taxonID	type	description	language	source
03B98B27BC07FF84FF101457FD75FE00.taxon	description	Figs. 3, 4 A, 4 B, 5 A, 6 A, 6 C, 7 A, 7 D, 8, 9, 10 A, 10 B, 10 C, 11	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	synonymic_list	Synonymy: Pomacea sp. 2 — Ramírez et al. 2022, fig 2 B.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	materials_examined	Type material: PERU. Holotype male MUSM 5692 H. Paratypes: MUSM 5692 P, SMF 348816 (ex MUSM 5692 P), MZSP 122665 (ex MUSM 5692 P). LORETO, Maynas prov., Iquitos, flooding area of Nanay River, Nueva BellavistaNanay, 03 ° 42 ' 35.0 ” S, 73 ° 14 ' 55.2 ” W, leg. R. Chujutalli, 25 / vi / 2018, 8 specimens; MUSM 5709 P, LORETO, Ucayali prov., Lake Huimbacocha, near Santa Rosa hamlet, Cruz Muyuna Lake, Ucayali River, 6 ° 43 ' 60 ” S 75 ° 05 ' 60 ” W, leg. I. Carrillo, 17 / ii / 2018, 5 specimens; MUSM 5624 P, LORETO, Loreto prov., Saramuro Island in Marañon River, 4 ° 43 ' 18.5 ” S 74 ° 57 ' 58 ” W, A. Ampuero and V. Borda, 14 / iii / 2011, 2 specimens; MUSM 5713 P, LORETO, Loreto prov., Pahuachiro Creek near Mullaca oxbow lake, Marañon River, 4 ° 35 ' 10.93 ” S 73 ° 36 ' 32.24 ” W, leg. J. Lomas, 11 / vii / 2021, 5 specimens; MUSM 5703 P, LORETO, Maynas prov., Timicurillo near Amazon River-Indiana, 3 ° 31 ' 54.5 ” S 73 ° 03 ' 45.2 ” W, leg. R. Chujutalli, 05 / i / 2020, 2 specimens; MUSM 5695 P, LORETO, Maynas prov., Iquitos Island in Itaya River, 03 ° 46 ' 20.4 ” S 73 ° 14 ' 17.4 ” W. Leg. J. Mozombite, 23 / x / 2019, 3 specimens; MUSM 5708 P, LORETO, Ucayali prov., Contamana, in flooded areas by Ucayali River, 07 ° 20 ' 52.6 ” S 75 ° 0 ' 49.3 ” W, R. Ramírez et al., 08 / ii / 2018, 5 specimens. Mitochondrial COI sequences are deposited into GenBank under the accession numbers MZ 351334 – MZ 351346, as well as 16 S rRNA, MZ 354657 – MZ 354662.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	etymology	Etymology: This species is named after Dr. Mario Peña, a prolific Peruvian malacologist, to honor his work on marine Mollusca in northern Peru.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	materials_examined	Additional material examined: PERU. MUSM 5702, SAN MARTIN, San Martin prov., fish farming pond near San Juan de Cumbaza, 06 ° 32 ' 06.5 '' S 76 ° 22 ' 14.3 '' W, leg. B. Mera & Y. Saavedra, 15 / vi / 2018, 5 specimens; MUSM 5698, LORETO, Maynas prov., Allpahuayo-Mishana National Reserve-IIAP pond used for fish farming, 03 ° 58 ' 02.7 '' S 73 ° 25 ' 05.8 '' W, leg. K. Mejia & E. Aquituari, 14 / v / 2018, 5 specimens; MUSM 5623, LORETO, Maynas prov., Belen Market in Iquitos, 03 ° 46 ' 01 '' S, 73 ° 14 ' 48 '' W, A. Ampuero & V. Borda, 18 / iii / 2011, 1 specimen; MUSM 5690, LORETO, Maynas prov., Iquitos, Bellavista Market, 3 ° 42 ' 12 '' S 73 ° 14 ' 54 '' W, R. Ramírez, 19 / ix / 2017, 5 specimens; MUSM 5696, LORETO, Maynas prov., Model market, 3 ° 44 ' 26.6 '' S 73 ° 14 ' 34.4 '' W, leg. R. Chujutalli, 12 / v / 2019, 1 specimen; MUSM 5693, LORETO, Maynas prov., Moronillo oxbow lake, near Nanay River, 03 ° 43 ' 15.1 '' S 73 ° 15 ' 55.0 '' W, leg. R. Chujutalli, 09 / iv / 2018, 4 specimens; MUSM 5694, LORETO, Maynas prov., Cocha in tributary of Nanay River, 03 ° 40 ' 0 '' S 73 ° 17 ' 06 '' W, leg. J. Mora, 18 / ix / 2017, 2 specimens; MUSM 5697, LORETO, Maynas prov., Ushpa Caño near Itaya River, 3 ° 46 ' 34.16 '' S 73 ° 14 ' 12.22 '' W, leg. J. Pacayo & R. Chujutalli, 12 / v / 2019, 1 specimen; MUSM 5701, LORETO, Alto Amazonas prov., Pampa Poza near Huallaga River, El Naranjal community, 5 ° 13 ' 29.58 '' S 75 ° 40 ' 21.58 '' W, leg. J. Sucumbio & R. Chujutalli, 21 / iv / 2019, 1 specimen; MUSM 5704, LORETO, Maynas prov., Santa Victoria Community, 3 ° 35 ' 1.1 '' S 73 ° 7 ' 4.3 '' W, leg. E. Andi & R. Chujutalli, 04 / ix / 19, 2 specimens; MUSM 5705, LORETO, Maynas prov., Yanayacu Ravine near Amazonas River, 3 ° 33 ' 36.46 '' S 73 ° 3 ' 42.59 '' W, leg. R. Chujutalli, 28 / iv / 2018, 5 specimens; MUSM 5706, LORETO, Maynas prov., Choro Chico oxbow lake near Los Invencibles rural village, mouth of Napo River, 3 ° 26 ' 15.71 '' S 72 ° 42 ' 7.12 '' W, leg. O. Noriega & R. Chujutalli, 20 / vi / 2019, 2 specimens; MUSM 5707, LORETO, Requena prov., Carachama community near Ucayali River, 5 ° 44 ' 26.52 '' S 74 ° 33 ' 17.27 '' W, leg. R. Chujutalli, 05 / vii / 2019, 5 specimens; MUSM 5710, UCAYALI, Coronel Portillo prov., Chanajao village in Yarinillal Cocha, near, Ucayali River, 8 ° 30 ' 43 '' S 74 ° 26 ' 20 '' W, leg. L. Cirano, 17 / ii / 2018, 5 specimens; MUSM 5712, MADRE DE DIOS, Tambopata prov., ITA Inkaterra in " El Tubo " trail, 12 ° 31 ' 58.2 '' S 69 ° 03 ' 01.3 '' W, leg. R. Ramírez, C. Ihuaraqui, 23 / xii / 2017, 6 shells; MUSM 5700, LORETO, Loreto prov., Mullaca oxbow lake, Nauta, Marañon River, 04 ° 35 ' 10.93 '' S 73 ° 36 ' 32.24 '' W, leg. J. Lomas & J. Icomena, 23 / x / 2019, 3 specimens.