The Egyptian German Society for Zoology. The Journal of Basic & Applied Zoology.

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The Journal of Basic & Applied Zoology (2012) 65, 214 219 The Egyptian German Society for Zoology The Journal of Basic & Applied Zoology www.egsz.org www.sciencedirect.

M. Riad Department of Zoology, Faculty of Science, Cairo University, Egypt Received 1 August 2012; accepted 25 August 2012 Available online 17 November 2012 KEYWORDS Intertrabecular bar; Cupolae;

1 The Journal of Basic & Applied Zoology (2012) 65, The Egyptian German Society for Zoology The Journal of Basic & Applied Zoology Studies on the ontogeny of Streptopelia senegalensis aegyptiaca (latham, 1790). 4-Post hatching development of the cartilaginous nasal capsule Total body length of nestling 64 mm M.M. Zaher *, A.M. Riad Department of Zoology, Faculty of Science, Cairo University, Egypt Received 1 August 2012; accepted 25 August 2012 Available online 17 November 2012 KEYWORDS Intertrabecular bar; Cupolae; Concha nasalis; Postprofundal process; Processus paraseptalis posterior Abstract In the post hatching stage of the chondrocranium of Streptopelia, resorption of the intertrabecular bar can now be observed along its whole length. It is most conspicuous in the anterior tip of the prenasal process. The nasal septum is affected by the resorption process. It is now a thin plate. Anteriorly, there is no evidence for the formation of a fenestra septi nasi. The dorsal border of the nasal septum merges with the parietotectal roof above it and with the medial walls of the cupolae. The development of a well chondrified blunt process projecting freely backwards from the posterior margin of the vestibular solum nasi at a point that lies in close vicinity to the intertrabecular bar. The concha nasalis (=paranasal cartilage) becomes extensive and more deep. The postprofundal process shows an upward elongation. The processus paraseptalis posterior becomes much more developed. ª 2012 The Egyptian German Society for Zoology. Production and hosting by Elsevier B.V. All rights reserved. Abbreviations: B.G. PT. C., back growing parietotectal cartilage; C. AN., cupola anterior of the nasal capsule; C. RDG., thick cartilaginous ridge of the back growing roof of the parietotectal cartilage and its anterior latero-dorsal process; CAV. CN., cavum conchale; CN. NAS., concha nasalis; CON. PAR. N. & MAX. TU.C., connection between paries lateralis nasi and maxilloturbinal cartilage; DEN., dentary; FEN. NA., fenestra narina; FEN. CRA. FAC., craniofacial fenestra; ITR. B., intertrabecular bar; L.NA. CAV., lateral portion of the main nasal cavity; LAC., lacrymal bone; LAC. G., lacrymal gland; M.C., Meckel s cartilage; M.NA. CAV., median portion of the main nasal cavity; M.S.L. MAX. TU. C., medially directed secondary lamella of maxillo turbinal cartilage; MAX. TU. C., maxilloturbinal cartilage; MX., maxilla; NA. VES., nasal vestibule; ORB. SIN., orbital sinus; P.L. MAX. TU. C., primary lamella of maxilloturbinal cartilage; P. NA. MAX., nasal process of maxilla; P. PMX. NA., premaxillary process of the nasal; PAR. N., paries lateralis nasi; POSTPROF. P., postprofundal process; PT. C., parietotectal cartilage; S. N., nasal septum; SOL. N., solum nasi. * Corresponding author. Tel.: address: drmostafa133@yahoo.com (M.M. Zaher). Peer review under responsibility of the Egyptian German Society for Zoology. Production and hosting by Elsevier ª 2012 The Egyptian German Society for Zoology. Production and hosting by Elsevier B.V. All rights reserved.

Studies on the ontogeny of Streptopelia senegalensis aegyptiaca (latham, 1790).

