Geographic variation and taxonomy of Liophis almadensis (Wagler) (Serpentes: Colubridae), and description of a new species of Liophis from Argentina and Bolivia.
The taxonomy of Liophis almadensis is, at best, chaotic. The species has been reported from widely scattered localities in South America in more than 40 citations and from more than 30 books and journals over the past 165 years. Recently, Vanzolini (1981) discussed the itinerary of the Spix and Martius 1817-1820 trip to Brazil, and designated the precise type locality of Liophis almadensis as Almada, Bahia, Brazil, 14[degrees] 39'S 39[degrees] 10'W. Additionally, Hoogmoed and Gruber (1983) examined the Spix and Wagler type material in the Natural History Museum of Munich and Leiden. They were able to establish ZSMH 2688/0 as the lectotype of Natrix almadensis. They also cited the spelling discrepancy (almada as opposed to almadensis) in the original description, as discussed by Vanzolini (1947) and Peters and Orejas-Miranda (1970). They concluded that the correct spelling, almadensis, was settled by the first revisor (Fitzinger, 1826).
In addition to the problem of spelling, it was stated in the original description that the number of ventrals ranged from 140 to 152, and the subcaudal number simply was given as "90." Of 245 individuals examined, the lowest number of ventrals is 143, and the highest number of subcaudals, 74. The lectotype is probably a newborn snake. It is only 170 mm in total length, and the ability to count subcaudals on a hatchling snake would be difficult today with poor magnification.
Liophis almadensis (Wagler, 1824)
Natrix almada Wagler, 1824:30. Type locality, Almada, Bahia, Brazil.
Natrix almadensis Wagler, 1824: pl. X, fig. 3.
Coronella almadensis, Fitzinger, 1826:895.
Liophis almadensis, Wagler, 1830:188.
Liophis conirostris Gunther, 1858:46. Type locality, "Brazil" and "Bahia."
Liophis wagleri Jan, 1859:274 (in part). Type locality, unknown.
Liophis (Lygophis) y-graecum Peters, 1882:129. Type locality, Villa de Guaratinqueta, Sao Paulo, Brazil.
Trigonocephalus scolecomorphus Bacque, 1906:111. Type locality, Asuncion, Paraguay.
Comments on synonyms. -- All populations of L. almadensis have a pronounced white or yellowish mark, edged with black, along the inner margins of the parietal scales. This mark may or may not extend anteriorly to the anterior edge of the frontal and supraocular scales. Dixon (1980), following Peters and Orejas-Miranda (1970), placed L. verecundus Jan in the synonomy of L. almadensis. The type has not been found and is presumed lost. However, a detailed analysis of the color pattern presented by Jan suggests that L. verecundus belongs in the synonomy of L. poecilogyrus. Jan's description of L. verecundus made no mention of the distinctive light parietal mark nor of the posterior dorsolateral light stripes present in L. almadensis. Jan did mention the basal shadowing of the ventrals, two rows of dark spots that form dorsolateral dark lines posteriorly, and a lateral row of regular dark spots that continue to the level of the vent, which are pattern characters found in L. poecilogyrus. In addition, the number of scale rows, subcaudals, and ventrals given by Jan fall within the range of variation of both species. Gunther (1858) made no mention of the distinctive light parietal mark in his description of L. conirostris, but upon examination, the mark is present in both syntypes.
Bacque's (1906) original description of Trigonocephalus scolecomorphus suggests that the color pattern is similar to that of L. almadensis. Bacque (1906) described the taxon as having two dorsolateral light lines, distinctive light parietal mark, rose colored belly, and a greenish gray dorsum with many smaller dark spots arranged in regular shapes and position. However, one could interpret the scales as possessing keels when Bacque stated that (paraphrased) "scales small and dense, free at the top, covering the entire back, and ... covered with a very marked keel." Bacque also stated that the "body, when fully extended, of a uniform thickness, rarely greater than that of an earthworm. The same goes for the length, which reaches 30-50 cm." The latter characters do not agree with those of L. almadensis. However, Serie's (1916) review of Bacque's work suggests that Bacque was not a herpetologist, confused many of the characters, and included many of the folklore stories that had little or no bearing to reality. Serie strongly suggested that T. scolecomorphus was identicial to one of the two common snakes in the area, L. almadensis and L. poecilogyrus. He also suggested that L. almadensis more closely fit Bacque's description because of the light parietal mark and the paired dorsolateral light lines on the body and tail, and I concur.