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	materials_examined	Type locality (Fig. 2 A): PERU. LORETO, Maynas prov., Iquitos, flooding area of Nanay River, Nueva Bellavista-Nanay (03 ° 42 ' 35.0 '' S, 73 ° 14 ' 55.2 '' W).	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	diagnosis	Diagnosis: Prostate broad proximally. Basal penis gland elongated, in dorsal sheath surface, covers medial and basal regions of the penis sheath. Shell (Fig. 3): Shell globose, medium to large (70 – 110 mm), solid, varied from depressed to elevated body whorls, this latter more noticeable in smaller individuals. Suture channel slightly pronounced. Sculpture marked by fine growth lines, often alternated by growth arrests, and with malleations in some specimens. Elliptical to ovate aperture, inner lip and columella ochre to orange, with interior ochre iridescent. Periostracum greenish brown to dark brown, with light brown spiral bands. Umbilicus wide and deep. Measurements in mm. Holotype, height: 77.76, width: 63.68, aperture height: 58.14, aperture width: 43.32. N = 182, (average ± standard deviation, minimum – maximum) height: 73.9 ± 17.4, 43.6 – 114.4; width: 62.1 ± 15.8, 35.0 – 98.4; aperture height: 53.7 ± 12.6, 33.5 – 83.0, aperture width: 37.9 ± 10.1, 21.9 – 61.7.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	description	Operculum (Figs. 4 A, 4 B): Operculum horny and reniform, covering entire opening. Outer surface with concentric growth lines left to marginal central area. Smooth inner surface, insertion mark elliptical, close to central left margin; occupying ~ 3 / 5 of the inner surface. External anatomy (Figs. 5 A, 8 B): Dark to light grey foot and sole, some specimens with beige sole. Cephalic tentacles elongated; ommatophores with wide base ~ 1 / 6 length of tentacles, short with dark eyes. Snout broad, with pair of lateral labial palps, with half of length of cephalic tentacles. Right lobe wide and short, left lobe about 4 times longer than previous one, with margins folded dorsally (siphon). Foot dorsoventrally flattened. Glandular furrow, with fine edges, occupying ~ 3 / 5 of anterior pedal margin. Columellar muscle covering ~ 1 / 4 of body whorl. Mantle cavity (Figs. 6 A, 8 A, 8 C): Mantle (Figs. 6 A, 8 A) border thick with undulations. Mantle cavity occupying almost half of body whorl. Osphradium (Fig. 8 C) elongated and laterally compressed, base broad and elevated at its proximal region. Lung sac long and wide, occupying ~ 1 / 3 of mantle cavity, with muscular walls, and internal walls highly vascularized. Ctenidial vein between gill and lung sac, thickening near pericardial region. Ctenidial filaments triangular, distal region slightly curved to left. Rectum lateral to ctenidium. Anterior kidney on posterior region of mantle cavity. Gonoduct parallel to rectum, under urinary gutter. Right margin of mantle cavity in males occupied by penis sheath (Fig. 9). In females, a bulging portion located on right margin, corresponding to pallial oviduct. Circulatory and excretory systems (Figs. 6 A, 8 A, 8 D): Ctenidial vein, short with thin walls (Fig. 8 A). Pericardium cavity small. Auricle short and slender. Ventricle triangular with wall thicker than the auricle. These two with a dull orange coloration and connected by a short vessel. Bulbous aorta emerging from posterior ventricular region, bifurcating to the posterior and anterior aorta. Anterior aorta broad, connecting to the light-yellow ampulla. Esophageal artery originating from ampulla, close to anterior aorta. Dark brown posterior kidney (Fig. 8 D), slightly concave and highly vascularized, with acute curvature and narrow and blunt anterior region. Anterior kidney beige colored, broad and triangular, connected