Rhinella exostosica, Ferrão & Lima & Ron & Santos & Hanken, 2020

Ferrão, Miquéias, Lima, Albertina Pimentel, Ron, Santiago, Santos, Sueny Paloma dos & Hanken, James, 2020, New Species of Leaf-litter Toad of the Rhinella margaritifera Species Group (Anura: Bufonidae) from Amazonia, Copeia 108 (4), pp. 967-986 : 970-980

Rhinella exostosica , new species urn:lsid:zoobank.org:act:FDE2D615-4BA5-4A19-969D-EF1B- BA5BFE09

Figures 1–3 , 4A–D , 5A–B , 6–9 , 10A–C , 11 ; Tables 1–2

Bufo typhonius : Duellman and Salas (1991); Duellman (2005). Bufo sp. (margaritifer complex): Moravec and Aparicio (2000). Bufo sp. II (margaritifer complex): Moravec and Aparicio

(2005). Rhinella cf. margaritifera 5: Pramuk (2006); Moravec et al.

(2014). Rhinella margaritifera ( ¼ typhonius ): Mendelson et al. (2011). Rhinella cf. margaritifera : Moravec et al. (2014); Santos et al.

(2015); Cusi et al. (2017).

Rhinella sp. ( Rhinella margaritifera complex): Moravec et al. (2016).

Rhinella margaritifera : Ávila et al. (2018).

Holotype.— INPAH 41323 (field number APL 19973), adult male from the Jac´ı Direito Sampling Module, 09827 0 44 00 S , Paratopotypes.— Five adult specimens, all collected by A. P. Lima, same locality as the holotype: INPAH 41317 (field number APL 17632), female, 25 May 2011 ; INPAH 41321 (field number APL 19688), male, 25 May 2013 ; INPAH 41322 (field number APL 19697), female, 25 March 2013 ; INPAH 41326–27 (field numbers APL 21152–53), male and female (respectively), 11 November 2014 .

Paratypes.— 15 adult specimens. 7 specimens, Jac´ı Novo Sampling Module, 09824 0 45 00 S, 64826 0 33 00 W, all collected by A. P. Lima: INPAH 41318–19 (field numbers APL 19409–10), females, 13 February 2013 ; INPAH 41320 (field numbers APL 19650), female, 22 March 2013 ; INPAH 41324 (field number APL 20029), male, 13 November 2013 ; INPAH 41325 (field number APL 21133), male, 8 November 2014 ; INPAH 41328– 29 (field numbers APL 21154–55), females, 12 November 2014 . 5 specimens, Morrinhos Sampling Module, 09804 0 34 00 S, 64814 0 46 00 W: INPAH 41312 (field number APL 15907), female, A. P. Lima, 9 November 2010 ; INPAH 41313 (field number APL 16422), male, R. Fraga , 13 January 2011 ; INPAH 41314 (field number APL 16468), female, R. Fraga , 14 January 2011 ; INPAH 41315–16 (field numbers APL 16473–74), male and female (respectively), A. P. Lima, 14 January 2011 . 3 specimens, Três Praias Camp, 09827 0 11 00 S, 64825 0 04 00 W, all collected by A. P. Lima and M. Prestes: INPAH 41330 (field number APL 21984), 11 January 2017 ; INPAH 41331–32 (field numbers APL 22030–31), males, 31 January 2017 .

Referred material.— Six adult specimens. Bolivia: NMP6 V 70687–8 , 2 males, 5–6 km NE of Riberalta , Beni, 11800 0 S, 66805 0 W, J. Moravec and J. Aparicio, 26–27 November 1999; CBF 5800 , male, Bolpebra, 10857 0 S, 69834 0 W, Nicolas Suaréz , Pando, J. Moravec, and J. Aparicio, between 30 January and 3 February 2005. Peru: NMP6 V 74915 , male, Regional Conservation Imiría , 17.4 km S of Masisea, 08836 0 18 00 S, 74818 0 23 00 W, Ucayali, J. Moravec and I. A. Tuanama, 27 September 2011; KU 215145–46 , 2 unsexed specimens, 15 km E Puerto Maldonado , 12832 0 38 00 S, 69803 0 23 00 W, Cusco Amazónico, Madre de Dios, W. E. Duellman, 1989 .

