Part 14
I have adopted Professor Parker's classification in the genera, only substituting _Cela_ Reichenbach for _Mesapteryx_ Hutton, which is a synonym of _Megalapteryx_ Haast. As to the species I have used my own judgment; I felt obliged to name a number of species acknowledged by Parker and Lydekker but not named, because this system of indicating species by the letters A, B, C, &c., which has crept into our nomenclature, will make all understanding impossible, as not always the same species is denoted by the same letter. A few of these species will naturally later have to be sunk, as some have been founded on skulls and others on leg bones, or so, which, when we get perfect individual skeletons may prove to be identical, but I do not think these will be many.
Besides a number of imperfect eggs, particulars of which will be found in Dr. A. B. Meyer's article in the Ibis, 1903, pp. 188-196, there are known two perfect Moa eggs and one almost perfect one.
1. Otago Museum. Molyneux River, 1901. _Pachyornis pondorosus_.
2. Tring Museum. Molyneux River, 1901. _Megalapteryx huttoni_.
3. Rowley Collection. South Island, 1859. _Dinornis novaezealandiae_.
{191}
DINORNIS.
The skull is broad and much depressed, with a comparatively wide, somewhat pointed and deflected beak. Breadth at the squamosals twice the height at basi-temporal. It has a flattened frontal region, and a wide median ridge on the upper surface of the praemaxillae. The mandible is in the form of a narrow U, with the angle much inflected, no distinct anticular process, and the symphysis moderately wide, narrowing anteriorly, with a prominent and broad inferior ridge, widest in front. The quadrate is elongated, with a very large pneumatic foramen. The sternum is nearly as long as broad, very convex, with distinct coracoidal facets, 3 costal articulations, very small and reflected costal processes, the lateral processes very broad and widely divergent, and a wide xiphisternal notch. The pelvis is narrow with a high ilium, in which the inferior border of the postacetabular portion is flat, and does not descend as a sharp ridge below the level of the anterior postacetabular vertebrae. The pubis has a small pectineal process; and the ventral aspect of the true and postacetabular vertebrae is very broad and much flattened.
The distal extremity of the tibio-tarsus is not inflected. A hallux is present in some species. The tibio-tarsus and tarso-metatarsus are long and slender, the length of the latter equalling and more often exceeding the length of the femur, and also exceeding half the length of the tibio-tarsus. The femur is comparatively long and slender, with a short neck, the head rising but slightly and projecting only a small distance, the linear aspera in the form of a long irregular line, the outer side of the distal extremity moderately expanded, the popliteal depression small, deep, and sharply defined, the profile of the inner condyle semi-ovoid and narrow, and the interior trochlear surface nearly flat. The phalangeals of the pes are long and comparatively slender, the proximal surface of the terminal segments not being trefoil-shaped. In the vertebral column the middle cervicals are long and narrow, with the postzygapophyses directed much outwardly and separated by a very deep channel, and the posterior face of the centrum low and wide. The dorsals have short transverse processes and neural spine, the anterior and middle ones (those with a haemal spine or carina) having a large anterior pneumatic foramen between the nib-facet, the foramen being triangular in shape. All the species of this genus are of comparatively large size, and include the tallest members of the family.
Type of the genus: _Dinornis novaezealandiae_ (Owen).
Number of species: 7. {192}
DINORNIS MAXIMUS OWEN.
_Dinornis maximus_ Owen, Trans. Zool. Soc. VI. p. 497 (1868).
_D. excelsus_ Hutton, Trans. N.Z. Inst. XXIV. p. 110 (1892).
_D. giganteus_ Haast, Trans. N.Z. Inst. I p. 88, No. 20 part.
This is the largest species of Moa, the tibio-tarsus being from 37.5 to 39.2 inches in length, while that of the largest _D. giganteus_ does not exceed 35 inches, but by far the largest number of the latter are considerably shorter.
The type bones were obtained in Glenmark Swamp, Middle Island of New Zealand, and were sent to Professor Owen by Major J. Michael of the Madras Staff Corps. Casts of these bones are in the British Museum, No. A 161 in the Palaeontological Department.
