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TRANSACTIONS
OF
THE ZOOLOGICAL SOCIETY
OF LONDON.
VOLUME XV.
LONDON:
PRINTED FOR THE SOCIETY :
SOLD AT THEIR HOUSE IN HANOVER-SQUARE ;
AND BY MESSRS. LONGMANS, GREEN, AND CO., PATERNOSTER-ROW.
1901.
PRINTED BY TAYLOR AND FRANCIS.
RED LION COURT, FLEET STRERT.
CONTENTS.
I. Report on the Collection of Fishes made by Mr. J. E. 8. Moore in Lake Tanganyika
during his Expedition, 1895-96. By G. A. BouLencsr, F.RS., F.Z.S. With
an Appendix by J. E. 8. Moorz, A.R.C.S. (Plates I.-VIII.) . . . . page 1
Il. On the Marine Copepoda of New Zealand. By G. Srnwarpson Brapy, M.D., D.Sc.,
PIGS) habeas nC MVMEZiS cau (lates) Xe Xels) se an) ee seme oll
III. On the Extinct Birds of Patagonia.—1. The Skull and Skeleton of Phororhacos
inflatus Ameghino. By C. W. Anpruws, B.Sc., F.Z.S., Assistant in the British
Museum (Natural History). (Plates XIV.-XVII.) ...... =. =. 98d
IV. Second Contribution to the Ichthyology of Lake Tanganyika.—On the Fishes obtained
by the Congo Free State Expedition under Ineut. Lemaire in 1898. By
G. A. Boutencmr, /.#.S., F.Z.S. (Plates XVITI-XX.) . ..... 8
V. On a Specimen of Cervus beigrandi Lart. (C. verticornis Dawk.) from the Forest-
Bed of East Anglia. By Stoney F. Harmer, Sc.D., FRS., Superintendent of
the University Museum of Zoology, Cambridge. (Plate XXI.) . . . . 97
VI. A Monograph of the Genus Casuarius. By the Hon. Waurer Roruscuip, Ph.D.,
F.ZS. Witha Dissertation onthe Morphology and Phylogeny of the Paleognathe
(Ratitee and Crypturi) and Neognathe (Carinate). By W.P. Pycrarr. (Plates
ONDE OUNTS)) Gy Boe tee ne cee eee eee 5 1(OKY
VII. Notes on the Transformations of some South-African Lepidoptera. By Lneut.-
Colonel J. Maucotm Fawcett. (Plates XLVI-XLIX.) .... . . 291
VILL. On some Points in the Anatomy of Polypterus. By J. S. Buperrr, WA., F.Z.S.,
Trinity College, Cambridge. (Plates L.-LIL) . ....... . . 828
Histroimcthesbapersicontained sim Vole XV.) 9. ai 2) ee ener ooo
ING EXSOTASPECLES te CCOMM ne ae olen Mira, 6. 4 Gul le Uae ey sete: sa Caen 4
TRANSACTIONS
OF
THE ZOOLOGICAL SOCIETY
OF LONDON.
Vout. XV.—Parr 1.
LONDON: |
PRINTED FOR THE SOCIETY,
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TRANSACTIONS
OF
THE ZOOLOGICAL SOCIETY
OF LONDON.
I. Report on the Collection of Fishes made by Mr. J. E. S. Moore in Lake Tanganyika
during his Expedition, 1895-96. By G. A. Bounencnr, F.R.S., ZS. With
an Appendix by J. E. S. Moors, A.R.C.S.
Received and read June 21, 1898,
[Puates I.-VIIT.]
THE examination of the collection of fishes brought home from Lake Tanganyika by
Mr. Moore, and with which I have been entrusted by him, has not yielded any such
startling results as have already been announced after study of the Invertebrates! Of
the two series in the fauna of Tanganyika, distinguished by its explorer as the normal
freshwater and the halolimnic groups?, the latter is unrepresented in the collection.
This may be due either to the origin of the present fish-fauna not reaching so far back
in time as that of the Mollusca and other Invertebrates, or to the incompleteness of
the series brought home. ‘The latter explanation may be the correct one, since, owing
to the lack of suitable dredging-apparatus, and to the difficulties of preserving, to
which Mr. Moore has alluded ?, only a small proportion of the fishes of the Lake
could be collected, mostly littoral forms found about the rocks; of the deeper-water
fishes, which were observed to be abundant both in species and in individuals, we are
still almost entirely ignorant; it is therefore to be expected that further collections,
1 J, E.S. Moore, “On the Zoological Evidence for the Connection of Lake Tanganyika with the Sea,”
Proc. R. Soe. lxii. 1898, pp. 451-458.
2 J. E. S. Moore, ‘“‘ The Molluses of the Great African Lakes,” Q. Journ. Micr, Sci. 1898, pp. 159-204,
3 L.c. See also‘‘The Physiographical Features of the Nyasa and Tanganyika Districts,’ Geogr. Journ
1897.
VoL. XV.—PART I. No. 1.—Deceniber, 1898. B
2 MR. G. A. BOULENGER ON A COLLECTION
made under more favourable circumstances, would considerably modify our views on
the general character of this most important section of the fauna.
How incomplete the collection is may be partly realized from the fact that, of the
six species previously described from Tanganyika, only one was rediscovered by
Mr. Moore. His series of sketches executed from fresh specimens, free use of
which he has kindly given me, thus enabling me to represent some of the new
species in their natural colours, also indicate the existence of several fishes which are
unrepresented in the collection. Large fishes, owing to the impossibility of preserving
them, had to be left behind, and the difficulties of transport by carriers resulted in the
loss of several jars containing spirit-specimens. Nevertheless, small as it is, and
though deficient in any but typically African freshwater forms, the collection is of great
interest, and Mr. Moore well deserves the thanks of all zoologists for the manner in
which, amid so many difficulties, he has succeeded in affording us a first glimpse at a
fish-fauna which has so long remained a mystery.
A study of the freshwater fishes of Africa has hitherto led to the assumption that
the bulk of the fauna originated in the region of the great lakes, whence they have
radiated towards the Mediterranean and the Atlantic and Indian Oceans—a view based
on the close affinity, often amounting to specific identity, of the fishes of the Nile, the
Niger, the Congo, and the Zambesi. The homogeneity of the fauna is absolutely
opposed to the conception of the great watersheds having been stocked from the sea,
within the geological period of which these fishes are representative, this being evident
even in the case of such forms as Zates and Tilapia, which are known to enter salt water.
Nothing contrary to this theory is brought to light by an examination of the fishes
obtained by Mr. Moore in Lake Tanganyika. The striking character of the Tanganyika
fish-fauna, as revealed by Mr. Moore’s collection, is the extraordinary variety of the
Cichliidw). This is a natural group distributed all over Africa, including Madagascar,
but, although rich in species, nowhere else showing within a limited area anything like
the modifications of structure described in this report, which have necessitated the
establishment of nearly as many, new genera as were previously known from the whole
of Africa. The generalized characters of some of the Cichlidw occurring in Lake
Tanganyika, regarding as indicative of generalization the greater development of the
anal fin, consistently with the system followed in classifying their more primitive
allies the Centrarchidw?, and the extent of the lateral lines, both of which are
complete in some of the newly-discovered forms®, afford further support to the
1 See P. Z. 8. 1898, p. 182. * Cat. Fish. 2nd ed. 1. p. 2 (1895).
$ The lateral line has usually been described as “ interrupted” in the Cichlidw and other groups in which
it has a similar disposition. ‘This expression conveys a serious misconception, and I have replaced it in the
diagnoses of the Cichlide and the Serranide allied to Plesiops by “ lateral lines two,” the necessity for this
change being particularly obvious in the case of some of the species described in this report. Fishes had no
doubt originally a greater number of lines of sense-organs along the body, as we may still observe in many of
OF FISHES FROM LAKE TANGANYIKA. 8
proposition enunciated above. Lake Tanganyika might be regarded as the centre of
origin of all the African genera of Cichlide.
Unfortunately, nothing reliable is yet known of this family in a fossil condition, with
the exception of a few American Tertiary types, which do not differ materially from
those inhabiting the same part of the world at the present time. There is no reason for
referring to it the Cretaceous fishes from Mount Lebanon described as “ Chromides”’
by Heckel! and by Davis. All we can gather from Heckel’s description is that
Pycnosterynz is based on some ‘ Pharyngognath Acanthopterygian,” and that it does
not belong to the Cichlide is conclusively proved by his statement: “ Rippen kurz,
dtinn, die hinteren auf langen Querfortsitzen ansitzend,’ a sentence that has been
translated by Davis as “ Ribs short and slender, the posterior ones supporting long
transverse apophyses”*. An examination of some of the specimens of Pycnosteryna’
exhibited in the British Museum suggests to me special relation to the Berycide, with
which they are provisionally associated by Mr. Smith Woodward, and certainly no
affinity to the Cichlide. Considering that coalesced lower pharyngeal bones occur at
the present day in such widely different groups as the Serramde, Gerride, Scienide,
Cichlide, Pomacentride, Embiotocide, Labride, Scaride, Pleuronectide, and Scombre-
socidc, this character, if correctly ascertained by Heckel, would in itself be no serious
objection to the allocation of Pycnosterynz among the Berycide. Giinther* also
doubts the correctness of Heckel’s determination when he states that “the position
of Pycnosterynx is uncertain,’ but we are not enlightened as to its affinities by the
further remark that “it approaches certain Pharyngognaths.” It appears certain that
true Perciform fishes, to which large group the Cichlidw belong, have not yet been
described from pre-Tertiary beds.
In order to show what is now known of the fish-faunas of the great lakes, lists are
appended of the fishes of Lakes Nyassa, Tanganyika, Victoria Nyanza, and Rudolf.
These lists must of course be taken as giving a very inadequate idea of the fauna,
owing to the incompleteness of the collections on which they are based; but, imperfect
as they are, they nevertheless will prove useful as a basis from which to judge of the
relation existing between the piscine inhabitants of these lakes. Of the other lakes,
unfortunately, nothing can be said at present, the occurrence of an undetermined species
of Haplochilus in the Albert Nyanza being all the information we possess.
the lower forms. In the Teleosteans, as a general rule, the lateral lines are reduced to one on each side,
extending uninterrupted from the head to the root of the caudal fin, although among this order there is
hardly a single large group that does not offer exceptions such as are shown in the Cichlidw. In these, both
upper and lower lateral lines may be complete ; or the upper may be complete and the lower reduced, or the
reverse; or the upper may be incomplete posteriorly and the lower anteriorly, which latter disposition has led
to the misconception involved in the term “ interrupted.”
1 Abbild. u. Beschr. n. Thier. Syrien, p. 235 (1843). * Tr. R. Dublin Soe. (2) iii. 1888, p. 531.
2 1D, @ i BBL * «Study of Fishes,’ p. 200 (1880).
BZ
4 MR. G. A. BOULENGER ON A COLLECTION
I. Lake Nyassa},
CicHLIDz. SILURIDA.
1. Paratilapia robusta Gthy. 24. Bagrus meridionalis Gthr.
2. fs afra Gthr. 25. Anoplopterus platychir Gthr.
3. ie modesta Gthr. 26. Synodontis zambesensis Gthyr.
4., 5) livingstonit Gthr,
De BS intermedia Gthr, Coro
6. a dimidiata Gthr.
7. x longiceps Gthy. 27, Labeo mesops Gthr.
8. Corematodus shiranus Bley. 28. Barbus trimaculatus Ptrs.
9. Tilapia shirana Blgr. 29. Barilius guentheri Blgr.
TO, 45 qoseomlibon ins. 30. Engraulicypris pinguis Gthy,
11. ,, kirkii Gthr. 31. Pelotrophus microlepis Gthr.
12. 4, == squamipinnis Gthr, 82. 2 microcephalus Gthr.
13. 4, vrendalli Bigr.
14. 4, ~~‘ dateristriga Gthr. @HARACINIDE:
15. 4, subocularis Gthr.
16. 4, johnstont Gthr.
17. ~=4, ~——‘lethrinus Gthr.
18. ,, — ¢tetrastigma Gthr. CyPRINoDONTIDs.
19. 4, ~ callipterus Gthr. 34. Haplochilus johnstoni Gthr.
20. 4, williamsi Gthr.
21. 4, aurata Bigr.
22. Docimodus johnstoni Blgr.
33. Alestes imberi Ptrs.
MorMyRID&.
35. Mormyrus discorhynchus Ptyrs.
MAstTAacEMBELIDA. 36. 35 catostoma Gthr.
23. Mastacembelus shiranus Gthr. 37. Mormyrops zambanenje Ptrs.
Il. Lake Tanganyika ?.
SERRANIDZ. 9. Telmatochromis temporalis Blgr.
1, Lates microlepis Blgy. 10. Julidochromis ornatus Blgy.
ll. Paratilapia pfefferi Blgr.
CicHLIps. 12. sy macrops Blgr.
2. Lamprologus fasciatus Blgr. 13. aa ventralis Blgr.
3. $ compressiceps Blgy. 14, 53 furcifer Blgr.
4. i moori Blgr. Nie i leptosoma Blgy.
5. 90 modestus Blgr. 16. Ectodus descampsii Blgr.
6 = elongatus Blgr. 17. 4, + melanogenys Bigr.
7 os furcifer Blgr. 18. Bathybates ferox Blgr.
8. Telmatochromis vittatus Blgr. 19. Eretmodus cyanostictus Blgr.
’ Of. Giinther, P. Z. 8. 1864, p. 307, 1898, p. 616, and Ann, & Mag. N. H. (6) xvii. 1896, p.397; Boulenger
P. Z.8. 1896, p. 915, and Ann. & Mag. N. H. (6) xix. 1897, p. 155, and (7) i. 1898, p. 254.
* Cf. Giinther, P. Z. 8. 1893, p. 628.
OF FISHES FROM LAKE TANGANYIKA,
Cicuiip# (continued).
20. Tilapia tanganice Gthr.
al, » Ourtoni Gthr.
22. a5 labiata Blgr.
23. Tropheus moorii Bley.
24, Simochromis diagramma Gthy.
25. Petrochromis polyodon Blgr.
26. Perissodus microlepis Bley.
27. Plecodus paradozus Bley.
MasraceMBrLipa.
28. Mastacembelus moorii Blgr.
29. B tanganice Gthr,
30. 5 ophidium Gthr.
SILuRIDA,
31. Clarias anguillaris L.
32. ,, ~—_ liocephalus Blgr.
33. Anoplopterus platychir Gthr.
34, Auchenaspis biscutata Geoftr.
35. Synodontis multipunctatus Blgr.
36. Malapterurus electricus Gm,
CyPRINID&.
37. Labeo, sp.
CHARACINIDA,
38. Alestes macrolepidotus C. & V.
39. ,, = macrophthalmus Gthr.
40. Hydrocyon forskalit Cuv.
CyPRINODONTID&.
41. Haplochilus tanganicanus Blgr.
PoLyPrERID&m.
42. Polypterus bichir Geoffr, (?).
LEPIDOSIRENIDA,
43. Protopterus annectens Ow.?’.
Ill. Lake Victoria Nyanza?.
Cicuiipe.
- Paratilapia longirostris Hilg.
99 cavifrons Hilg.
% retrodens Hilg.
. Tilapia nilotica Cuv.
» nuchisquamulata Hilg.
» Ssaduvagii Pfeff,'
» obliquidens Hilg.
ow GD) £8 E> $9 8S IS
MastaceMBELIDA,
8. Mastacembelus, sp.
SILURIDZ.
9. Clarias, sp.
10. Synodontis afrofischeri Hilg.
CyprinIp@.
ll. Labeo forskalii Riipp.
12. Labeo rueppellit Pfeff.
13. Barbus pagenstecheri Fisch.
14, 4, trimaculatus Ptrs.
CHARACINIDA,
15. Alestes rueppellit Gthr.
CyYPRINODONTIDA.
16. Fundulus teniopygus Hilg.
Mormynripa&,
17. Mormyrus oxyrhynchus Geoftr.
18, fe longibarbis Hilg.
LeEpIDUSIRENIDE,
19. Protopterus annectens Ow.
' Reported from Lake Tanganyika by Sir H. H. Johnston, Brit. C. Afr. p. 362 (1897).
2 Hilgendorf, Sitzb. Ges. naturf. Fr. Berl. 1888, p. 75; Pfeffer, Thierw. O.-Afr., Fische (1896); Giinther,
Ann. & Mag. N. H. (6) xvii. 1896, p. 397.
6 MR. G. A. BOULENGER ON A COLLECTION
IV. Lake Rudolf.
CicH1iip2. CHARACINIDA.
1. Tilapia DURE Cae 6. Citharinus geoffroyi Cuv.
2. 5, tristrami Gthr. 7. Alestes rueppellii Gthr.
SILURIDE. 8. Distichodus rudolphi Gthy.
3. Synodontis schal Bl. Schn.
: 4, ” smithit Gthr. PouyPrEeRIp”®.
Cyprinip&. 9. Polypterus senegalus Cuy.
5. Barbus, sp.
The fishes of Lake Nyassa are, with two exceptions, specifically distinct from those
of the Nile, as pointed out by Dr. Giinther ?, while seven out of forty-three species
represented in Lake Tanganyika occur both in the Nile and in the rivers flowing into
the Atlantic. And as the Mormyride, which furnish the two species common to
Nyassa and the Nile®, have not yet been recorded from Tanganyika, while, with the
exception of a small stream-Siluroid, not one of the species described from the former
lake has been rediscovered in the latter, it follows that, although similar in general
character, the fish-fauna of the two lakes shows no trace of community so far as
specific forms are concerned, as might have been expected from the absence of direct
communication between them.
Before concluding these prefatory remarks, I wish to express my thanks to
Messrs. J. H. Gardiner and J. Green for their kind assistance in supplying me with
sciagraphs of the new fishes, which, supplementing the skeletons prepared by
Mr. Groenvold, have enabled me to add some notes on the osteological characters of
the genera described as new.
SERRANIDA.
1. Laves MICROLEPIS, sp. n. (Plate I. fig. 1.)
Body elongate, its depth 33 times in the total length. Length of head 3 times in
total length; upper profile nearly straight ; diameter of eye equal to length of snout,
33 times in length of head; lower jaw projecting ; maxillary extending to below centre
of eye, the width of its distal extremity not quite half diameter of eye; pree- and
suborbitals finely serrated; cheeks, opercles, -and occiput covered with small scales ;
preopercular border forming nearly a right angle, finely toothed on its vertical limb,
with two or three widely-separated spines on its lower limb, and with one or two very
1 Giinther, P. Z. 8. 1896, p. 217. > Study of Fishes,’ p. 230.
3 Tf, as seems probable, the distinction between Mormyrops zambanenje and M. anguilloides should not be
maintained. On the other hand, the Nilotic specimens referred to Mormyrus discorhynchus may prove to
be specifically separable. Lake Tanganyika might thus ultimately possess no species of fish in common with
the Nile.
OF FISHES FROM LAKE TANGANYIKA. U
strong spines at the angle; opercular spine as much developed as the latter ; clavicle
with a group of 3 or 4 strong spines. 16 gill-rakers on lower part of anterior arch.
Dorsal VII, II 11; first and second spines short, third very strong, longest, ? length
of head; longest soft rays 3 length of head. Pectoral } length of head. Anal III 8;
spines short, second and third equal. Caudal truncate. Caudal peduncle twice as
long as deep. Scales 100-110 ee Body silvery, spotted and marbled with brown ;
dorsals spotted with brown ; caudal with irregular brown bars.
Total length 155 millim.
Two young specimens from Kinyamkolo.
This species is closely allied to the widely-distributed Lates niloticus Hasselq.
Compared to young specimens of the latter, it differs in the smaller scales, the higher
spinous dorsal, the longer caudal peduncle, aad the shape of the caudal fin, which is
truncate instead of rounded.
CICHLIDA.
2. LAMPROLOGUS FASCIATUS, sp. n. (Plate I. fig. 2.)
A few moderately large curved canine teeth in front of each jaw, followed by a
narrow band of minute teeth; lateral teeth very small. Depth of body 4 times in
total length, length of head 3. Snout as long as the diameter of the eye, which is
3 times in length of head and equals 13 interorbital width; maxillary extending to
below anterior border of eye; cheeks naked; opercles and occiput scaled. Gill-rakers
short, 12 on lower part of anterior arch. Dorsal XIX 8; spines increasing in length
to the last, which measures 2 length of head and nearly equals longest soft rays.
Pectoral 4 length of head. Ventral reaching vent. Anal X 6; spines increasing in
length to the last, which slightly exceeds longest dorsal. Caudal rounded. Caudal
peduncle as long as deep. Scales 46 2s lat. 1. — Yellowish, with 11 dark brown
bars, the first across the vertex ; fins greyish, dorsal and anal edged with blackish.
Total length 70 millim.
A single specimen from Kinyamkolo.
3. LAMPROLOGUS COMPRESSICEPS, sp. n. (Plate I. fig. 3.)
A few moderately large curved canine teeth in front of each jaw, followed by a
narrow band of minute teeth; lateral teeth very small, curved. Depth of body 25 to
23 times in total length, length of head 23? to 23. Head very strongly compressed,
with concave upper profile; snout a little longer than diameter of eye, which is
4 times in length of head and equals 14 interorbital width ; maxillary extending to
below anterior border of eye; cheeks naked; opercles and occiput with small scales.
Gill-rakers moderately long, 15 on lower part of anterior arch. Dorsal XX-XXI 6;
8 MR. G. A. BOULENGER ON A COLLECTION
spines increasing in length to the sixth, which measures half length of head, the
posterior a little shorter; longest soft rays a little longer than longest spines.
Pectoral 4 to 3 length of head. Ventral produced into a filament. Anal X 5; spines
increasing in length to the last, which equals the last dorsal ; longest soft rays 2 length
of head. Caudal rounded. Caudal peduncle as long as deep. Scales 32-33 2;
12)
lat. 1. a . Brown, with indistinct traces of five darker vertical bars; pectoral bright
yellow, other fins blackish towards the border.
Total length $3 millim.
Two specimens from Kinyamkolo.
4, LAMPROLOGUS MOORI, sp. n. (Plate I. fig. 4.)
9 or 10 equal, moderately large conical teeth in front of each jaw, followed by a
narrow band of minute teeth; lateral teeth very small. Depth of body 24 times in
total length, length of head 3 to 33. Snout as long as diameter of eye, which is
3 to 33 times in length of head and equals interorbital width; maxillary extending
to below anterior border of eye; cheeks with small, deciduous scales; opercles and
occiput scaled. Gill-rakers short, 9 or 10 on lower part of anterior arch. Dorsal
XIX-XX 8-9; spines slightly increasing in length to the last, which measures nearly
half length of head; middle soft rays prolonged, at least # length of head. Pectoral
2 to $ length of head. Ventral prolonged into a long filament. Anal VII-VIII
6-7 ; spines increasing in length to the last, which is a little longer than the longest
dorsal ; middle soft rays prolonged into filaments. Caudal rounded. Caudal peduncle
; lat. 1. 2°. Dark brown ; fins blackish.
2? 9-13"
99 9
as long as deep. Scales 33-35
Total length 95 millim.
Several specimens from Mbity Rocks and Kinyamkolo.
5-7
11-1
5. LAMPRGLOGUS MODESTUS, sp.n. (Plate I. fig. 5.)
A few large curved canine teeth, tipped with brown, in front of each jaw, followed
by a band of minute teeth; lateral teeth very small. Depth of body 3% to 32 times
in total length, length of head 3 to 33. Snout a little longer than diameter of eye,
which is 384 to 4 times in length of head and equals interorbital width; maxillary
extending to below anterior border of eye; cheeks naked; opercles and occiput scaled.
Gill-rakers very short, 7
increasing in length to the last, which is not quite half length of head; middle soft
rays prolonged, % to # length of head. Pectoral about $ length of head. Ventral
prolonged into a short filament. Anal V 6-7; spines increasing in length to the last,
which is as long as middle dorsals; middle soft rays prolonged, like the dorsals.
on lower part of anterior arch. Dorsal XX 8-9; spines
Caudal truncate. Caudal peduncle as long as deep. Scales 36-40 = lat. 1. 7.
OF FISHES FROM LAKE TANGANYIKA. 9
Uniform brown; soft dorsal and caudal fins with round black spots between the
rays.
Total length 75 millim.
A single specimen from Mbity Rocks, and another from Kinyamkolo.
6. LAMPROLOGUS ELONGATUS, sp. n. (Plate I. fig. 6.)
6 to 8 large canine teeth in front of each jaw, followed by a broad band of minute
villiform teeth ; lateral teeth very small. Depth of body 4 times in total length,
length of head 2% to 2$. Snout twice as long as diameter of eye, which is 5 times in
length of head and equals interorbital width ; maxillary extending to below anterior
border of eye; cheeks naked; opercles and occiput scaled. Gill-rakers moderately
long, 12 on lower part of anterior arch. Dorsal XVIII 10-11; spines slightly
increasing in length to the last, which measures 4 length of head; longest soft rays
half length of head. Pectoral half length of head. Ventral reaching vent. Anal
V 8; spines increasing in length to the last, which equals longest dorsal. Caudal
truncate. Caudal peduncle 14 as long as deep. Scales 90-95 aoe) lat. 1. =
Brown, with darker spots having a tendency to form cross-bars ; caudal fin with round
dark spots between the rays.
Total length 115 millim.
One specimen from Mbity Rocks, and one from Kinyamkolo.
7. LAMPROLOGUS FURCIFER, sp. n. (Plate II. fig. 1.)
A few large curved canine teeth in front of each jaw, followed by a moderately
broad band of minute villiform teeth ; lateral teeth very small. Depth of body 4 to 44
times in total length, length of head 2$ to 3. Snout as long as or a little longer than
diameter of eye, which is 54 to 3% times in length of head and exceeds interorbital
width; maxillary extending to below anterior fourth of eye; cheeks and opercles with
deciduous scales. Gill-rakers short, 16 on lower part of anterior arch. Dorsal
XX-XXI 7-8; spines increasing in length to the last, which measures > length of
head; middle soft rays produced, 3 to ? length of head. Pectoral # length of head.
Ventral reaching vent or origin of anal. Anal VI-VII 6; spines increasing in length,
the last nearly as long as last dorsal; middle soft rays produced. Caudal deeply
notched, crescentic. Caudal peduncle 14 as long as deep. Scales 50-54 ae lat. 1.
a lower lateral line often nearly complete. Dark brown, with very indistinct
blackish bars on the body; dorsal and caudal with round black spots between the
rays; tips of the caudal lobes whitish.
Total length 125 millim.
Three specimens from Kinyamkolo, and one from Mbity Rocks.
VOL. XV.—PaART I. No. 2.—December, 1898.
10 MR. G. A. BOULENGER ON A COLLECTION
The genus Lamprologus was known from a single species L. congoensis Schilthuis,
discovered a few years ago in the Congo. The species now described raise the number
to 7, distinguishable by means of the following key :—
I. Caudal rounded or truncate.
A. Anal with 10 spines.
D. XIX 8; Sc. 46 a depth of body 4 times in total length. . . . . . 1. L. fasciatus.
D. XX-XXI 6; Se. 32-33 & ; depth of body 23 to 2? times in total length. 2. L. compressiceps.
B. Anal with 6 to 8 spines.
D. XIX-XX 8-9; Sc. 33-35 © ; depth of body 23 times in total length . 3, L. moorit.
11-12’
D. XVITI-XIX 8-10; Se. 42-53 wats depth of body 3} to 4 times in total length.
4. L. congoensis.
C. Anal with 5 spines.
D. XX 8-9; Sc. 36-40 56. depth of body 3} to 32 times in total length. 5. L. modestus.
11-14”
D. XVIII 10-11; Se. 90-95 sat depth of body 4 times in total length . 6. L. elongatus.
II. Caudal deeply notched, crescentic. :
D, XX-XKI 7-8; A. VI-VII6; Sc.50-54 77. 2 2 2 1 1 1. . 7. LL. furcifer.
TELMATOCHROMIS, g. 1.
Body more or less elongate; scales ctenoid. Jaws with a series of conical teeth,
followed by a broad band of minute tricuspid teeth; lateral teeth small, conical.
Maxillary exposed. Dorsal with 20 to 22 spines, anal with 6 or 7. Vertebre 33
(16+17).
This genus is closely allied to Lamprologus, differing in the small teeth forming a
band behind the outer row being tricuspid instead of conical. It therefore stands
somewhat in the same relation to Lamprologus as Tilapia does to Paratilapia.
8. TELMATOCHROMIS VITTATUS, sp. n. (Plate II. fig. 2.)
12 to 16 enlarged conical teeth, tipped with brown, in the outer row in each jaw.
Depth of body 43 to 4% times in total length, length of head 4. Snout descending in
a strong curve, as long as or a little longer than the diameter of the eye, which is
32 to 4 times in length of head and equals interorbital width ; maxillary extending to
below the nostril; head naked, opercle with a few deciduous scales. Gill-rakers very
short and tew. Dorsal XXI-XXII 8; spines increasing in length to the last, which
equals } length of head; soft rays a little longer. Pectoral ¢ length of head. Ventral
produced into a short filament, reaching origin of anal. Anal VII 5-6; spines
increasing in length to the last, which equals last dorsal. Caudal rounded. Caudal
peduncle as long as deep. Scales 45-02 +3 Natale ae Yellowish, with a dark
OF FISHES FROM LAKE TANGANYIKA, 11
brown lateral stripe from the upper lip, through the eye, to the base of the caudal,
where it expands into a spot; another dark brown stripe from the vertex along the
base of the dorsal; a few brown spots on the dorsal; anal edged with dark brown; a
black bar at the base of the pectoral, which is white.
Total length 78 millim.
‘Two specimens from Mbity Rocks.
9. TELMATOCHROMIS TEMPORALIS, sp. n. (Plate IT. fig. 3.)
8 to 12 enlarged conical teeth, tipped with brown, in the outer row in each jaw.
Depth of body 3 to 33 times in total length ; length of head 3 to 3. Snout descending
in a strong curve, 14 as long as the diameter of the eye, which is 44 times in length of
head and a little less than interorbital width; maxillary extending to below anterior
border of eye; head naked, or with a few deciduous scales on the opercles. Gill-
rakers very short and few. Dorsal XX-XXI 6-7; spines increasing in length to the
last, which equals 3 to } length of head; middle soft rays produced, % to $ length of
head. Pectoral 3 length of head. Ventral produced into a filament, extending beyond
origin of anal. Anal VI-VII 6-7; spines increasing in length to the last, which
equals or slightly exceeds last dorsal; soft rays produced, like the dorsals. Caudal
rounded. Caudal peduncle as long as deep. Scales 43-46 <3 lat. 1. om Brown,
with small round darker spots between the dorsal, anal, and caudal rays; a dark
horizontal streak behind the eye; a dark bar at base of pectoral.
Total length 85 millim.
Three specimens from Kinyamkolo, and one from Mbity Rocks.
The deep anterior groove of the skull, in which the ascending processes of the pre-
maxillaries are received, extends to the anterior third of the orbits. and the strong
occipital crest is prolonged forward to it; parietal crests are entirely absent; the
chain of suborbital bones is very slender. None of the ribs are sessile, being inserted
on a step at the back of the transverse processes of the vertebrae at a short distance
from the centre; all bear epipleurals ; only the last pracaudal vertebra has a hemal
bridge.
JULIDOCHROMIS, g. n.
Body elongate ; scales ctenoid. Jaws very narrow, with a few curved canines in
front, the outer of which are very large and tusk-like, followed on the sides and behind
by minute conical teeth forming a narrow band. Maxillary exposed. Dorsal with 22
to 24 spines, anal with 8 or 9. Vertebre 34 (17-+-17).
This genus represents an exaggerated type of Lamprologus. ‘The very specialized
dentition and the general aspect remind one of the Julidine Labridw, whence the
name proposed.
G
bo
12 MR. G. A. BOULENGER ON A COLLECTION
- 10. JuLIpocHRoMIS orNATUS, sp. n. (Plate II. fig. 4.)
4 or 6 canines in each jaw, tipped with brown. Depth of body 4 to 43 times in
total length, length of head 34 to 34. Snout 14 to twice as long as diameter of eye,
which is 44 to 5 times in length of head and 1} in interorbital width ; maxillary
extending to below nostril ; cheeks naked ; opercles scaled. Gill-rakers very short and
few. Dorsal XXII-XXIV 5; spines equal from the eighth or tenth, 4 length of head ;
longest soft rays 4 to 3 length of head. Pectoral about 3 length of head. Ventral
produced in a filament, reaching origin of anal. Anal VIII-IX 4-6 ; spines increasing
in length to the last, which equals 2 length of head. Caudal rounded. Caudal peduncle
as long asdeep. Scales 45-50 are: lat. 1. aoe. Yellowish, with three dark brown
stripes on each side, the lower from the end of the snout to the base of the caudal, the
upper along the base of the dorsal; a large round dark-brown spot on the base of the
caudal; a small black spot at the base of the pectoral; anal edged with brown;
caudal brown.
Total length 85 millim.
Five specimens from Mbity Rocks.
The premaxillary groove is deep and extends to the anterior third of the orbits ; the
occipital crest is low and continued forward to the premaxillary groove; parietal
crests are present, but very feeble; the chain of suborbital bones is replaced by a
ligament ; the mandible is very massive, the lower surface flat, projecting as a keel on
the sides. The ribs are inserted on a step of the transverse processes, at a short
distance from the centre; all bear epipleurals; only the last preecaudal vertebra has a
hemal bridge.
11. PARATILAPIA PFEFFERI, sp. n. (Plate ITI. fig. 1.)
Teeth small, in 3 series in each jaw, forming a narrow band, outer largest. Depth
of body equal to length of head, 2% times in total length. Snout with straight upper
profile, as long as eye, the diameter of which is 33 times in length of head and equals
Id interorbital width ; maxillary extending to below anterior border of eye ; 3 series of
scales on the cheek; large scales on the opercle. Gill-rakers rather long, 11 on lower
part of anterior arch, the larger spatulate. Dorsal XVI &; spines increasing in length
to the sixth, which measures a little less than } length of head and nearly equals
longest soft rays. Pectoral $ length of head. Ventral reaching origin of anal. Anal
III 7; third spine longest, as long as longest dorsal. Caudal feebly emarginate,
Caudal peduncle 1 as long as deep. Scales very finely denticulate on the edge, 332 ;
lat. 1. — Pale olive above, silvery beneath, with seven darker vertical bars; fins
greyish brown.
Total length 76 millim.
A single specimen from Kinyamkolo.
OF FISHES FROM LAKE TANGANYIKA, 13
This species, which I have much pleasure in naming after the distinguished zoologist
of Hamburg, Dr. G. Pfeffer, who has much contributed to our knowledge of East
African ichthyology, is nearest allied to P. intermedia Gthr., from Nyassa.
12. PaRaTILAPIA MACROpS, sp. n. (Plate III. fig. 2.)
Teeth small, in 3 series in each jaw, forming a narrow band, outer largest. Depth
of body three times in total length; length of head 22 to 8. Snout with straight upper
profile, a little shorter than the eye, the diameter of which is 2% times in length of
head and exceeds interorbital width; maxillary extending to below anterior border of
eye; 2 or 3 series of scales on the cheek; large scales on the opercle. Gill-rakers
short, 11 on lower part of anterior arch. Dorsal XVI 10-12; spines increasing in
length to the sixth, which measures a little less than 3 length of head and equals
longest soft rays. Pectoral as long as head. Ventral extending a little beyond origin
of anal. Anal III 6-7; third spine longest, a little shorter than longest dorsal,
Caudal with crescentic emargination. Caudal peduncle as long as deep. Scales very
finely denticulate on the edge, 33-34 oe latayele oa} upper lateral line complete,
extending to base of caudal. Pale brownish above, silvery beneath; a series of five
indistinct dark blotches on each side of the body; spinous dorsal edged with brown
above.
Total length 70 millim.
Two specimens from Kinyamkolo, and one from Mbity Rocks.
Closely allied to the preceding species; distinguished by the larger eye, the complete
upper lateral line, the longer pectoral, and the more strongly emarginate caudal.
Connects the preceding species with the following, which represents a more aberrant
type.
13, PARATILAPIA VENTRALIS, sp. n. (Plate IIT. fig. 3.)
Teeth very small, in 2 series in both jaws, the outer larger and tipped with brown.
Depth of body 22 to 3 times in total length, length of head 3, Snout with curved
upper profile, a little shorter than the eye, the diameter of which is 2 to 27 times
in length of head and exceeds interorbital width; maxillary extending to below
anterior fourth of eye; 2 or 3 series of scales on the cheek; large scales on the
opercle. Gill-rakers rather long, lanceolate, 17 to 19 on lower part of anterior arch.
Dorsal XII-XIII 13-14; spines increasing in length to the eighth or ninth, which
measures nearly 4 length of head and } or $ longest soft rays. Pectoral a little longer
than head. Ventral much produced, extending far beyond the origin of the anal,
especially in the males, in which it may reach the end of the caudal. Anal III 9-10;
third spine longest, about 2 length of head; middle soft rays produced, nearly as long
as head. Caudal deeply emarginate, crescentic. Caudal peduncle 1} as long as deep.
14 MK. G. A. BOULENGER ON A COLLECTION
on? lat. 1. —
complete or nearly so, usually reaching base of caudal. Grey-brown above, with or
without irregular darker spots, silvery below ; fins grey-brown, the ends of the ventral
filaments white.
Total length 92 millim.
8 specimens from Kinyamkolo, and one from Mbity Rocks.
‘This species is remarkable among its congeners from continental Africa for the low
number of its dorsal spines combined with the large eye, the crescentic caudal, the
extremely produced ventrals, and the upper lateral line extending on the caudal
peduncle.
Scales finely denticulate on the edge, 34-36 upper lateral line
14, PARATILAPIA FURCIFER, sp. n. (Plate IV. fig. 1.)
Teeth very small, in 3 series in both jaws, the outer largest and tipped with brown.
Depth of body equal to length of head, 3 times in total length. Snout with curved
upper profile, a little shorter than the eye, the diameter of which is 22 to 23 times in
length of head and slightly exceeds interorbital width ; maxillary extending to below
anterior border of eye; 2 or 3 series of scales on the cheek; large scales on the
opercle. Gill-rakers rather long, lanceolate, 15 or 16 on lower part of anterior arch.
Dorsal XIII 13-14; spines increasing in length to the ninth, which measures a little
less than } length of head; some of the soft rays produced, nearly as long as head.
Pectoral a little longer than head. Ventral much produced, extending nearly to
caudal. Anal III] 9; third spine longest, 3 length of head; middle soft rays
produced, like the dorsals. Caudal deeply emarginate, crescentic, the rays at the
angles produced. Caudal peduncle 14 as long as deep. Scales finely denticulate on
the edge, 60-63 on lat. 1. — upper lateral line nearly complete, extending on the
caudal peduncle. Bluish above, white beneath ; a few ill-defined yellow streaks along
the body; some yellow marblings on the postocular part of the head; fins white,
with some yellow streaks on the dorsal and anal and between the ventral and caudal
rays (according to a sketch by Mr. Moore).
Total length 110 millim.
‘lwo specimens from Kinyamkolo.
Closely allied to the preceding; distinguished by much smaller scales.
15. PaRATILAPIA LEPTOSOMA, sp. n. (Plate III. fig. 4.)
Teeth small, in 3 series in the upper jaw, in 4 in the lower, outer largest. Depth
of body 4 to 43 times in total length; length of head 3. Snout with straight upper
profile, as long as or a little longer than the eye, the diameter of which is 33 to 33
times in length of head, and equals interorbital width; maxillary extending to below
anterior border of eye or not quite so far; 2 series of scales on the cheek; opercle
OF FISHES FROM LAKE TANGANYIKA. 15
covered with scales. Gill-rakers long, slender, and close-set, 20 on lower part of
anterior arch. Dorsal XII 14-15; spines increasing in length to the last, which
measures § length of head, and is nearly as long as the soft rays. Pectoral # length
of head. Ventral reaching origin of anal. Anal III 10-12; third spine longest, a
little shorter than longest dorsal. Caudal feebly emarginate. Caudal peduncle twice
ee lat. 1. a
Brown, lighter beneath ; dorsal and anal with or without brown longitudinal streaks ;
caudal spotted with brown or black at the base.
Total length 87 millim.
Two specimens from Kinyamkolo, and two from Mbity Rocks.
Approaches P. longiceps Gthr. in the elongate body, but differs greatly in the
dentition, the shape of the snout, and the ftin-formula. Only the Malagassy species of
the genus Paratilapia were known to have fewer than 13 spines in the dorsal fin; but
one of the species described above (P. ventralis) has only 12 or 13.
as long as deep. Scales very finely denticulate on the edge, 59-40
BATHYBATES, g. n.
Body elongate ; scales cycloid, small and irregular. Several rows of large fang-like
teeth in the jaws. Maxillary exposed. Dorsal with 14 spines, anal with 3.
Vertebre 36 (17+19).
This genus is closely related to Paratilapia, with which it is to a certain extent
connected by P. longiceps Gthr., from Nyassa. However, the more formidable
dentition, coupled with the character of the scales on the body, appear sufficient to
warrant the establishment of a new genus. ‘The unique species appears in most
respects more specialized than Paratilapia longiceps, although more primitive than
any known species of that genus in the longer anal fin and the complete upper
lateral line.
16. BAaTHYBATES FEROX, sp. n. (Plate IV. fig. 2.)
Teeth long and sharp, fang-like, wide apart, in 4 series in the upper jaw, in 3 in
the lower. Depth of body 4 times in total length, length of head 3 times. Snout
long and strongly compressed, with convex upper profile; eye large, its diameter
12 times in length of snout, 34 in length of head, and a little greater than interorbital
width ; maxillary not quite reaching to below anterior border of eye; 5 series of small
scales on the cheek; large scales on the opercle. Gill-rakers moderately long and
slender, 13 on lower part of anterior arch. Dorsal XIV 16; spines rather feeble,
slender, subequal from the fifth, which measures 7 length of head; longest soft rays
4 length of head. Pectoral ? length of head. Ventral not reaching vent. Anal IT]
16; spines short and feeble. Caudal deeply forked, middle rays not hi lf as long as
outer. Caudal peduncle nearly twice as long as deep. Scales small and irregular,
16 MR. G. A. BOULENGER ON A COLLECTION
especially below the lateral lines, 78 oe lat. 1. E ; the upper extending from the opercle
to the caudal, the lower from below the last dorsal spines to the caudal. Pale bluish
green, iridescent above, white below; dorsal and anal bluish grey; ventrals and caudal
yellowish ; pectorals yellow ; two dark streaks on the dorsal.
Total length 275 millim,
A single specimen from Kinyamkolo, taken at a depth of 400 feet. The specimen
is a female with ripe ova; these are of large size, measuring 53 millim. in diameter.
The stomach contains a small, partially-digested fish of the genus Paratilapia, as
first ascertained by a sciagraph kindly prepared by Messrs. Gardiner and Green,
which has enabled me to compare the structure of the vertebral column with that of
other members of the family Cichlidw. The insertion of the ribs is typical, viz.,
sessile, except on the last three preecaudal vertebra ; the transverse processes are short
and in front of the ribs. ‘The number of vertebra agrees with that of the more elongate
species of Paratilapia.
ERETMODUS, g. n.
Body moderately elongate; scales ctenoid. Jaws with rather large spatulate teeth
with truncated crowns disposed in oblique transverse rows of two or three. Lips much
developed; maxillary entirely concealed when the mouth is closed. Dorsal with
23 to 25 spines; soft rays reduced to 3 to 5; anal with 3 spines. Vertebre 30
(15+15).
A strongly aberrant form, remarkable for its dentition and the structure of the dorsal,
formed almost entirely of spines.
17. ERermopus cyanostictus, sp. n. (Plate III. fig. 5.)
8 or 10 transverse series of teeth in each jaw, the crowns reddish brown. Depth of
body equal to length of head, 3 times in totallength. Profile of snout curved ; length
of snout 14 to twice diameter of eye, which is 43 to 5 times in length of head, and
a little less than interorbital width; mouth extending to below nostril; cheeks and
opercles naked. Gill-rakers short, 9 or 10 on lower part of anterior arch. Dorsal
XXIII-XXV 3-5; spines subequal from the sixth, $ length of head, a little shorter
than soft rays. Pectoral 3 length of head. Ventral reaching vent, Anal III 6-7;
third spine longest, a little longer than dorsals; soft rays about 3 length of head,
22-23
6-9 °
Caudal rounded. Caudal peduncle deeper than long. Scales 32-35 aos lat. 1.
Blackish brown, with scattered pale blue dots; belly yellowish.
Total length 75 millim.
Five specimens from Mbity Rocks, and three from Kinyamkolo.
The toothed portions of the premaxillary and mandible are much developed in depth,
in a manner suggestive of the Sparidw and Scaride, and the teeth are implanted in
sockets. ‘The deep triangular groove for the reception of the premaxillaries extends
OF FISHES FROM LAKE TANGANYIKA. Uf
to between the orbits, and the occipital crest is prolonged to it ; the parietal crests are
produced on the frontals ; the preeorbital is large, and the chain of suborbitals very
slender. ‘The pharyngeal teeth have long slender shafts and conical brown cusps. The
ribs are attached to the back of short transverse processes; all but the last one support
epipleurals ; the last four preecaudal vertebrae have a hemal bridge.
18. Tinapia LaBiaTa, sp. n. (Plate V. fig. 1.)
Outer teeth rather large, feebly notched ; inner teeth very small, tricuspid, in 3 or
4 series. Depth of body equal to length of head, 22 to 22 times in total length.
Snout with straight upper profile, 14 to 1} diameter of eye, which is 34 to 44 times in
length of head and equals interorbital width; maxillary not extending to below
anterior border of eye; 3 or 4series of scales on the cheek; large scales on the opercle ;
lips very strongly developed, both produced into a large triangular lobe in front. Gill-
rakers moderate, 15 on lower part of anterior arch. Dorsal XVIII 10; middle dorsal
spines longest, about ? length of head, and a little shorter than longest soft rays.
Pectoral 3 to length of head. Ventral reaching origin of anal. Anal III 6-7; third
spine longest, as long as longest dorsals, slightly shorter than longest soft rays.
Caudal truncate. Caudal peduncle slightly longer than deep. Scales finely
denticulate on the border, 33-35 ee lat. 1. aa Pale olive, with 10 more or less
distinct darker cross-bars; fins greyish-brown ; dorsal sometimes with oblique dark and
light streaks ; caudal with numerous round dark spots between the rays,
Total length 170 millim.
Four specimens from Kinyamkolo.
This species is easily recognizable by the extraordinary development of the lips,
which bears a curious resemblance to that observed in the Central American Heros
labiatus*. It appears to be nearest allied to Ctenochromis nuchisquamulatus Hilg.,
and C. sauvagii Pfeffer, from the Victoria Nyanza.
TROPHEUS, g. 0.
Body moderately elongate ; scales ctenoid. Jaws angularly bent at the sides, with
bands of minute tricuspid teeth, an outer row of bicuspid teeth, and enlarged conical
teeth at the sides of the premaxillary; mouth transversely linear when closed;
maxillary concealed under the preorbital. Dorsal with 21 spines, anal with 6.
Vertebre 33 (17-+16).
This genus differs from Zi/apia in the angularly-broken premaxilla armed with
enlarged conical teeth at the sides, and the higher number of anal spines.
The name proposed (zpo¢etc, one who rears, brings up, educates) refers to the
peculiar mode of nursing the eggs described hereafter.
1 Giinther, P. Z. 8. 1864, pl. iv. fig. 1.
voL. XV.— Part I. No. 3.—December, 1898. D
18 MR. G. A. BOULENGER ON A COLLECTION
19. TRopHEUS MooRI, sp. n. (Plate V. fig. 2.)
Teeth minute, those of the outer series tipped with brown and numbering about 50.
Depth of body 23 to 22 times in total length, length of head 34 to 33%. Snout
descending in a strong curve, as long as or a little longer than the diameter of the eye.
which is 34 to 4 times in length of head, and equals $ to ? interorbital width ; mouth
extending to below anterior border of eye; 4 series of scales on the cheek ; large scales
on the opercle. Gill-rakers short, 11 or 12 on lower part of anterior arch. Dorsal
XXI 5-6; spines increasing in length to the sixth, which measures not quite half
length of head; longest soft rays 2 to # length of head. Pectoral as long as head.
Ventral produced into a short filament, reaching beyond origin of anal. Anal VI 5-6:
spines increasing in length to the last, which slightly exceeds longest dorsal. Caudal
OOF)
slightly notched. Caudal peduncle as long as deep. Scales 30-32 as lets Il, Fe
Dark brown; a large bluish-white blotch on each side; belly reddish brown ; fins
blackish.
Total length 110 millim.
Five specimens from Kinyamkolo.
The occipital crest is very strong and the parietal crests are produced on the frontals.
The insertion of the ribs is as described in the preceding genera Telmatochromis ana
Eretmodus.
The mouth and pharynx of one of the specimens contains four eggs of very large
size, the vitelline sphere measuring 4 millimetres in diameter, with an embryo in an
advanced stage of development. ‘The egg of the Fifteen-spined Stickleback ( Gasterosteus
spinachia), hitherto regarded as the largest Teleostean egg in proportion to the size
of the animal, measures only 3 millimetres in diameter’. Besides the Siluroids of
the genera Arius and Galeichthys, which have very large eggs, at least two species
of Tilapia were known to give shelter to their eggs in the manner noticed above, viz..
T. simonis Gthr. (Chromis paterfanilias Lortet), as observed by Prof. Lortet in
Lake Tiberias, and 7’. nilotica Cuv., as noticed by me ina specimen collected by Canon
Tristram in the same lake. But these eggs, produced by fishes of the size of our
common Perch, are very numerous, and measure only about 2 millim. in diameter. It
has besides been observed in these Tilapia, as well as in the Siluroids, that the function
of protecting the eggs devolves on the male sex, while, to my surprise, the
Tanganyika fish proved on autopsy to be a female. Whether this is constantly so, or
whether either parent takes to the nursing duties, remains to be ascertained by
examination of a larger number of specimens. Iam all the more disposed to think
the latter possibility will be confirmed, from the fact thata specimen of Z%lapia nilotica
1 The absolutely largest known Teleostean egg is that of the Siluroid Arius commersonz, which measures 17
millim. in diameter. The size of the Arius-egg is much underrated by Giinther (‘Study of Fishes,’ p. 160) and
by McIntosh and Masterman (‘ British Marine Food Fishes,’ p. 12), who ascribe to it a maximum of 10 millim
OF FISHES FROM LAKE TANGANYIKA. 19
with the pharynx filled with embryos belongs to the female sex, while Dr. Lortet’s
observations on 7. simonis had led to the belief that specimens carrying eges in that
manner are invariably males.
It is here necessary to recall the observation contained in Livingstone’s ‘ Last
Journals,’ vol. ii. p. 17, that the “ Dagala” or “ Nsipé,” of Lake Tanganyika, a small
fish 2 or 3 inches long, and very like Whitebait, is said to emit eggs by the mouth.
The comparison of this fish to Whitebait excludes the possibility of the one here
described being the “Dagala” or “Nsipé,” which will probably prove to be a
Cyprinodont, if not actually the Haplochilus tanganicanus described below.
SIMOCHROMIS, g. n.
Body moderately elongate; scales ctenoid. Jaws with broad bands of minute
tricuspid teeth, with an outer series of larger bicuspid teeth, and a single series of
conical teeth at the sides of the pramaxillary; maxillary concealed under the pre-
orbital. Dorsal with 17 or 18 spines, anal with 8. Vertebree 31-32 (15+16-17).
The type species differs from the other members of the genus to which it was
originally referred in the differentiation of the teeth at the sides of the pramaxillary,
a character in which it agrees with the genus Zropheus.
20. SIMOCHROMIS DIAGRAMMA Gthr.
Chromis diagramma Ginth. Proc. Zool. Soc. 1893, p. 632, pl. lviii. fig. 3.
The bands of teeth in both jaws consisting of 8 to 10 series, gradually decreasing in
size from the outer border. Depth of body 25 to 2% times in total length, length of
head 34 to 85. Snout descending in a strong curve ; diameter of eye 3 to 34 times in
length of head, 14 in interorbital width ; mouth extending to below anterior border of
eye; 4 series of scales on the cheek ; large scales on the opercle. Gill-rakers short,
12 or 13 on lower part of anterior arch. Dorsal XVII—XVIII 9-10; spines increasing
in length to the sixth, which measures } to ¢ length of head; longest soft rays about 3
length of head. Pectoral as long as head. Ventral reaching vent or origin of anal.
Anal III 7-8; third spine longest, as long as longest dorsal. Caudal truncate. Caudal
peduncle as long as deep. Scales 31-33 z; lat. 1. a Olive, whitish beneath ;
young with 9 rather indistinct vertical bars ; gill-membrane sometimes with dark brown
spots; fins greyish; a dark brown stripe may be present along the spinous dorsal,
which has a black edge.
Total length 105 millim.
Four specimens from Kinyamkolo.
20 MR. G. A. BOULENGER ON A COLLECTION
PETROCHROMIS, g. n.
Body moderately elongate; scales ctenoid. Jaws with very broad bands of minute,
bristle-like teeth, with incurved bi- or tricuspid crowns; maxillary concealed under the
preorbital. Dorsal with 17 or 18 spines, anal with 3. Vertebree 32 (16416).
Distinguished from Tilapia by the very broad bands of minute teeth with incurved
crowns, from Simochromis by the absence of conical teeth in the preemaxillary.
21. PETROCHROMIS POLYODON, sp. n. (Plate VI. fig. 1.)
Crowns of teeth brown. Depth of body 21 to 22 times in total length, length of head
93 to 3 times. Snout with convex upper profile, 14 to 1% diameter of eye, which is 4
to 41 times in length of head and 14 in interorbital width; mouth hardly extending
to below anterior border of eye; 4 or 5 series of scales on the cheek ; large scales on
the opercle. Gill-rakers very short, 12 or 13 on lower part of anterior arch. Dorsal
XVII-XVIII 8-9 ; spines increasing in length to the sixth or seventh, which measures
about 2 length of head; longest soft rays $ to 3 length of head. Pectoral nearly as
long as head. Ventral reaching vent or origin of anal. Anal III 7-8 ; third spine
longest, as long as longest dorsal. Caudal truncate. Caudal peduncle as long as deep.
3-4 29-94 : as ‘
Scales 32-34 s543; lat. 1. {34,. Olive-brown, whitish beneath ; fins grey or blackish.
Total length 185 millim.
Two specimens from Kinyamkolo, and two from Mbity Rocks.
The premaxillary and mandibular bones are very massive, and the maxillary is much
reduced in size; the ascending processes of the premaxillaries extend to between the
anterior borders of the orbits and are received in a deep excavation, to which the strong
occipital crest extends; the parietal crests are produced forwards as far as the frontals ;
the preorbital is large, and the chain of suborbitals very slender. Only the first rib is
absolutely sessile, the following being attached to the back of the transverse processes
at a short distance from the centre ; the epipleurals extend to the twelfth rib; the last
two preecaudal vertebree form a hemal bridge.
PERISSODUS, g. N.
Body elongate; scales cycloid. ‘Teeth rather large, unequal in size, few, with swollen
bases and low slightly-notched crowns, compressed transversely to the axis of the jaws,
disposed in a single series; maxillary exposed. Dorsal with 18 spines, anal with 3.
Vertebre 35 (17+ 18).
The extraordinary dentition which characterizes this new genus may be looked upon
as an extreme specialization of that exhibited by Ztlapia, a specialization in an
opposite direction from that attained by Petrochromis.
OF FISHES FROM LAKE TANGANYIKA. a1
22. PERISSODUS MICROLEPIS, sp.n. (Plate VI. fig. 2.)
10 teeth on each side of the premaxillary, 9 on each side of the mandible. Depth
of body 3% times in total length, length of head 31. Snout a little longer than
diameter of eye, which is 4 times in length of head, and almost equals interorbital
width ; lower jaw projecting; maxillary extending to below anterior border of eye;
3 series of scales on the cheek ; large scales on the opercle. Gill-rakers rather long,
14 on lower part of anterior arch. Dorsal XVIII 10; spines increasing in length to
the last, which measures } length of head; longest soft rays 3 length of head.
Pectoral # length of head. Ventral not reaching vent. Anal III 8; third spine
longest, nearly as long as longest dorsal ; longest soft rays half length of head. Caudal
truncate. Caudal peduncle 13 as long as deep. Scales 65 a8 lat. 1. a Uniform
dark reddish brown ; a blackish opercular spot.
Total length 100 millim.
A single specimen from Mbity Rocks’.
1 T think it desirable to allude here briefly to two remarkable genera of Cichlidw from Lake Tanganyika,
which I have founded on specimens obtained by Capt. Descamps and sent to me for study by the
Administration of the Congo Free State. The specimens are in so bad a condition that it will never be
possible to draw up from them complete definitions of the species, but their dentitions, which indicate
remarkable new genera, are well preserved, and I am sure the annexed figures together with the short notes
here given will be sufficient to ensure their future recognition :—
Ecropus, g. n.—Teeth very small, conical, in two series in both jaws, the outer larger; outer mandibular
teeth pointing outward, perpendicular to the others; maxillary concealed under the preorbital when the
mouth is closed. Scales rather large, ctenoid. Dorsal with 14 spines, anal with 3.
Ectodus descampsi, sp.n. (Fig. 1, a.)\—D. XIV 14; A. IIL 8. Depth of body 34 times in total length,
length of head 3. Eye very large, 24 in length of head, 12 interorbital width. Two series of scales on the
cheek. Sc. 84 4; lat. 1.2. Total length 95 millim.
Ectodus melanogenys, sp.n. (Fig. 1,0.)—D. XIV 16; A. III 13, Depth of body 5 times in total length,
Fig. 1.
+
(A
ZF A
ll Zz
a Gis b,
length of head 34. Eye 4 times in length of head, 14 in length of snout, 14 interorbital width, Three series
of scales on the cheek. About 35 scales in a longitudinal series. A black spot on the chin, Total length
95 millim.
bo
bo
MR. G. A. BOULENGER GN A COLLECTION
MASTACEMBELIDA.
23. MASTACEMBELUS MOORII, sp.n. (Plate VII. fig. 1.)
Depth of body 14 times in total length, length of head (without rostral appendage)
3 times; vent equally distant from end of snout and base of caudal; length of
head 24 to 3 times in its distance from vent, and 4 in its distance from first dorsal
spine. Snout 3 times as long as eye, ending in a trifid dermal appendage which is a
little longer than eye; cleft of mouth extending to below centre of eye; no pre-
opercular spines. Vertical fins united with the rounded caudal. Dorsal XX V-XXVII
70-80 ; spines very short. Anal II 70-80. Pectoral + length of head. Scales very
small, 30-35 between origin of soft dorsal and lateral line. Brown, tail with a
wide-meshed blackish network; dorsal and anal whitish, with a vertical series of
blackish spots or vertical bars; anal and caudal edged with blackish.
Total length 440 millim.
Two specimens from Mbity Rocks.
This species has been compared with J/. marmoratus, Perugia, from the Congo,
which differs in the more slender body (its depth contained 20 times in the total
length) with larger scales, the shorter tail, the longer rostral appendage (measuring
twice the diameter of the eye), and the higher number (30) of dorsal spines.
I think as many as 13 African species of Mastacembelus may be distinguished, as
shown in the following synopsis :—
PLEcoDUs, g. n.——Teeth large and few, in a single series, dilated at the base, truncated at the end, compressed,
slightly grooved in front, curved and directed backwards; 14 teeth in the upper jaw, 12 in the lower;
maxillary exposed. Scales moderate, cycloid. Dorsal with 19 spines, anal with 3.
Plecodus paradowus, sp.n. (Fig, 2..—D. XIX 14; anal III 12. Depth of body 4 times in total length,
length of head 41. Eye large, longer than snout, 4 length of head, 17 interorbital width ; maxillary extending
5 lat. 1. °. Total length 90 millim.
to below anterior third of eye; 3 series of scales on the cheek. Sc. 65 7;
OF FISHES FROM LAKE TANGANYIKA, 23
I. Vent nearly equally distant from the end of the snout and the base of the caudal fin.
A. Two strong preopercular spines.
D. XXIH-XXIV 75; A. Il 75-76; length of head (measured to extremity of opercle and without
rostral appendage) hardly twice in its distance from vent AS 6 times as great as its
distance from the first dorsal spe . . . oe a elVarmanchi s alive
D. XXIV 100; A. II 100; length of head 34 reat in its distance ro vent, equal to its distance
from the first dorsal spine. . . See - . . 2. M. eryptacanthus Gthr.
D. XXVII85; A. III 92; length of Tiel On in its Fivcnes from vent, 3 times as great as its
distance from first dorsal spine . . . . =... =... 3. M. congicus Blgr."
D. XXVI-XXX 80; A.TI180... . Bo 6 0 0 0 0 Sy WE inooaaGdS Wiksee
D. XXIX-XXX 117-130; A. II 117- 17: length of head nearly 4 times in its distance from
vent, and nearly twice as great as its distance from the first dorsal spine.
5. M. loennbergii Blgr.’
B. Preopercular spines minute (young) or absent.
D. XXVII-XXVIII 70-80 ; A. IT 80-90; length of head 8 to 4 times in its distance from vent,
equal to or a little less than its distance from the first dorsal spine.
6. M. flavomarginatus Blgy.’
D. XXVII 80-82; A. II 60-65 ; length of head equal to its distance from the first dorsal spine.
7. M. niger Sauv.
D. XXVII-XXIX 65-70; A. II 68-75 ; length of head 24 to 3 times in its distance from the
vent, and 4 in its distance from the first dorsal spine. . . 8. MZ. shiranus Gthr.
D, XXIX 95; A. II 85; length of head 3 times inits distance from the vent, aud 4 in its distance
' from the first dorsal spme. . . . . . 9. M. nigromarginatus Blgr.’
D. XX V-XXVII 70-80; A. II 70-80 ; see of heed twice in its distance from the vent, and
nearly 4 in its distance from the fir st dorsal spme. . . . 10. ML moorii Bler.
II. Vent much nearer the caudal fin than the end of the snout; one or two preopercular
spines.
ID), 2OOS C3 AG INGO oo 6 68 5 4 685 6 og 6 JU Whoa Eee
D. XXXITI-XXXV 50-60; A.IT50-60. . . . .. . . 12. MU. tanganice Gthr.
III. Vent much nearer the end of the snout than the caudal fin.
1D, SOOM ROOOIL NOs AG INNG 5 6 2 6 6 6 eo 5 5 co Meh WE Caen Cisne
‘ A species from the Upper Congo founded on a single specimen with mutilated tail and regenerated caudal
fin. I have now a perfect specimen.
* M. marchii Steind. nec Sauy.—Liberia.
* M. cryptacanthus Loennberg nec Gthr.—Cameroon.
+ A new species founded on specimens from the Gaboon previously referred to WM. cryptacanthus.
5 A new species founded on a single specimen from Ashantee.
° Type examined.—Vent equally distant from the head and from the caudal; head once and a halt as toug
as its distance from the first dorsal spine; 17 scales between origin of soft dorsal and lateral line,
24 MR. G. A. BOULENGER ON A COLLECTION
SILURID.
24. CLARIAS ANGUILLARIS L.
This species occurs in the Nile as well as in the Niger and the Ogowe. It is one
of those Siluroids which are known to live a considerable time out of water and to
make excursions on dry land after the manner of Hels.
25. CLARIAS LIOCEPHALUS, sp.n. (Plate VII. fig. 2.)
Vomerine teeth in a narrow band, without posterior process. Depth of body
52 times in total length, length of head 5 times. Head smooth, covered with sott
skin, slightly longer than broad; occipital process very short, angular; diameter of
eye 3 times in length of snout, 6 times in interorbital width; maxillary barbel as long
as the head, nasal barbel a little shorter; inner mandibular barbel 2 length of head.
Dorsal 70. Anal 50. Caudal free. Pectoral $ length of head, not extending to the
vertical of origin of dorsal fin. Uniform blackish brown.
Total length 80 millim.
A single specimen from Kinyamkolo.
96. ANOPLOPTERUS PLATYCHTR Gthr.
Two specimens from marshes near Mbity.
I have recently recorded the occurrence of this species in Lake Nyassa, and it has
also been described by Vaillant as Chimarrhoglanis leroy, from Mrogoro torrent,
Urugaru Mountains, East Africa.
97. AUCHENASPIS BIScUTATA Geoffr.
Known from the Nile and the Senegal.
28. SYNODONTIS MULTIPUNCTATUS, sp.n. (Plate VIII.)
Premaxillary teeth in 5 or 6 irregular transverse series; mandibular teeth in a
single series of 16, feebly curved, simple, measuring hardly 3 diameter of eye.
Depth of body 34 times in total length, length of head 33. Head scarcely longer
than broad, slightly convex on the crown; snout rounded, less than half length
of head, twice as long as eye; eye supero-lateral, well visible from above, its
diameter 44 times in length of head, twice in interorbital width. Gill-cleft very
narrow, not extending below base of pectoral. Maxillary barbel simple, reaching a
little beyond anterior third of pectoral spine; mandibular barbels strongly fringed,
outer a little more than twice as long as inner, and half as long as maxillaries.
Dorsal II 7; spine strong, a little shorter than the head, strongly serrated behind in
its distal half. Adipose fin low, a little shorter than the head, twice as long as its
distance from the dorsal. Humeral process simply granulate, sharply pointed, not
OF FISHES FROM LAKE TANGANYIKA.
bo
n
extending quite so far as the occipito-nuchal shield, which is 14 as long as broad and
reaches the first soft ray of the dorsal. Anal III 7. Caudal deeply bifurcate. Skin
smooth. Pale reddish brown above, with very numerous blackish-brown round spots,
which are smallest on the head and nape; lower parts and barbels white, unspetted ;
ventrals yellow ; dorsal and caudal broadly edged with yellow.
Total length 240 millim.
A single specimen from Sumbu.
29, MALAPTERURUS ELECTRICUS Gin.
The Electric Cat-fish is widely distributed, being on record from the Nile, the
Niger, the Ogowe, and the Congo.
CYPRINIDS.
30. LaABEo, sp. inc.
An unmistakable figure of a Zabeo is among Mr. Moore’s sketches, and although
the specimen is not represented in the collection entrusted to me for description, I
have included the fish as being the only representative of the Cyprinoids yet found
in Tanganyika. The specimen from which the drawing was made was of large size,
and therefore could not be preserved under the difficult circumstances to which
allusion has been made in the introduction to this memoir.
CHARACINIDA,
31. ALESTES MACROLEPIDOTUS C. & V.
Occurs in the Nile, the Senegal, the Niger, and the Congo.
32. ALESTES MACROPHTHALMUS Gthr.
This fish was known only from the Ogowe.
33. Hyprocyon FORSKALI Cuy.
Occurs in the Nile, the Senegal, and the Niger.
CYPRINODONTID4.
34. HAPLOCHILUS TANGANICANUS, sp.n. (Plate VI. fig. 3.)
Body compressed, its depth 4 times in total length; length of head 4% times.
Snout depressed ; lower jaw projecting beyond the upper; eye a little longer than
snout, a little shorter than interorbital width, 3 times in length of head. Dorsal 13,
originating at equal distance from the head and the caudal fin; the first ray corresponds
to the 18th scale of the lateral line; posterior rays longest, § length of head. Anal 28,
ariginating below extremity of pectoral. Pectoral $ length of head, extending far
beyond root of ventral, Caudal feebly emarginate. Caudal peduncle twice as long as
VoL. Xv.— PART I. No. 4.—December, 1898. E
26 MR. G. A. BOULENGER ON A COLLECTION
deep. Scales 42 in a longitudinal line, 11 in a transverse line. Silvery, with a darker
lateral stripe; dorsal and anal with greyish horizontal streaks.
Total length 80 millim.
A single specimen from Mbity Rocks.
PoLYPTERIDA.
35. PoLyprerus BICHIR Geoffr. (2).
Mr. Moore informs me that a Polypterus occurs in moderate abundance in the lake,
and although he did not bring home specimens, he feels tolerably confident, from his
recollection of them, that the fish belongs to the typical P. dichir, which occurs in
the Nile, the Niger, and the Congo, and not to any of the closely-allied species that are
often confounded with it.
APPENDIX. By J. E. S. Moors.
In the foregoing description of the fishes which I succeeded in bringing through
from Tanganyika to the coast, Mr. Boulenger has already alluded to the difficulties
that were experienced in transporting them in spirit, through several hundred miles
of often trackless, always scorching, forest, and of the inevitable losses which this
entailed. ‘The difficulties of transport, however, were by no means all. It must be
nearly impossible for anyone who has not visited the African lakes to realize their
huge size and oceanic character.
One must be as heavily equipped for dredging in these waters as would be required
for effective operations in the open sea. It will easily be understood, therefore, how
incomplete our knowledge of the deep-water fauna of these lakes must be considered,
when it is remembered that on Tanganyika I was of necessity forced to work with
native dug-out boats, and with nothing better than the natives themselves as motive
power.
In the case of the fishes, moreover, there are no sources of collateral evidence from
which we may obtain any insight into the nature of the deep-water forms, for dead
fishes, unlike molluscs, leave no shells behind them, to be thrown upon the beaches of
the lake, whereby, in the case of the molluscs, we gain some knowledge of the
existence of forms which have not been seen alive; nor can we make use of the
knowledge of the inhabitants in this matter, for the best of the Tanganyika natives
are but wretched fishermen, merely using either surface traps, or light and inefficient
drag-nets, which are thrown out a short distance from the shore and then hauled in
to the land. Such nets are necessarily used only on smooth sandy beaches, and
consequently the fishes caught in them are only of those s; ecies which inhabit places
OF FISHES FROM LAKE TANGANYIKA. 27
of this description, and although they are numerically abundant, they belong to half
a dozen species at the most.
Nearly all the new forms which I obtained were killed by dynamite from the
craggy ledges of the west coast of the lake, where the water was deep enough, about
20-35 feet, but not too deep for my men to dive and procure the greater number of the
fishes, which, after every shot, were invariably found on the bottom. The number
of fishes in such situations is really surprising, and on several occasions, after firing a
single cartridge, I obtained more than two tall negroes could well carry, when slung
in a bag between them on a pole.
Only one fish was obtained from anything approaching to the deeper water of
Tanganyika, about 400 feet, which was accidentally entangled in a basket dredge,
and this was the specimen of the new species Bathybates feroa described on p. 15.
Of the fishes which exist on the great mud-plains, which are to be found about
1000 feet below the surface in many portions of the lake, or of those which may occur
in the vastly deeper portions, absolutely nothing is known.
The fishes which have been collected, moreover, were obtained almost exclusively
from the south-western extremity of Tanganyika, and to suppose that the fishes which
occur in this locality are in any way representative of the fishes, even in similar
situations, throughout the whole lake, would be as absurd as if we were to suppose
that the shoal-water fish of the whole English Channel could be obtained from a few
miles of rocky beach about the Needles and the Isle of Wight. ‘That the fishes differ
in different portions of the lake I have myself observed, and it has also been shown
in a most striking manner by the fact that of the six species previously described
from ‘Tanganyika, which were collected by Captain Hore near Ujiji, I have only re-
discovered one.
The fishes which have been already obtained in ‘Tanganyika, or in Nyassa for that
matter, must therefore be regarded as merely the surface skimmings of these lakes,
and as in no sense representing the whole fish-fanna they may contain.
Really deep water, 300 fathoms and upwards, is to be fonnd over large areas in
Nyassa, and to what depth the water in Tanganyika may reach no one knows; but
it is obvious that where water of such depth exists, if the lakes in which it is con-
tained have been permanent for any great length of time, there ought to be modified
forms of the freshwater fishes to suit such depths, and, with the exception of Bathybates
ferox, these have not yet been obtained.
In the case of Tanganyika, the interest which attaches to the future exploration of
its deeper water is particularly great, since I have elsewhere shown that the majority
of the Halolimnic animals, most of the molluscs, sponges, and so forth, are exclusively
deep-water forms. In fact, it was only when my dredging capacities, so to speak,
were giving out, that the more interesting of these forms, the genera T'yphobia,
Bathanalia, and the like, were beginning to come in.
28 MR. G. A. BOULENGER ON A COLLECTION
That there are no Halolimnic representatives among the fishes which have hitherto
been obtained is no evidence that other fishes of a widely different and possibly of a
Halolimnic type may not in future be secured. On the other hand, the fact that the
Teleostean fishes now existing in Lake Tanganyika should not correspond with the
Molluscan section of the Halolimnic group is really what one would expect; for, as I
have recently shown ', the facies of the Molluscan section of the Halolimnic group is
almost, if not quite, indistinguishable from that of the Jurassic seas. Except the
Herring-like Leptolepide, few, if any, Teleostean fishes are represented in Jurassic beds,
and we should therefore expect the piscine accompaniments of the Halolimnic molluscs
to be entirely composed of Ganoids and the like. J found a species of Polypterus,
which I took to be P. dichir, abundant on the southern shores of Lake Tanganyika,
and it is quite possible that some of the active carnivorous fishes which inhabit the
open water may be Ganoids too. What the fish is that so much surprised Glaive,
when he crossed the lake, by attacking the paddles of his boat, is quite unknown, but
I myself saw these same fishes attack the paddles of my own boat, not 20 miles from
the spot where Glaive described them, on the west coast of the lake.
Judging, therefore, from the incomplete character of our knowledge of the fish-
fauna of Lake ‘Tanganyika, and from the vast antiquity of the lake, as evidenced by
the Jurassic facies of its molluscan shells, it is only natural to expect that future
exploration may reveal, among these fishes, forms that are of the highest scientific
interest from a morphological point of view. I mean that Tanganyika and its
neighbourhood present one of those few localities where it is legitimate to expect that
we may discover many forms that in most places have become extinct.
Our very slight acquaintance with the surface-forms existing in Nyassa and ‘Tan-
ganyika, and the complete absence of all knowledge of the contents of the deep
waters of these lakes, unfortunately by no means exhausts the sum of our present
ignorance of these matters. We have no real knowledge of the extent of the
Halolimnic fauna, beyond the lake in which it was originally discovered. I have
shown only that this fauna is not present in Nyassa nor in Shirwa, nor yet even in the
little lake Kela, not more than 20 miles from the Tanganyika shore, and that it
is extremely probable that it does not extend to Mwero and Lake Bangweolo to
the west. But for anything that can actually be shown to the contrary, it may be
present in Lake Rukwa to the east, and in the Albert and Albert Edward Nyanzas
to the north, and it is perhaps almost to be expected that some representatives
of this stock should be found in Lake Kivu, which lies in the same great valley not
more than 50 miles to the north of Tanganyika. But what is the relation of this
lake, the effluent of which flows south into Tanganyika, to the Albert and Albert
Edward Nyanzas, with effluents flowing into the Nile, towards the north? All that
‘ “ On the Hypothesis that Lake Tanganyika represents an old Jurassic Sea,’ Quart. Journ, Microse, Sci.
xli. no. 162, June 1898.
OF FISHES FROM LAKE TANGANYIKA, 29
we know at present is that the watershed between these lakes is situated in their
common valley, and that the whole series of lakes, from Tanganyika in the south to the
Albert Edward in the north, appears to lie in one continuous valley-system, which owes
its origin to a single geological movement, that has affected, although it may be
at very different times, an immense area of ground, and reaches at the present time
actually to the shores of the Red Sea.
The lakes which lie in these valleys immediately to the north of Tanganyika are
therefore of especial zoological interest, for it may be that in these isolated sheets of
water there exist animals analogous to those which constitute the Jurassic Halolimnic
fauna of Tanganyika itself. The similarity of the Halolimnic gastropods to those
of the Jurassic seas is, as I have insisted elsewhere, so complete and so remarkable
that no effort should be spared which may throw light on the real nature and origin
of these forms.
The only geographical fact which is at present available is this, that there exists a
certain similarity in the fish-fauna of Tanganyika to that of the lower Nile, and it
may be that this is an indication that at some time the Nile watershed was further
south, and drained out of Tanganyika to the north, in which case Halolimnic animals
will without doubt be found among the lakes towards the north. All this is, however,
little more than mere conjecture, but it will help my present purpose if it renders
more apparent the extreme desirability of becoming better acquainted with the zoology
of the lakes in the Rift valleys immediately to the north of Tanganyika.
The presence of a Halolimnic fauna in Lake Tanganyika, as something superadded
to the normal freshwater stock of the lake, is at present the central fact, and if the
expectations which the morphological characters of those Halolimnic animals already
known have raised should be fulfilled, we have something here which may completely
change our views respecting the past history of the continent, and provide morphology
with some most unique and valuable comparative material.
EXPLANATION OF THE PLATES.
PLATE 1.
Fig. 1. Lates microlepis, p. 6.
Fig. 2. Lamprologus fasciatus, p. 7.
Fig. 3. compressiceps, p. 7.
Fig. 4. 5 moori, p. 8.
Fig. 5. Be modestus, p. 8.
Fig. 6. _ elongatus, p. 9.
PLATE IL.
Fig. 1. Lamprologus furcifer, p. 9.
Fig. 2. Telmatochromis vittatus, p. 10.
VoL. XV.—PaRtT 1. Nu. 5.—December, 1898. P
ON A COLLECTION OF FISHES FROM LAKE TANGANYIKA.
Fig. 8. Telmatochromis temporalis, p. 11.
Fig. 34. Ne es dentition.
Fig. 3 0. Fe MS skeleton.
Fig. 4. Julidochromis ornatus, p. 12.
Fig. 44. . oF dentition.
Fig. 44. 2" a skeleton.
PAE lle
Fig. 1. Paratilapia pfeffert, p. 12.
Fig. 2. a macrops, p. 13.
Fig. 3. Ba ventralis, p. 15.
Fig. 4. im leptosoma, p. 14.
Fig. 5. Hretmodus cyanostictus, p. 16.
Fig. da. of ~ dentition.
Fig. 5 2. es a skeleton.
PLATE IV.
Fig. 1. Paratilapia furcifer, p. 14.
Fig. 2. Bathybates feroa, p. 15.
Fig. 2a. o ee centition.
PLATE V.
Fig. 1. Tilapia labiata, p. 17.
Fig. 2. Tropheus moorit, p. 18.
IB Wis > » dentition.
1B Boop ,, Skeleton.
PLATE VI.
Fig. 1. Petrochromis polyodon, p. 20.
Fig. La. die 45 dentition.
ies 170: es = skeleton.
Fig. 2. Perissodus microlepis, p. 21.
Joie, BOS 5 es dentition.
Fig. 3. Hapiochilus tanganicanus, p. 25.
PLATE VII.
Fig. 1. Mastacembelus moorit, p. 22.
Fig
to)
Fig. 2a.
+9
39
2. Clarias liocephalus, p. 24.
upper view of head and nape.
PLATE VIII.
Magnified (x3).
Synodontis multipunctatus, p. 24, with lower view of head.
Sram, 00l. Soe Vel NV GCL.
J.Green del. et hth. Mintern Bros.imp
LATE S MICROLEPIS- 2.LAMPROLOGUS FASCIATUS. 38.LAMPROLOGUS COMPRESSICEPS.
4 LAMPROLOGUS MOORII . 5.LAMPROLOGUS MODESTUS. 6. LAMPROLOGUS ELONGATUS.
Trans. Loo. Soe NOAV ICH |
3. 3b.
WN Sal [;
AR
RA
A BN
SAR oe ip /
Dee
Aa.
ba.
NS,
Lh.
Mintern Bros im
1.LAMPROLOGUS FURCIFER. 2.TELMATOCHROMIS VITTATUS.
3.TELMATOCHROMIS TEMPORALIS. 4. JULIDOCHROMIS ORNATUS.
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CONTENTS.
I. Report on the Collection of Fishes made by Mr. J. E. 8. Moore in Lake Tanganycha
during his Expedition, 1895-96. By G. A. Bounenesr, F.RS., F.ZS. With
an Appendix by J. B.S. Moore, A.R.C.S. (Plates I-VI.) . . . page 1
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Il. On the Marine Copepoda of New Zealand.
By G. Stewarpson Brapy, W_D., D.Sc., LL.D., F.R.S., CM.ZS.
Received February 9, 1899, read March 21, 1899.
[Puates IX.—XIII.]
IN this paper I complete, so far as possible, an account of two collections of New
Zealand Entomostraca submitted to me by Mr. G. M. Thomson of Dunedin, and
Drs. Meinert and H. J. Hansen of Copenhagen. A former contribution (already
published in the ‘ Transactions ’ of this Society, vol. xiv. p. 429) deals with the Ostracoda;
the present paper refers to the Copepoda. It is rather remarkable that no Cladocera
were noticed in any of the numerous gatherings represented in these collections, and
it is also unfortunate that of some of the most interesting species only one or two
imperfect examples were found,—in some cases so extremely imperfect that I have
made no attempt to describe or figure them. In some other cases of specimens
preserved in picric acid, the opacity and extreme brittleness of the subjects made
dissection very difficult, and recognition of anything beyond the coarser external
features quite impossible. The collections were made for the most part by means of
the tow-net at the surface and down to a depth of 6—8 fathoms; a few were from
littoral alge. Not much systematic investigation of the marine microzoa of New
Zealand appears hitherto to have been made; for the work already done we are
indebted chiefly to Mr. G. M. Thomson, who has published papers on the Ostracoda
and Copepoda in the ‘Transactions’ of the New Zealand Institute. There can be no
doubt that a rich harvest awaits any observer who may be able to devote time and
labour to the task.
The memoirs quoted in this paper are as follows. They are indicated in the
synonymy by the numbers assigned to them in the list :—
Borcg, Axet. (1).— Oversigt over de ved Norges Kyster iagttagne Copepoder henhérende til
Calanidernes, Cyclopidernes og Harpactidernes Familier. (Vidensk.-Selskabs Forhandlinger,
1864.)
Brapy, G. S. (1).—A Monograph of the free and semi-parasitic Copepoda of the British Islands.
(Ray Society, 1878-80.)
(2).—Report on the Copepoda collected by H.M.S. ‘Challenger’ during the years
1873-76. (1883.)
(3).—Description of a new Species of Cyclops. (Sixth Annual Report of the
Fishery Board for Scotland.) 1888.
(4).—A Revision of the British Species of Freshwater Cyclopide and Calanide.
(Natural History Transactions of Northumberland, Durham, and Newcastle-upon-Tyne,
vol. xi.) 1891.
VoL. xv.— Part ul. No. 1.—August, 1899. G
52 DR. G. 8S. BRADY ON THE
Craus, C. (1).—Die frei lebenden Copepoden, mit besonderer Berticksichtigung der Fauna
Deutschlands, der Nordsee und des Mittelmeeres. Leipzig, 1863.
Dana, J. D. (1)—American Journal of Science, vol. viii. (1849).
(2).—Crustacea of the United States Exploring Expedition (1852).
Girssrecut, W. (1).—Systematik und Faunistik der pelagischen Copepoden des Golfes von Neapel
und der angrenzenden Meeres-abschnitte. Berlin, 1892.
Kramer, A. (1).—On the most frequent Pelagic Copepoda and Cladocera of the Hauraki Gulf.
(Transactions and Proceedings of the New Zealand Institute, 1894, vol. xxvii.)
Lussock, J. (1).—On two new Subgenera of Calanide. (Annals & Magazine of Natural History,
March, 1853.)
Scorr, THomas (1t).—Report on Fntomostraca from the Gulf of Guinea, collected by
John Rattray, B.Sc. (Transactions of the Linnean Society of London, 2nd ser. Zool.
vol. vi. January, 1894.)
Tuomson, G. M. (1).—On the New Zealand Copepoda. (Transactions and Proceedings of the
New Zealand Institute, vol. xv.) 1882.
Section I. GNATHOSTOMA ‘Yhorell.
Family CALANID.
Genus Cauanus Leach.
1. CaLaNus FINMARCHICUS (Giinner).
Taken in the surface-net, Otago Harbour.
Genus Paracauanus Boeck.
1. ParacaLanus parvus (Claus). (Plate IX. figs. 1, 2.)
In the surface-net, Otago Harbour and Port Chalmers; Hauraki Gulf (Dr. Kramer).
Genus CLAUSOCALANUS Giesbrecht.
1. CLAUSOCALANUS ARCUICORNIS (Dana).
1849. Calanus arcuicornis Dana (1).
1852. Calanus arcuicornis Dana (2), p. 1056, pl. Ixxii. figs. 7 a, 5.
1892. Clausocalanus arcuicornis Giesbrecht (1), pp. 185, 1938, pl. i. fig. 14, pl. x. figs. 1-19,
pl. xxxvi. figs. 29-31.
1894. Clausocalanus arcuicornis Scott (1), p. 78, pl. vill. figs. 88-47.
Taken in the surface-net, Port Chalmers. Recorded also by Dr. Kramer from the
Hauraki Gulf.
This species has been sufficiently figured by Giesbrecht and Scott (oe. cit.).
2, CLAUSOCALANUS FuRCATUS (G. S. Brady). (Plate IX. figs. 3-7.)
1883. Drepanopus furcatus G. S. Brady (2), p. 77, pl. iv. figs. 1, 2, pl. xxiv. figs. 12-15.
1892. Clausocalanus furcatus Giesbrecht (1), p. 194, pl. xxxvi. figs. 32, 33, 35.
MARINE COPEPODA OF NEW ZEALAND. 33
The genus Drepanopus which, as proposed by the present writer, included two
species, D. furcatus and D. pectinatus, has been divided by Dr. Giesbrecht into
two genera—Drepanopus and Clausocalanus, to the latter of which D. furcatus is
transferred. The original description dealt with females only, no males having been
observed in the ‘Challenger’ collections. I therefore give here figures of some of the
more important diagnostic structures. In some of the New Zealand gatherings both
of these species occur together, and when associated with C. areuicornis and Paracalanus
parvus in various stages of growth it becomes by no means an easy matter to
separate them. The most conspicuous characters are to be found in the fusions of the
antennal joints, the relative lengths of the abdominal segments, and in the structure
of the fifth pair of feet in both sexes.
The anterior antenne of the female in C. furcatus reach somewhat beyond the
extremity of the cephalothorax and are 24-jointed, having all the joints perfectly
distinct; in the male (tig. 5) the first and second joints are coalescent, so also are the
eighth, ninth, and tenth ; all the rest are distinct. ‘The fifth pair of feet in the female
(fig. 4) are very short, two-jointed and ‘alike on both sides ; in the male (fig. 7) the foot
of the right side is five-jointed, and about half as long as the abdomen (considerably
shorter than in C. arcuicornis), that of the left side is two-jointed and only as long as
the first joint of the right foot. The abdomen is four-jointed in both sexes (figs. 3, 7),
the first segment being very short, the remaining segments nearly equal in the
female, while in the male the third is rather reduced and the fourth rather increased
in length.
Hab. Taken plentifully in the surface-net in Otago Harbour and at Port Chalmers,
and in the net at 7 fathoms off Gisborne.
Genus AcaRrTIA Dana.
1. ACARTIA ENSIFERA, sp. nov. (Plate IX. figs. 8-15.)
Female.—Seen from above (fig. §) the cephalothorax is elongated, slightly narrowed
towards the front, which is rounded but obscurely angulated in the middle, posterior
angles rounded off and entirely without spines ; rostrum obtuse, not furcate ; abdomen
about one-third as long as the cephalothorax (fig. 9), composed of three segments,
first segment very wide in the adult and equal to the united lengths of the second and
third segments; caudal stylets slender, more than twice as long as broad, longer than
either of the two preceding segments. ‘The antennules reach slightly beyond the
posterior extremity of the cephalothorax. Fifth pair of feet (fig. 10) two-jointed,
basal joint broad, quadrate, its outer margin prominent and mucronate in the middie,
whence arises a single very long seta; second joint bulbous at the base and tapering to
a long setiform apex. ‘Terminal spines of the swimming-feet (fig. 11) very slender and
sword-shaped, with finely pectinate margins. Length 1-2 millim.
G2
34 DR. G. 8. BRADY ON THE
Male.—Abdomen composed of four segments (fig. 14), the first very short, second
twice as long, third and fourth rather shorter; caudal stylets rather longer than the
preceding segment. Antennules more nodose than in the female owing to the distal
dilatation of many of the joints (figs. 12, 13), some of which are more or less distinctly
coalescent. ‘The fifth foot of the right side is longer than that of the left (fig. 15) ;
the inner margins of the first and second joints have large lobose protuberances, and
the last joint is bent upon itself at a right angle, ending in a sharp point; on the left
side the first joint is broad and quadrangular and has a long seta attached to its
angulated outer margin; second joint rather longer and narrower, third dilated at the
base and bearing on the inner margin of the narrowed distal extremity three small
setiform lamine.
Hab. Plentiful in surface gatherings from Otago Harbour, Port Chalmers, Bay of
Islands, and Napier; also in the net at a depth of 7 fathoms off Gisborne. ‘These
gatherings consisted almost entirely of females, males being extremely rare; almost
all had attached spermatophores.
A form described in the ‘ Challenger’ Report under the name of Acartia denticornis
is very nearly allied to this species, but the antennules of A. ensifera possess no
well-marked teeth, though these are constantly present in both sexes of A. denticornis.
The antennules in A. denticornis are fully as long as the animal itself, while in
A. ensifera they reach scarcely further than the extremity of the cephalothorax. The
peculiar enlargement of the first segment of the abdomen in the female A. ensifera is
likewise characteristic.
Genus Temora Baird.
1. TEMORA TENUICAUDA, sp. nov. (Plate IX. figs. 16-23.)
Female.—Cephalothorax robust, width great in proportion to the length (fig. 16),
broadly rounded in front, abruptly narrowed behind ; angles of the posterior extremity
rounded off, not at all produced or spinous. Abdomen short, four-jointed, first
segment about twice as long as either of the following three, and in the adult having
a pouch-like process on the ventral surface; caudal stylets cylindrical, extremely long
and slender, from eight to ten times as long as broad, bearing one small seta on the
outer margin beyond the middle, and three at the apex, one of which is spathulate at
its base and nearly as wide as the stylet itself (fig. 18). Antennules sparingly clothed
with extremely short sete, 24-jointed, slender, about as long as the cephalothorax
(figs. 16,17). Feet of the fifth pair (fig. 19) simple, rudimentary, three-jointed, the
last joint nearly twice as long as either of the preceding two and ending in three
small claw-like sete. Terminal spines of the outer branch of the swimming-feet
(fig. 20) well-developed and stout, the principal one strongly serrated with about
fourteen teeth on its inner margin. Length 1°75 millim.
MARINE COPEPODA OF NEW ZEALAND. 35
Male.—Cephalothorax and abdomen as in the female except that the abdomen is
five-jointed ; none of the caudal sete have dilated bases (fig. 23). Eighteenth joint
of the right antennule (fig. 21) very faintly serrulated. Fifth foot (fig. 22) of the
right side strongly prehensile, basal joint large, subquadrate, and produced into a Jong
curved claw which opposes the three-jointed outer ramus; first joint of the outer
ramus about twice as long as broad, second and third not much longer than broad and
bearing respectively one and two short marginal spines ; fifth foot of the left side short,
simple, three-jointed, last joint forming a short curved unguis.
Hab. Taken plentifully in the surface-net in the Bay of Islands and Otago Harbour,
also at a depth of 6 fathoms in Auckland Harbour.
Of all the specimens examined not one was found in perfect condition, and all were
very brittle and opaque owing to the action of picric acid preservative. The species is
very closely allied to the well-known 7’. longicornis of European seas, but there is no
serration of the right male antennule, except on the eighteenth joint ; the widely-
dilated tail-seta of the female is characteristic, and the serratures of the spines of the
swimming-feet are much fewer and larger than in 7. longicornis.
Genus CENTROPAGES Kroyer.
1. CENTROPAGES DISCAUDATUS, sp. nov. (Plate X. figs. 1-7.)
? Centropages typicus Kroyer, var. aucklandicus Krimer (1), pl. xv. figs. 1-5, p. 217.
Female.—Antennules very slender, 24-jointed, reaching considerably beyond the
posterior end of the cephalothorax. Infero-posteal angle of the cephalothorax produced
on each side into a long spine. Abdomen (fig. 6) three-jointed; first joint dilated
proximally and bearing on its dorsal margin a slender plumose seta; second joint
longer, much dilated proximally ; third very short, constricted proximally ; furcal joints
short, not much longer than broad, almost square as seen from above (fig. 7), distally
truncated and bearing five rather short terminal sete, three of which are bulbously
dilated at the base. Outer branch of the fifth pair of feet (fig. 5) having its second
joint produced inwardly into a very stout and long spine.
Male.—Infero-posteal angles of the cephalothorax spinous, but not so strongly as in
the female; abdomen four-jointed, all the joints subcylindrical (fig. 1), furcal joints
cylindrical, about twice as long as broad, terminal sete not swollen at the base; the
first abdominal segment short, the following three nearly equal, not very much longer
‘han broad and altogether devoid of spines. Basal joints of the antennules (fig. 1)
almost devoid of marginal processes or teeth; joints from the 15th to the 18th much
swollen, the 15th and 16th each bearing a strong marginal spine; the three following
joints densely pectinated with fine seta (figs. 2, 3), which, at the base of the 17th joint,
are replaced by a series of small squared tubercles. Outer branch of the fifth pair of
feet of the right side (fig. 4) terminating in a chelate joint, the blades of which are
36 DR. G. S. BRADY ON THE
long, slender, only moderately curved, and not at all crenulated, the outer blade having
a markedly constricted median neck ; outer branch of the left foot devoid of marginal
Length 1°6 millim.
Hab. Taken in the surface-net at Port Chalmers and in Otago Harbour.
sete, simple, and terminating in two very small apical sete.
This may perhaps be the form referred to by Dr. Kramer as Centropages typicus,
var. aucklandicus, though in some respects it does not agree accurately with his figures
and description. It cannot, I think, be properly referred to any variety of C. typicus,
though the pectinated armature of the male right antennule agrees exactly with that
species. ‘To make my meaning clearer I give some of the more important characters
of the two species in a tabular form.
It is, however, very closely allied to C. brachiatus (Dana), and the females of the two
species may not be easily distinguishable, but the armature of the antennule, together
with the structure of the fifth pair of feet in the male, presents very sufficient distinctive
characters.
Basal joints of the antennules...
15th and 16th joints of the
right male antennule.
First abdominal somite in
female.
Caudal segments in female......
Prehensile branch of right fifth
foot in male.
2. CENTROPAGES PECTINATUS, Sp. NOV.
C. typicus.
First, second, and fifth joints in
both sexes have strong mar-
ginal spines.
Sixteenth joint has a strong
spine.’
Two slender spine-like set on
ventral and a shorter one on
dorsal aspect.
Twice as long as broad; setz
not swollen at base.
Inner claw slightly pectinated ;
outer not distinctly con-
stricted in middle.
C. discaudatus.
No well-developed spines on the
basal joints in either sex.
A strong spine on both joints.
One dorsal seta.
Scarecly longer than broad ;
sete swollen at base.
Inner claw not pectinated ;
outer distinctly constricted.
(Plate IX. figs. 24-27.)
Female.—Abdomen slender, four-jointed (figs. 26, 27), the first joint wider than
the rest, bearing on its dorsal surface a brush of several fine hairs and at the posterior
angle a single slender spine; caudal segments about thrice as long as broad, two or
three of the principal terminal sete dilated basally (fig. 27 0). Lateral spines of the
outer branches of the swimming-feet lancet-shaped, with very finely pectinated margins ;
median joint of the outer branch of the fifth pair (fig. 25) produced internally into a
‘ Giesbrecht figures a rather feeble spine also on the fifteenth joint, but I have not been able to find it in
any of my specimens.
MARINE COPEPODA OF NEW ZEALAND. 37
long falcate spine, the concave margin of which is distinctly pectinated; the basal
joint of the fourth (?) pair of feet bears also a very long falcate spine (fig. 24). Male
unknown.
This description is extremely imperfect, being based upon only two battered
specimens which were taken—one off Napier in the net at a depth of 8 fathoms, the
other at the surface in Otago Harbour.
The characters seem clearly to distinguish this from any described species.
Subfamily PoNTELLIN «A,
Genus Lasibocera Lubbock.
1. Lasrbocura cervi Kramer. (Plate X. figs. 8-14.)
Labidocera cervi Kramer (1), p. 218, pl. xvi. figs. 6, 7.
Female.—Cephalothorax subtrunecate posteriorly, the ventral angles produced and
mucronate (fig. 9). Abdomen four-jointed, the first, second, and third segments about
equal, fourth about half as long as the preceding; caudal lamine rather longer than
broad, obliquely quadrate (figs. 9, 15); terminal sete scarcely as long as the abdomen.
Antennules reaching to the posterior extremity of the cephalothorax. Fifth pair of
feet (fig. 10) alike on both sides, two-branched, the internal branch composed of one
small joint, the outer of one much larger joint which bears three moderately large
lateral teeth and two smaller and much more slender ones at the apex. In the immature
condition, however (fig. 11), the outer branch consists first of three and afterwards of
two joints. Length 2:5 millim.
Male.—The infero-posteal angles of the cephalothorax (fig. 15) are not quite so
strongly spined as in the female; the abdomen five-jointed (fig. 13). The armature of
the right antennule consists of a fine pectination of the seventeenth and eighteenth
and of the coalescent nineteenth, twentieth, and twenty-first joints: the twenty-second
joint is prolonged internally, forming a sharp process which extends beyond the
apex of the penultimate joint. The right foot of the fifth pair (fig. 14) is strongly
prehensile, the basal joint simple, second joint attached near the middle by a hinge-
articulation, its outer portion forming a slender flexuous process, its inner portion
forming a broad lamina and giving attachment at its apex to a long and slender falcate
joint which opposes the outer process of the second joint; the foot of the left side is
simple, slender, three-jointed, nearly as long as that of the right side, its terminal joint
having a few short apical teeth and near the distal extremity a diffuse pubescent patch.
Length 2°8 millim.
Hab. In the surface-net, from Otago and Akaroa.
Though my observations differ in some respects from those of Dr. Kramer, I hav«
38 DR. G. 8S. BRADY ON THE
no doubt that this is the species named by him Labidocera cervi. Dr. Kramer says
that the abdomen of the female is two-jointed, and in some specimens I have found
the abdominal segments so interfused that it is difficult to make out the number:
there may, indeed, be no visible separation, but in some there are distinctly four
segments. In the allocation of joints to the right male antennule, I have followed
Dr. Giesbrecht (Labidocera nerii, &c.), not having myself been able to make them out
distinctly in the New Zealand specimens.
Family CYCLOPID.
Genus Cyctoprs (O. F. Miiller).
1. Cyctors Ewart G.S. Brady. (Plate X. figs. 15-17.)
1888. Cyclops ewarti G. S. Brady (3), pl. vii. figs. 1-6.
1891. 55 » idem (4) pl. vu. figs. 4-7.
One specimen of a Cyclops which I cannot in any way distinguish from C. ewarti
was found in a surface-net gathering from Otago Harbour.
This capture is peculiarly interesting, inasmuch as the species was originally described
from specimens taken in the Firth of Forth, and it seemed doubtful whether they
might not have made their way thither from some neighbouring freshwater habitat.
No other instance is on record, so far as I know, of a true Cyclops having been found
living in the sea, and it is very remarkable that this New Zealand example, of the
purely pelagic character of which there can be no doubt, should belong to the same
species.
I have thought it well to figure here some of the parts of the Otago specimen.
The only difference between it and the Scottish examples is the greater width of the
abdomen, which may, however, be accounted for by pressure. An interesting peculiarity
of the species, in which both northern and southern forms agree, is the bipectinate
character of the terminal spine of the inner branch of the fourth pair of feet (Pl. II.
fig. 17a). In all other species these pectinations are represented by very fine sete.
Genus O1tHona Baird.
1, OITHONA SPINIFRONS Boeck.
Oithona spinifrons Boeck (1), p. 25.
9 » G. 8. Brady (2), p. 90, pl. xiv. figs. 1-9, pl. xxiv A. figs. 1, 2.
Frequent in surface gatherings from Otago Harbour, and in the tow-net at 7 fathoms
off Gisborne.
I cannot distinguish these specimens from those which I have already (doc. cit.)
described and figured under the above name. Dr. Giesbrecht disagrees with my
MARINE COPEPODA OF NEW ZEALAND. 39
reference and thinks that they belong to 0. spinifrons Claus, but in the length of the
antennules and some other characters they seem to me to agree more closely with
Boeck’s species.
Family HARPACTICID.
Subfamily LoNGIPEDIINA.
Genus Ecrinesoma Boeck.
1. EcTINosoMA AUSTRALE, sp. nov. (Plate X. figs. 18-25.)
Antennules seven-jointed (?); first three joints stout and short, the succeeding joints
much more slender (fig. 18); antenne (fig. 19) nearly as long as the antennules, three-
jointed, secondary branch three-jointed, the second joint very short. Mandibles
(fig. 20) with one large terminal tooth and four short spinules; palp long, two-jointed,
and bearing on the first joint a minute two-jointed branchlet. The mouth-organs
(figs. 21, 22) and swimming-feet (fig. 23) present no special features. Feet of the
fifth pair (fig. 24) much subdivided ; inner segment digitiform and bearing two apical
spine-like sete, one of which is long, the other very short; outer segment laciniated,
its inner lobe bearing, like the neighbouring segment, two unequal sete, external to
which are three short processes bearing single apical sete, the first and third short,
the second long and slender. Candal lamine (fig. 25) very short and wide; sete
slender, non-plumose, the central one about twice as long as the two principal laterals.
Length ‘66 millim.
Hab. Otago Harbour, between tide-marks.
The distinctions between many so-called species of Hvtinosoma are extremely slight,
and in this case the fifth foot is the only member which presents very definite characters.
In the present state of our knowledge it seems very doubtful whether this, as well as
some others, should not rank rather as varieties than as distinct species.
Subfamily TacHIDIIN &.
Genus Evurrerre Claus.
1. EvuvTerre craciuis Claus.
18638. Euterpe gracilis Claus (1), p. 110, pl. xiv. figs. 1-13.
1880. ea » G.S8. Brady (1), p. 22, pl. xl. figs. 1-16.
1892. », acutifrons Giesbrecht (1), p. 555, pl. xliv.
1852. ? Harpacticus acutifrons Dana (2), p. 1192, pl. Ixxxiil. figs. 11 a, 6.
This species was found plentifully in most of the gatherings. In the surface-net
from Otago Harbour and Port Chalmers; in the net at 6 fathoms, Auckland Harbour ;
7 fathoms, off Gisborne; 8 fathoms, off Napier.
VoL. XV.—PaRT 11. No, 2.— August, 1899. H
40 DR. G. §. BRADY ON THE
Subfamily AMYMONIN &.
Genus AmMyMonz Claus.
1. AMYMONE cLAUsII ‘Thomson.
1882. Amymone clausii Thomson (1), p. 98, pl. v. figs. 1-8.
Taken in the surface-net, Bay of Islands.
Subfamily CANTHOCAMPTIN &.
Genus LaopHontE Philippi.
1. LAOPHONTE SERRATA (Claus).
1863. Cleta serrata Claus (1), p. 123, pl. xv. figs. 18-20.
1880. Laophonte serrata G. S. Brady (1), vol. ii. p. 71, pl. Ixxiii. figs. 1-14,
Hab. On alge in Lyttelton Harbour.
2. LAOPHONTE MEINERTI, sp. nov. (Plate XI. figs. 1-10.)
Antennules of the female short, seven-jointed (fig. 2); the comparative lengths of
go Bio eo Oo G aid
CoS 23 6
slender ; hand narrow, its external margin obscurely angulated near the middle ; terminal
unguis slender, slightly curved, rather longer than the hand. First pair of feet (fig. 5)
very slender; outer branch three-jointed, scarcely more than one-third as long as the
inner branch, exclusive of the terminal unguis; inner branch much elongated, destitute
of setee, with a long and slender apical unguis. Swimming-feet (fig. 6) small and
slender. Feet of the fifth pair (fig. 8) small, foliaceous, each joint bearing several
marginal setee, one of which is much longer than the rest. Length ‘75 millim.
the joints as in the following formula:
Posterior foot-jaws (fig. 4)
Second joint of the male antennule produced posteriorly into a short conical spine ;
fourth joint very much enlarged and forming a quadrate vesiculiform swelling, from
the anterior border of which arises a stout curved sensory filament (fig. 3).
Hab. On alge in Lyttelton Harbour. Two specimens.
Two further specimens, which may perhaps belong to this species, were found in a
gathering from between tide-marks in Otago Harbour. Figures of the posterior foot-
jaws and of the first and fifth feet of these specimens are given in Plate XI. figs. 9, 10.
These do not altogether correspond with similar parts of LZ. meinerti, but the differences
may possibly be varietal only.
MARINE COPEPODA OF NEW ZEALAND, 4]
Subfamily HARPACTICINA.
Genus Dactyopus Claus.
1. Dacrytopus tisBorpgEs Claus.
1863. Dactylopus tishoides Claus (1), p. 127, pl. xvi. figs. 24-28.
1880. 0 #0 G. S. Brady (1), p. 106, pl. liv. figs. 1-16.
Hab. Between tide-marks, Otago Harbour.
2. DACTYLOPUS HANSENI, sp. nov. (Plate XI. figs. 11-18.)
Very similar to D. tisboides in general appearance (fig. 11), but rather larger.
Antennules of the female (fig. 12) eight-jointed, rather densely setiferous; fourth
joint bearing a stout sensory filament. Posterior foot-jaw (fig. 14 a) slender; hand
bearing on each margin a single seta and a fringe of minute cilia; terminal claw
slender, about as long as the hand itself. Terminal claws of the first pair of feet
(fig. 15) long and slender, as are also the spines of the outer branch; penultimate
joint of the inner branch bearing near the distal extremity a single small plumose seta.
Fifth pair of feet of the female (fig. 18) large, the inner lamina considerably shorter
than the outer one ; apex finely ciliated, subtruncated, and bearing two long and two
short sete; distal extremity of the outer lamina bearing six sete of various lengths
and widely separated, the lateral margins ciliated; no cilia between the large sete.
Length -85 millim.
Male.—The antennule (fig. 13) is geniculated, but none of the joints are very tumid ;
terminal joint of the inner branch of the second pair of feet (fig. 16) forming two
curved finger-like processes, from near the base of which spring two long plumose sete :
fifth pair small (fig. 17); inner lamina shorter than the outer, with two short spine-
like apical sete; outer bearing six sete, the two apical ones much longer than
the rest.
Hab. Lyttelton Harbour: on alge, 1 specimen; tow-net at 6 fathoms, 1 specimen.
The nearest allies of this species seem to be D. cinctus (Claus) and D. stromii
Baird, but both of them differ distinctly in minor points.
Genus FLAVIA, gen. nov.