Introduction In three, previous publications submitted by the authors (Zaher and Riad, 2009, 2012a, in press), the study of the development of the early and intermediate stages as well as the optimum

In any nestling stage of Streptopelia, all the different regions of the neurocranium show no important difference as compared with those of the optimum stage, except the cartilaginous nasal capsule

That is why it seems, superfluous to go through a full description of the nasal capsule of a post hatching stage of the chondrocranium of Streptopelia.

2 Studies on the ontogeny of Streptopelia senegalensis aegyptiaca (latham, 1790). 215 Figure 1 Transverse section passing through the anterior most region of the nasal capsules of stage VIII (optimum stage) of Streptopelia senegalenis Section No.180 of the series. Introduction In three, previous publications submitted by the authors (Zaher and Riad, 2009, 2012a, in press), the study of the development of the early and intermediate stages as well as the optimum stage of the chondrocranium of Streptopelia was thoroughly dealt with. The present article is the fourth of the series. In any nestling stage of Streptopelia, all the different regions of the neurocranium show no important difference as compared with those of the optimum stage, except the cartilaginous nasal capsule in which some important changes are still taking place. That is why it seems, superfluous to go through a full description of the nasal capsule of a post hatching stage of the chondrocranium of Streptopelia. The description is curtailed only to the main changes occurred as compared with the optimum stage. Materials and methods The material for this investigation comprised three nestling stages of total body lengths; 64 mm, 70 mm and 89 mm the heads were transversely cut, after staining in bulk with Borax Figure 3 Transverse section passing through the middle region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially section is No carmine, the sections were counterstained with picroindigocarmine. The nasal capsule of the 64 mm nesting stage has been reconstructed in ventral and lateral views. Here, the cartilaginous entities seem to assume the stabilized condition when ossification and pneumatization set in rapidly. Results In the present nestling stage, ossification and pneumatization are well marked and are further progressed. However, as far Figure 2 Transverse section passing through the anterior most region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially section is No Figure 4 Transverse section passing though the middle region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially section is No. 266.

3 216 M.M. Zaher, A.M. Riad Figure 5 Transverse section passing through the middle region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially the section is No Figure 6 Transverse section passing through the posterior region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially the section is No as the chondrogenesis is concerned, it is not proposed to give a complete account of it at this stage. All the different regions of the chondrocranium show no important differences as compared with those of the optimum stage, except the nasal

4 Studies on the ontogeny of Streptopelia senegalensis aegyptiaca (latham, 1790). 217 capsule in which some changes are still taking place. However, its description is curtailed only to these main changes. Resorption of the intertrabecular bar (Figs. 1 8, ITR.B.) can now be observed along its whole length. It is most conspicuous in the anterior tip of the prenasal process which is now in the form of scattered cartilaginous fragment on its way to be completely absorbed. Undoubtedly, this has been induced by the ossification of the premaxillae. However, the best preserved part of the intertrabecular bar is that found in the region of the cupola anterior (Figs. 1 and 2, C.AN.). The nasal septum (Figs. 3 and 7, S.N.) has also been affected by the resorption process. It is now a thin plate. Anteriorly, there is no evidence for the formation of a fenestra septi nasi. It is also absent in the 73 mm nestling stage. The postero-dorsal wedge-shaped portion of the nasal septum has also been greatly affected and diminished in breadth when seen in a plane (Figs. 4 6). In the posterior region of the nasal septum, at the transverse level of the postprofundal process, a small craniofacial fenestra is formed by a process of resorption in its middle portion (Fig. 8, FEN. CRA. FAC.). In the 73 mm nestling stage more resorption is observed in the region of the fenestra which becomes remarkably large, oval in shape and extends along the whole length of the nasal septum (Figs. 1 and 3, S.N.) with its long axis dorso ventrally directed. The intertrabecular bar (Figs. 1 and 2, ITR.B.) forms its ventral border, while its dorsal border is the ventral margin of the wedge-shaped portion of the septum. This fact speaks in favor of the possible future formation of a craniofacial fissure when the intertrabecular bar at this point becomes completely regressed in more older nestling stages. In the optimum stage of Streptopelia, it has been stated that the medial walls of the two cupolae anteriores (Fig. 1, C.AN.) fuse imperceptibly together and with the parietotectal cartilage (PT.C.) over the nasal septum (S.N.) to which they are closely applied with a definite line of demarcation. In the present nestling stage, this line of demarcation is completely obliterated and the dorsal border of the nasal septum merges with the parietotectal roof above it and with the medial walls of the cupolae. This gives the impression that the nasal septum has participated in their formation in this particular area (Fig. 2, see also Fig. 1). One of the novelties of the present nestling stage is the development of a well chondrified blunt process projecting freely backwards from the posterior margin of the vestibular solum nasi at a point that lies in close vicinity to the intertrabecular bar (Fig. 2, SOL. N. P.). The back growing roof formed by the parietotectal cartilage is further developed. Its cartilaginous ridges (Figs. 6 and 8, C. RDG.) with their outer lateral processes become very thick, more accentuated and conspicuously dorso-laterally elevated to support the frontal bones. The maxilloturbinal cartilage (Fig. 3, MAX. TU. C.) extends far anteriorly. It gradually changes its topographic relation to the paries lateralis nasi (PAR.N.) from being fused to the medial lower edge of the paranasal cartilage posteriorly Figure 7 Transverse section passing through the posterior region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially the section is No. 323.

