4 research outputs found
Micrurus tikuna Feitosa, Jr, Pires, Zaher & Prudente, 2015, sp. nov.
Micrurus tikuna sp. nov. Figs. 1–2. Micrurus ornatissimus — Silva Haad 1994: 82. Micrurus ornatissimus — Campbell & Lamar 2004; Plate 159. Holotype. Adult female, MPEG 18199 (formerly Hospital Regional de Leticia-HRL 10101), collected by Juan Silva Haad, on 1991 at INCRA neighborhood, municipality of Tabatinga (04º 14 ’ 36 ’’S, 69 º 54 ’ 15 ’’W; ca. 80 m above sea level, hereafter asl), state of Amazonas, Brazil (Figs. 1–2). Paratype. Adult male, ICN 10853, collected by Maria Cristina Ardila, (field number MC 8531), on November 23 1996 at Km 9.5 from Leticia-Tarapacá road, Quebrada La Arenosa (02º 57 ’ 58 ’’S, 69 º 48 ’ 58 ’’ W, ca. 110 m asl), municipality of Leticia, department of Amazonas, Colombia. Diagnosis. Micrurus tikuna can be distinguished from all congeners by the combination of the following characters: (a) white scales or a white transversal band posterior to parietal scales present; (b) dorsum of head black, forming a conspicuous cephalic-cap; (c) black nuchal collar absent; (d) ventral part of the head black with irregular white blotches followed by red scales on the gular region; (e) ventral scales 205 in the single male and 225 in the single female; (f) subcaudal scales 47 in the male and 38 in the female; (g) 27 to 31 black body rings in both sex (with 3 to 4 dorsal scales long) bordered by narrow, black tipped white rings (with 1 / 2 to one dorsal scale long), alternated by 27 to 31 wider red body rings (4 to 6 dorsal scales long); (h) tricolor tail with six to seven black rings (four to six dorsal scales long) bordered by narrow irregular white rings (1 / 2 dorsal scale long) alternated by six to seven red rings of the same length as the black rings; (i) lateral view of head with black coloration covering first four supralabials and postoculars and the last three supralabials, temporal and occipital regions red (Figs. 1 – 2). Comparison with other congeners from western Amazon (Figs. 3–7, Table 1). Twenty-four species of Micrurus are known to occur in western Amazon. Micrurus filiformis (Günther, 1859), M. hemprichii ortoni Schmidt, 1953, M. lemniscatus helleri Schmidt & Schmidt, 1925, M. spixii spixii Wagler, 1824, M. spixii obscurus Jan, 1872, and M. surinamensis (Cuvier, 1817) are readily distinguishable from M. tikuna by their triadal pattern (vs. monadal pattern) (Table 1). Comparisons between Micrurus tikuna and the other 18 monadal species (including the two of the three currently recognized subspecies M. annellatus which present geographical distribution simpatric with M. tikuna) are given below with meristic characters summarized in Table 1 (character states for M. tikuna are shown in parenthesis). M. albicinctus Amaral, 1925, M. a. annelatus (Peters, 1871), M. margaritiferus Roze, 1967, M. medemi Roze, 1967 and M. putumayensis Lancini, 1962 (Fig. 3) differ from M. tikuna by having bicolor dorsal aspect on the body and tail (vs. tricolor dorsal aspect on the body and tail). Micrurus steindachneri steindachneri (Werner, 1901), M. s. orcesi Roze, 1967 and M. petersi Roze, 1967 differ from M. tikuna by having cephalic-cap surrounded by red scales, yellowish rings on the body, and heavily melanic dorsum (vs. cephalic-cap surrounded by white scales; white rings on the body and tail; and light colored dorsum). Micrurus mertensi Schmidt, 1936 and M. peruvianus Schmidt, 1936 differ from M. tikuna by having bicolor tail, black nuchal collar, and white rings with two scales long (vs. tricolor tail; nuchal collar absent; and white rings with one scale long) (Figs. 3 J, 5 B). Micrurus annellatus bolivianus Roze, 1967 differs from M. tikuna by having 15–24 black rings on the body, prefrontal scales completely black, white rings on body up to two scales long, and white transversal cephalic band (vs. 27–31 black body rings; white-spotted prefrontal scales; narrow white body rings 1 / 2 scales long; and absence of transversal white cephalic band) (Fig. 5 C). Micrurus averyi Schmidt, 1939 differs from M. tikuna by having head completely black, 8–12 black rings on the body, and bicolored tail (vs. black and white head; 27–31 body black rings; and tricolor tail) (Fig. 5 D). Micrurus catamayensis Roze, 1989 differs from M. tikuna by having black accessory rings isolating white rings from red rings on the body (vs. white rings contacting red rings) (Fig. 5 E). Micrurus langsdorffi (Wagler, 1824) differs from M. tikuna by having a black nuchal collar, yellow body rings, and red body rings up to four scales long (vs. absence of nuchal collar; white body rings; and red rings more than four scales long) (Fig. 5 F). Micrurus ornatissimus (Jan, 1858) differs from M. tikuna by having black rings 2–3 and red rings 3–6 scales long on the body and tail, 32–66 black rings on the body, 3–15 black rings on the tail, and black nuchal collar (vs. black and red body rings with 4–6 and 4–8 scales long, respectively; 27–31 black rings, 6–7 black tail rings; and absence of black nuchal collar) (Fig. 5 G). Micrurus pacaraimae Carvalho, 2002 differs from M. tikuna by having a black nuchal collar, snout region stained with white, and red rings 8–12 scales long (vs. nuchal collar absent; snout entirely black; red rings 4–6 scales long) (Fig. 5 H). Micrurus paraensis Cunha & Nascimento, 1973 differs from M. tikuna by having a black nuchal ring, prefrontal scales completely black, and first white ring on the cervical region (vs. absence of black nuchal ring; white-spotted prefrontal scales; red cervical region, with first white ring far caudally placed beyond neck) (Fig. 5 I). Micrurus remotus Roze, 1987 differs from M. tikuna by having a mostly bicolor tail, black nuchal collar, entirely black prefrontal scales, and the region between cephaliccap and nuchal collar (vs. tricolor tail; absence of black nuchal collar; white spotted prefrontal scales; and region between cephalic-cap and nuchal collar uniformly red) (Fig. 5 J). Description of the holotype (Figs. 1–2). Adult female, 633 mm (SVL) and 66.5 mm (TL), tail length 9.5% of SVL; head length 17.11 mm, distinct from body, corresponding to 2.8% SVL; body slightly wider than high. Rostral shield visible from above (width 4.5 mm and length 2.3 mm); internasals wider than long; prefrontals wider than long, in contact with nasals, internasals, supraoculars, preoculars, and