OF

CARNEGIE

MU

M
28 AlXlUST 19 7

VOl.. 56. AIITI

D GEOGRAPIllC V RI TIO OF
T OOMY
RELATED GREE PE IE OF
LJOPHI TYPHL S
OUTHAMERlC (SERPENTE : COLUBRID E)
JAMES

R Dr

The gr n peci of Liophi are reviewed... Liophis I)ph/us con iSlS of three ub pecies. L I. IJph/Lis
ma:r..on L. I. e/aeoides Chaco). and L I. bracllJ'lInlS ( crrado). Liophis l'iridi consi lS of two
sub pecies. L. I'. I'iridis grcste and Atlantic rain fore 1) and L. I'. prasillUS (CaaLinga). Liop"isjaegeri
i hown 10 con . I of two ub pcc1CS, L jaegeri jaegeri. east of Ihe Rio Parana, from ao Paulo area
of Brazil. 10 ruguay and rgenlina and L j. coraf/il'elllris. from the Rio Paraguay basin. Liophis
gr./t!/III/{m i a valid species from Ihe dry central Chaco of Argentina Bolivia. and P ragua . The
ch,llueters of Ih recentl described species, L Qlriw'lller and L marye/lellQe arc ummari.zcd. ke.
i provided for all pecies of "green" Liopltis.

INTRoDucno

Th tudy oflhe genus Liophis, a long and difficult task i now nearing completion. The ix pedes pre ented berein (L. typhlus. L. guentheri, L. jaegeri. L.
viridis. L. mar) ellenae. L. o/rb'enter) form a loosely connected comple of species
that ha c one feature in common they are aU some shade of'green. The green
color rna be dull to bright chlorophyll green to leaf gre n or with a deep olive
cast. Two taxa occasionaUy ha e an obscure mid-dorsal stripe of ariable width
that vari s in color from light brown to reddish and occasionally secondary dark
line as well.
When samples were adequate, each of the si species wa e amined for geographic variation. No additional specimens of two recently describ d taxa (L.
a/riven/el', L. mar) eilenae) 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
comm nts about pertinent literature.
SYSTEMATIC ACCOUNT

Liophis typhlus (Linnaeus)

oll/bertyphlus Linnaeus, 1758. Syntypes (nOI examined)-Royal Museum, lOckholm. Type-local
ity-Indiis (in crror),

Xe/lodO/l iso/epis ope, 1870. Holotype ANSP (losI). Type-Iocality-Pebas, Ecuador (=PerU).
Oph/!omorp/ws brachyunts Cope, 1887. Syntypts ANSP 11202...{)J. Type!ocality- hupada, Mato
Oro 0, Brazil.

.'

Uophis e/aeoides Grimn. 1916. Holotype CM R32. Type-Iocalily-Prov. del ara, Bolivia.
Uophis lIIacrops Werner, 1925. Holotype NMW 23420. Typ -locality-Paramaribo. urinam .
o177enclalUra/ comments. -Several names have been assigned mistakenly to
the synonomy of L. /yph/us. Jan (l863a, 1863b) and Jan and ordelli (1866)
Depanment of Wildlife and Fisheries
77843-2258.
ubmillcd 16 January 1987.

ciences Texas

173

&M University.

ollege

tation Texas

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suggested (hat L. typhllls was composed of three variclies in addition 10 (he

nomin:Hc race. prasillils. gasfros,icllIs. and o/il'acells. J have dClcrmincd that no
holotypes exist for any ofthcsc varietal names. An examination of Jan"s descriplions and Jan and Sordclli's illustrations (1866. Livr. 18, P1DIe IV. Fig. 3) suggest
that the name prasil/lls belongs to Liophis ,'irit/is. The illustr.:llion shows 69 sub
cauda Is. considcf<lbly more than the maximum known number of61 for L. tYVhllls.
but well within the known range of 63 to 84 for L. I';,-idis. An illustration of a L.
Iyphills by Jan and Sordclli (1866. Livr. 18, Pia Ie IV, Fig. 2) from Pernambuco."
Milan Museum. may represent the varicty gasfrosticlUS. of Jan (18630. 1863b).
The illustration shows 47 subcaudals. well within the known range of variation
in subcaudals for L. l)"phlIlS. However. the closest localities I,f L. Iyphlus to
Pernambuco are 1300 km to the cast 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 pot.'Cilog)TIIS from Pernambuco. and I suggest gaslrostialls belongs
to that species. The Jan name oliroce1lS is without description or illustration. It
appears in Jan's (18630, I 863b) list and key to Liophis as a varietal name of L
IJphllls and should be considered a "omell lluduII/.
Another misapplied name is Wagler's (1824) Nalrix forslC'ri. Hoge (1964) first
applied it to L. 1.11)hl1IS and suggested that the Surinam race be recognized as the
nominate taxon and the Brazilian fonn (by inference) L.1J'phlllsforslC'ri. The typclocality of L.{orsleri is Salvador. Bahia. IJrazil, a locality of considerable distance
from any known sample of L.l,rp!lJlIs. In addilion. the illustration and description
offorsler; suggests that this name is best applied to L. poedlog,rrus. The description
lists 60 subcaudals for the type of L. forsleri. The greatest number of subcaudals
recorded in 73 eastern Brazilian individuals of L. IJp!lJIlS is 48. The description
of the color pallem of forslni is unlike that of L. (rphllls. and more similar to
that of L. poecilogYrlIs. Hoogmocd and Gruber (1983) examined the holot)pe of
N.lorstC'ri in the Munich Museum and gave the subcaudal number as 58/58+ I.
Although they placed NforslC'r; as a synonym of L. l,rp!lJuS. I consider N.forsler;
to be a senior synonym of L. poecilog,rrus.
Parker( 1928)on comparing one ofthe paratypcs of L. gllemlteri(BM 1946.1.5.69)
to S<1mplcs of L. 1.I1)!lJlIs in the British Museum (Natural History). concluded that
it "failed to reyeal any consequential differences between these two species" ilnd
placed L. g//('III!1er; in the synonomy of L. typhllls. Parkcr erred in his analysis
because the ventrals of the specimen he examincd (and L. glll'IIt!leri in gcneral)
vary from 187 to 197, while ycntrals of L. lypli/us vary from 1)3 to 172. Liophis
glll'lIfheri is a vlliid species that is more closely related to L. piridis than to L.
lyphillS.
Dislribution. - Uop!lis t,rphll/s occurs in Amnzonian rainforests to an elevation
of about 1500 m along the southern and Castcrn side of the Andes in Colombia.
Ecuador and Peru. the rainforests of southern and eastern Vcnezucla. and similar
forests of Guyana. Surinam ilnd French Guianil. and Cerrado and Chaco forests
ofSE Bolivia. N ParaguilY, and SC Brazil (Fig. I).
Dt'scripIlOIl.- The maximum lotallength is 740 mm in males and 853 mm in
femnles. 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
\'cntrals 68 and 103 (.f = 82.2). Scale rows 7+8 or 8+9 usually fuse between
,"entrals 65 and 104 (.f = 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

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DtXON-SOUTH

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175

Fig. I.-Di Iribuli n or Liophis ryphills (dots) and L. gllellfhe,.i (diamond. tippled area) in South
America. Blaek line enclose numbered am pies used in the geographi variation analy e .

of331 Sp cimens numbered 133-172 (_x = 157.1) the number ofsubeaudals of

307 speei mens 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 nout I ngth of 127
adult vari d from 0.524-0.886 (x = 0.682). Maxillary teeth Cor 222 pecimens
aried from 19-28 (_\ = 22.4). The anal plate is divided and lh lor al iogl in
all p cimen xamincd.
Head hi Ids vary as follows: supralabials 4-5 (I) 7-7 (3). 7- (2) 8-8 249),
-9 (5)' upralabials entering orbit 4th only (1), 4+5 (252), 3+4/4+5 (2), 4+5/
5+6 (4) 4+5/ +5+6 (I); infralabials 5-7 (I) 8-8 (I), 9-9 (10 9-10 (_3).10-10
(216) 10-11 (5) II-II (2) 11-12 (I): preoculars 0-0 (I), I-I (_56); po loculars
I-I (3).1-2(1), _-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).
Th ill illl length for 69 hcmipenes varied from 6 to 13 _\ = 9.0) ubcaudals.
The organ is slightly bilobed, with each lobe aboul two subcaudals long. The
sulcu sp rmaticus usuall bifurcates about one-third the distanc from th base

