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Herpetologica, 66(1), 2010, 80–85
E 2010 by The Herpetologists’ League, Inc.

         A NEW SPECIES OF SIBON (SQUAMATA: COLUBRIDAE) FROM
                           WESTERN PANAMA
                      GUNTHER KOHLER1,3, SEBASTIAN LOTZKAT1,2,
                               ¨                                              AND   ANDREAS HERTZ1,2
 1
     Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany
       2
        Johann Wolfgang Goethe-University, Institute for Ecology, Evolution and Diversity, BioCampus—Westend,
                                 Siesmayerstr. 70, 60323 Frankfurt am Main, Germany

                                                                                                            ´
            ABSTRACT: We describe a new species of Sibon from the Reserva Forestal La Fortuna, Chiriquı Province,
          Panama. The new species is distinguished from all species presently assigned to Sibon by having 17 rows of
          dorsal scales at midbody and 15 rows one head length anterior to cloaca (vs. 13 or 15 rows of dorsal scales at
          midbody without reduction on posterior portion of body in all other species of Sibon), and dorsal scale rows
          that change orientation from the sides of body toward middorsum (vs. maintaining more or less the same
          orientation). A key to the Lower Central American species of Sibon is provided.

              RESUMEN: Describimos una nueva especie de Sibon procedente de la Reserva Forestal La Fortuna,
                               ´        ´
          Provincia de Chiriquı, Panama. La nueva especie se diferencia de todas las especies actualmente asignadas al
            ´
          genero Sibon por tener 17 hileras de escamas dorsales en la mitad del cuerpo y 15 hileras una longitud de
                                           ´                                                                  ´
          cabeza anterior a la cloaca (13 o 15 hileras de escamas dorsales en la mitad del cuerpo sin reduccion en la
                                                       ´                       ´
          parte posterior del cuerpo en todas las demas especies de Sibon), ası como hileras de escamas dorsales que
                                 ´
          cambian su orientacion desde los flancos del cuerpo hacia el centro del dorso (conservan su orientacion  ´
                                         ´
          longitudinal en todas las demas especies de Sibon). Presentamos una clave para las especies de Sibon que
                                           ´
          ocurren en el sur de Centroamerica.
            Key words: Colubridae; New species; Panama; Sibon; Squamata


   THE GENUS Sibon is a moderately diverse                          took additional data for the Lower Central
group of small to moderate-sized snakes that are                    American species of Sibon from McCranie
distributed from Nayarit and Veracruz, Mexico,                      (2006), Savage and McDiarmid (1992), and
throughout Central America, and well into                               ´
                                                                    Solorzano (2002). Abbreviations for museum
South America to Ecuador and Brazil. Recently,                      collections follow those of Leviton et al. (1985)
                                   ´
McCranie (2006, 2007) and Solorzano (2002)                                                       ´
                                                                    except for CRB (Coleccion de la Reserva
described new species of Sibon, bringing the                                   ´
                                                                    Bartola, Rıo San Juan, Nicaragua). Specimens
number of Central American species in this                          in the Appendix labeled with SL field
genus to 11 (Ko  ¨hler, 2008). Three fieldwork                      numbers will be deposited in the Universidad
seasons in Panama (January 2006, January and                             ´                   ´      ´
                                                                    Autonoma de Chiriquı, Davıd, Panama, and
February 2007, and May to August 2008)                              those labeled with JS field numbers will be
produced several specimens of Sibon annulatus                       deposited in the Museo de Ciencias Naturales
and S. nebulatus. Among the specimens collect-                      de la Universidad Centroamericana (UCA),
ed was a single adult female of Sibon that differs                  Managua, Nicaragua. We follow Dowling
strikingly from all its congeners in the number                     (1951) for the method of counting ventrals
and arrangement of dorsal scale rows. Compar-                       and Savage (1973) for the terminology of
isons with the known species of Sibon from                          scales in the loreal region of the head. The sex
Central America demonstrated that this speci-                       of the holotype was determined by visual
men represents an undescribed species.                              examination of relative tail length and basal
                                                                    girth. Head and scale measurements were
          MATERIALS AND METHODS                                     made to the nearest 0.1 mm with dial calipers
  The format for the description of the new                         held under a dissecting microscope. Snout–
species generally follows that of McCranie                          vent length and tail length measurements
(2007). We provide a list of the specimens                          were taken to the nearest millimeter alongside
examined for comparison in the Appendix. We                         a meter stick. Terminology for measurements
                                                                    is abbreviated as: snout–vent length, SVL; tail
     3
         CORRESPONDENCE: e-mail, gkoehler@senckenberg.de            length, TL; head length, HL; snout length,

                                                               80
March 2010]                                   HERPETOLOGICA                                              81




   FIG. 1.—Holotype of Sibon perissostichon (SMF 88716)
in life, SVL 458.0 mm.


SL; and head width, HW. Eye length equals
the horizontal distance across eye. Scale
dimensions were measured at the longest or
widest points along the longitudinal or per-
pendicular axis of the body, respectively. The
capitalized colors and color codes (the latter in
parentheses) are those of Smithe (1975–1981).
Drawings of the head were made using a
camera lucida attachment for a dissecting                   FIG. 2.—Head of holotype of Sibon perissostichon
microscope. A small tissue sample of For-                 (SMF 88716). Scale bars 5 2.0 mm.
schungsinstitut Senckenberg (SMF) 88716
was taken from the edge of a ventral scale                rows one head length anterior to cloaca (vs. 13
about midbody and deposited in the tissue                 or 15 rows of dorsal scales at midbody without
sample collection of the Senckenberg For-                 reduction on posterior portion of body in all
schungsinstitut und Naturmuseum, Frankfurt,               other species of Sibon), and dorsal scale rows
Germany.                                                  that change orientation from the sides of body
                                                          toward middorsum (vs. maintaining more or
              SPECIES ACCOUNT                             less the same orientation). Additionally, S.
                                                          perissostichon differs from its congeners by
        Sibon perissostichon sp. nov.
                                                          the following characteristics: subcaudals 108
                 (Figs. 1–4)                              in S. perissostichon vs. 113–126 in female S.
   Holotype.—SMF 88716, adult female as                   annulatus. Light interspaces between dark
judged by the shape of the base of the tail,              bands on body Sayal Brown (223C) in S.
collected near Lost and Found Ecohostel,                  perissostichon vs. interspaces orange to red-
Reserva Forestal La Fortuna, 8u 40.479 N, 82u             orange middorsally, grading to cream laterally
12.979 W, 1434 m elevation, by Sebastian                  in S. anthracops. Enlarged penultimate su-
Lotzkat on 14 May 2008. Original field                    pralabial not in contact with eye in S.
number SL 145.                                            perissostichon vs. this scale in contact with
   Diagnosis.—Sibon perissostichon can be                 eye in S. argus and S. longifrenis. Also, dark
distinguished from its congeners by having                dorsal bands or spots reach well onto venter or
17 rows of dorsal scales at midbody and 15                even across venter in S. perissostichon vs. dark
82                                            HERPETOLOGICA                                       [Vol. 66, No. 1




