Loricaria luciae, Thomas & Rodriguez & Cavallaro & Froehlich & Corrêa, 2013

Thomas, Matthew R., Rodriguez, Mónica S., Cavallaro, Marcel R., Froehlich, Otávio & Corrêa, Ricardo Macedo, 2013, Loricaria luciae, a new species of whiptail catfish (Siluriformes: Loricariidae) from the Paraguay and lower Paraná River basins of southeastern South America, Zootaxa 3745 (3), pp. 365-378 : 366-375

Loricaria luciae , new species

( Figs. 1A , 2–4 )

Holotype. ANSP 182408 , 187.0 mm SL; Argentina, Paraná basin, Corrientes Province, rio Paraná (left bank) at private park (Club San Martin) near town of Perichon , N of Rt. 12, NE of Corrientes, 27°25’6.94”S, 58°45’27.47”W, M.H. Sabaj, M. Mirande, and ornamental fishermen, 11 April 2005.

Paratypes (83). Argentina: Paraná basin: Corrientes Province: ANSP 193290 (3, 103.8– 179.3 mm SL), INPA 38777 (ex. ANSP 182408 ; 2, 151.3–175.0 mm SL), CI-FML 5874 (ex. ANSP 182408 , 160.0 mm SL; ex. ANSP 193290, 172.5 mm SL); same data as holotype . Bolivia: Paraguay basin: Santa Cruz Department: UF 82235 (3, 68.1–144.0 mm SL), CBF 13309 (ex. UF 82235 ; 2, 120.0–124.0 mm SL), rio Tucavaca 25 km NE of Santiago de Chiquitos (rio Paraguay drainage) , 18°12’54”S, 59°28’05”W, 24 June 1986, F.G. Thompson. Brazil: Paraguay basin: Mato Grosso State: MNRJ 27483 (1, 127.8 mm SL), ribeirão Baús (rio Cuiabá), Baús District, Município Acorizal , on the bridge of the old road , 15°14'27"S, 56°17'24"W, 22 July 1983, G.W. Nunan and W.D. Bandeira; MZUSP 78778 (1, 105.2 mm SL), rio Jaurú , bridge on Araputanga / Jaurú Road , 15°28’58”S, 58°35’53”W, 28–29 February 2002, F.A. Machado, F.C. T. Lima, G. Borges, and J. Nakagaki. Mato Grosso do Sul State: LIRP 5560 (4, 129.3– 168.3 mm SL), rio Formoso, at Morada do Rio farm, about 15 km from the mouth, Município of Bonito , 21°04’45”S, 56°20’21”W, 29 August 2004, Bodoquena Expedition Team; LIRP 5604 (19, 47.1–148.1 mm SL), Corrego Jenipapo, tributary of Bacuri, rio Miranda basin, Município of Bonito , 20°59’07”S, 56°27’11”W, 5 September 2004, Bodoquena Expedition Team. Paraguay: Paraguay basin: Amambay Department: UMMZ 206772 (4, 58.0–120.0 mm SL), rio Aquidaban in Parque Nacional Cerro Cora ca. 32km WSW of Pedro Juan Caballero , 22°38'12"S, 56°02'60"W, 26 July 1979, J.N. Taylor, T.W. Grimshaw, R. Myers, Creighton, R.M. Bailey, G. R. Smith. Central Department: UMMZ 208113 (29 + 1 CS, 151.8–189.1 mm SL), rio Paraguay overflow ca. 1.0 km S of Puente Remanso bridge, rio de la Plata drainage, 15 September 1979, J.N. Taylor, E. Koon, G. & B. Smith. Concepción Department: UMMZ 207830 (10, 116.2– 162.5 mm SL), rio Aquidaban at Paso Horqueta, ca 24 km NNW of Loreto , 23°03’48”S, 57°23’00”W, 5 September 1979, J.N. Taylor, G. & B. Smith, E. Koon, R. Myers. Paraguarí Department: UMMZ 205676 (2, 70.0–130.0 mm SL), Salto de Pirareta, 400+ m below the falls, ca 5.8 km S of Piribebuy ( rio Tebicuary drainage) , 25°41'60"S, 57°32'60"W, 26 May 1979, J.N. Taylor and T.W. Grimshaw .

