Brachyhypopomus draco, a new sexually dimorphic species of Neotropical electric fish from southern South America (Gymnotiformes: Hypopomidae)
Giora, Julia
Malabarba, Luiz R.
Crampton, William
Neotropical Ichthyology
2008
2008-12-31
6
2
159
168
MWM8
Giora & Malabarba & Crampton, 2008
Giora & Malabarba & Crampton
2008
[247,525,1512,1535]
Actinopterygii
Hypopomidae
Brachyhypopomus
Animalia
Gymnotiformes
2
161
Chordata
species
draco
sp. nov.
Figs. 1, 2
Holotype. MCP41540, 1(male, 137.3 mm LEA), Brazil, Rio Grande do Sul, Parque Estadual de Itapuã, lagoa Verde( 30°22’52.4"S, 51°01’25"W), 12 Jan 2004, D. Cognato, L. R. Malabarba, C. E. Machado & R. Q. Carvalho. Paratypes.All from Brazil, Rio Grande do Sul. Lagunados Patosdrainage: MCP41539, 1(male, 147.1 mm LEA), collected with the holotype. UFRGS 8888, 1(male, 126.4 mm LEA), same locality of the holotype, 18 Jan 2006, J. Giora, A. P. Dufech& J. Ferrer. MNRJ 30916, 2( 1 male, 151.8 mmLEA, 1 female, 128.4 mm LEA), same locality of the holotype, 9 Feb 2004, D. Cognato, M. Azevedo, A. Schaan& C. Hiroshi. MCP41538, 2( 1 male, 88.4 mmLEA, 1 female, 91.7 mm LEA), same locality of the holotype, 30 Jun 2004, D. Cognato& A. Schaan. MNRJ 30917, 1(male, 150.2 mm LEA), MZUSP 94429, 1(male, 139.1 mm LEA), same locality of the holotype, 9 Sep 2004, D. Cognato, M. Schossler& A. Schaan. MCP41537, 1(female, 108.4 mm LEA), same locality of the holotype, 10 Oct 2003, D. Cognato& A. Schaan. MNRJ 30918, 1(male, 138.2 mm LEA), same locality of the holotype, 11 Nov 2003, D. Cognato, J. Giora, J. Anza& A. P. Dufech. UFRGS 732, 1(female, 78.0 mm LEA), Viamão, arroio Alexandrina( 30°6’S, 50°57’W), 7 Sep 1980, L. R. Malabarba. UFRGS 820, 1(male, 93.7 mm LEA), Guaíba, açude near BR290highway ( 30°07’S, 51°20’W), 4 Nov 1979, L. R. Malabarba, C. F. M. Souto & P. A. Buckup. UFRGS 822, 1(male, 92.5 mm LEA), Guaíba, açude in BR290highway, near junction with BR116highway ( 30°03’S, 51°21’W), 18 Jan 1980, P. A. Buckup& C. Souto. UFRGS 6526, 1(male, 102.2 mm LEA), UFRGS 6525, 1(male, 100.6 mm LEA), Eldorado do Sul, artificial canal near BR290highway ( 30°2’55"S, 51°23’34"W), 10 Dec 2003, W. Crampton& L. R. Malabarba. UFRGS 4317, 1(male, 50.0 mm LEA), Viamão, arroio Itapuã( 30°15’S, 51°02’W), 1 Mar 1988, F. Becker& L. R. Malabarba. Rio Tramandaídrainage: UFRGS 6486, 1(male, 85.3 mm LEA), UFRGS 6488, 1(male, 98.1 mm LEA), UFRGS 6489, 1(male, 100.8 mm LEA), UFRGS 6490, 1(female, 95.5 mm LEA), UFRGS 6491, 1(male, 80.9 mm LEA), Capão da Canoa, creek parallel to Estrada do Marroad ( 29°43’9"S, 50°56’0"W), 20 Nov 2003, L. R. Malabarba, J. Giora, J. Anza & D. Cognato. MNRJ 30915, 2( 1 male, 69.9 mmLEA, 1 female, 85.2 mm LEA), Cidreira, lagoa Fortaleza( 30°8’58"S, 50°14’30"W), 11 Jul 2003, L. R. Malabarba. UFRGS 6671, 3( 2 males, 85.7-107.0 mm LEA, 1 female, 96.3 mm LEA), Terra de Areia, creek into rio Três Forquilhas, along the road RS486 ( 29°33’22"S, 50°4’19"W), 20 Nov 2003, L. R. Malabarba, J. Giora, J. Anza & D. Cognato. Rio Uruguaydrainage: UFRGS 6748, 1(male, 90.8 mm LEA), Rosário do Sul, Sanga do Jacaré, BR290( 30°12’4"S, 55°3’17"W), 18 Jun 2004, D. Cognato, W. Crampton, J. Giora& D. Rocha. MZUSP 94428, 8( 2 males, 95.6- 126.7 mmLEA, 6 females, 51.9-103.94 mm LEA), UFRGS 6753, 1(male, 88.3 mm LEA), Rosário