Like Dactylopus, except that the inner branch of the second pair of feet consists
of one greatly elongated joint, and is entirely destitute of spines or claws (ELE X.
fig. 30). Anterior and posterior foot-jaws simple, prehensile, with long terminal
ungues.
H 2
42 DR. G. S. BRADY ON THE
1. FLAVIA CRASSICORNIS, sp. nov. (Plate X. figs. 26-31.)
Antennules very short and stout, two-jointed, moderately setiferous (fig. 26). Feet
of the first pair as in Dactylopus, the terminal ungues thick and hook-like (fig. 29):
inner branch of the second pair (fig. 30) longer than the outer branch, very slender,
with truncate unarmed apex; near the base there is on each margin an angular
prominence, from which springs a minute seta, and near the distal extremity on the
outer margin a rounded non-setiferous tubercle, nearly opposite to which, from the
inner margin, arise two long and delicate plumose sete. Anterior foot-jaw simple,
elongated, with a slightly curved terminal unguis (fig. 27); posterior somewhat
similar, the hand armed with a long slender marginal spine and a strong curved
terminal unguis (fig. 28).
A single specimen of this species occurred in a gathering from among alge in
Lyttelton Harbour. The posterior antenne, mandibles, and maxilla were not seen
distinctly enough to permit of their being figured, but in general conformation are
similar to those of most Harpacticide.
Genus PHROS0, gen. nov.
Body slender; in general appearance like Canthocamptus. First four pairs of feet
with both branches three-jointed ; feet of fifth pair foliaceous, unbranched. Internal
branch of antenne two-jointed. Mandible strongly toothed, palp two-branched.
Anterior and posterior foot-jaws simple, with large terminal ungues and no palp.
1. PHROSO GRACILIS, sp. nov. (Plate XI. figs. 19-27.)
Female.—Antennules eight-jointed, rather densely setiferous ; first four joints stout,
last three much more slender (fig. 20): lengths of the joints as in the following
15 Oe BO. hoe.
NPGS eyhaney | fy) 4 AEN Sy
Antenne (fig. 21) nearly as large as the antennules, three-jointed ; to the basal joint is
attached a two-jointed internal branch. Mandibles well developed (fig. 22), with sharp
slender teeth and a palp composed of a large basal joint and two single-jointed branches.
Maxille (fig. 23) small, lobose (imperfectly seen). Anterior foot-jaw (fig. 24) slender,
simple, geniculated, with a stout terminal unguis and three plumose sete attached to
the second joint. Posterior foot-jaw (fig. 25) larger than the anterior, composed of a
single stout joint which bears a long falcate, terminal unguis. Basal joint of the fifth
pair of feet (fig. 27) short and broad, and bearing on each lateral angle a long seta;
terminal joint subovate, twice as long as broad; apex narrow and truncate; external
formula : from the fifth joint arises a stout sensory filament.
margin ciliated above, and below the middle having two long widely separated sete.
Length 1:3 milliim. Male unknown.
Hab. One specimen taken in the surface-net, Port Chalmers.
MARINE COPEPODA OF NEW ZEALAND. 45
Genus THALESTRIS Claus.
1. ‘THALESTRIS FORFICULA Claus.
1863. Thalestris forficula Claus (1), p. 181, pl. xvii. figs. 7-11.
3 o Thomson (1), p. 104, pl. x. figs. 16-21.
Otago Harbour, between tide-marks; “ dredged abundantly in Dunedin Harbour ”
(Thomson).
2. THALESTRIS CILIATA, sp. nov. (Plate XI. figs. 28-33.)
Robust ; lateral margins of the abdominal segments clothed with short sete (fig. 33).
Antennules nine-jointed (fig. 28); joints subequal, the length represented by the
; | LO Bes BeB07 58.0 TA eae ;
following formula: 3 S66 46-33 q Posterior foot-jaw forming a subovate,
elongated hand, the anterior margin of which bears a continuous fringe of short,
closely-set sete ; ‘terminal unguis stout, falcate, more than half as long as the hand
(fig. 29). Outer branch of the first pair of feet (fig. 30) with densely pectinated
margin: one long and stout spine at the apex of the first and one on the middle of the
second joint; the last joint has two fine marginal sete and one very long, slender
seta at the apex, between which are two stout faleate claws with hook-like projections at
their bases: the outer branch has a single seta near the distal extremity of the first joint,
and at its apex are two ungues, one large and one small. The marginal spines of the
swimming-feet (fig. 51) are long and slender, and the outer edges of the joints of both
branches are ciliated. The two lamine of the fifth pair of feet (fig. 52) are of equal
length; the outer lamina ovate, with finely ciliated margins, four long sete at the apex
and two on the outer edge; the inner lamina has five principal sete, two marginal and
three apical, the intervals between which are ciliated. Male unknown.
Hab. Akaroa Harbour. One specimen only.
3. THALESTRIS AUSTRALIS, sp. nov. (Plate XII. figs. 1-5.)
Autennules nine-jointed, slender, the comparative lengths of the joints as in the
following formula: : a! : : : : ; : : : i : : : : (fig. 1).
(fig. 2) subovate, bearing near the middle of the anterior margin a single spine-like
seta, terminal unguis moderately stout and reaching a little beyond the middle of the
hand. Outer branch of the first pair of feet (fig. 5) shorter than the inner, stout,
its inner margin bearing rather in front of the middle a stont plumose seta, outer
margin a large median spine and four apical ungues which successively increase in
length from first to last; the inner branch has a long. slender, apical unguis and a
much shorter seta; anterior margins of the larger ungues of both branches finely
pectinated. Marginal spines of the swimming-feet (fig. 4) long and slender, with
Hand of the posterior foot-jaw
44. DR. G. S. BRADY ON THE
blunt apices ; external margins of all the joints ciliated. LLamine of the fifth pair of
feet (fig. 5) broad, subovate, nearly equal in length; margins of the external lamina
finely ciliated, and bearing six subequal apical sete; inner lamina with five sete on
its subtruncate apical margin. Abdominal segments slightly overlapping each other,
but altogether devoid of sete or serrulations Male unknown.
Hab. Otago Harbour, between tide-marks. One specimen only.
Genus Harpacticus Milne-Edwards.
1. HARPACTICUS GLABER, sp. nov. (Plate XII. figs. 6-11.)
Female.—In general appearance like H. chelifer. Antennules (fig. 6) slender,
moderately setiferous, eight-jointed ; lengths of the joints as in the following formula :
Wee Be Bo Gav S
IO i Gat Ba
a small two-jointed branch, extremity of the last joint abruptly truncated and bearing
two long, rigid, finely pectinated spines and four slender geniculated sete ; posterior
foot-jaw (fig. 8) almost exactly similar to that of H. chelifer. Outer branch of the
first pair of feet elongated, two-jointed, terminating in three subequal, blunt, slightly
curved and very finely pectinated claws; outer margins very sparingly ciliated ; each
joint bearing within a short distance of its distal end a single short marginal seta:
inner branch terminating in a strong slightly curved claw; the long first joint slightly
ciliated and having near its apex a slender seta. Feet of the fifth pair two-jointed
(fig. 10), basal joint wide, its inner portion distally produced and giving attachment
to three long sete; terminal joint much narrower, margins ciliated, with three long
sete at the apex and one on the outer edge. ‘The spines of the swimming-feet, like
those of the first pair and of the antenne, are minutely pectinated. ‘The margins of
the abdominal segments (fig. 11) are smooth except for a few minute hairs at the
lateral angles and on the caudal lamin; principal tail-setee very long.
Hab, Otago Harbour. One specimen taken in the surface-uet.
From H. chelifer this species may be distinguished by the number and comparative
lengths of the joints of the antennules, by the armature of the antenne, the non-
spinous margins of the first pair of feet, the different build of the fifth pair, and the
almost smooth margins of the abdominal somites. There seem to be points of
difference also in the mandibles and maxille, but these I have not clearly made out.
Antenne (fig. 7) stout, nearly as long as the antennules, bearing
Genus Pevripium Philippi.
1. P<IDIUM NOVAI-ZEALANDIM, sp. nov. (Plate IX. figs. 28, 29; Plate XII.
figs. 12-15.)
This is in general build and appearance exactly like the common European species
P. interruptum, Goodsir, the only important differences being found in the posterior
MARINE COPEPODA OF NEW ZEALAND. 45
foot-jaw and fifth pair of feet. The manus of the second pair of foot-jaws (Pl. IX.
fig. 29) is ovate, elongated, the anterior margin fringed with a continuous series of
short sete ; terminal unguis long and slender. Feet of the fifth pair (PI. XII. fig. 14)
stout, simple, with truncate apex, which bears three short, stout, and very blunt spines,
the outermost of which is attached at a little distance from the apex: the inner
margin bears beyond the middle a single seta; the outer margin has also a single seta
much nearer the base, and is slightly pubescent between that point and the apex.
The shell is extremely hard and thick, and closely beset with small circular puncta.
Colour dark brown. Length ‘85 millim.
Hab. Taken in the surface-net, Port Chalmers and Otago.
Subfamily Ipyin &.
Genus Ipya Philippi.
1. Ipya Furcata (Baird).
Tisbe furcata Claus, Lilljeborg.
Canthocamptus furcatus Baird.
Idya furcata Boeck, Brady, Thomson.
On alge in Lyttelton Harbour. ‘Common in shore-kelp and rock-pools near
Dunedin ; also on kelp in Paterson’s Inlet” (Zhomson).
Genus ScUTELLIDIUM Claus.
1. ScUTELLIDIUM PLUMOSUM, sp. nov. (Plate XII. figs. 16-21, 25-25.)
In general form and appearance indistinguishable from the British S. tishoides.
The setose investiture of the first foot (fig. 20) is, however, more dense, and the
spinous armature of the last joint of the outer branch of the swimming-feet (fig. 21)
is also different, there being on the outer margin towards the apex two plumose
lancet-shaped spines with intermediate small hairs, while in S. tésbotdes (fig. 22) there
are three stout, non-plumose, dagger-shaped spines with no intermediate hairs. Fifth
foot in the female (fig. 25) elongated, slender, subcrescentic, pointed at the extremity,
with setose margins, but not pubescent on the surface; in the male (fig. 23) small,
subquadrate, with obliquely truncated apex, which bears three long and _ several
shorter setee. ‘The antennules in the male are geniculated (fig. 16): from the second
joint springs a small biarticulate peduncle, which bears three stout sensory filaments,
and to the antepenultimate joint is attached a much longer and stouter sense-organ.
Length :88 millim.
Hab. Vhis species occurred sparingly in surface gatherings from Otago and Port
Chalmers, and in gatherings from seaweeds in Akaroa and Lyttelton Harbours.
46 DR. G. S. BRADY ON THE
The males of §. plwmosum seem, like those of S. tisboides, to be scarce. I have seen
only one specimen, some parts of which are here figured. Of S. tisboides I have
hitherto seen only one imperfect male specimen; and as Dr. Claus has only very
partially described or figured it, one may suppose that his experience is similar.
Subfamily PoRCELLIDIIN A.
Genus PorcELLIDIUM Claus.
1. PorceLLipium FULVUM Thomson.
1882. Porcellidum fulvum Thomson (1), p. 107, pl. vi. figs. 10, 11, pl. vu. figs. 8-13.
On alge in Lyttelton Harbour.
Section I]. PH&CTLOSTOMA Thorell.
Family CORYCAIDAi Dana.
Genus Corycaus Dana.
1. Corycamus Ropusta Dana (?).
Specimens which I refer doubtfully to this species were taken in the surface-net off
Port Chalmers and in Otago Harbour.
Genus PatRocopg, gen. nov.!
Antennules five-jointed ; antenne three-jointed, simple; mandibles simply toothed ;
posterior foot-jaw forming a prehensile hand with two slender terminal ungues. ‘I'wo
pairs of swimming-feet, each foot consisting of two one-jointed branches.
1. PavRocopE RoBusta, sp. nov. (Plate XIII. figs. 1-9.)
Antennules (fig. 3) short, five-jointed, sparingly setiferous, joints nearly equal in
length. Antenne nearly as long as the antennules, simple, three-jointed, last joint
shorter than the preceding, subquadrate, with six sete arising from the subtruncate
apex, and on the proximal half of the posterior margin a series of about six small,
equal, pellucid, lancet-shaped sete; on the distal half two short and two rather longer,
stout, curved, plumose sete (fig. 4). Trophi very small; mandibles without a palp (?),
‘ quvpos, few; Kw, an oar.
MARINE COPEPODA OF NEW ZEALAND. 47
apex broad and bearing an even series of small teeth (fig. 5). Maxillee composed of four
digitiform lobes (fig. 6), which are armed with numerous spine-like divaricate seta.
Anterior foot-jaws laminar, bearing numerous lateral sete, but not divided into distinct
lobes (fig. 7): posterior foot-jaws two-jointed, basal joint short and stout, with three rigid
spiniferous sete at the anterior distal angle; second joint forming a strong subtriangular
hand, to the anterior angulated margin of which are attached two long stout sete,
to the narrow distal extremity are hinged two long, biarticulate, slender ungues, which
are armed with spine-like marginal sete (fig. 8). Two pairs of swimming-feet (fig. 9),
two-branched, each branch consisting of a single broad lamina; distal and outer
margins of the inner branch setiferous; outer branch with four large lancet-shaped
spines on the outer margin, at the apex a larger spine, and on the inner margin three
long sete. The cephalothorax is equal in length to the abdomen, and consists of
three segments only, the first two being coalescent (figs. 1,2); the abdomen is two-
jointed and suddenly narrower than the cephalothorax: to the posterior angles of the
last thoracic segment are attached two small sete ; angles of the first abdominal
segments produced backwards, second segment twice as long as the preceding, con-
stricted in front; caudal segments very short. Length ‘88 millim.
Hab. Of this very interesting species two specimens were found in a surface-net
gathering from Otago Harbour, and one in the net at 7 fathoms from off Gisborne.
The specimen described by Mr. T. Scott under the generic name Saphirella seems
to be, except as to the mouth-organs, very similar to the present, and I regret that,
owing to the Saphirella having passed out of Mr. Scott’s hands, I have not been able
to compare the actual specimens. Mr. Scott's descriptions may be found in his paper
on Entomostraca from the Gulf of Guinea (Transactions of the Linnean Society,
2nd ser. Zool. vol. vi. pt. 1, 1894).
[Closely allied to Pauwrocope is a form represented in these gatherings by only two
imperfect specimens, which, like Pawrocope, were taken at the surface in Otago
Harbour. I am unable, for want of suitable material, to give a complete account of
this species, and it seems not unlikely that it may really represent the male of
Paurocope, the general resemblance being very close, more especially in the swimming-
feet and caudal lamine. ‘The antennules and antenne are, however, very different,
but this difference, to a lesser extent, is likewise seen in the closely related genera
Sapphirina and Coryceus. A very remarkable character is found in the presence of
two simple lenses (ocelli) at the bases of the second pair of foot-jaws. Figures of the
animal and some of its appendages are given in Plate XIII. figs. 10-17.
Provisionally I propose the name Centromma thomsoni for this species. ]
VOL. Xv.— Part u. No. 38.—August, 1899. I
48 DR. G. S. BRADY ON THE
Section III. STPHONOSTOMA Thorell.
Family ENTOMOLEPID A, fam. nov.
Genus ENToMOLEPIS ', gen. nov.
Dorsal surface covered by a thin, oval, scale-like shield, beyond which the extremities
of the limbs project only slightly. Antennules slender, simple; antenne prehensile ;
swimming-feet two pairs (?). Abdomen short, slender, and, like the cephalothorax,
completely covered by the scale-like dorsal investment.
1. ENTOMOLEPIS OVALIS, sp. nov. (Plate XIII. figs. 18-21.)
Outline, as seen from above, oval, widest in the middle, width equal to more than
half the length (fig. 18). Antennules ten-jointed, slender, the penultimate joint
bearing a long and stout sensory filament (fig. 19); the comparative lengths of the
sie 5 9 boo Bot oO oOo To Oo Wo NO
joints as in the following formula: 5545-74-35 7 7 10 8°
(fig. 20), the basal joint bearing at its apex a minute secondary branch ; second joint
with a row of fine cilia on its distal half; third joint small and having a stout terminal
unguis. Maxilla simple, two-jointed, the second joint pubescent and bearing three
long terminal sete. Two pairs of foot-jaws (figs. 18 d, ¢), stout, prehensile, like those
of Dyspontius. Siphon slender and of moderate length; swimming-feet two pairs (?)
two-branched, each branch composed of three joints (fig. 18 /). Length 1:3 millim.
Antenne three-jointed
One specimen only was found among mud from a depth of 1-0 fathoms in Lyttelton
Harbour.
The foregoing description is incomplete, owing to some of the parts having been
imperfectly seen, and it is to be hoped that some future observer may be enabied by
the help of further specimens to describe more fully what seems to be a very remarkable
form.
Family ARTOTROGID A.
Genus Artotrocus Boeck,
1. ArtTotTroaus ovatus Thomson.
Artotrogus ovatus Thomson (1), p. 118, pl. x1. figs. 11-14.
In the surface-net off Port Chalmers. One specimen.
2. ARTOTROGUS BREVICAUDATUS, sp. nov. (Plate XII. figs. 27-29; Plate XIII.
figs. 22-26.)
Cephalothorax much narrowed in front, widest behind the middle; abdomen very
short, only about one third as long as the cephalothorax (Plate XII. fig. 22); siphon
* &vropor, an insect; dems, a scale.
MARINE COPEPODA OF NEW ZEALAND. 49
long and slender (fig. 24). Antennules very short, fourteen-jointed (2) (fig. 23), the
basal joints very indistinctly marked; comparative lengths of the joints as in the
following formula: 3 ; - ae ee = : - : = : = : = : = : = First segment of the
abdomen much longer than the following segments, inclusive of the furca (fig. 26).
The mouth-organs and swimming-feet present no distinctive characters. Length
1-1 millim.
Hab, Akaroa Harbour, 6 fathoms.
Genus Cariaus.
1. CaLiGus LONGICAUDATUS, sp.nov. (Plate XII. fig. 26.)
A single specimen of a Caligus, which seems distinct from any described species,
occurred in a surface-net gathering from Port Chalmers. The much elongated
abdomen, the abruptly angular carapace, together with the slenderness and length of
the last pair of feet, are the chief distinctive characters. Length 6 millim,
EXPLANATION OF THE PLATES.
jd BF. Ml Oe BG
Paracalanus parvus, p. 32.
Fig. 1. Antennule of male, x 140.
Fig. 2. Foot of fifth pair of male, x 210.
Clausocalanus furcatus, p. 32.
. Female, seen from right side, « 84.
Fifth pair of feet of female, x 250.
. Antennule of male, x 84.
. Fifth pair of feet of male, x 210.
. Abdomen and foot of fifth pair, ¢, seen laterally, x 165.
eo)
S
og
ID Or oo
Acartia ensifera, p. 33.
Fig. 8. Female, seen from above, x 84.
Fig. 9. Abdomen of female, seen from right side, x 110.
Fig. 10. Fifth pair of feet of female, x 210.
. 11. Terminal spine cf one of the swimming-feet, x 210.
Fig. 12. Right antennule of male, x 140.
Fig. 15. Left antennule of male, x 140.
Fig. 14. Abdomen of male, x 110.
Fig. 15. Fifth pair of feet of male, x 250.
DR. G. S. BRADY ON THE
Temora tenuicauda, p. 34.
. Female, seen from left side, x 50.
. Antennule of female, x 84.
. Abdomen of female, x 84.
. Foot of fifth pair, 29, X 210.
. Terminal spine of swimming-feet, x 300.
. Right antennule of male, x 84.
. Fifth pair of feet of male, x 110.
. Abdomen of male, seen from front, x 210.
Centropages pectinatus, 2, p. 36.
. Foot of fourth pair (?), x 84.
. Foot of fifth pair, x 210.
. Abdomen, seen from front, x 84.
. Do oh AIL SS eee
a. Sete of first segment, more highly magnified.
6. Sete of tail, more highly magnified.
Peltidium novee-zealandie, p. 44.
. Antennule of female, x 140.
, 2A),
Posterior foot-jaw, X 210.
PLATE X.
Centropages discaudatus, p. 35.
. Male, seen from right side, x 65.
. Armature of 17th segment of right antennule, ¢, x 210.
= 17th, 18th, and 19th segments of same, x 210.
Fifth pair of feet of male, x 84.
female, x 100.
99 99
. Abdomen of female, seen from side, x 100.
. Furca of female, x 100.
Labidocera cervi, p. 37.
. Antennule of female, x 50.
. Abdomen of female, seen from below, x 50.
> Hoot of fifth pain.) 95) >< 100!
. The same, immature, x 84.
. Right antennule of male, X 50.
. Abdomen of male, x 50.
. Fifth pair of feet of male, X 50.
Fig. 15.
Fig. 16.
. Fourth foot of same, x 140.
23)
(eye)
Oo OMS OP OO De
MARINE COPEPODA OF NEW ZEALAND. 51
Cyclops ewarti, p. 38.
Antennule of female, x 140.
Abdomen of same, x 84.
a. Terminal spine of inner branch of the same, x 300.
Ectinosoma australe, ¢ , p. 39.
. Antennule, x 250.
. Antenna, x 210.
. Mandible, x 250.
. Anterior foot-jaw, x 250.
. Posterior foot-jaw, x 250.
. One of the swimming-feet, x 210.
. Foot of fifth pair, x 210.
d. Caudal segments and sete, x 210.
Flavia crassicornis, 2, p. 42.
. Antennule, x 250.
. Anterior foot-jaw, x 250.
. Posterior foot-jaw, x 250.
. Foot of first pair, x 210.
second pair, x 210.
99
. Abdomen, x 100.
PLATE XI.
Laophonte meinerti, p. 40.
Female, seen from left side, x 84.
. Antennule of female, x 300.
ms male, x 300.
. Posterior foot-jaw, x 300.
. Foot of first pair, x 210.
. One of the swimming-feet, x 210.
. Inner branch of foot of second pair, ¢, 300.
. Foot of fifth pair, 9, x 210.
first pair, X 226.
fifth pair, variety, x 210.
39
39
52 DR. G. 8S. BRADY ON THE
Dactylopus hanseni, p. 41.
Fig. 11. Female, seen from left side, x 100.
Fig. 12. Antennule of female, x 210.
Fig. 13. na male, x 210.
Fig. 14. Inner branch of antenna, x 210.
Fig. 14 a. Posterior foot-jaw, x 280.
Fig. 15. Foot of first pair, xX 210
Fig. 16. Inner branch of foot of second pair, ¢, x 300.
Fig. 17. Foot of fifth pair, ¢, Xx 210.
ate 1 ». 85% MO,
Phroso gracilis, 2, p. 42.
Fig. 19. Female, seen from right side, x 68.
Fig. 20. Antennule, x 210.
Fig. 21. Antenna, x 210.
Fig. 22. Mandible, x 210.
Fig. 23. Maxilla, x 210.
Fig. 24. Anterior foot-jaw, X 250.
Fig. 25, Posterior foot-jaw, x 250.
Fig. 26. Foot of fourth pair, x 120.
Fig. 27. a) wuithypair, x 210:
Thalestris ciliata, 2, p. 43.
Fig. 28. Antennule, x 210.
ig. 29. Posterior foot-jaw, xX 260.
. Foot of first pair, x 210.
Se thirdipairs livia:
sy sed od joys SIMON),
. Abdomen, seen from front, x 84.
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BS
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PLATE XII.
Thalestris australis, 9, p. 43.
Fig. 1. Antennule, x 210.
Fig. 2. Posterior foot-jaw, x 219.
Fig. 3. Foot of first pair, X 175.
Fig. 4. One of the swimming-feet, x 175.
Fig. 5. Foot of fifth pair, x 130.
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MARINE COPEPODA OF NEW ZEALAND. 55
Harpacticus glaber, 2, p. 44.
. Antennule, x 200.
. Antenna, x 200.
. Posterior foot-jaw, x 210.
. Foot of first pair, x 140.
, fifth pair, x 210.
. Abdomen and caudal sete, x 84.
Peltidium nove-zealandie, ¢, p. 44.
. Foot of first pair, x 210.
peeetourth: pair, << 120:
tthe pair ><) 20,
. Abdomen, x 130.
Scutellidium plumosum, p. 46.
. Antennule of male, x 300.
. Antenna of male, x 300.
. Anterior foot-jaw of male, x 210.
. Posterior foot-jaw of male, x 210.
. Foot of first pair of female, x 210.
. Outer branch of one of the swimming-feet, x 300.
i a a (D. tisboides: Trish specimen), x 300.
. Foot of fifth pair of male, x 250.
. First abdominal somite of male, «x 210.
. Abdomen and tail of female with fifth pair of feet, x 140.
Caligus longicaudatus, p. 49.
. Female, seen from below, x 16.
Artotrogus brevicaudatus, p. 48.
. Mandible (?), x 120.
. Maxilla, x 210.
. Foot-jaw, x 120.
PLATE XIII.
Paurocope robusta, 2, p. 46.
. Animal, seen from left side, BA
; s) ss above, :
. Antennule, x 140.
oon o oe
10.
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16.
Sy,
LS:
ON THE MARINE COPEPODA OF NEW ZEALAND.
. Antenna, x 210.
. Mandible, x 500.
. Maxilla, x 500.
. Anterior foot-jaw, xX 500.
. Posterior foot-jaw, < 210.
. Foot of first pair, x 210.
Centromma thomsoni, p. 47.
Animal, seen from below, x 140.
Antennule, <x 210.
Antenna (?), x 210.
Mandible (2), x 250.
Anterior foot-jaw, x 210.
Posterior foot-jaw, x 210.
Foot of first pair, X 210.
Abdomen and caudal lamine, x 210.
Entomolepis ovalis, p. 48.
Male (2), seen from below, X 84.
a. Siphon; 6. Antenna; ¢. Mandible-palp?; d. Anterior foot-jaw ,
e. Posterior foot-jaw ; f Swimming-foot.
. Antennule, x 210.
. Antenna, xX 210.
. Mandible-palp (?), x 210.
Artotrogus brevicaudatus, p. 48.
. Outline of animal, seen from below, x 9d.
. Antennule, x 210.
. Siphon, x 120.
5. Posterior foot-jaw, x 120.
. Abdomen, X 140. |
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CONTENTS.
Il. On the Marine Copepoda of New Zealand. By G. Srmwarpson Brapy, M.D., D.Sc.,
Lb. Di TLRS. OMS. (Plates: UX) ae ee pagelok
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I. On the Eatinet Birds of Patagonia.—l. The Skull and Skeleton of Phororhacos
inflatus Ameghino. By C. W. Annrews, B.Sc., F.Z.8., Assistant in the British
Museum (Natural History).
Received March 21, read April 18, 1899.
[Puares XIV.—XVII.]
INa paper read at the Meeting of the British Association at Ipswich, in 1895, and
subsequently published in the ‘ Ibis’ !, the present writer gave a brief review of a very
interesting memoir by Dr. Florentino Ameghino on some remarkable fossil birds from
the Tertiary deposits of Patagonia. Since that time, the specimens described by
Ameghino have come to the Geological Department of the Natural History Museum,
so that an opportunity of examining the bones themselves has arisen ; and in the case
of the most remarkable of the specimens, the skull and skeleton of Phororhacos
inflatus, the careful removal of the adherent matrix has revealed for the first time
many points of great importance, e.g. the structure of the palate. It seems, therefore,
desirable to give as complete an account as possible of the osteological characters
of this bird, in order, if possible, to arrive at some conclusion as to its precise
relationship to recent forms; and although the account given in Ameghino’s excellent
paper is an accurate one as far as it goes, still it seems best, even at the risk of some
repetition, to describe in detail all the parts of the skeleton available, in order to
facilitate the comparison of them with recent types.
It may be at once stated that if the remaius described by Ameghino really formed
parts of one individual skeleton (and there seems to be no reason to doubt this), then
this bird presents a most extraordinary combination of characters, to which no close
parallel can be found among recent forms. A brief summary of the various opinions
that have been expressed relative to the systematic position of the Stereornithes will
be found in the above-mentioned review.
The present paper will be confined to a description and comparison of the skeleton
of Phororhacos inflatus Ameg., that species being represented by much the best
series of remains.
SKULL AND SKELETON OF PHORORHACOS INFLATUS AMEGHINO.
Of this species there is a fine series of bones, including the skull and mandible,
scapula, coracoid, bones of wing, pelvis, and bones of hind limb, as well as some
vertebra: these are said to belong to a single individual. ‘There is also a number of
1 « Remarks on the Stereornithes, a Group of Extinct Birds from Patagonia,” Ibis, 1896, p. 1,
VOL. XV.—PART UI. No. 1.-—October, 1899. K
56 MR. C. W. ANDREWS ON THE SKULL AND
odd bones referred to this species, but the following description is founded as far as
possible on the associated set, which is the type-specimen.
The Skull and Mandible. (Plates XIV. & XV.)
The skull and mandible are in a wonderfully perfect state of preservation, and, now
the matrix has been removed, all the important details of their structure can be made
out. The skull has been slightly crushed, so that there is some dislocation of the
bones of the roof, and also a slight compression of the beak, but otherwise it retains
its original form.
Occipital surface of skull of Phororhacos inflatus Ameghino, # natural size.
Lv., lambdoidal ridge ; m.¢., mammillary tuberosities; oc.c., occipital condyle ; oc.f., occipital foramen ;
p-p., paroccipital processes ; sf.r., supraforaminal ridge; q., quadrate.
The Occipital Region (text-figure, above).—The occipital surface is remarkable for its
flatness, its great width from side to side, and the distinctness with which it is marked
off from the other regions. This last peculiarity is due to the great development of
the lambdoidal ridge (/.r.), particularly towards the sides. In its middle portion this
ridge merely forms the angle (of about 100°) between the occipital and dorsal regions,
but laterally it is produced outward, forming prominent lateral crests which are
continuous ventrally with the broad paroccipital processes (p.p.). These latter are
convex posteriorly, both from above downward and from side to side; their inferior
angles extend somewhat below the occipital condyle, and are formed by the ventral
prolongations of the very well-marked supraforaminal ridges. These ridges run
downward and somewhat outward from the upper border of the foramen magnum (oc.f.),
and form the inner borders of the paroccipital processes, separating them from a
SKELETON OF PHORORHACOS INFLATUS. 57
depressed area lying between them and the foramen magnum. ‘This valley-like
depression is closed above by the meeting of the supraforaminal ridges (sf.r.), but
opens ventrally on either side of the occipital condyle. The inferior angles of the
paroccipital processes are united to the mammillary tuberosities (m.t.) of the basi-
temporal platform by prominent buttresses of bone, behind and internal to which the
vagus foramina open.
The foramen magnum (oc.f.) is relatively very small; it is oval in outline, the long
axis being vertical. The plane of the opening is inclined backward, making an angle
of rather more than 45° with the long axis of the skull. The transverse diameter of
the opening is 9 mm., the vertical 16 mm.
The slightly pedunculate occipital condyle (oc.c.) is oval, or rather reniform, ils
upper border being nearly straight, with a vertical groove running down from its
middle point nearly to the centre of the condyle. There is a small but deep pre-
condylar fossa (pe.f., Plate XV. fig. 1).
The base of the Skull (Plate XV. fig. 1)—The basitemporal platform (0.f.) is a
triangular area which is very slightly elevated, and below the level of which the
occipital condyle projects. It is concave from side to side, and the postero-lateral
angles are produced downward into the very prominent mammillary tuberosities
(m.t., fig. 1), which, as mentioned above, are united to the inferior angles of the
paroccipitals by buttresses of bone. ‘The posterior margin of the platform is defined
by a shallow groove separating it from the occipital region. The lateral borders are
thickened ridges, forming anterior prolongations from the mammillary tuberosities
(m.t.), and converging so that they appear to have met in front in a rounded median
angle. ‘This, no doubt, lies immediately below the common opening of the eustachian
tubes, which are immediately dorsal to the thickened ridges just described, and open
posteriorly by wide apertures into the tympanic cavity. In front of the basitemporal
platform the narrow rostrum (7.) is seen for a short distance, but it is here badly
preserved, and anteriorly is completely invested by the bones of the palate.
On the sides of the basitemporal platform, rather high up and behind its anterior
angle, are a pair of basipterygoid facets, with which the pterygoids articulate by means
of stout processes borne on their inner surfaces rather behind their middle point (5.p7.).
Below and behind the articular facets for the pterygoids the basitemporal forms a thin
plate of bone (pretemporal wing), which looks outward and downward; the lower edge
of this forms the floor of the eustachian canal, while its upper portion is the outer
wall of the large pretympanic recess, which opens widely into the tympanic cavity
immediately above the posterior aperture of the eustachian tube.
The side of the Skull (Plate XI1V.).—The tympanic cavity (ty.) is relatively small ;
posteriorly it is bounded by the concave anterior face of the paroccipital process; the
inferior portion of its inner wall is formed by the buttress of bone which joins the
paroccipital to the mammillary tuberosity (m.t.). Its outer edge is formed by a
downwardly-projecting flange of the squamosal, the anterior edge of which runs down
K 2
58 MR. C. W. ANDREWS ON THE SKULL AND
between the anterior and posterior quadrate facets. Of these, the antero-external (0.f.)
is very large, oval in outline, and concave in all directions, but more deeply so from
before backward; its outer edge forms a prominent lip on the lower edge of the
squamosal prominence. The other facet (¢.f.) is placed internal and posterior to the
Jast; in form it is a very elongate oval, the long axis being antero-posterior, in which
direction it is deeply concave: it locks outward and downward. The opening of a large
pneumatic fossa separates it from the downwardly projecting process of the squamosal
and also from the anterior facet. Mesiad of the posterior facet, but separated from it
by a narrow flat ledge of bone, is a very deep pocket-like depression, into which open
numerous foramina. ‘The most posterior of these seems to be the fenestra ovalis; the
others are probably pneumatic. Ventrally the tympanic cavity communicates by wide
openings with the anterior tympanic recess and the eustachian canal.
The squamosal region (Plate XIV., sq.) is deeply concave from before backward,
and slightly convex from above downward. Posteriorly it forms the anterivr face
of the prominent lambdoidal crest; ventrally, as already mentioned, it is produced
downward into a pointed process, which helps to form the outer boundary of the
tympanic cavity. Anteriorly its ventral edge is formed by the prominent outer lip of
the anterior quadrate facet (0.f.). About 7 mm. above this there is a prominent
elongated tuberosity, the long axis of which is directed downward and inward; this I
take to be the zygomatic process (zy.). From its upper end a slight ridge runs upward
and backward to the lambdoidal crest (d.7.), and forms the lower boundary of the
temporal fossa. Internal to the zygomatic process, and connected with it by a short
ridge, is another angular projection, the pretympanic process; from this also a slight
ridge runs upward and backward, but it soon dies away without reaching the
main temporal ridge.
As might be expected from the great massiveness of the mandible, the temporal
fosse are very large and deep. Their lower opening is bounded posteriorly by the
zygomatic, anteriorly by the postorbital process (p.ord.p.), which approach to within
17 mm. of each other. The fosse are much deeper than broad, and greatly constrict
the side of the cranium. Superiorly they are bounded by the temporal ridges, which,
as above mentioned, run upward and backward from the zygomatic process till they
meet the lambdoidal crest (/.7.), with which they are confluent for 28 mm. Leaving
the lambdoidal ridge, they sweep forward, first towards the middle line, then outward,
terminating anteriorly on the upper surface of the postorbital process. On the roof
of the skull the fossee are separated one from another by an interval of about
12 mm. only.
On the side of the cranium, immediately below the constriction caused by the
temporal fosse, and about on the same level as the quadrate facet, there is a deep
infundibuliform depression, at the bottom of which the trigeminal foramen opens.
This depression is marked off from the orbit by a ridge which runs upward and
SKELETON OF PHORORHACOS INFLATUS. 59
outward, terminating above in a prominent cristiform ridge on the ventral surface of
the postorbital process.
The orbits are very large; they appear to be divided one from another by a complete
bony septum (7.0.s.), or there was at most a small vacuity in front of the optic foramen.
These latter appear to be confluent, and lie on exactly the same level as the trigeminal
foramina.
In the upper posterior region of the orbit there is a deep depression, which no
doubt lodged a gland; from this a shallow groove runs forward along the upper edge
of the interorbital region. This groove marks the course of the olfactory nerve, and
immediately above it the great penthouse-like roof of the orbit arises. The whole of
the inner and posterior part of this is formed by the orbital plates of the frontals,
while externally it is completed by the enormous supraorbital plate of the lachrymal,
which in this specimen has been somewhat displaced downward on both sides, so that
the skull, seen from above, appears as if it possessed large supraorbital depressions,
such as occur in the Petrels, &c.; though, as a matter of fact, no such depressions
existed (see Plate XIV. and Plate XV. fig. 2).
The lachrymal (/ac.), as just mentioned, bears a very large supraorbital plate, which
extends nearly to the postorbital process, terminating posteriorly in a blunt rounded
angle. Its inner border was closely applied to, if not united with, the outer border of
the frontal, and its outer border forms the upper margin of the orbit. Anteriorly its
ventral surface is deflected, and the body of the bone turns inward and downward, and
is produced ventrally into a nearly vertical bar of bone, which in its ventral portion is
flattened laterally to a thin plate, and at its lower extremity articulates with the upper
border of the jugal. A little above its middle point this bar has a small backwardly
projecting process, but there does not seem to be any trace of a separation into two
elements at this point, such as was formerly suggested!. The form and relations
of this structure are almost precisely the same as in Serpentarius, or even more
Accipitrine, as in the Eagles, e.g. the Harpy Eagle; and I can see no justification for
Ameghino’s description of the orbit as open anteriorly in Phororhacos any more than
it is in the birds just mentioned, in which also this process reaches the jugal.
In front of the preorbital process of the lachrymal is the antorbital fossa, which
is triangular in outline, its antertor and upper angles being rounded off. Its anterior
border seems to be formed by a downgrowth of the nasal meeting an upward process
from the maxilla; there are, however, a number of cracks in this region, which render
its interpretation difficult, but by comparison of the two sides some degree of certainty
can be attained.
The nasals consist of a posterior body, which unites behind with the frontals and
supraorbitals ; but it cannot be determined whether the two meet in the median
| < This,’ 1896, p. 6.
60 MR. C. W. ANDREWS ON THE SKULL AND
line or whether they are separated by the facial process of the premavxille, the
junction with which is quite indistinguishable. It appears very probable that the
premaxille ran in between the nasals and formed at least the upper portion of the
high arch of the culmen between the nostrils, while the upper borders of those
openings are formed by processes of the nasals; these bones also send processes
downward and forward, forming the posterior border of the nares, and meeting the
maxilla below as already mentioned. The angle between the anterior and inferior
processes is rounded and has a thickened edge. The form and relations of the nasal
are best seen on the left side of the specimen.
Immediately within the anterior border of the antorbital fossa, and united with the
inner face of the downward process of the nasal, is the base of a broad band of bone (eth.),
which runs vertically down the middle of the fossa parallel to the downward process
of the lachrymal. The upper end of this band of bone widens out into a fan-shaped
expansion, the upper edge of which is slightly overlapped by the anterior margin of
the lachrymal. ‘The lower end, also somewhat expanded, is cancellous, and unites
with the upturned inner edges of the maxillo-palatine plates, with which the anterior
ends of the palatines are indistinguishably fused. The narrowest point of this bar of
bone is about its middle, below which it bears a backwardly-directed triangular
process. A similar and similarly-situated bar in the ethmoidal region occurs in
Cariama and also in some Accipitrine birds: e. g., in a modified form, in Serpentarius.
The jugal arch is oval in section posteriorly, but in front it is compressed so as to
form a broad, thin, vertical bar of bone. On the inner side of its posterior end
there is a rounded knob which fits into a corresponding pit on the outer surface of the
quadrate. Its anterior end seems to overlap the posterior prolongation of the maxilla.
The downward process of the lachrymal articulates with the upper edge of the jugal
close to its anterior end.
The form and relations of the maxilla to the surrounding bones are very difficult to
make out. As just mentioned, its posterior extremity seems to be overlapped by the
jugal, and in front of this it no doubt formed the edge of the beak for some distance.
On its inner side it widens out suddenly, the expanded portion being directed
downward and inward, and forming the large maxillo-palatine plate, with which the
anterior end of the palatine unites completely. The maxillo-palatine plates have a
thickened hinder border and are highly convex from side to side, and their upturned
inner margins unite with the downwardly-directed bar of the ethmoid. ‘The posterior
border of each maxillo-palatine plate forms a conunuous curve with the lower edge of
the jugal arch. On either side of the middle line of the palate they form a long ridge
which projects considerably below the tomium ; the fused anterior ends of the palatines
must also take part in the formation of this ridge. In the middle line they unite
for a space of about 3°5 cm., in front of which they are separated by a narrow cleft
2-5 em. long; how far they extend in front of this cannot be determined, nor is it
SKELETON OF PHORORHACOS INFLATUS. 6]
possible to make out the extent and relations of the facial parts of the maxille, and it
is therefore uncertain what share, if any, they may take in forming the boundary of
the nostrils.
The premazille no doubt form the greater part of the enormous beak, but their
exact boundaries are not distinct. _ Superiorly they send out fused nasal processes, which
form at least the upper part of the high bar of bone between the nostrils and perhaps
extended to the frontals. Anteriorly the bodies of the premaxille form the high laterally-
compressed anterior section of the beak, the tip of which forms a downwardly-directed
hook, about 3:5 cm. long and small compared to the bulk of the whole beak. From
the sides of the base of this hook a pair of parallel ridges run back on to the palate ;
their posterior portion is separated from the tomium by a deep groove, into which the
edge of the mandible fits when closed ; on their inner side also there is a deep channel,
divided by a median ridge, which just behind the decurved hook rises into a prominent
tuberosity, at first described as a tooth.
The cutting-edge of the beak forms a double curve; it commence a little behind the
point and first forms a strongly convex, prominent edge, sharp in front, but more
rounded behind ; posteriorly it is very slightly concave, and is a continuation of the
line of the lower edge of the jugal arch. The anterior convex portion is no doubt
formed by the premaxille ; the posterior is maxillary. In the premaxillary region the
narrow palate is closed, except for a few small median foramina, and is deeply
concave from side to side.
The beak, as a whole, is very deep from above downward, strongly compressed
laterally, and its whole surface covered with angular and vascular impressions, which
indicate that in life it was covered with a very thick horny sheath ; the whole must have
formed an extremely formidable weapon, whether for attack or defence.
The upper portion of the cranial region of the skull is much flattened (Plate XV.
fig. 2). Posteriorly it is greatly constricted by the great temporal fossa, which are
separated in the middle line by an interval of about 12 mm. only. In front of these
it widens out and reaches its greatest width at the level of the postorbital processes.
In front of these it again narrows, and anteriorly the frontals join the nasals, but their
limits are not very clear. The orbital borders of the frontals have thick roughened
edges, and probably the great supraorbital plates of the lachrymals, which now lie
slightly below the level of the frontals, may, in their normal condition, have united
with them throughout their whole extent; but, since their position is symmetrical
on the two sides, it is possible they may be in their natural position. As already
mentioned, this depression of the suprasrbital plates gives the skull, when looked at
from above, the appearance of having possessed deep supraorbital fossi.
The Palate (Plate XV. fig. 1).—The most important result of the careful removal
of the matrix from the skull is that the structure of the palate is completely revealed,
the bones being perfect and very little displaced. The taxonomic importauce of this
62 MR C. W. ANDREWS ON THE SKULL AND
region is very great, and examination of it in this bird on the whole tends to support
the views previously put forward.
The pterygoids (pt.) are stout, laterally compressed bones, of considerable depth. At
their posterior end they bear a cup-like depression, which receives a corresponding
tuberosity on the inner border of the quadrate, and on their upper border, one-third
of their length from the posterior end, there is a stout process, oval in section and
projecting backward, upward, and inward; it terminates in a facet, which articulates
with a corresponding surface on the base of the skull. These upper processes of the
pterygoids occupy the position, and perform the function, of the basipterygoid processes
of some birds. Anteriorly the pterygoids converge and just meet below the rostrum:
their anterior ends are abruptly truncated at right angles to their long axis, and form
facets for articulation with the palatines; these latter are very large bones; posteriorly
they bear short, broad pterygoid processes, which meet in the middle line and articulate
with the pterygoid facets just mentioned. The median union of the palatines extends
forward about 3 cm., and is marked by a slight ridge: in front of this their median
borders diverge and are notched by the hinder borders of the internal narial slits. In the
interval between the palatines no trace of the vomer is preserved, except that close to the
anterior end, where they join the maxillo-palatine plates, there are traces of a median
plate, which is probably a portion of that bone. The body of the palatine is very large,
and consists of two lamella separated by a deep valley and projecting almost vertically
downward. Of these the outer is much the larger, and forms a great ventrally-
directed flange. Its inferior border is somewhat thickened, and meets. the posterior
border, which also is thickened, nearly at right angles. The angle thus formed is slightly
everted, and a short distance above it the posterior border slopes gently backward to
the pterygoid process. The inner lamella is very small; its ventral border is thickened
and is continued backward to meet its fellow on the middle line, forming the median
ridge above noticed.
‘The anterior portion (maxillary process of the palatine is formed by the widening out
of the external lamella; this fuses with the maxillo-palatine plates, and together with
them forms the prominent longitudinal ridge projecting below the level of the tomium,
as before described. ‘The bird in which the palate most nearly resembles the fossil
in this respect is Yiomedea, in which the disposition and relative proportions of these
ridges and the form of the cutting-edges of the beak are very similar. Dorsally
the palatines send up a thin triangular plate of bone on either side of the rostrum and
embracing it.
The quadrate (Plates XIV. and XV. fig. 1, also text-figure, p. 63) articulates with
the skull by two distinct condyles sharply separated one from another by a deep pit,
at the bottom of which is a large pneumatic fossa, hollowed out in the neck of
the bone. ‘The antero-external capitulum (O.H.) is much the larger; it is criangular
in outline, convex from above downward, and slightly concave from side to side ‘The
SKELETON OF PHORORHACOS INFLATUS. 63
inner capitulum (I.H.) is borne on a thin, backwardly-directed buttress of bone; it has
two facets, one nearly flat, looking upward and inward, the other convex and directed
downward. ‘The neck of the bone is deeply excavated by pneumatic fossz, both on its
anterior and posterior face, and also on the inner side at the base of the orbital process
(O.P.). This latter is large; its upper and lower borders are nearly parallel, and it
terminates in a roughened surface; its length is about 17 mm.
Fig, 2.
Left quadrate of Phororhacos inflatus Ameghino.
1. From outer side.
2. Articular surface for mandible.
3. From front.
LH., inner head; I., inner articular surface for mandible ; J.C., cup for end of jugal arch; O., outer articular
surface for mandible; O.H., outer head; O.P., orbital process; P.T., tuberosity for articulation
with the pterygoid.
The lower end is massive and bears two articular surfaces (I. and O.) for the mandible,
which are separated by a shallow groove running outward and forward from the postero-
internal border. ‘The form of the facets will be best understood from the figure (fig. 2,
above). On the inner border of the bone, just beneath the base of the orbital process,
there is an elongated tubercle (P.T.) for union with the hinder end of the pterygoid ;
and on the outer side, immediately above the surface for the mandible, is the deep
cirenlar cup (J.C.) with clearly defined border, which receives the posterior end of the
quadrato-jugal.
The mandible (Plate XIV.) is specially remarkable for its very massive structure.
The relatively narrow symphysis is about 9 cm. long; its ventral border is convex, its
dorsal nearly straight, and but slightly upturned at the top. The upper surface
VoL. XV.—ParT 11. No. 2,—October, 1899. I
64 MR. ©. W. ANDREWS ON THE SKULL AND
is deeply concave from side to side, giving the symphysial region a spout-like form.
Behind the convexity of the symphysial region the ventral edge of each ramus is
anteriorly slightly concave and posteriorly rather strongly convex. The upper borders
of the rami are at first convex and sharp-edged, behind this, as far as the small coronoid
process, they are straight and blunt. Behind the coronoid process the edge slopes
down to the articular cup for the quadrate. The facets on this are two in number, one
external and elongated, one inner and rounded in outline ; both are concave from before
backward, and they are separated by a deep fossa, but no pneumatic foramina are
present. There is a large and very prominent internal angular process; the inferior
angle is broken away, but it can only have formed merely a slight projection. Just
beneath and in front of the coronoid process the mandible is perforated by a very large
vacuity.
Comparison of the Skull of Phororhacos with that of other Birds.
Comparison of the skull of Phororhacos with that of the Struthious birds at once
excludes the possibility of any affinity between it and them. The form of the palate,
in which, as already described, the palatines and pterygoids both articulate with the
rostrum, the reduction or absence of the vomer, and the double head by which the
quadrate articulates with the skull, are all points not occurring in any Ratite bird.
It then remains to determine what are the closest allies of Phororhacos among the
Carinate, a question of which any satisfactory solution is very difficult.
Taking the palate first for comparison, we find that it presents in some respects a
very remarkable resemblance to that of certain ‘Tubinares, especially the Albatross
(Diomedea). In this bird, as in Phororhacos, the maxillo-palatine plates together with
the anterior region of the palatines form on either side a very prominent ridge,
projecting considerably below the level of the tomium, the form of which also is very
similar in the two birds, The palatines and pterygoids are also similar to those of the
fossil in their relations one to another and to the rostrum ; but the palate differs much in
form, the internal lamina being much the larger, while in Phororhacos the reverse is the
case. There are a number of other important differences: thus, in Diomedea the palate
is schizognathous and the anterior end of the large vomer appears upon it; there are
large supraorbital fossee. In Phororhacos the palate is desmognathous, the vomer is
inconspicuous, and there seem to have been no supraorbital fossee whatever, although
slight displacement of the large supraorbital plates of the lachrymal at first sight
gives the skull the appearance of having possessed them. Moreover, the form of the
antorbital fossa and that of the quadrate are different in many respects. In spite of
a certain similarity between the skulls, it does not seem that any close affinity exists
between Phororhacos and the Tubinares, and in fact, as will be seen below, the structure
of the limb-bones and pelvis lends no support whatever to any such relationship.
Comparing the fossil with the skulls of various desmognathous types, one is at once
SKELETON OF PHORORHACOS INFLATUS. 65
struck with its Accipitrine aspect. This, of course, depends chiefly on the form of the
beak, but is heightened by similarity in several points of detail, e. g. the form of the
lachrymal and of the antorbital fossee. Closer examination shows, however, that the
structure of the palate is very different, particularly in the relations of the anterior
end of the palatines to the maxillo-palatine plates; the form of the articulation
of the quadrate with the mandible is also widely different. Of the Accipitrine birds,
Serpentarius approaches the fossil most closely both in structure of the anterior region
of the palate and in the existence of an articulation between the pterygoids and the
basis cranii. In the Storks the palate is in several respects like that of the fossil,
particularly in the relations of the palatines to the maxillo-palatine plates; the distal
articulation of the quadrate is also similarly constructed, but is relatively much wider
from before backward. On the other hand, the lachrymal bears neither a long
descending antorbital process nor a large supraorbital plate, and the antorbital fossa
is different.
Although the Gruiformes are typically both schizognathous and schizorbinal, it is
remarkable that Cariama and Chunga, which in external appearance and habits so
much resemble Serpentarius, should have become both desmognathous and holorhinal,
like Accipitrine birds. The existence of these characters, therefore, in the skull of
Phororhacos cannot be regarded as a bar to the relationship between it and the
Gruiformes, suggested in a former paper and supported by many structural peculiarities
of the pelvis and hind limb. In fact, since the general line of specialization of Phoro-
rhacos is similar to that which produced the Cariamide, the occurrence of these
characters might have been expected. In some points, indeed, ¢.g. in the form of the
quadrate, Phororhacos differs less from the typical Gruiformes than Cariama does.
Further discussion of the affinities of this remarkable form is best deferred till after
the remainder of the skeleton, as far as we know, has been described.
The dimensions of the skull are :—
cm.
Length from anterior end of premaxillze to paroccipital process. . . . 337
5 555 to centre of occipital condyle . 30:0
\RVicliln eae MOC OMA OROIGESITIS 6 co 56 o 6 09 G9 6 6 o 0 o Go Oo Ane
5 Ci Gli a Womyporell OSES 6 o o 5 0 6 op oo oe oO
5 Gs WOStOnomnll ROGENSES 9 co np » 0 0 6 5 0.8 oc 0 0 oo EK
5, (least) between temporal fosse@ - 5 2 2 s © « es ow es (OO
» Opposite posterior edge of nostril . . . . . «. - » «. - « 388
Depth of upper jaw at middle of nostril » . . . . «. ~~ « - ~« ILS
* i ay is Tmale OWNES 6 6 6 50 0 60 0 6 6 6 oF Ailey
ILE Or Quays Orns ¢ @ s 96 o 6 ¢ 60005 60 0 Zl
WadthvoL upper aw,atuiid dleypornitien sien) utc ci onnte! tli EER OsO
SIGN OF /ORHMED UG oo 9 0 09 0 06 9 Po OK CNY
Witch or clistall Gael or @uachenti® 5 5 56 o 6 56 6 06 6 6 5 6 Go Bh
L2
66 MR. C. W. ANDREWS ON THE SKULL AND
Fig. 3.
My
Ip Zp
ORL Paya
SB,
Uf D
Ny ah:
Poe Be
Diagram of skeleton of Phororhacos inflatus Ameghino, to show the relative proportions of the bones.
The portions of the skeleton preserved are shaded.
(About 3 natural size.)
SKELETON OF PHORORHACOS INFLATUS. 67
The dimensions of the mandible are :—
cm.
WEG 6 6 1G eR 6 Boe 2 de 6 oo oe UY
IDEN OMNIS 5 6 5 6 6 16 6 6 56 0 6 oD oo a) 6D
Depth of ramus at coracoid process . . . . . ..... 49
Wichin@renmonle emt og ¢ 6 o 6 6 6 6 6 6 oO 8
The hyoid (text-figure, below) is very well preserved, a most exceptional, if not
unique, circumstance in a fossil bird. The anterior element, the entoglossal
(ENT.GL.), is very large and clearly has originated from the union of two rod-like
elements, the ceratohyals, which are united by a thin plate of bone. On the ventral
surface the division between the rods is marked by a deep furrow, deepening posteriorly,
but the dorsal surface seems to have been regularly convex from side to side. ‘The
anterior end is somewhat broken; the posterior is deeply divided by a double
Fig. 4.
Hyoid of Phororhacos inflatus Ameghino, slightly reduced.
1. From below.
2. Basihyal from side.
2. Section of entoglossal at point A.
ENT.GL., entoglossal ; B.Hy., basihyal (first basibranchial) ; c.pr., ceratobranchial.
articulation for the basihyal (basibranchial 1) (B.uy.), and laterad to this the outer
angles are produced into short backwardly-directed processes.
The first basibranchial (B.Hy,) (basihyal) is a short bar of bone, the thinner lateral
borders of which seem to have been broken away. Its dorsal surface is concave from
before backward, and on its anterior half from side to side also; ventrally it is convex
from before backward. ‘The anterior end bears a double surface for articulation with
the entoglossal, separated by a slight ridge, which at its ventral extremity is produced
into an anteriorly-projecting prominence. Posteriorly there are two irregular surfaces
for the ceratobranchials, meeting at an angle of about 120°. The first basibranchial is
not prolonged backward into a urohyal, nor does it, as far as can be seen, bear any
surface for the articulation of a distinct median element: if such were present it must
68 MR. C. W. ANDREWS ON THE SKULL AND
have been small and have articulated with the bases of the ceratobranchials (c.BR.).
These are thin, slightly curved rods of bone, the anterior ends of which are enlarged
and bear the surface for the articulation with the first basibranchial. The posterior
ends are also slightly expanded and terminate in convex facets, apparently for union
with another element.
In the Albatross the entoglossal is relatively very much smaller, the first basibranchial
(basihyal) broad and short, and prolonged backward into a long urohyal, so that the
articular surfaces for the ceratobranchials are widely separated.
In Grus the entoglossal is very long, slender, and unossified ; the first basibranchial
(basihyal) is also long and slender, but, as in the fossil, the ceratobranchials articulate
with two contiguous surfaces on its hinder end, and the uroglossal is distinct and
articulates between the bases of the ceratohyals.
In Cariama the two elements forming the entoglossal are partly separated by a
median foramen and are prolonged into postero-lateral processes behind the articulation
with the first basibranchial. his is a short bar ef bone terminating in a pair of
surfaces for the ceratobranchials. The urohyal is very small and is not in contact
with the first basibranchial.
In the Birds of Prey the ossified portion of the entoglossal is, as a rule, deeply bifid
anteriorly, the two elements forming it being united for a short distance only. The
first basibranchial is somewhat elongate and is prolonged backward into a uroglossal
process.
In the Storks the entoglossal is imperfectly ossified, and the first basibranchial is
similar to that of the Raptores.
On the whole, the hyoid of Cartama approacnes the fossil most nearly.
The Coracoid. (Plate XVII. fig. 1.)
The coracoid is particularly remarkable for its great length and slenderness, and for
the reduction of the acrocoracoid (a¢.) process.
At the sternal end the bone is almost 32 mm. wide; its anterior face is convex
from side to side, and the posterior flat or rather concave. ‘The articular facet for the
coracoid groove of the sternum is large and well defined; it extends from side to side
of the distal end of the bone both on the anterior and posterior face. At about
its middle it is overhung by a very prominent tuberosity, from which a shelf-like
projection runs to the inner border of the bone; the union with the sternum must
have been very strong.
There is no distinct processus lateralis (p.l.), its place being occupied by the convex
outer border (p./.) of the expanded distal end. On the anterior surface a linea aspera
runs from the inner angle upward and inward, joining the ridge marking the outer
border at about + of the length of the bone from its distal end. ‘The shaft towards its
upper end is slightly compressed laterally. ‘The total length of the bone is 157 mm.
SKELETON OF PHORORHACOS INFLATUS. 69
The proximal end of the bone is peculiar. The acrocoracoid is almost completely
absent, the upper end of the bone rising scarcely at all above the scapular surface.
From the acrocoracoid region a ridge runs down the antero-internal border of the bone
and forms the inner wall of a deep channel, which is bordered externally by the
prominence formed by the surface for the scapula and the small procoracoid process.
On this descending ridge there is, at some distance from its upper end, an elongate flat
facet with which probably the clavicle articulated.
The surface for the scapula is a large, deeply concave area, nearly circular in outline ;
below its internal border arisesa small process or hook of bone (broken in the specimen
figured), which overhangs the groove above described and is the procoracoidal process.
The glenoid surface is a smooth, slightly convex area, the lower border of which
forms a prominent overhanging lip on the postero-external side.
The coracoid just described is one of the most remarkable boues of the skeleton.
Its extreme elongation and the peculiar structure of its upper end, particularly the
almost complete suppression of the acrocoracoid, are paralleled, so far as I am aware,
only in the coracoid of the flightless Aptornis. The coracoid of this bird differs,
however, in several important particulars, ¢.g.in the large size of its procoracoidal
process, the presence of a supracoracoid foramen, and in the complete fusion of the
lower extremity with the sternum.
Fiirbringer has regarded the absence of the acrocoracoid as diagnostic of the Ratite,
and has suggested that the term “ Platycoracoidee” should be used for them in
contradistinction to the Acrocoracoidee, or Carinate. The oceurrence of this
character in two such distinctly Carinate types as Apfornis and Phororhacos, however,
invalidates this distinction, and tends to show that the presence or absence of the
acrocoracoid, like that of the keel of the sternum, is in some way dependent on the loss
of flight, and probably also on the increase in the bulk of the body.
The articulation with the sternum is particularly well developed, and in no bird could
the prominent tubercle and shelf above described be found developed to the same
extent. No trace of the sternum is preserved, but, judging from the coracoids, it was
probably of considerable size.
Scapula. (Plate XVII. fig. 2.)
Of the scapula no perfect specimen is preserved, but the most complete, here figured,
wants the distal portion of the blade only. The surface for union with the coracoid is
highly convex and roughly circular in outline; it is much roughened, and the union of
the two bones must have been a very close one. The glenoid surface, which is slightly
convex, is raised on a prominence which projects considerably below the inferior border
of the blade. There is a very large acromium process. The portion of the blade
preserved is somewhat rounded externally and flat internally.
The width of the proximal end of this bone is 30 mm.
70 MR. C. W. ANDREWS ON THE SKULL AND
The Bones of the Wing.
The exact length of the wing cannot be determined, owing partly to the incomplete
state of the humerus and partly to the absence of the phalanges ; but it is certain that,
in proportion to the bulk of the bird, it had undergone extreme reduction and must
have been entirely useless for purposes of flight. At the same time the stoutness of
the bones and the distinctness of the muscle-impressions indicate that it was in some
way functional, and the presence of a series of tubercles along the posterior border of
the ulna shows that it was provided with a series of large quill-feathers, the remiges.
It may have been employed to assist the bird in running, or possibly as a shield,
somewhat in the same way as the Secretary-bird is said to use its wings, although for
this latter purpose it would appear to have been too small.
Of the humerus (Plate XVII. fig. 3) only the distal end is preserved in the type-
skeleton. Init the radial (r.) and ulnar condyles (w.) are of the ordinary avian form,
but are less distinctly separated than usual, the intercondylar groove being almost
obsolete. The surfaces of the condyles are rough, and probably a considerable pad ot
cartilage intervened between them and the bones of the forearm. The distal border of
the bone is very oblique, owing to the presence on the ulnar side of a pointed process
projecting some distance beyond the ulnar condyle (w.). ‘The anterior face of the bone
above the condyles is deeply concave, and the portion of the concavity close to the ulnar
border of the bone contains the well-marked elongated impression of the brachialis
anticus (b.a.). Posteriorly there is a shallow olecranon fossa.
The ulna (Plate XVII. fig. 4) is a short, very stout bone ; its shaft is slightly curved
and triangular in section, the sharpest angle forming its posterior border. At its
proximal end the impression of the brachialis anticus is not very clearly marked, but
the surface for the humero-cubital ligament is very distinct. There is a well-developed
olecranon process (o0/.), the end of which bears a rugose surface for the insertion of the
triceps tendon, and from this a slight ridge runs to another rugosity (for the insertion
of the long head of the triceps) lying immediately below the edge of the radial
glenoidal surface. On the postero-internal border, immediately below the articular
surfaces, is a tuberosity for the insertion of the biceps tendon. The sharp posterior
border of the bone bears a series of well-marked tubercles, indicating the existence of
large quill-feathers (secondary remiges) in the wing: there are about seven of these
tubercles.
Towards its distal end the shaft loses its triangular form and becomes circular in
section.
The distal articular head is large, and on its outer surface there are impressions
marking the origin of various flexor muscles of the digits, and a groove for the passage
of tendons. ‘The length of the ulna is approximately 110 mm.
The radius (Plate XVII. fig. 5) is represented by the proximal end only.
In the metacarpus (Plate XVII. fig. 6) the proximal articular surface is broad and
SKELETON OF PHORORHACOS INFLATUS. 71
only slightly grooved at its posterior end; there is no fossa into which the cuneiform
fits when the manus is flexed upon the forearm; the so-called pisiform process is
almost absent. The prominence formed by the fused first metacarpal is stout and
short, and terminates in a roughened surface which may possibly have formed the base
of a short horny spur or knob. Distally the first metacarpal terminates in a tubercle
for articulation with the phalangeal. The second metacarpal (me. 2) is short and
thick, compressed laterally and slightly curved. On its upper surface there is a nearly
longitudinal, faintly marked groove, and on the anterior surface of the distal end is
another short groove for tendons.
The third metacarpal (me. 3) is much more slender than the second; it is strongly
curved and so compressed that it forms a mere band of bone. On its ventral surface at
the proximal end is a prominent tubercle (¢.), which will be referred to below ; distally
it is fused with metacarpal 2 in the usual mauner. The distal extremities of the fused
metacarpals bear each an articular surface for their phalangeals, as in most birds. The
length of the metacarpus is 76 mm.
The reduction that has been undergone by the bones of the wing makes their
comparison with those of other birds difficult,
The distal portion of the humerus differs widely from the Albatross, in which the
impression of the drachialis anticus is in a different position and there is a large
ectepicondylar process. In the Birds of Prey the depression on the anterior face above
the articular surfaces is always much less than in Phororhacos, and the impression
of the drachialis anticus is more median ; in some the distal border is somewhat oblique
and there is a trace of the distal process.
In Ciconia the distal border is not oblique and the position of the drachialis impres-
sion is quite different.
In Grus the differences are the same as in Ciconia.
In both Cariama and Psophia also the distal border is only slightly oblique (in
Cariama there is a trace of a blunt distal process on the ulnar side).
The humerus to which I find most resemblance in the form of the distal end is that
of Diaphorapteryx, the large extinct Rail of the Chatham Islands. In this the distal
border is oblique, there is a fairly prominent distal process, and the form and position
of the drachialis anticus impression is similar. The humerus of Apftornis is also some-
what like the fossil, but in it reduction has gone further, the division between the radial
and ulnar condyles being still more indistinct and the distal process small.
The short, stout ulna is not very much like that of any of the birds referred to in the
comparison of the humerus, but approaches that of Psophia most nearly.
The proximal end of the right radius is preserved. Its articular surface is oval in
outline, and the bicipital tuberosity is strongly developed. It is almost identical with
the same element in Cartama, but the characters of the radius are not sufficiently different
in the various groups to be of much importance in determining affinities.
VOL. XV.—PaRT 11. No. 3.—October, 1899. M
12 MR. C. W. ANDREWS ON THE SKULL AND
The metacarpus differs widely from that of Diomedea, Grus, and Ciconia, and is most
like that of Psophia and Cariama.
In both these birds, as in Phororhacos, there is on the ventral edge of the third
metacarpal, close to its base, a small process forming a projection (¢., fig. 6, Plate X VIL.)
on the palmar aspect of the manus. I have not observed this in any but these birds,
and of them it is most strongly developed in Cariama, in which, however, there is a
prominent pisiform process.
On the whole, the evidence of the wing-bones seems to point to relationship to the
aberrant Gruiform birds Cariama and Psophia: but in the humerus there is resemblance
to such Rails as Ocydromus, Diaphorapteryx, and Aptornis, in which also the wings
have undergone more or less reduction; in this case the similarity may merely be due
to convergence in degeneration.
Sacrum and Pelvic Girdle. (Plate XVI.)
The vertebre uniting to form the so-called sacrum (fig. 3) (synsacrum, Parker) are
about fifteen in number. The most anterior articulated with the last free dorsal by the
broad saddle-shaped surface of the centrum and by very large anterior zygapophyses.
The neural spine of this vertebra is high and its summit is overlapped by and fused
with ossified fascize which form a forward extension of the supero-anterior angles of the
ilia; looked at from the side a great part of the spine is exposed to view, the anterior
borders of the ilia only just overlapping its hinder edge; between the spine and the
ilia there is on either side a chink-like ilio-neural canal.
Tubercular and capitular facets for a free rib are present, and both are roughly
semicircular, the convexity being directed upward and forward. The capitular facet,
borne on a slight parapophysial elevation, is close to the anterior end of the centrum
and at the lower end of a ridge which runs upward and backward, bearing near its
upper end the tubercular facet. From this a bar of bone, apparently equivalent to the
zygapophysial bar of a free vertebra, runs back and unites with the antero-inferior
margin of the ilium; from the tubercular facet a metapophysial ridge runs forward on
to the zygapophysis. Just behind and above the capitular facet there is a pneumatic
fossa of moderate size. The ventral surface of the centrum is pinched up so as to form
a sharp median ridge.
In the second vertebra the neural spine is completely concealed by the ilia, and
running upward and backward from the anterior end by the centrum there is a thick
ridge which, near its lower end, bearsa facet for the capitulum of a rib, and at its upper
end abuts against the lower border of the ilium; here also there is a median ventral
ridge. These vertebre are probably thoracic.
The centra of the next few vertebra have been destroyed, but opposite the acetabulum
they are again preserved. In this region the median ventral ridge is replaced by
a median groove defined by slight ridges, and disappearing opposite on the second of
the true sacrals As far as opposite the middle of the acetabulum the vertebre may be
SKELETON OF PHORORHACOS INFLATUS, 73
regarded as lumbar; the last of these bears a broad parapophysial process which abuts
against the inner border of the ilium. Behind this there seem to be two or three
Jumbo-sacrals (Mivart) in which no parapophyses are present and, except perhaps on
the hindermost, no diapophyses, unless these are directed dorsad. In this region the
pelvis is very narrow, so that the lateral acetabular or anterior renal fosse are extremely
small.
Behind the lumbo-sacrals are the true sacrals, two in number. Both these possess
ventral processes, which, no doubt, are formed by parapophyses + sacral ribs. In the
first the process is stout, much expanded at its outer end, and directed outward, upward,
and somewhat forward. In the second this process is very slender and quickly fuses
with the middle of the upper process (diapophysis), which in this vertebra is stout and
directed outward and forward. The diapophysis of the anterior sacral is slightly
in front of, and about 1 cm. above, the parapophysis ; it is directed backward, outward,
and upward. ‘The two diapophyses are separated by a nearly circular space, and their
expanded outer ends, together with that of the ventral process of the anterior sacral
fuse into a common mass of bone, which abuts against and fuses with the inner surface
of the ilium immediately behind the acetabulum.
On the vertebral centra, at the level of the second sacral rib, there is a median
hemal ridge, which is continuous as far as the third of the urosacrals which bear
transverse processes.
Behind the last sacral there is an interval in which no transverse processes are
present: then they reappear, the first being very slender and directed backward at a
very acute angle with the vertebral column; at its distal end it fuses with the succeeding
process. ‘This also is directed backward, but is much stouter than the last and greatly
expanded at its distal end, where it abuts against the inflected portion of the ilium
forming the floor of the pocket-like renal fossa. The next transverse process is very
broad, and likewise unites externally with the ilium ; it is separated from the processes
in front of and behind it by oval foramina. Behind this there are four pairs of similar
processes borne by vertebree with long, narrow centra having a slight hemal ridge. The
free end of the last fused centrum is flat or slightly convex. On the dorsal surface the
neural spines of the postacetabular “ sacrals” form a prominent median ridge, separated
from the ilia, anteriorly at least, by a narrow groove ; posteriorly there are several pairs
of slit-like interosseous foramina.
The pelvis, seen from above (Plate XVI. fig. 1), appears very long and narrow, the
postacetabular region not being expanded to any great extent. Along the middle
line runs a prominent ridge, formed in front by the fused upper edges of the ilia and
behind by the united neural spines of the postacetabular vertebre. Opposite the
hinder border of the acetabulum is a transverse ridge which, at its outer ends, rises
into a pair of very prominent supra-trochanteric crests; these, with the anti-trochanters,
form prominent lateral projections. On each side of the median ridge in the post-
acetabular region there is a narrow groove separating it from the upper edges of the
M2
74 Mk. C. W. ANDREWS ON THE SKULL AND
ilia, and at the bottom of this there are traces of one or two pairs of interosseous
foramina. The ilia terminate posteriorly in prominent ilio-caudal processes, which
project some distance beyond the last fused caudal. From these processes there runs
inward and forward on either side a ridge which, after continuing a short distance
parallel to the vertebral axis, runs downward and outward to the upper angle of the
ischiadic foramen; this ridge seems to be the hinder part of the ilio-lateral ridge,
which is so strongly developed in many birds, é. g. Rails. Anterior to it the dorsal
surface of the ilia are evenly convex from side to side.
Viewed from the front, the most conspicuous points are (a) the relatively large size
of the centra of the sacral vertebre; (8) the extreme lateral compression, the ilia
meeting in the iliac crest at an angle of not more than 10°; (y) the very narrow slit-
like opening of the ilio-neural canals; (8) the very prominent supra-trochanteric crests
which, from this point of view, completely hide the whole of the dorsal surface of the
postacetabular region.
In a posterior view the supra-trochanteric crests and anti-trochanters completely
conceal the whole pre-acetabular region. Behind them we have first the median ridge
of the sacrum, on each side of this a shallow groove, and external to this again the
ilia, the regular downward curve of which forms an are of a circle, and below these
the ischia curving outward and diverging one from another. This portion of the pelvis
has been crushed laterally, but not to any great extent.
Looking at the pelvis from the side (Plate XVI. fig. 2) it will be seen that the
acetabulum is considerably in front of the middle of the pelvis, and measured from
the middle of the acetabulum the pre-acetabular portion of the ilium is 140 mm. in
length, the postacetabular 265 mm., or roughly as 10 to 19. ‘The acetabulum itself is
nearly circular in outline, its antero-posterior diameter being slightly the greater
(acet.); through it the arches of the lumbo-sacral vertebre are visible. Its antero-
superior border forms a projecting lip, and the anti-trochanter, the lower border of
which is excavated by a narrow fossa, is large and very prominent, and, except for a
narrow groove, is continuous above with the supra-trochanteric crest. Anterior to the
acetabulum is the slightly concave and nearly vertical gluteal fossa of the ilium, below
the edge of which the centra of the pre-acetabular sacral vertebre are visible.
The ischiadic foramen (¢s.f.) is very large and roughly ovoid in outline. Just within
its antero-superior angle, and bordered anteriorly by the anti-trochanter and superiorly
by the edge of the ilium, there is a pocket-like fossa, which internally is separated by a
slight ridge from the mass of bone formed by the fusion of the processes of the true
sacrals. The anterior edge of thei schiadic foramen is continued downward as a ridge
crossing the shaft of the ischium and terminating below in a prominent process
(styliform process), against the inner side of which the pubis was in close contact, thus
enclosing an obturator foramen (0.f.) which forms a narrow oval opening extending from
beneath the posterior half of the acetabulum as far back as the front of the ischiadic
foramen.
SKELETON OF PHORORHACOS 1INFLATUS. 75
Posteriorly the pelvis ends in the sharp hinder angle of the ischium, which is separated
by a semicircular bay from the ilio-caudal process of the ilium.
Seen from below (Plate XVI. fig. 3) the narrowness of the pelvis is even more
noticeable than in the dorsal view. ‘There is practically no internal iliac fossa, the
centra of the anterior “sacral” vertebre being very large and projecting far below
the edges of the ilia. The anterior renal fosse are extremely narrow, and are scarcely
visible from this point of view; they are separated by the processes of the two sacrals
from the enormous posterior renal fossee which, as in Rails and some other birds, are
prolonged backward into long pocket-like extensions, floored by an ingrowth of the
ilium. The only bird in which this ventral ingrowth of the ilium is developed to
anything like the length seen in Phororhacos is Fulica, in which the broad transverse
processes of three or four urosacral vertebre unite with it, while in the fossil there are
six or seven such vertebre.
The Ilium. (Plate XVL., il.)
In its pre-acetabular region the dorsal border of the ilium is convex, the ventral
concave, and in front the two are united by a nearly straight anterior border which is
slightly inclined forward. The ilio-pectineal process is broken, but seems to have
been small.
As already mentioned, the postacetabular region is considerably longer than the
pre-acetabular, from which it is sharply separated by the supra-trochanteric crest. It
is nearly equal in width throughout its length. Posteriorly it terminates in the
prominent ilio-caudal process, the lower part of which, however, may be formed by
the ischium: from the end of this process runs a ridge which seems to mark the
junction of the two bones, and certainly is continuous in front with the suture
between them. A second ridge, commencing at the hinder border of the bone close
to its sacral border, runs forward and then downward to join that just described
close to the ischiadic foramen.
The Ischium. (Plate XVI., zs.)
The share which this bone takes in the formation of the acetabulum cannot be
determined, owing to the complete fusion of the pelvic elements in that region.
Beneath the ischiadic foramen it forms a bar of bone 15 mm. wide, which, near its
proximal end, bears on its ventral edge a short, stout process which touches the pubis,
thus enclosing an obturator foramen. Beneath the ischiadic foramen the ischium
expands into a broad plate of bone, the outer surface of which is concave from
above downward, and the inner traversed by a prominent rounded ridge marking
the prolongation of the axis of the bone. The upper edge unites closely with the
ilium, the lower curves downward and outward and terminates posteriorly in an
angular process which projects slightly further back than the ilio-caudal process,
76 MR. C. W. ANDREWS ON THE SKULL AND
The Pubis. (Plate XVI. fig. 2, pw.)
The pubis is probably imperfectly preserved ; all that now remains is a slender rod
of bone arising beneath the middle point of the acetabulum, and running back and
terminating against the inner side of the styliform process of the ischium, thus closing
an obturator foramen as above mentioned. Whether when complete it extended further
back in the usual way cannot be determined.
Comparison with the Pelvis of other Birds.
Comparison of this pelvis with that of the Ratitee shows at once that Phororhacos
certainly does not belong to that group, though it may have been, and probably was,
‘‘Ratite” in the strict sense of the term. It is true that in its length, narrowness, and
the large development of the supra-trochanteric processes there is some similarity
to the pelves of Dromeus and Struthio, a similarity which, no doubt, is merely the
consequence of adaptive modification due to a like mode of progression. On the other
hand, in essential points of structure, such as the form and relations of the ischia and
pubes to each other and to the ilia, the structure of the ‘‘ synsacrum,” particularly
in the distinctive form of the two true sacrals and in the form of the renal fossz, the
pelvis in the fossil is very different from that of any Struthious bird. In Apteryx alone
the sacrum shows slight points of similarity.
Among the Carinate birds the pelves which show most resemblance in general outline
to the fossil are those of the Grebes and Hesperornis; but in both of these the pre-
acetabular portions of the ilia do not unite with the spines of the “sacrals ” to form an
ilio-neural crest, but remain separated from them by a considerable interval, while, on
the other hand, the postacetabular portions approach one another very closely, and
may even unite in the middle dorsal line; the exact reverse is the case in Phororhacos
and most other birds.
To the pelves of the Cranes and Rails the similarity is in many ways remarkable, and
probably indicates a real relationship with those birds. The form and character of
the renal fosse, particularly the peculiar pocket-like prolongations of the posterior
fossee, and the general structure of the “ synsacrum,” especially the distinctness of the
true sacrals, are almost identical with those described in the fossil. One difference,
however, must be pointed out, viz. that while in Phororhacos the postacetabular region
is the longer, in nearly all the Gruiformes the reverse is the case. In Fulica, however,
the post- and pre-acetabular regions are of nearly equal length, and in Cariama the
postacetabular portion is the longer, as in Phororhacos. -
The pelvis of Cariama is also similar in the form of its posterior border, the presence
of prominent supra-trochanteric processes, and, so far as can be ascertained, in the
relations of the pubes. It has, however, a well-developed pelvic escutcheon, the whole
postacetabular region being relatively wider than in Phororhacos. On the whole, so
far as the pelvis is concerned, I see no reason for changing the opinion expressed in a
SKELETON OF PHORORHACOS INFLATUS. ul
former paper ', that Cariama is probably a not very remote modern representative of the
extinct type.
From the Birds of Prey the chief points of difference are, that in them the post-
acetabular region is very short and is deflected, and there is no ilio-caudal process to
the ilium. In Serpentarius, however, the first two of these characters are much less
marked, and in this respect, therefore, it approaches the fossil, these characters being
probably merely correlated with the fact that it is a much better walker than the other
Accipitrine birds.
In the Storks the pre-acetabular portions of the ilia meet at a very obtuse angle,
the postacetabular region is very wide, and there are no recesses to the post-renal fosse.
In the Herons these fossee, though present in some, are small, the ilio-neural canals
are widely open behind, and the pelvic escutcheon is wide; certain Galliformes,
e. g. Phasianus and Francolinus, in which renal pockets are present, differ from the
fossil in much the same respect.
In the account of the skull some points of similarity with that of the Albatross were
referred to, but in the pelvis no similarity whatever can be detected. Among the more
notable differences may be mentioned the complete absence of pocket-like extensions of
the renal fossee, almost complete absence of supra-trochanteric processes, the backward
prolongation of the ischia and their mode of union with the ilia, and the indistinctness
of the true sacral vertebra.
Ameghino, in his manuscript catalogue, distinctly states that this pelvis and other
bones were found associated with the skull above described, and formed part of one
individual; and if this is the case, as there is no reason to doubt, the structure of
the pelvis seems completely to} outweigh whatever evidence of relationship with the
Albatrosses may be found in the skull.
The dimensions of the pelvis are :—
cm.
Hxtremeylen ge chimenne nc Wew col bls eeTh MCT MUliaayleg 200 (sm faa (om ata cee aol (MLO
Length ofilium .. . : Sy AS iota RM GLiccm ee athei cae ween ele)
Longitudinal diameter of menedie fone tieht A aE MEN er itt) Bae Ryser on NOs)
Wertical diameter ofaschiadie foramen.) 4) 4 =
Length of obturator foramen. . . . . Fae ey incites 2: eet ESO
Greatest height from hemal ridge of sacrum to ton Oealbeyocxess 6 6 o hs)
WAGED Oo EMOTES 5 5 6 6 6 6 6 9 6 0 0 6 9 o oo oo
op lb SURO NMSe OAEIS Go 5 6 6 0 0 0 5 6 0 4 Oo OD 10°6
» between upper edges of acetabula . . . ...... =. . «495
5) (least)) behind! supra-trochantericicrests) 50 3. 9) «| =.) 1) 638
r (UGASD) jorenestonlemmrio 6 6 5 5 5 6 5 8 bp oo oo BY
pe icentrum oftrstitused sacral vertebray. |e ll) tell leeks OLS
anetin @e emvemin 5b go 6 op ole eo Opole. Oo BSD
1 «The Ibis,’ 1896, p. 1; see also ‘Science Progress,’ vol. vy. (1896) p. 398.
78 _ MR. Cc. W. ANDREWS ON THE SKULL AND
Vertebre.
Of the vertebra, other than those forming the synsacrum, only three are preserved in
this specimen. Of these one is the last free dorsal and the others anterior free caudals.
The last dorsal has a short centrum, terminated by the usual saddle-shaped articular
surfaces, of about equal height and breadth. The ventral portion of the anterior two-
thirds of the centrum is pinched up to form a prominent heemal ridge, the summit of
which has been broken away. Laterally the anterior end of the centrum is widened
out by stout parapophysial prominences, bearing each an oval cup for the head of the
rib. The diapophysis is broken away: immediately beneath its base is a large
pneumatic fossa, and another occurs on the side of the centrum. The zygapophyses
are of the usual form; the neural spine is very massive, and has its anterior and
posterior surfaces roughened for ligamentous union with the spine in front and behind
it: its upper portion is broken away.
The caudal vertebree (Plate XVII. fig. 7) have centra about as long as their anterior
face is wide, slightly constricted in the middle, and bearing towards their hinder end a
pair of strong, backwardly-directed transverse processes, only the bases of which remain.
The anterior articular face is wider than high, the posterior about equal in the two
directions; the anterior surface is slightly concave, the posterior convex, but with a
median pit, which probably marks the primitive position of the notochord. The neural
arch does not extend quite to the hinder end of the centrum, and the neural spine is
greatly thickened at its upper end, which forms a flat bilobate surface, which was
evidently connected with the vertebr before and behind by tendons, probably more
or less ossified. —
The Femur. (Plate XVII. fig. 8.)
The head of the femur rises to a marked degree above the level of the trochanter,
the upper portion of the elevation being formed by a large blunt-pointed process, which
partly divides the deep pit for the ligamentum teres into two parts. Ventrally also the
articular surface is divided by a shallow groove into an anterior and a posterior lobe, so
that when looked at from the inner side the head appears to be imperfectly trilobate.
The neck is much hollowed out on its ventral surface, and the head is clearly peduncu-
late. The articular surface of the trochanter is continuous with that of the head,
at least posteriorly.
On the anterior, and particularly on the outer, face of the trochanter are a
number of extremely distinct impressions of muscle-insertions. The shaft is as nearly
as possible straight, there being only a slight forward convexity in its lower portion.
In the middle it is cylindrical in section, but just above the condyles it is somewhat
flattened from before backward. The postero-internal border is defined by an
extremely conspicuous linea aspera, which, as Ameghino remarks, is a veritable
projecting lamina of bone. It runs from end to end of the shaft, terminating below
in the upper angle of the inner condyle. The anterior face of the shaft is likewise
SKELETON OF PHORORHACOS INFLATUS. 79
marked by an intermuscular ridge running obliquely downward and inward from the
outer edge of the trochanter: in its lower third it forks, one branch running to the
upper anterior angle of each of the condyles; in its upper portion this line no doubt
separated the surfaces for the crurwus and the vastus eaternus.
There is a deep popliteal fossa, which is separated by a prominent bar of bone from
the intercondylar fossa, which is marked by two deep pits for ligaments.
The outer condyle projects considerably below the inner; its fibular ridge is very
prominent, and the surface for the fibula slightly concave from side to side. On the
lower end of the outer condyle is a very distinct facet for the tendon of the outer head
of the tibialis anticus. Anteriorly the condyles project considerably, and the rotular
channel is comparatively deep.
Comparing the femur of Déomedea with the fossil, we find that it differs, among other
points, in the shallowness and form of its popliteal fossa and in the great antero-lateral
compression of the lower end of the shaft. ‘The head, moreover, is much less distinctly
pedunculate.
The femur of Grus differs in the somewhat more curved shaft, the elevation of the
trochanter above the head, the somewhat shallower popliteal fossa, and the smaller
degree of obliquity of the distal articulation.
In Psophia the trochanter is high and the popliteal fossa shallower than in the fossil.
In Cariama the shaft is rather curved and the trochanter is somewhat raised above
the head, which is very similar to that of the fossil: the distal end differs only in the
somewhat greater shallowness of the popliteal fossa.
In the Rails the shaft of the femur is always much curved, and the trochanter rises
considerably above the head.
In the Birds of Prey (including Serpentarius) the femur is pneumatic, the trochanter
rises above the head, and the neck, as a rule, is very short.
The dimensions of the femur are :—
mm.
er ot tues Myr semerpeme saree rea Seats Cole Voile tap, cms Py US Kel Sey cus. [etaiecicicietee fh
Windih at ronal’ Gl 6 9 G6 5 6 8 oe bb ee Go 8 HY
Pearavadistalven Gemma: wea mere MLN ue McK Mc aagl jl Ow
oy | Geramaloval? ee gg a) oo la a he lela 6 Bian GD
Cmenmtaanca orrmnilla Organ 56.5 5 6 68s 8 o 0 8 5» 5 © ell
The Tibio-tarsus. (Plate XVII. figs. 9 & 10.)
The tibio-tarsus is a comparatively long and slender bone. The shaft is sonewhat
flattened on its anterior and posterior surfaces, and tapers slightly towards its distal end.
It is straight, but the median border near its distal end curves slightly iuward, so
that the inner condyle projects considerably inward ; the outer condyle is ina line with
the outer border of the shaft. The anterior surface is bordered internally by a strongly-
marked Jinea aspera, which is continuous above with the procnemial crest, and towards
you. xv.— Parr m1. No. 4.—October, 1899. N
80 Mk. C. W. ANDREWS ON THE SKULL AND
the lower end forms the inner border of the very deep groove for the extensor tendons ;
it terminates at the inner end of the extensor bridge. This latter lies obliquely ; near
the upper border of its inner end there is a deep pit for the attachment of the oblique
ligamentous sling through which the tendon of the tibialis anticus passes: the other
end of the ligament is inserted lower down on a well-marked tubercle just above the
outer condyle; the lower border of the bridge passes externally into a prominent
tuberosity, such as occurs in the Storks, &c., and serves for the insertion of the ligament
bridging the extensor tendons.
The condyles are about equal in size, and are separated by a deep intercondylar gorge
which opens above into a median concavity lying in the middle line immediately below
the last-mentioned tuberosity, and, when the leg is flexed, receiving the large inter-
condylar tuberosity of the metatarsus. Posteriorly the intercondylar groove is short and
shallow, but is still sharply defined, being bounded by the prominent posterior projections
of the condyles. Looked at from the side the outer condyle is nearly circular in
outline, while the inner is much elongated from before backward ; both are concave
on their outer surfaces,
The upper end of the bone is relatively small, but the pro- and ectocnemial crests
are fairly well developed and rise considerably above the proximal articular surface,
which slopes upward and forward. The inner glenoid surface is flat or only
slightly convex; posteriorly it forms a considerable projection and is separated by
a notch from the outer or fibular surface, which is small and very convex in all
directions.
There are no pneumatic foramina. ‘The centre of the shaft is occupied by a large
smooth-walled cavity, and the thickness of the bone in the middle of the shaft does
not exceed 4 mm.
The tibio-tarsus of Diomedea differs from that of the fossil in the much greater
breadth of the intercondylar fossa, the nearly transverse direction of the extensor
bridge and the absence of a tuberosity at its outer end, and in possessing a much
Jarger cnemial crest.
In Grus the tibia differs widely in the relatively greater width of the distal
articulation, in which the condyles are small and the intercondylar groove broad; on
the other hand, there is a median tuberosity.
In Psophia the extensor bridge is much more transverse, and the outer condyle.
considerably the larger, as in Rails.
In Cariama the condyles are relatively smaller and the intercondylar groove
shallower and broader.
In Ciconia the tibia presents considerable resemblance to the fossil, but the
articular surfaces for the femur are less distinctly separated, the extensor bridge is
less oblique, and the groove for the tendons occupies the whole width of the shaft.
In the Accipitrine birds, including Serpentarius, the cnemial crests are very small,
SKELETON OF PHORORHACOS INFELATUS. 81
the groove for the extensor tendons nearly median, there is no distinct tuberosity
at the outer side of the extensor bridge, and the condyles are comparatively small
and separated by a broad intercondylar gorge; while, in correlation with the small
size of the intercondylar tuberosity of the metacarpus, there is no depression at the
upper end of the intercondylar groove, into which it fits when the bones are flexed
one upon another.
The dimensions of the tibio-tarsus are :—
em,
Length, including cnemial crest 895
s without cnemial crest . 37-5
Width at distal end . Sige Wee Gn oly ces Meh yeu Came a Meant 3
Width of shaft at narrowest point, 7. e. about 3 cm. above extensor bridge. 2:7
Circumference at same point . 8:2
Width from front to back at same point 2-0
The Metatarsus. (Plate XVII. figs. 11-17.)
The glenoidal facets for articulation with the tibia are deeply concave; the outer (0.)
is slightly below the inner, and has a bluntly pointed process on its outer border.
There is a prominent rounded intercondylar process (¢.c.), which on the outer side,
at its base, bears a shallow pit for the insertion of the outer semicircular ligament.
On the outer side of the bone, immediately below the anterior angle of the outer
glenoid facet, is a rough surface for the insertion of the lateral tibio-tarsal ligament.
Posteriorly the hypotarsus (hy.) forms a broad projecting mass, consisting of an
outer broad ridge and a narrow inner one, separated by a shallow groove; the outer
and inner surfaces of the hypotarsus are also slightly grooved, but there are no closed
or nearly closed channels for tendons. The whole hypotarsus is short, and the two
ridges constituting it converge below into a simple median ridge, which soon dies
away on the hinder surface of the shaft. In the slight depressions on either side
of the hypotarsus are the posterior openings of the interosseous foramina, the inner
being slightly higher up than the outer.
The anterior face of the shaft is deeply concave in its upper part, so that the
interosseous foramina open anteriorly at the bottom of a deep fossa, the outer a little
above the inner. Immediately below these foramina is a pair of oval rugosities, the
outer slightly above the inner, for the insertion of the tibialis anticus. Internal to
these is a deep groove, over which the inner border of the bone forms an overhanging
lip; this groove is for the tendon of the ealensor communis digitorum.
The concavity of the upper portion of the anterior face is continued as a broad
channel extending down about the upper half of the shaft; but there is no specimen
in which this region is complete, so that the exact length of the bone is unknown.
The posterior surface is occupied at its upper end by a ridge forming a prolongation
Na
82 MR. C. W. ANDREWS ON THE SKULL AND
of the hypotarsus. Lower down the shaft it is narrowed and flattened, its outer
border being formed by a prominent ridge, which is continued upward and outward,
and terminates above in the process which forms the outer border of the external
glenoid fossa ; its inner border is formed by a less prominent ridge, which is not, as in
Cariama, continuous above with the inner ridge of the hypotarsus.
The outer face of the shaft is bordered by the ridges which form the outer edges
of the anterior and posterior faces. This surface widens out towards the middle of
the bone, and is flat or even slightly concave. The outer surface is slightly convex,
and is not very sharply marked off from the posterior surface.
As above mentioned, the shaft is incomplete in both the metatarsi of the type-
skeleton, but the distal portion (three or four inches) of the right metatarsus is very well
preserved. The upper portion of the shaft preserved in this specimen is very slender
and roughly triangular in section, one angle of the triangle being a rugose ridge on
the posterior surface. A short distance above the inner trochlea, and at the lower
end of the ridge just mentioned, is the surface for the attachment of the hallux (ha/.).
Below this the posterior surface is slightly concave from side to side. The anterior
surface has near its outer side a shallow groove deepening distally and having at its
lower end the foramen for the tendon of the adductor digitt extern. ‘This foramen
leads into two channels, one opening between the outer and middle trochlea, the other
on the posterior surface of the bone immediately above the trochlee.
The distal trochlee are arranged in a slightly curved line (fig. 17). The median
trochlea (tr. 3) is very large and its articular surface is sharply delimited, both
anteriorly and posteriorly; the median groove is much deeper behind than in front.
The inner trochlea (¢. 2) is relatively small, and is considerably shorter than the
middle one. Its articular surface is evenly convex from side to side, except on its
posterior side, where it is slightly grooved. It is situated a little posterior to the
middle trochlea, behind which it projects considerably. The outer trochlea (¢r. 4) is
rather larger than the inner and is slightly longer; it also projects considerably
behind the median, and its articular surface is grooved posteriorly only. The lateral
surfaces of all the trochleew are deeply excavated by pits for the insertion of the
ligaments of the toes.
In comparing the metatarsus of Phororhacos with that of other birds, it must be
remembered that it has probably been profoundly modified in correlation with the
loss of the power of flight and the consequently exclusively pedestrian progression
of the bird. Among the peculiarities which may probably be thus accounted for
are the simple hypotarsus, the relatively great length of the bone, and the disposition
of the distal trochlee.
The metatarsus of Diomedea differs in several respects, the chief of which are
complete absence of all trace of a hind toe, relatively large size of the articular ends
compared with the shaft, breadth and complication of the hypotarsus, and absence of
a pit for ligament at the outer side of the base of the intercondylar tuberosity.
SKELETON OF PHORORHACOS INFLATUS. 83
In Grus the wetatarsus differs in the complication of the hypotarsus and the position
of the inner (2nd) trochlea, which is high up and much deflected backward.
In Psophia there is a close canal in the hypotarsus, otherwise the bone is much
like the fossil.
In Cariama the metatarsus in most respects closely resembles that of Phororhacos,
but it is proportionately more slender, and at its distal end the lateral trochlez
(particularly the inner 2nd) are less reduced. The metatarsus of Ciconia is similar in
many points, but at the proximal end the hypotarsus is longer, its crests much more
prominent, and it is situated further below the glenoid surfaces than is the case in
Phororhacos. At the distal end the chief difference is that the lateral trochlez are larger
in proportion to the median one.
In most Accipitrine birds the metatarsus is widely different, both in the form of the
hypotarsus and the deep excavation of the hinder surface of the shaft for the various
muscles of the toes and the arrangement of the trochlee. Serpentarius, however, has
undergone such modification in correlation with its terrestrial habits that it approaches
Phororhacos very much more closely than do the rest of the group: this is chiefly the
result of the elongation of the shaft and the alterations in the form of the talon and in
the arrangement of the trochlee. Nevertheless, there are several points, such as the
almost complete absence of an intercondylar tuberosity, in which it differs widely
from the fossil.
The dimensions of the metatarsus are :— carl
\ninin om jain gind) eg 5 th 6 oo G8 eG oo gt 8 Se
Pee atistaliendayey ace ciate nowy 0 lap isy RU ty fe oe emcee vane ak
» Obmickile Gromit 5 oo 56 0 0 6 6 6 0 6 69 6 6 9 o ily
Only one of the phalanges of the hind foot belonging to the skeleton above described
is preserved, but there is a nearly complete left pes of another individual of the same
species. In thisit is seen that the hallux was well developed and terminated ina strong
hooked claw: the metatarsal articulates with the tarso-metatarsus by a simple knob,
and terminates distally in a trochlear surface with a deep pit for ligament on its outer
side. The total length of this toe is about 53 mm. In the second digit there are, as
usual, three phalanges, the terminal (3rd) one being sharp-pointed and slightly hooked; the
remaining two, though shorter than the phalanges of the middle digit, are nearly as stout.
The lengths of the phalanges are: Ist, 31 mm. ; 2nd, 32 mm.; 3rd (ungual), 36 mm.; the
total length of the toe is about 93 mm. The middle toe consists of four phalanges, the
first of which is nearly as long as the two proximal phalanges of the second digit together ;
both it and the two succeeding digits are somewhat compressed from above downward.
The ungual phalangeal is a powerful hooked claw. The lengths of the phalanges are:
Ist, 55 mm.; 2nd, 38 mm.; 5rd, 27 mm.; 4th (ungual), 44 mm.
Of the inner toe only the ungual and one of the other phalanges is preserved; the
length of the former is about 27 mm.
§4 MR. C. W. ANDREWS ON THE SKULL AND
ConcLupING REMARKS.
In the preceding pages only a few of the types to which the fossils have been compared
are mentioned, they being the only types to which any resemblances pointing to possible
affinities could be made out. And even among these there are some to which the
similarity is so slight that they also might perhaps have been omitted. For instance,
in the case of Diomedea, it is only in the structure of the palate and one or two other
points in the skull that any similarity with Phororhacos can be detected, the rest of
the skeleton being strongly against any such relationship. In fact, it seems to the writer
that the only groups that really come into question are the Falconiformes and aberrant
Gruiformes, Cariama, Chunga, and to a less degree Psophia.
It is toa relationship with the former group that the general appearance of the skull
and feet of Phororhacos seems to point ; but, as above shown, the general structure of the
skeleton, particularly of the pelvis and hind limb, is opposed to this, while, on the other
hand, it is strongly in favour of affinity with the Gruiformes. In this, however, the
skull at first sight seems to oppose a number of serious difficulties, differing widely
from the skull of the typical Crane both in general appearance and in many points of
structure, ¢.g. in its holorhinal nares and desmognathous palate ; but it is precisely in
these points that the Cariamide also differ from the typical members of the group and
therefore resemble the fossil. Still the differences between the skull of Phororhacos and
that of Cariama are so great that some explanation of them is necessary if the close
affinity of the two forms is to be maintained. This explanation seems to be found in
the extraordinary size of the beak in the fossil, a specialization that has led to great
modifications in several regions of the skull. ‘Thus the size and weight of the upper
jaw have led to changes in the palate in the direction of greater rigidity, resulting in
the extensive union in the middle line of the maxillo-palatine plates and their
complete fusion with the anterior ends of the palatines (this occurs also in ‘Toucans
and Hornbills). Moreover, the development of the peculiar prominent ridges formed
by these combined elements, and, as already described, projecting below the tomium on
either side the middle line, also tends greatly to increase the rigidity of the beak, and
possibly the modes of union of the pterygoids with the basis cranii may still further
add to it.
Again, with the great massiveness of the mandible is correlated the large size of
the temporal fosse, and the weight of the skull as a whole has led to an increase
in the area of the occipital surface by the extension of the lambdoidal and paroccipital
crests, to which the muscles supporting the head are attached. ‘The characters thus
accounted for give the skull of Phororhacos its peculiar form, so different from that
of Cariama, and would be evidence of a high degree of specialization even if the
rest of the skeleton were unknown.
It was no doubt this high degree of specialization that brought about the extinction
of this giant bird as soon as the conditions to which it was adapted underwent some
SKELETON OF PHORORHACOS INFLATUS. 85
change, while the smaller more generalized Cariama and Chunga, or rather their
ancestors, survived. In fact, it appears that Phororhacos stands in somewhat the same
relation to the Cariamide that such forms as Glyptodon and Panochthus stand to the
modern Armadillos.
All the specimens described are from the Santa Cruz Beds of Monte Observacion,
Patagonia. The age of these deposits is doubtful, but they probably correspond to
some part of the Miocene of the Northern Hemisphere.
EXPLANATION OF THE PLATES.
PLATE XIV.
Skull and mandible of Phororhacos inflatus Amegh., p. 56. From side. # nat. size.
PLATE XV.
Skull of Phororhacos inflatus Amegh., p. 56.
Fig. 1. From below. # nat. size. Fig. 2. From above. 3 nat. size.
b.pt. Process of pterygoid articulating | o0.g. Orbital process of quadrate, p. 63.
with base of the skull, p. 57. pal, Palatine, p. 62.
6.t. Basitemporai platform, p. 57. pe,f. Precondylar fossa, p. 57.
eth. Ethmoid (?), p. 60. p.mez. Premaxilla, p. 61.
if. Facet for inner head of quadrate, p.orb.p. Postorbital process, p. 58.
p. 58. | pt. Pterygoid, p. 62.
i.0.s. Interorbital septum, p. 59. g. Quadrate, p. 62.
lac. Lacrymal, p. 58. gj» Quadrato-jugal, p. 60.
lr, Lambdoidal ridge, p. 58. r. Rostrum, p. 57.
m.t. Mammillary tuberosities, p. 57. sq. Squamosal, p. 58.
max.pal. Maxillo-palatine plates, p. 62. | sup.o. Supra-orbital plate, p. 59.
n. Nasals, p. 59. | ty. Tympanic cavity, p. 57.
oc.c. Occipital condyle, p. 57. zy. Zygomatic process, p. 58.
of. Facet for outer head of quadrate,p.58. |
PLATE XVI.
Pelvis of Phororhacos inflatus Amegh., p. 72. } nat. size.
Fig. 1. From above, Fig. 2. From side. Fig. 3. From below.
acet. Acetabulum, p. 74. o,f. Obturator foramen, p. 74.
a.t. Ante-trochanter, p. 73. p. Pubis, p. 76.
il. Ilium, p. 75. s. Sacral vertebree, p. 72.
as. Ischium, p. 75. u.s. Urosacral vertebre, p. 73.
is,f. Ischiadic foramen, p. 74.
86
ac,
b.a.
hyp.
mMC.2, MC.3.
oO.
ON THE SKULL AND SKELETON OF PHORORHACOS INFLATUS.
PLATE XVII.
Limb-bones of Phororhacos inflatus Amegh. 3 nat. size.
ics!
—
ge
AAO on FP oh
. Left femur, p. 78.
co co
. Right coracoid, anterior surface, p. 68.
. Right scapula, inner surface, p. 69.
. Distal end of left humerus, p. 70.
. Right ulna, olecranon restored from left side, p. 70.
. Right radius, proximal end, p. 70.
Right metacarpus, palmar surface, p. 70.
. Caudal vertebra, p. 78.
. Right tibio-tarsus, p. 79.
Tene, WD6 ep . distal end, p. 80.
Fie. 11. Left tarso-metatarsus, postero-internal view of proximal end, p. 81.
Fig. 12. + PA antero-external view of proximal end, p. 81.
Fig. 13. 5 5 anterior view of proximal end, p. 81.
Fig. 14. Right tarso-metatarsus,
Fig. 15. oe a
Fig. 16. # 3
Fig. 17. 99 99
Acrocoracoid, p. 69.
Impression of brachialis anticus,
p. 76.
. Inner articular facet at proximal
end of metatarsus, p. 81.
. Intercondylar tuberosity, p. 81.
Inner condyle of tibia, p. 80.
. Surface for hallux, p. 82.
Hypotarsus, p. 81.
Second and third metacarpals, p. 71.
Outer articular surface at proximal
end of metatarsus, p. 81.
posterior view of distal end, p. 82.
anterior view of distal end, p. 82.
upper articular surface, p. 81.
lower articular surface, p. 82.
o.c. Outer condyle of tibia, p. 80.
ol. Olecranon process of ulna, p. 70.
p.l. Processus lateralis of coracoid,
p- 68.
r, Radial condyle of humerus,
p. 70.
t. Tuberosity at base of the third
metacarpal, p. 71.
tr.2, tr.8, tr.4. Trochles of 2nd, 3rd, and 4th
metatarsals, p. 82.
u. Ulnar condyle of humerus, p. 70.
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CONTENTS.
Ill. On the Extinct Birds of Patagonia.—I. The Skull and Skeleton of Phororhacos
inflatus Ameghino. By C. W. Anprews, B.Sc., F.Z.S., Assistant in the
British Museum, Natural History. (Plates XIV.—XVIIL). . . . page 95
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IV. Second Contribution to the Ichthyology of Lake Tanganyika.—On the Fishes
obtained by the Congo Free State Expedition under Lieut. Lemaire in 1898. By
G. A. Bouuencer, F.R.S., F.Z.8.
Received May 13, read May 16, 1899,
[Puates XVIII.-XX.]
THe extraordinary richness of Lake Tanganyika in Perciform fishes of the family
Cichlide, first revealed by Mr. Moore’s collection, reported upon in these ‘Transactions,
vol. xv. 1898, pp. 1-30, pls. i.—viii., is further shown by the collection made in July—
August 1898 by the Lemaire Expedition at Moliro, at the southern extremity of the
Lake, already explored by Mr. Moore. Although consisting of no more than 34
specimens, the collection entrusted to me for description by the Government of the
Congo Free State contains the types of ten new species, three of which warrant the
establishment of new genera. These new generaare of further interest in emphasizing
a feature of the Tanganyika Cichlidw to which I have drawn special attention in my
previous contribution, p. 2, viz. the extent of the lateral lines in many of the forms
discovered by Mr. Moore. ‘The knowledge of forms with three lateral lines, a
condition previously unrepresented in the Cichlidw, adds force to the remarks I have
made with regard to the morphological significance of the so-called ‘interrupted
lateral line.”
The Lemaire collection is further valuable for the care with which coloured sketches
of most of the fishes have been taken on the spot by M. Dardenne, the excellent artist
attached to the expedition. These sketches have enabled me to represent some of the
more strikingly coloured forms in chromolithography.
In addition to the definitions of the new species, I am able to draw up descriptions
of the two species of Hctoedus of which very incomplete definitions were given in the
previous contribution, owing to the bad condition of the type-specimens. I have also
added notes on the known species of which specimens are contained in the Lemaire
collection, or of which coloured drawings, accompanied by an indication of the native
names, have been supplied by M. Dardenne.
VOL. XV.—PArT Iv. No. 1.— December, 1899. )
88 MR. G. A. BOULENGER ON A COLLECTION
SERRANID &.
1. Lates microueris Blgr.
Originally described from young specimens, 155 millim. long. The adult, measuring
800 millim. and weighing 14 Ibs., has, as could be expected, very different proportions
and is of a uniform dark silvery colour. The eye is contained 9 times in the length
of the head, which is 4 of the total; longest dorsal spine 4 length of head. Caudal
peduncle and caudal fin as in the young.
Native name: “ Sangala.”
CICHLID4.
