thalia, 193.

Acridotheres cristatellus, 353.

Acrocephalus orientalis, 352.

Actinia abyssicola, sp. n. (Pl. XLY.),
297; gelatinosa, sp. n., 298.

Actiniade, 295.

Actiniaria, general remarks on, 303.

, on new forms of, dredged in

the deep sea, 295.

, Pelagic, surface-swimming
species of, 295.

Actinine, 297.

Adolias adonia, 539, 564; asoka,
539, 565; garuda, 539, 564;
jama, 539, 564; laverna, 539,
565; Iudekingii, 539, 565; mo-
nina, 539, 565; ramada, 565.

Agiale, 204.

Kollari (Pl. XXIX.), 205.

ABgialitis dubia, 353; Geoffroyi,
353 ; mongolica, 353.

Aigithognathe, facial morphology of,
107.

, palate, varieties of, 111.

A®gocera amalthea, 202.

INDEX.

Athopyga magnifica, Sharpe, 342,
353; Shelleyi, Sharpe, 342, 353.

Aetomorphee, structure skull of, 128.

Agama colonorum, sacral plexus of,
(Pl. LXVI.), 517.

Agarista, position of, 157.

agrius, 204; alethe, 203;
Batesii, 203; Doleschallii, 203 ;
eriopis, 204; Lindigii, 202; Moorei,
203; pales, 204; pedasus, 204;
Semperi, 203; simyra, 204.

Agathia discriminata, 562.

Aglaope infausta, venation wings of,
160.

Alaptus excisus, Westw., n. sp. (Pl.
LXXIII.), 586, 587; fusculus, 587;
minimus, 587; pallidornis, 587.

Alcedininze wanting vomer, 127.

Alcedinide, 317.

Alcedo bengalensis, 317, 351.

Alcyone cyanipectus, 351.

Alligator, sacral nerves of, 523.

Allman, Prof., on the structure and
systematic position of Stephanoscy-
phus mirabilis, the type of a new
order of Hydrozoa (Pl. XIV.), 61.

Amage auricula, 508.

Amathusia phidippus, 538, 564.

Amaurornis olivacea, 354.

Amblypodia adatha, 548, 566; am-
phimuta, 548, 566; anthilus, 548,
566; antimuta, 548, 566; apida-
nus, 549, 566; aroa, 548, 566;
atosia, 548, 566; centaurus, 548,
566; Diardi, 549, 566; epimuta,
548, 566; eumolphias, 548, 566;
hypomuta, 549, 566; inornata,
566 ; metamuta, 548, 566 ; vibara,
566.

Amesia juvenis, n. sp., Butler, 559 ;
pexifascia, 559.

Ammotrypane arctica, M‘Intosh, n.
sp. (Pl. LXV.), 505; aulogaster,
Rathke, 504.

Amphiporus lactifloreus, 73 ; hastatus,
73; spectabilis, 77.

Amphitrite cirrata, 508.

Anzeretes parulus, yomerine cartilages
of, 110.

Anas luzonica, 354. :

Ancistrosyllis greenlandica, M‘Intosh,
n. sp. (Pl. LXY.), 502.

Anderson, Prof. J., on the osteology
and pterylosis of the Spoon-billed
Sandpiper (Eurynorhynchus pyg-
meus, Linn.), (Pl. XXXY.), 213.

Anisopoda, members of, from Atlan-
tic, 23.

Annelida obtained during cruise of
H.MS. ‘ Valorous’ to Davis Strait
(1875), 499.

Annelids, on mode of preservation
(footnote), 499.

of Greenlandic seas, Dr. M<‘In-
tosh on, 499.

Anops malayica, 546, 566; sperthis,
546, 566.

Anous stolidus, 354.

Anthreptes chlorigaster, sp. n., 342,
353; malaccensis, 342, 353; ma-
laccensis, os uncinatum of, 111.

Anthus gustavi, 338, 352.

Apaturide, modifications apical seg-
ment of (Pl. LVI.), 360.

harpagones of, 363.

tegumen of, 362.

, values genital armature of, in

family, genera, and species, 364,366.

+}

596

Aphneeus lohita, 549, 566.

Apical segment in European Rhopa-
locera, modifications in various fa-
mnilies of, 359, 364.

Apoderus curculionoides, 590.

Appendices anales of Lepidopteron,
357; of Trichopteron, 357.

Appias cardena, 551, 567 ; eleonora,
551, 567; figulina, 551, 567; na-
thalia, 551, 567; nero, 551, 567 ;
panda, 551, 567; plana, n. sp.,
Butler, 551, 567.

Apseudes cxca, Willemoes-Suhm, 24.

Aquiline, 313.

Arachnothera dilutior, 341, 352.

Arborophila, sp. ? 353.

Arcania granulosa, sp. n. (Pl.
XXXVIIL), 240.

Archzostomata compared with Deu-
terostomata, 209.

Archaster of W. Indies, 50.

Arctiidee, 557.

Arcturus of W. Indies, 50.

Ardea purpurea, 354.

Ardeide, 349.

Ardetta cinnamomea, 354 ; flavicollis,
354; sinensis, 354.

Argina cribraria, 557.

Argynnis, 159.

Aricia Cuvieri, 504; foetida, 504;
ercenlandica, M‘Intosh, n. sp. (Pl.
LXV.), 504; Kupfferi, 504; La-
treillii, 504. ;

Armature, male genital, of the Euro-
pean Rhopalocera, 357.

Artamide, 323.

Artamus leucorhynus, 323, 351.

Ascidians, on new forms of deep-sea,
287.

Askonema setubalense, spicules of,227.

Aspidogaster, comparison of 'Trema-
tode with, 210:

Astacus, deep-sea blind species, 48.

fluviatilis, 46 ; pellucidus, 49 ;

zaleucus, n. sp. (PI. X.), 49.

, anatomy of, 48.

, characters of, 49.

, colour and mode of life,

50.
, compared with the mam-
moth cave species, 49.
Astictopterus armatus, 555, 568 ;
diocles, 555, 565; gemmifer, n. sp.,
Butler, 555, 668; jama, 555, 568;

INDEX.

sindu, 555, 568; stellifer, n. sp.,
555, 568; zanites (Pl. LXIX.),
55 5,568.

Astrogonium of W. Indies, 50.

Astur soloénsis, 350; trivirgatus,
350.

Asynactia, 590.

Atella phalanta, 544, 565; sinha,
544, 565.

Athis japyx, 178.

Athyma amhaia, 540, 565; clerica,
n. sp., Butler (Pl. LXIX.), 540,
565; idita, 541, 565; kresna,
540, 565; laryma, 540, 565; leu-
cothoé, 540, 565; nivifera, n. sp.,
Butler (Pl. LXIX.), 540, 565;
pravara, 541, 565; sabrata, 540,
565; urvasi, 560.

Atlantic Crustacea from ‘ Challenger ’
Expedition, 23.

Attacus atlas, 561.

Atya, 27.

Auditory ossicula(Pls.LVIII.-LXIY.).

—— of Mluride, 392, 398;

of Anthropoidea as compared with

man, 379; of Antilocapride, 424;

of Antilopidea, 427; of Artiodac-

tyla, 421; of Baleenoidea, 456,464 ;

of birds, 490; of Bovide, 425,

432; of Camelide, 424, 431; of

Canide, 394, 395; of Carni-

vora, 390; of Castoride, 410; of

Cebide, 383; of Centetidee, 440,

444; of Cervide, 429, 482; of

Cetacea, 450-464; of Chevrotains,

423; of Chimpanzee, 378; of

Chinchillide, 416 ; of Chiromyide,

388, 390; of Chiroptera, 445—

449; comparison of monkeys and

lemurs, 389; of Cryptoproctide,

392, 394 ; of Cynomorpha or tailed _

Old-World monkeys, 380-383; of
Dasyproctide, 416; of Dasyu-
ride, 483, 487; of Delphinide,
459, 464; of Didelphia, 477-487 ;
of Didelphyide, 480, 487; of Dipo-
didwe, 414; of Duplicidentati, 419;
of Echidna, 488, 490 ; of Kdentata,
470-477; of Emballonuride, 449 ;
of Entomophaga, 472; of Equide,
420,431; of Erinaceide, 438, 444 ;
of Felide, 391 ; of Fissipedia, 390 ;
of Fossorial Rodents, 418 ; of Galeo-
pithecus, 442, 443 ; of Galaginide,

387, 389; of Giraffe, 429, 432;
of Gorilla, 377; of Hapalide or
Marmosets, 385; of Hyznide,
391, 392; of Hylobates, 379; of
Hyraces, 432; of Hystricide, 414;
of Hystricomorpha, 418; of Indri-
sine, 386; of Insectivora, 432—
444; of Lemuride, 386; of Lepo- .
ride, 417; of Loricata, 472, 476 ;
of Macropodide, 478, 487 ; of Ma-
eroscelids, 440,444; of man, 372;
of Manide, 474; of Marsupialia,
477-487 ; of Monotremata, 487—
492; of Muride, 410; of Muste-
lide, 395, 396 ; of Myogalide, 434,
444 ; of Myomorpha, 418; of My-
oxide, 409; of Nycticebide, 387,
389; of Octodontide, 415; of
Orang-utan, 378 ; of Ornithorhyn-
chus, 491, 493; of Otariide, 400,
407 ; of Peramelide, 486 ; of Peris-
sodactyla, 419 ; of Phalangistidiv,
483, 487 ; of Phascolomyidee, 485,
487; of Phocide, 404, 407; of Phyl-
lostomide, 449; of Physeteride,
456,463; of Phytophaga, 470; of
Pinnipedia, 399-407 ; of Platanis-
tide, 462, 464; of Proboscidea, 430;
of Procyonide, 392, 397; of Rhi-
nocerotidie, 419 ; of Rhinolophide,
446; of Rodentia, 407-419; of
of Saccomyide, 413; of Sciuride,
407, 408 ; of Sciuromorpha, 418 ;
of Simiide, 377; of Sirenia, 464—
470; of Soricide, 433, 444; of
Spalacide, 412; of Suide, 422,
431; of Talpide, 436, 444; of
Tapiride, 420; of Tragulide, 423,
431; of Trichechide, 401, 407;
of Tupaiide, 441, 444; of Ungu-
lata, 419-431; of Urside, 398,
399; of Vespertilionide, 448; of
Viverride, 392; of Ziphiide, 457,
464.
Axiothea catenata, 507.