do Sul, stream tributary of arroio Gueromana, BR290( 30°1’00"S, 55°23’18"W), 19 Jun 2004, D. Cognato, W. Crampton, J. Giora& D. Rocha. UFRGS 6785, 4( 2 males, 79.3-105.1 mmLEA, 2 females, 84.5-85.3 mm LEA), UFRGS 6494, 1(male, 92.5 mm LEA), UFRGS 6497, 1(male, 90.1 mm LEA), Rosário do Sul, sanga do Jacaré, BR290( 30°12’42"S, 55°3’17"W), 26 Nov 2003, L. R. Malabarba, J. Giora, D. Cognato, G. Neves & J. Ferrer. UFRGS 6789, 4( 2 males, 106.5-126.3 mmLEA, 2 females, 71.5-80.0 mm LEA), Rosário do Sul, stream tributary of arroio Gueromana, BR290( 30°1’0"S, 55°23’18"W), 26 Nov 2003, L. R. Malabarba, J. Giora, D. Cognato, G. Neves & J. Ferrer. UFRGS 6507, 1(male, 92.9 mm LEA), UFRGS 6509, 1(male, 92.5 mm LEA), UFRGS 6511, 1(male, 98.3 mm LEA), São Gabriel, stream tributary of arroio Piraí, BR290( 30°18’56"S, 54°24’22"W), 26 Nov 2003, L. R. Malabarba, J. Giora, D. Cognato, G. Neves & J. Ferrer. Non-type material.The following specimens were used to record the distribution of the new species, but are not used in the species description and are not part of the typeseries. However, they do not differ in morphometric or meristic characteristics from the typespecimens. Brazil, Rio Grande do Sul: MCP20215, 5, and MCP20217, 3, Eldorado do Sul; MCP20711, 1, Osório; UFRGS 7686, 30, Viamão, and UFRGS 7707, 3, Viamão; UFRGS 8475, 4, Pantano Grande - Rio Pardo; UFRGS 8938, 1, rio Tramandaí; UFRGS 6780, 11, São Gabriel; UFRGS 6750, 14, Rosário do Sul- Alegrete; UFRGS 8263, 28, Charqueadas. Paraguay : USNM 181483, 5, Pueblo Ybytymi, Departamiento Paraguarí; ANSP 170412, 1, Deparatamiento Missiones; ANSP 175180, 1, Departamiento Missiones. Uruguay: ZVC-P 2727, 1, Departamiento Artigas, lagoa Redonda.
Diagnosis. Brachyhypopomus dracocan be distinguished from all described congeners by the extreme broadening of the distal portion of the caudal filament in males during the reproductive period to form a distinct paddle shaped structure ( vs.moderately broadened distally in B. pinnicaudatusand B. brevirostrisinto paddle-shaped structures, and versus broadened along most of the length of the caudal filament in B. occidentalis, B. diazi, B. beebei,and B. janeiroensis). Brachyhypopomus dracocan be further diagnosed from other species of the genus by the following characters: caudal filament length 17.3-35.2% of LEA ( vs.36.5- 50.0% in B. jureiae; 33.0-50.0% in B. janeiroensis, and 13.3- 20.0% in B. bombilla); pectoral fin ray length 4.1-5.8% of LEA ( vs.6.7-7.9% in B. pinnicaudatus); body depth 8.8-12.2% of LEA ( vs.12.1-16.2% in B. pinnicaudatus); snout length 20.7- 30.0% of HL ( vs.31.7-32.2% in B. jureiae); gape width 9.5- 16.5% of HL ( vs.17.8-23.1% in B. pinnicaudatus); interorbital distance 22.7-35% of HL ( vs.14.9-20.5% in B. bombilla); branchial aperture 16.5-26% of HL ( vs.24.9-31.0% in B. pinnicaudatus); head width at operculum 48.4-65% of HL ( vs.60.0-73.0% in B. pinnicaudatus); head width at eyes 31.3- 45.6% of HL ( vs.24.7-31.7% in B. bombilla); number of anal fin rays 155-198 ( vs. 188-211 in B. jureiae; 251-295 in B. brevirostris; 214-228 in B. beebei, and 200-240 in B. occidentalis); upper jaw equal to lower jaw ( vs.upper jaw slightly longer than lower jaw in B. bombilla, B. occidentalis, B. diazi, B. brevirostrisand B. jureiae).
