Micrurus potyguara, Pires, Matheus Godoy, Jr, Nelson Jorge Da Silva, Feitosa, Darlan Tavares, Prudente, Ana Lúcia Da Costa, Filho, Gentil Alves Pereira & Zaher, Hussam, 2014

Pires, Matheus Godoy, Jr, Nelson Jorge Da Silva, Feitosa, Darlan Tavares, Prudente, Ana Lúcia Da Costa, Filho, Gentil Alves Pereira & Zaher, Hussam, 2014, A new species of triadal coral snake of the genus Micrurus Wagler, 1824 (Serpentes: Elapidae) from northeastern Brazil, Zootaxa 3811 (4), pp. 569-584 : 571-578

Micrurus potyguara sp. nov.

Micrurus lemniscatus carvalhoi Roze, 1967: 33 (partim).

Holotype. Adult male, UFPB 4359, collected by Gentil Alves Pereira Filho on August 22, 2006 at Mata do Buraquinho (7°8' 42.08"S, 34° 51'42.40"W), municipality of João Pessoa, state of Paraíba.

Paratypes. Five specimens: adult female ( Figure 1 ), UFPB 4355, collected on April 11, 2006; adult male, UFPB 4358, collected by Gentil Alves Pereira Filho on August 21, 2006; adult female, UFPB 4361 collected by Gentil Alves Pereira Filho on September 6, 2006 from Mata do Buraquinho, João Pessoa. NRM 1685, immature male, donated by N. E. Forssell to the museum in 1847 from municipality of Recife (8°3'S, 34°53'W), state of Pernambuco; and adult female, CAS 49297, collected by H. Heath on November 30, 1910 from Lagoa de Paparí (6°7'38.13"S, 35°9'54.64"W), municipality of Nísia Floresta, state of Rio Grande do Norte.

Diagnosis. Micrurus potyguara is distinguished from all other triadal species of Micrurus by combination of the following characters: a) parietals entirely black; b) head conspicuously broader than neck; c) posterior nasal scale black; d) frontal shield longer than distance from its anterior margin to the tip of the snout; e) infralabials mostly black; f) middle black rings of triads shorter or as long as the outer ones; g) gular region red with small disperse black blotches on posterior region of scales; h) tip of snout entirely black (rostral, internasals and nasals); d) black rings with similar length or central black ring slightly shorter than the anterior and posterior black rings of the triads.

Comparisons between species of the triadal group of coral snakes from northeastern Brazil. Three triadal coral snakes species are known to be sympatric or, at least, to approach the distribution of Micrurus potyguara . These are: M. lemniscatus carvalhoi , M. ibiboboca and M. brasiliensis . Differences between these species and M. potyguara are given below and summarized in Table 1 (character states in M. potyguara are shown in parenthesis).

M. potyguara (n=6) M. ibiboboca (N=536) M. l. carvalhoi (N=129) M. brasiliensis (N=31)

Range  SD Range  SD Range  SD Range  SD VE ♂ 231–237 234.7 3.2 198–247 222.5 9.1 224–263 237.6 8.3 210–243 226.5 7.8 ♀ 253–263 257.3 5.1 188–254 226.0 10.1 222–267 253.4 9.3 219–237 226.9 5.9 SC ♂ 34–38 36.0 2.0 17– 33 23.7 2.6 25 – 39 31.7 3.0 17– 28 22.5 2.7 ♀ 29–35 33.0 3.5 16 – 34 22.7 2.7 26 – 38 30.1 2.9 14 – 36 23.3 5.7 TriB ♂ 9–11 10 1. 0 7– 13 9.1 1.1 10 – 15 12.7 1.3 10 – 16 11.6 1.7 ♀ 11– 12 11.7 0.6 7– 12 9.1 1.1 9– 16 12.5 1.6 9– 13 10.8 1.1 TriT ♂ 1 2/3 * * 2/3–2 * * 1 1/3–2 * * 1–1 2/3 * * ♀ 1 1/3–1 2/3 * * 2/3–1 2/3 * * 1 1/3–2 * * 1–1 2/3 * * Micrurus lemniscatus carvalhoi ( Figures 2 b, 3i, 4b) shows a broad color and meristic variability along its widespread distribution in central and eastern regions of Brazil. However, when compared with M. potyguara ( Figures 2 a, 3a–f, 4a), M. l. carvalhoi has a black head cap that does not extend over all the surface of the parietal shields (vs. extended over all the parietal surface), the head is indistinct from the neck (vs. head distinct from neck), and the median black rings are never shorter than the anterior and posterior black rings on the same triad (median black rings longer than outer black rings); the hemipenis has a straight sulcus spermaticus (vs. curved), with longitudinal rows of spines along its margin (vs. lacking longitudinal rows of spines along the sulcus spermaticus), a capitular region longer than half of the total length of the organ and ornamented with very long, straight spines (capitular region shorter than half of the organ's length and ornamented with comparatively stouter, shorter spines), the base of the organ is ornamented proximally with longitudinal rows of slender spinules, retains a long spineornamented region on the hemipenial body, and a long basal naked pocket (vs. base of the organ with irregularly arranged spinules proximally, with a shorter spine-ornamented region, and a shorter basal pocket).