Non-type specimens (144). Brazil: Paraguay basin: Mato Grosso State: USNM 326340 (2 of 5, 92.7–129.5 mm SL), rio Paraguay in Caceres and surroundings, 11 August 1991, R.E. Reis, L. R. Malabarba, N.A. Menezes, M. Weitzman, and F.A. Machado. Mato Grosso do Sul State: FMNH 10859 (2 of 4, 83.6–90.4 mm SL), stream near 7 km of Rochedo, toward Correguinho (rio Negro drainage), 29 August 1998, O. Oyakawa, N. Menezes, A. Zanata, F. Lima, and others; LIRP 5558 (2, 42.1–57.0 mm SL), rio Salobra, close to Cachoeira Toca da Onça, Município of Bodoquena, 20°44’17”S, 56°44’05”W, 22 August 2004, Bodoquena Expedition Team; LIRP 5559 (1, 52.9 mm SL), Rancho Branco, rio Salobra drainage, Córrego Salobrinha, Municipality of Bodoquena 20°40’57”S, 56°46’36”W, 2 September 2004, Bodoquena Expedition Team; LIRP 5621 (2, 116.6–129.0 mm SL), Córrego da Fazenda, at bridge of MS-345 road, Taquaruçu-Bacuri-Miranda drainage, Municipality of Bonito, 20°53’40”S, 56°22’17”W, 4 September 2004, Bodoquena Expedition Team; MZUSP 59308 (8, 66.4–90.1 mm SL), Riacho da Chácara da Portela, tributary of rio Aquidauana, 19°48'40"S, 54°50'39"W, 28 August 1998, H. Ortega, M. T. Piza, R. Lourival, F.C. T. Lima, and others; MZUSP 59668 (1, 104.1 mm SL), rio Abobral, main channel, 19°25’73”S, 57°03’68”W, 7 September 1998, A. Machado and B. Chernoff; ZUFMS-PIS 1028 (2, 119.9– 146.1 mm SL), Córrego Salobrinha, 20°41’05.2”S, 56°46’41.8”W, 11 August 2001, D. Silva, M. R. Cavallaro, and L.S. Inocencio; ZUFMS-PIS 1036 (1, 115.0 mm SL), Córrego Salobrinha, 20°40’58.1”S, 56°46’39.4”W, 16 June 2001, M. R. Cavallaro and O. Froehlich; ZUFMS-PIS 1056 (2, 40.4–122.6 mm SL), Córrego Azul and rio Salobra, 20°45’35.8”S, 56°45’08.3”W, 9 April 2000, O. Froehlich and L.P.C. Lopes; ZUFMS-PIS 1081 (2, 153.0– 159.5 mm SL), rio Salobra, 20°40’16.3”S, 56°45’34.4”W, 12 August 2001, D. Silva, M. R. Cavallaro, L.S. Inocencio, and O. Froehlich; ZUFMS-PIS 1501 (1, 104.0 mm SL), Córrego Salobrinha, 20°40’58.1”S, 56°46’39.4”W, 13 April 2001, O. Froehlich, and L.S. Inocencio; ZUFMS-PIS 1511 (2, 49.1–94.3 mm SL), rio Salobra, 20°40’09.9”S, 56°45’38.5”W, 20 May 2000, O. Froehlich; ZUFMS-PIS 1559 (1, 124.1 mm SL), Córrego Salobrinha, 20°41’05.2”S, 56°46’41.8”W, 13 August 2001, D. Silva, M. R. Cavallaro, L.S. Inocencio, O. Froehlich; ZUFMS- PIS 1565 (2, 111.3– 123.9 mm SL), Córrego Salobrinha, 20°41’07.5”S, 56°46’41.4”W, 13 August 2001, D. Silva, M. R. Cavallaro, L.S. Inocencio, and O. Froehlich; ZUFMS-PIS 2498 (2, 153.9–188.0 mm SL), Córrego Salobrinha, 20°41’07.5”S, 56°46’44.3”W, 2 September 2005, M.J.A. Vilela, O. Froehlich, O.C. Forster, and R.D. Vargas; ZUFMS-PIS 2499 (1, 98.8 mm SL), rio Salobra, 20°46’49.6”S, 56°44’31.5”W, 3 September 2005, M.J.A. Vilela, O. Froehlich, O.C. Forster, and R.D. Vargas; ZUFMS-PIS 3212 (8, 65.8–132.1 mm SL), Córrego Salobrinha, 20°40’59.1”S, 56°47’11.6”W, 27 July 2006, O. Froehlich, F. Severo-Neto, M.C. Teixeira, M.O. Tanaka, M. R. Cavallaro, M. Beccari, and T. T.M. Taveira; ZUFMS-PIS 3213 (19, 61.7–141.8 mm SL), Córrego Salobrinha, 20°40’12.6”S, 56°46’12.1”W, 26 July 2006, O. Froehlich, F. Severo-Neto, M.C. Teixeira, M.O. Tanaka, M. R. Cavallaro, M. Beccari, and T. T.M. Taveira; ZUFMS-PIS 3214 (4, 62.3–135.5 mm SL), Córrego Salobrinha, 20°40’23.2”S, 56°46’24.5”W, 31 July 2006, O. Froehlich, F. Severo-Neto, M.C. Teixeira, M.O. Tanaka, M. R. Cavallaro, M. Beccari, and T. T.M. Taveira; ZUFMS-PIS 3215 (32, 36.1–139.8 mm