A new species of <i>Aequidens</i> (Cichliformes: Cichlidae) from the rio Paraguai basin, Brazil

Morphological and molecular data support the description of a new Aequidens species from the upper rio Correntes, considered herein as endemic to the upper rio Paraguai basin in the Cerrado biome in Brazil. The new species is distinguished from all congeners, except from A. plagiozonatus by having anteriorly oblique dark brown flank bars vs. vertical flank bars, and is additionally distinguished from some congeners by showing a discontinuous lateral band and presence of a dark cheek spot. The new species differs from Aequidensplagiozonatus by having the profile of the dorsal part of head almost straight (in lateral view), with a conspicuous concavity at the interorbital, and by the longer length of upper and lower jaws. Furthermore, delimitation analyses based on mitochondrial data provide additional support for the validity of the species. Our study data also revealed the occurrence, and consequently the first record, of A.plagiozonatus in the upper rio Araguaia basin, which was most likely driven by headwater capture events.

Keywords:
Cerrado biome; DNA barcode; Molecular data; Morphological data; Species delimitation

Authorship

Rianne Caroline de Oliveira

Formal analysis

Writing-original draft

Writing-review and editing

Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Centro de Ciências Biológicas, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900 Maringá, PR, Brazil. (RCO) rianne.oliveira@gmail.com (corresponding author), (GCD) gabrieldepra@gmail.com, (WJG) weferson@nupelia.uem.br.Universidade Estadual de MaringáBrazilMaringá, PR, BrazilNúcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Centro de Ciências Biológicas, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900 Maringá, PR, Brazil. (RCO) rianne.oliveira@gmail.com (corresponding author), (GCD) gabrieldepra@gmail.com, (WJG) weferson@nupelia.uem.br.

Programa de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Departamento de Biologia, Centro de Ciências Biológicas, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900 Maringá, PR, Brazil.Universidade Estadual de MaringáBrazilMaringá, PR, BrazilPrograma de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Departamento de Biologia, Centro de Ciências Biológicas, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900 Maringá, PR, Brazil.

https://orcid.org/0000-0001-7832-8522

Luiz Fernando Caserta Tencatt

Conceptualization

Formal analysis

Funding acquisition

Project administration

Writing-original draft

Writing-review and editing

Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Av. Fernando Corrêa da Costa, 2367, Boa Esperança, 78060-900 Cuiabá, MT, Brazil. (LFCT) luiztencatt@hotmail.com.Universidade Federal de Mato GrossoBrazilCuiabá, MT, BrazilDepartamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Av. Fernando Corrêa da Costa, 2367, Boa Esperança, 78060-900 Cuiabá, MT, Brazil. (LFCT) luiztencatt@hotmail.com.

https://orcid.org/0000-0002-8437-4354

Gabriel de Carvalho Deprá

Formal analysis

Writing-original draft

Writing-review and editing

https://orcid.org/0000-0002-5520-9763

Ricardo Britzke

Formal analysis

Project administration

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Writing-review and editing

Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Av. Gral. Antonio Alvarez de Arenales 1256, Jesús María 15072, Lima, Peru. (RB) rbritzke@unmsm.edu.pe.Universidad Nacional Mayor de San MarcosPeruLima, , PeruMuseo de Historia Natural, Universidad Nacional Mayor de San Marcos, Av. Gral. Antonio Alvarez de Arenales 1256, Jesús María 15072, Lima, Peru. (RB) rbritzke@unmsm.edu.pe.

https://orcid.org/0000-0002-1042-2544

Claudio Oliveira

Formal analysis

Funding acquisition

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Writing-original draft

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Departamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista, R. Prof. Dr. Antonio C. W. Zanin 250, 18618-689 Botucatu, SP, Brazil. (CO) claudio.oliveira@unesp.br.Universidade Estadual PaulistaBrazilBotucatu, SP, BrazilDepartamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista, R. Prof. Dr. Antonio C. W. Zanin 250, 18618-689 Botucatu, SP, Brazil. (CO) claudio.oliveira@unesp.br.

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

Weferson Júnio da Graça

Conceptualization

Formal analysis

Funding acquisition

Project administration

Writing-original draft

Writing-review and editing

Programa de Pós-Graduação em Biologia Comparada, Centro de Ciências Biológicas, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900 Maringá, PR, BrazilUniversidade Estadual de MaringáBrazilMaringá, PR, BrazilPrograma de Pós-Graduação em Biologia Comparada, Centro de Ciências Biológicas, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900 Maringá, PR, Brazil

https://orcid.org/0000-0002-8925-5629

Edited-by

Hernán López-Fernández

Correspondence

Rianne Caroline de Oliveira rianne.oliveira@gmail.com

Competing Interests

The author declares no competing interests.

