A new species of burrowing snake (Serpentes: Dipsadidae: Apostolepis) from the state of Mato Grosso, Central-West region of Brazil Author Santos, Fernanda Martins dos Author Entiauspe-Neto, Omar Machado Author Araujo, Jailini da Silva Author Souza, Moises Barbosa de Author Lema, Thales de Author Struessmann, Christine Author Albuquerque, Nelson Rufino de text Zoologia 2018 35 1 10 http://dx.doi.org/10.3897/zoologia.35.e26742 journal article http://dx.doi.org/10.3897/zoologia.35.e26742 1984-4689-35 FD43851ED39C48319459A2CCD2EBE7C6 FF88CF197E69055BFFF4FF94903BD44D 1290102 Apostolepis kikoi sp. nov. Figs 1 , 2-3 , Table 1 Apostolepis sp. - Struessmann 2000 : 163. [Cresonymy] Apostolepis sp. 1 - Martins and Lema 2015 : 102; Lema and Renner 2016 : 71. [Cresonymy] Apostolepis sp. 3 - Martins and Lema 2015 : 102 ( partim ). [Cresonymy] Apostolepis aff. borellii - Lema and Campbell 2017 : 28 ( partim ). [Cresonymy] Holotype. A female (MCP 12096) collected in 2000 at the Manso multi-use reservoir and hydroelectrical power plant - locally known as APM Manso - constructed at the confluence of the rivers Manso and Casca, Chapada dos Guimaraes ( 15°27 '39" S , 55°45 '00" W ; 811 m.a.s.l.), Mato Grosso, Brazil, by the faunal rescue team. Paratypes. three males and one female, same locality as the holotype: MCP 14524 (male), MCP 14525 (male) and MCP 11372 (female), date of collection unknown, collected by the faunal rescue team, and UFMT-R 1933 (male) collected on 1 December 1999 by the faunal rescue team. Diagnosis and comparison with other species. Apostolepis kikoi sp. nov. can be distinguished from all other Apostolepis by the combination of having five dorsal stripes (vs. dorsal stripes absent in A. ambiniger , A. ammodites , A. assimilis , A. breviceps , A. cearensis , A. dorbignyi , A. flavotorquata , A. multicincta , A. roncadori and A. tertulianobeui ; the presence of seven stripes on the dorsum in A. gaboi and A. niceforoi ; three stripes on the dorsum in A. cerradoensis , A. goiasensis , A. nigrolineata , A. quirogai and A. tenuis ; a pair of narrow lateral stripes in A. barrioi ; a pair of wide lateral stripes in A. albicollaris , A. dimidiata and A. polylepis ); the presence of a white nuchal collar (vs. white nuchal collar absent in A. ambiniger , A. barrioi , A. breviceps , A. christineae , A. goiasensis , A. intermedia , A. lineata , A. longicaudata , A. niceforoi , A. nigrolineata , A. polylepis , A. quinquelineata , A. rondoni , A. serrana , A. striata , A. thalesdelemai and A. vittata ); the presence of a triangular blotch covering portions of the third, fourth, fifth and sixth supralabials (vs. a light lateral spot below the eye, usually occupying the third and fourth supralabials in A. mariae ); 15 scale rows at midbody (vs. 17 in A. polylepis ); six supralabials (vs. five in A. breviceps , A. christineae , and A. vittata ); second and third supralabials entering orbit (vs. only the third supralabial entering orbit in A. breviceps ); preocular contacting nasal (vs. nasal and preocular separated by prefrontal in A. ammodites , A. arenaria , A. assimilis , A. breviceps , A. cearensis , A. dorbignyi , A. gaboi , A. goiasensis , A. intermedia , A . multicincta , A. phillipsi , A. polylepis , A. quirogai , A. tenuis and A. tertulianobeui ); temporals absent (usually 0 + 1 in A. ammodites , A. assimilis , A. cearensis , A. mariae , A. niceforoi , A. nigrolineata , A. quirogai , A. tertulianobeui and A. thalesdelemai , and 1 + 1 in A. flavotorquata and A. quinquelineata ); seven infralabials (vs. five in A. breviceps ; five to six in A. nelsonjorgei and A. vittata ; six in A. christineae , A. intermedia , A. multicincta ; eight in A. gaboi and A. quirogai ); four infralabials contacting the first pair of chinshields (vs. three infralabials contacting the first pair of chinshields in A. dorbignyi , A. intermedia , A. multicincta , A. tenuis and A. vittata ); a higher number of ventrals than A. arenaria and A. striata (203-209 vs. 168-181 and 202, respectively); fewer ventrals than A. christineae , A. intermedia , A. longicaudata , A. nelsonjorgei , A. niceforoi , A. phillipsi , A. polylepis , A. serrana , A. tertulianobeui , A. thalesdelemai and A. vittata (203-209 vs. 211-248 in the latter eleven species); fewer subcaudals than A. borellii , A. intermedia , A. longicaudata , A. nelsonjorgei , A. serrana and A. tertulianobeui (26-30 vs. 32-55 in the latter six species); a higher number of subcaudals than A. lineata , A. niceforoi and A. polylepis (26-30 vs. 24, 23 and 20-25, respectively) and fewer maxillary teeth than A. longicaudata and A. phillipsi (4 + 2 vs. 5 + 2 in the latter two species). Preserved specimens of Apostolepis kikoi sp. nov. are most similar to A. borellii , A. lineata , A. nelsonjorgei , A. nigroterminata , A. serrana and A. underwoodi in its general pholidosis and coloration pattern. However, the new species is distinguished from A. borellii by having its paraventral sides cream and unblemished (vs. blackish), a pair of triangular blotches covering portions of the third, fourth, fifth and sixth supralabials (vs. a small, trapezoidal blotch covering only the posterior half of the third and the entire fourth supralabial); tip of the tail conical (vs. rounded) and fewer subcaudals (vs. 26-30 vs. 32). Apostolepis kikoi sp. nov. can be distinguished from A. nigroterminata by having a triangular blotch covering portions of the third, fourth, fifth and sixth supralabials (vs. an irregular blotch on the posterior margin of the third to the anterior margin of the fourth supralabial - see also Harvey 1999 : 401, fig. 7 in Lema and Renner 2016 , and Figs 4-5 below), a blackish blotch on rostral scale adjacent to anterior border of prefrontals (vs. blotch absent), darker parietals (vs. light blotches irregularly distributed on parietal scales) and a distinct number of maxillary teeth (4 + 2 vs. 3 + 2). Further, it differs from A. nigroterminata in having its background color beige (in living specimens) (vs. background color red-orange); paravertebrals distinct (vs. paravertebrals indistinct); first and fifth stripes wider, covering upper half of third and lower half of fourth row on each side (vs. first and fifth stripes also wider, but covering first and about 50% of fourth row of scales on each side). Apostolepis kikoi sp. nov. can be distinguished from A. underwoodi by having a vertebral stripe one scale wide (vs. vertebral stripe narrow, running on the medial line of each vertebral scale); paravertebral stripes covering sixth row on each side (vs. paravertebral stripes covering half of fifth and half of sixth rows). Finally, Apostolepis kikoi sp. nov. is distinguished from A. lineata , A. nelsonjorgei and A. serrana by having its fourth supralabial scale rectangular (vs. triangular in A. nelsonjorgei ), 4-6 supralabials contacting parietals (vs. 5-6 contacting parietals in A. nelsonjorgei ), terminal scale black dorsolaterally (vs. terminal scale entirely white in A. lineata and A. nelsonjorgei ), anterior chinshields longer than posterior (vs. anterior and posterior chinshields of about the same size in A. nelsonjorgei ), the presence of a white nuchal collar (vs. white nuchal collar absent in A. lineata and A. serrana ) and a distinct number of subcaudals (27-30 vs. 40-46 in A. nelsonjorgei and 33 in A. serrana ). Description of holotype ( Figs 1 - 3 ). A small female, possibly juvenile, SVL 262 mm, TL 26 mm (9.92% of SVL). Body subcylindrical. Tail very short with tip conical and laterally compressed. Terminal scale pointed. Head slightly distinct from neck, narrower than diameter of midbody. Head length from quadrate-articular jaw joint to tip of snout (in lateral view) 6.56 mm (2.5% of SVL), 4 mm at widest point (60.9% of head length). Snout rounded in dorsal and lateral views, slender and slightly projected beyond jaws; snout length from tip of snout to anterior margin of right orbit 2.37 mm (36.12% of head length ). Rostral wider than high, visible from above; portion of rostral trapezoidal from above, 1.52 mm wide; length of rostral visible from above 0.63 mm. Prefrontal-rostral contact broadly