Acta ornithologica musei zoologici polonici tom I. Warszawa, 27. VII. 1935. Nr 14




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ACTA ORNITHOLOGICA MUSEI ZOOLOGICI POLONICI

Tom I. Warszawa, 27.VII. 1935. NR "14

Wladyslaw RYDZEWSKI*
Studies on Notarium in the Birds

(Tab. VIII - XI)

Z ' Zakladu Anatomji Porôwnawczej Uniwenytetu Warszawskiego).
The term notarium refers to the welding of the particular vertebral elements (bodies of the vertebrae, dorsal and ventral spines, transverse processes) of the dorsal area of spinal column of certain Birds.

The problem of the notarium has not been dealt with in a detailed way until now in the literature. It is true that many authors, while speaking about this formation, give quite vague descriptions, but none them are interested in it a specific way. Even Gadow (3) does not give more precise information concerning the various species of Birds.



Abel (1) covers this subject like a curious problem described as: “It is interesting to determine the presence of the Notariums with individual birds.”

Having at my disposal a rather great number of bird skeletons in the Institute of Comparative anatomy of the University of Warsaw, I dealt specifically with the study of


the notarium, especially at Galliformes, which is is more richly represented in the collections of the Institute, and of which I could to obtain more easily certain species apart from these collections.

Before giving the results of my work, I should thank professor Jan de Tur, director of the Institute, who testifies for my work, and likes it for its invaluable indications.

404 Wl.Rydzewski 2

The starting point to define the order of the provision of the vertebrae forming notarium presents a certain difficulty. In general one regards as dorsal vertebrae, those whose ribs are linked with the sternum (3, 4), the others, having false ribs, count in the number of cervical vertebrae (5). However, in the formation of the notarium only one vertebra has a false rib, in particular in Galliformes - the penultimate cervical vertebra.

Consequently, I take for starting point the first vertebra having an articulation with the false rib, otherwise I cannot determine if this vertebra belongs to the dorsal or cervical vertebrae.

Within the limits of Galliformes the dorsal vertebrae (not welded with the pelvis) one counts 5 - 7, of which 3 or 4 enter the formation of notarium.

The first two vertebrae (i.e. the first and the second) always have false ribs, the remainder have true ribs.

Before describing the particular species, I will initially try to present the description of a typical notarium. It is a formation which one currently sees only in Aves and only in certain groups and species of Aves.

The particular vertebrae taking part in the formation of the notarium, are welded not only by the bodies but in the same way their neural spines, which are strongly welded by forming a continuous osseous blade - the dorsal plate (lamina dorsalis notarii). The transverse processes are linked by the side osseous lamelles by forming a kind of grid - the transverse plate (lamina transversalis notariï). The holes of the transverse plate (Foraminae laminae transversalis) can be more or less covered with membrane, cartilage or osseous lamelles.

The hemal spines (ventrale Dornen, 5) of some of the first vertebrae (two or four) can be welded together, by forming a surface for the insertion of the strong muscles. Consequently the notarium is presented in general, to be formed by completely motionless components.

By leaving the conclusions for the end,
Studies on notarium in the Birds. 405

Fig.1


a - the dorsal plate, b - the transverse plate, c - foramina of the transverse plate, d - the side plates.
I pass now to the description of this formation within particular species. Initially I will describe the families Phasianidae and Tetraonidae because I had the occasion to examine all the species of Poland, pertaining to these families.
Phasianidae.

Perdix perdix (Flaxes).

In the adult specimens that I had at my disposal, there is not any difference between the males and the females in the structure of notarium. In the formation of the notarium consists of vertebrae 2-3-4-5. The neural spine of 5-me vertebra¦ is linked by a ligament with the same spine of 6-me free vertebra, and the latter is joined same manner with 7-me vertebra, already welded with the pelvis. The strongly ossified dorsal plate, at the top right edge, is a little thickened, thanks to the additional osseous lamelles. In Partridges the neural spine of 1-re vertebra and the last vertebra of the neck relatively broad, are raised and, linked between them by ligaments, form a kind of prolongation flexible of notarium. The transverse processes are welded. The lateral lamelles are thin and delicate. The Foraminae laminae transversalis are always six are irregular, not very large, covers most of the membrane. The transverse plate of notarium includes vertebrae 2-3-4-5 except the 6 – me

406 Wl.Rydzewski 4
is free, while giving only in its direction of short forward extensions of the lateral lamelles. The transverse processes of 6-me vertebra, relatively strong, widen towards their ends in the shape of range. The ventral apophyses of vertebrae 2-3-4-5 andpartly of 5-me, are welded so that 2-3-4 apophyses are linked in an osseous blade having of large foramina, covered almost completely with the membrane. The other apophyses are linked in the lower part of the blade by the cartilage. This welding in vertebrae 2-3-4 shows a broad transverse triangular surface compared to the ventral apophyses, being used for insertion of the muscles.

