A new miniature species of <i>Priocharax</i> (Characiformes: Characidae) from the upper rio Ipixuna, Purus drainage, Brazil

A new species of miniature fish of the characid genus Priocharax is described from a small lake near the rio Ipixuna, rio Purus drainage, Amazonas State, Brazil. It is distinguished from all congeners except P. pygmaeus by the lower number of teeth on the maxilla and dentary. It differs from P. pygmaeus by the presence of two postcleithra and 22–27 branched anal-fin rays (vs absence and 19–22). The new species is further distinguished from other species of Priocharax by a combination of characters involving the number of pelvic-fin rays and branched anal-fin rays, the number of postcleithra, the shape of postcleithrum 3, and the absence of the claustrum. Molecular evidence based on COI sequences of all valid species of Priocharax also corroborates the validity of this new species.

Keywords:
Biodiversity; Miniaturization; Ontogenetic truncation; Osteology; Taxonomy

Authorship

George Mendes Taliaferro Mattox

Laboratório de Ictiologia de Sorocaba, Departamento de Biologia, Universidade Federal de São Carlos – campus Sorocaba, Rod. João Leme dos Santos, km 110, Itinga, 18052-780 Sorocaba, SP, Brazil. (GMTM) gmattox@ufscar.br.Universidade Federal de São CarlosBrazilSorocaba, SP, BrazilLaboratório de Ictiologia de Sorocaba, Departamento de Biologia, Universidade Federal de São Carlos – campus Sorocaba, Rod. João Leme dos Santos, km 110, Itinga, 18052-780 Sorocaba, SP, Brazil. (GMTM) gmattox@ufscar.br.

https://orcid.org/0000-0003-4748-472X

Camila da Silva de Souza

Departamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista – campus Botucatu, Rua Prof. Doutor Antonio Celso Wagner Zanin, 250, 18618-689, Botucatu, SP, Brazil. (CSS) camilasvsouza@gmail.com; (CO) claudio@ibb.unesp.br.Universidade Estadual PaulistaBrazilBotucatu, SP, BrazilDepartamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista – campus Botucatu, Rua Prof. Doutor Antonio Celso Wagner Zanin, 250, 18618-689, Botucatu, SP, Brazil. (CSS) camilasvsouza@gmail.com; (CO) claudio@ibb.unesp.br.

https://orcid.org/0000-0002-3878-2373

Mônica Toledo-Piza

Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, 101, 05508-090 São Paulo, SP, Brazil. (MT) mtpiza@usp.br.Universidade de São PauloBrazilSão Paulo, SP, BrazilDepartamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, 101, 05508-090 São Paulo, SP, Brazil. (MT) mtpiza@usp.br.

https://orcid.org/0000-0001-7596-1410

Claudio Oliveira

https://orcid.org/0000-0002-7010-8880

Edited-by

Brian Sidlauskas

Correspondence

George Mendes Taliaferro Mattox gmattox@ufscar.br.

Author's Contribution

George Mendes Taliaferro Mattox: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Validation, Visualization, Writing-original draft, Writing-review and editing.

Camila da Silva de Souza: Data curation, Formal analysis, Investigation, Methodology, Software, Validation, Writing-original draft, Writing-review and editing.

Mônica Toledo-Piza: Conceptualization, Formal analysis, Investigation, Methodology, Supervision, Validation, Writing-original draft, Writing-review and editing.

Claudio Oliveira: Data curation, Funding acquisition, Investigation, Methodology, Resources, Software, Supervision, Validation, Writing-original draft, Writing-review and editing.

Competing Interests

The authors declare no competing interests.

Ethical Statement

Sampling for this study was authorized by Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) through permit number SISBIO/MMA 45429 and it is in accordance with the National Council for the Control of Animal Experimentation (CONCEA) approved by UNESP Ethics Committee on Use of Animals (CEUA), protocol number 1058.

SCIMAGO INSTITUTIONS RANKINGS

Figures | Tables

Figures (9)
Tables (3)

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FIGURE 1 |
Priocharax britzi, A. Holotype, male, MZUSP 125822, 13.7 mm SL; Brazil, Amazonas, Canutama, small lake approximately 400 m from the rio Ipixuna, after bridge crossing this river, on the right side of the Transamazônica Road (BR–230) going from Humaitá to Lábrea, rio Ipixuna drainage. B. Live paratype photographed immediately after capture.