SL), Córrego Salobrinha, 20°40’12.7”S, 56°46’12.1”W, 25 March 2007, O. Froehlich, O. Camilo, F. Severo-Neto, J. Souza, M.J.A. Morales, and T. T.M. Taveira; ZUFMS-PIS 3216 (7, 64.0– 119.1 mm SL), Córrego Salobrinha, 20°40’33.5”S, 56°46’34.1”W, 30 July 2007, O. Froehlich, F. Severo-Neto, M.C. Teixeira, M.O. Tanaka, M. R. Cavallaro, M. Beccari, and T. T.M. Taveira; ZUFMS-PIS 3217 (10, 42.4–117.7 mm SL), Córrego Salobrinha, 20°40’23.2”S, 56°46’24.5”W, 26 March 2007, O. Froehlich, O. Camilo, F. Severo-Neto, J. Souza, M.J.A. Morales, and T. T.M. Taveira; ZUFMS-PIS 3218 (5, 63.2–138.0 mm SL), Córrego Salobrinha, 20°40’23.2”S, 56°46’24.5”W, 30 July 2006, O. Froehlich, F. Severo-Neto, M.C. Teixeira, M.O. Tanaka, M. R. Cavallaro, M. Beccari, and T. T.M. Taveira. Paraguay: Paraguay basin: Alto Paraguay Department: MZUSP 54220 (1, 67.6 mm SL), unnamed riacho, about 200 m from the junction between rio Paraguay and rio Miranda, 20°40'60"S, 57°58'60"W, 6 September 1997, M. T. Piza, K. Awbrey, J. Thomsen, D. Moskovits, and others. Amambay Department: UMMZ 206734 (1, 41.0 mm SL), Arroyo Piky at bridge on dirt hwy. (Rte 5) ca. 47 km WSW of Pedro Juan Caballero (rio Aquidaban drainage), 22°41'60"S, 56°12'00"W, 24 July 1979, J.N. Taylor, T. Grimshaw, R. Myers, Creighton, and R.M. Bailey. Caaguazú Department: UMMZ 206900 (11, 45.0–80.0 mm SL), Arroyo Guazu at bridge on hwy. (Rte 2) ca. 23.1 km E of Coronel Oviedo jct. (rio Tebicuary drainage), 25°28'07"S, 56°11'04"W, 3 August 1979, J.N. Taylor, T. Grimshaw, R. Myers, Creighton, and R.M. Bailey. Canindeyú Department: UMMZ 206319 (1, 52.0 mm SL), Arroyo Itandey, tributary to Arroyo Carimbatay, ca. 7.6 km WSW (by dirt road) of Curuguaty (Arroyo Curuguaty- rio Jejui-Guazu drainage), 24°30'47"S, 55°43'25"W, 6 July 1979, R.M. Bailey, J.N. Taylor, T. Grimshaw, R. Myers, and Creighton; UMMZ 206577 (1, 106.0 mm SL), rio Corrientes and adjacent flood pool, 32.4 km W of turnoff to Curuguaty (60 km NE of Mbutuy intersection); rio Jejui-Guazu drainage, 24°22'54"S, 55°55'60"W, 21 July 1979, J.N. Taylor, T. Grimshaw, R. Myers, Creighton, and R.M. Bailey. Concepción Department: UMMZ 208009 (1, 92.0 mm SL), Arroyo Guaireno at bridge on dirt road, 33.0 km S of jct. with Rte. 5, at Yby-Yau (rio Ypane drainage), 23°15'18"S, 56°30'00"W, 11 September 1979, J.N. Taylor, E. Koon, R. Myers, G. Smith, and T. Creighton. Cordillera Department: UMMZ 205780 (2, 60.0–147.0 mm SL), small tributary of rio Piribebuy, 1.6 km S of Tobati (=Arroyo Tobati), 25°18'30"S, 57°04'00"W, 3 June 1979, R.M. Bailey, J.N. Taylor, and T. Grimshaw; UMMZ 205813 (2, 49.0–51.0 mm SL), Arroyo Tobati ca. 0.1 km E of Tobati (rio Piribebuy drainage), 25°16'54"S, 57°02'60"W, 4 June 1979, J.N. Taylor, T. Grimshaw, and R. Myers. Presidente Hayes Department: UMMZ 207137 (1, 119.0 mm SL), Riacho Je-e at bridge on paved highway, ca 67.5 km NW of Benjamin Aceval (Aguaray-Guazu drainage), 24°34'54"S, 58°01'00"W, 15 August 1979, G. R. Smith, J.N. Taylor, T. Grimshaw. San Pedro Department: UMMZ 207992 (1, 122.0 mm SL), rio Ypane at ferry crossing, ca 2.0 km S by dirt road to Belen, ca. 18 km ESE of Concepción, 23°29'30"S, 57°15'00"W, 10 September 1979, J.N. Taylor, G. & B. Smith, E. Koon, and R. Myers; UMMZ 208028 (1, 128.0 mm SL), rio Ypane on S bank at bridge on dirt hwy, 52.7 km S of jct with Rte 5 at Yby-Yau, 23°27'18"S, 56°31'00"W, 11 September 1979, J.N. Taylor, G. & B. Smith, E. Koon, and R. Myers.