to posterior kidney by duct located in left of latter. Two sets of lamellae inside anterior kidney. Nephropore in posterior region of anterior kidney. Digestive system (Figs. 6 C, 7 A, 7 D, 8 A, 8 E, 8 F): Buccal bulb pyriform with red musculature (Figs. 6 C, 8 E). Radula (N = 4) taenioglossate, with ~ 26 – 41 rows (Figs. 7 A, 7 D). Rachidian tooth rectangular with concave base and lateral margins; central cusp rounded with protruding pointed tip; lateral cusps rounded and of ~ 2 / 7 length of central peak. Lateral teeth, with triangular central cusp, and 2 small outer lateral cusps around it. Inner marginal teeth with two triangular cusps, internal cusp of ~ 1 / 5 length of outer cusp; outer marginal cusps with bulged tips. After dorsal folds, lateral to esophagus, two broad and short esophageal pouches, slightly rounded at its distal region (Figs. 6 C, 8 E). Pair of salivary glands alveolate and compacted, lateral to digestive tract. Salivary ducts inserted into bulb dorsal surface, ending in anterior region of longitudinal folds. After these pouches, esophagus widening forming crop, bulged in right side. Gastric muscle (Fig. 8 F) elongated. Stomach divided into dorsal and ventral chambers by reduced septum. Ventral chamber twice size of dorsal one, receiving two ducts of digestive gland in its ventral surface. Walls of ventral chamber with red muscles, possess series of transverse folds. Stomach pouch divided internally by transverse folds posterior to dorsal chamber. Intestinal groove whose left and right margins show major and minor typhlosole respectively. At its origin, major typhlosole distinguishable by having small and rectangular fold. Reduced sorting area lateral to this, slightly expanded. Distal region of intestinal groove with elongated caeca bordering sphincter. Intestine U-turned and bordering kidney cavity to enter by left margin. Within cavity, intestine with 4 loops connected to right side of posterior kidney wall by connective tissue. Rectum (Fig. 8 A) running longitudinally along right side of the mantle cavity, parallel to urinary gutter and gill, ending close to rectal papillae, located near left lobe of mantle. Reproductive system: Female (Figs. 6 A, 8 A). Ovary ivory-white, with agglomerated acini. Acini join forming visceral oviduct, connected to pallial oviduct in its ventral anterior third. Pallial oviduct (Fig. 6 A) embedded in light olive parenchymal tissue of albumen gland-capsule gland complex. Visceral oviduct connecting seminal receptacle (Fig. 8 A), reniform and large. Albumen gland lumen flattened, expanded proximally and distally. Copulatory bursa, ~ 1 / 3 of volume of seminal receptacle, adjacent to this and joining it by narrow channel. Ventral channel joins seminal receptacle and running through capsule gland. Seminal receptacle and albumen gland lumen connected to proximal lumen of capsule gland. This latter presents spiral disposition, occupying ~ 2 / 3 of pallial oviduct. Capsule gland lumen tapering distally. Vagina (Figs. 6 A, 8 A) narrow, thick-walled and red muscular folds inside, with small dorsal channel throughout its length, running longitudinally along right side of mantle cavity and passes beneath urinary gutter. Female opening small, adjacent to rectal papilla. In some specimens, vestigial penis sheath, elongated, with small groove, and penis like structure occurs in mantle cavity. Male (Fig. 9). Testis beige colored, occupying approximately first 2 ½ whorls. Spermiduct in basal region of testis, near pericardium region, connected to proximal region of cylindrical and flattened seminal vesicle at its left margin. Prostate (Fig. 9 C) broad in its anterior region, running longitudinally along mantle cavity, decreasing its diameter toward posterior region. Prostate inserted below anal papilla, continues with small outgrowth of tissue. Penis pouch (Figs. 9 A, 9 B) composed of translucent membrane, and penis rolled seen through. Penis bulb solid, occupies ~ 2 / 3 of penis pouch. Penis cylindrical and long, slightly decreasing diameter at distal portion. Penis pouch losing its thickness and forms membrane at base of penis sheath, which occurs in left anterior region of mantle cavity. Penis sheath elongated, slightly curved to right, with bulged base. Two glands in dorsal