Micrurus brasiliensis ( Figures 2 c, 3g, 4c) can be easily distinguished from M. potyguara by its red parietals and white snout and genial regions (vs. black parietals, snout and anterior genial region), the absence of the preocular white band (vs. preocular white band), triads with white rings significantly larger than the black rings (vs. white rings smaller than black rings), a smaller number of ventral and subcaudal scales (see Table 1), and a hemipenis with a sulcus spermaticus bifurcating below the level of the crotch of the lobes (bifurcates at the level of the crotch), with a capitular region that corresponds to two-third of the length of the organ that is ornamented with irregularly arranged and stouter spines and delimited by a shallower, less evident capitular groove (capitular region about half of the length of the organ, ornamented with slender spines, evident capitular groove).

Micrurus ibiboboca ( Figures 2 d, 3h, 4d) can be distinguished from M. potyguara by a lower number of ventral and subcaudal scales ( Table 1), white triadal rings as long as or longer than the black ones (vs. white rings shorter than black ones), snout scales often marked with white, a genial black blotch restricted to mental region (vs. region of the snout anterior to white preocular band is completely black and extensive black horseshoe shaped blotch over infralabials), a stouter hemipenis, with a capitular region longer than half of the total length of the organ and ornamented irregularly with small, less densely arranged spines (capitular region shorter than half of the organ's length and more densely ornamented with larger spines), lobes very short, with less than one-fifth of the total length of the organ (vs. about ¼ of the total length of the organ), and a basal region ornamented with small irregularly distributed spines (vs. spines arranged into longitudinal rows).

Description of the holotype. UFPB 4359, adult male, TTL 934.5 mm, HL 18.5 mm HL, SVL 840 mm, TL 76 mm, TL/SVL ratio 0.0 9. Dorsal scales 15–15–15, VE 237, SC 36 (pairs), anal plate divided, TriB 9, TriT 1 2/3, AB and PB rings ~ 5 scales long, MB ~ 4 scales long, AW and PW ~ 3 scales long, red rings slightly longer or as long as AB and PB rings. Head broader than neck, body wider than high ( Figure 5 ). Rostral wider than high; prefrontals slightly longer than internasals; frontal 1.5 times longer than its distance from snout; parietals twice as long as wide, as long as their distance from snout; frontal hexagonal, 1.5 times longer than wide. Temporals 1+1, Supralabials 7 (3rd & 4th bordering the eye), infralabials 7, preocular 1, postoculars 2. Snout black [rostral; nasals; internasals; prefrontals (anterior margins); supralabials 1 and 2 (dorsal half)]. White transversal prefrontal band [anterior ¼ of the frontal, prefrontals, preoculars (anterior ⅓), supralabials 2 (posterior half), 3 and 4 (anterior ⅓) at the left side and 2, 3, 4 and 5 (anterior margin) at the right side of head]. Black cephalic cap [frontal; supraoculars; parietals; preoculars (posterior tip); left supralabials 3 (posterodorsal corner), 4 (dorsal half), 5 and 6 (anterodorsal corner); right supralabial 5 (posterior half)]. Nuchal surface of the head red with black-tipped scales ( Figure 6 ). AB ring of anterior TriB separated from black cephalic cap by 3 red scales at middorsum. Genial and gular regions mainly red with horseshoe-shaped black perioral mark [mental, infralabials 1, 2, 3 and 4 (anterior part), anterior genials (anteriot half), posterior genials (lateral margins at the contact with infralabials 4]. Preventrals 1,2 and 3 and adjacent scales mottled with small irregular black blotches (posterior margins). AB ring of 1st TriB at the level of posterior margin of 1st VE.

