A new species of Moenkhausia (Characiformes: Acestrorhamphidae) from the Amazon basin at Serra do Cachimbo, Pará, Brazil

José Igor da Silva¹, Yasmim de Santana² and Manoela Maria Ferreira Marinho²

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Associate Editor: George Mattox

Section Editor: Bruno Melo

Editor-in-chief: José Birindelli

Abstract​

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Introduction​

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The genus Moenkhausia Eigenmann, 1903 comprises a diverse assemblage of small-sized characiform fishes, characterized by a wide range of morphological forms, body sizes, and vivid coloration, and broadly distributed throughout major South American River systems (Fricke et al., 2025). The genus is traditionally defined by a combination of features, including the presence of two rows of premaxillary teeth, with five teeth in the inner row, completely pored lateral line, and cau-dal-fin partially scaled (Eigenmann, 1917). However, such pre-cladistic definition resulted in a large, non-monophyletic group encompassing about 90 valid species (Melo et al., 2024; Fricke et al., 2025). Phylogenetic studies have confirmed the polyphyletic nature of the Moenkhausia (Mirande, 2010, 2019; Mariguela et al., 2013) and have made significant changes in its classification (Melo et al., 2024).

Recently, Moenkhausia was transferred to the Acestrorhamphidae, a family resurrected by Melo et al. (2024) to name a large clade of species formerly classified as Characidae. Also, eleven species previously assigned to Moenkhausia were transferred to the genera Bario Myers, 1940, Makunaima Terán, Benitez & Mirande, 2020 and Ramirezella Fernández-Yépez, 1949, but the remaining species continue to form a paraphyletic assemblage (Melo et al., 2024). Within this large group, the type species M. xinguensis (Steindachner, 1882) was recovered as closely related to M. britskii Azevedo-Santos and Benine, 2016, M. grandisquamis (Müller & Troschel, 1845), M. pankilopteryx Bertaco & Lucinda, 2006, M. surinamensis Géry, 1965, and M. restricta Soares & Benine, 2019. This clade is sister to a more inclusive group including other species of Moenkhausia as well as taxa currently placed in Hasemania Ellis, 1911, Hemigrammus Gill, 1858, Nematocharax Weitzman, Menezes & Britski, 1986,and Stichonodon Eigenmann, 1903.

Despite being recognized as an artificial group, new species continue to be described based on Eigenmann’s definition of Moenkhausia (Marinho, Dagosta, 2023; Lima et al., 2023; Reia et al., 2024) pending a more comprehensive resolution of the phylogenetic relationships within Acestrorhamphidae. In this context, we present the description of a new species of Moenkhausia collected in ichthyological surveys in underexplored regions of the Amazon basin in Pará, Brazil. This species is endemic to the headwaters of the Curuá River (Xingu River basin) and the Teles Pires River (Tapajós River basin), both draining the Serra do Cachimbo, an ancient geological formation located in southwestern Pará.

Material and methods

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Counts and measurements were taken according to Fink, Weitzman (1974) and Menezes, Weitzman (1990), except for counts of the horizontal scale rows below the lateral line, which were counted to the pelvic-fin insertion. Count of horizontal scale rows between dorsal-fin origin and lateral line does not include scale of median predorsal series. Standard length (SL) is presented in mm while all other measurements are presented as proportions of standard length, except subunits of head, which are presented as proportions of head length. In the descriptions, the frequency of each count is given in parentheses after its respective count. An asterisk indicates counts of the holotype. Counts of supraneurals, procurrent caudal-fin rays, branchiostegal rays, gill-rakers, dentary teeth and tooth cusps were taken from three cleared and stained paratypes (c&s), prepared according to Taylor, Van Dyke (1985). Vertebrae of the Weberian apparatus were included in the vertebral counts as four elements and the compound centrum of the caudal region as a single element. Color in life description was based on photography of freshly collected material and aquarium specimens. Institutional abbreviations follow Fricke, Eschmeyer (2025).

Results​

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Moenkhausia solaris, new species

urn:lsid:zoobank.org:act:6283798D-3CEC-42C5-87FC-D6E09D167B7F

(Figs. 1–3; Tab. 1)

Holotype. MZUSP 130975, 47.9 mm SL, Brazil, Pará, Novo Progresso municipality, stream affluent of 13 de Maio River crossing dirt road near PCH Salto do Curuá, Curuá River, Xingu River basin, 08°44’42.2”S 55°02’04”W, 7 Aug 2015, F. C. P. Dagosta, M. M. F. Marinho, P. Camelier & V. Giovanetti.