2. LAMPROLOGUS LEMAIRI, sp. n. (Plate XVIII. fig. 1.)
A few moderately large curved canine teeth in front of each jaw, followed by a
narrow band of minute teeth; lateral teeth very small. Depth of body 33 times in
total length, length of head 22. Snout slightly longer than the diameter of the eye,
which is 34 times in length of head and nearly double interorbital width; maxillary
extending to slightly beyond vertical of anterior border of eye; cheeks and occiput
naked; a few small deciduous scales on the opercles. Gill-rakers short, 9 on lower part
of anterior arch. Dorsal XIX 7; spines equal from the fifth, which measures a little
more than 4 length of head and ? longest soft rays. Pectoral $ length of head.
Ventral reaching origin of anal. Anal VIII 5; spines increasing in length to the last,
which slightly exceeds longest dorsals. Caudal rounded. Caudal peduncle a little
“pt
edged; a blackish oblique bar from below the anterior third of the eye to the maxillary ;
a large blackish opercular spot ; three oblique blackish bands, descending forwards, on
each side of the back, extending on the base of the dorsal; dorsal and anal tipped
with blackish.
Total length 107 millim.
A single specimen.
This species takes its place in the series between Z. moorti and L. congoensis.
longer than deep. Scales 48 lat. 1. > Pale brown, most of the scales dark-
3. TELMATOCHROMIS TEMPORALIS Bler.
Native name: ‘“ Muchétila.”
4, PARATILAPIA DEWINDTI, sp.n. (Plate XVIII. fig. 2.)
9
Teeth very small, in 5 or 4 series in both jaws, the outer scarcely larger and not
tipped with brown. Depth of body equal to length of head, nearly 5 times in total
OF FISHES FROM LAKE TANGANYIKA. 89
length. Snout with curved upper profile, shorter than the eye, the diameter of which
is 23 to 22 in length of head and exceeds interorbital width; maxillary extending
to below anterior fourth of eye ; 2 or 5 series of scales on the cheek; large scales on
the opercle. Gill-rakers rather long, lanceolate, 17 or 18 on lower part of anterior
arch. Dorsal XII-XIII 12-13; spines increasing in length to the eighth or ninth,
which measures 2 or 4 length of head and ? longest soft rays. Pectoral a little
shorter than head, extending as far as origin of anal. Ventral produced into a long
filament, extending beyond origin of anal. Anal III 9; third spine longest, 7 length
of head; middle soft rays produced, as long as or a little shorter than head.
Caudal deeply emarginate, crescentic. Caudal peduncle 14 as long as deep. Scales
0
finely denticulate on the edge, 37-58 ng lat. 1. ae upper lateral line not reaching
base of caudal. Grey above, white beneath; four yellowish stripes along each side ;
pectorals yellowish ; other fins dark grey or blackish.
Total length 100 millim.
Three specimens.—Native name: “ Likuko.”
This species is named in memory of the distinguished young geologist, Dr. De Windt,
attached to Lieut. Lemaire’s expedition, who was accidentally drowned in Lake
Tanganyika. It is very closely allied to P. ventralis Blgr., from which it differs in the
dentition and in the shorter pectoral fin.
5. BAatHYBATES FEROX Bley.
Native name: ‘‘ Musupa.”
‘TREMATOCARA, g, Nn.
Closely allied to Paratilapia, but with the teeth very minute, in a narrow band in
both jaws, the maxillary entirely concealed under the preorbital when the mouth is
closed, the muciferous cavities of the top and sides of the skull and of the mandible
extremely large, as in Acerina, and with a single, incomplete, lateral line.
6. TREMATOCARA MARGINATUM, sp. n. (Plate XIX. fig. 1.)
9
Depth of body 34 times in total length, length of head 23 to 8. Snout with curved
upper profile, shorter than the eye, which is 24 in length of head and exceeds inter-
ocular width ; mouth extending to below anterior border of eye; cheek naked ; a few
deciduous scales on the opercle; nasal, frontal, pre- and suborbital, preopercular, and
mandibular bones with very large and deep cavities separated by narrow septa and
02
90 MR. G. A. BOULENGER ON A COLLECTION
covered with a thin skin. Gill-rakers short, 10 on lower part of anterior arch. Dorsal
X 11; spines? length of head, a little shorter than longest soft rays. Pectoral acutely
pointed, as long as head. Ventral reaching origin of anal. Anal III 10; third spine
nearly as long as dorsals. Caudal with deep crescentic notch. Caudal peduncle 14 as
long as deep. Scales cycloid, 302; lateral line reduced to a few (6 or 7) short tubes
in the upper series. Pale brownish above, white beneath ; a bluish lateral stripe; fins
yellowish, dorsal and anal edged with blackish.
Total length 65 millim.
Two specimens.—Native name: “ Lilowé.”
GRAMMATOTRIA, g. 0.
Body elongate; scales ctenoid; three incomplete lateral lines. Jaws narrow, with
a band of minute conical teeth and an outer series of rather longer ones ; maxillary
concealed under the preorbital when the mouth is closed. A large papillose pad on
each side of the pharynx, between the gills. Dorsal with 15 spines, anal with 3.
7. GRAMMATOTRIA LEMAIRI, sp. n. (Plate XVIII. fig. 3.
Depth of body 4 times in total length, length of head 3. Snout with slightly
convex upper profile, 14 diameter of eye, which is 52 in length of head and equals
interocular width ; maxillary extending to between nostril and eye; 3 series of
scales on the cheek ; opercle covered with scales. Gill-rakers short, 12 on lower part
of anterior arch. Dorsal XV 14; spines slender, equal from the fifth, which measures
nearly 3 length of head; soft rays not longer than the spines. Pectoral acutely
pointed, nearly as long as head, extending as far as origin of anal. Ventral reaching
vent. Anal III 10; third spine a little stronger and shorter than longest dorsals.
Jaudal with deep crescentic notch. Caudal peduncle 24 as long as deep. Scales
48-52
6-7 i ;
55 yg; lat. 1. 26. Pale brown above, yellowish beneath; a small dark brown
13-15
opercular spot ; a round brown spot on caudal peduncle at root of caudal fin; dorsal
fin greyish, the soft portion with round white spots; other fins yellowish.
Total length 175 millim.
A single specimen.—Native name: ‘“ Murungi.”
8. Ecropus! pescampsi Bler. (Plate XIX. fig. 2.)
Depth of body 54 times in total length, length of head 3, Snout short, with curved
* To the generic characters as given in the original description must be added the presence of a large dermal
pad in front of the upper part of the gill-arches, same as in Pelmatochromis and Ohromidotilapia.
OF FISHES FROM LAKE TANGANYIKA. 91
upper profile, slightly shorter than the eye, the diameter of which is 24 in length of
head and equals 14 interorbital width; maxillary extending to between nostril and
eye; two series of scales on the cheek; opercle naked. Gill-rakers short, 11 on lower
part of anterior arch, Dorsal XIII 13 ; spines slender, increasing in length to the last,
which is about 3 length of head and little shorter than the soft rays. Anal III 8;
third spine slightly shorter than longest dorsals. Pectoral obtusely pointed, 2 length
of head. Ventral reaching vent. Caudal emarginate. Caudal peduncle 14 a slong
as deep. Scales 35 fe ati I = Pale brown above, yellowish beneath ; fins yellow ;
a round blackish spot on the hinder part of the spinous dorsal.
Total length 60 millim.
A single specimen.
9. Ecropus MELANoGENYS Bler. (Plate XIX. fig. 3.)
Depth of body 5 to 54 times in total length, length of head 34. Snout long, with
nearly straight upper profile, 13 diameter of eye, which is 4 times in length of head and
equals or slightly exceeds interorbital width; maxillary extending to between nostril
and eye; 3 series of scales on the cheek; deciduous scales on the opercle. Gill-rakers
short, 12-13 on lower part of anterior arch. Dorsal XIII-XIV 17; spines slender,
increasing in length to the last, which is about 2 length of head; last soft rays
produced, at least half length of head. Anal III 13; third spine 3 length of
head. Pectoral pointed, a little shorter than head. Ventral reaching origin of anal.
34,
Caudal deeply emarginate. Caudal peduncle twice as long as deep. Scales 43-44 75 3
lat. 1. —o Grey above, white below; dorsal scales with a pale blue central spot; a
blackish opercular spot ; chin and branchiostegal membrane blackish ; dorsal grey, with
whitish streaks and spots, and a large oval blackish spot in the middle of the spinous
portion; anal grey, streaked with whitish ; pectoral and caudal yellowish, the latter
with crescentic dark bands; ventral blackish at the end.
Total length 110 millim.
Two specimens.—Native name; ‘ Losorella.”
10, Ecropus LONGIANALIS, sp. n. (Plate XIX. fig. 4.)
Depth of body 5 times in total length, length of head 3. Snout long, with slightly
convex upper profile, 14 diameter of eye, which is 3} times in length of head and
nearly 14 interorbital width ; maxillary extending to between nostril and eye; 3 series
of scales on the cheek; deciduous scales on the opercle. Gill-rakers short, 12 on
lower part of anteriorarch. Dorsal XY 15; spines slender, increasing in length to the
92 MR. G. A. BOULENGER ON A COLLECTION
last, which is 4 length of head; soft rays slightly longer, the last not produced. Anal
III 17; third spine 4 length of head. Pectoral pointed, $ length of head. Ventral
nearly reaching origin of anal. ae deeply emarginate. Caudal peduncle twice as
long as deep. Scales 442; lat. 1. aaa Brownish above, whitish beneath; a blackish
opercular spot; a lateral series of rather indistinct dark spots; dorsal greyish; other
fins yellow.
Total length 97 millim.
A single specimen, with the mouth and pharynx full of advanced embryos.
The characters of the 5 species of Hcetodus may be contrasted as follows :—
EMdescampsi. WO, Xe anya 14.) VAC ANS eaSqegs4= 35h mle
Depth of body 33-54 in total length. Snout slightly shorter than ae
EL. melanogenys. D. XIII-X1IV 16-17. A.TII 13. Sq. 43- ME Nie, jl, eeno
7 3 14-16
Depth of body 5—54 in total length. Snout much longer than eye.
Fi. longianalis, D.XV 15. A.TIL17. Sq. 443; lat. 1. roe Depth of body 6 in
total length. Snout much longer than eye.
XENOTILAPIA, g. 0
Body moderately elongate; scales ctenoid; three lateral lines. Mouth small, very
protractile; upper jaw with series of minute conical teeth ; two mandibular teeth
turned outwards, in a single series; maxillary concealed under the preorbital when
the mouth is closed. A large papillose pad on each side of the PETS between the
gills. Dorsal with 14 or 15 spines, anai with 3.
11. XENOTILAPIA SIMA, sp.n. (Plate XIX. fig. 5.)
Depth of body 4 times in total length, length of head 33. Snout very short and
deep, with very steep convex upper profile; eye very large, oval, its diameter ? length
of head and much greater than interorbital width ; mouth nearly straight, Non soneall
extending to below anterior border of eye; 3 or 4 series of scales on the cheek;
deciduous scales on the opercle. Gill-rakers very short, broad, truncate, 9 on lower
part of anterior arch. Dorsal XIV—XV 12; spines subequal from the fifth, } length
of head; longest soft rays ? length of head. Pectoral acutely pointed, as long as head.
Ventral with the inner ray produced and reaching a little beyond origin of anal.
Anal III 11; third spine 4 length of head. Caudal with deep Creseeuve notch.
82-36
Caudal peduncle twice as longasdeep. Scales strongly ciliated, 40-41 55 oh 3; lat. 1. 21-22.
OF FISHES FROM LAKE TANGANYIKA. 93
Pale brownish, with a few round darker spots; a blackish opercular spot; a shining
golden spot on the subopercle ; dorsal greyish, other fins yellowish.
Total length 105 millim.
Two specimens.—Native name: * Lufuina.”
12. Trnapia LaBrata Bler.
Native name: ‘ Kobo.”
13. TILAPIA DARDENNH, sp. n. (Plate XX. fig. 1.
] gs
Teeth very small, in 4 or 5 series in both jaws, outer bicuspid, separated from
the series of smaller tricuspid teeth by a rather wide interspace. Depth of body 5 to 53
times in total length, length of head 54 to 53. Snout with strongly curved upper
profile, little longer than the diameter of the eye, which is 35 times in length of head
and equal to or a little Jess than interorbital width ; mouth small, 2 width of head,
extending to between nostril and eye; 5 or 6 series of scales on the cheek; large scales
on the opercle. Gill-rakers short and thick, 13 on lower part of anterior arch.
Dorsal XIX 10; sixth to ninth spines longest, not quite half length of head, a little
shorter than longest soft rays. Pectoral acutely pointed, as long as or slightly shorter
than the head, not extending to origin of anal. Ventral reaching vent or not quite so
far. Anal III 8; third spine as long as and much stronger than longest dorsals.
Caudal feebly emarginate. Caudal peduncle 1% or 1# as long as deep. Scales mostly
24-25
17-19"
dorsal cross-bars, the first between the eyes; yellowish streaks along the series of
ctenoid, 57 eg lattes Yellowish olive above, silvery beneath, with 10 or 11 dark
scales ; fins yellowish, dorsal with some olive marblings, pectoral and anal red at the
base.
Total length 155 millim.
Two specimens.—Native name: “ Sangani.”
14. TILAPIA RUBROPUNCTATA, sp. n. (Plate XX. fig. 2.)
Teeth very small, in 4 or 5 series in both jaws, outer bicuspid, separated from the
series of smaller tricuspid teeth by a rather wide interspace. Depth of body 3 times
in total length, length of head 22. Snout with straight upper profile, twice as long as
diameter of eye, which is 43 times in length of head and equals interorbital width ;
mouth large, # width of head, extending to between nostril and eye; a few deciduous
scales on the cheek ; large scales on the opercle. Gill-rakers short, rather slender, 12
or 13 on lower part of anterior arch. Dorsal XVI 9; spines equal in length from the
94 MR. G. A. BOULENGER ON A COLLECTION
sixth or seventh, measuring + length of head and 2 longest soft rays. Pectoral obtusely
pointed, % length of head, not extending to origin of anal. Ventral reaching vent.
Anal III 7; third spine a little shorter than longest dorsals. Caudal truncate.
Caudal peduncle as long as deep. Scales mostly cycloid, a few on the sides of the
body below the lateral line finely denticulate, 32-33 a lat. 1. ae Olive-brown
above, pale yellow beneath ; dark cross-bars on the back; each scale of the back and
sides with a central vermilion spot; head spotted and marbled with dark purplish
brown; lower jaw and lower part of opercular region bright yellow ; dorsal and caudal
fins bright yellow, spotted with dark brown; pectoral yellow; branchiostegal
membrane, pectoral region, outer edge of ventrals, and anal vermilion-red.
Total length 120 millim.
Two specimens of this handsomely-coloured fish, which bears the native name
“Kasanga Malengi” on M. Dardenne’s coloured sketch.
15. TILAPIA MICROLEPIS, sp. n. (Plate XX. fig. 3.)
Teeth very small, in 4 series close together in both jaws, outer larger, bicuspid, with
a principal and a small lateral cusp. Depth of body 33 to 4 times in total length,
length of head 3. Snout with straight or slightly convex upper profile, once and a half
the diameter of the eye, which is nearly 4 times in length of head and equals inter-
orbital width ; mouth moderate, its width 2 that of the head, extending to between
nostril and eye; 7 or 8 rows of scales on the cheek; larger scales on the opercle.
Gill-rakers short, rather thick, 13 or 14 on lower part of anterior arch. Dorsal XVI-
XVII 14-15; spines subequal in length from the fifth or sixth, measuring ¢ length of
head and alittle shorter than longest soft rays. Pectoral acutely pointed, ? length
of head, not extending to origin of anal. Ventral widely separated from vent. Anal
Ill 9; third spine as long as and a little stronger than longest dorsals. Caudal with
deep crescentic notch. Caudal peduncle twice as long as deep. Scales cycloid, 80-90
om: lat. 1. —- Pale olive-brown above, white below; faint dark bars across the
back and four round dark spots on each side, the last at the root of the caudal; fins
yellowish.
Total length 1165 millim.
Two specimens.—Native name: ‘‘ Mocupi.”
As in Tilapia desfontainesi, the dentition of this species may be regarded as
connecting Tilapia with Paretilapia.
16. TILAPIA GRANDOCULIS, sp. n. (Plate XIX. fig. 6.)
Teeth very small, in 4 or 5 series in both jaws, of outer series larger, bi- or tricuspid,
OF FISHES FROM LAKE TANGANYIKA. 95
and very obtuse. Depth of body 32 times in total length, length of head 3. Snout
short, with rounded upper profile; eye very large, a little longer than the snout, its
diameter 24 in length of head, and slightly greater than the interorbital width ; mouth
small, $ width of head, extending to between nostril and eye; a few deciduous scales
on the cheek; larger scales on the opercle. Gill-rakers very short, rather thick, 17 on
lower part of anterior arch. Dorsal XIII 14; spines slender, equal in length from the
seventh, measuring ¢ length of head, and a little shorter than longest soft rays.
Pectoral falciform, slightly longer than the head, extending as far as origin of anal.
Ventral prolonged in a long filament, extending beyond origin of anal. Anal III 10;
third spine shorter but stronger than longest dorsals. Caudal with deep crescentic
notch. Caudal peduncle a little longer than deep. Scales mostly ctenoid, 63 =>;
25
lat. 1. — Brown above, with ill-defined darker spots, whitish beneath ; pectorals
yellowish ; other fins blackish towards the edge.
Total length 115 millim.
A single specimen.
SILURIDA.
17. AUCHENASPIS BISCUTATA Geoffr.
Native name: “ Porocco.”
18. CurysicHTHYS CRANCHII Leach.
Two young specimens are in the collection. A larger specimen is represented in
the series of drawings made by Mr. Moore.
19. SynopontTIs MuLTIPUNCTATUS Bler.
The young (150 millim.) have the spots on the head and body larger and fewer.
Native name: “ Katétia.”
CHARACINIDSA.
20. ALESTES MACROLEPIDOTUS C, & V.
Native name: “ TIlala.”
21. ALESTES MACROPHTHALMUS Gthr.
Native name: “ Laala.”
VOL. XV.—PART Iv. No. 2.—December, 1899. P
96 ON A COLLECTION OF FISHES FROM LAKE TANGANYIKA.
CYPRINODONTIDA.
22. HAPLOCHILUS TANGANICANUS Bley.
Native name: “ Mohanga.”
> ore © bo
EXPLANATION OF THE PLATES.
PLATE XVIII.
. Lamprologus lemairii, p. 88.
. Paratilapia dewindti, p. 88.
. Grammatotria lemairii, p. 90.
PLATE XIX.
. Trematocara marginatum, p. 89. 1a. Side view of head;
1 6. Lower view of head, enlarged.
. Ectodus descampsi, p. 90.
» meélanogenys, p. 91.
oe longianalis, p. 91.
. Xenotilapia sima, p. 92.
. Tilapia grandoculis, p. 94.
PLATE XX.
. Tilapia dardennit, p. 93.
os rubropunctata, p. 93.
» mécrolepis, p. 94.
Trans.Loct, Soc Wt AV. F¢, XVM.
De
OOOO
ree
J.Green del.et kth.
Mintern Bros.imp.
I1LAMPROLOGUS LEMAIRII. 2.PARATILAPIA DEWINDTI. 3.GRAMMATOTRIA LEMAIRITI.
ej he
Suse eh
i.)
Troms. Loot. Soc. WA AV. G¢_ATL.
oe
a,
ir)
J Green del.et lth. Mintern Bros.imp.
LTREMATOCARA MARGINATUM. 2ECTODUS DESCAMPSI. 3.ECTODUS MELANOGENYS.
4.ECTODUS LONGIANALIS. 5.XENOTILAPIA SIMA. 6.TILAPIA GRANDOCULIS.
Trand. Look Soo Vt AVG, XX.
dsGreen del.et lth. : Maintern Bros.Chromoe.
1.TILAPIA DARDENNII. 2.T. RUBROPUNCTATA. 3.T.MICROLEPIS.
9
THE FOREST-BED OF HAST ANGLIA. 103
III. Arias. (Nearly complete, but wanting the end of the left transverse process. The
second series gives the measurements of the articulated specimen of O. giganteus
(Irish) in the Woodwardian Museum at Cambridge.)
C. be'grandi C.giganteus( Wood-
(Pakefield). wardian Museum),
inches, millim, | inches. millim.
ContrumMvereatestalenethvor meee cie ieee ere 2°5 64 BRT 68
Neural arch, length in median dorsal line..............-...-- 25 64 3-0) 77
Greatest wih of vertebra (estimated at twice the width of el 7.3 185 8-7 99]
PAM). cococcooegorosegoeHUnaNnDODE bUnDOODUON SE
Width between outer edges of facets for condyles of skull ...... 4-0 102 4-5 115
Width between outer edges of facets for odontoid process ...... 4-0 102 45 115
Front edge of facet for condyle to middle of lateral part of facet for 40 102 45 115
odontordiprocessineraie mirc cir rier en iran iar:
Tip of neural spine to a point vertically below it on the ot 3-0 76 3. Bl 93
HUI) Ot WING CMI Uhagcqscgageccg0ndc nade goavabodes
Tip of BYyPaDOD ELS veRneehy to the level of the hinder edge of 1 30 76 29 74
themeuralearchynethesmiddlcilinenassscwrt-tsiteriacieacnormetene tata f
Deepest point in the depression behind the ventral nerve-foramen \ 6 ral 18 4G
to the dorsal edge of the facet for the odontoid process........
Width of centrum between the two depressions .............. 19-21 47-53 2:8 72
Wadthkotmeuralycanalsnmeironbariiersclesicterlcicicnciercneeiotorene tts ie7/ 44
as 3 ame OCHUNGL Matera cnc yatgersrnsie varie arses ors 17 44
Vertical height of neural canal, in front .................... 12 3
9 5 behind weerrrcriiccrra tenner Weg 44
IV. Axis. (Practically complete.)
C. belgrandi C. giganteus ( Wood-
(Pakefield). wardian Museum).
inches, millim, | inches. millim.
Centrum, greatest length (without odontoid process) .......... 4:4. 112 o1 129
ie 53 io (with a pea) men ei wore Byrd 137
Neural arch, length at the top of the neural canal.............. 4-0 102 4:7 120
«9 SONA, LEMS OF ooogoobvovcancso0g00000 cad0DDga0N 5:0 127 Bess) 140
Hacotmoratlasswid thiol ieieitincthrsi icin nrenoiaeiclon: 3:9 100 44 112
Odontordyprocess;widthvat) tips errseeraciycileerienierioiie crn 16 40
Top of neural canal to bottom of facet for atlas .............. 2-9 74 3:2 82
Length of facet for anterior zygupophysis of 8rd vertebra ...-.. 16 40 1:9 45
Greatest vertical height (at hind end) ................-+.00- 54 137 6:2 157
a swadthy (ean) aiid tend) Mareintrrsrrcrsstrtrieicraiekceercre si 4:0 102 6:8 173
Posterior zygapophyses, width across............+0seeeeeeees 3:0 76 4-0 102
Posterior end of centrum, vertical height of articulating surface... 1°9 48
Top of neural spine to top of neural arch ....-........+.0000- 2-4. 61 3:6 92
vou. XvV.—PART Iv. No. 3.—December, 1899. Q
104 DR. S. F. HARMER ON CERVUS BELGRANDI FROM
Tue AFFINITIES OF CERVUS BELGRANDI.
It is now generally accepted that the Irish Deer (C. giganteus)! and its allies are
members of the Damine group. The particular position which C. belgrandi should
occupy in this group has not hitherto received adequate consideration, in consequence
of the imperfect condition of all the previously described examples. It appears to me
that Cervus belgrandi combines in a most interesting manner the characters of
C. dama aud its fossil allies (C. brown, &c.) and of the series of forms including the
irish Deer. Many specimens of C. dama agree precisely with C. belgrandi in
the number and position of the tines and in the general character of the palmation.
The principal differences between the two forms are: (1) the more horizontal position
of the beam in C. belgrandi; (2) the position and characteristic downward curve
of the first tine in that species; and (3) the existence of a scalloped edge of the
crown in QC. belgrandi, as distinguished from that of C. dama, which.is prolonged
into more definite tines. Both species agree in the absence of points on the
anterior edge of the crown. The palmated region includes the back-tine in C. dama,
whereas in C. belgrandi there is a distinct constriction between the back-tine and the
palmation. ‘The comparative suddenness with which the expansion begins in this species
is probably the reason why its real character has not hitherto been suspected.
Right antler of Cervus dama, believed to be of preglacial age, from Belzig, 10 miles 8.W. of Berlin ;
seen from the inner side. (After Keilhack.)
1 «Manuel d’Hist. Nat., traduit de l’allemand de J. F. Blumenbach,’ ii. 1803, p. 407.
THE FOREST-BED OF EAST ANGLIA. 105
A most interesting specimen, referred to Cervus dama, has been described by Keilhack }
from Belzig, 10 miles south-west of Berlin. It would not be easy to find a specimen
affording a more complete transition from CO. belgrandi to the modern C. dama than
this one, and I have accordingly reproduced one of Keilhack’s figures of the right antler
(woodcut, p. 104). The specimen far exceeds in size all the Fallow-deer measured by
Keilhack, the base of the beam being no less than 180 mm. in circumference, as com-
pared with 132 mm., the mean of the measurements of the recent individuals given by the
same autbor. It further approaches C. belgrandi in the scalloped edge of the posterior
part of the palmation and in the downward curvature of the large brow-tine, which,
however, originates immediately above the burr. The brow-tine appears from the
figures to spring from the beam somewhat on its posterior side, as in the Forest-Bed
form. ‘The affinity of Keilhack’s specimen to C, dama is, however, shown by the angle
made by the antlers with the skull, the direction being much less horizontal than in
C. belgrandi, and by the inclusion of the back-tine in the palmation, from which it
projects in much the same way as in the recent Fallow-deer. The beam terminates in
a short but distinct snag at the distal and anterior edge of the crown, but the interval
between this and the back-tine has a scalloped edge closely resembling that of the
Pakefield specimen. Nehring’, in remarking on the Belzig specimen, ina paper which
preceded Keilhack’s memoir, expresses the opinion that it must be regarded, if not as
an actual C. dama, at least as a direct ancestor of that species.
The Belzig specimen indicates in the clearest way the probable derivation of the
modern C. dama from forms resembling C€. belgrandi, the principal changes which
have taken place being apparently the diminution in size, the approximation of the
crowns of the two antlers by the assumption of a less horizontal position, the inclusion
of the back-tine in the palmation, and the replacement of the serrations of the crown
by the snags which characterize the recent Fallow-deer. The discovery of the Belzig
specimen and that of the form here described go far towards filling up the gap between
C. giganteus and C. dama, to the existence of which attention is called by Riitimeyer *,
in insisting on the affinity of these two forms. It may be noted that the immature
antlers 4 of C. dama have the posterior edge of the crown denticulated in a way strikingly
suggestive of C. belgrandi.
The resemblance of C. belgrandi to C. giganteus is no less striking. Pohlig® has
' K. Keilhack, ‘‘ Ueber einen Damhirseh aus dem deutschen Diluvium,” Jahrb. k. Preuss. geolog. Landes-
anstalt (1887), 1888, p. 283, pl. x1.
2 « Ueber das fossile Vorkommen von Cervus dama .... m Norddeutschland,” SB. Ges. naturf. Berlin,
1883, p. 69.
. * “ Beitrige zu einer natiirlichen Geschichte der Hirsche,” Abhandl. Schweiz. pal. Gesellsch. x. 1883, p. 111.
* Of. Cuvier, ‘‘ Recherches sur les Ossemens Fossiles,” 3rd ed. t. iv. 1826, pl. ui. figs. 28, 29.
5 « Tie Cerviden des thiiringischen Diluyial-Travertines, mit Beitrigen tiber andere diluviale und iiber recente
Hirschformen,” Paleontographica, xxxix. 1892, p. 215.
106 DR. S. F. HARMER ON CERVUS BELGRANDI FROM
recently defined several forms of the latter species, which are accepted with certain
modifications by Lydekker!. ‘Taking the nomenclature of the latter, we have
1. Cervus giganteus carnutorum Laug., of which C. verticornis is regarded as a synonym.
C. verticornis occurs typically in the Forest-Bed (early Pleistocene), and C. carnutorum in
beds of equivalent age in France. Boyd Dawkins records the occurrence of one specimen
of C. verticornis in the Pliocene (Red Crag).
2. C. giganteus belgrandi Lart., from the Mid-Pleistocene of France and Germany.
3. C. giganteus italie Pohl., from similar and later deposits of Italy, Hungary, &c.
4. C. giganteus ruffi Nehr. (=C. euryceros germanie Pohl.), the typical German race, found
also in other parts of the Continent and in England, contemporaneously with the Cave-Bear,
although first recorded from the Mid-Pleistocene.
5. C. giganteus typicus Lyd. (=C. euryceros hibernie Pohl.), the Irish Deer, the most recent form.
The conclusion that C. verticornis is a synonym of C. carnutorum appears to me in
the highest degree doubtful. The latter, found at Saint-Prest, near Chartres, was
described by Laugel? from some upper molars, some portions of the antlers, a metatarsal,
and a lumbar vertebra, which were all believed to belong to one species, comparable in
point of size with the Irish Deer. Laugel does not figure the antlers, but he calls
attention to the fact that their bases are remarkably approximated. The crown was
probably palmated, and is believed to have borne about five curved snags, the longest
of which measured 26 cm. ‘These characters are quite sufficient to show the difference
between Laugel’s species and C. verticornis, even without the illustrated account of the
Saint-Prest fossils which has been given by Gervais®. rom this memoir it appears that
the vertex of the skull of C. carnutorum is much raised, so as to resemble that of the
* Antilope Bubale” ; this being an important difference between it and the Forest-Bed
Stag. Theantlers are closely approximated at their bases (cf. pl. xvi. fig. 4 of Gervais)
and diverge from one another at an acute angle. An equally striking feature of this
species is that, when looked at from in front, the brow-tine appears to originate from
the outer outline of the beam, while that of C. belgrandi, viewed from the same
aspect, appears to spring from the inner outline,
Professor Boyd Dawkins and Mr. E. T. Newton* have, however, referred several
fragments from the Norwich Crag, and possibly from the Forest-Bed, to C. carnutorwm.
I am not prepared to express any opinion with regard to these specimens.
Tam indebted to Professor Boyd Dawkins for a letter in which he informs me that
he is inclined to regard his Cervus verticornis (1872) as identical with C. belgrandi
(Lartet) Belgrand (1869), and for the further information that the type-specimens of
the latter are no longer in existence. C. belgrandi was described in Belgrand’s work
1 T. cit. pp. 184-141. * Bull. Soc. Géol. France, (2) xix. 1862, p. 711.
* “ Zool. et Paléont. Générales.—Nouy. Recherches sur les Vertébrés Vivants et Fossiles,” 1° sér, 1867-1869,
p- 84, pl. xvi. figs. 3-8,
* For references, see EH. T. Newton, “The Vertebrata of the Pliocene Deposits of Britain,” Mem. Geol.
Survey, 1891, p. 26.
THE FOREST-BED OF EAST ANGLIA. 107
‘La Seine ’’!, parts of two frontals, with the bases of the antlers, being figured. ‘These
fossils appear to me to agree in all essential respects with the Pakefield specimen.
The inclination of the antlers, the distance from one another of the pedicles, and in
particular the position of the brow-tine agree in the two forms; the measurements
correspond with a sufficient degree of accuracy; while it further appears from
Belgrand’s plate xix. that the second tine of Cervus belgrandi does not originate in the
same line with the brow-tine, but at a point 90° further forward than the line
containing the base of the brow-tine, in exactly the way that is characteristic of
C. verticornis. Polilig? regards the rudimentary condition of the brow-tine as a leading
characteristic of C. belgrandi. Had he, however, copied Belgrand’s representation of
the right antler in pl. xviii., instead of that of the left antler, this condition would
have been much less apparent; and in the specimen 3 from Yaubach which he himself
figures (pl. xxiv. fig. 1) the brow-t’ae has essentially the characters of that of the
Forest-Bed form.
Cervus giganteus italie also shows some interesting resemblances to the Forest-Bed
form, The specimens in the Museums of Milan and Arezzo figured by Pohlig + have
a palmated crown which shows a distinct resemblance to the Pakefield specimen, from
which they differ, however, in the larger development of the points of the crown and in
having the brow-tine in the position characteristic of the typical C. giganteus. Part of
the definition of this form given by Lydekker—the upward inclination of the palmation,
the front border of which is curved inwardly so that much of the outer surface is seen
in a front view, and the occurrence of the points of the crown near the summit of the
palmation—will apply equally well to C. delgrandi. The general direction of the
antlers, the beams of which spread out at first nearly horizontally, the palmated part
then curving upwards, is again strikingly suggestive of that form.
C. megaceros ruffii was named and figured by Nehring®, who calls attention to the
absence of points on. the anterior edge of the palmated part as an indication of affinity
to C. dama. Although agreeing in this respect with C. belgrandi, it has already
acquired long tines springing from the distal border of the crown in a manner
suggestive of the Irish Deer. Some of the specimens of this race, for instance that
from the Worms Museum figured by Pohlig (p. 222, figs. 4¢ and 4d), show a much
closer approach to the Forest-Bed species, in the way in which the crown is set on to
the beam and in the slight development of the terminal snags.
The general conclusion seems to be that C. belgrandi is the most central type of the
Giant Deer, showing as it does a distinct affinity to the Fallow-deer as well as to the
1 « Hist. Gén. de Paris. La Seine.—I. Le Bassin Parisien,” par E. Belgrand. ‘ Planches de Paléontologie,’
1869, p. 13, pls. xvili.—xx. 2 2. ct. p. 232:
* I do not feel conyinced that this really belongs to O. belgrandi. * T. cit. p. 228.
* “Uber eine besondere Riesenhirsch-Rasse aus der Gegend yon Kottbus,” Sitzungsber. Ges. naturf. Fr.
Berlin, 1891, p. 151. See also papers by the same author, “ Neue Notizen iiber Cervus megaceros, var. ruffir,
Nhrg.,” ibid. 1892, p. 3; and ‘ Deutsche Jiger-Zeitung,’ xxxii. 1899, p. 631; xxxiii. 1899, p. 413.
VOL. XV.— PART Iv. No. 4.—December, 1899. R
108 ON CERVUS BELGRANDI FROM EAST ANGLIA.
Cervus giganteus series. The large size of the Pakefield specimen and the condition of
the sutures of the skull preclude the idea that it can be a young form, and there is every
reason to suppose that the characters shown by it are those of the adult animal. The
crown has become conspicuously palmated, but the long tines found in C. giganteus are
at present indicated merely by a slight serration of its distal border. Another point
which seems to me to deserve special notice is the direction of the vascular grooves on
the beam of the antler. In the Forest-Bed specimens these run in an accurately longi-
tudinal course up the posterior and ventral surface of the beam, while in the Irish Deer,
as well as in the specimens of C. giganteus from the Barrington gravel, the grooves have
an elongated spiral course. This may indicate a torsion of the whole antler. Itisa
noteworthy fact that the palmated crown of the Irish Deer is much more horizontally
placed than that of C. belgrandi, in which the direction is very oblique, the dorsal
surface looking somewhat backwards. If the anterior edge of the crown in C. belgrandi
were depressed by the torsion of the beam, so as to bring the palmation into a
horizontal position, this new position would not only agree with that found in the Irish
Deer, but it would result in a torsion of the vascular grooves, which would thereupon
assume the spiral course characteristic of the latter. C. dama agrees with the Irish
Deer in this respect. Although the brow-tine of the Irish Deer may have the same
curvature at its base as that in C. be/grandi, it comes off immediately above the. burr
and from the extreme front edge of the beam. ‘he second tine of the Irish Deer is
more nearly opposite the back-tine than in C. belgrandi, and the palmation often
begins soon enough to include it, instead of commencing above the back-tine, as in
that species.
The geological occurrence of C. belgrandi agrees well with the conclusion above
indicated. Its occurrence in the Forest-Bed is itself an evidence of its relative
antiquity, while the existence of the species in the Red Crag (Boyd Dawkins) carries
this form further back than any of the other races of Giant Deer.
EXPLANATION OF PLATE XXT.
1
Fig. 1. Cervus belgrandi.—Skull and antlers, from behind. X 7. ‘The specimen was
obtained at Pakefield, near Lowesteft. a, burr; J, first tine; d,d’, back-tine.
The tines c, which are not visible in this position, come off from the front
edge of the beam, midway between 6 and d.
. View of the same specimen from the right side. ‘The figure having been drawn
from a photograph, the parts of the right antler appear much larger than
they really are. 0, first tine ; ¢, second tine; d, d’, back-tine.
. Atlas vertebra of the same specimen, from the ventral side. ‘The end of ‘the
right transverse process is partially obscured by the matrix.
Fig. 4, Axis vertebra of the same specimen, from the right side. ‘The end of the neural
spine is partially obscured by the matrix.
I
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‘TIQNVYOTHE SNAXHO
EDOM ITIP B PL VO Ys
V. On a Specimen of Cervus belgrandi Lart. (C. verticornis Dawk.) from the Forest-
Bed of East Anglia. By Stoney F. Harwer, Sc.D., FR.S., Superintendent of
the University Musewm of Zoology, Cambridge.
Received April 24, read June 6, 1899.
[Puate X XT. ]
THE specimen which forms the subject of the present paper is from the Forest-Bed
series of Pakefield, near Lowestoft, where it was found as a result of the great
destruction of the cliff due to the high tides of November 1897. The pieces were
obtained from the men who had dug it out by my assistant, Ernest Lane, to whom is
due the credit of recognizing the importance of the specimen, which is now in the
University Museum of Zoology at Cambridge.
I have been unable to find a record of any specimen belonging to this species which
will compare with the present one in completeness. The posterior part of the skull,
including the whole of the brain-case, is in excellent condition. The anterior parts,
from the orbital region, are wanting. The left antler is nearly complete, and the parts
which are missing are to some extent represented in the right antler, a large part of
which is also present. ‘The atlas and axis were found associated with the skull.
The specimen throws considerable light on the real characters of the antlers of the
form usually known as Cervus verticornis; and in particular it demonstrates the
incorrectness of the diagnosis which is usually given of this species. C. verticornis was
originally described by Professor W. Boyd Dawkins ?, and a fuller account of it was
given by the same author in a later publication *®. The species is commonly represented
by basal parts of the antlers only, in collections from the Forest-Bed, and is readily
recognized by its massive cylindrical beam, with the brow-tine (usually situated about
2 inches above the burr) curving forwards and downwards. ‘The restorations given by
Boyd Dawkins indicate an antler which slightly widens out immediately above the
back-tine, and ends in a short bifurcated tip, which was not, however, really represented
in any of the specimens. Professor Dawkins’s opinion, stated in his fuller account, that
he cannot recognize any affinity between the antlers of C. verticornis and those of the
1 The abstract of this paper which appears on pp. 715, 716 of the ‘ Proceedings’ for 1899 was published
without giving me an opportunity of correcting the proof. he locality of the fossil and my conclusion with
regard to the nomenclature are incorrectly stated in it—S. F. H., Nov. 13, 1899.
2 « On the Cervidee of the Forest-Bed of Norfolk and Suffolk,” Quart. J. Geol. Soc. xxviii. (1872), p. 406.
8 «British Pleistocene Mammalia.—VI. British Pleistocene Cervidie,” Palazontographical Society, 1887.
1
98 DR. 8S. F. HARMER ON CERVUS BELGRANDI FROM
Irish Deer (C. giganteus) was no doubt the result of this inadequate restoration, which
is also responsible for such statements as those of Flower and Lydekker! that the
antlers are very short and thick, or of Lydekker? that their palmation is generally
slight.
The Pakefield specimen shows, on the contrary, that C. verticornis belongs to a type
comparable with the Irish Deer not only in its size but in the amount of the palmation
of the antlers, though differing from it in the absence of prominent snags springing
from the palmated extremity. It fully bears out the conclusion stated by Lydekker ®
that this form indicates the ancestral stock of C. giganteus.
The specimen measures 6 feet in a straight line between the extreme points of the
antlers, but this measurement hardly gives a correct idea of its real size. ‘The crown of
the right antler rises nearly vertically, while that of the left antler is much more
horizontally expanded, besides being more perfect. The distance of the distal extremity
of the left antler from the middle line of the skull is thus no less than 4 feet 5 inches.
If both antlers had been equally well preserved and if the direction of the right antler
had been as horizontal as that of the left antler, the total span, in a straight line,
would have been 7 feet 3 inches. The more complete left antler is no less than 52°6
inches, measured along the curve ; its pedicle is 2:9 inches in diameter (dorso-ventral)
and the burr is 3:6 inches. The brow-tine (b) is 2 inches above the burr and is
perfectly typical, springing from the back of the beam in such a way that its posterior
surface is level with the posterior surface of the beam. ‘The anterior surface of its base
is hollowed out, and beneath it there is a slight tubercle corresponding with the
accessory tines or with the “ offers” described by Boyd Dawkins‘ in the same position.
This is hardly indicated in the right antler. The beam is nearly cylindrical and is
strongly furrowed by grooves for blood-vessels, which run in a longitudinal direction
without showing any tendency to become spiral. ‘Two inches above the brow-tine, the
beam measures 2°6 inches in greatest diameter, its anterior part soon afterwards
becoming flattened in a dorso-ventral direction to form the second tine (¢), which is only
represented by its base, the proximal edge of which is about 7 inches from the distal
edge of the base of the brow-tine. Beyond the second tine, the beam nearly regains
its cylindrical shape, although having a low ridge on its anterior aspect. It has hitherto
been nearly straight, though slightly concave below and pointing somewhat backwards;
but beyond the second tine it curves rapidly upwards, expanding and becoming flattened.
At the origin of the back-tine (d) it is convex on its under surface and slightly concave
above. ‘The back-tine is fairly long (6 inches, though incomplete at its tip) and is
directed backwards and inwards; the distance of the proximal part of its base from
> “Yntroduction to the Study of Mammals,” 1891, p. 323.
* «The Deer of all Lands,” 1898, p. 140.
Se Lxcutspy Lak Pale SOG Compa onal envalea tig wleNO nr.
THE FOREST-BED OF EAST ANGLIA. we)
a point on the beam opposite the distal end of the base of the second tine is about
7 inches ; and from the posterior edge of the burr, in a straight line, about 19 inches.
Opposite the back-tine, the beam is 2°1 inches in greatest dorso-ventral diameter; and
it is still 1:9 inches thick, becoming flat above, but remaining convex below, at a point
9 inches above the middle of the base of the tine. The front edge of the antler is
wanting from a point 6°5 inches above the tine ¢, and for 11 inches from its lower end
the fractured edge varies from 1°4 to 1:7 inches in thickness. Posteriorly the edge has
been getting thinner from the back-tine, about 10 inches above which it expands into
a large palmated crown, the plane of which is different from that of the part which
precedes it. ‘The beam, from the second tine to the base of the palmation, was extended
upwards in a plane about 45° from the median vertical plane, its edge being seen in a
view of the skull from the front. The posterior edge of the crown is, on the contrary,
rotated outwards and downwards, so that much of its lower surface becomes visible from
in front. The posterior edge of the base of the crown is very concave, and passes
obliquely upwards and backwards, the palmation extending further backwards than the
end of the back-tine. The crown is quite complete except in front, and is everywhere
thin, scarcely exceeding an inch in thickness in any part; its distal border is gently
scalloped, about 7 points or rounded lobes being distinguishable, none of them being
more than mere denticulations. The thinness of the palmated crown at its antero-
distal extremity indicates that no other tines were present in this region. The part
between the main palmation and the back-tine is, however, thick (1:7 inches) on its
fractured anterior border; and another anterior tine may have been present in this
region, or the beam may have terminated in a distinct point, as in the Belzig specimen
(see woodcut, p. 104), mentioned below, or as in C. dama. From a comparison with
the latter, I do not, however, think it likely that an anterior tine was present.
The right antler agrees closely with the left, except that the region above the secornt s
tine turns dorsally much more sharply than that of the other side. It is indeed almost
at right angles to the beam, whereas that of the left side forms a much more open
curve with the beam. ‘The first and second tines agree almost exactly in position with
those of the left side, but the second tine is much better represented, measuring 7:5
inches in length and being practically complete. At its base it is flattened and is a
good deal broader than thick (breadth 2°8 inches, thickness 1‘Sinches). ‘The back-tine
is lost. ‘The anterior edge of the antler is complete for about 13:5 inches (measured
round the curve) above the base of the second tine, and the antler is prolonged for
another 10 inches above the point where the anterior edge ceases to be complete. Its
thinness makes it practically certain that no other tine could have existed.
The part of the skull which is preserved measures 9 inches in greatest length,
and extends as far forwards as the posterior portion of the orbit. The occipital
region and the base of the skull are in good condition. ‘The interval between the
100 DR. 8S. F. HARMER ON CERVUS BELGRANDI FROM
pedicles is wide (3 inches) and the pedicles are much more horizontally placed than
in C. dama, to which the species appears to be related ; they pass with an even curve
into the top of the frontal, whereas in another specimen of C. belgrandi in the Cam-
bridge collection they make a distinct angle with the frontal and are somewhat more
vertical. The pedicles are nearly cylindrical above, but have a sharp ridge descending
from their outer borders to the posterior margin of the orbit; their bases are deeply
hollowed out behind, as in other species of Cervus. Below the pedicles, the skull has
a median ridge, with a marked depression on either side of it, thus differing from a
part of a skull described by Boyd Dawkins’. The posterior part of the parietals
(apparently just in front of the suture with the occipital) has a round foramen nearly
in the middle line, and a smaller one slightly to the left side of it and rather further
forwards (the other side being here injured). I have found a similar vacuity in the
roof of the skull in another Forest-Bed specimen which I refer to C. savini Dawk.
The transverse occipital crest is strongly marked, in correlation with the great size of
the antlers, and the occipital surface is nearly vertical. Ventrally there is a very deep
emargination between the condyles. There was apparently a large supra-orbital
foramen, but the skull is deficient in this region.
The specimens of the Irish Deer (C. giganteus) in the Cambridge collection differ
from the above in having much shorter and less marked pedicles, in the comparatively
slight median ridge on the frontals and their more depressed character in this region,
and in the smaller size of the supra-orbital foramen. The vertex of the skull, between
the two pedicles, is less raised in C. belgrandi, the roof of the skull of which is practi-
cally flat from the front of the occipital crest to a point between the two pedicles. In
the Irish Deer, on the contrary, the skull between the pedicles is much higher than
the most prominent part of the occipital crest, from which it is separated by a distinct
concavity. A striking difference, which appears to me of some importance, is that
whereas in C. giganteus the vascular grooves on the beam of the antlers have a markedly
spiral course, those of C. delgrandi run quite straight up the beams. The race of
C. giganteus found in the Barrington River-gravel agrees with the Irish Deer in this
respect.
The occurrence of the atlas and axis vertebre associated with the skull is of special
interest. Their size is not far short of that of the corresponding vertebre in an
articulated specimen of the Irish Deer in the Woodwardian Museum at Cambridge,
the spread of whose antlers is 8 feet 4 inches. This is brought out in the tables of
measurements which follow. I have noted the following differences between the two
forms, although some of them may be no more than individual peculiarities.
In the Artas:—The neural arch is flatter above in C. giganteus, while it rises
externally more abruptly on the dorsal surface in C. belgrandi. The tip of the
1 Pal. Soe., ¢. cit. p. 28.
THE FOREST-BED OF EAST ANGLIA. 101
rudimentary neural spine is at the middle of the upper surface in the former, and in
front of the middle in the latter. The dorsal nerve-foramina of C. giganteus are
constricted into a figure-of-8 shape (and completely divided by a bridge in a specimen
from the Barrington gravel in the Woodwardian collection), while they are long slits
in C. belgrandi. ‘The neural arch of C. belgrandi has a wider sinus behind on its dorsal
side ; its anterior dorsal emargination is more square-cut: the ventral depression behind
the nerve-foramen is much deeper; the hypapophysis is most prominent behind, with a
median ridge passing towards the anterior end of the vertebra (that of the Woodwardian
specimen of C. giganteus having no ridge). In some of these latter points the Barrington
specimen holds an intermediate position between the two others.
In the Axis :—The dorsal edge of the neural spine is thinner in C. belgrandi; the
transverse processes, which are incomplete, were probably less projecting ; the posterior
zygapophyses are narrower (1:05 inches, as compared with 1:5-1:8 inches) and more
raised; there is a distinct emargination between the pesterior zygapophysis and the
neural spine; and the ventral median ridge is distinctly higher and bounded by
deeper grooves at its sides than in C. giganteus.
In the tables which follow, the measurements have been taken in such a way as to
permit of easy comparison with those given by Boyd Dawkins’, Belgrand?, and others.
I. SKULL.
inches. | millim.
Greatest length of the part of the skull preserved .................. > | BRO 228
Ghrsaicss Dmealiin OF ne wl, cose coocngecenc0p oes en oodneuocecaaon 81 206
Occipital crest to fronto-parietal suture ........-... see cece cere eee 4] 104
is 5 asl (Ghowasn Gis) Gocobbeaocaedonccodupos a 3°0 76
iGavel lxanyeem joeClales, cooogbhooudoacpeducunDasoc0K bp acho sco Rr 3:0 76
bp o supra-orpitaly foraminay) -e2 jee se eee eos 3 99
From burr to burr (shortest distance)...........--2- sees eee eee 54 137
i, 3 (distance between outer edges).......-.-.-----++- 10-4 265
Top of foramen magnum to top of occipital crest ..............---45. 3 | 76
Foramen magnum, height of .... 2.2... 2... eect e ee cet ee ilhealt 28
3 5 SU AKONHIaYtop2 | yin Blo, io orosBlatausiese core e Meceer eee tece ea necetuG ot canic 14 35
Frontal width, just below pedicles ....-.-..... 66.6, reels on baits 00d O 73 | 185
Deshdle, Mangia OPogosoodneodcnvccyegognobaoce dean eomuog c000 000% 18 46
re circumiferenceyOf sero i) SERN ESE TSUN arae runt apar Leientae te 8:5 | 215
,. antero-posterior diameter of .... 2... 6s se cece eee eee eee ee 24 | 61
,», dorso-ventral diameter of .........- 211s ee eect tee eee eine 2:9 74
Interval between bases of R. & L. brow-times. ....--.- ee eee eee eens 9:5 | 250
Brain-case, diameter (external) behind pedicles...........+..-+0e ses +6 > ality
Distance between outer edges of condyles ........ ++ ese eee eee eres +8 i
1 Pal. Soc., t. cit. pp. 25-28.
2 « Ta Seine.—I. Le Bassin Parisien. Planches de Paléont.,” 1869, p. 13.
102 DR. S. F. HARMER ON CERVUS BELGRANDI FROM
II. ANTLERS.
Left. | Right.
inches. millim. | inches. millim.
Circumference of baselof beam! Banas: eeee caine 9:1. 231 8:9 226
Diameter ae RESIN Cha PRR MA Abe Cloths bra Boley 2°9 74.
Diameter OF ULI. tis siaicvenese alee eheic tele eevee SCO ROR: 36 91
Circumferencesoigburrepmeee eee eee eee 11:0 280
Bure ito tame Wy (G)i issue swereneiereos iartolerstorn ceeraern rate rere 2:0 51
Burr to fork of tine 1 (measured to distal side of base) ........ 4:0 102
Circumference of beampaboyve tine sme cree arclereiiiccietoiae 8:2 208
Tine a ibasalicircumierencevo Lane ener 6:8 173
Tine 1 to tine 2 (c) (distal side of 1 to proximal side of 2)...... 7-0 178
Tine 2ucircumferencelorbaselolamee ene eee ae oe 7:0 178
Tine: 2) lon eth obs ce caraeicuesokce nae bsten evar womans Shoes rae Ie a Ss 75 190
Circumference of beam abovetine 20. .0: 0.15 j.es ese see nsec 9°5 241 10°5 261
iBurratoydistaltsidevotebaseiofatinesse meen eee ec eine 14-0 356 14:0 356
Tine 2 (¢) to tine 3 (d) (point opposite distal side of base of 2 to
IDaSO/O8 13) tke a iietee ciStande erausuieropeu tava eden reer ese eae 7:0 178
Posterior edge of burr to tine 3 (in a straight line) ............ 19-0 482
Burritoydistalisidevorpasclontinelol een eioeiee 22:0 560
ines Saccircumberencerotmbaselo Leaner eee 6:5 165
Miners Men othioie weymeree yee Coe ee ea Tae. 6-0 152
Circumference of beam above tine 8 ..............20cceeeees 12:2 310
Motalilenstha(Gntagstraightiline) ence een cr 50°5 1283 30:0 762
Hotalvlensthy(a@loneicunye) emer 52:6 1335 38:0 965
: we ————.\-—__—-—_—~Y
Total span (straight line) ............ VAG ah ace omaoad 6 feet. 1830 millim.
The variability in the position of the tines in this species is illustrated by the
following measurements of the interval between tine 1 and tine 2, taken from four
other specimens in the Cambridge collection :—
Catalogue No. inches. millim.
WIGAN GN) aerate peo etctapeuerenepatercvencees tae 3:5 89
MSA. ecient taa teleneponertencteeeiee arrester 4:0 102
GANG crevite ie teeatareterertetelarerawiousts tere i eet 4:5 114
ISO (ob BaddovodCboe US 6G00000040 5:0 127
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CONTENTS.
IV. Second Contribution to the Ichthyology of Lake Tanganyika —On the Fishes
obtained by the Congo Free State Lupedition under Lieut. Lemaire in 1898.
By G. A. Bovtunerr, F.R.S., 7.Z.8. (Plates XVIIL-XX.). . . page 87
V. On a Specimen of Cervus belgrandi Lart. (C. verticornis Dawk.) from the Forest-
Bed of Hast Anglia. By Sipyuy F. Harmer, Sc.D., Y.R.S., Superintendent of
the University Musewm of Zoology, Cambridge. (Plate XXI.). . . page 97
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VI. A Monograph of the Genus Casuarius. By the Hon. Wauter Roruscuitp, Ph.D.,
F.ZS. Witha Dissertation on the Morphology and Phylogeny of the PALMOGNATHA
(Ratira and Crypturi) and Neognatua (Carinata). By W. P. Pycrart.
Received and read June 20, 1899.
[Puates XXII.—XLV. |
Part I.—A Mownocraru or THE Genus Casvarius.
By the Hon. WatteR Roruscainp, Ph.D., F.Z.S.
Genus CASUARIUS Briss.
1790. Casuarius Brisson, Orn. v. p. 10.
1801. Rhea (non Latham, 1790) Lacépéde, Mém., de l’Inst. i. p. 519.
1842. Hippalectryo Gloger, Hand- u. Hilfsbuch, pp. xxxxili, 452.
The genus Casuarius is the typical genus of the group Casuarii of the Palwognathe
(Ratite of former authors), and inhabits the Papuan subregion, 7. ¢., New Guinea with
the islands in Geelvink Bay, Salwatty, New Britain, the islands of the Aru group,
Northern Queensland, and the island of Ceram in the Moluccas. The Cassowaries are
evidently separable into a great many local forms, apart from the few very distinct good
species ; but owing to their large size, to the uncertainty about the localities of the
living specimens that are brought to Europe, and to the disappearance after death of
the most characteristic coloration and structure of the bare skin on the head and neck,
our knowledge of the species is doubtless still limited. The material which I have
been able to study is very large compared with that available in most Museums. For
several years I have tried to procure a great number of living specimens and of skins
as well, and I have been able to place numerous fresh bodies and skeletons before
Mr. Pycraft, whose appended paper will, I trust, be of great value for the study of the
anatomy of the Palwognathe.
Outwardly the genus Caswarius may easily be recognized by the following
peculiarities :—
The bill is short, generally shorter than the head, laterally compressed, strong, and
with the culmen curved downwards near the tip. ‘The nostrils are placed in the
middle of the maxilla, in a wide membranous groove, and open anteriorly. Head and
neck bare and mostly brightly coloured, the skin much carunculated and wattled in
various places. ‘The forehead is ornamented with a more or less developed helmet of
various shapes. Wings quite rudimentary, with about five round black shafts, without
any indication of webs in the place of remiges; no rectrices apparent. The metatarsus
VOL. XV.—ParRT v. No. 1.—December, 1900. s
110 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
_ is shorter than in most other Ratite, very robust, and covered in front with hexagonal
scutes, and transverse ones near the toes. ‘Toes three, the two outer ones with obtuse,
curved, and short claws; the inner toe with a long, straight, powerful, pointed claw,
which is a dangerous weapon. The body is covered with stiff, hair-like feathers, in
which the after-shaft is as long as the principal shaft. The old birds are black, the
young ones brown, the nestling, when hatched, is striped longitudinally above.
The eggs are about six to eight in number, with a strong and very coarsely granulated
surface. When fresh they are evidently all of a light green colour, but when exposed
to the light they become first more bluish, then greyish, and at last almost cream-
coloured. The male alone incubates.
Herr Schalow (Journ. f. Orn. 1894) has attempted to make a key to distinguish the
eggs of the various species; but his key is a total failure and misleading, being
principally based on the colour, which is the same in all, and on the transparency of the
shell, which is not specifically different.
All Cassowaries are inhabitants of forests, while the rest of the large living Palwo-
gnathe (or Ratitw) ave denizens of steppes and deserts. Their food seems to consist of
all sorts of vegetable matter and fruits ; but they also pick up insects and any creeping
thing that comes in their way. In captivity, at least, they kill and devour chicks and
small birds when they come across them. ‘They also, like Ostriches, Rheas, and others,
swallow quantities of stones and gravel to assist digestion. ‘They are entirely diurnal,
sleeping from sunset till morning.
The voice of the Cassowaries is a curious sort of snorting, grunting, and bellowing,
usually not very loud, and differing according to the species.
Their temper is generally sullen and treacherous, and they are extremely pugnacious,
even the different sexes often fighting at other seasons than the breeding-season.
Exceptions are rather rare, but there is an adult Cassowary, now in the Society’s Gardens,
which is quite tame, and was always so, since I had him; while another young bird used
to follow the keeper who fed him like a dog when I had him in Tring, but has since
become somewhat shy.
It seems that the Dutch navigators in 1596 were the first Europeans who became
acquainted with the Cassowary, at least there does not appear to be any indication
that the Portuguese, who visited the East-Indian Archipelago long before, came across
the bird. For more than two and a half centuries there was no certainty as to
the existence of more than one species, but in 1854 Mr. Thomas Wall procured a
specimen of Caswarius casuarius australis near Cape York, which was described by his
brother, for the first time, in the ‘ Hlustrated Sydney Herald’; but the description
(made from memory) is as bad asit can be. In 1857 Gould described the New Britain
Cassowary, in 186) Blyth and Sclater described two new species, and since then
the described forms have constantly been increasing, till they now number 20. The
best general accounts are those by Salvadori in his monograph of the genus in the
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. ea
‘Memorie della Accademia di Torino,’ and in his famous ‘ Ornitologia della Papuasia,’
iil. pp. 473-503.
The Cassowaries are easily divided into three groups, as follows:—1. The typical
group, containing two species, Caswarius bicarunculatus and C. casuarius with its seven
subspecies ; 2. ‘The one-wattled Cassowaries, containing two species, C. philipi and
C. unappendiculatus, with four subspecies; 3. The Mooruks, which number four
species—C. papuanus with two subspecies, C. picticollis with two subspecies, C. bennetti
with two subspecies, and C. /orie. These three groups are readily separable externally
and anatomically. Mr. Pycraft’s memoir will detail the anatomical features. The
external characters may be roughly designated thus :—
Casque compressed laterally, two wattles on fore-neck . . . . Typicat Group.
Casque depressed posteriorly, sometimes compressed laterally, one
wattle on fore-neck. . . ..'...’. . .. .'. . » ONE-waTTLED CAssowaRizs.
Casque depressed posteriorly, no wattle on fore-neck . . . . . Tuer Mooruxs.
In the typical group the species Casuarius casuarius and its subspecies vary
enormously in the height and slope of the casque; and although in the “ Key” to the
species I have stated that certain forms are distinguished by a high or low casque, the
fact remains that we often have greater differences among individuals of one subspecies
than between two subspecies. Also we often find that males of one given subspecies
show, as a rule, much higher and more erect casques, while the females have the easque
lower and sloping back over the occiput. ‘There is no doubt that certain differences
exist in these two characters among the subspecies and species of the Casuarius
casuarius section; but too much weight cannot and must not be placed on these
differences: first, because there is considerable individual variation ; secondly, because
there is sexual variation; and, thirdly, because the period of growth seems variable
(at least in captivity). I have seen young birds: still in brown plumage with
enormous casques, while fully adult birds with black plumage showed hardly any
casque at all.
I have to thank Drs, Finsch, Gestro, Giglioli, and Heck, Count Salvadori,
Dr. Sclater, Mr. Blaauw, and others, for their kind help in lending me specimens,
drawings, &c.
Key to the Species and Subspecies of the Genus Casuarius.
( Plumage striped. Chicks.
1 { Plumage brown. Immature.
{ Plumage black. Adult. Seen ae 2.
/ Two wattles on fore-neck ; casque not depressed posteriorly,
| lone Conajnressecl leery 5 6 56 56 6 60 o 6 0 6 Sk
24 One wattle on fore-neck ; casque depressed posteriorly or
| compressed laterally
\ No wattle on fore-neck, casque depressed posteriorly . . 11.
112 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
| Wattles wide apart on sides of neck .
Wattles close together on front ofneck. . . . . .
( Size large; wattles very large, above 5 inches long .
Size large ; wattles medium, 3 inches or less, or small .
4.4 Size small; casque low, sloping, and less erect; sides of
neck anteriorly bluish purple, posteriorly red
!
( Size small; casque very high . sui 6
Sides of neck anteriorly bright blue, posteriorly soanlet
Sides of neck anteriorly bluish purple, posteriorly scarlet .
( Casque low but very large, wattles joined at base ; sides of
| neck anteriorly blue, posteriorly scarlet . . 6
6 { :
' Casque low, not very large, wattles separate at base ; sides
L of neck purplish mauve
Sides of neck entirely blue . : :
Sides of neck anteriorly blue, nostaniorle fcaler
Fore-neck blue, wattle crimson and blue; casque very
high, compressed laterally, depressed posteriorly . .
84 Fore-neck blue in front, orange-scarlet at the sides;
wattle blue
| Fore-neck orange or yellow, wattle yellow or iid
/ Casque depressed posteriorly, not compressed laterally .
9 { Casque compressed laterally, fore- and hind-neck and sides
( of the neck uniform orange . 5
Casque low, the blue not extending below ai, faved ile
| of neck yellow veined with red . 5
| Casque very high, the blue extending on to the Hone nalred
t sides of neck crimson .
u { Fore-neck blue . Hh
"| Fore-neck red, throat pink .
10 Hind-neck scarlet .
“™ | Hind-neck blue : Baa Petavek dire eet
| Large black patch on side of rent below ear surrounded
13 by abroad pink band. . .. .
No black and pink patch on sides of throat
( Head and neck blue, sides of neck black with pink Between
the;wriukles 20) — 2) : 5 0 ¢ 50
Head and neck blue, round seitth on fore! neck nl sides
14 of neck rosy purple. Pi ete ae
Head whitish blue, fore- and hide neck Whe sides of
nN neck purplish red
Fore-neck entirely blue .
Fore-neck with red patch in centre
C. bicarunculatus.
4
5.
6
C. casuarius.
We
C. casuarius sclateri.
C. casuarius australis.
C. casuarius beccarii.
C. casuarius violicollis.
C. casuarius intensus.
C. casuarius salvadorit.
C. philipi.
C. unappendiculatus rufotinctus.
9.
10.
C. unappendiculatus aurantiacus.
C. unappendiculatus.
C. unappendiculatus occipitalis.
12:
C. lorie.
13.
14.
C. papuanus edwardsi.
C. papuanus.
C. bennetti.
C. bennetti maculatus.
15.
C, picticollis hecki.
C. picticollis.
Py
Ie
1603.
1605.
1635.
1666.
1666-
1770.
1676.
1726.
1738.
' eo
NNN WNW
ov or oF
=): x
—) 4
ov Ol
oo
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 115
CasvaRIvs casuaRtus (L.). Common or Ceram Cassowary. (Plate XXII.)
Avis Eme Aldrovand., Orn. iii. p. 541 (Banda, brought to Holland by Schellinger, who
obtained one in Java: very bad figure).
Emeu Clusius, Exotic. lib. v. p. 98 (good figure, taken from same specimen).
Emeu Nieremberg, Hist. Nat. p. 218 (fig.).
Casuar Olearius, Gottorffische Kunstkammer, p. 23, pl. xiii. (alive in Gottorff; figure
copied from Clusius),
1699. Casoar Perr. & Duvern., Mém. Ac. Se. depuis 1666-1699, iii. 2, p. 155, pls. 56, 57
(teste Salvadori) ;
Buffon, Hist. Nat. Ois. i. p. 464.
Emeu Ray & Willughby, Orn. p. 105, pl. xxi.
Kazuwaris (Ceram), Valentyn, Oud en nieuw Oost-Indien, iii. (Amboina), p. 298.
Cassowary or Emeu Albin & Derham, Nat. Hist. B. ii. p. 56, pl. Ix. (alive at George Tavern,
Charing Cross).
. Emeu Barrére, Orn. spec. nov. class. 3, gen. xxxviii. 1.
. Casearius Klein, Hist. Av. Prodr. p. 17.
. Cela Moehring, Avium Genera, no. 56.
. Emeu Worm, Mus. Worm. p. 292.
. Avis Emeu Johnston, Theatr. Univ. de Avibus, p. 180, pl. 56.
. Struthio casuarius Linneus, Syst. Nat. ed. x. i. p. 155 ;
Id., Syst. Nat. ed. xii. 1. p. 265 ;
Gmelin, Syst. Nat. 1. p. 726, no. 2.
Casoar Brisson, Orn. v. p. 10, pl. i. fig. 2.
Casuarius Frisch, Vorstell. Vog. ix. Hauptart. 1, pl. 105, 105 @ (good fig. bird and egg).
Casuarius emeu Latham, Ind. Orn. ii. p. 664 ;
Dum., Dict. Sc. Nat. vii. p. 199;
Lesson, Voy. Coq., Zool. i. pt. 2, p. 711 (partim !) ;
Id., Traité d’Orn. p. 7, pl. 11. fig. 1 (partim !) ;
Thienemann, Fortpfl. ges. Vogel, p. 2, pl. i. fig. 1 (egg) ;
Schlegel, Mus. P.-B., Struthiones, p. 9.
Casuarius galeatus Bonn. Enc. Méth. 1. p. 4, pl. iv. fig. 2;
Vieillot, Nouv. Dict. Hist. Nat. v. p. 345, pl. B. 11. fig. 1;
Merrem, Abhandl. Berl. Akad. p. 179;
Stephens in Shaw’s Gen. Zool. xiv. 2, p. 432, pl. 29 ;
Vieillot, Gal. Ois. 11. p. 77, pl. 225 ;
Merrem in Ersch & Gruber’s Encycl. xv. p. 348;
A. v. Homeyer, Journ. f. Orn. p. 365 ;
Sclater, Proc. Zool. Soc. Lond. pp. 210, 250 ;
Des Murs, Tr. d’Oologie, p. 362 ;
Rosenberg, Journ. f. Orn. p. 45;
Crisp, Proc. Zool. Soc. Lond. p. 137 ;
Sclater, Trans. Zool. Soc. Lond. iv. p. 358, fig. a, p. 8360 (note), pl. 71;
114 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
1862. Schlegel, Jaarb. zool. Genootsch. Nat. Art. Mag. p. 196,
1863. Sclater, Proc. Zool. Soc. Lond. p. 234 ;
1864. Schlegel, Dierent., Vogels, p. 238;
1866. Sclater, Proc. Zoo]. Soc. Lond. p. 168;
1867. Id., op. cit. p. 179 (hatched in Society’s Gardens) ;
1869. Wallace, Malay Archip. n. p. 156 ;
1871. Flower, Proc. Zool. Soc. Lond. pp. 82-35 ;
1873. Garrod, Proc. Zool. Soc. Lond. p. 644;
1873. Rosenberg, Journ. f. Orn. p. 390 ;
1875. Sclater, Proc. Zool. Soe. Lond. pp. 86, 87 ;
1877. Harting, Ostriches & Ostr. Farm. p. 103, plate ;
1879. Rosenberg, Malay. Arch. p. 323 ;
1881. Sclater, Voy. ‘Challenger,’ Birds, p. 64;
1881. Salvadori, Mem. R. Ac. Se. Torino, (2) xxxiv. p. 188, pl. i. fig. 2;
1882. Id., Orn. Pap. e Moluce. iii. p. 479 ;
1891. Thébault, Bull. Soc. Phil. (8) iii. pp. 198-210;
1891. Salvadori, Mem. R. Ac. Sc. Torino, (2) xlii. p. 213 ;
1894. Schalow, Journ. f. Orn. p. 18 (egg) ;
1895. Salvadori, Cat. B. Brit. Mus. xxvii. p. 592 ;
1896. Oustalet, Nouv. Arch. Mus. Paris, (3) vill. p. 263 (one menagerie bird only in Museum).
1811. Casuarius casuarius Miger, Prodr. Mamm. et Av. p. 247.
1842. Hippalectryo indicus Gloger, Hand-u. Hilfsb. p. 452 ;
1844. Casuarius orientalis 8. MuJl., Land- en Volkenk. p. 109 (partim !).
1872. Hippalectryo casuarius Sundev., Tentamen, p. 152.
1875. Casuarius gavanensis (sic!) Gulliv., Proc. Zool. Soc. Lond. pp. 478, 488.
1879. Casuarius beccarit (non Sclater!) Pelzeln, Ibis, 1879, pp. 376, 377 ;
1881. Id., Ibis, p. 401 (egg).
Native name: called Suwuri by the Malays; but it is not certain if this isa local name on
Ceram, or, what is more likely, a word used for all Cassowaries in the trade and of Papuan origin.
Adult. Bill long; casque large, not very high, and somewhat sloping backwards,
compressed laterally, of a dark brownish horn-colour. Head and occiput Nile-blue,
becoming darker on the upper part of the hind-neck. Lower two-thirds of hind-neck
scarlet. Chin, throat, and fore-neck dark blue. On the lower fore-neck are two large
lappet-shaped wattles, joined at their base, rugose, granulated, and of a deep pink
colour. Naked lower sides of neck bluish purple anteriorly and bright scarlet
posteriorly. Lores black ; iris dark brown ; legs brownish grey. Plumage black. Size
medium, :
Total length about 13 m., bill from gape 98 mm., tarsus 250 to 275 mm., claw of
inner toe 70-78 mm.
Young (two-fifths grown). Plumage of various shades of yellowish brown to rufous
brown. Head and fore-neck dull leaden blue, hind-neck dull yellow, naked lower
sides of neck a mixture of dull blue and livid purple.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 115
Young (three-fifths grown). Plumage yellowish or rufous brown mixed with black.
Fore-neck dull indigo-blue ; head and occiput pale dull blue; hind-neck dull orange-
red, naked lower sides of neck blue, posteriorly mixed livid purple and dull red.
Young (full-grown). Plumage black, mixed with a few rufous feathers. Fore-neck
indigo-blue; head and occiput pale blue ; hind-neck scarlet, naked lower sides of neck
blue anteriorly, passing through plum-purple to scarlet posteriorly.
Chick. Head and neck rufous, paler below; three broad stripes on back, one
irregular stripe on each side reaching from the anus to the wing, and two other lateral
stripes extending to the thighs, the last two broken up into irregular blotches.
Hab. Ceram.
As shown in the synonymy, the Ceram Cassowary is the oldest known member of the
genus. In the year 1595 a number of merchants in Amsterdam formed a company,
and sent out four vessels to open communication with the Eastern Archipelago and to
bring home spices and other valuable merchandise. In December 1596 the ships were
anchored at Sydayo, in Java, and it was there that Jan Jacobsz Schellinger, the skipper
of the ship ‘ Amsterdam,’ was presented with a Cassowary, which had been brought to
Java from Banda Island. This was a day or two before Schellinger was murdered by
the chief of Sydayo. Although the ship ‘Amsterdam’ was left and burnt, the
wonderful “ large fowl” was brought on board of one of the other vessels and was
landed alive at Amsterdam in 1597. It was at first exhibited to the public for some
months, then came into the hands of Count George Everard Solms, of s’Gravenhage,
who kept it for a long time at the Hague, and afterwards presented it to the Elector,
Prince Ernestus of Cologne, who, again, subsequently gave it to the Emperor Rudolphus
the Second. Count Solms, before parting with the bird, had an excellent coloured
picture made of it, from which the very good figure in Clusius, representing the bird,
one of its feathers, and an egg, was taken. ‘This figure has been copied into several
other works. A wretched figure of the same bird was also published in the diary of
the long and dangerous voyage during which it was first observed by Europeans, and
this figure is reproduced by Aldrovandus. In 1666 Olearius, in his somewhat
despicable work ‘ Die Gottorffische Kunstkammer, assures us that one had been kept
alive at Gottorff in Schleswig; but he does not say how it was procured. Olearius’s
description is very amusing, but short and not very accurate. Valentyn (1726) seems to
have been the first to give the actual home of this bird, which is Ceram, He says that
some Dutchmen found it sitting on three eggs on that island as long ago as 1660.
Valentyn also says that the Cassowary occurring in Aru differs from that of Ceram.
Other writers have stated Sumatra, Ceylon, and all sorts of places to be inhabited by
Cassowaries, and even Linneus said: ‘‘ Habitat in Asia, Sumatra, Molucca, Banda.”
Up to the present day erroneous statements respecting the home of Caswarius casuarius
have often been made, but it is evident that of all the Moluccan islands only Ceram is
inhabited by a Cassowary.
116 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
Linneus, in 1758, bestowed our present specific name on this bird, calling it
Struthio casuarius, and diagnosing it as “ Struthio pedibus tridactylis, vertice paleari-
busque nudis.’ He then quotes Olearius, Albin, Dodart, Ray, Clusius, Bontius,
Willughby, Aldrovandus, and finally adds: “Callus elevatus quasi diadema verticis.
Palearia duo sub collo; collum a tergo nudum. Ungue intermedia majore ferit ; ova
punctis excavatis.”
The species is admitted by Wallace and Rosenberg to be not at all rare in the
interior of Ceram, but all travellers say that it 1s extremely shy and difficult to
approach, and it seems that no European naturalist has ever seen it in its wild state.
All we know about its life-history comes from native sources, and may be more or
less trustworthy, but it seems certain that it feeds principally on fallen fruit. ‘The
eggs are said to be three to five in number; but we have had, at Tring Park, six to
eight laid by one female, so that we cannot believe that its clutch can regularly contain
three eggs only. A young Casuarius casuarius was hatched in the Society's Gardens
in 1867, but observations respecting the time of incubation and other interesting points
were apparently not made. I have never succeeded in getting a male that would
pair, although I have had a female which laid many eggs. The egg described by
Schalow (Journ. f. Orn, 1894, p. 19) is faded. ‘There is, I am convinced, not a single
form of Cassowary that lays ‘‘ pale greyish-green ” eggs with “ dark grey-green” glazed
granulations. In any case Caswarius casuarius lays light apple-green eggs, with darker
bright green corrugations. ‘They are elliptical, the thickest zone about the centre,
though very often a thick and a thin end can be distinguished. The elevated
glazed granulations are all more or less connected, forming a labyrinthic network.
The measurements are:—150 : 94, 142: 93, 140: 90, 131: 90, 129: 87, 123: 84,
146: 85 mm.
The eggs are excellent food. Owing to their size it is difficult to boil them well in
the shell, and besides it requires a number of persons to finish one; but for scrambled
eggs, omelettes, cakes, and other purposes they are, like those of the Rhea and Emu,
most excellent.
bo
. CASUARIUS CASUARIUS BECCARII Scl. Beccari’s Cassowary. (Plate XXIII.)
1875. Casuarius beccarii Scl., Proc. Zool. Soc. Lond. p. 87, figs. 1, 2 on p. 86 (Vokan, Aru
Islands !) ;
1877. Forbes, Proc. Zool. Soc. Lond. pp. 307, 316 (anatomy) ;
1878. A. B. Meyer, Journ. f. Orn. p. 300; Salvad. Ann. Mus. Civ. Genoa, xii. p. 421;
1881. Salvad., Mem. R. Ac. Se. Torino, (2) xxxiv. p. 197 (partim: Aru only) ;
1882. Id., Orn. Papuas. e Moluce. iii. p. 484 (partim: Aru) ;
1894. Schalow, Journ. f. Orn. p. 21 (egg, Aru) ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 596 (partim: Aru).
1875. (7) Casuarius bicarunculatus (non Scl.!) Beccari, Ann. Mus. Civ. Gen. vii. p. 717 (Aru).
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. ALY
1881. (?) Caswarius galeatus (non Bonn.!) Scl., Zool. Voy. ‘Challenger,’ Birds, p. 64 (young,
bought in Ternate—seem to be young of C. c. beccarii from the Aru Islands; ef. Cat.
B. Brit. Mus. xxvii. p. 597).
1884. (?) Caswarius sp. A. B. Meyer, Zeitschr. ges. Orn. i. p. 296 (egg from Aru—as there are,
however, several Cassowaries inhabiting the Aru Islands, there is no certainty whatever
about an egg without history !).
Adult. Plumage black; casque very high, broad and massive, dark brown; bill long
and not pointed. Head and occiput pale greenish blue. Upper part of hind-neck
deeper blue, followed by a space of mixed blue and red, producing a somewhat livid
pale purple effect. Lower part of hind-neck scarlet. Chin, throat, and fore-neck
dark blue. Wattles large, broad, and very conspicuously joined at base, of a deep
fleshy-red colour, which colour runs up in a narrow line nearly to below the ear.
Naked lower sides of neck mixed red and blue. A very large bird, but I have not
been able to measure a fully adult dead example.
Young (three-fourths grown). Fore-neck indigo-blue, head and occiput pale blue ;
hind-neck orange-yellow, with a purple patch between blue and orange. Naked lower
sides of neck, anterior half dull blue, posterior half orange-yellow. Plumage brown.
Young (about full-grown). Same as adult, only colours less bright and plumage
brown.
Hab. Vokan, Aru Islands.
Dr. Sclater (P. Z. 8. 1875, p. 87) first recognized the Aru Cassowary as a distinct
species, and gave a woodcut, showing a side view of the head and the wattles trom in
front, accompanied by the following diagnosis :—
“ Casuarius similis C. australi et crista part modo elevata: sed caruncula cervicis una
media, ad apicem divisa.
“ Hab. Wokan, inss. Aroensium (Beccari).
“ Mus. civit. Genoeusis.”
In the same volume our author also mentions a Cassowary from the south
coast of New Guinea, of which he believes that it ‘* probably is of the same species
as that of the Aroo Islands,” namely C. beccarii. ‘This specimen was afterwards
described as C. sclateri, but later on reunited with C. beccarii, and so it has remained
also in the ‘Catalogue of Birds.’ However, I am fully convinced that the form from
Southern New Guinea can be separated from that from the Aru Islands, where it
inhabits, so far as we know, only the northern island of the Vokan group.
I am not able to give descriptions of well authenticated eggs of Beccari’s Cassowary,
but there is an enormous egg in the British Museum from the Aru Islands, which is
named C. bicarunculatus, which, however, from its large size I am more inclined to
believe is that of either C. ¢. beccarii or C.c. violicollis. It measures 140 : 98 mm.
Its granulations are low.
Schalow (/. c.) describes an egg brought from the Aru Islands by C. Ribbe, measuring
VOL. XV.—PaRT Vv. No. 2.— December, 1900. T
118 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
128: 89 mm. It is, however, not ascertained that it is an egg of C. ¢. beccarw, which,
as we know, is not the only Cassowary living in the Aru group. That the eggs of any
Cassowary are more transparent than those of any other species is evidently not the case.
3. CASUARIUS CASUARIUS SCLATERI Salvad. D?’Albertis’s Cassowary.
1844. Casuarius sp. S. Miill., Land- en Volkenkunde, p. 22 (Utanata, Princess Marianne Strait).
1875. Casuarius beccarii (non antea, p. 87, quod Aru!) Scl., Proc. Zool. Soc. Lond. p. 527,
pl. lviii. (S. New Guinea) ;
1880. Casuarius beccarii (non Scl., P. Z. S. 1875, p. 87!) D’Albertis, Nuova Guinea, pp. 494, 588 ;
1881. Salvadori, Mem. R. Ac. Sc. Torino, (2) sxxiv. p. 197 (partim: New Guinea only), pl. i. fig. 5 ;
1882. Id., Orn. Papuas. e Moluce. iii. p. 484 (partim: New Guinea only !) ;
1895. Id., Cat. B. Brit. Mus. xxvii. p. 596 (partim: New Guinea only).
1877. Casuarius australis (non Wall!) D’Albertis, ‘Sydney Mail,’ p. 143; id., Ann. Mus. Civ.
Genova, x. p. 19 (Fly River) ; id., Ibis, p. 372.
1878. Casuarius sclateri Salvad., Ann. Mus. Civ. Gen. xii. p. 422 (coast opposite to Touan or
Cornwallis Island, S.E. New Guinea) ; A. B. Meyer, Journ. f. Orn. p. 300 ;
1879. Sharpe, Ibis, p. 116 (type now preserved in Brit. Mus. !).
Adult. Casque dull brown, very high and much compressed laterally, very thin, and
when fully adult curved over to one side. Bill long and stout, black. Wattles very
large and long, sometimes over 7 inches, but in the old birds they get torn and
damaged either in fighting or when creeping through the thick scrub, so that often
they are very short or split into a number of ragged ribbon-like appendages. ‘This is
well illustrated in the series of figures of D’Albertis’s fine specimens in Count Salva-
dori’s monograph. This is the largest form of the C. caswarius group aud also the
largest Cassowary, standing, when erect, as much as 6 feet high and the body being
fully half as big again as that of an Emu (Dromq@us). Plumage black, bead and
occiput pale whitish blue. Fore-neck and nape brilliant indigo-blue. Hind-neck of
a most intense bright scarlet. Naked lower sides of the neck, posterior 3 brilliant
scarlet, anterior 4 bright blue. Total length about 1600 mm., bill from gape 130 to
140 and even 145 mm., tarsus 280 to 300 mm., inner claw 80 to 95 mm.
Juv. (two-thirds grown). Plumage dark brown, often nearly as black as adult, also
occasionally pale yellowish brown. Fore-neck leaden blue; head and occiput pale
blue. Hind-neck dull orange-yellow. Naked lower sides of neck, anterior 4 blue,
posterior % yellow.
Chick. Head and neck dark rufous, variegated with black bands and _ spots.
Longitudinal dorsal bands very broad, about 13 inches wide, black, slightly variegated
with rufous, and 5 in number—~z. é., a central one and two lateral pairs; one band on
flank extending down the thigh on each side, and a second one indicated by a number
of dark brown irregular spots.
Hab, Southern New Guinea from Port Moresby to the Princess Marianne Strait,
and perhaps even up to McCluer’s Inlet.
I have had several examples of this bird alive, though none quite adult. They
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. UY)
appear to be delicate and never to live long, and none ever seem to reach the size in
captivity which the species attains in a wild state. Adult wild specimens of this race
are the largest Cassowaries known. The fine adult bird from which the description and
accompanying photograph were taken was shot by Mr. Emil Weiske on the Brown River
in 8.E. New Guinea.
Fig. 1.
Casuarius casuarius sclatert.
Sclater’s Cassowary was at first thought to belong probably to Beccari’s Cassowary ;
it was then separated, and afterwards united with the latter by Count Salvadori. ‘The
original diagnosis of this form is as follows :—
‘“‘ Medius, niger, casside parum alta, crassa, superne rotundata, latere posteriore fere
verticali, brevissimo, crasso; paleare unico, mediocri, ad apicem diviso ; ungue digiti
interni breviuscula ; loris et mento violaceis ; capite et gula ceruleis, paleare rubro.
“Long. tot. 1™ 500, rostri hiat. 147 mm., tarsi 300, ungu. dig. int. 80 mm.”
TZ
120 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
A most magnificent series of this bird was procured on the Fly River by D’Albertis,
and a number of woodcuts of the heads and necks of them are given in the Annals of
the Genoa Museum and in Count Salvadori’s splendid monograph of the genus
Casuarius, and a coloured figure, from a sketch of D’Albertis, on plate i. (fig. 4) of
the same work.
Some eggs from near Port Moresby, which I received together with a chick, are
covered with glossy glazed granulations, which are not at all connected, but stand
more or less separated from each other. They are bright green, and measure 150: 103,
146: 100, 146: 97, and 146: 95 mm.
4, CASUARIUS CASUARIUS SALVADORII Oust. Salvadori’s Cassowary. (Plate XXIV.)
1875 ?. Casuarius tricarunculatus Beccari, Ann. Mus. Civ. Genova, vii. p. 717 (Warbusi) ;
1878. Salvad., Ann. Mus. Civ. Genova, xii. pp. 419, 420 ;
1881. Salvad., Mem. R. Ac. Sc. Torino, (2) xxxiv. p. 184;
1882. Salvad., Ann. Mus. Civ. Genova, xviil. p. 413 ;
1882. Salvad., Orn. Pap. e Moluce. ii. p. 473 ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 591 ;
1896. Oustalet, Nouv. Arch. Mus. Paris, (3) vill. p. 263 (type in Mus.).
1878. (Feb. 28rd.) Casuarius salvadorii Oustalet, Bull. Assoc. Sc. de France, Bull. no. 539, p. 350
(“ Warbusi ”’”) ;
1878. Scl., Proc. Zool. Soe Lond. pp. 213, 214, fig. 213 (Wandammen; head and neck); A. B.
Meyer, Journ. f. Orn. p. 202; Salvad. Ann. Mus. Civ. Genova, xii. p. 420;
1879. Salvad., Ibis, p. 105;
1881. Salvad., Mem. R. Ac. Se. Torino, (2) xxxiv. p. 204, pl. 1. fig. 5 ;
1882. Salvad., Orn. Papuas. e Moluce. ii. p. 488 (footnote) ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 595.
1878. Casuarius altijugus Scl., Nature, xvil. p. 875 (Wandammen) ;
1879. Salvad., Ibis, p. 105.
Adult. Plumage black; casque high, brownish black; bill thick, long, and not
pointed. Head and upper part of hind-neck pale greenish blue, rest of hind-neck
scarlet. Naked lower sides of neck blue anteriorly, purplish crimson posteriorly.
Chin, throat, and fore-neck deep blue. Wattles smal] and widely separated, base and
inner half blue, rest dark fleshy pink. ‘Total length about 13 m., tarsus 300 to
316 mm., bill from gape 150 to 190 mm., inner claw about 80 mm.
Chick. Head and neck rufous; rest of body creamy white or pale buff, with three
longitudinal dorsal black bands variegated with rufous. The three lateral stripes
along the flanks and thighs are broken up into a number of irregular brown spots
and patches.
Hab. North-west New Guinea.
Casuarius casuarius salvadorii was originally described by Oustalet and Sclater as
C, salvadorit and C. altijugus respectively, from skins obtained by Laglaize near
Wandammen on the south-west coast of Geelvink Bay, in North-west New Guinea ;
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 121
and to this day we know nothing absolutely certain as to the colour of the naked parts
of wild-shot Arfak birds; but there are a certain number of birds sent alive to Europe
as coming from New Guinea, and which agree neither with Caswarius casuarius from
Ceram, nor with C. ¢. beecarti from Aru, nor C. c. sclatert from Southern New Guinea
in colour, and these birds I, for the present, take to be Casuarius casuarius salvadorii
as they agree in other respects.
The bird from which I made the above given description lived two years at Tring
and one year in the Society’s Gardens. It is fully adult, and a male. I have had
several younger ones, but none lived to maturity.
The bird shortly mentioned by Beccari in his ornithological letter (Ann. Mus.
Genova, vii.) as agreeing in the form of the helmet with C. caswarius, but having
three wattles, namely, two lateral ones, and below these a smaller median third one,
was a young bird bought at Warbusi, a little to the south of Dorei. It was left at
Ternate to grow up, where Laglaize saw it still alive in July 1877, but it was
eventually lost. It must, in my opinion, have been an aberration of C. ¢. salvadorii.
Oustalet says that the type of C. salvadorii came also from Warbusi, at the bottom of
Geelvink Bay. Warbusi, however, is near Dorei, while Wandammen is the place
at the southern end of the Bay whence the other specimens from Laglaize were said to
come. Therefore Salvadori (1881) thinks that Warbusi might be a mistake for
Wandammen; but I see no reason why it should not be correct, since C. tricaruncu-
latus 1s not a species according to my judgment, as I have a similar aberration of
C.¢. sclateri. Laglaize probably bought his specimens from Bruijn or his hunters.
When excited these birds extend the neck horizontally and emit five or six very
loud grunting and grating sounds; frequently, when very angry, they spit several times,
elevate their wing-rudiments and the feathers of the hind-quarters, and peck and kick
most viciously.
5. CASUARIUS CASUARIUS INTENSUS Rothsch. Blue-necked Cassowary. (Plate XX VII.)
1898. Casuarius casuarius intensus Rothsch., Bull. B. O. Club, viii. (nos. lviii., Ixiv.), pp. xxi, lv.
Adult. Casque very high and laterally compressed, dark brown; bill pointed, short,
black. Plumage black. Size small. Occiput, nape, upper half of hind-neck, chin,
throat, and fore-neck deep brilliant blue. Lower half of hind-neck brilliant scarlet ;
between the blue and red of the hind-neck an irregular black patch. Naked lower
sides of neck of an intense blue. Two wattles on fore-neck medium-sized, oval, close
together, but separated their entire length, blue outer edge, and a few small spots pink.
Bill from gape about 110 mm., tarsus 275 mm., greatest length of helmet 130 mm.
Young (half-grown). Plumage brown; head and neck dull blue; lower hind-neck
dull orange-yellow. Lower naked sides of neck blue.
Young (three-fourths grown). Plumage brown, mixed with a few black feathers.
Fore-neck dark blue; head and occiput indigo-blue, paler round ear. Lower
hind-neck orange-red. Lower naked sides of neck bright blue.
122 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
Young (full-grown). Plumage black, mixed with brown feathers. Head and neck
deep blue, paler around the ear ; lower hind-neck scarlet. Naked lower sides of neck
very bright blue.
Hab. unknown.
In addition to the type I have had five others alive, and they all agreed in having
the intense blue colour and the entirely blue naked lower sides of the neck; but this
Cassowary remains still a very little-known form, and will do so till its habitat is
discovered. The original description is as follows :—
« ¢ ad. The casque differs from that of C. caswarius in being very high and much
more erect ; the wattles almost entirely blue instead of dark red; the blue of the head
and neck uniform and very dark; the orange of the hind-neck much restricted and
separated at the upper end from the blue by a black crescent-shaped patch ; naked
sides of lower neck entirely uniform blue instead of red, bordered anteriorly with
blue.”
6. CASUARIUS CASUARIUS VIOLICOLLIS Rothsch. Violet-necked Cassowary. (Plate XX VI.)
1899. Casuarius casuarius violicollis Rothsch., Bull. B. O. C. viii. (mos. lix., Ixiv.) pp. xxvii, lv.
Adult. Casque low, sloping back very much, laterally compressed, horny brown,
green at base. Bill very long, straight and pointed. Plumage black. Face bluish
green. Occiput and upper hind-neck pale greenish blue, throat and fore-neck brilliant
deep blue. Hind-neck orange-scarlet. Naked lower sides of neck bright violaceous
mauve, bordered anteriorly with a narrow line of blue, posteriorly with a narrow line
of scarlet. Two wattles on fore-neck large, round, and close together in female; small,
long, and wide apart in male, blue at base, rest fleshy pink.
Juv. (half-grown). Plumage brown; neck and head dull leaden blue; hind-neck
dull gamboge-yellow; lower naked sides of neck dirty flesh-pink.
Juv. (three-fourths grown). Plumage brown, with a few black feathers; head and
occiput pale dull blue; fore-neck dull indigo; hind-neck orange; lower naked sides
of neck dark pink streaked with purplish red.
Hab. ¢ Trangan Island, Aru Islands.
The type of the species was imported along with two specimens of C. bicarunculatus
from Aru, and is a female, for she has laid an egg; but I have since received seven
others of this form, and they all agree in their characters. As the other islands of the
eroup are inhabited by different forms of Cassowaries, I am inclined to think that
Trangan Island must be the home of this interesting form.
I described this form at the January meeting of the British Ornithologists’ Club in
1899 as follows :—
“This apparently undescribed form of Cassowary is most nearly allied to Caswarvus
casuartus salvadorii, but differs conspicuously in the colour of the naked parts and in
the very large size, which fully equals that of C. c. australis.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 123
‘Bill much longer and straighter than in other Cassowaries. Casque horny brown,
green at base. Face and a broad band running down the side of the bill bluish green.
Base of lower mandible dark blue, with a yellow line running along one-third of the
length of the mandible on each side. Wattles at base of fore-neck very large, round,
and short, 3 x 2% inches, pale blue at base, otherwise pink all over, entirely separate
for their whole length, but close together. Auricular orifice larger than in any other
Cassowary. Throat and fore-neck bright ultramarine-blue. Occiput and upper hind-
neck pale greenish or eau-de-Nil blue. Lower hind-neck brilliant orange-scarlet.
Naked lower sides of neck magenta-purple, bordered anteriorly with ultramarine-blue,
posteriorly with orange-scarlet; the magenta-purple space deeply carunculated and
sharply cut off from the red and blue borders, which are plain and smooth.”
The “ song” of this bird is an extraordinary performance, generally as follows: —
It lowers its head and neck and remains in this position—with head and neck
stretched out straight in front—for about fifteen seconds, with the bill open and
gradually inflating its neck, without making a sound; then, bowing and jerking its
head so that the bill and wattle clap together, it emits some barking grunts, apparently
with great effort, the skin of the neck being then so much inflated that the wrinkles
become quite obscure.
The type specimen laid an egg last year, which is, needless to say, bright green;
but its shell is not properly developed, and a description of the surface and size is
therefore better omitted.
7. CASUARIUS CASUARIUS AUSTRALIS Wall. The Australian Cassowary. (Plate XXV.)
1854. Casuarius australis Wall, Illustr. Sydney Herald of June 3rd ;
1857. Gould, Proc. Zool. Soc. Lond. pp. 269, 270 ;
1858. G. R. Gray, Proc. Zool. Soc. Lond. p. 196;
1860. Scl., Proc. Zool. Soc. Lond. p. 210; Ibis, p. 310 ;
1862. Scl., Trans. Zool. Soc. Lond. iv. p. 360; Schleg. Jaarb. zool. Genootsch. Nat. Art. Mag.
p. 200;
1865. Gould, Handb. B. Austr. ii. p. 206;
1866. Schleg., Zool. Gart. p. 180; Scl. Proc. Zool. Soc. Lond. p. 557 ;
1867. Scl., Proc. Zool. Soe. Lond. p. 242 (C. johnsoni=australis) ; Bennett, Proc. Zool. Soc.
Lond. p. 473 ;
1868. Scl., Proc. Zool. Soc. Lond. p. 376; Ibis, p. 348 ;
1868. Ramsay, Proc. Zool. Soc. Lond. pp. 381, 388 ;
1869. Gould, B. Austr., Suppl. pls. 70, 71; Krefft, Ibis, p. 348 ;
1870. Editors, Ibis, pp. 119, 120 ;
1871. G. R. Gray, Hand-list B. iil. p. 2, no. 9851; Flower, Proc. Zool. Soc. Lond. p. 32
(skeleton) ; Scl., Proc. Zool. Soc. Lond. p. 547 (in confinement) ;
1873. Schleg., Mus. Pays-Bas, Struthiones, p. 9;
1874. Ramsay, Proc. Zool. Soc. Lond. p. 320 ;
1875. Scl., Proc. Zool. Soc. Lond. p. 85 ;
124 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
1876. Ramsay, Proc. Zool. Soc. Lond. p. 119 (life-history) ; Rams., Proc, Linn. Soc. N. 8. Wales
i. p. 186;
1877. Rams., Proc. Linn. Soc. N.S. Wales, ii. pp. 196 (distribution), 376 (woodcuts of head) ;
Harting, Ostr. & Ostr. Farming, p. 95; Salv. & Sel., Ibis, p. 327 ;
1881. Salvad., Mem. R. Ac. Se. Torino, (2) xxxiv. p. 192, pl. 1. fig. 3 (head) ;
1882. Salvad., Orn. Pap. e Moluce. iii. p. 473 ;
1888. Rams., Tabul. List Austr. B. p. 19 (distribution) ;
1889. Lumholtz, Among Cannibals, pp. 98, 99;
1890. North, Nests & Eggs Austr. B. p. 294, pl. xv. (egg) ;
1891. Sharpe, Cat. Osteol. Spec. R. Coll. Surg. ii. p. 439;
1892. Schalow, Zeitschr. f. Ool. p. 11;
1894, Schalow, Journ. f. Orn. p. 24;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 594;
1896. Meston, Proc. Roy. Soc. Queensland, x. pp. 59-64.
1866. Casuarius johnsonii F. Miller, Australasian of Dec. 15th (cf. Proc. Zool. Soc. Lond. 1867,
pp. 241, 242) ; ;
1867. Carron & Bennett, Proc. Zool. Soc. Lond. pp. 473, 474; Krefft, Proc. Zool. Soc. Lond.
p. 482; Diggles, Orn. Austr. pl. & text ;
1868. A. Newton, Ibis, p. 348 (critical note on Diggles) ;
1869. Krefft, Ibis, p. 348 (maintains that johnsoni is correct name) ;
1870. A. Newton, Ibis, p. 120 (contends with others that the name australis is to be used).
1873. ? Casuarius regalis (nomen nudum) Rosenberg, Journ. f. Orn. p. 390.
Adult. Casque very high and massive, compressed laterally, deep brown; bill long
and stout, black. Next to C. casuarius sclateri, the largest of the forms of Caswarius
casuarius. Plumage black. Sides of head, occiput, and upper hind-neck pale greenish
blue; lower hind-neck scarlet, running a little into the blue of upper portion. Chin,
throat, and fore-neck deep indigo-blue. Lower naked sides of neck mixed red and
blue, giving a purple hue. ‘Two wattles on fore-neck very large, deep pinkish red,
mottled with paler pink, blue at base.
Young (quarter-grown). Head and hind-neck chestnut rufous. Chin, throat, and
fore-neck densely covered with short downy feathers of a brownish-buff colour. Rest
of body clayey brownish yellow mixed with dark rufous.
Young (two-thirds grown). Plumage brown. Head and occiput dull pale blue;
lower hind-neck orange; fore-neck leaden blue; lower naked sides of neck blue,
mixed with livid purple.
Chick. Head and hind-neck pale rufous; fore-neck yellowish buff; rest of body
yellowish buff. From base of neck to end of tail along the back run three broad
longitudinal black bands, variegated with rufous, and each about seven-eighths of
aninch wide. On the sides are three irregular wavy black bands extending from the
shoulder-girdle down the sides to the beginning of the metatarsus.
Hab. Northern Queensland.
I have kept the adult female, described above, since 1890: she laid some 40 or 50
eggs at Tring, and has now lived nearly two years in the Society's Gardens.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 125
The first discovery of the fact that a Cassowary existed in Australia is due to
Mr. Thomas Wall, who, like so many other explorers, left his bones in the waterless
deserts of the Australian continent. Wall procured a specimen, but this was lost, and
only a poor description, evidently made from memory, was published in the ‘ Illustrated
Sydney Herald’ of June 3rd, 1854, by Thomas Wall’s brother, William Sheridan Wall.
The account ran as follows :—
“The first specimen of this bird was procured by Mr. Thomas Wall, naturalist to
the late expedition commanded by Mr. Kennedy. This was shot near Cape York, in
one of those almost inaccessible gullies which abound in that part of the Australian
continent. The Cassowary, when erect, stands about 5 feet high. The head is
without feathers, but covered with a blue skin, and, like the Emu, is almost without
wings, having mere rudiments. The body is thickly covered with dark brown wiry
feathers. On the head is a large protuberance or helmet of a bright red colour, and to
the neck are attached, like bells, six or eight round fleshy balls of bright blue and
scarlet, which give the bird a very beautiful appearance. The first, and indeed the
only, specimen of the Australian Cassowary was unfortunately left at Weymouth Bay, and
has not been recovered. Mr. Wall being most anxious for its preservation had secured
it in a canvas bag and carried it with him to the spot where, unfortunately for himself
and for science, it was lost. In the ravine where the bird was killed, as well as in
other deep valleys of that neighbourhood, they were seen running in companies of
seven or eight. On that part of the North-eastern coast, therefore, they are probably
plentiful, and will be met with in all the deep gullies at the base of the high hills. The
flesh of this bird was eaten, and was found to be delicious; a single leg afforded more
substantial food than ten or twelve hungry men could dispose of at one meal. The
Cassowary possesses great strength in its legs, and makes use of this strength in the
same manner as the Emu. ‘Their whole build is, however, more strong and heavy than
that of the latter bird. They are very wary, but their presence may be easily detected
by their utterance of a peculiarly loud note, which is taken up and echoed along the
gullies; and it would be easy to kill them with a rifle.”
Most incorrect as this description is, there can be no doubt that it actually refers to
the Australian Cassowary, and we can therefore, in accordance with all other authorities,
accept the name of australis for this bird, and need not substitute that of johnsonii
for it.
This form is most closely allied to the Ceram Cassowary and to Sclater’s Cassowary.
The key, descriptions, and plates, however, will show the differences.
This bird is known only from Northern Queensland, from Cape York south to
Rockingham Bay, Of no other Cassowary are we so well acquainted with the life-
history. The most complete description of the latter is that of Ramsay in the Proc.
Zool. Soc. Lond. 1876, pp. 119-123. It runs as follows :—
“ One of the chief objects of my visit to Rockingham Bay was to become acquainted
VOL. XV.—PaRT y. No. 3.—December, 1900. U
126 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
a
with the habits of this noble bird. In 1872 I had sent my collector, Edward Spalding,
to this district for this purpose, but with very poor results.
“While in Brisbane on my way up I purchased by telegram a fine young living
specimen, the first that had been obtained and reared, and ultimately succeeded in
bringing it alive to Sydney and shipping it to the Society, where I am glad to hear it
arrived safely. I found also that several very young Cassowaries had been obtained,
and, for the first time, a nest and eggs had been found. ‘This was great news; and I
need not relate how I made all haste to the Herbert River Police Camp, where I was
most hospitably entertained and welcomed by Inspector Johnstone, who was the first
to rediscover and bring under the notice of others the existence of this remarkable
species. I found Inspector Johnstone a true sportsman, as well as an ardent lover of
nature, a zealous and energetic naturalist, and a careful observer. I am indebted to
this gentleman for much valuable information respecting the manners and customs
of the aborigines, and notes on the habits of many birds and animals new to me, and
especially for information on the present species.
“The Australian Cassowary is a denizen of the dense dark scrubs scattered over the
district of Rockingham Bay, and extending as far north as the Endeavour River. It
was tolerably plentiful only a few years ago even in the neighbourhood of Cardwell ;
but since the advent of sugar-planters, etc. on the Herbert River and adjacent creeks,
these fine birds have been most ruthlessly shot down and destroyed for the sake of their
skins, several of which I saw used for hearth-rugs and door-mats. Formerly they were
easily enough procured; but latterly so wary have they become, and their numbers so
decreased, that it is only with the greatest amount of patience that even a stray shot
can be obtained. I know of no bird so wary and timid; and although their fresh tracks
may be plentiful enough, and easily found in the soft mud on the sides of the creeks
or under their favourite feeding-trees, yet the birds themselves are seldom now seen.
During the day they remain in the most dense parts of the scrubs, wandering about
the sides of the watercourses and creeks, diving in through the bushes and vines at
the slightest noise. Toward evening and early in the morning they usually visit
their favourite feeding-trees, such as the native figs, Leichardt-trees, and various
species of Acmena, Jambosa, Davidsonia, &c.; they appear to be particularly fond of
the astringent fruit of the Leichardt-trees and of a species of Maranta, which produce
bunches of large seed-pods filled with juicy pulp, resembling in appearance the inside
of a ripe passion-fruit (Passiflora edulis). Fruits and berries of all kinds are eagerly
sought after; the tame semiadult bird which I had the pleasure of forwarding to the
Society (1875) became so fond of the fruit of the Cape-Mulberry that he would allow
no one to come near the tree he had taken possession of. ‘This bird has frequently
devoured at a time as much as three quarts of ‘loquats’ (fruit of Hriobotria japonica),
and several fair-sized oranges whole, besides its usual amount of bread per diem (about
3 pounds). In nature, I found that in the afternoons they frequently came out and
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 12%
walked along the scrubs, or along the sides of the river or creeks, and swallowed large
quantities of pebbles and small rough-edged stones. In confinement, plantains and
sweet potatoes (in large pieces, which they swallow whole) are a favourite food, while
nothing seems to come amiss to them—grasshoppers, spiders, earthworms, cockroaches,
caterpillars of all kinds, dough, and even raw meat. ‘They ascertain the flavour of their
diet by first taking it up in the tip of their bill and giving it a slight pinch ; and if not
suitable, they throw it away.
“J found they invariably refused green loquats, but always picked them up in the
bill first to try them. In confinement they become very tame, and may be allowed to
walk about the place without restraint, coming when called, or more often running after
and following any one accustomed to feed them. If disappointed or teased, they not
unfrequently ‘show fight’ by bristling up their feathers, and kicking out sideways or in
front with force sufficient to knock a strong man down—a feat I have witnessed on more
than one occasion. ‘These birds are very powerful, and dangerous to approach when
wounded. On more than one occasion a wounded bird has caused a naturalist to take
toa tree; the sharp nail of the inner toe is a most dangerous weapon, quite equal to
the claw of a large Kangaroo, and capable of doing quite as much execution.
