Anotosaura collaris Amaral 1933

Rodrigues, Miguel Trefaut, Jr, Mauro Teixeira, Vechio, Francisco Dal, Amaro, Renata Cecília, Nisa, Carolina, Guerrero, Agustín Camacho, Damasceno, Roberta, Roscito, Juliana Gusson, Sales Nunes, Pedro M. & Recoder, Renato Sousa, 2013, Rediscovery of the Earless Microteiid Lizard Anotosaura collaris Amaral, 1933 (Squamata: Gymnophthalmidae): A redescription complemented by osteological, hemipenial, molecular, karyological, physiological and ecological data, Zootaxa 3731 (3), pp. 345-370: 349-359

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Anotosaura collaris Amaral 1933


Anotosaura collaris Amaral 1933  

( Figs. 1–3A,B View FIGURE 1 View FIGURE 2 View FIGURE 3 , 4A View FIGURE 4 , 5 View FIGURE 5 )

Anotosaura collaris Amaral 1933: 69   ;

Anotosaura collaris   — Dixon 1973: 8;

Anotosaura collaris collaris   — Dixon 1974: 14;

Anotosaura collaris   — Vanzolini 1976: 120.

Holotype. MZUSP 788 View Materials . Type Locality: “ Villa Nova , Bahia, Brasil ”  

Additional material. 16 specimens (8 females, 8 males): MZUSP 103834 View Materials (field number MTR 24561 View Materials ), MZUSP 103836–103840 View Materials ( MTR 24563–24567), MZUSP 104210 View Materials , ( MTR 24568 View Materials ), MZUSP 103841–103844 View Materials ( MTR 24805–24808), MZUSP 103845 View Materials ( MTR 25010 View Materials ), MZUSP 103846 View Materials ( MTR 25022 View Materials ): Alto da Rainha: Senhor do Bonfim, Bahia (10°26'24.56"S, 40°10'32.19"W, 715m a.s.l.), 13–31 th December 2012. MZUSP 103835 View Materials ( MTR 24562 View Materials ): Missão do Sahy : Antonio Gonçalves, Bahia (10°31'3.66"S, 40°14'54.48"W, 895m a.s.l.), 13 th December 2012. MZUSP 103832 View Materials , 103833 View Materials ( MTR 24620 View Materials , 24621): Morro do Cruzeiro : Campo Formoso : Bahia (10°30'56.12"S, 40°18'31.51"W, 779m a.s.l.), 15 th December 2012. All specimens collected by the authors GoogleMaps   .

External morphology. Rostral broad, wider than high, contacting first supralabial, nasal and frontonasal. Frontonasal heptagonal, slightly wider than long, contacting rostral, nasal, loreal and prefrontals. Prefrontals hexagonal, as long as wide, in broad contact at midline. Frontal heptagonal, twice longer than wide; anterior margin angulose, indenting prefrontals, lateral margins almost parallel, slightly divergent posteriorly, contacting broadly the second and less the third supraocular; posterior margins diagonally contacting parietals and in straight contact with interparietal. Frontoparietals absent. Interparietal rectangular, longer than wide, as long as and narrower than frontal, narrower than parietals. Lateral margins of interparietal slightly concave. A pair of very large irregularly hexagonal parietals in straight lateral contact with interparietal, anteriorly contacting frontal and laterally in broad contact with third supraocular. External part of parietals slightly rounded, contacting two large temporal and two smaller occipital scales; in some specimens parietal contact sixth supralabial. Three supraoculars, first the smallest, second the largest, as large as prefrontal; first supraocular contacting only prefrontal, loreal, and second supraocular, separated by the frontal by the contact between second supraocular and frontal; third supraocular in broad contact with parietal and an enlarged postocular. Nasal above first and second supralabials, and also contacting loreal, frontonasal and rostral; large, longer than high, with nostril centrally placed in the lower part of scale. Loreal posterior to nasal, narrower and diagonally oriented; contacting frontonasal, prefrontal, first supraocular, first supraciliar, the frenocular, and second supralabial. Frenocular small, quadrangular, followed posteriorly by two to four suboculars, the second being the largest and located under the eye. Six supralabials, second and third subequal and smallest, fourth under the eye, sixth the highest and the longest, fifth supralabial separated from parietal by an enlarged postocular. Sixth supralabial followed posteriorly by a minute and elongate granule which contacts granules in the ear depression. Two to four superciliaries (usually two), the first being slightly longer, their suture indented by second supraocular. First superciliar contacting loreal, first and second supraoculars, frenocular, second superciliary, and an elongated, almost granular, preocular. Central part of eyelid with a single semitransparent disc surrounded by small, slightly pigmented, and irregularly shaped smooth granules. Lower eyelid with seven to nine strongly pigmented palpebrals. Temporal region with four large, smooth and juxtaposed scales between parietal, sixth supralabial, and the ear; two of them diagonally contacting the sixth labial. First temporal contacting parietal, second and third temporals, fifth and sixth supralabials and postocular. External auditory meatus absent, tympanum indistinct. Region of ear opening situated in the middle of a vertical incomplete fold covered by a series of small and juxtaposed granules extending to the gular region. Granules corresponding to the ear area contacting anteriorly two large temporals, and two much smaller, elongated, and juxtaposed flat granules. Lateral surface of neck with smooth scales and occasionally granules, irregular in size and shape, varying from juxtaposed to slightly imbricate, and arranged in regular transverse series between ear and shoulder. All head scales smooth and juxtaposed.