Here dimensions in millimetres:




A

25

25,5

24,5

24

24

B

6

6,5

6

6

6

C

26

27

26

25

24,5

D

9,5

9

9

9

8,5

E

6

-

5.5

6

5,5

- Nb a) the length of the higher edge of dorsal plate of the notarium including vertebrae 2-3-4-5.,

b) the height of the dorsal plate close to the 4-me vertebra,

c) the length of transverse plate, including the same ones, vertebrae that in a),

d) the maximum width of the transverse plate close to 5-me vertebra,

e) the minimal width of the transverse plate close to 2-to vertebra.
Among the Partridge specimens I found two of them with notaria built in a different way. Here vertebrae 2-3-4 are welded completely, and is welded to 5-me with 4-me by the centrum and by the higher edge of the dorsal plate, while the interior of the apophysis is filled by cartilage. This was probably in connection with the ossification of notarium. The edge of the dorsal plate is almost right, 6-me vertebra is linked by a ligament with vertebrae 5-me and 7-me. The transverse plate is also visible here only within the limits of the vertebrae 2-3-4. It thus forms fours foramina in the transverse plate.

Studies on notarium in the Birds. 407


The welding of the ventral apophyses includes vertebrae 2-3-4. Besides this, it has no other difference. Dimensions:

a

19,5

19

b

6

6

c

19

17,5

d

8

8

e

5,5

5,5

Nb. a) the length of the higher edge of the dorsal plate of notarium. including vertebrae 2-3-4,

b) the height of the dorsal plate close to 4-me vertebra,

c) the length of the transverse plate, including/understanding the same ones vertebrae that in a),

d) the maximum width of the transverse plate close to vertebra 4-me

e) the minimal width of the transverse plate close to vertebtra 2.

Alectoris Barbara (BON.)

This is an adult specimen, coming from Algeria - sex undetermined. The Notarium is developed identically with that of Perdix perdix (LINN.). The same degree of ossification.

Dimensions:

a) 26,5

b) 8

c) 26


d) 15

e) 6 (Same significances as for Perdix perdix page 4)


The difference between these two species consists only in the width of the transverse plate, which at Perdix petrosa is appreciably broader close to the 5-me vertebra. And for this reason its form differs from that from Perdix perdix (LINN.).
Coturnix coturaix (LINN.).

The structure of notarium of Caille is most delicate and lowest in the family of Perdicinae, although completely

408 Wl.Rydzewski 6
regular. In correlation with dimensions of these two species their notarium does not present substantial differences. The higher edge of the dorsal plate is slightly bent outwards and can be thickened, as with all other Galliformes. It is linked by an osseous lamelle with the neural spine of the 6-me vertebra which is also welded to the body of vertebra with the other vertebrae, and only the center of the spine is filled by a ligament. The lateral lamelles are very delicate. The ventral apophyses are welded at vertebrae 2-3-4.

Dimensions:

a) 19

b) 4,5

c) 18

d) 8,5


e) 7 (Even significance that for lost Perdix: p. 4),

Phasianus sp.

Adult specimens. Vertebrae 2-3-4-5 are welded, 6-me and 5-me are linked with 7-me by a ligament The dorsal plate is higher on the right edge and is strongly thickened. The transverse plate is covered with a fine membrane. The ventral Apophysis is welded in vertebrae 2-3-4.

Dimensions:







Male

Female

Male




a

42

40

39,5




b

10,5

9

10




c

"44

33,5

38




d

2l

20

iS




e

17

-

15

(Even significance that for Perdix perdix p. 4).

The difference between the male and the female: the transverse plate is shorter in the female.


Pavo cristatus lInn.

(Tabl. VIII, phot.I and 2).