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FIGURE 2 |
Priocharax britzi, paratypes, MZUSP 125823, c&s; anterior region of head in lateral view detailing the orbital region. A. 12.5 mm SL. B. 12.9 mm SL, image flipped. Antorbital and infraorbitals delimited by black line for clarity. Ant = antorbital; IO2 – 3 = infraorbitals 2 – 3. Scale bar = 0.2 mm.

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FIGURE 3 |
Priocharax britzi, paratype, MZUSP 125823, 12.9 mm SL, c&s; jaws in lateral view. Ana = anguloarticular; Cm = coronomeckelian; De = dentary; MC = Meckel’s cartilage; Mx = maxilla; Pmx = premaxilla; Ra = retroarticular. Scale bar = 0.2 mm.

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FIGURE 4 |
Shoulder girdle in lateral view of A. Priocharax britzi, paratype, MZUSP 125823, 12.9 mm SL, c&s. Image flipped. B. Priocharax cf. ariel, UFRO 15521, 11.8 mm SL, c&s. Cl = cleithrum; PecRdC = pectoral-fin radial cartilage; Pcl = unidentified postcleithrum; Pcl3 = postcleithrum 3; Pt = posttemporal; ScCoC = scapulocoracoid cartilage; Sucl = supracleithrum. Arrows point to middle fissure on pectoral-fin radial cartilage. Pcl3 slightly displaced ventrally from original position in A. Scale bars = 0.5 mm.

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FIGURE 5 |
Priocharax britzi, paratype, MZUSP 125823, 12.9 mm SL, c&s; Weberian apparatus in lateral view. Exoc = exoccipital; Int = intercalarium; NA3 – 4 = neural arches 3 – 4; NS4 = neural spine 4; OsS = os suspensorium; Sca = scaphium; Sn3 = supraneural 3; Soc = supraoccipital; T = tripus. Scale bar = 0.2 mm.

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FIGURE 6 |
Priocharax britzi, holotype, MZUSP 125822, 13.7 mm SL. A. Left lateral view of anal fin showing bony hooks. B. Ventral view of pelvic fin showing bony hooks. Scale bars = 0.5 mm.

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FIGURE 7 |
Maximum likelihood tree of five species of Priocharax, based on the COI gene (642 bp). Bars represent the number of species obtained by the ABGD and bPTP analyses. Numbers near nodes represent bootstrap values.

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FIGURE 8 |
Map of South America with a close-up of the upper and central portions of the Amazon basin, Brazil, illustrating the distribution of the five species of Priocharax: P. britzi described herein from the rio Purus basin (green losangle), P. varii (red dot), P. ariel (black dots), P. nanus (blue dot), and P. pygmaeus (yellow dots). Colors are the same as in the Fig. 7. Some dots may represent more than one lot.

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FIGURE 9 |
Small lake approximately 400 m from the rio Ipixuna, Canutama, Amazonas, Brazil, the type locality of Priocharax britzi.

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TABLE 1 |
Species, lots, vouchers, basins, locality information, and GenBank accession numbers of samples used in this study. In the locality column, Brazilian states are abbreviated as follows: AM = Amazonas, RO = Roraima.

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TABLE 2 |
Morphometric data of Priocharax britzi. N = number of specimens; SD = standard deviation. Range includes the holotype. [Note: In the recent description of Priocharax variiMattox et al., 2020Mattox GMT, Souza CS, Toledo-Piza M, Britz R, Oliveira C. A new miniature species of Priocharax (Teleostei: Characiformes: Characidae) from the Rio Madeira drainage, Brazil, with comments on the adipose fin in characiforms. Vertebr Zool. 2020; 70(3):417–33. http://dx.doi.org/10.26049/VZ70-3-2020-11
http://dx.doi.org/10.26049/VZ70-3-2020-1... :423, table 2), data of anal-fin length is incorrect. The correct information is as follows: Holotype = 22, range = 20–24, mean = 21.8, SD = 0.9, with n = 48 including the holotype].

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TABLE 3 |
Pairwise K2P genetic distances (and standard deviation) within and among species of Priocharax. Intraspecific genetic distances are highlighted in bold. Values are presented in percentages followed by standard deviation. Number of sequences: P. ariel = 6, P. britzi = 5, P. nanus = 1, P. pygmaeus = 9 and P. varii = 4.

