Two new species of snail-eating snakes of the genus Tropidodipsas (Serpentes, Dipsadidae) from southern Mexico, with notes on related species Author Gruenwald, Christoph I. https://orcid.org/0000-0001-5994-555X Biencom Real Estate, Carretera Chapala-Jocotepec # 57 - 1, C. P. 45920, Ajijic, Jalisco, Mexico. & Biodiversa A. C., Avenida de la Ribera # 203, C. P. 45900, Chapala, Jalisco, Mexico. & Herp. mx A. C. C. P. 28989, Villa de Alvarez, Colima, Mexico. cgruenwald@switaki.com Author Toribio-Jimenez, Sarahi Centro de Ciencias de Desarrollo Regional, Universidad Autonoma de Guerrero. Calle Pino s / n. Col. El Roble. C. P. 39640. Acapulco, Guerrero, Mexico. Author Montano-Ruvalcaba, Carlos Biodiversa A. C., Avenida de la Ribera # 203, C. P. 45900, Chapala, Jalisco, Mexico. & Herp. mx A. C. C. P. 28989, Villa de Alvarez, Colima, Mexico. Author Franz-Chavez, Hector Biodiversa A. C., Avenida de la Ribera # 203, C. P. 45900, Chapala, Jalisco, Mexico. & Herp. mx A. C. C. P. 28989, Villa de Alvarez, Colima, Mexico. Author Penaloza-Montano, Miguel A. Facultad de Ciencias Quimico Biologicas, Universidad Autonoma de Guerrero. Avenida Lazaro Cardenas # 88, Ciudad Universitaria Sur, C. P. 39074, Chilpancingo de los Bravo, Guerrero, Mexico. Author Barrera-Nava, Eduardo Y. Facultad de Ciencias Quimico Biologicas, Universidad Autonoma de Guerrero. Avenida Lazaro Cardenas # 88, Ciudad Universitaria Sur, C. P. 39074, Chilpancingo de los Bravo, Guerrero, Mexico. Author Jones, Jason M. Biodiversa A. C., Avenida de la Ribera # 203, C. P. 45900, Chapala, Jalisco, Mexico. & Herp. mx A. C. C. P. 28989, Villa de Alvarez, Colima, Mexico. Author Rodriguez, Christopher M. Los Angeles Zoo and Botanical Gardens, 45333 Zoo Drive, Los Angeles, P. C. 90027, California, USA. Author Hughes, India M. Department of Biology, University of South Alabama, 5871 USA Dr. N., Mobile, P. C. 36688, Alabama, USA. Author Strickland, Jason L. https://orcid.org/0000-0002-1927-7259 Department of Biology, University of South Alabama, 5871 USA Dr. N., Mobile, P. C. 36688, Alabama, USA. Author Reyes-Velasco, Jacobo https://orcid.org/0000-0002-8787-9980 Biodiversa A. C., Avenida de la Ribera # 203, C. P. 45900, Chapala, Jalisco, Mexico. & Herp. mx A. C. C. P. 28989, Villa de Alvarez, Colima, Mexico. text Herpetozoa 2021 2021-11-11 34 233 257 http://dx.doi.org/10.3897/herpetozoa.34.e69176 journal article http://dx.doi.org/10.3897/herpetozoa.34.e69176 2682-955X-34-233 FFCB4A62CCBA5446923C8F77B4D3A4DF Tropidodipsas papavericola sp. nov. Figs 4 , 5 Paratypes/ (5) (Fig. 5 ) INIRENA 2802 (original field number CIG 1496, Fig. 5C ). Sub-adult male, collected AOR 18.1 km S of Puerto del Gallo on road from Nuevo Dehli to Puerto del Gallo, Municipio de Atoyac de Alvarez (17.4342°, -100.191°, datum=WGS84, 1654 m a.s.l.), Guerrero, Mexico on June 29th, 2019 by Christoph I. Gruenwald , Miguel A. Penaloza-Montano , Eduardo Y. Barrera-Nava, Karen I. Morales-Flores and Janelle Morales-Flores. INIRENA 2803 (original field number CIG 1502). Sub-adult male collected DOR at 5 km S of La Laguna, on road from San Luis La Loma to Bajitos de la Laguna, Municipio de Tecpan de Galeana (17.5137°, -100.7737°, datum=WGS84, 1686 m a.s.l.), Guerrero, Mexico on June 29th, 2019 by Jason M. Jones and Carlos Montano-Ruvalcaba . INIRENA 2804 (original field number CIG 1632, Figs 5A-B ). Adult male, collected at Bajitos de la Laguna, Municipio de Tecpan de Galeana, Guerrero, Mexico on October 30th, 2019 and retrieved by Alejandro Lara and donated to Jason M. Jones. INIRENA 2805 (original field number CIG 1457). Sub-adult male, collected at Jaguar Research Facility, Municipio de Tecpan de Galeana, Guerrero, Mexico on June 10th, 2019 by Miguel