This bird was the tallest of all known birds, though it must have been considerably exceeded in bulk by _Aepyornis ingens_ and _Aepyornis titan_ of Madagascar.
Locality: Glenmark Swamp, Middle Island, New Zealand.
DINORNIS ALTUS OWEN.
_Dinornis maximus_ Owen, Ext. Birds N.Z. p. 253 (Dr. Lillie's specimen) (1879).
_D. altus_ Owen, Ext. Birds N.Z. (1879) p. 361.
_D. giganteus var maximus_ Owen, Trans. Zool. Soc. VI p. 497 (1868).
Only known by a tarso-metatarsus, femur and tibio-tarsus from the Middle Island, New Zealand. The bones at once noticeable by their great length, and are more slender than the same bones in _D. maximus_. This form must therefore, till further material comes to hand, be treated as a separate species.
Locality: Middle Island, New Zealand. Collected by Dr. Lillie. {193}
DINORNIS GIGANTEUS OWEN.
_Dinornis giganteus_ Owen, Trans. Zool. Soc. III p. 237 (1843) and p. 307 (1846).
_Moa giganteus_ Reichenbach, Nat. Syst. der Vög. p. XXX (1850).
_Dinornis maximus_ (non _D. maximus_ Owen of 1867!) Trans. Zool. Soc. X p. 147 (1877).
_D. validus_ Hutton, Trans. N.Z. Inst. p. 111 (1892).
This is, as regards size, one of the more variable forms in the tarso-metatarsus, while the tibio-tarsus is remarkably constant. The tibio-tarsus is almost invariably 35 inches in length, while the tarso-metatarsus varies from 17.5 to 19 inches in length.
The type of _D. giganteus_ Owen is from Poverty Bay; the type of _D. validus_ is from Glenmark.
Habitat: North and Middle Islands, New Zealand.
Portion of skeleton in Tring Museum, from Kopua Swamps, Canterbury, New Zealand.
DINORNIS INGENS OWEN.
(PLATE 42.)
_Dinornis ingens_ Owen, Trans. Zool. Soc. III p. 237 (1843).
_Movia ingens_ Reichenbach, Nat. Syst. der Vög. p. xxx (1850).
_D. ingens var. robustus_ Owen, Trans. Zool. Soc. III p. 307 (1846).
_Palapteryx robustus_ Owen, Trans. Zool. Soc. III p. 345 (1848).
_D. firmus_ Hutton, Trans. N.Z. Inst. XXIV p. 114 (1892).
_D. potens_ Hutton, l.c. p. 115.
_D. ingens_ shows considerable variation in size, but the inter-gradation is so complete that it seems impossible to retain the four species _ingens_, _firmus_, _potens_ and _robustus_, which Captain Hutton admits. This form was widely distributed over the North and Middle Islands. The type skull of _P. robustus_ came from Timaru, the type of _firmus_ from Wanganui, that of _ingens_ from Poverty Bay, while that of _potens_ is quoted from the East side of Middle Island, without specific type locality.
Habitat: North and Middle Islands.
The plate of this species was reconstructed by Mr. Frohawk from the skeleton and feathers in my museum, and the feathers found with the skeleton now in the York Museum. The only criticism that might be made in connection with this picture is that the feathers are drawn a little too much like those of _Apteryx australis_, but this is not of any consequence, as the Moa feathers in the Tring Museum and elsewhere vary considerably in appearance, though being more or less coloured like _Apteryx_ feathers.
There is an almost perfect skeleton in the Tring Museum. {194}
DINORNIS GRACILIS OWEN.
_Dinornis gracilis_ Owen, Trans. Zool. Soc. IV (1855) p. 141.
_D. torosus_ Hutton, Trans. N.Z. Inst. XXIV p. 117 (1892).
If we acknowledge that _D. novaezealandiae_ occurs both on the North and Middle Islands, then I feel sure that the distinctness of _D. gracilis_ and _D. torosus_ cannot be maintained, as the measurements intergrade completely.
