ZooKeys 361: 61–72, doi: 10.3897/zookeys.361.5738
Description of the hemipenial morphology of Tupinambis quadrilineatus Manzani and Abe, 1997 (Squamata, Teiidae) and new records from Piauí, Brazil
Marcélia Basto da Silva 1, Geraldo Rodrigues de Lima-Filho 1, Áurea Aguiar Cronemberger 1, Leonardo Sousa Carvalho 2, Paulo Roberto Manzani 3, Jânia Brito Vieira 4
1 Museu Paraense Emilio Goeldi, Departamento de Zoologia, Av. Perimetral, 1901, Bairro Terra Firme, CEP 66077-530, Belém, PA, Brazil
2 Universidade Federal do Piauí, Campus Amílcar Ferreira Sobral, BR 343, KM 3.5, Bairro Meladão, s/nº, CEP 64800-000, Floriano, PI, Brazil
3 Universidade Estadual de Campinas, Instituto de Biologia, Departamento de Biologia Animal, Rua Monteiro Lobato, 255, Distrito de Barão Geraldo, CEP 13083-862, Campinas, SP, Brazil
4 Universidade Federal do Piauí, Campus Ministro Reis Veloso, Avenida São Sebastião 2819. CEP. 64202-220, Parnaíba, PI, Brazil

Corresponding author: Marcélia Basto da Silva (marceliabasto@gmail.com)

Academic editor: J. Penner

received 3 June 2013 | accepted 27 September 2013 | Published 12 December 2013

(C) 2013 Marcélia Basto da Silva. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: da Silva MB, de Lima-Filho GR, Cronemberger ÁA, Carvalho LS, Manzani PR, Vieira JB (2013) Description of the hemipenial morphology of Tupinambis quadrilineatus Manzani and Abe, 1997 (Squamata, Teiidae) and new records from Piauí, Brazil. ZooKeys 361: 61–72. doi: 10.3897/zookeys.361.5738


Few data are available on the morphology of the hemipenis of teiid lizards, especially those of the recently-defined genus Tupinambis, a widely-distributed group of large-bodied lizards. This study provides an illustrated description of the hemipenis of Tupinambis quadrilineatus, which is similar to that of other representatives of the Tupinambinae subfamily. New records of the species from the state of Piauí, in northeastern Brazil, are also presented.


Hemipenis, systematics, Tupinambinae, Tupinambis, new records


The genus Tupinambis Daudin (Teiidae) comprises a group of large (maximum SVL of 400 mm) Neotropical lizards, which are distinguished from all other teiids by the combination of smooth dorsal scales, a single loreal, a gap in the granular scales separating the femoral from the abdominal pores, and a cylindrical tail with complete annuli alternating with annuli divided on the dorsal and lateral sides (Harvey et al. 2012). In a recent review of the Teiidae, Harvey et al. (2012) resurrected the genus Salvator Duméril and Bibron to refer to the species of the “southern clade” (sensu Fitzgerald et al. 1999) previously included in Tupinambis. These species are now known as Salvator merianae (Duméril & Bibron, 1839), Salvator rufescens (Günther, 1871) and Salvator duseni (Lönnberg, 1896). According to this scheme, the genus Tupinambis currently includes only the four species of the northern or “Amazonian” clade (sensu Fitzgerald et al. 1999) – Tupinambis longilineus Avila-Pires, 1995, Tupinambis palustris Manzani & Abe, 2002, Tupinambis quadrilineatus Manzani & Abe, 1997, and Tupinambis teguixin (Linnaeus, 1758). The genus Tupinambis is found in Colombia, Venezuela, Trinidad and Tobago, the Guyanas, the Amazon basin, and the savannas of Bolivia and Brazil (Harvey et al. 2012). Despite the conspicuous size of these lizards, zoogeographic data are sketchy, and new localities have been recorded recently for some species, such as Tupinambis longilineus (Lima and Pimenta 2008, Costa et al. 2008) and Tupinambis quadrilineatus (Ferreira et al. 2009, Silveira 2009).