Diagnosis. Loricaria luciae is distinguished from all other species of Loricaria , except L. coximensis , L. holmbergi , and L. pumila by having large and tightly joined plates confined to median abdominal space between lateral abdominal plates, with pectoral girdle entirely naked ( Fig. 1A ) or occasionally with small isolated plates near base of pectoral fin, vs. plates covering the entire median abdominal space and pectoral girdle, usually with a V -shaped anterior margin ( Fig. 1B ). It is distinguished from L. pumila by having a larger adult size (130–190 mm vs. less than 85 mm SL), iris operculum present (vs. absent), and larger eye size (minimum orbital diameter 16.2– 19.4% vs. usually less than 13.3% of HL). Loricaria luciae is most similar to L. coximensis and L. holmbergi , from which it is distinguished by having a larger adult size (130–190 mm SL vs. usually less than 130 mm SL) and larger post-ural plate length (17.0–20.3% vs. 7.4–14.2% of HL). It is further distinguished from L. coximensis by having a higher number of total lateral plates (32–33 vs. 28–31).

Description. A medium-sized species of Loricaria , reaching 190 mm SL, based on specimens examined. Morphometric data and selected meristic counts are presented in Tables 1 and 2, respectively. General body form is depicted in Figures 2–3 . Head and trunk relatively wide and depressed; caudal peduncle moderately long and depressed. Eyes large, minimum orbital diameter 16.2–19.4% of HL; iris operculum present. Postorbital notch well developed and angular; maximum orbital diameter 18.7–23.2% of HL.

Dermal plates on dorsum of body from snout tip to dorsal-fin origin with moderately developed odontode keels. Odontodes weakly developed on lateral margins of head from snout tip to opercle and along anterodorsal margin of orbit. Supraoccipital with two separate, but closely aligned parallel keels, continuing on two predorsal plates but more spaced apart. Nuchal plate without median keel, or if present, weakly developed. Dorsal plates between compound pterotic and dorsal-fin origin each with median keel. Lateral surface of exposed cleithrum with small median keel.

Maxillary barbel long, reaching or exceeding medial end of ventral gill opening, and with multiple secondary branches. Fringe barbels along margin of lower lip simple and relatively short. Filiform papillae covering both lobes of lower lip numerous and short, each less than half the length of fringe barbels. Premaxillary teeth 1–5 (modally 3) per ramus. Buccal papillae behind upper jaw long and slender, consisting of three central papillae, two lateral papillae, and 8–12 additional papillae behind premaxillary teeth (4–5 papillae per ramus and one between each ramus). Dentary teeth 6–10 (modally 7–8) per ramus.