Mental broad, wider than high. Postmental heptagonal, wider than long. Three pairs of genials, all contacting infralabials; first the smallest, third the largest, first and second in broad contact at midline; third separated at midline by a small granule and deeply indented by a pair of flat, enlarged, and diagonally disposed scutes. Five infralabials, first the smallest, all others subequal. Gulars smooth, imbricate, quadrangular, laterally juxtaposed, irregular in size, in five regular transverse rows; third and fifth rows with the longest scales. Gulars followed by a distinct interbrachial region with eight larger and elongate scales. A distinct collar fold characterized by some granules and reduced scales in the second row of gulars preceding the interbrachial row, another one between interbrachials and the last row of gulars.

Dorsal scales disposed in regular transversal rows, anteriorly smooth, imbricate, rounded in the occipital region, becoming progressively narrower, more elongate and rectangular towards the arm level, being progressively hexagonal with lateral sides almost juxtaposed posteriorly. Twenty-eight or 29 transverse rows of dorsals between interparietal and the posterior level of the hindlimbs. Lateral scales about the same size as dorsals, rectangular, imbricate laterally, not acuminate and more diagonally oriented than dorsals; those closer to ventrals, the larger. An irregular series of transversally arranged scattered granules in the skin separates transverse series of lateral scales. A distinct area with granular scales surrounds the area of arm insertion. Twenty three to 25 scales around midbody. Ventral scales smooth, laterally juxtaposed, slightly imbricate antero-posteriorly, rectangular, and rounded posteriorly; first rows almost squared becoming gradually longer, about twice as long as wide; 17–19 transverse rows from interbrachials (excluded) to preanals. Five scales in precloacal region, the central one being the smallest. Males usually show four small inconspicuous precloacal pores and two small femoral pores. Pores absent in females.

Tail scales are shorter and more imbricate antero-posteriorly than midbody dorsals, otherwise identical to them, disposed in regular transverse circles, lanceolate; those from ventral part of tail base wider, becoming gradually identical in size around tail. Tail regenerated with rectangular, juxtaposed and smooth scales.

Forelimbs extremely robust with large, smooth and imbricate scales, larger and flat dorsally; those from ventral part of brachium smaller. Forearm as long as thick. Anterior and ventral parts of hind limbs with irregularly large, smooth and imbricate scales, largest scales ventrally. Posterior part of hindlimbs with granular and juxtaposed scales. Carpal and tarsal scales large, imbricate; supradigital lamellae smooth, imbricate. Palmar and plantar surfaces with smooth, small granules; infradigital lamellae mostly divided, 6–8 on Finger IV and 12–16 on Toe IV. Fingers and toes clawed, respectively with the following relative sizes: 1 <2 = 5 <4 <3 and, 1<5 <2 <3 <4.