separating nasals. Suture between prefrontals 58.4% of length of frontal. Prefrontals almost as wide as long. Frontal hexagonal, longer (2.09 mm) than wide (1.42 mm at widest point). Nostril-orbit distance in right lateral view 1.23 mm (16.49% of head length). Interorbital width (shortest distance between dorsomedial margins of orbits) 2.51 mm (66.57% of head width). Nasal entire, contacting preocular. Preocular rectangular, about 34% of length of nasal. Loreal absent. Parietal almost twice as long (3.08 mm) as wide (1.44 mm at widest point); suture between parietals 1.97 mm; length of suture between parietals 94.2% length of frontal. Supraoculars subretangular, almost twice as long as wide. Maximum diameter of eye 0.70 mm, pupils round. Postoculars pentagonal, higher than wide. Temporals absent. Five occipitals wider than long; median occipital positioned between posterior tips of parietals, smaller than adjacent vertebral; two pairs of lateral occipitals twice as large as dorsals, contacting posterior edges of sixth supralabials. Supralabials 6/6, 1 contacting rostral, 1-2 contacting nasal, 2 contacting nasal and preocular, 2-3 entering orbit, 3-4 contacting single postocular, 4-6 contacting parietal. Supralabials in ascending order of size, with sixth supralabial higher and longer (1.46 mm long, 1.29 mm high) than remaining supralabials. Infralabials 7/7, 1-4 contacting anterior chinshields, 4-7 contacting posterior chinshields. Mental subtriangular, wider (1.14 mm) than long (0.82 mm), separated from anterior chinshields by contact between first infralabials. Anterior chinshields elongated, longer than posterior chinshields. Suture between chinshields 1.82 mm. Chinshields separated from ventrals by four gulars and two preventrals. Gulars in four rows between last supralabial and first preventral. Dorsal scales smooth in 15 rows at midbody. No apical pits. 205 ventrals. 26 paired subcaudals (tail complete). Anal scale divided. 4 + 2 maxillary teeth. Coloration of holotype in life ( Fig. 2 ). Head dorsally blackish with light blotches irregularly distributed on rostral, nasal, prefrontal, frontal, supraocular and parietal scales. Suture between prefrontals brownish. Dorsally, rostral scale has a blackish blotch adjacent to anterior border of prefrontals; ventrally, rostral scale has a blackish spot in its convex portion. Posterior upper margin of first and fourth supralabials blackish. Second supralabial blackish. Anterior portion and upper half of third supralabial blackish. Blackish head cap covers upper half of fifth and sixth (except for its lower anterior portion) supralabials. Posterior portion of nasal, preocular and postocular scales blackish. Infralabials and chinshields cream, except for small blackish spots on third and fourth scales. Brown pigment of throat region restricted to more laterally positioned gular scales (i.e., evidencing an incomplete gular band). White nuchal collar two and half to three dorsals long. Black nuchal collar absent. Background color beige, with five black stripes; vertebral stripe one scale wide; paravertebral stripes covering sixth and tenth rows, less distinct than remaining dorsal stripes; first and fifth stripes wider, covering upper half of third and lower half of fourth row on each side. Paraventral sides and venter unblemished. Black band on tail extends for nine scales dorsally; seven subcaudals are black. Terminal scale with band of melanophores dorsolaterally and entirely clear ventrally. Color of holotype in preservative ( Fig. 3 ). Head dorsally brownish with light blotches irregularly distributed on rostral, nasal, prefrontal, frontal, supraocular and parietal scales. Dorsally, rostral scale brown pigmented; this blotch covers anterior border of prefrontals; ventrally, rostral scale with brownish spot on its convex portion. Anterior and posterior parts of prefrontals brown. Anterior and posterior parts of suture between prefrontals brown. Posterior upper margin of first and fourth supralabials brown. Second supralabial brown. Anterior portion and upper half