frontal; left prefrontal contacting both internasals; frontal pentagonal, nearly as long as wide, 1.5 times longer than its greatest width; parietal longer than wide, corresponding to 18.9% of head length; supraoculars longer than wide; nasal shield divided above nostril and below in contact with first two supralabials, internasals, prefrontals, and preocular; preocular in contact with supraocular, prefrontal, nasal, and third supralabial; two postoculars, upper postocular longer than lower; temporals 1 + 2; supralabial seven, with third and fourth bordering the orbit; infralabial seven, first pair in broad contact behind symphysial; first four infralabials contacting chinshields; posterior chinshields longer and wider than the anterior chinshields. Dorsal scales rows 15 / 15 / 15, smooth, without apical pits; three preventrals, ventrals 225, cloacal plate divided, and paired subcaudals 38. Color Scales Coloration of the holotype in preservative (Figs. 1–2). Dorsum of body with 31 black rings (three to four scales long) bordered by narrow black tipped white rings (four to six dorsal scales), alternated by 31 wider red rings (four to five scales long); tricolor tail with six black rings bordered by narrow irregular white rings, alternated by six red rings slightly narrower than black rings (Fig. 1 A); ventral surface of body and tail with immaculate red rings (four to five ventral scales long) (Fig. 1 B); head black dorsally (from rostral to first dorsal scale), forming a conspicuous black cephalic-cap; prefrontal scales with white spots posteriorly; white scales bordering posterior margin of black cephalic-cap; region between cephalic-cap and first white body ring red (Fig. 2 A); laterally, black coloration entirely covers first three supralabials, dorsal portion of fourth supralabial, and postocular scales; ventral portion of fourth to seventh supralabial red; temporal and occipital regions, towards the first white body ring, red with scales mostly black tipped (Fig. 2 C). Symphysial and first four infralabial scales black with irregular white markings; first pair of chin shields white with irregular black markings; second pair black with white spots on the anterior one-fourth. Gular region red mostly scattered with black spots (Fig. 2 B). Color of the holotype in life (Fig. 8). The description of color in life was based on drawings and photographs from the holotype provided by Silva Haad (1994). Head with black cephalic-cap covering parietals, frontal, prefrontals, internasals, rostral, preoculars, postoculars and from first to fourth supralabial scales, may reach up to the first dorsal scales. Some specimens may have a narrow white strip separating this region's red hooded head post parietal. Prefrontal scales with white mark. Black cap separated from first white body ring by four to six dorsal scales. Laterally, head black from first to third supralabials and postocular, with black coloration including upper portion of fourth supralabial; head red from fourth supralabial until first white body ring, with most scales black tipped. In some specimens this red region is interrupted ventrally with prolonged white coloration on the gular region; anterior region of chinshields white, irregularly marked with black (including infralabial scales); posterior chinshields region mostly black; in some specimens red ring incomplete ventrally by invasion of white pigment on gular and first ventral scales; black body rings (three to four dorsal scales in length) followed by narrow white rings (0.5 scale in length) with black tipped scales and wider red rings (four to six dorsal scales in length); tricolor tail with black rings (four to six dorsal scales in length) followed by narrow white rings (0.5 scale in length), and red rings with same width or narrower than black rings. Ventrally red rings on the tail are incomplete. Description of the paratype (Fig. 9). Adult male, 540 mm (SVL) and 94.5 mm (TL), tail length is about 14.8% of SVL; head length 16.64 mm, distinct from body, corresponding to 2.62% of SVL; frontal pentagonal, nearly as long as wide, 1.4 times longer than its greatest width, parietal longer than wide, corresponding to 21.3 % of the head length. Preventrals three, ventrals 205, and paired subcaudals 47. The other body and cephalic characters were invariable with respect to the holotype. Hemipenis morphology (ICN 10853). Both hemipenes were everted in situ. Fully everted but partially expanded right hemipenis is slender, deeply bilobed and non-capitate. Sulcus spermaticus deep, bifurcated at base of lobes and running centripetally along lobes and reaching apexes; long lobes (about 40 % of total hemipenial length), tapering distally into pointed tips; distal half of lobes ornamented by small to medium-sized spines densely arranged in irregular longitudinal rows, decreasing in size toward apex of lobes; proximal half of lobes less densely ornamented by dispersed larger and slender spines; lobular spines thinner and more numerous on the asulcate side of the organ; distal half of hemipenial body ornamented by small, irregularly arranged, spines, gradually increasing in size proximodistally; proximal region of hemipenis naked, except for presence of large basal pocket, extending along most of lateral surface of hemipenial body; asulcate surface of hemipenial body with spines restricted near lobes bifurcation. Etymology. The specific epithet “ tikuna ” is a name in apposition derived from Tupi indigenous word “tacouma,” meaning a men with their face or nose painted with black (Gregório 1980). Tikuna also refers to the native Indian nation that originally occupied the western Amazon region along the upper Solimões River near the boundaries between Brazil, Colombia and Peru. Similar to the Tikuna Indians, the new species of Micrurus has a head predominantly black. Geographic distribution (Fig. 10). Micrurus tikuna is only known from Leticia, department of Amazonas, Colombia and Tabatinga, state of Amazonas, Brazil.Published as part of Feitosa, Darlan Tavares, Jr, Nelson Jorge Da Silva, Pires, Matheus Godoy, Zaher, Hussam & Prudente, Ana Lúcia Da Costa, 2015, A new species of monadal coral snake of the genus Micrurus (Serpentes, Elapidae) from western Amazon, pp. 538-554 in Zootaxa 3974 (4) on pages 539-548, DOI: 10.11646/zootaxa.3974.4.5, http://zenodo.org/record/23433
Micrurus paraensis