                                                            FIG. 4.—Scalation (middorsal portion of body) of (a)
                                                          Sibon perissostichon (SMF 88716) and (b) S. annulatus
   FIG. 3.—Holotype of Sibon perissostichon (SMF 88716)   (SMF 88715). Scale bars 5 2.0 mm.
at its capture site.
                                                          no dark dorsal bands and venter immaculate
dorsal bands or spots reach at most to outer              in S. manzanaresi and S. sanniolus. Ventrals
edges of ventrals, although dark shading may              186, subcaudals 108, bands solid dark brown,
be present on venter in S. argus and S.                   darker than interspaces in S. perissostichon vs.
longifrenis. One anterior temporal present in             ventrals 168–171 in females, subcaudals 95–
S. perissostichon vs. no such scale in S. carri.          100 in females, bands dark brown ventrally,
Therefore, penultimate supralabial not in                 their centers paler brown, paler than inter-
contact with parietal in S. perissostichon vs.            spaces in S. miskitus. Postmental present;
these scales in contact in S. carri. Bands solid          therefore, first infralabials not in contact with
dark brown, darker than interspaces in S.                 each other, vertebral row not enlarged in S.
perissostichon vs. bands dark brown ventrally,            perissostichon vs. first infralabials in contact
their centers paler brown, paler than inter-              with each other, no postmental, vertebral row
spaces in S. dimidiatus. Dorsal surface of head           moderately enlarged in S. nebulatus.
with black markings, ground color of body                    Description of the holotype.—Adult female
brown, postmental single in S. perissostichon             (Figs. 1, 2); SVL 458 mm; TL 210 mm; HL
vs. dorsal surface of head with green or yellow           15.0 mm; HW 9.0 mm; SL 4.2 mm. Head very
as well as red and black markings, ground                 distinct from neck; snout short, blunt in dorsal
color of body green, and postmental divided in            and lateral outline; rostral wider (2.6 mm) than
S. lamari. Ventrals 186, loreal in contact with           high (2.0 mm), not extending posteriorly
eye in S. perissostichon vs. ventrals 155, loreal         between internasals, its length visible from
separated from eye in only known female                   above one-quarter length of median internasal
specimen of S. linearis. Dark dorsal bands                suture (1.2 mm); internasals shorter (1.5 mm)
present, reaching well onto venter or even                than wide (2.0 mm), two-thirds length of
across venter in S. perissostichon vs. usually            prefrontal suture (2.6 mm); prefrontals longer
March 2010]                               HERPETOLOGICA                                           83


(2.6 mm) than wide (2.2 mm), their median            Brown (223B) with Jet Black (89) blotches;
suture two-thirds length of frontal; prefrontals     ventral surface of head dirty white with Raw
bordering eye above loreal; frontal longer           Umber (223) mottling and Jet Black (89) chin
(4.1 mm) than wide (3.5 mm), widest anteri-          shields and first ventral scale; iris Mars Brown
orly, length two-thirds that of parietal, in         (223A). Color in preservative (70% ethanol)
contact with prefrontals, supraoculars, and          similar to color in life.
parietals; supraoculars longer (3.6 mm) than            Etymology.—The name perissostichon is a
wide (1.6 mm), nearly as long as frontal;            noun in apposition and is derived from the
parietals longer (5.9 mm) than wide (3.8 mm),        Greek words perissos (beyond the regular
widest anteriorly, their median suture (4.4 mm)      number or size) and stichos (row, line),
slightly longer than frontal; parietals in contact   referring to the high number of dorsal scale
with frontal, supraoculars, upper postoculars,       rows in this species.
anterior and posterior temporals, and eight             Natural history notes.—The holotype was
nuchal scales; nasal divided, in contact with        encountered at 2053 h in secondary Premon-
first two supralabials, loreal, prefrontal, inter-   tane Forest while it was active within the
nasal, and rostral; loreal single, longer (1.6 mm)   dense, fog-moistened undergrowth about 0.5 m
than high (0.7 mm), its length less than one-        above the forest floor (Fig. 3). At the time of
half length of snout (tip of snout to anterior       encounter, the snake was moving about on the
edge of eye) and one-half length of eye              inclined trunk of a young tree covered with
(3.3 mm); preoculars absent; postoculars three       foliose hepatics. The undergrowth surround-
(right) and two (left); temporals one plus three,    ing this trunk consisted of herbaceous as well
anterior one above fifth supralabial, lowest         as young woody plants, among which we could
posterior one above sixth supralabial; suprala-      distinguish different ferns as well as members
bials six, with third and fourth (right) or fourth   of the families Rubiaceae and Melastomata-
(left) bordering eye (enlarged penultimate           ceae. The type locality is situated about 20
supralabial separated from eye by lower              altitudinal meters below a narrow ridge on a
postocular); pupil vertically elliptical; eye        very steep slope (approximately 45u inclina-
length two times length of loreal; infralabials      tion) facing the southern Pacific lowlands.
seven, first pair separated posterior to mental      Most trees’ diameters at breast height do not
by a single, triangular postmental, first infra-     exceed 50 cm. The dense canopy layer extends
labial in contact with postmental, infralabials      from about 10 m aboveground upward.
2–5 in contact with anterior chin shields;           Detached rocks are scattered among the
anterior chin shields paired, longer (5.6 mm)        scenery, and there are no streams nearby.
than wide (1.6 mm), longer than posterior chin       Air parameter measurements conducted after
shields; posterior chin shields longer (3.4 mm)      the capture showed a temperature of 17.7 uC
than wide (2.5 mm), paired, in broad contact         and 87% relative humidity. In the vicinity of
medially; mental groove present; ventrals 186;       the type locality of S. perissostichon, we
cloacal scute single; subcaudals 108, divided        recorded the following species of amphibians
except for third and fourth pairs that are fused;    and reptiles: Craugastor podiciferus, Dia-
ventrals plus subcaudals 294; dorsal scales          sporus diastema, Pristimantis cruentus, P.
smooth, in 15–17–15 rows (anterior–mid-              ridens, Anolis capito, A. fortunensis, A.
body–hindbody), without apical pits or supra-        humilis, Ptychoglossus plicatus, Dendrophi-
cloacal ridges; dorsal scale rows changing           dion paucicarinatum, Geophis sp., Imantodes
orientation from the sides of body toward            cenchoa, Liophis epinephelus, S. annulatus,
middorsum; vertebral row not enlarged.               Trimetopon slevini, and Urotheca sp.
   Color in life.—Dorsal ground color Sayal
Brown (223C) with Sepia (119) blotches                                DISCUSSION
reaching onto venter, mostly continuous                Aside from the new species described
across venter, some ventrally offset; ventral        herein, four species of Sibon are known to
surface between dark blotches dirty white,                                        ¨
                                                     occur in western Panama (Kohler, 2008): S.
suffused with Buff (124) and mottled with            annulatus, S. argus, S. longifrenis, and S.
Sepia (119); dorsal surface of head Verona           nebulatus. During our fieldwork, we have
84                                                                                                                              HERPETOLOGICA                                                                                                                      [Vol. 66, No. 1


collected eight specimens of S. annulatus                                                                                                                3(b).    Enlarged penultimate supralabial not in
(including a specimen collected the same                                                                                                                          contact with eye; dark dorsal bands or
                                                                                                                                                                  spots reach well onto venter or even
night as the holotype of S. perissostichon just                                                                                                                   across venter                                   5
                                                                                                                                                                                                          _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _


260 m WSW of the type locality; see SMF                                                                                                                   4(a).   181–201 ventrals; 112–121 subcau-
numbers in Appendix). All eight specimens of                                                                                                                      dals                                 Sibon argus
                                                                                                                                                                              _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _




S. annulatus have a single postmental scale,                                                                                                             4(b).    151–173 ventrals; 82–106 subcau-
                                                                                                                                                                  dals                            Sibon longifrenis
whereas this species is said to have usually a                                                                                                                                _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _



                                                                                                                                                          5(a).   First infralabials in contact with each
pair of postmentals; the presence of a single                                                                                                                     other behind the mental          Sibon nebulatus                          _ _ _ _ _ _ _ _


postmental is considered typical for S. dimi-                                                                                                            5(b).    First infralabials separated from each
          ¨
diatus (Kohler, 2008; Savage, 2002). However,                                                                                                                     other by one or two postmentals behind
on the basis of characteristics of dorsal                                                                                                                         the mental                                      6
                                                                                                                                                                                                  _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _




coloration, we refer these specimens to S.                                                                                                                6(a).   Bands dark brown ventrally, their cen-
                                                                                                                                                                  ters paler brown, paler than interspaces;
annulatus as opposed to S. dimidiatus. They                                                                                                                       usually one postmental present                                                                _ _ _ _ _ _ _ _ _ _ _ _


all have more or less solid dark brown bands                                                                                                                                                      Sibon dimidiatus
                                                                                                                                                                  _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _



without paler centers, reaching well onto                                                                                                                6(b).    Bands dark brown ventrally and lateral-
venter. Our Panamanian specimens of S.                                                                                                                            ly, their centers not paler brown to
                                                                                                                                                                  cream laterally; lateral portion of bands
annulatus have 184–193 ventrals and 103–                                                                                                                          darker than interspaces; one or two
118 subcaudals in males (n 5 2), and 170–185                                                                                                                      postmentals present                             7           _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _



ventrals and 118–130 subcaudals in females (n                                                                                                             7(a).   Dorsal surface of head with green or
5 4). Two juveniles (SMF 88713–14) have                                                                                                                           yellow as well as red and black markings;
177–184 ventrals and 108–114 subcaudals.                                                                                                                          ground color of body green          Sibon lamari                                    _ _ _ _ _ _ _ _



                                                                                                                                                         7(b).    Dorsal surface of head with pale and
Thus, our specimens show slightly greater                                                                                                                         dark brown markings; ground color of
variation in the number of ventral scales than                                                                                                                    body usually olive to pale brown                                                                  _ _ _ _ _ _ _ _ _ _



has been documented for this species so far                                                                                                                                                        Sibon annulatus
                                                                                                                                                                  _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _




(McCranie, 2007). Within the Dipsadini
(sensu Cadle, 2007), S. perissostichon is                                                                                                                 Acknowledgments.—Collecting and exportation permits
unique in having 17 dorsal scale rows at                                                                                                               were provided by A. Salazar, Y. Hidalgo, and J. Garcıa, ´
midbody and a reduction to 15 on the                                                                                                                   Autoridad Nacional del Ambiente, Panama City, Panama.
posterior portion of body. Reduction of the                                                                                                            Q. D. Fuenmayora and V. Martinez, Panama City,
                                                                                                                                                       Panama, provided valuable assistance with acquisition of
number of dorsal rows on the posterior body                                                                                                            these permits. For field assistance, we thank L. Stadler
is unusual in the genus Sibon. Only S.                                                                                                                 and N. Hamad, as well as A. Bennett, who also provided
nebulatus is known to occasionally have                                                                                                                logistical support. For the loan of or access to specimens,
‘‘reductions from 15 to 13’’ (Peters, 1960:174).                                                                                                       we thank L. Ford, C. J. Raxworthy, D. R. Frost, American
                                                                                                                                                       Museum of Natural History (AMNH), New York; C. J.
                                                                                                                                                       McCarthy, The Natural History Museum (BMNH),
     KEY TO THE SPECIES OF SIBON FROM LOWER                                                                                                            London; W. E. Duellman and J. E. Simmons, University
                CENTRAL AMERICA                                                                                                                        of Kansas, Natural History Museum (KU), Lawrence; R.
                                                                                                                                                       W. McDiarmid and W. R. Heyer, National Museum of
                                                                                                                                                       Natural History (USNM), Washington, DC. This paper is
     1(a).   Seventeen rows of dorsal scales at mid-
             body and 15 rows one head length                                                                                                          based on work funded to AH by the FAZIT-STIFTUNG,
             anterior to cloaca; dorsal scale rows                                                                                                     and to SL by the Studienstiftung des deutschen Volkes.
             changing orientation from the sides of                                                                                                    An earlier version of this manuscript was read by J. R.
             body toward middorsum (Fig. 4a)                                                                       _ _ _ _ _ _ _ _
                                                                                                                                                       McCranie, L. D. Wilson, and an anonymous reviewer who
                                        Sibon perissostichon
             _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
                                                                                                                                                       all helped to improve the paper with their comments and
                                                                                                                                                       corrections.
     1(b).   Thirteen or 15 rows of dorsal scales at
             midbody without reduction on posterior
             portion of body; dorsal scale rows
             oriented longitudinally (Fig. 4b)             2                                                 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
                                                                                                                                                                                      LITERATURE CITED
     2(a).   Thirteen dorsal rows at midbody                                                                     _ _ _ _ _ _ _ _ _                     CADLE, J. E. 2007. The snake genus Sibynomorphus
                                            Sibon anthracops
             _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _                                                       (Colubridae: Dipsadinae: Dipsadini) in Peru and
     2(b).   Fifteen dorsal rows at midbody                3                                               _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _     Ecuador, with comments on the systematics of the
     3(a).   Enlarged penultimate supralabial in                                                                                                         Dipsadini. Bulletin of the Museum of Comparative
             contact with eye; dark dorsal bands or                                                                                                      Zoology 158:183–283.
             spots reach at most to outer edges of                                                                                                     DOWLING, H. G. 1951. A proposed standard system of
             ventrals, although dark shading may be                                                                                                      counting ventrals in snakes. British Journal of Herpe-
             present on venter                             4       _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _     tology 1:97–99.
March 2010]                                       HERPETOLOGICA                                                       85