of third supralabial brown. Brown head cap covers upper half of fifth and sixth (except for its lower anterior portion) supralabials. Posterior portion of nasal, preocular and postocular scales brown. Infralabials and chinshields cream, except for small brown spots on third and fourth scales. Brown pigment of throat region restricted to more laterally positioned gular scales (i.e., evidencing an incomplete gular band). White nuchal collar two and half to three dorsals long. Black nuchal collar absent. Background color light brown, with five brown stripes; paravertebral stripes light brown and less distinct than remaining dorsal stripes. Venter immaculate. Dark brown band on tail extends for nine scales dorsally; seven subcaudals are dark brown. Terminal scale with band of melanophores dorsolaterally and entirely clear ventrally. Variation. Measurements and morphological variation are summarized in Table 1 . Largest male (MCP 14524) SVL 310 mm, TL 38 mm. Largest female (MCP 12096, holotype) SVL 262 mm, TL 26 mm. Smallest specimen measured is female (MCP 11372) with SVL 152 mm and TL 16 mm. Fifth and sixth supralabials in contact with parietal in MCP 14524, MCP 14525 and UFMT-R 1933; fourth, fifth and sixth supralabials in contact with parietal in MCP 11372. Head length ranges from 7.44-8.01 mm ( x- = 7.7 +/- 0.3) in males, 5.83-6.56 mm ( x- = 6.2 +/- 0.5) in females. Number of ventral scales ranges from 203-209 ( x- = 206.7 +/- 3.2, n = 3) in males and 205-207 ( x- = 206 +/- 1.4, n = 2) in females. Subcaudals range from 29-30 ( x- = 29.3 +/- 0.6, n = 3) in males and 26-28 in females ( x- = 27 +/- 1.4, n = 2). Preventrals 1 (n = 3) or 2 (n = 2). First infralabial fused with first chinshield in MCP 14524 on right side. Suture between prefrontals entirely pigmented by brown blotch in UFMT-R 1933. White nuchal collar two and a half to three dorsals long (MCP 11372, MCP 14524 and UFMT-R 1933) or two to three dorsals long (MCP 14525). Black nuchal collar vestigial in MCP 11372, MCP 14524 and MCP 14525 and apparent in UFMT 1933. First dorsal stripe, vertebral stripe and fifth dorsal stripe brown; paravertebral stripes light brown and less distinct than remaining dorsal stripes in all specimens. Caudal band extends for length of 7-10 scales dorsally and 3-8 subcaudals. In MCP 11372 fourth supralabial on each side entirely clear; it has a narrow, vestigial band of melanophores on the anterior portion of the terminal scale (dorsally) and small black blotches can be observed on its 22nd, 24th, 25th, 26th and 27th pair of subcaudal scales. Terminal scale with band of melanophores dorsolaterally and entirely clear ventrally in MCP 11372, MCP 14524 and UFMT-R 1933. Hemipenial morphology ( Figs 5 - 7 ). Retracted organs extend for length of nine subcaudals. Everted hemipenes subcylindrical, unilobed, unicapitate and noncalyculate. Basal region on sulcate side bears numerous spines of similar size. Several moderate-sized spines present on lateral region of hemipenial body in its absulcate side. Sulcus spermaticus bifurcates about two-thirds before end of organ; branches - which extend centrolineally - reach distal tip of lobe. Basal region on absulcate side also bears numerous spines of similar size, but these are abruptly replaced by two larger spines in middle region of hemipenial body, larger than those disposed on lateral region of hemipenial body. Capitulum confined to sulcate side. Distal region of absulcate side bears transverse papillate flounces, without calyces, whereas a small number of papillae are concentrated above flounces. Distribution ( Fig. 8 ). All individuals of the type series of Apostolepis kikoi sp. nov. were obtained in the area presently occupied by the Manso multi-use reservoir and hydroelectrical power plant (APM Manso), in most part situated in the municipality of Chapada dos Guimaraes , Mato Grosso, Brazil. With nearly 428 km2, the reservoir of APM Manso inundated - from December 1999 - many different Cerrado physiognomies (see Conceicao 2000 ), established over terrains up to 287 m.a.s.l. from the