The type of _D. gracilis_ came from Wanganui, while that of _D. torosus_ is a nearly perfect skeleton found in a cave at Takaka, near Nelson.
Habitat: New Zealand.
There is an imperfect skeleton in the Tring Museum, from a limestone cave at Takaka, near Motueka, Province of Nelson, New Zealand.
DINORNIS DROMIOIDES OWEN.
_Dinornis dromioides_ Owen, Trans. Zool. Soc. III. p. 235 (1843).
_Palapteryx dromioides_ Reichenbach, Nat. Syst. der Vög. p. XXX (1850).
_Palapteryx plenus_ Hutton, Trans. N.Z. Inst. XXIV p. 122 (1892).
This form also inhabited both islands, but was probably one of the rarest. The type of _D. dromioides_ came from Poverty Bay, and that of _P. plenus_ from Glenmark.
Habitat: New Zealand.
DINORNIS NOVAEZEALANDIAE OWEN.
_Dinornis novaezealandiae_ Owen, P.Z.S. (1843) p. 8.
_D. struthioides_ Owen, Trans. Zool. Soc. III p. 244 (1844).
_D. strennus_ Hutton, Trans. N.Z. Inst. XXV p. 8 (1893).
Professor Owen changed the name of this form, but we cannot accept this change, as it is against the laws of nomenclatorial priority, though we all appreciate the motive the Professor had in making this change. The type came from Poverty Bay, but the bird inhabits both islands.
This species had wings.
Habitat: New Zealand.
A nearly perfect skeleton in the Tring Museum from Waitomo district, Auckland, New Zealand.
{195}
MEGALAPTERYX HAAST.
Originally distinguished by Haast from the _Dinornithidae_ as an ancient form of the _Apterygidae_, but afterwards united by Lydekker with the _Dinornithidae_. Mr. Lydekker's diagnosis of the genus is as follows:--
"Distinguished from _Dinornis_ by the extreme slenderness and length of the femur and tibio-tarsus, and the relatively shorter tarso-metatarsus, of which latter the length is considerably shorter than that of the femur. The pelvis is much narrower than in Dinornis, with the ventral surface of the postacetabular sacrals ridged and narrower, and a more developed pectineal process to the pubis. The femur is markedly curved forwards, with the distal extremity moderately expanded, the popliteal depression larger and less defined, the linea aspera narrower and sharper, and a more distinct anterior intermuscular ridge."
The following additional diagnostic characters are taken from Mr. Charles W. Andrews' description of the complete skeleton of _Megalapteryx tenuipes_ in the Tring Museum (Nov. Zool. IV, pp. 188-194, fig. 1-2 in text and pl. VI):--
Width of cranium at paroccipital processes less than half the length of the basis cranii. Length of premaxilla less than two-and-a-half times that of the basis cranii. Body of the premaxilla pointed and slightly decurved; its length and breadth less than the basis cranii. The occipital plane slightly declined backwards. Occipital condyle projecting slightly beyond the paroccipital processes. Anterior and posterior lambdoidal ridges separated by a very narrow interval in their middle region only. Width at squamosals slightly more than double the length of the basis cranii. Mammillary tuberosities not very prominent. Margin of tympanic cavity evenly curved. Temporal fossae very large. The distance between the temporal ridges about four-fifths the width of the cranium at the fossae. The posterior temporal ridge confluent with the lambdoidal ridge. Post-temporal fossae very large.
The inferior temporal ridge is strongly marked, and there is a pretympanic process. The zygomatic process is well developed. Rostrum dilated towards its anterior end, compressed and carinate beneath the large presphenoid fossae. Mandible very slender. Posterior angular process small. Sternum very convex, and with a very nearly straight anterior border between the tuberosities for the coracoscapular ligaments. Costal processes short but large, with distinct {196} coracoidal facets. The lateral processes are long and distally expanded. The sternum is just as wide as it is long. There are three costal articulations. The most notable character is the enormous length of the toes, the middle one being longer than the tarso-metatarsus. The ungual phalanges are peculiarly long, narrow and curved, instead of being comparatively short and broad, as in most other Moas.