<i>Micrurus paraensis</i> Cunha & Nascimento (1973) <p> <i>Micrurus psiches [psyches] paraensis</i> Cunha & Nascimento, 1973: 276. Type-locality: Icoaraci, Belém, Pará, Brasil.</p> <p> <i>Micrurus donosoi</i> Hoge, Cordeiro & Romano, 1978: 71. Type-locality: São Felix do Xingu, Pará, Brasil.</p> <p> <i>Micrurus psyches paraensis</i> - Cunha & Nascimento, 1978: 164; Roze, 1982: 334; Campbell & Lamar, 1989: 140.</p> <p> <i>Micrurus paraensis</i> - Hoge & Romano, 1981: 400; Cunha & Nascimento, 1982: 18; Nascimento <i>et al</i>. 1987: 57.</p> <p> <i>Micrurus psyches donosoi -</i> Roze, 1982: 334; Campbell & Lamar, 1989: 140.</p> <p> <i>Micrurus paraensis debruini</i> Abuys, 1987: 215. Type-locality: Kwamalasamutu, Suriname.</p> <p> <i>Micrurus paraensis</i> - Cunha & Nascimento, 1993: 116; Roze, 1994: 178; Roze, 1996: 207; Carvalho, 2002: 186; Campbell & Lamar, 2004: 185; Frota, 2004: 13.</p> <p> <b>Diagnosis.</b> <i>Micrurus paraensis</i> can be distinguished from other species of <i>Micrurus</i> by the following combination of characters: black cap beginning on rostral, passing from the first to fourth supralabials, behind the eyes on postoculars and half of first temporal; white rings beginning on 5, 6 and 7th supralabials, sometimes beginning on half of fourth supralabial, through temporals towards parietals; black cap joined to first black body rings (nuchal collar) to end of parietals; black body rings range from 12 to 21, bordered by a short white rings (comprising one, half scale, with black posterior margin), 12–20 red rings range from, about 4 to 6 times longer than black rings; body rings do not form triads; 2–11 black tail rings alternated by 1–11 white rings; black tail rings with white spots on dorsal and ventral surfaces; and males have no supraanal tubercles.</p> <p> <i>Micrurus paraensis</i> can be distinguished from the eastern Amazonian coralsnakes (<i>M. filiformis, M. hemprichii hemprichii, M. lemniscatus lemniscatus, M. spixii, M. surinamensis</i>) by the monadal color patterns, long tail, long and bifurcated hemipenis (<i>vs.</i> triad color patterns, short tail, short and bilobed hemipenis). In other Amazonian regions, <i>M. paraensis</i> can be distinguished from monadal color patterned species (<i>M albicinctus</i> and <i>M. langsdorffi</i>) by the tricolor pattern and lower number of black body rings. Bicolored patterns with higher number of black body rings (67 to 90 in males; 77 to 91 in females) on <i>M. albicinctus</i>, white spots on frontals and supraoculars, and white gular region with black spots. <i>M. langsdorffi</i> has a tricolor monadal patterns, higher number of black, white and red body rings (18 to 38 black, 38 to 57 white and 19 to 38 red in males; 29 to 47 black, 46 to 60 white and 20 to 46 red in females) with short red rings (3 to 4 dorsals and ventrals long).</p> <p> Finally, it can be distinguished from the <i>M. psyches</i> group (<i>M. circinalis, M. mademi, M. psyches</i> and <i>M. remotus</i>) by the black body rings and the number of ventrals and subcaudals. <i>M. psyches</i> has black body rings ranging from 22 to 29 in males and 27 to 41 in females (if the black and the original red rings that are blackish are counted as black, the males have 44 to 57 and the females have 51 to 81), original red and black rings are 3 to 5 dorsals and ventrals long, and ventrals ranges from 188 to 196 in males and 203 to 212 in females. <i>M. circinalis</i> has black body rings ranging from 21 to 31, with short red rings (4 to 7 dorsals and ventrals long), and ventrals ranges from 174 to 193 in males and 192 to 209 in females. <i>M. medemi</i> has black body rings ranging from 15 to 25 (if the black and the original red rings that are blackish are counted as black, has 35 black body rings), and ventrals ranges from 193 to 198 in males and 211 to 218 in females. <i>M. remotus</i> has black body rings ranging from 25 to 29 in males and ranging from 29 to 40 in females, and ventrals ranges from 202 to 203 in males and 214 to 225 in females.</p> <p> <b>Description of Holotype.</b> MPEG 0851; adult male; SVL 390 mm; TAL 85 mm (17.8 % of TL); and HL 12.5 mm (2.6 % of TL). Head indistinct from the body. Eyes small, pupil round. Body cylindrical. Snout rounded. Tail long. Dorsal scales smooth, in 15 rows, without reduction, no apical pits; 192 ventrals; 52 subcaudals; anal divided; 7 supralabials, 3 and 4th in contact with the orbit; 7 infralabials, 1 to 4th in contact with the first pair of genials; nasal half-divided; one preocular; two postoculars, the superior is higher; 1+1 temporal. White lateral ring beginning on parietals from inferior region of 4th supralabial; 5, 6 and 7th supralabials white, the 6th is higher and the posterior edge of 7th supralabial is black; little anterior part of temporal is black, the remainder of the anterior temporal and the posterior temporal is white; parietal totally black joining the black cap to the nuchal collar, this is interrupted by white spots on the right side of ventral surface; 1 to 3th infralabials black with white spots; 4 to 6th infralabials white, the 4th one is higher; the first pair of mentals is black with white spots, the second pair is white with black top. 15 black body rings (2 to 3 dorsals and ventrals long); 14 red body rings (7 to 14 dorsals long with black top); 9 black tail rings, the second black ring has a white spot on the ventral surface; 4th and 7th black rings with a spot on the ventral-lateral surface, alternating with by 8 short white rings; a ventral black spot between the 5 and 6th black body rings and two between 7th and 8th black body rings (Table 1).</p> <p> <b>Description.</b> Rostral scale broader than high, with a triangular shape in frontal view and visible from above. Internasals shorter than prefrontals, which do not enter the orbit. Frontal longer than broad. Supraoculars aproximately as broad as the prefrontal. Parietals about two times longer than wide. Nasal half-divided. One preocular, longer than high. Two posoculars, the superior is higher. One anterior and one posterior temporal. Thirth and fourth supralabial in contact with the orbit. First pair of infralabials in contacts behind symphysial, and 1st to 4th pairs contact first pair of genials. Fourth infralabials in contact with second pair of genials. Two pairs of genials (Figure 1). Dorsals scales in 15 rows, smooth, without reduction. Anal plate divided.