Tupinambis quadrilineatus is endemic to the Cerrado savannas of central Brazil. The species was described in 1997, based on four specimens from Goiás, Mato Grosso, and Tocantins (Manzani and Abe 1997, Silveira 2009). Almost simultaneously, Colli et al. (1998) described the same form under the junior-synonym Tupinambis cerradensis. A number of other specimens were collected subsequently in the Brazilian states of Goiás, Minas Gerais, Mato Grosso, Maranhão, Tocantins, Piauí, Pará and the Distrito Federal (Barreto et al. 2007, Ferreira et al. 2009, Silveira 2009, Dal Vechio et al. 2013). The geographic range of the species is extended further in the present study.

The hemipenis of Tupinambis quadrilineatus is also described here for the first time. The hemipenial morphology of teiid lizards is poorly known (Harvey et al. 2012). Cope (1896) analyzed the hemipenis of the genera Dracaena, Tupinambis, Ameiva, and Cnemidophorus and concluded that the morphology of these typical teiid species consist of numerous delicate, imbricate, transverse laminae, which are closely attached to one another. Dowling and Duellman (1978, figure 83.2) published an illustration of the sulcate surface of the hemipenis of a species referred to as Tupinambis nigropunctatus Spix, 1825, however they did not provide a museum number, nor did they describe the organ. Presently, Tupinambis nigropunctatus is considered as a synonym of Tupinambis teguixin (Linnaeus, 1758), and its drawing exihibited a slightly bilobed and relatively long hemipenis, with distal laminae. In addition, the hemipenial morphology of 13 teiid species was described by Böhme (1988), but the author did not examine nor describe the hemipenis of Tupinambis.

Besides, Harvey et al. (2012) reviewed the taxonomy and phylogeny of the teiids and included descriptions of the hemipenes of a number of species of the subfamily Tupinambinae, including Crocodilurus amazonicus Spix, 1825 and Salvator merianae. This study shows that the hemipenis in the Tupinambinae can be characterized as an organ with transverse laminae, a pair of apical awns, and catchment folds. Awns are usually prominent subcylindrical structures, rounded at their distal ends, located at the apex of the lobes. The most elaborate sulcate catchment fold can be observed in Crocodilurus and Salvator, in which the portion of the fold closest to the sulcus projects outward as a prominent triangular flap. A summary of the hemipenial characters for the Teiidae subfamilies presented by Harvey et al. (2012) is shown in Table 1. The hemipenial morphology of Tupinambis nevertheless remains unknown, and the present paper provides a first detailed description of the organ in this genus.

Table 1.

Hemipenial characters of teiid lizards of three subfamilies of Teiidae (Harvey et al. 2012).

Subfamilies Proximal laminae Distal laminae Discontinuous laminae Awns Apical sulcate
Apical asulcate structures
Callopistinae 6 19 Present Absent Large Flat
Teiinae 0–50 5–24 Absent (in most species) Present Catchment Fold/ Papillate/ Subtriangular or Rounded Lobes Rounded Lobes/
High semicircular or Straight Ridges (Flap)/ Subtriangular
Flaps/ Papillate/ Rounded Lobes
Tupinambinae 27–40 44–71 Absent Absent/Styloid Catchment Folds with Triangular Flaps Rounded Lobes

Specimens were collected from a locality in the Cerrado savanna of the state of Maranhão and different phytophysiognomies in Piauí. The material examined is deposited in the herpetological collections of the Coleção de História Natural of the Universidade Federal do Piauí, Floriano, Piauí (CHNUFPI, curator: L. S. Carvalho) and the Museu Paraense Emílio Goeldi, Belém, Pará (MPEG, curator: A. L. C. Prudente). Museum abbreviations follow Levington et al. (1980). Scale counts, body measurements, and color pattern are based on the schemes of Manzani and Abe (1997) and Colli et al. (1998). The sex of the specimens was determined by the presence or absence of a hemipenis verified through an incision at the base of the tail. Hemipenis terminology follows Savage (1997), Myers et al. (1993) and Harvey et al. (2012), and the specimens were prepared as in Pesantes (1994), Manzani and Abe (1988) and Zaher and Prudente (2003).