Balabae, birds of, 310.

Basilan, birds of, 310.

Baza magnirostris, 350.

Belenois cynis, 551, 567.

Bendidex, 561.

Bermudas, on a Nebalia from, 26.

Birds’ palate, Aigithognathous varie-
ties of, 111.

Birds, sacral nerves of, 528; skull,
structure and development of, 99.

Birnara, n. g., nubila, Butler, 560.

Blind deep-sea Astacus, 48; Deca-
pods, 50; Tanaid, 23.

Blyth, Mr., on Sus cristatus, 255.

Botidide, 563.

Botys pterophoralis, 563; sp. ?, 563.

Brachista, 590.

Brachypodide, 334.

Brachysticta (syn.), 590.

Brachyurus erythrogaster, 308, 352 ;
propinquus, sp. n., 330, 352 ; sor-
didus, 309, 331, 352; Steerii, sp. n.
(Pl. XLIX.), 329.

Branchiz in Cidaride, 569; Echi-
noidea, 569.

Brown Linnet, facial arches of, 108.

Bryozoa of W. Indies, 50.

Bubo philippensis, 350.

Bubulcus coromandus, 349, 354.

Buceros hydrocorax, 322, 351.

Bucerotide of Philippines, 322;
structure face of, 126.

Buchanga cineracea, 324, 351.

Budytes viridis, 352. ©

Buntings, pterygoids of, 110.

Burgena transducta, 203.

Butalis manillensis, 326, 351.

Butaster indicus, 313, 350.

Butler, A. G., on the butterflies of
Malacca, 533.

Butorides javanica, 349, 354.

Butterflies of Malacca, 533.

Buzzards, structure face of, 136.

Cacatua hematuropygia, 312, 350.

Cacomantis merulinus, 320, 351.

Calappa hepatica (ftnote), 238.

Calleptiles latipennis, 559 ; Walkeri,
589.

Callidula abisara, n. sp., Butler, 558.

Calliope camschatkensis, 352.

Callipleea diocletianus, 535, 564; Le-
dereri, 536, 564; vestigiata, 535,
564.

Calobates melanops, 352.

Calcenas nicobarica, 308, 353.

Calornis chalybeus, 343, 353; pana-
yensis, 343, 353.

Camiguin, birds of, 309.

Campephagide, 323.

Cancer lunaris, 244; plicatus, 240;
victor, 243.

INDEX.

Canis familiaris, 254; lupus, 254.
Caphyra rotundifrons (ftnote), 238.
Capitella capitata, 507.

Caprimulgus europzeus, structure face
of, 113; double vomer of, 125.
griseatus, 351; manillensis, 351.

Caraphractus, 588.

Carcinas meenas, zoéa of, 47.

Cardinalide, pterygoids of, 110.

Cardisoma guanhumi, 46; sp. (San
Jago), develop. of, 46.

Cariama, structure face of, 129.

Carinella annulata, 78, linearis, 73.

Carpophaga senea, 346, 353 ; Picker-
ingii, 311, 353.

Carter, H. J., remarks on Dendro-
spongia Steeru, 231.

Castalius elna, 566; ethion, 547,
566; rosimon, 546, 566; roxus,
547, 566.

Castnia, charac. by Boisduval, 166 ; of
Westwood, 167.

, Monograph of, 155; acreoi-

des (Pl. XXIX.), 193; actino-

phorus, 193; actor, 169; albofas-

ciata, 184, 185; amycus?, 181,

182; ardalus, 171; argus, 179;

atymnius, 172; Besckei, 174; Bois-

duvalii, Becker, MS., 173; Bre-

courti, 171; cacica (Pl. XX VIII.),

169; chelone, 180; chremes (Pl.

XXIX.), 185; ciela, Botsduval,

MS.,179 ; clitarcha (Pl. XXXT.),

176; cochrus (Pl. XXYVIII.),

177; columbina, 190; cononia

(PI. XXXIT.), 188; cratina, 188;

cronida, 183 ; cronis, 183; cycna,

189, 191; Dalmannu (Pl. XXX.),

171, 179 ; daedalus, 167; decus-

sata, 175; dionea, 177; diva

(Pl. XXIX.), 187; ecuadoria (PI.

XXXII.), 189; eudesmia (Pl.

XXVIII.), 177; euphrosyne, 181;

evalthe, 181; Fabricii, 178, 185;

Fonscolombei, 178; futilis, 173 ;

galinthias, 172; geron, 168; Go-

dartii, 169, 175; Grayii, 171;

harmodius, 172; hecate, Botsduval,

MS., 174; hegemon, 171, 178,

179; Herrickii, Boisduval, MS.,

174; hesperiaris, 204; Hubner

(Pl. XXVITI.), 182; Humboldti,

180; icarus, 170; var. invaria,

170; inca, Boisduval, MS., 174;

SECOND SERIES.—ZOOLOGY, VOL. I.

597

inornata, 184; imvaria, 170;
japyx, 178 ; juturna (Pl. XXIX.),
182, 183; Kirstenii, 178; Langs-
dorfi, 170; Latreillii (PI.
XXVIII.), 161, 168; licas, 172,
173; licoides, 184; linus (Pl.
XXIX.), 192; lorze, Boisdu-
val, MS., 183; Marcelserresii
(Pl. XXIX.), 186; maris, 178;
meditrina, 182; Menetriesi, 180;
mimica (Pl. XXXIT.), 191; mor-
phoides, 185, 186 ; musarum, Bois-
duval, MS., 177; mygdon (Pl.
XXVIII.), 179, 180 ; orestes, 174 ;
palatina, 176; Pallasia, 171; pa-
pagaya (Pl, XXX.), 170; papili-
onaris (Pl. XXXI.), 175; para-
doxa, 163; paradoxa, Bozscduval,
MS., 176; penelope, 184; per-
sonata, 183; phalaris (Pl. XXX.),
177, 180; procra, 169; pylades,
169; rutila, 178; salasia, 181;
salvina (Pl. XXXIT.), 190; sa-
trapes (Pl. XXXI.), 175, 176;
Schreibersii, 168; simulans, 187 ;
Spixu, 172 ; Sternbergii, 182 ; stri-
gata, 174, 175; subvaria, 177;
thais, 185; thalaira, 185; the-
rapon (Pl. XXTX.), 176; tricolor,
187; truxilla, 162, 190 ; umbrata,
174; unifasciata, 194; veraguana
(Pls. XXVIII. & XXX.), 168;
Viryi, 181; zagrea, 189; zeryn-
thia, 170.

Castniz and allied genera, 164; ar-
rangement veins of hind wings,
162, 163.

as related to the Heliconiide,

158.

, Classifications of, 165, 166 ;
geographical range of, 164; Ma-
dame Merians and others, obser-
vations on transformations of, 164;
palpi of, 164; remarks on venation
wings of, 158, 161, 162, 163, 164;
transformations of, 164.

Casyapa irava, 553, 567 ; thrax, 553,
567.

Catapecilma, n. gen., 547; elegans,
548, 566.

Catopsilia catilla, 551, 567; chryseis,
561, 567; crocale, 550, 567;
scylla, 551, 567.

Cecropis daurica, 351.

AK

598

Celeomorphe (Woodpeckers), rank as
a zoological group, 20; structure,
face of, 2.

Centrobia, 590.

Centrococeyx viridis, 322, 351.

Centropine, 322.

Centrostomum ocellatum (Pl. XIX.),
88, 97; punctatum, 88, 97.

Ceratiocaris, 28.

Ceretes chremes, 185.

charac. by Boisduval, 166.

Cerianthide, 302.

Cerianthus bathymetricus, sp. n. (Pl.
XLY.), 302; membranaceus, 303.

Cethosia, 157; hypsina, 543, 565;
methypsea, n. sp., Butler, 543.

Ceyx melanura, 317, 351 ; philippen-
sis, 851; tridactyla, 351.

Cheetostricha, 590; dimidiata, 589.

Chalaraspide, 45.

Chalaraspis, n. gen., type of new
schizopod family (Pl. VIIL.), 37;
anatomy of, 38; characters, 39;
range of, 37; systematic position
of, 39.

unguiculata, n. sp., 40; mode
of life and colour, 40.