Paratypes. All from Brazil, Pará. Xingu River basin, Iriri River drainage, Curuá River drainage: MZUSP 119995, 7, 22.3–30.3 mm SL (4, 25–30.3 mm SL), Altamira municipality, 13 de Maio River, upstream PCH Salto Três de Maio, 08°44’59” S 55°01’58.8”W, 7 Aug 2015, F. C. P. Dagosta, M. M. F. Marinho, P. Camelier & V. Giovanetti. MZUSP 128862, 4, 45.1–55.2 mm SL, Altamira municipality, stream tributary of 13 de Maio River, at PCH Três de Maio 08°44’32.4”S 55°01’52.1”W, 12 Oct 2017, O. Oyakawa, M. M. F. Marinho, P. Camelier & R. Burger. MZUSP 128882, 4, 51.9–57.8 mm SL, same data as the holotype. MZUSP 128883, 17, 23.9–52 mm SL (10, 35.4–52 mm SL), 2 c&s (42.4–44 mm SL), Altamira municipality, stream tributary of 13 de Maio River at PCH Três de maio, upstream of the dam, 08°45’37.7”S 55°01’39”W, 12 Oct 2017, O. T. Oyakawa, M. M. F. Marinho, P. Camelier & R. Burger. ZUFMS 9281, 6, 41.2–49.1 mm SL, Altamira municipality, 13 de Maio River, Iriri basin, upstream of the dam, near the gate of PCH, Cachoeira da Serra, 08°45’6”S 55°02’05” W, 20 Sep 2019, E. A. Oliveira, F. M. C. Sassi & A. D. Silva. Tapajós River basin: MZUSP 122912, 18, 20.2–56.7 mm SL (11, 20.2–56.7 mm SL), 1 c&s (38.9 mm SL), Novo Progresso municipality, stream in dirty road perpendicular to BR-163 road, ahead Força Aérea Brasileira road, MT-PA direction, tributary of Braço Norte River at Vavareta farm, Teles Pires River drainage, 54°49’13.4”W 09°16’31.7”S, 8 May 2015, F. C. P. Dagosta, M. M. F. Marinho, P. Camelier & V. Giovanetti.

FIGURE 1| Holotype of Moenkhausia solaris, MZUSP 130975, 47.9 mm SL, Brazil, Pará State, Novo Progresso, stream affluent of 13 de Maio River crossing dirt road near PCH Salto do Curuá.

Diagnosis. Moenkhausia solaris is distinguished from all congeners, except M. cambacica Marinho, Ohara & Dagosta, 2021, M. clorophthalma Sousa, Netto-Ferreira & Birindelli, 2010, M. lineomaculata Dagosta, Marinho & Benine, 2015, M. parecis Ohara & Marinho, 2016, M. petymbuaba Lima & Birindelli, 2006, and M. plumbea Sousa, Netto-Ferreira & Birindelli, 2010, by the presence of a dark blotch on the anterior portion of each scale of the second to seventh longitudinal series (vs. pigmentation absent or, when present, concentrated at the posterior margin of scales, forming a reticulate pattern). Moenkhausia solaris can be distinguished from all the aforementioned species, except M. petymbuaba, by having the adipose fin and the base of caudal-fin lobes red (vs. hyaline in M. lineomaculata and yellowish in M. cambacica, M. clorophthalma, M. parecis and M. plumbea). It is distinguished from M. petymbuaba by the absence of a wide, midlateral dark stripe on the body (vs. presence). Moenkhausia solaris can be further distinguished from M. parecis and M. lineomaculata by having the caudal-peduncle blotch extending to tip of middle caudal-fin rays (vs. restricted to the base of rays), from M. clorophthalma by having hyaline adipose fin, with only scattered dark chromatophores (vs. base of adipose fin black) and 30–33 perforated lateral-line scales (vs. 25–28), from M. plumbea also by having 30–33 lateral-line scales (vs. 33–37) and by having seven branched pelvic-fin rays (only one side of one out of 29 specimens with six branched rays) (vs. six; only two out of 34 specimens with seven rays (Sousa et al., 2010), from M. petymbuaba by having 17–21 branched anal-fin rays) (vs. 22–24), and from M. cambacica by having the upper portion of eye dark and ventral portion green in life, without a dark line crossing it (vs. eyes with golden coloration with dark shaded line crossing the eye horizontally).