confluence of the Casca and Manso rivers (approximately at 14°52'S , 55°48'W ) upwards. Manso is the main tributary to the Cuiaba River, a major tributary of the left bank of the Paraguay River. The local climate is generally hot and semi-humid (classified as "Aw climate" in the Koeppen's climate classification map for Brazil, see Alvares et al. 2014 ), with a well-marked seasonality and rains concentrated in the summer, from October/November to April/May. Mean annual precipitation is 1350 mm; mean annual temperature is around 26 °C. There are four to five months of drought (May to September), and relative air humidity may drop to less than 30% from July to September ( Struessmann and Mott 2009 ). The Manso River basin is included in the morphostructural domain of the Parana River sedimentary basin. It is cut across two distinct lithostratigraphic units in nonconformity: the Permeable Mesozoic sandstones belonging to the Sao Bento (or Botucatu) Group and the inclined Precambrian phyllites and gneisses of the Cuiaba Group, frequently with a lateritic horizon at or near the surface ( Barros et al. 1982 ; Vieira Jr. et al. 2012 ). Figure 1. Line drawings from holotype of Apostolepis kikoi sp. nov. (MCP 12096) from the reservoir of Manso hydroelectrical power plant, Chapada dos Guimaraes , Mato Grosso, Brazil. Scale bar: 2 mm (Eduardo Melloni Lucchesi del.). Figure 2-5. ( 2) Holotype of Apostolepis kikoi sp. nov. (MCP 12096) from the reservoir of Manso hydroelectrical power plant, Chapada dos Guimaraes , Mato Grosso, Brazil (photo by Marcos Di Bernardo). (3) Lateral view of holotype (preserved) of Apostolepis kikoi sp. nov. (MCP 12096) from the reservoir of Manso hydroelectrical power plant, Chapada dos Guimaraes , Mato Grosso, Brazil (photo by Marcos Di Bernardo). Dorsal (4) and lateral (5) views of the specimen of Apostolepis nigroterminata (UFAC 504) from Campus Universitario da Universidade Federal do Acre, Rio Branco, Acre, Brazil (photo by Danyella Paiva da Silva). Figures 6-7. Hemipenis of Apostolepis kikoi sp. nov. (MCP 14524) from the reservoir of Manso hydroelectrical power plant, Chapada dos Guimaraes , Mato Grosso, Brazil, showing sulcate (6) and absulcate (7) sides (photo by Fernanda Martins dos Santos). Scale bar: 1 mm. Table 1. Variation in characters of the type series of Apostolepis kikoi sp. nov. Supralabials (SL), infralabials (IL), snout-vent length (SVL), tail length (TL), snout length (SL), head length (HL), preventrals (PV), ventrals (VE), subcaudals (SC), number of dorsal scales of black caudal band (DBB) and number of subcaudal scales of black caudal band (SBB). Measurements in mm.

-	MCP 11372	MCP 12096 (holotype)	MCP 14524	MCP 14525	UFMT-R 1933
Sex	F	F	M	M	M
SL	6/6	6/6	6/6	6/6	6/6
IL	7/7	7/7	7/7	7/7	7/7
SVL	152	262	310	304	263
TL	16	26	38	37	35
SL	1.74	2.37	2.70	2.61	2.46
HL	5.83	6.56	8.01	7.56	7.44
PV + VE	2 + 207	1 + 205	1 + 209	2 + 208	2 + 203
SC	28	26	29	29	30
DBB	10	9	7	8	9
SBB	6	8	5	3	5

Figure 8. Map of South America, illustrating the type locality (star) of Apostolepis kikoi sp. nov. - reservoir of Manso hydroelectrical power plant - Chapada dos Guimaraes ( 15°27 '39" S , 55°45 '00" W , 811 m a.s.l.), Mato Grosso, Brazil, and locality records for Apostolepis nigroterminata (circles) in the Brazilian state of Acre and Peru based upon the material examined. Square indicates the type locality of Apostolepis nigroterminata . Etymology. The specific epithet honors Francisco Luis Franco ( "Kiko" ), a specialist in Brazilian snakes, as a tribute to his relentless friendship, dedication and enthusiasm as curator of Herpetological Collection Alphonse Richard Hoge of Instituto Butantan, Sao Paulo, Brazil (partially and tragically destroyed by fire on 15 May 2010). Remarks. MCP 12096 was selected as the holotype because the general color in life of A. kikoi was described from the live holotype before it was euthanized. Apostolepis nigroterminata was described by Boulenger (1896) after a single specimen from "Cayaria" (= Callaria, Departamento de Ucayali), eastern Peru. Harvey's (1999) placement