Adult specimens. - male. Notarium is very strongly developed

7 Studies on notarium in the Birds. 409
The dorsal plate is high and thick, with the thick higher edge and bent slightly outwards. Vertebrae 2-3-4-5 is entirely welded, vertebra 6-me is free. The thickening of the edge of the dorsal plate extends the osseous lamelles stretching to vertebra 5-me to the sacral vertebrae, but is not welded with the osseous neural spine of vertebra 6-me. The neural spine of vertebra 5-me links by a ligament with the neural spine of vertebra 6-me. The transverse plate of the notarium is strongly developed. It is characterized by the absence of regular foramina of the transverse plate: one sees only the small ones there, irregularly dispersed holes, because of the broad transverse plates, strongly developed, which link the transverse processes, and the holes of the transverse plate are covered by gross osseous lamelles. The ventral apophyses are welded in vertebrae 2-3.

Dimensions:

a) 53

b) 18


c) 54,5

d) 33


E) 25 (Even significance that for Perdix perdix p. 4 -)

Gallus gallus (LINN.).

Adult specimens. The greatest variations in the development of notarium is seen in domestic Hens. It is rather easy to understand, being given the great number of races produced by the poultry breeders, and consequently, the differences in the head of these birds, the thickness of their bones, the constitution of all the body. The notarium is developed here in quite different ways. The dorsal plate is composed in theory of vertebrae 3-4-5-6 or vertebrae 2-3-4-5. Apart from these two cases, one can still see other variations like the welding of three vertebrae only, or welding partial parts of the vertebrae in the dorsal plate . In the first case vertebra 6-me is normal with vertebra 7-me (not welded with the pelvis) by a ligament, in the second vertebra 5-me is joined in the same manner with free vertebra 6-me,

410 Wl. Rydzewski 8


and the latter with vertebra 7-me already welded with the pelvis. The neural spine of vertebra 2-me, (in the first case) and of 1-re (in the second) broad and high, are linked by a ligament and sometimes by additional osseous lamelles with the vertebrae. One also sees partial welding of the vertebrae in the centra and the higher edge of the neural spines are welded, while the center of the dorsal plate is filled by a ligament. The higher edge of the dorsal plate for the majority is bent outwards, and is slightly thickened various ways. In any case in domestic Hens the thickening of the edges of the dorsal plate increases in the direction of the pelvis, by forming the surface of the articulation (with comp. page 16).

The welding and the ossification of the transverse plate shows various degrees of development. Here in the same way one meets two principal types, i.e. the welding of vertebrae 3-4-5-6, or vertebrae 2-3-4-5. The transverse plates are strongly developed and provided with irregular thickenings. The irregular formaina of the transverse plate, are covered with a manner varied of osseous lamelles or membranes. Under this report one can establish in hens no definite type. Since the large foramina are only covered with the membrane, in all a series of modifications, we arrive finally at the foramina covered almost completely by large osseous lamelles. All things considered we have here the type brought closer to the Peacock.

The ventral apophyses are welded in vertebrae 2-3-4-5 entirely or partially.

Dimensions:




a

43,51)

50, 5l)

32-412)

37- 432)

403)

413)

433)

b

13,5

15

12

13,5

13

12

14

c

44,5

48

40,5

42

41,5

41

44

d

25,5

27,5

24

25

22

23

24

e

19,5

19,5

18

19,5

17

17

20

Nb. 1) includes vertebrae 3-4-5-6.

2) includes vertebrae 3-4-5. The 6-me is welded by the body vertebra. The first measurement for vertebrae 3-4-5, the second for vertebrae 3-4-5-6, the transverse plate includes vertebrae 3-4-5-6.

3) includes vertebrae 2-3-4-5.

a), b), c), d), e) - order and significance as for Perdix perdix p. 4.

Studies on notarium in the Birds. 409
Tetraonidae.

Tetrao urogallus linn.

Adult specimens. males and a female. Vertebrae 2-3-4-5 are welded. Vertebra 6-me is linked with 5-me and 7-me by a ligament. The higher right edge of the dorsal plate is slightly thickened. The transverse plate of the “Guinea fowl” has foramina that are large and regular (with comp. page 11), but covered with a fine membrane. The ventral apophyses of vertebrae 2-3 are welded and linked by a ligament with 4-me.

Dimensions:






Fem

Male

Male

Male

Male




a

44,5

57

57

57

56




b

9

15

16

16

15




c

42,5

64

64

65

62




d

25

34

34

36

34




e

19,5

29

30

29

28,5

(Even significance that is lost for Perdix perdix p. 4).



Lyrurus tetrix (LINN.).

Adult Specimens. Welded vertebrae 2-3-4-5, 6-me are linked by a ligament with vertebrae 5-me and 7-me, which is welded with the pelvis. The right higher edge of the dorsal plate and is thickened. The strongly developed transverse plate, has foramina less regular than in the Cock of heather, and partially covered by osseous lamelles. The ventral apophyses of vertebrae 2-3-4-5 are not welded between them, and are linked only by ligaments.