</p> <p> <b>Color Pattern</b> - Black cap beginning on rostral, passing from 1st to 4th supralabials, behind the eyes on postoculars and half of temporal anterior, a white ring beginning on 5th and 6th supralabials, sometimes beginning on half of 4th supralabial, passing by temporals towards parietals; middorsally, the black cap is joined to the first black body ring (nuchal collar) by the end of parietals (Figure 1). First to third infralabials black; 4th to 5th and half of sixth infralabials white, mental white with black spots; 12 to 21 black body rings (3 to 4 dorsals long), 12 to 17 in males and 13 to 21 in females, short white rings formed by a half scale with black top; 12 to 20 red body rings (6 to 20 dorsals long, with top 2/3 of scale black), ranging from 12 to 17 in males and 12 to 20 in females; 2 to 11 black tail rings (ranging from 6 to 12 in males and 2 to 11 in females) alternating with 1 to 12 white rings (ranging from 6 to 12 in males and 1 to 11 in females); the black tail rings can present white spots on dorsal and ventral surface, the spots can sometimes form bands or rings. There is a tendency toward melanism on the dorsal surface. The ventral surface can be spotted (Figure 2).</p> <p> <b>TABELE 1.</b> Meristic and mophometric data (mm) for <i>Micrurus paraensis</i> holotype (0851) and paratypes (0909, 0 951, 0 973, 1424, 1496 and 2665) (Legend: - = no data).</p> <p> <b>Hemipenis.</b> The hemipenis is up to 15 subcaudals long, and the bifurcation starts at around the 10th or 11th subcaudal. The base of the organ is naked up to the level of the 5th or 6th subcaudal where tiny spines appear on the sulcus ridge. Spinules and a few small spines begin at the 6th subcaudal and gradually increase in size until 9th subcaudal below the point where the sulcus bifurcates. The sulcus divides around the 8th or 9th subcaudals.</p> <p>At the level, large calcified spines appear and continue on each fork of the hemipenis but gradually diminish in size again toward the apex. A large lateral naked fold runs from the base of the organ to the level of the 8th subcaudal (Figure 3).</p> <p> <b>Variation and sexual dimorphism</b> (N = 64) (Tables 2 and 3). <i>Micrurus paraensis</i> presents significant sexual dimorphism in four morphometric characters (SVL (Snout-vent length); TAL (Tail length); TL (Total length); frontal width; distance between eyes) and six meristics (Ventral; Subcaudal; number of black and red body rings; number of white and black tail rings), as follows: largest male 463 mm SVL (= 362.5; SD = 64.2; N = 31) and 85 mm TAL (10 to 21% of TL); largest female 590 mm SVL (= 422.0; SD = 101.7; N = 22) and 77 mm TAL (9 to 18% of TL) (t = 3.7; df = 22; p> 0.01); frontal longer than wide, in male occupies 17 to 28 % of the HL and in females occupies 20 to 32 % the HL (t = -2.7; df = 56; p> 0.01); distance between eyes is 2.9 to 4.8 mm (= 4.0; SD = 0.4; N = 29) in males (43 to 73 % of the WH) and 3.2 to 5.8 mm (= 4.3; SD = 0.6; N = 22) in females (52 to 64 % of the WH) (t = -2.2; df = 56; p <0.05); ventrals 184 to 197 (= 191.3; SD = 3.6; N = 35) in males, and 186 to 217 (= 205.1; SD = 7.9; N = 27) in females (t = -7.9; df = 23; p> 0.01); subcaudals 36 to 54 (= 47.3; SD = 3.4; N = 35) in males, and 30 to 52 (= 38.0; SD = 6.6; N = 27) in females (t = 6.7; df = 23; p> 0.01); black body rings 12 to 18 (= 14.8; SD = 1.4; n = 35) in males; and 13 to 21 (= 17.1; SD = 2.2; N = 26) in females (t = -4.7; df = 59; p> 0.01); red body rings 12 to 17 (= 14.0; SD = 1.3; N = 34) in males, and 12 to 20 (= 16; SD = 2.1; N = 24) in females (t = -4.1; df = 58; p> 0.01); black tail rings 5 to 12 (= 8.2; SD = 1.7; N = 34) in males, and 2 to 11 (= 6.4; SD = 1.9; N = 23) in females (t = 3.9; gl = 55; p = 0.01); white tail rings 5 to 12 (= 8.2; SD = 1.7; N = 34) in males, and 1 to 11 (= 6.0; SD = 2.0; N = 23) in females (t = 4.2; df = 55; p> 0.01).</p> <p>Among the meristics characters, that showed differences were: 7 supralabials, (3 and 4th in contact with the eye) (N = 63) and 7 (3, 4 and 5th) (N = 01) and temporals 1+1/1+1 (N = 43), 1+2/1+2 (N = 09), 1+1/1+2 (N = 02), and 1+2/1+1 (N = 03).</p> <p> <b>Geographic range.</b> Tropical rainforest in southern Suriname and Brazil. In Brazil, it is present in the states of Pará, western Maranhão, southwest Mato Grosso and Rondônia (Figure 4).</p> <p> <b>Comments.</b> The variation of color patterns and the existence of melanic populations caused some taxonomic problems related to the proper identification of <i>M. paraensis</i>. The material analyses permitted to identification of melanic populations along the geographic distribution of <i>M. paraensis</i> in the states of Pará at Carajás, Melgaço and Mato Grosso state, Brazil. According to Roze (1994), this color variation is very common among coralsnakes belonging to the <i>psyches</i> group. In <i>M. paraensis</i> this fact is not related to sex, geographic distribution or age of individuals. Roze (1994) used the term “amazonic melanism” to characterize the melanin for the species from the <i>psyches</i> group, other species of <i>Micrurus</i> and colubrids. The variation on the number of black body rings does not differ from variation presented by Cunha & Nascimento (1978; 1982; 1993), Campbell & Lamar (1989) and Roze (1996).</p> <p> The variation of the distribution of black and white tail rings of <i>M. paraensis</i>, cannot be treated as a diagnostic character, as considered by Abuys (1987) to describe the subspecies <i>M. paraensis debruini</i>. This character utilized by the author seems to be very common in <i>M. paraensis</i>, being observed in individuals from Mato Grosso and Pará states in Brazil.</p> <p> According to Roze (1996) sexual dimorphism on the number of ventrals and subcaudals is common among species of <i>Micrurus</i>. Cunha & Nascimento (1982) noted sexual dimorphism in the number of ventrals and subcaudals in <i>M. paraensis</i>. Aside from, the characters presented by Cunha & Nascimento (1982), sexual dimorphism was detected in six meristics and four morphometric characters (Tables 2 and 3).</p> <p> The variation presented by Cunha & Nascimento (1978; 1982; 1993), Campbell & Lamar (1989; 2004), Roze (1996) and Carvalho (2002) of the number of ventrals and subcaudals in <i>M. paraensis</i> were: males have 188 to 200 ventrals and 45 to 52 subcaudals; females have 194 to 213 ventrals and 30 to 47 subcaudals. Based on analyses of a larger number of specimens, these characters presented the following ranges of variation: males have 184 to 197 ventrals and 36 to 54 subcaudals; and females have 186 to 217 ventrals and 30 to 52 subcaudals.