Description. Morphometric data of the holotype and paratypes presented in Tab. 1. Body compressed, moderately elongated. Greatest body depth located slightly anterior or at dorsal-fin origin. Dorsal profile of head convex from anterior tip of upper jaw to vertical through anterior nostril; straight or slightly concave from later point to tip of supraoccipital spine. Predorsal profile slightly convex from tip of supraoccipital spine to dorsal-fin origin, straight from near dorsal-fin base to adipose fin; slightly concave along caudal peduncle. Ventral profile of head and body distinctly convex from lower lip to anal-fin origin; straight along anal-fin base, and concave between along caudal peduncle.

TABLE 1 | Morphometric data of Moenkhausia solaris. Range includes the holotype (H; MZUSP 130975). Specimens from rio Xingu are from MZUSP 119995 (4), 128862 (4), 128882 (4), 128883 (10), and ZUFMS 9281 (6), and from rio Tapajós are from MZUSP 122912 (9). N = number of specimens; SD = Standard deviation.

	Rio Xingu basin					Rio Tapajós basin
	N	H	Range	Mean	SD	N	Range	Mean	SD
Standard length (mm)	29	47.9	25.0–58.0	–	–	9	28.3–55.7	–	–
Percentage of standard length
Depth at dorsal-fin origin	29	36.5	30.0–37.9	32.7	1.9	9	28.9–39.6	35.3	3.7
Snout to dorsal-fin origin	29	54.0	51.4–58.2	55.0	1.6	9	52.5–55.9	53.9	1.0
Snout to pectoral-fin origin	29	29.4	25.5–30.4	27.7	1.3	9	25.5–29.3	27.4	1.3
Snout to pelvic-fin origin	29	49.8	46.0–50.9	48.4	1.3	9	45.3–49.8	47.2	1.4
Snout to anal-fin origin	29	66.1	63.4–75.4	66.3	2.2	9	60.5–65.3	63.7	1.7
Caudal-peduncle depth	29	12.6	10.5–13.0	11.8	0.7	9	9.7–12.6	11.8	0.8
Caudal-peduncle length	29	12.0	10.5–15.5	13.0	1.3	9	10.0–14.4	12.5	1.5
Pectoral-fin length	24	22.3	15.0–22.5	19.6	1.6	9	18.2–21.4	20.3	1.1
Pelvic-fin length	29	15.7	11.9–16.0	14.0	1.1	9	13.3–17.1	15.8	1.3
Dorsal-fin length	22	26.2	22.5–26.9	24.4	1.3	9	24.8–29.3	26.8	1.5
Dorsal-fin base length	28	13.3	9.6–15.1	12.6	1.0	9	12.5–15.1	13.7	0.8
Anal-fin length	28	19.9	11.5–20.2	15.8	2.4	9	14.8–19.5	17.2	1.7
Anal-fin base length	29	28.4	21.4–28.4	24.5	1.6	9	25.2–29.7	28.0	1.4
Eye to dorsal-fin origin	29	38.4	37.8–46.3	40.0	1.9	9	35.1–41.5	39.7	2.0
Dorsal-fin origin to caudal-fin base	29	49.7	42.8–51.0	47.4	2.2	9	45.6–52.6	50.0	2.2
Head length	29	29.5	26.1–32.2	28.8	1.6	9	24.9–29.3	27.2	1.6
Percentage of head length
Horizontal eye diameter	29	36.4	30.7–38.8	33.9	2.2	9	30.6–37.6	33.6	2.0
Snout length	29	22.2	20.2–28.5	24.2	2.0	9	17.3–27.5	21.8	3.3
Interorbital width	29	32.8	26.6–36.3	32.0	2.5	9	31.1–36.9	33.5	2.2
Upper jaw length	27	50.2	44.3–53.7	49.0	2.0	9	44.4–48.5	46.8	1.4