region of sheath: apical gland, ~ 1 / 3 of length of sheath, with rough surface; basal gland elongated, covering also middle region. Basal ventral gland embedded in sheath, with small opening slit on outer sheath face. Notch in middle of penis sheath. Throughout penis sheath occurs groove covered by thin membrane originating on its left side. Eggs (Figs. 10 A, 10 B, 10 C): Clutch (N = 4) elongated, calcareous eggs disposed in a tight cluster, with polygonal shapes, similar to honeycomb, light to lime green colored newly hatched, turning with the time to pale green; laid on vegetation or substrate above water; number of eggs approximately 400, average egg diameter ~ 4.24 mm. Shell variability (Fig. 11): Observed shells (N = 200) varied from an elongated body whorl morphotype to a compressed one. PCA for the 19 landmarks did not show geographical differentiation, with all groups overlapping in the morphospace. These did not group by locality, where every locality contained most of the shell shapes, a case of extreme intrapopulation variability. First and second principal components explained together 38.32 % of the variation. Shape variation across the first principal component showed an expanded aperture relative to the body whorl and a less pointed shell at the positive end of the axis; towards negative end of axis, shells have higher spires and elongated apertures. Meanwhile, second principal component displayed a rounded aperture, depressed body whorl and immersed spire at the positive end of the axis, and an elongated aperture and body whorl at the end of the negative axis. From here, we can categorize two types of shells: one group with pointed spires and other one with immersed spires. Procrustes ANOVA showed significant differences among localities (F: 4.2457, p <0.001). Pairwise comparisons found non-significant differences in most cases, except for following pairs: Amazonas / High Ucayali, Amazonas / San Juan de Cumbaza, High Ucayali / Nanay, Nanay / San Juan de Cumbaza (p <0.05) (Supplementary Table 1 provides the pairwise results).	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	distribution	Distribution and habitat: Commonly found in temporary flooded areas near Amazon River tributaries, and oxbow lakes. Snails found at the bottom of water bodies, even near populated areas. Fishers find them in their fish traps left overnight at an approximate depth of one meter in small rivers.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC07FF84FF101457FD75FE00.taxon	discussion	Remarks: Pomacea penai sp. nov. has been identified as Pomacea haustrum (Reeve, 1856) since Weyrauch’s collections (1940 s) from Ucayali River (NMW. Z. 1981.118.03479) (Pain 1960); Haas (1947) identified them as P. haustrum immersum (MUSM 520). Rawlings et al. (2007) sequenced one specimen from Ucayali River (16 S rRNA) housed in the Field Museum (FMNH 223530) and found it to cluster with specimens of one of the introduced species from Florida, USA, which laid green eggs. The following authors highlighted the latter apple snail as a different species from P. haustrum (Hayes et al. 2012; Cowie et al. 2017). Ramírez et al. (2022) found in a COI phylogeny a monophyletic group formed by Rawling’s specimen along with samples of P. penai sp. nov. from around Iquitos, Peru. Furthermore, P. penai sp. nov. internally differs from P. haustrum in possessing a longer penis sheath gland that occupies middle and basal region, a bulged proximal region of prostate, and laying green eggs, in contrast with the pink ones of P. haustrum described by Lopes (1955). Pomacea penai sp. nov. could be confused with Pomacea maculata Perry, 1810 as it shares many conchological characters, mainly in large specimens. Pomacea penai sp. nov. shells usually have a higher spire than P. maculata. Here we should add that the immersed spire morphotype is not a condition only found in P. penai sp. nov., due to its presence in other species, as P. reevei sp. nov., described in this work, and also P. maculata (Hayes et al. 2012). In addition to the anatomical characters mentioned below, P. penai and P. maculata were separated by a genetic distance ranging from 13.4 to 14.6 % (COI). Regarding the internal anatomy, P. penai sp. nov. has developed apical and basal penis sheath