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Vol. 30, No. 1, pp. 71-82, 1985
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EVOLUTIONARY TRENDS OF THE HISTOLOGICAL
PATTERN IN THE TEETH OF EDENTATA (XENARTHRA)
J. FERIGOLO
Department of Palaeontology and Stratigraphy, Federal University of Rio Grande do Sul and Museum
of Natural Sciences, Zoobotanical Foundation, Dr Salvador Franpa, 1427, 90.000-Porto Alegre, RS,
Brazil
Summary-In a comparative study of the dental structure of Edentata, the central tissue was identified
as a modified orthodentine, except in the Glyptodontidae where an osteodentine was found. Some
evolutionary trends of the tissues in these teeth may have been related with the extinction of ground sloths.
Comparative studies of the tissues in mammalian teeth seem to be good tests for systematics.

Cabassous unicinctus (MCN-MA 951)
Tolypeutes matacus (MCN-MA 962)
Dasypus novemcinctus (MCN-MA 954,955,956)
Dasypus hybridus (MCN-MA 957)

INTRODUCTION

The structure
of the teeth of the Edentata
(Xenarthra) has been studied several times since the
pioneer work of Retzius (1837), but mostly or exclusively in respect of the presence or absence of enamel
(e.g. Tomes, 1874; Pouchet and Chabry, 1884; R&e,
1892; Ballowitz, 1892; Spurgin, 1904; Martin, 1916).
Although some works deal with the different kinds of
dentine with which I am here concerned, no one has
attempted a general comparative study. For edentate
teeth, Owen’s Odontogruphy (1840-45) is still the
basic reference. More recent findings in polarized
light are available, for armadillos, in Keil and
Venema (1963) and, for tree sloths, in Schmidt and
Keil (1971).
As concerns enamel proper, I have not detected
it in unabraded teeth of Dasypus novemcinctus,
Euphractus sexcinctus and Tolypeutes matacus, nor
in those of a young Bradypus sp. but, as an enamel
organ exists in the Edentata in general, both the
absence of enamel or its presence as a thin layer
indicate a regressive (Simpson, 1932) or an autapomorphic character (McKenna, 1975).

Fam.: Glyptodontidae
Glyptodon? sp. (MCN-PV OO6*)
Fam.: Megatheriidae
Megatherium sp. (MCN-PV 004*)
Fam.: Mylodontidae
Lestodon sp. (MCN-PV 003*)
Scelidodon sp. (MCN-PV 005*)
Fam.: Bradypodidae
Bradypus inficatus (MCN-MA 964)
Bradypus sp. (MCN-MA 965,966,967,968,969)
Choloepus didactylus (MCN-MA 970)
Choloepus sp. (MCN-MA 971, 972, 973, 974,
975)
Scaeopus torquatus (MCN-MA 976, 977, 978)
All fossil specimens, marked *, came from Tarija
(Bolivia, Pleistocene); the recent ones came from
South or Central America. Histological nomenclature
follows in general the works by Mrvig (1951), Schmidt
and Keil (1958, 1971), Bradford (1967) and Boyde
(1971) except for the modified orthodentine.

MATERIALS AND METHODS
FINDINGS

Teeth of extant and extinct Edentata were studied
in normal and polarized light, as well as in the
scanning electron microscope. For light microscopy,
ground sections were prepared in the usual way. For
the SEM studies, ground sections and fractured
surfaces were used. All the sections, as well as some
of the fractured surfaces, were sometimes treated with
2.5-5.0 per cent HCl for 3O-60min, before being
covered with gold-palladium
and examined in a
JEOL (C 35) SEM at 15 kV.
List of species examined (MCN, Museu de Ciincias Naturais):

Dasypodidae (armadillos)
D. novemcinctus, D. hybridus. In Dasypus, the teeth
consisted of cementum,
orthodentine and modified
orthodentine
(Plate Fig. 1). The cementum was thick

and cellular, in some teeth composed of concentric
layers with enclosed cementocyte lacunae. These
lacunae were usually more abundant in the layer
adjoining the orthodentine, where they were more
rounded and had fewer canaliculi than elsewhere. In
the outer layers, they were usually more fusiform with
the canaliculi directed mainly towards the external
surface of the tooth. In some teeth, there was also a
thin layer of cementum with few or no lacunae. The
cementum became thinner towards the intra-alveolar
part of the tooth, but a reduction in its thickness
towards the occlusal surface as described by Keil and
Venema (1963) could not be observed. Internal to the
cementum there was a hard tissue similar to other
mammalian orthodentines. The central portion of the

