Alex. Agassiz.

l^ibrarn of tfj£ glusium

OF

COMPARATIVE ZOOLOGY,

AT HARVARD COLLEGE, CAMBRIDGE, MASS. JFaunSeS bg ptitiate sufastrtptton, in 1861.

Deposited by ALEX. AGASSIZ.

^0. Olf I Cl_

THE

TEANSACTIONS

OF

THE LINNEAN SOCIETY

OP

LONDON.

SECOND SERIES— VOLUME I. ZOOLOGY.

LONDON:

PEIHTED BY TATLOE AlfD FEANCIS, EED LION COUET, FLEET STEEET:

SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE ; AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER-ROW.

M.DCCC.LXXIS.

CONTENTS.

PART I.— Septembee, 1875.

I. On the Morphology of the Shell in the Woodpeckers (Picidse) and Wrynecks (Yungidse),

By Prof. W. K. Parker, F.B.S. (Communicated 6y H. T. Stainton, Usq., Sec. Linn. Soc.) (Plates I.-V.) page 1

II. On some Atlantic Crustacea from the ' Challenger ' Expedition. By Br. R. v.

Willemoes-Stjhm, Naturalist to the Expedition. (Communicated by Professor "Wyvlle Thomson, F.B.S. , F.B.S.) (Plates VL-XIII.) 23

III. On the Structure and Systematic Bosition of Stephanoscyphus mirabilis, the Type of

a new Order o/Hydrozoa. By Professor Allman, M.B., BL.D., F.B.S., 8fc., Pres. Linn. Soc. (Plate XIV.) 61

PART II.— December, 1875.

IV. On the external Anatom,y of Tanais vittatus, occurring with Limnoria and Chelura

terebrans in excavated Bier-wood. By John Denis Macdonald, M.B., F.B.S., Staff-Surgeon B.N., Assistant Brofessor of Naval Hygiene, Netley Medical School. {Communicated by W. T. Thiselton Dyer, M.A., F.B.S.) (Plate XV.) . . 67

V. On Valencinia Armandi, a new Nemertean. By W. C. M'Intosh, M.B., F.B.S. E.,

F.B.S. (Plate XVI.) 73

L iy ]

PART III.— July, 1876.

VI. On thirty-one Species of Marine Planarians, collected partly by the late Dr. Kelaart,

F.L.S., at Trincomalee, and partly by Dr. Collingwood, F.L.S., in the Eastern Seas. Dy Dr. Cuthbert Collingwood. (Plates XVII.-XIX.) . , page 83

VII. On the Structure and Development of the Bird's Shull. By "W. K. Parker,

F.B.S., F.L.S., ^G. (Part II.) (Plates XX.-XXVII.) 99

PART IV.— January, 1877.

VIII. A Monograph of the Lepidopterous Genus Castnia and some allied Groups. By Prof. J. O. Westwood, M.A., F.L.S., 8fc. (Plates XXVIII.-XXXIIL). . 155

IX. On a new Genus q/" Trematoda, and some neio or little-knoion Parasitic Hh^udinei.

By John Denis Macdonald, M.D., F.B.S., Deputy Inspector- General B.N., Professor of Naval Sygiene Army Medical School. [Communicated by G. E. DoBsoN, M.A., M.B., F.D.S.) (Plate XXXIV.) 209

X. On the Osteology and Ptei^ylosis of the Spoon-billed Sandpiper (Eurynorliynchiis

pygmseus, Linn.). By John Anderson, M.D., F.B.S.F., F.L.S., F.Z.S., Sfc, Curator of Bidian Museum and Professor of Comparative Anatomy at Medical College, Calcutta. (Plate XXXV. and 2 Woodcuts.) 213

PART v.— October, 1877.

