ANNALS OF CARNEGIE MUSEUM

Vol. 56, Article 8, Pp. 1 73-- 191 28 August 1987

TAXONOMY AND GEOGRAPHIC VARIATION OF

LIOPHIS TYPHLUS AND RELATED “GREEN” SPECIES OF
SOUTH AMERICA (SERPENTES: COLUBRIDAE)

James R. Dixon 1

Abstract
ThegreenspeciesofLiophisarereviewed.Liophistyphlusconsistsofthreesubspecies,L.t.typhlus
(Amazon), L. t. elaeoides (Chaco), and L. t. brachyurus (Cerrado). Liophis viridis consists of two
subspecies,L.v.viridis(AgresteandAtlanticrainforest),andL.v.prasinus(Caatinga).Liophisjaegeri
is shown to consist of two subspecies, L. jaegeri jaegeri, east of the Rio Parana, from Sao Paulo area
of Brazil, to Uruguay and Argentina and L. j. coralliventris, from the Rio Paraguay basin. Liophis
guentheri is a valid species from the dry central Chaco of Argentina, Bolivia, and Paraguay. The
charactersoftherecentlydescribedspecies,L.atriventerandL.maryellenaearesummarized.Akey
isprovidedforallspeciesof“green”Liophis.

Introduction
The study of the genus Liophis, a long and difficult task, is now nearing com-
pletion. The six species presented herein (L. typhlus, L. guentheri, L. jaegeri, L.
viridis, L. maryellenae, L. atriventer) form a loosely connected complex of species
that have one feature in common, they are all some shade of “green.” The green
color may be dull to bright, chlorophyll green to leaf green, or with a deep olive
cast. Two taxa occasionally have an obscure mid-dorsal stripe of variable width
that varies in color from light brown to reddish, and occasionally secondary dark
lines as well.
When samples were adequate, each of the six species was examined for geo-
graphic variation. No additional specimens of two recently described taxa ( L .
atriventer, L. maryellenae) are known, but the species’ essential features have been
summarized. Of the remaining four species detailed discussions include, where
appropriate, the currently recognized name, its synonyms, a discussion of the
synonyms, a description of the species, geographic variation, distribution, and
comments about pertinent literature.

Systematic Account
Liophis typhlus (Linnaeus)
ColubertyphlusLinnaeus,1758.Syntypes(notexamined)—RoyalMuseum,Stockholm.Type-local-
ity—Indiis(inerror).
XenodonisolepisCope,1870.HolotypeANSP(lost).Type-locality—Pebas,Ecuador(=Peru).
OpheomorphusbrachyurusCope,1887.SyntypesANSP11202-03.Type-locality—Chupada,Mato
Grosso,Brazil.
Liophis elaeoides Griffin, 1916. Holotype CM R32. Type-locality— Prov. del Sara, Bolivia.
LiophismacropsWerner,1925.HolotypeNMW23420.Type-locality—Paramaribo,Surinam.
Nomenclatural comments. — Several names have been assigned mistakenly to
the synonomy of L. typhlus. Jan (1863a, 1863Z?) and Jan and Sordelli (1866)

1 Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, Texas
77843-2258.
Submitted16January1987.

173
174 Annals of Carnegie Museum vol.56

suggested that L. typhlus was composed of three varieties in addition to the

nominate race, prasinus, gastrostictus, and olivaceus. I have determined that no
holotypes exist for any of these varietal names. An examination of Jan’s descrip-
tions and Jan and Sordelli’s illustrations (1866, Livr. 18, Plate IV, Fig. 3) suggest
that the name prasinus belongs to Liophis viridis. The illustration shows 69 sub-
caudals, considerably more than the maximum known number of 6 1 for L. typhlus,
but well within the known range of 63 to 84 for L. viridis. An illustration of a L.
typhlus by Jan and Sordelli (1866, Livr. 18, Plate IV, Fig. 2) from “Pernambuco,”
Milan Museum, may represent the variety gastrostictus, of Jan ( 1 863a, 18636).
The illustration shows 47 subcaudals, well within the known range of variation
in subcaudals for L. typhlus. However, the closest localities of L. typhlus to
Pernambuco are 1 300 km to the east and 800 km to the south. The illustration
shows dark smudges along the lateral edges of the ventrals, and scattered dark
scale edges on the dorsum as well. The illustration and description more closely
resemble L. poecilogyrus from Pernambuco, and I suggest gastrostictus belongs
to that species. The Jan name olivaceus is without description or illustration. It
appears in Jan’s (1863a, 18636) list and key to Liophis as a varietal name of L.
typhlus and should be considered a nomen nudum.
Another misapplied name is Wagler’s (1824) Natrix forsteri. Hoge (1964) first
applied it to L. typhlus and suggested that the Surinam race be recognized as the
nominate taxon and the Brazilian form (by inference) L. typhlus forsteri. The type-
locality of L. forsteri is Salvador, Bahia, Brazil, a locality of considerable distance
from any known sample of L. typhlus. In addition, the illustration and description
of forsteri suggests that this name is best applied to L. poecilogyrus. The description
lists 60 subcaudals for the type of L. forsteri. The greatest number of subcaudals
recorded in 73 eastern Brazilian individuals of L. typhlus is 48. The description
of the color pattern of forsteri is unlike that of L. typhlus, and more similar to
that of L. poecilogyrus. Hoogmoed and Gruber (1983) examined the holotype of
N. forsteri in the Munich Museum and gave the subcaudal number as 58/58+1.
Although they placed N. forsteri as a synonym of L. typhlus, I consider N. forsteri
to be a senior synonym of L. poecilogyrus.
Parker (1 928) on comparing one of the paratypes of L. guentheri (BM 1 946. 1 .5.69)
to samples of L. typhlus in the British Museum (Natural History), concluded that
it “failed to reveal any consequential differences between these two species” and
placed L. guentheri in the synonomy of L. typhlus. Parker erred in his analysis
because the ventrals of the specimen he examined (and L. guentheri in general)
vary from 187 to 197, while ventrals of L. typhlus vary from 133 to 172. Liophis
guentheri is a valid species that is more closely related to L. viridis than to L.
typhlus.
Distribution. —Liophis typhlus occurs in Amazonian rainforests to an elevation
of about 1500 m along the southern and eastern side of the Andes in Colombia,
Ecuador and Peru, the rainforests of southern and eastern Venezuela, and similar
forests of Guyana, Surinam and French Guiana, and Cerrado and Chaco forests
of SE Bolivia, N Paraguay, and SC Brazil (Fig. 1).
Description. — The maximum total length is 740 mm in males and 853 mm in
females. Body scales are smooth, without apical pits and typically in 19-19-15
rows. Scale rows 3 + 4 or 4 + 5 typically fuse on each side of the body between
ventrals 68 and 103 (x = 82.2). Scale rows 7 + 8 or 8 + 9 usually fuse between
ventrals 65 and 104 (x = 83.0). A sample of 48 females and 25 males from Brazil
and 49 males and 62 females from other Amazonian localities were analyzed for
sexual dimorphism in all quantitative characters and none was found. Ventrals
1987 Dixon— South American Liophis 175

Fig. 1.— Distribution of Liophis typhlus (dots) and L. guentheri (diamonds, stippled area) in South
America.Blacklinesenclosenumberedsamplesusedinthegeographicvariationanalyses.