5 218 M.M. Zaher, A.M. Riad Fig. 8 Transverse section passing through the posterior region of the nasal capsules of the 64 mm. Young nestling stage of Streptopelia senegalensis. Serially the section is No (Fig. 6) to be attached anteriorly to the upper medial surface of the paries lateralis nasi of the posterior portion of the anterior half of the nasal capsule (Fig. 3). Accompanied with the change of its topographic position, the maxilloturbinal cartilage also shows a new change in its configuration. It elongated medio ventrally and is deeply intruded between the medial and lateral portions of the nasal cavity (Figs. 4 6, M.NA.CAV. & L.NA. CAV.). Anteriorly, it ends with a swollen knob but posteriorly the knob becomes flattened and represents a medially directed secondary lamella. Forwards, this secondary lamella is directed medio-dorsally (Fig. 4) but backwards it is directed medio-ventrally (Fig. 5). Thus the maxilloturbinal cartilage of the previous stage represents its primary lamella. The concha nasalis (Figs. 6 and 7, CN. NAS.) (=paranasal cartilage) becomes extensive and more deep. Its connection with the maxilloturbinal cartilage is persistent. The dorsal connection between the paranasal cartilage and anterior maxillary process of the planum antorbitale is lost. The processus paraseptalis posterior becomes much more developed and acquires an antero-medial projection which is vertically disposed and abuts onto the trabecular bar. Discussion In the present stage of Streptopelia (Post hatching stage, total body length 64 mm) as well as in the 73 mm nestling stage no resorption process is present in the nasal septum. Therefore, there is no evidence for the formation of a fenestra septi nasi. However, the mentioned septi is present in the majority of described birds where it is developed before the optimum stage. The fenestra is developed by a resorption process in the anterior margin of the nasal septum at the transverse level of the fenestra narina of Pyromelana (Engelbrecht, 1958), Upupa & Merops (Mokhtar, 1975) and in Pterocles (Mokhtar et al., 1983), Corvus (Zaher et al., 1993), Coturnix (Abd El-Hady, 2008) and Columba (El-Shikha, 2011). The process of the vestibular floor present in the present stage of Streptopelia is also present in Merops and Upupa (Mokhtar, 1975). Such a process projects at a point that lies lateral to the intertrabecular bar in pterocles (Mokhtar et al., 1983), Gallinula & Bubulcus (Abu-Taira, 1996, 1997). The behavior of the paries lateralis nasi in the present stage of Streptopelia is similar to the condition described by Mokhtar (1975), Mokhtar et al. (1983), Zaher et al. (1991), Abu-Taira (1996, 1997) and Abd El-Hady (2008) in Upupa & Merops, Pterocles, Passer, Gallinula & Bubulcus and Coturnix, respectively. It should be admitted that the secondary lamella of the maxilloturbinal cartilage of the present nestling stage of Streptopelia is a rudimentary one when compared with that of the other described birds where a well medially directed secondary lamellae are of general occurrence e.g., Corvus (Zaher et al., 1993), Gallinula (Abu-Taira, 1996), Coturnix (Abd El-Hady, 2008) and Columba (El-Shikha, 2011). Undoubtedly, the processus paraseptalis posterior of the present stage of Streptopelia is homologous with the anteriorly directed middle turbinal of the planum antorbitale of Pyromelana (Engelbrecht, 1958), Rhea (Mu ller, 1961), Fulica (Til Macke, 1969), Gallinula (Abu-Taira, 1996), Coturnix (Abd El-Hady, 2008) and Columba (El-Shikha, 2011) where its edge is directed anteriorly and rests on the nasal septum. References Abd El-Hady, S.I., Full description of the neurocranium of the optimum stage of Coturnix coturnix japonica (Phasianidae, Galliformes). J. Egypt. Ger. Soc. Zool. 55 (B),