“T found the Cassowaries to be excellent swimmers, and frequently tracked them
across a good-sized creek or river. On Hinchenbrook Island, situated about 14 mile
from the mainland, they have been frequently met with; and I have myself heard
them calling at night and early in the morning as I passed up the channel, at a distance
of a least two miles from them. Mr. Johnstone informs me he met with one swimming
across a river of considerable width during his explorations while on the ‘ North-east
Coast Exploring Expedition.’ Their note, most usually emitted by the male, is a series
of harsh guttural prolonged croakings quickly repeated, and continued for about three
minutes; it is very loud, and may be detected across the water at a distance of at
least three miles on a still night. I have listened to it resounding through the scrubs
at a distance of a mile anda half on land, and then thought it close and one of
the most unearthly noises I ever heard. They breed during the months of August and
September. ‘The first nest procured was found by some of Inspector Johnstone’s
black troopers, from whom Mr. Miller, a settler on the Herbert River, purchased
some of the eggs. One which he kindly presented to me is of the light green variety
mentioned hereafter. The nest consists of a depression among the fallen leaves and
débris with which the ground in the scrubs is covered, with the addition of a few more
dry leaves. The place selected is always in the most dense part, and well concealed
by entangled masses of vegetation. The eggs were five in number in the only two
instances recorded; and in both cases one of the eggs in each set differed from the
others, being of a light green colour, and having a much smoother shell. The others
all have a rough shell, covered rather sparingly with irregular raised patches of dark
but bright green on a lighter green and smooth ground. In the pale (No. 1) variety
u 2
128 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
these raisings on the shell are closer together, and not so well developed; in both
varieties they are more thinly spread over the central portion than at the ends. On
the whole they closely resemble the eggs of Caswarius bennettii, in which similar
variations are noticeable; but they are larger and of a greater diameter, being greatest
in the middle. I am indebted to Inspector Robert Johnstone for the fine series of the
eges of this species which at present grace my collection.
“The following are measurements of some of the specimens of the eggs of both
species :—
“ Casuarius australis.
Length in inches. Width in inches.
“No. 1. Light-green smooth shell . . . . 5:33 x 3°73
No. 2. Dark-green rough shell . . . . . 53 x 3°88
“ Casuarius bennettii.
“No. 1. Light-green smooth shell . . . . 5°65 x 3°54
No. 2. Light-green rough shell . . . . . 5°82 x 3°31
No. 8. Light-green rough shell . . . . . 5:34 x 3°4
No. 4. Dark-green rough shell . . . . . 52 x 3°32
“The young of Caswarius australis are of a dull rusty brown, the feathers having
frequently a blackish shaft-stripe, giving to the back a streaked appearance. After the
first year the plumage takes a deeper, lighter brown hue, and black feathers begin to
appear mixed with brown, some being parti-coloured. After the second season, at the
age of from 18 to 24 months, the black feathers predominate, and the helmet, which has
hitherto been undeveloped, more like the shield of a Coot (fulica), begins to show a
keel or ridge in the centre, which rapidly increases in height. ‘The skin round the head,
on which still remain a few brownish hair-like feathers, begins to become wrinkled
and coloured, varying from bluish-green to orange on the lower part, and bright blue
on the sides of the neck, the wattles becoming carmine. ‘The helmet still remains
comparatively small and undeveloped long after the wattles and naked parts of the
neck become coloured. I believe that the helmet does not attain its full size until
the fourth or fifth year at least.
“Tn traversing these scrubs the head is carried low to the ground, and the vines and
branches of trees striking the helmet slide over it on to the back. Otherwise, in the
dense vine-scrubs bordering the Herbert River and elsewhere, progress would be
greatly impeded; but as it is, the Cassowaries traverse the scrubs with wonderful speed,
jumping over fallen trees and logs when in the way. <A young bird (the identical
specimen, I believe, forwarded by His Excellency the Marquis of Normanby to the
Society), while in the possession of Inspector Johnstone, during my visit succeeded in
jumping out of its yard over a fence more than 6 feet in height. I measured the fence,
and found it 6 feet 6 inches to the top rail, on which its feet-marks were plainly visible ;
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 129
the length of the yard was only 12 x 12 feet. I found the adult Cassowaries in full
moult in March; but the new feathers had not all made their appearance in May.
During these months specimens in confinement were remarkably irritable and frequently
sulky, even refusing their food (which they invariably do when unwell), and were at
times very spiteful, even attacking their keepers; but strangers chiefly come in for a
share of their dislike. At all times I have noticed they are very fond of bathing; the
semiadult bird before alluded to, which I forwarded to the Society, was remarkable in
this respect, and might frequently be seen waiting at the pump in the yard until some
one came for water, when he would sit down quietly under a copious shower, stretching
out his neck and ruffling his feathers up to allow the water to reach the skin. They
do not like any exposure, and always endeavour to get out of the sun. In the wild
state they seldom leave the scrubs, and certainly never do so in the heat of the day
unless hard pressed; but on the whole they are remarkably hardy, and bear confinement
well. In February last (1875) I purchased four fine young birds about 6 months old,
which were obtained from some settlers in the Herbert River district; these also I
forwarded to England during the same month.”
The eggs of C.c. australis cannot be distinguished from those of C. caswarius,
although most of them, but not all, are slightly larger. Eggs laid in my aviary
at Tring measure :—140:93, 131:87, 136: 88, 138:89, 1380: 93, 135: 95, 132: 95,
128:91, 128: 92, 129: 92, 121: 91 mm.; two in the British Museum measure 130: 84
and 143:91 mm.
8. CASUARIUS BICARUNCULATUS Scl. The Two-wattled Cassowary. (Plate XXVIII.)
1860. Casuarius bicarunculatus Sclater, Proc. Zool. Soc. Lond. pp. 211, 248, 249, fig. (young:
locality unknown) ;
1862. Id., Trans. Zool. Soc. Lond. iv. p. 358, fig. 6, pl. Ixxi.;
1862. Crisp, Proc. Zool. Soc. Lond. p. 187 (gall-bladder) ;
1862. Schlegel, Jaarb. zool. Genootsch. Nat. Art. Mag. p. 198 ;
1864. Id., Dierent., Vogels, p. 239;
1866. Scl., Proc. Zool. Soc. Lond. p. 168 ;
1866. Schleg., Zool. Gart, p. 178; id., Nederl. Tijdschr. Dierk. iii. pp. 250, 347 ;
1867. Rosenberg, Reis naar Zuidoostereil. p. 52;
1869. Scl., Proc. Zool. Soc. Lond. p. 149 (young, alive);
1872. Scl., Proc. Zool. Soc. Lond. pp. 150, 495, pl. xxvi. ;
1878. Garrod, Proc. Zool. Soc. Lond. pp. 470, 644 (anatomy) ;
1873. Schleg., Mus. Pays-Bas, Struthiones, p. 10;
1875. Rosenberg, Reist. Geelvinkb. p. 117 ;
1878. Salvad., Ann. Mus. Civ. Genova, xii. p. 316 (in Bremen Mus.) ; Rosenb. Malay. Arch. p. 373;
1881. Gould, B. New Guinea, v. pl. 73; Salvad., Mem. R. Ac. Se. Torino, (2) xxxiv. p. 185, pl. 1.
fig. 1 (head) ;
1882. Id., Ann. Mus. Civ. Genova, xviii. p. 414; id., Orn. Pap. e Moluce. ii. p. 475 ;
1894, Schalow, Journ. f. Orn. p. 19 (error: egg, Fly River !!) ;
130 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 591.
1857. (?) Casuarius galeatus Wallace, Ann. Mag. Nat. Hist. xx. p. 477 (Aru Islands !).
1858. (?) C. emeu (non Latham!) G. R. Gray, Proc. Zool. Soc. Lond. p. 187 (sternum from Aru).
1861. (?) Casuarius kaupi (non Rosenberg!) G. R. Gray, Proc. Zool. Soc. Lond. p. 438 ;
1863. Rosenberg, N. Tijdschr. Ned. Ind. xxv. p. 252 (non p. 251) (Aru); id., Journ. f. Orn.
p- 1385 (non p. 184, quod unappendiculatus !).
1866. Casuarius aruensis Schleg., Nederl. Tijdschr. Dierk. i. p. 347.
1867. Casuarius galeatus (non Vieillot!), Rosenberg, Reis naar Zuidoostereil. p. 52.
1884. (?) Casuarius sp. A. B. Meyer, Zeitschr. ges. Orn. 1. p. 296 (egg from Aru).
Native name: ‘ Kudari,” in the Aru Islands (fide Rosenberg).
Adult. Plumage black ; casque compressed at sides, not flattened at back, low and
deep, horn-brown. Bill long, pointed, black. Head and occiput pale greenish blue.
Upper part of hind-neck deeper blue, rest of hind-neck scarlet ; fore-neck dark blue.
On each side of the neck is a long pendent wattle, blue at base, deep pinkish red for
the rest of the surface. Lower naked sides of neck deep blue. 1
Young (three-fourths grown). Plumage brown. Hind-neck dull orange ; fore-neck
and head dull blue; wattles pale pink.
Young (half-grown). Plumage yellowish brown. Hind-neck and wattles dull whitish
flesh-colour ; fore-neck and head dull greyish blue.
I have received from Dr. Finsch, among a number of chicks of Cassowaries in the
Leyden Museum, one brought by von Rosenberg in 1866 from Kabroor Island, Aru
group, which is clearly C. dicarunculatus. It appears to be much faded, but is very
different from any other chicks I have seen; it is rufous all over, and has five dorsal
bands and two bands of irregular patches on the thighs of a darker, more brownish
rufous colour,
Hah. Wwammer and Kabroor, Aru Islands.
The two birds on which the descriptions here given are based have been kept alive
by me, both at Tring and in the Society's Gardens, and the older of the two has been
in England five years.
The first notice that appeared about this Cassowary was the following note by
Dr. Sclater on a young individual without locality (P. Z. 5. 1860, p. 211):—
“Casuarius bicarunculatus, a name I propose to apply to a Cassowary of which I have
recently obtained a young example for the Society in exchange from the Zoological
Gardens at Rotterdam. It is easily distinguishable by the throat-caruncles being
placed far apart on the sides of the throat, lighter colouring, &c. As the bird itself will
shortly arrive in this country, I hope to be able to give full particulars concerning this
new species at the next Meeting of the Society.” This was done (p. 249), and the throat-
wattles were figured. Adult individuals were first brought to London in 1872.
Valentyn (Oud en nieuw Oost-Indien, iii. p. 299) first mentioned that a Cassowary
existed in the Aru Islands, and that it differs from C. caswarius, but he gave no
HON, W. ROTHSCHILD ON THE GENUS CASUARIUS. 131
description of it. Wallace (1857) tells us that a Cassowary (which he calls C. galeatus)
is not at all uncommon in the Aru Islands, that the young are brought in numbers to
Dobbo, where they soon become tame, running about the streets, and picking up all
sorts of refuse food. It is probable that the birds he saw were mostly C. bicarun-
culatus. A sternum only was procured by Wallace.
Our first knowledge of the actual home of this bird is due to von Rosenberg,
who (‘ Reis naar Zuidoostereil.’) actually shot a Cassowary on the Aru Islands on April
15th, 1865, which he called C. galeatus, but which proved to be C. bicarunculatus. He
tells us (from native sources) that the pairing-season is in June and July, and that the
sexes live separate except at that time of the year: that the female makes a sort of
rough uest and sits on the eggs for about 28 days; but both these latter reports are
doubtless erroneous, as the observations in the Zoological Gardens have proved that
it is the male alone which sits, and that the time of incubation is much longer.
Rosenberg also tells us that not more than five eggs are found in the nest, and that
some more eggs are placed outside the nest to serve as food for the young when
hatched! ‘This ridiculous story—which is also told of the Ostrich—is, it is needless to
say, just as reliable as the former assertions. In spite of his want of knowledge of the
habits of the birds, however, Rosenberg deserves full credit for discovering the home
of C. bicarunculatus, which he obtained in the northern Aru Islands, Kabroor and
Wammer. ‘The native name of this bird in Aru is “ Kudari.” Specimens have from
time to time arrived alive in Europe, and I have now two alive, both received when
almost chicks.
Eggs from the Aru Islands have been described as those of C. bicarunculatus, but as
two or three Cassowaries are found in that group of islands, there is not @ priori any
absolute certainty about any of these eggs, which, however, do not seem to differ very
remarkably from those of C. casuwarius and its allies. ‘The egg identified as C. bicarun-
culatus in Nehrkorn’s collection is said by Schalow (Journ. f. Orn. 1894, p. 19) to be
from the Fly River, but in Nehrkorn’s ‘Katalog der Kiersammlung’ it is said to
be from the Aru I[slands!! It is therefore not an egg to depend on. Nehrkorn says
that it is so dark blue-green that it resembles the eggs of Dromeus, and that it
measures 135:90 mm. According to Schalow its glazed granulations are less
continuous and less numerous than those of C. casuarius, but this may be an individual
character. Schalow also distinguishes between Cassowary eggs that are * durch-
scheinend”’ and ‘ undurchscheinend ” when held against the light; but this is by no
means a specific difference, being mostly due to the way in which the eggs are emptied,
and the thickness of the egg-shell varies also considerably.
Four eggs in the Leyden Museum from Kabroor, collected by Rosenberg, have more
or less lost their original green colour, but agree in other respects. ‘The corrugations
are more or less interrupted and more or less elevated. ‘The measurements are:
LAV Sie seriou. MNS 2129786) nome
132 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
9. CASUARIUS UNAPPENDICULATUS Blyth.! One-wattled Cassowary. (Plates XXIX. &
XXX.)
1860. Casuarius sp., Blyth, Ibis, p. 193, and Sclater, P. Z.S. p. 210 (reprint Ann. & Mag. Nat.
Hist. ser. 8, vol. vi. p. 145) ; Casuarius unappendiculatus, Blyth, Journ. As. Soc. Beng.
vol. xxix. p. 112 (juv., no locality), reprint Ann. & Mag. Nat. Hist. ser. 3, vol. vii. p. 113;
C. unoappendiculatus, Blyth, Ibis, p. 307; C. uniappendiculatus, Bennett, Ibis, p. 403,
pl. xiv. (head and neck, from live bird in Amsterdam).
1861. C. unappendiculatus Blyth, Journ. As. Soc. Beng. vol. xxx. p. 193.
1862. C. wniappendiculatus Sclater, Trans. Zool. Soc. Lond. vol. iv. p. 359, pl. 74 (juv., fig. mala,
typus) ; Schleg., Jaarb. zool. Genootsch. Nat. Art. Mag. p. 198, pl.; id., Dierentuin,
p. 239, cum fig. ;
1866. Scl., P. Z. S. p. 84 (first descr. of egg); Schleg., Zool. Gart. p. 179; id., Ned. Tijdschr.
Dierk. vol. iit. p. 250 (Salwatti and opp. coast) ;
1869. Gould, Suppl. B. Austr, pls. 74, 75 ;
1871. Schleg., Nederl. Tijdschr. Dierk. vol. iv. p. 53; Gray, Hand-list, ii. p. 2, no. 9852 ;
1873. Schleg., Mus. Bays-Bas, Struthiones, p. 10;
1874, A. B. Meyer, Sitzb. der k. Ak. Wiss. Wien, vol. lxix. p. 218; Scl., Ibis, p. 417, note;
1875. Scl., P. Z.S. p. 583 (exact locality, Threshold Bay); Rosenberg, Reist. Geelvinkb, p. 117 ;
1877. Forbes, P. Z. 8. pp. 307, 313, 314, 316 (bursa Fabr.) ;
1878. Salvad., Ann. Mus. Civ. Genova, xii. p. 846 (chick, Sorong); Rosenb., Mal. Arch. p. 396
(kaupi=uniappend., woodcut) ;
1881. Salvad., Mem. R. Ac. Se. Tor. (2) xxxiv. p. 205, pl. i. fig. 6;
1882. Salvad., Ann. Mus. Civ. Gen. xviii. p. 414, no. 5; id., Orn. Pap. e Mol. iui. p. 490;
1886. Beddard, P. Z. 8. p. 145 (anatomy) ;
1893. A. B. Meyer, Abh. Mus. Dresd. no. 3, pl. ii. (egg) ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 597 ;
1896. Oust., Nouv. Arch. Mus. Paris, (3) vill. p. 264.
1861. C. kaupi Rosenb., Natuurk. Tijdschr. Nederl. Ind. xxii. p. 43, pl.; id., Journ. f. Orn.
p. 44, pl. i.;
1862. Schleg., Jaarb. zool. Genootsch. Nat. Art. Mag. p. 199;
1863. Rosenb., Natuurk. Tijdschr. Ned. Ind. xxv. p. 251;
1864. Id., Journ. f. Orn. p. 186;
1866. Scl., P. Z.S. p. 168 (=uniappendiculatus) ; Schleg., Zool. Gart. p. 180 ;
1867. Rosenb., Reis naar Zuidoostereil. p. 52;
1873. Schleg., Mus. Pays-Bas, Struthiones, p. 12; Rosenb., Journ. f. Orn. p. 390 (definite assertion
of identity with C. wniappendiculatus).
1894. C. unoappendiculatus Schalow, Journ. f. Orn. p. 23 (egg),
Native names: “ Mambeba” at Sorong, “ Kallo” in Salwatti (Rosenberg).
Adult. Casque low, very much flattened behind, black ; bill short, thick, and black.
Head and upper part of hind-neck pale blue, with a greenish tinge. Cheeks dark
* By an error, on the four Plates of the various forms of C. unappendiculatus, as well as on the distribution
map, the name is spelt uniappendiculatus. The original spelling, however, is without the 7.—W, R.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 133
blue; face-wattles lopg, broad and thin, very bright dark blue. Chin dark blue;
throat and a small part of hind-neck dark dull olive-yellow; fore-neck and rest of
hind-neck dark golden-yellow. A golden-yellow transverse patch on occiput. Naked
lower sides of neck dark orange-yellow. Plumage black ; wattle dirty mauve. Size
very large. Height from casque to ground about 5 feet 6 inches when walking, 6 feet
8 inches when erect.
Total length about 1655 mm., bill from gape 133, tarsus 280, claw of inner toe 84.
(After Salvadori, JZ. c.)
Young (half-grown). Casque brown; plumage yellowish brown; neck and naked
lower sides of neck pale yellowish flesh-colour. Head, chin, and occiput dirty bluish
green ; legs yellowish.
Chick. Head and neck uniform rufous; throat buff; body creamy buff, with five
distinct brown longitudinal bands on the body and a line of broken brown patches along
the thighs.
Hab. Salwatty and opposite coast of N.W. New Guinea (‘Threshold Bay, Tangion-
Ram, Sorong).
The first communication referring to the One-wattled Cassowary is the following
editorial note in ‘ The Ibis’ for 1860, p. 193:—* Mr. Blyth in his last letters (dated
Calcutta, Jan. 8th & 21st) speaks of an apparently new species of Cassowary (Casuarius)
in the aviary of the Babu Rajendra Mullick of Calcutta: ‘It has a yellow throat, a
single yellow throat-wattle, and a long strip of naked yellow skin down each side of the
neck. In its present (first) plumage, it is of a much lighter colour than the young
of the Common Cassowary of the same size, two of which are kept along with it;
and from the size of the legs, it is easy to perceive that when full-grown it is a
much smaller species.’ ”
In the ‘Journal of the Asiatic Society of Bengal,’ Blyth first published the
name :—
“Casuarius unappendiculatus, nobis, n.s., from its peculiarity of having but a
single pendulous caruncle in front of the neck. Specimen apparently more than
half-grown, and much paler in the colouring of its plumage than specimens of the
same age of the common C. galeatus, two fine examples of which are associated with
it in the same paddock. In lieu of the two bright red caruncles of the latter, the new
species has but a single small oblong or elongate oval yel/ow caruncle, and the bright
colours of the naked portion of the neck are differently disposed. The cheeks and
throat are smalt-blue, below which is a large wrinkled yellow space in front of the
neck, terminating in front in the oval button-like caruncle, and its lower portion being
continued round behind, while on the sides of the neck the yellow naked portion is
continued down to its base, the bordering feathers more or less covering and concealing
this lateral stripe of unfeathered skin ; on the hind part of the neck the bare yellow
skin is not tumous and corrugated as in the Common Cassowary, where also this part
VOL. XV.—ParT Vv. No. 4.— December, 1900. x
134 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
is bright red. The casque is about equally developed at this age in the two species.
The legs of the new species are smaller, from which I doubt if it attains to quite so
large a size as the other.”
The next we hear concerning this species is the arrival of an immature bird in the
Zoological Garden of Amsterdam ; this grew up, and thus was afterwards the first adult
described and figured by Schlegel. Also in this case Rosenberg was the first to
discover the actual home, for he obtained it on the island of Salwatti, and afterwards
specimens became known from several places on the opposite coast.
The bird in Amsterdam laid a pale green egg, ‘‘ thickly covered with raised spots of
dark green, and measuring 136: 89 mm.”
I have an egg in my collection (from the Buckley collection, bought from T. Cooke,
1871, who had it in exchange from ‘Mr. Franks”: this means probably that it was
from the Amsterdam dealer Frank, who had it from the Zoological Gardens, for it has
on its label “ Maart 1865” (or 6, the 5 being obliterated), this being about the time
when the bird laid eggs there). This egg differs from other Cassowary-eggs in having
numerous single glazed round knobs, like pin-heads, few of them being connected
with each other. The descriptions of the two eggs of C. unappendiculatus by Schalow
(J. c.), both from different sources, agree with mine—the peculiar isolated pin-head-like
elevations being well described and discussed as distinguishing the wnappendiculatus-
egg from all others. One of those in the British Museum, about the origin of which
there is no doubt, as it is one of the eggs laid in Amsterdam, agrees also with these in
having the separate knobs, while two others, from Frank, and one said to have come
from Salwatti, are more like those of Casuarius casuarius. ‘The one with the isolated
knobs measures 137: 88 mm., my own 151: 81, Schalow’s 146: 96, 148-0: 49°5 mm.,
while Sclater gives only 136: 89. Sclater also speaks of raised “spots” only. On the
other hand, seven eggs in the Leyden Museum, sent by Bernstein and Rosenberg, and
said to be from Sorong and Salwatti, do not show these characteristic isolated pin-head-
like elevations, but rather a more connected network of glazed granulations. ‘They
measure 145: 90, 145: 91, 145: 90, 145: 91, 147: 92, 142: 95,145:96 mm. They
are all very large.
It must therefore be left to future research whether the peculiarly isolated knobs on
some well-authenticated eggs of C. unappendiculatus are a specific character; but at
present this view cannot be taken, unless the identity of all the eggs in the Leyden
Museum and of some of those in the British Museum is doubted.
As usual, we know nothing definite respecting the biology of this bird in a wild state.
tosenberg tells us that an immature bird he got in Ternate was running about freely
and was much attached to people, while it was a fierce enemy to cats and dogs. When
it got angry it put up its feathers and emitted a peculiar blowing cry, followed by a
grunting like that of a young pig.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 135
10, CASUARIUS UNAPPENDICULATUS OccrIPITALIS Salvad. Jobi Island One-wattled Casso-
wary. (Plate XX XI.)
1875. (?) Casuarius papuanus Rosenberg, Reist. Geelvinkb. p. 117 (Jobi) ; Casoar di Jobi Beccari,
Ann. Mus, Civ. Gen. vii. p. 718.
1875. Caswarius occipitalis Salvad., ibid. (footnote) ;
1876. Scl., Ibis, p. 245 (note) ;
1878. Salvad., Ann. Mus. Civ. Gen. xu. p. 423; Scl., Ibis, p. 482; A. B. Meyer, Journ. f. Orn.
pp. 203, 300 ;
1881. Salvad., Mem. R. Ac. Se. Tor. (2) xxxiv. p. 209, pl. ii. fig. 7 (head) ;
1882. Salvad., Ann. Mus. Civ. Gen. xviii. p. 414, no. 6; id., Orn, Pap. e Mol. iii. p. 494 ;
1893. Meyer, Abh. Mus. Dresd. no. 3, p. 29, pl. ii. fig. 4 (egg) ;
1894. Schalow, Journ. f. Orn. p. 22 (egg) ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 598 ;
1896. Oust., Nouv. Arch. Mus. Paris, (3) viii. p. 264, pl. xiv.
1892. Casuarius westermanni Rey (nec Sclat.), Zeitschr. f. Ool. p. 19.
Native names : “ Orawei,” Jobi (Rosenberg) ; ‘ Orawai” at Ansus on Jobi Island (4. B. Meyer).
Adult. Casque very high, conical, flattened posteriorly. Beak long and very stout.
Iris deep brown. Face, head, chin, throat, and upper half of neck deep blue, rather
paler on hinder part of neck. Face-wattles long, thin, pendent, and much swollen at
their base. Lower half of neck deep yellow or pale orange. Pear-shaped wattle
yellow, more or less suffused with blue. Naked sides of lower neck deep crimson.
On the occiput is a transverse patch of dull dirty orange.
Plumage black and coarse; legs brownish grey; bill and casque black in most fully-
adult specimens, but some occasionally retain the greenish-horn coloured or pale olive
helmet of the immature bird. Legs dark olive.
The measurements of the type (cf. Salvadori, J. ¢.) are as follows:—Total length
about 13 m., bill from gape 137 mm., tarsus 280 mm., nail of inner toe 70 mm.
Young (full-grown). Plumage brown; face-wattles absent ; naked parts dull bluish
green where deep blue in adult ; dull yellow instead of pale orange on lower half of
neck, and dirty red mixed with yellow on lower sides of neck. Casque dull green.
Young (half-grown) similar to that of C. wnappendiculatus, but casque bright green.
Chick. Head and hind-neck pale rufous; body pale buff; back with five black bands
variegated with rufous; flank and thigh stripes broken up into irregular spots.
This form differs from C. wnappendiculatus principally in its more slender build and
its greater height, the greater extension of blue on the neck, the crimson lower sides
of the neck, and the very high casque.
It inhabits the island of Jobi and perhaps also the opposite coast of Geelvink Bay.
There occasionally occurs in this form, and probably also in all the other forms of
C. unappendiculatus, a curious melanistic variety, where nearly the whole neck is blue
and only a small spot of yellow remains at the base of the hind-neck. ‘This was
Xx 2
136 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
described by Oustalet as a distinct species under the name of Caswarius laglaizei; it
may stand as
CASUARIUS UNAPPENDICULATUS OCCIPITALIS aberr. LAGLAIZEI.
[1893. Cusuarius laglaizei Oust., Bull. Soc. Philom. (8) v. no. 9, pp. 1-8 (Jobi Island) ; 1896.
Oust. Nouv. Arch. Mus. Paris, (3) viii. p. 265, pl. xv. (Specimen killed on Jobi and
purchased by Laglaize; type of C. daglaizei, in Paris Museum. Careful description
and measurements and plate to show the supposed specific value, in which, however, I
do not believe at all.) |
It is most probable that it was C. unappendiculatus occipitalis of which A. B. Meyer
(Sitzber. Ak. Wiss. lxix. p. 217) says that it occurs on Jobi, where it is called
“Orawai,’ and of which Rosenberg had received some reports from the natives when
stating (‘Reistochten,’ p. 117, and Malay. Arch. p. 563) that Casuarius papuanus
occurred also on Jappen—the usual Dutch name for Jobi Island in Geelvink Bay.
Beccari brougbt a grand adult male from Ansus, on Jobi Island, to Italy, and this was
at once recognized by Count Salvadori as an undescribed form. In a footnote appended
to Beccari’s interesting ornithological letter in the Ann. Mus. Civ. Genova, vii. p. 718,
our learned friend diagnosed this form as follows :—
‘ CASUARIUS OCCIPITALIS mihi.
‘“‘Casside compressa, postice depressa; appendice colli antici, ut videtur, nulla ;
capite, gula et parte superiore cervicis ceeruleis; macula occipitali et colli parte nuda
inferiore flavis; area nuda laterali colli imi carnea.” ‘This diagnosis is followed by a
more detailed Italian description. Needless to say, the Count always recognized the
close relationship this bird had to Casuarius unappendiculatus from Salwatti. In the
‘Catalogue of Birds,’ xxvii. p. 599, he says that it is ‘‘ very similar to C. uniappendi-
culatus, but differs in having a triangular yellow patch on the occiput, the posterior
surface of the casque narrower, the single caruncle smaller.”
The measurements of the type are, according to Salvadori:—Total length about
1 m. 670 mm., bill from gape 137 mm., tarsus 280 mm., claw of inner toe 70 mm.
One egg obtained by Doherty on Jobi Island, and doubtless belonging to this form,
unless more than one occur on that island, shows a good many rather isolated knobs,
thus closely resembling those of C. unappendiculatus, which show this character.
It measures 144: 95 mm.
1]. CasvUARIUS UNAPPENDICULATUS AURANTIACUS Rothsch. Eastern One-wattled Casso-
wary. (Plate XXXII.)
1899. Casuarius uniappendiculatus aurantiacus Rothsch., Bull. B. O.C. vol. viii. p. 1 (17th May),
p. lvi (21st June); id., P. Z. S. 1899, p. 774 (June 20th ; published Oct. Ist).
Adult. Face, head, chin, and nape of neck bright turquoise-blue, darker round eyes
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 137
and on face-wattles. Rest of fore- and hind-neck and naked lower sides of neck deep
bright orange, without any red. ‘Throat-wattle blue. On the occiput a very wide and
large transverse patch of bright orange. Face-wattles large and very thick, balloon-
shaped when inflated by the bird during anger. Bill short and thick. Casque low,
compressed laterally, and not flattened posteriorly or hardly so. Plumage black and
coarse; legs brownish grey; iris pale brown. Build slender and tall, but not so tall as
either C. wnappendiculatus or C. u. occipitalis.
Young and chick unknown.
This form differs both from C. wnappendiculatus and C. u. occipitalis in its slender
build and paler blue colouring, but principally in the uniform orange of the neck, the
laterally compressed and posteriorly not flattened casque, and in the thick swollen
face-wattles, which are very similar to those of the following species, Casuarius philipi.
Hab. German New Guinea.
This form is known from one individual living at present in the Zoological Garden
of Berlin, where Mr. Keulemans made the accompanying picture. It is most interesting ©
to find the one-wattled group of Cassowaries extending to Kaiser-Wilhelmsland, and I
have no doubt whatever that my aurantiacus is a very good subspecies.
My original description (/. ¢.) is as follows :—
“« Face, cheeks, and occiput pale sky-blue ; throat dark blue. Occipital patch, fore-
neck, hind-neck, and lower sides of the neck deep reddish orange. Casque horny
green, and much more compressed laterally than in C. wunappendiculatus. Long cheek-
wattles absent, but the sides of the face distended as in C. philipi.”
12. CASUARIUS UNAPPENDICULATUS RUFOTINCTUS, subsp. nov. Red-tinted One-wattled
Cassowary.
Casque greenish horn-colour, compressed laterally and not depressed posteriorly ;
plumage black, mixed with brown, as the bird is not quite in full adult plumage, though
otherwise in full colour. Head, face, occiput, and upper hind-neck bright pale blue ;
on the occiput is a large patch of orange-brown. Chin and throat deep indigo-blue,
which colour runs down the centre of the fore-neck in a triangular band to the base of
the wattle, where it is about 1 inch wide. Wattle blue. Sides of fore-neck orange-
scarlet, changing to crimson when excited and the throat distended. Lower naked
sides of neck deep crimson, with a narrow anterior yellow border.
Hab. unknown.
I had this bird as a chick in striped plumage, and it was reared as a pet at ‘Tring,
but since it has been in this Society’s Gardens (15 months) it has come into colour and
is very different from any other form of C. wnappendiculatus. It is now 3 years old,
158 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
13. Casuartus puinirt Rothsch. Sclater’s Cassowary. (Plate XXXIIL.)
1898. Casuarius philipi Rothsch., Nov. Zool. v. p. 418.
Almost adult. Casque very high, compressed laterally and at the same time depressed
posteriorly, thus combining both forms of casque; the colour of the casque pale
whitish brown behind, darker in front ; bill thick and apparently rather short.
Plumage black ; feathers long and much curled on the body, the webs being close
and not disintegrated as in other Cassowaries, more like the tip of a freshly-moulted
feather of Dromeus. Feathers of the rump and tail enormously lengthened, some
of the tail-feathers when fresh-moulted trailing on the ground. Head, occiput, face,
and upper part of hind-neck pale greenish Nile-blue; fore-neck and uppersides of
neck and sides of nape deep indigo-blue; lower hind-neck lemon-yellow, flecked
with a few reddish spots on the edges. Lower naked sides of neck bright cherry-
crimson. Cheek-pouches, when extended and inflated during anger, of a dark
lavender-blue colour. The feathers of the neck come very high up and are
very thick; on the lower part of the fore-neck is a single round flat wattle, upper
two-thirds cherry-red, lower third blue. On the occiput is a broad patch of dull
clay-brown. Cry unlike all the other Cassowaries, being very loud and resembling
a deep roar.
This, quite the most remarkable Cassowary, is-not only distinguished by the
colour of its naked parts, its curious casque and feathers, but also by its extra-
ordinary shape. It is quite as short from the ground to the level of its back as
Casuarius bennetti, but in every other respect it is a giant; in fact, it can only
be compared with the extinct New Zealand Dinornis (Pachyornis) elephantopus as
regards proportions.
Hah. unfortunately unknown. :
It was shipped to London from Calcutta, and when it first came to the Society’s
Gardens was in brown plumage, and the colours of the head and neck were not
developed. It was thought to be a young C. wnappendicuiatus, but as soon as it began
to come into colour, I saw it was something quite new, in fact it is the most distict of
all the species of Cassowary.
I originally described this bird when it was much younger. My description was as
follows :—
“This new species is founded on a bird now living in the Zoological Gardens,
London, which, though far from adult, being almost brown in plumage, appears to be
full-grown, and the naked parts are fully coloured. It is closely allied to C. wni-
appendiculatus, but differs much in colour.
** Plumage when adult evidently black. Casque as yet undeveloped, pale yellowish
horn-colour. Throat and fore-neck deep purplish blue. A single small wattle on
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 139
fore-neck, round and flat, not pear-shaped as in C. uniappendiculatus ; upper third of
wattle purplish red, rest dark blue. Hairy feathers of neck very thick and reaching
high up the neck. Head, occiput, and upper half of hind-neck very pale greenish blue ;
lower half of hind-neck pale orange-yellow. Naked skin on lower sides of neck deep
crimson, fading into cherry-red on the edges. Legs very stout and short; body set
very low on the legs and very bulky, giving the bird the exact shape Dinornis
elephantopus must have had.
‘** Hab. Probably Eastern German New Guinea!.
“Named in honour of Dr. Philip Lutley Sclater.”
14, Casuarius papuanus Schleg. Westermann’s Cassowary. (Plate XXXIV.)
1871. Casuarius bennetti (non Gould, 1857) Schleg., Nederl. Tijdschr. Dierk. iv. p. 53.
1871. Casuarius papuwanus Schleg. (ex Rosenberg MS.), t. ¢. p. 54;
1873. Schleg., Mus. Pays-Bas, Struthiones, p. 11 (Andai) ; Rosenb., Journ. f. Orn. p. 390;
1875. Rosenb., Reist. Geelvinkb. pp. 84, 117,144, pl. 17 ; Salvad. Ann. Mus. Civ. Gen. vii. p. 717
(Andai) ;
1878. Rosenberg, Malay. Archipel, p. 563; A. B. Meyer, Journ. f. Orn. pp.-200, 203 ;
1879. Salvad., Mem. R. Ac. Sc. Tor. (2) xxxiv. p. 210, pl. ii. fig. 8 (head) ;
1882. Salvad., Ann. Mus, Civ. Gen. xvii. p. 414, no. 7; id., Orn. Pap. e Mol. iii. p. 495 ;
1894. Schalow, Journ. f. Orn. p. 20 (egg).
1871. Casuarius kaupi Sclat. (non Rosenberg, 1861), P. Z. 8. p. 627 (Mansinam) ;
1872. Sclat., P. Z. S. pp. 147-150, pl. ix. (Mansinam) ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 599;
1896. Oust., Nouv. Arch. Mus. (3) vii. p. 265.
1872. Casuarius papuensis Sclater, P. Z. 8. pp. 149, 150.
1874. Casuarius westermanni Scl., P. Z. 8. p. 248; id., Ibis, p. 417 (footnote) ;
1875. Scl., P. Z. 8. pp. 85, 87, 380, pl. xix. ;
1877. Gould, B. New Guinea, v. pl. 15;
1878. A. B. Meyer, Journ. f. Orn. pp. 200-203.
1874, ? Casuarius sp. A. B. Meyer, Sitzber. k. Ak. Wiss. Wien, Ixix. p. 216.
Native names: the following are given by Rosenberg ; but it is, of course, doubtful whether
they all apply strictly and only to this species, or to several, or whether they are meant for any
form of Cassowary: “ Mswaar” at Andai, “ Nhamdia” at Hatam, “ Wonggé ” at Dorei.
Adult. Bill short and pointed ; casque low and much flattened posteriorly, so as to
make it quite triangular in shape. Face behind eye, and head white. Nape, throat,
and fore-neck deep blue. Hind-neck scarlet. Anterior half of naked lower sides of
neck livid magenta-purple, posterior half scarlet.
In the place where in the Caswarius casuarius and C. unappendiculatus groups
* There is, however, no certainty about its home.
140 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
the throat-wattles are attached there is sometimes a small round knob of the same
colour as the rest of the fore-neck. Plumage black; feathers long and more silky
than in the species previously described, those over the tail being some 12 to
15 inches long and very thick, forming a sort of pendent train. Legs brown, bill and
casque black.
Total length (according to Salvadori) about 1 m. 400 mm., tarsus 260 mm., bill from
gape 120 mm., inner claw 78 mm.
Young (full-grown). Plumage brown, all colours on naked parts duller, the blue
being of a dull leaden shade, and the red replaced by orange-yellow.
Young (half-grown). Plumage dark chestnut-brown or bay-brown, each feather
crossed by several strongly defined black bars. Head and neck not bare.
Chick. Uniform dark brown, with two or three narrow longitudinal white bands on
the body, and several similar lines on the neck.
This bird inhabits the Arfak Peninsula, and has been said to come also from Jobi,
but this is not supported by any evidence.
Rosenberg discovered this Cassowary near Andai, in the Arfak Peninsula of New
Guinea, where it is called “ Mswaar” or “‘ Meswaar” by the Papuans. When first
mentioning it, in 1871, Schlegel called it C. bennett?, but stated that it differed from
the latter in the coloration of the upperside of the neck, which, however, he did not
consider of any importance, and remarked that Rosenberg believed it to be a new
species which he wished to call Casuarius papuanus.
Rosenberg tells us that the first glance at the type of his C. papwanus, which was
shot by his hunter Achmat (Achmed), convinced him that his former belief that the
Cassowaries of New Guinea were the same as those from Salwatti, the C. wnappendi-
culatus, was erroneous. Rosenberg further states that the adult female was shot
during the breeding-season, which lasts from February to April, and that the nest
consists of a depression on the ground, lined with leaves. He had all this information,
however, from natives, and he repeats the old fable with which he also treated us when
describing the Ceram Cassowary, that some eggs (five, he says) are placed in the nest,
others (two) outside, so as to serve as food for the chicks, and that they are hatched in
25 days. Of the eggs he says that they are in every respect like those of C. casuarius.
‘The adult bird shot by his hunter was at first only wounded, and attacked the man, who
had not a little trouble in keeping it off, and subsequently killed it with his hunting-
knife. Bruijn sent skins to Europe said to be from Andai and Emberbaki, or Amber-
baki. Laglaize purchased one from Amberbaki, and A. B. Meyer obtained it near
Dorei. Beccari says it occurs also on the mountains of Arfak, where he saw foot-
prints of a Cassowary at a considerable elevation.
The type of C. westermanni belongs certainly to C. papuanus. Sclater informs us
that it was said by the missionary who sent it to come from Munsinam (= Mansinam),
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 141
near Andai, where also Rosenberg obtained C. papuanus. ‘There is no reason to
doubt this statement. Meyer (Sitzber. k. Ak. Wiss. Wien, Ixix. p. 216) says that
Mansinam is a small island near Dorei, where no Cassowary could live in a wild state,
and that therefore the locality “ Mansinam” is incorrect. However, there is also a
village called Mansinam (or Mansiman) a few miles south of Andai, and, moreover, as
“C. westermanni” does not differ from C. papuanus, it is evidently quite unnecessary
to suppose that the Mansinam, whence the type of C. westermanni came, is the island
of Manaswari (which is, according to Meyer, the proper name, its principal village
alone bearing the name of Mansinam). It is more likely the Mansiman of the maps
(cf. the map of Rosenberg and many others), which was quite recently found still to
exist and was visited by William Doherty.
The eggs of C. papuanus in the British Museum are rather smoother than most egos
of the C. caswarius group and others, the granulations being rather flat. It is possible
that this flatness of the granulations is the rule in the eggs of the C. papuanus group.
Those in the British Museum measure 134 : 86 and 130: 84 mm.
Unfortunately, by carelessly reading Dr. Oustalet’s description of C. papuanus edwardsi,
I erroneously gave Salwatty as a further habitat of this species, and it is thus marked
on the maps (Plates XL. & XLI.) which show the distribution of the genus.
15. CASUARIUS PAPUANUS EDWARDSI Oust. Milne-Edwards’s Cassowary. (Plate
XXXV.)
1878. Casuarius edwardsi Oust., P. Z. S. p. 389, pl. xxi. (Dorey) ; Salvad., Ann. Mus. Civ. Gen.
xi. p. 425; Cab., Journ. f. Orn. p. 203 (editorial note); A. B. Meyer, Journ. f. Orn.
Pp. 299)
1895. Casuarius papwanus (partim!) Salvad., Cat. B. Brit. Mus. xxvii. p. 600.
Adult. Plumage black, feathers on rump and tail much elongated. Casque much
flattened posteriorly and black; bill long, pointed, and black. Occiput, cheeks, and
sides of face white; below ear and cheeks a large black patch surrounded by a broad
pink band, this pink band extends right round the back of the head below the white
occiput, but is concealed in the folds of the skin unless the bird is violently enraged.
Chin, throat, and fore-neck deep blue, in centre of fore-neck usually a round black
wart with green centre. Nape greyish black, rest of hind-neck scarlet. Naked
lower sides of neck dull scarlet washed with livid pinkish purple, the spaces between
the folds black.
Juv. (half-grown). Reddish brown, feathers on fore part of body and back faintly
barred transversely with black. Occiput and sides of head bluish white, the occiput
being whiter, Face and part of head round casque dark leaden grey, naked lower
sides of neck pale flesh-colour; legs yellowish olive.
VoL. Xv.—PakT V. No. 5.—December, 1900. y
142 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
Juv. (three-fourths grown). Casque not yet developed. Crown of head and face
black. Occiput dirty brownish white. Fore-neck and upper half of hind-neck dull
blue. Large irregular patch between ear-hole and angle of gape buffy pink. Lower
half of hind-neck rosy salmon-red. Centre of naked lower sides of neck scarlet-mauve,
bordered with a colour mixed orange and cherry-rose. Plumage brown, curiously
variegated with black.
Hab. Low country of N.W. Dutch New Guinea.
My living specimen performed as follows :—
It first extends its neck horizontally, then contracts it short and bends it, at the
same time inflating it, and then makes a series of at least twelve short and deep
grunting notes. Sometimes, when very angry, it, like other Cassowaries, spreads the
feathers of the back and hind-quarters out and snarls viciously.
The young three-quarter grown bird described above is not yet old enough to
perform these antics.
Milne-Edwards’s Cassowary was first described by Oustalet from a male killed
near Dorei, and brought to Paris by Messrs. Raffray and Maindron. ‘he differences
noticed by M. Oustalet are principally in the form of the helmet and the
coloration of the naked parts of the head and neck. ‘The latter are, indeed,
very conspicuous ; while the form of the helmet is more likely to be an individual
character.
I cannot believe that C. edwards? is ‘‘identical” with C. papuanus; but if it is a
subspecies of the latter, then the two forms must inhabit different areas, and it is
very likely that one of them frequents the lowlands, while the other lives among the
hills.
An egg laid by a living bird in my possession has the network of granulations rather
flat, like those of C. papuanus in the British Museum. It is rather pointed and seems
abnormally small, measuring only 125 : 86 mm.
16. Casvartus Lort@ Rothsch. Loria’s Cassowary. (Plate XX XVIII.)
1895. Casuarius picticollis Salvad. (nec Sclater), Cat. Birds Brit. Mus. xxvii. p. 600 (description !).
1898. Casuarius lorie Rothsch., Novit. Zool. p. 518.
1899. Id., Bull. B. O. C. viii. p. lvi (June 2lst); id, P. ZS. p. 775 (June 20th; appeared
Oct. Ist).
Adult. Bill short and pointed, casque low and much flattened posteriorly. Throat
and mesial line of neck dull pink, with three narrow longitudinal lines of blue, rest of
fore-neck scarlet. In front of the ear a small patch of livid magenta, above the ear
blue. Head and hind-neck bright blue. Anterior portion of naked lower sides of
neck livid purple, posterior portion scarlet, the whole surrounded with blue. Plumage
black, long and silky. Legs horny grey. Iris hazel.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 143
The total length of the fully adult male in the Museum of Genoa is about 14 m.,
the bill from gape measures 120 mm., the tarsus 244 mm., claw of inner toe 117 mm.,
according to Salvadori. My adult bird from the Upper Brown River (collected by
Weiske) measures :— Bill from gape 122 mm., tarsus 246 mm., claw of inner toe
70 mm.
Young (half-grown). Rufous-brown, each feather marked with narrow transverse bars
of brownish black.
Hab. Highlands of S.E. New Guinea.
Casuarius lorie was found by Signor Loria at Moroka in the highlands of the
Owen Stanley Mountains in British New Guinea. The fully adult specimen obtained
by Loria is now in the Museo Civico of Genoa. Mr. Loria made a coloured sketch of
the head and neck, which was kindly lent me by Dr. Gestro, and from which our plate
is taken.
It is this same bird which is described in the ‘ Catalogue of Birds,’ vol. xxvii. p. 601.
Another adult bird, a skin which shows the red of the neck well, was sent to me by
Mr. Weiske from the mountains on the Upper Brown River. I cannot possibly con-
found these red-necked individuals with Caswarius picticollis, which has the neck blue
with a small red spot in the centre of the fore-neck, and a red chin when young.
C. lorie evidently replaces C. picticollis in the highlands, the latter being, according
to our present knowledge, only found in the coast-region.
The second example mentioned in the list of specimens in the ‘ Catalogue of Birds’
(7. c.) I could not find in the collection, at least not under that name.
17. Casuarivs PrcticoLLis Scl. Painted-necked Cassowary. (Plate XXXVI.)
1874. Casuarius picticollis Scl., Rep. Brit. Assoc. p. 138.
1875. Scl., P. Z. S. p. 84, pl. xvii. (Discovery Bay) ; ibid. p. 349 (pullus, Milne Bay) ;
1877. Gould, B. New Guinea, v. pl. 14; Forbes, P. Z. 8. pp. 307, 315, 316 (anat.) ;
1881. Salvad., Mem. R. Ac. Se. Tor. (2) xxxiv. p. 213, pl. ii. fig. 9 (head) ;
1882. Salvad., Ann. Mus. Civ. Gen. xviii. p. 415, no. 8; Salvad., Orn. Pap. e Mol. iii. p. 499 ;
1893. (?) A. B. Meyer, Abh. Mus. Dresd. no. 3, p. 29, pl. ii. figs. 1, 2 (egg) ;
1894. (?) Schalow, Journ. f. Orn. p. 25 (egg) ;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 600.
1881. Casuarius kaupi (non Rosenberg !) Sharpe, Ibis, p. 500.
Adult. Plumage black ; casque high and depressed posteriorly, black; bill pointed,
black. Occiput and auricular region pale greenish blue. Chin, throat, fore-neck, and
nape of neck deep bright blue. In centre of fore-neck a deep red patch. Lower hind-
neck pale blue. Lower naked sides of neck bright rose-purple.
Total length about 1 m. 400 mm., tarsus 245 mm.,, bill from gape about 120 mm.,
inner claw 125 mm.
Ww
144 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
Immature. The immature bird varies considerably, and I describe the two most
different :—
(1) Chin and throat dull red, nearly joining the red patch in centre of fore-neck ;
rest of naked parts as in adult, only duller in colour; plumage black, completely
mingled with brown feathers.
(2) Plumage brown, slightly mixed with black feathers. Chin, throat, fore- and
hind-neck, and occiput dull blue, base of hind-neck orange. In centre of fore-neck a
large horseshoe-shaped red patch, lower naked sides of neck dull purple.
Juv. (one-third grown). Uniform pale brown.
Hab. Coast-region of British New Guinea.
In 1875 (P. Z. S. p. 85) Dr. Sclater characterized this bird as follows :—
“In OC. westermanni (pl. xix.) the throat is blue and the hinder part of the neck
deep orange-red. In the new species, which I propose to call C. picticollis (pl. xviii.),
the middle of the throat is red, and the hinder part of the neck bright blue. There
are, besides, minor differences, which will be evident on comparing the two drawings.
Now, so far as I know, these colours in the naked parts of the Cassowaries are quite
constant; and I can hardly doubt therefore that we have here to deal with different
species.” The original specimen came from Discovery Bay. It died, after having
lived for about twenty-nine months in the Society's Gardens, and is now preserved
in the British Museum. Other individuals are known from Milne Bay. The
description of C. picticollis in the ‘ Catalogue of Birds in the British Museum’ is
that of a mountain form which I consider to be specifically distinct, and which I
have separated as Casuarius loriw, as | had been able to examine several specimens
of it.
18. CAsuARIUS PICTICOLLIS HECKI Rothsch. Heck’s Cassowary. (Plate XX XVII.)
1899. Casuarius picticollis hecki Rothsch. Bull. B. O. C. viii. p. xlix (May 17th), p.lvi (June 21st) ;
id., P. Z. S. p. 775 (June 20th ; appeared Oct. Ist).
Adult. Bill short and peinted; casque low and flattened posteriorly. Occiput Nile-
blue, rest of head and neck deep indigo-blue. Naked lower sides of neck bright
crimson, bordered with indigo-blue. Cheek-wattles large and balloon-shaped when
extended. Legs brown. Bill and casque black.
Young and chicks unknown.
This form differs trom C. picticollis in the absence of red on the throat and chin, in
the much darker blue. of the head and neck, and in the bright crimson instead of
purplish-red naked lower sides of neck.
Hab. German New Guinea.
HON. W, ROTHSCHILD ON THE GENUS CASUARIUS. 145
Nothing is known of this form except one living individual in the Zoological Garden
of Berlin. My original preliminary description of it (J. ¢.) is as follows :—
“This bird bears the same relationship to C. picticollis that C. papuanus edwardsi
does to C. papuanus. ‘Uhe throat and hind-neck are deep indigo-blue. Occiput pale
greenish blue. A small round black wattle on the fore-neck. Lower sides of neck
dark crimson. Casque and plumage similar to those of C. picticollis.”
This form is named in honour of Dr. Heck, Director of the Zoological Garden of
Berlin, who kindly permitted Mr. Keulemans to make a drawing of the bird.
19. CasuaRiIus BENNETTI Gould. The Mooruk. (Plate XXXIX.)
1857. Casuarius bennetti Gould, P. Z. 8. p. 269, Aves, pl. 129 ;
1858. Gould, Ann. & Mag. Nat. Hist. (3) i. p. 299 ;
1858. J. E. Gray, P. Z. S. p. 271, Aves, pl. 144 (egg) ;
1859. Bennett, P. Z. 8. p. 32; Gould, B. Austr., Suppl. pls. 72, 73; Scl., Ibis, pp. 212, 235 ;
1860. Bartlett, P. Z. S. p. 205, Aves, pl. 162 (egg); Scl., P. Z. 8. p. 210; Bennett, Gath.
Naturalist in Australas. p. 243, pl. iv. ;
1862. Scl., Trans. Zool. Soc. Lond. ix. p. 359, pl. Ixii.; Schleg., Jaarb. zool. Genootsch. Nat. Art.
Mag. p. 198;
1863. Scel., P. Z. 8S. p. 234 (incubation), p. 518 (descr. of pullus), pl. xlii.; Jouan, Mém. Ac. Soe.
Nat. Cherb. ix. p. 323;
1864. Scl., P. Z. 8S. p. 271 (successfully reared) ;
1865. Gould, Handb. B. Austr. 11. p. 561 ;
1866. Schleg., Zool. Gart. vil. p. 178;
1871. G. R. Gray, Hand-list, ii. p. 2, no, 9850;
1873. Schleg., Mus. Pays-Bas, Struthiones, p. 4; Benn. & Sclat., P. Z.S. p. 519 (corr. of locality) ;
Garrod, P. Z. 8. pp. 470, 644 (anatomy) ;
1875. Scl., P. Z. S. pp. 85, 87; Scl., Rep. Brit. Assoc. p. 112;
1876. Ramsay, P. Z. 8. p. 122;
18795 Scley BS Zs Sap: ols
1880. Layard, Ibis, p. 303; Powell, P. Z. S. p. 493;
1881. Salvad., Mem. R. Ac. Se. Tor. (2) xxxiv. p. 214, pl. ii. fig. 10 (head) ;
1882. Salvad., Ann. Mus. Civ. Gen, xvii. p. 415, no. 9; id., Orn. Pap. e Mol. iii. p. 501;
1891. Evans, Ibis, p. 84 (period of incubat.); Sharpe, Cat. Osteol. Spec. R. C. Surg. iii. p. 438 ;
1892. Ost, Zeitschr. Ool. pp. 15, 94; Schalow, ibid. p. 11;
1894. Id., Journ. f. O. p. 20 (egg) ;
1896. Oust., Nouv. Arch. Mus. Paris, p. 256.
1869. Cassowary from the Solomon Islands (error !), Hutton, Ibis, p. 352.
Native name: “ Moorup” in New Britain (corrupted: ‘ Mooruk’’).
Plumage black. Baill long, arched and pointed. Casque high, sharply depressed
posteriorly, the various ridges being sharply defined. Back of head, occiput, and
146 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
upper hind-neck black-blue; lower hind-neck dark blue. Chin, throat, and sides of
fore-neck deep bright blue, rest of fore-neck blackish blue. Below the ear-hole and
reaching to the gape is a large irregular patch of very pale blue. Upper half of the
naked Jower sides of neck black, with purplish-pink streaks between the wrinkles,
lower half purplish pink with black spots.
Total length about 1 m. 330 mm., bill from gape 110 mm., tarsus 240 mm., inner
claw about 73 to 90 mm.
Chick. Head and hind-neck bright rufous mingled with black spots. Chin
and throat yellowish buff. Fore-neck and sides of neck rufous buff mingled
with black. Rest of body creamy buff. Five longitudinal black bands variegated
with rufous running along the back, and two black bands down the flanks and
thighs.
Hab. New Britain.
The adult bird here described was sent me four years ago by Captain Cayley Webster ;
but it is impossible to say if the bird was adult then or not, for though the plumage
was black, the casque was undeveloped, and the colours were dull.
The Mooruk is an inhabitant of the island of New Britain (now most unreasonably
and unscientifically officially renamed ‘‘Neu Pommern”). It was for the first time
made known in a letter from Mr. George Bennett, communicated to this Society
by Gould, who proposed the name Casuarius bennetti for the new bird, and reproduced
a drawing made by Mr. G. F. Angas from the live bird, which was not quite mature.
‘The letter reads as follows :—
“My dear Gould, I send you an account of a new species of Cassowary recently
brought to Sydney by Captain Deolin in the cutter ‘Oberon.’ It was procured from
the natives of New Britain, where it is known by the name of ‘ Mooruk.’ The height
of the bird is 3 feet to the top of the back, and 5 feet when standing erect; its colour
is rufous mixed with black on the back and hinder portions of the body, and raven-
black about the neck and breast: the loose wavy skin of the neck is beautifully
coloured with iridescent tints of bluish-purple, pink, and an occasional shade of green,
quite different from the red and purple caruncles of the Caswarius galeatus; the feet
and legs, which are very large and strong, are of a pale ash-colour, and exhibit a
remarkable peculiarity in the extreme length of the claw of the inner toe on each
foot, it being nearly three times the length which it obtains in the claws of the
other toes. This bird also differs from C. galeatus in having a horny plate instead of
a helmet-like protuberance on the top of the head, which callous plate has the
character of and resembles mother-of-pearl darkened with black-lead. The form
of the bill differs considerably from that of the Emu (Dromeus nove-hollandia),
being narrower, longer, and more curved, and in having a black and leathery cere
at the base, and behind the plate of the head a small tuft of black hair-like
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS. 147
feathers, which are continued in greater or lesser abundance over most parts of the
necks,
The original specimen was sent to London by Dr. Bennett, and proved itself to be a
female by laying eggs. It was, like most Mooruks, much more gentle and tame than
most other Cassowaries ; and this seems to be a peculiarity of the Mooruk, which is said
to be often kept as a pet by the natives of New Britain. It bears itself less upright,
and its neck seems to be shorter than that of other Cassowaries.
There is a very long and amusing account of its habits in captivity in Dr. Bennett’s
‘Gatherings of a Naturalist in Australasia,’ accompanied by a figure by Wolf.
In 1863 the Mooruk hatched a nestling in the Society’s Gardens. he female
began to lay in the middle of March, and laid half a dozen eggs at intervals of about a
week, The incubation, according to Dr. Sclater, lasted once 49 days, the next time
52 days. The young did not live more than a day. The eggs were for the first
time described (and figured, P. Z. S. 1858) as being of a pale olive colour with darker
olive tubercles, but the eggs laid in London show that they are when fresh of a bright
apple-green colour (see figure, P. Z. 8. 1860) with darker green glazed granulations.
They do not differ perceptibly from the eggs of Caswarius casuarius and allies, though
the glazed caruncles are somewhat less continuous and not so thickly and equally
distributed over the entire surface of the egg. The following measurements have been
obtained :—137: 88, 128: 81, 141 to 131:90 to 83, 150:88, 185:89, 136: 90,
152:87 mm. The full fresh-laid egg weighs about 22 ounces. It will thus be seen
that there is hardly any difference between the eggs of C. bennetti and C. casuarius.
Schalow’s description of the colour is not from fresh eggs—all he saw were evidently
faded ones. ‘The smooth ones (see P. Z. S. 1858 and 1860) are first-laid eggs, the
roughly granulated ones the properly developed eggs.
The notes of my live specimen—evidently a male—are usually a low and short
piping note, reminding one of that of a large chicken, and when a little excited
this note is louder and more plaintive. When much excited it makes its neck quite
short, and utters first some higher, then some lower barking sounds followed by some
snarling ones.
The eggs of Casuarius bennetti in the British Museum have a very different surface,
hardly two being alike. They are of different shades of brown, but, needless to say,
that is due to exposure, as we know the fresh eggs are green like all other
Cassowaries’ eggs. One has very few, large and highly raised glazed granulations,
one is almost smooth (like first-laid eggs of Cassowaries generally), one exactly
like the eggs of C. papuanus. ‘They measure:—141:89, 127:86, 146°5:90, and
137:90 mm. Schalow gives 137:88 and 128°5:81°5 mm. Ramsay gives 141 to
131: 90 to 83 mm.
148 HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
20. CASUARIUS BENNETTI MACULATUS, subsp. nov.
Casque high, much depressed posteriorly, and sharply ridged. Plumage black;
bill long, arched, and pointed ; inner claw long. (This bird was first identified by me
‘as a second specimen of C. picticollis hecki.) Occiput and hind-neck bright indigo-
blue; sides of head, face, and the fore-neck paler and brighter blue. On the lower
part of fore-neck is a large round patch, 14 inches in diameter and of a rosy-purple
colour. Naked lower sides of neck rosy purple: from and below the ear-hole
extending to the gape is a large white patch washed with rose-colour.
Hab. unknown.
This form may eventually prove to be only a colour-aberration of C. bennetti; but,
so long as we do not know the exact habitat of every species and subspecies of
Cassowary, and the full extent of their individual variation, I consider it right to name
and describe this bird as a subspecies of C. bennetti. ‘The type is now living in the
Society's Gardens, and was fully adult when it was imported. It seems to be a male,
and is not so wild as my specimens of true C. bennetti usually were.
Lf ue]
Part II.—On the Morphology and Phylogeny of the Paleognathe (Ratite and Crypturi)
and Neognathe (Carinatz).
By W. P. Pycrart.
TasLE oF ConreNnts.
Piterylosis Page Nervous System and Sensory Organs. Page
CWIGONE SooosdGussonogeadanoSdadouwo 151 TALI er suetey svettsyereycteqt onsysscuaeersie crap ae eae 247
s mbryorandeNestlingsns i elaeciecieell: 159 TIVO! scvep cre sraiinen eer casas teased neste ese soars 247
Structure of Feathers ................-; 162 Jacobson’s cartilages..............-0 0005 247
BheyNeossoptilesmercstasrt-eeerer yells 162 Alimentary Canal and Appendages.
Adult and Nestling Remiges of Caswarius . 165 Buccal cavity; wes scveye yey atee ase eRe: 248
Heleop tiles teararrriratntnriir dt ntor rier 166 Intestinal convolutions ................ 248
Summary of Pterylological Characters 168 Liver and Gall-bladder ................ 251
Respiratory Organs.
Osteology. TUB Aiysie ayevete: ors orca armen curred tents 252
SMU OPRAGME Sooopodcunvedgaclc Go edce 171 AT rRasacd ssc ulee op ub ree tant peer area 252
RPE NGStIN SR irda rs tert ccctetensye sven eiysia's 195 WAN G pl pO} sievoi44 cay okies vonniorel aera mek ete 254
Some points of significance in the Struthious Choansey istcs ta a/paeistesnanea cet oer 256
Palate res reatecachre tetera Pers cere ero eis 206 Circulatory System,
WertebralaC@olumnime-reiariretirracirnc 208 PRC arb 7 Baereoscusvestue'c cunt henner hereaeeentenony ae 256
RAIDS ersatkcrsovehe taunts oousnnie mstentecn saeecsey 215 Carotidsstecuen Vereen aniane inet bic iceeere 257
Sternum and Pectoral Arch.............. 216 Cloaca, Bursa Fabricii, and Urino - genital
PelyicrAnchiryaqcmipestecvscierpte cistron 225 System,
Pectoral@limbmerrraryeacretserrir ctl acer 233 (CIE hen corse mare cote ano iinels dammo ood on 257
JRA IGN) Gooocoocod sopoogccuDSnoéoa 235 BUESAI “sheyelsvareterara stesteru-vey yess gah ence 258
CopulatonysOre antarctic eceere 259
Myology. Development errno eer 259
Nkin=musclesmanrrrrriecicrt nicer racers 238 Phylogeny of the Paleognathe .............. 259
Muscles of Wing and Shoulder-girdle 238 SUMMATY Mra te eter 266
Thigh- and Leg-muscles ...............- 242 Keys to Genera and Species .............+.. 267
INTRODUCTORY. »
THE present investigation was undertaken at the request and with the co-operation
of the Hon. Walter Rothschild, M.P., who has furthermore furnished the bulk of the
material, and in numerous other ways greatly aided my work.
The collection
of
skeletons in the British Museum of Natural History has proved a source of great
wealth.
I am besides indebted to Mr. Beddard, Dr. Gadow, Dr. Forsyth Major, and
Prof. Howes for the loan of specimens in their possession or charge, and I take this
opportunity of recording my thanks.
The object of this investigation has been to ascertain, as far so possible, the rela-
tions of the Casuartide to the remaining “ Struthious” forms, and the position of these
vou. xv.—Part Vv. No. 6.—December, 1900.
Z
150 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
with regard to the “ Carinate.” That this attempt has not been fruitless will probably
be admitted.
From the nature of the aim of this paper, it will I trust be agreed that the scheme
of investigation adopted is about the best possible. The desire has been to select those
characters which bear directly or indirectly upon the problem of the phylogeny of the
group. Moreover, to have dealt in detail upon the anatomy of the group would have
been to repeat a vast body of facts already well known and ably treated in the memoirs
enumerated at the end of this work, and from which much has been extracted to
serve the ends I had in view. Hence the majority of the omissions which may be
noted in these pages are not accidental but of design. The memoirs of Garrod, Gadow,
Fiirbringer, Meckel, Cuvier, D’Alton, Parker, Beddard, Lydekker, and others are the
milestones which mark the progress of the knowledge which has been accumulated on
this group. There seemed to me to be no reason or end to be gained in transcribing
the information which these individually impart, but rather it appeared more expedient
to set myself the task of recording the substance of their achievements, and to endeavour
to add a few courses more to the structures which they have reared.
The rejection of the old terms Ratite and Carinate, which I now propose, seems to
me to be warranted. The Ratite condition is admittedly a secondary one, and the name
is objectionable, in that Hesperornis and some “ Carinatw” are also “ Ratite.’ The
terms now suggested are mutually exclusive, and based upon a primitive character. On
this account they are to be preferred. Furthermore, as will be shown, the change is not
to be attributed to a desire to effect a change for change sake: it would not have been
suggested but for the need, and it seems to me a real need, to include the Crypturt with
the group hitherto known as “ Ratitw.” To have retained the old terms would not
perhaps have been so very contradictory, for, since some “ Carinate” are really “ Ratite,”
there is no reason why some Ratite should not be “ Carinate”! <A further discussion
of the subject will be found in the summary.
In conclusion, I wish to record my grateful thanks to Mr. Rothschild for having
entrusted this work to my hands.
-PTERYLOSIS.
The description of the pterylosis of the Palewognathe is most easily accomplished
by an enumeration of the apteria. Though these are but few in number, they are
nevertheless of considerable importance, inasmuch as they disprove the prevalent
notion that the feathers in the “ Ratite” are evenly distributed over the body. It
has, however, long been known that apteria occurred in embryos of certain forms,
e. g. Struthio, Rhea, and Apteryx.
The descriptions of the Otl-gland, Rhamphotheca, Podotheca, and of the structure of
the feathers, both macro- and microscopical, will be found in this section.
PHYLOGENY OF THE PALAHOGNATHA AND NEOGNATHA, 151
a. Pterylosis of the Adult.1
CASUARIIDA.
Casuarius casuarivs Linn.
Apteria :—
A. spinale.—This was wanting.
A. mesogastrei.—This includes the characteristic and well-marked sternal callosity,
and extends backwards to within a short distance of the cloacal aperture. It is
widest on the abdomen.
A, trunci laterale-—Represented by the naked under surface of the wing and a
small truncal area at its base.
The head and more or less of the neck are, in the adults of this and other
species of the genus, bare, and brilliantly coloured. The extent and nature of this
coloration have already been dealt with by the Hon. Walter Rothschild in the first
part of this Memoir.
Pteryle :—
P. alaris (P\. XLV. fig. 10).—It is possible to distinguish in this both remiges and
coverts as in Meognathe. The former are represented by the characteristic
elongated porcupine-like quills. These are six in number and distributed
as follows :—-mid-digital (1) and metacarpals (2) representing the primaries, and
cubitals or secondaries (3).
The cubital series, at first sight, appears not to extend backwards to the elbow as
in Neognathe, the last of the spine-like remiges being attached not farther down
the ulna than its proximal fourth. Beyond this, however, are three moderately long
feathers occupying the position of remiges, but bearing each an aftershaft. Similarly
at the distal end of the series, we have, in front of the mid-di gital, a feather occupying
the position of a remex—possibly representing the addigital—but small and bearing
an aftershaft. This, like the three cubitals, appears to have a major covert, as also
have the other spine-like remiges. The mid-digital is very small; the other remiges
are of considerable length. The longest of these are the Ist metacarpal and the Ist
cubital.
The remiges are remarkable in that they are open at the free end (see page 166).
They are cylindrical, of irregular thickness and more or less crooked.
The tectrices are not distinguishable from the rest of the body-feathering. All bear
aftershafts. Coverts representing the tectrices majores of the Neognathe are certainly
’ As“ adult”? we have included nearly full-grown as well as full-grown specimens, 7. e. those which have not
yet attained the characteristic adult livery.
Zz 2
152 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
present, and bear the same relation to the remiges, Other coverts—t. mediw, nuinores,
and marginales—are not distinguishable.
There is no ala spuria.
Neither in this, nor in any other species of the Palewognathe are there any semi-
plume, plumule, or filo plumule. The absence of these last is of more significance
than of the two former. ‘There are no rectrices. ‘The oil-gland is wanting.
Fyelashes are present and well developed.
The Rhamphotheca of the upper jaw is compound, being composed of a small median
rhino- and small lateral gnathotheca. The nasal fossa is not protected by the rhampho-
theca, but is covered only by the integument. The anterior nares are impervious,
slit-like, slope obliquely forward, and placed at the extreme end of the fossa.
The Podotheca (fig. 2 A, p. 155) has the acrotarsium clothed, below with broad scutes,
above with large granular scales; the acropodium with scutes. The claw of the inner
toe is greatly elongated, to serve as a weapon of offence.
The wing is not armed with the blunt claw found in many other species.
C. casuarius beccarit (Sclater).
The pterylosis of this subspecies agrees with that of the typical form C. caswarius.
In one specimen I found 6 remiges, the additional quill representing a primary.
C. casuarius sclateri.
‘The pterylosis of this agreed with the two foregoing. In one half-grown specimen
the apterium spinale could be faintly traced from the inner surface of the skin. ‘The
wing bore a long blunt claw.
C. casuartus salvadorit.
Apter. spinale——Though not of great extent, in this species was well defined. It
extended* cephalad as far as a point corresponding with the exit of a vertical line
passing through the body from the posterior end of the sternal callosity, and caudad to
the middle of the pelvis above the acetabulum. Its total length was about 8 inches,
its width about 4 inch.
A. mesogastr@i.—Very narrow, and bounded by degenerate feathers, indicating a
tendency to increase the width of this space, and recalling similar conditions in the
Carinate. It could be traced backwards to within a short distance of the cloacal
aperture.
A. trunci laterale.—Represented by the naked under surface of the wing.
Pieryla alaris.—As in C. casuarius.
The wing was armed with a claw.
PHYLOGENY OF THE PALZOGNATHAZ AND NEOGNATHA. 153
C. CASUARIUS VIOLICOLLIs.
C. UNAPPENDICULATUS.
A. spinale.—As in C, casuarius salvadorii.
A. mesogastrei—This was much more clearly defined than in C. casuarius. Its
width exceeded that of the apterium spinale.
I have not been able to examine spirit-specimens of nestling Cassowaries. Con-
sequently I am unable to say whether there is an oil-gland, as in Dromwus (p. 154) ;
or whether the general pattern of the body-coloration is continued on the podotheca,
and the integument covering the nasal fossa: features which obtain in Dromeus.
One conspicuous difference between the nestling and the adult Caswarius is the fact
that the head and neck are densely feathered in the former. ‘The casque is indicated
by a horny plate covering the roof of the skull.
DRoMAUS NOVA-HOLLANDI&. (Nestling.)
Apteria :—
A. spinale.—This is wanting.
A. mesogastrai.—tn torm and relations this agrees with Caswarius, but is relatively
much smaller.
A, trunci laterale. —As in Casuarius.
Pteryle :—
P. alaris.—There are 17 remiges in all. Of these, 10 are cubitals and 7 metacarpo-
digitals. These remiges in the adult are peculiar, lacking the calamus and
possessing an aftershaft. Re
The tectrices are arranged in obliquely transverse rows, as in Rhea (p. 156), but they
are not sharply separated one from another as in that genus. Furthermore, they are
not separable into major, median, minor, and marginal series as in Neognathe, They
extend outwards on to the manus. All the primaries apparently belong to the meta-
carpal series. Digit II. is free. That is to say, so much of this vestigial digit as
remains projects freely beyond the feathers, and bears a claw as in Archwopterya,
Opisthocomus, and the Gallinw. In these last the claw appears in the embryo.
There is no ala spuria; and rectrices are also wanting.
Rhamphotheca.—That of the upper jaw resembles that of Caswarius. The rhinotheca,
caudad, is sharply distinguished from a thin cere in which the nostrils, which are
impervious, are placed. The position of the nostrils is about the middle of the
beak. They are not protected by an operculum.
The sheath of the lower jaw is also composed, as in Caswarius, of three pieces—a
median and two lateral. The tomium bears slight traces of denticulations similar to
s
154 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
those described by myself in the Tinamous [81]!. The young of Casuarius bear traces
of similar denticles.
Podotheca.—The acropodium is made up of small reticulate plates which gradually
fuse to form ten large transverse scutes, covering its distal third. These are widest in the
middle of the series. The border region, between the acrotarsium and the acropodium,
is clothed with tiny granulate plates; these extend downwards so as to clothe the
proximal half of both outer and inner toes—the remaining region of these, like the
whole of the middle toe, being protected by large scutes. The inner claw is not
elongated as in Casuarius. ‘The planta is covered with small reticuiate plates.
It is interesting to note that the podotheca and the cere are mottled with black
pigment, so as to carry out the general scheme of coloration which characterizes the
feathers. This mottling is most distinctly marked ina ripe embryo (fig.2.B). Ina stage
earlier than this, and in the nestling, they are not nearly so distinct.
The Uropygium is distinctly traceable in this nestling: in the ripe embryo it is
much more easily demonstrable. Here it is represented by a small rounded oval mass
partly embedded in the skin and lying immediately above the cloacal aperture, as in
Apteryx.
Neither in this nestling nor in the embryos is there any trace of a separate, detachable,
ege-tooth, as is found in the Neognathe ; but the region of the beak in which this is
usually borne is, in the embryos, produced into a small conical elevation of a whitish
colour. Perhaps we should be correct in regarding it as a non-detachable egg-tooth
becoming absorbed after hatching. In the Meognathe it is detachable, and falls off
after this event.
RHEID4.
RHEA AMERICANA.
Apteria :—
A. spinale.—In one adult and a three-months old nestling this was divided into two
portions—more or less distinctly—an anterior and a posterior; thus recalling the form
of this tract in some Tinamous, e. g. Calodromas elegans. The anterior space was
cordiform in shape, with its base towards the head. Its greatest width was nearly
2 in., its greatest length about the same. A more or less perfectly feathered area
of about 4 in. in length divided this from the posterior apterion which is continued
to the tail. It was at first about 2 in. wide, narrowed to 14 in., contracting again to
1 in. for the rest of its course. In another adult and three-months old nestling the
anterior apterion was very small.
fn other adults, and seven young birds ranging from embryos to half-grown
specimens, the anterior space was wanting.
1 Figures in brackets refer to Bibliography at end of paper.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA, 155
Fig. 2.
Oe
x
A.—Left foot of a nestling Casuarius casuarius sclateri, to show the arrangement of the scutes of the podotheca.
B.—Left foot, anterior aspect, of Caswarius loric, for comparison with A, to show the large hexagonal
plates clothing the proximal portion of the tarso-metatarsus.
C.—Left foot of a newly hatched Dromeus nove-hollandice, to show the arrangement of the scutes of the
podotheca, and the curious continuation of the pattern of coloration of the body down to the toes.
D.—Left foot of a ripe embryo of Rhea americana, to show the arrangement of the scutes. Note the presence
of feathers (f) on the tarso-metatarsus—temporary in this species, permanent in R. darwint.
E.—Left foot of an embryo Apterya australis mantelli, to show the form of the scutes of the podotheca.
156 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
A. trunci laterale—This occupies the whole under surface of the wing, extending
backwards to a point corresponding to the level of the free end of the scapula. Its
width is considerable. It extends dorsad, above the scapula, about | in., and ventrad,
below it, about 4 in. The feathers over the front and upper end of the thigh are
small and sparse, suggesting the formation of an apterion, thus tending to form a
distinct pteryla femoralis as in the Neognathe.
A. mesogastrei.—tThis includes the sternal callosity, which is 3# in. long and
2 in. wide. Immediately caudad of this callosity the space narrows to 1} in., and
again to about 4 in., which width is sustained for the rest of its length till it ceases
about 3 in. in front of the cloacal aperture.
Pteryle :—
P. capitis—The region of the eye is surrounded by a bare space; so also is the
external aperture of the ear. The rim of the aural orifice is surrounded by feathers
which serve to close the aperture. The throat is bare along each side of the mandible.
There are well-developed eyelashes.
Pt. alavris—The wing is eutaxial and of great size. Remiges 28. Cubitals 16.
Metacarpo-digitals 12 = metacarpals 7; addigital 1; mid-digitals 2; pre-digitals 2.
The arrangement of the quills and coverts in the wing of the Rhea appears to be
unique, at least in so far as the arrangement of the cubital remiges and the coverts
are concerned. ‘The pollex only bears a claw.
The cubital remiges are spaced about as wide apart as in the wing of a Neognathine
of about the same relative size, ¢. g. Stork or Eagle. There is one point, however, in
which they differ considerably : rows 1 to 6 do not rest, as the others, on the ulna,
The Ist and 6th almost reach it, the 3rd and 4th are most distant from it. They
seem to have been carried away by the tension of the ala membrana posterior, stretcied
in the angle between the manus and forearm.
The cubital remiges are rather shorter than their major coverts: those of the hand
are longer. ‘This difference in length between the remiges and their major coverts is
probably an indication of degeneracy. It is interesting to note, in this connection, the
fact that in the young chick the first cubital flight-feathers are not remiges, but major
coverts.
The metacarpo-digital remiges are arranged much as in the Neognathe, but
somewhat farther apart, so that they tend to form pairs with their major coverts.
Like the cubital remiges, they are of great length, but differ therefrom in that they are
longer—by an inch instead of shorter by that amount—than their coverts. The Ist
metacarpal remex is distinctly smaller than the rest of the series: see p. 157.
The ventral aspect of the wing is bare.
The coverts of the dorsal aspect are arranged in obliquely transverse series, the
feathers of each series being piled as it were one upon another, from six to nine
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 157
such coverts being included in each of these bundles. ‘The bases of these coverts
form a series of partitions, dividing the post-axial border of the wing into a number of
wide and deep compartments.
Carpal covert and remex. (PI), XLV. fig. 12.)
Two different sets of feathers seem to claim identification with these; one or other
of which is invariably present in the Neognathw. The 1st metacarpal remex appears to
me to represent the carpal remex of the Neognathe. ‘This is smaller than the rest
of the metacarpal series and has a large major covert. In its position and its relations
to the 2nd remex, it closely resembles the carpal remex of many Neoguathe.
The second set of feathers, referred to as possibly representing the Meognathine carpal
covert and remex, lie immediately pre-axiad of the base of the major covert of the
1st metacarpal remex ; or, in others words, pre-axiad of the base of that feather which
we have just tried to show may represent the carpal covert (¢.c., Pl. XLV. fig. 12).
The identification of the covert must stand or fall by the quill.
The possible carpal remex, ¢.7’., as we have just remarked, lies immediately
pre-axiad of the supposed carpal covert (c.c., Pl. XLV. fig. 12). Furthermore, for the
purposes of identification, we may mention that this new remex, **, is seen to be
serial with the ¢. medie@ of the forearm on the one hand, and is the terminal feather
to a transverse row of six on the other. In other words, it and its covert represents
the Ist median and minor covert of the manus. In position, that is to say with regard
to its relation to the Ist metacarpal remex—which is also to be regarded as possibly
representing the true carpal remex, ¢.r. fig. 12,—it very closely agrees with the relation
between the carpal remex and the Ist metacarpal remex of Neognathe (er,
Pl. XLV. fig. 12).
It does not seem possible to settle this question definitely. It may be that these
two feathers now under discussion may, in the Neognathe, represent sometimes one,
sometimes the other of these two sets. This seems the more probable when we
remember the varying development and relations to the primary remiges which obtain
in wings of Neognathe.
If the first and most favoured interpretation of the identity of the carpal remex is
correct, then it would seem that it represents, in Neognathe, the Ist metacarpal, not the
Ist cubital remex, as was contended by Mitchell [62]. If the second interpretation is
correct, then the carpal remex of the Neognathe is represented in Rhea by a median
and minor covert of the metacarpus.
T. majores.—On the cubitus these are slightly longer than their remiges, about 1 in.
On the manus they are slightly shorter. It is significant to note that in the young of
many JVeognathe the cubitals are at first shorter than their coverts; the coverts
performing for a brief time the function of the quills, as in the case of young Galline.
T. medie.—The median coverts of the cubitus offer no features for discussion other
VOL. XV.—ParT V. No. 7.—December, 1900. 2A
158 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
than those included in the general remarks on the arrangement of the coverts. There
are only two median coverts in the hand, and, as already hinted, one of these may
represent the carpal remex.
T. minores—There are 6 to 7 rows of minor coverts on the cubitus. They are
wanting in the manus.
T. marginales.—A broad bare space separates these from the minor coverts. They
are arranged along the pre-axial border of the wing, in groups of 3, just as in some
Neognathe. They are wanting in the hand.
Ala spuria.—There are 5 thumb-quills, as in the Meognathe.
Parapteron and Hypopteron.—These humeral remiges are wanting in Fhea.
The feathers along the pre-axial border of the humerus are of great length.
The whole under surface of the wing is perfectly bare, a probably degenerate
character.
Pt. caude.—There are no rectrices.
Uropygium.—Absent in adult (p. 159.)
The Rhamphotheca, like that of Dromwus, is made up of three pieces, both in the
sheath of the upper and lower jaws. ‘There is no trace of the denticulations found in
Dromeus, either in the adult, nestling, or embryo. The aperture of the nostrils,
which are pierced through the cere, is lateral.
The Podotheca of Rhea americana (fig. 1 D), according to Dr. Gadow [25], resembles
that of A. darwini, and differs from that of R. macrorhyncha in that the whole of the
acrotarsium is clothed in large scutes, whilst in R. macrorhyncha these scutes are
confined to the distal extremity. The leg of this last thus resembles that of Casuarius
and Dromeus. FR. darwinii is furthermore peculiar in that the feathering of the
tibio-tarsus is continued downwards on to the tarso-metatarsus for some distance.
It is interesting and significant to note, however, that a few feathers occur in this
region in the embryo of R. americana. Indeed, in this latter their extension may be
traced downwards to within a short distance of the toes, as will be seen in fig. 2, p. 155.
The planta is also invested by large scutes. In the 3-grown nestling and adult
these are arranged in the form of a series of paired plates. In the nestling and ripe
embryo they form a series of single transverse scutes. Along the inner side of these
large scutes will be found a number of smaller, wedged in between the larger. These
apparently gradually increase in size, and thus give rise to the secondary arrangement
of paired plates.
The claws of the toes are laterally compressed, and form a strong median dorsal
ridge or keel, not met with in other Palewognathe. This is most marked in the
outer and inner toes,
PHYLOGENY OF THE PALZOGNATHZ AND NEOGNATH 2. 159
b. Pterylosis of the Embryo and Nestling.
The ripe embryo and the nestling Rhea americana appear to differ from the adult
only in that the anterior portion of the apterion spinale is absent, and that there is a
distinct oil-gland.
I find no trace of an “ egg-tooth” in the ripe embryo, which I myself removed from
the egg. ‘This is also wanting in the embryo Dromeus and the nestling Caswarius.
Concerning Dromeus, however, see p. 154. The late Prof. Jeffery Parker failed to
find it in Apteryx at any stage of its growth.
STRUTHIONIDA.
STRUTHIO CAMELUS.
1 have not been fortunate enough to secure either nestling or adults of Struthio ;
consequently I am obliged to fall back upon the published accounts of others. Nearly
Fig. 3.
A.—Lateral view of an embryo of Struthio, after Lindsay, to compare with Apteryx. Note the absence of a
lateral cervical apterion.
B.—Lateral view of an embryo of Apterya australis, right side (after Parker), to show the pterylosis.. Note
the distinct pteryla colli ventralis and dorsalis and the pteryla humeralis.
all that can be gathered, moreover, concerning the pterylosis of the trunk of Sérut,io
concerns embryos only. The wing of the adult Ostrich has been ably described by
2a2
160 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Wray [97]; the figure illustrating his paper has been reproduced here by the kind
permission of the Zoological Society. The pterylosis of the trunk of the embryo has
been figured by Lindsay [48], and is reproduced here (fig. 34).
Apteria :—
Apt. spinale.—Very narrow, longer relatively than in Caswartus, which it resembles.
Apt. mesogastrei.—According to Hildebrand [37] this extends from the beak to the
cloaca, but judging from Miss Lindsay’s [48] figures it did not extend beyond the
sternal callosity, as in other flightless Palwognathe.
Apt. trunci laterale——This would appear in size and form to closely resemble that
of Apteryx, and to differ from all the remaining Palwognathe.
Hildebrand also describes an apterion on the crown of the head.
Pteryle :—
Pt. alaris (Pl. XLV. fig. 11).—According to Wray [97] there are 16 metacarpo-
digitals, distributed as follows:—metacarpals 8; addigital 1; mid-digitals 4; pre-
digitals 3. ;
According to Wray’s figure, reproduced on Pl. XLV. fig. 11, the primaries are
placed wide apart and almost at right angles to the long axis of the hand. The
number of the metacarpals is higher than in any other bird—8. In khea and some
Neognathe, i.e. Flamingo, there are 7 metacarpals. In no other bird, save Struthio,
are there 4 mid-digitals and 3 pre-digitals. The arrangement of the coverts appears
to differ materially from that of Rhea. The ala spuria is made up of 4 remiges.
I await a spirit-specimen before attempting the search for the carpal remex and
its covert.
It would be interesting to know whether the flexor carpi ulnaris resembled that of
Rhea in the large fleshy portion for the attachment of the remiges (fig. 5 a, p. 240)
and what is the condition of the vinculum tendinee. Assuming that the disposition of
these parts in Rhea represents a proto-carinate condition, we should expect to find the
wing of Struthio very similar in these particulars.
Both pollex and index are armed with a claw.
Lihaiwphotheca.—Compound as in Rhea
Podotheca.—Acropodium and lower portion of acrotarsium with broad transverse
scutes. Only 3rd and 4th toes are present; of these in the adult only the 3rd bears
a claw.
Uropygium wanting.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH®. 161
APTERYGIDA.
APTERYX AUSTRALIS MANTELLI.
Apteria:—
A. spinale.—This is wanting.
A, mesogastrei.—This is a well-marked space, extending forwards from the cloacal
aperture, along the middle line, as far as the base of the neck, to a point corresponding
to the interclavicular region of the Meognathe. Its greatest width (1 in.) is on
the abdomen; in the middle of the sternum it is nearly 4 in.
A. trunci laterali is of very considerable extent. It runs from a point immediately
over the head of the femur downwards and forwards as a conspicuously broad bare
space of about 1 in. in width; terminating at the base of the neck. At the wing it
gives off a branch downwards and backwards between the thigh and the pteryla
ventralis as usual, and terminates about 24 in. from the cloacal aperture.
Pteryle :—
Pt. capitis.—This is very dense, and bears numerous rictal bristles of great length.
Pt. spinalis—Forms a broad saddle-shaped tract blending posteriorly with the
femoral tract, and anteriorly passing into the pt. colli.
Pt. ventralis is well defined. Anteriorly it blends with an incipient pt. colli ventralis,
and is continued backwards as a sharply defined tract bounded on the one side by the
apterion trunci laterale and the other by the apt. mesogastrai.
The pt. colli is almost divided into a dorsal and ventral tract by the forward extension
of the apt. trunci laterale.
Pt. alaris (Pl. XLV. fig. 13).—Remiges 13. Cubitals 9. Metacarpo-digitals 4=
metacarpals 1; addigital 1; mid-digitals 2.
The innermost cubital is very weak, and is counted a remex solely on account of its
position. Yet I think that most would agree with me in this decision. The 3rd
metacarpal remex (?mid-digital) is cramped in position, and forced dorsalwards almost
into the position of a major covert. The calamus of these quills is very long in pro-
portion to the length of the whole feather, and is fusiform in shape. The form of
these quills is exactly reproduced, as will be seen in the figure (Pl. XLV. fig. 15), in
the first quills of the nestling Cassowary (see p. 165).
The proximal end of the forearm, over the region of the elbow-joint, is bare, as in
many Neognathe. ;
The arrangement of the coverts is that of obliquely-transverse rows, as in JVeo-
gnathe. Only tectrices majores, however, can be certainly made out. There appears
to be but one major covert on the manus.
The index-finger bears a large claw. ‘There is no pollex.
Pt. femoralis.—Unitormly feathered, passing forwards into a sharp, cone-shaped
point.
162 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The Uropygium is a large, compressed, naked, conical gland lying at the bottom of a
conspicuous depression—formed by the great development of the gluteal muscles,—
and immediately above the cloacal aperture. A figure of the uropygium will be found
in Mr. Beddard’s recent paper [8].
Rhamphotheca.—Apparently made up of the same elements as in the other Palwo-
gnathe ; but this is masked by the greatly elongated form of the beak. ‘There is a
distinct cere at its base, but this is not pierced, as in Rhea, Dromeus, and Struthio, by
the nostrils. These, in Caswarius, open near the top of the beak, and in Apteryx at
its extreme tip, where they will be found as a pair of minute slit-like apertures.
Podotheca (fig. 2 £).—The acrotarsium may be either made up of large rounded
scales, or of large scutes as in Rhea americana. ‘he latter form occurs, so far as I
have been able to make out, only in A. australis mantelli. ‘The planta in this species
is covered with large coarse papille. In an embryo of A. australis mantelli the fusion
of small plates into scutes in the acrotarsium is well shown (fig. 7).
CRYPTURI.
The pterylography of this group of the Palwognathe has been described in con-
siderable detail by myself [81]. It will suffice here to say that the apterta are more
marked than in the flightless forms. In the form of the apt. spinale, Calodromas
elegans, one of the Crypturi, closely resembles Rhea. Unlike the flightless members of
the group, filo-plumes occur—though very sparingly; and down-feathers. These last,
however, are restricted to the pteryla alaris.
The neossoptiles, in the large size of the aftershaft, differ from those of the flightless
forms ; but in the remarkable development of the radii and in the elongated flattened
shape of the rami, the Crypturi closely resemble the Struthiones (Pl. XLV. fig. 2).
The denticulated tomium of the nestling Calodromas recalls that of Dromeus.
Tue STRUCTURE OF THE FEATHERS.
The Neossoptiles.
The neossoptiles of the Palewognathe consist of prepenne only, but these present
many features of great interest.
The prepenne of Caswarius have a moderately long rhachis, bearing some 3—4 pairs
of rami. The radii are unsegmented, of relatively medium length, and bear minute
fila, sparsely distributed (Pl. XLV. fig. 5). The aftershaft is apparently represented
by about 6 rami standing at the base of the main shaft, and not seated along a separate
rhachis, as in the Fowl or Tinamou.
The prepenne of Dromeus (Pl. XLV. figs. 4, 4a) appear to be distinguishable from
those of Casuarius chiefly, and probably only, on account of their rather greater length ;
PHYLOGENY OF THE PALHZOGNATHA AND NEOGNATHZ. 163
the rhachis bears about 6 or 7 pairs of rami. The radii are slightly stouter, relatively.
As in Casuarius, fila are minute and rare.
The nestling-down of Apteryx (Pl. XLV. fig. 6) is easily distinguishable from that
of the remaining Palwognathe. Like that of Dromeus and Casuarius, it has a
well-developed shaft, but it differs from these in its greater length and in the very
numerous rami, From Rhea and Struthio it differs entirely, in that, in these, the
neossoptiles are umbelliform.
The most striking feature perhaps of the nestling-plumage of Apteryx is its great
length. In texture we appear to get three gradations. A. australis mantelli and
australis very closely resemble one another and differ from the remaining species in
that the feathers are coarser and slightly harsh to the touch. A. oweni and A. haasti
stand at the other extreme and are peculiarly soft. Some individuals of A. australis,
however, seem to stand between the tivo.
A. haasti is the most markedly distinct of all. The feathers clothing the head, neck,
and thighs are of a peculiarly downy character, recalling, in texture, the down-feathers
on the breast of the adult Duck. On the back and breast they are of a peculiar
“woolly” texture, resembling very strongly the intermediate plumage or pseudo-
definitive feathers, (mesophyles of Palmer), which follow the prepenne in the Owls.
There is no trace of an aftershaft to these prepenne.
The rami are closely set, very numerous (18 to 20 pairs), and moderately long. The
radii are relatively short, and have the appearance of being formed of a number of
short, laterally compressed joints, the dorsal and ventral angles of the anterior ends of
which are produced forwards to form a pair of minute fila.
In the length of the shaft and the great number of the rami the prepenna of Apteryx
stands alone amongst the Palwognathe ; moreover it is further peculiar in that it is not
produced beyond the most distal radii into long filaments.
A peculiarly interesting feature of the prepenne, and of the first definitive feathers,
is that the latter do no¢ thrust out the former, as they appear to do in all other known
birds. Exactly how this is avoided is a point for further investigation. I hope to be
able to explain this mystery shortly!. That this process of thrusting-out is avoided
can be still seen in a downy nestling of A. haastii in the Rothschild Collection. Here
the first definitive feathers are about halfway through the skin, and show no trace
whatever of a prepenna surmounting the tips, as do the other Palwo- and Neognathe.
Furthermore, they are peculiar in that they are not invested by the sheath common to
all developing feathers. This appears to break away as soon as it reaches the surface
of the skin.
? Tt may possibly turn out that this nestling-down plumage may represent preplumee instead of prepenns ;
this would account for their non-ejectment by the teleoptiles. The fact that plumule do not o.-2r in the
adult is of no importance. Zander [101] has described “ pinsel-artig” down-feathers in the embryo Ithea, but
this 1s probably an error.
164 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The structure of the adult contour-feather does not differ much from that of the
prepenna. ‘The chief distinction—apart from size—whereby the two differ appears to
be in this:—that in the adult the rami are arranged in pairs, and not alternately up
the shaft; that the radii are confined solely to the rami, and do not grow along the
shaft in the inter-ramal spaces ; and, thirdly, the rami are not so distinctly divisible into
nodes and internodes; but there is not much difference in the form and length of
the fila.
Casuarius, Dromeus, and Apteryx all agree in that the prepenna has an elongated
main shaft, and no distinct aftershaft.
Rhea and Struthio agree one with another, and differ from the above-mentioned, in
that the prepenna is umbelliform (Pl. XLV. figs. 38, 3a). The main shaft appears to
be represented by 3 rami conspicuously stronger than the rest with which they are
associated. The prepenne are borne upon the tips of the definitive feathers for a very
considerable time, giving a quite peculiar appearance to the first plumage.
In Rhea the bases of all the rami, 9 in number, are seated around the upper
umbilicus. The main axis of the feather—the rhachis—as just remarked, is represented
by 5 elongated rami, bearing radii from the base upwards for a short distance, and
produced beyond these into a long slender filament. The aftershaft is represented by
some half-dozen rami, furnished with numerous delicate radii. These last are
moderately long, strap-shaped filaments, with scarcely the faintest trace of fila.
In Struthio the typical neossoptile (Pl. XLV. figs. 7, 7a) has the three rami repre-
senting the main shaft produced far outward beyond the tip of the feather; the
projecting region of each taking the form of a hardened, glistening, ribbon-shaped
lamina, hollowed in its ventral surface. In this particular it closely agrees with the
Crypturi. The radii are short, and run along the rami as a series of relatively broad,
flattened, and pigmented lamine, set obliquely on to the ramus, and overlapping one
another much as in the normal adult MNeognathe contour-feather. Moreover, each
radius bears numerous but small, probably degenerate, fila and hooklets. The forma-
tion of the remarkable “ eye-spots” seen in these radii of many Meognathe are worthy
of notice and further examination.
The aftershaft rivals the main shaft in size, numbering some 12-15 rami, which do
not, however, differ from those of the main shaft, save only in that the rami are not
produced into elongated horny bands, whilst the radii are larger than those of the three
rami representing the main shaft.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH. 165
The Structure and Homologies of the Adult and Nestling
Remiges of Casuarius.
The Nestling.—The youngest nestling in which these can be studied is that of a
stuffed C. casuarius salvadorii (Rothschild Mus.).
In the dried wing of this there are 6 remiges. These are widely spaced. and
seated in a distinct posterior wing-membrane as in Neognathe. ach consists of a
moderately long calamus and a long and tapering rhachis bearing from 4-5 pairs of
symmetrically disposed rami set very far apart. There is but the feeblest trace of the
fusion between the prepenna and the displacing definitive feather now under discussion.
In C. casuarius sclateri the growth ot the wing-spines has proceeded a stage farther.
The definitive feather is seen to bear rami as well as prepenne.
In Casuarius lorie the quill-spines have grown very long and have lost the prepenna.
The 5th or innermost spine, representing the 3rd cubital remex, yet bears a few rami.
In this stage, and in the last described (C. c. selateri), we get an inkling into the
developmental history of these remarkable wing-spines.
This is not the place for a recapitulation of the developmental history of a feather,
yet it would be well to recall one or two facts concerned with this for the better under-
standing of what is to follow.
The axis of a feather, then, is divisible into two parts—the calamus or quill, and the
rhachis or shaft. The latter appears to be a continuation forwards of the dorsal region
of the tubular calamus. This being so, it is obvious that this drawn-out portion must
have two lateral free edges. ‘These in the normal feather turn inwards towards the
middle line and finally meet, leaving a very fine seam to indicate the line of junction.
But the calamus, it will be remembered, is a thin, dense, horny, and semitransparent
structure and perfectly hollow, whilst the rhachis in section is solid, being composed of
an outer layer continuous with that of the calamus, and an inner, medullary portion or
pith—a white, soft, cellular structure resembling elder-pith. This pith seems to be
added for the sake of supporting the outer horny membrane, thus insuring lightness,
toughness, and sirength. ‘The whole forms the shaft.
The dorsal elongated region of the shaft, now swollen out by pith, forms a sort of
plug to the otherwise open mouth of the calamus. The ventral edge of this mouth is
always visible. It forms the boundary of the upper umbilicus.
The aftershaft, so conspicuous a feature in the feathers of the Cassowaries, represents
an elongation of the ventral region of the calamus exactly similar to that which obtains
on the dorsal. Thus we have a dorsal and ventral rhachis.
In the remiges of C. lorie and C. ¢. sclateri we have only the dorsal rhachis, and
this is of great interest as only its outer layer is present, there being no pith-cells.
This layer takes the form of a delicate scroll-like, ribbon-shaped lamina whose free
edges turn inwards, but from the absence of the packing material, or pith, they fail to
meet in the middle line (Pl. XLV. fig. 14 a).
VoL. xv.—Part v. No. 8.—December, 1900. 2B
166 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The fate of the prepenna which remains attached to the young definitive feather can
be conjectured with probable certainty. It is shed or rubbed off as in Carinate, The
definitive rhachis with its rami appears to be carried for a short while longer, and then
to break off from the calamus by which it was supported. The fracture takes place at
the upper umbilicus, and leaves only the calamus. This appears to go on growing and
thickening its walls, so that in course of a short time it becomes the solid, elastic spine
so familiar in, and peculiar to, the Cassowaries.
In the wing of the adult Apteryx we have an almost similar series of wing-spines.
These differ, apparently, from those of Caswarius only in that they have retained the
rhachis and its rami, and in that the calamus is yet hollow. In one point, however,
they differ markedly, inasmuch as they have what is possibly a vestigial aftershatft.
This can be readily seen as an elongation of the central lip of the calamus, the sides
of which bear numerous rami.
The Teleoptiles.
In the Palwognathe there is but one form of teleoptile, corresponding to the
contour-feathers of the Neognathe, semiplumule, plumule, filoplumule, and powder-
down are all absent. Rictal bristles, such as occur in Apterya, and eye-lashes, such
as are found in all other Palwognathw save Apterya, are modifications of contour-
feathers.
Meijere claims to have discovered filoplume in Struthio, Rhea, and Casuarius, and
gives figures of each. I cannot, however, confirm this. I have found degenerate
feathers in the wing of Caswarius similar to that he figures from the neck of
C. papuanus : these, however, I regard as degenerate contour-feathers. I failed to find
anything in Rhea corresponding to his figure; Struthio I have not had an opportunity
of examining. He failed to find them in other Palwognathe.
In Casuarius the teleoptiles are, on the lower region of the back, of great length,
and relatively larger than in Dromeus. Only the rami of the lower part of the feather,
from wherever taken, bear radii, and these appear to be entirely destitute of fila. The
rami throughout the greater part of the length of the feathers are long and hair-like,
reminding one of egret-plumes.
The hyporhachis is nearly or quite as long as the main shaft, and does not differ
therefrom structurally, as in Neognathe.
The Casuariide are remarkable for the enormous spines, 5 in number, which are
borne by the wing. These are degenerate remiges, and are dealt with above.
The plumage of the Dromeide differs conspicuously from that of the Casuariide by
its greater softness of texture. In the latter it is harsh and coarse.
The rami are closely set, as in typical contour-feathers of Neognathe. In a feather
taken from the middle of the pteryla spinalis the radii are of great length, and give
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 167
the surface of the feather a very soft, silky appearance ; towards the tip they decrease
in length, and finally disappear. The distal end of the feather—about one-sixth of its
whole length—is peculiar, in that the rami bear no radii, but fuse more or less com-
pletely into one common laminate mass. Separate rami may, however, be distinguished
here and there under the microscope. The radii are very long, filamentous, and bear
only minute pointed fila, for the most part unpaired.
On the neck the feathers are of a looser structure, the radii being shorter, and
disappearing much sooner than in the back-feathers.
The contour-feathers of the Apterygide appear at first sight to differ but little from
those of many Neognathe, the vane of the feathers, seemingly, being continuous. The
fact that it is discontinuous is only proved after microscopic examination.
The rami are paired, set close together, and moderately long. The radii are long,
filiform, more or less distinctly jointed, and armed with numerous minute fila, set in
pairs, following close one upon another. These run in this fashion through almost the
entire length of each radius, only the extreme proximal end lacking such appendages.
According to Parker [71] they are longer in the embryo than in the adult. The entire
absence of hooklets at once distinguishes this from a laminate feather.
There is no aftershaft. (See also p. 163.)
In the Struthionide the adult contour-feather more nearly resembles that of the
Neognathe than is the case with any other of the Palwognathe.
The rami are set closely together, precisely as in the typical Carinate feather. The
radii take the form of narrow blade-shaped bands, directed upwards and outwards, so
that the rami from which they spring form the bottoms of V-shaped troughs. The
distal end of each radius bears fila disposed as in normal Meognathe, but none are
sufficiently elongated to form hooklets: they are probably, however, vestiges of these.
Moreover, the radii differ from those of more perfect feathers in that the distal and
proximal radii do not differ one from another: both series are alike, long and blade-
shaped. Thus, the vane is discontinuous—a probably degenerate condition.
In the Rheide the contour-feathers of the trunk are characterized by the markedly
discontinuous character of the vane; the rami appearing as a number of hard lines,
between which radii are only faintly traceable.
The remiges are of very considerable size, though far inferior to those of the Ostrich.
The rami bear numerous short and fairly conspicuous radii.
The radii of the track-feathers, under the microscope, appear to have degenerated
rather less than have those of the remiges. They approach those of the Ostrich. At
its proximal end, each radius is laterally compressed, the resultant lamina bearing some
4 or 5 long fila, probably vestiges of hooklets, and of the processes which occur on the
ventral edge of the lamina in the proximal radii of Neognathine feathers.
The radii of the remiges are filiform, but distinctly flattened: from side to side. ‘The
fila suggest vestiges of hooklets; they are frequently bifid.
2B2
168 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
In the Dinornithide the rami and radii are short. The latter, from the middle of
the feather to its base, are filiform and divided into a series of numerous short joints,
the anterior ends of which are produced into two or three pointed fila. Thus these
radii most nearly approach those of Apteryx, but they may be at once distinguished
therefrom by the relatively great length of the fila.
The distal ends of the rami bear no radii.
The feathers of Dinornis were first described by Dallas [15]. His description is
appended below :—
“The structure of the web is somewhat different from that which occurs in the Emu
and the Cassowary. ‘Towards the base of the shaft the barbs spring in groups of four
or five together from nearly the same spot, and thus this part of the web assumes a
tufted aspect. As we advance towards the apex this arrangement speedily ceases; the
number of barbs springing from the shaft gradually diminishes until each side bears
only a single series of these appendages. ‘The barbs consist of slender, flattened fibres,
bearing long, silky, and very delicate barbules, without any trace of barbicels, and
presenting a distinctly beaded appearance when examined by a simple lens. Under
the microscope..... the barbule appears merely divided by faint transverse partitions
into a series of cells, some of which towards the apex exhibit small tooth-like projections
representing rudiments of barbicels..... The barbs nearest the base of the feather,
both in the main web and accessory plume, are destitute of barbules for some distance
from their base; but this distance gradually decreases until the barb is furnished with
barbules throughout its whole length.”
As Dallas remarks, these fragments still leave us in ignorance of many points,
such as whether the basal rami (barbs) possessed the hair-like tips of the Emu and
Cassowary, and whether the apical portion of the feather supported simple rami as
in these last.
SUMMARY OF PTERYLOLOGICAL CHARACTERS.
Perhaps the most interesting pterylological result of the present enquiry has been
the light thrown upon the history of the remarkable wing-spines or vestigial remiges
of Casuarius. ‘The comparison of these with the remiges of the nestling on the one
hand, and with the wing of Apteryx on the other, leaves little or no doubt but that
the explanation, or rather interpretation, of the nature of these degenerate feathers is
correct.
The form and disposition of the apteria have been described at greater length than
ever before, and some new facts added. The existence of apteria has long been known—
at least in Struthio and Apteryx; though these facts had not yet found their way into
the text-books, which still repeat the old error that the feathers of the Palwognathe
were evenly distributed over the body.
The form and structure of the nestling-down has been worked out in considerable
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH&. 169
detail. Prepenne only are represented. In the Crypturi ouly is there a distinct after-
shaft, and this is of great size, equal, indeed, to the main shaft. In these particulars,
and in the complexity of the main shaft, it is unique. In Casuartus, Dromeus, and
Apteryx, these havea well-developed rhachis. An aftershaft appears to be represented
only in the two former, and here it consists only of a few sessile rami without any trace
of arhachis. The remiges of the adult Apteryx show how the loss of this may have
come about (Pl. XLV. fig. 13). In Rhea and Struthio the prepenna is umbelliform, and
thereby differs from that of the other Palwognathe: the aftershaft is represented, as in
Casuarius, by a few sessile rami. In Struthio, however, the number of these is very
great. In both Rhea and Struthio the main shaft is represented by 3 thickened rami,
which in the latter are produced forwards beyond the rest of the feather to form broad,
hollow, ribbon-shaped lamina, recalling in form the nestling-down of the Crypturi
(Pl. XLV. fig. 2).
The following point involves a mystery which I am anxiously endeavouring to solve.
The prepenne are regarded by some as nothing more than portions of the distal
extremities of the developing rami of the teleoptiles below. If this is so, how comes
it that the prepenne of Caswarius and Dromeus have ascarcely recognizable aftershaft,
whilst in the teleoptile it is of such great length as to be hardly distinguishable from
the main shaft? In the Tinamous these relations are exactly reversed. The aftershaft
in the adult feather is very small or wanting, and in the nestling it is as long as the
main shaft!
A further most serious objection to the probability of the truth of this view is the
fact that in Apteryx the nestling-down feathers are not driven out by the teleoptiles.
These arise at the side of the prepennew, the ultimate fate of which my series of
nestlings is not large enough to show. It is probable that they are shed as soon as the
definitive feathers have completed their growth. The peculiar downy nature of these
feathers does not seem to have been recorded before.
_ The discovery of a uropygium in Dromeus and Rhea is a point of some interest.