of A. borellii in the synonymy of A. nigroterminata resulted in the inclusion of the latter species in a former Brazilian list of reptiles (see Costa and Bernils 2015 ). However, it should be noted that Harvey's (1999) nomenclatural act was based on the analysis of the holotype of A . borellii (a specimen from Urucum massif, Mato Grosso do Sul, Brazil), one specimen collected at the confluence of Rio Araguaia and Tapirape , Tapirape Village, Mato Grosso do Sul (AMNH 87942) - that was subsequently re-identified as A. phillipsi by Martins and Lema (2015) - and several other specimens from Bolivia and Peru. Recently, Lema and Renner (2016) removed A. borellii from the synonymy of A. nigroterminata and restricted the distribution of the latter to some localities in Peru. However, these authors also listed a specimen of A. nigroterminata from "Brazil: Acre: Rio Branco" in Appendix as "UFAC w/n". The species was also included in an updated list of Brazilian reptiles and referred to occur in the states of Acre, Mato Grosso, and Para ( Costa and Bernils 2018 ). Besides the unvouchered mention to Acre in Lema and Renner (2016) , Lema et al. (2017) presented a picture of a specimen from Acre without a clear locality description or voucher number, which also provides little evidence for the occurrence of this species in Brazil. The record for Para is also unvouchered ( Maschio et al. 2012 ), while the specimen (UFMT 10672) from Nobres, Mato Grosso, referred to the species by Santos et al. (2011) was examined and is reidentified here as Apostolepis sp. Therefore, we argue that the specimens examined herein (UFAC 383, UFAC 397, UFAC 504) represent not only three locality records of A. nigroterminata for the state of Acre, Brazil, but also the first documented record of the species for the country. The specimens collected in Rio Branco (UFAC 397, UFAC 504) extend the geographic distribution of A. nigroterminata about 760 km northeastward from Callaria. In particular, the specimen depicted in Figs 4-5 matches the original description of Boulenger (1896) and that given by Lema and Renner (2016) in most details of scalation and color pattern of this species (see Table 2 ). Table 2. Variation in characters of five specimens of Apostolepis nigroterminata . Snout-vent length (SVL), tail length (TL), ventrals (VE) and subcaudals (SC). The UFAC specimens presumably are males; the FMNH 39646 specimen is a male. *Holotype, data from Lema and Renner (2016 : 67). Measurements in mm.

-	BMNH 1946.1.9.77*	FMNH 39646	UFAC 383	UFAC 397	UFAC 504
Sex	M	M	M	M	M
SVL	199	301	210	261	238
TL	18	26	17	19	18
VE	213	234	209	216	219
SC	26	28	24	27	25

Two other species of Apostolepis - A. lineata and A. vittata - were also described from Chapada dos Guimaraes ( Cope 1887 ). Although the only existing syntype of A. lineata is in very bad condition, Harvey's (1999) redescription is sufficiently complete to allow it to be unambiguously distinguished from Apostolepis kikoi sp. nov. Together with A. assimilis ( Fig. 9 ) and A. vittata ( Fig. 10 ), the description of Apostolepis kikoi sp. nov. increases the number of species of Apostolepis reported to occur sympatrically in the Manso reservoir area to three. An attempt was made to identify all the specimens of Apostolepis kikoi sp. nov. using Nogueira et al.'s (2012) key. However, the specimens could not be characterized beyond couplet 1, because of the many overlapping characters presented in the couplets. The assignment of Apostolepis kikoi sp. nov. into a formal group should await a more comprehensive phylogenetic arrangement than is available for the genus. Figure 9. Apostolepis assimilis . A specimen deposited at the Universidade Federal de Mato Grosso (UFMT-R 11088) from district of Sao Vicente, Santo Antonio do Leverger, Mato Grosso, Brazil (photo by Christine Struessmann ). Figure 10. Apostolepis vittata . A specimen deposited at the Universidade Federal de Mato Grosso (UFMT-R 12259) from the National Park of Chapada dos Guimaraes , Cuiaba , Mato Grosso, Brazil (photo by Christine Struessmann ).