glands, whereas in Pomacea bridgesii (Reeve, 1856) group species such as Pomacea scalaris (d'Orbigny, 1835) and Pomacea diffusa Blume, 1957 (Hylton-Scott 1957; Cowie et al. 2006), apical gland is absent and basal gland is reduced. P. maculata and Pomacea curumim Simone, 2004 penis sheath glands arrangement resembles P. penai sp. nov. (Simone 2004; Hayes et al. 2012), from which it differs in the elongated basal sheath gland, compared with the smaller basal and medial glands in P. maculata and P. curumim respectively. Pomacea penai sp. nov. has similarities with the proximal region of prostate and the curved penis sheath of specimen determined as P. haustrum illustrated by Lopes (1955) but differing in the proximal bulky region of the prostate. Polygonal eggs also have been reported in other Pomacea species from P. bridgesii group, such as P. scalaris and P. diffusa, but differ in their orange to brown colors (Cowie et al. 2006). Green colored eggs are described for Pomacea glauca (Linnaeus, 1758), Pomacea pyrum (Philippi, 1851), Pomacea decussata (Moricand, 1836), Pomacea nais Pain, 1949 (Cowie 2002), and Pomacea nobilis (Reeve, 1856) (Ramírez et al. 2022). Circulatory and excretory system of P. penai sp. nov. resembles Pomacea canaliculata (Lamarck, 1822), both with a broad anterior kidney and the independent origin of esophageal artery from ampulla; anterior kidney of P. penai sp. nov. is similar in shape to P. curumim. Concerning radula, concave rachidian base of P. penai sp. nov. is similar to P. maculata, and stout central cusp and lateral denticles to P. canaliculata. Compared to P. maculata, P. penai sp. nov. has a reduced stomach septum.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	description	Figs. 4 C, 4 D, 5 B, 6 B, 6 D, 7 B, 7 C, 7 E, 10 D, 10 E, 10 F, 10 G, 12, 13, 14, 15, 16 E	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	synonymic_list	Synonymy: Pomacea sp. 1 — Ramírez et al. 2022, fig 2 E.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	materials_examined	Types: PERU. Holotype male MUSM 5723 H. Paratypes: MUSM 5723 P, MZSP 122666 (ex MUSM 5723 P), SMF 348817 (ex MUSM 5723 P), LORETO, Maynas prov., Kichwa Cocha, near Napo River, Rango Isla populated center, 2 ° 21 ' 42.55 '' S 73 ° 52 ' 11.27 '' W, leg. R. Chujutalli & & S. Perez, 18 / viii / 2022, 7 specimens; MUSM 5716 P, LORETO, Maynas prov., Pucayacu, near Napo River, 2 ° 12 ' 13.26 '' S 74 ° 5 ' 11.55 '' W, leg. R. Chujutalli, 29 / xi / 2018, 5 specimens; MUSM 5715 P, LORETO, Maynas prov., Puerto Alegre near Lagartococha, Napo River, 2 ° 37 ' 20 '' S 73 ° 31 ' 40.1 '' W, leg. R. Chujutalli, 17 / viii / 2017, 5 specimens; MUSM 5720 P, LORETO, Alto Amazonas prov., Cocha Naranjillo near Huallaga River, Naranjal community, 5 ° 13 ' 27,56 '' S 70 ° 54 ' 32.1 '' W, leg. J. Sucumbio et al. 06 / iii / 2020, 5 specimens; MUSM 5724 P, LORETO, Ucayali prov., Hermanos Lake, near Cruz Muyuna Lake, Ucayali River. 6 ° 44 ' 31.8 '' S 75 ° 01 ' 40.1 '' W, leg. R. Chujutalli, 05 / iii / 2022, 1 specimen; MUSM 5719 P, LORETO, Alto Amazonas prov., Shirui Cocha near Huallaga River, Naranjal community, 5 ° 13 ' 29.58 '' S 75 ° 40 ' 21.58 '' W, leg. J. Sucumbio, et al. 27 / v / 2019, 1 specimen; MUSM 5627 P, MADRE DE DIOS, Tambopata prov., Chacra Gamitana station, near Madre de Dios River, 12 ° 30 ' 05.7 '' S 68 ° 58 ' 34.2 '' W, A. Ampuero et al., 17 / vi / 2011, 4 specimens; MUSM 5718 P, LORETO, Maynas prov., Bandeja Isla in Monte Verde oxbow, near Napo River, 2 ° 8 ' 2.78 '' S 74 ° 13 ' 16.91 '' W, leg. R. Chujutalli & G. Monchoa, 21 / xii / 2019, 4 specimens.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	etymology	Etymology: The specific epithet of the type species is in honor of the English malacologist Lovell Augustus Reeve, who contributed with several illustrated works in malacology.