The podotheca appears to be of some slight value for systematic purposes. Dr. Gadow
[25] long since pointed out the differences between the three species of Rhea. I have
not succeeded in finding any appreciable difference between the different species of
Cassowary, or in distinguishing that of Caswarius from Dromeus. Casuarius lorie
seems to differ from the other species in this respect and to form a type of its own
(fig. 23, p. 155), just as Apterya australis mantelli seems to differ from the other
Apteryges in having, as a rule, the acrotarsium clothed with transverse scutes in place
of small rounded plates.
Dromeus represents the less specialized of all the Palwognathe. Casuarius
undoubtedly comes next ; though in the brilliant coloration of its head and ueck, its
remarkable casque, spine-like remiges, and elongated claw on its inner toe it has made
a distinct advance upon Dromaus.
170 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Apteryx should be placed next. It has lost the aftershaft to the feathers; the
prepenna has a well-developed rhachis. In the large size of the apteria and in the
possession of an oil-gland it resembles the Carinatw; so also it does in the softness of
the neossoptiles.
Rhea and Struthio are undoubtedly the most highly specialized of the group. One
of the chief pieces of evidence of this, from the point of view of pterylological
characters, is the fact that the prepenne has neither aftershaft nor rhachis. The fact
that the wing is larger in these two than in the remainder of the group, only means
that the need for this organ remained longer, and is not to be regarded as an advance
upon that of Casuarius or Apteryx, as the case may be. In them the decline of the
wing set in earlier.
The structure of the teleoptiles in these last two is more complex than in the other
groups: hooklets are feebly represented. These have lingered longer, just as the power
of flight.
We may briefly sum up the salient features of the pterylosis of the Palwognathe
as follows :—
All the flightless Palwognathe agree, and differ from the Meognathe, in having
the under surface of the wing naked !, and in that all the feathers have discontinuous
vanes.
A. ADULTS.
Dromeus, Casuarius, and Apteryx have no ala spuria. Lacking a pollex.
The teleoptiles of Dromeus and Casuarius have a large aftershaft. There is no oil-gland.
Dromeus has numerous though greatly degenerate remiges—17 in all.
Casuarius has few remiges—6 in all. These are degenerate and peculiarly modified
to form elongated porcupine-like quills, representing only the calamus of
ordinary feathers.
The teleoptiles of Apteryx have no aftershaft and a large oil-gland; it has few remiges—
13 in all—and these have a long calamus. Apteria are larger. Rictal bristles are of
great length.
Rhea and Struthio have a large ala spuria.
The teleoptiles of Rhea and Struthio have no aftershaft: there is no oil-gland. Remiges are
numerous and large.
The remiges of Rhea number 28; metacarpo-digitals 12. There are no rectrices.
The remiges of Struthio number 32 ; metacarpo-digitals 16. There are large rectrices.
Apteryx and the Crypturi are the only Paleognathine lacking a sternal callosity.
1 The naked under surface of the wing of the flightless Struthious birds is probably a secondary feature due
to the suppression of the feathers by the long continued application of the wing to the side of the body.
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATEH.E. viel
B. NeEstiines.
The neossoptiles consist of prepenne only.
In Dromeus, Casuarius, and Apteryx the prepenna has a well-marked rhachis and numerous
rami, The aftershaft is feeble or wanting.
In Dromeus the rhachis bears about 8 pairs of rami: the rami are produced into
points beyond the distal radii.
In Casuarius the rhachis does not bear more than 5 pairs of radii.
In Apteryx the rami exceed 8 pairs; the rami are not produced beyond the radii ;
the prepennz of some parts recall the adult down of Ducks.
In Rhea and Struthio the prepenna is umbelliform ; the main.shaft is represented by 3 rami,
the aftershaft by numerous rami.
In Rhea the rami of the main shaft are not greatly elongated or flattened.
In Struthio the distal ends of the rami of the main shaft are much elongated and
peculiarly flattened and grooved, as in Tinamide.
In Crypturi the prepenna is very complex ; the aftershaft equals the main shaft in length.
OSTEOLOGY.
THE SKULL OF THE ADULT.
The skull of the Palwognathe—which, as we have elsewhere shown, includes the
so-called Ratite and Tinamous—differs from all other known birds in that, in the adult,
the pterygo-palatine connection is by symphysis or anchylosis, and not by an arthrosis.
The skull of the Tinamous, as will be shown presently, closely resembles that of
Rhea.
The Occipital Region (Pl. XLII. fig. 7).—The occipital condyle is sessile, and the
plane of the occipital foramen is inclined backwards, in all the Palewognathe save
Dinornis, Aipyornis, and Apteryx, and in the Tinamous. In Dinornis the plane of the
occipital foramen is nearly vertical; in Apterya the occipital condyle is pedunculated.
In Dinornis and Atpyornis this condyle projects beyond the dorsal margin of the
foramen. In Dromeus and Apteryx the supra-occipital, immediately above the foramen
magnum, presents the concavo-convex “cerebellar dome” so characteristic of the
Tubinares and Spheniscit. There is a well-marked supra-foraminal ridge (Pl. XLII.
fig. 8) running laterad on either side to become lost on the paroccipital process in all
save Apteryx and the Tinamous. In Apteryx the ridge ends abruptly in the form of
two pendent tubercles on either side of the foramen at about halfway down. In the
Tinamous the ridge terminates in the same region, but is without tubercles. The
occipital crest is most strongly marked in Rhea. The lambdoidal ridge runs outwards
and downwards as a sharply-defined ridge, terminating in a pair of strongly compressed
paroccipital processes, in all save Dinornisand Struthio. In these the ridge has a more
or less sinuous outline, and is not so easily followed. Only in Dinornis does the
lambdoidal ridge delaminate, as it were, in the region above the occipital crest, to
form anterior and posterior lambdoidal ridges.
172 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The paroccipital processes pass gently forwards at their base into the zygomatic
process, in all save in the Tinamous and Apteryx, Rhea, and Dinornis. In Apteryx and
the Tinamous they are separated from the zygomatic process by a notch more or less
deep. In Dinornis they pass forward into the squamosal prominence, and there lose
themselves. In Rhea only, externally, they are continued upwards as a strong ridge to
join the posterior boundary of the temporal fossa, and, internally, sweep round towards
the middle line in the form of a thin, curved, laminated plate of bone to form the inner
boundary of the tympanic recess.
The Roof of the Cranium (Pl. XLII. figs. 1-4).—The roof of the cranium is
produced laterally, in front of the temporal fossa, into a pair of pendent post-orbital
processes, in all save Apteryx and the Crypturi. In the former, post-orbital processes
are entirely absent, and in the latter they are reduced to a mere prickle. The anterior
region of this roof, in all save Dinornis, 4pyornis, Apteryx, and Crypturi, is charac-
terized by a pair of very long outstanding and backwardly-directed spines, being the
elongated supra-orbital process of the lachrymal. In Séruthio these are continued
backwards, by means of fused supra-orbital ossicles, to join the frontals ; thus enclosing a
space between the ossicles and the combined frontal and nasals (fig. 8).
In Dinornis the lachrymal passes backwards insensibly into the frontal to form a
broad overhanging ledge to the orbit. In the Crypturi it projects conspicuously on
either side of the skull, but is without the backwardly-projecting spines. In Apteryr
the lachrymal has become reduced to a mere vestige fused with the alinasal.
In all save Apteryx and the Tinamous the frontal is produced laterally to form an
overhanging pent-house to the orbit. In Apteryx, the frontal passes downwards and
inwards in the form of a smooth convex infolding of the bone, leaving the orbit without
any sharply defined boundary ; and in the Tinamous this region is deeply hollowed out,
almost to the middle line, so that the interorbital region of the skull is very narrow.
Young skulls of Rhea and Dromeus resemble the Tinamous in this respect.
There is a conspicuous cerebral dome in pyornis and Dromeus ater. In profile,
in Dromeus ater, the skull slopes rapidly downwards and forwards in front of this dome;
in Apyornis the gradient is much less marked.
The inter-orbital region in Dromeus is very wide, in Rhea very narrow: thus, taken
in connection with the development of the supra-orbital processes of the lachrymals,
this forms a useful character for systematic purposes.
In the Tinamide the frontals bear shallow supra-orbital grooves.
The Base of the Skull (Pl. XLII. figs. 5-8).—Casuarius, Dromeus, Struthio, and the
Tinamide all agree, more or less closely, in the form of the basitemporal platform. In
all, viewed below, it takes the form of a flattened, slightly tumid area, roughly penta-
gonal in shape, the base being caudad. Its postero-lateral angles are not produced into
mammillary processes, nor are there, save the Eustachian apertures, any sharply defined
boundaries forwards. Struthio and Dromeus somewhat closely resemble one another
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH. 173
in the form of this platform. In Dromeus ater it is very sharply defined, presenting
a steep face to the pre-condylar fossa ; in the other species of Dromeus and in Struthio
this face slopes gently upwards and backwards. Thus, in all save D. ater, the pre-
condylar fossa is shallow and moderately wide.
The pre-temporal platform is very sharply defined in Anteryx, Dinornithide, and Rhea.
In Apteryx and Dinornithine amongst the Moas, it is roughly triangular in form,
its postero-lateral angles are produced downwards into moderately or very large
mammillary processes. These in some of the Dinornithine coalesce in the middle
line, and present a deep vertical face when the skull is seen from behind. In the
Emeine the platform takes a laterally expanded pentagonal form, sharply defined along
its antero-lateral angles by a raised ridge helping to form the Eustachian groove.
The anterior basicranial fontanelle is marked by a deep groove. In Apyornis the
basitemporal platform is much shortened antero-posteriorly. The mammillary tubero-
sities run in the form of a low rough ridge along the whole of each postero-lateral side
of the pentagon.
In Rhea the platform has the shape of an elongated pentagon presenting steep
faces on all sides, though anteriorly it may slope gradually into the rostrum. The
Eustachian apertures lie immediately below the bases of the basipterygoid processes.
The Eustachian grooves are short, widely separated one from the other in the middle
line, and, in all except Dinornis and Casuarius, arched over by bone. In these last the
groove remains permanently open. In Rhea, Struthio, and Dromeus the mouth of the
aperture of each is conspicuous, lying on either side of the base of the basipterygoid
processes. In Apterys they lie nearer the middle line and are inconspicuous. In the
Tinamous these apertures are very wide apart, opening immediately behind the basi-
pterygoid processes.
The paroccipital notch is more or less wide in all save Apteryax, where it is narrow
and deep. As in #hea, it is more or less completely bounded on its outer side by a
much compressed, forwardly directed lamina of bone from the paroccipital process
(p. 172).
Within this notch, in Struthio, Rhea, Dromeus, Casuarius, Apteryx, and the Tinamous,
occurs the vagus foramen for the ninth and tenth nerves, the condyloid foramina for
the exit of the twelfth nerve lie behind it, between it and the occipital condyle.
In the Dinornithine the vagus foramen occurs behind the notch and much nearer to
the condyloid foramina, forming, with them, a little cluster of apertures, that of the vagus
being much the longest of the group. The carotid foramen in the Dinornithide lies
immediately in front of the notch, in all the other forms it occurs some distance further
forwards,
Traces of both anterior and posterior basicranial fontanelles occur in many of these
Paleognathine skulls, sometimes both are indicated in the same skull. The former
is the small pit in the middle line between the anterior ends of the basipterygoid
VoL. xv.—Part v. No. 9.—December, 1900. 2¢
174 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
processes ; the latter is more often slit-like and lies in the middle of the basitemporal
platform.
The parasphenoidal rostrum, which is of great length, is thin and rod-like in
Apteryx, Casuarius, Rhea, Dromeus, more or less inflated in the Dinornithide, Struthio,
and ‘Tinamous.
The Lateral Surface of the Cranium.—The tympanic cavity is moderately deep in
all save Apteryx, in which it is comparatively shallow. The mouth of the cavity is
somewhat lozenge-shaped in Dinornithide, Aipyornis, and more or less circular in the
other forms.
It is bounded posteriorly, in every instance, by the base of the paroccipital processes,
save only in the Tinamous, where, on account of the greater relative smallness of the
process, the tympanum is stretched along its free outer border. It is bounded
externally by the free inferior border of the squamosal. In all save Rhea and the
Tinamous, this border represents the posterior free edge of the zygomatic process.
In Rhea this process has shifted somewhat further forwards, and its place is taken by
a second but much shorter. ‘This second process may be regarded as a flange-like
downgrowth of the squamosal. The posterior region of this cavity lodges the apertures
of the internal ear; the anterior, in front of this, is continued forwards as a long,
spacious, tunnel-like pneumatic cavity to form the anterior tympanic recess. This
aperture is largest in Rhea, Dromeus, and Casuarius; it is much smaller in dpyornis.
The pre-temporal or alisphenoidal wing of the parasphenoid bounds the aperture of
this recess in front, and the pro-6tic forms its roof, the basi- and exoccipitals share in
the formation of its posterior and ventral walls. ‘The basitemporal plate and the
highly pneumatic basisphenoid receive the final termination of this recess beneath
the pituitary fossa, as is shown when the skull is seen in section. ‘The bones taking
part in the formation of this cavity can of course only be made out in the skulls
of very young nestlings. Each recess curves gently forward to meet its fellow of the
opposite side.
In Apteryx this recess is exceedingly small, and appears to lie, for the most part, if
not entirely, within the basisphenoid. There is no intercommunication between the
two recesses.
The roof of the tympanic cavity is formed for the most part by the large articular
surface of the quadrate. Behind this is a conspicuous cavity, the roof of which is
perforated by numerous pneumatic apertures leading into the diploé between the pro-
otic and squamosal. It represents the superior tympanic cavity. Mesiad of this is
the fenestral recess containing the fenestra ovalis and rotunda and the aperture of
the posterior tympanic recess. In the larger Dinornithide there is a small superior
tympanic recess.
The Squamosal Prominence.—This is bounded in front by the temporal fossa, behind by
the paroccipital process. It passes dorsad into the parietal, and is continued downwards
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHZ. 175
to form the zygomatic process. In Casuarius, Dromeus, Aipyornis, and Rhea this process
is of considerable length, and runs down to within a short distance of the cup-shaped
fossa for the articulation of the quadrato-jugal bar. In Rhea it runs forwards and
downwards to overhang this articulation.
In Dromeus, except D. ater, the anterior border of the squamosa! prominence bears
a strong protuberance.
In Struthio and Apteryx the zygomatic process is very short.
In Dinornithide the middle region of the squamosal is much inflated, making the
squamosal prominence very conspicuous.
The temporal fossa is more or less conspicuous in all the Palwognathe, save Apterya.
In Casuarius and Dromeus it is narrow and deep. In the latter its limits are
sharply defined.
In the Dinornithide the size of the fossa varies, but is generally wide and deep, its
general direction is backwards.
In Rhea it is wide and shallow, and with a linguiform impression. In Crypturi it
resembles that of Rhea, but is shallower.
In Struthio narrow, moderately deep, and slopes obliquely backwards.
Aspyornis has the same type of fossa as Struthio, but deeper and much more sharply
defined.
In Apteryx it is represented by a shallow but sharply defined depression extending
far forwards. It is peculiar from the fact that it is not bounded anteriorly by a post-
orbital process.
The post-orbital process in Caswarius is strongly developed ; it forms a broad
linguiform process extending downwards to within a short distance of the quadrato-
jugal bar. In Dromeus nove-hollandie and D. ater its downward extent is much less.
In ELmeus, amongst the Dinornithide, it is even larger than in Casuarius.
In khea it is relatively small, somewhat pointed, and directed slightly backwards.
In Struthio it is but slightly developed, and runs forward to blend with a shelf-like
projection of the frontal overhanging the orbit.
In Crypturi it is represented only by a minute prickle.
The trigeminal foramen is a conspicuous aperture lying, in Casuarius, at the bottom
of the temporal fossa at the point where the outer angles of the alisphenoid d
pre-temporal wing meet, and in the same transverse plane as the basipterygoid pro-
cesses. Immediately behind this, in Dromeus, and less distinctly in Apteryx and Rhea,
is a smaller aperture leading into the tympanic cavity.
The orbits in Casuarius, Dromeus, and Rhea are all more or less alike in form and
size. All are strongly walled-in behind by overhanging and laterally projecting post-
orbital processes, and are in front protected and increased by long backwardly directed
spurs from the lachrymal.
Struthio differs markedly from the above in that the spurs of the lachrymal over-
2c2
176 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
arching the orbits are short and blunt, but are connected with the frontals by a more
or less imperfect chain of supra-orbital ossicles, as in the Crypturi. Generally it
would seem, from the number of skulls which I have examined, that these supra-
orbitals fuse completely with one another and the lachrymal and frontal so as to
obliterate all trace of their independence. <A specimen in the Rothschild Museum,
however, has this chain on one side perfectly developed (Pl. XLII. fig. 3). The
existence of this supra-orbital chain seems to have escaped the notice of all recent
writers. On looking up the subject I find it was first described by Cuvier [14] in 1799.
Hildebrand [37] in 1806 seems to have rediscovered them. Meckel [56] and other
writers at the beginning of the century were also aware of their existence.
There is a distinct, but small, pre-sphenoid fossa. The optic foramina are distinct
and raised high above the rostrum, as in Apyornis.
Apyornis and Dinornithid@ agree in the form of the supra-orbital ledge. The pre-
orbital region of this is formed externally by the lachrymal, which has completely
fused with the frontal to form one uniform plate of bone. This is just what would
happen in Struthio if the fossa between its backward spur and the nasal were
filled up.
In Apyornis, as in Struthio, the optic foramen is raised high above the level of the
parasphenoidal rostrum. Below the lacerate fossa, and between it and the trigeminal
foramen, the surface of the combined alisphenoid and pre-temporal wing is much
inflated and roughened by numerous elongated spine-like processes for the attachment
of muscles.
In Dinornithide the optic foramina are deeply overhung by the outstanding orbito-
sphenoids; in this respect resembling Apterya.
In Apteryax the orbits are small and tubular, without post- or pre-orbital processes,
and the interorbital region of the frontals reduced to its smallest possible limit.
In Crypturi there is no lacerate or pre-sphenoid fossa or post-orbital process; the
pre-orbital backward spurs of the lachrymal are wanting, and the interorbital region
of the frontal is deeply notched, being cut away to within a short distance of the
interorbital septum.
The lacerate fossa in some Dinornithide takes the form of a deep pit; generally,
however, it is represented only by a shallow depression. In and around this lie certain
of the cerebral nerve-apertures worthy of note.
In Dromeus and Apyornis these are very distinct. In the former those of the
first division of the v. (orbito-nasal) and the vi. (abducent) nerves lie, the latter below
and slightly mesiad of the former, and both external to the rest. That of the tv.
(pathetic) lies mesiad of and slightly above the 111., being divided only by a narrow bar
of bone from the 11. (optic). The 111. oculo-motor lies directly below the rv. ; like the
Iv. it is only separated from the 1. by a narrow bony bar. Lowest of all, and forming a
triangle with the m1. and vi. foramina, lie the foramen for the arteria ophthalmica interna.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 177
In many Dinornithide these apertures lie closely packed together down the mouth
of a deep lacerate fossa.
In most of the other Palwognathe more or fewer of these foramina become more
or less confluent one with another and with the optic foramen.
The Ethmoidal Region.—The mesethmoid, in all but Dinornithide and Apteryx,
forms the thin median septum known as the interorbital septum. It is continued
forward into the pre-lachrymal fossa as far as the cartilaginous septum nasi. Its
ventral border rests upon the parasphenoidal rostrum ; its dorsal border affords support
to the frontal and nasals. This it does by means of paired, lateral, horizontal, ecto-
ethmoidal plates. These, the superior aliethmoids, curve outwards and downwards,
as the inferior aliethmoids, to form the walls of the olfactory chamber. It is
perforated superiorly in the dried skull so as to place the two olfactory chambers in
communication in all but Apteryx, Casuarius, and Crypturi.
In Dromeus and Casuarius the whole of the outer wall of the aliethmoidal region—
the only region that ossifies—is deeply invaginated. The lower region of this in-
vaginated wall forms a large, outstanding, antorbital plate, to the outer free edge of
which the lachrymal is closely applied (Pl. XLIV. fig. 1a). The olfactory chamber is
thus reduced to a narrow slit-like cavity, whose outer wall is scroll-shaped, the scroll
being formed by the invagination before mentioned.
This invagination lodges the Harderian gland, and is called therefore the Harderian
fossa. Opening above and behind this is a small supra-orbital fenestra.
In Rhea the invagination of the aliethmoidal wall is less sharply marked, and the
Harderian fossa, though larger, passes gently backward into the orbit. Except in
very old specimens, the greater portion of the wall of this fossa remains unossified.
The supra-orbital fenestra, when ossification is complete, is large.
In Struthio only the superior aliethmoidal wall is ossified. The postero-inferior
region forms the antorbital plate; this is continued upwards and forwards to form
the roof of the olfactory chamber. ‘There is a supra-orbital fenestra.
In Crypturi the antorbital plate stands out at right angles to the mesethmoid as
a narrow bar of bone. ‘The upper portion of the aliethmoidal wall resembles that
of Rhea in being deeply invaginated, so much so as to force it inwards almost on to
‘he mesethmoid, thus forming a huge Harderian fossa and reducing the olfactory
chamber to the smallest possible limits. ‘There is a well-marked supra-orbital fenestra.
In all the Palwognathe, save the Crypturi, the ossified ectoethmoidal plate extends
backwards, on either side of the dorsal border of the interorbital septum, as far as the
brain-case. Thus only the lower region of this septum is visible in the skeleton, not
its whole extent as in Neognathe. This feature is most noticeable in the Dinornithide
and in Apterya. In these the olfactory chambers are of enormous size, extending
backwards nearly or quite as far as the optic foramina, thus so far encroaching
upon the orbit as to obliterate the interorbital septum. ‘This last is traceable only
178 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
in the Dinornithide as the vertical plate dividing the pre-sphenoid fosse. There is a
well-marked supra-orbital fenestra.
The Dinornithide are apparently peculiar in the development of a pair of well-
developed triangular processes, forming a horizontal plate projecting from the ventral
border of the mesethmoid, and extending outwards to the level of the lachrymal.
In Apteryx the ossified ectoethmoidal plates form two oblong, much inflated bulle
on either side of the head, nearly filling up the very small orbit, the horizontal plates
of the mesethmoid sweeping outwards, downwards, and inwards, the ventral edge
fusing with the mesethmoid immediately above the parasphenoidal rostrum. Forwards,
the part corresponding to the antorbital plate affords support to the lachrymal, as in
Dromeus.
As the late Prof. T. J. Parker has pointed out [71], ‘‘One of the most striking
characteristics of the skull of Apteryx is the extreme complexity of the turbinals. When
the mesethmoid is removed there are seen in the olfactory chamber proper four well-
marked obliquely vertical folds, while a fifth is continued into the narrow or respiratory
portion of the nasal cavity. The three hindmost of these folds are perfectly distinct
one from another, and I propose to call them, respectively, the anterior, middle, and
posterior turbinals. ‘The fourth and fifth folds are intimately connected with the
anterior turbinal: I call the uppermost of the two, which forms part of the olfactory
region, the anterior accessory turbinal, and that which extends forwards, and has no
olfactory function, the ventral accessory turbinal.
“ All the turbinals, with the exception of the ventral accessory, are covered by
Schneiderian membrane, and are therefore analogous to the ethmo-turbinals of a mammal.
The ventral accessory turbinal is covered with ordinary mucous membrane, and belongs
to the merely respiratory portion of the nasal chamber ; it may be compared with the
maxillo-turbinals of mammals.
“On each side of the ventral edge of the mesethmoid, in the vomerine region, .... .
is a slender rod of cartilage, imbedded in connective tissue, and lying parallel to, and
either immediately dorsad or slightly laterad of the dorsal edge of the trough-like
VOMETe eens it can be readily prepared in a well-macerated skull by carefully
removing the vomer. It is obviously the vestigial cartilage of Jacobson’s organ .. .”
The ventral accessory turbinal does not appear to ossify in the adult Apterya skull.
There are no ossified turbinals in the other Palwognathe.
The /achrymal in Casuarius is crescentic in form, with convex border forwards.
Its upper limb, when visible, is very broad and deeply notched. It forms a broad
overhanging ledge to the orbit. Its lower limb is flattened from before backwards and
extends downwards to within a short distance of the quadrato-jugal bar. At the
junction of the upper and lower limbs it is perforated by a large foramen for the
passage of the lachrymal duct. In the adult, the inferior inner lateral border is fused
with the antorbital plate.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH. 179
In Dromeus the outer fork of the >-shaped notch of the upper limb is greatly
developed, forming a long, backwardly directed, spine-like supra-orbital process
(Pl. XLII. fig. 2). The inferior limb is notched below the lachrymal foramen.
In D. nove-hollandie the inner fork of this notch is short and confluent with the
free edge of the ventral border of the antorbital plate. The outer fork is produced
downwards far beyond this, and rests upon the quadrato-jugal bar. In D. irroratus
and D. ater the outer and inner forks of the prong are subequal, and do not quite
reach the level of the ventral border of the antorbital plate, which is continued
downwards by means of a slender column on to the maxillo-palatal process, where it
expands.
In Struthio (Pl. XLII. fig. 3) the outer fork of the upper limb is comparatively
short and blunt, and projects more laterally than in the above mentioned. Further-
more, it differs in that it is joined by a series of large bony scales lying between its
posterior extremity and the supra-orbital ledge of the frontal, recalling the supra-
orbital chain of bones in the Tinamou. Late in life the scales fuse with the lachrymal
in front and the frontal behind, thus enclosing a large supra-orbital foramen. The
lower limb of the lachrymal takes the form of a slender backwardly curved rod, whose
free end is curved sharply outwards to join the inner border of the quadrato-jugal bar.
To the inner side of this outwardly directed free end is attached a larger ossiculum
palatinum, which in the adult fuses with the lachrymal on the one hand, and the
antorbital plate on the other. Below the ventral border of its inner end comes in
contact with the palatine.
AEpyornis and Dinornis have the lachrymal so completely fused with the frontal
and antorbital as to be with difficulty distinguishable. ‘The upper limb in both has
completely merged into the supra-orbital ledge. The lower in Dinornithide is much
flattened antero-posteriorly. The lachrymal foramen is imperfect, its outer bar being
wanting. Insome, e.g. Anomalopteryz, it is perfect. Its ventral extremity is squarely
truncate and articulates with the maxillo-jugal bar.
In the only Apyornis skulls I have been enabled to examine, this lower limb has
been broken away.
In both these groups the anterior border of the lachrymal rests against a well-
marked, spine-like, maxillary nasal process.
In Apteryx the lachrymal is described by Parker [71], from young specimens, as “a
small irregular bone consisting of a shell-like central portion applied to the surface of
the aliethmoid, and of an ascending portion which articulates with the descending
process of the nasal. It is perforated obliquely by the lachrymal foramen.” In the
adult it is completely fused with the aliethmoid. Its anterior border is often traceable
as a thin suture between it and the maxillary process of the nasals.
In the Crypturi the lachrymal is indistinguishably fused with the nasal, and forms
therewith a prominent outstanding process bounding the orbit in front. There are no
backwardly projecting spines. The ventral limb fuses with the antorbital plate.
180 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The Cranial Cavity.—The metencephalic fossa in all, save Dinornis and Afpyornis,
takes the form of a moderately deep basin, the floor of which is tilted upwards toa
very considerable extent. The anterior tilted portion forms the posterior wall of the
pituitary fossa. Posteriorly it is continued backwards on to the occipital condyle. Its
postero-lateral border, immediately below the pro-6tic is pierced by the vagus foramen,
to the inner side of, and slightly posterior to, which lie one or two condyloid foramina.
Mesiad of the internal auditory meatus, and anterior to the same, is the abducens
foramen (Vv1.).
In Zpyornis and Dinornis the floor of this fossa is almost flat.
The ceredellar fossa is roofed by the parietal and supra-occipital bones. ‘The pro-
dtics bound it laterally, ventrally it passes into the metencephalic fossa. It is bounded
behind by the free edge of the occipital foramen, and in front by the median portion of
the tentorial ridge. The late Prof. Jeffery Parker, in his masterly monograph, states
that in Dinornithide the supra-occipital region of this fossa is marked by “ transverse
_grooves corresponding with the gyri of the cerebellum.” Immediately above the
internal auditory meatus lies the floccular fossa. The size of this, according to Parker,
appears to vary individually in Dinornithide. It is of considerable depth in the
other Palwognathe and in 'Tinamous.
The mesencephalic fossa is a deep basin-like depression in all save Dinornithide and
Apterys.
It appears to be more sharply defined in Casuarius than in any other member of this
group. It is bounded above by a very prominent shelf of bone, forming the lateral portion
of the tentorial ridge, behind by the pro-dtic. It extends forwards considerably
beyond the level of the pituitary fossa which bounds it in the middle line. ‘The
trigeminal foramen (v.) leaves by a large aperture excavated out of the ventral border
of this fossa, between it and the pro-6tic, and is of considerable size. Just within the
mouth of this foramen, below the rim of its anterior border, lies the aperture of the
orbito-nasal nerve (v.'), which has its exits in an almost obsolete lacerate fossa. In
Dromeus the trigeminal and orbito-nasal foramina are distinct. The tentorial ridge,
bounding the fossa superiorly, is almost as strongly developed as in Casuarius.
In Struthio this fossa is relatively smaller and shallower, neither are its boundaries
so sharply defined. The apertures of the trigeminal foramen and the orbito-nasal
nerve lie close together.
Rhea and Crypturi have the fossa somewhat more sharply defined than in Struthio,
but in none of these does that portion of the tentorial ridge bounding the fossa
superiorly form anything more than a low ridge.
In Dinornithide this fossa is relatively ill-defined and shallow. The aperture for
the trigeminal foramen steals away a large portion of its external wall. The orbito-
nasal aperture opens as in Casuarius, is tunnel-like, and bounds the fossa anteriorly,
separating it from the pituitary fossa.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 181
In Apteryx it appears to be reduced to the vanishing point. It is extremely small,
oblong in shape—its long axis horizontal,—and more or less sharply defined, when
seen at its best. It lies entirely behind the level of the pituitary fossa. It is
bounded above by a low tentorial ridge, in front by a ridge of bone dividing it from the
pituitary fossa, below by the rim of the metencephalic fossa, and behind by a huge
projecting shelf-like ledge from the tentorial ridge, The trigeminal foramen opens at
its postero-dorsal border, the orbito-nasal in its anterior border.
In A. haasti the outer wall of this fossa is almost entirely absorbed by a large aperture
common to the trigeminal and orbito-nasal nerves.
The pituitary fossa in Casuarius is a deep pocket-shaped depression. Its mouth is
directed slightly forwards. The rim thereof forms the dorsum sell@ behind, and the
pre-pituitary ridge in front. Within the mouth of this fossa (Dinornis) or on its rim
(Dromeus) is the oculo-motor (11t.) foramen. The pathetic (1v.) foramen lies above and
to the outer side of the oculo-motor, often it serves also for the passage of the internal
ophthalmic artery. Anterior to the prepituitary ridge in the middle line is a narrow
knife-like optic platform, on either side of which are the confluent optic foramina,
The optic platform is continued upwards vertically and again expands to form the
pre-optic ridge, which in Dromeus, and slightly less obviously so in Casuarius, joins
the tentorial ridge.
In Struthio the pituitary fossa is of great depth; the pre-pituitary ridge and optic
platform are well marked.
In Dinornithide the fossa is comparatively shallow, but of considerable circumference.
The optic platform is very wide.
In Apteryx the pre-pituitary ridge is but feebly developed. ‘The optic platform is
not distinguishable from the pre-pituitary ridge, which divides the optic foramina one
from another. ‘The pre-optic ridge is well developed and overhangs the fossa, the
mouth of which, in consequence, has a semicircular outline. It should be mentioned
here that the bottom of the fossa is produced forwards into a yet deeper, almost
spherical, depression, overhung by the pre-pituitary ridge.
In Tinamous this fossa is deep and tubular in form.
The cerebral fosse are sharply defined in all the Palwognathe save Apteryx. In all,
with the exception just mentioned, the cerebral lies entirely in front of the cerebellar
fossa. In Apteryx, and to a lesser extent in the Crypturi, the cerebellar contour is
scarcely distinguishable from the cerebral. ‘The tentorial ridge, which serves to cut off
the cerebral from the remaining fosse, arises from the level of the pre-optic ridge and
trends outwards and backwards to the level of the top of the pro-otic; from this point
it turns sharply upwards and forwards to meet in the mid-dorsal line, over the region
corresponding with a vertical line passing upwards from the dorsum selle, That
portion of the ridge which forms the upper boundary of the mesencephalic fossa in
Casuarius and Dromeus is of great depth, and projects into the cranial cavity like a
voL. Xv.—Part Vv. No. 10.—December, 1900. 2D
182 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
deep shelf. This is less marked in Rhea. In Dinornis and Crypturi it is but feebly
developed.
The cerebral fosse in Dinornithide do not taper forwards to finally pass into the
infundibuliform olfactory fosse, but, on the contrary, are of great size and width in this
region, reducing the olfactory fossz to the dimensions of shallow pits.
The median bony falx, continued forwards from the tentorial ridge to the crista
galli, is extremely well developed in Casuarius and Dromeus, less so in Rhea,
and only feebly in Struthio and Dinornis and Tinamous. ‘There is no trace of it
in Apteryx.
In Casuarius and Dromeus the fossee bear distinct and deep impressions for the
temporal and frontal lobes, a low ridge indicating an indistinct Sylvian fissure.
The olfactory fosse in all, save Dinornis and Apteryx, are paired, more or less
tubular, infundibuliform chambers, separated one from another by a crista galli.
Anteriorly they are closed by a more or less perforated plate for the passage of the
olfactory nerves.
In Dinornis the olfactory fossee take the form of shallow pits lying at the anterior
end of the large cerebral fossa. The cribriform plate is large.
In Apterya these fossee are of great size, and pass insensibly backwards into the
cerebral fosse. They are not, in the dried skull, shut off anteriorly from the turbinals
by a cribriform plate as in other Radtitw, but continued forwards directly into the
olfactory chamber.
The Premaailla.
The premaailia in all the Palewognathe is peculiar, in that the nasal process is
median, and not paired as in all other birds (Pl. XLII. figs. 1-3). In all save
Struthio there are well-developed palatine proceses.
In Casuarius the beak is more or less laterally compressed. The body—that
portion of the premaxilla from which the nasal, maxillary, and palatine processes are
given off—is short, and more or less deeply grooved on either side, so that the distal
end of the nasal process seems to be continued forwards into a strongly marked ridge.
The maxillary processes extend backwards as a pair of slender splints to the level of
the lachrymal on either side. ‘The palatine processes are a pair of short narrow
lamine extending backwards to a point corresponding with a vertical line passing
upwards behind the top of the free end of the median nasal process. They enclose
anteriorly a small chink-like palatine notch. In C. australis there is a small pre-
narial septum, corresponding to that described by Parker in Dinornis, as the ‘“ strong
ascending keel, formed posteriorly of paired plates, but solid in front and gradually
diminishing in height towards the top of the beak.” This keel is derived from the
dorsal surface of the palatine processes.
The nasal process is rod-like, its free end passes backwards into a tunnel-shape
in the inflated, mesethmoid mass forming the characteristic casque of these birds.
PHYLOGENY OF THE PLZOGNATHA AND NEOGNATHA. 183
In Dromeus the beak is flattened dorso-ventrally. The premaxillary body is larger
than in Caswarius, and, as in this genus, the rod-like nasal process seems to be
continued forwards to the tip of the beak by reason of paired lateral grooves which
divide it from the maxillary processes.
The nasal process is relatively longer than in Caswarius. Posteriorly it is received
into a shallow groove excavated out of the median borders of the nasals. The tip of
its free end rests upon the mesethmoid.
The maxillary processes are splint-like, and extend backwards to the level of the
anterior region of the orbit—beyond the body of the lachrymal.
The palatine processes are short pointed lamelle, rather more widely separated one
from another in the middle line than in Caswarius. The chink-like anteriorepalatine
foramen is closed caudad, as in Caswarius, by the vomer. ‘This process extends
backwards as far as the level of the anterior border of the posterior narial aperture.
In Casuarius, it should be mentioned, it falls short of this.
In Dinornis the median nasal process is markedly flattened and expanded dorso-
ventrally. ‘The maxillary processes are very short, not extending backwards beyond
the level of the hinder end of the antorbital fossa.
The palatine processes take the form of flattened lamine, extending backwards, as in
Dromeus, to the level of the anterior end of the posterior nares. The alveolar borders
are deeply pitted.
The anterior palatine foramen is small and cordiform. It is closed above by the
parasphenoid rostrum, and posteriorly by the vomer. In Hmeus it is more completely
closed than in any other Palwognathe by a pair of processes from the postero-internal
angles of the palatine processes, which meet in the middle line, serving at the same
time as articular surfaces for the anterior ends of the probably paired vomers.
The prenarial septum is most clearly marked in Dinornis. Its postero-ventral
border is bilaminate, but the lamine rapidly fuse to form the single median vertical
septum.
In Apyornis the nasal process rises sharply from the body of the bone, giving the
beak a peculiar conical form. ‘The alveolar border is deeply pitted. The backward
extensions of the palatine and maxillary processes cannot be made out in either
the British Museum skull or that belonging to the Hon. Walter Rothschild. In the
British Museum skull there is a prenarial septum, but the lamine composing this
have not fused to form a median plate as in Dinornis. Instead, as indicated, they
remain widely separated, and enclose between them a long tunnel-like cavity extending
forwards to the tip of the beak.
In Rhea the premaxillary body is of greater length than in any other Ratite, and
much depressed dorso-ventrally. It is grooved dorsally on either side of the distal end
of the nasal process, as in the other forms.
The nasal process is very broad and flattened, but tapers more or less caudad
9
‘*)
4 ~
134 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The maxillary, which is confluent for about half of its length with the palatine
process, is styliform, and terminates in the anterior region of the antorbital fossa,
The palatine process is of great size, extending beyond the maxillary process, and
terminating at, or falling but little short of, the anterior border of the posterior nares.
I this it agrees with Dromeus.
Struthio differs from all the other Ratite in the absence of a palatine process, and the
consequent enormous size of the palatine foramen.
The whole premaxilla has a unique triradiate form ; the radii are of great length,
so that the body of the bone is reduced to the smallest possible dimensions.
The nasal process is broad and flattened, tapering somewhat abruptly posteriorly.
The maxillary processes are long and styliform, terminating at the level of the
anterior region of the orbit.
In Crypturi the form of the premaxilla is precisely similar to that of Rhea. The
free end of the palatine process is brought to the level of the anterior border of the
posterior nares, and interlocks with the distal end of the palatine. In this last point
it differs from Rhea. The palatine foramen, as in Rhea, is long and narrow.
Apteryx is the most abnormal of all the Palwognathe. The whole beak is
produced forwards into an elongated probe, curved slightly downwards, in the tip of
which the external nostrils he.
In the adult skull the fusion of the separate elements, not only of this region of the
beak alone, but of the skull generally, is so complete, that it is impossible to
make out their boundaries.
The body of the premaxilla is reduced to its smallest possible limits, being com-
posed only by the confluent end of the rod-like nasal process, and of the combined
maxillary and palatine processes, which for this reason are but delicate rods.
The nasal process takes the form of a long slender rod, grooved ventrally and
running backwards to the mesethmoid. Its posterior region rests upon the elongated
nasal bones.
The maxillary and palatine processes are fused throughout the greater part of their
length, and form elongated flattened rods, divided in the middle line by a narrow groove
representing the palatine foramen. Posteriorly there is given off a short splint-like
palatine spur, representing the palatine process. The extremest limit of this falls
considerably short of the anterior end of the antorbital fossa.
The maxillary process can be traced backwards splint-wise as far as the level of the
foot of the lachrymal. Thus the maxillary processes are of about equal length,
‘The nasal, and nasal process of the premaxilla above, and the maxillary process of
the premaxilla below, serve to form a long narrow groove leading forwards into the
anterior nasal fossa, this last having shifted forwards to the extreme end of the beak,
This nasal furrow bears a strong resemblance to that of the Ibises, the difference being
that the nasal fossa hes behind instead of in front of the groove, as in Apteryx. It
seems to suggest that the shifting forward of the nasal aperture has followed the course
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA., 185
of the orbito-nasal nerve, accompanying it to its tip, so as to bring the sense of smell
to aid the sense of touch found in the Ibises.
It is interesting to note that in the very early embryo, as shown in Parker’s figures,
the position of the external nares is near the middle of the beak. Its forward shifting
seems to be involved in the growth of the elongating beak.
The Maxillo-jugal Arch.
The mazilla in Casuarius takes the form of an elongated flattened splint. Its
interior.half has the outer and inner borders turned upwards to meet in the mid-dorsal
line so as to form a long thin-walled tunnel tapering forwards to a point. This is the
antrum. The rim of the mouth of this is produced backwards into two short spikes ;
the outer forms a strong dorso-lateral and the inner a lateral spike, conspicuous when the
skull is viewed ventrally. The floor of this tunnel, viewed ventrally, is twice as wide as
the posterior region of the maxilla. This wider region—the floor of the antrum,—
which in the ventral view presents a deeply hollowed free posterior border forming the
anterior boundary of the posterior narial aperture, is the mavillo-palatine process.
The whole inner border of this process is grooved for the reception of the vomer.
The maxilla extends forwards above the maxillary and palatine processes of the pre-
maxilla. The latter process is received into a shallow depression in the maxillo-
palatine, a raised ridge of which bone can be seen lying between the palatine and
maxillary process. The backward extension of the maxilla is considerable. It runs
below the quadrato-jugal bar on the outer side and the palatine on its inner side to a
point slightly beyond the level of the free border of the hinder end of the posterior
nares, or, to take another standard, to within a short distance of the base of the vomer.
The quadrato-jugal fossa is the name which I have applied to indicate the large
space enclosed by the quadrato-jugal bar on the outside, the palatine and pterygoid in
the inside, and the hinder end of the maxilla in front.
In Casuarius it does not extend forwards beyond the hinder end of the posterior
narial aperture.
In Dromeus the maxilla is relatively much wider than in Caswarius, and does not
extend so far backwards. The maxillo-palatine processes are wider but very much
shorter antero-posteriorly than in Caswarius. The antrum, in consequence, is much
reduced in length and capacity. Its dorsal wall, moreover, is fenestrated. The mesial
borders of the maxillo-palatine processes rest upon the vomer.
The quadrato-jugal fossa extends forwards to within a short distance of the middle
of the posterior narial aperture.
In Dinornithide the maxillo-palatine region of the maxilla is short and broad. Its
anterior end is wedged in between the maxillary and palatine processes of the pre-
maxilla. Ventrally it appears as an oblong flattened plate. Its postero-internal
border bounds the anterior end of the posterior nares. Its posterior border is either
notched or hollowed and forms the anterior boundary of the quadrato-jugal fossa.
186 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
From its dorsal surface in Dinornis there arises a large hollow shell of bone, the inner
wall of which embraces the triangular plate of the rostrum. ‘The cavity of this bony
shell—the antrwm—opens posteriorly by a large foramen into the quadrato-jugal fossa.
It articulates with the palatine by a short recurrent process from its postero-internal
angle.
In Emeus there is no antrum. What corresponds to its inner wall in Dinornis exists
as a flat concavo-convex vertical plate applied to the anterior border of the mesethmoid
triangular process and the rostrum.
In Rhea the maxilla is almost entirely represented by the maxillo-palatine process.
Viewed dorsally it is seen to take the form of a flat, triangular, perforate plate of bone,
with its sharply pointed end running forwards to within a short distance of the
anterior end of the external narial aperture. ‘This more pointed region rests upon the
palatine process, and is bounded externally by the maxillary process of the premaxilla.
Its postero-mesial border is bounded by the vomer. Its hinder end presents free
postero-internal and external borders. The former lies more or less markedly forward
than the latter, and bounds the posterior nares anteriorly. The latter lies more or less
considerably caudad, and bounds the anterior end of the quadrato-jugal fossa. Its
postero-internal angle is produced backwards into a long spine, which is closely
applied to the outer border of the palatine, extending backwards for a considerable
distance. Its postero-external border is also produced backwards spine-wise, and
underlies the quadrato-jugal bar. From the dorsal surface of the maxillo-palatine
plate there arises a long slender style which, directed upwards and backwards, ultimately
articulates with the anterior angle of the lachrymal, and thus serves to divide the
lachrymo-nasal (antorbital) from the nasal fossa (Pl. XLIII. fig. 9). In all other
birds this is done by the descending maxillary process of the nasal. The antrum is
represented only by a minute aperture at the base of the vertical rod—the lachrymo-
nasal process of the maxillo-palatine—just described.
The guadrato-jugal fossa extends forwards nearly as far as the level of the anterior
end of the posterior nares.
In Struthio the maxilla is splint-shaped, produced posteriorly into two spine-like
processes, the inner of which runs dorsad over the distal end of the palatine, the outer
running ventrally below the quadrato-jugal bar. Anteriorly it extends forwards
nearly as far as the body of the premaxilla. The maxillo-palatine processes are small
and hatchet-shaped ; their mesial borders clamp the vomer, which is grooved laterally
for their reception. There is a small shallow antrum, the roof of which is fenestrated.
The anterior border of the maxillo-palatine process bounds the enormous palatine
foramen posteriorly.
The ossified style, described in Rhea as dividing the lachrymo-nasal from the nasal
fossa, is here represented by a ligament springing from the external dorsal angle of the
mouth of the antrum and inserted into the short descending process of the nasal.
PHYLOGENY OF THE PALEZOGNATHAH AND NEOGNATHA. 187
The quadrato-jugal fossa extends forwards to within a short distance of the anterior
border of the posterior nares.
In Crypturi the maxilla resembles that of Rhea, but its maxillo-palatine process is
much narrower and is unfenestrated. There is no antrum. ‘The lachrymo-nasal fossa
is bounded anteriorly by a descending process of the nasal and not by a spur from the
maxillo-palatine as in Rhea.
In Apterya the maxilla is of great size, its maxillo-palatine processes, seen ventrally,
extending forwards to the middle of the elongated beak, and backwards along the outer
border of the palatine to within a short distance of the level of the free end of the
orbital process of the quadrate. They are separated one from another in the middle line,
forwards, only by the narrow vomer. Posteriorly there is an external lateral spur for
articulation with the quadrato-jugal bar. The proximal mesial border is closely fused
with the external border of the short palatine. ‘There is no antrum. Immediately in
front of the distal end of the palatine the maxillo-palatine process occasionally sends
inwards a very short process to articulate with the vomer, which apparently
corresponds with the much larger vomerine process found in Struthio.
The quadrato-jugal is so reduced in size in Casuarius as to be represented by little
more than a nodule of bone, scarcely projecting beyond the limits of the glenoid cup
of the quadrate.
The jugal in Casuwarius is connected by a close vertical suture with the quadrate
portion, and projects above it at this part in the form of a more or less prominent
spine. It extends forwards as far as the middle of the lachrymo-nasal fossa.
In Dromeus, seen from the outside, the quadrato-jugal scarcely exceeds that of
Casuarius (Pl. XLIII. fig. 4a). On the inner side, however, it extends forwards along
the jugal for about one-third of its length. In Casuarius it does not extend beyond
the rim of the quadrate cup on the inside. The forward extent of jugal is about the
same as in Casuarius.
Rhea and Struthio resemble Dromeus in the size and relations of these two bones.
In Apterya the quadrato-jugal is a long slender style, reaching as far forwards as the
middle of the zygomatic arch. The jugal is long and slender, extending forwards
along the dorsal aspect of the quadrato-jugal as far as the level of the anterior
extremity of the palatine, which corresponds with a line drawn transversely across the
vomer about its middle.
In Crypturi the quadrato-jugal is as much reduced as in Caswarius. The mesial
border of the proximal end of the jugal articulates, as in this form, with the quadrate.
The Vomer, Pterygoid, and Palatine (see also p. 206). (Pl. XLII. figs. 5-8.)
In Casuarius the vomer is of great length and slenderness. In C. ¢. salvadorii,
C. ec. sclateri, and C. c. beccarti it is cleft from behind forwards for more than half its
188 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
length into two slender rods grooved dorsally; in C. bennetti only about one-fourth is so
divided: the cleft not extending more than half the distance of the posterior nares, in
the other forms it is continued forwards beyond this point. C. c. australis differs from
both types, the cleft of the vomer being rather less than half the total length and
terminating in the region of the anterior end of the posterior narial aperture.
Anteriorly the vomer becomes laminate and wedged in between the maxillo-palatine
processes, and passing forwards rests upon the palatine processes of the premaxilla.
The palatine foramen by the great forward extension of the vomer is almost obliterated.
In a ventral view—before complete anchylosis has taken place—the feet of the
vomer appear to be wedged in between the pterygoid and palatine. Removal of this
latter, or a dorsal view, however, reveals the true relations of these parts. It will then
be seen that the right and left limbs are closely applied to the external ventral border
of the twisted pterygoid, extending backwards to a point very near its middle. The
pterygoid can be traced forwards along the mesial border of the vomer as a slender
spike for some considerable distance. On a dorsal view, the right and left grooved
limbs of the vomer are well displayed, and show that each has been twisted outwards
so that its inner free border comes to look dorsally, and the latter ventrally. The
flattened surface of this twisted region is closely-applied to the ventral surface of the
anterior end of the pterygoid, so that vomer and pterygoid form one common rod
articulating with the quadrate. The dorsal border of the twisted end of the vomer is
slightly thickened and, traced backwards, trends downwards beneath the outer border
of the pterygoid. From the vomer backwards, the sutural surface is continued as a
shallow groove along the outer ventral border of the pterygoid, terminating at the
level of the basipterygoid processes. It is this ventral union of palatine and
pterygoid that gives the impression that the vomer is wedged in between these two
bones, when the skull is viewed from this surface.
In Dromeus (Pl. XLIT. fig. 7) the “feet” of the vomer extend forwards for about
one-third of its total length. Anteriorly it expands into a broad oval plate tapering
again forwards to terminate, as in Caswarius, near the body of the premaxilla. The
maxillo-palatine process overlaps the distal end dorsally; the palatine processes
underlie it ventrally. Its relations with the pterygoids and palatines posteriorly are
precisely similar to those which obtain in Caswarius. As a matter of fact, it should be
remarked, the palate of Dromeus is even simpler in type than in Casuarius, and
consequently represents, as we shall see, the simplest arrangement of these bones of
all the Palwognathe.
After these, the simplest palate amongst the remaining Paleognathine forms is that
of Struthio.
The vomer of Struthio (Pl. XLII. fig. 8) is a degenerate structure. Anteriorly
it does not extend beyond the middle of the enormous palatine foramen ; posteriorly
it is cleft, but has entirely lost all connection with the pterygoid, falling short thereot
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATHA. 189
by some very considerable space. It has assumed a trough-like form, the hollow of
the trough closely embracing the parasphenoidal rostrum. On either side the vomer
is closely embraced by the maxillo-palatine processes, which fit into specially roughened
facets. A similar, but less perfect, arrangement of these parts obtains in the
Cassowary.
If the vomer were produced backwards on the pterygoid we should have a
type of palate more nearly resembling that of Casuwarius and Dromeus than any
other Ratite.
In Rhea the vomer is cleft posteriorly for rather less than half its length, and is pro-
duced anteriorly into two large blade-like lamine, widely separated in the median line,
and each terminating in a pointed extremity. These anterior lamine are embraced
between the palatine processes of the premaxilla and the maxillo-palatines. The
parasphenoidal rostrum appears in the median line, between these processes,
terminating at the level of their free anterior ends. In Casuarius, Dromeus, and
Struthio the anterior end of the rostrum is entirely concealed by the vomer. In the
two former the free end of the vomer projects far beyond the rostrum, in the latter the
free end of this and of the vomer terminate at the same point. The body of the vomer
takes the form of a pair of vertical lamin, fused anteriorly, and moulded on to the
rostrum, upon which it slides; the forward anterior horns lie in a horizontal plane ;
the feet are obliquely truncated, their external lateral borders suturating with the
pterygoid (Pl. XLII. fig. 5).
The true relations of the vomer to the pterygoid cannot be made out except from
a study of the dorsal view; they will then be seen to be precisely what obtains in
Casuarius or Dromeus, the foot of the vomer underlying the anterior end of the
pterygoid. This last is in Ahea rod-shaped in form, but the sutural surface is
dorso-ventrally compressed and closely applied to the vomer; at least in young birds,
in adults it seems to become less compressed and yet more pointed. Ina ventral view
of the skull the vomer would appear to suturate with the palatine only and not the
pterygoid, the former being apparently thrust in between them.
Examined from the dorsal surface, however, it will be seen that the palatine—the
exact form of which will be discussed presently—underlies the foot of the vomer on
either side, and is continued backwards along the pterygoid in the form of a long
pointed spur, concealing all but the proximal end of the pterygoid, so that, seen
ventrally, as already remarked, the vomer and pterygoid would seem to be shut off one
from the other.
This is a fact of considerable significance, inasmuch as this isolation of the vomer is
a point wherein all the Neognathew appear to agree, and differ from the Palwo-
ynathe. But, as I have elsewhere shown, when discussing the palatine of Rhea
(p. 206), this is only an apparent difference.
In Crypturi the vomer is precisely similar in form and proportions to that of Rhea.
VOL. xvV.—ParT V. No. 11.—December, 1900. 22
190 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Furthermore, it agrees with the same in the nature of its relations with the pterygoid.
The palatine, in the part played by it with regard to the pterygoid and vomer, differs
but little from that of Rhea. Its chief difference consists in the fact that it is much
narrower from side to side, and that it terminates in a pair of spines, an outer and an
inner, instead of an outer pterygcidal only as in Rhea. The form of the palatine, as
a whole, will be described latter.
In Dinornithide, e.g. Hmeus (Pl. XLII. fig. 3), the vomer takes the form of a
pair of perfectly distinct vertical lamine embracing the parasphenoidal rostrum.
They are continued forwards only for a very short distance beyond the level of the
anterior end of the posterior narial aperture. It differs from that of Rhea (1) in the
greater vertical extent of the paired lamin, (2) in that these are often free
throughout, and (3) in that there are no anterior horns running forward between the
palatine processes of the premaxilla and the maxillo-palatine. In Himeus, the palatine
processes send inwards a pair of flattened spurs to embrace the sides of the anterior
end of the vomerine plate.
Posteriorly, the feet of the vomer suturate with the pterygoid exactly as in Rhea.
The relations between these bones and the palatine is also the same. Indeed, the
inturned under-flooring region of the palatine in Hmeus is more strongly developed
than in Lhea, projecting backwards to the level of its pterygoidal spur.
In Apteryx the vomer is relatively short, and cleft from behind forwards for about
half its length. Its form and relation are described on p. 205.
The pterygoid in Casuarius is, seen ventrally, au elongated, crescentic rod of bone
dorso-ventrally compressed, with convex surface directed towards, and partly resting
on, the parasphenoidal rostrum, and its concavity, for the most part, articulating with
the palatine. Anteriorly, it runs forwards along the inner side of the hinder limb of
the vomer—which is at this point wedged in between the pterygoid and palatine—and
terminating thereon in a sharp spine. Its proximal end articulates with the quadrate,
below the orbital process; mesiad and anterior to this articulation is a second for
articulation with the basipterygoid process. Seen dorsally, the pterygoid is fusiform,
not crescentic, a large median tube overlapping that part of the palatine which ona
ventral view appears within a crescentic hollow. Thus the true shape of the pterygoid
cannot be seen without the removal of the palatine, which discloses the dorsal
overlapping lip.
In Dromeus (Pl. XLII. fig. 7) the form and relations of the pterygoid are simpler.
Its relative length is about the same, and it terminates anteriorly ina point. The
anterior region of its external lateral border is directed upwards in the form of a thin
vertical plate affording a sutural surface for the external vertical plate of the vomer.
The posterior region of this border is straight and affords attachment to the palatine.
Its internal, mesial border is convex, the convexity facing the angle between the
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 191
parasphenoidal rostrum and the basipterygoid processes. The palatine is but loosely
attached to this bone. The free end of the vomer underfloors its anterior end.
In Struthio the form of the pterygoid (Pl. XLII. fig. 8) closely resembles that of
Dromeus: the points wherein it differs are points wherein it is also degenerate.
It lacks the upstanding vertical plate from its antero-external lateral border, and it
has fused with the palatine.
In Khea the pterygoid (Pl. XLII. fig. 8) has undergone a very considerable reduction
in size, and now takes the form of a \— -shaped rod. The short upper limb bears
facets for articulation with the quadrate and basipterygoid processes. The longer is
laterally twisted around its own axis, and continued forwards till it reaches the
parasphenoidal rostrum. Ventrally it is grooved. This grooved surface is applied,
anteriorly, to the superior border of the vomer, and posteriorly to the upper surface of
the postero-external angle of the posterior extremity of the palatine. Thus the
posterior end of the palatine comes to underlie the posterior extremity of the vomer.
In the Dinornithide, e.g. Hmeus, the form of the pterygoid very closely resembles
that of Rhea. The points wherein it differs are such as must be regarded as repre-
senting a more primitive condition: the chief of these lies in the greater width dorso-
ventrally; the pterygoid of Hmeus possessing a broad ledge along its mesial border.
Its relations with the vomer and palatine are practically the same as in Rhea.
In Crypturi the pterygoid is rod-like as in Rhea, relatively longer, much straighter,
and more slender.
In Apteryx the pterygoid is elongate, laminate, and pointed in front where it rests
upon and fuses with the vomer. It anchyloses very early and completely with the
palatine, so that the precise form of the pterygoid as a whole cannot be made out in
the adults (see p. 204). Proximaily, it is tightly wedged in between the basipterygoid
process on the one side and the orbital process of the quadrate on the other, the
pterygoid having the appearance of being “‘morticed” in between these two points
of contact.
The palatine in Casuarius takes the form of a roughly triangular plate. Its base
articulates with the pterygoid and vomer, and the anterior half of its outer side with
the maxilla. Its connection with the pterygoid is by means of an overlapping suture,
the pterygoid underlying a broad shelf of bone given off from the palatine. The
vomerine suture is that of a simple apposition of contiguous edges, as also is the
junction with the maxilla, In the adult the palatine and pterygoid become indistin-
guishably fused proximally.
In Dromeus the palatine resembles in form that of Casuarius. It is, however,
fenestrated. Its suturations differ only in that there is no overlapping shelf above
the pterygoid. ‘here is no fusion between pterygoid and palatine as in Caswarius.
In Rhea the palatines (Pl. XLII. tig. 6) are broad thin lamine, roughly quadrangular
2E2
192 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
in form, with the external border produced backward into a pointed spur which runs
along the pterygoid, and forwards for a considerable distance along the maxilla. Its
mesial border is cut away anteriorly so as to embrace the external border of the vomer
instead of underlying it, as is done by the remaining posterior portion. Its form is
seen best from the ventral aspect of the skull.
In the Dinornithide, e. g. Emeus, the form of the palatine resembles that of Rhea,
but by a little tension the body of the bone has come to be vertical instead of hori-
zontal in position, thus giving a rod-shaped superficial appearance.
In the Crypturi the main body of the palatine is, as in Rhea, horizontal, but the
anteriorly produced external lateral border is of much greater length, and takes the
form of an elongated and slender rod suturating anteriorly with the maxilla.
In Struthio the palatine (Pl. XLII. fig. 8) resembles that of many Carinate in that
the main body of the bone is twisted into the form of an almost vertical scroll. It is
peculiar, however, in that it is fused at its base with the overlapping pterygoid. The
external lateral border is produced forwards, as in Tinamous, into an elongated slender
rod, but which fuses anteriorly with the base of the hastate maxillo-palatine process.
In Apteryx the palatine (Pl. XLII. fig. 5) takes the form of a more or less flattened
rod, differing in its relations from all the other associated forms. Anteriorly, it
is fused with the maxillo-palatine process. The greater part of its external border is
anchylosed with the maxilla.
The remaining posterior end of this bone anchyloses with the external border of
the pterygoid. What corresponds with the posterior region of the mesial border in
Dromeus is here free, and runs in a backwardly transversely oblique direction across
the skull. For the precise relations of the bone however, see p. 204.
The Quadrate. (Pl. XLITI. figs. 10-12.)
The quadrate in Caswarius has its various processes very sharply defined.
The ofie process is very slender, tapering gradually downwards into the body of the
bone. At its base, where it merges into the main body, isa large pneumatic foramen,
it opens on the posterior surface. The head for articulation with the pro-6tic and
squamosal is single, oblong in form, with its long axis directed backwards and inwards.
The inner and outer mandibular condyles join mesially, the two projecting backwards
to form a conspicuous spur. The difference in the level of the two condyles is but
slight.
The orbital process is well defined, moderately large and free, with a sinuous
superior border. At its extreme base, and on its inner side, is a small demi-facet for
the pterygoid.
‘The outer surface of the quadrate is very closely applied to the long stout zygomatic
process of the squamosal, which terminates only a short distance above the level of the
cup-shaped quadrato-jugal fossa.
PHYLOGENY OF THE PALHOGNATHA AND NEOGNATHA, 195
The quadrate of Dromeus differs but slightly from that of Caswarius. The outer
and inner mandibular condyles are on distinctly different levels, the inner being
ventrad of the outer, shorter from without inwards and wider from before backwards.
The backwardly projecting condylar spur is less marked and formed entirely by the
outer condyle.
The orbital process is less hollowed above.
In the Dinornithide, e. g. Emeus, the quadrate resembles that of Dromeus in the
difference of level between the outer and inner mandibular condyles and in the form of
the orbital process. This process is, however, peculiar in that its whole inner surface is
requistioned for the articulation of the pterygoid. The inner mandibular condyle is
peculiar in that the middle region of its articular surface is produced downwards, whilst
the outer condyle looks more directly downwards than in Dromeus.
The pneumatic foramen is mesiad, and not posterior, A posterior foramen is, however,
sometimes present in the Dinornithide.
In Struthio the inner and outer mandibular condyles are widely separated ; otherwise
they closely resemble those of Dromeus. The orbital process differs from that of the
other forms so far described, in that its distal end turns sharply upwards, and in that
there projects from its inner ventral border a sharply defined articular facet for the
pterygoid. ‘The ofc process is relatively longer than in the other forms. Both the
mesial and posterior foramina are present, though the latter may apparently be
sometimes absent.
In Aipyornis the mandibular condyles resemble in form and disposition those of
Struthio, and in some respects Dinornis. The form of the orbital process seems to have
differed from all other genera in its great breadth. Its free end is missing, but from
the sharp edge of the proximal end it is probable that it articulated with the pterygoid
as in Struthio. 'The pneumatic foramen is mesial. The specimen from which this
description is taken was kindly lent me by Dr. Forsyth Major.
In Rhea the quadrate resembles that of Dromeus rather closely. The external
mandibular condyle is broken up into two distinct articular facets. The inner is
similarly subdivided but not so distinctly ; of these two last the outer is vertical and
looks forwards and outwards, the inner ventral. The otic process is very short. Its
articular head has its long axis at right angles to the long axis of the skull. The
posterior pneumatic foramen lies immediately below the articular condyle. The
orbital process is very short, blunt, and very wide, it does not project beyond the level
of its articulation with the quadrate.
In Crypturi the form of the quadrate is quite peculiar in the development of a
distinct cingulum projecting backwards above the external mandibular condyle. The
otic process is bent backwards at a sharp angle, and forms with the orbital process a
Y-shaped bone. The external form of the orbital process resembles that of Struth/o
but is more slender; the method of articulation with the pterygoid closely resembles
194 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
that of Dinornis. The form of the outer and inner mandibular condyles is not unlike
that of Dinornis. The inner condyle is, however, more nearly spherical than in any of
the other forms herein described.
Apteryx possesses a readily distinguishable quadrate. ‘The height of its vertical axis
is less than that of a line passing backwards through the orbital process, which is very
long.
The articular condyles for the mandible present the same ground type as in Dromeus
and Casuarius, but the external condyle presents a tendency to split into two facets,
the region nearest the quadrato-jugal bar being scooped out. The zygomatic process
of the squamosal is closely applied to the outer side of the quadrate. In Struthio and
Rhea, it should be mentioned, there is a considerable space between these two points.
In all the Palwognathe the quadrate is peculiar in that the pterygoidal articulation
extends upwards on to the orbital process for more than half of its length.
In Apteryx the articular surface for the pterygoid is rather more sharply defined
than in the other Palwognathw. Arising at the base of the internal mandibular
condyle, it extends outwards on to the inferior border of the base of the orbital process,
taking the form of a deep hollow into which the pterygoid closely fits.
The otic process has two fairly distinct articular facets. Superiorly transversely
elongated, its mesial extremity is expanded into an almost circular head and articulates
with the pro-6tic. Its external extremity runs obliquely down to the shaft towards the
middle line, and affords an articular surface for the alisphenoid and squamosal.
The pneumatic posterior aperture is sometimes absent.
The separation into pro-6tic and squamosal heads in the other Palwognathe is only
very feebly indicated.
The exoccipital does not contribute towards the articular surface for the quadrate in
Apteryx.
The Mandible. (Pl. XLIIL. fig. 9.)
The mandible in Casuarius is long, slender, and sigmoidally curved. Posteriorly, it
is expanded to afford an articular surface for the quadrate. This is moderately deep,
and with sloping sides meeting at a sharp angle at the bottom of the fossa. Internally,
it is produced into a blunt ¢nternal angular process. The surface for the external condyle
forms a long narrow facet running parallel with the external border of the cup ; that
for the inner forms a band-shaped facet, hollow from within outwards. Immediately
without the inner end of this internal facet is a pneumatic foramen. ‘The posterior
angular process is only feebly developed.
In Dromeus the internal and external angular processes are larger than in Casuarius.
The rami are almost straight, and the symphysis is very broad. In other respects the
jaw agrees with that of Casuarius. In Dinornithide, e. g. Dinornis, the articular
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA., 195
surface for the external condyle of the quadrate is cup-shaped. The internal and
external angular processes are feebly developed.
In Rhea the form and disposition of the articular surfaces agree with those of
Dromeus. The internal angular process is large. The symphysis is very broad, The
ramus bears a small, perforate posterior lateral vacuity.
Struthio in the form of its mandible rather closely resembles Dromeus. It differs,
however, in its greater relative slenderness, in that it is sigmoidally curved, in the
greater relative size of the expanded posterior portion, and in the elongated form of
the articular facet for the external condyle of the quadrate.
Aipyornis also resembles Dromeus. The rami are, however, more laterally com-
pressed, have an elongated anterior lateral vacuity, a broad dentary border, and the
articular facet for the internal condyle of the quadrate situated at the bottom of a deep
cup. The internal angular process is well developed and bears a large pneumatic
foramen.
In Crypturi the mandible resembles that of Rhea. The posterior angular process is,
however, longer in Tinamous, and the posterior lateral vacuity appears to be wanting.
Aptery« differs from all the other forms in the great length of the symphysis, which
is about half the length of the ramus. The internal angular process is well developed.
The coracoid in all the Paleognathine forms is small and rod-shaped. Sometimes, as
in Struthio, it is reduced to a mere vestige. The dentary suture is always distinct.
The Hyoid.
Except in Apteryx, only the cerato-branchials appear to ossify in the Paleognathine
forms. Epibranchials and the true hyoid elements appear to remain permanently
cartilaginous. In Apteryx the basihyal appears to ossify at least occasionally.
Tue SKULL or THE NESTLING.
The Cartilage-bones.
The youngest of the Palzognathine skulls in my possession is that of an embryo
Dromeus. The cartilage-bones are in all the stages herein described completely
ossified.
The basi-occipital, in the young Dromeus mentioned above, is perfectly distinguishable
from the neighbouring bones (Pl. XLIII. fig. 5).
It is bounded antero-laterally, and in front, by the basi-temporal plate of the para-
sphenoid, and postero-laterally by the exoccipital. It is produced caudad to form
the median portion of the occipital condyle. It is interesting to note, however, that
it forms only the ventral part of this. It is entirely shut off from the occipital foramen
196 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
by the exoccipital portions of this condyle, which meet in the median line above it.
Seen dorsally, the basi-occipital is cut off from the pro-étic by a broad patch of cartilage.
Anteriorly it only just touches the basisphenoid.
In the youngest Cassowary and Rhea skulls in the Museum collection, the sutures
of the basi-occipital have almost closed. It takes, however, a slightly larger share in
the formation of the occipital condyle, inasmuch as it takes a small share in the
formation of its free border.
In a half-grown Apterya australis mantelli it agreed very closely with that of
Dromeus, and, as in this form, the exoccipitals meet in the middle line above the basi-
occipital.
The exoccipital, externally, is relatively large. The upper portion of its posterior
(inner) border bounds the infero-lateral border of the epiotic (fig. 4). The lower
region of this border is hollowed to form the lateral boundary of the foramen magnum.
The share which it takes in the formation of the occipital condyle is considerable: in
Dromeus the right and left plates meeting in the middle line behind the basi-occipital.
Its dorso-lateral border is bounded by the parietal and squamosal in Dromeus, Casu-
arius, and Apteryx, and by the squamosal only in Rhea-—this last, in Rhea, extending
backwards to join the supra-occipital. The lower portion of the antero-lateral border
is free, and in the shape of a large paroccipital process bounds the tympanic recess
posteriorly. Its internal dorsal angle lies within the tympanic cavity, and helps to form
the inferior pro-6tic articulation for the quadrate.
Internally, the exoccipital is almost entirely concealed by the opis- and pro-dtics.
The supra-oceipital, in Dromewus and Rhea, runs upward and forward in the form of a
blunt cone; in Apterysx the superior border is gently hollowed. In Dromwus, Casuarius,
and Apteryx it is cut off from the squamosal by the exoccipital ; in Rhea it runs up-
wards to join the squamosal. That portion of its outer (inferior lateral) border which
comes into contact with the exoccipital really belongs to the epiotic. In the Penguin
this last is easily distinguishable, but in the forms now under description it can only
be made out on a reference to the interior of the skull.
The epiotic is not indicated externally in the peculiarly distinct manner seen in the
Penguins ; even in the skull of the youngest of the embryo Apteryxes in our collection
its only indication is the venous canal which points the boundary line between this and
the supra-occipital. Internally, it is fairly distinct in all save Apteryx. In all the
skulls of this collection save those of the embryo Apterya and Dromeus, it has fused
with the pro-dtic.
The pro-otic, even in the youngest Apteryx (Pl. XLIII. fig. 3) and Dromeus
(Pl. XLITT. fig. 5) skulls, is not visible externally above the tympanic cavity, being
concealed by the squamosal. In this it differs markedly from the young Penguin,
in which a large portion of this bone remains uncovered until the bird is quite
half-grown (82).
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH#. 197
The pro-6tic completely shuts out the squamosal from the interior of the skull in all
but Apteryxr. In this type a large, roughly triangular plate appears immediately above
the pro-étic and wedged in between the parietal and alisphenoid.
The floccular fossa is deep, and oblong in all but Apterya, the long axis running
vertical to that of the skull. In Apterya it is represented by a small but deep and
perfectly circular pit.
Externally, the pro-6tic appears only within the tympanic cavity and affords two more
or less distinctly separated articular surfaces for the quadrate: the one lying slightly
above and in front of the other. Distad of the superior glenoid surface lies a similar
articular surface furnished by the alisphenoid; caudad of the inferior lies a second
facet furnished by the exoccipital. These appear to be most distinctly traceable in
Casuarius. Externally, articular surface for the quadrate is completed by the squamosal
Thus there are two pro-6tic, one exoccipital, one alisphenoid, and one squamosal
articular facet for articulation of the quadrate.
The opisthotic is seen as a small oblong nodule of bone wedged in between the pro-
otic and the exoccipital, and bounded above by the inferior end of the epiotic. It is
not visible externally.
The foramen for the vagus lies at its ventral extremity and between it and the exoccipital.
The dasisphenoid has in every case fused completely with the underlying basi-temporal
plate, and is therefore traceable only from the superior aspect of the skull. It is
moderately thick in section, the body of the bone being thickened by a mass of diploé.
It is bounded behind by the basi-occipital, laterally by the pro-dtic and alisphenoid.
Anteriorly, in Rhea, it is produced forward into a thin vertical plate of bone, to join the
cartilaginous presphenoid; in the other forms this vertical plate is much shorter. It
serves to form the inner half of the circumference of the foramen for the orbito-nasal
nerve (V.), the outer portion being contributed by the alisphenoid. Out of its
anterior region is scooped the pituitary fossa. The abducent foramen pierces it on
either side near its postero-lateral angle. It forms the floor of the anterior region of
the metencephalic fossa.
In Apteryx it is almost quadrangular. In Casuarius and Dromeus, and in Rhea, it is
expanded laterally to form a pair of wings, and is narrower behind than in front.
The alisphenoid is bounded by the parietal and squamosal behind, the orbital plate of
the frontal above, and the combined basisphenoid and alisphenoid as rings of the
parasphenoid.
In Dromeus, Casuarius, and Rhea it bears a share in the formation of the post-orbital
process.
The alisphenoid lodges the greater part of the mesencephalic fossa, and contributes
a share towards the formation of the anterior wall of the cerebral fossa.
The orbitosphenoid is represented only by cartilage and is much reduced in size.
The presphenoid is represented by a thin vertical plate of cartilage resting upon the
VoL. xv.—Part v. No. 12.— December, 1900. 2 &
198 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
parasphenoidal rostrum and wedged in between mesethmoid in front and the anterior,
laterally compressed vertical plate of the basisphenoid just described. It extends
upward and backward to the orbital plate of the frontal. From this region it sends
outwards and backwards a pair of cartilaginous wings, which represent the orbito-
sphenoids.
The mesethmoid, in all but Apteryx, in common with the presphenoid region, forms
a conspicuous interorbital septum. In the nestling Dromwus, Caswarius, Rhea, and
VOstrich, and in the adult Tinamou, this is pierced by a large interorbital fenestra.
This fenestra appears to be excavated partly out of the large, backwardly directed
plate which unites with the orbital plate of the frontal, and partly (caudad) out of the
anterior border of the presphenoid. In the nestling, the antero-ventral border is more
or less linguiform, and terminates at the level of the anterior border of the lachrymal.
From this, there runs forward a narrow, triangular, cartilaginous septum nasi. In the
adult Casuarius, Dromeus, Dinornithide, and 'Tinamous, ossification extends forwards
almost to the end of the rostrum, and terminates in a vertical truncate border, beyond
which there is a small cartilaginous septum nasi. In Rhea and Séruthio the ossification
extends quite to the free end of the rostrum, 7. ¢. relatively further forwards, and
terminates in a sharp point instead of a vertical truncated border. In these, the
cartilaginous septum nasi is wanting or greatly reduced.
The dorsal border of the mesethmoid, in Dromeus, Caswarius, and Dinornithide, is
of considerable thickness and very slightly convex ; it forms a complete floor to the
olfactory fossa, extending outwards on either side to the orbital plate of the frontal
(Pl. XLIV. fig. 4). In this particular, as will be seen later, it differs conspicuously
from Apteryx, wherein this region forms a thin median partition between two extensive
cavities. At the anterior end of the olfactory fossa, both in Dromeus and Casuarius,
it sends upwards, and then backwards, a strong median pillar deeply grooved on either
side for the passage of the olfactory nerves. This corresponds to the tegmen cranii
described in Apteryx by 'T. J. Parker, and referred by Kitchen Parker to the tegmen
cranii of the lower vertebrates. Thus the evista galli,in these two forms, is furnished
entirely by this tegmen cranii, the “ tegminal process ” of Jeffery Parker.
At the apex of this tegminal process, as has been well described by Parker, the
mesethmoid reaches the outer surface of the skull; here it throws out lateral horizontal
ecto-ethmoidal plates—the aliethmoids, as Jeffery Parker proposed to call them. In
Dromeus the upper surface of these plates forms a lozenge-shaped area, bounded by
the frontals behind, the nasals laterally, and the nasal process of the premaxilla in
front. The free edges of these right and left aliethmoidal plates in the nestlings turn
downwards and inwards (Pl. XLIV. fig. 4), and in the adult become further extended
and intricately folded to form the lower region of the aliethmoid and the antorbital
plates. In the nestling Dromeus it should have been remarked that the median vertical
portion of the mesethmoid is pierced by a large vacuity, the cranio-facial fissure.
PHYLOGENY OF THE PALZOGNATHZ AND NEOGNATHA. 18)
Casuarius differs conspicuously from Dromeus and the other forms associated here-
with in that, that portion of the mesethmoid which in Dromeus, for instance, reaches
the outer surface of the skull to form the lozenge-shaped plate already described, is
surmounted by an independently ossifying mass forming the centre of the characteristic
casque of the adult—the lateral portions of the casque being furnished by the inflation
of the nasals and that portion of the frontal bone which comes into juxtaposition with
the nasal and median masses (Pl. XLIV. figs. 8 & 4.a—b). The median portion often
extends so as to cover the parietal.
In the adult this casque is seen to be built up of a homogeneous mass of cancellated
tissue of great delicacy, and protected in the living bird by a horny sheath. The form
and development of this casque varies considerably, and is one of the factors employed
in the determination of species.
The exact homology of this casque has yet to be made out. That the lateral
portions are simply inflations of the nasal and frontal bones there can be no doubt, but
the origin of the median portion is not so certain. It appears to be an independent
structure superimposed upon the mesethmoid, rather than an outgrowth of that element.
Stages in the growth and composition of this casque are shown in Pl. XLIV. figs. 3
and 4 a—0.
In Rhea and the Tinamous, as in Dromeus and Casuarius, the dorsal border of the
mesethmoid comes into close relation with the orbital plate of the frontal, but the
relationship is even closer, this border being narrower, so that the crista galli is
formed only by the tegminal process. But the passage for the olfactory nerves is
restricted to an exceedingly narrow space lying on either side of this process. In
Dromeus and Casuarius this passage, though formed in precisely the same way, is
much larger.
In Apterya the dorsal border of the mesethmoid is exceedingly narrow and knife-
like, and slightly concave. It is, moreover, relatively much shorter antero-posteriorly
than in Caswarius or even Dromeus. ‘The tegminal process is less well-developed. In
the embryo, it appears on the surface of the skull as in Dromeus, and also, as in this
form, sends out horizontal plates which form a lozenge-shaped shield between the frontal
aud nasal bones. Later in life all trace of this shield is obliterated.
The quadrate in the embryo of Dromeus differs in shape from that of the adult in
the lack of an ossified orbital process, and in that the external mandibular condyle
is <-shaped instead of transversely oblique.
In Apteryx that of the embryo appears to differ in nowise from that of the
adult.
The other skulls in the Museum collection are too advanced to afford any intermediate
characters.
The articular is a pyramidal nodule of bone, bounded cephalad by the coronoid
the base of which conceals its anterior face; externally (in Rhea) by the supra-
2F2
200 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
angular. It furnishes the articular surface for the quadrate. In Dromeus and Apterys,
both angular and supra-angular take part in the protection of its external face.
The stapes is represented in the dried skull only by the bony base. It does not
apparently offer any characters of value from a systematic point of view. The extra-
columella has been already admirably described by Gadow, Parker, and others.
The Membrane-bones.
The parietal, in Casuarius, Dromeus, and ? Struthio, is a transversely oblong plate
of bone, the anterior and outer borders of which are produced forward into a small
but sharp point which is wedged in between that portion of the frontal which over-
hangs the post-orbital process and the process itself. ‘This parietal spur thus helps to
form the base of the post-orbital process, the main body of which is furnished by the
alisphenoid.
In Rhea and ?'Tinamous this antero-external parietal angle is not produced forwards,
and fails entirely to reach the postorbital process (Pl. XLITI. fig. 9). Its postero-
external angle is overlapped by an upstanding process of the squamosal.
Internally the parietal carries on the tentorial ridge from the alisphenoid upwards to
its junction with the falx. Behind this ridge it is gently scooped out to complete the
roof of the cerebellar fossa, in front it is also hollowed out to form the posterior wall
of the cerebral fossa.
In Apteryx, as in the other forms, the parietal is irregularly four-sided. Its external
border is bounded entirely by the squamosal. In all the other forms, it will be noted,
it extends forward beyond this.
Internally, it is found to lack the tentorial ridge. Its hinder and external borders
pass insensibly the one into the other. ‘The former skirts the supra-occipital and a
portion of the pro-6tic beyond, the latter rises gently from the hinder region of the
superior border of the pro-6tic—where the hinder border may be said to cease—upwards
to skirt the upper’ border of the small triangular area of the squamosal which enters
into the formation of the inner wall of the skull. It is entirely cut off from the
alisphenoid. In the other forms, e. g. Rhea, Dromeus, the superior borders of the
pro-6tic and the alisphenoid form the boundaries ventrad of this bone.
The frontal extends backwards so as to form the greater part of the roof of the
cerebral fossa. Its hinder border is more or less sinuously curved, and traced from
within outwards will be found, in Caswarius and Dromeus, to run transversely from the
middle line to the base of the post-orbital process, from which, however, it is actually
cut off by a very slender spur from the parietal. In Rhea, on reaching the antero-
external angle of the parietal, it turns abruptly upwards and outwards, so as to rest
upon the base of the post-orbital process itself. Externally, the frontal is bent in upon
itself to form a deep orbital plate which articulates caudad with the alisphenoid and
cephalad with the mesethmoid, eventually terminating in a long spike, in 2hea, resting
upon the mesethmoid, and overlapped by a greatly elongated posterior extension of the
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 201
nasals and the free end of the inner spur of the lachrymal. In Dromeus the relations
are similar, but the nasal spur is shorter.
In Casuarius the forward extension of the frontal is curtailed, since it terminates
with the mesethmoidal region of the orbital plate, at the level of the origin of the
antorbital plate. The greater part of its mesial border is much thickened and inflated ;
this region being involved in the formation of the casque, and representing its hindmost
postero-lateral boundary. In some species this inflation of the bone extends backwards
as far as the parietals.
Furthermore, it is significant to note the fact that the frontals fail to meet in the
middle line anteriorly, where they under-floor the median ossification of the casque
(Pl. XLIV. fig. 4). Thus a portion of the ventral border of the casque comes to take
part in the formation of the roof of the cerebral fossa. It would seem, from this, as
though this secondary ossification was gradually engrafting itself more and more
completely upon the skull, by absorption of the underlying bone in much the same way
as bone has replaced cartilage in other cases in the cranial skeleton.
The extreme anterior end of the mesial border of the frontal is gently hollowed out
so as to form the external lateral boundary of the superficial lozenge-shaped plate of
the mesethmoid.
In Apteryx the frontal presents one or two points of difference from that of the
remaining forms associated herewith.
Its mesial border joins its fellow of the opposite side, and its hinder border is
bounded by the parietal, as usual. Its postero-ventral or lateral border joins the
aliethmoid. Immediately in front of this, the frontal sends downwards and inwards a
narrow plate of bone which, eventually, meets its fellow of the opposite side in the
middle line, immediately behind the crista galli. From the outside, this orbital plate
of the frontal gives off a short slender plate to the aliethmoid.
Cephalad of this orbital plate is a deep notch, the swpra-orbital fontanelle. This
fontanelle is bounded anteriorly by a narrow bar of bone, called the descending process
of the frontal, which at its ventral extremity joins the aliethmoid. Mesiad of this
bar—between its upper and the outer wall of the aliethmoid—the orbito-nasal nerve
passes from the orbit to the nasal cavity.
The sgwamosal, in all the forms herein dealt with, entirely conceals the pro- and
opisthotics. In the embryo Dromeus, which forms the subject of this paper, it is almost
quadrangular, its antero-ventral angle is produced into a blunt process which extends
downwards to the level of the quadrato-jugal articulation. ‘This downward process of
the squamosal is thickened on its inner surface and so closely applied to the quadrate
as to render it immovable.
In the embryo skulls of Dromeus, the squamosal and the parietal are connected
by means of a close suture, the juxtaposition of the two elements forming a perfectly
level surface. Posteriorly the squamosal rests upon the outer border of the exoccipital,
and later fuses therewith. Similarly the squamosal, anteriorly, rests upon the ali-
202 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
sphenoid, anchylosis therewith taking place later. ‘Thus it is that the pro-6tic comes to
be entirely concealed.
Tn all this, Dromeus and Casuarius seem to be less primitive than the Penguin. Ina
skull of this group recently described and figured by me [82] the squamosal was attached
to the dried skull simply by its articulation with the parietal. Its hinder border was
deeply emarginate and free, exposing the pro-6tic. Its anterior border was also free,
not extending to the alisphenoid, but exposing between itself and this last a small
portion also of the pro-dtic.
There is one great point of difference, however, between the squamosal of Dromeus
and that of the Penguin. In the former, its antero-ventral angle is produced
downwards into a large “‘ zygomatic process,” to embrace the quadrate; in the latter,
this spine is replaced by two small spines, as in all the Neognathe.
Except in Apteryx the squamosal takes no part in the formation of the inner wall of
the skull.
In Apteryx the squamosal has entered into a slightly closer relation with the
alisphenoid and exoccipital than in the forms described above, resembling in this
respect the Neognathe. ‘Thus the anterior border is concave and fits the corresponding
convex border of the alisphenoid; whilst the posterior border, though separated by
a narrow line of cartilage from the exoccipital, yet gives signs of effecting a close union
therewith immediately.
The squamosal appears on the inner surface of the skull as a small triangular plate
above the pro-6tic.
The nasal in Dromeus is holorhinal. The maxillary process is greatly reduced,
especially so in the adult, where it exists only as a vestige. It is produced caudad into
a long slender process resting on the frontal, and projecting beyond the lozenge-shaped
plate of the mesethmoid, to which it binds externally. Its premaxillary or internal
process is of considerable length, extending forwards almost the whole length of the
nasal process of the premaxilla, which protects them from above.
Dinornis is holorhinal.
A skull of Megalaptery«, in the collection of the Hon. Walter Rothschild, retains
sufficient of the traces of sutures in this region to indicate the form of the nasal in the
Dinornithide.
In this skull, it is seen to be relatively much shorter than in Dromeus. Its backward
extension does not pass the level of the horizontal mesethmoid plate and is not so
pointed as in Dromeus. Its maxillary process is wanting. ‘The ossified aliethmoidal
plate projects beyond its lateral border.
In Casuarius the nasal is schizorhinal. ‘The maxillary process is vestigial, and the
premaxillary process long, as in Dromeus. It is peculiar in that the body of the bone
is greatly inflated and takes part in the formation of the casque as already described
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH 2. 203
In Rhea the nasal must be regarded as holorhinal, though the maxillary process is
absent. Inasmuch as this process, when present, serves as the boundary, caudad, of
the anterior nares, its place must be regarded as being filled by a vertical spike of bone
arising from the maxillo-palatine process to the anterior border of the lachrymal. In
Struthio this vertical spike—in Rhea separating the anterior nares from the lachrymo-
nasal fossa—is represented by a ligament which is attached dorsad to the free end of
the maxillary process of the nasal.
In Struthio the nasal is holorhinal (Pl. XLII. fig. 3). Its maxillary process is
well developed, and continued downwards on the maxillo-palatine by ligament.
The Zinamide may be regarded as holorhinal, though the nasal cleft has extended
backwards so far as to lie level with the free end of the nasal process of the premaxilla.
The maxillary process is not wanting asin Rhea, though reduced to a mere vestige.
The nasal cleft is bounded externally by a vertical, bony rod, representing the
maxillary spike found in Rhea. It differs therefrom in being rod-like, very slender,
and is perfectly free. Above, it articulates, by means of a short inturned process, with
the anterior border of the lachrymal and the vestigial maxillary nasal process, whilst
ventrally it articulates with the extreme postero-external border of the maxiilo-palatine
process where it joins with the rod-shaped portion of the maxilla. This vertical rod
was regarded by Kitchen Parker as the maxillary process of the nasal. It is, however,
in all the skulls I have examined, quite distinct from this.
The lachrymal, in Dromeus, sends backwards a very long, curved, rod-like process
to overhang, and thereby greatly increase the size of the orbit (Pl. XLII. fig. 2). It
sends downwards a large antero-posteriorly flattened limb, the outer border of which
is produced still further downwards into a short rod-like spine which articulates with
the quadrato-jugal bar. The body of this ventral process of the lachrymal is perforated
by a large foramen for the passage of the lachrymal duct. The inner border of this
ventral process articulates with the antorbital plate.
In Casuarius the lachrymal differs from that of Dromeus in the greater relative
shortness of its external orbital process, which is scarcely longer than the internal. A
further point of difference lies in the fact that the external orbital process passes
insensibly forwards and downwards into a laterally compressed plate, which, after
forming the boundary caudad of the posterior narial aperture, turns sharply backwards,
and becomes the antero-posteriorly compressed ventral process bounding the lachrymo-
nasai fossa as described in Dromeus. From this it differs in that the laminated portion
is continued downwards to the level of the quadrato-jugal bar, instead of stopping short
of this, and sending downwards thereto a spine from its outer border. The foramen for
the lachrymal duct is smaller than in Dromeus. Internally it articulates with the
antorbital plate.
In Rhea the lachrymal resembles that of Dromeus. ‘The supra-orbital process is
large. The lachrymo-nasal process is also antero-posteriorly compressed, ‘The foramen
204 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
for the lachrymal duct is very large. This lachrymo-nasal process stops short of the
quadrato-jugal bar and is connected therewith by a large subquadrate ossiculum
lachrymo-palatinum. Internally the last eventually fuses with the antorbital plate.
In Struthio the lachrymal resembles that of Rhea. Its supra-orbital process is,
however, relatively shorter and thicker. In two nearly adult skulls (one of which is
from the collection of the Hon. Walter Rothschild) this orbital process supports one
end of a chain of irregularly shaped ossicles which is continued backwards along the
frontal as far as the post-orbital process. Normally this chain of ossicles is so completely
fused as to be indistinguishable from the frontals on the one hand, and the lachrymal
on the other. There is a large supra-orbital fenestra bounded in front by the lachrymal
and behind by the frontal ; externally by a portion of this chain of supra-orbitals.
The lachrymo-nasal process is continued downwards as a sigmoidally curved rod to
terminate on the inner side of the quadrato-jugal bar. On the inner side of this
extremity of the lachrymal there lies a subcylindrical ossiculum lachrymo-palatinum
articulating mesially, in very old specimens, with the antorbital plate. The lachrymal
foramen is represented only by a deep notch.
In Apteryx the lachrymal is vestigial, and takes the form of a small roughly triangular
plate of bone closely applied to the outer surface of the aliethmoid and the maxillary
process of the nasal. It is perforated by the lachrymal foramen.
The premaailla, maxilla, jugal, quadratojugal, vomer, palatine, and pterygoid differ
in no important respects from that of the adults already described (p. 187).
The vomer, pterygoid, and palatine of Apterya demand, however, a further examina-
tion, as these in the adult are too completely fused with the neighbouring bones to
render it possible to make out their boundaries.
The pterygoid is bifid cephalad, being divided into two moderately long, pointed and
widely separated prongs. The inner prong, which constitutes the main body of the
bone, is deeply grooved along its mesial border, producing two shelf-like projections
one above the other: these converge near the tip of the prong to form a slender pointed
style resting in a groove on the outer border of the posterior extremity of the vomer as in
other Palwognathe, and as in the hemipterygoid of Neognathe, e. g. Impennes, Colymbi.
The outer limb is closely applied to the ventral and external aspect of the extreme
hinder end of the maxillo-palatine process, extending as far forwards as the point from
which the maxillary portion of the quadrato-jugal bar is given off. In the cleft between
the outer and inner limbs the palatine is interposed, as will be seen presently.
The palatine is short and broad, with a slightly bifid free extremity. It can only be
seen in its entirety from above. It is closely applied, throughout its entire length, to
a backwardly projecting plate from the mavxillo-palatine process. This it entirely
conceals from above. Its proximal extremity sends off, upward and inward, a flange-
like plate to articulate with, and underlie, the extreme hinder end of the vomer.
Thus this last has its right and left limbs sandwiched, on either side, between the
PHYLOGENY OF THE PALZOGNATH® AND NEOGNATHA, 205
pterygoid above and the palatine below. A portion of this flange-like plate of the
palatine helps to fill up the cleft between the pterygoid fork already described. As
mentioned above, the palatine rests upon a backwardly directed plate from the maxillo-
palatine process, the whole of which it conceals from the dorsal aspect. Ventrally,
this process is found to run along, and below, the external palatine border, and to
terminate near its extreme posterior end. ‘The forward extension of the palatine on
the ventral surface corresponds with the level of the tip of the vomer. This extension
is derived from the ventral surface-layers of the main body, the palatine terminating
abruptly on the dorsal aspect in a V-shaped notch at the level of the middle of the
vomer. ‘This ventral process rests in a groove along the mesial border of the maxillo-
palatine process.
The relations of the vomer have already been more or less indicated. Suffice it to
say, that the free end of each posterior limb is wedged in between the spine-like
extremity of the pterygoid above and the flange-like inward projection from the dorsal
surface of the palatine below.
The palate of Apteryx will be found, on a close comparison with that of the remaining
Paleognathe, to have departed trom the typical Struthious form in the direction of
that taken by Rhea, the relations between palatine and vomer being similar in both.
The torm of the pterygoid differs from that of all other birds, in that it is bifid, or, in
that it sends off a separate limb whose mesial border passes downwards to underlie the
external border of the palatine. ‘This external, palatine limb is elsewhere unrepre-
sented. ‘The internal, vomerine limb may be regarded as a flattened plate folded upon
itself, the edge being inwards. ‘The superior edge, or upper lip, must be regarded as
representing the mesial border of an originally flattened pterygoid such as that of
Dromeus.
The dentary is of great length. In Dromeus, Casuarius, Rhea, Apteryx, and
‘Tinamous it extends backwards to within one-fourth of the whole length of the ramus.
It is relatively less in Struthio. Caudad each ramus divides into dorsal and ventral
limbs, of which the ventral is the larger. In Rhea the dorsal limb is paired.
The splenial is of great length in all, extending forwards to the symphysis of the
jaw, and backwards to within a short distance of the level of the glenoid cavity for the
quadrate. It is rod-like in Struthio, laterally compressed in the remaining forms.
The angular, in all save Apterya, articulates, proximally, with the supra-angular,
being embedded caudad in its external border; distally, it twists downwards so as to
form the ventral border of the ramus. In Casuarius and Dromeus its extreme posterior
and inner border comes into contact with the articular. In Rhea and Struthio, it falls
considerably short of this.
In Apteryx, the angular articulates proximally with the ventral border of the
articular, which is hollowed out to receive it. The supra-angular rests upon its
dorsal border.
vou. xv.—Part vy. No. 13.—December, 1900. 26
206 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The supra-angular, in all, articulates proximally with the articular. Its dorsal
border is received between the right and left moieties of the superior limb of the
dentary. Its ventral border descends caudad to the level of the inferior border
of the angular; distad it rises gradually so as eventually to cross the dorsal border of
the same.
The coronary is in all a short, slender, and rod-like bone. In many Neognathe it
is often of considerable size and laterally compressed.
In all the Palwognathe it appears to extend backwards so as to embrace the
inner border of the articular and to form the internal boundary of the articular surface
for the quadrate. In Rhea and Apterys it is very feebly developed distad, and may be
described as vestigial. In Casuarius and Dromeus it is much larger. Struthio is
intermediate. In no case does it extend forwards beyond the posterior third of the
ramus.
Such characters of the skull as are of use for systematic purposes will be found in
the ‘ Keys” at the end of this paper.
More material is needed before these can be made complete, since a “ Key” to the
species has not been possible in the case of Caswarius. Whether the characters
considered to be specific in the case of the other genera will stand the test of time
remains to be seen. In several instances I have been obliged to rely upon the characters
of a single skull, as in the cases of the skulls of Khea macrorhyncha, for instance.
I feel justified in reviving Dr. Sclater’s Struthio meridionalis [go| on account of the
many marked peculiarities which the skull and the trunk of this skeleton presents.
The work of distinguishing the skulls of the four species of Apteryx has been a very
difficult one. Indeed, as Mr. Beddard has already remarked [8], “the very slightest
differences, apart from those of size, distinguish the Apteryges.”
Some Pornts oF SIGNIFICANCE IN THE StRUTHIOUS PALATE,
The following points appear to be worthy of special consideration and may fitly be
discussed here.
In comparison with the palatines of the Neognathe, it will be remarked that in
Rhea, as in the other Palewognathe, the palatines are widely separated one from
another by the laterally expanded feet of the vomer. In the Neognathe the palatines
meet in the middle line, thrusting the vomer forwards, so that it now appears to
articulate not with the pterygoid but with the palatines. In hea and Crypturi we
have probably a clue to the way in which this has come about. ‘Turning once again
to the skull of Dromeus, we notice that the palatines are only loosely attached to the
outer border of the vomerine feet, whilst in Rhea (Pl. XLII. fig. 5) the palatines have
moved inwards so as to overlap these feet and approach one another mesially. Their
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 207
relation to the pterygoid has also changed, for they no longer are connected with its
outer border but by this same inward movement have come to underlie it.
In the Neognathe, the inward movement has attained its maximum, the palatines
meeting one another mesially, as we have just remarked, thrusting the vomer forwards
in so doing. As a matter of fact, however, as we have already shown in earlier papers
[82], this is not altogether the case. In the young bird the pterygoid extends
forwards in a spike-like form, much as in Rhea, so as to articulate with the vomer,
though but by the slightest contact. ‘The palatines have succeeded in moving inwards
beneath these anterior pterygoid ends so as to all but entirely sever the original
relations between them and the vomer.
Later in life (soon after hatching) the severance is complete. At this stage, the
anterior end of the pterygoid fractures at a point corresponding with the free end of
the palatine. The fracture later becomes a true joint, and the anterior end of the
pterygoid resting upon the palatine gradually merges with this bone so as to obliterate
all traces of its original existence. Thus the free pterygoid of the Neognathw is a
secondary feature, the palato-pterygoid connection in the late embryo not differing
materially from that of the Palwognathew. Further, the apparent isolation of the
vomer from the pterygoid in the Neognathe is seen to be a coenogenetic character,
so that the palate of this group is brought into close relation with that of the
Paleognathe.
A further point of interest in this comparison between the Palzo- and Neognathine
skull is the change which the vomer in the latter has undergone in relation to the
parasphenoidal rostrum, a change which indicates a shortening both of vomer and
rostrum.
The vomer in the Weognathe rarely extends backwards beyond the base of the
antorbital plate, in the Palwognathe it may reach nearly as far as the basipterygoid
processes. This is an undoubted proof of the shortening of the vomer.
That the rostrum has also undergone a considerable shortening is shown by the fact
that in the Palwognathw it extends forwards for a very considerable distance beyond
the level of the lachrymo-nasal fossa, in the Meognathew it commonly ends in the
region of the antorbital plate. :
Yet other evidences of shifting and modification of the dromeognathous palate
reveal themselves in the Meognathw when we come to closely compare them, and
whilst these show how closely the two groups are related they show still more the
lower grade of type persistent in the Palwognathe.
In the Palwognathe the free ends of the basipterygoid processes articulate with
the extreme proximal end of the pterygoid quite close to the articulation with the
quadrate. In Neognathe these processes, when present, have shifted forwards on to
the rostrum, so as to articulate with the middle of the pterygoid.
The forward shifting of the Neognathine vomer, which we have already noticed—a
262
208 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
shifting which brings its proximal end more or less immediately under the antorbital
plate—appears to have been accompanied by a similar shifting of the palatines, since
these no longer are connected with the main body of the pterygoid bone but with its
distal end. ‘This, as we have already shown, ultimately fractures and fuses with the
palatine, a joint forming at the line of fracture. This connection of the palatine and
pterygoid by means of a joint is a point of great difference between Paleo- and
Neognathine skulls.
In the Tinamidw we have an intermediate stage between the Palwo- and the Neo-
gnathe, The vomer is undoubtedly relatively shorter posteriorly than in Rhea, its
free end lying midway between the level of the antorbital plate and the basipterygoid
processes. The pterygoid has increased in length, so that the vomer and palatine
articulate with its distal extremity only. The quadrato-jugal fossa has also increased
in length relatively, extending forwards now beyond the level of the antorbital plate
as far as the vestigial maxillo-nasal process.
A careful study of these points will greatly facilitate the conviction that the
egithognathous and schizognathous skulls are but modifications of the dromeognathous
type. The desmognathous is a further modification of the schizognathous palate.
The single-headed otic process of the quadrate in the Palwognathe, upon which
so much stress has hitherto been laid, appears to have less importance than the points
to which attention has just been drawn, for in Apéeryx the otic process is two-headed,
as in Neognathe.
THe VERTEBRAL CoLuMN.
The memoirs of Owen, Mivart, and T. J. Parker on the vertebral column of the
struthious skeleton render it quite unnecessary to do more than briefly comment
thereon here.
a. The Presynsacral Vertebre.
All the presynsacral vertebre of the Palwognathew are heterocelous, and all of
this region are free except in the Yinamide, in which certain of the thoracic
vertebree fuse.
The vertebra of Dromcus are the least specialized in type; those of Caswarius are
very similar.
Both in Dromeus and Casuarius the cervical vertebre are conspicuously shortened
antero-posteriorly. In Cuswarius the neural spine, traced from the head backwards,
undergoes considerable change of form. Anteriorly it is little more then a median
tubercle rising from the centre of a flat neural plate. At about the 6th vertebra it
sends backwards a pair of low ridges which terminate in a pair of hyperapophyses.
At about the 10th vertebra, the spine has exchanged its <-shape for a transversely,
crescentic form. ‘This, for the next 4—5 vertebra, becomes broken across in the middle
PHYLOGENY OF THE PALAIOGNATHA AND NEOGNATH A. 209
line to form two high curved ridges sloping downwards and backwards to terminate in
metapophyses. At about the 15th vertebra the two ridges again coalesce and form a
very high vertical transverse neural spine. From the 18th backwards this gradually
becomes more and more laterally compressed, so as to pass insensibly into the typical
neural spines of the thoracic vertebra. Every neural spine, from that of the axis
backwards, bears a fossa at its base, both anteriorly and posteriorly. The latter is the
deeper. The vertebre from the middle of the neck backwards have these fossz of
very considerable size. They lodge a ligament.
From 6-10 in C. casuarius and C. ¢. australis the diapophysis sends back a bar of
bone to the hyperapophysis.
The vertebrie are all pneumatic. In the hinder cervicals there is a large pneumatic
aperture dorsad of the interzygapophysial ridge. In the thoracic there are several
very large ones—one below the transverse process, oue between the transverse process
and the postzygapophysis, and one dorsad, lying between the anterior zygapophysis
and the base of the neural spine. This last is represented by a deep fossa in Dromeus.
The aperture ventrad of the anterior zygapophysis is feebly developed or wanting in
Dromeus.
The cervical ribs (pleurosteites) and hypapophyses resemble those of Dromeus.
In Dromeus the vertebre are less specialized than in Casuarius. The high
transversely expanded neural spines are wanting, though the ligamental neural fosse,
especially that caudad of the neural spine, are very deep. ‘The pneumatic fossa in, or
above, the interzygapophysial ridge is very deep, as also is that lying at the base and
in front of the neural spine. The sides of the fosse, moreover, are smooth, not obscured
by cancellated tissue as in Casuarius.
In Struthio the centra of the vertebree are relatively much longer than in Dromeus or
Casuarius. The neural spines of the anterior cervicals are long, low, and rise to form
a sharp median ridge. The ligamental fossee are narrow grooves channelled out of
this ridge. The posterior cervicals have the neural spines wider and shorter, antero-
posteriorly, and they are deeply hollowed for the ligament.
The cervical ribs, as in Casuarius and Dromeus, are long, but more slender than in
these. As in Dromeus they fuse with a plate of bone depending from the diapophysis
and a lateral parapophysial outgrowth from the anterior end of the centrum below
the prezygapophysis. The presence of this rib serves to enclose a canal for the
vertebral artery. One great point of difference between this region of the vertebra and
that in Drom@us and Casuarius lies in the fact that in Struthio the lamella depending
from the diapophysis, and with which the rib articulates, is continued backwards along
the centrum for a considerable distance, forming an extensive and tunnel-like passage
for the artery.
The pneumatic apertures are not so conspicuous as in Dromeus. There is no
aperture in the interzygapophysial ridge of the anterior ceryicals as in Dromeus. In,
210 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
the hinder cervical and thoracic vertebre there is a large aperture beneath the
diapophysis, opening in the roof of the tunnel for the vertebral artery, or in the thoracic
vertebra below the prezygapophysis and between the dia- and parapophysial articula-
tions of the tuberculum and capitulum. The deep fossa at the base of the anterior
border of the neural spine of the hinder thoracic vertebre is here yery shallow.
In Rhea the vertebree have departed still more from the normal type. The cervicals
are very long and cylindrical. Neural spines are present only on the 2nd to 5th
vertebre. ‘The free hinder borders of the postzygapophyses of the 6th vertebra are
received into a pair of shallow pits on the 7th, lying mesio-caudad of the prezyga-
pophyses. Cervical ribs are only feebly developed. From the 2nd to 11th vertebre
they are represented only by short styliform processes; from this vertebra backwards
they are represented only by the extreme anterior articular end of the rib which serves
to form the band-shaped external wall of the vertebrarterial canal. As in Struthio, the
vertebrarterial canal is roofed by a backward extension of the lamelliform plate
depending from the diapophysis and serving as an articulation for the cervical rib.
This plate is described and figured as the “rudimentary rib” by Mivart [63]:
doubtless by a slip.
In Apterya the form of the atlas and axis vertebre bears some resemblance to
Dromeus. The rest of its vertebra differ conspicuously from those of the forms
herewith associated. The neural spines never bifurcate. On the anterior vertebre
they form compressed blades from the 8th to the 12th; they are columnar and
vertical in form and central in position with regard to the neural lamine. The
diapophyses of the 6th and 7th vertebra project outwards and backwards beyond
the level of the prezygapophysis; from the 7th to the 13th backwards and down-
wards ; from this vertebra caudad they project directly outwards, and merge gradually
into the transverse processes of the thoracic vertebra. The neural spines of the
thoracic vertebre are equal in size, of great breadth antero-posteriorly, and interlock
along the dorsal ridge by anterior and posterior bifurcations as in many Veognathe.
There are no pneumatic apertures in either cervical or thoracic vertebra. Cervical
ribs are vestigial.
In Crypturi the thoracic vertebrae are anchylosed and bear hypapophyses. In the
Paleognathe these occur only on the posterior cervicals (cervico-thoracic). The
penultimate is free, the ultimate fused with the synsacrum. The interzygapophysial
ridge is perforated by pneumatic apertures. In the cervical vertebre pneumatic
apertures open into the roof of the vertebrarterial canal.
The diapophyses of the cervicals project forwards beyond the prezygapophyses.
The neural arch is constricted in the middle, immediately behind the prezygapophyses.