SCIMAGO INSTITUTIONS RANKINGS

Figures | Tables

Figures (9)
Tables (2)

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FIGURE 1 |
Aequidens pirilampo, NUP 23543, 94.7 mm SL, holotype, Brazil, State of Mato Grosso, municipality of Itiquira, Comprido Creek, tributary of Correntes River, rio Paraguai basin.

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FIGURE 2 |
Head profile (A) convex in Aequidens plagiozonatus, NUP 13372, 76.6 mm SL, horizontally inverted, and (B) triangular in A. pirilampo, NUP 23543, 94.7 mm SL holotype. Arrow indicates the groove between the eyes.

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FIGURE 3 |
Aequidens pirilampo, living specimens photographed just after capture in the rio Comprido, rio Paraguai River basin, 17°32’04.8”S 54°25’36.6”W, uncatalogued. Photos by L. F. C. Tencatt.

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FIGURE 4 |
Coloration and ontogenetic variation in Aequidens pirilampo, NUP 23544, (A) 87.2 mm SL, (B) 65.7 mm SL, (C) 24.6 mm SL, (D) 22.8 mm SL, (E) 22.5 mm SL, and (F) 14.6 mm SL, collected with holotype. Bar 9 includes the infraorbital bar in smaller specimens (C-F).

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FIGURE 5 |
Ceratobranchials 5 (lower pharyngeal jaw) of Aequidens pirilampo, NUP 23547, paratype, 59.2 mm SL, in the occlusal plane, with anterior portion upwards.

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FIGURE 6 |
Partial map of South America showing the distribution of Aequidens pirilampo, in the rio Paraguai basin (yellow shade), Brazil. Purple circle, type-locality in the córrego Comprido, rio Correntes basin; red diamonds, additional localities, rio Correntes basin; orange triangle, sinkhole; black star, locality of A. plagiozonatus in the rio Boiadeiro, rio Araguaia basin (red shade).

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FIGURE 7 |
Collecting sites of Aequidens pirilampo, showing (A) the ribeirão Comprido, its type-locality, (B) a stream with unknown name, and (C) the córrego de Cima, all tributaries of the rio Correntes in the border region of Mato Grosso and Mato Grosso do Sul states, Central Brazil. Photo (A) by LFCT, (B) by Heriberto Gimênes Jr., and (C) by Hans Evers.

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FIGURE 8 |
General morphology and color pattern of Aequidens plagiozonatus of (A) an uncatalogued specimen captured in the Ponte de Pedra Reservoir, rio Correntes, rio Paraguai basin, a couple of days after preservation, and (B) an uncatalogued living specimen captured in the rio Boiadeiro/córrego Gordura, rio Araguaia basin, both specimens with about 70.0 mm SL. Photo (A) by LFCT and (B) by Hans Evers.

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FIGURE 9 |
Maximum-likelihood tree based on the cytochrome oxidase c subunit I gene (COI) partial sequence and the species delimitation analyses evidencing the Aequidens pirilampo, clade (rio Paraguai basin) and the presence of A. plagiozonatus in the rio Paraguai (rios Taquari and Itiquira), and rio Araguaia basin (rio Boiadeiro). Morph.: morphological analysis. Notes labeled with numbers represent bootstrap support.

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TABLE 1 |
Morphometric data of Aequidens pirilampo. N = total number of specimens examined; SD = Standard deviation.

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TABLE 2 |
Interspecific Kimura 2-parameter mean distance of Aequidens and Cichlasoma species (below diagonal) and standard errors (above diagonal) were obtained by a bootstrap procedure.

FIGURE 1 | Aequidens pirilampo, NUP 23543, 94.7 mm SL, holotype, Brazil, State of Mato Grosso, municipality of Itiquira, Comprido Creek, tributary of Correntes River, rio Paraguai basin.

FIGURE 2 | Head profile (A) convex in Aequidens plagiozonatus, NUP 13372, 76.6 mm SL, horizontally inverted, and (B) triangular in A. pirilampo, NUP 23543, 94.7 mm SL holotype. Arrow indicates the groove between the eyes.

FIGURE 3 | Aequidens pirilampo, living specimens photographed just after capture in the rio Comprido, rio Paraguai River basin, 17°32’04.8”S 54°25’36.6”W, uncatalogued. Photos by L. F. C. Tencatt.