A. Penaloza-Montano and Eduardo Y. Barrera-Nava. INIRENA 2810 (JRV 0362, Fig. 2D ). Juvenile male, collected DOR at 4.2 km S of La Laguna, on San Luis San Pedro - La Laguna Rd., Municipio de Tecpan de Galeana (17.5173°, -100.7715°, datum=WGS84, 1620 m a.s.l.), Guerrero, Mexico on July 17th, 2020 by Jason M. Jones. Diagnosis. Tropidodipsas papavericola sp. nov. is placed in the genus Tropidodipsas based on phylogenetic evidence (Fig. 6 ). It belongs to the Tropidodipsas fasciata group as defined by Kofron (1987) based on possessing a laterally compressed body, head distinctly wider than neck, protruding eyes of moderate to large diameter, vertebral and paravertebral scales not wider than other dorsal scales, 17 maxillary teeth, 22 dentary teeth, postmental scale absent. The species differs from all described species of Tropidodipsas recognized herein based on the following combination of characters: (1) 15/15/15 rows of smooth dorsal scales with no enlarged vertebral row; (2) prefrontal broader than long, not entering orbit; (3) loreal almost square, slightly longer than broad, not entering orbit; (4) 179-189 ventral scales in males, unknown in females; (5) 69-76 divided subcaudals in males, unknown in females; (6) 25-34 pale bands on body and 10-16 pale bands on tail; (7) 26-36 dark bands on body and 10-16 on tail, irregular with faded pale centers in adult individuals; (8) TL/SVL 0.30-0.35 in males, unknown in females; (9) iris dark chocolate-gray in life. Comparisons. Tropidodipsas papavericola sp. nov. is most similar to T. philippii , T. fasciata , T. guerreroensis and T. tricolor sp. nov. It is the only species of Tropidodipsas in Mexico with a nebulated pattern of obscured body rings, although the nearly sympatric Sibon nebulatus exhibits a similar pattern as do several species of snail-suckers in northwestern South America ( Arteaga et al. 2018 ). It is distinguished from other Mexican snail-suckers such as the Geophis chalybeus species group, the Geophis omiltemanus species group, G. sartorii new comb., G. annuliferus new comb., Tropidodipsas fischeri (Fig. 9C,D ), S. carri , G. sanniolus new comb. and S. linearis by the laterally compressed body shape and and the nebulated pattern of broken up dorsal bands on a gray or brown background. It differs from S. anthracops (character states in parenthesis) by possessing 15 scale rows at midbody (vs. 13), possessing a nebulated dorsal color pattern (vs. tricolor banded pattern), incomplete banding on venter (vs. complete), and lacking a loreal that enters the orbit (vs. loreal entering orbit). It may be distinguished from S. dimidiatus by lacking a post-mental scale (usually one or two post-mental scales present), and possessing less than 80 subcaudals in males (vs. more than 95). Distinct from the superficially similar Sibon nebulatus (Fig. 9A, B ) by vertebral row of dorsal scales same size as other dorsal scales (vs. vertebral row of dorsal scales 1.25-1.35 times larger than other dorsal scales), smaller adult body size of 450-560 mm in males (vs. up to 890 mm in males), fewer dark body bands, 26-35 on body (vs. 36-47 on body), head predominately dark brown or black (vs. head pattern speckled with dark and light marking) and absence of pink speckling in the dorsal pattern (vs. pink speckling present). It differs from Dipsas gaigeae by having 15 dorsal scale rows at midbody (vs. 13), possessing 179-189 ventral scales (vs. 155-169), loreal that does not enter orbit (vs. loreal enters orbit), and 26-35 pale body bands (vs. 7-12). Distinct from Dipsas brevifacies by possessing 179-189 ventral scakes (vs. 162-180), prefrontal and loreal not in contact with orbit (vs. variable in both), usually possessing 1+2 temporals (vs. usually 2+3), usually possessing one pair of infralabials in contact after the mental (vs. usually two pairs of infralabials in contact after the mental). Within Tropidodipsas , T. papavericola sp. nov. differs from T. fasciata and T. guerreroensis by possessing 15 smooth dorsal scale rows (vs. 17 keeled scale rows). Tropidodipsas papavericola sp. nov. differs from T. tricolor sp. nov. by lacking the tricolor outline in the dorsal bands (vs. possessing a tricolor outline in dorsal bands), 26-35 dorsal dark bands on body (vs. 19-22), by possessing a prefrontal which does not enter orbit (vs. prefrontal entering orbit), 2 preoculars (vs. 1), 5-7 supralabials (vs. 7-8), 6-7 infralabials (vs. 8-9), less subcaudal scales, 69-76 in males (vs. 78-79 in males), by possessing a pale ventral coloration of head with irregularly scattered dark spots, and a black and white mental (vs. pale ventral coloration of head with heavy dark stippling anteriorly and black mental), and by a smaller eye - head length ratio of 17-21% (vs. 25%). It is most similar to T. philippii (Fig. 8C, D ), from which it can be distinguished by having smooth dorsal scales (vs. slightly keeled), a longer tail which is 30-35% of SVL in males (vs. 25-29% in males), and less subcaudal scales, 69-76 in males (vs. 76-90 in males). Genetic divergence in a 1,072-bp long fragment of the mitochondrial cytb gene between T. papavericola sp. nov. and T. philippii is 12-14%; between T. papavericola sp. nov. and T. guerreroensis , 10-13%; between T. papavericola sp. nov. and T. fasciata is 10-12%. Description of holotype. (Fig. 4 ) INIRENA 2801. Adult male. SVL 421 mm; TL 137 mm; TotL 558 mm. HL 13.8 mm (from tip to posterior border of parietal) and 13.5 mm (from snout to posterior commissure of mouth); HW 9.8 mm, head distinct from body, approximately 2 times wider than neck. Snout long, SL 5.7 mm, contained 2.4 times in HL, rounded in dorsal and lateral profile. Loreal region tall, canthus rostralis angular. Rostral 1.2 times as broad as high (3.3 mm wide, 2.7 mm high); internasals broader than long (1.7 mm length, 2.8 mm width), rounded anteriorly, in lateral contact with anterior and posterior nasals; prefrontals longer than broad, (3.8 mm length, 2.9 mm width), rectangular, in median contact with each other and in lateral contact with postnasal, loreal, upper preocular and supraocular; frontal 1.1 times longer than broad, (4.4 mm long, 3.9 mm broad), pentagonal with angular tip posteriorly, in contact with prefrontals, supraoculars and parietals. Two moderately large preoculars on each side, upper larger than lower. Single supraocular large, in contact with upper preocular, prefrontal, frontal, parietal, and upper postocular. Two moderately large postoculars on each side, upper one larger than lower. Parietal 1.4 times as long as wide (6.0 mm long, 4.2 mm wide), the length of parietal 61% of HL, the common suture between parietals 4.4 mm, equal to frontal length. Nasal divided, prenasal slightly taller than postnasal, with combined length of both nasals 2.6 mm, longer than loreal. Loreal small, slightly longer than high and almost square, 1.4 high, 1.7 mm long on left side, 1.6 mm high, 1.8 mm long on right side, not reaching orbit. Eye moderately large, 3.1 mm, 23% of HL. Supralabials 7 on both sides, first in contact with nasals, second and third in contact with loreal, fourth and fifth entering orbit, sixth largest and in contact with lower postocular, anterior and posterior temporals. Temporals 1 + 2. Five nuchal scales in contact with parietals. Table 3. Meristic variation of Tropidodipsas tricolor sp. nov. and Tropidodipsas papavericola sp. nov.

Species	T. tricolor sp. nov.		T. papavericola sp. nov.