Description.Morphometric and meristic data are presented in Table 1. Head conical, nearly triangular in lateral view, snout short, eyes small, mouth terminal with upper jaw equal to lower jaw, no teeth in both jaws. Body slender, slightly laterally compressed. Dorsal profile straight from snout to supraoccipital, slightly convex posteriorly to caudal filament. Ventral profile gently convex from lower jaw to anal-fin origin, slightly convex to nearly straight along anal-fin base. Highest body depth located posterior to anal-fin origin. Body depth increasing smoothly from head to body region near the 30th anal-fin ray, clearly gradually decreasing from that point to caudal filament. Caudal filament moderately short, slender and laterally compressed in females, juveniles and males in non-reproductive period; caudal filament paddle-shaped in sexually mature males (see sexual dimorphism, below). Cycloid scales covering uniformly the body, except head and fins. Scales smaller on all anterior quarter portion of body; posterior scales at dorsal and medial body regions two to four times larger in diameter than anterior ones; smallest scales covering region of anal-fin pterygiophores. First anterior perforated scale of lateral line above pectoral fin origin and lateral line extending to caudal filament tip, hardly discernible. Branchial aperture small and slightly anterior to pectoral fin origin. Anus with the presence of urogenital papilla in males and females, although less developed in juveniles. Pectoral fins rounded with pigmented rays and perpendicular insertion; pectoral-fin rays i-ii + 13-15 (15-17 total pectoral-fin rays, n = 47, median = 16). Anal fin relatively long with vii-xii + 148- 186 rays (155-198 total anal-fin rays, n = 41, mean = 181.3) which are pigmented. Anal-fin origin located posterior to posterior edge of pectoral fin. Precaudal vertebrae 21-23 (20-22 anterior, 1-2 transitional; n= 6). Coloration in life.General body color brownish varying from dark brown to yellowish brown, dark brown near dorsal mid-line and clearing ventrally. A highly variable number of dark brown bands, well delineated or not, nearly perpendicular or oblique to longitudinal body axis, running posterodorsally from base of anal-fin rays to nearby lateral line and occurring from head to tip of caudal filament. Anal and pectoral-fin rays speckled brown, with hyaline inter-radial membranes.Adults and juveniles exhibit the same coloration pattern. Table 1.Morphometric data for Brachyhypopomus draco. Character Holotype n Minimum Maximum Mean L1 L2 SD 95% confidence limit Length to the end of anal fin (mm) 137.3 44 50 151.9 96.9 24.55 Percents of length to the end of anal fin Head length 10.1 44 9.4 14.2 11.4 11.08 11.69 1.00 Caudal filament length 29.1 34 17.3 35.2 25.1 23.32 26.86 5.15 Snout to anal fin origin 18.3 44 17.9 23.0 20.3 19.93 20.70 1.27 Depth of caudal filament 2.7 44 1.4 3.1 2.1 2.01 2.24 0.38 Longest anal fin ray 3.7 43 2.9 4.4 3.6 3.52 3.73 0.34 Longest pectoral fin ray 4.1 41 4.1 5.8 5.0 4.85 5.16 0.50 Body depth 9.4 44 8.8 12.2 10.8 10.52 11.08 0.92 Percents of head length Snout length 26.5 50 20.7 30.0 25.6 25.12 26.08 1.68 Gape width 14.5 50 9.5 16.5 13.7 13.32 14.14 1.45 Orbital diameter 9.5 50 9.4 14.4 12.3 11.90 12.64 1.30 Interorbital distance 23.4 50 22.7 35.0 28.4 27.60 29.25 2.91 Posterior nare to eye 3.4 49 2.9 5.4 4.2 4.04 4.34 0.52 Branchial aperture 23.3 50 16.5 26.0 19.9 19.31 20.44 1.99 Head width at operculum 54.1 50 48.4 65.0 57.0 55.96 57.96 3.50 Head width at eyes 37.2 50 31.3 