Mouth terminal, jaws equally aligned. Posterior limit of maxilla almost reaching vertical through middle of orbit. Premaxillary teeth in two rows, outer tooth row with 3(5), 4*(25) or 5(6) teeth with three or five cusps directed inward; inner tooth row with 4 (8) or 5*(28) teeth with five cusps, except symphyseal teeth, with four cusps, all cusps directed outward. Maxilla with 2(20) or 3*(16) teeth at its anterodorsal margin, bearing three to five cusps; anterior teeth usually largest. Dentary with 4*(26) or 5(10) larger teeth with five cusps, followed by one (3) smaller tooth with three cusps and a series of 8(1) or 9(2) diminutive teeth, with one to three cusps considerably smaller than the anterior larger teeth. Cusps of larger dentary teeth directed inward. Central cusp of all teeth more developed than lateral cusps (Fig. 2).

FIGURE 2| Lateral view of left side of premaxilla, maxilla, and dentary of Moenkhausia solaris, MZUSP 128883, 39.9 mm SL, paratype. Scale bar = 1 mm.

Scales cycloid, circuli exposed over whole area of scales. Lateral line completely pored, slightly curved ventrally, with 30(8), 31(12), 32*(11) or 33(6) perforated scales. Longitudinal scale rows between dorsal-fin origin and lateral line 4(2), 5*(35), 6(1). Longitudinal scale rows between lateral line and pelvic-fin origin 3*(26), 4(11) or 5(1). Median series of scales along dorsal line between tip of supraoccipital spine and dorsal-fin origin 9*(10) or 10(28). Horizontal scale rows around caudal peduncle 10(3) or 11*(35). Caudal with small scales present on proximal portion of upper and lower caudal-fin lobes.

Dorsal-fin rays ii*(32) or iii(6), 8(10) or 9*(28). Base of posteriormost dorsal-fin ray approximately at vertical through anal-fin origin. Supraneurals 4(3) or 5(1), slightly wider dorsally. First dorsal-fin pterygiophore posterior to neural spine of 10^th(2) or 11^th(1) vertebrae. Adipose fin present. Pectoral-fin rays i*(38) 9(12), 10*(22) or 11(4). Tip of adpressed pectoral fin reaching vertical through pelvic-fin origin or falling just short of that point. Pelvic-fin rays i,7*(38) (one specimen from MZUSP 119995 with i,6 in one side and i,7 in the other side of body). Anal-fin rays iv(3), 17(3), 18(13), 19(9), 20*(8) or 21(4). Posterior unbranched and anterior branched anal-fin rays longest, subsequent rays gradually decreasing in size. Principal caudal-fin rays i,9,8,i*(32) or i,9,9,i(6). Twelve (1) or 13(2) dorsal procurrent caudal-fin rays and 10(1), 11(2) ventral procurrent caudal-fin rays. Total vertebrae 32(1) or 33(2). Precaudal vertebrae 16(3) and caudal vertebrae 17(1) or 16(2). Branchiostegal rays 4(3). First gill arch with two (3) gill-rakers on hypobranchial, 8(3) on ceratobranchial, and 8(3) on epibranchial.

Coloration in alcohol. Ground color tan, with chromatophores spread the entire head and body, except for the abdomen, densely concentrated on the dorsal portion from upper jaw to caudal peduncle (Fig. 1). Margin of upper lip dark. Vertically elongate humeral blotch present. Presence of a dark blotch on the anterior portion of each scale of the second to seventh longitudinal series, fading towards scale margin. Blotches above the lateral line larger than blotches below the lateral line. Thin line formed by subjacent chromatophores at the horizontal septum. Caudal peduncle with a horizontal oval blotch, frequently extending to the tip of middle caudal-fin rays. Lower portion of caudal peduncle with a clear area. All fins with scattered dark chromatophores on interadial membranes. Distal portion of interadial membranes of dorsal-fin and anal-fin with concentration of dark chromatophores.

Coloration in life. Melanophores pattern as described in Coloration in alcohol section. Humeral blotch less intense in live coloration. Dorsal portion of head and body golden yellow, ventral half of head and body pale. Dorsal portion of eye dark, ventral portion green. Anterior portion of dorsal and anal-fin orange to yellow, remaining rays yellowish to hyaline. Caudal-fin reddish, more intense at its anterior half. Adipose fin red. Base of pelvic and pectoral fins yellowish (Fig. 3).