Order: Edentata (Xenarthra)
Fam.: Dasypodidae
Chaetophractus villosus (MCN-MA 953)
Chaetophractus vellerosus (MCN-MA 952)
Euphractus sexcinctus (MCN-MA 958, 959, 960;
MCN-PV 007*)
Zuedyus pichiy (MCN-MA 963)
Priodontes giganteus (MCN-MA 961)
71

In the direction of the intra-alveolar part. this tissue presented less dentinal tubules and some few vascular canals which increased in number in the same direction. continuous with those of the orthodentine. some special features were found. In the modified orthodentine. Mylodontidae and Megatheriidae (ground sloths) Scelidodon sp. it gradually became thicker and cellular. and vascular canals as in the modified orthodentine (Plate Fig. vellerosus. villosus. Euphractus and Tolypeutes). or in the lateral part of the teeth towards the external one. The central tissue. the cementum was similar. The cementum (thin. closer to the centre. The included vascular canals had a hair-pin shape but no direct communication with the vessels of the pulp cavity. Between the central modified orthodentine and the external orthodentine. In Megatherium it was reduced to a thin layer interposed between cementum and modified orthodentine (cf. Between the external orthodentine and the internal modified orthodentine. giganteus. there were the same tissues as in other Dasypodinae. The thick cementum was formed as appositional layers in Scelidodon. The orthodentine was also similar to that of other mammals. the vascular canals were regularly distributed. 4. their convexity being directed towards the occlusal surface. though there were some differences. towards the intraalveolar part. Between the external orthodentine and the internal modified orthodentine. It was not possible to identify this tissue in Scelidodon (Fig. but was more strongly developed in Lestodon (Plate Fig. P. at the centre of the crown but were bent slightly more outwards laterally. E. 1938. Bradypodidae (tree sloths) B. but more peripherally they diverged slightly towards the external surface (Plate Fig. In Scelidodon. S. Choleopus sp. an intermediary tissue corresponding again to the orthovasodentine of Arsuffi (1938). 9). It showed a larger number of included vascular canals than that of other Dasypodinae. novemcinctus. Bradypus sp. The central axis of the tooth had the same slightlymodified orthodentine without vascular canals as was observed in adult and young specimens of Euphractus. however.. C. the vascular canals (Plate Figs 9-12) were more numerous and longer than in the homologous tissue of the tree sloths. As evidence of immaturity. matacus (still having conical teeth) was studied. it was a highly-developed vasocementum presenting a complex pattern of vascular canals. The cementum was rather thick but acellular (Plate Fig. The poorly-mineralized dentine present in immature Zaedyus. C. the cementum was sometimes thin. Chaetophractus and Euphractus was not found in young specimens of armadillos (D. torquatus. T.72 J. FERIGOLO tooth was formed by a modified orthodentine with a different birefringence and fewer dentinal tubules than the adjacent orthodentine. the central tissue was a poorly-mineralized dentine. was suggestive of that of Dasypus. in all these armadillos the skull bones were still unfused. The cementum was thin and acellular. as shown by the polarizing microscopy. but not in the adult stage. there was an intermediary tissue. this tissue seemed to retain its thickness. injiiscatus. sexcinctus. and in some cases the central dentine was almost amorphous with only a few vascular canals or none. In the specimens studied. Megatherium sp.. it was not possible to be positive about the adult tooth structure. pichiy. In the teeth of a young Bradypus sp. In the modified orthodentine. for Bradypus tridactylus). The cementum was thin and acellular in the cusp tip but. there was another tissue showing both dentinal tubules. 12) and Lestodon. Young specimen of Bradypus sp. pichiy. Although in the teeth of Choloepus the structure was usually similar to that in Bradypus and Scaeopus. the most striking feature was the presence of a great number of vascular canals. plate 83). didactylus. Only one young specimen of T. the teeth had a similar structure (with minor irrelevant differences). being specially numerous near the long axis of teeth (as described by Arsuffi. matacus. just below the orthodentine of the cusp tip there were irregular dentinal tubules (Plate Fig. the tissues were exactly the same as those in the recent adult specimen. 8) was of similar structure. unicinctus. C. In the teeth of these species. The dentinal tubules were directed axially. but were also peculiar in some respects compared with other Edentata. Lestodon sp. Near the limits of the pulp chamber it again became thiner but was still cellular. The vascular canals had no apparent connection with the pulp cavity vessels. acellular) and the orthodentine were similar to those of the armadillos just described. there was in Megatherium (Fig. They were more or less regularly disposed. Although thicker than in Dasypus. Because the specimens of Zaedyus and Chaetophractus were immature ones. Sometimes a layer of acellular cementum covered the cellular one. C. vellerosus. . the tissues were similar to those in other armadillos. and possibly in Lestodon. but had only a few vascular canals. 7) and Megatherium. and adult specimens of Cabassous as well. 5). The orthodentine was similar to that in the teeth of the adult specimens. the modified orthodentine being. They were always parallel to the adjacent dentinal tubules. 2). and that of Tolypeutes was young. had few dentinal tubules or none at all. 6) but no vascular canals. They were formed mainly by tissues homologous with those of the Dasypodinae. In the teeth of these ground sloths. In the modified orthodentine. C. highly developed. The tissues in the teeth of these armadillos were similar to those in the others.. 3). lying in a homogeneous mineralized matrix which. similar to that found in the adult tree sloths (Arsuffi’s orthovasodentine). in some cases containing a few vascular canals (Plate Fig. In these sloths. The slightly modified orthodentine was like that of the other Dasypodinae. Z. In immature Z. Owen. nevertheless. a similar tissue was found. orthovasodentine of Arsuffi. In this species. In the teeth of an adult specimen of Euphractus from Tarija. In all specimens. but there were also differences. the tissues closely resembled those in the tree sloths. except that it had no vascular canals. parallel with the longitudinal axis of the tooth in the central part. Particularly in Megatherium. In one immature specimen of Euphractus. the orthodentine (Plate Fig. 1938). (skull 48 mm long). a slightly modified orthodentine. 1840-1845. except for those that were close to the pulp. villosus and C.