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of the organ. Large spines arc present on the asulcale surface of the basal two
thirds of the organ, usually becoming much smaller on the lobed part Oflhc organ.
A smooth apical disk is present on the outer edge of each lobe and a naked basal
pocket is usually prescnt.
Color ill 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.
Thejuvcnile dorsal pattern is variable with somejuveniles having distinct black
chevrons with a wide black nuchal band. The nuchal band usually fades !o an
obscure greenish black mark at a total length of 21 mm. Other juveniles from
155-255 mOl in totallcngth have a pair of blackish nuchal spots that begin on
or al Ihe posterior edge oflhe parietals, and slant postcroventrally. 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 mriafioll. - There arc three distinct geographic populations of L.
fJ1J"'IIS that correspond to forest refugia postulated by Vanzolini and Williams
(1970), and to some extent, to those of Haffer (1974). The three geographic populations were dctcrmined by comparison of Ihe numbers of maxillary teeth. ventrals. subeaudals, the ratio of tail length to total length. and the ratio of the eye
diameter to snout Icngth. Six samples were used to detennine 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 specimens 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 specimens from the Cerrado of southeastem Brazil (Fig. I). The Student's T tesl was
utilized to determine significance of pair-wise comparisons between samples. A
Siudent's T tesl value of6.314 or greater is at 95% level of significance or grealer.
Pair-wise tests for differences between samples I and 2, I and 4, 2 and 3, 2 nnd
4 were nOI signiftcanl for any oflhe characters listed above. Samples I and 3 were
significantly different from each other in numbcr of vcntrals and tail/lOtallength
ratio. However, samples 1 and 3 are separated by sample 2, and sample 2 did
not differ significantly from either I or 3. The significance noted between I and
3 is probably an artifact of distance, with samplc 2 representing the middle of a
cline. Samples 3 and 4 were not signiftcantly different in any characters except
vcntrals. 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 (sec Fig. I). This
suggests Ihat a hiatus may exist between samples 3 and 4 in Amazonian Bolivia.

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

high numbers of ventrals from samples I to 4, I to 2, 2 to 4, 2 to 3. The samples
form an almost closed circle of demes (Fig. I), with the bottom of the circle open,
representing thc diffcrences in the Bolivian samples mentioned abovc. It seems
dear that the Amazon samples arc closely related, but with a trend to divide the
samples into castern and western demes.
Pairwise comparisons of samples 4 and 6 suggest a strong differentiation in all
characters examincd. The differences in the numbers of vcntrals, subcaudals, and
tail/total length ratios are highly significant between thc twO samples. with T
values of23.8. J 9.1. and J 5.8 respectively. The number of maxillary teeth differed
significantly(T value'" 7.0) but eye diameter/snout length ratios did not (Tvalue =
2.14). The data suggest that the Brazilian Amazon forest and deciduous mesophytic forest samples represent different aJlopatric taxa. Comparison of samples
3 and 6 shows thc same trend, except the number of maxillary teeth is not
significant (T - 4.91). Pair-wise comparison of samples 4 and S shows significant
differences in the number of ventrals and maxillary teeth (f - 22.03. 10.54.
respectivcly). but not in the number of subcaudals. tail/total length. or eye diamcter/snout length ratios (T = 2.66, 4.59, 4.62, respectively). Pair-wise comparison ofsamples 3 and 5 shows significant differences in the number of ventraIs.
maxillary teeth, and subcaudaIs (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 sam pic from Bolivia is morc closely
related to the Amazon forest sample than to thc deciduous mesophytic forest
sam pic of llra~tjl. However. both non-Amazon forest samples are very distinct
from the Amazon forest samples.
A pair-wisc comparison of samples 5 and 6 shows significant differences in the
number of subcaud.als. m.axillary teeth, and t.aiVtot:lI length ratios (T = 21.05,
6.43, 12.9, respectively), but not in the number of ventra Is or cye diameter/snout
length ratio (T - 1.36,3.59). Thesc samples represent dilTerenttaxa, evcn though
they .arc rel.atively close geographically (Fig. I).
The sc.ale count d.ata and Icngth rntios suggest three distinct populations of
LioplJis Iyphlus. Samples I, 2, 3, and 4 represent the Amazon forest form, s.ample
5 the northern Chaco form, .and s.ample 6 lhc Cerrado form.
The color patterns of s.amples 1,2,3, and 4 arc similar, and the pattcrns of
samplcs 5 and 6 are idcntical. The dorsum and head of adult individuals from
snmples 5 and 6 arc chlorophyll green. The young and juveniles have black nape
spots .and some h.ave four rows ofdorsal dark spots. Adults of the Amazon samples
have a Icafgreen to bright green dorsum with di.agonal reddish brown to blackish
chevron ffi.arks along each side of the body. The head m.ay be greenish to bright
blue. Young and juveniles are greenish dorsally with bold dark chevron marks
and a rel3livcly l.arge bl.aek nuchal blotch.
Diagnoses .and distributions of the three taxa follow:
Lioph;s f}'ph/us f}'ph/us

(Linn.acus)

CoIubt'r /Jphfus linn:lCu5. 1758.

),'(',wdon isol('pis Cope. 1870.
"'OCTOPS Werner. 1925.

uopnu

Diagnosis.-Dorsum usually green with distinct, reddish brown to black chevrons on e.ach side. occasionally fading posteriorly. The dorsal surface of the head

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may be greenish to bright blue. The venter is while to light yellow, usually without
dark markings. Juveniles have a large black nuchal blotch and dark chevrons.
The number of ventraIs varies from 133 to 163 (.t = 147.3), subcaudals vary from
47 to 61 (.t' = 54.2), m<lxillary teeth vary from 20 to 28 (x = 23.6), lailftolallength
ratios vary from 0.160 to 0.210 (J' = 0.184), and eye diameter/snout length ratios
vary from 0.510 to 0.861 (.t' = 0.736).
lJiSlri!JwiOI1.- Throughout the Amazon Basin in rainforest, to about 1000 m
elevation (sec specimens examined).

Liophis tJ'phlus elaeoides Griffin

Liopllis e1uroidrj Griffin. 19 J 6.

Diagllosis. - Dorsally adults are uniform chlorophyll green with a light yellow
venter. Juveniles have four ro\ys of dorsal dark spots on a green background, and
a pair of black nuchal spots. The number of ventrals varies from 158 10 172 (.~ =
163.5), subcaudals vary from 49 to 56
= 51.9). maxillary teeth vary from 17
to 23 (.~ = 19,9), tail/towllength ratios vary from 0.160 to 0.200 (S = 0.171),
and eye diametcr/snout length ratios vary from 0.520 to 0.740 (.f = 0.620).
Dislribmiotl.-Liophis typhlus elaeoides is known only from the upper Rio
Paraguay Basin, which includes the mesic Chaco forests of soulheastem Bolivia,
nonhern Paraguay, and western Malo Grosso, Brazil (see specimens examined).

<,t

Liophis typhlus bmchYllYIIs (Cope)

Opl1l'olllorplw.t hrachJ"llrl1S Cope. 1887.

Diagllosis.-Adults are uniform green dorsally with a light yellow venter. Juveniles have four rows of dark dorsal spots on a green background and a pair of
black nuchal spots. The number orventrals vary from 151 to 171 (X' = 162.2),
subcaudals vary from 40 to 49 (.f = 44.4), maxillary teeth vary from 18 to 24
(x = 21.5). tail/total length ratios vary from 0.140 to 0.160 (S = 0.149), and eye
diameter/snout length ratios vary from 0.55 to 0,81 (.t = 0.675).
Dislriburioll.-Liophis I. brachYllrus occurs in deciduous mesophytic forests of
southeastern Brazil, and in the Campos Cerrado forests of east-central Brazil (see
specimens examined).
Commellfs. - Two recent papers by Miranda and Couturier (1983. 1984) comment on lhe presence and geographic variation of L. typhllls in Argentina. I have
neither examined their specimens, nor toea led specimens of L. Iyphills rrom
Argentina in other museums. I believe they may have erred in identification of
their specimens. Photographs presented by Miranda and Coulurier (1984) suggeSI
that Ihey may have conrused L. llIi1iol'is and L. poecilogyrus wilh L. fyphlus.
Short works on the distribution and taxonomy of L. Iyphlus are Hoge (1964),
Amaral (1931. 1935, 1949), Gans (1960). Parker (1928, 1935), Petcrs (1960).
Peters and Orejas-Miranda (1970), Roze (1966), and Prado and Hoge (1948).
Peters (1963) compared the maxillary teeth of L. lyphlllS with those of other
species of Liophis.
SpeclmlllS~xumim>d.-(UopltiJfyplt!lufyph!lu) BOLIVIA. Be/r;: Ixiamus AMNii 22457; Rio Ikni
AMNH 22270. I~RAZIL. Act'l": Alto Purus MZUSP 2498: Pono Walter MZUSP 7390. Amapa:
Cuidade Oiapoque IB 13780. 13783; Serro do Navio KU 9787J-97875. Amu::olllu; Uarrcim do
l\-lalupiri. on lhe Rio Madcirn MZUSP 5912: Carvociro AMNH 36167: Cosln Allamim on the Rio
JJpura MZUSP 6600: lago Alcxo Mel 3290: Manaus MZUSP 3051. 3797; Rcscn'a INPA ~'!lUSP
7606. 7619: SJnla Isabel on the Rio Negro USNM 83532: sao Paulo de Olivcnen AI\lINH 53311:

1987

DIXON-SouTH AMERICAN

LfOI'1I1!i

179

Tap.1u:l MZUSP 5770: no specifIC locality UZM 601213. Marallhatr. Akki.a Aracu 19ar:ropc GllrupiUna MZUSP 4303. 4826. PorQ: As Prom. on the Rio Cumini-Miri MZUSP 5103: Bcltm 18 15667;
C~ninde.. on lhe RioGurupi MZUSP4267. 4285: near Maraba. Seml.ndo None KU 124608: Oxiximina
MZUSP47%: Uru1. ParqU( Nac. da Am:u:onia... on the Rio Tapajos MZUSP 7838. Rondo"'Q: Porto
\'elho MZUSP 3690. COLOMBIA .\IQQmal: Rio Gara;oa Mel 27339. Mtfa; \'iIIa\icc:ncio 188589.
IiJllpn: Timbo UTA R3805; Yapima UTA RID33. ECUAOOR. Napo-P(Ula:a: Alpa)"'K'a FMNH
4069. UMMZ 89020-22: Ando:15 AMNH 41949; DaOOs+C~nc:1oI AMNH 35892: Cl\am~I.a-Nor
mandi.a-Rio Damba AMNH 359Z9-359JQ: Jara)<ICU AMNH 28796; M01QS i10d Vicinity AMNH
35838-35839: 82 km ESE Macas AMNH 114616: Rio Ibmba AMNH 15209. 15213.23296_ 23302.
28848: Rio PaSt:llii MCZ 36966: Santa Rosa. EI Tigrt" AMNH 49165. FRENCH GUIANA (CAYENNE). MOIna USNM. 6172:Saul Mel 149399: ncar Sophie Mel 77510-77511. GUYANA. BartoPOllltO road 11M 1954.1.3.64; 82 km S BaniGI. BM 1934.11.1.125: lkrbice 8M RR 1964: IXmcran
Rh-cr. Lama CRX'1r: AMNH 36106; Essntllibo. near Lethc:m USNM 146376; Kanabo AMNH 21335.
98197-98198: Matah AMNii 61542: Rupunllni District. ootth of Acara.i Mountains. est of Ne....
Rh'er KU 69826-69828: Rllpununi Di!ilrict. nonh of Atarai Mountains. "'-esl of Ne Ri\'cr KU
69826-69828: no specific locality M HG 279.38. PERU. Ama:OIlas: Caterpi7.a. Rio Calerpila USN""
(RWM field series) 14995. I SOB. 14185-15186. Jill/in: Rio Percne MCZ 42434. 1.()rI'IO: Balta LSUMZ
14584; Cenlro Union TCWC 44682; Cerros de Contaya. on the Rio Tapiche AMNH 53376: Conlamana on the Rio UC3)'Olli AMNH 52130: Estirim on the Rio Ampi)'3CU MZUSP 4394: Iquitos
AMNH 52734: 53118. 53284. 53667. 53696. 53735. 53771. 53773. 53876. 53923. 53949, 54321,
5.\]54.5.\48].5-'894.56109: Moropon TCWC ]8049. 44294; <mllana (Reforma) AMNH 54957.
USNM 127124: P:lmpa ~lrrmosa on Ihe Rio Cushabala)' AMN~I 55409. 5541 5. 55442, 55791,55885.
56003: Pam)'Ol AMNU 53249: Rcqocna (/'.Ioole Carmelo) AMNH 55600. 55626: RO~Olbo}Ol AMNH
52.\83.53110.53288.55695: San Antonio on the Rio ha)'a AMNH 52920. 53667. 51693. 53696.
5)735: Shiriar:ro. on the Rio Nana)' AMNH 56015; Trapichc-UIIXjllinia AMNii 52195: YlInam<HlO
TCWC 4()542-40543. Madrrtk Diar. Rio ~Icalh. 50 km S Puerto Pardo LSUMZ 36778-79; mouth
of Rio To~ on the Rio TlImoopata L'iUMZ ]9.\244. l>asro: liC01.2Zin Valky LA01 76805. San
.lla,wr. MO)'obamba liM 74.8.459.94_8..\.64. SURINAME: Ch:u1esbul'l AMNH 104624: Coppcname.' Ri\"l:'r Mel 152203. 152205. 1526]3--152634: Jarawq TCWC 60543. 60756: Panmaribo AMNH
8146.8682. MCZ 16401: Polribo CM 44302; Sipali...;ni Aintrip CM 84667. Mel 152636: Zanderi
(Airport) MCZ 152635. VENEZUEU.. Ama:onar. ArabopO UMMZ 85279; Aroroima CiiOOs Mel
385.\1: Mount Duid:l JqlOO AMNii ]6617_ 36620. Sol/far; EI /'.'1:Intcco TCWC 60168: no spcrilic
Iocalit) MHNLS 1632.
(lJOp/IIS t)phhu Morouln) BOLIVIA: CochQmoomoo: Villa Tunari UMMZ 15J095. Samo eru::
Buena\'istaCM R2696. R2698. R2701. R2704. R2869. R2886. R293O, R2931; UMMZ67967-67968.
67969(2).67970(3): lJucna\ista. near Rio Colorado CM R2860. R2865. R2955: Santa Crul. LSUMZ
11825: Santa Crul. de la Sierra CM RJ:! (holol)"JlC). R44. R59, R91. R94. R95. R97. R98. R99. R102.
MCZ 11860. MZUSP 6474: San Jose de Chiquitos CM 34842. No specific locali1r: eM R29]8R29]9. FMNU 195898. I-ICD 2820. 2822. Tavc 55290. BRAZIL: '\/alO Grosso: Corumbii, near
Urueum Mounl3insCM ]4841; Fa7.enda Bela Vista ofthe [Ilia Insu:l MZUSP 7264: Falenda Vaeun7.ni
Oflhl' Rio Paraguai MZUSI) 7266: Generalso Ponce. Corumba 1325954: Maleta IIJ 14975: I'ono
Murtinho IB 26177-78.
(Uopilis l)'lIhllIS hrachYllfIIs) [)R,\ZI L: HoMo: Mira Serra. 41 km from Morro de Chopco MZUSP
7554. Galli:: Araguari [ll 685 I: Falcnda Lucushac, 011 the Rio Verde 11.113060. MaiO Grlmo: Ampua
III 9939; AquidilUilIl3 MZUSI' 33589: Chapada ANsr I 1202-1 1203. IlM 92.420.13: T(lUGM~ dll Semi
Hl 24543: Tauna)' IB 7674; Urucum. ncar Corumb;i BM 1928.1.12.3. ,\-Ii/laS Gerais: Hono ForcSlal
I[) 10493: Irma lllidur 1lJ 6933: Itamlx: do Mato Dcnlro MZUSP 8061: Juil de Fora 111 25 I90. 26685:
Machado IIJ 1630]: Sant:l Rila do EXlrema J() 5540. 5562. H.iodlJrmdro: Nova Friburgo IIJ 10516.
Slio Paulo: ArnrJguara 11I231: Atit>aia JB 21376. 27081. 27201: Barra Assuj;nui lB 290'*2: Barrctos
[Il 5030: liaurcri [13 7212: Ilauru MZUSP 823: llornceia MZUSP 4227: Iknueiltu MZUSP 2410. 2412:
Bun IB 6718. 6763. 12313: Campo Limp!) IB 80S. 6452. 10265-10267, 24903--24904: Capao IJonito
1323465: CaS(';l\'el MZUSI' 824: Conde do Pinllal IB 9975: COlia III 5743. 6935. 7140. 9906. 19906,
23236.23377.24542.32627: Curupa IB 7061: Dona Catarina III 22598: Elias Fausto IB 232. 319.
392. 578. 791. 10461: Elihu Root IB 234; Engenheiro Cesar de SoU7.ll IB 8338: Ferrn de Vasconcelos
MZUSP 2487: Guarllihos IIJ 23726-23727: Hono Flol1;$lal IIJ 10493: lbiuna 18 12252. 23794:
lpanema IB 230. 235: ltagua 104813: ltapccerica da Serra 18 7794. 19682.25040.25043: Itapctininga
III 2.\565: Ilapolis 18 7956: ltaqller.1 IB 30758: Ilaquaqll(:('Cluba IIJ 18349: Ilirapina 18 6655: Jal"lliua
IIJ 21381:Jarinu 18 10292: Lauro Mulkr 18 10533. 10556. 10559: I..cmc 18 236. 547: MlIinfKloc IU
7373. 7461: Mairipora III 28603: Magi das Cruza IB 12592. 19658, 23547. 24881. 28284. MCZ
39415. MZUSP 6460: Osa.sco 10 12500_ 27910--27911: Pcdemciras 18 6938: Ptndamonhanpba 10
1257; Pira.jll MCZ 39416: Promiss30 18 9969: Ribcin;lS Pires 18195%; Rio Daro ID 7277.10296:

voo..>.

ISO

Santa Adelia 18 6909: Santa Emestina 18 21969. MZU5P 2479; sao Carlos 18 10016; Sio Jose do
Rio Pardo 18 5444: Sio Jose dos Campos IB 21348; Sio Paulo 18 797, 7142. 12980. 12993. 16223.
16900. 18854, 252.95. 27497. 28458. MCZ 17956. MZUSP 817-820, 2388, ])]8-3JJ9. 3695. USNM
692J8-69::!:39: SU1.ano 18 19104: Valinho 18 6719; Vintw:do 18 291S: no spccifie locality 18 233. 237,
797. 840. 4145. 5491, 5642. 69.so. 9775. IOS81. 12112. 17777.

Liophis guenthl!l'i Pcracca

Liophis gllrntl/l'ri Peracrn. 1897. Six synlypcs (Fh'e 10SI ?), Mus. Compo Anal.. Torino. Type localit)'Caiza. Bolivia.

Commellls. -Liophis gllcllIheri has resided as a synonym of L. typhlus. Parker

(1928) compared one of the syntypes (now 8M 1946.1.5.69) with a specimen of
L.lyphlus. and suggested it "failed to reveal any consequential differences between
the IWO 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 syntypcs that were housed in the Museum of Zoology and Comparative
Anatomy in Torino.
Dislributioll.-Uophis gliemheri apparently is restricted to the central Chaco
of Bolivia. Argentina and Paraguay (Fig. I).
Descriplioll. - The following description is based on six adult females and three
adult males. The maximum total length of males examined is 655 mm, offemales
723 mOl. Peracca (1897) gave 890 mOl as the maximum total length but did not
indicate the sex. Scale rows arc 19-1915, smooth. without apical pits; a reduction
of the third and fourth scale row occurs between ventrals 98-1 18 (X = 106.9),
and reduction of the seventh and eight scale rows between ventrals 99-112
(.t= 106.1). The number of ventrals varies from 187 to 197 (X' - 191.6), the
number of subcaudals from 53 to 57 (.f = 54.6) (only five with complete tails).
The number ofmaxiUary teeth varies from 20 to 22 (X = 20.9). Head scales arc
as follows: supralabials 78 (I), 8-8 (7), 89 (I); supralabials entering orbit 4+5
(8),4+5/5+6 (I); ,nf",labials 9-9 (I), 9-10 (1),10-10 (5),10-11 (2); preoculars
11, postoculars 22, lorcals 11, temporals 1+ I (I), 1+211 + I (I), 1+2 (7). The
anal plate is divided. The tail/total length ratio varies from O. I 52-0.168 (.f0.159); eye diameter/snout length ratios vary from 0.467 to 0.578 (.t = 0.502).
The hemipenis is nine subcaudals long, with the sulcus spcrmatieus dividing
at the level of the third subcaudal. slightly bilobed (about one and a half subcau
dais), and very spinose. The smooth apical disks are pronounced.
The general dorsal color is light blue in preservative, but Peracea (1897) slates
that living specimens arc immaculate green dorsally and yellowish while ventrally.
The ventrals and subcaudals arc usually immaculate white in preservative. In two
preserved specimens there arc dark flecks and/or spots on the ouler edges of the
ventrals. Aecks cover all ventrals in one specimen and three founhs of the ventrals
in the other.
Sp"iml',ul'xamilU'd.-ARGENTINA: Salfa: Hic:km:l.nn IML lOS. 30]: Luna Muc:na IML 301:
FomlO1tl: IneenicroJuarez IML422. 490. BOLIVIA: no specific loc:iIIlily 10 tl17. 8M (unnumbered):
Chaco: Cail.a 8M I946. 1.S.69. PARAGUAY: Chaar. Lorna Plata KU 734:54.

Liophis ,iridis Gunther

Uophis l';rid,s GUnther. 1862. Hok>typc. BI\.I 1946.1.S.69. Typcloc:iIIlily-Bnu:il, PcmamblKO. no
specific 10C11;ly.
Liophis I}'ph/IIS prosino J;:1n and Sorddli. 1866. Hololype (lost?). Typc.locality-Brn7.il.

DIXO -SoUTH AMERI

19 7

I I

40

.......

.'

,
I~~ . . . . .,..... ; ,

}
I
(

'

"

~.... t, ... _.'~.:f'

..",

' ..

o
o

@(j)~

,"

_----:'
.;

,,

35

.'D

d)

I~

Fig. 2.- Distribution of Lio/1hi I'iridis in northeastern Brazil. DOIS wilhin the diagonal lines ( gre Ie
plant formation) and lightly stippled area (Atlantic rainforesl formation) represent the Agresl -Atlantic
Rainforest sample. Dots within the dashed line (Olatinga plant formation) represent the aatinga
sample. Densely stippled areas within the Caatinga formation represent enclaves of the en-ado plant
formation.

Distribution-Known only from the Caatinga Agrcste, and Atlanti rainforest

vegetation communiti of northeastern Brazil (Fig. 2).
Description. - The follo\ ing description of L. viridi is based on 159 individual
from throughout the range. The maximum total length of males is 595 mm and
of females 670 mm. Th scale fO\ s arc 19-19-17" smooth, and with one apical
pit. The redu tion to 17 ro\ occurs through a fusion ofscaJe rows three and four
between entral 98 and 1 1 (x = 110.0).
ual ariation \ as not d t ctabl
e en in large sample" therefore all scale data \ re combined for intra ample
comparisons. eotral ary from 169 to _02 (of = 1 6.0); subcaudals from 63 to
4 (.t = 74.1). Tai total length ratios ofaduh ary from 0.193 to 0.2 9 (x =
0.221). The e c/ noul ratios of adults ary from 0.-00 to 0.6
(x = 0.562.

182

ANNALS OF CAIll'EGIE r.IUSEUM

VOl..