Sexual dimorphism. No sexual dimorphic feature was observed in the examined specimens.

FIGURE 3| Live specimens of Moenkhausia solaris, aquarium specimens, not preserved. Photo by Oliver Lucanus.

Geographical distribution. Moenkhausia solaris is known from headwaters of the Curuá River drainages, Xingu River basin, and from the upper Teles Pires River drainage, Tapajós River basin, restricted to the Serra do Cachimbo plateau, Pará State, Brazil (Fig. 4).

Geographical variation. Comparative analysis of morphometric and meristic data taken from populations from Xingu and Tapajós River basins did not reveal any significant differences, with the exception of the relative proportion of the dorsal-fin length, which is higher in the specimens of Tapajós River 24.8–29.3%, n = 9 (vs. 22.5–26.9%, n = 22 specimens of Xingu River). However, it should be mentioned that part of the specimens had damaged fins.

Ecological notes. The new species occur in small streams with preserved riparian vegetation. One of the localities from the Curuá River drainage (MZUSP 128883) is a shaded area with shallow water and sandy bottom, covered with dense vegetation (Fig. 5).

FIGURE 4| Distribution map of Moenkhausia solaris in the Serra do Cachimbo, Pará, Brazil. Type-locality is represented by the red dot. White line indicates the watershed between the Tapajós and Xingu River basins.

Etymology. The specific epithet solaris comes from Latin, meaning sunny or pertaining to the sun. In allusion to the bright gold color of body and red color of the adipose and caudal fin present in live specimens. An adjective.

Conservation status. Moenkhausia solaris occurs in the headwaters of the Curuá River drainages, Xingu River basin, and in the upper Teles Pires River drainage, Tapajós River basin, restricted to the Serra do Cachimbo plateau. Despite the restricted distribution, it occurs in relatively preserved environment with no identifiable threats. Therefore, Moenkhausia solaris should be classified as Least Concern (LC) following the International Union for Conservation of Nature (IUCN) categories and criteria (IUCN 2024).

FIGURE 5| One of the localities where Moenkhausia solaris was collected, MZUSP 128883, a small stream tributary of 13 de Maio River at the area of PCH Salto Três de Maio, upstream of the dam.

Discussion​

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Moenkhausia solaris is morphologically very similar to a group of species of Moenkhausia with endemic distribution, each confined to headwater streams of the River Madeira, Tapajós, and Xingu River basins: M. cambacica, M. clorophthalma, M. parecis, M. petymbuaba, and M. plumbea. They share the presence of dark blotches on scales of the second to seventh longitudinal series of the body, formed by a higher concentration of chromatophores on the anterior portion of scales and colored eyes, along with large head, round dorsal-fin profile, and a relatively short anal-fin base (Ohara, Marinho, 2016). Such a distinctive pigmentation pattern is unique among Acestrorhamphidae and may indicate a close relationship among them (Sousa et al., 2010; Ohara, Marinho, 2016; Marinho et al., 2021). A similar longitudinal series of blotches is also observed in Moenkhausia lineomaculata and some populations of M. cotinho Eigenmann, 1908 (Dagosta et al., 2015). Moenkhausia lineomaculata and M. cotinho, however, exhibit a distinguished reticulate color pattern formed by pigmentation on the distal margins of scales, which is absent in M. solaris.Such reticulate pattern appears to have evolved independently twice within Acestrorhamphidae, once in the species of Bario, to which M. lineomaculata is likely closely related, and once in Ramirezella, a genus possibly related to M. cotinho (Melo et al., 2024).Therefore, a close relationship among M. solaris, M. lineomaculata and M. cotinho is unlikely.

The restricted distribution of M. solaris in the Serra do Cachimbo is notable and biogeographically significant. The Serra do Cachimbo is considered an area of exceptionally high ichthyofaunal endemism, characterized by low overall species richness, and with few examples of taxa with broad distribution across the Amazon basin (Dagosta, de Pinna, 2019). Such a biogeographic pattern, named “Extreme Shield” by Dagosta, de Pinna (2019), is related to elevated geomorphological formations, including rapids and waterfalls, which act as natural barriers to dispersal and gene flow.