Tupinambis quadrilineatus Manzani & Abe, 1997


Tupinambis quadrilineatus Manzani & Abe, 1997: 2 (adult male holotype deposited in the Museu de Zoologia of the Universidade Estadual de Campinas, ZUEC 1963, type-locality: Fazenda Bandeirantes, Municipalty of Baliza, Goiás, Brazil (16°13'S, 51°25'W, SAD69), not examined).
Tupinambis cerradensis Colli, Péres & Cunha, 1998: 479 (adult male holotype deposited in the Coleção de Herpetologia of the Universidade de Brasília, CHUNB 00468, type-locality: Rosário Oeste, Mato Grosso, Brazil (14°50'S, 56°25'W, SAD69), not examined).
Tupinambis quadrilineatus; Taylor 2003: 44, Langstroth 2005: 106, Silva Jr. et al. 2005: 81, Vitt et al. 2005: 8, Werneck and Colli 2006: 1987, Guimarães et al. 2007: 353, Recoder and Nogueira 2007: 270, Silveira 2009: 442, Ferreira et al. 2009: 355, Moreira et al. 2009: 187, Recoder et al. 2011: 275.

Tupinambis quadrilineatus differs (see Table 2 for scale counts and measurements) from the other species of the genus in the presence of 11–18 femoral pores (15–18 in Tupinambis teguixin, 20–22 in Tupinambis longilineus, 18–26 in Tupinambis palustris), 94–118 scales around the mid-body (94–124 in Tupinambis teguixin, 90–98 in Tupinambis longilineus, 112–119 in Tupinambis palustris), 113–138 dorsal scales (102–126 in Tupinambis teguixin, 110–121 in Tupinambis longilineus, 111–122 in Tupinambis palustris) and the coloration. In Tupinambis quadrilineatus, the upper lateral stripe is well-defined along the flanks, whereas in other species, it is indistinct or absent (Avila-Pires 1995, Colli et al. 1998, Manzani and Abe 2002, Harvey et al. 2012).

Table 2.

Scale counts of the specimens of Tupinambis quadrilineatus analyzed in the present study and the known range of values for the species, according to Manzani and Abe (1997) and Colli et al. (1998).

Character CHNUFPI 0036 CHNUFPI 0037 CHNUFPI 0038 MPEG 16817 MPEG 16845 MPEG 30139 MPEG 30140*** MPEG 30141 Known range of values
Sex Male Male Male Female Female Male Immature male Male
Femoral pores* 11 10 12 10 10 11 11 11 11–18
Pre-cloacal pores* 10 8 10 8 9 8 8 8 5–11
Dorsal scales 127 118 119 115 116 109 111** 117 113–138
Scales around midbody 116 105 120** 112 116 90** 103 98 94–118
Ventral scales in a transverse row 24 24 25 25 26 23 24 25 20–28
Lamellae under fourth finger 15 13 15 14 15 14 15 14 12–17
Lamellae under fourth toe 29 30 30 33 34** 27 32 27 26–33
Loreal scale 1 1 1 1 1 1 1 1 1
Supralabial scales* 17** 15 17** 16 16 - - - 13–16
Infralabial scales* 14 14 16 14 14 - - 14 13-17
Snout-vent length (mm) 260 260 227 245 235 260 135 255 88-270
Body width (mm) 49.56 54.48 48.36 59.98 51.27 58.66 - 55.98 17.92-61.86
Body height (mm) 36.58 37.20 30.46 42.09 36.03 44.59 - 40.66 13.95-51.98
Head length (mm) 55.66 55.92 51.29 52.05 52, 80 54.32 - 52.39 24.10-62.04
Head width (mm) 42.73 44.98 34.98 41.61 33.91 39.44 - 40.22 15.16-44.28
Head height (mm) 30.20 28.37 28.26 35.47 30.35 36.23 - 29.69 11.60-38.40

* Total number on both sides.