however. 1971). contrary to what was maintained by Retzius and Owen. The central tissue of the teeth of Zaedyus. of different stages of a modified orthodentine agreed with the opinion of Schmidt (1924) on the origin of this tissue in B. but not completely with those of Owen (184&l 845) dealing with armadillos in general. which can be seen by direct inspection. vellerosus and of the immature specimen of Euphractus. surrounded by concentric incremental lines. forming the prominent ridges that have been used for systematic purposes. Although it is not clear from his description. there were in the sloths all degrees of transition from a typical orthodentine to a vasodentine (i. 1884). Keil and Venema (1963) using microhardness tests found that the modified orthodentine is softer than the surrounding orthodentine. this interpretation is more accurate than that of Arsuffi. In his classification of dentines. but could only study the walls of the vessels near the pulp chamber. In both armadillos and tree sloths. 758). p. villosus.. Orvig (1951) included the “orthovasodentine” of Arsuffi (1938) among the orthodentines. however. The central tissue. With regard to the armadillos and sloths (with the possible exception of Orophodontoidea). there are two main reasons for considering the central tissue of the teeth of the tree and ground sloths as modified orthodentine. The structure in Megatherium and Scelidodon substantially agreed with the descriptions by Owen (1840-1845 pp. not all armadillos have the same structure nor a “hard substance. the part played by the tissues in the ridges of the teeth of armadillos is not clear. the tissue considered by Owen (184Gl845. see also his plates 83 and 84) for Megatherium and Scelidotherium. The description given by Owen (184&l 845) for a supposed young Bradypus disagrees in several points with what was found in the specimen here studied. 1938. Thus. findings usually used to characterize the vasodentines. In the intermediary tissue in the tree sloths. According to him. 324) supposed. seems to be actually an oblique section of an already worn tooth (as already pointed out by Pouchet and Chabry. and the lack of the occlusal part of the tooth (absence of cementum and orthodentine). The orthodentine showed the usual features. connective tissue and what she called pulpareticulum cells. the dentinal tubules persisted in adult growing teeth together with vascular canals. which can be seen by the presence both of vascular canals in the modified orthodentine and of cellular cementum (see Owen plate 85). no OB3”*1t 13 direct connection was found between the vascular canals and the dentinal tubules (as also pointed out by Schmidt and Arsuffi). p. Contrary to what was said by Owen (p. closely resembling bone” (see also his plate 85) and by Arsuffi (1938) as a “kanalisiertes Dentin”. was recognized as independent of the surrounding orthodentine by Keil and Venema (1963) using mainly polarized light. Arsuffi also described inside these vascular canals some odontoblasts. In the intermediary tissue of the teeth of a young Bradypus sp. the odontoblasts should during dentine formation achieve increasing contact with the tissues of the pulp. because in the same specimen some teeth have a ridge including all tissues and others have no ridge at all. Concerning Euphractus and Cabassous. because the “vasodentines” and the “orthovasodentines” are only different stages of a modified orthodentine. It is also possible to interpret such tissue as the most peripheral part of the central modified orthodentine. and. The structure of the teeth of tree sloths in general agreed with the descriptions by Retzius (1837) Owen (184&1845). but the main point here is not what name should be applied to this tissue. The absence of odontoblast processes is secondary. (b) an intermediary tissue found between orthodentine and modified orthodentine in the tree sloths and some ground sloths (“Ortho-Vasodentin” of Arsuffi. . The interpretation in sloths. corresponding to the slightly-modified orthodentine of the adult ones. homologous with that of armadillos: (a) a slightly modified orthodentine in the central portion of the teeth of a young Bradypus sp. the dentinal tubules were still present (Fig. 338. 6). Here I found striking differences from other Edentata. modified orthodentine). As far as I can see. but to recognize it as intermediary between the typical orthodentine and the modified orthodentine. was not a modified orthodentine but contained a great number of well developed vascular canals. these processes are present at the earlier stages of development of the vasodentine but disappear more or less completely at later stages”. the real nature of cementum was uncertain. the specimen figured on his plate 82. is a poorly-mineralized dentine which seems to be formed in the growing teeth of immature specimens. except that. before full mineralization takes place. for. finally. DISCUSSION ModiJed orthodentine In armadillos. As far as I can see. Owen probably described the teeth of Dasypus as if they were of Euphractus. C. As shown by dentinal-tubules orientation in the superior portion. tridactylus. and has its cavity lined by a layer of unvascular dentine”. 342-344. Contrary to what Owen (p. which is covered by cement. probably due to degeneration of tissues after losing communication with the pulp vessels. As the available specimen was not well preserved. the findings agreed with the observations of Retzius (1837) and Keil and Venema (1963). It was not possible to find any lacunae for cells (osteocytes) nor dentinal tubules in these layers (Plate Figs 13 and 14). 342) as a “hard substance. 343) said “It is true that vasodentine differs from ordinary orthodentine in that it is without dentinal tubes and contains numerous capillaries. closely resembling bone” in the central part of the teeth.e. As Mrvig (p. the teeth of young Bradypus were not formed “chiefly of the hard dentine. the . Schmidt (1924) Arsuffi (1938) and Schmidt and Keil (1958. C. 329). the ridges seem to depend more on the direction of the masticatory movements and on the position of teeth in occlusion than on the arrangement of tissues. and some ground sloths..Histological pattern in the teeth of Edentata Glyptodontidae Glyptodon? sp. regarded here as being intermediary between typical orthodentine and the modified orthodentine of the adult specimens.