Dorsal surfaces of body and tail light brown with an irregularly distributed dark brown reticulate pattern. Lateral parts of body and tail dark brown with an irregular light brown to cream reticulum. An irregular light line about one scale width extends dorsolaterally from posterior part of head to the middle of tail and becomes inconspicuous toward the tip. Ventral parts of body and tail cream but mottled by a scattered irregular dark brown pattern; tail become gradually darker distally. Dorsal parts of head with an irregular reticulate dark brown pattern. Lateral parts of head predominantly dark brown with scattered cream spots concentrated in the suture of labials; ventral parts of head lighter, cream, strongly mottled by an irregular dark brown pigmentation, especially in the external parts. Limbs dark brown, with scattered light brown marks, cream ventrally. Palmar and plantar surfaces light brown.

Maximum SVL: 44 mm (females); 41 mm (males). Tail length about 1.5 times SVL in adult males and females.

Hemipenial description. The left hemipenis of MZUSP 103845 ( Fig. 4A View FIGURE 4 ) is evidently bilobed, relatively small, extending along approximately five rows of subcaudals (approx. 3.5 mm). The hemipenial body is roughly Y-shape. The lobes are detached from the body by a marked basal constriction and are completely nude, presenting a round tip (one of the lobes is partially everted getting and artifactual sharp tip).

The sulcus spermaticus is a relatively broad channel originating at central region of the base of the organ and emarginated by distinctive thickened lips in the basal region. From the base the sulcus spermaticus proceeds in a straight line towards the lobular crotch. At the lobular crotch the sulcus is divided in two shallow branches that run through the internal faces of the lobes and end slightly before the lobular tips. The hemipenial body is ornamented by 14 transversal nude flounces that lack any vestige of mineralized spines or spinules, even after their immersion on Alizarin Red solution for 24 hours. The five more basal flounces are continuous and occupy all the central region of the asulcate face, whereas the remaining and more apical flounces are shorter and interrupted by a nude region in the sagittal region of asulcate face. In both lateral faces of the organ only the four more basal flounces are continuous, whereas the ten remaining are separated by a nude area that gets broader in direction of the distal region. There is a puncture in the asulcate face of the organ, in the region between the first and fifth transversal flounces, probably made accidentally during the hemipenial eversion or filling.

The hemipenis of Anotosaura collaris   is similar with the organ of A. vanzolinia   described and illustrated by Nunes (2011). However, the hemipenis of A. collaris   lacks the evident calcified and hook-shaped spines present in the 10 more apical flounces of the sulcate face and the vestiges of calcified structures present in some flounces of the asulcate face of the hemipenis of A. vanzolinia   ( Fig. 4B View FIGURE 4 ).

Osteology. ( Fig. 5 View FIGURE 5 ) The premaxilla is short, approximately as long as wide ( Fig. 5A View FIGURE 5 ). Its alveolar plate bears 10 unicuspid pleurodont teeth and contacts laterally the anteromedial process of the maxilla ( Fig. 5B View FIGURE 5 ). The short nasal process extends dorsally and overlaps the anterior end of the nasals.

The palatal shelf of the maxilla bears 10 teeth and is overlapped posteriorly by the maxillary process of the palatine. Its orbital process supports the maxillary process of the jugal. The facial process extends dorsally to meet the nasal, frontal, and prefrontal ( Fig. 5A View FIGURE 5 ).

The vomer ( Fig. 5C View FIGURE 5 ) has a well-developed dorsal crest that holds, together with the premaxilla and maxilla, the Jacobson's organ. It extends in an anterior process that rests on a pit in the palatal plate of the premaxilla. Posteriorly, it contacts the ventral surface of the palatine.

The dome-shaped septomaxilla, located between the vomer and the nasal process of the premaxilla, covers the Jacobson's organ dorsally and laterally. A lateral maxillary process contacts the dorsal surface of the palatal shelf of the maxilla. The nasal rests on the dorsal surface of the septomaxilla.

The palatine has a wider anterior portion composed of a medial vomerian process meeting the vomer and a lateral maxillary process contacting the palatal shelf of the maxilla; a pronounced concavity between these processes forms the choanal canal ( Fig. 5C View FIGURE 5 ). The palatine ends in a pterygoid process.

The large prefrontal ( Fig. 5A View FIGURE 5 ) forms the anterior margin of the orbit. It is overlapped by the facial process of the maxilla anteriorly, and extends posteriorly in a well-developed posterodorsal process, delimiting part of the dorsal margin of the orbit. A large lacrimal foramen and a short lacrimal flange are present. The lacrimal was not observed.

