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Coniophanes fissidens (GÜNTHER, 1858)

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Higher TaxaColubridae (Dipsadinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesConiophanes fissidens convergens SHANNON & SMITH 1949
Coniophanes fissidens dispersus SMITH 1941
Coniophanes fissidens fissidens (GÜNTHER 1858)
Coniophanes fissidens proterops (COPE 1860)
Coniophanes fissidens punctigularis (COPE 1860) 
Common NamesE: Yellowbelly Snake
S: Panza Amarilla
E: Eduardo’s forest snake [eduardoi]
S: Hojarasquera de Eduardo [eduardoi] 
SynonymCoronella fissidens GÜNTHER 1858: 36
Coniophanes fissidens — COPE 1860: 248
Tachymenis fissidens — GARMAN 1884: 62
Erythrolamprus fissidens — ANDERSSON 1901: 23
Coniophanes fissidens obsoletus MINTON et al. 1960
Coniophanes fissidens — VILLA et al. 1988
Coniophanes fissidens — PETERS & OREJAS-MIRANDA 1970: 67
Coniophanes fissidens — LINER 1994
Coniophanes fissidens — SAVAGE 2002
Coniophanes fissidens — MATA-SILVA et al. 2015
Coniophanes fissidens — WALLACH et al. 2014: 176
Rhadinaea eduardoi MATA-SILVA et al. 2019

Coniophanes fissidens convergens SHANNON & SMITH 1949
Coniophanes fissidens convergens — LINER 2007

Coniophanes fissidens dispersus SMITH 1941
Coniophanes fissidens dispersus SMITH 1941: 106
Coniophanes fissidens dispersus — LINER 2007

Coniophanes fissidens fissidens (GÜNTHER 1858)
Coronella fissidens GÜNTHER 1858: 36
Coniophanes fissidens fissidens — BAILEY 1939: 14
Coniophanes fissidens fissidens — LINER 2007

Coniophanes fissidens proterops (COPE 1860)
C[oniophanes] proterops COPE 1860: 249
Tachymenis proterops — GARMAN 1884: 62
Coniphanes fissidens proterops — TAYLOR 1949: 210
Coniophanes fissidens proterops — LINER 2007

Coniophanes fissidens punctigularis (COPE 1860)
Coniophanes punctigularis COPE 1860: 248
Dromicus chitalonensis MÜLLER 1878: 407
Coniophanes punctigularis — MÜLLER 1880
Coniophanes fissidens punctigularis — SMITH 1941: 107
Coniophanes fissidens punctigularis — MERTENS 1952: 60
Coniophanes fissidens punctigularis — LIVEZEY & PECKHAM 1953
Coniophanes fissidens punctigularis — PETERS & OREJAS-MIRANDA 1970: 67
Coniophanes fissidens punctigularis — LINER 2007 
DistributionMexico (Veracruz, Puebla, Chiapas, Quéretaro, San Luis Potosí, Tamaulipas, Hidalgo, Oaxaca), Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, NW Ecuador, Peru, Colombia [Castro,F. (pers. comm.)]; elevation (Honduras): 800-990 m

convergens: Mexico (Veracruz); Type locality: Mexico: 6 mi NW Tihuatlan, near Castillo de Teayo, Veracruz; W.L. Burger; April 20, 1949.

dispersus: Mexico (Guerrero, Michoacan, probably Nayarit to Oaxaca); Type locality: El Limoncito, Guerrero.

eduardoi (invalid): Mexico (Oaxaca); Type locality: Mexico, Oaxaca, municipality of Santa Catarina Juquila, El Obispo, 1,320 m (UTM 681141.99, 1789988.05 [= 16.183573, -97.305614, datum WGS 84]

fissidens: Type locality: “Mexico”.

obsoletus (invalid): Costa Rica; Type locality: Costa Rica: 1 mi E Volcan de Buenos Aires, Puntarenas; Sherman A. Minton, Jr.; July 22, 1957.

proterops: Mexico (Veracruz); Type locality: vicinity of Jalapa, Veracruz.