Chalcophaps indica, 348, 353.

Chalcosia coliadoides, var. latifasciata,
559.

Chalcosiide, 558.

Challenger Exped. Actinaria, 295 ;
Atlantic Crustacea of, 23; Deep-sea
Ascidians, 287.

Chameleo vulgaris, sacral nerves of
(Pl. LXVI.), 514.

Charadriide, 349.

Charadrius fulyus, 349, 353.

pluvialis, skull compared with
Eurynorhynchus, 214.

Charaxes hebe, 539, 564; jalysus,
539, 564; Schreiberi, 539, 564.

Chauliodus, W. Indian fish, 50.

Chelidon urbica, stylo-hyal arches of,
106; postoral arches of, 105.

Chelonians, comparisons palate of,
with Birds, 130.

Cheluria terebrans, 67.

Chick’s Skull, researches on, 101.

Chone Duneri, 509; infunduliformis
(Pl. LXY.), 508.

Chrysococeyx basalis, 320, 351.

Chrysocolaptes erythrocephalus, sp. n.
(Pl. XLVI.), 315, 350; maculi-

INDEX.

ceps, sp. n., 314, 350; xanthoce-
phalus, 314, 350.

Chrysocolaptes hematribon, 350; lu-
cidus, 350.

Cicéntide, 349.

Cidaride, on certain organs of, 569.
Cidaris nutrix, 571; tribuloides (Pl.
LXX.), 571; of W. Indies, 50.
Cinnyris speratus, 341, 353; jugu-

laris, 341, 353.

Circus melanoleucus, 350; erugi-
nosus, 308; spilonotus, 308, 350.

Cirratulus cirratus, 507. ;

Cirrochroa bajadeta, 543, 565; jo-
hannes, 543, 565; orissa, 543,
565; rotundata, n. sp., Butler,
543, 565.

Cistenides granulata, 507; hyper-
borea, 507.

Cisticola, sp.?, 336, 352; semirufa,
352,

Cittocinela Iuzoniensis, 352; nigra,
sp. n. (Pl. LIL.), 335, 352.

Clarke, R., and Mivart, St. G., on
sacral plexus and sacral vertebra,
Lizards and Vertebrata (Pl. LXYVI.
& LYII.), 513.

Cleosiris catamita, 558.

Clerome faunula, 564; gracilis, 564.

Cobalus elia, 554, 567; ciliatus, n.sp.,
Butler, 554, 567.

Coccothraustes, pterygoids of, 110.

Coceygomorphe, face of, 126.

Coelites humilis, Butler, 564.

Collingwood, Dr., on marine Plana-
rians, collected partly by Dr.
Kelaart at Trincomalee, and partly
by Dr. Collingwood in Eastern
Seas (Pls. XVII. to XTX.), 83.

Collocalia troglodytes, 320, 351.

Columbee of Philippines, 346.

Columbide, 347.

Comparison of Trematoda and Hiru-
dinei, 209.

Copsychus mindanensis, 352.

Coraciide, 316.

Corallimorphus, gen. noy., 299 ; pro-
fundus, sp. n. (Pl. XLY.), 300;
rigidus, sp. n., 301.

Corone philippina, 343, 353.

Corvide, 343.

Corvus monedula, stapedials of, 107.

Corybantes nicon, 185.

Corydalla lugubris, 337, 352.

Cosmocoma, 588; =Polynema, 583.

Cossus, 158, 164.

Cotuza drepanoides, 562.

Craniorrhinus leucocephalus, 309,
322, 351; Waldeni, 322, 351.

Crateropus caudatus, 352.

Criniger frater, sp. n., 834, 352.

Crossurus vittatus, 67.

Crustacea, notes on Oxystomatous,
235; on some Atlantic, 23; tabu-
lar view, how often got in ‘ Chal-
lenger’ Exped., 56.

Cryptobranchus, sacral nerves of,
524,

Cryptocnemus Grandidieri, 241;
Holdsworthi, sp.n.(Pl. XX XYIIL.),
241; pentagonus, 241.

Cuculide, 320; on structure and de-
velop. of face in, 112.

Cujo, birds of, 310.

Cyclopides salsala, 555, 568.

Cyclopspitta lunulata, 312, 350.

Cyclosia panthona, 559.

Cynthia erotella, n. sp., Butler, 544,
565.

Cyornis philippensis, sp. n., 325, 351.

Cypselidee, 319.

Cypselus apus, interorbital fenestrze
of, 104.

Cyrestis rahria, 544, 565; nivea,
545, 565.

Cystisoma Neptunus, 24.

Cystisomidz, a new family of Crus-
tacea, 25.

Dactylethra, sacral nerves of, 525.

Damalis egens, 558.

Damias dichroa, 203; elegans, 203 ;
melaxantha, 203 ; varia, 203.

Danaidx, modifications apical seg-
ment of, 360.

Danaine, 534.

Danais crocea, 536, 564; grammica,
536, 564; melaneus, 536, 564;
melanippus, 536, 564; plexippus,
586, 564; septentrionis, 536, 564;
vulgaris, 536, 564.

Dasylophus superciliosus, 321, 351.

Deep-sea Ascidians, 287.

Deidamia crucifera= Willemoesia cru-
cifera, 50; leptodactyla, 49, 50.
Deidamia, n. gen. of Crabs, but a

synonym, 50.
Delias dione, 550, 567; metarete,

n. sp. Butler, 550, 567; ninus,
567.

Dendrobiastes basilanica, sp. n. (PI.
LITI.), 332, 352.

Dendrocygna vagans, 454.

Dendrophila frontalis, 338, 352 ;
cenochlamys, sp. n. (Pl. LIII.),
338, 352.

Dendrospongia Steerii, Muriec, gen. &
sp.n.(Pls. XXXVI. & XXXYVILI.),
228, acerate spined spicula of, 226 ;
affinities of, 229; sections of, 221 ;
history of, 219; microscopic struc-
ture of, 224; Mr. Carter’s remarks
on, 232; scopuline spicula of, 226 ;
sexradiate dermal spicula of, 227 ;
spicular rosettes of, 227; vitreous
fibre of, 225.

Dentalium of W. Indies, 50.

Descriptions of some minute Hyme-
nopterous insects, by Prof. J. O.
Westwood (Pl. LX XIII), 583.

Desmognathous palate, varieties of,
112.

Deudorix domitia, 549, 566; jarbas,
549, 566; petosiris, 549, 566;
pheretima, 549, 566.

Deuterostomata compared with Ar-
cheostomata, 209.

Development face of Yungidx, 17.

—— of Land Crabs (PI. XI.), 46.

Diadematidx, 569.

Diceum dorsale, 340, 352; hema-
tostictum, 339, 352; hypoleucum,
339, 352; papuense, 352; retro-
cinctum, 352; trigonostigma, 340,
352.

Dicholophus cristatus, structure skull
of, 128.

Dicruride, 324.

Dicrurus balicassius, 351; mirabilis,
324, 351.

Digonopora (or moncecious Turbel-
laria) subtribe, 86, 87.

Diprotodont Marsupial Hypsiprym-
nodon, Prof. Owen on, 573.

Discophora menetho, 538, 564 ; tullia,
538, 564.

Dithyrocaris, 28.

Ditrypa arietina, 509; greenlandica,
M‘Intosh, n. sp., 509.

Doleschallia pratipa, 539, 564.

Domestic Swine, Prof. Rolleston’s
views on, 253.

INDEX.

Doran, Alban H. G., on morphology
of mammalian ossicula auditus
(Pls. LVIII.-LXIY.), 371.

Dorocidaris papillata, branchie of,
569; compasses, muscles, and liga-
ments of, 570; genital gland of,
571.

Drepanophorus, 77, 79.

Dryococeyx, Sharpe, n. g., 321.

Harringtoni, sp. n., 321, 351.

Dysporus piscator, 354; sula, 354.

Eagles, on palatine bones of, 137.

Kar-bones, small, of Mammals, A. H.
G. Doran on, 371.

Echinide, 569.

Echinoidea, branchiz of, 569.

Kchinometridze, 569.

Kechinus of W. Indies, 50.

Edwardsia coriacea, sp. n. (Pl. XLY.)
299.

Elanus hypoleucus, 350.

Elasmodes obtusus, n. sp., 88, 93.
Elymnias lutescens, 564; mehida,
538, 564; nigrescens, 537, 564.

Elymniine, 537.

Encyrtus, 588.

Epinephele Hyperantus, as type geni-
tal armature (Pl. LY.), 359.

Ergolis ariadne, 545, 565; merione,
545, 565.

Erycinide, 545,

—— modifications apical segment of
(Pl. LYI.), 359.

, harpagones of, 363.

, tegumen of, 361.

, values genital armature of, in
fam., gen., and species, 364, 365.

Eryon arctiformis, 55.

Eryontide, gen. of fossil crustaceans,
to which Willemeesia, n. gen., deep-
sea, is probably allied, 50.

Erythrea pheenicura, 353.

Hsacus magnirostris, 353.

Estheria, 27.

Kteone arctica, 502;
pusilla, 502.

Euchone analis, 508.

Kudamus? yucce not a Castnia, but
a Hesperian, 205.