5). Furthermore in ground sloths there was the highest degree of development of the modified orthodentine with the greatest number of vascular canals . Although Arsuffi (1938) had given a detailed description of the modified orthodentine (her “vasodentine” or “calcified pulp”). The possibility of a synapomorphy between Glyptodontidae and Orophodontidae should be considered. these would be the only groups of Edentata (with the possible exception of the Pampatheriinae) that share this character. as suggested by Ameghino (1920) Castellanos (1937) James (1957) and Paula Couto (1980). as far as I can see her interpretation is not entirely correct. The continuously-growing and abrasion processes should be considered as normal ones in the teeth of the Edentata [as already pointed out by Arsuffi (1938) for B. thinking that it is a kind of mineralized pulp formed to avoid the pulp to be exposed in the mouth. For instance. Pampatheriinae. 325) gave a full description of dental tissues in Glyptodon and other Edentata. a welldeveloped modified orthodentine was found. with a even greater number of vascular canals. Owen did not show any lacunae for cells (osteocytes) between these layers. which could be another important character shared with the Glyptodontidae. He went on to say that “The medullary canals are surrounded by fine compact concentric strata. This opinion seems to be corroborated by the facts above. In my view. but frequent enough in lower vertebrates (see e. because. 325. as far as I know. the name vasodentine is not appropriate mainly because it has been used frequently to describe several different varieties of orthodentine (lilrvig. it would be more appropriate to classify the pampatheres as an independent family. The interpretation of the modified orthodentme as resulting from the fast growing and abrasion processes is not appropriate mainly because a portion of a fast growing and typical orthodentine was still preserved in all teeth examined. p. . mainly because in the young Bradypus sp. 190). was found in the tree sloths. 1976). He described the teeth of Glyptodon as having their central portion formed by “vascular osseous texture”. Owen observed in these teeth a tissue similar to. Glyptodontidae and Orophodontidae. triductvlus]. Sasso and Della Serra (1965) were the only authors who interpretated the modified orthodentine (of Brndypus tridactylus) as a variety of orthodentine (their “vascularized orthodentine”).74 J. tree and ground sloths. in the specimen discussed here. uncommon in mammalian teeth. Because of this. Palaeopeltis is supposed to have had an osseous dermal armour. in the abrasion process. 1967. Osteodentine Concerning the histological pattern of the glyptodontid. vascular dentine (probably osteodentine). included in the dentine. the structures in Glyptodon described both by Owen and here (Glyptodon?) could be interpreted as bone (osteons) or as dentine (denteons). he did not realize the fundamental difference between osteodentine and modified orthodentine of the teeth of Glyptodon and armadillos: “In the Glyptodon the vascular osseous texture occupies a larger proportion. 1) was thick and cellular and the modified orthodentine showed the largest number of vascular canals in the armadillos studied. Nevertheless. This point could be illuminated by a comparative study of teeth tissues including the Dasypodinae. it should be more appropriate to interpret it as belonging to some Glyptodontidae incertae sedis as did Simpson (1945. is not quite correct. The absence or scarcity of dentinal tubules he supposed to be due to reduction. p. have in the central part of their teeth a “. see also his plate 86) except for certain points. The abrasion process was here in its beginning. The presence of such supposed dentinal tubules suggests an interpretation of this tissue as a typical osteodentine. If Pakzeopeftis actually has an osteodentine in its teeth. 75) or as an aberrant glyptodontid as did Paula Couto (1979. Cementum The degree of development of cementum may not be independent of the formation of the modified orthodentine but closely related to the development of orthodentine as a compensatory mechanism for abrasion.g. If this is correct. In armadillos. 595) pointed out that the Orophodontoidea. discussed here. Nevertheless. one should deduce that both the Glyptodontidae and the Orophodontoidea must have osteodentine in their teeth. ‘*. and once more a greater apposition of cellular cementum (Fig. Hoffstetter (1958. a greater apposition of cementum was found in species where there was a greater development of the modified orthodentine or a trend towards an increase in the vascular canals in that tissue. In accord with this interpretation. Although Owen (p. p. p. a significant feature is that in plate 86 of Owen’s Odontogruphy. I do not agree. which probably correspond to dentinal tubules. a still greater development of modified orthodentine. considered by him as a lateral branch in the evolution of the ground sloths. 1951. 1967). but are wider than in true bone”. or the same as. radially disposed. Modified orthodentine has a lower degree of resistance to wear than the adjacent orthodentine. Orvig. the features agreed with Owen (184&l 845. FERIGOLQ odontoblasts surpass the vascular loops (of the pulp) that are in this way. One possible representative of the Orophodontoidea. Similarly. considering this and other characters. this tissue is clearly different from the modified orthodentine of other Edentata. if his assumption in what concerns the Pampatheriinae is correct (considering only this character) this group could be more closely related to the Glyptodontidae (and Orophodontidae) than we have so far believed. than in the small Armadillos”. the concentric layers of hard tissue surrounding the vascular canals are traversed by thin lines. their intimate texture and composition are essentially the same”. the assumption of Hoffstetter (1958) that the structure of the teeth of the Orophodontoidea is similar to that of the Cingulata. Forming prominent ridges in the teeth. 1963). because in the Dasypodinae here studied the central tissue of the teeth consisted of a modified orthodentine and not of an osteodentine. This structure recalls that in the majority of Cingulata especially that of Pampatherines” (my translation). 1951. On the other hand. as shown by microhardness tests (Keil and Venema. the cementum in Dasypus (Fig.