412 Wl. Rydzewski 10


Dimensions:





Fem

Male

Male




a

39,5

41

39




b

7,5

8

7,5




c

44

45,5

43,5




d

23

21

21




e

16

15,5

15,5

(even significance that for Perdix perdix p. 4).


Lagopus sp.

( VIII, phot . 3).



Adult specimen. unspecified sex . Vertebrae 2-3-4-5 are welded. Vertebra 6-me is free, and linked with vertebrae 5-me and 7-me by a ligament. The vertebra 7-me is welded with the pelvis. The higher edge of the dorsal plate - slightly thickened and is slightly bent outwards. The transverse plate is strongly developed with foramina covered with rather strong osseous lamelles, for this family. Ventral apophyses of the vertebrae 2 - 3 are welded, others - free.

Dimensions:

a) 31,5

b) 7


c) 32

d) 17


e) 7,5 Même significance, that for Perdix perdix p. 4).

Tetrastes bonasia (LINN.).

Adult specimen. unspecified sex . In comparison with the size of bird, the notarium is developed here rather strongly. Welded vertebrae 2-3-4-5. The vertebra 6-me is free, linked by a ligament with vertebrae 5-me and 7-me, the latter welded with the pelvis. The higher right edge of the dorsal plate and is thickened. The transverse plate includes vertebrae 2-3-4-5. The lateral lamelles which are slightly developed stretch in extensions until the transverse processes of vertebra 6-me, but are not welded with the latter. The foramina of the transverse plate of unimportant size and irregular, are covered

11 Studies on notarium in the Birds. 413


by a thin osseous plate. The ventral apophyses are welded in vertebrae 2-3-4.

Dimensions:

a) 24

b) 5


c) 25,5

d) 12, 5

e) 9 (Even significance that for Perdix perdix p. 4

Numididae.

Numida meleagris (linn.) ( IX, phot . 4 and 5).

Adult specimens. females . The vertebrae 2-3-4-5 are welded. Vertebra 6-me is linked with 5-me and 7-me by a ligament. The vertebra 7-me welded with the pelvis. The higher edge of the dorsal plate is the right. In the transverse plate the transverse lamelles are very definitely developed, form large and regular foramina of the transverse plate which are not covered with membrane or osseous lamelles. The ventral apophyses are welded in vertebrae 2-3-4 giving relatively one surface limited well for the insertion of the muscles.

Dimensions:




a

41,5

39




b

10

10




c

41

40,5




d

23

23




e

16

17

(Même significance that for Perdix perdix p. 4) -



Meleagridae.

Meleagris gallopavo lINN.

( IX and X, phot . 6 and 7).



Adult specimens. males and females. In the Turkey breadth notarium is developed most strongly among all

414 Wl. Rydzewski 12


species which I studied. Vertebrae 2-3-4, and partially the 5-me, are welded. 6-me is already in the area of the pelvis (but is not welded with it) and links itself with vertebra 5-me and 7-me by a ligament. Vertebra 5-me is welded with the preceding ones either completely, or by the centrum and the higher edge of the dorsal plate, which strongly is thickened and bent in outward. In this last case there remains a slit in the higher part of the dorsal plate. The strongly developed transverse plate, the lateral lamelles link the transverse processes of vertebrae 2-3-4-5.

In the Turkeys one finds formations which are hardly encountered in the other species of Galliformes. They are in particular ossifications going obliquely of the dorsal plate to the transverse plate. They irregular, more or less are ossified, welded in their basal parts with the transverse processes by an osseous element of spongy consistency. In addition in the studied specimens on the dorsal plate were additional osseous lamelles going along notarium (see Ci afterwards). The ventral apophyses are developed in all the vertebrae, except, are welded to 5-me in vertebrae 2-3-4 and are linked by ligaments with 1-re vertebra and the last cervical.



Dimensions:


a

59

54

57,5

59

58

50

63,5

68,5

54

54

59

62,5

b

21, 5

16

22

22

22

17

23

23

17

18

21

22

c

60

54

59

60

61

51

65

7l

54

54,5

57

65,5

d

37

33

36

37,5

34,5

31

40

40,5

32,5

32,5

36

38,5

c

31

25

29 . 5

29,5

31

24,5

24,5

32

24

25

29

29

(Even significance that fear Perdix perdix p. 4).
Cracidae.