of 331 specimens numbered 133-172 (x = 157.1), the number of subcaudals of

307 specimens varied from 41-61 (. x = 49.5). Tail length divided by total length
for 105 adult males varied from 0.145-0.206 (x = 0.169), and for 160 adult
females 0.137-0.210 (x = 0.162). Eye diameter divided by snout length of 127
adults varied from 0.524-0.886 (x = 0.682). Maxillary teeth for 222 specimens
varied from 19-28 (x = 22.4). The anal plate is divided and the loreal single in
all specimens examined.
Head shields vary as follows: supralabials 4-5 (1), 7-7 (3), 7-8 (2), 8-8 (249),
8-9 (5); supralabials entering orbit 4th only (1), 4 + 5 (252), 3+4/4 + 5 (2), 4 + 5/
5 + 6 (4), 4 + 5/4 + 5 + 6 (1); infralabials 5-7 (1), 8-8 (1), 9-9 (10), 9-10 (23), 10-10
(216), 10-11 (5), 11-11 (2), 11-12 (1); preoculars 0-0 (1), 1-1 (256); postoculars
1-1 (3), 1-2 (1), 2-2 (253), 2-3 (3); temporal condition 1 + 1 (3), 1+2 (242), 1 + 1/
1+2 (12), 1 +2/1+3 (3), 0 + 2/1+2 (1).
The in situ length for 69 hemipenes varied from 6 to 13 (x = 9.0) subcaudals.
The organ is slightly bilobed, with each lobe about two subcaudals long. The
sulcus spermaticus usually bifurcates about one-third the distance from the base
176 Annals of Carnegie Museum vol.56

of the organ. Large spines are present on the asulcate surface of the basal two
thirds of the organ, usually becoming much smaller on the lobed part of the organ.
A smooth apical disk is present on the outer edge of each lobe and a naked basal
pocket is usually present.
Color in life.— The dorsal color of some adults is light blue on the head and
various shades of green on the body and tail. Amazon samples have reddish
brown to black chevron marks over most of the lateral (occasionally middorsal)
surfaces of the body. These dark marks are usually prominent in young and
juveniles, less so in adults. Some individuals from throughout the range have
scattered white scales mixed with the green. Other individuals are uniform green
and lack dark chevron marks and blue heads. The venter is usually white or light
yellow without darker markings. However, a few individuals have dark smudges
or distinct marks ventrolaterally, and the subcaudals may have dark spots or lines.
The juvenile dorsal pattern is variable with some juveniles having distinct black
chevrons with a wide black nuchal band. The nuchal band usually fades to an
obscure greenish black mark at a total length of 210 mm. Other juveniles from
155-255 mm in total length have a pair of blackish nuchal spots that begin on
or at the posterior edge of the parietals, and slant posteroventrally. Occasionally
a secondary pair of medium sized dark spots occur on the neck, followed by two
rows of paravertebral dark spots and two rows of lateral dark spots to above the
vent. Sometimes, the body appears to be reticulated with darker lines on a greenish
ground color. One young individual (SVL 225 mm) has distinct spots on scale
rows one, two, four, six, and seven, and occasional dark spots along the edges of
the ventrals.
Geographic variation.— There are three distinct geographic populations of L.
typhlus that correspond to forest refugia postulated by Vanzolini and Williams
(1970), and to some extent, to those of Haffer (1974). The three geographic pop-
ulations were determined by comparison of the numbers of maxillary teeth, ven-
trals, subcaudals, the ratio of tail length to total length, and the ratio of the eye
diameter to snout length. Six samples were used to determine variation. The
samples were chosen by the proximity of individuals to each other and by natural
vegetation types, as follows: Sample 1-27 specimens from Surinam and French
Guiana; Sample 2-24 specimens from Guyana and Venezuela; Sample 3-53 spec-
imens from Amazonian Colombia, Ecuador, Peru, western Brazil, and Bolivia;
Sample 4-29 individuals from Amazonian Brazil; Sample 5—45 specimens from
the Chaco of Bolivia and western Mato Grosso, Brazil; and Sample 6 — 87 spec-
imens from the Cerrado of southeastern Brazil (Fig. 1). The Student’s T test was
utilized to determine significance of pair-wise comparisons between samples. A
Student’s T test value of 6.3 14 or greater is at 95% level of significance or greater.
Pair-wise tests for differences between samples 1 and 2, 1 and 4, 2 and 3, 2 and
4 were not significant for any of the characters listed above. Samples 1 and 3 were
significantly different from each other in number of ventrals and tail/total length
ratio. However, samples 1 and 3 are separated by sample 2, and sample 2 did
not differ significantly from either 1 or 3. The significance noted between 1 and
3 is probably an artifact of distance, with sample 2 representing the middle of a
cline. Samples 3 and 4 were not significantly different in any characters except
ventrals. This may also be an artifact of distance. However, two males from
reasonably close localities in Amazonian Bolivia had ventral counts of 140 and
157. Their ventral counts fall near the average of the samples with which they
were associated, 143.5 for sample 4, and 154.2, for sample 3 (see Fig. 1). This
suggests that a hiatus may exist between samples 3 and 4 in Amazonian Bolivia.
1987 Dixon —South American Liophis 111

A statistical analysis of ventral number between samples appears to show low to

high numbers of ventrals from samples 1 to 4, 1 to 2, 2 to 4, 2 to 3. The samples
form an almost closed circle of demes (Fig. 1), with the bottom of the circle open,
representing the differences in the Bolivian samples mentioned above. It seems
clear that the Amazon samples are closely related, but with a trend to divide the
samples into eastern and western demes.
Pair-wise comparisons of samples 4 and 6 suggest a strong differentiation in all
characters examined. The differences in the numbers of ventrals, subcaudals, and
tail/total length ratios are highly significant between the two samples, with T
values of 23.8, 19.1, and 15.8 respectively. The number of maxillary teeth differed
significantly (T value = 7.0) but eye diameter/snout length ratios did not (T value =
2.14). The data suggest that the Brazilian Amazon forest and deciduous meso-
phytic forest samples represent different allopatric taxa. Comparison of samples
3 and 6 shows the same trend, except the number of maxillary teeth is not
significant (T = 4.91). Pair-wise comparison of samples 4 and 5 shows significant
differences in the number of ventrals and maxillary teeth (T = 22.03, 10.54,
respectively), but not in the number of subcaudals, tail/total length, or eye di-
ameter/snout length ratios (T = 2.66, 4.59, 4.62, respectively). Pair-wise com-
parison of samples 3 and 5 shows significant differences in the number of ventrals,
maxillary teeth, and subcaudals (T = 9.07, 8.49, 6.36, respectively), but not in
the tail/total length and eye diameter/snout length ratios (T = 2.96, 6.0). The
relative differences between the Amazon samples (1-4) and the two non- Amazon
samples (5-6) suggest that the Chaco forest sample from Bolivia is more closely
related to the Amazon forest sample than to the deciduous mesophytic forest
sample of Brazil. However, both non-Amazon forest samples are very distinct
from the Amazon forest samples.
A pair-wise comparison of samples 5 and 6 shows significant differences in the
number of subcaudals, maxillary teeth, and tail/total length ratios (T = 21.05,
6.43, 12.9, respectively), but not in the number of ventrals or eye diameter/snout
length ratio (T = 1.36, 3.59). These samples represent different taxa, even though
they are relatively close geographically (Fig. 1).
The scale count data and length ratios suggest three distinct populations of
Liophis typhlus. Samples 1, 2, 3, and 4 represent the Amazon forest form, sample
5 the northern Chaco form, and sample 6 the Cerrado form.
The color patterns of samples 1, 2, 3, and 4 are similar, and the patterns of
samples 5 and 6 are identical. The dorsum and head of adult individuals from
samples 5 and 6 are chlorophyll green. The young and juveniles have black nape
spots and some have four rows of dorsal dark spots. Adults of the Amazon samples
have a leaf green to bright green dorsum with diagonal reddish brown to blackish
chevron marks along each side of the body. The head may be greenish to bright
blue. Young and juveniles are greenish dorsally with bold dark chevron marks
and a relatively large black nuchal blotch.
Diagnoses and distributions of the three taxa follow:

Liophis typhlus typhlus (Linnaeus)

ColubertyphlusLinnaeus,1758.
XenodonisolepisCope,1870.
LiophismacropsWerner,1925.
Diagnosis. — Dorsum usually green with distinct, reddish brown to black chev-
rons on each side, occasionally fading posteriorly. The dorsal surface of the head
178 Annals of Carnegie Museum vol.56

may be greenish to bright blue. The venter is white to light yellow, usually without
dark markings. Juveniles have a large black nuchal blotch and dark chevrons.
The number of ventrals varies from 1 33 to 1 63 (x = 147.3), subcaudals vary from
47 to 61 (x = 54.2), maxillary teeth vary from 20 to 28 (x = 23.6), tail/total length
ratios vary from 0.160 to 0.210 ( x = 0.184), and eye diameter/snout length ratios
vary from 0.510 to 0.861 (x = 0.736).
Distribution. — Throughout the Amazon Basin in rainforest, to about 1000 m
elevation (see specimens examined).