Kudamus (Megathymus) yucce, vena-
tion wings of, 160.

Eudrepanis pulcherrima, 341, 353.

Eudynamis mindanensis, 320, 351.

flava, 502;

599

Eulabes javanensis, 353.

Eulabetine, 344.

Kulacura osteria, 541, 565.

Eulalia problema, 502; viridis, 502.

Eulophus, 588.

Eumyias panayensis, sp. n., 326,
351.

Eunoa Crstedi, 500.

Kupalamides ctesiphon, 169. |

Euphausia simplex, 45,

Kuphausiide, 45.

Euphrosyne armadillo, Sars, 500;
borealis, 500 ; foliosa, 500.

Eupolynoé occidentalis, M‘Zntosh,
n. sp. (PI. LXY.), 501.

Kuplectella cucumer, 232.

Kuplea, 157; Bremeri, 535, 564;
chloé, 335, 564 ; Grotei, 535, 564 ;
margarita, 585, 564; Ménétriési,
535, 564; midamus, 535, 564;
Ochsenheimeri, 535, 564; phos-
bus, 535, 564 ; Pinwilli, n. sp. (Pl.
LXIX.), 535, 564.

Kuripus euploeoides, 565.

European Rhopalocera, on male geni-
tal armature of, 357.

Wild Boar, 254.

Eurylemide, 344.

Eurylepta (Pls. XVII.-XIX.) affinis,
87,96; atraviridis, 87, 95; cere-
bralis, 87, 96; dulcis, 87, 96;
fusca, 87, 95; Kelaartii, 87, 92;
purpurea, 87, 96; striata, 87, 97;
undulata, 87,95; violacea, 87, 96;
viridis, 87, 96; zeylanica, 87, 97.

Euryleptide, 87.

Kurynorhynchus pygmeus, 213; com-
parison skeleton of, with Tringa,
217; hyoid arch of, 216; pelvis
and limb of, 216; shoulder-girdle
of, 216; skull of, 213; vertebral
column of, 215 ; viscera of, 216.

Kurystomus orientalis, 316, 351.

Kuschema subrepleta, 559.

Kusemia pales, 204.

Eusthenelais abyssicola, M‘Intosh, n.
sp., 501; hibernica, 501.

Eyarne impar, 501.

Excalfactoria chinensis, 353.

Face, structure of, in Buzzards, 136;
in Caprimulgus europeus, 113; in
Coccygomorphe, 126 ; in Cuculine
Birds, 112; in Dicholophus cris-

4K 2

600

tatus, 128; in Falco tinnunculus,
132; in Giant Goatsucker, 124;
in Harriers, 136; in Kites, 136; in
Podargus, 124; in Scythrops novee-
hollandiz, 120; in Trochilide, 116.

Facial Arches in Passerine types, on
structure and development of, 104.

Falconidx, 313.

Falco severus, 350.

Fern-Owl, structure face of, 113.

Flabelligera affinis, 506.

Fringillide, preoral arches of, 108.

Galathea of W. Indies, 50.

Gallicrex cinerea, 353.

Gallinee of Philippines, 348.

Gallinago megala, 354; scolopacina,
B54.

Gallinula chloropus, 353.

Gallus bankiva, 348, 353.

Gastropacha repanda, 160.

Gavia ridibunda, structure skull of,
138; 1st stage, 142; 2nd stage,
145; 3rd stage 146; 4th stage,
148; 5th stage, 148; 6th stage,
150.

Gazera as charac. by Boisduval, 166.

columbina, 190; personata,
188; simulans, 187; zagrosa, 189.

Gecarcinus lateralis, 46.

Gelasimus annulipes, var. (footnote),
238 ; Gaimardi (footnote), 238.
Genital armature, male, European

Rhopalocera, 357.

Genusa bigutta, 563.

Geograpsus crinipes (footnote), 238.

Geometride, 562; wings of, 160.

Geopelia striata, 353.

Gerydus Horsfieldi, 546, 566; ni-
yalis?, 546, 566; symethus, 546,
566.

Gerygone simplex, 352.

Gigantic Hydroid, 287.

Glareola orientalis, 353.

Glycera capitata, 503; setosa, 503.

Glyphodes amethysta, n. sp., 563.

Gnathophausia, n. gen., 36; gigas,
N. sp., carapace of, 28; characters
of, 37; gnathopods, periopods, and
branchie of, 30; oral apparatus
of, 29; pleon and appendages of, 31.

gracilis, colour and mode of life,

34; systematic position of, 35;

zoéa, 0. sp., structure of, 32, 37.

INDEX.

Goatsucker, structure face of, 124.

Gorsachius melanolophus, 308, 349,
354.

Gouride, 348.

Gracula javanensis, 344.

Gralle of Philippines, 349.

Grammatophora barbata, sacral plexus
of (Pl. LXVI.), 516.

Grapsus cruentatus, 46; terrestrial
habits of, 46; pictus(footnote), 238.

Graucalus striatus, 323, 351; suma-
trenesis, 323, 351.

Green Woodpecker, skull of, 4.

Guimaras, Birds of, 309.

Gull, on structure skull of, 138.

Gymnorhina, palate of, 111 ; tibicen,
stylohyal of, 107.

Halcyon chloris, 319, 351; coro-
mandi, 351; gularis, 317, 351;
hombroni, 318, 351; Lindsayi,
351; pileata, 351; Winchelli, sp. n.
(Pl. XLVIL.), 318, 351.

Haliaetus leucogaster, 350.

Harmothoé imbricata, 500.

Harpactes ardens, 316, 350.

Harpagones, European Rhopalocera,
structure of, 362.

Harriers, structure face of, 136.

Hebomoia glaucippe, 551, 567.

Hecatesia, gen. as characterized and
defined by Westwood, 199; exultans

. (Pl. XXXIIT.), 200; fenestrata,
200; thyridion, 200.

Heliconiide as related to the Cast-
nie, 158.

Helotarsus ecaudatus, mesopterygoid
of (Pl. XXY.). 130.

Hemiphaga poliocephala, 347, 353.

Hepiali, 158.

Hepialus humuli (Pl. XXIX.), 163.

Herodias garzetta, 354; intermedia,
354.

Hespagarista, 203; interjecta, 203.

Hesperia badra, 554, 567; harisa,
554, 567; malayana, 567 ; sophia,
White, 194; vitta, 554, 567.

Hesperide, 553.

» modifications apical seements

of (Pl. LVII.), 360.

’ harpageones of, 363.

—, tegumen of, 362.

, value genital armature of, in

fam., gen., and species, 364, 366.

Hesperide, venation wings of, 160.

Hestia linteata, n. sp. (Pl. LXIX.),
531, 564; lynceus, 536, 564.

Heterocera, 557.

Heterocerous Lepidoptera, venation
wings of, 160.

Hexactinellid group of Spongida,
new genus of, 219.

Hierococcyx pectoralis, 351; stre-
nuus, 351.

Himantopus autumnalis, 353; leu-
cocephalus, 353.

Hind foot, Prof. Owen on its deyelop-
ment in yarious groups, 581.

Hirudinei, on some new or little-
known parasitic, 209.

Hirundinide of Philippines, 328.

Hirundo gutturalis, 352; rustica, 328,
352.

Hornbills, structure face of, 126.

House-Martin, postoral arches of, 105.

Hulodes caranea, 561.

Humming Birds, face of, 116.

Hyalincecia tubicola, 499.

Hybleea constellata, 561.

Hybleeidee, 561.

Hydrochelidon leucopareia, 354.

Hydrophasianus chirurgus, 354.

Hydrozoa, type new order of, 61.

Hyla, sacral nerves of, 525.

Hyloterpe philippensis, 351.

Hymenopterous insects, descriptions
of some minute (Pl. LX XIII.), 583.

Hypobythius, nov. gen., 287; caly-
codes, sp. n. (Pl. XLIV.), 287.

Hypochrysops elegans, synonym, 548.

Hypolimnas bolina, 543; incom-
moda, n. sp., Butler, 543, 565.

Hypolyzna erylus, 549, 566 ; etolus,
549, 566.

Hypotenidia philippensis,
striata, 353 ; torquata, 354.

Hypothymis azurea, 325, 351 ; super-
ciliaris, sp. n., 326, 351.

Hypsa egens, 558; heliconia, 557 ;
subsimilis, 557.

Hypsine, 557.

Hypsipetes philippensis, 335, 352;
rufigularis, sp. n., 335, 352.

Hypsiprymnide, 573.

Hypsiprymnodon moschatus, bones
&e., limbs, 578,579; dentition of,
580; descrip. exterior of, 575; feet
of, generally, 576; habits of, 579;

354 ;

male scrotum and female pouch,
579 ; skull of, described and com-
pared with other marsupials, 577 ;
sole of hind foot of, 574. _

Tanthenas griseigularis, 353.
Ichneumon atomos, 583; punctum,
--583.

Icteridee, pterygoids of, 110.

Ideopsis daos, 536, 564.

Iguana tuberculata, sacral nerves of
(Pl. LXVII.), 517.

Incus of Mammalia (see under Au-
ditory ossicula), 371-497.

Indian Wild Hog, 254.

Internal ear-bones of Mammalia,

’ Mr. Doran on, 371.

Intersacral Nerve, definition of, 514.