The anterior margin of the frontal overlaps the posterior end of the nasals. A distinct anterolateral process is observed laterally to the nasal. The posterior margin of the frontal bears two fronto-parietal tabs that overlap corresponding surfaces of the parietal ( Fig. 5A View FIGURE 5 ). A posterolateral process contacts the postorbitofrontal.

A stout postorbitofrontal is present ( Figs. 5A, B View FIGURE 5 ), probably corresponding to the fused postorbital and postfrontal. This element forms part of the posterodorsal margin of the orbit and, together with the squamosal, forms the lower margin of the upper temporal fenestra ( Fig. 5A View FIGURE 5 ). The posterior end of the squamosal fits into a notch in the tympanic crest of the quadrate.

The parietal meets the postorbitofrontal anterolaterally. Posteriorly, the parietal forms the anterior margin of the posttemporal fenestra, the posterior margin of which is delimited by the supraoccipital. The postparietal process, which extends posteriorly along the transverse ridge of the supraoccipital, meets the squamosal and supratemporal. A long and slender descending process of the parietal approaches the epipterygoid ( Fig. 5B View FIGURE 5 ).

The jugal forms the lower and part of the posterior margins of the orbit, contacting the maxilla and postorbitofrontal ( Fig. 5B View FIGURE 5 ). A large maxillary foramen is seen in its medial portion. The jugal bears a medially directed ectopterygoid process that contacts the ectopterygoid.

The large Y-shaped pterygoid ( Fig. 5C View FIGURE 5 ) contacts, at its anterior end, the palatine and the stout ectopterygoid. Medially it contacts the basipterygoid process from the parabasisphenoid and posteriorly, the quadrate. The rod-like epipterygoid has an expanded dorsal end, and its ventral end rests on the dorsal surface of the pterygoid ( Fig. 5B View FIGURE 5 ).

The quadrate ( Fig. 5B View FIGURE 5 ) has a cephalic condyle that contacts the supratemporal, squamosal, and the paroccipital process, a lateral conch with a slight tympanic crest, and a mandibular condyle that articulates with the mandible.

The suborbital fenestra is delimited by the maxilla, vomer, palatine, pterygoid, and ectopterygoid. The upper temporal fenestra is delimited by the postorbital, postfrontal, parietal, and squamosal.

The chondrocranium is composed of the supraoccipital, basioccipital, exoccipital, opisthotic, otooccipital, and parabasisphenoid. The supraoccipital closes the roof of the braincase and forms the dorsal margin of the foramen magnum. Laterally, it contacts the prootic-otoccipital complex. The posterior and horizontal semicircular canals are observed within this bone.

The prootic-otooccipital complex is located laterally, forming the otic capsule. The prootic is the anterior element and the otooccipital is the posterior one, and both delimit the fenestra ovalis, to which the columella is inserted ( Fig. 5C View FIGURE 5 ). The anterior semicircular canal is seen in the alar process ( Fig. 5B View FIGURE 5 ) of the anterior margin of the prootic. The incisura prootica, below the alar process, forms a C-shaped exit for the trigeminal nerve. The facial foramen is located posteriorly to the incisura prootica. The crista prootica extends ventrally from this latter foramen to the parasphenoid. The vestibular and lagenar cavities of the prootic are distinct in the inner surface of the prootic.

The otooccipital is composed of the fused opisthotic and exoccipital; the occipital recess and vagus foramen are delimited anteriorly by the opisthotic and posteriorly by the exoccipital. The anterodorsal margin of the opisthotic forms the well-developed paroccipital process. The exoccipital contributes to the occipital condyle. Three foramina for the hypoglossal nerve are present.

The ventral basioccipital, which forms most of the occipital condyle, is fused to the parabasisphenoid, located anteriorly ( Fig. 5C View FIGURE 5 ), and contacts the prootic and otooccipital laterally. The tetraradiate orbitosphenoid is connected to the interorbital cartilages.