Distribution. -- Liophis almadensis is restricted to the south side of the Rio Amazonia, from Ilha Marajo along the Atlantic coast to the Brazilian state of Rio Grande do Sul, and westward to the Bolivian Chaco on the north and to the Paraguayan Chaco to the south (see Fig. 3).
Description of species. -- The following description is based upon 245 individuals from throughout the range of the species. An analysis of sexual dimorphism in a single large sample of individuals (98) from the state of Sao Paulo, Brazil, showed no statistical differences in the number of ventrals, subcaudals, or tail length/total length ratios; therefore, sexes were grouped for statistical analyses of those characters. The maximum total length of males is 524 mm, of females, 603 mm. Tail length/total length ratios of adults vary from 0.145 to 0.253 ([bar.x] = 0.217). Eye diameter/snout length ratios of adults vary from 0.563 to 0.778 ([bar.x] = 0.662). Scales are smooth, with one apical scale pit, and in 19-19-17 rows, never more than 19 scale rows at the tenth anterior ventral and at midbody. The reduction to 17 scale rows occurs through a fusion of scale rows 3 and 4 between ventrals 86 to 101 ([bar.x] = 91.4). The number of ventrals varies from 143 to 170 ([bar.x] = 156.6), number of subcaudals varies from 50 to 74 ([bar.x] = 63.6), and the number of maxillary teeth varies from 19 to 25 ([bar.x] = 21.6). The last two maxillary teeth are enlarged, without grooves, separated from the remainder of teeth by a diastema equal to the width of one or two prediastemal teeth; occasionally, the diastema is absent. Variation in head scales follow (number in parentheses is the number of individuals with that character): supralabials 7-7 (right/left) (5), 7-8 (20), 8-8 (218); infralabials 8-8 (1), 8-10 (3), 9-9 (24), 10-10 (190), 11-12 (1); supralabials entering orbit 3+4 (4), 3+4+5 (1), 3+4/4+5 (17), 4+5 (219), 4+5/5+6 (1), 4/4+5 (1); preoculars 0-0 (1), 1-1 (241, 1-2 (1); postoculars 1-1 (4), 1-2 (3), 2-2 (226), 2-3 (9), 3-3 (1); temporals 0+1 (1) 0+2 (1), 0+2/1+2 (1), 0+1/1+2 (2), 1+1 (11), 1+2 (210), 1+1/1+2 (19); invariate single loreal and divided anal plate.
The hemipenis (in situ) is 8 to 13 (x = 10.5) subcaudals in length. The sulcus spermaticus forks at the level of fifth or sixth subcaudal. The sulcate surface of the hemipenis is densely covered with small spines, whereas the asulcate surface is covered with large spines to the edge of the lobes (about two subcaudals from the tip), and the remainder of the lobe is covered with spinules to the edge of the smooth apical disk (Fig. 1).
Liophis almadensis is typically a grayish green to light olive brown snake whose body and tail has either distinct or obscure dark brown to black blotches or cross bands anteriorly and dorsally between scale rows five or six. The markings become more obscure posteriorly, frequently fading into dusky spots on the tail. The dorsal dark spots may appear as narrow crossbands between dorsolateral light stripes anteriorly. Posteriorly, there is a pale cream to yellowish tan line on scale row six and one-half of seven, or scale rows five and six, or one-half of five, all of six and one-half of seven. This line may be distinct from midbody to above the tail, or only evident on the posterior one-fourth of the body. The light line may be bordered below by a dark brown or black line that may be ill-defined in preservation. Occasionally, the entire dorsum may appear brownish with scattered light spots, which may appear as streaks or as zig-zag lines and affect only a few scales at a time. In younger specimens, the anteriormost nape band may be black, followed by two incomplete bands slightly lighter in color with narrow light interspaces one scale row wide. The ventral color is yellowish, pinkish, rose, or reddish orange with scattered black marks or smudges. The dorsal surface of the head is olive brown, blackish, or dark gray. The whitish or yellowish mark along the inner edge of the parietal scales (occasionally extending onto the frontal and supraoculars) may be shaped as a "W," "X," "Y," or "U," with dark edges (Fig. 2).