punctigularis: Mexico (Chiapas, isthmus of Tehuantepec), Guatemala, to Honduras; Type locality: Honduras.  
TypesSyntypes: BMNH 1946.1.9.61; BMNH 1946.1.3.2-3 (Coniophanes fissidens proterops - Two of the types of C. fissidens)
Holotype: ANSP 3742 [punctigularis]
Holotype: INHS (= UIMNH) 3821 [convergens]
Holotype: INHS (= UIMNH) 46573 [obsoletus]
Holotype: USNM 5285 [proterops]
Holotype: FMNH 100130 [dispersus]
Holotype. CH-CIB 5457 (given as CIB = Centro de Investigaciones Biológicas at the Universidad Autónoma del Estado de Hidalgo, Mexico, original field number VMS-2029), a subadult male, collected by Eduardo Mata-Silva on 6 June 2018 at 1800 hrs (Fig. 1) [eduardoi] 
DiagnosisDiagnosis (genus): Small to medium-sized xenodontine snakes; body moderate with proportionate tail; head distinct from neck. Dorsal scales smooth, without pits, in 17 to 25 rows reducing one to four times by the loss of the paravertebral rows (rarely the last reduction, third or fourth, takes place by loss of the third lateral rows). Anal and subcaudals divided. Head scales normal in number and arrangement; nasal divided or semidivided; 1 or 2 pre- and 2 postoculars; temporals 1-2; upper labials 7 or 8 (rarely 9). Pupil of eye round. Maxillary teeth 8 to 15, subequal, followed by a gap and 1 or 2 moderately enlarged grooved fangs; mandibular and palatine teeth subequal. The generalized color pattern consists of a dark middorsal stripe and a dark lateral stripe on either side on a grayish brown ground. Various modifications occur. A light stripe through the upper labials is present, and there is usually at least a faint trace of a light temporal stripe through the top of the eye. (Hemipenes described subsequently) (from Bailey 1939: 4).

Diagnosis (fissidens): Smith 1941: 104

This diagnosis works only for Central American species, not for South American ones (Ricardo Palacios, pers. comm., 26 March 2020).

Coniophanes is closely allied to Rhadinaea, differing chiefly in having the hind maxillary teeth grooved (Dunn 1944: 495).

COLOR AND PATTERN: The body is orangish brown to brown, weakly and somewhat variably striped or lined with dark brown or black. Usually there is a vertebral dark line or row of dots (one on each scale apex in the vertebral row) and, in addition, the median three or seven scale rows may be darker brown than the ground color on each side. Most Panamanian specimens that I have seen have a dark lateral stripe that lies principally on rows 4 and 5 and the lower edge of row 6, posteriorly dropping one row lower. The bottom side of this stripe tends to be ill defined, and in one individual the entire lower sides are darkened, as is frequent in specimens from Costa Rica. In part, this variation may be due to ontogenetic changes, as suggested by C. B. Fisher (in litt.). The upper margin of the lateral stripe is usually distinct because of a greater accumulation of dark pigment, and some individuals have only this dark line as a remnant of the broader stripe, which possibly is being lost in some populations. The upper margin of the lateral stripe, or line, usually is further emphasized by a series of small, white or pale tan dashes lying immediately above. These dashes are most distinct on the posterior part of the body, usually showing no traces elsewhere except on the neck, on which there is a short, white line several scales in length. Also on the neck is a pair of tiny, black-bordered white ocelli. Each ocellus is no larger than one or two scales and is situated between the posterior temporal plates and the anterior end of the short, white line on the sixth scale row. The top of the head is brown, usually with an inconspicuous pair of black dots, one on each side of the anterior part of the interparietal suture. In some cases there is a fine. white line or series of dots extending from the upper margin of the eye a short distance back toward the white ocellus on the side of the nape. The upper edges of the anterior supralabials, from the snout to the eye, may be black-bordered or not, but all available specimens have a black postocular line that extends obliquely down from the eye and across the corner of the mouth to the lower side of the neck. Adjacent to the lower edge of the postocular black line there may be an enamel-white line that, on a few specimens, skirts the lower edge of the eye and continues uninterrupted to the snout. Below the white line the supralabials are grayish white and usually intensely speckled with black. The white lip marking may be partly an attribute of age, as it is obvious on six adults but reduced or absent from four juveniles, in which the black specking is likewise reduced or absent. The under side of the head is speckled with black in most specimens. There is a black dot near the tip of each ventral plate, and the number of black specks and little spots varies over the rest of the light venter. Although the ventral spotting is largely irregular, there often is a tendency for some markings to form two rows, each about a fourth to a third of the width of a ventral from the edge of the belly. (Myers 1969)

COLOR IN LIFE: Slevin (1942) stated that two specimens from Boquete were light brown, with a pinkish tinge, in life. At least some individuals from central Panama have the body strongly tinged with orange. The white ocelli on the nape and, when present, the white labial line and the dorsal, white postocular line (or row of dots), are enamel-white and very conspicuous in a hand-held specimen, although on the forest floor these markings possibly serve as disruptive (hence concealing) coloration. The under side of the head was white in four specimens for which I recorded data, but W. E. Duellman (field notes for K.U. No. 75672) noted one in which the chin was pale orange. Two Canal Zone specimens and one from El Valle had a pale to medium orange suffusion on the sides of the belly, with a reduced median part of the ventrals and most of the subcaudals light yellow. Another individual from El Valle had a yellowish white venter. The iris color of specimens from central Panama has been noted as reddish orange (two), reddish brown (one), and rich brown (one). The tongue is brown or reddish brown, with the distal end of the stalk and the forks black. (Myers 1969)