** Exceeds maximum value recorded in previous studies.

***This specimen has a damaged head and part of the body, which prevented the withdraw of the remaining measures.

Hemipenial morphology.

The hemipenis of three specimens of Tupinambis quadrilineatus (CHNUFPI 0036, CHNUFPI 0038 and MPEG 30139) were prepared for analysis. The organ is relatively long, robust and slightly bilobed, with a total length of 5.0 cm and a width of 2.0 cm in the distal portion of the body (Figure 1). When inverted, the organ extends as far as the fifteenth subcaudal scale. Sulcus spermaticus bifurcated, deep and centripetal. Edge of the sulcus spermaticus pronounced along its entire length. The point of bifurcation of the lobes extends inwardly towards the central region of the styloid process. A pair of short and prominent lobes (about 16% of the total size of the organ) in the form of styloid process are present on either side of the sulcate and asulcate surface, with a pair of catchment folds (extensions of the lips of the sulcus, in the form of prominent sulcal flaps, with rounded edges) coating the styloid process. The region between the lobes is smooth on both the sulcate and asulcate surfaces. Naked sulcate and asulcate expansion pleat. Between 35 and 38 distal laminae (mean = 36 ± 1, n = 3), arranged in a transverse row on each side, extending from just below the apical folds to the base of the lobes. A lateral sulcus separates the distal laminae from the sulcate and asulcate surfaces. Fifteen to 17 proximal laminae (mean = 16 ±1, n = 3). Basal region smooth on the sulcate surface, and wrinkled on the asulcate surface. Discontinuous laminae and basal papillae absent.

Figure 1.

Right hemipenis of Tupinambis quadrilineatus (CHNUFPI 0036). A sulcate surface B asulcate surface C lateral region. Scale bar = 1 cm.

The hemipenial morphology of Tupinambis quadrilineatus is similar to that of other Tupinambinae in the ornamentation of the body, which are bilobed and have lamelae (Cope 1896, Dowling and Duellman 1978, Harvey et al. 2012). As in Salvator merianae, Tupinambis teguixin and Crocodilurus amazonicus (Dowling and Duellman 1978, Harvey et al. 2012), the hemipenis of Tupinambis quadrilineatus lacks the discontinuous distal laminae seen in Ameiva ameiva and Ameivula ocellifera. However, Salvator merianae, formerly considered to be a member of the genus Tupinambis, has a relatively long hemipenis, which lacks the lateral and medial expansion pleats and has more laminae (distal laminae: 56–71 and proximal laminae: 33–40) than other teiids (Harvey et al. 2012). See Table 1 for the differences in the hemipenial morphology of three subfamilies of Teiidae (Harvey et al. 2012). The morphology and ornamentation of the hemipenis play an important role in the diagnosis of species, and have proven to be an excellent indicator of the phylogenetic relationships among taxa (Cope 1896, Böhme 1988, Harvey et al. 2012). Harvey et al. (2012) concluded that the relationships among the genera of Tupinambinae, especially Tupinambis and Salvator, require further study, and that a more detailed analysis of hemipenial morphology, as well as muscles and osteology, may contribute to a more definitive understanding of the systematics of the group.


Based on eight specimens. Snout-vent length 135–260 mm (mean = 234.9 mm); body width 48.4–60.0 mm (mean = 54.0 mm), body height 30.5–44.6 mm (mean = 38.5 mm), head length 51.3–55.9 mm (mean = 53.6 mm), head width 33.9–45.0 mm (mean = 39.7 mm), head height 28.3–36.2 mm (mean = 31.2 mm). See Table 1 for a complete list of the measurements and scale counts recorded in the present study and those available in the literature (Manzani and Abe 1997, Colli et al. 1998, Silveira 2009).