KOHLER, G. 2008. Reptiles of Central America. Herpeton,
  ¨                                                           13.129 W, 1320 m: SMF 88715; Pianista trail, 8u 529 17.10
   Offenbach, Germany.                                        N, 82u 249 57.40 W, 1005 m: SMF 86411; Pianista trail, Casa
LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E.                   ´
                                                              de Ancon: SMF 88713–14; Veraguas: Cerro Mariposa,
   DAWSON. 1985. Standards in herpetology and ichthyol-       680 m: SMF 89596; Cerro Mariposa, 1000 m: SMF 89597.
   ogy: Part I. Standard symbolic codes for institutional     Sibon anthracops.—COSTA RICA: Guanacaste: Volcan
   resource collections in herpetology and ichthyology.       Rincon de la Vieja: SMF 77133; EL SALVADOR: No
   Copeia 1985:802–832.                                       additional locality data: SMF 68197; La Libertad: Santa
MCCRANIE, J. R. 2006. New species of Sibon (Squamta:          Tecla, Pinares de Suiza: SMF 81286; San Salvador: San
   Colubridae) from northeastern Honduras. Journal of         Salvador: SMF 79239; GUATEMALA: Zacapa: San Vi-
   Herpetology 40:16–21.                                      cente, Aldea Santa Elena: SMF 82594; NICARAGUA:
MCCRANIE, J. R. 2007. A second new species of Sibon                                                     ´
                                                              Carazo: Diriamba: KU 174287; Estelı: 15 km E Estelı,      ´
   (Squamta: Colubridae) from La Mosquitia, northeast-        800 m: KU 116975; Granada: Volcan Mombacho: SMF
   ern Honduras. Herpetologica 63:213–218.                    81024; Managua: 1 mi N Sabana Grande: KU 42299; 6 mi
PETERS, J. A. 1960. The snakes of the subfamily Dipsadinae.   WSW Sabana Grande: KU 42300; 8 km on South road: KU
   Miscellaneous Publications Museum of Zoology Univer-       174286; Casa Colorada, Las Nubes: KU 174282–85; Rivas:
   sity of Michigan 114:1–224.                                Morgan’s Rock, 11u 189 28.90 N, 85u 549 59.10 W, 35–265 m:
SAVAGE, J. M. 1973. A revised terminology for plates in the   JS field number 806, 838, SMF 88971–72. Sibon
   loreal region of snakes. British Journal of Herpetology    dimidiatus.—HONDURAS: Copa            ´n: Near Quebrada
   5:360–362.                                                 Grande, 1300 m: SMF 79119; NICARAGUA: Matagalpa:
SAVAGE, J. M. 2002. The Amphibians and Reptiles of Costa      Selva Negra, 12u 59.969 N, 85u 54.559 W: SMF 77800,
   Rica. A Herpetofauna between Two Continents,               77860, 79027–28; Matagalpa: BMNH 1894.7.26.45. Sibon
   between Two Seas. University of Chicago Press,             longifrenis.—COSTA RICA: Heredia: Puerto Viejo de
   Chicago, Illinois, USA.                                               ´
                                                              Sarapiquı, 50 m: SMF 79966; HONDURAS: Gracias a
SAVAGE, J. M., AND R. W. MCDIARMID. 1992. Rediscovery                                    ´       ´    ´
                                                              Dios: Reserva de Biosfera Rıo Platano: SMF: 86012;
   of the Central American colubrid snake, Sibon argus,
                                                              NICARAGUA: Jinotega: Bosawas, Kulum Kitang, 14u
   with comments on related species from the region.
                                                              19.89 N, 84u 56.29 W, 180 m: SMF 88182. Sibon
   Copeia 1992:421–432.
                                                              nebulatus.—COSTA RICA: Guanacaste: Volcan Arenal:
SMITHE, F. B. 1975–1981. Naturalist’s Color Guide.
                                                              SMF 77134; EL SALVADOR: Ahuachapa               ´n: Parque
   Part I. Color Guide. 182 Color Swatches. American
                                                              Nacional El Imposible, La Fincona, 720 m: SMF 81283;
   Museum of Natural History, New York, New York, USA.
                                                                                   ´
                                                              HONDURAS: Atlantida: La Ceiba: SMF 79879; NICARA-
SOLORZANO, A. 2002 (2001). Una nueva especie de
     ´
                    ´                                         GUA: Atla    ´ntico Norte: Wanı, Siuna: UCA 214; Big Falls,
                                                                                            ´
   serpiente del genero Sibon (Serpentes: Colubridae)
                                                                ´
                                                              Rıo Pis Pis: AMNH 12682; Rancho Alegre, 13u 399 47.50 N,
   de la vertiente del Caribe de Costa Rica. Revista de
           ´
   Biologıa Tropical 49:1111–1120.                            85u 019 38.90 W, 315 m: SMF 85510; Bosawas, Muru Lak,
                                                              14u 21.219 N, 84u 56.529 W, 185 m: SMF 87225; Parque
                                                              Nacional Saslaya, Cerro El Toro, 830 m: SMF 83122;
                .Accepted: 18 November 2009                       ´
                                                              Atlantico Sur: Kukra: AMNH 12680–81; betw. Kukra and
                .Associate Editor: Christopher Raxworthy                                                ´
                                                              Kanawa: AMNH 12679; Isla del Maız Grande: AMNH
                                                                                       ´
                                                              97068–70; Isla del Maız Grande, Mt. Pleasant: KU 174303;
                                                              Boaco: near Boaco: KU 174302; Granada: Volcan Mom- ´
                                                              bacho, near lower antenna, 11u 49.999 N, 85u 58.779 W,
                      APPENDIX I                              1100 m: SMF 79561, 79835; Managua: Casa Colorada, El
                                                              Crucero: KU 174295–99; Casa Colorada, Las Nubes: KU
            Comparative Material Examined                     174288–994, 174300; Nueva Segovia: 5 km N, 2.5 km E
                                          ´
  Sibon annulatus.—HONDURAS: Colon: Reserva de                                                 ´
                                                              Jalapa, 680 m: KU 112982; Rıo San Juan: Los Guatuzos,
  ´        ´     ´
Biosfera Rıo Platano, El Ocotillal, Cabeceras de Rıo    ´       ´    ´
                                                              Rıo Frıo, Fundeverde, 11u 049 37.00 N, 84u 449 55.70 W,
  ´
Platano, 370 m: SMF 86014; Gracias a Dios: Reserva de                                                       ´
                                                              45 m: SMF 87266; Los Guatuzos, along Rıo Papaturro,
  ´        ´     ´
Biosfera Rıo Platano, Pomokir, 150 m: SMF 86013;              11.0227u N, 85.0513u W, 40 m: JS field number 995; near
NICARAGUA: Atla    ´ntico Norte: Parque Nacional Saslaya,                                                          ´
                                                              El Castillo, finca of JOHN KINLACH: KU 174301; Rıo San
Campamento Los Monos, 13u 45.19 N; 85u 02.29 W, 850 m:        Juan, Boca de San Carlos, 10u 47.269 N, 84u 11.709 W, 20 m:
SMF 79655; Jinotega: Reserva Biosfera Bosawas, ca. 0.5 km                       ´
                                                              SMF 79836; Rıo San Juan, Bartola, 10u 58.379 N, 84u 20.359
SE Pueblo Wiso, 13u 59.609 N, 85u 19.609 W, 200 m: SMF        W, 30 m: CRB 011–13, SMF 79837; Camp Machado:
                               ´      ´         ´
78369; Matagalpa: Cerro Musun, Palan, Bilampı, 13u 009        USNM 19558; San Juan del Norte: USNM 5571, 62997;
41.00 N, 85u 149 11.60 W, 620 m: SMF 88180; PANAMA:                                  ´
                                                              PANAMA: Chiriquı: Pianista trail, Casa de Calisto: SMF
Bocas del Toro: Palo Seco, 8u 469 39.50 N, 82u 129 330 W,     88717; Los Algarrobos: Casa Culebra, 134 m: SL field
                                  ´
1030 m: SMF 85077–78; Chiriquı: near Lost and Found           number 165; Volante, 480 m: SMF 89599; Veraguas: Cerro
Ecohostel, Reserva Forestal La Fortuna, 8u 40.439 N, 82u      Mariposa, 933 m: SMF 89598.