SIZE AND SCUTELLATION: Coniophanes fissidens is a moderately robust snake. The largest measurable Panamanian specimen is a female 466 mm. in total length (preserved), of which the tail is 120 mm. The tail varies from 33.1 to 41.3 per cent of the total length in three males (including two juveniles) and from 25.8 to 38.5 per cent in six females (including two juveniles). Any possible ontogenetic variability in the tail length proportion is not obvious, because the extremes in the series of females are for two adults. The dorsal scales are smooth, except for anal ridges that are present on some females as well as males, and usually are in 21-21-17 rows, although there may be reduction to 19 by midbody and to 15 immediately in front of the cloaca. Ventrals range from 110 to 121 (mean 114.5) in four males, and 116 to 124 (mean 119.2) in six females. Subcaudals are 63 to 70 (mean 66.3) in three males, and 57 to 66 (mean 61.3) in six females. There is one, or in some cases two, preoculars and two postoculars. The general temporal formula is 1 + 2, although there may be some asymmetry or other disarrangement of the basic pattern. Supralabials are eight per side, with the second and third touching the loreal and the fourth and fifth in contact with the eye. Infralabials are 10, or in some cases nine, with the first five (or first four if there are only nine plates) touching the anterior genials and the fifth to sixth ( or fourth to fifth) touching the posterior genials. (Myers 1969)

Diagnosis (convergens). A race of Coniophanes fissidens related to C. f. proterops and similar to C. f. dispersus; apparently: dorsal scale rows 19; males with supraanal ridges; supralabials 7; ventral 121, caudal 80, ventrals-minuscaudals 41 in a single male; median border of dorsolateral light stripe well defined on tail, poorly defined on rear of body; dorsolateral light stripe disappearing about two head lengths posterior to head; spots on belly very small, scattered. (Shannon & Smith 1949)

Description of type (convergens). Portion of rostral visible from above 'one half length of median internasal suture; prefrontals much larger than internasals, their median suture 2 2/3 length of that of internasals; frontal a little less than 1½ times as long as broad, its length 5/4 its distance from tip of snout, which in turn is about equal to its distance from posterior (median) edge of parietal; sides of frontal somewhat convergent posteriorly; nasal completely divided; loreal small, slightly smaller than lower postoculars; 1-1 preoculars; 2-2 postoculars; temporals 1-2, the upper secondary twice as long as lower secondary; supralabials 7-7. Anterior chinshields ¾ length of and a little wider than posterior chinshields; latter separated from each other except for their anterior third; infralabials 9-9, the first ones in contact medially between chinshields and mental, the anterior five in contact with chinshields (two with the posterior, four with the anterior); head scales with a sprinkling of fine pustules absent on body. Dorsal scales in 19-19-15 rows; ventrals 121 (male); caudals 80; anal divided; total length 474 mm., tail 157 mm. Dorsal pattern very poorly defined, the whole surface nearly uniform, dull, light brown; sides slightly darker than back, with a sharply defined dark line marking the boundary between the two zones along the lower edge of the fifth (anteriorly) and upper edge of the fourth (posteriorly) scale rows; dorsum slightly lighter adjacent to this dark line than elsewhere, but no dorsolateral light stripes evident except (1) on nape, where it is whitish, well defined, occupying less than one scale row anteriorly at its point of margin at angle of jaws, and nearly all of two scale rows posteriorly at about 3 head lengths back of the head where it disappears completely, and (2) on extreme posterior part of body and on tail, where the stripe, little lighter than on body, has a well defined, black median edge. A narrow white line along sides of head extending from lower margin of orbit through the angle of the jaws, and bordered above by a dark line merging above with the ground color; one or two large, light bordered dark spots on each of the anterior 5 supralabials; latter otherwise stippled on a light (whitish) background; chin and throat rather closely stippled; numerous small black flecks on belly and tail, somewhat more numerous laterally. (Shannon & Smith 1949)

Diagnosis (dispersus): “Scales in 19 rows; males with supraanal tubercles; SU-
pralabials 8; ventrals 120 and 122 in two males; caudals 81 in one male; ventrals minus caudals 41 in one male; spots on belly small, scattered, not forming regular series; middle and posterior part of belly may be completely unspotted (except ends of ventrals) ; white dorsolateral stripes on nape short, diffuse; inner border of dorsolateral light stripe poorly defined on tail, not evident in front of anus; no spots or irregularities of pattern in dorsal area between lateral stripes; latter poorly defined, diffuse.” (Smith 1941: 106)

Description of holotype (dispersus): Smith 1941: 106

Diagnosis (eduardoi). A snake of the genus Rhadinaea that can be distinguished from all congeners by the following combination of morphological features: supralabials 7, with 3rd and 4th entering orbit; 120 ventrals; 71 subcaudals; one subpreocular (lower preocular); 17 dorsal scales throughout body; a head pattern lacking postorbital pale markings but having a pale line extending from the lower rear quadrant of the eye to the ultimate supralabial and slightly beyond, and a midbody dorsal color pattern of a lateral series of black dots in the lower apex of the scales of row V and a middorsal line confined to the middorsal scale row consisting of a series of disjunct spots on the posterior apex of otherwise dark brown scales.