</p> <p> In the hemipenis of species of <i>Micrurus</i> a large lateral naked fold is present, and the sulcus spermaticus is bifurcated (Slowinski, 1995; Roze, 1996). Hemipenial characters were also used to define two monophyletic groups (Roze, 1989, 1996; Campbell & Lamar, 1989; Slowinski, 1995). <i>Micrurus paraensis</i> shares with <i>M. albicinctus, M. averyi, M. langsdorffi, M. psyches</i> and others species with a monadal color pattern, long tail, long and strongly bifurcated hemipenis, with lobes elongated terminally and a large lateral naked fold; this may be a monophyletic group. According to Roze (1994), the <i>psyches</i> group form a natural group that share color pattern in monads, cephalic black cap, absence of supra-anal tubercles on males and presence of an elongated hemipenis, strongly bifurcated, and they can be differentiated by the number of ventrals and subcaudals, number of black and white body and tail rings. In the hemipenis of <i>M. paraensis</i>, described here for the first time, it is possible to observe the presence of characters considered to be synapomorphies of the <i>M. psyches</i> group (elongated organ and strongly bifurcated with lobes terminally elongated, and a large lateral naked fold, beyond, large calcified spines). Therefore, a systematic analysis in a phylogenetic paradigm could express the inferences of these characters.</p>Published as part of <i>Feitosa, Darlan Tavares, Prudente, Ana Lúcia Da Costa & Lima, Ana Caroline De, 2007, Redescription and variation of Micrurus paraensis Cunha & Nascimento 1973 (Serpentes: Elapidae), pp. 35-45 in Zootaxa 1470</i> on pages 36-44, DOI: <a href="http://zenodo.org/record/176669">10.5281/zenodo.176669</a>
Micrurus silviae Di-Bernardo, Borges-Martins & Silva, 2007, sp. nov.
Micrurus silviae sp. nov. (Figures 1 and 2) Type Material — Holotype: CRUPF 1488, adult male, from Passo Fundo Municipality, Miranda Stream (within the drainage basin of the Companhia Riograndense de Saneamento—CORSAN dam), in the W-NW of Rio Grande do Sul; collected on 21 March 2005 by Vilson Luiz Mello de Paulo. Paratypes: Rio Grande do Sul - IB 699 (M), Alegrete; MCN 4652 (M) and IB 52173 (M), Cacequí; IB 6072 (M), IB 16643 (M), IB 18689 (M), and IB 23418 (F), Carazinho; IB 53080 (M), Catuípe; MCN 5155 (M), IB 11122 (M), IB 11256 (M), IB 11432 (M), IB 15758 (M), IB 16604 (M), IB 18570 (M) and IB 40194 (F), Cruz Alta; IB 7357 (M), Friedhemi; IB 7846 (F), IB 8088 (M), and IB 23621 (F), Ijuí; IB 11615 (M), Itapeví, Alegrete; ZUFSM 0 86 (M), ZUFSM 2173 (F), and ZUFSM 2341 (M), Manoel Viana; CRUPF 581 (M), Mato Castelhano; CRUPF 149 (M), Nicolau Vergueiro; CRUPF 194 (M), Passo Fundo; IB 13640 (M), Rosário do Sul; IB 16055 (M), Santa Bárbara do Sul; MCN 3892 (M), Santa Maria; MCP 15983, Santo Augusto (M); MCP 12164 (F), Sarandi; MCN 8187 (F), Tupanciretã. Diagnosis — Micrurus silviae is diagnosable from all other South American triadal coralsnakes by the combination of the following fixed characters: a) snout and head mostly black colored with white bordered scales (anteriorly), including the rostral, internasals, nasals, prefrontals, preoculars, and postoculars; b) gular region white with few black markings; and c) middle black ring 1.5 or twice as long as the external ones. Additionally M. silviae is unique in possessing an anteromedial process in the frontal bones. Description of holotype —Rostral wider than high, prefrontals slightly longer than the internasals. Frontal longer than its distance from the snout; parietals as long as their distance from the snout. Dorsals in 15 - 15 - 15 rows; 1 + 2 temporals; 7 supralabials; 7 infralabials; no preocular; 2 postoculars; 226 ventrals, 5 preventrals, and 21 subcaudals (paired). Snout predominantly black with white bordered scales anteriorly. Top of the head black including the frontal, supraoculars and parietals. Anterior border of the parietals partially white, and anterior border of the supraoculars and frontal white. Anterior temporals red and heavily marked by black with a thin white border anteriorly; posterior temporals red marked with black posteriorly (inferior border of the posterior inferior temporal white). First six supralabials white with black markings along the posterior bor- der reaching the middle and superior part of the 2 nd, 3 rd, 4 th, and 5 th scales; 6 th supralabials mostly white (anterior half) with red posterior border, and black middle superior border; 3 rd and 4 th infralabials in contact with the orbit. Gular region white (mostly immaculate) with black markings between the anterior and posterior genials, and the posterior tip of the second genials. First five infralabials white; 6 th and 7 th infralabials red and all infralabials black marked posteriorly. 1 st and 2 nd ventral scales divided. The neck is red with first two to three scales heavily marked by black and the remainder of red scales black-tipped. The supraoculars are black with anterior white border. The first triad is separated from the parietals by two dorsal scales. There are 14 (13 + 1) black body triads, with the middle black ring two times longer than the external ones. White rings are blacktipped and shorter than the external black rings. Red rings shorter than the whole triad and all scales (red) are weakly black-tipped. Ventrally all black and white rings are irregular not always completing the circle around the body. Overall length of the holotype is 1,113 mm with a head length of 31 mm, a SVL of 1,050 mm, and a TL of 63 mm. The TL/SVL ratio is 0.06 (Figures 1 and 2). Variation —With the initial sample of 34 specimens males showed a range of 205 to 232 ventral scales, 18 to 26 subcaudal scales, and 10 to 15 triads (n = 27; Tables 1 and 2). Females showed a range of 211 to 234 ventral scales, 15 to 24 subcaudal scales, and 11 to 13 triads (n = 6; Tables 1 and 2). The first triad is separated from the parietal scales by two to five dorsal scales. The head (nasals, prefrontals, frontal, supraoculars and parietals) is black with thin white borders at the anterior suture of scales. Rarely there is a reduction of the black color at the parietals in