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Sibon perissostichon sp. nov. pan köhler et al. 2010

  • 1. Herpetologica, 66(1), 2010, 80–85 E 2010 by The Herpetologists’ League, Inc. A NEW SPECIES OF SIBON (SQUAMATA: COLUBRIDAE) FROM WESTERN PANAMA GUNTHER KOHLER1,3, SEBASTIAN LOTZKAT1,2, ¨ AND ANDREAS HERTZ1,2 1 Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany 2 Johann Wolfgang Goethe-University, Institute for Ecology, Evolution and Diversity, BioCampus—Westend, Siesmayerstr. 70, 60323 Frankfurt am Main, Germany ´ ABSTRACT: We describe a new species of Sibon from the Reserva Forestal La Fortuna, Chiriquı Province, Panama. The new species is distinguished from all species presently assigned to Sibon by having 17 rows of dorsal scales at midbody and 15 rows one head length anterior to cloaca (vs. 13 or 15 rows of dorsal scales at midbody without reduction on posterior portion of body in all other species of Sibon), and dorsal scale rows that change orientation from the sides of body toward middorsum (vs. maintaining more or less the same orientation). A key to the Lower Central American species of Sibon is provided. RESUMEN: Describimos una nueva especie de Sibon procedente de la Reserva Forestal La Fortuna, ´ ´ Provincia de Chiriquı, Panama. La nueva especie se diferencia de todas las especies actualmente asignadas al ´ genero Sibon por tener 17 hileras de escamas dorsales en la mitad del cuerpo y 15 hileras una longitud de ´ ´ cabeza anterior a la cloaca (13 o 15 hileras de escamas dorsales en la mitad del cuerpo sin reduccion en la ´ ´ parte posterior del cuerpo en todas las demas especies de Sibon), ası como hileras de escamas dorsales que ´ cambian su orientacion desde los flancos del cuerpo hacia el centro del dorso (conservan su orientacion ´ ´ longitudinal en todas las demas especies de Sibon). Presentamos una clave para las especies de Sibon que ´ ocurren en el sur de Centroamerica. Key words: Colubridae; New species; Panama; Sibon; Squamata THE GENUS Sibon is a moderately diverse took additional data for the Lower Central group of small to moderate-sized snakes that are American species of Sibon from McCranie distributed from Nayarit and Veracruz, Mexico, (2006), Savage and McDiarmid (1992), and throughout Central America, and well into ´ Solorzano (2002). Abbreviations for museum South America to Ecuador and Brazil. Recently, collections follow those of Leviton et al. (1985) ´ McCranie (2006, 2007) and Solorzano (2002) ´ except for CRB (Coleccion de la Reserva described new species of Sibon, bringing the ´ Bartola, Rıo San Juan, Nicaragua). Specimens number of Central American species in this in the Appendix labeled with SL field genus to 11 (Ko ¨hler, 2008). Three fieldwork numbers will be deposited in the Universidad seasons in Panama (January 2006, January and ´ ´ ´ Autonoma de Chiriquı, Davıd, Panama, and February 2007, and May to August 2008) those labeled with JS field numbers will be produced several specimens of Sibon annulatus deposited in the Museo de Ciencias Naturales and S. nebulatus. Among the specimens collect- de la Universidad Centroamericana (UCA), ed was a single adult female of Sibon that differs Managua, Nicaragua. We follow Dowling strikingly from all its congeners in the number (1951) for the method of counting ventrals and arrangement of dorsal scale rows. Compar- and Savage (1973) for the terminology of isons with the known species of Sibon from scales in the loreal region of the head. The sex Central America demonstrated that this speci- of the holotype was determined by visual men represents an undescribed species. examination of relative tail length and basal girth. Head and scale measurements were MATERIALS AND METHODS made to the nearest 0.1 mm with dial calipers The format for the description of the new held under a dissecting microscope. Snout– species generally follows that of McCranie vent length and tail length measurements (2007). We provide a list of the specimens were taken to the nearest millimeter alongside examined for comparison in the Appendix. We a meter stick. Terminology for measurements is abbreviated as: snout–vent length, SVL; tail 3 CORRESPONDENCE: e-mail, gkoehler@senckenberg.de length, TL; head length, HL; snout length, 80
  • 2. March 2010] HERPETOLOGICA 81 FIG. 1.—Holotype of Sibon perissostichon (SMF 88716) in life, SVL 458.0 mm. SL; and head width, HW. Eye length equals the horizontal distance across eye. Scale dimensions were measured at the longest or widest points along the longitudinal or per- pendicular axis of the body, respectively. The capitalized colors and color codes (the latter in parentheses) are those of Smithe (1975–1981). Drawings of the head were made using a camera lucida attachment for a dissecting FIG. 2.—Head of holotype of Sibon perissostichon microscope. A small tissue sample of For- (SMF 88716). Scale bars 5 2.0 mm. schungsinstitut Senckenberg (SMF) 88716 was taken from the edge of a ventral scale rows one head length anterior to cloaca (vs. 13 about midbody and deposited in the tissue or 15 rows of dorsal scales at midbody without sample collection of the Senckenberg For- reduction on posterior portion of body in all schungsinstitut und Naturmuseum, Frankfurt, other species of Sibon), and dorsal scale rows Germany. that change orientation from the sides of body toward middorsum (vs. maintaining more or SPECIES ACCOUNT less the same orientation). Additionally, S. perissostichon differs from its congeners by Sibon perissostichon sp. nov. the following characteristics: subcaudals 108 (Figs. 1–4) in S. perissostichon vs. 113–126 in female S. Holotype.—SMF 88716, adult female as annulatus. Light interspaces between dark judged by the shape of the base of the tail, bands on body Sayal Brown (223C) in S. collected near Lost and Found Ecohostel, perissostichon vs. interspaces orange to red- Reserva Forestal La Fortuna, 8u 40.479 N, 82u orange middorsally, grading to cream laterally 12.979 W, 1434 m elevation, by Sebastian in S. anthracops. Enlarged penultimate su- Lotzkat on 14 May 2008. Original field pralabial not in contact with eye in S. number SL 145. perissostichon vs. this scale in contact with Diagnosis.—Sibon perissostichon can be eye in S. argus and S. longifrenis. Also, dark distinguished from its congeners by having dorsal bands or spots reach well onto venter or 17 rows of dorsal scales at midbody and 15 even across venter in S. perissostichon vs. dark