[FIGURE 1 OMITTED]
[FIGURE 2 OMITTED]
Geographic variation. -- Ten samples, ranging from the states of Pernambuco to Rio Grande do Sul, Brazil, and from Bolivia to Paraguay (Fig. 3) were utilized for an analysis of geographic variation in the numbers of ventrals, subcaudals, maxillary teeth, and tail length/total length ratios. Standard univariate statistical tests and Student's t-test were used to determine significant differences between samples. Pair-wise comparisons were made for all samples in all combinations. As previously indicated, there were no significant sexual differences among characters analyzed for a single large sample and the characters were combined for the analysis.
Univariate analysis (Fig. 4) suggested that the Para/Pernambuco/Paraiba sample (Fig. 3, sample 1,) was significantly distinct from the remaining nine samples. However, in a pair-wise comparison, Student's t-test values indicated that sample one was not significantly distinct from sample two, nor sample two from three, and so on. However, sample one was retained as a single sample (A) because univariate analysis indicated that it was different from the other samples. Sample one also was retained for comparison with other combined samples from throughout the range of the species. The remaining samples were not distinct from another by Student's t-test values, but univariate analysis suggested some significance in mean values. The central samples--Bolivia (sample nine), Mato Grosso and southern Para, Brazil (sample eight), and Paraguay (sample 10)--deviated enough from mean values of other samples to warrant grouping as a single large sample (B). The remaining samples ranging from the Brazilian states of Bahia (sample two) to Rio Grande do Sul (sample seven) also grouped into a single large sample (C, Figs. 3 and 4). Again, when these three larger samples were compared for the number of subcaudals and ventrals there were no significant differences between the samples, agreeing with the Student's t-test. The result of this comparison was surprising, because these characters vary geographically in many other species of Liophis.
Similar species. -- Liophis carajasensis Cunha and Nascimento (1985) closely resembles L. almadensis. It is known only from the type locality in the Serra Norte, Para, Brazil (circled star, Fig. 3). I have examined three paratypes of L. carajasensis and, in most respects, they resemble the color pattern of L. almadensis, including the distinctive light mark on the parietals. However, the number of subcaudals in L. carajasensis varies from 67 to 72 in the type series of 28 specimens (Cunha and Nascimento, 1985). Even if the number of subcaudals in the type series averaged on the low end, it would exceed the mean of the two closest samples of L. almadensis by five to six subcaudals. I hesitate to place this species in the synonomy of L. almadensis because of the subcaudal differences between the two species and prefer to wait until the entire type series of L. carajasensis can be examined.
Another population of Liophis that resembles L. almadensis occurs in low to moderate elevations (600 to 2000 meters from the Bolivian State of Santa Cruz south to the Argentine state of Catamarca (squares, D, Figs. 3 and 4). The color and pattern of the venter and dorsal surface of the body is similar to that in L. almadensis, including the posterior dorsolateral light stripes and body blotches. However, this population differs by the absence of the distinctive light parietal mark, the presence of an extra scale row reduction, more ventrals, fewer subcaudals, and a shorter, wider head. There is no existing name for this taxon, and I propose it be known as
Liophis ceii, new species
Holotype. -- Texas Cooperative Wildlife Collection no. 53409; adult male, from near Tucuman, Tucuman, Argentina; taken by James R. Dixon on 18 May 1977.
Paratypes. -- IML 35, 568, 587, 608, 796 (3), 895, TCWC 63824-25, 66842, topotypes; IML 1174-75 El Manchao, IML 650 Rio Paleonsa, Catamarca, Argentina; IML 613 Capillas, IML 690 Valle Grande, IML 9 and 594 Yuto, Jujuy, Argentina; IML 572 Brealito, IML 657 (3) El Alisal, IML 480 Hickman, IML 610 Saucelito, Salta, Argentina; IML 518 Fisica, IML 926 San Javier, IML 927 Santa de Medina, IML 896 (2) Trancas, IML 784 Yerba Buena, Tucuman, Argentina; UMMZ 69551 (6) and 69552 (5) Comarapa, 2000 meters, Santa Cruz, Bolivia; IML 631 Iscayachi, Tarija, Bolivia.