Etymology.— The specific epithet exostosica is derived from the Latin ‘‘exostosis’’ and a reference to the strongly developed bony protrusion at the angle of the jaw of the new species.

Diagnosis.— Rhinella exostosica is a large-sized species of the R. margaritifera species group ( Fig. 1 ; Pramuk, 2006). The species is diagnosed by the following combination of characters: 1) SVL 63.4–84.7 mm in females, 56.3–72.3 mm in males; 2) snout subacuminate in dorsal view; 3) snout lacks pronounced fleshy proboscis; 4) upper jaw curved upward in lateral view; 5) strongly developed bony protrusion at angle of jaw; 6) tympanic membrane and tympanic annulus present and evident; 7) supratympanic crests well developed; 8) proximal portion of supratympanic crest same height or shorter than parotoid gland in lateral view; 9) canthal crests poorly developed; 10) large parotoid glands; 11) 3–6 dorsal vertebral apophyses; 12) divided distal subarticular tubercle on finger III; 13) relative length of fingers III. IV. II. I; 14) skin on dorsum granulated with conical tubercles; 15) advertisement call duration 295–394 ms (339626 ms), composed of groups of 7–9 (7.860.6) pulsed notes, with the last note consisting of 2–4 (2.960.8) pulses, and a dominant frequency of 1,012 –1,163 Hz (1,081663 Hz).

Comparisons.— We compare the new species with all nominal species of the Rhinella margaritifera species group, with particular attention to R. martyi and R. paraguayensis due to their morphological similarity and tentative phylogenetic placement, respectively. Rhinella exostosica can be distinguished from all members of the R. margaritifera species group by its combination of a strongly developed bony protrusion at the jaw angle, supratympanic crest shorter than or the same height as the parotoid gland in lateral view, and bifid distal subarticular tubercle on finger III. Diagnostic characters of compared species are enclosed in parentheses or brackets unless stated otherwise.

Maximum snout–vent length (SVL) of male Rhinella exostosica is 72.3 mm, which is much larger than R. acutirostris (holotype, 47 mm), R. alata (43.2 mm; Santos et al., 2015), R. castaneotica (41.9 mm; Caldwell, 1991), R. gildae (64.5 mm; Ávila et al., 2018), R. hoogmoedi (52 mm; Caramaschi and Pombal, 2006), R. lescurei (34.664.3 mm; Fouquet et al., 2007b), R. magnussoni (45.3 mm; Lima et al., 2007), R. parecis (53.5 mm; Ávila et al., 2020), R. proboscidea (54 mm; Lima et al., 2006), R. scitula (46.1 mm; Caramaschi and Niemeyer, 2003), R. sebbeni (59.7 mm; Vaz-Silva et al., 2015), and R. stanlaii (54 mm; Lötters and Köhler, 2000).

Rhinella exostosica differs from R. roqueana by having the upper jaw curved upward in lateral view and a divided distal subarticular tubercle on finger III (jaw straight and single distal subarticular tubercle; Melin, 1941); from R. yunga by having a tympanic membrane and annulus (absent), a divided distal subarticular tubercle on finger III (single distal subarticular tubercle), and dorsal vertebral apophyses (absent; Moravec et al., 2014); from R. dapsilis by lacking a pronounced fleshy proboscis on the snout (present), dorsal skin granulated with conical tubercles (dorsum smooth), supratympanic crests well developed (poorly developed), and maximum SVL 84.7 mm in females (77 mm; Myers and Carvalho, 1945; Hoogmoed, 1986); from R. sclerocephala by having a divided distal subarticular tubercle on finger III (single), the upper jaw curved upward in lateral view (straight), and maximum SVL 72.3 mm in males and 84.7 mm females (67.3 mm and 77.4 mm in males and females, respectively; Mijares-Urrutia and Arends, 2001); from the holotype of R. margaritifera by its subacuminate snout in dorsal view (truncate), proximal portion of the supratympanic crest the same height or shorter than the parotoid gland in lateral view (supratympanic crest higher than the parotoid gland; Fig. 4A, C, H ), and a strongly developed bony protrusion at the angle of the jaw (protrusion moderately developed).