  • 3. 82 HERPETOLOGICA [Vol. 66, No. 1 FIG. 4.—Scalation (middorsal portion of body) of (a) Sibon perissostichon (SMF 88716) and (b) S. annulatus FIG. 3.—Holotype of Sibon perissostichon (SMF 88716) (SMF 88715). Scale bars 5 2.0 mm. at its capture site. no dark dorsal bands and venter immaculate dorsal bands or spots reach at most to outer in S. manzanaresi and S. sanniolus. Ventrals edges of ventrals, although dark shading may 186, subcaudals 108, bands solid dark brown, be present on venter in S. argus and S. darker than interspaces in S. perissostichon vs. longifrenis. One anterior temporal present in ventrals 168–171 in females, subcaudals 95– S. perissostichon vs. no such scale in S. carri. 100 in females, bands dark brown ventrally, Therefore, penultimate supralabial not in their centers paler brown, paler than inter- contact with parietal in S. perissostichon vs. spaces in S. miskitus. Postmental present; these scales in contact in S. carri. Bands solid therefore, first infralabials not in contact with dark brown, darker than interspaces in S. each other, vertebral row not enlarged in S. perissostichon vs. bands dark brown ventrally, perissostichon vs. first infralabials in contact their centers paler brown, paler than inter- with each other, no postmental, vertebral row spaces in S. dimidiatus. Dorsal surface of head moderately enlarged in S. nebulatus. with black markings, ground color of body Description of the holotype.—Adult female brown, postmental single in S. perissostichon (Figs. 1, 2); SVL 458 mm; TL 210 mm; HL vs. dorsal surface of head with green or yellow 15.0 mm; HW 9.0 mm; SL 4.2 mm. Head very as well as red and black markings, ground distinct from neck; snout short, blunt in dorsal color of body green, and postmental divided in and lateral outline; rostral wider (2.6 mm) than S. lamari. Ventrals 186, loreal in contact with high (2.0 mm), not extending posteriorly eye in S. perissostichon vs. ventrals 155, loreal between internasals, its length visible from separated from eye in only known female above one-quarter length of median internasal specimen of S. linearis. Dark dorsal bands suture (1.2 mm); internasals shorter (1.5 mm) present, reaching well onto venter or even than wide (2.0 mm), two-thirds length of across venter in S. perissostichon vs. usually prefrontal suture (2.6 mm); prefrontals longer
  • 4. March 2010] HERPETOLOGICA 83 (2.6 mm) than wide (2.2 mm), their median Brown (223B) with Jet Black (89) blotches; suture two-thirds length of frontal; prefrontals ventral surface of head dirty white with Raw bordering eye above loreal; frontal longer Umber (223) mottling and Jet Black (89) chin (4.1 mm) than wide (3.5 mm), widest anteri- shields and first ventral scale; iris Mars Brown orly, length two-thirds that of parietal, in (223A). Color in preservative (70% ethanol) contact with prefrontals, supraoculars, and similar to color in life. parietals; supraoculars longer (3.6 mm) than Etymology.—The name perissostichon is a wide (1.6 mm), nearly as long as frontal; noun in apposition and is derived from the parietals longer (5.9 mm) than wide (3.8 mm), Greek words perissos (beyond the regular widest anteriorly, their median suture (4.4 mm) number or size) and stichos (row, line), slightly longer than frontal; parietals in contact referring to the high number of dorsal scale with frontal, supraoculars, upper postoculars, rows in this species. anterior and posterior temporals, and eight Natural history notes.—The holotype was nuchal scales; nasal divided, in contact with encountered at 2053 h in secondary Premon- first two supralabials, loreal, prefrontal, inter- tane Forest while it was active within the nasal, and rostral; loreal single, longer (1.6 mm) dense, fog-moistened undergrowth about 0.5 m than high (0.7 mm), its length less than one- above the forest floor (Fig. 3). At the time of half length of snout (tip of snout to anterior encounter, the snake was moving about on the edge of eye) and one-half length of eye inclined trunk of a young tree covered with (3.3 mm); preoculars absent; postoculars three foliose hepatics. The undergrowth surround- (right) and two (left); temporals one plus three, ing this trunk consisted of herbaceous as well anterior one above fifth supralabial, lowest as young woody plants, among which we could posterior one above sixth supralabial; suprala- distinguish different ferns as well as members bials six, with third and fourth (right) or fourth of the families Rubiaceae and Melastomata- (left) bordering eye (enlarged penultimate ceae. The type locality is situated about 20 supralabial separated from eye by lower altitudinal meters below a narrow ridge on a postocular); pupil vertically elliptical; eye very steep slope (approximately 45u inclina- length two times length of loreal; infralabials tion) facing the southern Pacific lowlands. seven, first pair separated posterior to mental Most trees’ diameters at breast height do not by a single, triangular postmental, first infra- exceed 50 cm. The dense canopy layer extends labial in contact with postmental, infralabials from about 10 m aboveground upward. 