the unique fashion observed in Micrurus altirostris. However, sometimes a reduction of black in the margins of the parietals and a varying degree of white in the snout is also observable. The temporal region has a variable degree of red and black. The red rings may attain the length of the whole triad and even be a 1 / 3 longer; black tips of scales may vary from moderate to absent. The position of the first triad (as separated from the parietals) is quite variable within a range of one and a half to six dorsal scales with one specimen with one and a half scales (holotype), six specimens with two scales (17.6 %), nineteen with three scales (55.9 %), five with four scales (14.7 %), two with five scales (5.9 %), and one with six scales (2.9 %). This is another example of the extreme values as 91.2 % of the specimens had between one and a half to four dorsal scales separating the first triad from the parietals. The gular region has no tendency to melanism. Maximum total lengths of specimens of M. silviae includes 1,506 mm (MCP 15983 M), 1,333 mm (UPF 194 M), 1,310 mm (IB 6072 M), and 1,294 mm (IB 13640 M). Color in preservative —All scales of the red rings are faded giving a cream color that is also visible on the head (behind and around the parietals). The black color of head, snout and triad rings are black and somewhat faded as a brownish color in old specimens. The white color of triad rings and gular region are light cream. The general morphological characters of head and body color, triad arrangement, and scale counts are also diagnosable (by the PAA criterion of Davis & Nixon 1992) by the following combination: a) head color pattern —mostly black in M. silviae, including the snout and all the head scales. In M. altirostris the black color is almost restricted to the supraoculars and frontal with the snout with varying degrees of melanism with a tendency to complete black scales (also interpreted as black snout with varying degrees of white markings). In most specimens, the parietals are almost completely red with a posterior black margin. In M. baliocoryphus the black color is mostly in parietals, frontal, and supraoculars with a white snout lightly marked with black; sometimes a white cross band might be present between the parietals and frontal and supraoculars. In M. frontalis, M. pyrrhocryptus and M. tricolor the head and snout are black with thin white markings between scales; very often the white markings are more evident in the snout of M. frontalis; b) gular region pattern —white in M. silviae (with very few black markings); black in the anterior part in M. altirostris with remaining posterior part red, which might also be interpreted as a red (including both pairs of gulars and 3 rd to 7 th infralabials) and white (anterior border of the 1 st pair of gulars an 1 st to 3 rd infralabials) with intense melanism. In M. baliocoryphus the gular region is mostly red with anterior part white (first three infralabials and anterior pair of genials), and red with varying black markings in M. frontalis, M. pyrrhocryptus and M. tricolor; c) number of triads —ranges from 11 to 16 in M. silviae, 12 to 20 in M. altirostris, 10 to 16 in M. baliocoryphus, 10 to 19 in M. frontalis, 6 to 14 in M. pyrrhocryptus, and 9 to 12 in M. tricolor; this is typical of several meristic characters that show considerable overlap among species (Table 1). For this character the most divergent species are M. altirostris and M. pyrrhocryptus (on both numerical extremes), but there is an obvious gradient among species when using means and extreme values. However, when using means and standard errors it is possible to separate the species that overlap with M. silviae and/or M. baliocoryphus (Figure 3). In M. silviae (range = 11–16 triads), 97.0% of the specimens (n= 33) are within the range of 11–15 triads, with one specimen having 16 triads (3.0%). In M. altirostris (12–20 triads) 98.8 % of the specimens (n= 574) are within the range of 13–18 triads, with six specimens with 19 triads (1.0%), and one specimen with 20 triads (0.2 %). In M. baliocoryphus (10–16 triads) 94.3 % of the specimens (n= 100) are within the range of 11 –15 triads, two specimens have 10 triads (1.9 %), and four specimens with 16 triads (3.8 %). In M. frontalis (10–19 triads) 98.8 % of the specimens (n= 565) are within the range of 11–17 triads, three specimens have three triads (0.5 %), and three specimens with 18 triads (0.5 %), and one specimen with 19 triads (0.2 %). In M. pyrrhocryptus (6–14 triads) 96.6 % of the specimens (n= 225) are within the range of 7 and 11 triads, four specimens have 6 triads (1.7 %), one specimen with 12 triads (0.4 %), two specimens with 13 triads (0.9 %), and one specimen with 14 triads (0.4 %). In M. tricolor (9–12 triads) 95.0% of the specimens (n= 19) are within 9 and 11 triads, and one specimen had 12 triads (5.0%). All species show a pattern of one complete tail triad and usually 1 / 3 and rarely 2 / 3 of an incomplete triad. The same assumptions of extreme values and standard errors are applicable to ventral and subcaudal scales (see below); d) position of the first triad —in M. silviae the first body triad is separated from the parietals by two to six scales, in M. altirostris it ranges from one to three scales, in M. baliocoryphus it ranges from three to six scales, in M. frontalis it ranges from one to seven scales (fusion of the black color of head and first triad is often observable); in M. pyrrhocryptus it ranges from five to eight scales, and in M. tricolor it ranges from seven to eleven scales. This character is subject to some variation but is useful for distinguishing these last two species (as a group) from the first four (as a group); e) pattern of triads —middle black ring one-and-one-half to twice as long as the external ones in M silviae with red and white dorsal scales usually black-tipped (weakly tipped on red rings) and white rings shorter than the external black rings; subequal black rings in M. altirostris (middle one can be a little longer), with white rings as long as the external black rings (sometimes a little shorter, and very short in some specimens from Uruguay) with dorsal scales black-tipped (a little less in the red rings); middle black ring 2 to 2.5 times longer than the external ones in M. baliocoryphus with white rings (immaculate in anterior 1 / 3 to 2 / 3 of body) as long as the