The mandible ( Figs. 5B, C View FIGURE 5 ) is formed by the dentary, splenial, angular, surangular, coronoid, and prearticulararticular complex. The dentary bears 15 teeth. Its labial surface ( Fig. 5B View FIGURE 5 ) contacts the splenial and angular ventrally, the surangular posteriorly, and overlaps the labial process of the coronoid. The lingual surface of the dentary is overlapped by the splenial and by the anteromedial process of the coronoid. The coronoid bears a high dorsal process ( Fig. 5B View FIGURE 5 ), and a posteromedial process that overlaps the prearticular, forming the anterior margin of the adductor fossa. The surangular is located posteriorly on the labial surface of the mandible, forming the external margin of the adductor fossa. The prearticular-articular complex forms most of the posterior end of the mandible and the floor and posterior margin of the adductor fossa.

The hyoid apparatus ( Fig. 5C View FIGURE 5 ) is composed of a short basihyal, a glossohyal extending anteriorly, and three pairs of visceral arches: the hyoid cornu and the associated epihyal originating from its mid portion, and the first and second ceratobranchials. The specimen analyzed has 28 presacral vertebrae. Nine intercentra are present, associated with the vertebral centrum of the most anterior vertebrae. Ribs are present from the 4th vertebra on, and the last presacral vertebra does not bear ribs.

The scapular girdle ( Fig. 5D View FIGURE 5 ) is composed of a stout clavicle that bears a large foramen on its medial end, a rod-like interclavicle, the scapulocoracoid with four foramina, and the suprascapula. A sternum with a large central foramen, and a rod-like xiphisternum are present. The sternum is associated with two pairs of sternal ribs and the xiphisternum is associated with a single pair of xiphisternal ribs.

The forelimb ( Fig. 5D View FIGURE 5 ) is composed of a humerus with well-developed proximal and distal heads, a stout ulna and a slender radius, and a series of carpals and phalanges. The carpal region is composed of ulnare, radiale, pisiform, palmar, central, and four distal carpals (from II to V). The phalangeal formula of the forelimb is 2:3:4:4:3.

The pelvic girdle ( Fig. 5E View FIGURE 5 ) is formed by the pubis, ischium, and ilium. The pubis has a large obturator foramen and a well-developed pectinal process. Epipubis, epiischium and hypoischium are present.

The hindlimb ( Fig. 5E View FIGURE 5 ) is formed by the femur, tibia, and fibula. A large astragalus-calcaneum and smaller distal tarsals III and IV are present in the tarsal region. Two ventral sesamoids are also present. The phalangeal formula of the hindlimb is 2:3:4:5:3.

Morphological variation and morphometry. Table 1 presents comparative data on external morphology of Anotosaura collaris   , A. vanzolinia   , Colobosauroides cearensis   , C. carvalhoi Soares & Caramaschi, 1998   , and Dryadosaura nordestina   . Table 2 shows morphometric variation in A. collaris   and comparisons among males and females. Sexes were not significantly different in SVL (P = 0.394) but sexual dimorphism was detected in relative size, with females presenting larger LBL (P <0.01) and males presenting larger HW (P <0.01) ( Table 2). Sexual dimorphism did not interacted with species differences (P> 0.05 for all characters) thus sexes were pooled for intrageneric comparisons. Although similar in body size, A. vanzolinia   had shorter limbs, with significantly smaller FEM, HUM and FAL (P <0.01), a narrower head with smaller HW (P <0.01) and it was slightly but significantly more elongated than A. collaris   , presenting proportionally larger LBL ( ANCOVA with SVL as covariate, P <0.01).

A discriminant analysis including only males of the two species of Anotosaura   plus C. cearensis   and D. nordestina   shows that the first discriminant function explains 93.5% of variation among species, with high and positive loadings for limb length ( FAL, HUM, TIB, and FTL) ( Table 3). In the morphological space defined by this axis, both Anotosaura species   are well differentiated from Colobosauroides cearensis   and Dryadosaura nordestina   ( Fig. 6 View FIGURE 6 ), presenting relatively shorter forelimbs and hindlimbs. The four species are correctly classified by the discriminant function with 100% of correct classification for Anotosaura collaris   and C. cearensis   , 91.7% for A. vanzolinia   (one individual was classified as A. collaris   ) and 87.5% for D. nordestina   (one individual was classified as C. cearensis   ). The second discriminant function explains only 4.9% of variation and present positive loadings for all variables ( Table 3).

Karyotype. The karyotype of Anotosaura collaris   is composed by 44 biarmed chromosomes, 20 macrochromosomes and 24 microchromosomes ( Fig. 7 View FIGURE 7 ). Pairs 1, 4, 5, 6, 8 and 10 are metacentric and pairs 2, 3, 7, and 9 are submetacentric.