2–5 in contact with anterior chin shields; Detached rocks are scattered among the anterior chin shields paired, longer (5.6 mm) scenery, and there are no streams nearby. than wide (1.6 mm), longer than posterior chin Air parameter measurements conducted after shields; posterior chin shields longer (3.4 mm) the capture showed a temperature of 17.7 uC than wide (2.5 mm), paired, in broad contact and 87% relative humidity. In the vicinity of medially; mental groove present; ventrals 186; the type locality of S. perissostichon, we cloacal scute single; subcaudals 108, divided recorded the following species of amphibians except for third and fourth pairs that are fused; and reptiles: Craugastor podiciferus, Dia- ventrals plus subcaudals 294; dorsal scales sporus diastema, Pristimantis cruentus, P. smooth, in 15–17–15 rows (anterior–mid- ridens, Anolis capito, A. fortunensis, A. body–hindbody), without apical pits or supra- humilis, Ptychoglossus plicatus, Dendrophi- cloacal ridges; dorsal scale rows changing dion paucicarinatum, Geophis sp., Imantodes orientation from the sides of body toward cenchoa, Liophis epinephelus, S. annulatus, middorsum; vertebral row not enlarged. Trimetopon slevini, and Urotheca sp. Color in life.—Dorsal ground color Sayal Brown (223C) with Sepia (119) blotches DISCUSSION reaching onto venter, mostly continuous Aside from the new species described across venter, some ventrally offset; ventral herein, four species of Sibon are known to surface between dark blotches dirty white, ¨ occur in western Panama (Kohler, 2008): S. suffused with Buff (124) and mottled with annulatus, S. argus, S. longifrenis, and S. Sepia (119); dorsal surface of head Verona nebulatus. During our fieldwork, we have
  • 5. 84 HERPETOLOGICA [Vol. 66, No. 1 collected eight specimens of S. annulatus 3(b). Enlarged penultimate supralabial not in (including a specimen collected the same contact with eye; dark dorsal bands or spots reach well onto venter or even night as the holotype of S. perissostichon just across venter 5 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 260 m WSW of the type locality; see SMF 4(a). 181–201 ventrals; 112–121 subcau- numbers in Appendix). All eight specimens of dals Sibon argus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ S. annulatus have a single postmental scale, 4(b). 151–173 ventrals; 82–106 subcau- dals Sibon longifrenis whereas this species is said to have usually a _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 5(a). First infralabials in contact with each pair of postmentals; the presence of a single other behind the mental Sibon nebulatus _ _ _ _ _ _ _ _ postmental is considered typical for S. dimi- 5(b). First infralabials separated from each ¨ diatus (Kohler, 2008; Savage, 2002). However, other by one or two postmentals behind on the basis of characteristics of dorsal the mental 6 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ coloration, we refer these specimens to S. 6(a). Bands dark brown ventrally, their cen- ters paler brown, paler than interspaces; annulatus as opposed to S. dimidiatus. They usually one postmental present _ _ _ _ _ _ _ _ _ _ _ _ all have more or less solid dark brown bands Sibon dimidiatus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ without paler centers, reaching well onto 6(b). Bands dark brown ventrally and lateral- venter. Our Panamanian specimens of S. ly, their centers not paler brown to cream laterally; lateral portion of bands annulatus have 184–193 ventrals and 103– darker than interspaces; one or two 118 subcaudals in males (n 5 2), and 170–185 postmentals present 7 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ ventrals and 118–130 subcaudals in females (n 7(a). Dorsal surface of head with green or 5 4). Two juveniles (SMF 88713–14) have yellow as well as red and black markings; 177–184 ventrals and 108–114 subcaudals. ground color of body green Sibon lamari _ _ _ _ _ _ _ _ 7(b). Dorsal surface of head with pale and Thus, our specimens show slightly greater dark brown markings; ground color of variation in the number of ventral scales than body usually olive to pale brown _ _ _ _ _ _ _ _ _ _ has been documented for this species so far Sibon annulatus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ (McCranie, 2007). Within the Dipsadini (sensu Cadle, 2007), S. perissostichon is Acknowledgments.—Collecting and exportation permits unique in having 17 dorsal scale rows at were provided by A. Salazar, Y. Hidalgo, and J. Garcıa, ´ midbody and a reduction to 15 on the Autoridad Nacional del Ambiente, Panama City, Panama. posterior portion of body. Reduction of the Q. D. Fuenmayora and V. Martinez, Panama City, Panama, provided valuable assistance with acquisition of number of dorsal rows on the posterior body these permits. For field assistance, we thank L. Stadler is unusual in the genus Sibon. Only S. and N. Hamad, as well as A. Bennett, who also provided nebulatus is known to occasionally have logistical support. For the loan of or access to specimens, ‘‘reductions from 15 to 13’’ (Peters, 1960:174). we thank L. Ford, C. J. Raxworthy, D. R. Frost, American Museum of Natural History (AMNH), New York; C. J. McCarthy, The Natural History Museum (BMNH), KEY TO THE SPECIES OF SIBON FROM LOWER London; W. E. Duellman and J. E. Simmons, University CENTRAL AMERICA of Kansas, Natural History Museum (KU), Lawrence; R. W. McDiarmid and W. R. Heyer, National Museum of Natural History (USNM), Washington, DC. This paper is 1(a). Seventeen rows of dorsal scales at mid- body and 15 rows one head length based on work funded to AH by the FAZIT-STIFTUNG, anterior to cloaca; dorsal scale rows and to SL by the Studienstiftung des deutschen Volkes. changing orientation from the sides of An earlier version of this manuscript was read by J. R. body toward middorsum (Fig. 4a) _ _ _ _ _ _ _ _ McCranie, L. D. Wilson, and an anonymous reviewer who Sibon perissostichon _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ all helped to improve the paper with their comments and corrections. 1(b). Thirteen or 15 rows of dorsal scales at midbody without reduction on posterior portion of body; dorsal scale rows oriented longitudinally (Fig. 4b) 2 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ LITERATURE CITED 2(a). Thirteen dorsal rows at midbody _ _ _ _ _ _ _ _ _ CADLE, J. E. 2007. The snake genus Sibynomorphus Sibon anthracops _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ (Colubridae: Dipsadinae: Dipsadini) in Peru and 2(b). Fifteen dorsal rows at midbody 3 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Ecuador, with comments on the systematics of the 3(a). Enlarged penultimate supralabial in Dipsadini. Bulletin of the Museum of Comparative contact with eye; dark dorsal bands or Zoology 158:183–283. spots reach at most to outer edges of DOWLING, H. G. 1951. A proposed standard system of ventrals, although dark shading may be counting ventrals in snakes. British Journal of Herpe- present on venter 4 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ tology 1:97–99.