by (Text Fig. The osteodentine of the Glyptodontidae is different from the modified orthodentine of other Edentata in its strut. (c) an even greater development of modified orthodentine (and of vascular canals) as in the tree sloths. .Histological pattern in the teeth of Edentata 9) and the heaviest apposition of cementurn (Fig. the structure of the teeth suggests two possibilities namely they may have evolved by one or by two evolutionary trends. If the teeth of an armadillo like Euphractus or are regarded as the primitive form. 75 and a reduction in the thickness of cementum (as in Euphractus and Cabassous). it is possible to see a unique trend for the derivation of the dental structure of other armadillos and sloths as well (Text Fig. a still greater development of these tissues. (Fig. 16): (a) an inclusion of a few vascular canals in the modified orthodentine. (C) by Euphractus. Scaeopus and Choloepus). although some speculations are possible. Considering armadillos. 15): (a) a reduction and disappearance of vascular canals in modified orthodentine. Evolutionary trends of the times Phylogenetic reconstructions based on the analysis of a few characters cannot obviously be made. Diagram of the first hypothesis. and a development of a slightly-thicker acellular cementum (as in Priodonres). cementum seemed to be thinner than in the sloths. ture. 15. A stasigenetic stage could be represented by Dasypus (and in a lesser degree by Priodontes). (A) represented by Dosypus. and has probably evolved in relation to the Orophodontidae and Pampatheriinae. 7). tree and ground sloths. the most extreme example being in the ground Cabassous Fig. all the teeth of the armadillos and sloths could easily derive from it. in ground sloths (as Lestodon. Not in scale. Scelidodon and mainly Megatheritun). (E) by Megatherium. In the glyptodontids in which there is an osteodentine which is highly resistant to abrasion. (b) the non-adaptive greater inclusion of vascular canals in modified orthodentine and a still greater deposition of cellular cementurn (as in Dasypus). (D) by the tree sloths. If the teeth of an armadillo like Dasypus is the primitive form. (B) by Priodontes. Interpret these extreme degrees of development of cementurn as physiological hypercementosis. (b) a non-adaptive trend towards greater development of modified orthodentine (with an increased development of vascular canals) and of cementum (as in Bradypus. if Hoffstetter (1958) is correct.