45.6 39.9 39.09 40.63 2.71 Secondary sexual dimorphism.During reproductive period, established through monthly variation analysis of male and female gonads in a B. dracopopulation, males undergo hypertrophy of the distal portion of caudal filament ( Fig. 2). This has been observed between August and December in a population studied from the type-locality (A. Schaan, J. Giora and C. Fialho, in preparation). Extremely vertical broadening and lateral compression give the caudal filament a paddle-like shape. After the reproductive period, this structure regresses until the caudal filament resembles those of females and juveniles. In addition, adult males are significantly larger than females. Electric organ discharge. Brachyhypopomus dracogenerates a continuous train of pulse-type EODs. Adult specimens (with developing or fully developed gonads) ( 72.6-105 mmLEA) exhibited the following EOD parameters. The mean EOD repetition rate (per individual) during the day ranged from 15.7-24.6 Hz (mean among all specimens, 19.6 Hz, SD 2.5, n = 11 individuals fishes), with a standard deviation (SD) of 0.4- 1.2 Hz (mean 0.7, SD 0.3, n = 11), and coefficient of variance of 1.8-7.0 % (mean 3.9 %, SD 1.88, n = 11). The lowest and highest absolute pulse rates recorded from all diurnal recordings were 13.7 and 29.2 Hz respectively. During the hours of peak foraging activity, 1-3 hours after sunset, the mean EOD repetition rate (per individual) ranged from 34.2-45.8 Hz (mean among all specimens 38.6 Hz, SD 5.2, n = 6) with a standard deviation of 2.8-10.8 Hz (mean 2.9, SD 1.1, n = 6) and a coefficient of variation of 8.3-27.3 % (mean 20.7 %, n = 6). The lowest and highest absolute pulse rates recorded from all nocturnal recordings were 8.4 and 80.2 Hz respectively. In sum, adult specimens of B. dracoexhibited a distinct increase in pulse rate from the resting day-time state (mean 19.6 Hz) to the nocturnal active state (mean 38.6 Hz). The coefficient of variation of pulse rate during nocturnal activity (mean 20.7 %) was considerably higher than during the day (mean 3.9%) reflecting the greater variability in pulse rate during foraging. Disturbances such as minor vibrations provoked novelty responses, sudden increases in the resting pulse rate from the baseline rate ( Fig. 3). The approximately stable resting diurnal EOD punctuated by novelty responses results in a right-skewed distribution of inter-pulse intervals in a histogram ( Fig. 3). Some specimens of B. dracodisplayed spontaneous complete cessations of the EOD for periods of up to 1 minute. Normal EOD activity was resumed after these interruptions. Fig. 2. Brachyhypopomus draco(holotype, MCP 41540, male, 137.3 mm LEA). Detailed images of head, above, and tail, below. TheEODs waveforms of adult specimens of B. dracocomprised two components of alternating polarity (P1 and P2) sensu Crampton& Albert(2006), and varied in duration from 1.172 -1.992ms (mean 1.578, SD 0.147, n = 23) ( Fig. 4). The Peak Power Frequency( PPF) (see Fig. 4) of these EODs varied from 0.6958 -0.8942kHz (mean 0.7707, SD 0.0534, n = 23). We did not encounter evidence for obvious sexual differences in the EOD waveforms of B. draco, despite evidence for this phenomenon in other species of Brachyhypopomus( e.g. B. occidentalis, Hagedorn, 1985, Shumway & Zelick, 1988; B. pinnicaudatus Stoddard et al., 2003; B. brevirostris, Kawasaki & Heiligenberg, 1989, Crampton, 1996). Where a sexual difference exists, this is manifest as an elongation of the second negative waveform phase, P2 and a corresponding