FIGURE 1 | Priocharax britzi, A. Holotype, male, MZUSP 125822, 13.7 mm SL; Brazil, Amazonas, Canutama, small lake approximately 400 m from the rio Ipixuna, after bridge crossing this river, on the right side of the Transamazônica Road (BR–230) going from Humaitá to Lábrea, rio Ipixuna drainage. B. Live paratype photographed immediately after capture.

FIGURE 2 | Priocharax britzi, paratypes, MZUSP 125823, c&s; anterior region of head in lateral view detailing the orbital region. A. 12.5 mm SL. B. 12.9 mm SL, image flipped. Antorbital and infraorbitals delimited by black line for clarity. Ant = antorbital; IO2 – 3 = infraorbitals 2 – 3. Scale bar = 0.2 mm.

FIGURE 3 | Priocharax britzi, paratype, MZUSP 125823, 12.9 mm SL, c&s; jaws in lateral view. Ana = anguloarticular; Cm = coronomeckelian; De = dentary; MC = Meckel’s cartilage; Mx = maxilla; Pmx = premaxilla; Ra = retroarticular. Scale bar = 0.2 mm.

FIGURE 4 | Shoulder girdle in lateral view of A. Priocharax britzi, paratype, MZUSP 125823, 12.9 mm SL, c&s. Image flipped. B. Priocharax cf. ariel, UFRO 15521, 11.8 mm SL, c&s. Cl = cleithrum; PecRdC = pectoral-fin radial cartilage; Pcl = unidentified postcleithrum; Pcl3 = postcleithrum 3; Pt = posttemporal; ScCoC = scapulocoracoid cartilage; Sucl = supracleithrum. Arrows point to middle fissure on pectoral-fin radial cartilage. Pcl3 slightly displaced ventrally from original position in A. Scale bars = 0.5 mm.

FIGURE 5 | Priocharax britzi, paratype, MZUSP 125823, 12.9 mm SL, c&s; Weberian apparatus in lateral view. Exoc = exoccipital; Int = intercalarium; NA3 – 4 = neural arches 3 – 4; NS4 = neural spine 4; OsS = os suspensorium; Sca = scaphium; Sn3 = supraneural 3; Soc = supraoccipital; T = tripus. Scale bar = 0.2 mm.

FIGURE 6 | Priocharax britzi, holotype, MZUSP 125822, 13.7 mm SL. A. Left lateral view of anal fin showing bony hooks. B. Ventral view of pelvic fin showing bony hooks. Scale bars = 0.5 mm.

FIGURE 7 | Maximum likelihood tree of five species of Priocharax, based on the COI gene (642 bp). Bars represent the number of species obtained by the ABGD and bPTP analyses. Numbers near nodes represent bootstrap values.

FIGURE 8 | Map of South America with a close-up of the upper and central portions of the Amazon basin, Brazil, illustrating the distribution of the five species of Priocharax: P. britzi described herein from the rio Purus basin (green losangle), P. varii (red dot), P. ariel (black dots), P. nanus (blue dot), and P. pygmaeus (yellow dots). Colors are the same as in the Fig. 7. Some dots may represent more than one lot.

FIGURE 9 | Small lake approximately 400 m from the rio Ipixuna, Canutama, Amazonas, Brazil, the type locality of Priocharax britzi.

TABLE 1 | Species, lots, vouchers, basins, locality information, and GenBank accession numbers of samples used in this study. In the locality column, Brazilian states are abbreviated as follows: AM = Amazonas, RO = Roraima.