XI. On Steer e's Sponge, a new Genus of the Seccactinellid Group of the Spongida. By

James Murie, M.D., F.L.S., F.G.S., ^c. (Plates XXXVI. & XXXVII. and 2 Woodcuts.) 219

XII. Notes upon the Oxystomatous Crustacea. By Edward J. Miers, F.L.S., F.Z.S.,

Assistant in the Zoological Department, British Museum. (Plates XXXVIII.- XL.) 235

XIII. On the Domestic Pig of Prehistoric times in Britain, and on the mutual Belations of this Variety of Pig and Sus scrofa ferus, Sus cristatus, Sus andanianensis, and Sus barbatus. By George Rolleston, M.D., F.B.S., F.L.S., Linacre Professor of Anatomy and Physiology, Oxford. (Plates XLI.-XLIII.) 251

[ V ]

XIV. On two new Forms of Deep-sea Ascidians, obtained during the Voyage of JEL.M.S. ' Challenger.' By H. N. Moselet, M.A., F.B.S., Fellow of Exeter College, Oxford. {Co77imunicated by Dv. J. MvniE, F.Z.S.) (Plate XLIV.) . page 287

XV. On new Foi^ms 0/ Actiniaria dredged in the Deep Sea ; with a Description of certain

Felagic Surface-swimming Species. By H. N. Moselet, M.A., F.B.S., Fellow of Fxeter College, Oxford, late Naturalist on board H.M.S. ' Challenger.' [Com- municated by Dr. J. Murie, F.L.8.) (Plate XLV.) 295

PAET VI.— November, 1877.

XVI. On the Birds collected by Professor J. B. Steere in the Philippine Archipelago. By R. BowDLER Sharpe, F.L.S., F.Z.S., 8fc., Senior Assistant Zoological Depart- ment, British Museum. (Plates XLVI.-LIV. and 2 Woodcuts.) .... 307

PAET VII.— August, 1878.

XVII. O71 the Male Genital Armature in the European Bhopalocera. By P. Buchanan White, M.D., F.L.S., ^c. (Plates LV.-LVII.) 357

XVIII. Morphology of the Mammalian Ossicula auditiis. By Alban H. Gr. Doran, F.B.C.S. {Commimicated by Professor Plower, F.R.S., F.L.S., ^c. (Plates LVIII.-LXIV.) 371

XIX. On the Annelida obtained during the cruise of E.M.S. 'Valorous' to Davis Strait in 1875. By W. Carmicuael M'Intosh, M.D., LL.D., F.B.SS. L. 8f E., F.L.S., 8fc. (Plate LXV. and 2 Woodcuts.) 499

XX. On the Sacral Plexus and Sacral Vertebrae of Lizards and other Vertebrata. By

St. George Mivart, Sec. Linn. Soc, and the Rev. Egbert Clarke, F.L.S. (Plates LXVI. & LXVII. and 9 Woodcuts.) 513

PAET VIII.— June, 1879.

XXI. The Btitterflies of Malacca. By Arthur Gardiner Butler, F.L.S. , F.Z.S., Senior Assistant in the Zoological Department, British Museum. (Plates LXVIII. & LXIX.) 533

[ vi ]

XXII. On certain Organs of the Cidaridce. By Ohaeles Stbwakt, Esq., F.L.S., M.E.C.S., Lecturer on Comparative Anatomy, St. Thomas's Hospital. (Plate LXX.) iv.c^^. f^--.' pj^ge 569

XXIII. On Hypsiprimnodon, Bamsay, a Genus indicative of a distinct Family (Pleopo- didge), in the Diprotodont Section of the Marsupialia. By Professor Owen, C.B., F.B.S., F.L.S., &c.. Superintendent of the Natural- History Departments in the British Museum. (Plates LXXI. & LXXII. witli 3 Woodcuts.) , ... 573

XXIV. Descriptions of some minute Hymenopterous Insects. By Prof. J. 0. Westwood, M.A., F.B.S., c&c. (Plate LXXIII.) 583

Index 595

EEEATA.