Tolaus longinus, 549.

Tora scapularis, 333, 352.

Trena cyanogastra, 352; melanochla-
mys, sp. n. (Pl. LI.), 334, 352;
Tweeddalii, Sharpe, 333, 352.

Trenidz, 333.

Isis of W. Indies, 50.

Ixus goiavier, 352; sinensis, 352.

Jungipicus maculatus, 350.

Junonia asterie, 545, 565; eudoxia,
565; iphita, 545, 565; laomedia,
545, 565; lemonias, 545, 565;
orithya, 545, 565.

Kangaroo, sole of hind foot of, 574.

Kelaart, Dr., Marine Planarians col-
lected by, 83.

Kettelia Lowii, n. gen. & sp., 560.

Ketupa ceylonensis, mesopterygoid of
(Pl. XXY.), 131.

Kites, on structure face of, 136.

Lacerta viridis, sacral nerves of, 515.

Lacertilia, nerves of lumbar, sacral,
and precaudal region of, 514.

Lalage dominica, 324, 351.

Lambrus nodosus (ftnote), 238.

Lampides agnata, 547, 566; almora,
547,566; aluta, 547, 566; beeticus,
547 ; beroé, 547, 566; cerula, 547,
566; kankena, 547, 566; macroph-
thalma, 547, 566; patala, 547,
566; pseudelpis, n. sp., Butler (Pl.
LXVIII.), 547, 566.

TLanassa Nordenskidldi, 508.

Land-Crabs, on development of, 46.

INDEX.

Lanice conchilega, 73.

Lanius lucionensis, 323, 351; nasu-
tus, 351; schach, 351.

Laomedea flexuosa, 65.

Laphania Boecki, 508.

Laridx, 350.

Larus, sp., 354.

Lathromeris, 590.

Laughing Gull, skull of, 140.

Laurion corculum, n. sp., Butler, 559.

Lezena abranchiata, 508.

Lebadea alankara, 541, 565; martha,
541, 565.

Lepidogrammus Cumingi, 351.

Lepidonotus subleyis, 501.

Lepidopterous genus Castnia, J. O.
Westwood on, 155.

Leptochelia, 68.

Leptoplana aurantiaca, Coll., n. sp.
(Pl. XVIII.), 88, 94; patellensis,
n. sp., 88, 93.

Leptoplanide, 88.

Lethe europa, 537, 564.

Leucocerca nigritorquis, 351.

Leucophasia sinapis, venation wings
of, 160.

Leucosia (Pl. XXVIII.) brunnea,
Miers, sp. n., 237; fuscomaculata,
sp. n., 236; neocaledonica, 236;
ornata, sp. n., 236; pallida, 239;
Perryi, sp. n., 238; pubescens, sp.
n., 238; pulcherrima, sp. n., 236;
reticulata, sp. n., 237; urania, 236;
Whitmeei, Miers, 238.

Leucosiide, descriptions new species
of, 235.

Leucotreron gironieri, 353.

Libytheidee, modifications apical seg-
ment of (Pl LVi.), 359.

; harpagones of, 363.

—, tegumen of, 362.

, values genital armature of, in
family, genera, and species, 364,365.

Limenitis procris, 541, 565.

Jimnoria terebrans, 67; xylophaga,70.

Linota cannabina, facial arches of, 108.

Linotrypane apogon, 505.

Liparidee, 560.

Liphyra brassolis, 546, 566.

ithosiide, 557.

Lizards, sacral plexus and sacral ver-
tebre of, 513; nervous and osseous
structures of sacral region in, 522.

Lophogaster typicus, 27.

601

Lophogastride, characters of, 36, 45.
Loriculus chrysonotus, 313, 350;
Hartlaubi, 350; occipitalis, 350 ;
philippensis,350; regulus, 313, 350.

Loxura atymnus, 550, 567.

Lubbock, Sir J., on aquatic Hymeno-
ptera, 583.

Luidia of W. Indies, 50.

Lumbriconereis fragilis, 503.

Lumbricus tubicola, obtained by Sir
John Ross, 499.

Luzon, Birds of, 309.

Lycena cagaya, 548, 566; karsandra,
548; lysizone, 548, 566; sangra
548, 566.

Lycenide, 546.

, modifications apical segment of
(Pl. LVII.), 359.

—, harpagones of, 363.

==F, tegumen of, 361.

, values genital armature of, in
family, genera, and species, 364,365.

Lyceenine, 547.

Lycenopsis haraldus, 546, 566.

Lycorea ceres, 187.

Lyncornis macrotis, 351.

Macdonald, Dr. J. D., anatomy of
Tanais vittatus, occurring with
Limnoria and Chelura terebrans in
excavated pier-wood (PI. XV.), 67.

, on new genus of Trematoda and
some new or little-known parasitic
Hirudinei (Pl. XXXIY.), 209.

M‘Intosh, Dr. W. C., on Annelida
obtained during cruise of H.M.S.
‘Valorous’ to Davis Strait (Pl.
LXY.), 499.

, on Valencinia armandi, a
new Nemertean (Pl. XVI.), 73.

Macroglossa proxima, 557.

Macroglossinz (of Malacca), 557.

Macronus straiticeps, sp.n., 331, 352.

Macropteryx comatus, 319, 351.

Macropygia tenuirostris, 347, 353.

Malacca, Butterflies of, 533.

Malamaui, Birds of, 310.

Maldane Sarsi, 507; triceps, 507.

Male genital armature in European
Rhopalocera, 357.

Malleus of Mammals (see under Au-
ditory ossicula), 371-497.

Mammalian internal ear-bones, A. H.
Doran on, 371.

602

Margarodide, 563.

Marine Leeches, branchie of, 210;
generative processes of, 211.

Planarians of eastern seas, 83.

Marsupialia, earliest forms pentadac-
tyle, 580; Hypsiprymnodon mos-
chatus, Prof. Owen on, 573; modi-
fication hind foot of, 573.

Matuta (Pls. XX X1X.—XL.) Banksii,
245; distinguenda, 247; doryphora,
244; eranulosa,sp.n., 245; Leseurii,
248 ; lineifera, sp. n., 245; lunaris,
247; maculata, sp. n., 246; ob-
tusifrons, sp. n., 247; Peronii, 243 ;
picta, 246; planipes, 246 ; revision
species of, 243; rubro-lineata, sp.
n., 244; victor, Mabr., 243; victrix,
243, victrix, var. crebrepunctata,
244,

Matutidee, historical summary of, 241.

Megalema asiatica, skull of, 122.

Megalurus palustris, 352.

Megapodiide, 348.

Megapodius Cumingi, 353; Lowi,
348, 353.

Megathymus, 205.

Melanitis leda, 537, 564.

Melanopelargus episcopus, 349, 354.

Meropide, 316.

Merops bicolor, 316, 350; philip-
pinus, 316, 350.

Messaras erymanthis, 544, 565.

Microhierax erythrogenys, 350.

Microma, 589.

Micronia astheniata, 563.

Microniide, 563.

Micrura fusca, 73.

Miers, HE. J., notes on Oyxstomatous
Crustacea (Pl. XXXVIII.), 235.

Milionia basalis, 558.

Mindanao, Birds of, 309.

Minyad, a larval (Pl. XLY.), 296.

Minyadineze, 295.

Mivart, St. G., and Clarke, R., on
‘sacral plexus and sacral vertebra
of Lizards and other Vertebrata
(Plates LXVI., LXVII.), 513.

Mixornis Woodi, sp. n., 331, 352.

Mca mirabilis, 73.

Monitor arenarius, sacral neryes of,
520.

Monocaulus, sp. ?, 287.

Monticola solitaria, 335, 352.

Mopsea of W. Indies, 50.

INDEX.

Morphine, 538.

Morphology of mammalian Ossicula
auditus, by A. H. G. Doran, 371.
Moseley, H. N.,on new forms of deep-
sea Ascidians obtained during
voyage H.M.S. ‘Challenger’ (Pl.

LXIYV.), 287.

, on new forms of Actiniaria
dredged in deep-sea; with descrip-
tion of certain pelagic surface-
swimming species (Pl. XLY.), 295.

Motacilla luzonensis, 352.

Motacillide, 337.

Mulleripicus funebris, 350.

Munia atricapilla, 345, 353; leuco-
gastra, 345, 853; jagori, 353;
minuta, 353.

Murie, Dr. J., on Steere’s Sponge, a
new genus of the Hexactinellid
group of the Spongida, 219.

Muscicapide of Philippines, 325.

Musophagide, os uncinatum of, 127.

Mycalesis fusca, 537, 564; hesione,
587, 564; janardana, 537, 564;
mineus, 537, 564; nautilus, 564;
polydecta, 537, 564.

Myina annulipes, 589; livens, 589.

Mymar pulchellus (Pl. LXXIIT.),
584; taprobanicus, Ward, 6 & 2
585; Wollastoni, Westw., n. sp.
(Pl. LXXIT.), 585.

Myra angulata, 239; carinata, 239 ;
elegans, 239; mammillaris (Fl.
XXXVIIL.), 239.

Myrina amrita, 550, 567; anasuja,
567; chitra, 549, 566; freja, 550 ;
lapithis, 549, 567 ; marciana, 549,
566; megistia, 549, 566; tharis,
550, 567; travana, 549, 566.