Rhinella exostosica differs from R. martyi in relative finger length III. IV. II. I (III. I. II. IV), the proximal portion of the supratympanic crest the same height or shorter than the parotoid gland in lateral view (higher than parotoid gland; Fig. 4A, C, E ), parotoid gland large (small) and thenar tubercle ovoid (round). Males of R. exostosica present wider IOD than males of R. martyi (IOD/SVL ¼ 0.1860.01 in R. exostosica ; IOD/SVL ¼ 0.1460.01 in R. martyi ). The advertisement call of R. exostosica is emitted in groups of 7.860.6 pulsed notes with a call duration of 339626 ms, and the last note is composed of 2–4 pulses (maximum 6 notes per call, call duration 295613 ms, and the last note has up to 6 pulses; Fouquet et al., 2007b).

Rhinella exostosica differs from R. paraguayensis sensu stricto by having the proximal portion of the supratympanic crest the same height or shorter than the upper limit of the parotoid gland in lateral view (higher; Fig. 4A, C, G ), maximum SVL 72.3 mm in males and 84.7 mm in females (52.6 mm and 53.3 mm in males and females, respectively), snout subacuminate in dorsal view (rounded), parotoid glands large (small), vertebral apophyses present (absent), strongly developed bony protrusion at the angle of the jaw (poorly developed and straight), and an advertisement call with mean dominant frequency of 1,081663 Hz (1,439671 Hz; Ávila et al., 2010). Rhinella exostosica is readily distinguished from R. cf. paraguayensis from Bolivian lowland (sensu Jansen et al., 2011) by having a strongly developed bony protrusion at the angle of the jaw (poorly developed), dorsal vertebral apophyses (absent), and snout subacuminate in dorsal view (rounded; specimens reported in Jansen et al., 2011).

The new species is easily distinguished from Rhinella aff. margaritifera from the west bank of the upper Madeira River ( Fig. 5 ) by having the proximal portion of the supratympanic crest the same height or shorter than the parotoid gland in lateral view (higher), a divided distal subarticular tubercle on finger III (single), a strongly developed bony protrusion at the angle of the jaw (poorly developed), canthal crests poorly developed (well developed), and parotoid glands large (small).

Description of holotype.— INPAH 41323 (field number APL 19973), adult male, SVL 68.7 mm ( Figs. 2 , 3 , 4A–B , 6 ). Head wider than long (HW/HL ¼ 1.1); HL 35% of SVL. Snout protruding in lateral view and subacuminate in dorsal view; dorsal surface slightly concave; nasal opening directed dorsolaterally; internarial distance 38% of interorbital distance. Canthus rostralis delimited by a poorly developed canthal crest; loreal region concave. Eye–nostril distance 114% of eye diameter, 150% of horizontal tympanum diameter, and 142% of upper eyelid width. Eyes protuberant, wider than tympanum (EL/TYMH ¼ 1.45; EL/TYMV ¼ 1.30); eye diameter 137% of UEW. Absence of projections on upper eyelid; UEW 44% of IOD. A strongly developed and curved bony protrusion at the angle of the jaw is visible in dorsal, ventral, and lateral views; distance between bony protrusions equals 116% of HW. Preorbital and canthal crests poorly developed; supraorbital, supratympanic, and parietal crests well developed; proximal portion of supratympanic crest shorter than the parotoid gland in lateral view; distance between supratympanic crests slightly larger than head width (POCD/HW ¼ 1.01) but smaller than distance between bony protrusions (POCD/BPD ¼ 0.87). Tympanum large, vertically oval (TYMV/TYMPH ¼ 1.12), with a distinct annulus. Parotoid gland well developed, subtriangular in dorsal view and elliptic in lateral view; in dorsal view, twice as long as wide (PGL/PGW ¼ 2.04); parotoid gland length 240% of POCL. Parotoid gland bordered by a line of small conical tubercles; a lateral line of large conical tubercles extends from the proximal corner of the parotoid gland to the groin. Two vertebral apophyses expanded dorsally. External choanae small, oval, and laterally positioned; separated by approximately four times their width. Tongue oval, four times longer than wide. Vocal slits present; vocal sac single and subgular.

Anterior limbs robust; forearm as robust as upper arm; a line of small conical tubercles borders the forearm. Hand long; HAND3 90% of UAL; relative lengths of fingers III. IV. II. I ( Fig. 3C ); lateral fringes developed, especially on fingers I, II, and IV, with small conical tubercles extending from the outer lateral of finger I to the external lateral of finger IV. Fingertips poorly expanded; palmar tubercle large, tear-shaped, flat and smooth; thenar tubercle ovoid, pronounced, approximately 65% of palmar tubercle length. Subarticular tubercles developed on all fingers, single on fingers I, II, and IV, divided on distal articulation of finger III, single on proximal. Supernumerary tubercles conical, varied in size, and irregularly distributed.