reduction in the PPF of the EOD. We observed no consistent EOD waveform shape differences. Likewise, we observed no significant difference between the PPF of five sexually mature males ( 0.7504 -0.8942kHz, mean 0.8178, SD 0.0675) and 2 sexually mature females ( 0.7385 -0.7949kHz, mean 0.7667, SD 0.0399) (2-sample T-test, df = 5, n = 7, P = 0.38). The observation of a lower PPF in mature females than males contrasts with cases of sexual EOD differences in Brachyhypopomus, where sexually mature males invariably display lower PPFs. Nonetheless, we did note that the Spectral Power Densities of the two sexually mature females exhibited a more rapid decline from the PPF than in the five sexually mature males ( Fig. 4). Males and females with developing gonads also exhibited no obvious difference in waveform shape, and no significant difference in the PPF (males: 0.6958 -0.8362Hz, mean 0.7518, SD 0.0460, n = 10; females: 0.7111 -0.8194kHz, mean 0.7642, SD 0.0422, n = 6) (2-sample T-test, df = 14, n = 16, P = 0.59). All recorded specimens had been held in social isolation for several days before they were recorded. This is known to minimize the hormonally-induced modulation of the P2 phase in sexually mature males ( Stoddard et al., 2003). Further investigation is required to explore whether EOD differences emerge in males that are exposed to normal social stimuli ( i.e.the presence of sexually mature conspecifics). All recordings presented here were made during the southern summer (December) when ambient water temperature is typically in the range 24 – 30º C. During the winter, water temperatures in Rio Grande do Suldecline to as low as 10º C. We noted that these seasonal temperature changes have a significant impact on the waveform duration and shape of the EODs of B. draco. The PPF of the EOD declines substantially, and the P2 phase becomes diminished in relative amplitude. We will present detailed observations on this phenomenon elsewhere. Loureiro & Silva (2006) discuss the effect of temperature on the EODs of B. bombilla.
Distribution. Brachyhypopomus dracois widely known from central, southern and coastal regions of Rio Grande do Sul state, Brazil, and Uruguay. It is known from three drainages: laguna dos Patos, rio Uruguayand rio Tramandaí, and was also found at two localities in Paraguay(ANSP 170412; ANSP 175180; USNM 181483).
Habitat. Brachyhypopomus dracoinhabits river edges, slowmoving creeks, lagoons and flooded areas with muddy or sandy bottom and abundant emergent or floating vegetation. The species was particularly abundant in the typelocality, a lagoon ( 30°22’52.4"S, 51°01’25"W) inside a state preserved area, Parque Estadual de Itapuã in Rio Grande do Sul, Brazil. The lagoon is approximately 4 hectares in area and reaches 1 meterin depth. It has a muddy bottom, abundant macrophytes and vegetal material in various stages of decomposition, and is surrounded by psammophilous forest with strong vegetational influence from Atlantic forest formations ( Fig. 6). At the typelocality, B. dracowas found occurring together with an undescribed Gymnotus(Crampton et al., in prep.).At other collecting sites throughout Rio Grande do Sul statethe species was collected along with the gymnotiforms Eigenmannia trilineata, B. bombillaand another undescribed Brachyhypopomusspecies(see Comparative Material).
Etymology.Name “draco”, from the Greek “drakon” meaning dragon, in reference to the shape of distal portion of caudal filament in mature males, similar to that illustrated in these imaginary creatures.