Species	Lot	Voucher	Basin	Locality	Coordinates	GenBank n.
Priocharax varii	LBP 28495	96981	Rio Madeira	Rio Preto, affluent of rio Jamari, Candeias do Jamari, RO, Brazil	08°52’53.5”S 63°37’50.8”W	MT754786
Priocharax varii	LBP 28495	96982	Rio Madeira	Rio Preto, affluent of rio Jamari, Candeias do Jamari, RO, Brazil	08°52’53.5”S 63°37’50.8”W	MT754785
Priocharax varii	LBP 28495	96984	Rio Madeira	Rio Preto, affluent of rio Jamari, Candeias do Jamari, RO, Brazil	08°52’53.5”S 63°37’50.8”W	MT754783
Priocharax varii	LBP 28495	96985	Rio Madeira	Rio Preto, affluent of rio Jamari, Candeias do Jamari, RO, Brazil	08°52’53.5”S 63°37’50.8”W	MT754784
Priocharax ariel	LBP 28442	98284	Rio Negro	Igarapé Tibarrá on left side of rio Negro, Santa Isabel do Rio Negro, AM, Brazil	00º26’28.1”S 64º56’57.5”W	MT754780
Priocharax ariel	LBP 28442	98285	Rio Negro	Igarapé Tibarrá on left side of rio Negro, Santa Isabel do Rio Negro, AM, Brazil	00º26’28.1”S 64º56’57.5”W	MT754781
Priocharax ariel	LBP 27704	98286	Rio Negro	Igarapé Tapage, rio Urubaxi, approximatelly 1 h from mouth of river, S. I. Rio Negro, AM, Brazil	00º30’05.3”S 64º49’11.7”W	MT754778
Priocharax ariel	LBP 27704	98287	Rio Negro	Igarapé Tapage, rio Urubaxi, approximatelly 1 h from mouth of river, S. I. Rio Negro, AM, Brazil	00º30’05.3”S 64º49’11.7”W	MT754782
Priocharax ariel	LBP 27704	98288	Rio Negro	Igarapé Tapage, rio Urubaxi, approximatelly 1 h from mouth of river, S. I. Rio Negro, AM, Brazil	00º30’05.3”S 64º49’11.7”W	MT754779
Priocharax ariel	LBP 25858	96383	Rio Negro	Igarapé Uacatuna, São Gabriel da Cachoeira, AM, Brazil	00°03'38.0''S 67°05'45.0''W	MT754777
Priocharax nanus	LBP 28490	98283	Rio Negro	Igarapé Tibarrá on left side of rio Negro, Santa Isabel do Rio Negro, AM, Brazil	00º26’28.1”S 64º56’57.5”W	MT754766
Priocharax pygmaeus	LBP 22464	96986	Rio Amazonas	Quebrada La Ponderosa, Letícia, Colombia	04°08’24.4’’S 69°56’53.4’’W	MT754771
Priocharax pygmaeus	LBP 22464	96987	Rio Amazonas	Quebrada La Ponderosa, Letícia, Colombia	04°08’24.4’’S 69°56’53.4’’W	MT754774
Priocharax pygmaeus	LBP 22464	96988	Rio Amazonas	Quebrada La Ponderosa, Letícia, Colombia	04°08’24.4’’S 69°56’53.4’’W	MT754769
Priocharax pygmaeus	LBP 22464	96998	Rio Amazonas	Quebrada La Ponderosa, Letícia, Colombia	04°08’24.4’’S 69°56’53.4’’W	MT754768
Priocharax pygmaeus	LBP 22739	96989	Rio Amazonas	Quebrada Pichuna, Letícia, Colombia	04°07’33.8’’S 70°00’28.9’’W	MT754772
Priocharax pygmaeus	LBP 22739	96990	Rio Amazonas	Quebrada Pichuna, Letícia, Colombia	04°07’33.8’’S 70°00’28.9’’W	MT754773
Priocharax pygmaeus	LBP 22739	96991	Rio Amazonas	Quebrada Pichuna, Letícia, Colombia	04°07’33.8’’S 70°00’28.9’’W	MT754776
Priocharax pygmaeus	LBP 22739	96992	Rio Amazonas	Quebrada Pichuna, Letícia, Colombia	04°07’33.8’’S 70°00’28.9’’W	MT754770
Priocharax pygmaeus	LBP 22739	96993	Rio Amazonas	Quebrada Pichuna, Letícia, Colombia	04°07’33.8’’S 70°00’28.9’’W	MT754775
Priocharax britzi	LBP 28493	98295	Rio Purus	Marginal lake to rio Ipixuna, Canutama, AM, Brazil	07°31’11.5’’S 63°20’59.6’’W	MW374298
Priocharax britzi	LBP 28493	98296	Rio Purus	Marginal lake to rio Ipixuna, Canutama, AM, Brazil	07°31’11.5’’S 63°20’59.6’’W	MW374297
Priocharax britzi	LBP 28493	98297	Rio Purus	Marginal lake to rio Ipixuna, Canutama, AM, Brazil	07°31’11.5’’S 63°20’59.6’’W	MW374296
Priocharax britzi	LBP 28493	98298	Rio Purus	Marginal lake to rio Ipixuna, Canutama, AM, Brazil	07°31’11.5’’S 63°20’59.6’’W	MW374300
Priocharax britzi	LBP 28493	98299	Rio Purus	Marginal lake to rio Ipixuna, Canutama, AM, Brazil	07°31’11.5’’S 63°20’59.6’’W	MW374299
Galeocharax humeralis	MAG ICT 0206	LAR217	Rio Paraná	Entre Rios, Rosario, Argentina	32°51’46.8’’S 60°38’27.6’’W	KU288918