Hind limbs robust. Thigh longer than tibia (THL/TL ¼ 0.91); thigh length 47% of SVL, tibia length 42% of SVL. Tarsus length 77% of FOOT4 and 27% of SVL. Foot relatively short; FOOT4 35% of SVL. Relative lengths of toes IV. III. V. II. I; lateral fringes present and developed on toes, with small conical tubercles extending from the distal portion of the inner metatarsal tubercle on toe I to external toe V; toes with moderate webbing, webbing formula I 1–2 þ II 1–2 1/2 III 1–3 1/2 IV 3 1/2 –1 1/2 V. Subarticular tubercles conical, single on all toes; outer metatarsal tubercle small and subconical; inner metatarsal tubercle large and ovoid, approximately twice the size of the outer metatarsal tubercle. Supernumerary tubercles present, varied in size, and irregularly arranged.

Skin granulated with conical and flat tubercles of varied size irregularly distributed on dorsum and flanks, granulated with small conical tubercles on limbs. Tubercles absent on interorbital region. Upper eyelids with small conical tubercles. Tiny conical tubercles on subocular region, lips, and on bony protrusions at jaw angle. Ventral surface granulated.

In life, dorsal surface of body and limbs brown; dorsal surface of head orangish brown; lateral surfaces of head brownish orange; crests orange; lateral line of conical tubercles orange; lateral fringes on toes and fingers orangish cream ( Fig. 6 ). Iris tan without black reticulations. Flanks orange tan. Chin, throat, and chest orange with inconspicuous light gray and cream blotches; ventral surface of arms cream; belly and ventral surface of thigh with conspicuous dark gray blotches irregularly distributed; ventral surface of tarsus dark gray. Palmar and plantar surfaces dark gray; thenar and palmar tubercles cream; subarticular and large supernumerary tubercles cream on hand; inner and outer metatarsal tubercles cream; subarticular and supernumerary tubercles dark brown on foot.

In preservative ( Fig. 3 , 4A, B ), the brown dorsal coloration becomes gray; dorsal surface of head and snout brown; lateral surface of head brownish cream ( Fig. 3A, B ). Flanks cream; lateral line of tubercles light gray. Lateral fringes on toes and fingers cream. Chin, throat, chest, and ventral surfaces of thigh cream with gray and light gray blotches; ventral surface of arms cream. Palmar and plantar surfaces dark gray; tubercles on hand cream to light gray; tubercles on foot light gray.

Color and morphological variation.— In preservative, dorsal color varies from grayish cream (44% of specimens), dark brown (25%), orangish cream (19%), and dark gray (6%) to light gray (6%). Dorsolateral line of tubercles is grayish cream (44%), light brown (37%), dark brown (13%), or gray (6%). A dead-leaf pattern with dark blotches is present in 69% of specimens ( Fig. 7A–C ) but faded or inconspicuous in the rest ( Fig. 7D–F ). A cream-colored vertebral line extending from the snout to the urostyle is present in 81% of specimens. Dorsal surface of hind limbs with dark blotches or bars is seen in 81% of specimens. Ventral surfaces of chin, throat, chest, and thighs are colored by different shades of cream with light to dark gray blotches (which range from scarce to densely concentrated) in 94% of specimens ( Fig. 8 ), but these surfaces are light gray with cream and gray blotches in the rest ( Fig. 8 ). Although coloration in life is more vivid, the basic pattern is generally retained in preservative ( Fig. 9 ).

Morphologically, all of the type series of Rhinella exostosica resembles the holotype, although the species exhibits sexual dimorphism in several characters ( Table 2). Females are larger than males ( SVL, t ¼ –3.9643, df ¼ 11.224, P ¼ 0.002) and have more vertebral apophyses ( APO, t ¼ –2.6784, df ¼ 13.705, P ¼ 0.018) and a longer finger I ( HAND1 / SVL, t ¼ –2.6222, df ¼ 8.7776, P ¼ 0.028). Conversely, males have longer supraocular crests ( SOCL / SVL, t ¼ 2.5907, df ¼ 13.766, P ¼ 0.021), wider tympanums ( TYMH / SVL, t ¼ 2.3596, df ¼ 7.9322, P ¼ 0.046), larger eyes ( EL / SVL, t ¼ 2.5296, df ¼ 9.0854, P ¼ 0.032), and wider upper eyelids ( UEW / SVL, t ¼ 3.2986, df ¼ 13.029, P ¼ 0.005) than females .