I had the occasion to study only two specimens, within this family. They are in the collection of Institute

13 Studies on notarium in the Birds. 415

of Comparative anatomy of the University of Warsaw. They are Mitu mitu (LINN.) and Penelope marail (P. L S . Müll) coming from Guyana. These species belong to two different genera.


Panelope marail gmel.

Adult specimen. unspecified sex. Vertebrae 2-3-4-5 are welded. Vertebra 6-me is free, linked by a ligament with 5-me and 7-me. The higher edge of the dorsal plate is slightly thickened and completely right. The well developed transverse plate, approaches with the type in the Guinea fowl being bored with the large regular foramina. The foramina of the transverse plate are covered partially with a delicate membrane. The ventral apophyses, which develop only in vertebrae 2-3, are welded.

Dimensions:

a) 38

b) 9


c) 30,5

d) 23,5

e) 15,5 (Even significance that for Perdix perdix p. 4)

Mitu mitu (LINN.).

Adult specimen. unspecified sex . Vertebrae 2-3-4-5 are welded. The 6-me is linked with 5-me and 7-me by a ligament. The higher edge of the dorsal plate slightly thickened and slightly bent outwards. The strongly developed transverse plate, with foramina lies covered with membrane . The ventral apophyses are well developed in vertebrae 2-3-4, are not welded.

Dimensions:

a) 57,5

b) 15,5

d) 31

e) 21 (Even significance that for Lost perdix p. 4),



416 Wl. Rydzewski 14

Opisthocomidae.

Opisthocomus hoazin (P. L S. mûll.)

( X, phot . 8).

Adult specimen coming from South America. unspecified sex . Vertebrae 2-3-4 are welded. 5-me and 6-me vertebrae are linked by a ligament with 4-me and 7-me, which is also welded. The higher edge of the dorsal plate - thickened and straight. The transverse plate, slightly developed, shows a completely specific structure. There are not here the lateral lamelles but only the transverse processes, very broad, which are linked with the bodies of the vertebrae by triangular osseous lamelles. The two free vertebrae, are 5-me and 6-me, are linked with very strong neural spines, with a flat rim - as well as vertebra 1-Re.

For lack of a more significant number of specimens of this species it was impossible for me to solve the problem if this welded feature is a constant character for this species, or is an exceptional case. It is possible that this skeleton had undergone artificial deformations during the preparation. It should be noted, all the same, that the configuration of this transverse plate differs appreciably from that of the other species.

There are ventral apophyses.

Dimensions:

a) 28,5

b) 10


c) the side plates do not exist.

d) 19


e) 8 (Mêm E significance that for Perdix perdix p. 4),

* ***********************************

The notarium is always welded in the Galliformes. It is developed and perfectly strongly ossified here, by including vertebrae 2-3-4-5. Vertebra 6-me is usually linked by a ligament with the 5-me and 7-me, and the latter is sometimes welded with the pelvis.

While the dorsal plate is not prone to the large

15 Studies on notarium in the Birds. 417
variations, the transverse plate can show highly varied aspects (Pavo, Meleagris, Numida). That can be connected with the various degrees of the development of the lateral lamelles, and the covering of the foramina of the transverse plate by thin osseous lamelles.

The development of the ventral apophyses is subjected in the same way to the various variations. In spite of these differences, Galliformes remain welded in all, in theory, the same way in its characters.

It is to be noticed, that in the area of welding special articulation is developed. It is quite especially marked at Gallus gallus (LINN.). This articulation has a thickening higher edge of the dorsal plate of the last vertebra of the notarium. The dimensions of the notarium is a concave surface of the articulation and on the edge corresponding of the neural spine of the following vertebra (either free or welded with the pelvis) - convex articular surface. This is, in principle, the only point of the notarium linked with the pelvis, while remaining mobile, this articulation plays the role of an element ensuring a special stiffness of the notarium in the transverse direction compared to the axis of the body.

The phenomenon of the same character, decides in the whole of osseous lamelles or of tendons, the energy of notarium with the sacral vertebrae. These lamelles do not develop at all the species. They are of the two dimensions of the dorsal plate in the higher edge. The whole of these lamelles is composed of 4 or 6 long lamelles, inserted into the neural spines of the last vertebra of notarium and stretching between the lower part of the illia beyond the sacral vertebrae. Moreover, a variable number of short lamelles extend in the same direction, like in the opposed direction, i. e. of the pelvis to notarium, by intertwining the neural spine of the free vertebra, which is between notarium and the pelvis. All these lamelles or are welded to the apophyses, or link themselves there by tendons. On photograph 2 (Pavo) one sees this lengthened lamelle, stretch along the neural spine.