Liophis typhlus elaeoides Griffin

LiophiselaeoidesGriffin,1916.
Diagnosis. — Dorsally adults are uniform chlorophyll green with a light yellow
venter. Juveniles have four rows of dorsal dark spots on a green background, and
a pair of black nuchal spots. The number of ventrals varies from 158 to 172 (x =
163.5), subcaudals vary from 49 to 56 (x = 51.9), maxillary teeth vary from 17
to 23 (x = 19.9), tail/total length ratios vary from 0.160 to 0.200 (x = 0.171),
and eye diameter/snout length ratios vary from 0.520 to 0.740 (x = 0.620).
Distribution. — Liophis typhlus elaeoides is known only from the upper Rio
Paraguay Basin, which includes the mesic Chaco forests of southeastern Bolivia,
northern Paraguay, and western Mato Grosso, Brazil (see specimens examined).

Liophis typhlus brachyurus (Cope)

OpheomorphusbrachyurusCope,1887.
Diagnosis. — Adults are uniform green dorsally with a light yellow venter. Ju-
veniles have four rows of dark dorsal spots on a green background and a pair of
black nuchal spots. The number of ventrals vary from 151 to 171 (x = 162.2),
subcaudals vary from 40 to 49 (x = 44.4), maxillary teeth vary from 18 to 24
(x = 21.5), tail/total length ratios vary from 0.140 to 0.160 (x = 0.149), and eye
diameter/snout length ratios vary from 0.55 to 0.81 (x = 0.675).
Distribution.— Liophis t. brachyurus occurs in deciduous mesophytic forests of
southeastern Brazil, and in the Campos Cerrado forests of east-central Brazil (see
specimens examined).
Comments.— Two recent papers by Miranda and Couturier (1983, 1984) com-
ment on the presence and geographic variation of L. typhlus in Argentina. I have
neither examined their specimens, nor located specimens of L. typhlus from
Argentina in other museums. I believe they may have erred in identification of
their specimens. Photographs presented by Miranda and Couturier (1984) suggest
that they may have confused L. miliaris and L. poecilogyrus with L. typhlus.
Short works on the distribution and taxonomy of L. typhlus are Hoge (1964),
Amaral (1931, 1935, 1949), Gans (1960), Parker (1928, 1935), Peters (1960),
Peters and Orejas-Miranda (1970), Roze (1966), and Prado and Hoge (1948).
Peters (1963) compared the maxillary teeth of L. typhlus with those of other
species of Liophis.
Specimensexamined.—(Liophistyphlustyphlus)BOLIVIA,Beni:IxiamusAMNH22457;RioBeni
AMNH 22270. BRAZIL, Acre: Alto Purus MZUSP 2498; Porto Walter MZUSP 7390. Amapa:
Cuidade Oiapoque IB 13780, 13783; Serro do Navio KU 97873-97875. Amazonas: Barreira do
Matupiri, on the Rio Madeira MZUSP 5912; Carvoeiro AMNH 36167; Costa Altamira on the Rio
Japura MZUSP 6600; Lago Alexo MCZ 3290; Manaus MZUSP 3051, 3797; Reserva INPA MZUSP
7606, 7619; Santa Isabel on the Rio Negro USNM 83532; Sao Paulo de Olivenca AMNH 53311;
1987 Dixon— South American Liophis 179

Tapaua MZUSP 5770; no specific locality UZM 601223. Maranhao : Aldeia Aracu Igarape Gurupi-
Una MZUSP 4303, 4826. Para: As Pedras, on the Rio Cumina-Miri MZUSP 5103; Belem IB 15667;
Caninde,ontheRioGurupiMZUSP4267,4285;nearMaraba,SerrandoNorteKU124608;Oxiximina
MZUSP 4796; Urua, Parque Nac. da Amazonia, on the Rio Tapajos MZUSP 7838. Rondonia: Porto
Velho MZUSP 3690. COLOMBIA, Macanal : Rio Garagoa MCZ 27339. Meta : Villavicencio IB 8589.
Vaupes: Timbo UTA R3805; Yapima UTA R5033. ECUADOR, Napo-Pastaza : Alpayaca FMNH
4069, UMMZ 89020-22; Andoas AMNH 41949; Banos+Canelos AMNH 35892; Chamala-Nor-
mandia-Rio Bamba AMNH 35929-35930; Jarayacu AMNH 28796; Macas and Vicinity AMNH
35838-35839; 82 km ESE Macas AMNH 114616; Rio Bamba AMNH 15209, 15213, 23296, 23302,
28848; Rio Pastaza MCZ 36966; Santa Rosa, El Tigre AMNH 49165. FRENCH GUIANA (CAY-
ENNE), Mana USNM, 6172; Saul MCZ 149399; near Sophie MCZ 77510-775 1 1 . GUYANA, Bartica-
Potaro road BM 1954.1.3.64; 82 km S Bartica BM 1934.11.1.125; Berbice BM RR1964; Demerara
River, Lama Creek AMNH 36106; Essequibo, near Lethem USNM 146376; Kartabo AMNH 21335,
98197-98198; Matali AMNH 61542; Rupununi District, north of Acarai Mountains, west of New
River KU 69826-69828; Rupununi District, north of Acarai Mountains, west of New River KU
69826-69828; no specific locality MNHG 279.38. PERU, Amazonas : Caterpiza, Rio Caterpiza USNM
(RWM field series) 14995, 15033, 1 4 185-1 5 1 86. Junin: Rio Perene MCZ 42434. Loreto: BaltaLSUMZ
14584: Centro Union TCWC 44682; Cerros de Contaya, on the Rio Tapiche AMNH 53376; Con-
tamana on the Rio Ucayali AMNH 52130; Estiron on the Rio Ampiyacu MZUSP 4394; Iquitos
AMNH 52734; 53118, 53284, 53667, 53696, 53735, 53771, 53773, 53876, 53923, 53949, 54321,
54354, 54483, 54894, 56109; Moropon TCWC 38049, 44294; Orellana (Reforma) AMNH 54957,
USNM 127124; Pampa Hermosa on the Rio Cushabatay AMNH 55409, 55415, 55442, 55791, 55885,
56003; Pamya AMNH 53249; Requena (Monte Carmelo) AMNH 55600, 55626; Royaboya AMNH
52483, 53110, 53288, 55695; San Antonio on the Rio Itaya AMNH 52920, 53667, 53693, 53696,
53735; Shiriara, on the Rio Nanay AMNH 56075; Trapiche-Utuquinia AMNH 52195; Yanamono
TCWC 40542-40543. Madre de Dios: Rio Heath, 50 km S Puerto Pardo LSUMZ 36778-79; mouth
of Rio Torre on the Rio Tambopata LSUMZ 394244. Pasco: Iscozazin Valley LACM 76805. San
Martin: Moyobamba BM 74.8.4.59, 94.8.4.64. SURINAME: Charlesburg AMNH 104624; Coppen-
ame River MCZ 1 52203, 1 52205, 1 52633-1 52634; Jaraweg TCWC 60543, 60756; Paramaribo AMNH
8146, 8682, MCZ 16401; Potribo CM 44302; Sipaliwini Airstrip CM 84667, MCZ 152636; Zanderi
(Airport) MCZ 152635. VENEZUELA, Amazonas: Arabopo UMMZ 85279; Arocoima Canos MCZ
38541; Mount Duida region AMNH 36617, 36620. Bolivar: El Manteco TCWC 60168; no specific
locality MHNLS 1632.
(. Liophis typhlus elaeoides ) BOLIVIA: Cochambamba: Villa Tunari UMMZ 153095. Santa Cruz:
Buenavista CM R2696, R2698, R270 1 , R2704, R2869, R2886, R2930, R293 1 ; UMMZ 67967-67968,
67969(2), 67970(3); Buenavista, near Rio Colorado CM R2860, R2865, R2955; Santa Cruz LSUMZ
11825; Santa Cruz de la Sierra CM R32 (holotype), R44, R59, R91, R94, R95, R97, R98, R99, R102,
MCZ 11860, MZUSP 6474; San Jose de Chiquitos CM 34842. No specific locality: CM R2938-
R2939, FMNH 195898, HCD 2820, 2822, TCWC 55290. BRAZIL: Mato Grosso: Corumba, near
Urucum Mountains CM 3484 1 ; Fazenda Bela Vista of the Ilha Insua MZUSP 7264; Fazenda Vacurizai
of the Rio Paraguai MZUSP 7266; Generalso Ponce, Corumba IB 25954; Maleta IB 14975; Porto
MurtinhoIB26177-78.
(. Liophis typhlus brachyurus ) BRAZIL: Bahia: Mira Serra, 41 km from Morro de Chapeo MZUSP
7554. Goaiz: Araguari IB 6851; Fazenda Lucushac, on the Rio Verde IB 13060. Mato Grosso: Arapua
IB 9939; Aquidauana MZUSP 33589; Chapada ANSP 11202-11203, BM 92.420. 1 3; Taugara da Serra
IB 24543; Taunay IB 7674; Urucum, near Corumba BM 1928.1.12.3. Minas Gerais: Horto Forestal
IB 10493; Irma Badur IB 6933; Itambe do Mato Dentro MZUSP 8061; Juiz de Fora IB 25190, 26685;
Machado IB 16303; Santa Rita do Extrema IB 5540, 5562. Rio de Janeiro: Nova Friburgo IB 10516.
Sao Paulo: Araraguara IB 231; Atibaia IB 21376, 27081, 27201; Barra Assugnui IB 29042; Barretos
IB 5030; Baureri IB 7212; Bauru MZUSP 823; Boraceia MZUSP 4227; Botucatu MZUSP 2410, 2412;
Buri IB 6718, 6763, 12313; Campo Limpo IB 805, 6452, 10265-10267, 24903-24904; Capao Bonito
IB 23465; Cascavel MZUSP 824; Conde do Pinhal IB 9975; Cotia IB 5743, 6935, 7140, 9906, 19906,
23236, 23377, 24542, 32627; Curupa IB 7061; Dona Catarina IB 22598; Elias Fausto IB 232, 319,
392, 578, 791, 10461; Elihu Root IB 234; Engenheiro Cesar de Souza IB 8338; Ferraz de Vasconcelos
MZUSP 2487; Guarulhos IB 23726-23727; Horto Florestal IB 10493; Ibiuna IB 12252, 23794;
IpanemaIB230,235;ItaguaIB4813;ItapecericadaSerraIB7794,19682,25040,25043;Itapetininga
IB 24565; Itapolis IB 7956; Itaquera IB 30758; Itaquaquecetuba IB 18349; Itirapina IB 6655; Jaragua
IB 21381; Jarinu IB 10292; Lauro Muller IB 10533, 10556, 10559; Leme IB 236, 547; Mairinque IB
7373, 7461; Mairipora IB 28603; Mogi das Cruzes IB 12592, 19658, 23547, 24881, 28284, MCZ
39415, MZUSP 6460; Osasco IB 12500, 27910-27911; Pedemeiras IB 6938; Pindamonhangaba IB
1257; Piraju MCZ 39416; Promissao IB 9969; Ribeiras Pires IB 19596; Rio Claro IB 7277, 10296;
180 Annals of Carnegie Museum vol.56