Myristicivora bicolor, 347, 353.

Myside, 45.

Myzanthe pygmea, 352.

Naidonereis quadricuspida, 504.

Nasal labyrinth of Woodpecker, 5, 9.

Nautactis purpureus, n.sp. (P1.XLY.),
295.

Nebalia, sp.?, from Bermudas, 26;
longipes, n. sp. (Pl. VI.), anat. of,
26; characters of, 27.

Nebaliide, 45.

Nectariniide, 339.

Negros, Birds of, 309.

Nemeobiine, 545.

Nemertean, on a new, 73.

Nemertes Neesii, 510.

Nemotois scabriosella, venation wings
of, 160.

Nephropsis Stewarti, 50.

Nephthys cxea, 501; ciliata, 501.

Neptis (Pls. LXVIII.—LX1X.) colu-
mella, 542, 565; dindinga, n. sp.,
542, 565; dorelia, n. sp., 542, 565;
gononata, n. sp., 541, 5€5; hor-
donia, 542, 565; leuconata, n. sp.,
541, 565; mamaja, n. sp., d41,
565; nata, 541, 565; paraka, n.
sp., 542, 565; tiga, 542, 565;
vikasi, 541, 565.

Neptunus sanguinolentus(ftnote), 238.

Neochera marmorea, 558.

Nereis pelagica, 503; zonata, 503.

Nettapus coromandelianus, 354.

Nicolea arctica, 508 ; zostericola, 508.

Nicomache lumbricalis, 507.

Ninox philippensis, 350.

Nothria conchylega, 503.

Notocirrus tricolor, 503.

Numenius pheopus, 304.

Nursia abbreviata, 240; Hardwick,
240 ; plicata? (Pl. XXVIIT.), 240;
sinuata, sp. n., 239.

Nychia Amondseni, 500; cirrosa, 500.

Nyctalemon docile, n. sp., Butler, 562.

Nyctemera coleta, 558; tripunctaria,
558.

Nyctemeride, 558.

Nycticorax griseus, 354; manillensis,
354.

Nyetipao crepuscularis, 561.

Nymphalide, 534.

——, modifications apical segment of

(Pl. LY.), 360.
: harpagones of, 363.
—-, tegumen of, 362.

, values genital armature of, in
family, genera, and species, 364, 366.
Nymphalinz, 539.

Oceanactis, gen. nov. (Pl. XLY.), 296;
rhodactylus, sp. n., 296.

Octacnemus, gen. noy. (Pl. XLIY.),
287 ; bythius, sp. n., 289.

Ocypoda hippeus, 46; rhombea, 46.

Oligosita, 590; collina, 589; nodi-
cornis, Westw., n.sp. (Pl. LX XIIZ.),
592; Staniforthii, n. sp.,591; sub-
fasciata, n. sp., 091.

ne

Ommatophoride, 561.

On Atlantic Crustacea from ‘ Chal-
lenger’ Expedition, by Dr. R. v.
Willemoes-Suhm, 23.

On certain organs of the Cidaride,
by Charles Stewart, 569.

On Hypsiprymnodon, Ramsay, a new
genus indicative of a distinct family
(Pleopodidee) in the Diprotodont
section of the Marsupialia, by
Prof. Owen (Pls. LXXJ-LXXII.),
573.

On morphology of skull in Wood-
peckers (Picide) and Wrynecks
(Yungidee), by W. K. Parker, 1.

Onychoprion anesthetus, 354.

Ophelia limacina, 505.

Ophideres fullonia, 561.

Ophideridee, 561.

Ophiomusium of W. Indies, 50.

Ophioneurus signatus, 589, 590.

Orgyia turbata, n. sp., Butler, 560.

Oriolide, 328.

Oriolus chinensis, 328, 352; philip-
pensis, 352; Steerii, Sharpe, 329,
352; suluensis, Sharpe, 329, 352.

Ornithoptera ruficollis, n. sp., Butler,
552, 567.

Orthia, gen. as characterized by Bois-
duyal, 166, and Boisdv. pars, 193.

augias, 167, 201; nexa, 201;

pelargus, 194.

, 167, untenable as a genus, see
Othria, Westw., 201.

Orthotomus castaneiceps, 337, 352 ;
cinereiceps, 337, 352; derbianus,
352; frontalis, 336,352; ruficeps,
337, 352.

Ortygometra cinerea, 308, 349, 353.

Os uncinatum in Passeres, 110; in
Sun-birds, 111.

Osmotreron axillaris, 346, 353 ; ver-
nans, 346, 353.

Ossicula auditus, on the morphology
of the mammalian, 371.

Osteology of the Spoon-billed Sand-
piper, on the, 213.

Othria, gen. as characterized by
Westwood, 201.

amalthea, 202; amazonica, 202;
augias (P]. XXIX.), 201; colum-
bina, 202; eucadorina, 202; Lin-
digii, 202; nexa, 201.

Owen, Prof., on Hypsiprymnodon mos-

INDEX.

chatus, 573; development of hind
foot in vertebrates, 581, 582.

Owenia filiformis, 507.

Owls, structure skull of, 138.

Oxycephalus piscator, 25.

Oxycerca jagori, 353.

Oxystomatous Crustacea, EH. J. Miers
on (Pls. XXXVITI-XL.), 235.

, literature of subject, 235.

Pachyrhamphus, vomerine cartilages
of, 110.

Padda oryzivora, 308, 345.

Palate of Hemilophus fulvus, 13 ;
of Eagles, 137; Picus analis, 13; P.
major, 14; P. minor, 15; P. viridis,
7; Picumnus minutus, 16; of Vul-
tures, 137.

Palate, summary of characters of the
Saurognathous type of, 19; varie-
ties of Desmognathous, 112.

Palawan, Birds of, 310.

Pamphila aria, 554, 568; augias,
554, 568 ; meesoides, nu. sp., Butler,
554, 568; maro, 555, 568; mat-
thias, 554, 568; nigrolimbata,
555, 568; pilanus, 554, 568.

Panay, Birds of, 309.

Pandita sinope, 542, 565.

Papilio achates, 553, 567 ; agamem-
non, 552, 567; amycus, 182 ;
antiphates, 552, 567; archeus,
156; arruana, 156; arycles, 552,
567; axion, 552, 567; bathycles,
552,567 ; Boisduvalii, 156; brama,
567; chremes, 185; clytia, 552;
cochrus, 177; croesus, 156; cronis,
183; cronius, 156; cyparissias,
168; dardanus, 181; Delessertii,
552, 567; demolion, 552, 567;
diphilus, 553, 567; dedalus, 168 ;
erebus, 567; Esperi, n. sp., Butler
(Pl. LXVIII.), 553, 567; evalthe,
181; evemon, 552, 567; harmo-
dius, 172; helenus, 553, 567;
icarus, 170; iswara, 553, 567 ;
licus, 173; linus, 172; Lydias,
156; malayanus, 552, 567; mestor,
553, 567; oceanus, 156; palati-
nus, 176; panthous, 156; paris
and local varieties of, 156; pega-
sus, 156; pelasgus, 194; phalaris,
177; pirrha, 172; polytes, 552,
567; poseidon, 156; prexaspes,

603

558, 567; priamus, 156, and local
var. of, 156; pronomus, 156 ;
pylades, 170; rama, 567; rich-
mondia, 156; sarpedon, 552, 567 ;
saturnus, 553, 567; syphax, 172;
thais, 185; triton, 156; Urvil-
liana, 156; yvaruna, 553, 567 ;
zagreeus, 189.

Papilionide, 550.

, modifications apical segment of

(Pl. LVI.), 359.

harpagones of, 363.

tegumen of, 361.

——, value genital armature of, in
fam., gen., and species, 364, 365.

Papilionine (subfam.), 552.

Paradisea papuana, palate of, 111.

Paramphinome pulchella, 502.

Parasa bandura, 561.

Parasitic Hirudinei, 209.

Paride, 338.

Parker, W. K., on Morphology of Skull
in Woodpeckers and Wrynecks, 1.

, on the Structure and De-
velopment of the Bird’s skull
(Plates XX.—XXVIT.), 99.

Parthenos lilacinus, n. sp., Butler,
544, 565.

Parus amabilis, sp. n. (Pl. LIJIT),
338, 352; elegans, 338, 352.

Passer montanus, 353.

Passeres of Philippines, 323.

Passerine types, on structure and
development of facial arches in, 104.

Patagonia gigas, face of, 116.

Pedicellariz of Cidaride, 571.

Pelargopsis gigantea, 311, 317, 351;
Gouldi, 351; leucocephala,317,351.

Pelecanoides urinatrix, interorb. fe-
nestre of, 104.

Pelecanus philippensis, 354; roseus,
354.

Penang fauna, to be accepted with
modifications, 534.

Penelopides manille, 351; panini,
309, 323, 351.

Pericrocotus cinereus, 351; igneus,
324, 351.

Petalophthalmus, n. gen., a Mysidi-
form Schizopod, 40; characters of
the genus, 43 ; systematic position
of, 43, 44; armiger, n.sp. (Pl. VII.),
40, 41, 44; anatomy of, 40; habitat
of, 40 ; habits and colour of, 44.

?

b)

604

Phabotreron amethystina, 353; leu-
cotis, 309, 353; nigrorum, sp. 2.,
346.