The vertebree of Dinornis resemble those of Caswarius. The fosse lying beneath
the transverse process of the thoracic vertebra, absent in Dromeus, were much larger
relatively than in Caswarius. The cervicals resembled those of some species of
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 211
Casuarius in having a bony bar running from the diapophysis backwards to the
hyperapophysis.
The thoracic vertebre call for no special remark here. They have already been
minutely and carefully described by Mivart.
The vertebra known as the cervico-thoracic should probably be regarded as true
thoracic vertebre, the ribs of which have lost their sternal segments through the
backward shifting of the sternum, This view has already been promulgated by the
late T. J. Parker. Writing of Apteryx, he says: “The occasional absence of
the cervico-thoracic uncinates, taken in connection with the vestigial condition of the
ribs in one case and with the fact that in the same skeleton the first thoracic rib
of the right side terminates ventrad in a blunt free end and has no sternal portion,
seems to point to an inclusion of anterior thoracic vertebre in the cervical region by
atrophy of their ribs.”
b. The Synsacral Vertebre. (Pl. XLIV. fig. 5.)
In Casuarius there are 8 (eight) pre- and 8 (eight) post-sacral vertebrae, with the
two true sacrals numbering eighteen synsacrals.
Of these, two are thoracic. ‘The 1st lumbar has large parapophysial processes; the
2nd to 3rd of this series are much smaller; the 4th and 5th become larger and
abut, the 4th against the pectineal process and the 5th against the ischium. Behind
these follow two lumbo-sacral. The true sacrals are closely crowded and have slender
parapophysial processes, which abut against the ilium and ischium immediately behind
the acetabulum. Behind these in C. dicarunculatus lie two pseudo-sacrals with large
parapophysial processes abutting against the ilium. But for the sacral plexus they
might readily be mistaken for true sacrals. They are really the first two caudals, behind
which follow six fused caudals with diminishing and backwardly directed transverse
processes. In other Cassowaries there is but one pseudo-sacral. ‘The sacral vertebrae
do not bear diapophyses.
In Dromeus the number of synsacral vertebrae is the same as in Casuwarius. The
6th lumbar, however, does not send out a parapophysis to abut against the pectineal
process.
The neural spines of the synsacral vertebre are all highly pneumatic, and made up
of loose cancellated tissue. ‘They remain separate for some time after hatching, but
in the adult are fused into one mass, all traces of separate vertebra being lost. In the
young bird the pre-sacral vertebre are just distinguishable between the closely
approximated pre-acetabular regions of the ilium, The post-sacral have the neural
spines laterally expanded so as to keep the post-acetabular ilium moderately widely
separated, so that between each vertebra is a deep loculus (Pl. XLIV. fig. 5).
In the adult the whole becomes roofed over by a thin bony plate extending across
from the post-acetabular ilium of one side to that of the other.
212 MR. W. P. PYCRAFL ON THE MORPHOLOGY AND
Seen dorsally in the half or even three-quarter grown birds the neural spines from
above the acetabulum backwards for some considerable distance are found to take
the form of vertical transverse plates, forming a series of loculi between the post-ilia.
‘Though further back in the series the transverse plates become columnar, the loculi
remain. In the adult these loculi are completely covered in by a thin sheet of bone,
formed by the fusion of a series of thin plates capping the neural spines.
In the embryo Dromeus (Pl. XLIV. fig. 6) it is worth noting that the lofty
transversely expanded neural spines of the sacral and anterior post-sacral vertebrae of
the adult are represented only by low cartilaginous ridges, and bounded on either side
by the free edge of the dorsal border of the ilium. In the dried skeleton the carti-
laginous portion shrinks up, revealing (1) a pair of ossified neuroids, which fail to
meet one another in the middle line, and (2) a low osseous ridge marking the
foundation of the similar cartilaginous ridges already described. The increase in
height of the neural spines is followed by a corresponding increase in depth of the
ilium (p. 227).
The centra of these synsacral vertebree in this early stage are, in the dried skeleton,
represented only by a flat bony plate, so that a single vertebra, with its neural arches,
is stirrup-shaped. ‘The thinness of the centrum makes the neural space of a relatively
much greater size than in the adult. ‘The nerve-apertures are single and not paired
as in the adult. This 1s well shown in Pl. XLIV. figs. 5-6,
In Struthio there are ten pre-sacral and eight post-sacral vertebre.
The series commences with the last two thoracic vertebre. Then follow seven
lumbar vertebre. The parapophysial elements of these are all short and thick.
The diapophyses of the lumbar vertebra from the 4th backwards extend upwards
on to the inner side of the ilium as forwardly directed spikes. Short and triangular
on the 4th, they gradually increase in length up to the 8th, where they form long
rod-shaped bars of bone. On the 10th, this process is short and rod-shaped.
The lumbo-sacral vertebra are possibly represented by a single vertebra which bears
vestigial parapophyses. In one skeleton in the Museum collection the lumbo-sacrals
were certainly unrepresented. ‘The last lumbar bore distinct parapophysial elements,
which on the left side extended backwards to join the first sacral rib.
‘The sacral vertebree are two in number and have large diapophyses. They lie more
caudad than those of Caswarius, resting against the hinder border of the downward
ischiadic or post-acetabular process of the ilium, and therefore conspicuously caudad
vf the acetabulum. In Caswarius the sacral ribs lie immediately behind the
acetabulum. ‘The outer free ends of the sacral ribs give rise to a large swollen process
from the dorsal surface, which extends upwards and backwards to reach the diapophysis
of the 2nd sacral, sometimes extending forward to that of the Ist. In one specimen
in the Museum collection the lumbo-sacral has developed a parapophysial process on
the left side.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. ies
Sometimes the former fuse with
The Ist caudal bears both para- and diapophyses.
those of the sacral. Behind this lie seven caudals without parapophyses, before the first
free caudal is encountered. The neural spines of these are not transversely expanded
as in Casuarius.
In Fhea there are ten pre- and five post-sacral vertebrae.
The pre-sacral comprise two thoracic, four lumbar, and four lumbo-sacral; the last.
of these may develop parapophysial processes.
The sacral ribs fuse with the ilium more dorsad than in Struthio, Caswarius, or
Dromeus. In the last three they abut more or less extensively against the post-acetabular
ascending process of the ischium. In Rhea they rise dorsad, so as to reach the level
of the centre of the acetabular are.
The Ist sacral is intermediate in position, with regard to the acetabulum, between
Casuarius and Struthio. In the latter, it will be remembered, it lies considerably behind
the acetabulum, in Caswarius level with its hinder border.
The sacral ribs ave very short and almost indistinguishably fused with their
diapophyses. In the possession of diapophyses these vertebre differ from those of
Casuarius and Dromeus, and resemble Struthio, in which, however, they are much
more extensively developed.
The first post-sacral, or lst caudal, bears both para- and diapophysial processes,
but they are almost indistinguishably fused. The 2nd candal bears diapophyses only.
The 3rd to 5th caudals are somewhat remarkable in that they are represented only by
atrophied vertebre. The centra of these are reduced to thin delicate rods, supporting
slender diapopbyses and feeble neural spines. ‘The 6th vertebra resembles the oth,
but has a stronger neural spine. It is the first vertebra to escape from the clasp of
the post-acetabular ilium. The 7th has a much stronger neural spine and slightly
thicker centrum, but like the 6th has lost its diapophyses. The 8th and 9th have
inflated cancellous centra and short neural spines; they fuse by their ventral aspects
with the ischia. The 10th is the first free caudal, of which there are eight in all:
these have short thick centra and short neural spines.
In Apteryx there are nine pre- and five or six post-sacrals, with the true sacrals
sixteen in all.
The pre-sacrals represent two thoracic, four lumbar, and three lumbo-sacral. The
first post-sacral (1st caudal) bears short para- and diapophysial processes. The post-sacral
are chiefly remarkable for the great lateral compression which they have undergone.
The synsacrum of the Dinornithide differs in many respects from that of all the
It resembles in many ways that of Dromeus and Caswarius
The pre-sacrals are apparently
There is an
other Paleognathe.
on the one hand, and that of Struthio on the other.
eleven in number, viz., two thoracic, six lumbar, and three lumbo-sacral.
indication of a fourth lumbo-sacral bearing parapophysial processes, which have almost
completely fused with those of the 1st sacral. There are five post-sacrals.
VOL. XV.—PART v. No. 14.—December, 1900. 2H
214 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The pre-sacrals resemble those of Dromeus and Casuarius in that they descend
below the level of the ventral border of the pre-ilium, but not greatly so. In Struthio
this feature is much more marked, the whole centrum and base of the neural spine
being exposed in side view. ‘The parapophysial processes of the first caudal are
directed backwards instead of forwards. From this cause the true sacral vertebre
are completely isolated.
In Struthio, Casuarius, Dromeus, and Apteryx, it will be remembered, the para
pophysial process of the first or even first and second caudals may combine with
those of the true sacrals to form a sacral mass, the true nature of which is difficult
to make out in the adult. The post-sacrals (caudals) resemble those of Struthio
rather than any other Ratite in the great development of the diapophysial process,
which extends upwards, outwards, and backwards to separate the post-acetabular
ium.
In Dinornis, however, this separation of the post-acetabular ilia is carried to a
greater length than in any other Ratite. The neural spines of the Dinornithide are
peculiar in that they combine dorsally to form a broad lozenge-shaped superficial plate
of bone which serves as a cover to the huge fossa that otherwise exists between the
upwardly directed transverse processes and the post-ilia.
The synsacrum of the Crypturi differs from that of all the other forms herein
described. Moreover, in the points wherein it differs from these it resembles the
Neognathe.
One of the most noticeable differences is the fact that the post-sacrals have lost their
neura] spines. In all the flightless Palwognathe these are of great length. ‘The
diapophyses of the post-sacral region rise from the top of the centrum, at the base of
a long, faint, median ridge marking the position of the neural spines in other forms.
These diapophyses develop thin plates of bone which fuse one with another to form a
lozenge-shaped plate, protecting the inter-diapophysial fosse, as is done by the similar
plate from the neural spines in Dinornis.
‘The synsacrum of the Tinamous includes one thoracic, four lumbar, five lumbo-sacral,
and five post-sacral. There are no parapophysial processes abutting against the
pectineal process as in the flightless Palwognathe. ‘This process, it should be
mentioned, has shifted further backward relatively to the sacral vertebra, so that a
line drawn across from the base of the anterior concave border of the one would pass
immediately in front of the first sacral.
The sacral ribs are very long, and articulate with the ilium immediately behind the
acetabulum by a long vertical roughened surface. The rib of the second sacral is
vestigial or wanting.
The free caudal vertebree in the Palwognathe do not exceed ten in number. Some-
times the 9th and 10th are more or less fused.
In Struthio the 10th is laterally compressed to form a kind of pygostyle.
PHYLOGENY OF.THE PALZOGNATHA AND NEOGNATHA, 215
In Rhea there are but seven free caudals. This is due to the great backward extension
of the ischia. Counting from the free end of the post-acetabular ilium backwards, as
in other forms, there are eleven post-iliac caudals.
Riss.
The anterior cervical ribs (pleurosteites) have already been described.
It is probable that the free cervical ribs (cervico-dorsal, Mivart) and the vertebre
to which they articulate really belong to the thoracic series. ‘This is indicated by the
fact that these ribs have lost their connection with the sternum, and have lost their
sternal ribs through the backward shifting of the sternal plate.
In Casuarius and Dromewus there may be three or four free cervicals. In Struthio
and Rhea, two. In Apterya one, which bears a large uncinate (in A. oweni there is
a shorter and more style-like rib anterior to this and wanting the uncinate).
In the Crypturi there are two free cervicals.
The thoracic ribs in Casuarius and Dromeus agree with Struthio, and differ from
Rhea, in the fewness and slight development of the uncinates.
In Casuarius the Ist five pairs of thoracic ribs articulate with the sternum by means
of sternal ribs. The 6th and 7th are long, but have no sternal segments. The 7th
pair are more or less overlapped by the pre-ilium. The 8th pair are very short and
underlie the pre-ilium. The 9th pair are represented by a scale-like vestige repre-
senting little more than the tuberculum and capitulum, which in the young bird can
still be made out (e. g. 1874 a Mus. Coll.) closely applied to the pre-ilium.
In C. bennetti there are but three free cervical ribs and only four pair of thoracic ribs
articulating with the sternum by means of sternal segments. ‘The 5th pair have very
long sternal segments, but they fail to reach the sternum, We may gather from this,
however, that they have not long lost their connection therewith.
The uncinates of the ribs of Caswariws are never large. Generally only two or three
appear to be present, and these may anchylose with their respective ribs. In a half-
grown C salvadorii I find five uncinates. These have been probably lost in maceration
in most of the skeletons which I have examined.
In Dromeus there are two free cervicals; the second is very long, and almost
articulates with the anterior lateral process of the sternum. Following these come
four pairs of thoracic ribs connected with the sternum by sternal ribs. These are
succeeded by a pair (7th) with very long sternal segments, but which fail to reach the
sternum. ‘The next two pairs have no sternal segments, and are overlapped by the
pre-ilium.
In Struthio there are two free cervicals and five pairs of thoracic ribs articulating
with the sternum by means of sternal ribs. The pair next behind these has long
sternal segments which do not reach the sternum. Jn a mounted specimen in the
collection of S. camelus this 6th sternal segment actually articulates with the sternum.
2H 2
216 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The following pair (7th thoracic) are long, but without sternal segments; the 8th
pair are short and fuse with the pre-ilium.
In Rhea there are two free cervicals, three pairs of thoracic articulating with the
sternum by sternal ribs, and provided with very long uncinates. ‘The 4th and 5th
pairs are long, but have no sternal segments. The 6th pair are very short and fuse
with the pre-ilium. ;
Amongst Dinornithide there appears to be some variation.
In Dinornis maximus there are three free cervicals, two thoracic connected with the
sternum by sternal segments, one long free rib, and three free ribs behind this
overlapped by the pre-ilium.
In Pachyornis there seems to have been but one free cervical or cervico-thoracic,
three thoracic articulating with the sternum, four long free ribs, and short free ribs,
these last two being overlapped by the ilium. The uncinates were very long.
In Apterya the ribs are remarkable for the great breadth and flatness of their
vertebral segments. ‘There may be one or two pairs of cervico-thoracic ; there are four
thoracic articulating with the sternum (in a skeleton of A. australis I found five on
one side, and in a skeleton of A. oweni, both in the Rothschild Coll., I found but
three thoracic vertebra connected with the sternum) and four pairs of free ribs behind
these, all of which are overlapped by the pre-ilium. ‘The last pair are often
exceedingly short. Six pairs of ribs bear uncinates, the series commencing in the
last cervico-thoracic. The uncinates are long and broad, extending backwards to the
hinder border of the rib next behind.
In the ribs of all the Palwognathe, save Apterya and some Tinamous, there is a
large pneumatic aperture lying in the cleft between the tuberculum and capitulum.
THE STERNUM AND PEcTORAL ARCH.
a. The Sternum.
As with the other portions of the skeleton so far examined, so with the sternum,
Dromeus possesses the most generalized type.
The sternum of the young Vromeus and that of the young Caswarius are very
similar in appearance; in the adult they are readily distinguishable.
In the adult Dromeus the anterior lateral processes are very long, slender, and
directed upwards and backwards. There is no spina externa nor interna, but a deep
emargination in their place. The ventral lips of the coracoid grooves are well
developed. ‘The articular surfaces for the coracoids are short, not exceeding in
length the distance between the articulation for the first and third sternal ribs.
These grooves differ from those of all the other Palwognathe in that they overlap,
causing the bases of the coracoids to cross one another in the middle line as in many
Carinate.
PHYLOGENY OF THE PALZOGNATHEZ AND NEOGNATH. 217
Fig. 4.
A.—Ventral aspect of the sternum of Dromceus nove-hollandie, to show the form and position of the coraciod
grooves and anterior lateral processes.
B.—Ventral aspect of the sternum of Dromeus ater. Note the degenerate condition of the coracoid grooves
and the well-marked protuberantia sterni.
C.—Ventral aspect of the sternum of Rhea macrorhyncha for comparison with that of Dromeus. Note the
difference in the position of the coracoid grooves, of the articulation for the sternal ribs, the large
protuberantia sterni, and the thin places indicated by shading denoting the position of former
fenestre or fissures.
D,.—Ventral aspect of the sternum of an embryo Apteryx after Parker, to show the possible vestigial keel.
a.s. = articulation for sternal rib.
c.g. = coracoid groove.
a.l.p. = anterior external process.
ps. = protuberantia sterni.
p.lp. = posterior lateral process.
k, = keel.
v.c.g. = ventral lip of coracoid groove.
218 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
The sternum of Dromeus ater is remarkable, possessing as it does characters in
common with both Casuarius and Rhea, but is at the same time Dromeine.
As in D. nove-hollandie, the anterior lateral processes are conspicuously large and
have the same form. The outline of the free edge of the corpus sterni from the
articulation of the last thoracic rib backwards is the same. ‘The coracoid grooves
are obsolete. That of the right side is represented by a portion of the inner angle of
the lower lip of the groove. ‘This, as is seen in the figure (4 B), persists in the form
of a median asymmetrical tubercle. The obsolescence of these grooves indicates that
the pectoral arch and wing were much more degenerate than in Dromeus nove-
hollandie or D. ivroratus. The corpus sterni differs from that of D. nove-hollandie in
the relatively much greater development of the protuberantia sterni, which resembles
that of Rhea, and in the relatively narrower width across the region of the incipient
posterior lateral processes. Four ribs articulate with the sternum, in D. nove-
hollandie only three.
The following measurements in millimetres are of interest :—
D. nove-hollandie. D. ater.
Greatestjlen thyme amie ee eer 160 95
Greatest width (across ant. lat. proc.) . . 140 90
Least width (across post. lat. proc.). . . 105 55
It should be noted that the metasternum of D. afer bears traces of fracture,
indicating a somewhat greater length than the above measurements. ‘The breadth of
the sternum nearly equals its length. In Caswarius it is much longer than broad.
The articular surfaces for the sternal ribs are divided one from another by pneumatic
apertures forming pockets of great depth, extending from without inwards towards
the middle line. ‘The hinder border, both in Dromeus and Casuarius, is shield-shaped.
No external lateral processes are present. The sternal plate is of great thickness and
highly pneumatic.
In the young Dromeus the anterior lateral processes are very short, and the ventral
margins of the coracoid grooves are almost confluent and not separated by a deep
median notch.
In Casuarius the sternum is relatively much longer than in Dromeus, the increase in
length being most noticeable from the articulation for the last thoracic rib backwards.
The coracoid grooves are much longer, and the anterior lateral process is almost
obsolete. In the middle line between the ventral lips of the coracoid groove in
Casuarius there opens forward a large pneumatic aperture; this runs backwards into
the body of the bone for some considerable distance, penetrating a low median ridge
or protuberance resembling a vestigial keel. This is wanting in CO. unappendiculatus,
a shallow groove taking its place, the sides of which further back become conspicuously
raised to form a pair of long low ridges.
PHYLOGENY OF THE PALZAOGNATHA AND NEOGNATHA, 219
The sternal plate of Caswarius is of great thickness and highly pneumatic.
Such specific differences as I have been able to find will appear in the “ Key.”
The sternum of Rhea resembles that of Dromeus and Casuarius in several respects,
but is yet very readily distinguishable from either.
The coracoid groove, though shallow, is sharply defined. The outer and inner lips,
though but feebly developed, are subequal in length. The groove of the left side is
separated from its fellow of the right by a smooth-lipped and gently hollowed plate
of bone representing the anterior border of the sternum, equal in length to the groove
itself. ‘The anterior lateral processes are large.
The articular surfaces for the ribs are closely crowded, and follow immediately
behind the anterior lateral process. Thus the distance from behind the articular
surface for the last rib to the hinder border is more than twice the distance of the
space separating the coracoid grooves; or, in other words, the articular surfaces of the
ribs occupy only one-sixth of the whole lateral sternal border, in Caswarius it occupies
about one-third.
The body of the sternum is notched posteriorly and tapers from before backwards.
It presents a deep basin-like hollow dorsad and is produced ventrad into a long,
swollen, ridge-like prominence. ‘The corpus sterni is much thinner than in Dromeus
or Casuarius.
The articular surfaces for the sternal ribs are each more or less distinctly divided
into two, the outer being the larger. The pocket-like interarticular spaces (incisure
intercostales) are of great depth.
Large pneumatic apertures open below the base of the anterior lateral process. For
further remarks on the sternum of /hea, see p. 224.
In Struthio the sternum is roughly about as broad as long. It resembles that of
Dromeus in some respects and hea in others. The form and position of its coracoid
grooves resemble that of Dromeus. ‘The ventral lip, however, is wider than the dorsal.
Both dorsal and ventral lips converge mesiad. The right and left grooves are separated
by a slight hollowing out of the anterior sternal border. The anterior lateral processes
are moderately well developed and project outward and forward.
The articular surfaces for the sternal ribs are of great width and occupy nearly the
whole lateral border of the sternum, thus differing from both Dromeus and Rhea.
The pocket-like pneumatic incisure intercostales are relatively less deep than in the
foregoing forms. The median portion of the bar serving for the articulation of the
ribs has lost its glenoid surface, which is preserved only at either end. In this point
Struthio and Rhea agree.
There are large posterior lateral and in some sterna incipient median lateral processes.
The ventral prominence of the sternum lies beliind the centre of the sternal plate,
and is broader and more flattened than in Ahea; in which, by the way, it lies in the
middle of the sternal plate.
220 ; MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Apteryx, in the form and position of its coracoid grooves, resembles Rhea more
closely than any other group. Like Rhea they are widely separated by a deeply
concave median portion of the anterior border of the sternal plate. The groove lies at
the base of the anterior lateral process. It is very short, and has a conspicuously
projecting lower lip.
The sternal plate differs from that of all the foregoing in being non-pneumatic.
Posterior lateral processes are well marked, and
Its breadth exceeds its length.
The articular
divided one from another and a median process by a pair of notches.
surfaces for the ribs, as in Struthio, extend along the greater part of the lateral
sternal wall.
Occasionally, ¢.g.,in an A. australis described by Owen, the sternum was fensterated.
Beddard describes a pair in A. australis mantelli. In another of this species were a
pair “ of pin-hole size.”
Parker directs attention to a distinct keel on the sterna of A. australis mantelli.
As Beddard remarks, though not easy to see, this is readily felt. He found traces in
A. haastii, and I think to these we might add A. oweni. <A similar keel, and much
more easily seen, often occurs on the sterna of Struthio.
The researches of T. J. Parker and Beddard, which I am able to confirm, show that
no really reliable specific characters can be obtained from the sternum of Apéteryz.
Roughly, as Parker and Beddard point out, “in A. australis the length of the
corpus sterni appears to be constantly more than half its breadth.”
In A. australis mantelli “ the length of the corpus sterni is—often considerably—
less than half its breadth.”
In A. oweni “the length of the body is less than half its breadth.”
A. haastii, according to Beddard, belongs to the same category as the last two.
The emargination of the anterior border of the sternal plate and the length of the
metasternal and posterior lateral processes vary so much that they cannot be relied
upon as specific characters.
The anterior emargination of the sternum seems to offer at least one more or less
constant character. Thus, both Parker and Beddard agree that this is usually a
semicircular incision, but that A. australis mantelli may be distinguished “ from
A. australis by the greater depth of the emargination in the former subspecies.”
Mr. Beddard finds at least one exception to this, in A. australis mantelli. Parker
points out that “in 4. oweni.. . . the emargination of the anterior border is slightly
sinuous, each side presenting a sigmoid curvature.” There would appear to be no
exceptions to this rule.
The sternum of Dinornithide more closely resembles that of Apteryx than of any
other form herein described. From this it differs in the following points :—That
portion of the anterior border of the sternal plate which separates the coracoid grooves
in Apterya is in the present group perfectly straight instead of hollow. Furthermore,
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH #%. 221
this region of the sternal plate is much thickened to form a prominent lip, looking
upwards. ‘The coracoid grooves are exceedingly shallow and only just visible. As in
Apterya, these grooves lie far apart, at the antero-lateral angle of the sternal plate.
The anterior lateral processes are short; the articular surfaces for the sternal ribs lie
crowded together immediately behind the anterior lateral process, as in Rhea. The
posterior lateral processes are of great length; the metasternum is often of great
length. The form of this and of the posterior lateral processes varies much in the
different groups into which the Dinornithes are divided. ‘The length of the meta-
sternum in Apterya does not extend beyond the posterior lateral processes.
In Aipyornis the sternum resembles that of Apteryx more nearly than any other
Paleognathe. As in Apteryx, the anterior border of the sternal plate is deeply
notched, the coracoid grooves are widely separated and very similar in form. The rib-
facets are wide and well spaced along the lateral border as in Apteryx, not crowded
together as in Rhea or Dinornis. There are short but well-marked posterior lateral
processes, but there is no metasternal element. As Andrews has pointed out in a
recent and admirable paper on the skeleton of this bird, the sternum of Apyornis
represents an embryonic stage.
As Mr. Andrews has pointed out [4], the ‘sternum of Apyornis consists of the two
primitive costo-sternal elements only, and in this respect corresponds to an embryonic
stage in the development of the sternum in the recent Ratitee.” More especiaily does
this seem to apply, he goes on to point out, to the embryonic sternum of Apterya.
The sternum of the Crypturi differs from that of the flightless Palwognathe
chiefly in the possession of a large keel. In general form the corpus sterni recalls that
of Anomalopteryx. This is chiefly due to the large posterior lateral processes, which
in Crypturi arise very far forwards. ‘The articular surfaces for the sternal ribs are
closely crowded and lie on either side of the coracoid grooves. The interarticular
spaces—incisure intercostales—form deep pocket-like apertures, as in Dinornis and Rhea,
for instance. ‘The crowding together and forward position of the articular surfaces for
the sternal ribs is another Rheine character.
The anterior lateral processes are very similar in form and position to those of Ehea,
which they still further resemble in that the ventral aspect of the base lodges a
pneumatic aperture.
The coracoid grooves in the Crypturi are separated by a deep gorge carved out of
the corpus sterni. These grooves differ from those of Ahea in that the upper and lower
lips are better developed, and in that their transverse extension is less while their depth
is greater relatively.
There is a large spina interna.
Turning once again to the corpus sterni and comparing that of hea macrorhyncha.
kindly lent me by Mr. Beddard, with that of a Tinamou, we notice a point of some
significance. ‘To wit, that in hea on either side of the great median protuberantia
VOL. XV.—PartT v. No. 15.— December, 1900. 21
222, MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
sterni, near the middle of its extent (fig. 4c), is an area of bone conspicuous on
account of its thinness. Now it seems more than probable that these thin areas are
the last traces of a once deep fissure which has gradually closed up, thus uniting to
the body of the sternum a pair of posterior lateral processes. Parker has noticed a
similar indication of the closure of these fissures in Apteryx.
Fusion probably first took place, in the case of Rhea, between the free ends of the
processes and the mesosternum by syndesmosis of the membrane, by which the fissure is
invariably closed. A slight expansion of the free ends of the processes and metasternum,
such as is indicated in Zinamide, would materially hasten this process of closing. It
is also to be noted that the membrane closing this fissure is generally thicker posteriorly
than anteriorly ; this also would contribute towards the work of obliteration. If, once
again, we imagine these fissures open, the resemblance between the sternum of Rhea
and Crypturi will be found to be significantly striking.
It has been already suggested by Gadow that the protuberantia sterni may represent
a degenerate keel. ‘This seems quite probable. Seen in profile its keel-like resemblance
is quite as close as that claimed for Apterya by Parker (fig. 4p). The sternal callosity
so conspicuous in the living bird is represented amongst the Neognathw in Opistho-
comus and is a quite secondary character.
b. The Pectoral Arch.
The shoulder-girdle of the flightless Palwognathe has undergone considerable
modification and bears evidence throughout of retrogression. The coracoid and scapula
are no longer free, except in the very young bird, but completely anchylosed, forming,
the one with the other, a very obtuse angle, instead of an acute angle as in modern
birds.
In Dromeus the sternal (epi-coracoid) border of the coracoid, though wide, is less than
the length of the whole coracoid. Only the inner half of this border is received within
the coracoid grooves ; its outer portion is free and underlies the anterior-lateral process.
The pro-coracoid is ligamentous. The supra-coracoid foramen is large and receives
several pneumatic apertures.
The scapula is slender, and looks like an outward, upward, and backward outgrowth
of the coracoid. It bears a strongly marked articular facet for the vestigial furcula.
The pectoral arch of Dromeus ater does not appear to differ materially from that of
D. nove-hollandiew. I gather this from a sketch by Dr. Henri P. Gervais, kindly
made for me at the request of Prof. Oustalet, to whom I am much indebted, as
well as to Dr. Gervais.
The following measurements (in millim.) accompanied the sketch of Dr. Gervais :—
Length from pro-coracoid process to tip of scapula............... 0°127
Greatestllengihvorqcoracaid yys) 9 eee eee eee 0-062
Width, epi-coracoid region
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH A. 223
In the sketch the supra-coracoid foramen is not indicated, neither is there any
indication of clavicles.
In Casuarius the coracoid is relatively much shorter than in Dromeus. Its breadth
equals its length. The whole sternal border rests within the coracoid groove. Unlike
Dromeus, the pro-coracoid is large and there is a small pro-coracoid fenestra. In the
young Casuarius the pro-coracoid is ligamentous. ‘The supra-coracoid foramen is large
and receives numerous pneumatic apertures. The scapula resembles that of Dromeus
in the general form and in the size of the acromion.
In Rhea the coracoid is long and comparatively slender, tapering rapidly from its
sternal border forwards to the supra-coracoid foramen. ‘The pro-coracoid is ligamentous,
leaving a conspicuous notch on the mesia) border of the macerated coracoid. ‘There is
generally a supra-coracoid foramen, but this never receives pneumatic apertures. The
whole sternal border rests in the coracoid grooves. At the base of the coracoid and on
its dorsal surface is a large pneumatic aperture. This does not seem to be represented
in any other Palewognathe except, and feebly, some Cryptur?.
The scapula is peculiar, curving from the coracoid first inwards then backwards.
The glenoid surface for the humerus lies at the distal end of the coracoid instead of at
its antero-lateral angle.
In Struthio the size of the pre-coracoid is enormous, extending backwards as a long
finger-like process ultimately to reach and fuse with the internal angle of the base of
the coracoid, enclosing in this way a huge fenestra. The sternal (epi-coracoid) border
of the coracoid is of great width. ‘The supra-coracoid foramen is wanting. We might
mention here that the pre-coracoid of S. molybdophanes appears to be peculiar in its
greater relative slenderness.
In Apteryx the pre-coracoid seems to be wanting only in A. oweni, and in this species,
it would seem, at all stages of its growth. In the remaining species it is represented in
the adult by ligament only ; so that in the macerated skeleton the pre-coracoid fenestra
is represented only by the deep notch on the mesial border of the coracoid. Ina
skeleton of A. australis mantelli this notch has been nearly filled up by a secondary
extension of bone in the coracoid of the left side. The supra-coracoid fenestra is
minute or absent in all but A. oweni, in which it is large. This point, remembering
the absence of the pro-coracoid in the latter and its presence in the remaining species,
is interesting.
In the Dinornithidw the coracoid has reached a state of extreme reduction, being
rod-like and merging almost insensibly with the scapula.
In the Aipyornithida, in the form of the pro-coracoid, it is intermediate between
Rhea and Dromeus.
The coracoid of the Crypturi differs from that of its congeners—the flightless Palwo-
gnathe—chiefly in that it has a well-developed acro-coracoid. ‘There is a well-marked
212
224 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
external lateral process at its base. The pre-coracoid is wanting. The scapula is free,
as in the young of the flightless members of this group.
The furcula is slender and U-shaped.
Tur DEVELOPMENT OF THE STERNUM AND PecTorRAL ARCH.
a. Sternum.
Miss Lindsay’s [48] researches have shown that during development the sternum
undergoes a process of shortening both anteriorly and posteriorly, at least in so far as
its costal elements are concerned; the posterior shortening is, however, more than
compensated by the addition of a long metasternum. That the shortening takes place
both anteriorly and posteriorly is proved by the existence of ribs, which in the embryo
are connected with the sternum, but which in the adult become divided therefrom and
atrophied. ‘The cause of the shortening is “ consequent on that posterior translation
of the shoulder-girdle which is at once expressed by the lengthening of the neck and
the shortening of the trunk in the avian as compared with the reptilian type.” This
sternal abbreviation is common to both Palwo- and Neognathe.
In Struthio “the 7 days’ embryo shows an anterior part which may probably be
compared to the manubrium sterni of mammals.” A study of a series of embryos shows
that the anterior lateral processes are an outgrowth of the costal sternum ; in the Neo-
gnathe these processes are to be regarded as vestiges of a former extension of the
costal sternum anteriorly.
The posterior lateral processes in the very early chick, according to Miss Lindsay,
“consist of the ends of the primitive costal bands, preserved and prolonged for the
attachment of the lateral part of the rectus, and subsequently modified ... to afford
attachment to the obliquus externus.”
These processes, as existing in the adult, may be considered a part of the meta-
sternum, since the primitive part bears but a small proportion to the later additions.
The metasternum, both in Struthio and Rhea, remains for the most cartilaginous
throughout life. The cartilaginous metasternum of Dromeus and Casuarius is
relatively somewhat smaller.
Posterior lateral processes are wanting in the sterna of Rhea, Casuartus, and
Dromeus.
In Apterya, according to Parker [71], “it appears certain that... each half of the
sternum is not formed by the antero-posterior union of the whole of the sternal ribs.”
In one stage of the series which he examined the sternum apparently increased in
length from before backwards to the level of the fourth sternal rib, which extended
mesiad to meet it.
The metasternum did not appear until after the posterior lateral processes had
attained nearly their full length. In one specimen the metasternum was double, and
PHYLOGENY OF THE PALZOGNATHZ AND NEOGNATH. 225
a
in another it was perforated by an oval foramen, which Parker suggests indicates a
paired origin.
No traces of a keel were observed, though such are undoubtedly present in some
adults (fig. 4p).
Ossification of the sternum, according to Gadow, begins during the last week of
embryonic life.
In Struthio and Apteryxv a pair of pleurostea only are represented ; in Rhea, Casuarius,
and Dromeus there are a pair of additional centres representing pro-ostea.
Although there is no positive trace of a keel in the developing “ Ratite” sternum, it
is generally assumed that such a structure was present in more remote ancestors, and
that its loss is the result of degeneration accompanied by decaying, and terminating in
absolute loss of the power of flight. It is possible, however, that the earliest birds did
not possess a carinate sternum. The Ratite sterna of the Palwognathw may well
represent this ancient type. The keel was never largely developed in the sterna of the
Pterodactyle, and is not always present among the Chiroptera.
b. Pectoral Arch.
In Struthio and Apteryx only is there a well-developed pre-coracoid.
Apteryx, however, presents, as Parker [72] points out, “ the remarkable circumstance
that in one species of Apterya (A. oweni) the coracoid is solid, presenting no coracoid
fenestra and therefore no procoracoid, while in another species (d. australis) the
procoracoid is present at a comparatively early age, and is frequently retained in the
form of a ligament in the adult. In other words, the differences between the shoulder-
girdle in two species of Apterya are of precisely the same nature as those distinguishing
the Struthiones from the other Ratite.”
In Dromeus and Casuarius and in Rhea the pre-coracoid is vestigial. It may even
be, as Lindsay has suggested, a new growth of the coracoid rather than the homlogue
of the pre-coracoid of Struthio.
Clavicles occur only in the adult Dromeus and embryo Caswarius.
Petvic ARCH.
The pelvic arch of the Palwognathe differs from that of the Neognathw chiefly in
two points :—(1) The great height of the neural spines of the synsacrum (Pl. XLIV.
fig. 5); and (2) the persistent ilio-ischiadic fissure. But though there can be no
difficulty whatever in distinguishing the Struthious from the Neognathine pelvis, yet
these cannot apparently be defined in mutually exclusive terms. There is no com-
prehensive definition that will cut off the one from the other. ‘To keep to the pelvic
arch of the Struthious birds: it is difficult to find a definition for this which
shall not be subject to any exceptions; for although the persistent ischiadic fissure is
226 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
undoubtedly a Struthious, because a primitive, character, yet the Rhea and the
Cassowary contradict this.
The long and narrow synsacrum, with its high neural spines, finds its counterpart in
the synsacrum of the Pygopodes ; and this is a point fraught with some significance to
this last group, inasmuch as we have already evidence to show that the Pygopodes are
a primitive type, and it would seem that this last character is to be regarded as a piece
of additional evidence to this end.
The synsacrum, however, of the Pygopodes differs from that of any Struthious bird
in (1) its much greater lateral compression ; (2) the absence of any pleurosteal clements
caudad of the last thoracic vertebra; and the exposure of the whole of the fused neural
crest lying between the pre-acetabular ilium. In Podicipidide the post-acetabular ila
meet dorsad of the neural crest, whilst in Colymbidw they do not rise beyond the level
of the base of this crest.
In the Pygopodes, again, the ischiadic fissure of the post-acetabular ilium is reduced
to a comparatively small foramen, and the post- greatly exceed the pre-acetabular ila
in length.
In the elongated synsacrum, the vertical ilia, and the comparatively high neural
spines of the synsacrum the Podicipides resemble the flightless Palwognathe.
It may be, the long, narrow synsacrum is primitive, and that this form of synsacrum,
which is characteristic only of the Pyyopodes (including Hesperornis) amongst the
Neognathe, was derived from a primitive ancestor which had not yet adopted the
more usual Neognathine form of pelvis. The fact that the much more recent and
highly specialized Auks, which have adopted the same mode of life as the ancient
Divers, have a distinctly Neognathine type of pelvis with widely separated post-
acetabular ilia supports the view that the synsacrum of the Pygopodes and Palwo-
gnathe is primitive in type.
In the flightless members of the Palwognathe the pre-ilium is always vertical in
position, and always meets its fellow in the middle line above the synsacrum.
Pleurosteal elements representing the sacral vertebre are always present.
The Apyornithide and Dinornithide, whilst they agree with the remaining Palwo-
gnathe in the great height of the synsacral neural spine, differ, as we shall see, in
one or two important features from the other Palwognathw, though these structures
are readily traceable back to the more primitive form,
In Dromeus the pre- and post-acetabular ilia are subequal in length, or the post-
acetabular is markedly the longest. The pre-acetabular ilium has a strongly arched
dorsal border, a sigmoidally curved ventral border, which caudad sends down a long
acetabular process to join the pubis and serves at the same time as the anterior boundary
of the acetabulum, ‘The depth from the dorsal to the ventral border 1s considerable,
this portion of the innominate bone forming a vertical blade very slightly hollowed in
its middle. The post-ilium is narrow cephalad and tapers rapidly from before back-
PHYLOGENY OF THE PALZOGNATHZ AND NEOGNATHA, 227
wards, this region of the pelvis in this respect resembling Struthio. In half-grown
specimens it is quite pre-caudad; in adult birds it becomes fused with the extreme
postero-dorsal angle of the ischium.
The ilium increases markedly in breadth during its growth. This increased breadth
accompanies a corresponding increase in the height of the neural spines of the sacral
and post-synsacral vertebree, which, as has already been described (p. 208), in the
embryo are represented only by low cartilaginous transverse ridges. In the embryo the
pre- and post-acetabular ilia are slightly convex superficially, in the adult they become
slightly concave.
The supra-trochanteric process marks the region, more or less, of the sacral vertebre.
It lies immediately above the anti-trochanter, but does not project outwards as in
Rhea. One of the most remarkable features of the pelvis of Dromeus and Casuarius
is the close embrace with which the post-ilia receive the post-sacral vertebre of the
synsacrum, almost entirely suppressing the para- and diapophysial processes.
In Dromeus and Casuarius the sacral vertebree extend outwards beneath the vertical
border of the ilium to articulate with the ascending process of the ischium at its
junction with the ilium.
The pelvis of D. ater differs from that of D. nove-hollandie chiefly in that the
development of the pectineal process is but slight—wherein it approaches Caswarius,—
and in that the obturator fissure is wider, the rotation backwards of the pelvis being
less. The crest of the pre-ilium is less arched.
The pelvis of Dromwus can scarcely be distinguished from that of Casuarius.
In the young bird (half-grown) the neural spines of the sacral region of the syn-
sacrum appear to be more transversely expanded than in Caswarius. In the adult the
pre-ilium appears to be relatively of greater breadth in Caswarius and the post-ilium is
relatively longer and narrower. ‘The strong <-shaped supra-trochanteric processes
and the width across from one side to the other are apparently rarely attained by the
Cassowary.
Judging from the skeletons in the Museum collections it would seem that in the
width across the supra-trochanteric process and the relatively long post-ilium, the
pelvis of Dromaus is readily distinguishable from that of Caswarius.
Unfortunately, however, these data do not appear to be really reliable, for in a pelvis
of C. casuarius the resemblance to these very characteristic points in Dromaus were so
close as to render it most difficult to distinguish between the two genera.
In C. bennetti the width across the supra-trochanteric process is as conspicuous as in
Dromeus, but the whole pelvis is of course much smaller than in Dromeus.
In the young (half-grown), both of Casuarius and Dromeus, the ischiadic fissure is
complete. Later in life it is closed by fusion of the extreme antero-dorsal angle of the
fore end of the ischium fusing with the ilium. In very old birds the width of this
fissure caudad becomes seriously encroached upon by the downgrowth from the post-
228 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
ilium of an exceedingly thin plate of bone with a jagged free edge. It arises in the
region of the 5th post-sacral vertebra, and is continued backwards from this point. From
the superior border of the ischium there arises a similar plate of bone of corresponding
length. In very old birds these two jagged-edged plates nearly meet, foreshadowing
the formation of the ilio-ischiadic foramen of the Neognathe.
The interobturator process, both in Casuarius and Dromeus, is well marked.
The tschia, both in Dromeus and Caswarius, are more or less rod-shaped and
very distinctly bent inwards towards the middle; this is a point of great significance
when considered in connection with the ischium of Rhea. The ischium is roughly
2-shaped. Its vertical process articulates with (in the young) and eventually
fuses with the pre-ilium, forming the lower half of the anti-trochanter. Antero-
ventrally it articulates with and eventually fuses with the pubis, the fusion with the
pubis taking place much earlier than is the case with the vertical process and the ilium.
The pubes are very slender relatively, with a strong outward curve in Dromeus and
Casuarius casuarius, straight or only slightly curved in the remaining species of
Casuarius. In the nestling they do not extend backwards as far as the level of the
free end of the post-ilium. In the adult they extend beyond this, reaching nearly to
the end of the ischia. They are always free posteriorly in the dried skeleton,
connected by cartilage in the living bird with the ischia.
The pectineal process seems to be formed for the most part by the descending process
of the ilium. It reaches its greatest development apparently in Dromeus, but in a
pelvis of Caswarius casuarius it is only slightly less in size. In the remaining species
of Casuarius it is but feebly developed.
The outline of the pre-ilium differs, but the series of adult skeletons in the Museum
collection is not sufficiently large to enable any reliable data to be drawn from
comparisons.
Specific differences in the form of the pelvis of Caswariws may possibly be found, but
I think it extremely doubtful. The collection of adult skeletons which forms the
subject of this paper is not nearly large enough to afford the material necessary for
such a task.
The pelves of the C. bennetti group seem distinguishable from the C. casuarius group
by reason of the relatively longer and narrower post-acetabular ilium. We have
no adult skeletons of the C. unappendiculatus group.
In Rhea the form of the pelvis is unique, but, as will be shown, is probably a direct
modification of the Dromeine type.
In the nestling the pre- and post-acetabular ilia are subequal in length. ‘The latter
is abruptly truncated caudad, and rests by its postero-ventral angle upon the closely
approximated ischia, to be described presently. The supra-trochanteric process is
but feebly developed, and the anti-trochanter is but slightly below the level of the
neural crest of the synsacrum. ‘The post-acetabular ilium is long and narrow and
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHAZ. 229
closely resembles, at this stage, the permament condition which this bone obtains
in Struthio.
The extreme antero-ventral angle of the pre-ilium is sharply defined and prodyced
outwards, overlapping the penultimate thoracic rib, with which it ultimately becomes
completely fused. The innominates meet in the middle line above the synsacrum,
cephalad and caudad, but leave the neural spines of the synsacrum exposed at the
bottom of a groove over the sacral region.
The ischia, which, it will be remembered, in Caswarius approached one another in
the mid-ventral line, in Fhea actually meet at a point corresponding with a line passing
downwards from behind the anti-trochanter. From this point backwards they remain
in close juxtaposition, later in life more or less completely fusing. ‘These approximated
ischia form a lcng backwardly projecting median bar, produced caudad far beyond the
level of the post-acetabular ilium. Thusa kind of false roof is formed to the abdominal
cavity and a floor to the renal fossa.
The pubes, as in Dromeus and to a lesser extent in Casuarius, are bowed outwardly
and terminate freely near the end of the ischia.
The pectineal process is formed in part by the descending acetabular process of the
ilium and partly by the pubis.
The adult pelvis differs in many respects from that of the nestling, carrying the points
wherein it is peculiar a stage further in development.
The supra-trochanteric process now forms a large overhanging ledge, directly above
the acetabulum. The innominate bones now meet in the mid-dorsal line throughout
the whole of their length. ‘The post-acetabular ilium fuses on either side by a broad
expanded foot with the ischia, and beyond this fusion—a point of contact only in the
nestling—is continued backwards a long pointed process. The pubes, turning suddenly
caudad and mesiad, fuse eventually with the ischia.
The sacral vertebree in Caswarius lay level with the ventral border of the innominate,
and extended outwards to articulate with the vertical acetabular process of the ischium.
In Fhea these vertebre are raised up so as to lie above the level of the acetabulum.
The post-acetabular ilium, in the nestling Grebe, is subequal in length with the pre-
acetabula, in the adult it is longer than this.
In Struthio the pelvis presents one or two points wherein it differs markedly from
that of the other Palwognathe.
Like Dromeus, the post-acetabular is longer than the pre-acetabular ilium, only the
difference is much greater in Struthio. ‘The post-acetabular region remains permanently
and widely separate from the ischium, and is so narrow in proportion to its length as
to be almost rod-shaped. ‘The pre-acetabular ilium is relatively as long as in Dromeus.
The fovea iliaca anterior is rather convex than concave. In this particular Struthio is
peculiar: since the pre-acetabular ilium instead of being concave externally, so as to,
on this account, more or less completely embrace the neural spines of the synsacrum,
VOL. XV.---PAR? V. No. 16.—December, 1900. 2K
230 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
is convex, and forms, with its fellow of the opposite side, a long tunnel divided into
right and left halves by the neural spines of the synsacrum, which acts as a median
partition.
The dorsal plane, which in Dromeus, Casuarius, and Rhea was restricted to a small
area between the supra-trochanteric precesses, here, in Struthio, extends backwards to
the extreme end of the post-acetabular ilium, though it is very narrow. Furthermore,
this plane slopes gently downwards on either side, so that its extremest outer angles
approach the anti-trochanter.
The pre-acetabular ilium meets in the mid-dorsal line above the synsacral spines: the
post-acetabular ilia remain widely separated one from another throughout the greater
part of their length; in old individuals they meet and fuse with the last two post-
sacral vertebre of the synsacrum. Down the median space, between the two post-
ilia, runs a long bony bar, formed by the fusion of anterior and posterior ossifications
of the neural spines. In Dinornithide, it will be remembered, it was pointed out [1]
the equivalent ossifications extended laterally, so as to reach the ilia on either side, and
thus close what would otherwise form a deep lateral trench lying on either side of the
synsacrum. ‘This trench in Struthio remains permanently open.
The ¢schium is very slender and rod-shaped. At its posterior extremity it sends
downwards a broad plate of bone which fuses with the pubis. In the young bird, it
should be mentioned, the ischium is free posteriorly. The post-acetabular illum and
the ischium remain widely separated throughout life.
The pubis is long, slender, and rod-shaped. In the adult it extends caudad far
beyond the post-ilium and ischium, and curving sharply downwards and forwards, meets
its fellow of the opposite side in the middle line, and becomes permanentiy fused
therewith in a long, forwardly directed symphysis. ‘This fusion of the pubes posteriorly
occurs in no other living bird.
As is well known now, the pubis of Struthio is further remarkable for a small bony
plate, attached to the middle of its ventral border by cartilage. In macerated
skeletons this is generally lost; in this case its position is indicated by a roughened
surface.
This bone appears to have deen originally described by Meckel [56], who held it to
represent the marsupial bone of the Marsupials. More than forty years later this bone
was rediscovered and described by Garrod and Frank Darwin [28]. They also regarded
it as probably homologous with the marsupial bone of the Kangaroo and its allies.
With this view Beddard [7] seems also to agree.
The pectineal process is long, slender, and curved, its free end being directed
upwards.
The ilio-ischiadic fissure remains permanently open in Struthio. In Casuarius and
Dromeus and in Rhea it closes later in life. The obturator fissure is closed, by fusion
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATIA- 231
of the free end of the pubis with the ischium, both in Rhea and Struthio, and only in
these.
The pelvic girdle of Apteryx resembles that of Dromeus and Casuarius in some
respects. This is what we should expect, since Dromaus probably represents the
ground form from which the other Palwognathe have been derived.
In the extraordinary relative length of the pre-acetabular ilium <Apterye stands
alone. It extends forward so as to overlap some four thoracic ribs) The post-
acetabular ilium is as short relatively as the pre-acetabular is long; it never exceeds
half the length of the pre-ilium. Asin Séruvthio, the post-ilium and ischium are widely
separated, but the separation is more marked since the backward rotation of the
ischium is somewhat less than in Séruthio.
The innominates meet in the mid-dorsal line as in Rhea, but with this difference:
whereas in /hea although the median borders meet, yet a dorsal plane is present as a
widening out and flattening of that portion of the dorsal aspect of the pelvis which lies
between the supra-trochanteric processes, in Apterya these dorso-median edges meet
along their whole length, save in the region of the sacrum proper, where a narrow
chink reveals the presence of some three neural spines. There is no supra-trochanteric
process, but the ilia meet in the middle line to form a long knife-like ridge. The
Jovea iliaca anterior extends backwards into the post-acetabular ilium, terminating
caudad of the anti-trochanter. Behind the acetabulum the vertebral column is bent
sharply downwards, and has undergone great lateral compression, so that the centra
of the vertebree have become vertically compressed plates but little thicker than their
neural spines. Furthermore, owing to the downward flexure of the posterior synsacral
vertebrae the post-acetabular ilia have come to embrace the neural spines of the
vertebrae instead of the centra. The synsacrum extends, moreover, by a single vertebra
beyond the level of the free (caudad) end of the post-acetabular ilium.
The tschium is a relatively long flattened blade, forming a wider angle with the
ilium than in any other of the Ratitw.
The pubis is slender, with a more or less well-marked ventral curve. Caudad, its
free end may touch the postero-ventral angle of the ischium, but it never fuses
therewith.
The obturator foramen and fissure are confluent.
The pectineal process is large and pointed, and formed in part by the descending
process of the ilium and in part by the pubis.
Specific differences in the pelves of Apterya are, as Beddard has already shown, very
slight. According to him the pectineal process is shorter in 4. owen? than in
A, australis. In our specimens if there is any real difference the reverse would appear
to be the case. I have managed to find sufficient differences to construct a key for the
species, though with difficulty. Mr. Beddard has already shown that the width across
Mie
232 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
the pre-ilium varies, but I think that this, in spite of variation, combined with other
characters may be relied on.
Apteryx australis seems to stand alone in the great width of the pre-acetabular
ilium, a width due to a highly arched dorsal border and a very considerable lateral
expansion of its antero-ventral border. A rather sinuous post-acetabular ventral
border and a sudden widening of the post-acetabular ilium caudad are combinations
apparently peculiar to this species. In A. haastw the pre-ilia are sharply truncated
forwards, and the pre-ilium is generally broader in proportion than in A. owent. The
post-acetabular ventral border appears to be concave.
A. australis mantelli appears, superficially, not to be very readily distinguishable
from A. oweni. The chief differences appear to lie in the smaller pectineal process and
broader ischia of A. australis mantelli.
The pelves of Dinornithidw and of the Apyornithide very closely resemble one
another, and differ from all other flightless members of the Palwognathe in that the
post-acetabular region of the pelvis is flattened out into a large, pentagonal plate, nearly
as broad as long. This is made up partly by the great length of the transverse
processes of the synsacral vertebre, and partly by the great widening of the dorsal plane
of the ilia—a widening only feebly represented among the Palwognathe elsewhere
in Struthio.
In the relations of the ischium and pubis the two pelves now under discussion most
nearly resemble Apteryx. Apteryx, however, differs in one respect, in that in this
genus the obturator fissure and foramen are confluent. In the Dinornithine pelvis
the foramen is shut off from the fissure. The pectineal process is large in Apterya,
very small and wanting in the Dinornithine pelvis.
In the Dinornithide the sacral are more or less easily distinguishable from the post-
sacral. In the Zpyornithide this is not the case.
In the Dinornithide the post-sacral neural spines lie in the middle of a deep fossa,
the floor of which is formed by the upwardly directed neural spines. In the adult this
fossa is closed more or less completely by a bony roof formed by tabular lateral
expansions from the crest of the neural spines. Caudad, however, this closure is not
quite complete: a pair of lateral slits run up on either side of the median neural
plate, from behind forwards; the extent of these slits decreasing with age, but never
entirely disappearing.
In 4&pyornis the roofing of the fossa is ample: a double row of foramina only
excepted, which run from behind forwards to the crista transversa.
The pelvis, both in Dinornithide and Aipyornithide, is relatively much shorter in
proportion to its width than in the other Palewognathe.
(Sh)
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA, 23
Tue PecroraL Lime.
The pectoral limb is in all the flightless Palwognathe a very degenerate structure.
Perhaps that of Rhea should be regarded as the least degenerate.
In the Rhea the length of the extended ante-brachium and manus is only very
slightly greater than that of the brachium. The length of the manus is greater than
the ante-brachium. ‘The length of the pollex with its claw is very nearly as great as
the 11. metacarpal. In a nestling Rhea I find on the m1. digit a vestige of a claw; a
similar vestige has been already detected by Wray in the Ostrich. The ulnare
possesses the typical Neognathine form—in Struthio this is a mere nodule. The Ist
phalanx of the second digit is broad and flat, the 2nd phalanx is pointed.
The ante-brachium is about one-third shorter than the brachium. ‘The post-axial
border of the ulna from the olecranon outwards for some considerable distance is
greatly compressed. The superficial appearance of the forearm resembles that of
many long-winged Weognathe.
In the humerus the pectoral crest is obsolete, as also is the crista inferior.
is no subtrochanteric fossa, and the fossa for the brachialis internus is only faintly
indicated.
In Struthio the length of the extended ante-brachium and manus is about one-third
less than the length of the humerus, and falls far short of that of Rhea, being but a
little more than two-thirds as long as the brachium.
In Struthio the manus is longer than the forearm, in Rhea the reverse is the case.
In Rhea the forearm is about two-thirds and in Struthio about one-third the length of
the humerus. The distal carpal mass only just reaches the base of the 1st metacarpal ;
in Rhea it extends pre-axiad so as to support the whole base of the pollex. The Ist
phalanx of the m. digit is relatively longer in Struthio than in Rhea. The 1. metacarpal
in Struthio is more or less rod-shaped, that of the m1. is rod-shaped and bowed. In
Rhea the distal end of metacarpal 1. is flabellate, the 11. metacarpal is very slender,
rod-shaped, and sigmoidally curved.
The ulna, in Struthio, is much expanded distally by a more or less triangular out-
growth of its pre-axial border. The olecranon is but feebly developed. The distal
end of the radius is grooved and much expanded in the direction of its articular surface
for the radiale.
The pectoral crest of the humerus is more conspicuous than in Rhea, forming a long,
low, swollen ridge. There is a wide and shallow fossa distad of the caput humeri,
possibly representing the sulcus transversus. There is also a shallow incisura capitis.
The crista inferior is wanting. ‘The subtrochanteric fossa is present but shallow, but
it receives no pneumatic apertures. The tuberculum medius is moderately well
developed.
The linea aspera for the triceps is raised into a long and sharp ridge, which reaches
234 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
its greatest development rather below the middle of the humerus. There is a feebly
developed ectepicondylar process.
The radiale is well-developed ; the ulnare is reduced to a small nodule.
In Dromeus, Casuarius, and Apteryx the wing has undergone extreme reduction.
In Dromeus the length of the whole wing is scarcely longer than the skull. The
length of the extended forearm and hand is about equal to the length of the humerus.
The manus is considerably shorter than the forearm. There is no distinguishable
carpal region, and the radius is anchylosed with the fused carpo-metacarpal mass. ‘The
metacarpals and phalanges have all fused into one indistinguishable rod of bone. The
pectoral crest is but faintly represented.
There is a small subtrochanteric fossa, into which opens several pneumatic foramina.
In the skeleton of D. ater which I have had for examination the wing is
wanting. Judging from the measurements given in Milne-Edwards’s memoir, how-
ever, it seems to have been smaller relatively than in D. nove-hollandiw. From the
vestigial character of the coracoid grooves one would have expected a much greater
difference.
In Casuarius the wing, strangely enough, though relatively smaller than in Dromeus,
is yet less degenerate. The proportions of the brachium, ante-brachium, and manus
are much the same as in Dromeus.
In the adult, the three metacarpals can be plainly made out. That of the pollex
is vestigial. Only one phalanx appears to be represented—that of the 1. metacarpal,
and this is but a vestige. The radiale has anchylosed with the radius, but is still
distinguishable. ‘There appears to be no ulnare. ‘The humerus has the pectoral crest
more strongly developed than in any other Palwoguathw. The subtrochanteric fossa,
smaller in Dromeus, is very large in Casuarius.
In a nestling Caswarius, a radiale, ulnare, and distal tarsal mass are plainly
distinguishable.
In Apteryx the degeneration of the wing has reached a stage somewhat intermediate
between that of Dromwus and Casuarius. In the relatively great length of the
humerus, however, it differs from both and resembles Rhea. ‘The extended ante-
brachium and manus are only a little more than half the length of the humerus.
The radius is shorter than the ulna.
The carpus and manus, as both Parker and Beddard have already pointed out, are
extremely variable. In A. haastii Parker found an ulnare, radiale, and a fairly distinct
metacarpale m1. In A. australis there are no distinct carpals, but traces are dis-
tinguishable of three metacarpals. The index has two or three phalanges, the last
terminating in a claw. In A. oweni the radiale appears to be always distinguishable.
The manus appears to vary most, according to Beddard, in the matter of degeneracy,
in A. australis mantelli.
In an embryo of Apteryx australis, T. J. Parker found, not only a separate radiale,
PHYLOGENY OF THE PALZOGNATHEZ AND NEOGNATHA, 235
ulnare, and distal carpal mass, but also an intermedium, an element which has so far
been recorded previously in Opisthocomus and Dendrecia.
The fact that in the adult Apteryx there is a distinct alar membrane, or patagium,
as has been pointed out by Parker, is strong evidence in favour of the view that the
wing of Apteryx is a degenerate structure, once capable of performing the duties of an
organ of flight.
In A%pyornis, apparently the wing was represented only by the humerus. A humerus,
believed to be that of an Apyornis, has been described by Andrews. Its length was
63 mm. ‘The upper end resembled that of the humerus of Aptornis, the lower that of
Casuarius, “but still further reduced, the radial and ulnar trochlee uniting completely
to form a single articular surface.”
Tue Petvic Lives,
The differences between the pelvic limbs of the existing Palwognathe are neither
many nor great.
In Dromeus the femur differs from all the other Palwognathe, save Struthio, in
that it is highly pneumatic, and bears on the ventral aspect of the femur a large
pneumatic foramen. It lies between the great trochanter and the head of the femur.
The popliteal fossa is of moderate depth.
The tidio-tarsus is not easily distinguishable from that of Caswarius. The chief
difference appears to rest in the slighter and less vertically extended procnemial crest.
This vertical direction is rather marked in Caswarius. The fibula extends to the lower
fourth of the shaft of the tibio-tarsus.
The tarso-metatarsus is as long as the tibio-tarsus, grooved in front, and has a
moderately developed median keel to the hypotarsus.
The proximal phalanx of digit 11. is relatively long, the 2nd very short; in digit Iv.
the proximal phalanx is long, the 2nd and especially the 5rd and 4th are very short.
The pelvic limb of D. ater closely resembles that of D. novw-hollandie. The
anterior metatarsal groove is, however, somewhat deeper.
The outer toe, in each foot, in this skeleton has lost one phalanx.
In Casuarius the femur is non-pneumatic. ‘The position of the pneumatic fossa in
Dromeus may be indicated by a minute aperture. ‘The tibio-tarsus, as already pointed
out, differs but slightly from that of Dromeus.
The tarso-metatarsus is much shorter than the tibio-tarsus, is very deeply grooved
anteriorly, and is pierced by a foramen dorsad of the scar for the tibialis anticus.
Thus it can readily be distinguished from the tarso-metatarsus of Dromeus.
The ungual phalanx of the inner toe is greatly elongated ; in other respects the size
of the phalanges closely agrees with those of Dromeus, but the proximal phalanx of
digit 11. is relatively shorter.
236 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
In Struthio the femur, as in Dromeus, is highly pneumatic, and, as in Dromeus,
bears a large pneumatic fossa between the head of the femur and the great trochanter.
The shaft is relatively much thicker than in Dromeus or Cassowary, and the fibular
condyle is much larger than the inner tibial condyle. ‘The groove dividing them is also
narrower and deeper than in Dromeus.
The tibio-tarsus, apart from its greater size, is peculiar in the relatively smaller
size and hooked form of the ectocnemial crest, and the great length of the fibula,
which extends to within a short distance of the upper border of the proximal tarsal
mass. The extensor groove is shallow. Furthermore, the lateral borders of the
posterior trochlear surface are produced backwards and upwards into a pair of
proximal ridges at right angles to the long axis of the shaft.
The tarso-metatarsus is of course at once distinguishable by the absence of ento-
trochlea. It is grooved anteriorly, and bears a pair of foramina dorsad of the scar for
the tibialis anticus.
The ungual phalanx of the outer toe is represented by a small nodule.
In Rhea the femur is non-pneumatic. It may be at once distinguished from that
of any other Palwognathe by the great depth of the popliteal fossa.
The tarso-metatarsus resembles that of Struthio in the feeble development and in
the form of the ectocnemial crest and in the backward projection of the condyle of
its distal extremity. The femur falls far short of the level of the tarsal mass. The
tarso-metatarsus has a shallow groove anteriorly, a deep fossa above the scar for the
tibialis anticus, and a very broad and low intercotylar process. ‘The hypotarsal keel
lies to the inner side of the median line. In Dromeus and Caswarius it is median in
position.
The Ist phalanx of digit 1. is conspicuously lengthened, the 2nd is very short;
of the tv. digit the proximal phalanx is long, the 2-4 very short.
In Aptery the femur is relatively large and more slender than in any other living
Palwognathe, save the Tinamous. It is non-pneumatic, and the popliteal fossa is
deep.
The tibio-tarsus proximally is broad and flattened from back to front, neither the
ecto- nor procnemial crests are much developed. The extensor groove is deep. The
intercondylar gorge is deep, and the ento-condyle of great size.
The tarso-metatarsus is shorter than the femur, much flattened from back to front ;
the intercotylar tubercle is well marked; there is a fossa above the scar for the tibialis
anticus; the 2nd and 4th trochlewe are widely separated from the meso-trochlea. The
hypotarsus bears a pair of low median ridges.
There are four digits. The digits contain no conspicuously shortened or elongated
phalanges, and therein differ from the other Palwognathe herein described.
In Dinornis the femur is relatively much shorter and thicker than in Struthio. Its
distal end is of great width; the intercondylar gorge is very wide and shallow and
PHYLOGENY OF THE PALZOGNATHEH AND NEOGNATHA, 237
extends proximad on to the shaft, terminating rather behind its middle. There are
no conspicuous pneumatic foramina. The great trochanter is very large; it is pro-
duced upwards above the head, and not backwards caudad of the head, being an
exaggeration of what obtains in Dromeus and Casuarius. ‘The popliteal fossa is well
defined, but not deep.
The tibio-tarsus is much flattened from front to back, as in Apterya and Apyornis ;
the ectocnemial crest is large, and gives the anterior view of the proximal end of the
bone a flabellate form; the procnemial crest is feebly developed. The shaft may or
may not be inflected—according to the genus. There is a strong extensor bridge and an
intercondylar tubercle.
In Atpyornis the femur is also very short. The great trochanter is very lofty and
produced caudad. The intercondylar gorge is wide and deep, and the popliteal fossa
shallow.
The tibio-tarsus has its proximal end much flattened from front to back; a feeble
procnemial and large laterally directed ectocnemial crest. Distally the shaft is
inflected. There is no extensor bridge, and the groove is shallow.
The tarso-metatarsus is short, wide, and grooved anteriorly, but the groove is shallow.
The ectotrochlea is widely separated from the mesotrochlea. There is a low, median,
hypotarsal ridge.
Phalanx 1 of digit 1. is moderately long, the 2nd is short; phalanges 2, 3 of
digit Iv. are conspicuously short, the 4th is shorter. The proportions of the phalangeals
agree closely with those of Dromeus.
In working out the development of Apteryax, Parker [71] found, in the earlier stages,
all five digits present—as in some Neognathe. The fifth digit was a short conical
cartilage attached by its proximal end to the fibulare, and by its pre-axial border to the
distale.
The fibula was of the same length as the tibia.
The tarsus contained the usual elements—a tibiale, fibulare, and distale. The
ascending process of the tibiale at no stage showed any sign of a separate origin,
comparable to an intermedium. Later, after hatching, there appears in the mesotarsal
articular pad a pair of centralia. Sometimes only one is present.
The procnemial crest ossifies, as usual, from a separate centre. Concerning this
ossification I would remark that it is doubtful whether it has any phylogenetic signi-
ficance. Rather it would seem to be comparable to the separate ossification-centre of
the great trochanter of the mammalian femur, and to indicate a position of great
strain. In a recent paper I, however, described it as an epiphysis, as also, though
unknown to me at the time, did the late W. K. Parker [79]. In my own case I may
claim some justification, since the base of this procnemial ossification—that of a
young Grebe—was sufficiently large to form a complete tibial cap, divided from the
shaft by cartilage, as is a true epiphysis. Moreover, the resemblance to a true
VoL. Xv.—Part y. No. 17.—December, 1900. 21
238 MR, W. P. PYCRAFT ON THE MORPHOLOGY AND
epiphysis is the more complete in that it affords an articular surface for the femur.
I will not comment further upon the matter here as I have the subject still under
investigation.
MYOLOGY.
The musculature of the Archwopalatine is now well known, thanks to the researches
of Beddard, Fiirbringer, Gadow, Garrod, Haughton, Meckel, Owen, Parker, and others.
So well have these been done, that, gleaning late in the day, I have but little to add.
It must not be supposed, however, from this, that the subject is now quite exhausted.
On the contrary, many points yet await investigation ; mention of these finds no place
here, however, either because material for this paper was lacking or the points appeared
to be of little or no value so far as it was concerned.
THe SKIN-MUSCLES.
The dermal muscles appear to be most strongly developed in Apteryx.
Casuarius and Dromeus appear to be peculiar in that they possess a strongly
developed dermo-dorsalis. In a Caswarius unappendiculatus this extended from the
base of the skull down the skin of the neck and back along the middle line. At the
base of the neck the muscle gives place to tendon, which, becoming rapidly stronger,
is inserted, by means of numerous long, coarse, tendinous fibres, along the back as far
as the pre-acetabular ilium.
Tur Muscues oF THE WING AND SHOULDER-GIRDLE.
To the loss of flight we may attribute the absence of many muscles found in
Neognathe. Furthermore we must regard this loss as one of considerable antiquity,
since the flightless members of the Neognathe exhibit no such reduction in the number
of the muscles concerned in this form of volition.
Mr. Beddard [7] has given an admirable summary of the missing wing-muscles of
the Palwognathe. They are the pectoralis propatagialis, biceps propatagialis,
deltoides propatagialis, deltoides nvinor (excluding Apteryx), scapulo-humeralis anterior,
and expansor secundariorum. Concerning the last mentioned, traces occur in the
wing of Rhea (see fig. 5 a, p. 240).
The form of flexor carpi ulnaris of Rhea is extremely interesting, as will be seen by
a comparison of the accompanying figures (pp. 240, 241).
It will be noticed that in Fhea (fig. 5a) this muscle is divided into two parts:
(1) a pre-axial ribbon-shaped and partly tendinous, which is inserted on to the ulnare,
and (2) a post-axial, much larger, strap-shaped, fleshy portion inserted on to the
Ist metacarpo-digital remex. ‘The post-axial border of this hinder strap-shaped
portion is bounded by a narrow thread of tendon—a rudimentary wnculum elasticum.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 239
In the Crypturi we have a second modification (fig. 5B, p. 240). The post-axial fleshy
moiety has here apparently degenerated so as to be represented only by the vestigial
muscular slips from the body of the pre-axial portion, which is thick and fleshy. The
vinculum elasticum represented in Rhea by a rudiment only—by rudimentary I mean
nascent, not vestigial—is here increased to form a great tendinous sheet. We may
suppose that this is an improvement on the proto-carinate form seen in Rhea.
In Opisthocomus (fig. 5c, p.241) both pre- and post-axial moieties are fleshy. The post-
axial shows the first indication of the complex vinculum elasticum of the Neognathe.
In Leptoptilus the pre-axial portion is almost entirely tendinous. Proximally, the
tendon divides, one branch running directly to the ulnare, and one fusing with the
anterior border of a muscular post-axial portion throughout the greater part of its
length, finally trending upwards to join the main pre-axial tendon on the ulnare.
This last, muscular, post-axial portion apparently represents the broad strap-shaped
portion of Rhea; it differs therefrom only in that it is inserted into the first cubital
instead of the first metacarpal remex.
There are numerous other modifications, varying in complexity, to be found amongst
the Neognathw, a study of which would probably yield important results.
The wing of Rhea represents, probably, as Dr. Gadow—to whom I showed these
facts—remarks, a proto-carinate form.
In the manus of Rhea, Mr. Beddard writes, ‘‘ we have more evidence of degeneration
than in Struthio. There are, in the first place, only twenty-one muscles at most, and
some of these are much simplified.
“The muscles that appear to be totally wanting are (1) the extensor digitorum
communis, (2) the pronator profundus.
“The extensor indicis is only represented by the belly arising from the wrist. The
Jlexor sublimis may possibly be represented by a slip of muscle arising from the
tendinous edge of the flexor metacarpi uluaris....
“The origin of the biceps is peculiar; it arises not only from the coracoid spine by
a rounded tendon, but also by a sheet cf tendon edged with muscle from the whole of
the coracoid and from just an adjacent bit of the sternum. It is inserted on to both
radius and ulna.
“ As in the Ostrich, the radio-metacarpalis ventralis arises from the ulna. In Rhea
there is a special peculiarity in the presence of a muscular slip running from the
tendon of the extensor metacarpi ulnaris near to its insertion to the extensor indicis.
Finally the ectepicondylo-ulnaris is distinct.”
Of Struthio, Mr. Beddard writes that, ‘in spite of the small size of the manus... .
relatively to that of flying birds, there is but little, if any, evidence of degeneration in
its musculature. On the contrary, indeed, for it might be said that the wing-mnscles
of Struthio are less degenerate, or at any rate less modified, than those of Curinates, in
that amount of muscle compared with tendon is greater. The complication of the
2u2
240 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Fig. 5,
A.
Rs
SA
conjoined flexors digitorum is highly suggestive of a walking or climbing animal.
It seems to be conceivable that the Ostrich branched off from the avian stem before
the power of flight was perfectly established.
“The biceps of Struthio arises from the spina coracoidea and is inserted on to the radius
and ulna and the membrane between them. The extensor metacarpi radialis is single.
The ectepicondylo-ulnaris is absent or fused with the extensor metacarpi ulnaris. The
extensor digitorum communis supplies only the index. The two pronators form only
one muscle. The flexor digitorum sublimis and profundus arise by a single head
from the flexor condyle of the humerus. The two muscles immediately divide; the
upper part (=sublimis) ends in two tendons, of which one is inserted on to (the)
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 241
A.—Inner view of a dissection of the wing of Rhea americana, to show the remains of the expansor secund-
ariorum and the ? proto-carinate condition of the flexor carpi ulnaris. Note the large, fleshy inferior
portion eventually inserted into the base of the 1st primary remex and the thin tendinous band which
runs along its free border, probably representing the vinculum elasticum.
B.—Inner view of a dissection of the wing of Oalodromas elegans. Note that the fleshy posterior portion of
the flewor carpi ulnuris has become reduced to slender slips of muscle to the bases of the secondary
remiges, whilst the vinculum elasticum has greatly increased in size to form a broad tendinous sheet.
C.—Inner view of a dissection of the wing of Opisthocomus cristatus, showing another phase in the develop-
ment of the vinculum elasticum and the persistence of a strongly developed posterior belly to the flewor
carpi ulnaris. Numerous intermediate stages occur amongst the Neognathe.
exp.sec. = eXpansor secundariorum,
fl.c.u. = flexor carpi ulnaris: '=anterior, ''= posterior portion.
uln. = ulnare.
v.é. = Vinculum elasticum.
radiale, the other fuses with the upper tendon of profundus, and also gives off two
slips which surround that tendon and, reuniting, fuse with the lower tendon of the
profundus. The lower part of the muscle (=profundus) gives off two tendons, of
which the upper ends on the first metacarpal, while the lower runs to the base of the
last phalanx of the index.
“The flexor metacarpi ulnaris ends fleshily on the ulnare, but is prolonged beyond
this bone, receiving also some fibres from it to the metacarpal.
“The radio-metacarpalis ventralis... . arises from the ulna and not from the radius.
“ The total number of muscles in the hand of the Ostrich is twenty-three, allowing for
_the absent ectepicondylo-ulnaris. ‘The additional muscle is a smal] pronator quadratus.”
242 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
In Apteryx the usual flexors and extensors of the forearm are present, and, as the
late Prof. T. J. Parker points out, a rather unusually large development of muscles
acting as pronators and supinators for so small and vestigial an organ.
The biceps of Apteryx is single-headed, arising only from the coracoid ; it is inserted
only on to the radius. Apteryx differs only from the other genera in the possession of
the Gallinaceous and ‘Tinamine entepicondylo-ulnaris and an accessory brachialis
anticus.
In Casuarius and Dromeus the wing seems to have reached its most vestigial
condition.
The biceps in Casuarius arises only from the coracoid, and is inserted by a single
tendon both upon the radius and ulna. In Dromeus, according to Beddard, the biceps
has the same peculiar origin to that of Rhea (p. 241).
‘** All the Struthious birds,” writes Beddard [7], “except Apteryx, have lost the
serratus metapatagialis, the latissimus dorsi metapatagialis, and the pectoralis
abdominalis.
“On the other hand, Apteryx has lost what the other Struthious birds have retained,
the latissimus dorsi anterior and the rhomboideus profundus; the latter muscle,
however, is not distinguishable in the Cassowary.
“Tt must be admitted, therefore, that Apteryx, so far as concerns the anterior
extremity, has diverged from the hypothetical ancestral condition in slightly different
lines from other Struthiones.”
The rhomboideus superficialis in Rhea and Struthio arises as in Neognathe. In
Rhea it is inserted on to both the coracoid and scapula, in Struthio on to the scapula
only.
In Apteryx, Casuarius, and Dromeus it arises from the ribs.
The rhomboideus profundus in Caswarius and Dromeus arises from the ribs. In
Caswarius it is with difficulty distinguishable from the serratus profundus, and on this
account Fiirbringer [22] inclines to disallow its existence as a separate muscle.
The serratus superficialis of Apteryx retains the pars metapagialis, which is wanting
in the other Palwognathe.
The coraco-brachialis internus of Casuarius is entirely tendinous; in Rhea largely so.
Furthermore, in this last genus it extends on to the sternum. In Struthio it is larger
than in any other Palewognathe.
THIGH- AND LEG-MUSCLES3.
Struthio and Apteryx only have the leg-muscles complete, which make up the
formula AB. XY +.
In the Struthiones generally the accessory femoro-caudal (pars iliaca m. caud-ilio-
femoralis) calls for special comment. In all save Apteryx it is characterized by its
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 243
great size, in Struthio and Casuarius being rightly described as enormous. ‘The femoro-
caudal, on the other hand, is relatively but feebly developed or absent.
Rhea and Dromeus both agree in having lost the femoro-caudal.
gf. mac. art.
gt. Wax. foost.
Left side view of a dissection of the thigh-muscles of Casuarius unappendiculatus, to show the accessory
Struthious adductor, the femoro-caudal, and accessory semitendinosus muscles. ‘The accessory adductor,
femoro- and accessory femoro-caudal=the caud-ischio-ilio-femoralis of Gadow.
gl.min, = gluteus minor.
Ace.ad. = accessory Struthious adductor.
maximus anterior.
b. = biceps. gl.mxani. = 4,
f.c. = femoro-caudal.
ace.f.c. = accessory femoro-caudal.
s.t, = semitendinosus.
ace.st. = accessory semitendinosus.
ad.l, = adductor longus.
cr, = crureus.
gl.med, = gluteus medius.
gl.mx.post. = 4, posterior.
giant. = ,, anterior.
Sar. = sartorius.
smb, = semimembranosus.
ect.c. = ectocnemial crest.
ad.l, = adductor longus.
p.b. = pubis.
The femoro-caudal muscles of Apteryx are peculiar and are differently determined
by Garrod and Beddard. Garrod’s view is adopted in the present contribution, after
independent dissections. Garrod [30] says :—‘‘ The accessory femoro-caudal is peculiar ;
244° MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
for, on removing the biceps cruris, its superficial portion is seen running obliquely
upwards and forwards to the whole length of the linea aspera of the femur, from its
usual origin. The sciatic artery and nerve are superficial to this muscle (adductor
longus of Owen) and parallel to its insertion, as in most birds; but they, as is not the
case except in the Struthiones and Crypturi, perforate it at the ischiadic notch, leaving
a small portion of the muscle (the adductor brevis of Owen) above them. The anterior
terminal fibres of this muscle are situated external or superficial to the accessory semi-
tendinosus.
“‘ After this muscle has been removed or turned back, there is seen a deeper muscle,
which, if the one described above did not exist, would be justly considered to be the
femoro-caudal and the accessory femoro-caudal, part springing from the iliac ridge and
part from the coccyx, whilst both are inserted into the posterior portion of the linea
aspera and have the nerve to the semi-membranosus situated between them and the
adductor magnus.” ‘This deeper muscle Garrod does not further distinguish. Probably
it should be regarded as representing the obturator externus.
The femoro-caudal, it should be remarked, is very slender; it lies deep of the semi-
membranosus. In Garrod’s paper he appears to associate it first with the superficial
and then with the deeper muscle.
Mr. Beddard apparently regards the obturator externus of this paper, and the deep
layer of muscle described by Garrod, as the muscle which would be justly considered
the accessory femoro-caudal, in the absence of the superficial layer just described.
The muscular mass embracing the emergence of the sciatic nerve and artery
should be regarded, I would suggest, as the Struthious adductor referred to by both
Garrod and Beddard, but fused distad with the gluteus anterior, and caudad with
the accessory femoro-caudal. A comparison of fig. 6, p. 243, will serve to show the
probability of the correctness of this interpretation.
The semitendinosus and its accessory in Casuarius, Dromeus, and Rhea are very
similar. In Fhea the former is inserted on to the tibia by means of a long tendon,
in the Cassowary at least of the two former genera it loses itself in the gastrocnemius.
In Struthio the accessory head is small. In <Apteryx, according to Beddard, the
accessory head shows a slight variation in its relations within the genus. ‘In
A. australis the accessory head of the semitendinosus was distinctly separated from,
though parallel to, the middle head of the gastrocnemius. In A. haasti no such
separation is obvious.”
The semimembranosus (ilio-flexorius, Gadow) of Apterya is two-headed, the second
head arising from the ischium; between these two heads lies the head of the femoro-
caudal.
In Dromeus the semimembranosus, according to Garrod, is peculiar in that it has
an aponeurotic connection with the middle of the linea aspera.
In Casuarius it is but feebly developed ; it passes rapidly into a thin flat tendon to
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 245
be inserted into the shaft of the tibia, below the neck. As it passes the middle head
of the gastrocnemius it gives off thereto, at right angles, a delicate tendinous band.
In khea it is small, sharply truncated forwards, terminating distad of the ventral
border of the accessory semitendinosus. Its antero-dorsal angle fuses with the tendon
of the semitendinosus, distad of its accessory head. Its postero-ventral angle sends
downwards a long slender tendon to the gastrocnemius, to be inserted a short distance
above the tarsal joint.
In Struthio, according to Garrod, the semimembranosus tendon fuses with that of
the accessory semitendinosus and then passes down the leg as a long thin tendon to
the tendon of the gastrocnemius. Thus closely resembling that of Rhea.
The ambiens is wanting only, among the Struthiones, in Dromeus. Casuarius,
however, is generally regarded as wanting this muscle.
In Struthio its origin is peculiar, being from the ilium instead of the pectinal
process of the pubis.
In Apteryx its origin is from the pectineal process of the pubis; as usual, it is
inserted into the flexor perforatus digiti 1. It does not, according to Beddard, give
off slips to the other two muscles of the perforatus complex. ‘“ Furthermore, these
small tendinous bands, accompanied by muscle-fibres, and forming a thin, flat sheet of
tissue, are continuous with the ambiens tendon above, and appear to be in connection
at the other end with the short arm of the biceps sling, and to arise from the fibula.
I look upon this sheet of muscle and tendon as a second head of the flexor in
question, and as corresponding to the fibular head of birds, such as Nycticorav. If
this be so, Mitchell’s contention that the ligamentous head in question is a rudimentary
ambiens is not so certain as it seemed to be, for both occur in Apteryx. I found the
same state of affairs in A. haasti.”
In Rhea americana, Gadow [25] found the ambiens to be typically developed. This
I am able to confirm from my own dissections.
T would remark that in an adult of this species the ambiens tendon, at the level of
the head of the fibula, sent upwards a tendinous slip to the fibrous tissue of the lower
border of the tendinous insertion of the crureus and origin of the fl. perf. d. m1. and
perf. et perf. d. 1., and a few fibres to the tendinous portion of the origin of the
common belly of the fl. 1.-1v. and perf. et perf. 11.
In R. macrorhyncha he found that “the muscle arose from the latero-dorsal aspect
of the pubic spine, and at the same time from the big crural vein, the muscular and
aponeurotic fibres of the muscle having firmly got hold of the ventral aspect of this
vein. .... The tendon of the muscle passed the knee in the typical way.
“In R. darwini, 2, the m. ambiens of either side arose from the pubic spine as
usual, but its tendon, before reaching the knee, became flattened out and attached
itself with a broad fan-shaped and very thin aponeurosis to the patella, in a similar
style as the median additional portion of the m. femoro-tibialis (m. vastus).” In a
VoL. xv.—Part v. No. 18.—December, 1900. 2M
246 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
male FR. darwini the ambiens of the left thigh stopped at the knee, as in the female,
but in the right thigh the muscle was typically developed and passed the knee with a
strong tendon.
In Casuarius the ambiens, according to Garrod, is wanting. Concerning this,
Dr. Gadow, in describing the ambiens of hea, a portion of which we have just
quoted, continues, “ This abnormal condition of m. ambiens, which seems to be pre-
vailing in Darwin’s /hea, is the intermediate stage between a typically developed
m. ambiens and such forms in which, as in Casuwarius, this muscle has lost still more
of its independence, and then only forms an additional head of the median part of
the portio media m. femori tibialis s. vasti. Without an elaborate examination
and comparison of the formation of these muscles with their nerve-supply,
we should, with Garrod, come to the conclusion that Caswarius did not possess an
ambiens muscle. The assumption of still further reduction of the distal portion of
the m. ambiens explains what I have observed in some Passerine birds, ¢. g., in a
specimen of Lanius bentet, in which the m. femori tibialis internus, besides being
strongly developed, received in its proximal part a thin spindle-shaped semitendinous
head from the pubic spine. This additional little slip is probably the last trace of
the ambiens muscle, which is now generally lost by the Passerine birds.
“The case above described is one way in which this muscle gets lost; in other cases,
e. g., Ciconia and Phenicopterus, the reduction does not begin by its tendon becoming
attached to the neighbouring tendons in the knee-region, but the whole muscle shows
a diminution of its tendon and belly to a mere thread, till at last this also disappears,
é. g., in Abdiniia and Xenorhynchus.”
The muscle-formule of the Palwognathe is as follows :—
Strut hio
TERED GIN TSC NE dy,
Casuarius
Crypturi
Rhea B.X.Y +.
Dromeus IB REX Ys
The deep flexors of the foot of the Struthiones belong to the types IJ. and IV. of
Gadow.
Apteryx alone represents type II. ‘The vinculum is strong, and runs directly
downward to join the flevor profundus. ‘The flexor hallucis tendon is slender.
Struthio, Rhea, Casaurius, and Dromeus all belong to type IV. The separate
tendons of the jflewor hallucis longus and of the fl. perforans s. profundus can
be distinguished as far as the middle of the tarso-metatarse, when they completely
fuse.
Hexor brevis digiti 11., which arises from the tendon of the flexor profundus and is
inserted into digit 111., occurs only in Rhea amongst the Struthiones.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA, 247
NERVOUS SYSTEM.
THe BRAIN.
Parker has shown that, compared with the brains of many Neognathw—Turkey,
Emu, Goose, Pigeon,—the cerebral hemispheres in Apterya are proportionately of
large size, and further peculiar is the way they overlap the cerebellum, in this respect
bearing a very close resemblance to the Passeres. This overlapping conceals the
grooves which mark the anterior region of the cerebellum, the hemispheres extending
as far back as the flocculi. The posterior grooves are visible superficially, as in other
brains. The pineal peduncle, in consequence of this overlapping, is inclined from
the vertical position backwards.
The position of the thalamencephalon is also peculiar, in that this region is tilted
backwards, the lamina terminalis looking upwards, the foramen of Monro becoming
postero- instead of antero-dorsal.
THE Kye.
The pecten is well developed in all the Palwognathe, save Apteryx. In
Dromeus the number of its folds is reduced to four; in Struthio the lamelle are from
14-20 in number.
In Apteryx the pecten is generally believed to be wanting in the adult, though
present in the embryo. Dr. G. Lindsay Johnson, by a series of drawings recently
exhibited at a soirée of the Royal Society, has shown that this is not really the case.
‘The pecten is present in Apterya, though it assumes a peculiar form, being columnar,
not fan-shaped, and plicated. In all other birds the pecten has two or more plications
and is never conical—usually forming a vertical, plicated, densely pigmented septum
dividing the disc into a right and left half, although its base expands over it so that
very little of the disc is seen. The eye is small. Hence the apex of the pecten
penetrates the vitreous to a distance nearly equal to that of the posterior surface of the
lens from the retina.
Dr. Johnson’s researches in this direction have related almost exclusively to the
mammalian eye; and they have been most fruitful in results, enabling him to demon-
strate a precisely similar “ pecten” in the eyes of several Mammalia, that of the
Golden Agouti—as he showed by means of drawings, exhibited at the above-
mentioned sozée in June of this year—bearing a most striking resemblance to that of
Apteryx.
JACOBSON’S CARTILAGES.
These, according to Parker, are represented by a pair of vestigial rods of cartilage
lying parallel to, and either immediately dorsad or slightly laterad of, the dorsal edge
2M 2
248 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
of the vomer. It can be readily prepared in a well-macerated skull by carefully
removing the vomer. It was first described by W. K. Parker [77] in Rhea.
THE ALIMENTARY CANAL.
Buccat Cavity.
The buccal cavity of the Palwognathe presents one noticeable feature wherein it
differs from that of the Neognathe (Pl. XLV. fig. 1): since, in the former, the
posterior nares, or choanze, form a wide, open, more or less cordiform aperture, divided
by a median septum into right and left lateral moities; in the Neognathw the
choane are slit-like. The Eustachian aperture lies immediately caudad of the choane.
In all the Palwognathe the tongue is more or less vestigial. The glottis is also
a relatively wider aperture than in the Neognathe. In Casuarius the tongue has a
denticulate free edge.
THE CONVOLUTIONS OF THE INTESTINE.
Dr. Gadow, in his paper “On the Intestinal Convolutions in Birds” [26], considered
the flightless members of the Palwognathe a very heterogeneous group, because of
the great diversity in the length and arrangement of the main gut and in the develop-
ment of the ceca. “In none of these birds has it come to the development of closed
and well-developed loops of the mid-gut (with the exception of the duodenum). In
this respect they represent the lowest type amongst the recent birds..... Their
connections with recent Carinate are distant. Nearest of them to the latter comes
Apterys through more defined loops, and the Cryptwri seem to represent the link. .
All the Archwopalatine agree with each other in having the second loop right-handed
and the third left-handed; this is a feature which occurs again only in the Crypturi,
Galline, Opisthocomus, and in the Cuculide.”
Amongst the Neognathe, it will be remembered, Gadow was enabled to demonstrate
a greater harmony, the conyolutions of the intestines being always referable to one of
seven types. ,
Dr. Gadow’s conclusions, it should be remarked, are based upon a study of the
coiling of the intestines within the body-cavity.
Mr. Mitchell [60], who approached the study of the intestinal tract from a different
point of view, viz., after its removal from the body, and the severance of certain
vessels and secondary connections, which reveal the nature of the convolutions of the
gut along its attachment to the ventral border of the dorsal mesentery, and the
position of the diverticulum cecum vitelli and its relation to the middle mesenteric
vein, entirely confirms Dr. Gadow’s conclusions as to the primitive nature of the
Palwognathe judged by this standard. He remarks:—‘‘It is plain that, so far as
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 249
degree of divergence of type in the alimentary canal goes, the Ratites deserve their
accepted place at the bottom of the avian scale.”
I have been enabled to examine the intestines of all the Palwognathe, save
Struthio. So far as my material goes it entirely confirms the observations of Gadow,
Mitchell, and Beddard.
Casuarius, Dromeus, and Apteryx all resemble one another rather closely.
In Casuarius and Dromeus the duodenal loop is wide, in Apéerya narrow. In
Casuarius, as Mr. Mitchell points out, the pancreatic and hepatic ducts open into a
diverticulum of the distal loop. In Dromeus and Apterya the duodenum is longer and
narrower, and the pancreatic and hepatic ducts open separately into the duodenum.
In other genera the small intestine is relatively short, and the convolutions therefore
are simple. The rectum is short and straight; in Caswarius and Dromeus its inner
lining is thrown into a number of strongly marked irregular rugous folds,
In Casuarius and Dromeus the ceca are relatively short, of a uniform thickness
throughout, and have a narrow lumen. In Apterya the ceca are relatively much
longer, exceeding 6 inches in length, whereas in the two former they do not exceed
5 inches. Furthermore, in Apterysx the terminal half becomes suddenly and markedly
dilated ; each cecum thus presents a sausage-shaped free end, borne on a narrow tube-
like stem.
There are no valvular appendages in the ceeca of these three forms.
Rhea is intermediate between Caswarius on the one hand and Struthio on the other.
The duodenal loop is narrow and closed. ‘The convolutions of the small intestine
resemble Casuarius, the posterior rectal portion Struthio. ‘The pancreatic and hepatic
ducts open separately. ‘The resemblance to Struthio consists in the relatively great
length of the rectum ; but it differs therefrom in that it is only thrown into a single,
and not a series of complex loops.
The ceca are relatively of enormous length, exceeding those of any other Palwo-
gnathe. They appear, however, to vary in this respect very considerably. In the
series of measurements given by Remouchamps [85] the variations in 2. macro-
rhyncha range between | foot 44 inches and 2 feet 9 inches; in &. americana between
2 feet 4 inches and 4 feet 8 inches. In a female of this latter species dissected by me
they attained a length of 5 feet 3 inches, accompanying the small intestine throughout
the greater part of its length. ‘The lumen was sacculated to within a short distance
of the extreme end of the cecum by means of a series of semilunar valves alternating
one with another. ‘The position of these valves was indicated on the outside of the
ceecum by means of a series of constrictions (fig. 7, p. 250).
The intestinal coils of Struthio I have not had the good fortune to examine. I
therefore append Mr. Mitchell’s description [60]. He writes :—‘* The first two parts
of the intestine and the three great veins were according to type, the yolk-sac
diverticulum being conspicuous in the circular loop opposite the end of the median
250 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Cxeca of Rhea americana.
s.2.=small intestine.
¢.= ceca.
1.i.=large intestine, forming 7.l.=rectal loop.
mesenteric vein. The distal limb of the duodenum presented a short lateral diver-
ticulum, and the ceca are relatively longer than in Caswarius, and, as has been
described frequently, were marked by the attachment of a spiral valve. But the rectal
part of the intestine, that supplied by the posterior mesenteric vein, is expanded into
an enormous coil swung at the circumference of a semicircular expansion of its
mesentery. Only in Chawna, and in the Eagles and Petrels, have I found the slightest
trace of a convergent resemblance to this feature, but in the latter the subsidiary rectal
loops, although supplied by the rectal vessel, lie above the ceca.”
According to Beddard the spiral valve of the ceca contains about twenty turns.
These ceca, he points out, are furthermore remarkable in that they open into the
rectum by a common orifice, thus differing from those of all other birds.
The ceca of the Tinamous resemble those of Casuarius ; Calodromas elegans,
however, affords a remarkable exception, differing as it does in this matter from all
other known birds. These caca have been described and figured by Beddard [6].
They are much wider than usual, and have the outer walls produced into numerous
prominent diverticula decreasing in size from behind forwards.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 251
The following table from Beddard [7] will give at a glance the known variations in
the proportions of the alimentary canal :—
| Small intestine. Large intestine. Creea.
ft ins. ft. ins, tty) inss ft. ins. |
LEVEE KTROROYLG, @ ncconcogcosece 2 i @ 1 43 Less) ne
; - ET eee sty Mt 58 il iil 2 9 |
f a AC hes 8. 5 0 1 4 2 0 |
Rhea americana ............. ewe 6 | i 868 Dh Al by 8) |
* a Oy nteenun Catenin as a) | 2 2 fh |
% EMR CVOUNG et: or ettees SOR me ioe nine off eons ||
SRO CHAE, go anccaoconcse 23 32 «(9 By 3} 2) Ail
ae I-10) ab, Sn 2) ae Aa ed 93 4 | 30 8 2D NOH
; aS ONS er vaarsitge Riareun oe rete Ate 23) 0 8} Den
| x i Gil aia eine sect eaets 24 «6 SLs 2 8 3
Kioto Pee etree FR Be aL 278): 23 0 24 9 1 a6)
are PRR OLB Ae ng eeepc aN > | * 6 | 8 2B 2 7
Casuarius unappendiculatus, 9.1... | a | 10 43 5
6 casuarius beccarit, Q «1... 4 8 | 1 0 42
55 POELCHOS, Ci b0000000000008 a0) | 10 4
o bicurunculatus ........-+..| a. OU | 0 7
3 UDOIGHON G+ piaamanenooeolos | 3 103 113 32 33
| Dromeeus nove-hollandia ............ 10 «6 0) 5
| Apterya australis 20.004. esses esses AA 47 7
a OW CRIS WII a TL oe 3. 4} 43 7
|
Tue Liver.
The liver, in all the Palwognathe, save Struthio, agrees with that of the
Neognathe in that it is divided into two lobes.
In Casuarius and Dromeus the right lobe is larger than the left. A Spigelian lobe
is represented.
Both Casuarius and Dromeus possess a gall-bladder. In the former the gall opens,
in common with the pancreatic ducts, into a diverticulum on the distal loop of the
duodenum.
In Apteryx the left liver-lobe is larger than the right; the gall-bladder is occasion-
ally absent.
In Rhea the right and left liver-lobes are equal in size. ‘The gall-bladder is
generally wanting.
252 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Struthio is remarkable in that, according to Beddard, the right and left lobes are
fused into a single heart-shaped lobe. There is a Spigelian lobe. The gall-bladder
is wanting; and there is but a single bile-duct, which opens near the pylorus.
Mr. Beddard found the free edge of the liver in Rhynchotus rufescens pierced by
three small vessels of the venal portal system—a fact which, as he points out, has a
very lizard-like appearance.
RESPIRATORY ORGANS.
a. THE Lunes (Pl. XLV. fig. 9).
The costo-pulmonary muscles in Caswarius are short, thick, and powerful, five pairs
in all; they arise from the thoraco-sternal articulation of the ribs and extended fanwise
on to the aponeurosis covering the lung. Those of Rhea are relatively longer than in
Casuarius. In Apteryx, according to Huxley [41], they form “ broad flat bands,” which
‘take their origin from the vertebral ribs, at some distance below the attachment of
the pulmonary aponeurosis, and, proceeding obliquely dorsad and forwards, spread
out and are inserted into that part of the aponeurosis which covers the posterior
inferior facet.
In the relative size and distribution of the ecto-, ento-, and para-bronchia, and in the
size of the vestibule, the flightless members of the Palwognathe do not appear to
differ much one from another or from the Neognathe.
In the lungs of a Casuarius unappendiculatus the bronchial rings extended backwards
as far as the 4th ento-bronchium. ‘The inner border of the anterior and posterior
inferior facets of the lung are deeply sculptured by a system of numerous parallel
grooves of open tubes, closely packed and running along the surface of the lung from
the mesial border outwards so as to converge towards, and open into, a spacious
chamber communicating posteriorly and ventrally with the Ist ento-bronchium
(Pl. XLV. fig. 9), further forwards and also ventrally into the anterior end of the
pre-bronchial ostium. These grooves give the ventral face of the lung, after the
removal of the aponeurosis, the appearance of having been bored by 'Teredos, the groove
representing a hemisection. ‘They are crossed and recrossed by transverse strands of
tissue and punctured by minute apertures.
They may be divided into two groups according to whether they are fed by a trunk
from the Ist or the 3rd ento-bronchium. Thus, as will be seen in fig. 9, all the
secondary grooves cephalad of the main trunk marked “ent. 1.” are fed from this, whilst
ento-bronchium 11. supplies all the grooves caudad thereof.
These grooves are converted into tubes by the pulmonary aponeurosis.
b. Tur Arr-sacs (Pl. XLV. fig. 8).
In Casuarius the pre-bronchial sac is paired, and extends forward on each side of
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATHA. 253
the cesophagus for a considerable distance beyond the level of the Ist cervico-
thoracic rib.
The sub-bronchial sac is divided into right and left loculi, enclosing a median
tunnel-like space between them, the roof of which is formed by the trachea. This
tunnel can be traced into a large pneumatic cavity in the sternum Opening on its
anterior border in the middle line between the coracoid grooves.
The intermediate sacs were three in number. A similar subdivision has been de-
scribed by Beddard in Podargus and “one or two Accipitres.” In the present instance
it would seem that this third sac is due to a subdivision of the anterior intermediate.
The posterior intermediate was large. Its roof was pressed inwards by the forward
extension of the posterior abdominal.
The posterior abdominal was of considerable size. That of the right side enclosed
the right kidney; on the left side the anterior lobe of the kidney was found to lie
without the sac and to be enclosed in a separate sac.
The intermediate and posterior abdominal sacs of the right side were markedly
smaller than those of the left.
The sacs are fed as follows:—The pre-bronchial and sub-bronchial by a common
ostium from ento-bronchium I. ; the anterior intermediate (a) from ento-bronchium II.,
and (6) from the anterior of two canals arising one immediately behind the other from
the ventral floor of the meso-bronchium immediately below the apertures of the
mn-Iv. ecto-bronchia. The ento-brenchial ostium opens into the anterior, the
ecto-bronchial into the posterior loculus. ‘The posterior intermediate by the hinder
of the two hinder ecto-bronchial canals. The posterior abdominal by the terminal
end of the meso-bronchium—the ostiwm posterius.
The 2nd ento-bronchium does not open, as described by Huxley in Apterya, into
the sub-bronchial ostium, but turns upwards and backwards, and terminates cecally
like the 4th.
The air-sacs in Rhea differ in no essential respect from those of Casuarius.
The sub-bronchial sac is unpaired. In one Rhea americana which I dissected the
anterior intermediate sac was fed by two ostia widely separated one from another. In
a second example the ostium of the anterior intermediate sac of the left side and the
septum dividing this sac from the posterior intermediate were both wanting.
The posterior abdominal sacs are much larger than those of the Cassowary.
Apteryx differs from all other birds, as was first pointed out by Huxley, in that,
owing to their smaller size, the posterior intermediate and abdominal sacs are enclosed
within the sub-pulmonary chamber. In all other birds a great increase in the size of
these two has driven out the posterior abdominal sac, so that it projects, like a
hernial sac, between the peritoneum and the parietes into the abdominal cavity.
The respiratory organs of the Palwognathe are singularly uniform in character
inter se and with the Neognathe.
VoL. XV.—PaRT v. No. 19.— December, 1900. 2N
254 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
c. THe WINDPIPE.
The most valuable of the numerous treatises on the windpipe of the Palwognathe
—from a systematic point of view—is that by Forbes [20]. We are concerned here
only with the thoracic end of the trachea, and the degree of specialization which it has
reached in the direction of the formation of a “ syrinx.”
Struthio and Apteryx, up to the time of Mr. Beddard’s paper [8], have been held
to represent the least specialized forms in this particular.
In Struthio there are no intrinsic muscles. There is a membrana tympaniformis
interna, but only a trace of a membrana semilunaris. ‘ Internally,” according to
Forbes, ‘the mucous membrane of the interior is greatly thickened, forming a vocal
cord, in the region of the last three tracheal rings and first two bronchial semirings.
The lateral tracheal muscles stop at the point of insertion of the sterno-tracheales.”
In Apteryx there is a broad membrana tympaniformis interna and a feeble
membrana semilunaris. As in Struthio there is no pessulus and, it was believed, no
intrinsic muscles.
Mr. Beddard’s dissections, however, showed that these were ‘especially plainly
developed in Apteryx australis.” In this species he found “a thin band of muscle
continuous above with the extrinsic muscle; this passes down the side of the trachea
and gradually thins out into fibrous tissue, which seems to die away upon the last
tracheal ring, hardly reaching the bronchi. Its state, in fact, is clearly one of degenera-
tion, and very much on a par with the state of the corresponding muscle in the
Hoatzin (Opisthocomus). In A. mantelli the same muscle was visible, but it did not
Seems ir to be anything like so well developed as it is in the species first referred to.”
It appears to be entirely wanting in A. haastii and A. oweni.
There is a feeble vocal cord, and the lateral muscles stop at the insertion of the
sterno-tracheales.
Casuarius, as Forbes remarks, shows one interesting peculiarity, the lower end of
the trachea becoming slightly dilated above the bronchi. Furthermore, the last
tracheal rings are incomplete behind, the median space to which this gives rise being
filled up by fibrous and elastic tissue arranged transversely. By the contraction of
this, after death, the ends of the rings are drawn inwards, so forming within the
tube a low longitudinal ridge, resembling the tracheal septum of certain Petrels
and Penguins.
There is no pessulus, the membrana tympaniformis interna passing over from one
bronchus into the other. A very slight thickening of this membrane in the middle
line between the two bronchi suggests a membrana semilunaris, but, as Forbes
remarks, there is nothing that can properly be called by this name. The outer vocal
cord is well developed.
Forbes describes, but so far I am not able to confirm this, “On the internal wall of
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 255
the! bronchus) 75.0 a second much slighter and less prominent fold, slightly concave
forwards, running somewhat obliquely backwards and downwards, and supported by
the anterior ends of the third and sixth bronchial semirings.”
There are no intrinsic muscles. The sterno-tracheales in Casuarius and in Dromeus
expand so as to meet in the middle line in front.
The tracheo-bronchial region of the trachea in the adult Dromwus has not yet been
described. In a young specimen dissected by Forbes there seemed to be no great
differences between it and Casuarius. The membrana semilunaris seemed to be better
developed.
The trachea of Dromwus is peculiar, in that its front wall is pierced by a long,
vertical, slit-like median aperture, caused by the deficiency of certain of the tracheal
rings in this region. The lining membrane of the tube projects from this opening
and forms a large dilatable sac, which can be filled with air at will. It is probably
concerned with the production of the drumming sound which this bird occasionally
makes. It has been described in detail and figured by Murie [63]; but seems to have
been first discovered and described by Peter Fremery [21] in 1819.
In Apterya, it will be remembered, the syrinx is but feebly developed, and the intrinsic
muscles are degenerate. The intrinsic muscles of Rhea were first correctly described
by Forbes [20]. The last four or five, or even six, tracheal rings fuse more or less
completely to form a box. The number of rings involved in this fusion can generally
be traced by the interannular sutures, which appear never to be completely obliterated.
There is a complete and cartilaginous pessulus and a membrana semilunaris. Forbes
describes, and I am able to confirm, ‘‘on the middle part of the membrana tympani-
formis interna a thickened fibro-cartilaginous area.” ‘This is triangular in form, its
base resting on the pessulus, and its apex terminating near the 4th bronchial semiring.
It would be interesting to compare this thickened tissue with certain fibrous pea-
shaped pads found in this region in Sula bassana, first described by Garrod, and
Catarrhactes chrysocome, described and figured by Watson in the ‘ Challenger’ Reports.
The broncho-trachealis (intrinsic syringeal) muscle in my dissections, as in Forbes’s,
arises from the last ten tracheal rings, and passes downwards to be inserted into the
anterior ends of the last four or five bronchial semirings and the membrane between.
The sterno-trachealis terminates just pre-axiad of the origin of the intrinsic muscle.
In the Casuariida, as Forbes has pointed out, the “ terminal part of the trachea tends
to develop into an expanded tympanum”’; the tracheal rings are softer and become
imperfect posteriorly, the junction of the trachea and bronchi is less marked, and the
arrangement of the sterno-tracheales is peculiar.
In Struthio and Apteryx the tendency of the trachea is to become narrower at its
termination, and the distinction between this and the bronchi is much sharper than in
Caswarius.
2Nn 2
256 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
d. THe Cnoana (Pl. XLV. fig. 1).
The choane of the Palwognathe, it is interesting to remark, differ in form from the
Neognathe in their larger size. In all the Neognathe the aperture is slit-like, whilst,
as will be seen in the figure, in the Palwognathe it is triangular or cordiform in
shape.
CIRCULATORY SYSTEM.
Tue Heart.
The heart of the Palewognathe does not differ materially in structure from that of
the Neognatiw. It has been already ably described and its most important features
pointed out by Owen [67], Lankester [45, 46], Beddard [5], Rolleston [87], and others,
so that little remains for me to say.
Fig. 8.
A. ace. t. SCp. w
weet
SS
A.—Dissection of the heart of Causuarius wnappendculatus, to show the right auriculo-ventricular valve and
the ‘‘ moderator band.”