Photography 9 represents these plates in the Turkey.

418 Wl. Rydzewski 16

In the Cock of bruyère, apart from the lamelles mentioned one finds two more widened lamelles, welded with illia (photo 10).

At side of the ossified lamelles we also meet strong tendons. All these lamelles are strongly developed in the quite old birds. They are probably formed in consequence of the ossification of the tendons. In Guinea fowl these lamelles are replaced by strong tendons, which, while crossing, also embrace the neural spine of the free vertebra.

In Galliformes the notarium is an ordinary formation, but not exclusively for this group of Birds. We also encounter the notarium in the Birds belonging to other systematic groups, but it then presents a particular phenomenon for separate species and cannot be generalized in all the representatives of a given group.

I present the description of notarium at the species, which were at my disposal, where it is more or less developed. We see that they belong to the various systematic groups.

I regard notarium of Galliformes as best developed in the contemporary Birds.
Falconidae.

Tinnunculus columbarius (LINN.) and Cerchneis tinnuuculus (LINN.) are the two raptors representatives that have a notarium. The dorsal plate, as well as the transverse plate, are developed in a normal way, as in Galliformes. The ventral apophyses are strongly developed and welded onto the first two vertebrae. Other vertebrae do not have ventral apophyses.
Tinnunculus columbarius (linn.).

Adult specimen. - unspecified sex. Vertebrae 2-3-4-5 are welded together. Vertebra 7-me is free, linked with 6-me and with the pelvis by a ligament. The higher edge of the dorsal plate is right and slightly thickened. The transverse plate is strongly developed and has 8 regular foramina

17 Studies on notarium in the Birds. 419

and not covered by the membrane. The ventral apophyses of the first two vertebrae are welded. Dimensions:

a) 26,5

b) 5

c) 27


d) 12

E) 9,5

Nb . a) the length of the higher edge of the dorsal plate of notarium,

b) the maximum height of the dorsal plate,

c) the length of the transverse plate,

d) the maximum width of the transverse plate,

e) the minimal width of the transverse plate.

Cerchneis tinnunculus (LINN.)

( XI, phot. II).

Adult specimen - female . The notarium is almost identical with that of Tinnunculus columbarius (linn). It differs from it only by the presence from 4 welded vertebrae, the transverse plate has the 6 formaina . Apart from that there are not other differences.

Dimensions:

a) 21

b) 4,5


c) 22,5

d) 12,5

e)10,5 (Even significance that for T. columbarius).
Columbidae.

In the Doves the notarium is developed very strongly. Its principal character consists of an almost total absence of holes of the transverse plate. They are covered by the osseous lamelles osseous, and thus this transverse plate roughly resembles that of the Peacock. The notarium is composed of three vertebrae.


Columba livia gmel.

( XI, phot. 12).

Adult specimen. Unspecified sex. Vertebrae 3-4-5 are welded.

420 Wl. Rydzewski 18


welded . 6-me vertebra - linked with the 5-me and the pelvis by a ligament. The higher edge of the dorsal plate is right and thickened. The transverse plate is strongly developed. One does not see here holes of the transverse plate quite distinct, because they are covered completely or almost by osseous lamellas. The lateral lamelles emit prolongations until the transverse processes of 6-me vertebra. The ventral apophyses, only developed on vertebrae 3-4 are welded.

Dimensions:




a)

18,5

19,5




b)

6

6




c)

21

21




d)

13, 5

13,5




E)

11

11

(Even significance that for T. columb. p. 17).



Columba rupestris pall.

Adult specimen coming from Siberia - sex undetermined. The notarium, developed as in Columbia livia gmel., contains the same vertebrae. But the holes of the transverse plate are covered by osseous lamelles. The holes of the tranverse plate are rather small and irregularly shaped.

Dimensions:

a) 17,5

b) 4,5

c) 20,5

d) 12,5

E) 9 (Even significance that for T. columb. P . 17).


Pteroclidae.
Pterocles alchata (LINN.).

Single specimen, coming from Algeria. Sex undetermined. Only the dorsal plate is developed. It is composed of vertebrae 2-3-4-5. 6-me vertebra is linked with the F 5-me to the pelvis by a ligament. The higher edge of this

19 Studies on notarium in the Birds. 421
plate is right and thickened. The ventral apophyses are weakly developed on vertebrae 2-3. Dimensions:

a) 28


b) 7

Nb . a) the length of the dorsal plate,

b) the maximum height of the dorsal plate.