Page 59, bottom line, for loupe read pocket-lens 240, line 15 from top, for N. plicata read N. sinuata 566, 5 from top, /or Altiisara reatZ Abisara 568, 18 from bottom, for Tagisdes read Tagiades

THE

TRANSACTIONS

OF

THE LINNEAN SOCIETY

OF

LONDON.

SECOND SERIES.-ZOOLOGY.

VOLUME I. PART THE FIRST.

LONDON:

PKINTED BY TAYLOR AND FEANCIS, BED LION COURT, FLEET STEEET :

SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE ; AND BY LONGMANS, GREEN, READER, AND DYER, PATERNOSTER- ROW.

M.DCCC.LXXV.

CONTENTS.

PAET I.— 1875.

I. On the Morfhology of the Skull in the Woodpeckers (Picidae) and Wrynecks

(Yungidae). By W. K. Parkbk, Esq., F.B.S. (Communicated by H. T. Stainton, Bsq., Sec. L.S.) page 1

II. On some Atlantic Crustacea from the ' Challenger ' Expedition. JBy Dr. R. v. Willemoes-Stjhm, Naturalist to the Expedition. ( Communicated by Prof. Wtville Thomson, F.B.S. 8f L.S.) 23

III. On the Structure and Systematic Position of Stephanoscyplms mirabilis, the Type of a new Order of Hydrozoa. By Prof. Allman, M.B., LL.D., F.B.S., Sj-c, Pres. Linn. Soc 61

TRANSACTIONS

THE LINNEAN SOCIETY.

I. On the Morphology of the Skull in the Wood2ieckers (Picidse) and Wrynecks (Yungidse). By W. K. Parkek, Esq., F.B.S. (Commumcated by H. T. Stainton, Bsq., Sec.Z.S.)

(Plates I.-V.)

Read April 2nd, 1874.

Introduction.

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

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

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

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

SECOND SERIES. ZOOLOGY, VOL. I. B

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

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

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

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

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

On the Structure and Development of the Face in the " Picidte (Celeomorphse," H.,

« Saurognathse," P.). The necessity for a morphological study of birds becomes manifest when it is re- membered that Coracias was for a very long time classed with the " Corvidse," and that,

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 3

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

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

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

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

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

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

b2

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

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

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

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

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

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 5

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

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

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

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

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

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

The angle below the centre for the pars plana is the substratum that should form the " OS uncinatum;" the mass in front of this part, and below the superior turbinal {u. tb), is the nasal gland {n.g) ; the " aliseptal" {al.s), or root of the inferior turbinal, is seen above; in front of this the " alinasal" fold {al.n) ; below this the nasal floor {n.f), and altogether in front the pedate trabecular cornu [c. tr). All these parts are brought into daylight by transverse sections (Plate I. figs. 5-8). In the foremost of these (fig. 5) the recurrent trabecular horns (<?. tr) are shown as two thick lobular masses of cartilage, with a binding isthmus formed between the ends of the " cornua." This is very similar to what is seen in an ordinary Mammal, e.g. the Pig. Hedging in the tuberous mass we see the nasal processes of the prsemaxillaries {n. px), the dentary part {d.px), the fore shaft of the maxillary {mx), the palatine spur of the prsemaxillary {p. px), and outside this the pra^palatine bar (p.pa). In the next section (fig. 6) the same bony sections are shown, with the exception of the palatine portion of the prsemaxillary. The " nasal nerves " {n. n) come into view here below the fore part of the septum {s. n), and at the top this wall has grown into a pair of roof-plates (aliseptals), which develop into a large thick half-scroU, the " alinasal turbinal " [al. th). Outside these valvular growths there is an infero-lateral development of cartilage, which at first is a toall and then 2i floor {n. ID, n.f). In the succeeding section (fig. 7) new parts come into view : the two arms of the nasal bone {n) ; the interpalatine spur {i.pa) ; and the fore part of the " inferior turbinal " {i. tb). The section of this arrested scroll is hammer-shaped, an outgrowth being developed inside as well as outside the root ; the outer coil is the rudiment of the scroll, which in Bhea and Tinctmus is coiled upon itself three times (" Ostrich skuU,"