Type of the genus _Megalapteryx hectori_, Haast.
Number of species 4. {197}
MEGALAPTERYX HECTORI HAAST.
_Megalapteryx hectori_ Haast, Trans. Zool. Soc. XII, p. 161 (1886); Lydekker, Cat. Fossil B. Brit. Mus., p. 252.
This form was described by Sir Julius von Haast as a gigantic _Apteryx_. This error arose from the absence of the skull. There is, however, no doubt now, since the skulls of _Megalapteryx_ are known, that although sufficiently aberrant to form a distinct sub-family, the birds included in this genus are _Dinornithidae_ and not _Apterygidae_.
Habitat: Middle Island, New Zealand.
MEGALAPTERYX HAMILTONI SPEC. NOV.
Lydekker, Cat. Fossil Birds in Brit. Mus., p. 252, under _M. tenuipes_ (1891).
The type is a left femur, No. 32145 in the British Museum. It is smaller and relatively narrower than the femur, of either _M. hectori_ or _M. tenuipes_. This is most noticeable at the distal extremity.
Habitat: North Island, New Zealand. (Type locality Waingongoro.)
Named after Mr. A. Hamilton, who did so much in discovering deposits of extinct New Zealand birds. {198}
MEGALAPTERYX TENUIPES LYD.
_Megalapteryx tenuipes_ Lydekker, Cat. Foss. Birds Brit. Mus. p. 251 (1891).
This species was described from the tibio-tarsus, which is longer and relatively more slender than in _M. hectori_. Its distal width is about one-ninth of its length, while in _M. hectori_ it is about one-seventh. The length of the tibio-tarsus is approximately 0.405 mm. = 16 inches, and width of distal extremity about 0.044 = 1.74 inches. Type specimens Nos. 49989 and 49990, British Museum.
Habitat: Middle Island, New Zealand, and perhaps North Island. (Type locality Lake Wakatipa, Queenstown, Otago.)
Complete skeleton in the Tring Museum.
Mr. Lydekker mentions also a right femur from the North Island, of the same proportions as those of _M. tenuipes_ and 0.255 m. (= 10.1 inches) long. It may probably belong to a different form, as we know _M. tenuipes_ otherwise only from the Middle Island. {199}
MEGALAPTERYX HUTTONII (OWEN).
(PLATE 41.)
_Dinornis huttonii_ Owen, Ext. Birds, N.Z., p. 430 (1879).
_Dinornis didinus_ Owen, Trans. Zool. Soc. XI, p. 257 (1883).
_D. didiformis_ Haast, (non Owen 1844) Trans. N.Z. Inst. I, p. 83, Nos. 5 & 6 (1869).
_Mesopteryx didinus_ Hutton, Trans. N.Z. Inst. XXIV, p. 129 (1892).
The synonymy of this form is somewhat confused, but I think it is clear that _huttonii_ of Owen is its proper name. Professor Owen (Ext. B. p. 430) says:
"In the collection from the Glenmark Swamp, South Island, are bones that scarcely differ, save in size, from the dimensions (? W.R.) of the type bones of _Dinornis didiformis_ from the NORTH ISLAND. They are noted as of a large variety of that species." Captain Hutton remarks: "The bones that I have arranged under the name _D. didiformis_ belong probably to a new species. The tibia is well marked and quite distinct, but the femur and metatarsus, that I have associated with it, pass almost into _D. casuarinus_, but are rather smaller. _D. casuarinus_ is undoubtedly a good species, easily distinguished by its tibia." Possibly the _Dinornis_ of the SOUTH ISLAND, with the tibia characteristic of _D. didiformis_ of the NORTH ISLAND, may need to be noted for the convenience of naming the bones as _Dinornis huttonii_.
When describing his _D. didinus_, Professor Owen failed to recognise its identity with his previously named _D. huttonii_, doubtless owing to the leg bones being hidden by the dry integument. This being the case, it is necessary to reinstate the name _huttonii_, as it has four years' priority over _didinus_.