Total plates in lateral series 32–33 (modally 32); anterior lateral plates 18–21 (modally 19) with two parallel odontode keels widely separated, becoming closely aligned on caudal peduncle posterior to distal margin of anal fin; posterior lateral plates (i.e., with closely aligned keels) 12–15 (modally 13, rarely 15). Post-anal plates 18–20 (modally 19). Lateral abdominal plates 6–8 (modally 7), rectangular and elongate. Median abdominal space between lateral abdominal plates with medium to large-sized polygonal plates, tightly joined; pectoral girdle entirely naked ( Fig. 1 ) or with isolated patches of small plates near base of pectoral fin ( Fig. 3 ). In immature specimens examined (42.1–59.9 mm SL), abdominal plates were less developed than in larger adults. Plates in lateral abdominal series and pre-anal region were fully developed in the smallest specimen examined (42.1 mm SL), with median abdominal space mostly or entirely naked. Median abdominal plates were partially developed, but small and variably spaced in specimens 47.1–59.9 mm SL, with pectoral girdle entirely naked. Pair of midventral plates immediately posterior to anus separated anteromedially, forming small roughly U- to V-shaped naked area partially surrounding anus; overall shape of anteromedial plate margins curved, acuminate ( Fig. 4A ) to acutely triangular ( Fig. 4B ) in females and males, respectively (see also Sexual Dimorphism).

Dorsal-fin spine length 19.3–26.5% of SL, even with or slightly longer than branched rays, reaching eighth plate posterior to its origin when adpressed; distal margin of fin slightly concave when erected. Adpressed pectoral fin reaching sixth lateral plate posterior to cleithrum; distal margin straight to slightly concave when erected. Pelvic-fin spine length 15.8–20.1% of SL, extending posteriorly to basal quarter of adpressed anal fin. Adpressed anal fin reaching seventh plate posterior to its origin; distal margin straight to slightly convex when erected. Caudal fin with concave distal margin, upper unbranched ray thickened and produced into a long trailing filament (broken in all specimens examined). Post-ural plate large (length 17.0–20.3% of HL).

Color in alcohol. Ground color light reddish-brown on dorsal and lateral surfaces, fawn or yellowish brown on ventral plated surfaces, and cream on unplated surfaces ( Fig. 2 ). Dorsum of head with large bronze to copper blotches; individual plates entirely or partially edged with dark brown to black pigment. Dorsum of trunk and caudal peduncle with six dark brown transverse saddles; first saddle narrow, crossing plates at level with anterior 2–3 branched rays of dorsal fin; second saddle widest, crossing three mid-dorsal plates immediately posterior to dorsal-fin insertion; remaining four saddles variable in width, situated from about level with posterior margin of anal fin to 2–3 plates anterior to end of caudal peduncle. Ventral surfaces plain, except for dark brown melanophores covering dorsal surface of upper lip and maxillary barbel, and naked area between upper lip and plated margin of snout.

Dorsal fin with small round to oval dark spots evenly spaced on spine and branched rays; interradial membranes lightly dusted with brown melanophores. Pectoral and pelvic fins with large dark-brown oval spots on spine and irregularly distributed on branched rays. Caudal fin with round spots on upper and lower unbranched rays including trailing filament; branched rays with irregularly distributed small dark spots. Post-ural plates covered with brown melanophores forming an elliptical blotch at base of caudal fin.

Color in life. Ground color pale brown to fallow on dorsal and lateral surfaces of body and pectoral fins. Dorsum of head marked with faint, dark grayish-brown to chocolate colored blotches, one extending from middle of orbits anteriorly through nares and along lateral margins of snout, leaving a pale median V-shaped mark on midline of snout anterior to nares; and a second extending posterodorsally from opercle and cleithrum to posterior portion of supraoccipital. As in alcohol preserved specimens, individual plates on head and predorsal region entirely or partially edged with dark brown or black pigment, only more accentuated to form a coarse, loosely reticulated pattern. Dorsal saddle patterns and distribution of spots on fins similar to preserved specimens, except more bold and contrasting. Ventral unplated surfaces on pectoral girdle, head, and surrounding anus and bases of pelvic fins white; abdominal plates with golden-yellow iridescence ( Fig. 3 ).