Santa Adelia IB 6909; Santa Emestina IB 27969, MZUSP 2479; Sao Carlos IB 10016; Sao Jose do
Rio Pardo IB 5444; Sao Jose dos Campos IB 21348; Sao Paulo IB 797, 7142, 12980, 12993, 16223,
16900, 18854, 25295, 27497, 28458, MCZ 17956, MZUSP 817-820, 2388, 3338-3339, 3695, USNM
69238-69239; Suzano IB 19104; Valinho IB 6719; Vinhedo IB 2915; no specific locality IB 233, 237,
797, 840, 4745, 5491, 5642, 6950, 9775, 10581, 12112, 17777.

Liophis guentheri Peracca

LiophisguentheriPeracca,1897.Sixsyntypes(Fivelost?),Mus.Comp.Anat.,Torino.Typelocality—
Caiza,Bolivia.
Comments. — Liophis guentheri has resided as a synonym of L. typhlus. Parker
(1928) compared one of the syntypes (now BM 1946.1.5.69) with a specimen of
L. typhlus , and suggested it “failed to reveal any consequential differences between
the two species.” However, no overlap in the number of ventrals occurs between
the two species. An unnumbered British Museum specimen from an unspecified
locality in Bolivia is similar to the numbered syntype in the collection of the
British Museum in its state of preservation, and may well be one of the five
missing syntypes that were housed in the Museum of Zoology and Comparative
Anatomy in Torino.
Distribution.— Liophis guentheri apparently is restricted to the central Chaco
of Bolivia, Argentina and Paraguay (Fig. 1).
Description. — The following description is based on six adult females and three
adult males. The maximum total length of males examined is 655 mm, of females
723 mm. Peracca (1897) gave 890 mm as the maximum total length but did not
indicate the sex. Scale rows are 19-19-15, smooth, without apical pits; a reduction
of the third and fourth scale row occurs between ventrals 98-118 (x = 106.9),
and reduction of the seventh and eight scale rows between ventrals 99-112
(x = 106.1). The number of ventrals varies from 187 to 197 ( x = 191.6), the
number of subcaudals from 53 to 57 (x = 54.6) (only five with complete tails).
The number of maxillary teeth varies from 20 to 22 (x = 20.9). Head scales are
as follows: supralabials 7-8 (1), 8-8 (7), 8-9 (1); supralabials entering orbit 4 + 5
(8), 4 + 5/5-F6 (1); infralabials 9-9 (1), 9-10 (1), 10-10 (5), 10-11 (2); preoculars
1-1, postoculars 2-2, loreals 1-1, temporals 1 + 1 (1), 1 +2/1 + 1 (1), 1+2 (7). The
anal plate is divided. The tail/total length ratio varies from 0.152-0.168 (x =
0.159); eye diameter/snout length ratios vary from 0.467 to 0.578 (X = 0.502).
The hemipenis is nine subcaudals long, with the sulcus spermaticus dividing
at the level of the third subcaudal, slightly bilobed (about one and a half subcau-
dals), and very spinose. The smooth apical disks are pronounced.
The general dorsal color is light blue in preservative, but Peracca (1897) states
that living specimens are immaculate green dorsally and yellowish white ventrally.
The ventrals and subcaudals are usually immaculate white in preservative. In two
preserved specimens there are dark flecks and/or spots on the outer edges of the
ventrals. Flecks cover all ventrals in one specimen and three fourths of the ventrals
in the other.
Specimens examined.— ARGENTINA: Salta : Hickmann IML 105, 303; Luna Muerta IML 301;
Formosa : Ingeniero Juarez IML 422, 490. BOLIVIA: no specific locality IB 1177, BM (unnumbered);
Chaco : Caiza BM 1946.1.5.69. PARAGUAY: Chaco : Loma Plata KU 73454.

Liophis viridis Gunther

Liophis viridis Gunther, 1862. Holotype, BM 1946.1.5.69. Type-locality— Brazil, Pernambuco, no
specific
locality.
LiophistyphlusprasinaJanandSordelli,1866.Holotype(lost?).Type-locality—Brazil.
1987 Dixon— South American Liophis 181

Fig.2.—DistributionofLiophisviridisinnortheasternBrazil.Dotswithinthediagonallines(Agreste
plantformation)andlightlystippledarea(Atlanticrainforestformation)representtheAgreste-Atlantic
Rainforest sample. Dots within the dashed line (Caatinga plant formation) represent the Caatinga
sample.DenselystippledareaswithintheCaatingaformationrepresentenclavesoftheCerradoplant
formation.