56

Maxillary teeth vary from 17 to 24 C~ = 20.2). with the lasl (Wo 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
cnlering orbit 4+5 (155). 4+5/5+6 (3), 4+5+6/5+6 (I); infralabials 8-9 (2),
S-IO(I), 9-9(2). 9-10(4).10-10(144),10-11 (4). II-II (2): prc<lCul.r> I-I (156).
1-2 (2). 22 (I): postoculars 1-1 (I). 2-2 (157). 2-3 (I): lorcals 1-1 (159): temporals
1+1 (12),1+1/1+1(21).1+2(145).1+211+3(1).
The hcmipcnis is 9 10 13 (.f = 10.8) subcaudals long (ill S;II1) and bilobed for
1.5 to 3 subcaudals. Large spines occur from the base to the proximal part orlhc
lobe. Spines about onc half the size of the basal spines occur from the lobe 10 the
distal end. Th~ sulcus spcrmaticus is divided ncar thc basc of the lobes. Thc
smooth apical disk is rdati\'c1y large,
The dorsum is bright green in life. Young and small juveniles may ha\'e 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 sc\'cn and eight. The number
of spots is variable in any row. varying from 36 to 60. especially in individuals
from 180 to no mm in total length. Some young havc only a nape band. immediately followed by onc pair of dorsal spots, Occasionally young havc a black
latcrnltail stripe. Spots onju\'~nilcs from 240 to 370 mm in totallcngth arc faded
postcriorly. ormally all individuals over 400 mm in total length arc uniform
green and the belly and subcaudals arc immaculatc white. Vanzolini et aL (1980)
give a v~f)' good color description of living individuals of this species.
Geographic l'oriotiOll.-A Caatinga sample and an Agrcste-Atlantic rninforest
sample were formed from three coastal subsamples and two inland subsamples
of L. \'iridis (Fig. 2).
Thc Caatinga sample is significantly distinct (T = 13.4) from the AgrcsleAtlantic rainforest sampl~ in having a higher number of ventrals, 181-202 (x =
189.8), versus 169-188 (J: = 179.6), and a higher number of ventra Is (T - 6.44)
at Ihe 19-1 7 reduction sit~ 102-124 (x = I 14.6) vcrsus 98-1 16 (.f .,. 106.6). Othcr
scale and tooth chamcters arc not significantly different belween samples.
One specimen from Jeremoabo, Bahia. and five specimens from Mina de Sao
Felix do Arnianto. Bahia. arc known from thc eastern parl oflhe Caalinga. Their
taxonomic featurcs more closely approximate those of the Agreste-Atlantic rainforest sample than others from the Caatinga. Although the samples arc small [
suspect that they may represent intergrades bctween two well-defined subspecies.
The Caatinga sample (Fig. 2) does not have a formal namc. However, Jiln
(1863a. 1983b) and Jan and Sordelti (1866) mention a varietal name associated
with L. typltlus that belongs to L. I'iridi~. It was originally cited as a varietal name
(without description) by Jan (I 863a. 1863b) but was illustrated by Jan and Sordelli
(1866) and their iconolype represents the only description. Jan (I 863a. 1863b)
retorded two specimens of the variety. one from "Fernambueo" in the Milan
Museum, and one from "Brasilc" in the Stuttgart Museum. Jan and Sordelli
illustrated Ihe laller specimen, thus it becomes the holotypc (by inference). However. neither specimcn now can be located. I cannot definilely assign thc name
to cither of the IWO populations that require re<::ognition. Howcvcr. the genus is
already ovcrburdcncd with excessive synonyms and I propose that Jan and Sol'
delli"s varictal name. prasillfis. as proposed in the combination L. 'HJh!lls prasill/IS.
be recognized as thc proper name for the Caatinga popultion of L 'iridis.

1987

DIXON-SovTII AMEIUCAN L,Or/IIS

18'

Liophis l'iridis l'iridis Gunther

Uophls l"Ir1du Gilnlhcr. 1862.

Diag1losis. - Dorsally adults arc leaf green to bright grecn, and pale cream to
while vcntrally. Juvenilcs ha\'e a black nape band followed by a various number
of paired black dorsolatcral spots. The number of ventraIs varies from 169 to 188
(.t' - 179.6). and reduction occurs between ventrals 98 and 116 (.t - 106.6).
Distribwio'l. -Liophis I'. I'iridu occurs in the Atlantic rainforest and the Agreste
forest from Recife. Pcrnambuco. east and south 10 Salvador. Bahia. Brazil.
Liophis riridis prasinus Jan and Sordclli
l.wphu tJphlus /NtlSutO Jan and SonkJli. 1866.

Diag1losis.-Adults arc bright green. wit.h an immaculate cream to white venter.

Juveniles ha\'c 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
postcrior. latcral black stripes that extend Onto the tail. The number of ventrals
varies from 181 to 202 (.t' = 189.8). with the reduction occurring between ventrals
102 and 124 (.t' - 114.6).
Disiriblllio'l.- Uoplus ". prasimlSoccurs from Jooo Pereira. Maranhao in northeastern Brazil. south to sao Francisco. Minas Gernis. thcn northeast to about
Parauagua. Rio Grande do orte. Brazil.
Commt>III.-Vitt (1983) found Hyla ruhra, Pll)'salol"nlllS cllIit>ri. and several
unidentified Icptodactylids in 14 stomachs of L. I'iridis. Vin cxamined 16 gravid
fcmales and dctermined that L. "iridis produces continuous clutches throughout
thc ycar. Ovarian clutch size varied from two to six (.t' - 3.69) eggs.
Short taxonomic works for L. \'iridis include Cordeiro and Hoge (1973). Gomes
(1918). and Schmidt and Inger (1951). Vanzolini ct al. (1980) give an excellent
summary of scale data. color or young and adults, and brief biological notes on
a Caating.1 sample.
Spt.-c;III('IIS C".\"(Im/1/C"r!.-"SQUTH A1I.'1ERICA"; eM R281. IlRAZIL: no specific localil)' FMNH
5694. I\'IZUSP 81 5. NHMW 20739: no specifiC Slate. Parauagua ZMUC 601250. "'/(/JUs: Quebrangulo
MZUSP 3453. /JuMa: No spccifll;' localily FMNH 5721. III 3200. MCZ 2946. 147217-147218. Assu
un Torre ]I) 980. Brreiras UMMZ 108712-108716, Ilom Jesus uc lllp.1 Cm R344. UI\'IMZ 10871 l.
Fciro de Santana KU 29478. [tnpoan MZUSP 5713. Jercmoabo MZUSP 5738, Mina de S:1o Feliz
do Amianto III 28170-28172, 28215-28216. Salvador 11M 90.1.27.2, 1924.9.20.26, MZUSP 28382839. ("/'{/I'I/: Aeudl' Amanari Maranguapc MZU51' 3450. Boa Vi:lgem III 20200. Ca.xueirH ZMUC
3342(3). Coluna MZUSP 5319-20. Limol,iro do Norte III 12776-12777, Unique [U 20 142. Murl/l1hilo:
L1goa uc Jo~o Pereira CM R316. ,\Ii/laS G('ruis: Januaria UMMZ 108705-108709. Siio Francisco
UMMZ 108710. /'lIrt1hyba: Corcmas MZUSP 345J. Joiio l)e5soa MZUSP 7976-7977. 7999. 8262.
UFFB 139, 141. 143. Mamanguapc MZU51' 3456. I'trt/a/llIJl/co: no specific localil)' MANH 4448,
BI\I 1946.1.2.69. 80.II.2S.<I. 80. [ 1.25.8. MC2 1447. 146945-146947, AQreslina MZUSP 4925. 4942.
4946,4965. 4970. E~u MZUSP 6693-6699. 6700-6718. 7071-7091. 5 kill E E~u MZUSP 6950.13
km Exu MZUSP 6940-6943. 5 km N E.xu MZU5P 6920-6921. II km S Exu MZUSP 7092. Ponta
de Pooms MZUSP 5177. Recife DEH 624. 517. 526. LSUMZ36787.1';lIIlr; Acude Peri Peri MZUSP
3424.3427-3429.3433.3435-3438.3441. 3443. 3445. TCfCzina III 360. 473. 121~1214. Valenca
MZUSP 5814. Rro GrUlmf'do Nortf". Ce-.U-3. Mirim CAS 49320. SiJo 1'01110; I....emc 10 543 (in crrorJ.
Srrglpt': Amcaju IIJ 22458. Maruim zr...l UC 601251. $anlO Amaro das Brollu. MZUSP 6985. 7306.

Liophl's jaeger; (Gunther)

CorolJrl/tl)Qt'gf'rI Gunler. 1858. Synlypes- 8M 1946.1.9.12. 1946.1.5.78. T}pcIOC<1ll1y-Br.nil, no
specifIC l0C2hly.

I 4

ALS OF CARNEGIE
\

OL.

EU I

56

..- -. ..

#'

..~.""

+-'- '.

(-' -ttl ~-

, :.
,f

Ii.

....

.~

1';-

,,/:"

25

,,'

,',1"'-'_\

+
30

+
60

ss

50

45

Fig. 3.-Distribution or Liophi.fJQe ('ri in southeaslern outh merica. Black lines enclose sample
ulilizcd in Lbe analyses or g' graphic ariation. The large bla k d t represents several 0 erlapping
localities.
Liophis dorsalis Pelers. 186 . Type-Iocalily-Brazil, no specific localily.
..tporophis cora!li~ell/ris Boulengcr. 1894. Type-locality-island norlh or Concepcion. ncar an

vador. nOrlh Paraguay.

lVUJdillaea iiI/cola Jensen, 1900. Type-locality: Taboleiro

ai-

rande. Minas Geiras, Brazil.