Specimen	INIRENA 2800	INIRENA 2798	INIRENA 2801	INIRENA 2802	INIRENA 2803	INIRENA 2804	INIRENA 2810	INIRENA 2805
Sex	Male	Female	Male	Male	Male	Male	Male	Male
Snout-vent Length	210	310	421	366	363	430	258	336
Tail Length	67	93	137	119	113	127	91	112
Total Length	277	403	558	485	476	557	349	448
Head length	8.30	13.62	18.97	15.25	15.83	18.58	13.03	14.92
Anterior Chinshield	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide
Posterior Chinshields	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide	Longer than wide
Gulars	3	3	4	4	5	4	5	5
Infras in contact with 1st CS	5	5	5	4	4	5	4	4
Ventral Scales	183	183	180	180	188	189	182	179
Subcaudal Scales	79 / 79	79 / 79	76	74 left / 74	73 / 73	69	74	73
Pre-oculars (Side)	1. 1	1. 1*	2. 2	2. 2	2. 2	2. 2	2.2	2. 2
Post-oculars (Side)	2. 2	3. 3	2.2	2. 2	2. 2	1. 1	2. 2	2. 2
Anterior Temporals (Side)	1. 1	1. 1	1. 1	1. 1	1. 1	1. 1	1. 1	1. 1
Posterior Temporals (Side)	2. 2	2. 2	2. 2	2. 2	2. 2	2. 2	2. 2	2. 2
Supralabials (Side)	8. 7	7. 8	7. 7	6. 6	6. 7	5. 6	7. 7	6. 6
Infralabials (Side)	8. 8	8. 9	7. 6	6. 6	6. 6	6. 6	7. 7	6. 6
Post-mentals	0	0	0	0	0	0	0	0
Pale Body Bands (Side)	19. 19	19. 20	-	32. 28	34. 35	29. 27	27. 27	25. 25
Pale Tail Bands (Side)	9. 9	8. 8	-	14. 12	-	11. 11	15. 16	13. 13
Black Body Blotches (Side)	19. 19	19. 21	32. 33	30. 31	35. 35	27. 27	29. 28	26. 26
Black Tail Blotches (Side)	8. 8	8. 8	14. 14	15. 15	-	10. 11	16. 14	13. 13
Total Pale Body Blotch Count (Body & Tail)	28. 28	28. 28	46. 47	45. 46	42. 42	37. 38	45. 42	39. 39
TL/TotL	0.24	0.23	0.25	0.25	0.24	0.23	0.26	0.25
TL/SVL	0.32	0.30	0.33	0.33	0.31	0.30	0.35	0.33
Internasal Length	1.00	2.34	1.70	1.34	1.50	1.46	0.90	1.44
Internasal Width	1.60	2.62	4.48	3.53	3.58	4.74	2.96	3.52
Loreal Length	0.9 / 1.0	1.19 / 1.10	1.73 / 1.70	1.46 / 1.38	1.36 / 1.34	1.26 / 1.22	1.0 / 1.0	1.40 / 1.38
Loreal Width	1.0 / 1.0	1.24 / 1.12	1.84 / 1.82	1.45 / 1.2	1.38 / 1.42	1.40 / 1.46	0.92 / 0.94	1.30 / 1.26
Loreal into orbit	No	No	No	No	No	No	No	No
Pre-frontal Length	1.70	2.34	3.39	2.81	2.89	3.26	2.48	2.36
Pre-frontal Width	2.00	2.64	5.75	4.69	4.69	5.06	3.84	4.33
Prefrontal into orbit	Yes	Yes	No	No	No	No	No	No
Frontal Length	2.80	2.88	4.23	3.58	3.71	3.93	3.31	3.60
Frontal Width	2.40	2.70	3.81	2.73	3.35	3.35	2.71	3.08
Parietal Length	4.30	5.54	6.58	6.82	5.51	6.26	4.61	5.28
Length Anterior Chinshields	2.4 / 2.6	3.49 / 3.63	4.73	4.03	4.31	5.72	3.33	3.58
Length Posterior Chinshields	1.9 / 1.8	2.28 / 2.42	3.72	1.94	2.25	2.67	1.81	2.15
Eye Diameter	2.10	2.80	3.20	3.02	3.11	3.56	2.71	2.85
ED/HL	0.25	0.21	0.17	0.20	0.20	0.19	0.21	0.19
Pre-Frontal Length/Width	0.85	0.89	0.59	0.60	0.62	0.64	0.65	0.55
HL/SVL	0.04	0.04	0.05	0.04	0.04	0.04	0.05	0.04
Midbody scale rows	15	15	15	15	15	15	15	15
Head scale rows	15	15	15	15	15	15	15	15
Caudal scale rows	15	15	15	15	15	15	15	15
Lateral compresion	Yes - Extreme	Yes - Extreme	Yes - Oval	Yes - Oval	Yes - Oval	Yes - Oval	Yes - Oval	Yes - Oval
Anal plate	Undivided	Undivided	Undivided	Undivided	Undivided	Undivided	Undivided	Undivided
Penultimate supralabial enlarged	Yes	Yes	Yes	Yes	Yes	Fused	Yes	Yes
First infraliabals in contact	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes
Pale nuchal band	Yes	Yes	Yes	Yes	Yes	Yes	Yes	Yes