Distribution.— Known only from the Caatinga, Agreste, and Atlantic rainforest
vegetation communities of northeastern Brazil (Fig. 2).
Description. — The following description of L. viridis is based on 1 59 individuals
from throughout the range. The maximum total length of males is 595 mm, and
of females 670 mm. The scale rows are 19-19-17, smooth, and with one apical
pit. The reduction to 1 7 rows occurs through a fusion of scale rows three and four
between ventrals 98 and 121 (x = 110.0). Sexual variation was not detectable
even in large samples, therefore all scale data were combined for intrasample
comparisons. Ventrals vary from 169 to 202 (x = 186.0); subcaudals from 63 to
84 (x — 74.1). Tail/total length ratios of adults vary from 0.193 to 0.249 (x =
0.221). The eye/snout ratios of adults vary from 0.500 to 0.684 (x = 0.562).
182 Annals of Carnegie Museum vol„56

Maxillary teeth vary from 17 to 24 (x = 20.2), with the last two teeth enlarged
and separated from the remainder by a decided gap. The anal plate is divided in
all specimens examined.
Head scales vary as follows: supralabials 8-8 (156), 8-9 (2), 9-9 (1); supralabials
entering orbit 4 + 5 (155), 4 + 5/5 + 6 (3), 4 + 5 + 6/5 + 6 (1); infralabials 8-9 (2),
8-10(1), 9-9 (2), 9-10(4), 10-10(144), 10-11 (4), 11-11 (2); preoculars 1-1 (156),
1-2 (2), 2-2 (1); postoculars 1-1 (1), 2-2 (157), 2-3 (1); loreals 1-1 (159); temporals
1 + 1 (12), 1 + 1/1+2 (21), 1+2 (145), 1 +2/1+3 (1).
The hemipenis is 9 to 13 (x = 10.8) subcaudals long (in situ) and bilobed for
1.5 to 3 subcaudals. Large spines occur from the base to the proximal part of the
lobe. Spines about one half the size of the basal spines occur from the lobe to the
distal end. The sulcus spermaticus is divided near the base of the lobes. The
smooth apical disk is relatively large.
The dorsum is bright green in life. Young and small juveniles may have black
bands in the nape region followed by a series of black spots arranged in four linear
rows, primarily along scale rows three and four, and seven and eight. The number
of spots is variable in any row, varying from 36 to 60, especially in individuals
from 180 to 220 mm in total length. Some young have only a nape band, im-
mediately followed by one pair of dorsal spots. Occasionally young have a black
lateral tail stripe. Spots on juveniles from 240 to 370 mm in total length are faded
posteriorly. Normally all individuals over 400 mm in total length are uniform
green and the belly and subcaudals are immaculate white. Vanzolini et al. (1980)
give a very good color description of living individuals of this species.
Geographic variation.— A Caatinga sample and an Agreste- Atlantic rainforest
sample were formed from three coastal subsamples and two inland subsamples
of L. viridis (Fig. 2).
The Caatinga sample is significantly distinct (T = 13.4) from the Agreste-
Atlantic rainforest sample in having a higher number of ventrals, 181-202 (x =
189.8), versus 169-188 (x = 179.6), and a higher number of ventrals (T = 6.44)
at the 19-17 reduction site 102-124 (x = 1 14.6) versus 98-1 16 (x = 106.6). Other
scale and tooth characters are not significantly different between samples.
One specimen from Jeremoabo, Bahia, and five specimens from Mina de Sao
Felix do Amianto, Bahia, are known from the eastern part of the Caatinga. Their
taxonomic features more closely approximate those of the Agreste-Atlantic rain-
forest sample than others from the Caatinga. Although the samples are small I
suspect that they may represent intergrades between two well-defined subspecies.
The Caatinga sample (Fig. 2) does not have a formal name. However, Jan
(1863a, 1983/?) and Jan and Sordelli (1866) mention a varietal name associated
with L. typhlus that belongs to L. viridis. It was originally cited as a varietal name
(without description) by Jan (1863a, 1863/?) but was illustrated by Jan and Sordelli
(1866) and their iconotype represents the only description. Jan (1863a, 1863 b)
recorded two specimens of the variety, one from “Pernambuco” in the Milan
Museum, and one from “Brasile” in the Stuttgart Museum. Jan and Sordelli
illustrated the latter specimen, thus it becomes the holotype (by inference). How-
ever, neither specimen now can be located. I cannot definitely assign the name
to either of the two populations that require recognition. However, the genus is
already overburdened with excessive synonyms and I propose that Jan and Sor-
delli’s varietal name, prasinus, as proposed in the combination L. typhlus prasinus,
be recognized as the proper name for the Caatinga popultion of L. viridis.
1987 Dixon— South American Liophis 183

Liophis viridis viridis Gunther

LiophisviridisGunther,1862.
Diagnosis. — Dorsally adults are leaf green to bright green, and pale cream to
white ventrally. Juveniles have a black nape band followed by a various number
of paired black dorsolateral spots. The number of ventrals varies from 169 to 188
(x - 179.6), and reduction occurs between ventrals 98 and 1 16 (x = 106.6).
Distribution. —Liophis v. viridis occurs in the Atlantic rainforest and the Agreste
forest from Recife, Pernambuco, east and south to Salvador, Bahia, Brazil.

Liophis viridis prasinus Jan and Sordelli

LiophistyphlusprasinaJanandSordelli,1866.
Diagnosis. — Adults are bright green, with an immaculate cream to white venter.
Juveniles have bright green dorsums with a single black nape band, or with four
parallel rows of 35 to 60 black spots per row. Occasional juveniles have a pair of
posterior, lateral black stripes that extend onto the tail. The number of ventrals
varies from 1 8 1 to 202 (x = 189.8), with the reduction occurring between ventrals
102 and 124 (x = 114.6).
Distribution. —Liophis v. prasinus occurs from Joao Pereira, Maranhao in north-
eastern Brazil, south to Sao Francisco, Minas Gerais, then northeast to about
Parauagua, Rio Grande do Norte, Brazil.
Comment.— Yin (1983) found Hyla rubra, Physalaemus cuvieri, and several
unidentified leptodactylids in 14 stomachs of L. viridis. Vitt examined 16 gravid
females and determined that L. viridis produces continuous clutches throughout
the year. Ovarian clutch size varied from two to six (x = 3.69) eggs.
Short taxonomic works for L. viridis include Cordeiro and Hoge (1973), Gomes
(1918), and Schmidt and Inger (1951). Vanzolini et al. (1980) give an excellent
summary of scale data, color of young and adults, and brief biological notes on
a Caatinga sample.
Specimens examined.—" SOUTH AMERICA”: CM R281. BRAZIL: no specific locality FMNH
5694, MZUSP 815, NHMW 20739; no specific state, Parauagua ZMUC 601250. Algoas : Quebrangulo
MZUSP 3453. Bahia: No specific locality FMNH 5721, IB 3200, MCZ 2946, 147217-147218. Assu
da Torre IB 980, Brreiras UMMZ 108712-108716, Bom Jesus de Lapa Cm R344, UMMZ 108711,
Feiro de Santana KU 29478, Itapoan MZUSP 5713, Jeremoabo MZUSP 5738, Mina de Sao Feliz
do Amianto IB 28170-28172, 28215-28216, Salvador BM 90.1.27.2, 1924.9.20.26, MZUSP 2838-
2839. Gear a: Acude Amanari Maranguape MZUSP 3450, Boa Viagem IB 20200, Caxueira ZMUC
3342(3), Coluna MZUSP 53 1 9-20, Limoeiro do Norte IB 1 2776-12777, Unique IB 20142. Maranhao:
Lagoa de Joao Pereira CM R316. Minas Gerais : Januaria UMMZ 108705-108709, Sao Francisco
UMMZ 108710. Parahyba: Coremas MZUSP 3451, Joao Pessoa MZUSP 7976-7977, 7999, 8262,
UFFB 139, 141, 143, Mamanguape MZUSP 3456. Pernambuco: no specific locality MANH 4448,
BM 1946.1.2.69, 80.1 1.25.4, 80.1 1.25.8, MCZ 1447, 146945-146947, Agrestina MZUSP 4925, 4942,
4946, 4965, 4970, Exu MZUSP 6693-6699, 6700-6718, 7071-7091, 5 km E Exu MZUSP 6950, 13
km Exu MZUSP 6940-6943, 5 km N Exu MZUSP 6920-6921, 1 1 km S Exu MZUSP 7092, Ponta
de Pedras MZUSP 5177, Recife DEH 624, 517, 526, LSUMZ 36787. Piaui: Acude Peri-Peri MZUSP
3424, 3427-3429, 3433, 3435-3438, 3441, 3443, 3445, Terezina IB 360, 473, 1210-1214, Valenca
MZUSP 5814. Rio Grande do Norte: Ceara Mirim CAS 49320. Sao Paulo: Leme IB 543 (in error).
Sergipe: Aracaju IB 22458, Maruim ZMUC 601251, Santo Amaro das Brotas MZUSP 6985, 7306.