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 7

plate X. figs. 14 & 15, and plate xv. fig. i. tb) ; in the Powl (" Fowl's skull," plate Ixxxvi. figs. 4-6, i. tb) it lias two coils. In the same figure (7) the aHnasal turbinal (al. tb) is seen to form one nearly perfect coil. It is somewhat more like the inferior turbinal of ordinary birds. In these things the Woodpecker is but little more specialized than a Lizard or Sea-Turtle.

Behind, the inferior turbinal (fig. 8) is continuous above with the aliethmoid (al.e), which infolds itself, is pinched in and crenate, and becomes a rudimentary superior tur- binal. This will be better seen in the thoroughly ossified parts of the adult (Plate V. fig. 2, a. tb) : where the inferior turbinal ends, running into the front of the antorbital plate, there a rudimentary " middle turbinal " is seen (ibid. m. tb). In the young bird this section (Plate I. fig. 8) shows the termination of the nasal floor (n.f), curving upwards to the partially ossified " perpendicular plate " (p.e) ; the nasal nerves (n. n), the palatine (pa), the nasals, frontals, and nasal processes of the prsemaxillaries (n,f, n.px) are also cut through, and a good view is obtained of the extent and relations of the " maxiUo-palatine processes" (mx.p). The condition of the nasal floor in this section is preparatory to a remarkable metamorphosis, to be described hereafter. The adult skull and face (Plate II.) can now be interpreted, notwithstanding the mixture there is in it of arrest and of an unusual degree of metamorphosis.

The palate (figs. 1 & 2) is a remarkable piece of basketwork ; and as in this species ossification is intense, the nasal labyrinth brings in its many bony parts to add to the complexity. The prseoral facial bones are thin and splintery, but are, notwithstanding, very strong. The strongest part is the solid fore end of the prsemaxillary ; yet it is quite hollow, forming scarcely any diploe except short side props at the junction of the lateral with the lower plate. The head is the hammer, and the bony beak the helve, to this curious combination of axe and chisel, the dense horny sheath forming an apt " suc- cedaneum " for steel. The bindings on and interlacements of the facial plates and bars have then." meaning in this, that they carry off the jar from the brain of this plumy artisan. Taking the parts in detail, we find that the " basipterygoid processes " {b.pg) are arrested, and yet the pterygoids cleave close to the converging cranio-facial base.

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

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

the prsemaxillary (figs. 1, 2, 3, p.])cc) comes, besides, under the fore part of the septum with its recurrent wings; it has anchylosed with the inner edge of the long lathy prse- palatine bars (j3.^«). There is also a slender " median palatine process" to the prse- maxillaries {m.p. px) ; in the specimens before me it is distinct from the nasal septum ; but in another of my preparations, now in the College of Surgeons, the two parts are anchylosed. The dentary region of the praemaxillary overlaps, and is completely fused with the maxillary ; the nasal processes (fig. 5, n.px) are well anchylosed to their nasal and frontal surroundings, with a small part of their own median suture evident.

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

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

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

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

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

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 9

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

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

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

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

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

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

SECOND SERIES. ZOOLOGY, VOL. I. C

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

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

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

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

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

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

The basihyal (Plate V. fig. 1, b.br) is If inch in length, the proximal part of the next arch (1st branchial) (5r. 1) 1 inch 2 lines, and the distal portion is about 4f inches in length ; each ramus of the 3rd postoral is therefore about 6 inches long. There is no " urohyal." In this the Woodpeckers agree with the Gannet, Balseniceps, Rhea, and some other " Microglossse."

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

THE SKULL IN THE WOODPECKERS AND WRYNECKS. 11

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

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

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