Hemipenis of the holotype. Everted and expanded hemipenis ( Figure 4 a) is bilobed, short and semicapitate (see discussion). Sulcus spermaticus curved, deep and bordered by small spines, bifurcated at the base of the lobes with diverging branches oriented medially and reaching the apexes. Capitular groove evident but not deep, transversely oriented on the asulcate surface, encircling the organ and bending bilaterally towards the base adjacent to the sulcus spermaticus at the sulcate surface. Lobes short (less than half of the capitular region length). Capitular region uniformly ornamented with oblique rows of stout, curved spines, decreasing in size towards the apexes of the lobes except in the lobular crotch, ornamented with smaller and less densely arranged spines. Basal region ornamented proximally with sparse and irregularly arranged spinules and small spines (smaller than the capitular ones) distally. Basal pocket deep, with a naked bottom and bordered with spinules.

♂ ♀

UFPB 4359 NRM 1685 UFPB 4358 CAS 49297 UFPB 4361 UFPB 4355 Variation ( Table 2 ). Largest specimen 916 mm TTL; largest female 751 mm TTL. Tail longer in males (~ 8.63% of TTL) than in females (~ 7.20% of TTL). Ventral scales 253–263 ( = 257.33; SD = 5.13) in females, 231–237 (? = 234.67; SD = 3.21) in males; subcaudal scales 34–38 (? = 36; SD = 2) in males, 29–35 (? = 33; SD = 3.46) in females. Males with 9–11 body triads and 1 2/3 tail triads; females with 11–12 body triads and 1 1/3–1 2/3 triads on the tail. Black irregular blotches over preventrals and adjacent scales of gular region. Anterior black ring of the first body triad separated from parietals by 2–3 red vertebral scales. Anterior and posterior black rings slightly longer or same length of middle black ring. White rings about half the length of middle black ring ( Figure 3 ). Anterior and posterior red rings of same length or longer than black rings. Black-tipped scales on Red and White rings rendering a finely reticulated aspect. Irregularly distributed black dots can be scattered on red rings. Remaining characteristics similar to holotype.

Geographic distribution and habitat. Micrurus potyguara is only known from tropical ombrophilous lowland coastal forests (tabuleiros) of the states of Paraíba, Pernambuco, and Rio Grande do Norte in Brazil, ( Figure 7 ). The Brazilian coastal rainforests (tabuleiros) occurs throughout northeastern region, extending from Rio Grande do Norte south as far as Espírito Santo and presents a compound of typical Atlantic Coastal Forests species ( Barbosa & Thomas 2008). The arboreal strata can reach 35 meters in altitude in forested and 10 meters in savanna areas. These savannas are located within forested areas and exhibits a sandy soil covered by grassy fields with scattered small trees and are locally called as tabuleiros ( Figure 8 ). The coastal forests of the states of Paraíba, Pernambuco and Rio Grande do Norte fall under this classification according to Barbosa (2008) and Silva & Rodal (2008).

Etymology. The specific epithet “ potyguara ” is a Tupi indigenous name, here employed as a noun in apposition (poty = shrimp;`war = eaters) (Gregório 1980). This name refers to the denomination of a native people that occupied historically the northeastern coast of Brazil, for which the shrimp fishing is one of the most traditional economic activities.

Statistical analysis. The PCA performed for separate sexes rendered two separate groups, M. brasiliensis + M. ibiboboca and M. l. carvalhoi + M. potyguara sp. nov. ( Figure 9 ). The most important data contributing for this distribution are presented at the Table 3 . The DFA also supported the same group formation ( Figure 10 ). Wilks’ Lambda, F e p values for the characters of DFA are presented at the Table 4 . The poor specific-level definition obtained by means of statistical analysis within the genus Micrurus is a well-known phenomenon, already demonstrated e.g. by Silva Jr. & Sites (1999), Di-Bernardo et al. (2007), Feitosa et al. (2007), is generated by the conservative nature and overlapping of the meristic and morphometric characters within the genus and reinforces the need to rely on qualitative characters over extensive samples to achieve most accurate taxonomic resolution.