Advertisement call.— The advertisement call of Rhinella exostosica is emitted in a series of 965 calls (5–12, n ¼ 7) with a call duration of 339626 ms (295–394 ms; n ¼ 16) and an inter-call interval of 4836209 ms (254–980 ms; n ¼ 16; Fig. 10A–C ). Calls are composed of 7.860.6 pulsed notes (7–9, n ¼ 16) with a note duration of 1868 ms (7–47 ms; n ¼ 48) and an inter-note interval of 3664 ms (28–45 ms; n ¼ 32). Overall, notes are formed by 2.360.8 pulses (1–4 pulses, n ¼ 48) with a pulse duration of 861 ms (7–12 ms; n ¼ 48). The number of pulses per note varies during the call; the first note (2.360.4 pulses [2–3 pulses, n ¼ 16]) and the last note (2.960.8 pulses [2–4 pulses, n ¼ 16]) usually contain more pulses than notes in the middle of the call (1.660.5 pulses [1–2 pulses, n ¼ 16]). Calls have a dominant frequency of 1,081663 Hz (1,012 – 1,163 Hz, n ¼ 16) and a bandwidth of 423617 Hz (409–452 Hz, n ¼ 16).

Tadpoles.— Tadpoles of Rhinella exostosica were described by Duellman (2005).

Distribution and natural history.— Rhinella exostosica inhabits forests of the eastern portion of the upper Madeira Basin in Brazil, Bolivia, and Peru ( Fig. 11 ). In Brazil, males and females of R. exostosica are active during the day on leaf litter within open lowland forest. At night, specimens are usually found on green leaves of shrubs or at the base of small trunks up to ~ 1 m high. Calling males were unsuccessfully sought close to small streams and temporary puddles within disturbed and intact forests at the sampling sites in Brazil. After seven years of field surveys, we finally came upon an explosive breeding event on 31 January 2017 in a bay of the Jací-Paraná River during heavy rain at the Três Praias Camp. Males began calling at ~ 1600 h sitting alongside the bank river or while floating within shallow water. We also found males calling on dense stands of macrophytes floating above deeper waters.

In Bolivia, Rhinella exostosica inhabits the forest of the Madre de Dios Basin and Acre Basin. In Riberalta (Beni Department), the species was recorded in a terra firme forest on the east banks of the Beni River close to the junction with the Madre de Dios River ( Moravec and Aparicio, 2000). In Bolpebra (Pando Department), a calling male of R. exostosica was collected by Moravec and Aparicio (2005) in a temporary pond surrounded by secondary forests along the east bank of the Acre River.

In Peru, Duellman (2005) recorded 450 individuals of Rhinella exostosica along a trail paralleling the Madama Stream, close to the junction with the Madre de Dios River (Cusco Amazónico, Madre de Dios Department). All individuals were found in a terra firme forest. Explosive breeding events also occur after heavy rains between November and February. Most calling males were found sitting on shallow backwaters or adjacent banks of the Madama Stream .

Figures

Fig. 1. Phylogeny of species of the Rhinella margaritifera species group based on one nuclear and three mitochondrial genes (2,997 bp) and reconstructed through Bayesian inference. Posterior probabilities above 0.80 are shown close to nodes and those greater than or equal to 0.95 are represented with an asterisk. Scale bar represents substitutions/site. Photographs by Santiago Ron, bioweb.bio (R. dapsilis, R. margaritifera), Anthony S. Ferreira (R. proboscidea) and Albertina P. Lima (Rhinella exostosica, new species). See Data Accessibility for tree file.

Fig. 2. Dorsal (A) and ventral (B) views of the male holotype of Rhinella exostosica, new species, INPAH 41323, SVL 68.7 mm.