external black rings (Table 2); subequal black and white rings in M. frontalis (sometimes the middle black ring may be slightly longer than the external ones) with varying degrees of black tipped scales; middle black ring 2 to 4 times longer and white rings shorter than the external black rings in M. pyrrhocryptus with all scales black tipped; and middle black ring 2 to 3 times longer than the external ones in M. tricolor with white rings (immaculate in anterior 1 / 3 to 2 / 3 of body) of the same length of the external black rings. The description of a red or white ring being immaculate means that they are without the heavy black markings on the tip of the dorsal scales, they are extremely variable among specimens of a given species, and they are sometimes restricted to the posterior 1 / 3 or 2 / 3 of body; f) number of ventral scales —ventral scales range from 205 to 232 in males (x= 220.52) and from 211 to 234 (x= 221.43) in females of M. silviae; 173 to 242 in males (x= 208.80) and from 185 to 223 in females (x= 208.85) of M. altirostris; 172 to 232 in males (x= 221.06) and from 208 to 233 in females (x= 220.10) of M. baliocoryphus; 190 to 250 in males (x= 225.07) and from 197 to 252 in females (x= 225.93) of M. frontalis; 202 to 244 in males (x= 228.33) and 213 to 251 in females (x= 232.09) of M. pyrrhocryptus; and 214 to 234 in males (x=226.00) and 208 to 231 in females (x= 221.12) of M. tricolor (Table 1). This character shows considerable variation among the six species with an overlap owing to extreme values, which can be better demonstrated with standard errors as some species are separable (i.e., low numbers in M. altirostris and high in M. pyrrhocryptus; Figure 4); g) number of subcaudal scales —subcaudal scales range from 18 to 25 in males (x= 22.30) and from 15 to 24 (x= 18.71) in females of M. silviae; 12 to 28 in males (x= 19.83) and from 11 to 25 in females (x= 18.04) of M. altirostris; 14 to 31 in males (x= 22.82) and from 14 to 23 in females (x= 20.15) of M. baliocoryphus; 14 to 27 in males (x= 21.94) and 14 to 28 in females (x= 19.86) of M. frontalis; 16 to 30 in males (24.56) and 19 to 26 in females (x= 21.97) of M. pyrrhocryptus; and 24 to 29 in males (x= 26.42) and 20 to 26 in females (x= 22.50) of M. tricolor (Table 1). This character shows the widest range of overlap among the six species but with standard errors both sexes for most species are separable (Figure 5). Sexual dimorphism —to demonstrate sexual dimorphism the ratio of TL/SVL is usually adopted with varying degrees of success (Roze 1996; Silva & Sites 1999; Campbell & Lamar 2004). For this group of triadal coralsnakes there is also a considerable overlap of values. However, with standard errors both sexes are easily separated (Figure 6). It can be also highlighted with the data of ventral and subcaudal scales (see also Figures 4 and 5). Hemipenial morphology —as in most of triadal coralsnakes the hemipenis of M. silviae is bilobed with uniform size spines and a bifurcated sulcus spermaticus (CRUPF 1488 — holotype, right organ; Figure 7). In the sulcate and asulcate surfaces the spines are large and uniform from the tip to near the base. The lobes are moderately long and conical; viewing from the sulcate side, the tip of the right lobe is more acute than the left. Measuring from the division point of the sulcus spermaticus, the lobes are 10.0 mm long, representing 48.3 % of the total length of the organ. Between the lobes (in the intersection of them) there is a poorly ornamented region, with few and small spines, better visible from the sulcate side. The fully everted organ is 20.7 mm long, reaching the level of the 7 th subcaudal. The distance from the base of the organ to the division point of the sulcus is 13.6 mm; the division occurs at the level of the 4 th subcaudal. The base of the organ is 5.3 mm long and naked. Cranial osteology —the premaxilla presents the posterior extension of the palatine process in both sides, a character shared with M. altirostris, M. baliocoryphus, M. frontalis, M. pyrrhocryptus, and M. tricolor. The dorsomedial process of the prefrontals are narrow and do not contact at midline. Only M. frontalis and M. pyrrhocryptus seem to present this condition. The anteromedial process of the frontals is present in M. silviae and absent in all other species. The parietal is longer and wider (measured at its widest point) in M. silviae. There is a dorsal crest at the midline of the parietal bone that might present a flat triangular tabular process anteriorly. This process might be discrete and small (as in M. pyrrhocryptus), reach the anterior 1 / 3 (M. silviae), or be wider and reach 2 / 3 of the entire length of the parietal bone (M. altirostris, M. baliocoryphus, M. frontalis, and M. tricolor). In M. silviae the parietal bone also presents a similar process at the posterior end of the parietal which is also pointed (as in M. pyrrhocryptus and M. tricolor). The suture of prefrontals + frontals is anterior to the maxilla + pterygoid suture in M. silviae, M. baliocoryphus, M. frontalis, M. pyrrhocryptus, and M. tricolor, and posterior in M. altirostris. In M. silviae there are seven to eight palatine, four to seven pterygoid, and ten to twelve dentary teeth. There are no noticeable differences in shape and proportions of the remaining cranial bones (Figure 8). However, there is an ongoing osteological study within the triadal coralsnakes that might contribute to better understand the fixed and variable conditions of this set of characters. Statistical summary of morphological variation — Tables 1 and 2 summarize the descriptive statistics for all meristic and morphometric data collected from the specimens used in this description and the comparative species. For all characters analyzed, it is possible to observe a large overlap of variation among species, as revealed by maximum and minimum values, but in general, the use of standard errors show statistical differences among the species. Wilk’s statistics was equal to 0.045 (F = 111.98, P <0.0000), indicating significant differences among species in the multivariate space of the characters analyzed. The first two canonical axes explain 95 % of the variation among species, which 81 % of this total amount being concentrated in the first canonical axis. This first axis is basically composed by number of triads (TRI) and length of middle black