Description of holotype. -- Adult male, total length 524 mm, tail, 94 mm, tail length/total length ratio 0.179, eye diameter/snout length ratio 0.774; scales smooth, with one apical pit, and in 19-19-15 rows; first reduction occurs through the fusion of scale rows 3+4 at the 99th ventral (right), 100th (left); second reduction occurs through the fusion of scale rows 7+8 over the 102nd ventral (right/left). Ventrals number 164, subcaudals 52, supralabials 8-8, infralabials 10-10, preoculars 1-1, postoculars 2-2, temporals 1+2-1+2, supralabials entering orbit 4+5-4+5, loreal 1-1, maxillary teeth 19-19, anal plate divided.
Hemipenis everted, six subcaudals in length (11 mm), covered with a dense layer of small spinules that are interspersed with larger spines. The sulcus spermaticus divides about 3 mm from the base of the hemipenis, and each fork of the sulcus curves towards the outer edge of each lobe and enters the smooth apical disk at the inner lateral edge. There are six or seven rows of large spines concentrated along the outer edge of each lobe, with smaller spines scattered above the sulcus spermaticus to the edge of the disk. The asulcate surface has large spines around one-fourth of the base, with smaller spines between the lobes.
Dorsal body color pattern consists of a series of obscure narrow crossbands that are outlined with dark edges about one scale row in width, from nape to above vent. The latter are separated from each other by a transverse row of scales that contain paired white dots on their bases. These light interspaces and dorsal dark bands range in a more or less regular pattern between the sixth scale rows. A lateral series of blackish dots occur along the fourth row of scales, more or less in a regular series, from the nape to the vent. These blackish spots are separated from each other by a grayish green to olive ground color. The white to pale tan dorsolateral line occurs on scale rows 5-7 anteriorly, and 4-6 posteriorly. The anterior part of the light dorsolateral line is obscured by the cross bands, but becomes distinct on the posterior one-third of the body, and throughout the tail. The top of the head is olive green to gray green, with darker areas along the edges of the prefrontals, frontal, supraoculars and parietals. The side of the head is dark from the upper edge of the snout to the nape, and extends downward onto the upper edges of the first six supralabials, and progressively lower on supralabials 7 and 8. The nape band is divided middorsally, edged with black on the outer edges, tending to make the middle of the band appear light in color. The ground color of the venter is rose in life, with the chin, throat and infralabials white. There are a series of lateral black marks on every other ventral, never extending completely across the venter. Some of these black marks are centrally located on the posterior part of the venter. The subcaudals are rose with an occasional black mark on the outer edge of the subcaudals.
[FIGURE 3 OMITTED]
[FIGURE 4 OMITTED]
Geographic variation. -- A series of specimens from above 2000 meters and below 1000 meters in Argentina and Boliva show no sexual dimorphism in numbers of ventrals and subcaudals but indicate significant differences (univariate analysis) between subcaudal numbers by elevation. The number of ventrals from specimens above 2000 meters varies from 158 to 168 ([bar.x] = 162.2), and subcaudals vary from 54 to 64 ([bar.x] = 59.7). The number of ventrals from specimens below 1000 meters varies from 157 to 170 ([bar.x] 165.1), subcaudals, 49 to 59 ([bar.x] = 53.3). I have not examined specimens from between 1000 and 2000 meters nor have I noted any conspicuous differences in color pattern between high and low samples. There is a strong possibility that the trend from low to high numbers of subcaudals is clinal. Dixon (1983) cited similar elevational trends in various populations of L. epinephelus.
Etymology. -- This species is named in honor of my friend and colleague, Dr. Jose M. Cei, who has contributed much to our knowledge of the amphibian and reptile fauna of Argentina and Chile.