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	materials_examined	Additional material examined: PERU. MUSM 5634, UCAYALI, Purus prov., Oxbow Lake near Cocha San Marcos, 9 ° 53 ' 47.9 '' S 70 ° 51 ' 69.5 '' W, 28 / vii / 2010, 4 shells; MUSM 5633, UCAYALI, Purus prov., Alto Purús River: Esperanza Surveillance Post, Cocha San Marcos, 9 ° 53 ' 48 '' S 70 ° 51 ' 69.5 '' W, leg. F. Chang, 09 / vii / 1994, 1 shell; MUSM 5635, UCAYALI, Purus prov., Cocha Livia, 9 ° 56 ' 58.2 '' S 70 ° 54 ' 32.1 '' W, 25 / vii / 2010, 8 shells; MUSM 5632, UCAYALI, Purus prov., Miguel Grau, Cocha Grau near Curanja River, 09 ° 57 ' S 70 ° 59 '' W, leg. F. Chang, 08 / ix / 1994, 1 shell; MUSM 5629, MADRE DE DIOS, Tambopata prov., Cocha Camp in Cuzco Amazonico-Biotrop (Inkaterra), Madre de Dios River, 12 ° 35 ' S 69 ° 05 ' W, R. Ramírez, 30 / iv / 1991, 3 shells; MUSM 5722, MADRE DE DIOS, Tambopata prov., ITA Inkaterra, Madre de Dios River, 12 ° 32 ' 04.9 '' S 69 ° 02 ' 50.8 '' W, leg. W. Ruiz, 23 / xii / 2017, 10 shells; MUSM 5631, MADRE DE DIOS, Tambopata prov., Miraya temporary lake, Palma Real River, ca. Palma Real Control Post, Palma Real Community, 12 ° 30 ' 59.1 '' S 68 ° 48 ' 44.3 '' W, leg. H. Ortega, 28 / vii / 1995, 2 shells; MUSM 5628, MADRE DE DIOS, Tambopata prov., Oxbow Lake near Valencia Lake, 12 ° 28 ' 11.9 '' S 68 ° 47 ' 32.8 '' W, A. Ampuero et al., 17 / vi / 2011, 1 shell. Mitochondrial COI sequences are deposited into GenBank under the accession numbers MZ 351319 – MZ 351332, as well as 16 S rRNA MZ 354652 – MZ 354656. Type locality (Fig. 2 D): PERU. LORETO, Maynas prov., Kichwa Cocha, near Napo River, Rango Isla populated center (2 ° 21 ' 42.55 '' S 73 ° 52 ' 11.27 '' W).	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	diagnosis	Diagnosis: Grey penis sheath wide with bulged tip. Penis sheath gland in the apical region, occupying the entire bulged tip and with a small notch.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	description	Shell (Fig. 12): Shell large (81 – 137 mm), globose, thin, some specimens with thick shells, with 5.3 to 5.6 whorls. Periostracum yellowish to chestnut; Sculpture similar to previous species. Spire slightly elevated to depressed, with a deeply sutural channel. Aperture wide and ovate. Interior cream to white; specimens from Madre de Dios and Purus basins with notorious dark spiral bands. Outer lip light brown to white, and as in the previous character, southern specimens different due to the presence of dark spots in outer lip. Umbilicus narrow and deep. Measurements in mm. Holotype, height: 114.10, width: 93.28, aperture height: 90.22, aperture width: 60.18. N = 181, (average ± standard deviation, minimum – maximum) height: 102.7 ± 21.6, 72.6 – 207.7; width: 85.8 ± 19.5, 59.7 – 185.4; aperture height: 81.8 ± 16.2, 60.1 – 164.2; aperture width: 54.4 ± 12.9, 37.1 – 120.7. Operculum (Figs. 4 C, 4 D): Features similar to P. penai with the following remarks. Operculum ovate semicircular, covering ~ 2 / 3 of aperture. Concentric grooves in outer surface, with mesomarginal nucleus. Smooth inner surface. Elliptical insertion mark occupying ~ 2 / 3 of internal area. External anatomy (Figs. 5 B, 13 B): Characters similar to P. penai with the following differences. Dark grey to black foot. Sole grey. Long cephalic tentacles. Short and broad ommatophores with dark eyes. Snout broad and elongated, with lateral labial palps, with ~ 2 / 3 of length of cephalic tentacles. Elongated siphon. Glandular furrow, occupying ~ 2 / 3 of anterior pedal margin. Broad and short columellar muscle, covering ~ 1 / 5 of the body whorl. Mantle cavity (Figs. 6 B, 13 A, 13 C): Similar characters of mantle to those of P. penai sp. nov. with following characters. Cavity (Figs. 6 B, 13 A) occupying almost half of body whorl. Osphradium (Fig. 13 C) in left side of cavity, posterior to border, comprised laterally, with curved posterior region. Base of osphradium, wide and slightly elevated at anterior region. Lung sac elongated and trapezoidal, occupying ~ 3 / 5 of mantle cavity, with thick muscular walls. Ctenidial vein between gill and lung sac, thickening near pericardium region. Ctenidial filaments slender and long with tip slightly curved to left. Circulatory and excretory systems (Figs. 6 B, 13 A, 13 D): Similar to those described for previous species, with following remarkable attributes. Pericardium cavity (Fig. 13 A) wide. Auricle broad. Ventricle pyramidal. Posterior kidney (Fig. 13 D) slightly concave and vascularized, broad, with anterior region slightly curved with blunt tip. Anterior kidney elongated and triangular. Digestive system (Figs. 6 D, 7 B, 7 C, 7 E, 13 A, 13 E, 13 F): Structures similar to those of P. penai sp. nov. Buccal bulb spherical and