6 Studies on the ontogeny of Streptopelia senegalensis aegyptiaca (latham, 1790). 219 Abu-Taira, A.M., Description of the neurocranium of gallinule Gallinula chloropus (Rallidae, Gruiformes). The optimum stage. Proc. Zool. Soc. A. R. Egypt 27, Abu-Taira, A.M., The development of the osteocranium of the common egret, Bubulcus ibis (Ardeidae, Ciconiformes). The membrane bones and the articular. J. Egypt. Ger. Soc. Zool. 29, El-Shikha, A.M., Anatomical and Histological Studies on the Development of the Skull of an Egyptian Bird, M.Sc. Thesis, Benha Univ., Benha. Engelbrecht, D., The development of the chondrocranium of Pyromelana orix. Acta Zool. 39, Mokhtar, F.M., Studies on the Development of the Avian Chondrocranium (order Coraciiformes), Ph.D. Thesis, Cairo Univ., Cairo. Mokhtar, F.M., Zaher, M., Hammouda, H.G., The development of the pin-tailed sandgrouse Pterocles alchata caudacutus (order Columbiformes). 1 the development of the early stages of the neurocranium. Bull. Fac. Sci.: Cairo Univ. 50, Mu ller, H.J., Die Morphologie and Entwicklung des Craniums von Rhea Americana Linné. I. Das knorpelige Neurocranium. Z. Wiss. Zool. 165 (3 4), Macke, Til., Die Entwicklung des Craniums von Fulica atra L. Morphol. Jahrb. 113 (2), Zaher, M.M., Abou-Tira, A.M., ABD El-Kader, I.Y., Abd El-Hady, S.I., Developmental studies on the neurocranium of Corvus corone sardonius (Hooded crow, Mediterranean race), order Passeriformes. J. Egypt. Ger. Soc. Zool. 11B, Zaher, M.M., Riad, A.M., Studies on the ontogeny of Sterptopelia senegalensis aegyptiaca, 1. Description of three early developmental stages. Emrpiokoriz <rn 18 (Bbg. 1 2), Zaher, M.M., Riad, A.M., 2012a. Developmental studies on the neurocranium of Streptopelia senegalensis Aegyptiaca (Latham, 1790), 2. Description of three intermediate developmental stage. J. Egypt. Ger. Soc. Zool. 64B, 47 71, Comparative, Anatomy & Embryology. Zaher, M.M., Riad, A.M., Studies on the ontogeny of streptopelia senegalensis aegyptiaca (latham, 1790), 3. Description of the optimum stage of the chondrocarnium. J. Egypt. Ger. Soc. Zool. vol. 64B, Comparative Anatomy & Embryology, in press. Zaher, M.M., Riad, A.M., Hassan, F.M., The development of the chondrocranium of Passer domesticus niloticus. (order Passeriformes. 1 the development of the early stages of the neurocranium. J. Egypt. Ger. Soc. Zool. 3,

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