Misapplied synonym. - Rhadinaea dichroa Werner (1899) has been in the synonomy of L. jaegeri sine 1972 (Peters and Orejas-Miranda, 1972). However an
examination of the type description reveals that the holotype has 19 midbody
cale rows 159 ventrals and 50 subcaudals. The dorsum is brown with the bases
of tJle scales white and the posterior edges black. It also has a tailItotal length
ratio orO.\85. The de cription is identical to that for Liophis poecilogyrus caesius.
a form common to th haeo of Bolivia. Argentina. Paraguay and Brazil, and 1
a ign it to that sp eie .
Distribution. - Liophisjaegeri occurs from about 19 latitude in Brazil to about
35 latitude in Uruguay and Argentina \ cst to about 61 longitude, along the
Rio Paraguay basin in rgentina, Paragua and Brazil (Fig. 3).
Description. - orne 41 individuals of L. jaegeri ere examined and full data
laken on 160. Tail/total I ngth ratios were recorded for 0 er 400 individuals. Th
ma.ximum total length of males is 539 mm, of females 616 mm. Dorsal seal s

1987

DIXON-SolTTH AMERICAN LIOI'IIIS

185

arc smooth, in 17 rows, normally without reductions, and without apical pits.
When reductions are present (15 individuals). they occur with a fusion of scale
rows three and four on one or both sides of the body. reducing to 15 or 16 between
ventrals 88 and 141 (,t = 121.2).
Sexual variation in the number of ventrals, subcaudals, maxillary leeth, and
taiVtotallength ratios is absent. therefore the samples were combined forstatistical
analysis. The numocr of vcntrals of 160 individuals varies from 146 to 169 (.f =
157.5). and sulx:audals from 52 to 75 (.f = 61.7). The number of maxillary teeth
varies from 22 10 29 (.t = 25.6). The laiVtotal length ratios varies from 0.181 to
0.268 (.t ". 0.221). The number of palatine and pterygoid teeth were recorded for
onc individual and lhcy wcre 10 and 25 respectively.
Hcad scales are as [ollows; sup",labials 6-7 (Il. 7-7 (Il. 7-8 (4). 88 (147l. 8-9(4).
99 (I); supralabials entering the orbit 3+4 (I). 3+4/4+5 (3), 3+4+5/4+5 (I).
3+4+5/4+5+6 (I). 4+5 (148), 4+514+5+6 (2). 4+5+6 (It. in[",labials 8-8
(3).8-9 (4). 8-10 (2), 9-9 (5), 9-10 (24l. 9-11 (I). 10-10 (112), 10-11 (5); preoculm
11 (155). 1-2 (2). 22 (I): po5toculars (2-2) and loreals (11). temporals I + I (5),
1-1112 (I I). 1+212+ I (I). 1+2 (141); eye diameter/snout length ralios of 20
adults varies from 0.533 to 0.844 (.t' = 0.655). The anal plate is divided in all
specimens. Hemipenial length varies from 7 to 13 (.t' - 10.3) subcaudals.
A hemipenis extending 12 subcaudals has the sulcus spermaticus divided at
the level of the 6th subcaudaL and thc lobes begin at the level of the 9th subcaudal.
Large spines occur on the asulcate side of the hemipcnis to the edge of th~ large,
smooth. apical disk. Calyces appear to be absent and only a weak basal naked
pocket is present.
The general dorsal color is dull grecn, olive grecn. 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 pans 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 arc presented by Miranda et al. (1982).
Geogl'QlJ/tic l'al"iafioll.- 160 individuals were grouped into nine geographic sam
pIes. Three samples were located in the Rio Paraguay basin in a north/south linc,
and six samples were scallcrcd from Uruguay to southeastern Brazil, morc or less
in a nort h/south Ii ne. Usi ng uni variate statistics, the num ber of ven trllls, maxillary
Iceth, 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
dcviation at'
I error, respectively. In addition, Student's T Test values
were used to ,
: significance at the 95% level, in a pair-wise sample com
parison. No significant differences in paiHvisc comparisons exist for any of the
samples arranged in the two north to south lines. There arc 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 numocr of ventra Is,
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 ( ,., 18) vary from 63 to 75 (.t = 68.5, 3.8, 0.9), while the
combined six eastern samples (N = 110) \'ary from 52 to 71 (X - 60.4. 3.4,0.3).

186

VOL.

56

the tail/total length ratios of the western samples val')' from 0.214 to 0.268 (.f 0.231. 0.29. 0.07) and those of the eastern samples from 0.193 to 0.248 (1' 0.221,0.13.0.01). This suggests that the subcaudals arc smaller in the western
sample. hence more subcaudals are present on a tail about the same length as in
[he eastern sample. The eastern and western samples are scparated by the Rio
Uruguay and Rio Parana basins and appear 10 be allopalric (Fig. 3). However.
additional collecting may reveal parapatry of the samples.
Since the number of subcaudals is significantly different between the two samples. I propose recognition of the following taxa:
Liophis jargeri corallil'e.ntris Boulcngcr

Diagllosis.-Dorsum ground color leaf green. somclimcs with a middorS31

brownish red stripe five to six scale rows wide, extending from the nape 10 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 ventrolaleral blackish marks on the edges of
the ventrals. Subcaudals vnry from 63 to 75 (.f = 68.5).
Distribution. -Known only from the Rio Paraguay Bnsin of Argentina. Paraguay. nnd Brazil (Fig. 3).
Lioph;s jaegeri jaeger; (Gunlher)
1...JOphu dona/u Pell'n. 1863.
1i~/ll J('1lsccn. 1900.

Rhadl/I(U."(I

Diagl/osis. -Dorsum olive green. grayish green or leaf green. often with a middorsal brownish or reddish stripe six to eight scale rows wide. sometimes with
dnrk daiS along scale row four. and occasionally along scale rows three and five.
Venter rose or reddish. wilh or without ventrolateral blaek marks on the edges
of Ihe vcntrals. Subcaudals vary from 52 to 7 I (.t' - 60.4).
Distriblllio". - Known only from the cast side of the Rio Parana Basin east to
the Atlantic coast. and from Uruguay nOl1h to 19"5 lalitude in Brazil (Fig. 3).
COII/mem.-Littie is known aboul the natural history of this species. I have
found 7 and 9 eggs in two specimens, and Miranda Ct aJ. (1982) indicated Ihat
Ihe avcrage numbcrofeggs forthis species isaboul 14. Miranda el al. also indicated
Ihat the species is diurnal. frequently found in humid places, and cats small frogs.
Notations on some field tags suggest that Ihe species also occurs in brushy arcas,
has been found "crossing dirt roads" and "in a puddle of a cow Irack in low
campo, 10:30 am, 84F. and in full sun:'
Dixon (1985) presented evidence that L. jaegeri and L. maryellellae hybridize
in an area near Uelo Horizontc, Minas Gerais, Brazil. Additional material from
the lalter area has 110t been located, and the hybrid zone remains undefined.
5"NXill/('l/s exammffl. -(I. jat'J;(''; corallil't'lI/ris). t\RGENTINA. Corrielltes: Gral Pa:r.. <:Slcros de
Iii Santa Lucia IML 802. ltu7.aingo 1~'1L 1180. Rosario: Rosario MNHP 1899.6-7. Sa/lla Ft.": Da":ados
del Rincon CM 39130, Duenos Aires MNHP 67650. Laguan Guadalupe eM 39128. URAZIL, MUlo
Grosso: Corumba 18 26S7S. Rio t\pa UMt\IZ 108761. upper Rio Pal1ln:tlRio Paraguay UNM 68831.
PARAGUAY. Asuncion BM 1930.11.27.208. Bunl3\'ist.a. NE ofCaacu3nu MCZ 34204. Isla Con
ttpclOO BM 1946.U.8S.I'rima\'cl1I 8M 1%O.1.2.84-8S. 1960.1.3.40. 1962.81.
(Uophis jrwgm )Qt'gm). BRAZIL ,Hmas Gnats: no specific klcality MCZ 20715-20716. lido
Horizoole UMMZ 109069. Campo AI~ IB 196. Coronel Fabriciano IB 22993. Imbuzdm USNM
100676. Itapc\"lI t8 41126-41127. JUIZ de Fa>:! III 30728. 33SOI. Mutum IB 3802. No\'a Badcm tB
192. Curo PretO III 897. 3311. Pusa Vintc IB 4968. PocosdcCakl.lIIlB 4714. Santa Rita de utrema
IB SS60. 8192. TaboiciroGrandeZMUC601253. Toledo IB 44S31. UNi IB 17308. ParoM:Araucaria
18 72S2. 7346. 13632. Balsa No\'J 18 18293.1939-22941. 2294J...22944. 23112, 1J1I~12I1S.