broad (Figs 6 D, 13 E). Radula (N = 3) with same configuration as previous species with the following differences: with ~ 26 – 41 rows (Figs 7 B, 7 C, 7 E). Rachidian tooth rectangular with slightly concave base and lateral margins; central cusp broad and rounded; lateral cusps rounded and of ~ 2 / 11 length of central peak. Lateral teeth, with triangular central cusp, and minuscule outer lateral cusps around it. Inner marginal teeth with two tiny triangular cusps, internal cusp of ~ 2 / 11 length of outer cusp. Pair of salivary glands compact (Figs 6 D, 13 E), lateral to digestive tract. Esophageal pouches adjacent to esophagus slender and short. Gastric muscle broad and long (Fig. 13 F). Septum lingulate and elevated. Proximal region of major typhlosole with elongated fold. Minor typhlosole origin with developed fold at margin of stomach pouch. Sorting area of same proportion as P. penai sp. nov. Elongated and wide intestinal caeca. Reproductive system: Female (Figs. 6 B, 13 A). Structures resembling previous species with the following differences. Ovary ivory, with several separated acini joining and forming the visceral oviduct. Seminal receptacle (Fig. 13 A) elongated. Parenchymal tissue of albumen gland-capsule gland complex orange colored. Albumen gland lumen expanded. Vagina broad (Figs. 6 B, 13 A), with thick walls, adjacent to right side of mantle cavity, beneath a short urinary gutter. Female opening wide. Male (Fig. 14). Visceral structures similar to previous species. Testis cream, occupying approximately first 3 1 / 2 whorls. Slender spermiduct in the basal region of the testis, over pericardium region. Seminal vesicle globose and reniform (Fig. 14 C). Prostate elongated with a strong curvature at the proximal region, running adjacent to rectum, and beneath urinary gutter in its distal portion. Prostate inserted below anal papilla. Penis pouch (Figs. 14 A, 14 B) massive, containing penis and penis bulb, this latter solid and occupying ~ 1 / 2 of penis pouch. Penis cylindrical and thick, abruptly decreasing diameter at distal portion. Penis sheath grey, wide, triangular, slightly curved to the right, and bulged tip. With two glands in right dorsal margin: one in distal region, occupying entire bulged tip and with a small notch; basal gland wrinkled, elongated and subtriangular to trapezoidal, near to sheath base and occupying ~ 1 / 3 or less of penis sheath. Basal outer gland embedded in the right region of the sheath, with a pore in the ventral surface. Eggs (Figs. 10 D, 10 E, 10 F, 10 G): Clutch (N = 3) with spherical eggs, slightly tight disposed; newly hatched white with a slime layer cover, turning with time to a cream color; number of eggs approximately of 300, average egg diameter near 6 mm. Shell variability (Fig. 15): Geometric morphometric analysis of the shells (N = 142) showed most specimens from Madre de Dios basin had a lower spire and expanded aperture. First and second principal component explained 48.65 % of the variation. Shape variation across the first principal component showed an expanded aperture relative to a depressed body whorl at the positive end of the axis; towards negative end of axis, shells have elevated body whorls. For the second principal component, shells displayed an elongated aperture and high spire at the end of the negative axis, and an expanded aperture and depressed spire at the positive end of the axis. Procrustes ANOVA showed highly significant differences among localities (F: 7.2221, p <0.001). Pairwise comparisons also found significant differences among several localities. In particular, Madre de Dios specimens had highly significant differences compared to other localities, except Amazonas (Supplementary Table 1 provides the pairwise results). Other significant pairs were Amazonas / Huallaga, Amazonas / Napo and Napo / Huallaga (p <0.05). axes.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	distribution	Distribution and habitat: Deep oxbow lakes near Napo, Huallaga, Ucayali, Madre de Dios and Purus Rivers. Fishers usually find them in their fish traps left overnight.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