Johnels and Dr B.-O.g. FERIGOLO sloths (like Lestodon. Diagram of the second hypothesis. Section of Palaeozoology. The vasocementum. Stolt. Acknowledgements-l am grateful Fig. Scelidodon. and Dr C. (D) by the tree sloths. The development. orthodentine and slightly-modified orthodentine) is closer to that of most mammals. 16. for providing me material. (E) by Megatherizun. of a failed experiment. (C) by Dasypus. would have been affected by natural selection. i. it is logical to assume the development of the modified orthodentine having taken place at the expense of the orthodentine itself. a faster rate of growth. to the technical staff. A stasigenetic stage could be represented by Euphractus and Cabassous. Brazil. Nevertheless. as seen in Megatherium. is noteworthy as is the thick cementum in these forms. Section of Vertebrate Zoology (SMNH). 1 should also express my gratitude to Dr A. except by forming a larger occlusal surface. in others there may not only have been greater development of modified orthodentine but at the same time a reduction in the thickness of the orthodentine itself. If in some ground sloths the relative thickness of the orthodentine (in comparison with other tissues) seems to be preserved (e. the vasocementum seems no more effective in compensating wear: apparently it did not increase resistance to wear of the teeth. was as easily abraded as the modified orthodentine. A trend towards the reduction of orthodentine and a large development of modified orthodentine (an extreme degree of which could be represented by Mega therium). Anderson for the artwork: Mr B. the second hypothesis seems the most appropriate one. such a trend could have remained unaffected by natural selection for a long time. (B) by Priodontes. Blom for his kind heln with the SEM: Mr L. Dental histology and the extinction of some ground sloths The development in the Edentata of a modified orthodentine. de Paula Couto. In the latter representatives of animals like Megatherium. and specially Megatherium). Section of Entomology (SMNH). Associated climatic/floral changes in Pleistocene may also have contributed. for example. by still greater apposition of the cementum. the condition in Euphractus and Cabassous should be relatively plesiomorphic. G. (A) represented by Euphractus. in continuously-growing teeth.Not in scale. in Megatherium. which has low resistance to abrasion. Federal University of Rio Grande do Sul. In this case. As the modified orthodentine seems to have evolved from a typical orthodentine. Mr U. Swedish Museum of Natural History (SMNH). in some ground sloths. If in this trend a greater wear of the teeth could be physiologically compensated in some way. bearing in mind the importance of mastication and the consequent wear of the teeth in these huge herbivorous animals. . Samuelson for photographic copies. Srelidodon). The orthodentine formed only prominent thin ridges interposed between these tissues. of a tissue less resistant to wear could have been a dangerous trend in the evolution of some ground sloths and could have contributed to their extinction.e. and in-particular to: Mr G. Dr L. for example. we could have had the deleterious grade of a non-adaptive trend.J. Brundin. ‘Bergman for the Plates. As the histological pattern of the teeth of armadillos such as Euphractus and Cabassous (thin acellular cementum.