Captain Hutton says that a few bones of this form have been obtained in the North Island at Poverty Bay and Te Aute; but I am convinced he is in error and that these bones are aberrant individual bones of _A. didiformis_ and that _M. huttonii_ is confined to the South or rather Middle Island. The plate of this species has been reconstructed by Mr. Lodge from the mummified remains which form the type specimen of _Didornis didinus_, and the feathers found in the alluvial sands of the CLUTHA RIVER. The type of _Dinornis didinus_ was found at Queenstown by Mr. Squires.
Habitat: Middle Island, New Zealand.
Mr. C. W. Andrews, in his description of my complete skeleton of _Megalapteryx tenuipes_ has shown that Owen's type specimens of his _Dinornis didinus_ are certainly of a species of the genus _Megalapteryx_, and closely {200} allied to _M. tenuipes_. Mr. Andrews, however, throws some doubt as to whether the pelvis and femora, referred to this species by Hutton, really belong to it.
A complete egg which I consider must be of this species is preserved in the Tring Museum. Its measurements are as follows:--
Large circumference, 21.4 inches = 535 mm. Small " 17.5 " = 437.5 mm.
This egg was dredged up on the Molyneux River, near Otago, during gold dredging operations in 1901; a second perfect egg was dredged up a few months before in the same river, and was referred by Dr. Benham to _Pachyornis ponderosus_.
{201}
ANOMALOPTERYX REICHENBACH.
The skull is narrow and vaulted, with a long, sharp and slightly deflected beak. Breadth at the squamosals 1½ times the height at basi-temporal, which has a constricted praemaxillary ridge, and the quadrate with a very small pneumatic foramen. The mandible is V-shaped, with a slight inflection of the angle, and a distinct postarticular process. The symphysis is very narrow and pointed, with a long and narrow inferior ridge, not expanding markedly at either extremity. The sternum is longer, flatter and narrower than in _Dinornis_, having no distinct xiphisternal notch, three costal articulations, long and narrow costal processes, slender lateral processes which are often elongated, and usually no coracoidal facets. The pelvis is wider and lower than in _Dinornis_, with the lower border of the postacetabular portion of the ilium descending as a sharp ridge much below the level of the sacral ribs, and without any distinct pectineal process. A hallux is present. The tibio-tarsus and tarso-metatarsus are relatively shorter and stouter than in _Dinornis_, the latter being shorter than the femur, which is usually stouter and relatively shorter than in _Megalapteryx_. The length of the tarso-metatarsus is less than half that of the tibio-tarsus. The femur, besides being usually relatively shorter is readily distinguished from that of _Dinornis_ by its more expanded extremities, the rather longer neck, and the much larger and ill-defined popliteal depression.
The vertebrae are of the general type of those of _Pachyornis_, but the anterior pneumatic foramen commences in the third dorsal. The phalangeals are intermediate between those of _Dinornis_ and _Pachyornis_. Haast considered that the coracoid was aborted and often absent in this genus, in _Emeus_, and _Pachyornis_. As additional characters of the skull it may be mentioned that there is a prominent supra-occipital protuberance, and a depression on the squamosal above the quadrate; the par-occipital processes are pointed, and the basi-occipital processes only slightly prominent; so that the posterior profile of the basi-occipital is nearly straight. The quadrate has a very short anterior process.
All the species of the genus are small, in fact _parvus_ is the smallest but one of the family.
Type of the genus: _Anomalopteryx didiformis_ (Owen).
Number of species: 4. {202}
ANOMALOPTERYX DIDIFORMIS (OWEN.)
_Dinornis didiformis_ Owen, Trans. Zool. Soc. III, p. 242 (1844).
_Anomalopteryx didiformis_ Reichenbach, Nat. Syst. der Vög. p. 30 (1850).
_A. didiformis_ Lydekker, Cat. Fossil B. Brit. Mus., p. 275.