APPENDIX -- SPECIMENS EXAMINED
Liophis almadensis. ARGENTINA. no specific locality "Niederlin's Expedition to Gran Chaco and Misiones" AMNH 17482. BOLIVIA. Beni: Lake Rogoagua AMNH 22458; San Joaquin, 20 km S. AMNH 104571; Santa Rosa, Rio Mamore AMNH 101872. Santa Cruz: Buenavista BM 19184.108.40.206, CM 2820, 2897, 2910, 2917, 2968; La Perdiz CM 2852; Santa Cong BM 19220.127.116.11; Santa Cruz de la Sierra CM 112; Santa Roca de la Roca UF 68510. BRAZIL. Bahia: no specific locality BM 1918.104.22.168, 1922.214.171.124, 19126.96.36.199; 188.8.131.52, 184.108.40.206-7, MCZ 3641, MZUSP 779; Barra IB 987; Camacari IB 960-61; Cruz das Almas MZUSP 2415-16; Fazenda Cabucu IB 23886; Mata do Sao Joao IB 1059, 1407, 1764-65; Salvador MZUSP 2832, 2836. Goias: Araguacema IB 12030; Campinas IB 6073, USNM 100746; Gurupi MZUSP 8008; Jatai, Fazenda Cachoeirinha MZUSP 3776; Rio Verde MZUSP 3343; Tocantinia MZUSP 3343; Tocantinia MZUSP 3831. Guanabara: no specific locality MCZ 4334, 17950, MZUSP 781. Mato Grosso: Aquidauana IB 19385; Buriti, Chapada dos Guimaraes MZUSP 5347; Campo Grande IIB 24667; Chapada BM 1903.3.26.28; Coxim, 35 km SW, on Rio Taquare USNM 163502; Cuiaba IB 29871, MZUSP 1735; Dourado, via Itahum IB 27907; Guavira IB 15005; Jupia MZUSP 4423; Jurema IB 1621-22; Miranda IB 15021; Ponta Pora IB 17278, 17312, 24637, 27749; Porto Vehlo, Rio Tapirapes MZUSP 3772; San Luis de Caceres IB 23846, 24154-55, MZUSP 3314; Sao Felix IB 12028; Tapirapes IB 12042, 12060; Utiariti MZUSP 4744-46; Xingu IB 28757. Minas Gerais: no specific locality MCZ 1795, 13957; Cerra do Cipo MZUSP 7585; Lima Duarte IB 16744; Machado IB 11173, 16759-61, 18555; Mariana IB 7324, MZUSP 777, 782; Penetenciaria Neves near Belo Horizonte UMMZ 109068; Pirapora IB 16477; Santa Barbara do Mato Dentro IB 2122, 5569-70; Sao Joa del Rei, Rio dos Mortes CM 352; Tres Santa IB 10329; Valadares IB 16763. Para: Cachimbo MZUSP 3315; Camaragibe IB 16624; Ilha do Marajo IB 17657. Paraiba: Joao Pessoa MZUSP 7998; Mamanguape MZUSP 3317. Parana: Dorizon AMNH 6489; Pameira IB 21958; Ponta Grossa IB 4333-34, 5350, 23980, MZUSP 5788; Porto Uniao (Vitoria) IB 13564. Pernambuco: no specific locality DEH 512; Bonji DEH 258, 757, 759; Camaragibe DEH 221; Jaboatao MZUSP 7429; Recife DEH 134, 155, 536, 623; Sao Lourenco da Mata DEH 102; Sao Severino do Ramos MZUSP 8035. Rio Grande do Sul: no specific locality BM 220.127.116.11-80; Capela IB 5904, 5938, 6371 Porto Alegre IB 6336, MZUSP 799; Rio Pardo IB 17196, 17227; Rosario do Sul IB 15942; Sao Leopoldo IB 5685, 5687, 5934, 5951, 11353-54. Santa Catarina: no specific locality IB 636. Sao Paulo: no specific locality MCZ 17948-49, 17951-53, UMMZ 62837-45, USNM 76394; Caixa d' Agua MCZ 39435; Funil MZUSP 786-87; Guaianases MZUSP 6350; Ipanema MZUSP 747; Itanhaem MZUSP 3991; Itapetininga MZUSP 4629, 4633; Jundiai MZUSP 800, 4033-35; Osasco MZUSP 3681; Perus MZUSP 751-52, 754-56, 758-60; Raiz do Serra MZUSP 748; Sao Bernardo do Campo MZUSP 4048, 4052-53; Sao Paulo MZUSP 746, 801-12, 2379-80, 2383, 2405, 2418, 2568-69, 