1987

Dllw:-;-SoUTH AMERICAN LIOl'IIIS

IS'

14343-24345.24466.26226.28785.32538. Campo Largo MINU 102257. Carambei III 7403. Castro

m6244-6247. 7931. 10392. Curilioo UMMZ IOS722. m197. 23006. 23439-23440. 29321. Dori:wn
18893. Guarapua\'a IIJ 24630. 24661. 24710. 24889. 26954. J3&uanah'a m 17842. Jooqulm Munmho
1118263. JoSo ElI&Cmo IB 13027. 15100. MallellB 18115. 19544. 19670. l'aul0 Frontin IIJ 2498624981.25321. Pirai do SullB 12425. Rio Azul III 21346. 22801. Tronco 111130. Rio ikJalK'lfO:
Friburgo MZUSP 210S-2107. Muri 18 21261. 219:?9. 24579. Peu'Opolis m 21-113. 21475. Rio de
J:U'lClro (l\.lorro cia Urn) 10 985. Rio Gralllk do Sill:. no spccilk Ioeali1y. 8M 82.10.4.60-82.1004.61.
6IA.l\.lCZ 5633. 17932-11933. MZUSP 1109. 1112-1113. 1127-1121. UMl\.lZ67241. ZMUC6012S4601255. Akundnnha MZUSP 54n. Cacequi 18 10025. Canda m 13526. Calliu do SuIIl164II.
6716.9316. Cachocil1l do Sui 23]15. 23317. CnuAlla 18 11]16. Eslado Ecolo&lca do Taim MZUSP
752-1. Gu:uba CASI96n. Parm No\'o 1Il7&t5. Passo Fundo IB 8049. 8060. 1212. rc-Iolas 1841057.
PonoAkvc 18 7264. 7572. 11015.21411.12509-225 II. 22565. 22132. 23 119. 23-& 19-23420. 23S43.
45935. Quebra [knIt-Maran 18 23302. Rcstin~ Stca 18 7305. 9831. S10 Joio MNHP 1194A.lO.
sao I.zopoIdo MSUSP 4100. 18 7300. 7352. 7776. 8357. Sonta Calanna: 8:mKaS 18 13319. Canoinhas. I\t;IIr Tauna)' 18 24612. haiOpolis 18 16669. Lap do None- 18 9700. Pono Unllo 18 25992.
sao JoaqUlm 18 45187. Tauna~ 18 16328. Tre:s Barras 18 7948. Silo Palllo: no qJCCIOC Ioc:Ihl~ MC2
207110-2071}. UMMZ62804. 108760. Agudos 18 22097-22100. Abcmc:uia 18 955]. AlunOpohs
18 19562. Araraquara 18 19413. Arpui 18 831. Atibaia 18 23 149. 23254. 26889. 26889. Anh:lnqlba..
KM 24 1823163. lJ:urinha 18 43D3. Barum 18 6987. lknlo Otnrino 18 20641. Boquenio da Praia
Gr.andr 18 11524. Boracria MZUSP ~98. 80lllCliU MZUSP 2406. IJr.apnt':l huhsu IB JO.l98.
CalC'iras 18 41061. CalGo O'AllU3 18 5459. 5682. 6716. Campo larao 4639. Campo l.1mpo 18 4738.
5309.5745.7349.7541. 18658. Campos; do Jord3o 18 8450. 30692. MZUSP 1122. Capiliu\'a 18
3070. Cotl3 1IJ 7504-H05. 24281. 2919S-29196. Embu km 28. 18 JO.UI. JO.l90-JO.l94. 305741.
Embu G~ 18 16617. 167796. ]1299. Estado Gabriel Pi7;1 IB 5428. Guarulhos 18 SOJI. 23556.
23724. 41961. MZUSP 4121. Ibiuna 18 26979. 32251. 32265. 40930. 42203. 42222. 42337. 43764.
Ilapellnlnp 18 10287-10288. Ilapcccrica da Serra 1841115. -121]8. 42942. 4]882-43883. 44116.
44189.44498. hanba 18 5551. 5586. 57.11. 2.118>'-24186. ltaquera IB 10189. 10197-10200. IIU 18
3223-1. Jac:uri IIJ 3293. 24926. 28502. Jarinu 18 12810. 27542. 30186. Jundlai IB 19636. L6bo 18
32B. macNonla IB 7026. Mairipora IB 2-1380. 269.19. 27958. ~Iairinque ID 6042. 6048. 6075. 6618.
~101i das Cruus m 10196. 10268. 10295. 10]18.21387.21395.22596. MorciraslB 17606.~
cmClras IB 5470. Pendbe' IIJ 5532. Pcrus llJ 3292. Pindamonhanpba 18 18820. Pinhcirinhas IB
32420. Piriluba IIJ 908. Pilar do SullO 32121. 3145". Quiljuna IIJ 6257-6258. Raiz da Scm MZUSP
1114-1116. Ribclr::lo Pires [0 10232. Ribeir.i.o Pr"IO 18 228]1. Rodovia Dos IJandciranlcsdnl 60.
m 34397. Sallo IB 4516. 4578-.1579. sao Bento do SaplK':li IB 3283. ~o Ikrnardo do Campo MZUSP
3289. 3612. 4463-4465. IU 22608. 42919. S.io Lourenco da Scm IB 45789. sao Paulo MZUSP 1117.
1119. 1950.2372. 2387. 2609. 2635. 2960-2961. 3355. 3688. 4095. 4130. 4756. au 191. 19]-95.
647.695.1315.1321-1322.1416-1417.3252.349-1.4643.5411. 5573, 6189. 7646. 8012. 8575.
9561.9842.9879.1019.... 10195.10252.10271.12214.12977.12986.17085.17871.18270.18405.
18920.22882.14178.24431. 24442. 26656. 27014. 27157, 27212. 27281. 27478. 27498. 27556,
27571. 27631. 28308. 28503-28508. 29222. 29236-20237. 29249. 29614. 29924. 30683. ]2446.
32504.32800.3280]. 32925.32999.34277.41 129.42109.42195.44162,45746.45751.~'0
Roquc
11} 5391. 44228. Serra dn Bocnina MZUSP 4912. Sorocaha 1I3 6865. SoUf.:IS IB 23151. Sutnnrc lB
1332. SU7ano [1120376. Tnpimi IB 28060. TrcmemlX: IH 10368. Vnlentim GcnlillB 28671. Vinhcdo
III 30320. URUGUA Y. ('(lIlI'loIlI'S: no specific localilY BM 84.2.23.39. FMN H 10215. Arroyo Solis
Gr:lI\dc CM 38959. 11lll1mlos de Carrasco C/l.l 55422-55423. Roehll USNM 89995. I{Ulll Inlcrbulc
neMi:t CAS 93100. A(OII/{'l'i(/('o: no SIX"l'irlC locality MNHP 171. Snlltiallo Va1.(jllC'f. LSUMZ 27753.
MNMHI' 168.

Liophis mar)'ellenae Dixon

LiO/I!li.f waf)Y'/II'IIfIt' Dillon.

1985. Hololypc-AM NH 62202. Typcloc:llily-Annllpolis. OO;;lS. Urazi!.

Descri/JtiOI/. - The following description is based on 10 specimens. Maximum

10lal length of males 435 mOl, of females 530 mOl. Tail/tolal length ratios vary
from 0.221 10 0.262 (X = 0.240). Diameter of eye/snoul lenglh ralios "ary from
0.605 10 0.844 (of = 0.686). Dorsal scale rows arc 19-19-17. smooth. with an
apical scale pit. Reduction 10 17 rows occurs belween ventrals 74 and 95 (.f =
82.8). The anal plate is divided. The number of venlrals varies from 144 to 159
(.f - 150.9). The number ofsubcaudals varies from 63 1082 (.f "" 67.9). The
number of maxillary tccth varies from 25 10 28 (.f ... 26.1). Head SC3lcs vary as

188

ANNALS OF CARNEGIE MUSEUM

YOLo

56

-"0

SCAL.[
~~o

.~(Io

6~O \oIIIU:i

.. IL..(;I ....

~T CII,~

SINUSOIDAL PROJeCTION

Fig. 4.- Distribution of Liophis alrivenle,. and L. maryellenae in southeastern Brazil. Dots represent
L. marJle/tenae; the square L. alril'en!er.

