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1987 Liophis - Green Species of South America - Dixon PDF
1987 Liophis - Green Species of South America - Dixon PDF
CARNEGIE
MU
M
28 AlXlUST 19 7
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
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
174
~'OL
56
19 7
DtXON-SOUTH
AMERICAN L'QPIfIS
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 .
176
VOL
56
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.
1987
DIXON-SoUTIi AMERICAN
L,opms
177
(Linn.acus)
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
178
VOL,
56
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).
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
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.
19 7
I I
40
.......
.'
,
I~~ . . . . .,..... ; ,
}
I
(
'
"
..",
' ..
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.
182
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
18'
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.
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
ai-
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
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
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
IS'
188
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.
1987
DtXON-SouTH AMERICA.....
189
L/OI'IIIS
Uop/rUQIfII't'ntl'r
HolotYJX'~f\IZUSP 5066.
T)"I)Clocality-80rattia (E$-
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.
c.
190
I express m>' appreciation 10 my wife 1I1Iar)". who has shared my pursuit of LioplJis and herpetology
1931. Studies ofncOIropical ophidia. XXI II. Additional nOll'S on Colombian snakes.
Bulletin Al1livcnin [ns\itUiC of America. 4;85-89.
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- - - . 193511. Collceta hCrprl0logica no cenlro do Ilrasil. Mcm<>rias do Irlstitulo (Jutantan. 9:235-
246.
BoUI.ENQER, A. G. 1894. List ofrcpliks and balr.ll.'"hi:lAs collected by Dr. J. Uohls ncar Asuncion.
1987
19'
r"KK(K. H. w. 1928. NOles on repliles and oolrachians from Mallo Grosso and E. Both'ia. Annals
and Mag:u.ine of Nalural History, (10)2:96--99.
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of London. 19l5:SOS-S.lO.
rr:.....cc". M. G. 1897. Vl:iwo dd Dou. Alfredo Bo~1Ii nd Oxo Boli\'Qno c ncl"" Rcpubblica
A~ntin:r.. Renili cd anfibi. Bollenino Musci di ZooIO!ia rei Anatomia Comparau. Tonno. 12:
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10 Bra7.ll rlCldiana: ZOO'OEY. 3 1 :~39-46:S.
V....~SI. P. E......-.tu E. E. WILLIAMS. 1970. Sourn Amttlcan anoks: the ~phicdilkrmtiatlOn
and C\'oIutton of lhe Anolis chryw/tpis group (Sauria. IpanKbc). Arquh'os de Zoo&ocia, $30
PaulO. 19:1-298.
v....~st. P. E.. A. M. M. RA....os-COST.... AND L J. VITT. 1980. Rcplris das Cutmps. Acadama
8n1silcin de Ciencias. Rio de Janicro. 161 pp.
Vrn. L J. 1983. Ea>logy of an anuran-eating guild oflcrrestnal troplal snakes. HC'fl)eloloca. 19:
S2-66.
W,\Clt.u, J. 1824. Sct'penlum brasihensium speciel; no\OII.' 00 hislOI~ nalu~11c des nprs noU\'elles
de serpens. rueillics 1.'1 obsc",recs pendanl Ie \'O}'agc dans rinterieur du Iknil dans Its allnccs
1817. 1818. 1819. 1820. cs:cnllc par onfIT' de sa Maple Ie roi de lb.\ie~. pubti prilr Jean de
Spu..... ecntc d'spre5 les nOles du \'O)-ageur par Jean Waglcr, Monachii. Franc. Seraph. Hubsch
mann. 7S pp.
WI:IINI:II. F. 1899. Ikseh~ibung dniger ncucrSehlangen und Balrachier. ZOok>&isc::her An:(Ci~r. 22:
114-117.
- - - . 1925. NI'Ul.' odcr wenig bck:lnnte schlangen ausdem wiener naturhiSlllrisc::hcn Slaalsmuscum.
SilZUnl5bcrichte dcr Akadcmk der WiSSCIIschaften in Wien. 1]4;45-66.
z.