5107378307
2004-01-12
MCP
D. Cognato & L. Malabarba & C. E. Machado & R. Q. Carvalho
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamão
2
161
Laguna dos Patos
MCP 41540
1
Rio Grande do Sul
holotype
5107378308
2004-01-12
Laguna dos Patos
MCP
D. Cognato & L. Malabarba & C. E. Machado & R. Q. Carvalho
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamão
2
161
MCP 41539
1
Rio Grande do Sul
paratype
5107378319
2006-01-18
Laguna dos Patos
UFRGS
J. Giora & A. P. Dufech & J. Ferrer
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamao
2
161
UFRGS 8888
1
1
Rio Grande do Sul
paratype
5107378333
2004-02-09
Laguna dos Patos
MNRJ
D. Cognato & M. Azevedo & A. Schaan & C. Hiroshi
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamao
2
161
MNRJ 30916
2
1
1
Rio Grande do Sul
paratype
5107378309
2004-06-30
Laguna dos Patos
MCP
D. Cognato & A. Schaan
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamao
2
161
MCP 41538
2
1
1
Rio Grande do Sul
paratype
5107378323
Laguna dos Patos
MNRJ
Brazil
2
161
MNRJ 30917
1
1
Rio Grande do Sul
paratype
5107378305
2004-09-09
MZUSP
D. Cognato & A. Schaan & M. Schossler
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamao
2
161
Laguna dos Patos
MZUSP 94429
1
1
1
Rio Grande do Sul
paratype
5107378341
2003-10-10
Laguna dos Patos
MCP
D. Cognato & A. Schaan
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamao
2
161
MCP 41537
1
1
1
Rio Grande do Sul
paratype
5107378336
2003-11-11
Laguna dos Patos
MNRJ
D. Cognato & J. Giora & J. Anza & A. P. Dufech
Brazil
-30.381222
Parque Estadual de Itapuã, Lagoa Verde
13
-51.02361
Viamao
2
161
MNRJ 30918
25
1
1
Rio Grande do Sul
paratype
5107378324
1980-09-07
Laguna dos Patos
UFRGS
L. R. Malabarba
Brazil
-30.01
arroio Alexandrina
-50.95
Viamao
2
161
UFRGS 732
1
1
Rio Grande do Sul
paratype
5107378320
1979-11-04
Laguna dos Patos
UFRGS
L. R. Malabarba & P. A. Buckup & C. Souto
Brazil
-30.17
açude near BR 290 highway
-51.33
Guaíba
2
161
UFRGS 820
1
1
Rio Grande do Sul
paratype
5107378312
1980-01-18
UFRGS
P. A. Buckup & C. Souto
Brazil
-30.05
açude in BR290 highway, near junction with BR116 highway
-51.35
Guaíba
2
161
UFRGS 822
1
1
Rio Grande do Sul
paratype
5107378329
Laguna dos Patos
UFRGS
Brazil
2
161
UFRGS 6526
1
1
Rio Grande do Sul
paratype
5107378303
2003-12-10
Laguna dos Patos
UFRGS
W. Crampton & L. Malabarba
Brazil
Eldorado do Sul
-30.048
artificial canal near BR290 highway
-51.392
2
161
UFRGS 6525
1
1
Rio Grande do Sul
paratype
5107378330
1988-03-01
Laguna dos Patos
UFRGS
F. Becker & L. R. Malabarba
Brazil
Viamão
-30.25
arroio Itapuã
-51.0333
Viamao
2
161
UFRGS 4317
1
1
Rio Grande do Sul
paratype
5107378317
[816,1459,786,807]
Rio Tramandaí Drainage
UFRGS
Brazil
2
161
UFRGS 6486
1
1
Rio Grande do Sul
paratype
5107378325
[816,1217,817,838]
Rio Tramandaí Drainage
UFRGS
Brazil
2
161
UFRGS 6488
1
1
Rio Grande do Sul
paratype
5107378342
Laguna dos Patos
UFRGS
Brazil
2
161
UFRGS 6489
1
1
Rio Grande do Sul
paratype
5107378326
[983,1377,847,868]
Rio Tramandaí Drainage
UFRGS
Brazil
2
161
UFRGS 6490
1
1
Rio Grande do Sul
paratype
5107378332
2003-11-20
Rio Tramandaí Drainage
UFRGS
L. R. Malabarba & J. Giora & J. Anza & D. Cognato
Brazil
-29.719