Molecular Phylogeny. Bayesian and Maximum Likelihood analysis recovered the same topology; Ecpleopus gaudichaudii Duméril & Bibron, 1839   as an early-diverging lineage within Ecpleopodini, sister to two distinct clades. One clade with ( Colobosauroides cearensis   + ( Anotosaura   + Dryadosaura   ), and the other grouping Marinussaurus   , Leposoma   and Arthrosaura   , with unresolved relationships. The analysis recovered Anotosaura   as monophyletic ( PP =1, ML bootstrap=98) ( Fig. 8 View FIGURE 8 ).

Uncorrected p -distances varied from 2.2 to 19.3% for 16S, 3.7 to 23.8% for 12S, 8.8 to 28.9% for ND4 and 4.0 to 27.8% for C-mos among Gymnophthalmidae   sampled. Distances between Anotosaura collaris   and A. vanzolinia   varied from 2.2 to 2.8% for 16S, 3.7 to 4% for 12S and 8.8 to 9.4% for ND4 and while distances among A. collaris   specimens were nearly zero (Table 4).

Distribution and natural history. Our sampling efforts in the present and as well as previous field trips covered a reasonably wide range of habitats and areas in the Caatinga. However, Anotosaura collaris   was only found in three localities: Senhor do Bonfim, Missão do Sahy, and Campo Formoso ( Fig. 9 View FIGURE 9 ). Maximum straight-line distance between them is about 15km. All sites are situated in isolated relictual mountains, emerging from the general Caatinga area, in the northern end of Serra de Jacobina, a small branch of Chapada Diamantina. Other lizards obtained at the same sites were the gymnophthalmids Acratosaura mentalis   and Psilophthalmus sp.   , the phyllodactylid Gymnodactylus geckoides   , the gekkonid Hemidactylus brasilianus   , the teiid Ameivula ocellifera   , and the tropidurids Tropidurus hispidus   and T. semitaeniatus   . This region has a basement formed by Paleoproterozoic and Neoarchean (2800–1600 MYbp) sedimentary metamorphic rocks ( Nápravník 2011) and rises up to 1200 m a.s.l. The soils occupied by A. collaris   are clayish, different from those occupied by all other species analyzed here, which tend to be more sandy (Table 5). These mountains are a barrier for the wet winds coming from the coast, which is congruent with the low precipitation seasonality at these sites (Table 5), and the presence of cloud forests at higher elevations. At the top, those mountains are often covered with campos rupestres, bearing extensive rock outcrops, whereas the lower flatlands are covered with typical thorny-shrubby Caatinga vegetation.

In Senhor do Bonfim, specimens of Anotosaura collaris   were found at Alto da Rainha, an isolated mountain at about 715 m a.s.l. elevation. In Missão do Sahy, an old indian Mission, specimens were found on the top of a mountain at 895 m a.s.l. and in Campo Formoso, at Morro do Cruzeiro (780 m a.s.l.). Two A. collaris   ’ shed skins were also found in Coqueiral (10°33'9.06"S, 40° 8'42.26"W), another isolated mountain at about 593 m a.s.l..

All specimens were collected by active search, using rakes. Anotosaura collaris   shows semi-fossorial behavior. Individuals were found only in the sub-superficial stratum, under leaf litter or small rocks. The maximum lizard density found was a group of two adults and two juveniles collected under a bush covering an area of approximately 32 m 2.


Museu de Zoologia da Universidade de Sao Paulo


Royal British Columbia Museum - Herbarium


Humboldt University Zoologisches Museum


Musee de Lectoure














Anotosaura collaris Amaral 1933

Rodrigues, Miguel Trefaut, Jr, Mauro Teixeira, Vechio, Francisco Dal, Amaro, Renata Cecília, Nisa, Carolina, Guerrero, Agustín Camacho, Damasceno, Roberta, Roscito, Juliana Gusson, Sales Nunes, Pedro M. & Recoder, Renato Sousa 2013

Anotosaura collaris

Vanzolini, P. 1976: 120

Anotosaura collaris collaris

Dixon, J. 1974: 14

Anotosaura collaris

Dixon, J. 1973: 8

Anotosaura collaris

Amaral, A. 1933: 69