FIGURE 4 | Coloration and ontogenetic variation in Aequidens pirilampo, NUP 23544, (A) 87.2 mm SL, (B) 65.7 mm SL, (C) 24.6 mm SL, (D) 22.8 mm SL, (E) 22.5 mm SL, and (F) 14.6 mm SL, collected with holotype. Bar 9 includes the infraorbital bar in smaller specimens (C-F).

FIGURE 5 | Ceratobranchials 5 (lower pharyngeal jaw) of Aequidens pirilampo, NUP 23547, paratype, 59.2 mm SL, in the occlusal plane, with anterior portion upwards.

FIGURE 6 | Partial map of South America showing the distribution of Aequidens pirilampo, in the rio Paraguai basin (yellow shade), Brazil. Purple circle, type-locality in the córrego Comprido, rio Correntes basin; red diamonds, additional localities, rio Correntes basin; orange triangle, sinkhole; black star, locality of A. plagiozonatus in the rio Boiadeiro, rio Araguaia basin (red shade).

FIGURE 7 | Collecting sites of Aequidens pirilampo, showing (A) the ribeirão Comprido, its type-locality, (B) a stream with unknown name, and (C) the córrego de Cima, all tributaries of the rio Correntes in the border region of Mato Grosso and Mato Grosso do Sul states, Central Brazil. Photo (A) by LFCT, (B) by Heriberto Gimênes Jr., and (C) by Hans Evers.

FIGURE 8 | General morphology and color pattern of Aequidens plagiozonatus of (A) an uncatalogued specimen captured in the Ponte de Pedra Reservoir, rio Correntes, rio Paraguai basin, a couple of days after preservation, and (B) an uncatalogued living specimen captured in the rio Boiadeiro/córrego Gordura, rio Araguaia basin, both specimens with about 70.0 mm SL. Photo (A) by LFCT and (B) by Hans Evers.

FIGURE 9 | Maximum-likelihood tree based on the cytochrome oxidase c subunit I gene (COI) partial sequence and the species delimitation analyses evidencing the Aequidens pirilampo, clade (rio Paraguai basin) and the presence of A. plagiozonatus in the rio Paraguai (rios Taquari and Itiquira), and rio Araguaia basin (rio Boiadeiro). Morph.: morphological analysis. Notes labeled with numbers represent bootstrap support.

TABLE 1 | Morphometric data of Aequidens pirilampo. N = total number of specimens examined; SD = Standard deviation.

	Holotype	Paratypes
	Holotype	N	Range	Mean	SD
Standard length (mm)	94.7	26	40.1–94.0	60.0	–
Percents of standard length
Body depth	43.7	26	41.9–46.6	44.2	1.4
Preanal distance	74.1	26	72.8–79.4	76.4	1.6
Prepelvic distance	45.5	26	43.9–48.5	46.4	1.1
Prepectoral distance	39.9	26	37.7–44.4	41.3	1.6
Predorsal distance	43.4	26	42.6–48.8	46.0	1.7
Distance from dorsal to caudal fin	70.2	26	63.9–68.8	66.2	1.2
Distance from dorsal to anal fin	56.6	26	52.9–57.6	55.5	1.2
Distance from dorsal to pelvic fin	42.8	26	42.2–46.6	43.9	1.2
Distance from dorsal to pectoral fin	26.5	26	26.5–31.4	28.4	1.2
Caudal peduncle depth	16.9	26	15.5–17.6	16.7	0.5
Caudal peduncle length (straight)	7.7	26	4.9–8.0	6.0	0.7
Caudal peduncle length (oblique)	12.0	26	10.2–12.3	11.0	0.5
Pectoral-fin length	30.6	25	28.5–33.6	31.0	1.4
Pelvic-fin length	32.8	26	28.1–39.6	32.2	2.7
Pelvic spine length	12.2	26	11.9–15.7	13.5	1.0
Dorsal-fin base length (spine)	42.1	26	40.3–46.9	42.8	1.6
Dorsal-fin base length (total)	61.3	26	54.6–61.4	58.6	1.6
Last dorsal-fin spine length	11.8	26	11.5–15.1	13.0	1.0
Anal-fin base length (spine)	5.2	26	4.2–6.5	5.3	0.5
Anal-fin base length (total)	19.5	26	16.7–19.6	18.2	0.8
Last anal-fin spine length	12.9	26	11.4–15.6	12.9	0.9
Head length	39.1	26	37.8–43.0	40.5	1.3
Head depth	38.9	26	34.7–39.9	37.1	1.7
Head width	18.7	26	17.6–20.9	19.3	0.9
Orbital diameter	9.0	26	10.5–15.0	12.7	1.3
Postorbital head length	14.9	26	14.4–15.5	14.9	0.3
Interorbital distance	13.1	26	9.2–19.2	11.0	1.8
Snout length	15.7	26	11.6–15.2	13.7	0.9
Cheek depth	11.0	26	8.7–11.8	10.5	0.8
Lachrymal depth	8.2	26	5.0–7.8	6.7	0.8
Upper jaw length	14.2	26	12.2–15.1	13.6	0.8
Lower jaw length	17.5	26	16.8–19.2	17.9	0.6
Midlateral spot length	9.4	26	7.9–10.7	9.5	0.7
Percents of head length
Head depth	99.5	26	69.8–100.5	91.5	4.8
Head width	47.8	26	45.2–51.1	47.6	1.7
Orbital width	23.0	26	23.0–36.5	31.4	2.6
Postorbital	38.1	26	35.3–39.2	36.8	0.9
Interorbital width	33.4	26	23.0–46.6	27.2	4.5
Snout length	40.2	26	24.9–40.2	33.7	2.2
Cheek depth	28.2	26	21.4–31.1	25.9	2.3
Lacrimal depth	20.9	26	12.4–20.9	16.5	2.3
Upper jaw length	36.3	26	29.7–38.0	33.6	2.3
Lower jaw length	44.7	26	40.2–46.9	44.1	1.6