B.—--Dissection of the heart of Rhea americana, to show the right auriculo-ventricular band and the abnormal
septal valve.
ace.t. = accessory tendon.
sep.v. = septal portion of right auriculo-ventricular valve.
s.v. = semilunar valves.
r.du.v. = right auriculo-ventricular valve.
p. = papillary muscle.
mod.b. = moderator band.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 257
My only point concerns the moderator band first described in Casuarius by
Rolleston. It seems to have been taken for granted by subsequent writers that this
was always present, if not in all the Palwognathe, at least in the Casuariide.
I have only succeeded in finding it once, and in the heart of a Casuarius casuarius
salvadorti, though I have examined specimens of C. wnappendiculatus, C. casuarius
heccarit, C. casuarius violicollis, C. casuarius sclateri, and Casuarius casuarius,
In addition to the large papillary muscle binding the right auriculo-ventricular
valve to the anterior wall of the ventricle, 1 find two or more very slender “ chorde
tendinez’’ extending from the outer surface of the auriculo-ventricular valve to the
anterior ventricular wall, recalling the similar structures described and figured by
Lankester [45] in the heart of Ornithorhynchus. In Rhea and Dromeus only one of
these threads was present. I have, however, only had the opportunity of examining
one Dromeus for this purpose.
The septal half of the valve in Casuarius is thick and fleshy. In Dromeus it is
relatively somewhat larger, and, in the only adult heart which I have had an
opportunity of examining, differs from Caswarius in that its free ventral or hinder
border is curved forwards at its insertion towards the floor of the left auricle,
forming by this means a sort of pocket-like flap on the wall of the inter-ventricular
septum.
In Rhea this same arrangement occurs again. In one adult which I dissected
(fig. 8B) this was remarkably well developed. The valve was very thin, and the
forward pocket-shaped extension into the anterior end of the ventricle was strongly
developed and was continued yet further in the form of a narrow slip of muscle
attached only at its ends.
In Dromeus, in addition to one or two delicate tendinous threads from the outer
surface of the auriculo-ventricular valve to the anterior ventricular wall, there were
numerous fleshy strands of muscle, interlacing one another, running parallel with the
valve, and from it to the anterior wall of the ventricle.
- Trabecule carne were well developed both in Rhea and Dromeus.
CAROTIDS.
All the Palwognathe, save Apterya, have normally two carotids. Occasionally, in
Rhea, the left only persists. In Apteryx there is only one carotid, the left.
CLOACA, BURSA FABRICI, AND URINO-GENITAL SYSTEM.
a. CLOACA.
The terminal end of the alimentary canal in the Palwognathe, as inthe Neognathe,
is divisible into a copro-, wro-, and proctodewm, as defined by Dr. Gadow [24].
In Casuarius, Rhea, and Apteryx the coprodeum passes gradually into the rectum,
but in Struthio and some Neognathe the coprodeum is preceded by an additional
258 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
chamber, though this is but small and not very sharply defined. In this succession of
chambers therefore, as Dr. Gadow has pointed out, Struthio resembles some Saurians
and young Crocodiles.
In Struthio, defecation and micturition are separate acts, and the urine, instead of
passing from the uro- to the coprodeum, is transferred to the bursa Fabricii, which,
unlike that of all other birds, is of considerable size and remains permanently open.
Its function, acting as a physiological dorsal bladder, is unique. A true allantoic
bladder, communicating directly with the ureters, is wanting in the Aves.
The cloacal aperture of the Palwognathe, as in Neognathe, is round, not transverse
as in Reptilia.
Fig. 9.
The terminal end of the alimentary canal of Caswarizs violicollis, to show the cloaca and its divisions
into copro-, uro-, and proctodeum and the form and position of the bursa Fabricii.
b.f. = bursa Fabricii.
c.d, = coprodeum.
u.d. = urodeum.
p.d. = proctodzeum.
cl, = clitoris.
ovid, = aperture of oviduct.
u. = aperture of ureter.
6. Tue Bursa.
The bursa Fabricii of the Palwognathw has been very carefully studied by Forbes
[20] and Wenckebach [97].
In the Paleognathe—save only in Apteryx—as Forbes [20 4] has shown, the
bursa in young birds is of considerable size. Furthermore, he held that it differed
PHYLOGENY OF THE PALZOGNATHA AND NBEOGNATHA. 259
fundamentally from that of the Neognathw, since in these last it opens by a more
or less constricted aperture into the general cavity of the cloaca, whilst in the
Struthiones the exact opposite was the case, the cloaca opening into the bursa.
oH rompthisewaryer it will be at once evident that the cloaca does not open directly to
the outer surface, but indirectly through the bursa by means of its large posterior and
inferior aperture.”
I find myself unable to confirm this point, and I think the accompanying figure will
justify me. The only difference which I can see between the Palwo- and Neognathe in
this particular is that in the latter the bursa opens into the roof of the proctodeum by
means of a minute aperture, in the former it opens by a very large one, so large as to
absorb nearly the whole roof of the proctodeum.
When these parts are dissected from spirit-specimens, in which hardening has taken
place in situ, Forbes’s description seems to be amply verified, as I can testify. The
accompanying figure was made from a specimen in which the hinder end of the gut
had been removed soon after death, the cloacal aperture set widely open, and the
whole plunged into formol and left for two or three days. The wall of one side was
then removed and the true relations revealed.
The function of the bursa is yet unknown.
c. CopuULATORY ORGANS.
The Paleognathe all possess a well-developed copulatory organ.
That of Struthio is a solid grooved organ containing erectile tissue and resembling
very nearly that of the Chelonia and Crocodilia. It can be retracted into a
pouch in the floor of the proctodeum. The m. protractor penis receives a slip from
the transversus analis, which descends from the pelvic bones. A pair of retractores
penis arise from the pelvis and are attached to the ventral aspect of the middle portion
of the organ.
In the remaining Palwognathe the penis resembles that of the Anseres amongst
the Carinatwe, being evertible and more or less markedly spirally twisted and grooved
dorsally.
The copulatory muscles of the Palwognathe, Dr. Gadow [24] points out, differ
from those of the Neognathe, inasmuch as the latter are derived from the sphincter ani
solely, whilst in the Palwognathe they are partly differentiated from muscles which are
still attached to the pelvis, and are therefore skeleto-genital.
DEVELOPMENT.
The egg of the Palwognathw, save in Apteryx and perhaps the Crypturi, differs from
that of the Neognathw in that the pore-canals are branched. In Apteryx, as in the
Neognathe, they are unbranched.
In Struthio, according to Nassonoy [64], and Apteryx, according to Parker [71], an
260 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
opercular fold grows backwards from the hyoid arch so as to cover the first and second
branchial clefts. This has been noticed in some Neognathe. Sushkin [94] has figured
and described it in Falco tinnunculus. ‘The first discovery of this operculum appears
to have been made by Rathke [84] in 1832.
PHYLOGENY OF THE PALAOGNATH A.
I enter upon the discussion of this subject with very considerable trepidation—not
only on account of the difficulty of the problem, but also because it brings me into
direct conflict with the opinions of those who speak after deliberation and long
experience, and are, for this reason, and rightly, accounted authorities amongst us.
It is now generally admitted that the Palwognathe must be descendants of an
Avian proto-carinate form capable of flight.
Parker [71] has given a number of characters which lend the strongest support to
this view. ‘To many they will be convincing. ‘The most important of these are :—
‘The fact that the skeleton of the fore limb is that of a true wing.”
“The presence of an alar membrane or patagium.”
“The presence of remiges and tectrices majores.”
The alar membrane is a structure of great significance. According to Parker, it is
the remnant of a once extensive scale-covered patagium. As these scales gradually
became transformed into feathers the size of the patagium decreased.
As to the mono- or polyphyletic origin of the group there is much divergence of
opinion.
Newton and Beddard consider the Palewognathe to be monophyletic; Fiirbringer,
Gadow, and Parker polyphyletic. With these last I also agree, but not entirely so.
There can be no doubt but that the Palwognathe represent a lower grade than the
Neognathe. Nevertheless the latter are closely related thereto.
It is probable that they have been derived as a diverging branch from that stock
which gave rise to the Rhetde, Dinornithide, and Atpyormithide. The diagram
(fig. 10, p. 265) is intended to make this clear.
The very close resemblance of the palate of the Crypturt to that of Rhea and
Dinornithide and the general resemblance in the form of the pelvis of Crypturi
and Dinornithide lend strong support to this view.
The structure of the pelvic arch in the Palewognathe is of considerable interest,
inasmuch as it bears upon the relation of this group to the Neognathe.
It retains its most primitive form in Archwopteryx, Dinornis, Aipyornis, Hesperornis,
Ichthyornis, Apteryx, and in the Crypturi. In all these the ischiadic fissure and free
pubes are retained throughout life. In Meognathe the ischiadic fissure becomes
converted into a foramen by synchondrosis between the ilium and ischium. In
Struthio the ischiadic fissure is permanent, but the pubes fuse in the middle line.
PHYLOGENY OF THE PALAOGNATHA AND NEOGNATH. 261
In Rhea the ischiadic fissure is closed; the ischia, furthermore, meet one another
below the vertebral column, and the pubes fuse with them posteriorly. Thus Séruthio
and Rhea are each unique in the form of the pelvic arch.
Struthio seems to be a stumbling-block. Thus Prof. Newton writes [65]: “Some
systematists think there can be little question of the Struthiones being the most
specialized and therefore probably the highest type of these Orders” (‘ Zatite”).
Others, in spite of the great amount of specialization which Struthio has undoubtedly
undergone, would regard this form as the most primitive, not only of living birds, but
of the Palwognathe living and extinct.
According to Furbringer, on account of the great number of its primaries, its
coracoid, and many of its shoulder- and arm-muscles, the form of its pelvis, two-toed
foot, ‘‘gewisse Eingeweide, etc., entfernen sie weiter von allen anderen lebenden
Vogeln, als diese in der Regel unter einander divergiren. Zugleich bietet Struthio
neben verschiedenen mehr vorgeschritten Specialisirungen progressiver und retrograder
Natur eine Anzahl héchst primitiver Charaktere dar, die ihre tiefer als die anderen
Ratiten stellen und durch welche diese sehr alte Form Blicke bis in die friihesten
Vorzeiten des Vogelstammes thun liisst.”
Beddard considers ‘‘that Struthio is removed far from the Dinornithide, as well as
from other Ratites, by the structure of its palate, which diverges much.” He does not,
however, go as far as Fiirbringer, for he continues further on: “There are really,
however, not a large series of characters in which they may be fairly said to be more
primitive than some other groups..... ” This last statement is evoked in answer to
Fiirbringer’s contention that Struthio is really very primitive.
I find myself in agreement with Beddard: more than this, I believe that we have to
look to Dromeus and Not to Struthio for the most primitive of living birds.
This opinion is based mainly upon a study of the bones of the palate in this and the
allied forms. These in Dromeus seem to represent the ground type from which, or
from some closely allied form, the palates of Struthio, Rhea, and Dinornis may be
derived. ‘The palate of Apterya must be considered by itself.
The palate of Struthio compared with that of Dromeus will be found by no means
so fundamentally different as seems to have been supposed. On the contrary, it is
contended that the palate of Struthio is a specialized and, in some respects, retrograde
form of that of Dromeus.
The palato-pterygoid connection is similar in both.
The palatine and the forward extension of the maxillo-palatine processes are both
considerably longer, both actually and relatively, in Séruthio than in Dromeus.
Struthio lacks palatine processes to the premaxilla, and has a degenerate, one might
almost say vestigial, vomer. Like all degenerate structures, however, it gives some
certain signs of a former perfection. Thus, in a skull which in this paper is attributed
to Struthio meridionalis the vomerine limbs extend backwards to within a very short
VOL. XV.—ParRT y. No. 20.—December, 1900. 20
262 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
distance of the pterygoid. If this proximal extension were carried back sufficiently far
to enable the vomer to join the pterygoid the resemblance to the Dromeine type would
be really striking.
If we turn now to the palate of Rhea, and compare this with that of Dromeus, we
shall find the differences more decidedly marked. The peculiar and almost complex
relations which obtain between the palato-pterygo-vomerine articulations have already
been described in these pages (p. 206), and are sufficiently distinct to render further
comparisons unnecessary.
But the palate of Rhea is curiously similar to that of Dinornis and the Tinamous.
So similar as to render it probable that these are related more closely one to another
than to Dromeus or Struthio.
The affinity of Rhea to the Dinornithide has already been hinted at by Beddard,
though upon different grounds. In discussing the position of Struthio, he remarks that
though “ this is removed far from the Dinornithide, as well as from other Ratites, by
the structure of its palate, which diverges much,..... it is not clear that Rhea is so
remote; the existence of an apparent homologue of the maxillo-nasal bone... .. isa
point of somewhat striking likeness to Hmeus.”
Nathusius, again, in studying the egg-shells of Rhea and Dinornis, was so impressed
with the likeness between them that he proposed to unite them in the same genus.
Parker [73], however, strangely enough, remarks: ‘I know of no character in the
skull of Rhea by which it definitely approaches the Moas..... ” T say strangely
enough advisedly, for I feel sure that had Parker’s attention been drawn to the points
to which attention is directed in this paper he would have grasped its significance and
written quite otherwise.
About the palate of Apyornis we as yet know nothing. In the roof of its skull it
resembles the Dinornithide, inasmuch as, just as in this group, there are no out-
standing supra-orbital processes to the lachrymal. In some other points it resembles
Struthio; but since in its pelvis and other points it closely resembles Dénornithida,
I am inclined to place it near this group.
The palate of the Tinamous is truly Dromwognathous, but approaches the Neo-
gnathine type, tending towards schizognathism. In its pterygo-palatine articulation,
as I have already pointed out (p. 208), it closely approaches the Meognathe, the
palatine having shifted from contact with the body of the pterygoid to form a
connection more or less intimate with its tip.
The palate of Apteryx in the peculiar forked pterygoid and the complexity of the
relations between it and the palatine and vomer seems to differ markedly from all the
other Palwognathe, or, indeed, from all living birds.
Here, again, is a conclusion diametrically opposed to that of Parker. Whether or
not I failed to appreciate the points of resemblance Parker claims must remain to be
seen. This discrepancy is certainly remarkable, and will seem to cast doubt upon the
PHYLOGENY OF THE PALZOGNATHEH AND NEOGNATHA, 263
4
validity of my conclusions rather than his, for those who know his monographs on the
development of Apteryx and the skull of the Dinornithide will agree that they are
monuments which mark an epoch in the history of this subject. In his work on the
skull of the Dinornithide he wrote: ‘The marked differences between the Moas and
Kiwis are certainly for the most part adaptive... .. The real affinities underlying
these differences are, however, shown by the striking similarity of the bones of the
palate in the two forms.”
The skull of Caswartus, it may be remarked here, differs in no essential respect from
that of Dromaus. In whatsoever it differs from Dromeus point to specialization, as,
for instance, the development of the casque upon the mesethmoid (p. 199, Pl. XLIV.
fig. 3) and the small quadrato-jugal fossa.
The skull of Dromeus, it would seem, must be regarded as the most generalized of
living birds, at least in so far as the bones of the palate are concerned.
Struthio somewhat nearly approaches Dromeus in the arrangement of these bones,
fthea, the Tinamous, the Dinornithide, and probably the Apyornithide agree more
closely one with another than with Dromeus.
Apteryx differs from all in the direction of increased complexity of these parts.
We have now Struthio and Dromeus with Casuarius opposed to Rhea, Dinornithide,
and Crypturi, and possibly Apyornis. ‘These last we may further subdivide by
means of the pelvis. This will separate AKhea from the remaining forms; inasmuch
as in Rhea the pelvis is long and narrow, with the post-acetabular ilia meeting in the
middle line, as in Apteryx.
In the Dinornithide, Mpyornis, and the 'Tinamous the post-acetabular ilia are
separated by the long transverse processes of the synsacrum.
It may seem that this character of the pelvis is a somewhat artificial one, but
reflection will show, I think, that it is probably not so. It does not seem to belong
to the category of adaptive characters, since Rhea and Apteryx in the general confor-
mation of the pelvis agree, though their habits are very different, and both bear some _
similarity to that of Struthio—similar in so far as the great length and transverse
width are concerned. In the Tinamide, Dinornithide, and Afpyornis the pelvis owes
its great breadth to (1) elongated transverse processes, and (2) the broad dorsal plane
of the post-acetabular ilium.
The pelvis of Struthio bears an undoubted resemblance to the Dinornithine form,
inasmuch as, like Dinornis or Atpyornis, the transverse processes bear the post-
acetabular ilia away from all contact with the neural spines of the synsacrum.
Fiirbringer’s view with regard to the Dromeide is not exactly in harmony with
the views adopted here, but it lends some support thereto nevertheless. He says
Dromeus and Casuarius “bilden zwei sehr nahe verwandte Familien..... welche
etwas hoher als die Struthionidw, aber tiefer als die Rheide stehen und im Ubrigen
sowohl von den anderen Ratiten ..... als von den Carinaten eine entfernte
202
264 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Stellung einnehmen,” and adds: “Ihre Vorgeschichte kommt mir am meisten dunkel
By this time the tendency of the foregoing will no doubt have revealed itself. Its
intention is to suggest that Dromeus and Casuarius—the Dromeide—are to be
regarded as the most primitive of the Palwognathe ; and that the Struthionide are
most probably the nearest approach thereto, for, in spite of the great specialization
which they have undergone, they undoubtedly retain many archaic characters.
The Rheide, Dinornithide, Aypornithide, and Crypturi are all probably more
nearly related one to another than to the Dromeidw. The Apterygide are the most
aberrant of all, but have affinities with the Dinornithide.
This implies that the Palwognathe are to be regarded as polyphyletic—probably
triphyietic. An attempt to express this relationship is given in the diagram on
p. 265.
I propose to discuss the details of this “tree” in so far as concerns the Neognathe
elsewhere. Of the Palwognathe I need only say I recognize 7 Orders, viz. :—
Casuaril.
Struthiones.
Rhee.
Crypturi.
Dinornithes.
Epyornithes.
Apteryges.
Their relation to the other members of the Class may be expressed as follows :—
Class AVES.
Subclass I. ARCHAORNITHES.
» Il. NEoRNITHEs.
Grade A. Palwognathe.
» B. Neognathe.
It would have been helpful here to discuss the further evolution of this group to its
final end in the various genera and species, but this is unfortunately impossible in
the present contribution owing to lack of material. The diagram will give a slight
indication of the writer’s views on this subject.
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATH &, 265
Fig. 10.
Pici f
f f Passeres
Trogones’ f f
Colii
Coraciss
Bucerotes
Accipitres
Cesena i Columbs
H | # Pterocles
Steganopodes Af i y Aisas
Musophagi if Cathartes 5
= Limicola,
Palamides
Gruidss
=
FS Rallida
S Hurypygide
Mesitides
V
Y
Y
VY
Y,
47
iY | LF
Yf WA Tubinares
‘GM A
UY
( A == Colymbt (including Hesperornis)
¢}
4
H
H
ESS
)
A
Yj Sphenisci
Z ~
IA,
LF
Dinornithes
»))
4 pyornithes
Z A
A“ Za
A
LY Za
tity LS”
4
Apteryges
Phylogenetic diagram showing the probable Relationships of the Palwo- and Neognathe.
|||| == Palewognathine stock. 38e = Eutaxic forms. Z = Diastataxic forms
Blank branches indicate eutaxic forms amongst a diastataxic group; blank spaces in a branch that both
eu- and diastataxic forms occur in the same genus or family,
266 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
SUMMARY.
The group of birds which we collectively designate the “ Ratita” is the same as that
to which this name was originally given’ by Merrem in 1813 (with the addition of
Apteryx, then unknown), on account of the fact that all agreed in the absence of a keel
to the sternum. They were “raft-breasted,” in contradistinction to the Carinate, or
keel-breasted !.
Nitzsch, Huxley, Newton, Sclater, and Sharpe have successively adopted Merrem’s
subdivision of the class Aves—adding the Saururee, unknown to Merrem.
Other workers, both before and since Merrem, have accorded to the “ Ratite” no
greater prominence than that of an order or suborder of the class Aves. Sometimes
even this was considered too great a distinction, and they were reduced to the rank of
a Family, including such forms as Bustards, Plovers, &c.
The “ Ratitee” of Merrem were apparently regarded by that author, as well as by
Huxley, Newton, Sclater, Sharpe, and other systematists, as a monophyletic group.
Fiirbringer, Gadow, and Parker stand prominently forward as advocates for a poly-
phyletic origin.
Gadow’s views are admirably set forth in his contribution to Bronu’s ‘'Thier- Reich ’
[26]. Briefly, he regards the “ Ratitee” as a morphologically monophyletic group,
standing in the same relation to the class Aves as the Anthropomorphe to the rest of
the Mammalia. /
With this view we entirely agree. We venture further, and include the Crypturt
within the group. This was done long since by Garrod [30] and Stejneger.
Beddard [7], later, has shown a strong inclination to follow suit. Thus, he says:—“ It
is unquestionably to the Struthiones that they showed the greatest number of important
likenesses, so much so, indeed, that their inclusion in one great group with them
would be by no means an unreasonable way of disposing of them.”
The Dromeognathous palate is sufficient justification for such an association.
Consent to this, however, renders inoperative the old term “ Ratitz,” which at best is
but a makeshift ; for many of the Carinate are Ratite, whilst the Tinamous have a
large keel to the sternum.
We propose instead the terms Palwognathey (Ratite + Crypturi) and Meognathe
(Carinatee—Crypturi). The adoption of these surmounts the difficulty indicated above.
Gadow regards the Struthiones as a group of primitive forms more nearly repre-
senting Proto-Carinate than any other living birds. Their retention under the common
name “ Ratitee” he regarded as convenient rather than an indication of close affinity.
Similarly we regard the Palwognathe as the unsuccessful descendants of proto-
carinate forms, but retaining probably but few of their characteristics unaltered. It
would be too much to regard any of the existing Meognathw as the direct descendants
of any of the existing or extinct Palwognathe known to us, but it seems not
* Blanchard in 1859 substituted for Merrem’s the terms Tropido- and Homalosterni.
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATHA. 267
improbable that their source may be traced to that stock which gave rise to the Rheo-
Dinornithine type of palate and pelvis.
Casuarius, which forms the nucleus of this research, is here regarded as only
generically distinct from Dromwus. These two forms are, it is here contended, closely
allied. Struthio is perhaps derived from the same ancestral stock. Hitherto, how-
ever, these two forms, Caswarius and Dromeus, have been regarded as more closely
allied to Dinornis and Apterya than any other forms. Apteryx we regard as quite
distinct. The interrelationships of this group are indicated in the diagram (p. 265).
The skeleton of Dromeus ater, procured at the last moment, at great trouble, by
Mr. Rothschild, has proved of considerable interest in one or two respects. It does
not differ much, however, from that of the larger species: wherein it differs it appears
to approach the Cassowaries. I have seen no authenticated skeleton of D. irroratus.
Skulls of adult Caswarius and Dromwus are badly needed; as also is the adult
skeleton of Rhea darwini. It is interesting to note that the characteristic feathered
tarso-metatarsus of &. darwini is temporarily reproduced in the embryo of R. americana
(fig. 2 D, p. 155).
Whether the four species of Ostrich indicated in this paper will be further confirmed
is a point which time alone will show. It seems probable.
I hope to be able to supply much that is Jacking in this paper by a further contri-
bution at no distant date in the shape of an Appendix. Since this was written the
British Museum has acquired a hind-limb of Genyornis. This is remarkable chiefly on
account of the fact that digit 1. was becoming absorbed. The proximal phalanx was
of great length and slenderness. In this we may trace the early stage in the evolution
of a second two-toed Struthious bird. As Messrs. Stirling and Zeitz have shown,
Genyornis was undoubtedly a near ally of Dromeus.
Key To THE OSTEOLOGY OF THE PALHOGNATHA,
based on the Characters of the Adult Skeleton.
A. SkuLt. (Plates XLIL—XLIV.)
The pterygoid free anteriorly or resting on the paired posterior ends of a large voiser ; connected
with the palatine either by the opposition of its outer border with the mesial border of the palatine
or by squamous suture, or by complete fusion, but never by means of opposed glenoid surfaces.
The vomer is always present, and paired, and is either fused with or tightly wedged in between
the maxillo-palatine processes ; basipterygoid processes always present, of great size, seated at
the base of the parasphenoidal rostrum, and articulating with the extreme proximal end of the
pterygoid; nasal processes of the premaxilla unpaired; parasphenoidal rostrum and mesethmoid
continued forward beyond the level of the lachrymal.
A. Beak short or of medium Jength, never probe-shaped.
a. Vomer deeply cleft posteriorly into widely expanded right and left limbs
running directly backwards beneath the pterygoids ; palatine articulating
268
MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
posteriorly by its mesial border with external border of the combined vomer
and pterygoid; voraer of great length, continued forwards beyond the
free end of the parasphenoidal rostrum; pterygoid laminate; orbital
process of quadrate decurved and articulating with the pterygoid at its
base ; anterior palatine foramen very small, and lying near the tip of the
beak beyond the vomer; anterior narial and lachrymo-nasal fossz, in
dried skull, confluent ; mesethmoid with a truncated anterior border ;
nasal process of premaxilla rod-shaped ; nasal with a vestigial external
process.
a’, Vomer expanded into a broad flattened plate between the maxillo-
palatines, and separated from the parasphenoidal rostrum; beak
dorso-ventrally depressed ; palatine fenestrated posteriorly, not anchy-
losed with pterygoid; horizontal plate of mesethmoid appearing on
the surface of the skull between nasals, but never inflated to form a
casque ; lachrymals fusing with nasals and frontals ; with long supra-
orbital processes
'. Vomer only very slightly enhonded between the arate palabiviesl Beal
laterally compressed ; palatine unfenestrated and anchylosed with the
pterygoid and vomer; horizontal plate of mesethmoid, nasals, and
more or less of frontals inflated to form a casque, sometimes of great
size ; quadrato-jugal fossa not extending forwards to level of antorbital
plate
6. Vomer large, ata teciieeal aeceies investing the par ee esoitel rostrum ;
distal end of pterygoid and hinder end of vomer hidden (in ventral aspect
of skull) by the palatine which underlies them; pterygoid rod-shaped.
c', Vomer deeply cleft anteriorly, exposing the parasphenoidal rostrum
between a pair of long spurs; anterior palatine foramen long and
slit-like.
a", Lachrymo-nasal fossa bounded anteriorly by a bony spike rising
upwards from the middle of the external border of maxillo-palatine ;
with large post-orbital processes ; orbital process of quadrate very
short, not extending beyond level of articulation with pterygoid ;
lachrymal with large orbital processes; mesethmoid visible as a
lozenge-shaped plate between nasals . 5
'. Lachrymo-nasal fossa bounded anteriorly by a deine rade articu-
lating above with the nasal and below with the hinder border of the
maxillo-palatine plate ; no post-orbital processes ; orbital process of
quadrate moderately long, with a free end projecting beyond articu-
lation with pterygoid; lachrymal without orbital processes ; nasal
schizorhinal; inter-orbital septum perforate ; mesethmoid not visible
on surface of skull
d', Vomer truncated anteriorly, not extending so oe ere as the aa of
the parasphenoidal rostrum; anterior palatine foramen small, closed
by anterior end of parasphenoidal rostrum ; large post-orbital processes ;
lachrymals fused with frontals and not readily traceable; orbital
Dromeus.
Casuarius.
Rhea.
Tinamide.
PHYLOGENY OF THE PALZOGNATHE AND NEOGNATHA. 269
process of quadrate long, articulating with whole inner border with
pterygoid ; olfactory cavity of great size . . . . . . . . . . Dinornithide.
c. Vomer deficient posteriorly, not extending backwards to reach the pterygoid,
anteriorly extending forwards, as in Dromeus, to conceal the parasphe-
noidal rostrum ; the body of vomer deeply hollowed, fitting closely on to
the rostrum, and embraced on either side by the maxillo-palatine pro-
cesses ; maxillo-palatine processes hastate; anterior palatine processes of
great size; pterygoid laminate; palatine rod-shaped, fused proximally
with external border of pterygoid ; premaxilla wanting a palatine process ;
lachrymals continued backwards to fuse with a chain of supra-orbitals
and with the frontals, so as to enclose a supra-orbital foramen ; lachrymal
with an osiculum lachrymo-palatinum ; nostrils holorhinal ; anterior narial
and Jlachrymo-nasal fossee confluent; quadrate with long recurved
process bearing a well-defined projecting facet for the pterygoid in
the middle of its internal ventral border . . . . . . . . . . . Struthio.
B. Beak elongated, decurved, probe-shaped ; anterior narial fossa minute, con-
fined to tip of beak ; lachrymo-nasal fossa enormous, extending forwards to
within a short distance of tip of beak ; skull rounded, without post-orbital
processes ; lachrymal vestigial ; olfactory chamber of great size, extending
backwards to optic foramen; vomer passing backwards to fuse with the
pterygoid and palatine, and, with these, helping to form the floor of a
tubularmecessropemingcimward sia mcnurny amie i) eens ane nnnEEA penta:
Key to the Species.
STRUTHIO.
a. Maxillo-palatine processes not rising dorsad, the anterior border of the
maxillo-palatine process at right angles to the long axis of the skull.
a’. Palatines projecting distad beyond anterior border of maxillo-palatine
process ; maxillo-palatine processes without any external lateral fenes-
trated plate é Porth, gti ener Ua BG as ety ae en S. australis,
6'. Palatines not reaching as far forward as the level of the anterior border of
the maxillo-palatine process.
a’, Vomer very short; parasphenoidal rostrum thick; maxillo-palatine
processes with an external lateral fenestrated plate. . . . . . . S.molybdophanes.
6’. Vomer very long ; maxillo-palatine processes without an external lateral
fenestrated plate; parasphenoidal rostrum slender ; roof of the skull
with deep depressions mesiad of the supra-orbital ledges; nasals
meeting in the middle line proximad of the nasal process of pre-
med 5 o 6 oo 0 6 © 9 6 6 6 6 6 BH 6 bh 8 0 (Sh aRHOAaOS”.
b. Maxillo-palatine processes rising dorsad ; anterior border of maxillo-palatine
processes sloping obliquely backwards ; vomer broad, with short, pointed,
posterior limbs; palatines projecting distad beyond maxillo-palatine pro-
CESSESP a sbucrie hint sad OO. wae veh SS ey th el then cea ke The tS ee Seecamelus:
1 We believe that Struthio massaicus (Neum.) is identical with this bird—W. R. & W. P. P.
VOL. XV.—Part v. No. 21.—WDecember, 1900. 2eP
270 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
APTERYX.
a. Vomer long, equal to or longer than the width across the paroccipital
processes at widest, notched anteriorly, aud with the median cleft not
exteading forwards beyond the middle - . . A, australis.
b. Vomer less than width across paroccipital processes.
a’, Vomer notched anteriorly, the median anterior portion nearly as long as
the paired posterior limbs; paroccipital processes well defined; basi-
temporal plate bounded antero-laterally by a pair of ridges passing
forwards into parasphenoidal rostrum . : 6 6 0 8 6 0-0 Zils AMIRI,
bo’. Vomer imperfect anteriorly, median portion much shorter than the paired
jee limbs.
= Size not less than.7 inches) 2) 953, 3 2 ee wl le el ee Aa mantelle:
. Size not less than 6} inches . . . . . . - . . . A. owen,
Raea.
A. Temporal fossa more or less sharply defined and extending backwards to
meet the lambdoidal ridge; lachrymal with elongated dorsal processes
arching backwards over orbit.
a. Posterior nares markedly wider anteriorly than posteriorly ; anterior vome-
rine cleft not extending backwards to the level of the hinder angle of the
lachrymo-nasal fossa; length of quadrato-jugal fossa less than width
across beak measured across the maxillary pillars bounding the lachrymo-
nasal fossa anteriorly . . . . . : 5 6 0 6 0 6 6 Li COAG
b. Posterior nares not wider anteriorly chan Heeiecionley anterior vomerine
cleft extending backwards to the level of the hinder angle of the lachrymo-
nasal fossa; length of quadrato-jugal fossa equal to width across beak at
the level of the maxillary processes bounding the lachrymo-nasal fossa. .macrorhyncha'.
B. Temporal fossa not sharply defined and not extending backwards to lambdoidal
ridge; lachrymals with short orbital processes; anterior vomerine cleft very
SINOTialins Meme Ge Seno W hol wot Moni dg lecd vaecoila Byetodia tego sie okt a. ye dali COORG
Dromzvs.
a. ae not exceeding 7 inches.
. Post-orbital processes short . . . . . .. . - . . . « .« D.nove-hollandie.
Q Post- orbital processes long, extending downwards to ihe a short distance
ES) Grecnechemosmeaal ime ss 5 5 6 5 6 oo 6 ce om 6 0 Jb Garena
b. Size not exceeding 5 inches; roof of skull with a very conspicuous cerebral
COM 6 6 6 so 6 8 o 6 0 6 59 0 0 2D, QuaP
1 The specific characters of this species have been drawn from a single adult skull, kindly lent me by
Mr. Beddard from the Zoological Society’s Collection.
2 This skull is from a skeleton in the Florence Museum. The bones of the palate are missing. The cranium
closely resembles that of D. nove-hollundi, differing therefrom chiefly on account of its very small size and
the prominence of the cerebral dome, whilst in the peculiar form of the inferior limbs of the lachrymal it
PHYLOGENY OF THE PALHZOGNATHA AND NEOGNATHA.
B. VerTEBR# (not including Crypturi) }.
2
‘
1
All the pre-synsacral vertebre are heterocelous and free; all the synsacral vertebre develop
high neural spines ; intercentra are wanting.
A. Hyperapophyses of axis well developed; cervical vertebre short; cervical
diapophysial lamelle large and conspicuous; at least one lumbo-sacral
vertebra ; sacro-caudal (post-sacral) vertebrze well ossified.
a. Hyperapophyses of atlas moderate; neural spine of axis rather high;
hyperapophyses of axis well developed ; cervical vertebra rather short ;
neural spines of cervical vertebrae well developed and greatly excavated
postaxially ; no neural spines laterally bifureating ; catapophyses com-
mence at about the sixth vertebra, and do not form a ring; hypapophyses
well developed, not much produced postaxiad; diapophysial lamella
extending towards postzygapophysis, large and conspicuous; cervical
ribs moderately long and styliform; thoracic and lumbar neural spines
not much axially extended, and not subequal in height; post-sacral
vertebre raised in part, more or less, above ventral margins of acetabula ;
sacral vertebre not plainly distinct from post-sacrals ; sacral and post-
sacral neural spines laterally expanded and separating the post-acetabular
ilium ; caudal vertebrie eight or nine; pygostyle cylindrical.
a’. Total number of vertebre fifty-four; cervical vertebra seventeen or
eighteen ; vertebre with ribs nine; post-sacral vertebre nine or ten ;
pre-axial articular surface of atlas moderately notched dorsally ; hypa-
pophysis of axis much elongated; hypapophyses developed from the
eighteenth or nineteenth to the twenty-third vertebra; a conspicuous
fossa on each side of the pre-axial part of the neural spines of dorsal
vertebrz ; post-sacral vertebra raised decidedly dorsad of ventral
Tinian OF acim 5s 56 vw 6 6 0 6 0 09 16 6 @ 5 4
b/. Total number of vertebre from fifty-five to fifty-nine ; cervical vertebra
fifteen or fourteen ; vertebre with (free) ribs eleven or twelve; sacro-
caudal (post-sacral) vertebre eight or nine; pre-axial articular surface
of atlas moderate, may become a foramen; hypapophysis of axis
extremely long; hypapophyses developed from eighteenth to twentieth
vertebra ; fossze beside pre-axial part of neural spines very small ; sacro-
caudal (post-sacral) vertebra raised scarcely at all dorsad of acetabula .
b. Hyperapophyses of atlas large ; neural spine of axis lofty ; hyperapophysis
of axis moderate or extremely large; cervical vertebrz short ; neural
spines of cervical vertebra generally more or less well developed ; cata-
Dromeus.
Casuarius.
resembles D. trroratus.
species.
In the conspicuous development of the cerebral dome it differs from both existing
I would remark here that the specific characters attributed to D. irroratus are based upon an examination
of a single unnamed skull in the British Museum Collection ; but these appeared to me to be sufficiently
marked to justify my determination of this skull as of D. wroratus.
1 This Key is compiled, with modifications, from Mivart’s memoir,
2P
9)
)
bo
bo
MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
pophyses may form a ring ; diapophysial lamella extending towards
postzygapophysis, large and conspicuous or extremely so; cervical ribs
moderate or small; sacro-caudal (post-sacral) slightly raised dorsad of
ventral margin of acetabulum ; cervical vertebre fifteen.
ce’. Total number of vertebre from forty-five to forty-seven ; post-synsacral
vertebre three ; pre-axial articular surface of atlas scarcely at all
notched dorsally ; hyperapophyses very long ; a minute hypapophysis
to atlas; no hypapophysis to axis; neural spine of axis very high ;
hyperapophysis of axis extremely large ; catapophyses commence at
the fifth vertebra, in the tenth and eleventh vertebre they form a
ring ; hypapophyses developed from the twelfth cervical to the third
thoracic vertebra; diapophysial lamella extremely large and con-
spicuous, developing conspicuous postaxially extending processes ;
cervical styliform ribs small; thoracic and lumbar neural spines
exceedingly extended axially and subequal in height; synsacrum with
sacral vertebrz plainly distinct from post-sacral ; post-sacral bent much
ventrad of ilia ; free caudal vertebr seven or nine; pygostyle conical.
’, Hyperapophyses of axis moderate ; neural spines of cervical vertebrae
well developed and deeply excavated postaxially ; neural spines of
fourth, fifth, sixth, and seventh cervical vertebre bifurcating laterally ;
catapophyses commencing at the fourth vertebra; hypapophyses
commence at fifteenth (last cervical) vertebra; diapophysial lamella
extending towards postzygapophysis, large and conspicuous; cervical
ribs probably moderately long and rather styliform; thoracic and
lumbar neural spines not much axially extended, and probably not sub-
equal in height ; synsacrum with post-sacrals not bent ventrad of ilia.
B. Axis vertebra with a hypapophysis, with or without well-developed hyper-
apophyses ; cervical vertebrz greatly elongated ; neural spines of cervical
vertebree small or almost obsolete; catapophyses commencing about the
sixth vertebra; catapophysis never forming a ring; diapophysial lamella
extending towards posterior zygapophysis, small and inconspicuous; a
conspicuous fossa on either side of pre-axial part of neural spines either of
cervical or thoracic vertebre; synsacrum with sacral vertebre plainly
distinct from post-sacral vertebre.
e’. Total number of vertebre fifty-six ; cervical vertebrae seventeen ; vertebre
with ribs ten; no lumbo-sacral vertebree ; synsacrum with eight post-
sacral vertebre ; pre-axial articular surface of atlas with a large dorsal
notch; hyperapophyses of atlas moderate ; hypapophysis of axis rather
small; neural spine of axis moderate ; neural spines of cervical vertebree
small, but not obsolete; no neural spines bifurcating laterally; no
neural spines aborted ; hypapophyses developed from the eighteenth
(first thoracico-dorsal) to the twenty-first (first thoracic) vertebre ;
hypapophyses but little produced pre-axially ; a conspicuous fossa on
each side of pre-axial part of neural spines of thoracic vertebr, but not
of cervical; cervical ribs long and styliform; thoracic and thoracico-
Apteryz.
Dinornithide.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA.
lumbar neural spines but little axially extended, but considerably
extended dorso-ventrally and not subequal in height ; synsacrum with
post-sacrals not at all raised dorsad of acetabula, but rather depressed
ventrad ; post-sacrals perfectly ossified ;
no lumbo-sacral vertebre; pre-caudal vertebre
sacral plainly distinct from
post-sacral vertebree ;
ten; pygostyle a laterally compressed plate . UM SON Ys Struthio.
f’. Total number of vertebre forty-nine; cervical vere fourteen ; pre-
axial articular dorsal surface of atlas with only a minute dorsal notch ;
no hyperapophysis to atlas; hypapophysis of axis not very large;
neural spine of axis not very high; hyperapophyses of axis well deve-
loped; neural spines of cervical vertebre almost obsolete, those of
fourth to seventh vertebre abort; neural spines of thirteenth to
sixteenth vertebre bifurcate laterally; hypapophyses developed from
last cervical to second dorsal vertebra; hypapophyses narrow and
pre-axially produced ; a conspicuous fossa on each side of pre-axial part
of neural spmes from twelfth to the fifteenth inclusive ; cervical ribs
short and not styliform; free cervical ribs nine ; synsacrum with post-
sacral vertebre raised quite dorsad of acetabula, also very imperfectly
ossified and much reduced; sacral plainly distinct from post-sacral
vertebre ; post-sacrals nine ; free caudal vertebre five ; pygostyle small
amGl q@ylbnchaenll g 5 6 6 o 540 0 Oo 6 Rhea.
The vertebral formula is :—
Synsac. eS
IDOE 65065000000 Cy. 18: Oy. Th. 3: Th.3+1+4+2: Lb.6: Lb.se.2: Se. 2: Cd. 949=55.
— eH
6 18
Synsac.
Casucriusteeieia Cy.15: Cv. Th. 4: Th.5+1+3: Lb.5: Lb.sce.2: Se. 2: Cd. 94+9=55.
—S SS —
9 18
Synsac.
poe
Struthto 28s secs selene Cv. 17: Cy. Th.3: Th. 542: Lb. 8: Se. 2: Cd.8+10 = 5).
a) nd
7 18
Synsac.
TOG go 009660000000 Cy. 14: Cy. Th.3: Th. 34241: Lb.3: Lb.se.4: Se.2: Cd. 94+8=49.
S— Sue
6 17
Synsac.
a i
ZVOTBED 650 0000606 . Cy.15:; Oy. Th.1: Th.44+341: Lb.4: Lb.se.4; Se. 2: Cd. 348=45.
(A. a. mantelli.) ame na fal =
274 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
C. Srernvum (fig. 4, p. 217) anp Pectorat Arca}.
The sternal plate is keelless; the coracoid anchyloses with the scapula, and lacks the acro-
coracoid, the coraco-scapular angle is very wide; furcula vestigial or absent.
A. Coracoid grooves not widely separated.
a. Metasternum pointed, no posterior lateral processes; pro-coracoid not
forming a distinct bar.
a’. Coracoid grooves and bases of coracoids overlapping ; anterior lateral
process long, directed upwards and backwards ; a pair of short recurved
processes lying on the anterior sternal border between the anterior
lateral processes ; with vestigial clavicles; sternal plate nearly as broad
as long; coracoid longer than broad . . . . . . . . . . )~Dromeus.
b’. Coracoid grooves not overlapping ; anterior lateral processes of sternum
short or obsolete; no clavicles; sternal plate much longer than broad;
@ormconel AS lownel as lkime 6 5 6 3 6 oo 6 6 0 6 oo fl lo 6(CRRITRINS.
6. Metasternum more or less pointed, with large posterior lateral processes ;
pro-coracoid of great size, forming a distinct bar of bone ; body of the
sternum with a conspicuous ventral prominence ; no clavicles ; articular
surfaces for ribs of great'width . . . . . . ..... =. . «. Struthio.
B. Coracoid grooves widely separated.
a. Corpus sterni basin-shaped with a marked ventral prominence ; posterior
lateral processes absent ; anterior lateral processes conspicuous, each with
a large pneumatic aperture at the base; articular surfaces for ribs
crowded together immediately behind the anterior lateral processes ;
coracoid grooves at the base of the anterior lateral processes and separated
one from another by the wide emarginate body of the anterior border of
the sternum ; coracoids long and slender ; pre-coracoid relatively small. Rhea.
b. Sternal plate flat, slightly or not at all pneumatic ; metasternal and posterior
lateral processes always present ; facets for ribs not separated by deep
pneumatic pits.
a’. Articular facets for ribs widely spaced, and lying entirely behind the
anterior lateral processes, and not separated by deep pneumatic pits ;
coracoid nearly as broad as long; pre-coracoid not conspicuous. . . <Apteryx.
’. Facets for ribs crowded, few in number, and partly underlying the base
of the anterior lateral processes ; coracoid extremely reduced . . . Dinornithide.
C. Sternal plate flat, pneumatic; metasternal element wanting ; facets for ribs
separated by deep pits; sternal plate exceedingly narrow from before
IQS. og 6@ Gao @ po O° 6 ob © 4 6 6 © 0 6 of 6. 6. 0 Lee,
* The sternum and pectoral arch of the Crypturi will be found described on pp. 221 & 263.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH., 275
Key to the Species.
StRUTHIO,
a. Middle region of anterior border straight or slightly arched; length of
coracoid groove greater than distance between articulations of sternal
ribs 1-5
a’. Posterior lateral processes long and spatulate, with an extensively ossified
metasternal region between the posterior lateral processes
6’. Posterior lateral processes moderately long; ossified metasternal region
between posterior lateral processes small . ; 6 oo 6
b. Middle region of anterior border of sternum sinuous ; posterior Tnteral pro-
cesses not spatulate; no projecting metasternal region between the
posterior lateral processes; length of coracoid groove less than the
distance between the articulations of sternal ribs 1-5.
c’. Width between the free end of the anterior lateral processes much greater
than the width across their bases . . . Fe te Aes nee
. Width across free end of posterior lateral processes about equal to the
width across their bases . ... . =. . «©
Ruea.
a. Anterior border of sternum between the coracoid grooves slightly excavated ;
width between right and left coracoid grooves greater than length of
articular surface for coracoid ; articular surfaces for sternal ribs 3
6. Outer border of sternum between the coracoid grooves deeply excavated ;
width between right and left coracoid grooves equal to or jess than length of
articular surface for coracoid; articular surfaces for sternal ribs 4
Dromzvus.
a. Length 64 in. Coracoid grooves distinct and the sternal protuberance but
by slightlyadevelopedianea rommeme ints) -rao- (oll 00 l- ) well co oe oe
b. Length not exceeding 3% in.; coracoid grooves obsolete ; the ventral lip of
. S. molybdophanes.
S. camelus.
S. meridionalis.
S. australis.
R. americana.
. R. macrorhyncha.
D. nove-hollandie.
right side only represented ; sternal protuberance conspicuous. . . . D. ater.
CasuaRIus.
A. With a pair of low, median, keel-like ridges lying immediately behind a line
drawn across the sternum immediately behind the coracoid grooves, without
median pneumatic cavity between the coracoid grooves . . . . . « C. unappendiculatus,
B. Without paired median ridges; with large pneumatic cavity between the
coracoid grooves.
a. Coracoid grooves deep, the lower lip strongly marked when seen from
above ; pneumatic cavity very large, extending backwards into a low
swollen protuberance lying in the middle line of the sternum between
the coracoid grooves.
276 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
a’. Width between inner angles of articulation for last sternal rib less than
width across metasternum at widest. . . . - + « . . . C, casuarius.
b/. Width between inner angles of articulation for last ternal rn greater
than width across metasternum at widest . . . . . . . . « « Cie. beccarii.
b. Coracoid grooves shallow, lower lip not strongly marked when seen from
above ; mouth of the pneumatic aperture produced forward into prominent
lips; anterior lateral process short and recurved. Metasternal region
dae long and narrow.
. Width across anterior lateral processes of sternum twice feu of
metasternum . . é Sh aette 5 6 Ae nse Clicennette:
. Width across anterior Tea process at’ sternum ese an twice width
AGO MNO 5 6 6 56 9 5 6 46 9 6 6 6 © 6 0.0 0 Oh ymin.
D. Pstvic Arcu!. (PI. XLIV. fig. 5.)
Synsacral neural spines of great height ; sacral vertebra, and sometimes the last lumbo-sacral,
and first and second caudals with conspicuous pleurosteal elements ; pre-acetabular ilia always
meeting in the mid-dorsal line above the neural spines of the synsacrum ; the synsacral neural
epines never projecting beyond the level of the post-acetabular ilia; ilio-ischiadic fissure either
permanently open or closed only by the fissure of the extreme ends of the ilium and ischium.
A. Pubes remaining permanently free caudad.
a. With the post-acetabular ilia more or less widely separated in the mid-
dorsal line; a more or less well-developed supra-trochanteric process.
a’. Post-acetabular ilia separated by the intervention of the synsacral neural
spines; ischia bowed inwards towards the middle line, and fused
posteriorly with the ilium; perforation of inner wall of acetabulum
constricted by ossification.
a’, Post-acetabular longer than pre-acetabular ilium, and tapering
gently . Nie rik rem eneittE Dereon carat se Gee Sater 15, 0a
. Post-acetabular shorter than arecaoetabular ilium, and arched
dorsally
Dromeus.
6 6 6 SU Aeneid eee MUSE uot MCOSUATIUSs
b’. Post-acetabular ilia aiaely separated by means of transverse processes,
leaving neural spines in middle of a deep fossa roofed in late in life by
a sheet of bone; ischia widely separated one from another and from
post-ilia ; post-acetabular with a very wide dorsal plane.
ce’, Median fossa dividing post-acetabular ilia only imperfectly roofed
in, leaving a more or less extensive cleft on either side of a broad,
flat neural plate Pout ° . Dinornithide.
d’’, Median fossa dividing post- acetabular ilia completely rooted oy hone
but perforated by a double row of foramina, one on either side of
the median line. Sih teu diee use Lie stub sOlreNE AUD YOTnNLS
» The pelvic arch of the Crypturi will be found described on p. 263. There was no need to include it
here.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA,
te
oa |
=]
b. Post-acetabular ilia closely approximated, the neural spines at most only
barely visible between the closely approximated innominates; no supra-
trochanteric process ; ischia widely separated from post-acetabular ilium;
obturator foramen and fissure confluent ; post-sacral vertebrae of the
synsacrum welded into a large laterally compressed mass, bent down so
as to bring the neural crest to the level of the ventral border of the post-
WHEN, 5 oo es oa dt toe Soar 0 5 6 > 0 6 6 9 AYiaIER
B. Pubes anchylosed candied mth one another or Beh ‘g@linn,
c. Post-acetabular ilia widely separated from ischia, and from one another
in the mid-dorsal line; pubes fused caudad (a) with the ischia and (d)
with one another in the mid-ventral line; the synsacral vertebre appear-
ing below the level of the ventral border of the ilium . . . oer ey SENULIIOS
d. Post-acetabular ilia anchylosed with ischia, and with one ETotiier in the
mid-dorsal line; pubes anchylosed with ischia caudad; ischia closely
united in the middle line one with another throughout almost entire
length (unique) ; the post-sacral vertebrae of the synsacrum atrophied . Rhea.
Key to the Species.
Ruea.
a, Pre-ilium not projecting forward beyond the last thoracic vertebra and rib,
which forms the limit of the synsacrum cephalad . . . . . . . . . WR. americana.
6, Pre-ilium projecting forward conspicuously beyond the last thoracic vertebra
pln) ¢ 5 6.6) Gg oS ob 6 0 6p a 59 6 6 5 6 6 o 6 Oo 5 JenmomeAahD.
APTERYX.
A. Width across anti-trochanter equal to or less than the width of pre-acetabular
ilium at widest.
a. Pre-acetabular ilium of great width, with a strongly arched dorsal border,
and large laterally expanded antero-ventral angles; width across pre-
acetabular ilia at widest part=to length of ischium, and far exceeding
length of post-acetabular ilium; post-acetabular illum with a sudden
widening at its free end; breadth of pre-ilium at widest=breadth across
AMT 5 o 0 0 0.0 0 6 0 6 6 0 0 9 6D . - A. australis.
B. Width across anti-trochanter always greater than width of pre- acetal
ilium at widest; post-acetabular tapering caudad; breadth of pre-ilium at
widest less than breadth of anti-trochanter.
6. Pectineal process large ; five lumbar vertebree.
a’. Post-acetabular ilium with a concave ventral border, and a slight notch
immediately behind the anti-trochanter . . . . . . . . . . A. haasii.
o'. Post-acetabular ilium with straight or convex ventral border ; no notch
behind the anti-trochanter . . . SD ee ey cig hind Lao hier EAR OWWCNE®
c. Pectoral process small ; four lumbar ver ayes Je Gh soy ideo oe echo oo) neh Chai
VoL. XV.—PAR?T v. No. 22.—December, 1900. 2a
278 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Drom us.
a. Length 17 in.; pectineal process well developed ; dorsal border of pre-ilium
much arched ; obturator fissure nearly closed ; pubo-ischiadic fissure nearly
Gogetil 5 o 6 6 5 0 6 ob oo eo ol ll 68 ll ol oO Di pipeiolianite,
6. Length 93 in.; pectineal process obsolete; dorsal border of pre-ilium not
arched ; obturator fissure wide ; pubo-ischiadic fissure very wide. . . . Dz ater.
EK. Prcrorat Lime.
A. Extended forearm and hand longer than the humerus; ulnare resembling
that of the typical Neognathe ; radius and ulna long, with but a narrow
chink between them; distal end of Me. II. flabelliform . . . . . . . Rhea.
B. Extended forearm and hand much shorter than the humerus.
a. Humerus with a shallow, but blind subtrochanteric fossa; a very strongly
developed linea aspera for the triceps, forming a conspicuous triangular
ridge; radius and ulna short and bowed; ulnare a simple nodule; distal
end of Me. II. not markedly expanded; manus as in Rhea, resembling
typical carimate form, though too small for purposes of flight . . . . Struthio.
6. Humerus much bowed, without subtrochanteric fossa or indication of
pectoral crest ; radius and ulna with but a very narrow chink between
them; a vestigial radiale and ulnare; a vestigial pollex bearing an
unequal phalanx, with the Me. III. incomplete and free distally . . . Apteryax.
C. Extended forearm and hand about equal to the length of the humerus; a
subtrochanteric fossa ; radiale and ulnare anchylosed with metacarpals.
a, Subtrochanteric fossa large ; ulnare indistinguishable ; radiale anchylosed
with radius and carpo-metacarpus, but distinguishable. . . . . . . Cuaswarius.
6. Subtrochanteric fossa very small; no carpal elements distinguishable ;
neither pollex nor third digit represented. . . . . . =. =. ..: . Dromeus.
F, Pretvic Lie.
When the bones are pneumatic only the femur is conspicuously so; save in the Dinornithide
the extensor bridge is wanting; the hallux, when present, is never large.
a. Toes three in number ; tarso-metatarsus far exceeding femur in length.
a’. Ectocnemial crest large, rivalling the procnemial in size ; ento-condyle with
a deep lateral pit; distal extremity of shaft above entocondyle (lateral
surface) with a deep groove.
a, Procnemial crest extending down the shaft; femur with a large
(SATTEAG CNOA GG 0 6 6 6 Go oo 5 ol ee ol LDR UD,
b!. Procnemial crest not extending down the shaft ; femur non-pneumatic;
ungual phalanx of inner toe greatly elongated . . . . . . . . Casuarius.
4!. Ketocnemial crest small, a mere tubercle of the procnemial; no ento-
condylar pit or groove on the lateral surface of the shaft ; a moderately
conspicuous extensor groove; popliteal fossa of the femur very deep;
ungual phalanges laterally compressed. «9. . . » = » 2 1-2.) Ried,
PHYLOGENY OF THE PALAOGNATHEX AND NEOGNATEHA. 279
6. Toes two in number ; tarso-metatarsus far exceeding the femur in length.
ce’. Ectoenemial crest small, a mere tubercle on procnemial ; fibula extending
to level of proximal tarsal mass; extensor groove obsolete; a deep ento-
condylar pit continued backwards into a groove; the lateral ventral
border of ento- and ecto-condyles continued backward and upward to
project spur-wise from the shaft .
ce. Toes four; tarso-metatarsus short.
d', Extensor bridge wanting.
ce", Femur long and slender; tibio-tarsus with a deep intercondylar gorge ;
entocondyle very large, projecting conspicuously in front of shaft in
lateral view ; none of the phalanges conspicuously shortened
d", Femur very short and thick, with enormous great trochanter; tibio-
tarsus without intercondylar gorge, ento- and ecto-condyles not being
clearly distinguishable: Ph. 2. D. I. short; Ph. 2, 3, 4. D. IV. short.
e’, Extensor bridge present ; femur short ; ento-condyle of tibio-tarsus large,
resembling that of Apterya
List oF WORKS REFERRED TO OR CONSULTED.
1, Aux, P. H. E.—Syrinx of Rhea. Bull. Soc. Philomat. 1874, p. 38.
Struthio,
Apteryx.
Aipyornis.
Dinornithide.
2. Autis, T.—On the Skeleton of the Apteryx. Journ. Linn. Soc., Zool. xi. 1873, p. 523.
3. Anprews, C. W.—Note on a complete Skeleton of Apyornis from Madagascar. Geol. Mag.
vol. iv. 1897, p. 241.
4. Anprews, C. W.—On the Skull, Sternum, and Shoulder-girdle of Apyornis. Ibis, 1896
p. 376.
5. Bepparp, F. E.—On the Heart of Apteryx. Proc. Zool. Soc. 1885, p. 188.
6. Bepparp, F. E.—On the Alimentary Canal of the Martineta Tinamou.
pp- 61-66.
Bepparp, F, E.—The Structure and Classification of Birds. 1898.
comY
. .
vol. vi. 1899, pp. 386-402.
eo]
Ibis, 1890, vol. iit.
Bepparp, F. E.—Notes on the Anatomy of the Genus Apteryx. Novitates Zoologice,
g. Beppaxp, F. E. and Mrrcnetx, P. C.—On the Structure of the Heart of the Alligator. Proc.
Zool. Soc. 1895, pp. 343-349.
10, Buumensacn, J. F.—Abbildung. naturhistor. vol. vii,
11. Borcxinc, A.—De Rhea americana. Diss. Inaug. Bonn, 1863.
12. Burcxuarpt, R.—Ueber Apyornis. Paleeontol. Abhandl. neue Folge, Bd. ii. Heft 2 (Jena,
1893).
13. Cunnincuam, R. O.—Notes on some Points in the Osteology of Rhea americana and Rhea
darwini. Proc. Zool. Soc. 1871, p. 105.
ZQy2,
280
14.
15s
16.
C7
18,
1g.
20.
MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Cuvier, Baron G.—Lecons d’Anat. Comp. t. ii. 1799-1800 (Ann. vin.), p. 69.
Datuas, W. S.—On the Feathers of Dinornis robustus. Proc. Zool. Soc. 1865, p. 265.
Ducuamp, G.—Observations sur ’ Anatomie du Dromeus. Ann. Sci. Nat. xvi. 1873.
Epwarps, A. Mitnr- and Oustater, E.—Notice sur quelques Espéces d’Oiseaux du Muséum
WHistoire Naturelle. Centen. Mus. d’Hist. Nat. Paris, 1893 (l’Emeu ou Hmou noir,
p-. 246). See also Mitwn-Epwarps.
Evans, F. P.—Note on the Carotids of Rhea americana. Ann. & Mag. Nat. Hist. xi. 1884,
p. 458.
Frower, Sir W. H.—On the Skeleton of the Australian Cassowary, Casuarius australis. Proc.
Zool. Soc. 1871, p. 32.
Forses, W. A.—On the Conformation of the Thoracic End of the Trachea in the Ratite
Birds. Proc. Zool. Soc. 1881, p. 778.
20A. Forpes, W. A.—On the Bursa Fabricii in Birds. Proc. Zool. Soc. 1877, pp. 804-318.
21.
22.
2s
24.
25.
26.
Dy).
28.
29.
30.
3I.
32.
33:
34:
35:
36.
37:
38.
39:
40.
Fremery, P. J.—Specimen Zoologicum sistens observationes, presertim Osteologicas de
Casuario nove-hollandie.
Fixsrincer, M.—Untersuch. zur Morphol. und System der Végel. Amsterdam, 1888.
Gapow, H.—Zur vergleich. Anat. der Muskulatur des Beckens und hinteren Gledmasse der
Ratiten. Jena, 1880.
Gapvow, H.—Remarks on the Cloaca and on the Copulatory Organs of the Amniota. Phil.
Traus. Roy. Soc. 1887, pp. 5-37.
Gapow, H.—On the Anatomical Differences in the Three Species of Rhea. Proc. Zool. Soc.
1885, p. 308.
Gapow, H.—Bronn’s Thier-Reich. Bd. vi. Vogel, 1891, Anatom. Theil.
Gavow, H.—Ibid. Syst. Theil, 1893.
Garrop, A. H., and Darwin, F.—Note on an Ostrich lately living in the Society’s Collection.
Proce. Zool. Soc. 1872, pp. 356-362.
Garrop, A. H.—On the Carotid Arteries of Birds. Proc. Zool. Soc. 1873, p. 457.
Garrop, A. H.—On certain Muscles of the Thigh of Birds and on their Value in Classifica-
tion: Part I. Proc. Zool. Soc. 1873, pp. 626-644.
Garrop, A. H.—Ditto: Part II. Op. cit. 1874, pp. 111=128.
Gnorrroy St. Hitarrn, E.—Compos. des Appareils génitaux, urinaires et intestinaux, d leurs
points de rencontre dans l’Autruche et dans le Casoar. Mém. d. Hist. Nat. tome ix.
1822, p. 438.
Gruser, W.—Ueber das Thranenbein der Straussartigen Végel. Bull. Ac. Sci. St. Pétersb.
1855, p. 161.
Haventon, S.—On the Muscular Mechanism of the Leg of the Ostrich. Proc. Roy. Irish
Acad. vol. ix. 1866. !
Havenron, 8S.—On the Muscular Anatomy of the Emu, Dromeus nove-hollandie. Ibid.
1868, pp. 487-497.
Haventon, S.—On Muscular Anatomy of the Rhea. Ibid. p. 497.
Hitprsranor, G. F.—Bemerkung iiber den Embryo des Strausses im Hie. Voigt’s Magaz. d.
Naturkunde, Bd. xi. 1806, pp. 97-126.
Hurron, IF. W.—The Moas of New Zealand. Trans. N. Z. Instit. vol. xxiv.
Houxtey, 'T’. H.—On the Classification of Birds. Proc. Zool. Soc. 1867.
Huxtey, I’. H—The Anatomy of Vertebrated Animals. 187 iy
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATH®. 281
- Huxiuy, T. H.—On the Respiratory Organs of Apteryx. Proc. Zool. Soc. 1882, p. 560.
- Kyox, R.—Observations on the Anatomical Structure of the Cassowary of New Holland.
Edinb. Philos. Journ. vol. x. 1824, p. 132.
- Kwox, R.—Additional Observations on the Structure of the Trachea in the Cassowary Emeu
of New Holland. Edinb. Philos. Journ. vol. x. 1823-24, p. 137.
. Layxester, E. R.—On the Right Cardiac Valve of Echidna and of Ornithorhynchus. Proc.
Zool. Soc. 1883, pp. 8-14, pls. iii., iv.
. Lanxesrer, E. R.—On the Heart described by Prof. Owen in 1841 as that of Apteryx. Proc.
Zool. Soe. 1885, pp. 289, 240.
» Lanxuster, E. R.—On the Right Cardiac Valve of the Specimens of Apteryx dissected by Sir
Richard Owen in 1841. Proc. Zool. Soc. 1885, pp. 477-482.
. Livpsay, B.—On tiie Avian Sternum. Proc. Zool. Soc. 1885, pp. 684-716, pls. xlii—xly.
. LypsekKker, R.—Cat. Foss. Birds, 1891.
- Macazister, A.—On the Anatomy of the Ostrich. Proc. Roy. Irish Acad. 1867.
. Macartney, J—An Account of the Organ of Voice in the New Holland Ostrich. Brit.
Assoc. Rep. Bristol, 1836.
. Mau, A. W.—Om Luftrér-sacken. (fv. K. Vet.-Ak. Férh. 1880, no. 7, p. 33.
. Marsnatt, W.—Beobachtungen iiber das Verhiltniss der Federn. Zool. Gart. vol. xvi.
1875, p. 121.
. Maver, A. F. J. C.—Analecten fiir vergleich. Anatom. 1839.
. Mecxet, J. F.—System der vergleich. Anatom., Abtheil. 11. 1825, p. 202.
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1830, p. 200; 1832, p. 273.
. Merere, J. H. ne.—Ueber die Federn der Végel. Morphol. Jahrb. xxiii. Bd. 4, 1895,
pp. 562-591.
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Rendus Acad. Sci. t. exviti. 1894, p. 122.
. Mitnz-Epwarps and Granpipier.—Sur les Ossements d’Oiseaux provenant des Terrains
récents de Madagascar. Bull. Mus. Hist. Nat. pt. 1895, p. 9. See also Epwarps.
. Mircuxztz, P. C.—On the Intestinal Tract of Birds. Proc. Zool. Soc. 1896, pp. 186-159.
. Mircnett, P. C.—On so-called Quintocubitalism. Journ. Linn. Soc., Zool. vol. xxvii.
. Mrvart, Sr. G.—On the Axial Skeleton of the Struthionide. Trans. Zool. Soe. x. p. 1.
. Morte, J.—On the Tracheal Pouch of the Emu. Proc. Zool. Soc. 1867, p. 405.
. Nasonov, N. V.—K istorii razvitiya Afrikanskagho Strausa. [On the history of the deve-
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. Newton, A.—Dictionary of Birds (Introduction), 1896, p. 109.
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Roy. Soc. 1891, vol. 182.
MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
. Parker, T. J.—Additional Observations on the Development of Apteryx. Phil. Trans. Roy.
Soc. 1892, vol. 183.
. Parker, T. J.—On the Cranial Osteology, Classification, and Phylogeny of the Dinornithide.
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. Parser, T. J.—Text-Book of Zoology, vol. ii. 1897.
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vol. 11. p. 740.
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Linn. Soe. xxvi. (1870), p. 609.
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Leyden, 1888.
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 283
98. Woopwaxp, A. S.— Vertebrate Paleontology, 1898.
99. Wray, R. S.—On some Points in the Morphology of the Wings of Birds. Proc. Zool. Soc.
1887, pp. 343-356.
100, Wunperticu, C.—Beitriige zur vergleichenden Anatomie und Entwickelungsgeschichte des
unteren Kehlkopfes der Vogel. Nov. Act. Ac. Leop.-Carol. xlviii. 1884.
1o1. Zanprer.—Ueber das Gefieder des amerikanischen Strausses. Schrift. phys.-dkonom. Gesell.
K@nigsb. xxix. 1889, p. 31.
EXPLANATION OF THE PLATES.
PLATE XXII.
Casuarius casuarius. Common Cassowary (p. 113). (Drawn from life from a bird
formerly at Tring, now living in Dublin.)
PLATE XXIII.
Casuarius caswartus beccarti. Beccari’s Cassowary (p. 116). (Drawn from life from
a bird formerly at Tring, now in the Society’s Gardens.)
PLATE XXIV.
Casuarius casuarius salvadorii. Salvadori’s Cassowary (p. 120). (Drawn from life
from a bird formerly alive at ‘Tring, now mounted in the Tring Museum.)
PLATE XXV.
Casuarius caswarius australis. Australian Cassowary (p. 123). (Drawn from life
from a bird that lived ten years at Tring, and is now living in the Society’s Gardens.)
PLATE XXVI.
Casuarius casuarius violicollis. Mauve-necked Cassowary (p. 122). (Drawn from
life from the type specimen formerly at Tring, now living in the Society's Gardens.)
PLATE XXVIL.
Casuarius casuarius intensus. Blue-necked Cassowary (p. 121). (Drawn from lite
from the type specimen formerly at Tring, now mounted in the Tring Museum.)
PLATE XXVIII.
Casuarius bicarunculatus. 'Two-wattled Cassowary (p. 129). (Drawn from life from
a bird formerly at Tring, now living in the Society’s Gardens.)
284 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
PLATE XXIX.
Casuarius unappendiculatus. One-wattled Cassowary (p. 132). (Drawn from life
from a bird in the possession of Mr. Blaauw.)
PLATE XXX.
Casuarius unappendiculatus. One-wattled Cassowary (p. 132). Whole birds, adult
and young. (The figure of the adult was made from Mr. Blaauw’s bird; the young
was drawn from a bird at Tring, the skin of which is preserved in the Tring Museum.)
PLATE XXXI.
Casuarius unappendiculatus occipitalis. Jobi Island Cassowary (p. 135). (The
head, neck, and naked parts were drawn from a bird now living in the Society’s
Gardens, the casque from the mounted specimen in the Paris Museum.)
PLATE XXXII.
Casuarius unappendiculatus aurantiacus. Kastern One-wattled Cassowary (p. 136).
(Drawn from life from the type now living in the Berlin Zoological Gardens.)
PLATE XXXII.
Casuarius philipi. Sclater’s Cassowary (p. 138). (Drawn from life from the type
now living in the Society’s Gardens. )
PLATE XXXIV.
Casuarius papuanus. Westermann’s Cassowary (p. 139). (Drawn from life from
two specimens formerly at Tring and the Society's Gardens, now preserved in the
Tring Museum.)
PLATE XXXV.
Casuarius papuanus edwardsi. Milne-Kdwards’s Cassowary (p. 141). (Drawn from
life from a specimen formerly at Tring, now mounted in the Tring Museum.)
PLATE XXXVI.
Casuarius picticollis, Painted-necked Cassowary (p. 143). (Drawn from the plate
in Gould’s ‘ Birds of New Guinea.’)
PLATE XXXVILI.
Casuarius picticollis hecki. Heck’s Cassowary (p. 144). (Drawn from life from
the type specimen living in the Berlin Zoological Gardens.)
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 285
PLATE XXXVIII.
Casuarius lorie. Loria’s Cassowary (p. 142). (Drawn from a sketch made from
a fresh-killed bird by Dr. Loria.)
PLATE XXXIX.
Casuarius bennetti. Bennett's Cassowary (p. 145). (Drawn from life from a bird
formerly at Tring, now in the Zoological Society’s Gardens.)
PLATE XL.
Map showing the geographical distribution of the six subspecies of Casuwarius
casuarius and of Casuarius bicarunculatus.
PLATE XLI.
Map showing the geographical distribution of the three subspecies of Caswarius
unappendiculatus, two of Casuarius papuanus, two of Casuarius picticollis, Casuarius
lorie, and Casuarius bennetti.
PLATE XLII.
The dorsal aspect of the Skull (figs. 1-4).
Fig. 1. The skull of Rhea americana (p. 270), to show the form of the temporal fossa,
the partly fused mesethmoid plate, the form of the lachrymals and _nasals,
and the broad nasal process of the premaxilla.
. The skull of Dromeus nove-hollandie (p. 270), showing the rod-like nasal
process of the premaxilla, the fused lachrymal, broad interorbital region, and
Les)
ore
| w)
widely separated temporal fossze.
. The skull of Struthio molybdophanes (p. 269), showing the supra-orbital ossicles
running backwards from the lachrymal to the frontal, and the broad nasal
process of the premaxilla. The mesethmoidal suture is obliterated.
Fig. 4. The skull of Apterya australis (p. 269), showing the small size of the lachrymal
and the absence of its supra-orbital processes, the complete fusion of the
=
or
gs
[Se
nasal process of the premaxilla, and the rounded form of the frontals, which
do not protect the orbit above.
Ventral aspects (figs. 5-8)
Fig. 5. The ventral aspect of fig. 1, showing the large size of the palatine
processes of the premaxilla and of the maxillo-palatine processes. The
peculiar relations between the vomer, palatine, and pterygoid (described on
p- 207), and the anterior end of the parasphenoidal rostrum between the
anterior cleft of the vomer.
VoL. XV.—PaRT v. No. 23.—December, 1900. QR
286 MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
Fig. 6. Ventral aspect of fig. 2, showing the broad vomer and its continuance
backwards on to the pterygoid. The palatines articulate with the external
border of the foot of the vomer and external border of the pterygoid. In
Rhea the articulation between pterygoid and vomer is concealed by the
palatine.
articulate with the pterygoid.
7. Ventral aspect of fig. 3, showing the vestigial vomer, which does not now
Note the absence of the palatine processes of
the premaxille. The palatine has fused with the external pterygoid border.
Fig. 8. Ventral aspect of fig. 4, showing the complete fusion of the vomer, pterygoid,
and palatine.
ADDITIONAL LETTERS.
ant.b,f. =anterior basi-cranial fontanelle.
ant.h.=antrum.
b.6.p.= basitemporal platform.
b.p.=basipterygoid processes.
hy for.= hypoglossal foramina, 1, 2, 3.
!.=lachrymal.
Lp.=lachrymo-nasal pillar.
ir. =lambdoidal ridge.
m.e.=mesethmoid.
mz.p.=maxillo-palatine process.
n.pmx.—=nasal process of premaxilla.
n.=nasal.
0.c.=occipita] condyle.
pa.=palatine.
pa.r.=parasphenoidal rostrum.
p.pmx.=palatine process of premaxilla.
p-p.=paroccipital process.
pt.=pterygoid.
s.=supra-orbital bones from lachrymal to
frontal.
v.=vomer.
vag for.=vagus foramen.
PLATE XLITI.
The immature Skull.
Fig. 1. Dorsal aspect of the palatine bones of Apteryx mantelli (p. 204), to show the
sutures, as yet unclosed, between the vomer, pterygoid, and palatine.
Fig. 1a. Ventral aspect of the same, to show similar sutures.
Fig. 2. Sagittal section of skull of Apteryx australis mantelli, after Parker, for the
purpose of comparison with Dromeus. Note the appearance of the squamosal
on the inside of the skull in Apteryz.
i)
ey
oe)
. 8. Ventral aspect of the skull of Emeus (p. 270), for the purpose of comparison
with Rhea. Note the similarity in the relations of the pterygoid and vomer.
Fig. 4. Sagittal section of cranium of embryo Dromewus nove-hollandie, to show the
separate bones.
Fig. 4a. Outer view of fig. 4. Note how completely the squamosal conceals and
protects the pro- and opisthotic bones. In the nestling Penguin the
squamosal is deeply emarginate posteriorly and exposes the otic bones.
Fig, 5. Ventral aspect of base of skull of Ahea americana, to show sutures,
PHYLOGENY OF THE PALZOGNATHA AND NEOGNATHA. 287
g. 5a. Lateral aspect of skull of Rhea americana, to show sutures, Note the
vertical spine (lachrymo-nasal pillar) from the maxillo-palatine process,
which replaces the maxillary process of the nasal; the remarkably small size
of the quadrato-jugal; and the outer and inner views of the lower jaw.
. 56. Lateral aspect of pterygoid of Rhea americana (p. 191), to show its relations
with the vomer.
. 5c. Outer aspect of mandible of fig. 5, to show sutures.
. 5d. Inner aspect of mandible of fig. 5, to show sutures.
g. 6. Quadrate of Bubo maximus, to show double, widely separate, squamosal articular
head. Nat. size.
Fig. 7. Quadrate of Dromeus irroratus?, posterior aspect, to show single articular
head. Nat. size.
Fig. 8. Quadrate of Apteryx owent, posterior aspect, to show double squamosal
head.
ADDITIONAL LETTERS.
a.=angulare. o.h.=otic head of quadrate.
als.=alisphenoid. o.r.=orbital process of quadrate.
ar, =articulare. p.=parietal,
ant.=antrum of Highmore. pf.=precondylar fossa.
b.s. = basisphenoid. pro. =pro-otie.
d,=dentary. $.a.=supra-angular.
cor.=coronoid. $.0.=supra-occipital.
ep.o.=epiotic. sp.=splenial.
ex. =exoccipital. sq.=squamosal.
ff.=floccular fossa. sq.h.=squamosal head of quadrate.
fr.=frontal. vag.f.=vagus foramen.
m.=meatus internus. vo.pr.=vomerine process of premaxilla,
mes. =mesethmoid. l.p.=lachrymo-nasal process of premaxilla.
op. = opisthotic.
PLATE XLIV.
Fig. 1. Dorsal aspect of the skull of an adult Dromeus ater (p. 270), to show the
conspicuous, embryonic, cerebral prominence and the fused lachrymals.
. la. Lateral view of the same skull (fig. 1), showing the cerebral prominence
and temporal fossze.
Fig. 2. Dissection of a skull of a young Dromeus nove-hollandia, lateral view. to
show the form and position of the mesethmoid for comparison with fig. 3
et seq.
Fig. 3, The mesethmoid of a young Casuarius unappendiculatus (p. 199), to show the
mesethmoid and the large median superposed element forming the centre of
the casque of the adult.
‘ig, 4a,
MR. W. P. PYCRAFT ON THE MORPHOLOGY AND
ig. 4. Lateral aspect of a portion of the skull of Casuarius sclatert, showing the
frontal, nasal, and median elements of the casque.
Dorsal aspect of fig. 3 a.
‘ig. 4b. Dissection of a skull of Casuarius sclateri=fig. 3 a, showing the mesethmoid
and a portion of the ventral border of the median ossification of the
casque interpolated between the mesial border of the anterior ends of the
frontal.
Fig. 5. Lateral view of the synsacrum of a young Casuarius wnappendiculatus after
removal of the right innominate, to show the great height of the neural
spines of the vertebree.
Fig. 6. Lateral view of the synsacrum of an embryo Dromeus nove-hollandie (p. 212),
prepared for comparison with fig. 5. Note that at this stage the neural
spines have not yet risen above the centra of the vertebre, but form only a
series of lateral wings. ‘
Fig. 6a. Dorsal aspect of a portion of the synsacrum (fig. 6), to show more clearly
that the neural spines do not rise above the centra as in the adult
condition (fig. 5).
Fig. 6 6. Lateral view of the same pelvis (fig. 6), to show the separate elements
of the innominate.
ADDITIONAL LETTERS.
ant. = anti-trochanter. o.p.fr.= orbital process of frontal.
jf.c.=frontal moiety of casque. | p-=pubis.
i. =ilium. p.p.=pectineal process.
il.is.f.=ilio-ischiadic foramen. pubo-isch.f.= pubo-ischiadic fissure.
is. =ischium. s.c.=sacral vertebra 1-2.
/b. \-6=lumbar vertebra 1-6. s.cd. =sacro-caudal vertebree. pyaar
lb, \-2=lumbo-sacral vertebrie 1-2. s.th.=sacro-thoracic ,, ; EESTI
me.c.= median element of casque. v. = vestigial thoracic rib.
n.c.=nasal element of casque. v.me.=veutral border of mesethmoid.
vb.=obturator fissure,
PLATE XLV.
Fig. 1. The buccal cavity of Rhea americana (p. 248), to show the large choane
characteristic of the Palwoguathe.
Fig. 2. A neossoptile of Calodromas elegans (p. 162), after Pycraft, to show the large
aftershaft, equalling the main shaft in size, the complex rami of the main
shaft, and their production forwards into horny filaments resembling the
grooved ribbon-shaped bands of the young Ostrich (fig. 7).
Fig. 2a. A radius from the distal end of the main shaft.
PHYLOGENY OF THE PALAOGNATHAZ AND NEOGNATH &. 289
Fig. 8. A neossoptile of Rhea americana (p. 164): neither the main nor the aftershaft
ig. 10.
ealull,
, te,
have a distinct rhachis. The main shaft is represented by three stiffened
rami, produced into filaments as in Tinamous and Ostrich, the aftershaft by
- numerous slender rami.
ig. 3a. A portion of a radius from fig. 3 (p. 164). Note the absence of fila.
e. 4, A neossoptile of Dromeus nove-hollandiev (p. 162). The aftershaft is barely
distinguishable ; the main shaft well developed: compare with figs. 3-7.
. 4a. A portion of a radius of fig. 4. Fila are yet represented but are small.
ge. 5. A portion of a radius of Casuarius casuarius, showing the still further
suppression of the fila.
. 6. A neossoptile of Apterya mantelli (p. 163). There is no aftershaft.
e. 6a. A portion of a radius of fig. 6. Note that fila are numerous and large,
indicating a less degenerate condition than figs. 4a and 5.
.7. A neossoptile of Struthio camelus, showing the three peculiarly modified
rami described on p. 164, and representing the main shaft, and the numerous
downy rami representing the aftershaft.
iad
. Ta. A portion of a radius of fig. 7. These are flattened as in Crypfuri (fig. 2).
Note the “ eye-spots.”
. 8. Dissection of the air-sacs of the left side of a Casuarius unappendiculatus
(p. 252), showing the three divisions of the anterior intermediate sacs and the
positions of the ostia throughout. Note the position of the anterior end of
the posterior abdominal air-sac—forcing down the roof of the posterior inter-
mediate. The posterior abdominal, furthermore, is seen to take the form of
a‘more or less sausage-shaped sac adherent to the inner wall of the posterior
intermediate. It encloses a part of the kidney.
g. 9. Dissection of the left lung of Casuarius unappendiculatus (p. 252), to show
the costo-pulmonary muscle, meso-, ecto-, and ento-bronchia and their
ramifications. Note that the ento-bronchia 1 and 3 between them supply all
the tubule of the ventral surface of the lung described on p. 252. The
systems fed by ento-bronchium 1 have been exposed by dissection, those fed by
ento-bronchium 2 lie caudad of these and are concealed by the aponeurosis.
Ventral aspect of a dissection of the wing of an adult Casuarius casuarius,
to show the hypertrophied calami described on p. 165.
Dorsal aspect of a preparation of the wing of an Ostrich. After
Wray.
. Dorsal aspect of a portion of the wing of Rhea americana (p. 156), to show
the remarkable distribution of the coverts in transversely oblique bundles.
Ventral aspect of the wing of Apteryx mantelli (p. 161), to show the number
of remiges and their remarkable resemblance to the young Cassowary described
on p. 165.
VOL. Xv.— PART v. No. 24.—December, 1900. 25
29) ON THE MORPHOLOGY ETC. OF THE PALZOGNATHA AND NEOGNATHA.
Fig. 14. A remex of a nestling Casuarius australis, dorsal aspect, before the loss of
the rhachis described on p. 165.
Fig. 14a. Ventral aspect of fig. 14, to show the peculiar rhachis.
Fig. 14}. Section of the rhachis of figs. 14-14 a (p. 165).
Fig. 15. A remex of Apteryax mantelli (p. 166), for comparison with that of the young
Cassowary described p. 165.
ADDITIONAL LETTERS.
a. =aftershatft.
a.int.1-3 =anterior intermediate air-sacs 1-3.
a.s.=ala spuria.
ap.=aponeurosis.
c.=calamus.
c.c,=carpal covert.
ch.=choane.
cl.=claw.
¢.p.m. = costo-pulmonary muscle,
e.r.= carpal remex.
c.r.=cubital remiges.
*—=hypothetical carpal covert.
OER a REELEINC Xs
e.=eye-spots.
ect.=ecto-bronchia.
ent. =ento-bronchia.
eu. = eustachian aperture.
f.=filum.
h.=heart.
k.=\kidney.
mb. =meso-bronchium.
m.d.r.=metacarpo-digital remiges.
o.=ostium posterius.
obl.sep.= oblique septa.
o.ent. 3=ostia of ento-bronchium 3.
o.mb, 1-2=ostia of outer and inferior lateral
branches (1-2) of meso-bronchium.
o.sub.=ostia leading into sub-bronchial sac.
DIMI 5p pre-bronchial sae.
p.0.=pre-bronchial air-sac.
2? d2
p.int.= posterior intermediate air-sac.
p.abd. = posterior abdominal air-sac.
p.a.=pulmonary artery.
para.b.=parabronchia.
p.v.=pulmonary vein.
rd.=yradius.
rh.=rhachis.
rh.""=rhachis of aftershaft.
rm. =ramus.
r.r.=ribbon-shaped prolongations of rami.
r.g.=rima glottidis.
t.=tongue.
i.f.=terminal filaments of the rami.
t.mj.=tectrices majores.
tmd= *,, medi.
i — minores.
tes — sae marginales.
3~——==passage from ento-bronchium 1 to
vestibule.
4.2 =traverses entire length ento-
bronchium 1 to vestibule.
5» =passes from ento-bronchium 1 to
vestibule.
6=—» =traverses whole length ento-
bronchium 4.
traverse lateral meso-bronchia into
inter-
anterior and posterior
mediate air-sacs.
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CONTENTS.
VI. A Monograph of the Genus Casvarius. By the Hon. WALTER Rorascuitp, Ph.D.,
PZS. With a Dissertation on the Morphology and Phylogeny of the Pauao-
anaroa (Rarira and Crypturi) and NwoenaTia (Cartnata). By W. P.
Pronamt.. (Plates*XXIE-XLV.). - 2 ee ee page 109
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feo)
VII. Notes on the Transformations of some South-African Lepidoptera.
By Lieut.-Colonel J. Maucotm Fawcerr }.
Received October 17, 1899, read February 6, 1900.
[Puates XLVI.-XLIX.]
THE following paper is the result of a collection and observations made during a
residence of a year at Ladysmith and Pietermaritzburg, Natal, during which particular
attention was paid to the habits and early stages of the Lepidoptera of the district.
CONTENTS.
I. Remarks on the Climatic and Natural Conditions of Natal: p. 291.
II. Observations on the Rhopalocera: p. 292.
III. Observations on the Heterocera: p. 302.
IV. Explanation of the Plates: p. 319,
T.—RemMarkKS on THE Curmmatic AND Natural Conpitions or NATAL.
The climate may be described in short as one marked by extreme and rapid changes
of temperature, except in what is known as the “Coast Belt.” On the higher
plateaux, about 3000 feet, a change of wind from west to east has, in the experience
of the writer, caused a sudden drop of 22° of the thermometer in one hour; while in
the colder months (June to September) the range between day- and night-temperatures
is frequently as much as 70°. Such rigorous conditions, taken in conjunction with the
want of covert, may account for the extremely poor butterfly-fauna of the higher levels.
The rainy season, here coincident with the hotter months, usually begins in November
and lasts till the middle of March, and is characterized by exceptionally severe
thunderstorms and heavy sea-fogs and rains from the south-east ; during the remainder
of the year the weather is dry, the days being warm and the nights very cold.
Natal may be divided, for the purpose of these Notes, into three well-defined regions
or zones, as the country rises in successive plateaux inland :—
1. The Coast Belt or Zone, from sea-level up to 1000 feet elevation. (Tropical
Zone.)
2. The Middle Zone, from 1000 to 5000 feet elevation. (Subtropical Zone.)
3. The High Levels; 5000 feet elevation and over, (Temperate Zone.)
? Communicated by R. Trimen, F.R.S., F.Z.8.
VOL. XV.—Part VI. No. 1.—April, 1901. 20
prri,
292 LIEUT.-COL. J. M. FAWCETT ON THE
1. The Coast Zone, in which Durban is situated, has a tropical climate, and abounds
in thick jungles or “ bush ” (as it is called in South Africa) of virgin forest, the open
spaces being mainly under cultivation for tea, pine-apples, sugar-cane, and bananas.
In this district Bellevue, some 12 miles out of Durban, and Pinetown are good
localities for Butterflies, while a fine tract of virgin forest, called the ‘* Berea,” extends
to the very outskirts of the city of Durban; but, as a rule, all the bush in this zone is
full of Butterflies and Moths of species characteristic of the tropics.
2. ‘The Middle Zone comprises (for the purpose of these Notes) the Maritzburg
district. The country here is much wore open, and consists of grassy downs with here
and there a little mealie cultivation, and a few plantations of wattle (Llephantorrhiza
burchellii) and blue gum (/ucalyptus globulus), imported from Australia.
About 70 miles inland a plateau runs north and south, parallel to the sea, and
attaining a height in some places of 5000 feet : on the eastern slopes of this range are
the only coverts of virgin forest in this zone; they are known as the “Town Bush ”
(above Maritzburg) and “ Karkloof” (about twenty miles north of it). In these coverts
alone are the rarer species of Rhopalocera to be found: the commoner species, the
larvee of which feed on more or less cultivated plants, such as orange, lemon, passion-
flower, vine, oak, &c., being found in Maritzburg and its vicinity. It is remarkable,
however, how many larve in Natal feed on imported plants.
3. The High Levels comprise the country above the 5000 feet plateau above alluded
to up to the ‘Transvaal and Orange Free State. It mainly consists of open grassy
downs (called “ veldt” by the Boers), with here and there rocky knolls (‘‘ kopjes” or
‘“‘koppies ”) covered with large boulders, entirely devoid of timber, with the exception
of a few imported blue gums planted round the farm homesteads ; from this district,
however, came but few of the forms mentioned below.
1I1.—OBSERVATIONS ON THE RHOPALOCERA.
Most of the species which have been observed are double-brooded, the larvee of the
first brood appearing in November and December, changing to pupe in January, and
the imagines appearing early in February; while the second brood is met with in
March, and remains in the pupal stage till the following October. ‘Those which are
single-brooded generally appear as imagines in October.
In the case of species which have already been figured and described, merely the
writer’s experiences and observations in rearing them are recorded, and- the reader is
referred to the published descriptions and figures.
Mr. J. F. Quekett, Curator of Durban Museum, and Mr. W. Haygarth, of Natal
Government Railways, have rendered much valuable assistance in the preparation
of these Notes, and have presented many of the larve described to the writer.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 293
Mr. J. Medley Wood, Curator of the Durban Botanic Gardens, has kindly identified
the various food-plants.
The larve are figured natural size, except where stated otherwise.
RHOPALOCERA.
Family NYMPHALIDE.
Subfamily DaNaAIN& (Bates).
1. Danais cHrysippus (Linneus).
The larva and pupa of this well-known species have been already figured by
Mr. Trimen, in ‘ Rhopalocera Africe Australis,’ plate i., 1862.
The South-African larva seems to differ considerably from De Nicéville’s description
of the Indian form in ‘ Butterflies of India, Burma, and Ceylon,’ where only “ two large
yellow patches” on the second and third somites are mentioned, and nothing is said of
the crimson bases to the “ dark retractile (?) horns.”
The larve reared in Maritzburg fed on “ Gomphocarpus fruticosus,’ a common plant
in marshy places, and also on the ‘ veldt” in Natal; on being cut the plant exudes a
bitter white milky substance, which may account for the bitter taste of the imago,
which is said to be its protection, ‘This larva is much infested by the maggots of a
very small hymenopterous insect, and out of many examples taken, only two survived
to hecome imagines. I have heard similar experiences detailed by other collectors.
The imago is, however, common all over Natal, and is the only butterfly to be seen
on the high levels during the colder months. I have noticed the alcippus, but not the
dorippus form in Natal.
Subfamily AcR a#1N & (Bates).
2. AcR#A ACARA (Hewitson).
The larva and pupa of this species have been already figured by Mr. Trimen in
‘South African Butterflies,’ i. plate i. figs. 1, 1 a.
Numerous examples of this larva were reared in Maritzburg on a species of Passi-
flora, which is a common plant on the verandahs of houses. ‘The period passed in the
pupal stage in January was only 11 days. Some of these larve were placed in a box
in which some of the same species had already changed into pups, and the Jarvie
272
294 LIEUT,-COL. J. M. FAWCETT ON THE
proceeded to devour the pup, although there was plenty of the food-plant in the box.
This is the only species in which I have noticed this carnivorous habit. I have not
obtained this species in the coast districts.
3. ACREA PETR#HA (Boisduval). (Plate XLVI. fig. 1, larva; figs. 2, 38, pupa.)
Larva. Ground-colour pale golden brown, with a dorsal and lateral black lines, and
a black transverse line on each segment bearing two largish white spots and six long
branched black spines, those on 3rd, 4th, and 5th segments being longer than the
remainder. Head large proportionately to body, black with a white bifid mark on
front. ‘Thoracic legs and claspers yellowish. The young larve reared were all blackish
in colour, and fed in companies on Oncoba kraussiana (Planch.).
Pupa appears to be dichromatic, some being pale brown and others ferruginous ;
in both forms the fine black lines and spots peculiar to Acrwa pup are much reduced.
The pupal stage lasted 15 days in January.
‘The imago is common in the Durban and Maritzburg districts.
4. ACR@A ENCEDON (Linneus) = A. lycra (Fabricius). (Plate XLVI. figs. 4, 4,
larva ; fig. 6, pupa.)
Larva, Slaty black, with a yellow lateral line above prolegs and claspers. On each
segment three deep black fine transverse lines enclosing two white patches dorsally and
two yellow patches laterally. On the centre black line of each segment are placed six
black spines (branched). Head, thoracic legs, and claspers black.
Pupa waxy white, with the usual fine black lines on the wing-covers and black spots
with orange centres on the abdominal segments.
The larva figured was not fully grown. It became one-fifth longer and propor-
tionately thicker before changing into a pupa.
It feeds on a species of Commelina, a common weed in gardens at Maritzburg.
The imago is one of the commonest butterflies in Durban, both on the Berea and
in the Back Beach Bush ; but is not so often seen in Maritzburg.
5. ACRHA RAHIRA (Boisduval). (Plate XLVI. fig. 7, larva; figs. 8, 9, pupa.)
Larva. Back and sides blackish; thoracic legs, claspers, and a line above them
chrome-yellow. A dorsal white stripe, and on each segment four yellow spots from
which spring four branched yellow spines, the lower pair springing from the yellow
spiracular line. These spines are shorter than in the majority of Acrwa larve. Head
yellow.
Feeds on a species of groundsel, Erigeron canadense (Linneus).
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 295
Two figures of the pupa are given: one pupa is waxy white and similar to the pup
of other Acrew, the other ferruginous. The ferruginous pupe had nearly always
been attacked by ichneumons, with which the larve were much infested. The imago
is a common butterfly in the coast district.
6. Acr#a BUxTONI (Butler). (Plate XLVI. fig. 10, larva; figs. 11, 12, pupa.)
Larva. Pale buff dorsally, deepening to pale green on the sides, with a buff lateral
spiracular line above thoracic legs and claspers, which are also buff. ‘Two dorsal pale
green stripes, interrupted on every segment by a pale yellow transverse stripe bearing
four black branched spines; below these are two buff-coloured spines springing from
the buff spiracular line. Head yellowish.
Feeds on a species of nettle, locally called “ pink hibiscus” (although it is not a
hibiscus at all). It is a common plant on the Berea, Durban, where I found the
larva, and has been identified for me by Mr. Medley Wood as Triumfetta rhomboidea
(Jacq.).
The larva is here figured, x $, to exhibit the detail.
Pupa waxy white, with the usual fine black lines and spots with orange centres,
beautifully gilded ; pupe formed in the dark, however, inside a box, are slaty black.
Imago fairly common in Durban district.
7. PLANEMA ESEBRIA (Hewitson).
This larva and pupa have been figured by Trimen, ‘South African Butterflies,’ i.
pl. i. figs. 2& 2a. I took it ona species of nettle (Urtica sp.), in the covert known
as the “‘ Back Beach Bush,” near Durban. My specimens resulted in two females of
the white variety A. The imago is a very common butterfly in the Durban district.
Subfamily NyMPHALIN& (Bates).
8. JUNONIA CEBRENE (Trimen). (Plate XLVI. fig. 15, pupa.)
This larva has been already figured by Trimen, ‘South African Butterflies,’
pl. 1. fig. 4.
1 reared the larva from an egg which I saw deposited by the female on Justicia
natalensis, a common plant amongst the grass of the veldt.
Pupa chocolate-brown, with four rows of small yellow spots on each segment, and a
submarginal row of white spots round the edge of the wing-covers, with a row of four
white spots inside them, ‘The specimens reared passed 13 days in the pupal stage
296 LIEUT.-COL. J. M. FAWCETT ON THE
(23rd Jan. to 4th Feb.). I reared one specimen from the egg which I saw deposited
by the female, but regret that I made no notes of the earlier stages, save that the
young larve were, during the first two moults, lighter and more violaceous in colour,
and the spines were proportionately much smaller.
The imago is common everywhere in Natal.
9. JUNONIA cLELIA (Cramer). (Plate XLVI. fig. 14, larva; fig. 15, pupa.)
Larva. Dark slaty black, covered with minute yellowish-white atoms, and each
segment bearing black branched spines as in the larva of J. cebrene, from which it only
differs in being lighter coloured and brownish underneath, and in having an interrupted
white lateral line above spiracles.
Feeds on Asystasia coromandeliana.
Pupa. Dark chocolate-brown ; wing-covers lighter ; yellowish-white spots on thorax
and each segment and wing-covers, as in pupa of J. cebrene. Duration of pupal stage
11 days (March 1 to 12).
Imago very common in Durban, more rarely seen in Maritzburg.
10. Pyramers carpur (Linneus).
The larva of this well-known species has been so often described that any description
here would be superfluous; 1 will therefore merely state that I reared it ona large
species of thistle named Stobawa discolor (Harv.). The larva was very similar to that
of Junonia clelia, but differed in being longer in proportion to its width ; the branched
spines were yellow instead of black, and the black dorsal line was more defined ; the
pupa was beautifully gilded.
11. Cuaraxes canprope (Godart). (Plate XLVI. fig. 16, larva; fig. 17, pupa.)
Larva. Grass-green, irrorated with minute yellow spots, which coalesce into a lateral
line below the spiracles, which ends on the 12th segment in a bifid tail. On the 6th
and 8th segments are placed three yellow spots with orange centres, one dorsal and
two subdorsal, the dorsal spot being larger than the subdorsal spots and more oval in
shape. Head dark green in front, brownish to pale yellow at the sides, with four light
brown serrated horns, two springing divergently from the top of the head and two
from the sides.
Feeds on a species of Croton, probably Croton sylvaticus (Hochst).
Pupa. Grass-green ; very short and thick, suspended by tail to stem of food-plant.
I am indebted to Mr. Walter Butcher for a specimen of this larva from the neighbour-
hood of Durban, where the imago is common on the Berea in the summer months.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 297
Family PAPILIONID.
Subfamily Prerin & (Bates).
12, Prmris severtNA (Cramer). (Plate XLVI. fig. 18, larva; figs. 19, 20, pupa.)
Larva. Dark green on back, covered all over with very fine whitish hairs. ‘Two
lighter subdorsal lines, and below them a row of yellow spots, one on each segment,
placed in a darker green line. A light yellow lateral line along spiracles above a light
green line just above thoracic legs and claspers, which, with under surface, are white.
Head dark brown with minute white spots; 1st segment light yellow anteriorly.
Feeds on Capparis corymbifera.
Pupa. Bluish green with two yellow spots on each segment subdorsally, and four on
each side of thorax. A fine black line along dorsal side of thorax, and two small white
tubercles with black bases at angle of wing-covers.
Imago common in Durban and Maritzburg.
This larva is described in ‘South African Butterflies’ as ‘Dull reddish sandy on
back, with a median longitudinal streak of violaceous grey.” It may be dimorphic ;
but I have never met with a specimen of this colour (‘S. African Butterflies,’ vol. iii.
p. 69)
13. Pipris HELLICA (Linneus). (Plate XLVI. fig. 21, larva; fig. 22, pupa.)
Larva. Yellow above, bluish green underneath ; a light blue dorsal stripe, and on
each side of it a broader greenish stripe, and below these a pale yellow spiracular
stripe; body covered by minute black papilla and some sparse short hairs; head
bluish, covered with black dots.
Feeds on a common weed in the grass of the veldt, of which JI did not get the name.
Pupa. Yellow above and green underneath, covered with minute black spots.
Pupal stage lasted only 7 days in February.
Common everywhere in Natal.
1“ My description (0. c.) of this larva was made from numerous Natalian specimens sent alive to me by
Colonel Bowker. I observed, on the page quoted, that in its earlier stages the larva is dorsally much tinged
with greenish, and it is of interest to find that the green tint commonly persists till the full-grown stage.”-—
t. TRIMEN.
298 LIEUT.-COL. J. M. FAWCETT ON THE
Subfamily PaPILIONIN& (Bates).
14. Papitto poticenes (Cramer). (Plate XLVI. figs. 23-28, larva; figs. 29, 50,
pupa; fig. 31, head of larva with tentacles.)
Larva. 1st stage. Chocolate-brown, each segment with a transverse yellow stripe ;
three yellow spines on thoracic segments, and a yellow bifid tail consisting of two
yellow divergent spines. The larva does not exceed 4 inch in length in this stage.
2nd stage. Ground-colour ferruginous red, with transverse black stripes as follows :-—
one between each two segments, and two on each segment enclosing a greyish-blue
streak hetween them. Ist and 2nd segments and bifid tail yellow with black points.
A black spiracular line divides the ferruginous upper portion of the body from the
lower portion, which is violaceous grey. ‘Three thin black spines with scarlet bases on
thoracic segments. Head ferruginous, thoracic legs and claspers pale grey.
3rd stage. Ground-colour has become green, fading to yellow at sides above the
black lateral line; the black line dividing the segments has become obsolescent, and
the two black lines on the segments enclosing the blue streak have become ferruginous.
Otherwise as in 2nd stage.
In this stage larva attains its full size.
4th (and final) stage. Entirely green above, with the exception of the three black
spines on the thoracic segmeuts, which still retain their scarlet bases, and are joined by
transverse black streaks; under surface violaceous grey. Head and Y-shaped tentacular
organ pale green.
Larva feeds on Uvaria caffra, called by the natives “ Maswinda,” a common bush in
the coverts of the coast-belt.
Pupa green, very broad, and very much flattened, with a very prominent thoracic
dorsal projection pointing forward; the lateral projections pale ferruginous; two
subdorsal lines of black spots on each side.
One pupa formed ina box was pale pinkish, with the subdorsal region strongly
marked with black.
Only the 2nd stage of this larva is described in ‘South African Butterflies’ (vol. iii.
p. 203).
The imago is common in the Durban district.
15. Papitio BRasipAs (Felder). (Plate XLVI. figs. 32-34, larva; figs. 35, 36,
pupa.)
Larva. st stage. Pale yellowish ferruginous, with reddish diagonal streaks on
sides; back of 3rd, 4th, and 5th segments occupied by an olive-green area, with a
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 299
median dorsal pink stripe from head to tail, bounded by an olive-green stripe on each
side. Ist segment reddish brown superiorly, with short tubercles on each side; a pair
of short pink tubercles dorso-laterally, bounded inwardly by dark olive-green, and
joined by a fine pink transverse line on 2nd and 35rd segments; tail bifid.
2nd stage. Pale green above, darker green underneath ; all the markings described
in Ist stage defined by slightly darker green, except the olive-green area on the back
of the thoracic segments, which has become dark green in this stage, while the pink
median dorsal line and transverse stripes have become a rather startling whitish cross.
3rd (and final) stage. Shortly before changing into a pupa the larva loses all its
distinctive marks, and becomes very pale yellow all over, almost diaphanous.
Feeds on Popovia caffra (Kaffir name, “small Maswinda”), a common shrub in the
coast district.
Pupa. Bright green. On each side of back, from apex of dorso-thoracic projection
to anal extremity, a narrow yellow elevated stripe; on summit of back a pair of similar
stripes commencing considerably apart near base of dorso-thoracic projection, and
converging till they meet at anal extremity. A series of diamond-shaped lozenges
down the back, medially, and a series of diagonal streaks subdorsally between the two
yellow stripes, and extending beyond them to the cephalic area. Duration of pupal
stage 21 days (24th January to 16th February).
Common in Durban and the coast-zone. I have taken the allied species Papilio
leonidas in the same coverts and at the same time. It is, I should say, doubtful if
they are distinct species. P. brasidas is the commoner species in the Durban district.
I have not met with either in the vicinity of Maritzburg.
16. Papitio morania (Angas). (Plate XLVI. figs. 37-39, larva; fig. 40, pupa.)
Larva (early stage). General colour grass-green on back, incisions of segments pale
blue; two dark chocolate-brown lateral stripes above spiracles (which are white),
Head, thoracic legs, claspers, and bifid tail brownish yellow. Ist segment yellow
dorsally, bearing two short tubercles on each side; 2nd and 5rd segments bearing
dorso-laterally a pair of black spines, short, with orange bases, and joined by two broad
transverse streaks of chocolate-brown; that on the 2nd segment having three brown
spots posteriorly, and that on 3rd segment five brown spots anteriorly, and being much
larger than the other, extending posteriorly over part of the next segment; from this
three dark brown longitudinal lines, one dorsal and two subdorsal, extend to the
11th segment, where the two subdorsal lines become two chocolate-brown spots joined
by fine transverse lines, while the dorsal line (which is much finer than the others)
extends to the tail. A double lateral line of black spots along the segments, two on
each segment along the upper line, and one on each segment in the lower line.
2nd stage. On casting its last skin, the larva emerges exactly the same as in the
VOL. XV.—Part vi. No. 2.—April, 1901. 2U
300 LIBUT.-COL. J. M. FAWCETT ON THE
early stage ; but in a very short time (perhaps half a day) it loses all its distinguishing
characters in that stage, and becomes uniform pale green; a yellow lateral line above a
darker green line being all that remains to represent the two very distinct brown lateral
lines of the early stage, and the under surface, thoracic legs, claspers, and head have
all become pale green. ‘The black spines on the thoracic segments, however, remain
the same, except that their bases become more reddish. In this stage the larva is very
similar to the larva of P. policenes in its green stage; but the latter always retains its
brown lateral streak and its blue-grey under surface.
3rd (and final) stage. Like the larva of P. brasidas, this larva also becomes pale
yellow and almost diaphanous just before pupating.
It feeds on Uvaria caffra (the large “ Maswinda”), like the larva of P. policenes.
Pupa. Grass-green and more rounded than in pupa of P. brasidas. Dorso-thoracic
prominence thicker and pointing more upward; a pair of raised yellow lines from
thoracic prominence along the sides to anal extremity, also a second pair of similar
lines run subdorsally from base of thoracic prominence to tail, where they converge
and meet.
Pupe formed on Ist February remain over the winter months in this stage.
The imago is common in the vicinity of Durban.
17. Papitio pEMopocus, Esper. (Plate XLVI. figs. 46-48, larva; figs. 49, 50,
pupa; fig. 51, head of larva with tentacles.)
Larva (early stage). Ground-colour black, with short yeilow tubercles on the thoracic
and 10th, 11th, and 12th segments. Top of Ist segment yellow between the two
tubercles ; a dorsal line of diamond-shaped lozenges commences grey on the 4th
segment, becomes white on 6th and 7th segments, and darkens again to grey and
brown on the remaining segments to the tail, where it is obsolescent. On 4th, 5th,
and 6th segments are white lateral spots, which converge and meet the dorsal line of
lozenges on 5th and 6th segments; similar white lateral spots on 10th, 11th, and
12th segments. ‘ail bifid and yellow, head reddish. ‘This larva, which apparently
mimics the droppings of birds, varies but little in this stage.
Full-grown stage. The ground-colour in this stage varies according to the colour of
the lemon-leaves on which the larva is feeding, from pale green almost yellow to dark
green; the transverse streaks on the sides also vary from light brown almost
ferruginous in the pale specimens to deep velvety black in the dark green examples.
The most common form has a grass-green ground-colour, with velvety black transverse
streaks on the three thoracic segments, and black diagonal streaks on the 7th, 8th,
and 9th segments, converging till they almost meet dorsally on Sth and 9th segments.
Under surface and claspers greyish white, and a broad white lateral line above spiracles.