Trochilidae.

In the two specimens of birds, which I had the occasion to study, the notarium is developed very specially. It is completely welded with the pelvis. The notarium extends on the two vertebrae which are apart from the pelvis.


Hylocharis sapphirina (gmel.)

( XI, phot . 13).

Specimen coming from South America - sex undetermined. Vertebrae 5 and 6 are entirely welded together and with the pelvis. The dorsal plate is slightly developed. The 4 foramina of the transverse plate are covered by very thin osseous lamelles, so that one cannot distinguish the limit between notarium and the pelvis.

Dimensions:

a) 3

b) 1


c) 3

d) 3


e) 3 (Even significance that for T. columb. p. 17).

Trochilus colubris linn.

Specimen coming from South America - finished indé sex . Notarium pronounced as with Hylocharis sapphirina (gmel.). The structure of the pelvis is stronger though. There are not other differences.

Dimensions:

a) 4


b) 1,5

422 Wl. Rydzewski 20

c) 3

d) 3


E) 3 (Even significance that for Pterocles alchata p . 19).

Alaudidae.

In the Larks we see a marked notarium in a quite special way: 1) it can be developed in the form of coalescences of the higher edges of the neural spines by osseous lamelles, and by the welding partial of the transverse processes by lateral lamelles, 2) the neural spines can be welded more or less together forming a regular dorsal plate.


Alauda arvensis

linn. ( XI, phot . 14).

Adult, but young specimen. Female. The vertebrae are not welded between them, which preserves their flexibility. It is not formed by the regular dorsal plate or the transverse plate. The higher edges of the neural spines emit osseous lamelles, embracing the neural spine of the vertebrae located behind. We see here the phenomenon resembling that which we meet at Galliformes (see Galliformes - supplementary lamelles). The transverse processes emit side lamelles which form the configuration of the transverse plate.


Alaemon alaudipes (desf.)

Specimen coming from Algeria; unspecified sex . Welded vertebrae 2-3-4 form the dorsal plate. The transverse plate is not developed.

Dimensions:

a) 10


b) 3 (Even significance that for Pterocles alchata p.19).
Ammomanes isabellina (temm.).

Specimen coming from Algeria - unspecified sex. Vertebrae 3-3-5, welded, form, the dorsal plate. The transverse plate

21 Studies on notarium in the Birds. 423
is not developed, only the transverse processes are linked between them by ligaments.

Dimensions:

a) 7,5

b) 2 (Even significance that for Ptetodes alchata p. 19).

* ***********************************

Until now no one sufficiently understood the formation and functional significance of notarium.

R. Martin (5) while speaking about the welding of the dorsal vertebrae in Columbiformes writes:

“Of the kind of the fusion of the vertebrae little can be said by us, since the system is limited to a high degree upon the animal’s age.¶ It is limited to the peacock ¦¦¦on the vertebral bodies, the neural spines and ventral apophyses¦, by consequence of the ossification of the ligament, ¦ ¦even still the Querfostsätse (lateral lamellas?) also changes in relationship.”

Is this thesis thus applicable to Galliforms? One will not be able to know until studies of development of the embryologic notarium are done. I propose to study this problem in the future.

eaton (3) considers the notarium, in all its parts to be the result of ossification of certain ligaments of the thoracic area of the spinal column, and the ossification of the tendons of muscles - longissimuss dorsi and socro-lumbales.

With regard to the problem of the functional role of notarium, the opinions of the authors are shared. For example, the welding of the ventral apophyses is very comprehensible as bases insertion of the strong muscles flexeurs of the neck (3) - as for the other parts of notarium the opinions are divided, and allow me to quote some of it here.

J. versluys (8) writing:

“A very strong sternum appears necessary, in order to give to the strong flight muscles a sufficient firm origin surface; thus in connection we find the thoracic vertebrae only a few among themselves mobile, they are often connected among themselves merging into one piece - the Notarium.”

424 Wl. Rydzewski 22
äöü

A. reichenow (6) says:

“The dorsal vertebrae, whose count 4-13, deformed with one another and give the strength necessary for flight to the trunk”

O. Abel (i) speaks:

“This feature exists in the fusion of a changing number of dorsal vertebrae segmented to the Notarium, whose function in the body is something similar to the pelvis in walking terrestrial vertebrates.”