  • 6. March 2010] HERPETOLOGICA 85 KOHLER, G. 2008. Reptiles of Central America. Herpeton, ¨ 13.129 W, 1320 m: SMF 88715; Pianista trail, 8u 529 17.10 Offenbach, Germany. N, 82u 249 57.40 W, 1005 m: SMF 86411; Pianista trail, Casa LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. ´ de Ancon: SMF 88713–14; Veraguas: Cerro Mariposa, DAWSON. 1985. Standards in herpetology and ichthyol- 680 m: SMF 89596; Cerro Mariposa, 1000 m: SMF 89597. ogy: Part I. Standard symbolic codes for institutional Sibon anthracops.—COSTA RICA: Guanacaste: Volcan resource collections in herpetology and ichthyology. Rincon de la Vieja: SMF 77133; EL SALVADOR: No Copeia 1985:802–832. additional locality data: SMF 68197; La Libertad: Santa MCCRANIE, J. R. 2006. New species of Sibon (Squamta: Tecla, Pinares de Suiza: SMF 81286; San Salvador: San Colubridae) from northeastern Honduras. Journal of Salvador: SMF 79239; GUATEMALA: Zacapa: San Vi- Herpetology 40:16–21. cente, Aldea Santa Elena: SMF 82594; NICARAGUA: MCCRANIE, J. R. 2007. A second new species of Sibon ´ Carazo: Diriamba: KU 174287; Estelı: 15 km E Estelı, ´ (Squamta: Colubridae) from La Mosquitia, northeast- 800 m: KU 116975; Granada: Volcan Mombacho: SMF ern Honduras. Herpetologica 63:213–218. 81024; Managua: 1 mi N Sabana Grande: KU 42299; 6 mi PETERS, J. A. 1960. The snakes of the subfamily Dipsadinae. WSW Sabana Grande: KU 42300; 8 km on South road: KU Miscellaneous Publications Museum of Zoology Univer- 174286; Casa Colorada, Las Nubes: KU 174282–85; Rivas: sity of Michigan 114:1–224. Morgan’s Rock, 11u 189 28.90 N, 85u 549 59.10 W, 35–265 m: SAVAGE, J. M. 1973. A revised terminology for plates in the JS field number 806, 838, SMF 88971–72. Sibon loreal region of snakes. British Journal of Herpetology dimidiatus.—HONDURAS: Copa ´n: Near Quebrada 5:360–362. Grande, 1300 m: SMF 79119; NICARAGUA: Matagalpa: SAVAGE, J. M. 2002. The Amphibians and Reptiles of Costa Selva Negra, 12u 59.969 N, 85u 54.559 W: SMF 77800, Rica. A Herpetofauna between Two Continents, 77860, 79027–28; Matagalpa: BMNH 1894.7.26.45. Sibon between Two Seas. University of Chicago Press, longifrenis.—COSTA RICA: Heredia: Puerto Viejo de Chicago, Illinois, USA. ´ Sarapiquı, 50 m: SMF 79966; HONDURAS: Gracias a SAVAGE, J. M., AND R. W. MCDIARMID. 1992. Rediscovery ´ ´ ´ Dios: Reserva de Biosfera Rıo Platano: SMF: 86012; of the Central American colubrid snake, Sibon argus, NICARAGUA: Jinotega: Bosawas, Kulum Kitang, 14u with comments on related species from the region. 19.89 N, 84u 56.29 W, 180 m: SMF 88182. Sibon Copeia 1992:421–432. nebulatus.—COSTA RICA: Guanacaste: Volcan Arenal: SMITHE, F. B. 1975–1981. Naturalist’s Color Guide. SMF 77134; EL SALVADOR: Ahuachapa ´n: Parque Part I. Color Guide. 182 Color Swatches. American Nacional El Imposible, La Fincona, 720 m: SMF 81283; Museum of Natural History, New York, New York, USA. ´ HONDURAS: Atlantida: La Ceiba: SMF 79879; NICARA- SOLORZANO, A. 2002 (2001). Una nueva especie de ´ ´ GUA: Atla ´ntico Norte: Wanı, Siuna: UCA 214; Big Falls, ´ serpiente del genero Sibon (Serpentes: Colubridae) ´ Rıo Pis Pis: AMNH 12682; Rancho Alegre, 13u 399 47.50 N, de la vertiente del Caribe de Costa Rica. Revista de ´ Biologıa Tropical 49:1111–1120. 85u 019 38.90 W, 315 m: SMF 85510; Bosawas, Muru Lak, 14u 21.219 N, 84u 56.529 W, 185 m: SMF 87225; Parque Nacional Saslaya, Cerro El Toro, 830 m: SMF 83122; .Accepted: 18 November 2009 ´ Atlantico Sur: Kukra: AMNH 12680–81; betw. Kukra and .Associate Editor: Christopher Raxworthy ´ Kanawa: AMNH 12679; Isla del Maız Grande: AMNH ´ 97068–70; Isla del Maız Grande, Mt. Pleasant: KU 174303; Boaco: near Boaco: KU 174302; Granada: Volcan Mom- ´ bacho, near lower antenna, 11u 49.999 N, 85u 58.779 W, APPENDIX I 1100 m: SMF 79561, 79835; Managua: Casa Colorada, El Crucero: KU 174295–99; Casa Colorada, Las Nubes: KU Comparative Material Examined 174288–994, 174300; Nueva Segovia: 5 km N, 2.5 km E ´ Sibon annulatus.—HONDURAS: Colon: Reserva de ´ Jalapa, 680 m: KU 112982; Rıo San Juan: Los Guatuzos, ´ ´ ´ Biosfera Rıo Platano, El Ocotillal, Cabeceras de Rıo ´ ´ ´ Rıo Frıo, Fundeverde, 11u 049 37.00 N, 84u 449 55.70 W, ´ Platano, 370 m: SMF 86014; Gracias a Dios: Reserva de ´ 45 m: SMF 87266; Los Guatuzos, along Rıo Papaturro, ´ ´ ´ Biosfera Rıo Platano, Pomokir, 150 m: SMF 86013; 11.0227u N, 85.0513u W, 40 m: JS field number 995; near NICARAGUA: Atla ´ntico Norte: Parque Nacional Saslaya, ´ El Castillo, finca of JOHN KINLACH: KU 174301; Rıo San Campamento Los Monos, 13u 45.19 N; 85u 02.29 W, 850 m: Juan, Boca de San Carlos, 10u 47.269 N, 84u 11.709 W, 20 m: SMF 79655; Jinotega: Reserva Biosfera Bosawas, ca. 0.5 km ´ SMF 79836; Rıo San Juan, Bartola, 10u 58.379 N, 84u 20.359 SE Pueblo Wiso, 13u 59.609 N, 85u 19.609 W, 200 m: SMF W, 30 m: CRB 011–13, SMF 79837; Camp Machado: ´ ´ ´ 78369; Matagalpa: Cerro Musun, Palan, Bilampı, 13u 009 USNM 19558; San Juan del Norte: USNM 5571, 62997; 41.00 N, 85u 149 11.60 W, 620 m: SMF 88180; PANAMA: ´ PANAMA: Chiriquı: Pianista trail, Casa de Calisto: SMF Bocas del Toro: Palo Seco, 8u 469 39.50 N, 82u 129 330 W, 88717; Los Algarrobos: Casa Culebra, 134 m: SL field ´ 1030 m: SMF 85077–78; Chiriquı: near Lost and Found number 165; Volante, 480 m: SMF 89599; Veraguas: Cerro Ecohostel, Reserva Forestal La Fortuna, 8u 40.439 N, 82u Mariposa, 933 m: SMF 89598.