Phacocheerus, 264.

Phagonia, 588.

Phalacrocorax, sp. ?, 354.

Phalangista, sole hind foot of, 574.

Phasianide, 348.

Philentoma albiventris, sp. n., 325,
351; cyaniceps, 309, 351.

Philippine Archipelago, on birds col-
lected by Prof. J. B. Steere in, 307.

, bird families of, 311.

— , distribution birds of, 308.

, table genera and spec. birds
of, 350.

Phlogcenas luzonica, 348, 353.

Pholoé minuta, 501.

Phyllacanthus baculosa, 571.

Phyllodoce greenlandica, 502.

Phyllornis palawanensis, sp. n.
(Plate L.), 333, 352.

Phyllornithide, 333.

Phylloscopus lugubris, 336, 352;
magnirostris, 352.

Picarie of Philippines, 314.

Picide, 314.

Picus analis, palatal structure of, 13 ;
major, 14; minor, 15; viridis,
development of skull, 4.

Pieride, modifications apical segments
of (Pl. LVI.), 359.

——, harpagones of, 363.

—— ——.,, tegumen of, 361.

——,, values genital armature of, in
fam., gen., and species, 364, 365.

Pierinee (subfam.), 550.

Pieris cronis, 183; cronissa, 183.

Pig, on the domestic, of prehistoric
times in Britain, 251.

, lachrymal bone, distinctively

characteristic, 259, 263.

, nasals characteristic of, 263.

——,, naso-frontal suture of, 262.

, Skull, measurements of lachry-

mals in different species, 269.

, third lower molar distinctive

between Sus scrofa and S. cristatus,

262, 278, 284.

, value as a domestic animal, 258.

Pigs, Aithiopian species of, 257 ;
African do., 256; Asiatic varieties
of, 256, 261; Bornean young, oc-
casionally striped, 280; Chinese

INDEX.

young, never striped, 280; Irish,
260; of Asia, 254.

Pig’s skull, difficulty of distinguishing
European and Indian kinds, Dr.
Gray on, 255.

Pipra auricapilla, vomerine cartilage
of, 110.

Pitta Rochi, Bruggemann, 309.

Pittide, 329.

Planaria, application of term, 85.

Planarians, Mr. Collmgwood on ma-
rine species of, 83.

Planocera thesia (Pl. XIX.), 88, 98.

Planoceride, 88.

Planularia of W. Indies, 50.

Plastingia callineura, 555, 568.

Platalea leucorodia, 217.

Pleopodide, n. fam., Owen, 573.

Plesioneura alysos, 556, 568; as-
mara, 556, 568; dan, 556, 568;
folus, 556, 568; Pinwilli, n. sp.,
Butler (Pl. LXVIIL.), 556, 568.

Plexus and sacral vertebree of Lizards
and Vertebrates, 513.

Ploceidx, 345.

Plotus melanogaster, 354.

Plyctolophide, 312.

Podargus, on structure of face in, 124,

Podiceps philippensis, 354.

Poliolophus urostictus, 334, 352.

Polychrus marmoratus, sacral nerves
of, 522.

Polynema fuscipes, 583 ; natans, 583 ;
natatorial habits of, 584.

Pontobdella, black species of (Pl.
XXXIYV.), 34.

Poreus madagascariensis, 255.

Poritia sumatree, 546, 566.

Poropoea, 589, 590.

Porphyrio pulverulentus, 353.

Porzana fusca, 353 ; pygmea, 353.

Potamocheerus africanus, 255.

Pourretia coarctata, 164.

Pratincola caprata, 335, 352.

Prato, branchie of, 211.

Praxilla arctica,507; praetermissa,507.

Prehistoric times, on the domestic pig
of, 251.

Presacral nerves defined, 514.

Prestwichia aquatica, Lubbock, 587.

Prioniturus discurus, 312, 350.

Prionochilus, sp. ?, 340, 352.

Prionospio Steenstrupi, 507.

Procerus Busku, Coll., n. sp. (Pl.

XVII.), 87, 91; concinnus, Coll.,
n. sp. (Pl. XVII.), 87, 90; Han-
cockanus, Coll., n. sp., 87, 91.
Prometheus casmilus, 178.
Prothoe caledonia, 564,
Pseudolalage melanoleuca, 351.
Psittaci of Philippines, 312.
Psittacide, 312.
Pteroptrix, 589.
Pterylosis of Spoon-billed Sandpiper,
217, 218.
Ptilocichla, n. gen., 332.
falcata, n. sp. (P1.I1.), 332, 353.
Ptilocarpa griseopectus, 353.
Puffinus leucomelas, 309, 354.
Pyrrhocentor melanops, 322, 351.

Querquedula multicolor, 354.

Rallide, 349.

Rallina fasciata, 354.

Ramphastos toco, skull of, 127.

Rana pipiens, stapedials skull of, 106.

, sacral nerves of, 525,

Remipes pacificus ({tnote), 238.

Rhabdornis mystacalis, 353.

Rhamphiculus occipitalis, 353.

Rhinopalpa fulva, 545, 566.

Rhipidura nigritorquis, 325, 351.

Rhopalocera, 535 ; on the male geni-
tal armature in European, 357.

Rhyacophilus glareola, 354.

Rhyncheea capensis, 304.

Rhynchites betule, 590.

Rodine Loveni, 507.

Rolleston, G., on domestic Pig of pre-
historic times in Britai, and on
the mutual relations of this variety
of Pig and Sus scrofa, var. ferus,
S. cristatus, S. andamensis, and S.
barbatus (Pls. XLI.—XLIII.), 251.

Rollulus, sp. ?, 353.

Rothia, n. gen., Westw., charac. of,
204; agrius, 204; eriopis, 204;
pales, 204; pedasus, 204; simyra,
204.

Sabella crassicornis, 508; pavonia, .
508; spetbergensis, 508.

Sacral plexus in Lizards and Verte-
brates, 513.

region, nervous and osseous

structures of, in Batrachians, 523 ;

in Birds, 526; in Mammals, 525;

in Lizards, 522.

Sacral vertebre in birds defined, 530 ;
in Lizards and Vertebrates, 513;
in Mammals, Reptiles, and Batra-
chians, 526.

Salamandrina maculosa, sacral nerves
of, 524.

Salpinx leucogonis, n. sp., Butler,
536, 564.

Samytha sexcirrata, 508.

Sandpiper, Spoon-billed, skeleton of,
213; pterylosis of, 213.

Sarcophanops Steerii, sp.n. (Pl. LIV.),
344, 353.

Sareops calvus, 344, 353; Lowii,
sp. n., 344, 353.

Saturniide:, 561.

Satyride, modifications apical seg-
ment of (Pl. LVI.), 360.

harpagones of, 363.

tegumen of, 362.

, value of genital armature of, in
fam., gen., and species, 364, 366.

Satyrinz, 537.

Saurognathse (Woodpeckers), deve-
lopment face of, 2.

Saurognathous type of palate, sum-
mary of characters of, 19.

Sealibregma inflatum, 506.

Scardia mediella, venation wings of,
160.

Scelostrix candida, 350.

Schizopods, comparison deep-sea and
shallow-water forms, 45; on some
genera of, with a free dorsal shield,
28; synopsis characters of, 44.

Sclerothamnus Clausii, Marshall, =
Dendrospongia Steerii, Wurie, 231.

Scolecolepis cirrata, 506 ; ? Jeffreysii,
MIntsh., n. sp. (Pl. LXY.), 506.

Seoloplos armiger, 504.

Scops megalotis, 350.

Scudder, Mr. §., Syst. Revis. Amer.
Butterflies, remarks on, by Dr. F.
B. White (ftnote), 365.

Seythrops novee-hollandiz, on struc-
ture of face in, 120.

Secusio mundipicta, 558.

Serpula ?. sp. of, 509.

Sesarma obesum (ftnote), 238 ; quad-
ratum (ftnote), 238,

Sesia, 164.

Setaria ruficauda, sp. n., 327, 351.

Sharpe, R. Bowdler, on the birds
eollected by Professor J. B. Steere

2

2

INDEX.

in the Philippine Archipelago (Pls.
XLVI.-LIYV.), 307.

Siphonium of Crow’s and Crocodile’s
skull, 106.

Sipunculus of W. Indies, 50.

Sittide, 338.

Skull of Birds, on the structure of, 99.

Accipiter nisus, 35; in Aetomor-

phe, 128; Dicholophus cristatus,

128; Falco tinnunculus, 132.

Gavia ridibunda, Ist stage, 142 ;

2nd stage, 145; 3rd stage, 146 ;

4th stage, 148; 5th stage, 148 ;

6th stage, 150.

Megalema asiatica, 122 ; Spar-
row-Hawk, 135; Strigide, 138;
Woodpeckers and Wrynecks, 1.

Sphenodon, stapedial processes of, 106.

Sphingide of Malacca, 557.

Sphyngiceps, n. g., Coll., 86; lacteus,
Coll. (Pl. XVII.), 87, 90.

Spilornis holospilus, 313, 350.

Spilosoma, n. sp., 557.

Spirorbis borealis, 509 ;
509 ; verruca, 509.

Spizaétus philippensis, 350.

Sponges, mode of fishing for, in Phi-
lippines (ftnote), 220.