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

10- LO (8), I I-I I (2); preoculars I-I (10); postoculars 2-2 (10); loreals 1-1 (I 0);
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 southeastern Brazil (Fig. 4). For specimens examined, see Dixon (1985).

1987

DtXON-SouTH AMERICA.....

189

L/OI'IIIS

Liophis atd.enter Dixon and Thomas

Dixon and Thoma~ 1985.

tado Biokl&iaJt, Silo !'auk>, Br:I7JI.

Uop/rUQIfII't'ntl'r

HolotYJX'~f\IZUSP 5066.

T)"I)Clocality-80rattia (E$-

Descriplio1J. -the following description is based on three specimens. Maximum

total length of male 505 mm in snout-vent length). TaiVtotal length ratios vary
from 0.195 to 0.214 (.f = 0.203). Dorsal scales are in 1919-17 rows. smooth.
without apical scale pits. The reduction to 17 rows occurs between ventrals 73
and 79 (.f - 76.7). The anal plate is divided. The number of ventraIs varies from
141 10 148 (.f "'" 144.0). Number of subcaudals from 49 to 56 (.t - 52.5); the
number of maxillary leeth from 23 10 24 (.f = 23.2). Head scales arc as follows:
supralabials 8-8 (3), supralabials entering orbit 4+5 (3), infralabials 10-10 (3).
preoculars 1- I (3). postoculars 2-2 (3). temporals 1+2 (3).
The hemipenis of the onJy known male is short. 5.5 subcaudals long (in situ).
and bilobed for the distal hair. The sulcus spermaticus divides about one third
of the dis13ncc from the base. The distal end of thc lobes contain smooth .apical
disks. Fivc large spines arc present along thc outer surface of cach lobe. ncar its
base. while the inner surface of the lobe is barco In addition to the larger spines.
many smaller ones are scattered over the shan.
In preservative. the dorsum is uniform olivc brown. The throat and first sevcn
vcntrals arc grayish while. and the remainder become progressively darker until
the posterior two-thirds of the ventcr is black. The subcaudals arc grayish with
OCC3sional dark smudges.
Distribution. - This taxon is known onJy from the type locality (Fig. 4). See
Dixon and Thomas (1985) for specimens examined.
Key to the "Gr(!f'Il" Liophis ofSowh America

I. Scale rows 19-19-15 or 19-19- I 7

Scale rows 17-17-17or 17-17-15
2. Scale rows 19-19-15
.
.
Seale rows 19~19-17
3. Ventrals 133-172. one scale pit
Ventrals 187-199, no scale pit..
.
4. Ventrals 141-159, maxillary teeth 23-26
Ventrals 169-202. maxillary teeth 17-24
5. $ubcaudals 62-82. belly yellowish orange
$ubcaudals 49-56, belly black

2
jaegeri
,................. 3
, ................ 4
lytJhlus

Kllelllheri

5
I'iridis
lIwlJ1elle/lae
all"il'ellfer

ACKNOWLEDGI'.'1ENTS
I thank the following eur:lIors who have indulged my requests for specimens over many )ears R.
L lJel.Y and J. W. Wright. Los Angeles County Museum of Natural History (!.ACM); J. A. Campbell.
Unh'ersity of TCl\:as. Arlington (UTA); C. L. S. Cordeiro. Instituto DUlanlan (ID); H.C. Drt.'SS:lucr.

(HCD); K. C. Drewes and A. E. Le\iton. California AaJdemy of Science (CAS); W. E. Duellman.

Uni\ersity of Kansas Mu:;c:um of Natural History (KU); W. R. HC)er. R. McDiarmid and G. Zug.
United StattS National Museum of Natural History (USNM); M. S. Hoogmoed. Rijksmuscum van
Natuurlijke Historic. (KMN Ht. r\. G. Kluge and R. A. Nussbaum, Uni\'crsity of Michigan Mu:;c:um
of ZooIOSY (UMMl); W. W. Lamar (WWL); V. MlIhncl1. Museum d'Histoire Naturellc. GCIlC:\c
(MNHG): E. Malnalc and T. U7..lell. AaJdcmy of NaLUrnl Sciences. Phillldclphia (ANS?); H. Man:,
Field Museum of Natural Hislory. ChiGlgO (FMNH); C. J. McCoy. Carnegie Museum (eM):
W.
M)"ers:md R. G. Zw~ifd,Amcrit3nMuseum ofNaLUral Hislory(AMNH);J. B. Rumussc:n. Zoologisk
Museum Uni\crsitt'lsparkcn. Kobenha\'n(UZM): D. A. Rossman. Louisiana State Uni\'crsity MlolKUm
o(Zoolocr(LSUMZ): R. Rom. Museum ational d'Histoire Naturellc, Paris(MHNP); A. F. Slimson.
British Museum (N;lIural Hislory)(BM); P. E. Vanzolini. Muscode Zoolocia. Uni,'crsi<bdc:S3o Pauk>,
(MZUSP. OEH); S. E. Williams. MlISC'um ofComparali,'C ZooIolY (MCZ).

c.

190

ANNALS OF UIlSEGIE MUSEUM

I express m>' appreciation 10 my wife 1I1Iar)". who has shared my pursuit of LioplJis and herpetology

in gene",l. for morc than 30 years.

LITERATURE CiTED
A~t.\RAl, A. 1.lO.

1931. Studies ofncOIropical ophidia. XXI II. Additional nOll'S on Colombian snakes.
Bulletin Al1livcnin [ns\itUiC of America. 4;85-89.

- - - , 1935(1, Estudos sobre ophidios ncotropicos XXXII. Ap0nlamentos sobre a fauna da Colombia. Mcm6rias do [nstitu[o UUlant:m. 9:208-216.
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246.
BoUI.ENQER, A. G. 1894. List ofrcpliks and balr.ll.'"hi:lAs collected by Dr. J. Uohls ncar Asuncion.

Paraguay. Annals and Mag:t7.inc of Natural l-iislOry. (6)13:342-348.

COPE. E. D. 1870. Seventh contribution 10 the herpclology of Tropical America. Pnx-ecdings American Philosophical Socicl). 11:1'l1-169.
1887. Srnopsis of lhe U:ural.'hia and Reptilia obtained br H. H. Smith in Ihe pm\'incc of
~lalO Grosso. Brnril. PTOI.-'dings American Philosophical Sociel). 24;44-60.
CORDORO. C. L. A1<1> A. R. HOGE. 1974. Conlribuido ao ronhccimenlo das scrpcnles do eslado
de Pernambuco. Mem6rias do InSlilulO lJut:mt:m. 37(1973):261-290.
CUl'Il..... A. R.. AND F. P. AOMEr-TO. 1976. Ofidios da i'\maz6nia. X. ,\s Ibm da ragiao lem' do
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DIXON. J. R. 1985. A new species oflhe ncolropical snake genus Uophis from Brazil. Proceedings
Biological Socicly Washinglon. 98:295-302.
DJXON. J. R.. ANll R. A. THOM,\S. 1985. A new species of South American W:lIer snake (genus
UOfJlu's) from southeastern Br.l7.i1. Hcrpclologil.'a. 41:259-262.
GANS. C. 1960. Notes on a hcrpcloiogiclil collecting trip through the southl.'astcm lowlands ofBoliv!:I.
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GO~I~.l. F. 1918. Contribuil;:io parn 0 conhecimento dos ophidios do Brasil. 11. Ophidios do Museu
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GRIF~lN. L E. 1916. A l.'atalog of the Ophidia from South America at present (june 19(6) contained
in the Carnegie Museum. withdescriptionsofsome new spt:dcs. Memoirs ofthe Carnegie Muscum.
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GU!'>TltElt. A. 1858. Calalogue of Ihe colubrine snakes in the collCClion of Ihe Brilish Museum.
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- - - . 1862. On new spccicsofsnakcs in the collection oflhe IJritish Museum. Annalsand Mag;uine
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!I....""'R. J. 1974. Avian speciation in tropical South America. Publication of lIN: NUliall OmithologjcaIOub.14:1-39O.
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in Ihe Nalural Musca in Munich (Gcrm:my) and Lciden (The Netherlands). Spixiana. Suppl. 9:
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JAI'<. G. l86]a. Enumcra7.ione sislemMic;1 degli oFldi appartcnenti al gruppo Coroncllidac. Arehivio
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JAN. G .. ANn F. SoRI>ElU. 1866. lconogmphie gcnernlc des ophidiens. Tome Second Volume. Livr.
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1987

DIXON-SouTH A.\tEJlICAN LIOI'IIIS

19'

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z.