Marinho, Dagosta (2024) discussed that most of the endemic fishes of the Serra do Cachimbo are confined to a single major hydrographic basin that drains the area, namely the Jamanxim River basin (Tapajós River basin), Teles Pires River basin (Tapajós River basin), and Iriri River basin (Xingu River basin). There are only two known exceptions, Hyphessobrycon cachimbensis Travassos, 1964, endemic to the Serra do Cachimbo and occurring in both Teles Pires and Iriri basins (Marinho, Dagosta, 2024) and M. solaris, which shows the same distributional pattern (Fig. 4). Such a pattern suggests a shared biogeographical history between the small tributaries of the Curuá River (Iriri/Xingu River basin) and the upper portions of the Braço Norte River (Teles Pires/Tapajós River basin) at the Serra do Cachimbo slope, probably by headwater capture events. The close proximity of these drainages, approximately 2 km apart near the areas where M. solaris was collected, also helps to support this hypothesis.

Moenkhausia solaris is sympatric with M. clorophthalma in the Xingu stretches,a morphologically similar species. They can be readily distinguished from each other by the coloration of the adipose fin (hyaline in alcohol specimens or red in life in M. solaris vs. deep black in M. clorophthalma) and by the number of body scales, which is 30–33 perforated lateral-line scales in M. solaris (vs. 25–28 in M. clorophthalma).

Comparative material examined. Material examined here are the same listed at Dagosta et al. (2015) and Ohara, Marinho (2016), with the addition of Moenkhausia plumbea: Tapajós River basin:MZUSP 119977, 10, 30.1–48.5 mm SL. Moenkhausia clorophthalma: Xingu River basin:MZUSP 107307, 10, 30.6–38.8 mm SL. MZUSP 128881, 26, 16.5–43.8 mm SL.

Acknowledgments​

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We thank Oliver Lucanus (McGill University) for kindly providing color in life photographs and Antônio da Silva (UFSCar), Ezequiel de Oliveira (Seduc-MT), Francisco Sassi (UFSCar), Fernando Dagosta (UFGD), Marcelo Cioffi (UFSCar), Priscila Camelier (UFBA), Osvaldo Oyakawa (MZUSP), Rafael Burger (UFBA), Victor Giovanetti (IB-USP) for helping in the field expedition.

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Authors

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José Igor da Silva¹, Yasmim de Santana² and Manoela Maria Ferreira Marinho²

^[1]    Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, Cidade Universitária, Castelo Branco, 58051-900 João Pessoa, PB, Brazil. (JIS) igor.cbio.ufpb@gmail.com.

^[2]    Instituto de Biologia, Universidade Federal do Mato Grosso do Sul, Bairro Universitário, 79070-900, Campo Grande, MS, Brazil. (YS) yasmim31sant@gmail.com, (MMFM) manoela.marinho@ufms.br (corresponding author).

Authors’ Contribution

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José Igor da Silva: Data curation, Formal analysis, Investigation, Methodology, Project administration, Software, Writing-original draft, Writing-review and editing.

Yasmim de Santana: Data curation, Formal analysis, Investigation, Methodology, Validation, Writing-review and editing.

Manoela Maria Ferreira Marinho: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, Writing-original draft, Writing-review and editing.

Ethical Statement​

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Not applicable.

Competing Interests

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The author declares no competing interests.

Data availability statement

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The authors confirm that the data supporting the findings of this study are available within the article.

Funding

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JIS and YSS were funded by the Conselho Nacional de Desenvolvimento Científico e Tecnológico PIBIC/UFPB/CNPq/2023–2024 and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) 88887.148413/2025–00, respectively. This research was supported by the project The South American Characiformes Inventory funded by Fundação de Amparo à Pesquisa do Estado de São Paulo (2011/50282–7) and INCT-Peixes, funded by MCTIC/CNPq (405706/2022–7).

How to cite this article

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Silva JI, Santana Y, Marinho MMF. A new species of Moenkhausia (Characiformes: Acestrorhamphidae) from the Amazon basin at Serra do Cachimbo, Pará, Brazil. Neotrop Ichthyol. 2025; 23(3):e250089. https://doi.org/10.1590/1982-0224-2025-0089

Copyright​

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This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Distributed under

Creative Commons CC-BY 4.0

© 2025 The Authors.

Diversity and Distributions Published by SBI

Accepted June 10, 2025

Submitted May 13, 2025

Epub September 08, 2025