Fig. 3. Dorsolateral (A) and lateral (B) views of the head, and ventral views of the hand (C) and foot (D) of the male holotype of Rhinella exostosica, new species, INPAH 41323. Scale bars, 5 mm.

Fig. 4. Lateral and dorsolateral views of the head of Rhinella exostosica, new species (A–D), R. martyi (E, F), R. paraguayensis (G), and R. margaritifera (H). (A, B) Male holotype, INPAH 41323, SVL 68.7 mm. (C, D) Female, INPAH 41320, SVL 77.1 mm. (E, F) Female holotype, MNHN 2006.2601, SVL 66.5 mm. (G) Male holotype, UFMT 7430, SVL 50.4 mm. (H) Holotype, ZISP 257.1. Photographs by Miquéias Ferrão (A–D), Fréderic Braux, MNHN (E, F), Robson Ávila (G), and Konstantin D. Milto (H).

Fig. 5. Dorsolateral (above) and dorsal (below) views of Rhinella exostosica, new species (A, B) and R. aff. margaritifera (C, D) from the upper Madeira River, municipality of Porto Velho, Rondônia, Brazil. (A) Adult female, INPAH 41322. (B) Adult male, INPAH 41321. (C) Adult female, INPAH 41333. (D) Adult male, INPAH 41334. Localities: A, B, Jac´ı Direito Sampling Module; C, D, west bank of the upper Madeira River. Photographs by Albertina P. Lima.

Fig. 6. Coloration in life of the adult male holotype of Rhinella exostosica, new species, INPAH 41323 (field number APL 19973). Dorsolateral (A) and ventral (B) views. Photographs by Albertina P. Lima.

Fig. 7. Variation in dorsal coloration of preserved Rhinella exostosica, new species, from Brazil. (A) Adult female, INPAH 41320, SVL 77.1 mm. (B) Adult male, INPAH 41321, SVL 56.7 mm. (C) Adult male, INPAH 41327, SVL 84.4 mm. (D) Adult male, INPAH 41313, SVL 72.3 mm. (E) Adult female, INPAH 41312, SVL 84.2 mm. (F) Adult female, INPAH 41329, SVL 72.1 mm.

Fig. 8. Variation in ventral coloration of preserved Rhinella exostosica, new species. (A) Adult female, INPAH 41314, SVL 83.4 mm. (B) Adult female, INPAH 41329, SVL 72.1 mm. (C) Adult female, INPAH 41327, SVL 84.4 mm. (D) Adult female, INPAH 41322, SVL 68.7 mm.

Fig. 9. Color variation in life of Rhinella exostosica, new species, from Brazil (A–D), Peru (E), and Bolivia (F). (A) Adult females: INPAH 41320 (above), SVL 77.1 mm; INPAH 41322 (below), SVL 68.7 mm; (B) Adult female, INPAH 41318, SVL 79.1 mm. (C) Adult female, INPAH 41322. (D) Adult female, INPAH 41318. (E) Subadult, NMP6V 74915. (F) Adult male, CBF 5800. Photographs by Albertina P. Lima (A–D) and Jiř´ı Moravec (E–F).

Fig. 10. Advertisement calls of Rhinella exostosica, new species (A–C), R. paraguayensis (D–E), and R. martyi (F–G). (A) Oscillogram of a series of eight-note calls of R. exostosica, new species. Detailed views of two (B) and one (C) calls of R. exostosica, new species. Detailed views of two (D) and one (E) calls of R. paraguayensis. Detailed views of two (F) and one (G) calls of R. martyi. Recordings: (A–C) INPAH 41331, Três Praias Camp, east bank of the Jací-Paraná River (affluent of the east bank of the Madeira River), Porto Velho, Rondônia, Brazil; (D–E) UFMT 2112, east bank of the Sepotuba River, Caceres, Mato Grosso, Brazil; (F–G) MNHN 2001.2006, Brownsberg Nature Park, Brokopondo district, Suriname.

Fig. 11. Geographic distribution of Rhinella exostosica, new species (stars, circles, and squares) and Rhinella aff. margaritifera BRA (triangles). Localities: 1, type locality of R. exostosica, new species, Jac´ı Direito Sampling Module; 2, Três Praias Camp; 3, Jac´ı Novo Sampling Module; 4, Morrinhos Sampling Module; 5, Riberalta; 6, Bolpebra; 7, Puerto Maldonado; 8, Masisea.