Academic Press. 3-34. 535-636. (I 967) Microanatomy and histochemistry of dentine. for the kind interest he has taken in this research. siideles Tandbenets struktur. and Keil A. 3. W. J. Am. McKenna M. Morph. In: Dental Morphology and Evolution (Edited by Dahlberg A. Am. 31. Bradford E. An. 27. Martin B. (1958) Xenarthra. 157-164. (1963) Struktur und Mikrohlrteuntersuchungen an Zlhnen von Giirteltieren (Xenarthra).). Castellanos A. Hist. Anat. Tomes C. Zool.) Vol. E. 2001. 1874. (1840-1845) Odontography. l-88. Mus. Orvig T. 40. Retzius A. 149-192. Paleont. 495-512. mikrosk. Boyde A. J. for financial support. and to the CNPq. W. 7. (1924) Ueber das Dentin von Bra&pus triaisctylus. organ in the armadillo. Acad. Anat. Arch. EntwGesch. Schmidt W. where this work was carried out. I. Carl Hanser. James G. 2. . G. und die Persistenz seines Keimrandes bei dem erwachsenen Thier. ArsulIi E. 527-531. (1951) Histologic studies of placoderms and fossil I . (1884) Contribution a l’odontologie des Matnmiferes. pp. 796808. EntwMech. Orvig T. Section of Palaeozoology (SMNH). (1957) An edentate from the Pleistocene of Texas. Am. (1920) Cobras(Edited by Torcelli A. brasil. 44-48. Academic Press. 149-160. Baillibre. 20. J. Fisiogr. Rose C. and Venema B. (1937) Anotaciones sobre la linea filogenetica de 10s Clamiterios. Zool. (1967) Phylogeny of tooth tissues: Evolution of some calcified tissues in early vertebrates. and Della Serra 0. Paula Couto C. In: Traitk de Paleontologie (Edited by Piveteau J. T. In: Structural and Chemical Organization -of Teeth (Edited by Miles A. for the fossil specimens. Owen R. Ballowitz E. S. Stockholm. Bull. and Darling A. 1: The endoskeleton with remarks on the hard tissues of lower vertebrates in general Ark.) Vol. La Plata. Brazil. Cienc. J. (1971) Comparative histology of mammalian teeth. J. Jl microsc.) Vol. Schmidt W. 2. Pergamon Press. 79-96. 5. University of Chicago Press. S. 45-l 10. In: Structural and Chemical Organization of Teeth (Edited by Miles A. 11. A. Am. Q. Anat. (1916) Tooth development in Dasypus nouemcinctus. Revta bras. 75-84. da S. DP. (1975)Toward a phylogenetic classification of the Mammalia. New York. Fe 8.) Vol. M. Publs Inst. Keil A. Sci. Paula Couto C. 58. Mus. Sasso W. P. F. Braunscheig. W. Physiol. ‘J. G. Anat. 81-94. Director. Anat. New York. (1892) Beitrlge zur Zahnentwickelung der Edentaten. and Chabry L. Finally. Beitr. I want to express my great indebtness to Dr Tor 0rvig. overleaf. C. 567. (1976) Paiaeohistological notes: 4:’ The interpretation of osteodentine with remarks on the dentition in the devonian diphoan Griphognathus. London. de (1980) Urn tatu gigante do Pleistocene de Santa Catarina. 2. Paris. pp. 1-4. E. Schmidt W. 133-156. nat. Miinchen. Masson. (1837) Mikroskopiska undersiikingar iifver Tandemes. 2. Orvig T. 647-691. 85. Anat. La Plata Taller impressiones Oficiales. Ser. Mammalial. 173-195. 6. and Szalav F.Histological pattern in the teeth of Edentata for his constructive criticism and advice. (1958) Die gesunden und die erkrankten Zahngewebe des Menschen und der Wirbeltiere im Polarisationmikroskop. In: Phylogeny of the Primates (Edited bv Luckett W. 2146. Pouchet G. (1945) The principles of classification and a classification of mammals. 108. 1.. l-350. REFERENCES Ameghino F. E.). (1874) On the existence of an enamel. Hoffstetter R. S. Novit. Biol. Plenum Press. I wish to thank the other authorities of the Swedish Museum of Natural History (Naturhistoriska riksmuseet). 52. Geol. Academia Brasileira de Citncias. and above all for his friendship and kindness when I was in Stockholm during the winter 1981/82. (1965) Observa9bes sobre as estruturas de dentes de xenartros pertencentes aos &erOS Daypus. Plates 14 Spurgin A. 9. 77 elasmobranchs. l-35. Simpson G. Rio de Janeiro.) DD. (1932) Enamel on the teeth of an Eocene edentate. J. Simpson G. seine Ausbildung im Embryo. Euphractus e Bradypus (Edentata. New York. 25. J. (1904) Enamel in the teeth of an embryo edentate (Dasypus novemcinctus Linn. and Keil A. KVA Handl. . (1892) Das Schmelzorgan der Edentaten. Anr. (1971) Polarizing Microscopy of Dental Tissues (Translated by Poole D. 97-107. Chicago. for stylistic corrections. 1836. de (1979) Tratado de Paleomastozoologia. 321-454.) pp. (1938) Beitrage zur Kenntnis des Vasodentins.