TABLE 2 | Interspecific Kimura 2-parameter mean distance of Aequidens and Cichlasoma species (below diagonal) and standard errors (above diagonal) were obtained by a bootstrap procedure.

	Species	1	2	3	4	5	6	7	8	9	10	11	12	13
1	Aequidens pirilampo		0.0096	0.0246	0.0215	0.0157	0.0156	0.0235	0.0227	0.0287	0.0213	0.0195	0.0218	0.0238
2	Aequide plagiozonatus	0.0307		0.0257	0.0213	0.0156	0.0155	0.0230	0.0214	0.0286	0.0211	0.0195	0.0219	0.0228
3	Aequidens diadema	0.1752	0.1861		0.0198	0.0218	0.0231	0.0082	0.0208	0.0314	0.0235	0.0239	0.0222	0.0256
4	Aequidens gerciliae	0.1480	0.1477	0.1274		0.0210	0.0217	0.0182	0.0112	0.0274	0.0200	0.0191	0.0199	0.0218
5	Aequidens tetramerus	0.0892	0.0890	0.1552	0.1465		0.0072	0.0204	0.0191	0.0265	0.0194	0.0182	0.0209	0.0206
6	Aequidens epae	0.0857	0.0855	0.1621	0.1483	0.0242		0.0214	0.0201	0.0271	0.0206	0.0194	0.0215	0.0214
7	Aequidens pallidus	0.1744	0.1714	0.0407	0.1216	0.1498	0.1532		0.0189	0.0269	0.0218	0.0216	0.0208	0.0239
8	Aequidens michaeli	0.1644	0.1575	0.1433	0.0467	0.1322	0.1355	0.1358		0.0261	0.0185	0.0181	0.0211	0.0206
9	Pterophyllum altum (outgroup)	0.2383	0.2350	0.2653	0.2263	0.2181	0.2239	0.2271	0.2119		0.0276	0.0275	0.0281	0.0289
10	Cichlasoma paranaense	0.1530	0.1493	0.1710	0.1328	0.1286	0.1364	0.1652	0.1214	0.2300		0.0079	0.0169	0.0192
11	Cichlasoma dimerus	0.1340	0.1305	0.1739	0.1235	0.1164	0.1242	0.1590	0.1173	0.2275	0.0264		0.0181	0.0186
12	Cichlasoma sanctifranciscense	0.1567	0.1597	0.1581	0.1344	0.1439	0.1466	0.1553	0.1489	0.2254	0.1038	0.1140		0.0192
13	Cichlasoma orientale	0.1845	0.1702	0.2023	0.1591	0.1491	0.1568	0.1935	0.1493	0.2348	0.1280	0.1206	0.1206

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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 species of <i>Aequidens</i> (Cichliformes: Cichlidae) from the rio Paraguai basin, Brazil

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[1] Edited-by

Hernán López-Fernández