Liophis jaegeri (Gunther)

Coronella jaegeri Gunter, 1858. Syntypes—BM 1946.1.9.12, 1946.1.5.78. Type-locality— Brazil, no
specific
locality.
184 Annals of Carnegie Museum vol.56

Fig. 3.— Distribution of Liophis jaegeri in southeastern South America. Black lines enclose samples
utilizedintheanalysesofgeographicvariation. The large black dot represents several overlapping
localities.

LiophisdorsalisPeters,1863.Type-locality—Brazil,nospecificlocality.
AporophiscoralliventrisBoulenger,1894.Type-locality—islandnorthofConcepcion,nearSanSal-
vador,northParaguay.
Rhadinaea lineata Jensen, 1900. Type-locality: Taboleiro Grande, Minas Geiras, Brazil.
Misapplied synonym. —Rhadinaea dichroa Werner (1899) has been in the syn-
onomy of L. jaegeri since 1972 (Peters and Orejas-Miranda, 1972). However, an
examination of the type description reveals that the holotype has 19 midbody
scale rows, 159 ventrals, and 50 subcaudals. The dorsum is brown with the bases
of the scales white and the posterior edges black. It also has a tail/total length
ratio of 0. 1 85. The description is identical to that for Liophis poecilogyrus caesius,
a form common to the Chaco of Bolivia, Argentina, Paraguay and Brazil, and I
assign it to that species.
Distribution. — Liophis jaegeri occurs from about 1 9°S latitude in Brazil, to about
35°S latitude in Uruguay and Argentina, west to about 61°W longitude, along the
Rio Paraguay basin in Argentina, Paraguay and Brazil (Fig. 3).
Description. — Some 415 individuals of L. jaegeri were examined and full data
taken on 160. Tail/total length ratios were recorded for over 400 individuals. The
maximum total length of males is 539 mm, of females 676 mm. Dorsal scales
1987 Dixon —South American Liophis 185

are smooth, in 17 rows, normally without reductions, and without apical pits.
When reductions are present ( 1 5 individuals), they occur with a fusion of scale
rows three and four on one or both sides of the body, reducing to 1 5 or 1 6 between
ventrals 88 and 141 (x = 121.2).
Sexual variation in the number of ventrals, subcaudals, maxillary teeth, and
tail/total length ratios is absent, therefore the samples were combined for statistical
analysis. The number of ventrals of 160 individuals varies from 146 to 169 (x =
157 . 5 ), and subcaudals from 52 to 75 (x = 61.7). The number of maxillary teeth
varies from 22 to 29 (x = 25.6). The tail/total length ratios varies from 0.181 to
0.268 (x = 0.221). The number of palatine and pterygoid teeth were recorded for
one individual and they were 10 and 25 respectively.
Head scales are as follows: supralabials 6-7 (1), 7-7 (1), 7-8 (4), 8-8 (147), 8-9(4),
9-9 (1); supralabials entering the orbit 3 + 4 (1), 3 + 4/4 + 5 (3), 3 +4 + 5/4+ 5 (1),
3+4 + 5/4 + S + 6 (1), 4 + 5 (148), 4 + 5/4 + 5 + 6 (2), 4+5 + 6 (1); infralabials 8-8
(3), 8-9 (4), 8-10 (2), 9-9 (5), 9-10 (24), 9-11 (1), 10-10 (112), 10-1 1 (5); preoculars
1-1 (155), 1-2 (2), 2-2 (1); postoculars (2-2) and loreals (1-1), temporals 1 + 1 (5),
1-1/ 1-2 (11), 1 +2/2+1 (1), 1+2 (141); eye diameter/snout length ratios of 20
adults varies from 0.533 to 0.844 (x = 0.655). The anal plate is divided in all
specimens. Hemipenial length varies from 7 to 13 (x = 10.3) subcaudals.
A hemipenis extending 12 subcaudals has the sulcus spermaticus divided at
the level of the 6th subcaudal, and the lobes begin at the level of the 9th subcaudal.
Large spines occur on the asulcate side of the hemipenis to the edge of the large,
smooth, apical disk. Calyces appear to be absent and only a weak basal naked
pocket is present.
The general dorsal color is dull green, olive green, or olive brown. The venter
normally is rose or coral red, with or without lateral dark marks on the edges of
the ventrals. In life there is a reddish brown to olive brown stripe covering scales
rows 8 through 12, and frequently parts of scale rows 7 and 13. The stripe varies
in width from 5 to 10 rows of scales. Occasionally dark brown spots occur on the
posterior ede of rows 3, 4, 5, and sometimes along scales rows 7 and 8. The upper
and lower lips, throat and anterior ventrals may be cream or yellowish. Color
descriptions also are presented by Miranda et al. (1982).
Geographic variation.— 1 60 individuals were grouped into nine geographic sam-
ples. Three samples were located in the Rio Paraguay basin in a north/south line,
and six samples were scattered from Uruguay to southeastern Brazil, more or less
in a north/south line. Using univariate statistics, the number of ventrals, maxillary
teeth, subcaudals, and tail/total length ratios were analyzed for intra and inter-
sample variation. The two numbers that follow a mean value are the standard
deviation and standard error, respectively. In addition, Student’s T Test values
were used to determine significance at the 95% level, in a pair-wise sample com-
parison. No significant differences in pair-wise comparisons exist for any of the
samples arranged in the two north to south lines. There are trends in both lines
to increased number of ventrals and subcaudals from north to south. The trend
for maxillary teeth is reversed, with numbers decreasing from north to south.
Pair-wise comparisons of the Student’s T Test values for the number of ventrals,
maxillary teeth and tail/total length ratios of eastern and western samples were
not significant. However, a significant difference between eastern and western
samples in the number of subcaudals exist. Subcaudals of the combined three
western samples (N = 18) vary from 63 to 75 (x = 68.5, 3.8, 0.9), while the
combined six eastern samples (N = 1 10) vary from 52 to 71 (x = 60.4, 3.4, 0.3).
186 Annals of Carnegie Museum vol.56

the tail/total length ratios of the western samples vary from 0.214 to 0.268 (x =
0.231, 0.29, 0.07) and those of the eastern samples from 0.193 to 0.248 (X =
0.221, 0.13, 0.01). This suggests that the subcaudals are smaller in the western
sample, hence more subcaudals are present on a tail about the same length as in
the eastern sample. The eastern and western samples are separated by the Rio
Uruguay and Rio Parana basins and appear to be ailopatric (Fig. 3). However,
additional collecting may reveal parapatry of the samples.
Since the number of subcaudals is significantly different between the two sam-
ples, I propose recognition of the following taxa:

Liophis jaegeri coralliventris Boulenger

Diagnosis.— Dorsum ground color leaf green, sometimes with a middorsal
brownish red stripe five to six scale rows wide, extending from the nape to the
tail, and with a dark brown line on scale row three on each side of the body.
Venter rose to red, with or without ventrolateral blackish marks on the edges of
the ventrals. Subcaudals vary from 63 to 75 (x = 68.5).
Distribution.— Known only from the Rio Paraguay Basin of Argentina, Para-
guay, and Brazil (Fig. 3).