creek paralel to Estrada do Mar road
-50.9333
Capão da Canoa
2
161
UFRGS 6491
1
1
Rio Grande do Sul
paratype
5107378334
2003-07-11
Rio Tramandaí Drainage
MNRJ
L. R. Malabarba
Brazil
-30.149
Lagoa Fortaleza
-50.2416
Cidreira
2
161
MNRJ 30915
2
1
1
Rio Grande do Sul
paratype
5107378345
2003-11-20
Rio Tramandaí Drainage
UFRGS
L. R. Malabarba & J. Giora & J. Anza & D. Cognato
Brazil
-29.556
Creek into rio Três Forquilhas along the road RS486
-50.0719
Terra de Arreia
2
161
UFRGS 6671
3
1
2
Rio Grande do Sul
paratype
5107378316
2004-06-18
Rio Uruguay drainage
UFRGS
D. Cognato & W. Crampton & J. Giora & D. Rocha
Brazil
-30.201
Sanga do Jacaré
-55.0547
Rosário do Sul
2
161
UFRGS 6748
1
1
Rio Grande do Sul
paratype
5107378301
Rio Uruguay drainage
MZUSP
Brazil
2
161
MZUSP 94428
8
6
2
Rio Grande do Sul
paratype
5107378338
2004-06-19
Rio Uruguay drainage
UFRGS
D. Cognato & W. Crampton & J. Giora & D. Rocha
Brazil
-30.01666
strea tributary of arroio Gueromana BR290
-55.388
Rosário do Sul
2
161
UFRGS 6753
1
1
Rio Grande do Sul
paratype
5107378328
Rio Uruguay drainage
UFRGS
Brazil
2
161
6785
4
2
2
Rio Grande do Sul
paratype
5107378346
Rio Uruguay drainage
UFRGS
Brazil
2
161
UFRGS 6494
1
1
Rio Grande do Sul
paratype
5107378304
2003-11-26
Rio Uruguay drainage
UFRGS
L. R. Malabarba & J. Giora & D, Cognato & G. Neves & J. Ferrer
-30.211
Sanga do Jacaré
-55.054
Rosário do Sul
2
161
UFRGS 6497
1
1
Rio Grande do Sul
paratype
5107378321
2003-11-26
Rio Uruguay drainage
UFRGS
L. R. Malabarba & J. Giora & D, Cognato & G. Neves & J. Ferrer
Brazil
-30.01666
strea tributary of arroio Gueromana BR290
-55.388
Rosário do Sul
2
161
UFRGS 6789
1
1
Rio Grande do Sul
paratype
5107378339
[816,1189,1607,1628]
Rio Uruguay drainage
UFRGS
Brazil
2
161
UFRGS 6507
1
1
Rio Grande do Sul
paratype
5107378310
UFRGS
Brazil
2
161
UFRGS 6509
1
1
Rio Grande do Sul
paratype
5107378302
2003-11-26
Rio Uruguay drainage
UFRGS
L. R. Malabarba & J. Giora & D, Cognato & G. Neves & J. Ferrer
Brazil
-30.315556
stream tributary of arroio Piraí BR290
-54.406113
São Gabriel
2
161
UFRGS 6511
1
1
Rio Grande do Sul
paratype
5107378311
[930,1402,1881,1902]
MCP
Brazil
2
161
MCP 20215
5
Rio Grande do Sul
5107378331
MCP
Eldorado do Sul
2
161
MCP 20217
3
Rio Grande do Sul
5107378343
[1085,1309,1911,1932]
MCP
Brazil
Osório
2
161
MCP 20711
1
Rio Grande do Sul
5107378337
UFRGS
Brazil
3
162
Viamão
2
161
UFRGS 7686
30
Rio Grande do Sul
5107378318
[286,524,760,781]
UFRGS
Brazil
Viamão
3
162
UFRGS 7707
3
Rio Grande do Sul
5107378340
UFRGS
Brazil
Pantano Grande - Rio Pardo
3
162
UFRGS 8475
4
Rio Grande do Sul
5107378315
[324,627,791,812]
UFRGS
Brazil
rio Tramandai
3
162
UFRGS 8938
1
Rio Grande do Sul
5107378344
UFRGS
Brazil
São Gabriel
3
162
UFRGS 6780
11
Rio Grande do Sul
5107378322
[305,768,821,842]
UFRGS
Brazil
Rosário do Sul - Alegrete
3
162
UFRGS 6750
14
Rio Grande do Sul
5107378313
[129,456,852,873]
UFRGS
Brazil
CHarqueadas
3
162
UFRGS 8263
28
Rio Grande do Sul
5107378306
USNM
Paraguay
Pueblo Ybytymi
3
162
USNM 181483
5
Paraguari
5107378335
ANSP
Paraguay
3
162
ANSP 170412
1
Missiones
5107378314
ANSP
Paraguay
3
162
ANSP 175180
1
Missiones
5107378327
ZVC-P
Uruguay
lagoa Redonda
3
162
ZVC-P 2727
1
Artigas