Sexual dimorphism. Several specimens examined ranging in size from 130–190 mm SL had the following secondary sexual characteristics that are similarly exhibited by mature males of other species of Loricaria ( Isbrücker 1981) : thickened pectoral spine with expanded distal portion, shortened and rounded premaxillary and dentary tooth cusps, and expanded lower lip with shortened filiform papillae and marginal fringe barbels. Sexual dimorphism previously unreported in Loricaria also was observed in the shape of the naked area posteriorly surrounding the genital papilla and position of the papilla within the naked area. In females, this area was broadly triangular to rounded, acuminate, with an anteriorly positioned papilla ( Fig. 4A ) vs. acutely triangular with a centrally positioned papilla in males ( Fig. 4B ). Those sexually dimorphic characteristics were also newly observed in Loricaria holmbergi and L. coximensis .

Distribution and habitat. Loricaria luciae is currently known from throughout the rio Paraguay basin of Bolivia, Brazil, and Paraguay, south to its confluence with the rio Paraná in Argentina ( Fig. 5 ). Most collections of this species are from the Paraguay ecoregion as delineated by Abell et al. (2008), which encompasses the extensive Pantanal wetland complex drained to the south by the rio Paraguay and many of its left-bank tributaries including the Negro, Miranda, Taquari, Piquiri, Cuiabá, Aquidauana, Apá, Ypané, and Tebicuary drainages. Based on material examined in this study, it appears to be less common in right-bank tributaries of the rio Paraguay draining the more arid Chaco ecoregion and in the lower Paraná ecoregion ( Abell et al. 2008). Individuals have been collected in flowing water habitats ranging from small, seasonally intermittent streams (arroyos) to large permanent rivers. Since 2000, this species has been collected repeatedly in small order streams with clear water in the upper rio Miranda drainage within the boundaries of the Parque Nacional da Serra da Bodoquena (M. Cavallaro, and O. Froehlich, pers. obs.). In the Aquidaban and Tebicuary drainages, several specimens were collected in and above riffles at depths of 0.3–1.5 m, where the species was reportedly common (J.N. Taylor et al. UMMZ, unpubl. field notes). In the rio Negro at the southern end of the Pantanal wetlands, specimens were collected in black water at the edge of the main river channel in swift current over sand and mud substrates (B. Chernoff, et al. FMNH, unpubl. field notes). In the lower rio Paraná, the holotype was collected with a seine at night along the shore over substrates of sand, silt and mud, at a depth of less than 1.5 m (M. Sabaj Pérez, pers. comm.).

Etymology. Named in honor of Dr. Lúcia H. Rapp Py-Daniel, Curator of Fishes at Instituto Nacional de Pesquisas da Amazônia, for her many contributions Neotropical ichthyology, particularly loricariid taxonomy and systematics.

Figures

FIGURE 1. Abdominal plate development pattern in adult specimens. (A) Loricaria luciae, paratype, UMMZ 207830, 152.4 mm SL. (B) Typical condition in Loricaria, as exemplified by L. simillima, lectotype, BMNH 1880.12.8.77, 163.1 mm SL. Note lack of plates across pectoral girdle in L. luciae.

FIGURE 2. Loricaria luciae, paratype, ANSP 182408, 178 mm SL, dorsal, lateral and ventral views of preserved specimen.

FIGURE 3. Live coloration of Loricaria luciae, holotype ANSP 182408, 187.0 mm SL, dorsal and ventral views. Photos by M. Sabaj Peréz.

FIGURE 4. Sexual dimorphism in the shape of the naked area surrounding the genital papilla in Loricaria luciae. (A) Female, paratype, LIRP 5604,148.4 mm SL. (B) Male, paratype, LIRP 5560, 129.4 mm SL.

FIGURE 5. Geographic distributions of Loricaria luciae (filled circles; open circle = type locality), L. coximensis (square), and L. holmbergi (triangle). Some symbols represent more than one lot or locality. Numbered locations: 1 = rio Paraguay; 2 = rio Cuiabá; 3 = rio Jaurú; 4 = rio Tucavaca; 5 = rio Taquari; 6 = rio Negro; 7 = rio Aquidauana; 8 = rio Miranda; 9 = rio Aquidaban; 10 = rio Ypane; 11 = rio Jejui-Guazú; 12 = rio Tebicuary.