TABLE 2 | Morphometric data of Priocharax britzi. N = number of specimens; SD = standard deviation. Range includes the holotype. [Note: In the recent description of Priocharax *varii***Mattox et al., 2020**Mattox GMT, Souza CS, Toledo-Piza M, Britz R, Oliveira C. A new miniature species of Priocharax (Teleostei: Characiformes: Characidae) from the Rio Madeira drainage, Brazil, with comments on the adipose fin in characiforms. Vertebr Zool. 2020; 70(3):417–33. http://dx.doi.org/10.26049/VZ70-3-2020-11
http://dx.doi.org/10.26049/VZ70-3-2020-1... :423, table 2), data of anal-fin length is incorrect. The correct information is as follows: Holotype = 22, range = 20–24, mean = 21.8, SD = 0.9, with n = 48 including the holotype].

	Holotype	N	Range	Mean	SD
Standard length (mm)	13.7	48	9.3–14.1	11.8	-
Percentages of standard length
Depth at dorsal-fin origin	23	48	20–24	22.1	0.9
Snout to dorsal-fin origin	53	48	52–56	53.9	0.9
Snout to pelvic-fin origin	38	48	38–43	39.9	1.0
Snout to anal-fin origin	52	48	49–55	51.8	1.0
Dorsal-fin length	28	46	23–28	25.4	1.0
Dorsal-fin base	12	48	9–14	11.7	1.0
Pelvic-fin length	12	47	9–12	10.3	0.7
Anal-fin length	22	46	21–25	22.8	1.0
Anal-fin base	35	45	30–37	34.3	1.3
Caudal-peduncle depth	8	47	7–8	7.5	0.4
Caudal-peduncle length	15	45	13–19	15.7	1.2
Head length	25	48	23–27	25.3	0.9
Percentages of head length
Orbital diameter	35	46	33–39	36.2	1.2
Interorbital distance	40	44	30–42	35.9	2.6
Snout length	19	46	16–22	19.5	1.2
Upper jaw length	50	47	46–58	50.9	2.7
Percentages of caudal peduncle length
Caudal-peduncle depth	53	45	39–56	48.4	3.6
Percentages of orbital diameter
Snout length	56	45	45–59	53.6	3.8

TABLE 3 | Pairwise K2P genetic distances (and standard deviation) within and among species of Priocharax. Intraspecific genetic distances are highlighted in bold. Values are presented in percentages followed by standard deviation. Number of sequences: P. ariel = 6, P. britzi = 5, P. nanus = 1, P. pygmaeus = 9 and P. varii = 4.

	1	2	3	4	5
1- P. nanus	-
2- P. ariel	18.8±2.3	0.2±0.1
3- P. pygmaeus	18.2±2.1	24.9±2.5	0.0±0.0
4- P. varii	17.9±2.1	19.3±2.0	20.2±2.1	0.2±0.1
5- P. britzi	18.6±2.2	21.0±2.1	19.7±2.1	20.4±2.0	0.1±0.1

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Sociedade Brasileira de Ictiologia Neotropical Ichthyology, Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Universidade Estadual de Maringá., Av. Colombo, 5790, 87020-900, Phone number: +55 44-3011-4632 - Maringá - PR - Brazil
E-mail: neoichth@nupelia.uem.br

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A new miniature species of <i>Priocharax</i> (Characiformes: Characidae) from the upper rio Ipixuna, Purus drainage, Brazil

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[1] Edited-by

Brian Sidlauskas