The present form is confined to the North Island. Owen's type was collected by the Revd. Wm. Williams, and came from Poverty Bay.
Habitat: North Island, New Zealand.
Portion of skeleton in Tring Museum.
ANOMALOPTERYX PARVUS (OWEN.)
_Dinornis parvus_ Owen, Trans. Zool. Soc. XI, pp. 233-256, pls. LI-LVII (1883).
_Anomalopteryx didiformis_ Hutton, Trans. N.Z. Inst. XXIV, p. 123 (1892), part.
_A. parva_ Lydekker, t.c., p. 278.
This small form is confined to the Middle Island. The type, a skeleton in almost complete condition, was dug up in a cave at Takaka, near Nelson, and is now in the British Museum. A much less perfect skeleton is in my museum at Tring.
Habitat: Middle Island, New Zealand.
ANOMALOPTERYX ANTIQUUS HUTT.
_Avian Remains_ Forbes, Trans. N.Z. Inst. XXIII, p. 369 (1891).
_Anomalopteryx antiquus_ Hutton, Trans. N.Z. Inst. XXIV, p. 124 (1892).
_A. antiquus_ was named by Captain Hutton from the photographs of bones described by Dr. Forbes in the above-quoted article. The evidence is very slight on which to found a species, but I prefer to treat it as one, for the bones were discovered in the Upper Miocene, a much older stratum than most remains of _Dinornithidae_ occur in.
Locality: Timaru, Middle Island, New Zealand. {203}
ANOMALOPTERYX FORTIS HUTT.
Anomalopteryx fortis Hutton, Trans. N.Z. Inst. XXV, p. 9 (1893).
This is the largest of the genus, and the type bones came from Glenmark. I append comparative table of Measurements:
+-------------------+------------------+------------- | Tarso-metatarsus. | Tibio-tarsus. | Femur. --------------+-------------------+------------------+------------- A. fortis | 8.0 inches. | 17.5 inches. | 9.8 inches. A. didiformis | 6.3 " | 13.3 " | 8.0 " A. parvus | 6.3 " | 13.7 " | 8.5 " --------------+-------------------+------------------+-------------
Locality of Type: Glenmark.
Habitat: Middle Island, New Zealand.
{205}
CELA REICHENBACH.
Skull convex, the temporal fossae very large. Breadth at the squamosals 1.6-1.7 times the height at the basi-temporal. Length from the supra-occipital to the nasals rather less than the breadth at the squamosals. Occipital condyle hidden by the supra-occipital. Ridge between temporal fossae and supra-occipital narrow. Beak short, slightly compressed and rounded at the tip, though more pointed than in _Anomalopteryx_. Lower mandible nearly straight and rather slighter than in _Anomalopteryx_, V-shaped. Sternum with coracoid pits faintly indicated or absent; length less than breadth. Costal processes well developed, lateral processes diverging at different angles.
Pelvis broader in proportion than in _Dinornis_, the acetabula set more forward. Tarso-metatarsus shorter than the femur, and less than half the length of the tibio-tarsus. Hallux present in some species. The smallest species of Moa is _Cela curtus_.
Type of the genus: _Cela curtus_.
Number of species: 5.
CELA CURTUS (OWEN.)
_Dinornis curtus_ Owen, Trans. Zool. Soc. III, p. 325 (1846).
_Cela curtus_ Reichenbach, Nat. Syst. der Vög. p. 30 (1850).
_Cela curta_ Hutton, Trans. N.Z. Inst. XXIX, p. 550, pl. XLVII, Fig. B.
This and the following are the two smallest species of _Moa_, having been about the size of a large turkey. It also is the most abundant species at Whangarei, and appears to have been most common in the North of the Island. The type is from Poverty Bay.
Habitat: North Island, New Zealand. {206}
CELA OWENI (HAAST).
_Dinornis oweni_ Haast, Trans. Zool. Soc. XII, p. 171, pl. XXXI, XXXII (1886).
_Cela curtus_ Hutton, Trans. N.Z. Inst., XXIV, p. 127 (1892), portion.