Head and thoracic legs reddish ; head surmounted by two reddish tubercles, between
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 301
which the ¥-shaped tentacular process is situated. The transverse black streaks on
the 2nd and 3rd segments have a line of ferruginous ocellate spots variegated with
light blue and yellow. The black diagonal streaks on 7th, 8th, and 9th segments are
mottled with rather indistinct purple ocellate markings, and edged outwardly with
fine lines of pale yellow. There are also short diagonal streak-like spots on the 9th
and 12th segments just above the white spiracular line. Two yellow tubercles over
the anal extremity.
The pupa also varies in colour according to its surroundings: the most common
form is light green, paling to two yellow subdorsal streaks; head and cephalic
prominences (which are well-marked) brown, as is also the thoracic prominence
posteriorly, from which a broad brown streak extends almost to the anal extremity.
Several dark green and brown spots on the segments.
I first bred this larva at Ladysmith, in the high level zone, in April 1898, on a plant
named Clausena inequalis; the larve began pupating on the 7th April, and the first
imago emerged on 12th October, 1898. I noticed that these specimens were smaller
than those which I subsequently reared on lemon-leaves at Maritzburg in December
and January following; these individuais only averaged a fortnight in the pupal stage.
This is the commonest and most widely distributed Papilio in S. Africa.
18. Papinio nireus (Cramer). (Plate XLVI. figs. 41-44, larva; fig. 45, head of
larva with tentacles.)
Larva. 1st stage. Ground-colour black, paling to greenish on the sides, the under
surface, thoracic legs, and claspers being greenish yellow; head, thoracic, 10th, 11th,
and 12th segments surmounted by orange-coloured tubercles in pairs ; the pairs on the
three thoracic segments being joined by transverse orange lines bearing two to three
tubercles each, smaller than those at the extremities. 7th, 8th, and 9th segments
ferruginous ; 5th, 6th, and 10th segments black superiorly with white sides. In this
stage the larva somewhat resembles that of P. demodocus. In some individuals the
black markings are replaced by very dark green.
2nd stage. Ground-colour grass-green superiorly, under surface greyish white. No
ferruginous markings on 7th, 8th, and 9th segments, and no lateral white spots on
dth, 6th, and 10th segments, these being replaced by a pale lemon-coloured lateral
streak. ‘The tubercles, which have become light lemon-coloured, are confined to the
thoracic and 12th segments.
3rd stage (final). Ground-colour generally dark green, but, like P. demodocus, varying
according to the colour of the leaves of the food-plant. A wide pale greenish-blue
oval area with a thin whitish line across its centre, on the summit of the 3rd and 4th
segments, bounded anteriorly and posteriorly by a transverse line of ocellate spots :—
the anterior series outlined in black on an ochreous transverse band, the ocelli at the
2u2
302 LIEUT.-COL. J. M. FAWCETT ON THE
extremity of the band being larger than the rest; the posterior series indistinct: four
ocelli in each series have light blue centres. A lateral white stripe along the spiracles
from 4th to 12th segments: tubercles on anal segment yellow, well-defined, and joined
by a raised yellow pale streak; tubercles on back of Ist segment pale and incon-
spicuous. ‘The Y-shaped tentacles in this species are bright crimson and much longer
than those of P. demodocus; the scent emitted is, however, similar.
Pupa. Bright green; point of dorso-thoracic prominence, two spots below it (at
abdominal base), and edge of lateral abdominal angles creamy reddish. In shape
somewhat attenuated anteriorly, cephalic processes short and directed laterally
outward, so that the frontal line of head is widened and but slightly concave.
Thoracic lateral angles moderately acute ; dorsal prominence also elevated acutely but
not inclined forward. Sides of abdomen widely flattened, and so extended as to form
a very marked angle on each side of third abdominal segment; whence abdomen
narrows very rapidly and greatly to the extremity. Infra-pectoral region, where wing-
covers meet, very strongly convex.
A marked constriction dorsally at junction of thorax and abdomen.
In its natural position, attached vertically or nearly so, head uppermost, the anterior
portion of this pupa is seen to be very much more bent backward than it is in
P. demodocus. J have taken this description of the pupa, with some alterations, from
‘South African Butterflies,’ as I thought it so well describes the peculiarities in shape
of the pupa.
The imago is not so often met with as P. demodocus, and seems to prefer wooded
country more. In its haunts, however, and in the coast district it is one of the
commonest species.
III.—OBsERVATIONS ON THE HETEROCERA.
Family SATURNIIDZ.
1. Acrias Mimos&. (Plate XLVII. figs. 1, 2, larva; fig. 3, cocoon.)
Larva. Ground-colour grass-green, with paired dorsal series of long conical humps
with yellow apices, surmounted by three or four short black hairs, and the same
number of longer yellow hairs, from 2nd to 10th somites inclusive ; the 11th somite
has only one similar dorsal hump, and the Ist somite no hump, with the black and
yellow hairs planted just above the head ; a subspiracular line of small tubercles with
similar hairs. Between each somite, from 3rd to 11th, a yellow transverse streak
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 305
folding over a blue transverse streak at the incisions of the somites. Head and
thoracic legs ferruginous, spiracles white, with ferruginous centre.
In the early moults this larva is ferruginons, the head and thoracic legs being black.
Feeds on Sclerocarya caffra (Sond.), called in Natal “the wild mango tree”—a
common tree in the coast districts, but not found higher up.
When about to change the larva spins a greyish-white silky cocoon, with a line of
ventilation-holes round the exit end, which it attaches longitudinally to a twig of the
food-plant. The pupa is chocolate-brown and of the usual shape. This moth appears
to be double-brooded, the larva appearing in November and February, and the perfect
insects in January and the following October.
Common at Sydenham, near Durban.
2. NUDAURELIA WAHLBERGI (Boisduval). (Plate XLVII. fig. 4, larva; fig. 5, pupa.)
Larva. Ground-colour deep velvety black, each somite bearing four branched
ferruginous spines with reddish bases, two subdorsally and two laterally, from 2nd to
12th somite inclusive, the spines on the 2nd somite having black bases. Between the
subdorsal and the lateral row of spines are placed a collection of small yellow spots on
each somite; the red bases of the subdorsal and lateral spines being joined on 10th
and 11th somites. Spiracles white: a subspiracular row of small yellowish tubercles
bearing a few whitish hairs, one on each somite. Head and legs concolorous with
body.
Feeds on English oak, but has also been taken in large numbers on peach-trees, in
gardens in Maritzburg—both these trees being imported species in Natal. When full-
fed it burrows underground, where it makes a sort of web !, inside which it undergoes
its transformation. Individuals which changed about 14th March reappeared as
imagines on 6th May and subsequent days.
Pupa dark chocolate-brown.
Common in Maritzburg.
3. Buna#A cAFFRARIA (Stoll). (Plate XLVII. fig. 8, larva.)
Larva. Ground-colour deep velvety black ; each somite, from 4th to 12th, bearing
eight yellow tubercular processes, two subdorsally, two laterally, and four (in two rows)
on each side subspiracularly. The 2nd somite bears four black processes, two sub-
dorsally and two laterally ; the 3rd somite bears four black processes, as in the 2nd,
«The group of Antherwa to which this species belongs—Nudaurelia—is noted for forming no cocoon,
and the closely allied species 4. (N.) cytherea, menippe, and tyrrhea, as well as Buncea caffraria, as far as I
have observed, simply bury themselves without any attempt at forming even a ‘ web.’”—R, Tren.
304 LIBUT.-COL. J. M. FAWCETT ON THE
and two small yellow processes on each side, in line with the subspiracular processes
on the other somites. Spiracles red; those on the 4th to 11th somites being sur-
rounded by an irregularly shaped red area. Head and legs concolorous with body.
Larva figured has not attained its full size.
Mr. Quekett writes: ‘‘The food-plants of this Bunea are, at Durban, Celtis kraus-
siana (Bernh.), and Ekebergia meyeri (Presl), and at and near Maritzburg a species of
Cussonia, on which I have taken the Jarve in the Botanic Gardens.”
This larva, which much resembles that of NV. wahlbergi, undergoes its transforma-
tion underground in a similar manner to that larva, and is double-brooded, the larvae
appearing in November and December, and the second brood in March and April;
these latter individuals remain pupe until the following October, my first specimen
emerging on 12th of that month.
Pupa dark reddish brown; very similar to that of Gynanisa maia (Klug).
4, GYNANISA MAIA (Klug). (Plate XLVII. fig. 6, larva; fig. 7, pupa.)
Larva. Ground-colour pale green, with paired dorsal series of humps ; each somite,
from 2nd to 12th, bearing four silver spikes with yellow points inclined backwards, one
subdorsally and one laterally on each side; also a number of small silver spots placed
irregularly over each somite; the first somite has no spikes, but is raised to a sharp
ridge, with a black edge, which may be of assistance to the larva in forcing its way
through the ground. Spiracles purple, and immediately beneath them a purple lateral
line having on its lower edge a yellow raised lateral line bearing a small orange-coloured
tubercle on each somite, and thickening considerably above anal claspers, where it has
a series of small black tubercles superiorly. Head green, with black side-streaks
defining the eye. Under surface darker green with minute white spots, and a row
of small red tubercles, one on each somite just above the claspers, which are green.
Thoracic legs pale brown banded with black.
Feeds on the common wattle (Elephantorrhiza burchellii, Bth.) and on several species
of thorny acacia. When full-fed the larva burrows underground, and there undergoes
its transformation, making little or no preparation in the way of a web.
The pupa is dark reddish brown, and is remarkable for the large size of the antenne
covers. ‘The moth is double-brooded, the larve appearing in November, and the
imagines emerging end of January ; the second brood of larvee appearing in March and
emerging in the following October. At Fort Napier, Maritzburg, the wattle-trees were
completely cleared of their leaves by the March brood this year.
If the collector possesses one female, any number of males can be secured by placing
the female in a box on the verandah. The writer has seen as many as twenty males at
one time fluttering round the box about 10 p.w. Next morning the verandah was
strewn with their wings, a cat, as was subsequently discovered, having caught and
devoured them.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 305
5. PSEUDAPHELIA APOLLINARIS (Westwood). (Plate XLVII. figs. 11, 12, larva;
fig. 13, pupa.)
Larva. Ground-colour bluish grey, each somite with a broad transverse indented
black streak, thickest on 2nd somite, and two finer black transverse lines across the
upper part of the body between the somites. Body broader in the middle than
anteriorly or posteriorly. Above anal extremity a sharp-pointed black horn—a fine
black spiracular line, and immediately below it a broad raised ferruginous line bearing
a small black tubercle crowned with some short yellow hairs on each somite. Under
surface and abdominal claspers pale yellow. Head, thoracic legs, and anal claspers
black. A tuft of short hairs on the summit of the 2nd somite.
Feeds on Jurrea heterophylla (Smith), and undergoes its transformation underground ;
the period passed in pupa state (in March) was 17 days.
Pupa dark red-brown, abdominal somites strongly marked ; chiefly remarkable for
the long sharply-pointed tail resembling the point of a thorn.
The imago is a common species, flying by day, in the Berea Bush, Durban.
6. Urota stvope (Westwood). (Plate XLVII. fig. 9, larva; fig. 10, pupa.)
Larva. Ground-colour velvety black, with a series of paired pale yellow humps formed
by a thick raised transverse yellow streak, bearing a pair of short tubercles crowned
with short yellow hairs dorsally on each somite, and a series of similar short tubercles
laterally at the lower end of each streak. Spiracles pale yellow, and below them an
interrupted, raised, subspiracular line covered with short yellow hairs. Head black,
surmounted by a yellow transverse band where it joins Ist somite, and crowned with
some yellow hairs. Thoracic legs black, abdominal claspers yellow banded with black,
anal claspers and extremity yellow, all covered with yellow hairs.
Feeds in companies on the “ Kaffir boom ” tree (EKrythrina caffra, Thb.).
Pupa dark red-brown and of the usual form, with a sharpish spur or horn at the
anal extremity. ‘The transformation is carried out underground.
Fairly common in Durban district.
7. Lupra sminax (Westwood). (Plate XLIX. fig. 6, larva; fig. 7, cocoon.)
Larva. Ground-colour rufous, with irregular pale blue spots surrounded by a thin
white line. On each somite a short tuft of black hairs, sarmounted by a tuft of longer
and finer hairs of same colour, and the first five and last somites covered with short
yellow hairs. Head, legs, and claspers brown.
This larva, which looks, through a microscope, like a piece of old china ware in
colour, is one of the most unpleasant larvee to handle that I have ever met with. The
short black hairs on each somite possess poisonous qualities, which produce on the
hand a white rash akin to that produced by a bad stinging from nettles. It forms a
306 LIEUT.-COL. J. M. FAWCETT ON THE
cocoon round itself, with its hairs, on the side of the box. Food-plant oak; but I
have also reared the larva on Jasmine (Jasminium pubigerum).
On quitting South Africa on a short leave of absence home in May 1899, I left two
specimens of this larva, in the pupal stage, together with all my larvee which had not
completed their transformations, in the care of a friend who was remaining on in
Pietermaritzburg, as I was afraid that, during the voyage, the heat of the tropics
would cause them to emerge prematurely ; but to prevent, as I thought, the possibility
of any mistake in identifying the imagines, I placed each larva or pupa in a different
box, with a small ticket, with a reference to the plate and figure of the larva in my
sketch-book, which I requested my friend to attach to the pin of the imago when it
emerged.
I subsequently returned to South Africa when war broke out, and on the relief of ©
Ladysmith was invalided home ; on passing through Pietermaritzburg in March 1900,
my box containing the emerged imagines (which had all been placed in a large cork
box which I had left with my friend) was returned to me, when I found my ticket
referring to the drawing of this larva pinned to an imago of Ludia smilax (Westw.).
On reaching England I forwarded a figure of the imago to Mr. R. Trimen, and asked
him for the name of the moth, as I was unacquainted with it. Iwas therefore much
surprised to receive the following reply from him :—‘I know the moth you figure in
your note of yesterday very well, it is Ludia smilax (Westw.); but I am much
surprised to hear that this species resulted from the ‘ old china’ caterpillar figured by
you, because I have by me figures of no fewer than three quite different larvee
attributed to ‘ smilav’—I mean different from each other, as well as entirely different
from yours. If you are quite certain about the moth resulting from the old china
larva, then this stage of L. smilav presents amazing variability. W. D. Gooch and
Dr. J. KE. Seaman made drawings of this caterpillar ; they are rather rough, but those
of the apparently more prevalent variety agree in representing a pale greenish larva
with broad black rings; Gooch thought that this was the male, but he does not seem
to have proved this, though he bred a male from one of this pattern. Gooch’s second
form of larva is quite like the other in shape, tubercles, hairs, &c., but the black rings
are almost everywhere broken up into mere scattered spots, and the ground-colour,
generally, is dull yellow; he thinks this is the female.
**Much more amazing is Dr. Seaman’s second form of smilax larva; it is black,
marbled with white, and with vivid scarlet rings! There is no doubt that some larve
seem to vary almost indefinitely, but if ths is a true case all through, I think it is
unsurpassed.”
it is therefore with some doubt that I append the description of this larva as the
larva of Ludia smilax (Westw.), inasmuch as I cannot vouch myself, from personal
observation, for its accuracy, though I have no reason to doubt the accuracy of my
friend, who is certain that no mistake has been made.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 307
Family SPHINGID.
Subfamily ACHERONTIIN A,
8. ACHERONTIA ATROPOS (Linneus).
No description is necessary of the larva of this well-known species; and 1 will
merely remark that I reared it on a species of Spathodia, an imported tree which is
often met with in Berea Bush, Durban. The larva underwent its transformation on
14th February, and the imago emerged on 17th March after a pupation of 38 days.
This is one of the instances of larvee feeding on imported trees in Natal.
I also reared the dark form (body uniformly fuscous, first three somites pink
subdorsally) at Maritzburg, on Jasminium pubigerum (also an imported plant in Natal).
This larva underwent its transformation on 5th April, and hibernated in the pupal
stage ; this form is much rarer than the green and yellow form.
Subfamily SMERINTHINA.
9. LopHostetaus puMoLINIT (Latreille)!. (Plate XLVIII. fig. 7, larva; fig. 8,
pupa.)
Larva. Ground-colour very pale green, a pair of blue-black steely branched spines
with pale yellow bases and basal areas subdorsally on each somite from 2nd to 10th.
The 1st somite has no spines, and the 11th has only one spine, thicker than the others
and replacing the horn in other species. A lateral row of smaller black spines
springing from the upper edge of a spiracular row of large yellow spots; a subspiracular
row of small black spines springing from lower edge of above-mentioned spots, and,
below these, two spines placed diagonally on the 4th, 6th, 7th, 8th, and 9th somites
just above the claspers ; the 5th somite having three spines, and the 2nd, 5rd, and
10th somites one spine each in this series. Abdominal claspers yellow, with black
extremities, each extremity bearing three short black divergent spines ; anal extremity
and claspers horny and rufous, with a broad black edging. Head pale green superiorly,
pale ferruginous inferiorly; two black vertical stripes on the face, ending with a
detached black spot above them. Sides of head black, as in the larva of Acherontia
atropos. ‘Thoracic legs pale ferruginous, banded with black.
1 «A brief description of the extraordinary spiny larva of this Hawk-moth—drawn up by me from a
coloured drawing by Mr. W. D. Gooch, and a coloured photograph by the late Dr. J. KE. Seaman—was
published by Prof. Meldola in his English edition of Weismann’s ‘ Studies in the Theory of Descent,’ vol. ii,
pp. 527, 528 (1882).”—R. Trimen.
VOL. XV.—PART V1. No. 3.—April, 1901. 2x
308 LIEUT.-COL. J. M. FAWCETT ON THE
Feeds on Hibiscus tiliaceus (Linneus). When full-fed the larva burrows under
ground, and forms a sort of chamber with a web, in which it undergoes its trans-
formation.
The pupa is dark reddish brown, and only distinguishable from that of A. atropos
by its greater thickness and the abdominal somites being more horny.
The specimens reared remained in the pupal state from February till the following
October.
The imago is not uncommon in Durban and the coast districts.
Subfamily CH HROCAMPIN A.
10. CHaRocaMPA CAPENSIS (Linneus). (Plate XLVIT. figs. 17, 18, larva; fig. 19,
pupa.)
Larva. Ground-colour pale green, thickly irrorated subdorsally with darker green
diamond-shaped spots, from 5th to 11th somites; these spots coalesce into a series of
diagonal streaks along the somites subdorsally and spiracularly. A paler green lateral
stripe from 5th to 11th somite, with a dark green stripe along its upper edge; horn
very short and pink ; a reddish “eye ”-like spot edged with white superiorly on 4th
somite. Head and claspers green, thoracic legs pink ; spiracles red.
When ready for its transformation the larva becomes dull pink flesh-coloured, the
lateral line only remaining green; the eye-like spot and diamond-shaped irrorations
become black, and the ground-colour of the dorsal region becomes dull yellowish.
The figure is taken from a larva in this final stage.
Feeds on common vine.
Pupa. Head, wing-covers, and dorsal region dark brown, with black markings ;
abdomen pale pink flesh-coloured. Time passed in pupa state about five weeks.
When about to change the larva spins a web amongst leaves, &c., on the surface of
the ground in some sheltered place, and undergoes its transformation inside.
The imago is fairly common in the evening in flower-gardens in Maritzburg.
I have also reared a red form of this larva, which may be described as follows :—
Ground-colour uniformly ferruginous; a pale subdorsal stripe from 2nd somite to
horn, white above, darkening to yellow beneath, bounded inferiorly by dark red diagonal
stripes on each somite from 4th to 10th. On 4th somite a conspicuous ‘eye ”-like
spot, black superiorly, white inferiorly.
Mr. R. Trimen writes to me of this larva:—* The red form of C. capensis larva is,
at Cape Town (where the species feeds on the cultivated vine), very much rarer than
the green one; the colour of the latter is decidedly protective on the cultivated vine,
but that of the red one was not so (except slightly, perhaps, when the larva is on
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA, 309
the older woody stems). I imagine, however, from one of the red ones which
I found on its native food-plant, the wild Cape vine (Cissus capensis), that this
tint was probably acquired in relation to the latter plant, which is densely clothed
with red down on the younger shoots and underside of the leaves. I have noticed
the common Cape Butcher-bird (fiscus collaris) taking capensis-larve from the
cultivated vines at Cape Town and spiking them on thorns; such persecution would
seem to show that these larve are in need of protective resemblance to their food-
plants.”
11. CuHarocampa osiris (Dalman). (Plate XLVIII. figs. 3, 4, larva; figs. 5, 6,
pupa.)
Larva. Ground-colour pale green, thickly irrorated with darker green diamond-
shaped spots as in C. capensis. A paler green lateral stripe from 5th to 11th somites,
ending at a very small yellow horn. ‘Two large “ eye ’-like spots (green with a light
blue centre, on which are placed six minute white spots), surrounded by a narrow light
yellow iris, on 4th somite; two smaller round yellow spots on 5th somite. Head,
spiracles, and claspers green ; mandibles and thoracic legs pink.
When ready for transformation the larva becomes pale reddish brown with a
violaceous suffusion; the intersections of the somites and lateral stripe become white,
while the diamond-shaped spots become black where they are most thickly placed near
the junction of the somites; the large “eye-like ” spots become brown with a green
iris, while the yellow spot becomes black. The effect of these changes is to give the
larva, when viewed dorsally, a very startling resemblance to a snake of the banded
Krait species.
Feeds on the common vine.
Pupa \ight brown dorsally, variegated with black spots and streaks defining head,
eyes, antenn, wing-covers, nervures, and abdominal somites; breast, wing-covers, and
underside of abdomen pale pink flesh-coloured.
This pupa is chiefly remarkable for its abnormally long snout, the palpi-covers being
produced to form a shield or covering for the haustellum.
The transformation takes place amongst leaves on the surface of the ground, without
much attempt at making a chamber with a web, the leaves being loosely strung
together by means of a silken thread.
This larva was received from Bellevue, a suburb of Durban ; the imago seems to be
rather uncommon.
12. CHHROCAMPA BALSAMIN® (Boisduval). (Plate XLVIII. fig. 1, larva ; fig. 2,
pupa.)
Larva. Ground-colour bright grass-green ; from 6th to 11th somites a brown inter-
2x2
310 LIEUT.-COL. J. M. FAWCETT ON THE
rupted dorsal streak, and traces of two double subdorsal streaks mainly formed by
small brown spots arranged in transverse rows across the somites. On the 4th somite
a pair of black “ eye ”-like spots with a white iris; on the 5th somite a pair of red
“eye ”-like spots with a white iris, and between these the dorsal streak above
mentioned is continued, green in colour, to the head; thoracic legs pink; head and
claspers green; horn light brown and straight, ending in a point.
Feeds on Jussiwa repens (Linneus).
Pupa. Head, thorax, and wing-covers yellowish brown, remainder of body reddish
brown, paler dorsally and abdominally; palpi-covers slightly produced and prominent.
The pupa is formed in a light web amongst leaves on the surface of the ground, in the
usual manner of the larvee of Chwrocampine.
The imago emerges in about a fortnight (in February).
Found in Durban district.
13. Cuarocampa ipricus (Drury). (Plate XLIX. fig. 1, larva; fig. 2, pupa.)
Larva. Ground-colour pale green; each somite, from 4th to 10th, with a pair of
triangular spots, brown superiorly, yellow inferiorly, at its anterior edge. The pair of
spots on 11th somite are lengthened into a brown and yellow streak, and end in the
horn, which is brown, and sharp-pointed like a thorn. The pair of spots on the 4th
somite are larger than the remainder, and bear on their yellow area an oval, black,
“eye ”-like spot with a white pupil near its upper edge ; each spot has three minute
white points on its brown area; spiracles white; a pinky-white subspiracular line, and
above it a collection of brown points at each intersection of the somites. Head and
claspers green, thoracic legs light brown.
Feeds on Spermacoce natalensis (Hochst.), a common herb among the grass of the
veldt.
Pupa. Pale golden brown, with small black spots defining the nervures on the wing-
covers, and sprinkled over the abdominal somites ; spiracles black. Period of pupation
about 23 days. Habits and transformation similar to other species of the genus.
The imago is very common, hovering over flowers in gardens in Maritzburg, from
the early afternoon till dusk.
The larva and pupa are here figured x $ in size on nature.
14. Dapayis neru (Linneus).
The transformations of this insect are so well known that figures and detailed
descriptions are unnecessary here; but it may be as well to note that numerous
specimens were reared from the larva on oleander, which plant is a common shrub in
gardens in Maritzburg. ‘The colour of the larva in Natal is very pale green (almost
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 311
yellow) dorsally, darker below; a white lateral line with numerous small silver-white
spots sprinkled over, above, and below it, the lower edge of the line is bounded by a
broad light blue area extending downwards as far as the spiracles from 6th to 10th
somites. On 3rd somite a pair of large, black, double-pupilled, eye-like spots, the
pupils being white, surrounded by a blue and a crimson iris.
Pupa pale ferruginous ; habits and transformation similar to those of C. capensis.
Duration of pupa state 10th or 11th February to 4th March.
Subfamily SPHINGINA.
15. ProroparcE Mauvriti, Butler. (Plate XLVIII. figs. 9, 10, larva; me, IL,
pupa ; fig. 12, larva, dorsal view of 1st four somites.)
I am informed at the British Museum that the name MJacrosila solani, by which this
species is usually known in South Africa, should be applied to the Mascarene form.
Larva. Ground-colour grass-green, under surface darker. Paired humps on Ist
and 2nd somites ; a purple dorsal stripe from 4th somite to horn; lateral oblique
purple stripes from 4th to 10th somites ; these stripes join the dorsal stripe on every
somite, and are defined inferiorly by parallel narrow white oblique stripes; horn
ferruginous, long, and beset with yellowish tubercles; spiracles small, red, with black
centres. Head green, with vertical black stripes on the face and sides, as in larva of
A. atropos. Thoracic legs black, claspers green.
Feeds on Duranta plumieri, a common shrub in gardens in Natal; but which,
according to Mr. Medley Wood, “is included in the flora of Natal by mistake, and is
most certainly not indigenous.”
I have also reared a dark form of this larva, which may be described as follows :—
Ground-colour very pale brown, the oblique purple stripes of the green form being
represented in this form by oblique stripes of dark violaceous grey covered with
blackish atoms. Head pink, with lateral and frontal vertical black stripes; first three
somites pink subdorsally, surmounted by paired dorsal diamond-shaped fuscous spots.
It will be observed from this description that the head and first three somites of this
larva bear a striking resemblance to the dark form of the larva of Acherontia atropos.
Spiracles black; prolegs and claspers pale brown with fuscous bands; horn yellow and
serrated.
The specimen figured fed on Dahlia variabilis (an imported plant in Natal),
Pupa. Dark reddish brown, with a long external sheath for the proboscis. ‘The
transformation is effected underground, where, as in the case of A. atropos, the larva
constructs a sort of chamber for the pupa.
Mr. Trimen states “ the case of M/. solani seems to me especially interesting, because
not only is the larva imitative of that of A. atropos in both forms, but the moth also
312 LIEUT.-COL. J. M. FAWCETT ON THE
is decidedly imitative of atropos-moth, so much so that I have taken it for atropos
when at rest on a tree-trunk. At Cape Town both the dark forms (of solani and
atropos \arve) are rare in comparison with the green ones.”
Subfamily MACROGLOSSIN&.
16. CEPHONODES HYLAS (Linneus). (Plate XLVIII. figs. 13-18, larva; fig. 19, pupa.)
This well-known insect has been reared from six different forms of larva in Natal,
which show almost every gradation from an almost wholly green larva with white
subdorsal stripes to an almost wholly black one, in which the subdorsal stripes are
replaced by very dark grey ones.
The various forms will now be described in detail.
Larva. 1st form. Ground-colour pale green, a white-bordered light blue dorsal line,
and a thin white subdorsal line bordered superiorly by a thin carmine line, the combined
lines ending in a yellow streak at the base of the horn. Spiracles white, bordered
with red edges, that on 11th somite surrounded by a rufous area. Head green, Ist
somite beset with yellow tubercles superiorly ; thoracic legs and claspers brownish ; horn
green, curved and pointed, and beset with yellow tubercles. Plate XLVIII. fig. 13.
This seems to be the Indian form described by Hampson in ‘ Fauna of British India.”
2nd form. Ground-colour pale green; a white-bordered blue dorsal line ; a subdorsal
white line defined in places superiorly and inferiorly by small black oval spots ; under
surface, legs, and claspers pale brown. Spiracles white, surrounded by oval red areas ;
a yellow subspiracular line, interrupted and defined inferiorly with black, which curves
up vertically on the posterior half of each somite from 6th to Yth, otherwise as in
Ist form. Plate XLVIII. fig. 14.
3rd form. Similar to 2nd form, but differing in the following characters :—The dorsal
line is dark grey ; the black oval spots defining the white subdorsal line inferiorly are
much more numerous and extend downwards between the spiracles till they meet a
black subspiracular line. The red areas surrounding the spiracles more enlarged.
Plate XLVIII. fig. 15.
4th form. Similar to 3rd form, but differing as follows :—Dorsal line darker, the
black oval spots defining the white subdorsal line inferiorly are coalesced into a con-
tinuous black lateral line; and the whole body below it is pale brown, with only an
isolated patch of green on each somite round the red spiracular areas. Ist somite
yellow, covered with small yellow tubercles. Plate XLVIII. fig. 16.
5th form. Dorsal and subdorsal stripes as in 4th form. Ground-colour black ;
a rufous area round each spiracle, and a few irregular yellow marks on each somite ;
a few traces of the yellow subspiracular line, and a yellow patch at base of horn;
claspers and anal extremity brown; head and Ist somite yellowish. Plate XLVIII.
Mes lis
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 315
6th form. Entirely black, with the exception of the red areas surrounding the
spiracles, and the head, base of horn, and anal extremity and claspers, which are dull
ferruginous ; dorsal line black; subdorsal line and Ist somite dark grey. Plate
XLVITI. fig. 18.
All these forms of larva were found at the same time feeding on Gardenia and also
on Kraussia lanceolata (Sond.). When ready for their transformations the larve
burrowed underground, and there underwent their change to pupe; the perfect insects
emerged in from 3 to 4 weeks. In order to test whether all the different forms of
larva produced the same imago, each form of larva was placed in a separate box
and carefully labelled, but when the imagines emerged it was impossible to detect any
difference between them.
Pupa. Dark reddish brown.
‘The imago seems common all over Natal.
Family ZYGANID.
17. Zycaya (AnteRIs) aMPpLA (Walker). (Plate XLIX. figs. 27, 28, larva; fig. 29,
cocoon.)
Larva. Ground-colour white, with pink stripes and black spots. A white dorsal
line bordered by two subdorsal rows of black spots, two on each somite; beneath these
a pink lateral line bordered inferiorly by a white spiracular line, on which is situated
a row of smaller black spots in groups of two on each somite. A subspiracular pale
fulvous line, and below it an interrupted line of black lunules, one on each somite ;
under surface and claspers white ; head black, with a white bifid frontal stripe.
General form of larva thick in middle, tapering towards extremities, and very similar
to larvee of Lyceenide in shape.
Feeds on a bush with very large leaves with spiny sedges (like holly) of which 1
have not been able to obtain the name.
The pupa is formed in a strongly-woven oval cocoon, usually pale fulvous, and
affixed to a leaf of the food-plant; time passed in pupal stage about 25 days (March
3rd or 4th to March 29th, 1899).
Found in large numbers on the food-plant in the Back Beach Bush, Durban.
Family LASIOCAMPID &.
18. Gonomera Postica (Walker). (Plate XLIX. figs. 3, 4, larva; fig. 5, cocoon.)
Larva. There are two common forms of this larva in Natal, one with long grey
hair and one with long fulvous hair.
1st form. Ground-colour deep velvety black, thickly covered on sides with long grey
hair ; a thin black dorsal line and paired black subdorsal patches of short black hairs
on each somite from 4th to 12th. Head black, covered with brownish-grey hairs ;
314 LIEUT.-COL. J. M. FAWCETT ON THE
spiracles, thoracic legs, and claspers red. The 2nd form is similar to the Ist, but with
the long hairs bright fulvous. Both forms feed on the same food-plant at the same
time, grow to the same size, and produce male and female imagines irrespectively.
This larva should not be handled without gloves, as its hair penetrates the skin of
the hand and causes much irritation. It appears in great numbers in November and
early December, feeding on common wattle (Elephantorrhiza burchellii) and also on a
species of thorny acacia ; the second brood appearing in March and April.
‘The pupa is formed inside a cocoon beset with the hairs of the larva, which cause a
very irritating effect to the fingers if it 1s grasped carelessly, and is affixed to a branch
of the food-plant, as shown in the figure.
Pup formed in April hybernate in this state, and emerge as imagines in the middle
of October.
In common with species of Saturniidz, the male of this moth seems to possess in a
high degree the power of discovering the whereabouts of the female, and the possession
of one female will enable a collector to secure a large number of males.
Common and widely distributed in Natal.
Family LYMANTRID A.
19. Duicuta FascrAta (Walker). (Plate XLIX. figs. 13, 14, larva.)
Larva. Ground-colour velvety black ; dorsal area white, thickly covered with white
hairs, except on the 4th, 5th, 11th, and 12th somites, which are surmounted by tufts
of long black hairs; a dorsal red spot on the 9th and 10th somites; a spiracular line
of red spots, and below this an interrupted white line bearing long white hairs.
Head black; thoracic legs and claspers red.
Feeds on species of Bauhinia, and also on rose-bushes and oak, the two latter being
imported plants in Natal.
When ready for its transformation, the larva forms a silky cocoon with its hairs,
inside which it changes and remains as a pupa for about 24 days (Dec. 30th to
Jan. 25rd).
Common in gardens in Maritzburg.
20. DASYCHIRA GEORGIANA, sp.n. (Plate XLIX. fig. 19, imago; fig. 20, larva ;
fig. 21, pupa.)
Allied to D. horsfieldii. Head and thorax white; branches of antenne fulvous.
Fore wing white, irrorated with black scales; the antemedial lines more angled than
in D. horsfieldii; the hind wing white, with some darker hairs on the internal area.
Abdomen orange, with a dorsal line of black spots on first four somites. Underside
white, no cell-spots.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 315
Larva. Ground-colour velvety black, with very long pale yellow hairs; each somite
with two red and four greyish tubercles, from which spring the long silky yellow
hairs, except the 4th, 5th, 6th, and 7th somites, on which the two dorsal tubercles
coalesce, and from these spring four thick dorsal tufts of shorter grey hairs, and
11th somite, which bears a tuft of shorter blackish hairs. Head red above, black
beneath ; thoracic legs and claspers red, and a red transverse band on 12th somite.
Feeds on common wattle (Hlephantorrhiza burchelli).
The pupa is formed in a slight web constructed from the hairs of the larva, and
is pale fulvous with black markings; wing-covers greyish, with veins delineated by
thin black lines. The descriptions are made from two specimens reared in Maritzburg
in February and March. There is a female specimen of the imago in the British
Museum unnamed.
Family HYPSID A.
21. Hypsa apuipas (Hopffer)=H. subretracta (Walker). (Plate XLVII. fig. 16,
larva.)
Larva. Chocolate-brown ; under surface greyish ; an indistinct dark dorsal line.
Three red tubercles, one dorsally and two subdorsally, on Ist to 3rd somites; paired
red subdorsal tubercles, four on each somite, on a black subdorsal line, from 4th to 12th
somite ; a broad white spiracular streak on 5th, 6th, 10th, and 11th somites. Head dark
brown ; thoracic legs and claspers fuscous. A few sparse brown hairs on the body.
Feeds on the common fig, both cultivated and wild; the specimen figured fed on the
cultivated fig, but I have reared others which I found as pupe in hollows on the
trunks of the wild fig-trees round the gardens in the square at Durban. When about
to change, the larva rolls up a fig-leaf in which it spins a web, and the same process
is gone through in the hollows of fig-tree trunks. ‘The specimen figured remained as
a pupa from 18th February to 42nd March.
Pupa black, with polished surface.
Imago fairly common in Durban and Maritzburg.
Family ARCTIIDA.
22. SPILOSOMA PUELLA (Druce). (Plate XLIX. fig. 15, larva.)
Larva. Ground-colour dark fuscous, each somite with six large and two small black
tubercles surmounted by tufts of black hairs. A pale ferruginous dorsal line, darker
on the summit of each somite; under surface paler; head and thoracic legs black,
spiracles ferruginous. ;
Feeds on the Grenadilla passion-flower.
vol. XV.— Part vi. No. 4.— April, 1901. 2Y¥
316 ’ LIEUT.-COL. J. M. FAWCETT ON THE
Pupa black, with polished surface, formed inside a cocoon constructed out of the
hairs of the larva. ‘The imago emerged in about 17 days (March 8-26).
Common in Maritzburg.
Subfamily ARCTIIN &.
23, PHISSAMA FLAVA (Walker). (Plate XLIX. fig. 8, larva.)
Larva. Ground-colour pale fuscous, each somite with a red-brown area superiorly,
on which is situated a dorsal line of white spots surrounded by black, one on each.
somite; also a series of black tubercles bearing tufts of black and reddish hairs.
Head, thoracic legs, and claspers ferruginous.
Feeds on Ornithogalum eckloni, a species of lily, and other plants.
When ready for its transformation the larva weaves a cocoon with its hairs and
remains a pupa for about 17 days (Ist to 17th December in the case of the specimen
figured).
The imago is common in Maritzburg district.
24, PHISSAMA SCREABILE (Walker). (Plate XLIX. fig. 9, larva.)
Larva. Ground-colour greyish white, the somites broadly black transversely and
beset with long black and white hairs; the white spaces between the somites traversed
by fine black transverse lines ; under surface black. Head, thoracic legs, and claspers
bright red.
Feeds on Ornithogalum eckloni, also on mealie: a larva reared on the latter plant
differed from the individual figured in its lighter parts being buff instead of white ;
this larva produced a female imago.
The pupa is formed inside a cocoon constructed with the hairs of the larva.
Duration of the pupal stage about three weeks (9th to 29th December, specimen figured).
Imago is common in Maritzburg district.
Subfamily NycTEOLIN &.
25. Farias INSULANA (Boisduval). (Plate XLIX. fig. 25, larva; fig. 26, cocoon.)
Larva. Ground-colour pale bluish-black, with deep black markings; Ist to 3rd
somites pale brown; 4th somite with a white transverse streak; an interrupted white
dorsal line from 4th to 8th somite; 9th to 12th somite pale brown dorsally. A sub-
dorsal line of paired short brownish spines with scarlet bases; a lateral line of similar
paler spines. Head and legs pale brown.
Feeds on Hibiscus.
‘The pupa is formed in a stout thickly-woven cocoon attached to stem of food-plant.
The larva is figured x $, to get in detail.
Common in gardens in Maritzburg.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA. 517
Family AGARISTID.
26. EUSEMIA BUTLERI (Walker). (Plate XLVII. fig. 14, larva; fig. 15, pupa.)
Larva. Ground-colour light bluish-grey, with fine black transverse stripes and spots
on each somite; a reddish spiracular line. Head, thoracic legs, and Ist and 11th
somites superiorly reddish ochreous; a few sparse hairs distributed over the somites ;
thoracic legs fuscous; ventral claspers blue-grey, anal claspers reddish ochreous.
Feeds on common vine (cultivated).
The larva undergoes its transformation underground, and remains a pupa about
one month.
Pupa reddish fuscous, very rugged in appearance, the abdominal somites very
strongly defined.
The imago has a very strong penetrating scent.
Not uncommon in Maritzburg.
Family NOCTUIDA.
Subfamily TRIFIN &.
27. GLoTTULA PANCRaTI (Cyr.). (Plate XLIX. fig. 17, larva; fig. 18, stem of food-
plant showing hole.)
Larva. Reddish brown, each somite with three small black warty spots, one dorsally,
two subdorsally, and two small pale yellow subdorsal spots; between each two somites
a transverse row of five oblong pale yellow spots, divided mesially by the intersection
of the somites. Head, legs, and claspers ferruginous; a few short fulvous hairs on
the somites.
This larva feeds on a species of lily, eating the flowers as well as the stems; also
making a burrow inside the stem (as shown in fig. 4¢). It can eat its way out of any
box, except tin. When full-grown it undergoes its transformation underground.
Pupa red-brown, with strongly-marked somites.
The tmago is noteworthy owing to its exceedingly small and short pale green
haustellum, which would seem to be of little use to it.
28. AGRoTIS SEGETIS (Schiff.). (Plate XLIX. fig. 16, larva.)
Larva, Ground-colour pale brown, with small black spots placed irregularly on the
somites ; fuscous dorsal and lateral lines, the former narrow, the latter rather broad
and bounded inferiorly by a narrow white spiracular line. Under surface, thoracic legs,
and claspers pale fulvons. Head ferruginous, the eyes defined by two crescentic black
frontal lines.
Feeds on the cultivated vine, and undergoes its transformation underground.
The pupal stage lasted about one month, in March.
Imago common in Maritzburg in February and March.
22
318 LIEUT.-COL. J. M. FAWCETT ON THE
Subfamily QUADRIFIN &.
99, PoLYDESMA GLAUCINANS (Guen.). (Plate XLIX. fig. 22, larva; fig. 23, pupa
in web.)
Larva. Ground-colour bright grass-green, with a broad white dorsal line from head
to 12th somite, which bears on its centre a narrow pale blue dorsal line ; an interrupted
white lunular lateral line, bordered with black superiorly. Head, thoracic legs, and
claspers green; the first pair of abdominal claspers aborted; the 12th somite tapering
to a point above the anal claspers.
Feeds on wattle (Elephantorrhiza burchellii).
Pupa pale brown, formed in a cocoon among the stems of the food-plant.
Imago very common in Maritzburg in March.
30. Tanropyea sytvina (Stoll), (Plate XLIX. figs. 10, 11, larva; fig. 12, pupa in
cocoon.)
Larva (early moults). Ground-colour buff; on each somite, except Ist and 11th, a
broad very dark crimson transverse band; on Ist and 11th somites he band is pale
ferruginous and bears some small black spots. In the final moult the above-mentioned
bands become deep velvety black, with the exception of those on the Ist and 11th
somites, which remain the same, the ground-colour becoming pale greenish. Head,
legs, and claspers pale ferruginous.
Feeds on Ornithogalum eckloni (Sch.), a species of lily common in swampy places.
When full-fed the larva burrows underground and constructs a chamber for the pupa,
which is surrounded by a strong hard envelope, of a consistency similar to cement, with
a small hole at one end for the exit of the imago.
Pupa deep red, with black antemedial and postmedial lines on the wing-covers.
The period of pupation lasts about five weeks.
The imago is not uncommon in Maritzburg in January and February.
Family GEOMETRID A.
31. Boarmra acactarta (Boisduval), (Plate XLIX. fig. 24, larva.)
Larva. Ground-colour pale reddish brown, inclined to olive-green subdorsally ; a
pair of dorsal processes with whitish centres on 5th somite, and a pair of dorsal white
spots on 11th somite. Head, legs, and claspers reddish.
Feeds on a small species of sunflower, which is a common plant in gardens at
Maritzburg.
The transformation to pupa is effected underground.
TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA, 319
My specimens of the imago were identified as B. acaciaria at the British Museum,
but they are much paler and have fewer transverse bands than the examples in that
collection, or the specimen figured by Hampson in ‘Fauna of British India,’ vol. iii.
p. 265.
IV. EXPLANATION OF THE PLATES.
PLATE XLVI.
Fig. 1. Larva of Acrea petrea (Boisd.), p. 294.
Figs. 2, 5. Pupa of ditto.
Figs. 4, 5. Larva of Acrwa encedon (Linn.), p. 294.
Fi . Pupa of ditto.
cao
Fig. 7. Larva of Acrwa rahira (Boisd.), p. 294.
“Figs. 8, 9. Pupa of ditto.
Figs. 11, 12. Pupa of ditto,
Fig. 13. Pupa of Junonia cebrene (Trimen), p. 295.
Fig. 14. Larva of Junonia clelia (Cramer), p. 296.
Fig. 15. Pupa of ditto.
Fig. 16. Larva of Charaxes candiope (Godart), p. 296.
Fig. 17. Pupa of ditto.
Fig. 18. Larva of Pieris severina (Cramer), p. 297.
Figs. 19, 20. Pupa of ditto.
Fig. 21. Larva of Pieris hellica (Linn.), p. 297.
Fig. 22. Pupa of ditto.
Fig. 23, Larva of Papilio policenes (Cramer), 1st stage, p. 298.
Figs. 24, 25. Ditto, 2nd stage (dorsal and lateral views).
Fig. 26. Ditto, 3rd stage.
Figs. 27, 28. Ditto, final stage (dorsal and lateral views).
Figs. 29, 30. Pupa of ditto (dorsal and lateral views).
Fig. 31. Head of larva with V-shaped tentacles protruded.
Fig. 32. Larva of Papilio brasidas (Felder), early moults, p. 298.
320 LIEUT.-COL. J. M. FAWCETT ON THE
Figs. 35, 54. Ditto, final moult (dorsal and lateral views).
Figs. 35, 36. Pupa of ditto (lateral and dorsal views).
Figs. 37, 38. Larva of Papilio morania (Angas), early moults (lateral and dorsal views),
p. 299.
Fig. 59. Ditto, final moult.
Fig. 40. Pupa of ditto.
Fig. 41. Larva of Papilio nireus (var. lyeus) (Cramer), Ist stage, p. 501.
Fig. 42. Ditto, 2nd stage.
Figs. 43, 44. Larva of Papilio nireus (var. lywus) (Cramer), final stage, p. 301.
Fig. 45. Head of larva with V-shaped tentacles protruded.
Fig. 46. Larva of Papilio demodocus (Esper), early moults, p. 300.
Figs. 47, 48. Ditto, final moult. Fig. 48 shows the attitude adopted by the larva
when the V-shaped tentacles are protruded.
Figs. 49, 50. Pupa of ditto (dorsal and lateral views).
Fig. 51, Head of larva with V-shaped tentacles protruded.
PLATE XLVII.
Fig. 1. Larva of Actias mimose, early moults, p. 302.
Fig. 2. Ditto, final moult.
Fig. 5. Cocoon of ditto, showing ventilating holes.
Fig. 4. Larva of Nudaurelia wahlbergi (Wallengren), p. 303.
Fig. 5. Pupa of ditto.
Fig. 6. Larva of Gynanisa maia (Klug), p. 304.
Fig. 7. Pupa of ditto.
Fig. 8. Larva of Bunea caffraria (Stoll), p. 305.
Fig. 9. Larva of Urota sinope, p. 805.
Fig. 10. Pupa of ditto.
Figs. 11, 12. Larva of Pseudaphelia apollinaris (Westwood) (dorsal and lateral views),
p. 305.
. 13. Pupa of ditto.
. 14. Larva of Eusemia butlert (Walker), p. 317.
. 15. Pupa of ditto.
Fig. 16. Larva of Hypsa aphidas (Hopff.), p. 315.
Fig. 17. Larva of Cherocampa capensis (Linneus) (green form, final stage), p. 308.
Fig. 18. Ditto, red form.
Fig. 19. Pupa of ditto.
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PLATE XLVIII.
Larva of Cherocampa balsamine (Boisduval), p. 309.
Pupa of ditto.
Larva of Cherocampa osiris (Dalman), p. 309.
Ditto, final change before pupation.
6. Pupa of ditto (ventral and lateral views).
. Larva of Lophostethus dumolinii (Latreille), p. 307.
. Pupa of ditto.
. Larva of Protoparce nauritt (Butler), green form, p. 511.
. Ditto, dark form.
. Pupa of ditto.
. Dorsal view of first four somites of larva, dark form.
. Larva of Cephonodes hylas (Linn.), Ist form, p. 512.
. Ditto, 2nd form.
. Ditto, 3rd form.
. Ditto, 4th form.
. Ditto, dth form.
. Ditto, 6th form.
. Pupa of ditto.
PLATE XLIX.
. Larva of Cherocampa idricus (Drury), X $, p. 310.
Pupa of ditto, x ¢.
. Larva of Gonometa postica (Walker), red form, p. 315.
Ditto, grey form.
Cocoon of ditto.
. Larva of Ludia smilax (Westwood), p. 305.
. Cocoon of ditto.
. Larva of Phissama flava (Walker), p. 516.
. 9. Larva of Phissama screabile (Walker), p. 316.
re, 1G),
. Ditto, final moult.
2. Pupa of ditto, in underground chamber.
. Larva of Dulichia fasciata (Walker), p. 514.
. Ditto, dorsal view.
. Larva of Spilosoma puella (Druce), p. 315.
. Larva of Agrotis segetis (Schiff.), p. 317.
. Larva of Glottula pancratii (Cyr.), p. 317.
Larva of Teniopyga sylvina (Stoll), early moults, p. 318.
os
322 TRANSFORMATIONS OF SOME SOUTH-AFRICAN LEPIDOPTERA,
Fig. 18. Stalk of food-plant, showing entrance to chamber made by larva.
Fig. 19. Dasychira georgiana, sp. n., p. 314.
Fig. 20. Larva of ditto.
Fig. 21. Pupa of ditto in web.
Fig. 22. Larva of Polydesma glaucinans (Guen.), p. 318.
Fig. 23. Pupa of ditto in web.
Fig. 24. Larva of Boarmia acaciaria (Boisduval), p. 318.
. 25. Larva of Earias insulana (Boisduval), x $, p. 316.
. Cocoon of ditto.
s. 27, 28. Larva of Zygena (Anteris) ampla (Walker), p. 313.
. 29. Cocoon of ditto.
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CONTENTS.
VII. Notes on the Transformations of some South-African Lepidoptera. By Lieut.-
Colonel J. Macoum Fawcerr. (Plates XLVI-XLIX.) . . . - page 291
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VIII. On some Points in the Anatomy of Polypterus.
By J. 8. Bupertt, W.A., F.Z.8., Trinity College, Cambridge.
Received and read May 8, 1900.
[Puares L.-LIL.]
ContENTS,
Page
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VI. General Considerations on the Structure and Growth of the
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XG xplanationvotthe Plates ran eenicisic serene cciciniariee 335
XIV. Explanation of the Lettering of the Figures ..............4- 338
‘I. Introduction.
HAVING obtained a large amount of material of both the species of Polypterus
found in the Gambia, in the spring of 1899, I have thought it advisable to attempt
to fill up some gaps in our knowledge of the anatomy of this most interesting of fishes.
A great deal of work has already been done in this direction by such eminent
zoologists as Agassiz, Geoffroy St.-Hilaire, Joh. Miiller, Leydig, and Hyrtl, as well as
by numerous other authors in recent years. The anatomy of the head of Polypterus
has been described in detail by Pollard, the brain by Waldschmidt, the skull by
Traquair and by Bridge, while Hyrtl has described the blood-supply of the external gill.
The foundation of this work of recent years had been laid, however, by Joh. Miiller
in his ‘ Bau und Grenzen der Ganoiden.’
In the present paper I have, I believe for the first time, described in detail the
urinogenital system of the male and female Polypterus, together with the later stages
in the development of these organs. I have also added some observations upon the
vascular system, the external gill, the abdominal pores, the anal fin, and the skull,
VOL. XV.—PaRT vil. No. 1.—April, 1901. 22
MR. J. S. BUDGETT ON SOME POINTS IN
[Sl)
bo
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Il. The Male Organs.
The following observations have been made on adult male specimens of Polypterus
senegalus and P. lapradei taken at the commencement of the breeding-season. Series
of the urinogenital organs of the young P. senegalus, 13 cm. and 9 cm. in length, have
also been carefully studied.
In the adult male a ridge of testicular tubules extends the entire length of the body,
lying parallel to and directly over the kidney on either side (PI. L. figs. 2, 3, ¢.r.). Each
testis-ridge is enlarged towards its anterior end, forming a conspicuous lobulated testis
(fig. 3, ¢.): that on the left side is situated more anteriorly than that on the right.
The testis and testis-ridge are covered by peritoneum, the two folds of which are
approximated at the base of the testis to form a mesorchium, in which spermatic veins
pass to the cardinals (fig. 3, sp.v.).
The tubules of the testis and testis-ridge open by very numerous short ducts into a
longitudinal canal extending the whole length of the gland lying in the mesorchium at
the base of the testis, and further back between the testis-ridge and the kidney (Pl. L.
fig. 3, t.d.; Pl. LI. figs. 10, 11, ¢.d.). Posteriorly this duct leaves the testis-ridge
as the vas deferens (PI. L. figs. 5, 6, v.d.), and passing backwards in the same sheath of
connective-tissue as the ureter, opens upon a papilla into the narrow neck of the urino-
genital sinus just before it opens to the exterior (PI. L. fig. 5; Pl. LI. figs. 12, 14, g.ap.).
This duct is of even calibre throughout the greater part of its length, but is
somewhat dilated in the region of the lobulated testis.
The tubules of the adult ripe testis are very numerous, and dilated with spermatozoa
and what appear to be sperm mother-cells. The tubules are embedded in lymphoid
tissue with deeply-staining nuclei (Pl. LI. fig. 10, ¢.). The spermatozoa are very small,
about the diameter of a red corpuscle in length, thickened anteriorly and tapered
posteriorly (Pl. LI. fig. 13).
The arrangement of the tubules of the testis-ridge is a simplification of that of the
testis-tubules. Here three or four longitudinal tubules are connected by numerous
transverse tubules with the testis-duct (Pl. LI. fig. 15, ¢.7.tbs.). The walls of these
tubules are lined with columnar epithelium, and are not surrounded by lymphoid
tissue as are the tubules of the testis, but by dense connective-tissue.
The tubules of this testis-ridge do not appear to be functional testis-tubules,
although amongst them were found what appeared to be traces of spermatozoa.
It seems possible that the tubules of this ridge assist in carrying away the sperm from
the testis to the vas deferens, but do not themselves actually produce spermatozoa.
In the youny Polypterus senegalus, 13 cm. in length, the tubules of the testis which
are embedded in lymphoid tissue are lined by a single layer of large-celled columnar
epithelium, while the lumina are small (PI. LI. fig. 16, ¢.tds.). The tubules of the testis-
ridge differ little from those of the adult male (Pl. LL. figs. 15, 17, ¢.r.tds.) ; the tubules,
THE ANATOMY OF POLYPTERUS. 325
however, are smaller as compared with the testis-duct in the young than in the adult.
It is noteworthy also that the vas deferens, which in the young is very thick-walled and
has a larger lumen, is much larger as compared with the ureter in the young than in
the mature individual (PI. LI. figs. 14, 18, v.d.).
In the young Polypterus the duct of the testis runs forward a short distance
anteriorly to the testis. This portion of the duct could not be traced with certainty in
the adult. No opening in the young male was found, however, into the body-cavity.
In the very young male, 9 cm. in length, the tubules of the testis are foreshadowed
by the nuclei being arranged in double rows, but there are no lumina; the duct,
however, has a wide lumen and is well formed (PI. LII. fig. 19, ¢.d.). At this stage the
tubules of the testis-ridge are not yet developed ; the duct, however, is here well formed,
as opposite the testis. Posteriorly the vasa deferentia end blindly in the wall of the
ureter (Pl. LIT. fig. 20, w.v.u.).
Ill. The Female Organs.
The following observations have been made on adult female specimens of
P. senegalus and P. lapradei taken at all times of the year, and also upon specimens
of the young female P. senegalus 9 cm. in length.
The funnel-like opening of the oviducts into the body-cavity (Pl. L. fig. 1, p.ap.od.)
were mentioned and figured by Joh. Miller ; the ducts were figured in more detail by
Hyrtl, and the ovaries were described. My own observations, however, do not in some
respects agree with those of the latter author. According to Hyrtl the two oviducts
unite to form a urinogenital sinus, into which the two ureters open by a common mid-
dorsal aperture.
I have carefully studied the adults of both species found in the Gambia, and I find
that the ureters are dilated posteriorly, lying closely approximated to each other, but
not communicating, except immediately before opening to the exterior (Pl. L. figs. 1,
4,u.; fig. 4, s.u.g.s.). Shortly before the ureters open to the exterior the oviducts open
into their lateral walls precisely as do the vasa deferentia in the male (PL. L. fig. 4, g.ap.).
Further, in the young female 9 cm. in length the course of these ducts has exactly the
same relation to the ureter as in the male, only that the oviducts are considerably more
dilated; they lie immediately over the genital ridge, which anteriorly is developed
into the ovary (Pl. LII. figs. 21, 22, 23, mes.o.).
The great difference between the sets of organs in the male and female is that in
the male the genital gland discharges directly into the duct, whereas in the female
the genital products are shed free into the body-cavity, and thence find their way to
the mouth of the duct. Were the outer wall of the duct in the female carried
forwards to enclose that side of the ovary from which the ova are shed, or were the
testis-duct in the male open anteriorly to receive the products of the testis from the
body-cayity, the arrangement would be precisely analogous in the male and female.
2272
4224
326 MR. J. 8. BUDGETT ON SOME POINTS IN
The latter appears actually to be the case in Polyodon folium, where, according to
Hyrtl, the duct of the testis, as well as the duct of the ovary, opens into the body-
cavity by means of a peritoneal funnel.
At what stage the opening of the oviduct into the ccelom is acquired in Polypterus I
cannot definitely say. It was, however, open in my youngest specimen 9 cm. in length
(Pl. LIT. fig. 22, p.ap.od.).
In young females 9 cm. in length the genital ducts have not a free opening into the
ureter, but, as in the male, the ducts end blindly in the wall of the latter (Pl. LI.
fig. 24, w.o.u.). The communication is complete, however, by the time the young
Polypterus is 12 cm. in length.
Though it would be unwise to form any definite opinion as to the significance of
these ducts until their primary origin shall have been made clear, yet, seeing that
Figs. 1-3.
Ovary of Polypterus.
Fig. 1.—Diagrammatic representation of the developing ovary.
Fig. 2.—The outer wall of the ovary between each septum beginning to bulge.
Fig. 3.—The outer wall thrown into deep folds, as in the ripe ovary.
e.0.w., external wall of ovary ; i.0.w., internal median wall of ovary ; 0', white side of ovum;
o''., black side of ovum ; 0.s., septum dividing ovary into compartments.
very immature specimens have been examined, it seems worth while suggesting that
the very high development of the genital ducts at an early stage in both sexes, and
their similar arrangements, point to their being homologous with one another and also
with the embryonic Miillerian duct. The discussion of the & priori objections and the
consequences of this conclusion need not here be discussed, in view of the probability
of the question being settled by a study of the early development of Polypterus.
THE ANATOMY OF POLYPTERUS. S27
The ovary in Polypterus develops as a genital ridge lying along the ventral surface
of the kidney, and separated from it posteriorly by the genital duct (Pl. LIL. figs. 21,
22, 23).
A large vein and artery traverse the median wall of the ovary (Pl. LIL. fig. 21, 6.v.).
The ovary becomes divided into numerous compartments, on the outer wall of which the
ova are developed (Pl. LII. fig. 21, 0.a¢., and text-fig. 1, p. 326).
With increased development of the ova in size and number, the outer wall becomes
greatly enlarged in surface, while the median wall is not thus enlarged (text-fig. 2).
It thus happens that the outer wall bulges out into great folds between the septa
(text-fig. 3).
As the ova develop they become deeply pigmented upon the pole of the ovum
which is attached to the outer wall of the ovary, Although the outer wall becomes so
much folded, nevertheless the ova retain their original “ orientation” at the extremities
of each fold, so that, when ripe, through the median wall of the ovary, which is
a thin smooth sheet, only the pigmentless poles are seen (text-fig. 3, 0’), while
Fig. 4.
oak
The ovarian ovum of Polypterus, with attachment (0.at.)
to the outer wall of a compartment of the ovary.
through the external much-folded wall only the pigmented poles are seen (text-
fig. 3, 0"). It has been stated that the outer wall of the ovary of Polypterus is without
a covering of peritoneum. It will be understood from the above-given description
of the growth of the ovary, and from the examination of text-fig. 4, that this is not
strictly correct.
IV. The Kidneys.
The kidneys are similar in male and female. They consist of two bands of excretory
tubules, glomeruli, and embedding lymphoid tissue (PI. LI. figs. 10, 11, 16) lying on
either side of the vertebral column between the peritoneum and the muscles of the
body-wall, and are constricted metamerically by the projecting myocommata, The
kidneys of the two sides do not unite (Pl. L. figs. 2, 3, £.).
The minute structure of the kidney of the young Calamoichthys has been described
by Lebedinsky. His description of the arrangement of the tubules in Calamoichthys
328 MR, J. 8S. BUDGETT ON SOME POINTS IN
will answer perfectly for that in Polypterus, except that in my youngest specimen
9 em. in length I can find no trace of the nephrostomes opening into the coelom
described by him in “ larve” 12 cm. in length. It seems, then, that Calamoichthys,
which is so much smaller in the adult than Po/ypterus, is in a more larval condition
at 12 cm. length than is the Polypterus at 9 cm.
It is possible that the nephrostomes never entirely close in Calamoichthys. However
this may be, I have found in Polypterus no trace of nephrostomes opening into the
coelom.
In my youngest specimens the uriniferous tubules were arranged in distinct
metameric masses, the metamerism disappearing in older specimens. ‘The glomeruli
did not appear to be thus arranged, there being a very indefinite number of these
structures to each metamere (PI. LI. figs. 10, 16, m.cps.). The openings of the tubules
into the ureter are far more numerous and irregular in the adult than in the young.
V. The Ureters.
The ureters lie along the whole length of the kidney between the outer ventral
edge of the latter and the body-wall (Pl. L. figs. 2,3; Pl. LI. figs. 10, 11, w.) (text-
figs. 5, 6). ‘They receive the kidney-ductules. The ureter on passing ventralwards
from the hind end of the kidney becomes dilated, and, in the male, joining its fellow of
the opposite side, forms a large urinary sinus (Pl. L. figs. 2, 5, 6; Pl. LI. fig. 12, w.s.).
The urinary sinus still passing ventralwards to a position just dorsal to the rectum
becomes constricted to a narrow neck and, just before opening to the exterior, receives
on either lateral wall the opening of the genital duct, then opens to the exterior in a
depression just posterior to the anus (PI. L. figs. 6, 7, 8, w.g.ap.).
In the female the ureters do not become confluent until immediately before they
open by a slit-lke aperture just posterior to the anus. Otherwise they resemble
entirely those of the male (text-fig. 6, p. 329).
VI. General Considerations on the Structure and Growth of the
Urinogenital Organs.
It having been shown by Balfour and Parker and also by Semon that the testis of
Lepidosteus, and perhaps also of Acipenser, was connected with the kidney-tubules,
it was of extreme interest to see whether this were the case also in Polypterus, in many
respects the most archaic of recent Teleostomes.
No such connections as a matter of fact exist. The products of the testis pass out
by a well-developed duct, which, running the same course as the ureter, opens into the
lateral wall of the latter close to its termination (text-fig. 5).
It is well known that the oviduct of the female Polypterus is short and is open
anteriorly by a wide peritoneal funnel, the ova being shed through the external wall of
the ovary into the body-cavity. This duct in the female runs a course similar to the
THE ANATOMY OF POLYPTERUS. 329
genital duct in the male, opening into the ureter on its lateral wall shortly before its
termination (text-fig. 6).
The resemblance of these ducts in the male and female is brought out still more
‘strongly by a study of the quite young fish 9 cm. in length. Apart from the fact
that the oviduct opens into the body-cavity, the arrangements in the two sexes are
identical.
Fig. 5. Fig. 6.
---OK%
SS Ss
a eee ee
——
as
p.ap.od.—e} *
i
<=
Si
oval --
Fig. 5.—Diagram of the urinogenital organs in the male Polyplerus.
Fig. 6.—Diagram of the urinogenital organs in the female Polypterus.
g-ap., genital aperture; &., kidney; ov., ovary; ovd., oviduct; p.ap.od., peritoneal aperture of oviduct ;
s.ug.s., septum of urinogenital sinus; t., testis; t.d., testis-duct; t7., testis-ridge ; t.7.d., duct of testis-
ridge; w., ureter; v.d., vas deferens.
‘The question as to whether these ducts are both homologous with the Miillerian
ducts of Elasmobranchs, Amphibia, and Amniota can of course only be settled by a
knowledge of their actual origin.
The fact remains, however, that in this most primitive fish the arrangement of the
genital ducts is nearly identical in the two sexes, and would seem to substantiate the
330 MR. J. S. BUDGETT ON SOME POINTS. IN
view which was latterly held by Balfour that primitively the testis had a duct of its
own, derived either from a Miillerian duct like that of the female Elasmobranch,
or a structure sui generis, as held by Howes and Max Weber, and that, on the other
hand, the connection of the testis with the tubules of the kidney found in the male
Elasmobranch, Amphibian, and Amniot is a secondary one.
There is, however, a difficulty with regard to the latter view in the fact that
Lepidosteus, the Ganoid fish which, as regards its ovary and oviduct, most closely
resembles the Teleostean arrangement, in having a closed ovary continuous with its duct,
is the very one which shows this supposed secondary connection of testis with kidney.
It is possible, however, that this acquirement is confined to Lepidosteus, while
other Ganoids retain the primitive condition ; and it may be that it is a feature which
has been frequently acquired independently. So that the Elasmobranch, Amphibian,
and Amniot are not necessarily a separate evolutionary line from the Crossopterygian,
Dipnoon, Ganoid, and Teleost, but the Amphibia may have acquired the Elasmobranch
arrangement after they split off from the Dipnoi, which have not acquired it.
That the arrangement in Lepidosteus is not primitive seems probable from the fact
that the testis-tubules open into a well-marked longitudinal collecting-duct, which
lies along the ureter in the same position as the testis-duct in Polypterus, and it is
from this longitudinal duct that the transverse tubules pass to the kidney.
That the arrangement in Polypterus is not secondary seems probable from the fact
that not only is it the simpler method of conveying the testis-products outwards, but
is, on the whole, closely similar to the arrangement in the female Polypterus, and we
can hardly suppose that in the primitive vertebrate the ova and spermatozoa found
exit by totally different means.
VII. Addominal Pores.
In both male and female, abdominal pores are present in Polypterus. ‘They have
been correctly described in the adult as fine canals opening to the exterior on either
side of the vent (Pl. L. figs. 7,8). In the young female 9 cm. in length there is a very
fine nucleated diaphragm cutting off the communication of the coelom with the exterior.
This is, however, extremely delicate (Pl. L. fig. 9, ad.p.).
In the young male 9 cm. in length the ccelom is completely shut off from the
exterior, the abdominal pores not being yet formed.
VIIL. The Anal Fin.
Traquair has already noted in Calamoichthys that the males have an enlarged anal fin.
In Polypterus, during the breeding-season at least, this difference is not merely one of
size but also of shape and form, as shown in PI. L., figs. 1 & 2.
The anal fin in the female is narrow and pointed, while in the male it is twice as
deep as in the female, and its surface is thrown into deep folds between the successive
THE ANATOMY OF POLYPTERUS. 331
fin-rays. ‘he muscle of the anal fin is greatly enlarged in the male, protruding as a
rounded mass into the celom. It is this mass which causes the ureters and genital
ducts to turn so abruptly ventralwards in the male (PI. L. figs. 2, 5, 6, an.f.m.).
I may mention that in an abnormal male specimen in which the anal fin was absent
this muscle was completely absent, and the excretory and genital ducts ran backwards
into an extension of the coelom, then forwards ventrally to open in the normal position
behind the vent.
This sexual character almost entirely disappears out of the breeding-season.
Leydig has suggested that there is internal fertilization in Polypterus from the fact
that in the ccelom of a female Polypterus he found masses of filaments which he took
for spermatozoa. That these filaments are not spermatozoa can at once be seen by
comparing the figure he has given with my figure of spermatozoa from the ripe testis
(PE Eee 13):
When the arrangement of the oviducts is considered, it seems extremely improbable
that the spermatozoa would find their way into a duct which opens into the urino-
genital sinus upon a papilla. It remains to be seen to what use the male Polypterus puts
this modified anal fin.
IX. The Vascular System.
The blood-supply to the external gill has been worked out by Hyrtl, while the main
roots of the arterial system were described by Joh. Miller. I would call attention,
however, to a few additional details. Having injected a male specimen with salt-
solution when killed, it was re-injected with a coloured gelatine in the laboratory.
The specimen was adult and had no external gill. The details of the blood-supply
to the external gill were made out on a young specimen in which it was possible to
inject the hyoidean artery with a coloured fluid. The figure of the arterial system was
made by a combination of these two dissections.
As Hyrtl has shown, the hyoidean artery arises at the anterior end of the ventral
aorta immediately in front of the first afferent branchial artery, and passes to the
base of the operculum, at the centre of which it meets the efferent hyoidean artery,
to run with the latter to the posterior edge of the operculum and thence to the
external gill.
The point I wish to call attention to is that the afferent and efferent arteries at
the extremity of the gill are continuous one with another, forming a drawn-out
loop. . From the afferent limb branches run to the pinne, at the extremity of
which they loop back to the main efferent limb; similar tertiary loops pass into the
pinnules.
At the root of the external gill there is a dorsal and a ventral muscle ; towards the
extremity of the gill these break up into numerous isolated bundles (PI. LIL. figs.
25, 26, 27). The whole arrangement is quite similar to that of an Amphibian or
Dipnoan external gill.
VOL. XV.—Part vil. No. 2.— April, 1901. 3A
332 MR. J. S. BUDGETT ON SOME POINTS IN
Hyrtl, from the arrangement of the main blood-supply to the external gill, argues
that this must be homologous with the pseudobranch of Actpenser, which has the
same structure as the succeeding gills.
The external gills of larval batrachians are borne upon the first two or three gill-
arches, there being to each of these arches an external epidermal gill and an internal,
probably endodermal, gill.
Kerr has shown that in larval Dipnoi which possess external gills there is to each
arch an internal, probably endodermal, gill and an external epidermal gill, both being
supplied by the same afferent artery.
It appears probable therefore that in the external gill of the hyoid arch in Polypterus
we have not the homologue of the internal endodermal pseudobranch of Acipenser,
but of the external epidermal gill of Dipnoi and Amphibia.
Moreover there is in Polypterus at the base of the operculum a stout branch from
the efferent artery (Pl. LII. fig. 25, hy.eff') which runs parallel with the afferent
artery. The presence of this branch is suggestive, as indicating the position of the
pseudobranch, corresponding to the pseudobranch of Acipenser, of which in Polypterus
there is no further trace.
Pollard states that he could find no trace of a connection between the last efferent
branchial artery and the dorsal aorta, the blood from the last hemibranch passing only
to the air-bladder. Part of the blood from this gill does, however, pass to a branch
joining the third efferent artery, which on the right side meets the coeliac artery
(6r.1V.eff.). The main part of the blood from the hemibranch of the IVth arch
passes to the air-bladder on either side and is returned from them by veins passing to
the hepatic veins, as shown by Joh. Miiller. The vein on the right side is of great
size, corresponding to the size of the right air-bladder, and posteriorly unites
with the caudal vein.
It is difficult to see how these air-bladder veins in Polypterus could get converted
into the pulmonary veins in Amphibia, It seems more probable that the great vein
of the right air-bladder corresponds to the anterior abdominal vein of Amphibia,
though it is notorious that veins frequently make secondary connections.
The subclavian artery of Polypterus passes outwards dorsal to the vagus and the
coeliac and pulmonary arteries, and curving ventralwards gives off, near the pericar-
dium, a branch on either side (text-fig. 7, d7.). The posterior branch, the brachial,
is a stout artery passing to the pectoral fin; the anterior branch runs dorsally
again parallel with the main subclavian, and divides to supply the muscles of the
shoulder-girdle (text-fig. 7, dr.'). After giving off these two branches, the sub-
clavian is continued as a small coronary artery along the sides of the pericardium,
at the anterior end of which it passes to the walls of the conus and ventricle (text-
MES (hy PS)
The blood from the conus and ventricle is returned to the ductus Cuvieri by a
THE ANATOMY OF POLYPTERUS. 333
thick-walled, deeply-pigmented, coronary vein, which runs free in the pericardial
chamber from the ventricle to the ductus Cuvieri.
From the dorsal side of the dorsal aorta just behind the junction with it of the
second efferent branchial artery there passes outwards on either side a small vertebral
artery, which curving dorsally round the spinal column appears to enter the spinal
canal.
The arterial system of Polypterus.
br., brachial ; 6r.', branch of subclavian to shoulder-girdle; Or., branch of subclavian which becomes the
coronary artery; br. Z, IT, II. IV.aff., branchial afferent I., I1., III., 1V.; br. 2. 17. 11, IV. ef-s
branchial efferent I., II., IIL, IV.; ca., carotid; cel., celiac; ev.g., external gill; hy.aff., hyoidean
afferent; hy.eff., hyoidean efferent; hy.eff.', hyoidean efferent branch; p.a., pulmonary; s.cl.
subelavian ; sp., spermatic,
Pollard has pointed out the extremely primitive condition of the dorsal arterial
system in Polypterus. The single efferent vessel from each gill-arch uniting with a
median dorsal aorta, resembles most the condition found in Chlamydoselachus and the
embryos of other Selachians.
3A 2
34 MR. J. S. BUDGETT ON SOME POINTS IN
(Sh)
X. The Cranium.
In this paper I have no intention of dealing with the anatomy of the head and
craninm of Polypterus, except that I wish to mention that the peculiar differences
in the condition of the sphenoid bone which Bridge has described in two specimens
which he examined, and called specimen A and specimen B, have been found to be
specific in the two species from the Gambia. The sphenoid bone of Bridge’s specimen
A, resembling that figured by Traquair, in which the lateral wings of the sphenoid did
not curve inwards and fuse anteriorly in the frontal region, resembles exactly that of
Polypterus senegalus, Cuy. The sphenoid (or “sphenethmoid”) bone of Bridge’s
specimen B, which he suspected of being Polypterus lapradei, Steind., resembles
exactly that species}.
XI. Conclusion.
If in this paper I have occasionally reiterated facts which have long been known,
I trust such has not been done without adding at least some details of interest. In
any case I hope I have always acknowledged the authority.
I feel that no apology is needed for having entered into minute details in some cases,
for an accurate knowledge of the anatomy of a creature of such surpassing interest as
the Polypterus seems te me in every way desirable.
In obtaining my material for this investigation I have been aided by a grant of £50
from the Balfour Fund of the University of Cambridge. The work has been done
in Mr. Adam Sedgwick’s laboratory. ‘To Mr. Sedgwick, Mr. Graham Kerr, and
Prof. Howes my thanks are due for much help and advice in my work.
XII. Bibliography.
Ayers: “Morphology of the Carotids,” in Bull. Mus. Comp. Zool. Harvard, vol. xvii., 1889.
Batrour: ‘Comp. Embryology,’ vol. 11. p. 606.
Batrour and Parker: “The Structure and Development of Lepidosteus.” Phil. Trans. Roy. Soe.
Lond. part u., 1882.
BrrpGe, T. W.: “ Some points in the Cranial Anatomy of Polypterus,’ in Birm. Phil. Soe. vol. vi.
part 1., 1886.
Garman: “ Chlamydoselachus anguineus,” in Bull. Mus. Comp. Zool. Harvard, vol. xii., 1885.
Howes: “On some Hermaphrodite Genitalia of the Codfish.” Journ. Linn. Soe., Zool. vol. xxiii.
Huxiry: “ Classif. of Devonian Fishes,’ in Mem. Geol. Survey, dec. 10, 1861.
“ On the Oviducts of Osmerus.” Proc. Zool. Soc. Lond. 1883, p. 182.
Hyrtt: “ Ueber d. Pori abdominales, d. Kiemen-Arterien, u.d. Glandula thyroidea d. Ganoiden.”
Sitz. d. Wiener Akad. Bd. viii., 1852.
* Since this paper was read, I have examined a young specimen of P. lapradei 12 cm. in length, and find
that at this age the sphenoid has not yet developed its specific character, but resembles that of the adult
P. senegulus.
THE ANATOMY OF POLYPTERUS. 335
Hyrti: “ Ueber den Zusammenhang der Geschlechts- und Harnwerkzeuge bei den Ganoiden.”
Denksch. der k. Akad. der Wiss., Bd. viii., Math.-naturw. Classe, Wien, 1854.
“Ueber die Blutgefasse der ausseren Kiemendeckel-Kieme von Polypterus lapradei
Steind.” Sitzungsb. d. k. Akad. d. Wiss. Wien, Math.-naturw. Cl., lx. Bd. i. Abth. 1869.
)
o 66 itrh rik 1 y 4 a . :
JuncErsEen: “ Beitriige zur Kenntniss der Geschlechts-Organe der Knochenfische.” Arbeit. zool.-
zoot. Inst. Wiirzburg, Bd. ix., 1890.
Kerr, J. Granam: “ The External Features in the Development of Lepidosiren paradowa, Fitz.”
Phil. Trans. Roy. Soe. vol. 192 B, 1900.
Lrsepinsxy: “ Ueber die Embryonalniere von Calamoichthys calubaricus.” Arch. mikr. Anat.
Bad. xliv. p. 216.
Lrypie: “ Histologische Bemerkungen tiber den Polypterus bichir.” Zeitschr. wiss. Zool. 1854.
Miter, Jonannes : ‘ Ueber den Bau und die Grenzen der Ganoiden.’ Berlin, 1846.
Potrarp, H. B.: “On the Anatomy and Phylogenetic Position of Polypterus”’ Zool. Jabrb.
Anat. u. Ont. Bd. v., 1892.
Semon, R.: “ Zusammenhang der Harn- und Geschlechtsorgane bei den Ganoiden.” Morph.
Jahrb. Bd. xvii.
STemnpacuner: “ Polypterus aus dem Senegal.” Sitzb, d. k. Akad. d. Wiss. Wien, Math.-
naturw. Classe, Bd. lx., 1869.
Traquair: “ Cranial Osteology of Polypterus.” Journ. Anat. & Phys. vol. v., 1871.
Traquair: “On the Anatomy of Calamoichthys.’ Proc. Roy. Soc. Edinburgh, vol. v., 1866.
Weser, Max: ‘Die abdominal Poren der Salmoniden, nebst Bemerkungen iiber die Geschlechts-
[S)
organe der Fische.” Morph. Jahrb. Bd. xii.
XIII. EXPLANATION OF THE PLATES.
PLATE L.
. General side view of the posterior part of the viscera in a female Polypterus
lapradet (p. 825), showing the course of the oviducts and the anal fin of a
female in the breeding-season.
General side view of the posterior part of the viscera in a male Polypterus
lapradei (p. 324) in which the ureters have been injected blue, showing the
testis-ridge and duct running along the ventral external wall of the ureter.
The anal fin of a male in the breeding-season is also shown with the
anal-fin muscle projecting into the posterior part of the ccelom.
. A median ventral view of the testis, the kidney, and their ducts of the same
male specimen dissected out (p. 524). The ureter, as in fig. 2, is injected
blue. Figs. 2 & 3 together give a complete view of the urinogenital organs
in the male.
. Ventral view of the urinogenital sinus in the female (pp. 325, 350), showing
to) dD
the openings of the oviducts upon papille, the septum of the urinogenital
sinus, the narrowness of the anal fin in the female, and the corresponding
336
Fig.
Fig.
Or
10.
ig. 15.
16.
MR. J. 8S. BUDGETT ON SOME POINTS IN
smallness in size of the anal-fin muscle. ‘The rectum has been partially cut
away and the ventral wall of the urinogenital sinus removed.
. A similar view of the urinogenital sinus in the male (pp. 324, 331), showing
the genital aperture, the narrow neck of the urinogenital sinus, the great
width of the base of the anal fin and the great development of its muscle.
. View from the right side of the urinogenital sinus in a large male (p. 324),
the anal fin and its muscle, showing the course of the vas deferens; these
organs, together with the rectum, having been dissected from the body.
. View from below of the anal region in a female (pp. 528, 330), showing the
slit-like urinogenital aperture and the abdominal pores.
29
. A similar view ina male (pp. 328, 330), showing the wide urinogenital aperture.
. A vertical section of a very young female Polypterus senegalus, 9 cm. in length
(p. 330), passing through the abdominal pores, showing a thin nucleated
diaphragm closing the abdominal pores, at the same time showing an
exudation of the ccelomic fluid. ‘The caudal vein and abdominal vein are
cut through just anterior to their junction.
PLATE LI.
A vertical section through the testis and kidney of an adult male (pp. 324,
327,328). The ureter is injected blue. The testis-tubules are shown opening
into the testis-duct, and a kidney-ductule opening into the ureter. The
magnification is 6 diameters.
. A similar section behind the testis (pp. 824, 327, 328), showing the relation
of the testis-ridge and duct to the ureter in an adult male.
. A similar section in the region of the urinogenital sinus (pp. 324, 328), showing
the duct of the testis opening upon a papilla into the urinogenital sinus.
. Spermatozoa compared with a blood-corpuscle (pp. 324, 331). Zeiss ocular 3,
objective E.
. An enlarged drawing of the right half of fig. 12 (pp. 324, 825), showing the
stout columnar epithelia] lining of the vas deferens.
Figs. 14 to 24 are all magnified about 70 diameters.
An enlarged drawing of the testis-ridge of fig. 11 (p. 324), showing the similar
appearance of the ductules of the ridge and the main testis-duct. They
are lined by a large-celled epithelium and embedded in connective-tissue
with a few small blood-vessels.
A vertical section through the kidney and testis of a young male P. senegalus,
13 cm. in length (pp. 324, 327, 828), showing the minute structure of the
testis. The tubules are lined by large glandular cells, and are embedded
in lymphoid tissue similar in appearance to the lymphoid tissue of the
relies
5 se
> Al,
5 all
ig. 22.
ie. 26,
THE ANATOMY OF POLYPTERUS, 337
kidney. ‘The section shows also the similar appearance of the testis-duct and
ureter. The uriniferous tubules of the kidney are embedded in a mass of
lymphoid tissue.
A similar section passing behind the testis (p. 325). When compared with
fig. 15 the figure shows the relatively great development of the main duct
of the testis-ridge, the small development of the ductules of the ridge in
number and size, and the small relative size of the ureter.
A similar section in the region of the urinogenital sinus (p. 525), showing the
opening of the vas deferens into the latter and its lining of large columnar
cells. The large relative size of the vas deferens is seen on comparing with
fig. 14.
PLATE LIT.
. A vertical section through the testis and ureters of a very young male
P. senegalus, 9 cm. in length (p. 325), showing the precocious development of
the testis-duct, the tubules of the testis not being yet formed.
A similar section through the posterior termination of the testis-ducts (p. 325),
showing that they are only just acquiring their connection with the urino-
genital sinus.
A vertical section through the developing ovary and ureter of a very young
female P. senegalus, 9 cm. in length (pp. 325, 326, 327), showing the ova
developing from the external wall of the ovary, the ovary divided into a
number of loculi, and the artery and vein in the median wall.
A similar section behind the ovary passing through the peritoneal opening of
the oviduct (pp. 325, 326).
. A similar section behind the peritoneal opening of the oviduct (pp. 325, 326),
showing the oviduct lying between the mesoarium or female genital ridge
and the ureter. ‘To be compared with fig. 17.
. A-similar section through the posterior termination of the oviduct (p. 526),
showing that they have not yet acquired their connection with the urino-
genital sinus. To be compared with fig. 20.
. Left operculum bearing external gill of Polypterus lapradei, 30 cm. in length
(pp. 831, 332), showing the blood-supply to the gill and the efferent artery
giving off a branch which runs parallel with the afferent artery. The
muscles of the operculum and the gill are also shown.
An enlarged drawing of the tip of the external gill (pp. 331, 352), showing
its minute structure and the continuation of the afferent into the efferent
artery at the end of the gill.
. A pinnule of the external gill highly magnified (p. 331).
338 ON SOME POINTS IN THE ANATOMY OF POLYPTERUS.
XIV. EXxXpLaANATION OF THE LETTERING OF THE FIGURES.
abp. Abdominal pore. ovd.ep. Epithelium of oviduct.
ab.p.c. Abdominal pore-canal. p.ep. Peritoneal epithelium.
pl. Pinuule.
aff.art. Afferent artery. pn. Pinna.
an.f. Anal fin.
an.f.m. Anal-fin muscle.
ab.v. Abdominal vein.
r. Rectum.
sp. Spermatozoa.
ao. Aorta. sp.art. Spermatic artery.
6.c. Blood-corpuscle. sp.ep. Spermatic epithelium.
b.v. Blood-vessel. sp.v. Spermatic vein.
b.w.m. Body-wall muscle. sw.bl. Swim-bladder.
cv. Cardinal vein. t. Testis.
cd.v. Caudal vein. t.d. Testis-duct.
con.tis. Connective-tissue. t.d.ep. Epithelium of duct of testis.
d.m. Dorsal muscle. t.d.sh. Testis-duct sheath.
e.ov.w. External wall of ovary. tr. Testis-ridge.
eff.art. Efferent artery. t.r.tbs. Testis-ridge tubules.
fb. Fat-body. t.tbs. Testis-tubules.
gap. Genital aperture. u. Ureter.
g-p. Genital papilla.
m.ov.w. Median wall of ovary.
k, Kidney.
kv. Kidney-vein.
u.ep. Epithelium of ureter.
u.g-.ap. Urinogenital aperture.
u.g.s. Urinogenital sinus.
u.s. Urinary sinus.
lt. Lymphoid tissue. u.sh. Sheath of ureter.
m.cps. Malpighian corpuscles. u.ths
mes.o. Mesoarium.
mes.t. Mesorchium.
. Uriniferous tubules.
v.d. Vas deferens.
v.d.ep. Epithelium of vas deferens.
n.o. Nucleus of ovum. v.m. Ventral muscle.
o. Ovum. w.o.u. Point where oviduct ends in wall of
o.at. Attachment of ovum. mmctes
op.m. Opercular muscle. w.v.u. Point where vas deferens ends in wall
ov. Ovary. of ureter.
ov.d. Oviduct.
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39
LIST OF THE PAPERS CONTAINED IN VOL. XV.
Awnprews, C. W., B.Sc., F.Z.8., Assistant in the
British Museum (Natural History).
On the Extinct Birds of Patagonia.—I.
The Skull and Skeleton of Phororhacos
inflatus Ameghino
Bounencer, G. A., F.R.S., F.Z.8.
Report on the Collection of Fishes made
by Mr. J. E. S. Moore in Lake Tangan-
yika during his Expedition, 1895-96.
With an Appendix by J. E. 8. Moors,
A.R.C.S.
Second Contribution to the Ichthyology of
Lake Tanganyika.—On the Fishes ob-
tained by the Congo Free State Expedi-
tion under ‘Lieut. Lemaire in 1898 ....
Brapy, G. Srewarpson, M.D., D.Sc., LL.D.,
F.R.S., C.M.Z.8.
On the Marine Copepoda of New Zealand .
Boneerr, J. S., M.A., F.Z.8., Trinity College,
Cambridge.
On some Points in the Anatomy of Poly-
pterus
Fawcerr, Lieut.-Colonel J. Matcorm.
Notes on the Transformations of some
South-African Lepidoptera ..........
VOL xv.—Part vil. No. 3.—Apri/, 1901.
Harmer, Srpnny F., Sc.D., F.R.S., Superinten-
dent of the University Museum of
Zoology, Cambridge.
On a Specimen of Cervus belgrandi Lart.
(C. verticornis Dawk.) from the Forest-
Bed of HastyAneliagyancrierercie tie
Pycrart, W. P.
A Monograph of the Genus Casuarius.
By The Hon. Watrer Roruscurc,
Ph.D., F.Z.8.
the Morphology and Phylogeny of the
With a Dissertation on
Paleognathe (Ratite and Crypturi)
and Neognathe (Carinate). By W. P.
Pycrarr
Roruscuitp, Hon. Watrer, Ph.D., F.Z.8..
A Monograph of the Genus Casuarius.
By The Hon. Watrer RornscHrcp,
Ph.D., F.ZS.
the Morphology and Phylogeny of the
With a Dissertation on
Paleognathe (Ratite and Crypturi)
and Neognathe (Carinate). By W. P.
PycraFrr
Page
97
109
INDEX OF SPECIES,
Abdimia, 246.
Acartia denticornis, 34.
ensifera, 33, 34, 49.
Acherontia atropos, 307.
Aerea acara, 293.
buxtoni, 295, 319.
encedon, 294, 319.
lycia, 294.
petraa, 294, 319.
rahira, 294, 319.
Actias mimose, 302, 320.
Aipyornis, 171, 172, 173, 174, 175, 176, 179, 180,
183, 195, 221, 232, 235, 237, 260, 2638, 274,
76, 279, 281.
Africa, on the Transformations of some Lepidoptera,
of, 291-322. |
_ Agrotis segetis, BG, Gell,
Alestes imber?, 4.
macrolepidotus, 5, 25, 95.
Pp ;
macrophthalmus, 5, 25, 95.
rueppellir, 5, 6.
Amymone claus, 40.
Andrews, C. W. On the Extinct Birds of Patagonia.
—I. The Skull and Skeleton of Phororhacos |
inflatus, 55-86.
Anomalopteryx, 179, 221.
Anoplopterus platychir, +, 5, 24.
Antherwa (Nudaurelia) cytherea, 303.
—— (——-) menippe, 303.
( ) tyrrhea, 308.
Apteryx, 76, 150, 154, 159-290.
australis, 159, 163, 220, 225, 232, 234,
251, 254, 270, 277, 285.
mantel, 155, 161, 162, 163, 169,
220, 223, 232, 234, 270, 273, 277, 286.
haastu, 163, 220, 232, 234, 244, 254, 270,
277.
244,
196,
EIC., IN VOL. XV.
Apterya mantelli, 254, 286, 289, 290.
owent, 168, 215, 220, 223, 225, 232, 234, 251,
254, 270, 277, 287.
Aptornis, 69, 71, 72.
Archeopteryx, 153, 260.
Arius commersonii, 18.
Artotrogus brevicaudatus, 48, 53, 54.
ovatus, 48.
| Auchenaspis biscutata, 5, 24, 95.
Bagrus meridionalis, 4.
Barbus sp., 6.
pagenstechert, 5.
trimaculatus, 4, 5.
Barilius quentheri, 4.
Bathybates, gen. noy., 15.
ferow, 4, 15, 27, 30, 89.
Boarmia acaciaria, 318, 322.
Boulenger, G. A. Report on the Collection of Fishes
made by Mr. J. E. 8. Moore in Lake Tan-
Expedition, 1895-96,
eanyika during his
1-30.
Second Contribution to the Ichthyology of
Lake Tanganyika.—On the Fishes obtained
by the Congo Free State Expedition under
Lieut. Lemaire in 1898, 87-96.
Brady, G. Stewardson. On the Marine Copepoda of
New Zealand, 31-54.
Bubo maximus, 287.
Budgett, J. S. On some Points in the Anatomy of
Polypterus, 323-838.
Bunea caffraria, 303, 320.
Calamoichthys, 327, 330, 335.
calabaricus, 334.
Calanus arewicornis, 32.
jfimmarchicus, 32.
Caligus longicaudatus, 49, 53.
Calodromas elegans, 154, 162, v41, 250, 288.
33B2
342
Canthocamptus furcatus, 45.
Cariama, 60, 65, 68, 71, 72, 76, 77, 79, 80, 82, 83,
84, 85.
Carinatze, on the Morphology and Phylogeny of the,
149-290.
Oasuarius, a Monograph of the Genus, 109-290.
sp., 117, 118, 130, 152, 139.
altyugus, 120.
—— uaruensis, 130.
aurantiacus, 137.
—— australis, 117, 118, 123, 124, 125, 128, 280,
290.
becearti, 114, 116, 117, 118.
— bennetti, 111, 112, 128, 188, 139, 140, 145,
146, 147, 148, 188, 227, 228, 251, 276, 285.
maculatus, 112, 148.
bicarunculatus, 111, 112, 116, 117, 122,
130) WSik Qi 25h 283285.
casuartus, 111, 112, 113, 114, 121, 122, 124,
180, 181, 134, 189, 140, 147, 151, 152, 209,
227, 228, 257, 276, 283, 285, 289.
australis, 110, 112, 122, 123,
209, 283.
beccan lil 2 UG SIA ile SONS i
251, 257, 276, 283.
intensus, 112, 121, 283.
salvadorit, 112, 120, 121, 122,
187, 257, 2838.
selatenv, V2> Ws, 119; 127, 124) 152)
155, 187, 257.
violicollis, 112, 117, 122, 153, 257, 283.
— edwardsi, 141, 142.
—— emeu, 113, 130.
galeatus, 118, 117, 130, 1381, 133, 146.
javanensis, 114.
johnson, 123, 124, 125.
kaupi, 130, 132, 139, 143,
laglaizei, 136.
—— lorie, 111, 112, 142, 148, 144, 155, 169,
285.
—— nowe-hollandice, 280.
oceipitalis, 185, 136.
orientalis, 114.
-—— papuanus, 111, 112, 135, 136, 139, 140, 141,
142, 145, 147, 284, 285.
edwardsi, 112, 141, 145, 284.
philipi, 111, 112, 137, 138, 284,
129,
129, 188,
152, 153,
INDEX.
Casuarius picticollis, 111, 112, 142, 143, 144, 145,
251, 276, 284, 285.
heckt, 112, 114, 148, 284.
regalis, 124,
salvadorii, 120, 121, 215.
sclatert, 117, 118, 288.
tricarunculatus, 120, 121.
unappendiculatus, 111, 112, 130, 1382, 138,
134, 1385, 136, 137, 1388, 1389, 140, 153, 218,
228, 238, 248, 251, 252, 256, 257, 275, 284,
285, 287, 288, 289.
aurantiacus, 112, 136, 284.
occipitalis, 112, 135, 186, 137, 284.
laglaizer, 136.
rufotinctus, 112, 137.
violicollis, 258.
westermanni, 135, 189, 140, 141, 144.
Catarrhactes chrysocome, 255, *
Centronma thomsoni, 47, 54.
Centropages discaudatus, 35, 50.
pectinatus, 36, 50.
typrcus, 30.
aucklandicus, 35.
Cephonodes hylas, 312, 321.
Cervus belgrandi, on a specimen of, from the Forest-
Bed of East Anglia, 97-108.
—— browni, 104.
carnutorun, LOG.
dama, 99, 100, 104, 105, 107, 108.
curyceros germane, 106.
haiberniw, 106.
giganteus, 98, 100, 101, 103, 104, 105, 107,
108.
belgrandi, 106.
carnutorum, 106.
italice, 106, 107.
, 106.
—— typicus, 106.
TU
megaceros ruffiz, 107.
verticornis, 97-108.
Cherocampa balsamine, 309, 321,
capensis, 308, 320.
idvicus, 310, 321.
osiris, 309, 321.
Charaxes candiope, 296, 319.
Chauna, 250.
Chlamydoselachus, 333.
INDEX. 543
Chlamydoselachus anguineus, 334.
Chromis diagramma, 19.
paterfamilias, 18.
Chrysichthys cranchit, 95.
Chaunga, 65, 84, 85.
Ciconia, 71, 72, 80, 83, 246.
Citharinus geoffroy?, 6
Clarias sp., 5.
anguillarvis, 5, 24.
liocephalus, 5, 24, 30.
Clausocalanus arcuicornis,
fureatus, 32, 33, 49.
Oleta serrata, 40.
Congo Free State Expedition, on the Fishes obtained
in Lake Tanganyika by the, 87-96.
Copepoda, on the New-Zealandian Marine, 31-54.
Oorematodus shiranus, 4.
215
32, 33.
Coryceus robusta, 4:6.
Crypturi, on the Morphology and Phylogeny of the,
149-290.
Ctenochromis nuchisquamulatus, 17.
Cyclops ewarti, 38, 51.
Dactylopus cinctus, 41.
hanseni, 41, 52.
strom, 41.
tishoides, 41, 53.
Danuis chrysippus, 293.
aleippus, 293.
dorippus, 293.
Daphnis nerii, 310.
Dasychira georgiana, 314, 322.
horsfieldvi, 314.
Diaphorapteryx, 71, 72.
Dinornis, 168, 171, 173, ae 180, 181, 182, 183,
186, 193, 194, 210, 214, 221, 260, 262
267.
maximus, 216.
— robustus, 280.
(Pachyornis) elephantopus, 138, 139.
Diomedea, 62, 64, 72, 79, 80, 82, 84.
Distichodus rudolphi, 6
5 263, |
Drepanopus furcatus, 32, 33.
pectinatus, 33.
Dromeus, 76, 118, 131, 188, 153-288.
ater, 172, 173, 175, 179, 217, 218, 222, 22
7,
234, 235, 267, 270, 275, 278, 287.
— irroratus, 179, 267, 270, 271, 287.
Dromeus novee-hollandic, 146, 153, 155, 175, 179,
217, 218, 929 297, 934. 935, 251, 270, 275;
278, 280, 285, 286, 287, 288, 289.
Dulichia fasciata, 314, 321.
Earias insulana, 316, 322.
Echidna, 281.
Ectinosoma australe, 39, 51.
Eetodus, gen. nov., 21.
descampst, 4, 90, 92, 96.
longianalis, 91, 92, 96.
melanogenys, 4, 21, 91, 92, 96.
Emeus, 183, 186, 190, 191, 192, 193, 262, 286.
DAenbone pinguis, 4.
Entomolepis, gen. nov., 48.
ovalis, 48, 54.
Eretmodus, gen. noy., 16.
cyanostictus, +, 16, 30.
Eusemia butleri, 317, 320.
Euterpe ucutifrons, 39.
gracilis, 39.
Extinet Birds of Patagonia, Andrews on the, 55-86.
Falco tinnunculus, 260.
Fawcett, Lt.-Col. J. Malcolm. Notes on the Trans-
formations of some South-African Lepidoptera,
291-322.
Flavia, gen. nov., +1.
crassicoriis, +2, 51.
Francolinus, 77.
Fulica,
Fundulus teniopygus, 5.
75, 76.
Gasterosteus spinachia, 18.
Genyornis, 267.
newtoni, 282.
Glottula pancratii, 317, 321.
Glyptodon, 85.
Fonometa postica, 313, 321.
Grammatotria, gen. nov., 90.
lemairri, 90, 96.
Grus, 68, 71, 72, 79, 80, 83.
12,
Gynanisa maia, 304, 320.
Haplochilus johnstoni, +.
-— tanganicanus, 5, 19, 25, 30, 96.
On a Specimen of Cervus bel-
grandi Lart. (C. verticornis Dawk.) from the
Forest-Bed of East Anglia, 97-108.
Harpacticus acutifrons, 39.
chelifer, 44.
Harmer, Sidney F.
544 INDEX.
Harpacticus glaber, 44, 58. Mastacembelus flavomarginatus, 23.
Hesperornis, 76, 150, 226, 260. —— liberiensis, 23.
Hippateetryo, 109. —— loennbergii, 23.
indicus, 114. —— marchit, 23.
Hydrocyon forskalii, 5, 25. —— marmoratus, 22, 23.
Hypsa aphidas, 315, 320. ——. moorit, 5, 22, 23, 30.
subretracta, 315. —— niger, 23.
Ichthyology of Lake Tanganyika, 1-30, 87-96. | —— nigromarginatus, 23.
Tchthyornis, 260. —— ophidium, 5, 23.
Idya furcata, 45. —— shiranus, 4, 23.
Julidochromis, gen. nov., 11. tanganice, 23.
ornatus, 4, 12, 30. Megalapteryx, 202.
Junonia cebrene, 295, 319. Moore, J. E. S., on the Fishes collected in Lake
clelia, 296, 319.
Labeo sp., 5, 25.
Tanganyika in 1895-96 by, 1-30.
Mormyrops anguilloides, 6.
forskalia, 5.
zambanenje, 4, 6.
—— mesops, 4. | Mormurus catostoma, 4.
rueppellia, 5. discorhynchus, 4, 6.
Labidocera cervi, 37, 50. longibarbis, 5.
nerti, 38.
oxyrhynchus, 5.
Lamprologus compressiceps, 4, 7, 10, 29. Neognathe, on the Morphology and Phylogeny of
congoensis, 10, 88. the, 149-290.
elongatus, 4, 9, 10, 29. | New Zealand, on the Marine Copepoda of, 31-54.
fasciatus, 4, 7, 10, 29. Nudaurelia wahlbergi, 303, 320.
furcifer, 4, 9, 10, 29. | Nyeticorax, 245.
lemairei, 88, 96. | Ocydromus, 72.
—— modestus, 4, 8, 10, 29. | Otthona spinifrons, 38.
moorii, 4, 8, 10, 29, 88. | Opisthocomus, 153, 239, 248.
Laophonte meinerti, 40, 51. | cristatus, 241, 282.
serrata, 40. Ornithorhynchus, 257, 281.
Lates microlepis, 4, 6, 29, 88. | Osmerus, 334.
niloticus, 7. | Pachyornis, 216.
Lemaire, Lieut., on the Fishes obtained in Lake | Palaognathe, on the Morphology and Phylogeny of
Tanganyika by the Congo Free State Expedi- the, 149-290.
tion under, 87-96. Panochthus, 85.
Lepidoptera, on the Transformations of some South- | Papilio brasidas, 298, 319.
African, 291-322. | —— demodocus, 300, 320.
Lepidosiren paradoawa, 334. —— leonidas, 299.
Lepidosteus, 328, 330, 334. —— morania, 299, 320.
Leptoptilus, 239. | —— nireus, 301.
Lophostethus dumolinii, 307, 321. —— , var. lywus, 320.
Tudia smilax, 305, 321. | —— policenes, 298, 319.
Macrosila solani, 311. Paracalanus parvus, 32, 49.
Malapterurus electricus, 5, 25. Paratilapia afra, 4.
Mastacembelus sp., 5.
cavifrons, 5.
dewindti, 88, 96.
dimidiata, 4.
congicus, 23.
eryptacanthus, 23. |
INDEX.
Paratilapia furcifer, 4, 14, 30.
intermedia, 4, 13.
leptosoma, 4, 14, 30.
——- livingstonii, 4.
—— longiceps, 4, 15.
longirostris, 5.
macrops, +, 138, 30.
modesta, 4.
—— pfefferi, 4, 12, 30.
retrodens, 5.
—— robusta, 4.
ventralis, 4, 13, 15, 30, 89.
Patagonia, on the Extinct Birds of,
55-86.
Paurocope, gen. nov., 46.
robusta, 46, 53.
Pelotrophus microcephalus, 4.
microlepis, +.
Peltidium interruptum, 44.
novee-zealandiw, 44, 50, 53.
Perissodus, gen. nov., 20.
microlepis, 5, 21, 30.
Petrochromis, gen. nov., 20.
polyodon, 5, 20, 30.
Phasianus, 77.
Phissama flava, 316, 321.
screabile, 316, 321.
Pheenicopterus, 246.
Phororhacos inflatus, on the Skull and Skeleton of,
55-86.
Phroso, gen. noy., 42.
gracilis, 42, 52.
Pieris hellica, 297, 319.
severina, 297, 319.
Planema esebria, 295.
Plecodus, gen. nov., 22.
paradowus, 5, 22.
Podargus, 253.
Polydesma glaucinans, 318, 322.
Polypterus, on some Points in the Anatomy of, 323—
338.
bichir, 5, 26, 28, 334.
laprade, 324-338.
senegalus, 6, 324-338.
Porcellidium fuluum, 46.
Protoparce mauritii, 311, 321.
Protopterus avnectens, 5.
Pseudaphelia apollinaris, 305, 320.
Pycraft, W. P.
vo
He
Or
Psophia, 71, 72, 79, 80, 83, 84.
On the Morphology and Phylogeny
of the Palseognaths (Ratitee and Crypturi) and
Neognathe (Carinate), 149-290.
Pyrameis cardui, 296.
Ratite, on the Morphology
149-290.
Rhea, 109, 150-289.
and Phylogeny of the,
americana, 154, 155, 1é :
245, 249, 250, 251, 253, 256, 267, 270,
219, 2, 279, 280) 285; 286; 287, 288;
289.
darwim, 155, 153, 245, 246, 267, 270,
279.
macrorhyncha, 158, 206, 217, 221, 245,
249, 251, 270, 275, 277.
| Rhynchotus rufescens, 252.
Rothschild, Hon. Walter. A Monograph of the
Genus Casuarius, 109-148.
Scutellidium plumosum, 45, 53.
tishoides, 45.
Serpentarius, 59, 60, 65, 77, 79, 83.
Simochromis, gen. noy., 19.
diagranma, 5, 19.
Spilosoma puella, 315, 321.
Struthio, 76, 150, 159-289.
australis, 269, 275.
camelus, 159, 215, 251, 269, 275, 289.
casuarius, 113, 116.
meridionalis, 206, 261, 269,
molybdophanes, 223, 269, 27
7).
285.
2
5;
Sula bassana, 255.
Synodontis afrofischeri, 5.
multipunctatus, 5, 24, 30, 95.
—— schal, 6.
smithii, 6.
zambesensis, +.
Teniopyga sylvina, 318, 321.
Tanganyika (Lake), on the Fishes collected by
Mr. J. E. S. Moore in, 1-30.
, on the Fishes obtained by the Congo Free
State Expedition in, 87-96.
Telmatochromis, gen. noy., 10.
temporalis, 4, 11, 30, 88.
vittatus, 4, 10, 29.
Temora longicornis, 35.
tenuicauda, 34, 50.
346
Thalestris australis, +3, 52.
ciliata, 43, 52.
forficula, 43.
Tilapia aurata, 4.
burtoni, 5.
— callipterus, 4.
—— dardennii, 93, 96.
—— desfontainesi, 94.
grandoculis, 94, 96.
johnstoni, 4.
kirkti, 4.
— luteristriga, 4.
— lethrinus, 4.
—— mucrolepis, 94, 96.
—— mossambica, 4.
——— milotrca, 5, 6, 18.
—— nuchisquamulata, 5.
—— obliquidens, 5.
— rendalli, 4.
labiata, 5, 17, 30, 95.
INDEX.
Tilapia rubropunctata, 93, 96.
—— sauvagii, 5.
——- shirana, 4.
——- simonis, 18.
—— squamipinnis, 4.
—— subocularis, 4.
—— tanganice, 5.
—— tetrastagma, +.
—— trisirami, 6.
—— williamsi, 4.
Tisbe furcata, 45.
Trematocara, gen. nov., 89.
marginatum, 89, 96.
Tropheus, gen. nov., 17.
moorit, 5, 18, 30.
Urota sinope, 305, 320.
Xenorhynchus, 246.
Xenotilapia, gen. nov., 92.
sima, 92, 96.
Zygena (Anteris) ampla, 318, 322.
END OF VOLUME XV.
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CONTENTS.
VILL. On some Points in the Anatomy of Polypterus. By J.S. Bupexntr, M.A., P.Z.8.,
Trinity College, Cambridge. (Plates L.--LIl.). . . . . - . « page 323
List of the Papers contained in Vol. XV. . . . . . . - . +. . . 339
Index ofSpecies zen Vol MVe eo
Titlepage and Contents to Vol. XV.
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