Figures

FIGURE 1. Live specimen of Micrurus potyguara paratype, UFPB 4355, from João Pessoa, state of Paraíba, Brazil. Photo by Claudio Sampaio.

FIGURE 2. Dorsal and lateral views of the head of the holotypes of Micrurus potyguara UFPB 4359 (a, e), Micrurus lemniscatus carvalhoi USNM 76341 (b, f), Micrurus brasiliensis UMMZ 108880 (c, g) and Micrurus ibiboboca AMNH 3937 (d, h).

FIGURE 3. Graphic expression of the proportional triad pattern of the holotype of Micrurus potyguara UFPB 4359 (a) and the paratypes UFPB 4358 (b), UFPB 4361 (c), CAS 49297 (d), NRM 1685 (e), UFPB 4355 (f) compared with the holotypes of M. brasiliensis UMMZ 108880 (g), M. ibiboboca AMNH 3937 (h), and M. lemniscatus carvalhoi USNM 76341 (i).

FIGURE 4. Sulcate (left) and asulcate (right) views of the hemipenis of Micrurus potyguara (a), M. lemniscatus carvalhoi (b), M. brasiliensis (c) and M. ibiboboca (d). Scale bars = 5 mm.

FIGURE 5. Dorsal (a) and ventral (b) views of Micrurus potyguara holotype, UFPB 4359, from João Pessoa, state of Paraíba, Brazil.

FIGURE 6. Dorsal (a), lateral (b), and ventral (c) views of head and first body triad of Micrurus potyguara holotype, UFPB 4359. Drawings by Darlan T. Feitosa.

FIGURE 7. Geographic distribution of Micrurus potyguara (yellow diamonds), Micrurus l. carvalhoi (red circles), Micrurus ibiboboca (orange squares) and M. brasiliensis (purple diamonds) in Central and Northeastern Brazil.

FIGURE 9. Bivariate plot for the first two axes derived from scores of PCA analysis for males (A) and females (B) for M. ibiboboca, M. brasiliensis, M. l. carvalhoi e M. potyguara sp. nov. (95 % confidence regions).

FIGURE 10. Bivariate plot for the first two axes derived from scores of DFA analysis for males (A) and females (B) for species M. ibiboboca, M. brasiliensis, M. l. carvalhoi e M. potyguara sp. nov. (95 % confidence regions).

FIGURE 8. M. potyguara type locality landscape. Tropical ombrophilous lowland coastal forests of Northeastern Brazil from Rio Grande do Norte to Espírito Santo states, known as “ tabuleiros ”.

Tables

TABLE 2. Meristic and morphometric variation of the Micrurus potyguara. Abbreviations = snout-vent length (SVL), tail length (TL), head length (HL), length of the anterior red ring of the median triad (AR), length of the anterior black ring of the median triad (AB), length of the anterior white ring of the median triad (AW), length of the middle black ring of the median triad (MB), length of the posterior white ring of the median triad (PW), length of the posterior black ring of the median triad (PB), length of the posterior red ring of the median triad (PR), number of triads on body (TriB), and number of triads on tail (TriT).

Meristic VE 237	231	236	256	263	253
SC 36	38	34	29	35	35
TriB 9	10	11	12	12	11
TriT 1 2/3	1 2/3	1 2/3	1 1/3	1 2/3	1 2/3
Morphometrics (mm) SVL 840	259	620	420	623	700
TL 76	27	55	33	50	51
HL 18.5	9.81	13.9	13,36	13.66	15.5
AR 21	3.53	13.2	10,16	13.1	10.5
AB 15.4	5.28	10.1	8,01	11	9.2
AW 7.4	2.1	5.6	3,84	5.3	4.6
MB 10.8	5.41	10.1	10,27	11.1	12.4
PW 7.4	1.93	5.6	4,75	5.3	4.6
PB 15.4	4.34	10.1	6,45	11	9.2
PR 21	5.57	13.2	11,75	13.1	10.5