note dentinal tubules and vascular canals (arrow) in (a) MCN-PV 004. showing natural cast of vascular canal (arrow) in modified orthodentine. MCN-MA 965. x 2060 Fig. SEM of etched tooth fragment showing modified orthodentine (arrow). modified orthodentine not shown. SEM of etched tooth fragment of Brudypus sp. note absence of an intermediary tissue between (a) and orthodentine (b). x 460 Plate 4. SEM of etched transverse section of a tooth of Glyplodon ? sp. orthodentine (b) and modified orthodentine (c). note dentinal tubules and vascular canals (arrows) in (a). x 37 Fig. SEM of longitudinally-fractured tooth of P. MCN-MA 963. 5. x 930 of Z. 4. Transverse thin section of a tooth of Gfypfodon ? sp. MCN-MA 968. MCN-PV 006. x 110 (a) and . SEM of etched transverse section of a tooth of Scelidodon sp. 3. Longitudinal thin section of a tooth of Bradypus sp. Fig. Longitudinal thin section of a tooth of D. 12. x 124 Fig. showing thick cellular cementurn (a). x200 Plate 2. Transverse thin section of a tooth of Megurherium sp. showing orthodentine osteodentine (b). SEM of etched transverse section of a tooth of Lesfodon sp. SEM of etched transverse section of a tooth of Lestodon sp. Fig. note concentrical layers in denteons (arrow). nmemcinctus. SEM of etched transverse section of a tooth of Megatherium sp. 10. showing natural casts of vascular canals (arrow) in modified orthodentine.78 J. 2. 7. showing vascular canals (arrows) in cementum. 14. 6. showing vascular canals (arrow) in modified orthodentine (a). MCN-PV 005. MCN-MA 966. MCN-PV 003. gigunfeus showing orthodentine cementurn (b). orthodentine (b) and vascular canals (arrow) in the modified orthodentine (c). x 100 tubules Plate 3. 11. x 460 Fig. showing irregular dentinal (arrow) and vascular canals in modified orthodentine (a). showing natural casts of dentinal tubules. FERICKILO Plate 1. x 390 Fig. Transverse thin section of a tooth of Scelidodon sp. orthodentine (b) and modified orthodentine (c) with vascular canals (arrow). showing the intermediary tissue (a) and orthodentine (b). x 160 Fig. showing cellular cementum (a). Fig. Longitudinal thin section of a tooth of young Bradypus sp. 1. 8. showing orthodentine (a) and osteodentine (b). Fig. x 1780 Fig. MCN-MA 956. MCN-PV 005. x 158 Fig. 9. MCN-PV 003. MCN-PV 006. 13. showing the intermediary tissue (a). MCN-MA 961. x 54 Fig. MCN-PV 004. pi&y (a) and acellular with vascular canal Fig.

19 .Histological pattern in the teeth of Edentata Plate 1.

FERIGOLO Plate 2. .80 J.

Histological pattern in the teeth of Edentata J..i i--F-~ Plate 3. 81 .

82 J. FERIGOLO Plate 4. .