Liophis jaegeri jaegeri (Gunther)

LiophisdorsalisPeters,1863.
RhadinaealineataJensen,1900.
Diagnosis. — Dorsum olive green, grayish green or leaf green, often with a mid-
dorsal brownish or reddish stripe six to eight scale rows wide, sometimes with
dark dots along scale row four, and occasionally along scale rows three and five.
Venter rose or reddish, with or without ventrolateral black marks on the edges
of the ventrals. Subcaudals vary from 52 to 71 (x = 60.4).
Distribution. — Known only from the east side of the Rio Parana Basin east to
the Atlantic coast, and from Uruguay north to 1 9°S latitude in Brazil (Fig. 3).
Comment. —Little is known about the natural history of this species. I have
found 7 and 9 eggs in two specimens, and Miranda et al. (1982) indicated that
the average number of eggs for this species is about 1 4. Miranda et al. also indicated
that the species is diurnal, frequently found in humid places, and eats small frogs.
Notations on some field tags suggest that the species also occurs in brushy areas,
has been found “crossing dirt roads” and “in a puddle of a cow track in low
campo, 10:30 am, 84°F, and in full sun.”
Dixon (1985) presented evidence that L. jaegeri and L. maryellenae hybridize
in an area near Belo Horizonte, Minas Gerais, Brazil. Additional material from
the latter area has not been located, and the hybrid zone remains undefined.
Specimens examined.— (L. jaegeri coralliventris). ARGENTINA, Corrientes : Gral Paz, esteros de
la Santa Lucia IML 802, Ituzaingo IML 1 180. Rosario : Rosario MNHP 1899.6-7. Santa Fe: Banados
del Rincon CM 39130, Buenos Aires MNHP 676-50, Laguan Guadalupe CM 39128. BRAZIL, Mato
Grosso : Corumba IB 26575, Rio Apa UMMZ 108761, upper Rio Parana/Rio Paraguay UNM 68831.
PARAGUAY, Asuncion BM 1930.11.27.208, Buenavista, NE of Caaguarzu MCZ 34204, Isla Con-
cepcion BM 1946.1.5.85, Primavera BM 1960,1.2.84-85, 1960.1.3.40, 1962.81.
(. Liophis jaegeri jaegeri). BRAZIL, Minas Gerais : no specific locality MCZ 20715-20716. Belo
Horizonte UMMZ 109069, Campo Alegre IB 196, Coronel Fabriciano IB 22993, Imbuzeiro USNM
100676, Itapeva IB 41 126-41127, Juiz de Fora IB 30728, 33501, Mutum IB 3802, Nova Badem IB
192, Ouro Preto IB 897, 331 1, Passa Vinte IB 4968, Pocos de Caldas IB 4714, Santa Rita de Extrema
IB 5560, 8192, Taboleiro Grande ZMUC 601253, Toledo IB 4453 1, Uba IB 17308. Parana: Araucaria
IB 7252, 7346, 13632, Balsa Nova IB 18293, 2939-22941, 22943-22944, 23112, 23114-12115,
1987 Dixon— South American Liophis 187

24343-24345, 24466, 26226, 28785, 32538, Campo Largo AMNH 102257, Carambei IB 7403, Castro
IB 6244-6247, 7931, 10392, Curitiba UMMZ 108722, IB 197, 23006, 23439-23440, 29321, Dorizon
IB 893, Guarapuava IB 24630, 24661, 24710, 24889, 26954. Jaguariaiva IB 17842, Joaquim Murtinho
IB 8263, Joao Eugenio IB 13027, 15700, Mallet IB 18775, 19544, 19670, Paulo Frontin IB 24986-
24987, 25321, Pirai do Sul IB 12425, Rio Azul IB 21346, 22801, Tronco IB 730. Rio de Janeiro :
Friburgo MZUSP 2705-2707, Muri IB 21261, 21929, 24579, Petropolis IB 21473, 21475, Rio de
Janeiro (Mono da Urea) IB 985. Rio Grande do Sul : no specific locality. BM 82.10.4.60-82.10.4.61,
61 A, MCZ 5633, 17932-17933, MZUSP 1 109, 1 1 12-1 1 13, 1 127-1128, UMMZ 6724 1 , ZMUC 60 1 254-
601255, Alexandrinha MZUSP 5472, Cacequi IB 10025, Canela IB 13526, Caxias do Sul IB 6418,
6716, 9316, Cachoeira do Sul 23315, 23317, Cruz Alta IB 18386, Estacao Ecologica do Taim MZUSP
7524, Guaiba CAS 89672, Pareci Novo IB 7645, Passo Fundo IB 8049, 8060, 8212, Pelotas IB 41057,
Porto Alegre IB 7264, 7572, 1 70 1 5, 224 1 1 , 22509-225 1 1 , 22565, 22832, 23 1 89, 234 1 9-23420, 23543,
45935, Quebra Dente-Maran IB 23302, Restinga Seca IB 7305, 9837, Sao Joao MNHP 1894.440,
Sao Leopoldo MSUSP 4100, IB 7300, 7352, 7776, 8357. Santa Catarina : Banacas IB 13319, Canoin-
has, near Taunay IB 24612, Itaiopolis IB 16669, Lagoa do Norte IB 9700, Porto Uniao IB 25992,
Sao Joaquim IB 45187, Taunay IB 16328, Tres Banas IB 7948. Sao Paulo : no specific locality MCZ
207110-20713, UMMZ 62804, 108760, Agudos IB 22097-22100, Abernessia IB 9553, Altinopolis
IB 19562, Araraquara IB 19413, Arpui IB 831, Atibaia IB 23149, 23254, 26889, 26889, Anhanquera,
KM 24 IB 23163, Baninha IB 43133, Barueri IB 6987, Bento Quirino IB 20641, Boqueirao da Praia
Grande IB 11524, Boraceia MZUSP 4498, Botucatu MZUSP 2406, Braganca Paulista IB 30498,
Caieiras IB 41061, Caixa D’Agua IB 5459, 5682, 6716, Campo Largo 4639, Campo Limpo IB 4738,
5309, 5745, 7349, 7541, 18658, Campos do Jordao IB 8450, 30692, MZUSP 1122, Capitiuva IB
3070, Cotia IB 7504-7505, 24281, 29195-29196, Embu km 28, IB 30488, 30490-30494, 305741,
Embu Guagu IB 16617, 167796, 34299, Estacao Gabriel Piza IB 5428, Guarulhos IB 5041, 23556,
23724, 42961, MZUSP 4721, Ibiuna IB 26979, 32251, 32265, 40930, 42203, 42222, 42337, 43764,
Itapetininga IB 10287-10288, Itapecerica da Sena IB 41115, 42138, 42942, 43882-43883, 44116,
44189, 44498, Itatiba IB 5551, 5586, 5741, 24185-24186, Itaquera IB 10189, 10197-10200, Itu IB
32234, Jacarei IB 3293, 24926, 28502, Jarinu IB 12810, 27542, 30886, Jundiai IB 19686, Lobo IB
3253, macedonia IB 7026, Mairipora IB 24380, 26949, 27958, Mairinque IB 6042, 6048, 6075, 6618,
Mogi das Cruzes IB 10196, 10268, 10295, 10318, 21387, 21395, 22596, Moreiras IB 17606, Ped-
erneiras IB 5470, Peruibe IB 5532, Perus IB 3292, Pindamonhangaba IB 18820, Pinheirinhas IB
32420, Pirituba IB 908, Pilar do Sul IB 32121, 32454, Quilauna IB 6257-6258, Raiz da Sena MZUSP
1 1 14-1 1 16, Ribeirao Pires IB 10232, Ribeirao Preto IB 22831, Rodovia Dos Bandeirantes dm 60,
IB 34397, Salto IB 45 1 6, 4578-4579, Sao Bento do Sapucai IB 3283, Sao Bernardo do Campo MZUSP
3289, 3682, 4463-4465, IB 22608, 42919, Sao Lourenco da Sena IB 45789, Sao Paulo MZUSP 1 1 17,
1119, 1950, 2372, 2387, 2609, 2635, 2960-2961, 3355, 3688, 4095, 4130, 4756, IB 191, 193-95,
647, 695, 1315, 1321-1322, 1416-1417, 3252, 3494, 4643, 5411, 5573, 6189, 7646, 8042, 8575,
9561, 9842, 9879, 10194-10195, 10252, 10271, 12214, 12977, 12986, 17085, 17871, 18270, 18405,
18920, 22882, 24178, 24431, 24442, 26656, 27014, 27157, 27212, 27281, 27478, 27498, 27556,
27571, 27631, 28308, 28503-28508, 29222, 29236-20237, 29249, 29614, 29924, 30683, 32446,
32504, 32800, 32803, 32925, 32999, 34277, 41129, 42109, 42195, 44162, 45746, 45751, Sao Roque
IB 5391, 44228, Sena da Bocaina MZUSP 4912, Sorocaba IB 6865, Souzas IB 23151, Sumare IB
1332, Suzano IB 20376, Tapirai IB 28060. Tremembe IB 10368, Valentim Gentil IB 28671, Vinhedo
IB 30320. URUGUAY. Canelones : no specific locality BM 84.2.23.39, FMNH 10215, Anoyo Solis
Grande CM 38959, Banados de Carrasco CM 55422-55423, Rocha USNM 89995, Ruta Interbale-
nearia CAS 93100. Montevideo : no specific locality MNHP 171, Santiago Vazquez LSUMZ 27753,
MNMHP168.