Spongide, on a new geuns of Hexac-
tinellid group of, 219.

Squatarola helvetica, 353.

Syuilla armata (ftnote), 238.

Stapes of Mammalia and Birds (see
under Auditory ossicula), 371-497.

Steatornis caripensis, structure face
ef, 123, 125.

Steere, Prof. J. B., on birds collected
by, in the Philippines, 307.

Steere’s Sponge, on, 219.

Sterna Bergii, 350.

Sternine, 350.

Stephanoseyphus, general remarks on,
64, 65; homology, 64; mirabilis,
n. sp. (Pl. XIY.), 61, 65.

Stewart, Charles, on certain organs of
the Cidaridse, 569.

Strigidee, on skull of, 138.

Strix stridula, mesopterygoid of,
130.

Struthionide, feet of, as compared
with some Marsupials, 580.

Sturnia violacea, 308, 343, 353.

Sturnids:, 343.

Stylochidee, 88.

spirillum,

SECOND SERIES.—ZOOLOGY, VOL. I.

605

Stylochoplana elegans (Pl. XIX.),
88, 98; meleagrina, 88, 98.

Stylochopsis malayensis, Coll., n. sp.
(Pl. XVIIT.), 88, 94.

Suidz, bronze statues of, 278, 279;
Etruscan, 278 ; odontograms teeth
of, 278 ; Prof. Riitimeyer on, 279;
Schutz, Herr J. W., on, 280; skull
measurements ‘of, 275, 276, 281,
282, 283, 284, 285, 286 ; twist in
tail of, 254.

Sulu Islands, birds of, 310.

Surniculus velutinus, sp. n., 320,351.

Sus andamanensis, 254, 262, 280,
283; barbatus, 254, 265, 272, 285,
286; barbatus, peculiarities skull
of, 265; barbirussa, 262; capensis,
257; celebensis, 253, 267, 274;
ceylonensis, 266; cristatus, 253,
254, 255, 256, 269, 284, 285;
europeus, 255; indicus, 253; lar-
vatus, 255; leucomystax, 253 ;
papuensis, 254; scrofa, var.
wgypti, 257; scrofa, var. ceramica,
271; scrofa, var. domesticus, 281;
scrofa, var. ferus, 252, 270, 281 ;
scrofa, var. palustris, 253; senna-
ariensis, 256, 277; sinensis, 256;
tawanus, 264; ternatensis, 276;
timorensis, 254; verrucosus, 254,
266, 367; vittatus, 253, 267:
zeylonensis, 266.

Swine, bibliography on, 272, 273;
parasites of, as elucidating spe-
cific distinctions, 268; relation-
ship of domestic to wild, views on,
252 species from Lofoten Islands,

Uc

Sylvicolidee, pterygoids of, 110.

Sylviide, 336.

Sympalamides mimon, 180.

Sympheedra dirtea, 540, 565; emalea,
565.

Synemon, gen. as used by Westwood,
194; catocaloides (Pl. XXXIIT.),
195, 196 ; directa, 197; gratiosa,
198 ; hesperioides, 198 ; icaria,
196; leta, 195; mopsa, 196;
nais, 195; notha, 197; nupta,
197; obscurella, 197; parthenoides,
196; plana, 196,197; selene, 195,
Sophia, 194; Sternbergii, 182;
theresa, 195; vagans, 198.

Syntomis basifera, 557.

AL

606

Tachytrypane Jefireysi, M‘Intosh, n.
sp. (Pl. LXV.), 505.

Tenia in Swine, 268; solium, 268 ;
mediocannellata, 268.

Tagiades calligana, n. sp., Butler (Pl.
LXIX.), 556, 568; gana, 556, 568;
layata, n. sp., Butler, 557, 568;
rayi, 556, 568; trichoneura, 568.

Tanaécia aruna, 540, 565; pulasara,
540, 565.

Tanagride, pterygoids of, 110.

Tanaid, on a blind deep-sea, 23.

Tanais vittatus, anat. of (Pl. XV.),67;
cavolini, 67; dubius?, 68; habits
of, 70.

Tanygnathus luzoniensis, 312, 350.

Tascina, gen. as defined by West-
wood, 199; orientalis, 199.

Tegumen of European Rhopalocera,
structure of, 360.

Teius teguexin, sacral plexus of, 516.

Telphusa fluviatilis, carries young,
and development with metamor-
phosis, 48.

Terebellides Stroemi, 508.

Terias formosa, 550, 567; hecabe-
cides, 550; imanata, 550, 567 ;
pumilaris, 550, 567; sari, 550,
567; senna, 567.

Terinos robertsia, 544, 565; teu-
thras, 544, 565.

Thaumantis noureddin, 538, 564;
pseudaliris, 538, 564.

Thaumops pellucida, a synonym, 25.

Theclinee (subfam.), 548.

Thecomedusx, a new order of Hydro-
z0a, 65.

Thelephus circinnatus, 508.

Thermesiidx, 562.

Thriponax javensis, 314, 350.

Thysanozoon Alderi, Coll., n. sp., 87,
88; Allmanni, Coll.,n.sp., 87, 89 ;
auropunctatum, 87, 94.

Tiga javanensis, 315, 350.

Timeliide, 331.

Tinea (footnote), 157.

Torfschwein, 254, 279, 282.

Tortoise, sacral nerves of, 523.

Totanus calidris, 309, 354;
cens, 304.

Trayisia Forbesii, 506; glandulosa,
M‘Intosh, n. sp. (P1. LXY.), 506.

canes-

INDEX.

Trematoda, new genus of, 209; n. sp.
of, described, 210.

Treron nasica, 396, 343.

Treronide, 346.

Trichobranchus glacialis, 508.

Trichogramma, 588, 590; erosicornis
(Aprobosca), n. sp. (Pl. LXXIIT.),
592; evanescens, 588 ; vitripennis,
589; Walkeri, 589.

Trichogramini, genera of, 590.

Tringa minuta compared with Ku-
rynorhynchus, 213, 217 ; ruficollis,
354; salina compared with Hury-
norhynchus, 214, 217.

Tringoides hypoleucus, 354.

Trochilide, structure face of, 116.

Troglodytes vulgaris, vomer of, 109.

Trogonide, 316.

Trophonia plumosa, 506.

Tube from Globigerina-ooze, Green- |

land seas, 510.
Turbellaria dendroccela, 85, 86, 87.
Turdide, 335.
Turdus chrysolaus,351; obscurus, 351.
Turnix ocellata, 353.
Turtur Dussumieri, 347, 353; humi-
lis, 347, 353.
Tympanic ring of Gorilla, 377.
Typhlolepta Byerleyana, Coll., n. sp.
(Pl. XVIL.), 88, 92.
Typhloleptide, 88.

Urania, 157.

Uraniide, 562.

Urapteride, 562.

Urapteryx marginipennis, n.
Butler, 562.

Urbanus licus, 184.

Sp.,

Valencinia Armandi, M‘Intosh, n. sp.
(Pl. XVI.), 73; cutaneous system
of, 73; digestive system of, 78 ;
muscles of, 76; nervous system of,
78; lineiformis, 73.

¢Valorous,’ Annelida
eruise of (1875), 499.

Vertebrata, sacral plexus of, 513.

Volvocivora czrulescens, 351.

Vomers of Prcus viridis, 4.

Vultures, the palatine bones of, 137.

obtained in

Westwood, Prof. J. O., descriptions

of some minute Hymenopterous in-
sects, 583.

Westwood, Prof. J. O., monograph of
the lepidopterous genus Castnia
and some allied groups (Pls.
XXVIII-XXXIIT.), 155.

White, Dr. F. Buchanan, on the
male genital armature in the Eu-
ropean Rhopalocera (Pls. LY.—
LYII.), 357.

Wild Boar, from what stock derived,
253.

Willemoesia, noy. gen.(Pls. X.—XII1.),
50; crucifera, n. sp., anatomy of,
52, characters of, 54, colour and
habits of life, 53 ; leptodactyla,n. sp.
(Pl. XIII.), 50, characters of, 54,

Willemoes-Suhm, Dr. R., on some
Atlantic Crustacea from the ‘ Chal-
lenger’ Expedition, 23.

Woodpeckers, on skull of, 1.

Wren, facial structure of, 109.

Wrynecks, morphology skull of, 1.

Xantholema hemacephala, 351;
rosea, dol,

Xantholestes, gen. nov., Sharpe,
327; panayensis, sp. u., 327,
351.

Xanthopygia narcissina, 327, 351.

Xanthotenia busiris, 538, 564.

Ypthima corticaria, nu. sp., Buéler,
537, 564; methora, 537, 564;
philomela, 537, 564.

Yunx torquilla, face of, 17; palate
of, 17.

Zanclopteryx saponaria, 563.

Zebu, Birds of, 309.

Zemeros albipunctata, 545, 566;
emesoides, 545, 566.

Zeocephus cinnamomeus, sp. n. (Pl.
XLVIII.), 352, 328; cyanescens,
sp. n., 328; rufus, 308, 352.

Zereniide, 563.

Zeuzera, 164.

Zeuxidia amethystus, 538, 564.

Zoéa-stages of Crabs, observations on,
46, 47.

Zosterops Meyeni, 352.

Zygenide, 557.

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