ring (MBL). The second axis is defined by ventrals (VE) and length of anterior black ring (ANTBL) (Table 3). The distribution of individuals in the bivariate space formed by the first two canonical axis reveal that M. altirostris and M. pyrrhocryptus appear in opposite regions of the space along the first axis. In the other hand, M. baliocoryphus, M. silviae, M. tricolor, are in a clear intermediate position between the two extremes (M. altirostris and M. pyrrhocryptus), whereas M. frontalis overlaps with M. silviae and M. altirostris and M. baliocoryphus. There is also an overlap of M. tricolor and M. pyrrhocryptus (Figure 9). These results of CVA are qualitatively similar when the sexes are analyzed separately (data not shown). Despite the overlap of M. silviae and M. baliocoryphus and M. tricolor and M. pyrrhocryptus on both canonical axes there are several fixed characters that separate these species. For M. silviae and M. baliocoryphus it includes: (a) length of middle black ring; (b) length of white rings; (c) absence or presence of melanism on anterior 1 / 3 to 2 / 3 of white rings; (d) color pattern of the gular region; (e) color pattern of the snout; (f) color pattern of the head; (g) cranial osteology features. For M. tricolor and M. pyrrhocryptus it includes: (a) length of middle black ring; (b) distance of first triad from parietal scales; (c) length of external black rings compared to the middle black ring; (d) cranial osteology features.The Hotelling’s T 2 -test indicated that all species differ significantly from each other considering the 12 variables used (p<0.0001) (Table 4). When using the Mann Whitney U-test for number of ventral, subcaudal scales and triads number separately many differences were significative even after the Bonferroni’s correction for multiple significance tests. However, for each of these three variables there were exceptions: (a) ventral scales—there were no significant differences between M. baliocoryphus and M. silviae (p= 0.306), M. baliocoryphus and M. tricolor (p= 0.142), M. frontalis and M. tricolor (p=0.600); and M. silviae and M. tricolor (p= 0.057); (b) subcaudal scales—there were no significant differences between M. baliocoryphus and M. frontalis (p= 0.070), M. baliocoryphus and M. silviae (p=0.400), M. frontalis and M. silviae (p= 0.840), and M. pyrrhocryptus and M. tricolor (p= 0.294); (c) number of triads—there were no significant differences between M. baliocoryphus and M. silviae (p=0.500), and M. baliocoryphus and M. frontalis (p=0.100) (Table 5). Etymology —the specific name is a noun in genitive case, in honor of Silvia Di Bernardo a promising herpetologist that died suddenly in July 2002. Geographic Distribution — Micrurus silviae is known from 17 localities in Western to Northern Rio Grande do Sul, Brazil. One specimen was photographed in one additional locality, the Municipality of São Francisco de Assis (not collected). It is possible that its range includes Misiones, part of NE-E Corrientes (Argentina) and SE-E of Departamento Itapua (Paraguay) (Figure 10). TABLE 4. T 2 -test between species of coralsnakes. Above diagonal are the significance value levels. Below diagonal are T 2 -test values. Legend: Species 1 – M. altirostris; Species 2 – M. baliocoryphus; Species 3 – M. frontalis; Species 4 – M. pyrrhocryptus; Species 5 – M. silviae; Species 6 – M. tricolor. Legend: Species 1 – M. altirostris; Species 2 – M. baliocoryphus; Species 3 – M. frontalis; Species 4 – M. pyrrhocryptus; Species 5 – M. silviae; Species 6 – M. tricolor. Micrurus silviae probably occurs in open habitats, as its distribution is associated to the grasslands areas of western and northern Rio Grande do Sul. The geographical range of known records agrees closely to the northern delineation of the Uruguayan Savannas ecoregion along its interfaces with the Paraná-Paraíba Interior Forests and Araucária Moist Forests (Olson et al., 2001; WWF, 2001). This region corresponds to the range of grasslands inter-laced with gallery woodlands (IBGE, 1986). Distribution occurs along different geomorphologic units, encompassing parts of the Peripheric Depression and the Brazilian Southern Plateau, extending from the western lower elevations in the Depression of the Ibicuí-Negro Rivers and the Uruguaiana Plateau to the northern Santo Angelo Plateau (IBGE, 1986). Altitudinal distribution accordingly ranges from about 100 to nearly 700 m (above sea level) in Mato Castelhano. Other sympatric species —Within the same geographical region there are records of one monadal spec
<i>EGLN1</i> involvement in high-altitude adaptation revealed through genetic analysis of extreme constitution types defined in Ayurveda
It is being realized that identification of subgroups within normal controls corresponding to contrasting disease susceptibility is likely to lead to more effective predictive marker discovery. We have previously used the Ayurvedic concept of
Prakriti
, which relates to phenotypic differences in normal individuals, including response to external environment as well as susceptibility to diseases, to explore molecular differences between three contrasting
Prakriti
types:
Vata, Pitta, and Kapha
. EGLN1
was one among 251 differentially expressed genes between the
Prakriti
types. In the present study, we report a link between high-altitude adaptation and common variations rs479200 (C/T) and rs480902 (T/C) in the
EGLN1
gene. Furthermore, the TT genotype of rs479200, which was more frequent in
Kapha
types and correlated with higher expression of
EGLN1
, was associated with patients suffering from high-altitude pulmonary edema, whereas it was present at a significantly lower frequency in
Pitta
and nearly absent in natives of high altitude. Analysis of Human Genome Diversity Panel-Centre d’Etude du Polymorphisme Humain (HGDP-CEPH) and Indian Genome Variation Consortium panels showed that disparate genetic lineages at high altitudes share the same ancestral allele (T) of rs480902 that is overrepresented in
Pitta
and positively correlated with altitude globally (
P
< 0.001), including in India. Thus,
EGLN1
polymorphisms are associated with high-altitude adaptation, and a genotype rare in highlanders but overrepresented in a subgroup of normal lowlanders discernable by Ayurveda may confer increased risk for high-altitude pulmonary edema.
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