Mental 2.4 times as broad as long (2.6 mm broad, 1.1 mm long), flat anteriorly, triangular posteriorly and separated from the anterior chinshields by the first pair of enlarged infralabials which are in contact with each other. Infralabials 7/ 8; on the left 1-5 in contact with anterior chinshields and fifth in contact with posterior chinshield; on the right 1-6 in contact with anterior chinshields and sixth in contact with posterior chinshield. Anterior chinshields irregular, both 2.3 times as long as wide (left 4.6 mm long and 2.0 mm wide, right 4.4 mm long and 1.9 mm wide). Left posterior chinshield 1.6 times as long as wide (3.5 mm long, 2.0 mm wide), and right posterior chinshield 1.3 times as long as wide (2.6 mm long, 2.0 mm wide). Four gular scales. Infralabials and scales in the chin region smooth. Dorsal scales in 15-15-15 rows, smooth throughout body; apical pits not evident. Ventrals 180; anal plate single; 76 paired subcaudal scales. Body laterally compressed. Pupil elliptical. Figure 4. Tropidodipsas papavericola sp. nov. Holotype (INIRENA 2801). Holotype (INIRENA 2801) in life (A); Ventral aspect of Holotype (INIRENA 2801) in preservative (B,E); Dorsal aspect of Holotype (INIRENA 2801) in preservative. Figure 5. Tropidodipsas papavericola sp. nov. Variation. Male (INIRENA 2804) from Municipio de Tecpan de Galeana, Guerrero (A-B); Male (INIRENA 2802) from Municipio de Leonardo Bravo, Guerrero (B); Juvenile male (INIRENA 2810) from Municipio de Tecpan de Galeana, Guerrero (D). Coloration in life of the holotype (Fig. 4A ). Dorsal coloration of head, body and tail brown with 32/33 irregular dark brown bands on the body and neck which all have pale centers that are stippled with paler brown and pale cream. Pale brown nuchal band two body scale rows long but reduced to one at the mid-dorsal line; not involving the posterior part of the parietals. Dark bands on body and tail 3-7 scale rows long, widest mid-dorsally, and alternating on posterior half of body. Head coloration predominately dark brown on dorsal surface and with light brown mottling and pale cream on rostral and anterior labials. First three supralabials bicolor pale cream and dark brown, posterior two supralabials bicolor dark brown with pale brown. Ventral coloration of head pale cream with irregular dark brown blotches. Ventral coloration of body pale yellowish cream, with dark and pale bands incomplete on venter. The ventral pattern consists of longitudinal rows of dark spots that connect to form irregular longitudinal lines. Anal plate bicolor, dark brown and pale cream. All 15 dark bands on tail incomplete ventrally, dark coloration forming longitudinal lines ventrally on tail. Tail tip pale cream with black tip. Pupil black, iris dark chocolate-gray. Coloration in preservative of the holotype (Figs 4B-E ). General coloration bicolor. Dorsal surfaces of head, body and tail predominately dark brown with darker brown-black bands. Pale bands brown-gray, but whitish in a few places where stratum corneum has been removed (snake was going into shed when preserved). The dark bands remain brown-black and the pale cream faded to white. Variation. Meristic variation of five available specimens given in Table 3 . Variation in color and pattern detailed below (Fig. 5 ). Color in life. A sub-adult male paratype from above Nueva Dehli, Municipio de Atoyac de Alvarez , Guerrero (INIRENA 2802) had a pale brown body coloration with 31/29 dark brown bands on body and neck posterior to pale nuchal band and 14 on the tail. This specimen has light centers in all dark body bands, albeit limited to the lateral portions only. This implies that the irregular dark banding with light centers may be subject to ontogenetic variation, as this specimen is intermediate in size and pattern between the juvenile INIRENA 2810 and the large adult holotype INIRENA 2801. Neither the pale body bands, nor the dark body bands, are complete ventrally, reaching only to the lateral edges of the ventrals. The venter was cream or pale tan mid-ventrally with two dark brown irregular longitudinal stripes running down the length of the entire venter. The lines run into the remnants of the dark brown dorsal bands on the edges of the venter (Fig. 5C ). Color in preservative. An adult male paratype from above