Diagnosis (proterops): Size rather small. Scales in nineteen longitudinal rows, thin, elongate, obtuse. Head scarcely distinct, short profile of muzzle not elevated. Anterior plates of the head small ; loreal a little longer than high. One pre-, two postoculars. Superior labials seven, third and fourth entering the orbit. Vertical plate elongate, lateral borders convergent, posterior angle acute. Occipitals long. Inferior labials nine ; geneials two pairs, nearly equal. Gastrosteges 130, anal one, divided, urosteges ? (tail badly mutilated.) Head and body 9 in. 7 lin. in length. The stump of the tail appears tetragonal in section. (COPE 1860)

Coloration (proterops) Above light brown, every scale densely punctulated with darker, especially near the margins. From the first to the fourth row of scales this is deeper, giving the sides a darker shade. The vertebral row of scales, from the occipitals to the end of the tail is also darker. Top of the head densely and obscurely vermiculated and punctulated. The dark shade on the fourth row of scales becomes a band anteriorly, and is bordered above and below with white on the neck. The lower white border is continued to the eye, and is bordered above on the labials with black. The upper white border is discontinued on the neck, but reappears as a spot, three scales back of the occipitals. Inferior half of rostral, upper and lower labials, chin, throat and belly, light brownish yellow, densely punctulated with brown. Each labial with a darker spot in the centre. Fewer punctulations on the urosteges. (COPE 1860)

Diagnosis (proterops): Smith 1941: 105

Diagnosis (punctigularis): Diagnosis.-Scales in 21 rows; males with supraanal tubercles; supralabials 8, rarely 7; ventrals 119 to 130 in females, 116 to 125 in males; caudals 71 to 85 in females, 80 to 91 in males; ventrals minus caudals 39 to 54 in females, 31 to 38 in males; spots on belly very small, scattered; belly sometimes unspotted (except ends of ventrals, dark as sides of body); median border of dorsolateral light stripes very distinct on posterior part of body as well as on tail; dorsolateral light stripes distinct on much of anterior part of body; a series of spots on each side of middorsal line, about halfway between lateral and middorsal stripe; spots fused with dorsolateral stripe in all except young specimens, but always distinct laterally; body not orange in young. (Smith 1941: 107)

Description of holotype (punctigularis): Smith 1941: 107 
CommentRelative abundance in Honduras: infrequent

Distribution: this species has been reported from Peru, but without specific locality (T. Doan, pers. comm. 30 Apr 2012). Not in Yucatan state (Mexico), fide Gonzalez-Sanchez et al. 2017. Not in Nayarit fide WOOLRICH-PIÑA et al. 2016 (who only report C. lateritius).

Type species: Coronella fissidens GÜNTHER 1858 is the type species of the genus Coniophanes HALLOWELL in COPE 1860: 248.

Synonymy: Bailey 1937 pointed out that the type of proterops is “a specimen of fissidens” without formally synonymizing the two. Palacios-Aguilar & García-Vázquez 2020 synonymized Rhadinaea eduardoi MATA-SILVA et al. 2019 with C. fissidens. Kaiser et al. 2013 rejected the subgeneric names Cottonserpens Hoser 2012, Daraninserpens Hoser 2012, Laidlawserpens Hoser 2012, Smythserpens Hoser 2012 invalid and rejected their use instead of Coniophanes.

Key: Myers 1969 has a key to the “southern” species of Coniophanes (bipunctatus, fissifens, piceivittis, joanae, and dromiciformis) 
EtymologyThe generic name is derived from the Greek words konio, meaning "dust" and phano, meaning "appearance." The specific name is derived from the Latin words fissus, a suffix denoting a division into two parts, and dens, meaning "tooth" (Lemos-Espinal & Dixon 2013).

Rhadinaea eduardoi was named in honor of Eduardo Mata-Silva, collector of the holotype. Eduardo is the younger brother of the senior author of the paper, is a resident of Río Grande, Oaxaca, and was a highly valued member of the field crew working in Oaxaca. He also outshined the rest of the crew when it came to finding snakes, as evidenced by his discovery of the holotype of the snake. 
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