2622, 4010, 4046, 4763, UMMZ 79667; Sao Paulo, Agua Funda MZUSP 3312; Sao Paulo, Ibirapuera MZUSP 2636-37; Sao Paulo, Ipiranga MZUSP 768-75, 783-85, 738-44, 761-66, 1362, 1646, 1681, 1702, 1744, 1768, 1882, 1919, 2781; Sao Paulo, Sacoman MZUSP 1830; Sao Vicente MZUSP 788-89, 4156, 4594; Tiete MZUSP 4038. PARAGUAY. no specific locality USNM 11263 (2); Carumbe FML 573, 575, 751; Nueva Asuncion, 28.8 km. W Fortin Madrejon LACM 126504; Primavera, Alto Paraguay BM 1918.104.22.168, 1922.214.171.124, 19126.96.36.199, 19188.8.131.52, 1962.63-67.
Liophis ceii. ARGENTINA. no specific locality IML 623 (3), 642, 800. Catamarca: El Machado IML 1174-75; Rio Paleonsa IML 650. Jujuy: Arroyo Quemado, Yuto IML 9, 594; Capillas IML 613; Valle Grande, Abra de Canas IML 690. Salta: El Alisal IML 657 (3); El Brealtio IML 572; Hickman IML 480; Saucelito, 50 km S Oran IML 610. Tucuman: San Javier IML 926; Santa de Medina IML 927; Trancas IML 896 (2); Tucuman IML 6-35, 568, 578, 608, 796(3), 895, TCWC 53409, 63824-25, 66042. BOLIVIA. Santa Cruz: Comarapa UMMZ 69551 (6), 69552(5). Tarija: Iscayachi, 34 km. WNW Tarija IML 631.
This paper would not have been possible without the help of museum curators such as R. Zweifel, C. J. McCoy, A. Stimpson, E. Williams, C. L. Cordeiro, P. E. Vanzolini, A. Kluge, R. W. McDiarmid, J. W. Wright, F. W. King, and R. F. Laurent.
I especially thank P. E. Vanzolini, C. L. Cordiero, and R. F. Laurent for their hospitality during our visits to Brazil and Argentina.
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______. 1983. Systematics of the Latin American snake, Liophis epinephelus (Serpentes, Colubridae). Pp. 132-142, in Advances in herpetology and evolutionary biology (A. G. J. Rhodin and K. Miyata, eds.), Mus. Comp. Zool., Harvard, xix + 725 pp.
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Serie, P. 1916. Sobre tres supuestos nuevos trignocephales del Paraguay. Physis, 2 (10):171-174.
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______. 1981. The scientific and political contents of the Bavarian Expedition to Brasil. Pp. ix-xxix, in Herpetology of Brazil (J. B. von Spix and J. G. Wagler), SSAR Facs. Reprint in Herpetology.
Wagler, J. 1824. Serpentum brasiliensium species novae ou Histoire naturelle des especes nouvelles de serpens, recueillies et observees pendant le voyage dans l'interieur du Bresil dans les annees 1817, 1818, 1819, 1820, excute par ordre de Sa Majeste le Roi de Baviere, publiee par Jean de Spix,... ecrite d'apres les notes du voyageur par Jean Wagler. Monachii, Franc. Seraph. Hubschmann, viii+75 pp.
______. 1830. Naturliches System der Amphibien mit vorangehender Classification der Saugetiere und vogel. Munchen, Stuttgart und Tubingen, i+354 pp.
JAMES R. DIXON
Department of Wildlife and Fisheries Sciences, Texas A & M University, College Station, Texas 77843
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|Author:||Dixon, James R.|
|Publication:||The Texas Journal of Science|
|Date:||Aug 1, 1991|
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