03B98B27BC0CFF9FFF1017B6FDABF980.taxon	discussion	Remarks (Fig. 16): Pomacea reevei sp. nov. matches with the illustration of Delessert (1841: pl. 31, figs. 3 a, 3 b) as Ampullaria canaliculata (Lamarck, 1822). This author described the specimen (Fig. 16 A, coded MHNG-MOLL- 33489) as a large-shelled individual claiming that it once belonged to Lamarck’s collection. Our concept of P. reevei sp. nov. includes Ampullaria gigas (non Spix in Wagner, 1827) as presented by Reeve (1856: pl. 1, fig. 3) in his Conchologia Iconica. Reeve described it in detail based on shell material (NHMUK 20110205), with type locality of Solomon’s River (possibly referring to Solimıes River, the upper section of Amazon River located in Brazil) (Fig 16 B). Sowerby (1909) and Alderson (1925) disregarded Delessert specimen as A. canaliculata based on its size, and considered it was Ampullaria gigas (Spix in Wagner, 1827), synonymizing also with A. gigas (Reeve, 1856). Later, Pain (1960) proposed A. gigas of Spix and Reeve to be synonyms of P. maculata (Cowie & Thiengo 2003). This could be demonstrated by P. reevei sp. nov. specimens in Pain collection housed in Wales Museum (NMW. Z. 1981.118.00486; NMW. Z. 1981.118.00201) determined as P. maculata (Figs. 16 C, 16 D). We should add that previous authors such as Pain or Alderson described the shell of their P. gigas as thin, a trait present in our material, and did not mention Spix’s holotype, probably as much of his material was lost or destroyed (Alderson 1925). Hayes et al. (2012) assigned a neotype for P. gigas, and rejected the possible type status of Delessert’s shell arguing that there are still doubts regarding the labelling process of this specimen. Based on observations of collected material for the present research, we propose that the material used by Delessert and Reeve could be P. reevei sp. nov. Likewise, we compared P. reevei sp. nov. with Pomacea maculata described by Hayes et al. (2012). Shells of P. reevei sp. nov. are frequently thin and fragile even in most larger specimens, compared to the thickness of P. maculata. Deeply narrow umbilicus and angled channeled suture are shared with Pomacea crosseana (Hidalgo, 1871) (syntypes examined, MNCN 15.05.1047 and 15.05.11485), where P. reevei had a more prominent and expanded body whorl. Male reproductive system of P. reevei sp. nov. resembles Pomacea paludosa (Say, 1829) (Hanning 1979), but with following exceptions: apical penis sheath gland covers entire sheath tip including the notch, and basal internal gland does not cover most of penis sheath gland. It is noteworthy that in COI phylogeny, P. reevei and P. paludosa were closely related (genetic distance ranged from 13.6 – 14.3 %), although with low support, within P. canaliculata clade; one species of this clade, P. maculata, had a distance of 12.1 – 12.8 % to P. reevei (Ramírez et al. 2022). Pomacea reevei sp. nov. could be distinguished from P. maculata and P. canaliculata by possession of its apical penis sheath gland located in the bulged tip, as opposed to enlarged apical gland that occupies more than 1 / 3 of the penis sheath (Sachwatkin 1920; Hayes et al. 2012); this character is shared also by Pomacea lineata (Spix, 1827) (Thiengo 1987). Penis pouch proportion in P. reevei sp. nov. is similar to Pomacea dolioides (Reeve, 1856) and differs from this species by its smaller internal penis sheath glands, compared to the developed basal internal gland in P. dolioides (Tillier 1980). Kidney of P. reevei sp. nov. has characters similar to those of P. maculata, differing by its anterior broad set of lamellae in anterior kidney, as is present in P. canaliculata (Hayes et al. 2012). Ampulla vessels have the same pattern of P. canaliculata (Hayes et al. 2012). Regarding eggs, P. reevei shares spherical eggs shape and mucus matrix with P. paludosa (Snyder & Snyder 1971; Rawlings et al. 2007). Spherical shape also appeared in species from this clade like P. canaliculata, P. maculata and P. lineata (Thiengo 1987; Cowie et al. 2006; Hayes et al. 2012). Also, P. reevei sp. nov. eggs are characterized by their highest individual average diameter among all described Pomacea species.	en	Ampuero, André, Ramírez, Rina (2023): Description of two new species of apple snail (Ampullariidae: Pomacea) from Peruvian Amazonia. Zootaxa 5258 (1): 76-98, DOI: 10.11646/zootaxa.5258.1.3, URL: http://dx.doi.org/10.11646/zootaxa.5258.1.3