Liophis maryellenae Dixon

LiophismaryellenaeDixon,1985.Holotype—AMNH62202.Type-locality—Annapolis,Goias,Brazil.
Description. — The following description is based on 10 specimens. Maximum
total length of males 435 mm, of females 530 mm. Tail/total length ratios vary
from 0.221 to 0.262 (x = 0.240). Diameter of eye/snout length ratios vary from
0.605 to 0.844 (x = 0.686). Dorsal scale rows are 19-19-17, smooth, with an
apical scale pit. Reduction to 17 rows occurs between ventrals 74 and 95 (x =
82.8). The anal plate is divided. The number of ventrals varies from 144 to 159
(x = 150.9). The number of subcaudals varies from 63 to 82 (x = 67.9). The
number of maxillary teeth varies from 25 to 28 (x = 26.1). Head scales vary as
188 Annals of Carnegie Museum vol.56

Fig.4.—DistributionofLiophisatriventerandL.maryellenaeinsoutheasternBrazil.Dotsrepresent
L.maryellenae;thesquare,L.atriventer.

follows: supralabials 8-8 (10), supralabials entering orbit 4 + 5 (10), infralabials

10-10 (8), 11-11 (2); preoculars 1-1 (10); postoculars 2-2 (10); loreals 1-1 (10);
temporals 1+2(10).
The hemipenes vary between (in situ ) 9.5 to 12 (x = 10.5) subcaudals long.
The structure is slightly bilobed, each lobe about two subcaudals in length, and
their distal ends have a smooth apical disk. The sulcus spermaticus divides about
the middle of the shaft. The entire structure is spinose, with the spine length
decreasing gradually from the base to the apex. A basal naked pocket is present.
The dorsum is usually some shade of green, grayish to olive green, with some
indication of a brown or reddish brown mid-dorsal stripe. Other dark lines may
be present laterally; some indication of dark flecks may occur over the body. In
life, the venter is yellowish orange. For more details of color, see Dixon (1985).
Distribution. — This species apparently is restricted to the tablelands of south-
eastern Brazil (Fig. 4). For specimens examined, see Dixon (1985).
1987 Dixon— South American Li o phis 189

Liophis atriventer Dixon and Thomas

LiophisatriventerDixonandThomas,1985.Holotype—MZUSP5066.Type-locality—Boraceia(Es-
tacaoBiologica),SaoPaulo,Brazil.
Description. =the following description is based on three specimens. Maximum
total length of male 505 mm in snout-vent length). Tail/total length ratios vary
from 0.195 to 0.214 (x = 0.203). Dorsal scales are in 19-19-17 rows, smooth,
without apical scale pits. The reduction to 17 rows occurs between ventrals 73
and 79 (x = 76.7). The anal plate is divided. The number of ventrals varies from
141 to 148 (x = 144.0). Number of subcaudals from 49 to 56 (x = 52.5); the
number of maxillary teeth from 23 to 24 (x = 23.2). Head scales are as follows:
supralabials 8-8 (3), supralabials entering orbit 4+5 (3), infralabials 10-10 (3),
preoculars 1-1 (3), postoculars 2-2 (3), temporals 1+2 (3).
The hemipenis of the only known male is short, 5.5 subcaudals long (in situ),
and bilobed for the distal half. The sulcus spermaticus divides about one third
of the distance from the base. The distal end of the lobes contain smooth apical
disks. Five large spines are present along the outer surface of each lobe, near its
base, while the inner surface of the lobe is bare. In addition to the larger spines,
many smaller ones are scattered over the shaft.
In preservative, the dorsum is uniform olive brown. The throat and first seven
ventrals are grayish white, and the remainder become progressively darker until
the posterior two-thirds of the venter is black. The subcaudals are grayish with
occasional dark smudges.
Distribution. — This taxon is known only from the type locality (Fig. 4). See
Dixon and Thomas (1985) for specimens examined.

Key to the “Green” Liophis of South America

1. Scale rows 19-19-15 or 19-19-17 2
Scale rows 17-17-17 or 17-17-15 jaegeri
2. Scale rows 19-19-15 3
Scale rows 19-19-17 4
3. Ventrals 133-172, one scale pit typhlus
Ventrals 187-199, no scale pit guentheri
4. Ventrals 141-159, maxillary teeth 23-26 5
Ventrals 169-202, maxillary teeth 17-24 viridis
5. Subcaudals 62-82, belly yellowish orange maryellenae
Subcaudals 49-56, belly black atriventer

Acknowledgments

I thank the following curators who have indulged my requests for specimens over many years R.
L. Bezy and J. W. Wright, Los Angeles County Museum of Natural History (LACM); J. A. Campbell,
University of Texas, Arlington (UTA); C. L. S. Cordeiro, Instituto Butantan (IB); H.C. Dressauer,
(HCD); R. C. Drewes and A. E. Leviton, California Academy of Science (CAS); W. E. Duellman,
University of Kansas Museum of Natural History (KU); W. R. Heyer, R. McDiarmid and G. Zug,
United States National Museum of Natural History (USNM); M. S. Hoogmoed, Rijksmuseum van
Natuurlijke Historic, (RMNH); A. G. Kluge and R. A. Nussbaum, University of Michigan Museum
of Zoology (UMMZ); W. W. Lamar (WWL); V. Mahnert, Museum d’Histoire Naturelle, Geneve
(MNHG); E. Malnate and T. Uzzell, Academy of Natural Sciences, Philadelphia (ANSP); H. Marx,
Field Museum of Natural History, Chicago (FMNH); C. J. McCoy, Carnegie Museum (CM); C. W.
Myers and R. G. Zweifel, American Museum of Natural History (AMNH); J. B. Rasmussen, Zoologisk
MuseumUniversitetsparken,Kobenhavn(UZM);D.A.Rossman,LouisianaStateUniversityMuseum
of Zoology (LSUMZ); R. Roux, Museum National d’Histoire Naturelle, Paris (MHNP); A. F. Stimson,
BritishMuseum(NaturalHistory)(BM);P.E.Vanzolini,MuseodeZoologia,UniversidadeSaoPaulo,
(MZUSP, DEH); S. E. Williams, Museum of Comparative Zoology (MCZ).
190 Annals of Carnegie Museum vol.56

IexpressmyappreciationtomywifeMary,whohassharedmypursuitofLiophisandherpetology
ingeneral,formorethan30years.

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