Bajitos de la Laguna, Municipio de Tecpan de Galeana, Guerrero (INIRENA 2804) has 29/27 dark bands on the body posterior to the pale nape band and 12 on the tail. This specimen differs from the holotype because it has a relatively well-defined pattern of dark and pale bands with regular edges. Furthermore, the first well-defined dark band on the neck does not have a pale center, and on the second dark body band the pale center is interrupted mid-dorsally. The venter is white and covered with irregular black spots that do not form longitudinal rows of spots nor are fused into lines and dashes (Fig. 5A, B ). A juvenile male paratype from above San Luis La Loma, Municipio de Tecpan de Galeana, Guerrero (INIRENA 2810) has 28/29 dark bands on the body and neck posterior to the pale nuchal band and 16 on the tail. This specimen lacks any light centers in the dark body bands of the tail, suggesting that this color pattern may be subject to an ontogenetic shift from juveniles to adults. It has black stippling in the light interspaces reminiscent of T. tricolor sp. nov. Both the pale body bands and the dark body bands are not complete ventrally, reaching only onto the lateral edges of the ventrals. The venter (in preservative) is white mid-ventrally, with two irregular longitudinal stripes running down the length of the entire venter. The lines run into the remnants of the dark dorsal bands on the edges of the venter (Fig. 5D ). An adult male paratype from above San Luis La Loma, Municipio de Tecpan de Galeana, Guerrero (INIRENA 2803) has 35 dark bands on body and neck posterior to the pale nuchal band. None of the dark bands have pale centers mid-dorsally, some have the pale centers on the lateral portions of the body and others lack them altogether. Ventral coloration white with black blotches forming an irregular "checkerboard" pattern. No pale or dark body bands complete on venter. A sub-adult male paratype, from above Tecpan de Galeana, Guerrero (INIRENA 2805) has 26 dark bands on body and neck posterior to the pale nuchal band, all with faded pale centers in the dark body bands, most of them extensive but incomplete mid-dorsally. It has 12 dark bands on the tail and a dark tail tip. Ventral coloration white with dark gray mottling, with no dark or pale dorsal bands complete on venter, but the dense mottling makes the dark dorsal bands on posterior portion of body connect in an alternating undulated pattern. Dentition. The holotype (INIRENA 2801) appears to have 14 maxillary teeth visible, however we did not remove the maxillary arch to avoid damaging the specimen. We observed in situ the maxillary arch and dentition of the holotype (INIRENA 2801): it has 14 maxillary teeth and 20-22 dentary teeth. An adult male paratype (INIRENA 2805) has 17 maxillary teeth (counting empty sockets) on the right side. We counted 22 dentary teeth on the right dentary of this same specimen. Distribution, habitat and ecology. This species appears to be restricted to moderate elevations in the Sierra Madre del Sur in central Guerrero in two distinct areas (Fig. 10 ). Habitats include cloud forest, mesic pine-oak woodland, tropical evergreen forest, tropical semi-deciduous forest and combinations of these assemblages. Known from around 1600-2200 m a.s.l. in the Sierra de Tecpan de Galeana, Sierra de Atoyac and above Acapulco, in the foothills in the area encompassed between Acahuizotla and El Treinta. Specimens were found at night during the rainy season on low to medium (1.5-3.0 m) vegetation or crossing the road. Etymology. The specific epithet papaver + cola refers to the living among poppy plants (genus Papaver ) which are illegally planted throughout the range of this species for the extraction of opium gum.