Geographic distribution.

The Tupinambis specimens available in Brazilian collections were examined together with the eight Tupinambis quadrilineatus specimens collected during the present study, in Maranhão and Piauí (Figure 2). The localities reported here represent the northernmost known records of Tupinambis quadrilineatus, and extend the known distribution of the species at least 500 km from the nearest locality, in Balsas, Maranhão (Barreto et al. 2007). This is the northernmost record of the occurrence of the species.

Figure 2.

Known localities for Tupinambis quadrilineatus in Brazil. Distrito Federal (DF): Brasília, Gama (1). Goiás (GO): Iaciara (2) Minaçu (3) Mara Rosa (4) Santa Terezinha de Goiás (5) Pirenópolis (6) Aragarças (7) Baliza (8). Maranhão (MA): Balsas (9) São Raimundo das Mangabeiras (10). Mato Grosso (MT): Primavera do Leste (11) Chapada dos Guimarães (12) Rosário Oeste (13) Cáceres (14). Minas Gerais (MG): Chapada Gaúcha (15) João Pinheiro (16). Piauí (PI): Guadalupe (17) Lagoa Alegre (18) Altos (19) Monsenhor Gil (20) Amarante (21), Ribeiro Gonçalves (22) Uruçuí (23). Pará (PA): Santa Maria das Barreiras and Redenção (24). Tocantins (TO): Gurupi (25) Mateiros (26). The localities recorded in the present study are represented by red squares. The type-locality of Tupinambis quadrilineatus is shown as an asterisk, the type-locality of its junior-synonym (Tupinambis cerradensis) is shown as a star and remaining records from the literature are shown as blue circles (Manzani and Abe 1997; Colli et al. 1998; Guimarães et al. 2007; Silva Jr. et al. 2005; Vitt et al. 2005; Mesquita et al. 2006; Recoder and Nogueira 2007; Ferreira et al. 2009; Silveira 2009; Recoder et al. 2011; Dal Vechio et al. 2013). The Cerrado savanna biome is highlighted in gray.

Five Tupinambis quadrilineatus specimens were examined in the collection of the Goeldi Museum. In 1993, specimen MPEG 16817 was collected in Balsas, Maranhão (reported by Barreto et al. 2007), and specimen MPEG 16845 was captured in the municipality of Lagoa Alegre, Piauí. In 2009, three specimens were collected during the Parnaiba Project in Ribeiro Gonçalves (MPEG 30139), and Uruçuí (MPEG 30141), in the state of Piauí, and São Raimundo das Mangabeiras (MPEG 30140), in Maranhão.

The herpetological collection of the Universidade Federal do Piauí provided specimens or records of Tupinambis quadrilineatus from a number of sites in Piauí. Specimen CHNUFPI 0036 (Figure 3A) was collected in 2010 in the Palmares National Forest (05°02'55"S, 42°35'59"W, SAD69), in the municipality of Altos. The vegetation of this area is semi-deciduous tropical forest typical of the Cerrado, an ecotonal region between Cerrado and Amazonia biomes, similar to that found in Lagoa Alegre. Tupinambis quadrilineatus occurs in syntopy with Salvator merianae in this area, as recorded at a number of other sites (Colli et al. 1998, Silveira 2009). Also in 2010, a roadkilled specimen of Tupinambis quadrilineatus (CHNUFPI 0037) was collected in the municipality of Monsenhor Gil (05°39'56"S, 42°35'28"W, SAD69). In May 2011, the third and final Tupinambis quadrilineatus specimen held in the collection (CHNUFPI 0038; Figure 3B–D) was collected in a pitfall trap installed in the vicinity of a small stream within an area dominated by Cerrado savanna (sensu strictu) in the municipality of Guadalupe (05°2'55"S, 42°35'59"W, SAD69). Two other specimens were observed in the municipality of Amarante (06°14'43"S, 42°46'46"W and 06°2'1"S, 43°3'40"W, SAD69) in 2009 and 2011, but specimens were not collected. In this area, the vegetation was dominated by secondary semi-deciduous tropical forest, mixed with patches of Cerrado sensu strictu.