Finally eaton (2) writes:

“Since this consolidation of dorsal vertebrae in birds, though usually associated with increased power of flight, is not brought about by ossification of the tendons of any muscles of the fore limb, but rather by the ossification of the tendinous origins and insertions of muscles of trunk and neck, it seems possible that the extent to which such development is carried may depend in some slight degree upon the physiology of cervical region."

It seems to me that the opinion that the notarium is dependent on flight in General is not justified. It can be in connection with a certain kind of flight, because, as I had already the occasion to note, one sees it the best developed in Galliformes, which have well known specific flight.

In addition, the Birds whose flight appears to us most perfect, like the raptors, do not have any trace of notarium.

In Pteranodon, which has the best developed notarium of all animals (2), one notices that the former area of the body is very strongly developed, while the pelvis is developed only slightly. That appears, supposes ABEL (1) because:.. the major task ... is taken over with the pterosaur by the arms.”

It is necessary to suppose that the notarium appears in proportion with the movements of runners, specific to all Galliformes. However, the representative types of this manner of being. e.g. the Ostriches, Kazuars etc do not even have a trace of notarium (7).

23 Studies on notarium in the Birds. 425

Two possibilities still remain for us to consider, namely the manner of feeding and specific types of flight.

I have the impression that with regard to Galliformes, the development of the notarium can be connected with the specific manner in which these birds take food, i.e. wedging. With a relatively short neck, a strong scaffolding of the thorax is essential, more especially as all the weight of the body rests on the pelvis, and, consequently, the backbone is immobilized to provide assistance to maintain balance.

Finally the specific type of flight. Indeed, at all Galliforms we observe a flight trémoussé (Flatterflug) and in all the other birds, which have the same type of flight we find more or less notarium developed. And, for example, Cerchneis tinnunculus (LINN.) and Tinnunculus columbarius (LINN.) have notaria very distinctly developed. The Larks and some of the Bird-flies show a degree different from the welding of the dorsal vertebrae.

In addition Columbiformes, whose flight cannot be regarded as a typical trémoussé flight, have notarium rather strongly developed.

While summarizing I believe that it is not still possible to explain the functional significance of notarium. All the same the most acceptable assumption would be to admit that it is a formation especially dependent on the “trémoussant” flight type.

BIBLIOGRAPHY

1. Abel O Grundzùge der Palaeobiologie der Wirbeltiere . Stuttgart 1912.

2. eaton G F Osteology of Pteranodon . Memoirs of tea Connecticut Academy Ci Arts and Sciens, vol. II. 1910.



3. gadow H. Die Vôgel;in broun' S Klassen und Ordnungen Tierreichs.

4. ma . rsiul . l W DER Beam der Vôgel, Leipzig 1895.

5 Martin R. Die vergleichende Osteologie to dîr Golumbiformas. Zool . Jahrb . XX. Heft III Jena, 1904.



6. reicbenow A. Die Vôgel . Stuttgart 1913 - 14.

7. shufeldt R. W. One tea classification of unquestionable tea groups of birds . AmenCâa
Naturalist XXXVII. Boston 1903.

8. versluts J, Das Skelet; in . G. chu. hirrsch Vergleichende Anatomy der Wirbeltiere, Berlin, 1927.
426 Wl. Rydzewski. 24

EXPLANATIONS OF THE PHOTOGRAPHS.

1. Pavo cristatus linn. - side view dorsal.

2. Pavo cristatus linn.- side view. One sees an additional plate étirant notarium with the pelvis.

3. Lagopus sp.-' dorsal side view.

4. Numida meleagris (LiNN.)-sight on dorsal side.

5. Numida meleagris (linn.)- side view.

6. Meleagris gallopavo linn. - side view dorsal. One sees ossifications étierant itself obliquely of the plate dorsal-with the transverse plate.



7. Meleagris gallopavo linn. - side view.

S. Qpisthocomus hoazin (P . L . S. mxjll.)- side view dorsal.

9. Meleagris gallopavo linn. - • additional plates.

10. Tetrao umgallus linn. - additional plates, welded with ossa ilei.

11. Cerchneis tinnunculus (linn.) - side view dorsal.

12. Columba livia gmel.- side view dorsal.

13. Hylocharis sapphirina (ûmel.)- side view dorsal. X 2.

14. Alauda arvansis linn. - side view. X 2.


RÉSUMÉ


…. Not Translated ….

* Original citation: Rydzewski, W. 1935. Etudes sur le notarium chez les Oiseaux. Acta Ornithologica Musei Zoologici Polonici 1:403-427. Translated by Chris Organ, Montana State University, August 2003.


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