Figure 3.

Adult male Tupinambis quadrilineatus. A specimen collected in the Palmares National Forest, Altos, Piauí (CHNUFPI 0036; Scale 5cm) B specimen collected with pit-fall traps at Guadalupe, Piauí (CHNUFPI 0038) C lateral view of the head and D dorsal view of the anterior region of the body (CHNUFPI 0036).

These findings expand the geographic distribution of Tupinambis quadrilineatus is northwards, and encompass the the region between the states of Piauí and Maranhão, which is dominated by Cerrado sensu strictu and/or forested patches of the Cerrado–Amazon ecotone. In this region, Tupinambis quadrilineatus also occurs in syntopy with Salvator merianae, which was previously classified as a member of the genus Tupinambis.


We are grateful to the Chico Mendes Institute for the Conservation of Biodiversity (ICMBio) for specimen collection permit #23561-1/SISBIO. We are especially indebted to A. L. C. Prudente (Museu Paraense Emílio Goeldi) for the loan of MPEG specimens and making working space available, A. O. Maciel for providing locality data for the specimens collecting during the Parnaiba Project and to S. Ferrari for the English revision of this paper.

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Material analyzed
Table of localities.

Tupinambis quadrilineatus: Maranhão: MPEG 16817 (Rio Matões, right bank tributary of the Rio Balsas, Balsas), MPEG 30140 (São Raimundo das Mangabeiras), Piauí: CHNUFPI 0036 (Palmares National Forest, Altos), CHNUFPI 0037 (Monsenhor Gil), CHNUFPI 0038 (Fazenda São Pedro, Guadalupe), MPEG 16845 (Lagoa Alegre), MPEG 30139 (Ribeiro Gonçalves), MPEG 30141(Estação Ecológica de Uruçuí-Una, Uruçuí).

Collection Collection number Family Genus Species Locality Municipality State Latitude, Longitude Coordinate origin
MPEG MPEG 16817 Teiidae Tupinambis quadrilineatus Near Matões River, tributary of the Balsas Rive Balsas Maranhão 7°31'S, 46°2'W Google Earth
MPEG MPEG 30140 Teiidae Tupinambis quadrilineatus São Raimundo das Mangabeiras Maranhão 7°1'S, 45°28'W A. O. Maciel
CHNUFPI CHNUFPI 0036 Teiidae Tupinambis quadrilineatus Palmares National Forest Altos Piauí 05°02'55"S, 42°35'59"W GPS
CHNUFPI CHNUFPI 0037 Teiidae Tupinambis quadrilineatus BR 343 Monsenhor Gil Piauí 05°39'56"S, 42°35'28"W GPS
CHNUFPI CHNUFPI 0038 Teiidae Tupinambis quadrilineatus São Pedro Farm Guadalupe Piauí 5°2'55"S, 42°35'59"W GPS
MPEG MPEG 16845 Teiidae Tupinambis quadrilineatus Lagoa Alegre Piauí 4°26'S, 42°35'W Google Earth
MPEG MPEG 30139 Teiidae Tupinambis quadrilineatus Ribeiro Gonçalves Piauí 7°33'S, 45°14'W A. O. Maciel
MPEG MPEG 30141 Teiidae Tupinambis quadrilineatus Uruçuí-Una Ecological Station Uruçuí Piauí 7°14'S, 44°33'W Google Earth
Not collected Not collected Teiidae Tupinambis quadrilineatus BR 343 Amarante Piauí 6°14'43.49"S, 42°46'46.02"W GPS