A state-of-art review of Loricariidae (Ostariophysi: Siluriformes) cytogenetics

Francisco de Menezes Cavalcante Sassi¹, Marcelo de Bello Cioffi¹ and Orlando Moreira-Filho¹

Abstract

Loricariidae is a Neotropical fish family divided into six subfamilies and ranking in third among the most biodiverse fish groups. This study conducts an updated review of the cytogenetic investigations within the family, discussing the trends in chromosomal evolution and the main gaps and future directions for studies. Covering 125 publications that analyzed 234 species from all subfamilies except Lithogeninae, corresponding to about 21% of the valid species diversity, our study revealed samples from six different river basins in Argentina, Brazil, Ecuador, Paraguay, and Venezuela. There was a dearth of data for northeast Brazil, the Western Amazon, the Guianas Shield, and other Neotropical countries. In loricariids, there are seven different sex chromosome systems and a variety of diploid numbers (2n) ranging from 33 to 96 as a result of different chromosomal rearrangements such as fusions, fissions, translocations, and inversions. We recorded more simple nucleolar organizer regions (Ag-NOR) compared to multiple ones, and the fundamental number (FN) varied between 34 and 142. Populational studies have been conducted only in a few taxa, but a remarkable karyotype variation that includes B chromosomes is shown. Despite continuous efforts, cytogenetics still does not adequately capture the diversity of Loricariidae.

Keywords: Ag-NOR, B chromosomes, Karyotype, Plecos, Sex chromosomes.

Loricariidae é uma família de peixes Neotropicais dividida em seis subfamílias e ranqueada na terceira posição dentre os grupos mais biodiversos de peixes. Este estudo conduz uma revisão atualizada das investigações citogenéticas na família, discutindo as tendências evolutivas cromossômicas e as principais lacunas e direções futuras para estudos. Cobrindo 125 publicações que analisam 234 espécies de todas as subfamílias, exceto Lithogeninae, correspondendo a aproximadamente 21% da diversidade de espécies válidas, nosso estudo revelou amostragens em seis bacias hidrográficas distintas na Argentina, Brasil, Equador, Paraguai e Venezuela. Há uma escassez de dados para o nordeste do Brasil, o oeste da Amazônia, o escudo das Guianas e outros países Neotropicais. Em loricariídeos, há sete sistemas de cromossomos sexuais distintos e uma variedade de números diploides (2n), variando de 33 a 96 como resultado de rearranjos cromossômicos diferentes como fusões, fissões, translocações e inversões. Nós registramos mais regiões organizadoras de nucléolo (Ag-RON) simples quando comparado com múltiplas, e o número fundamental (NF) variou entre 34 e 142. Estudos populacionais foram realizados em poucos organismos, mas uma extraordinária variação cariotípica que inclui cromossomos B é demonstrada. Apesar de esforços contínuos, a citogenética ainda não captura adequadamente a diversidade de Loricariidae.

Palavras-chave: Ag-RON, Cariótipo, Cascudos, Cromossomos B, Cromossomos sexuais.

Introduction

Among vertebrates, the paraphyletic group of fish outstands because of its high richness and diversity of morphology and ecology (Nelson et al., 2016). They can be found all over the Earth in freshwater and marine environments, and humans exploit them widely as primary food sources or as ornamental species for the aquarium trade (Evers et al., 2019; Novák et al., 2020, 2022). Such diversity is observed in about 36,000 valid species, with almost half of them belonging to the 450 families that make up the infraclass Teleostei (Fricke et al., 2023). More than 18,000 species are found in freshwater environments, the majority of which are found in tropical and subtropical regions, including the Amazon River Basin, tropical Africa, and southeast Asia (Lévêque et al., 2008; Nelson et al., 2016). The monophyletic order Siluriformes (popularly known as catfishes), is home to a large portion of this variety (Fink, Fink, 1981, 1996; Arratia et al., 2003; Saitoh et al., 2003; Sullivan et al., 2006). This order is composed by the suborder Loricarioidei, which consists of six families that are endemic to South America, and Siluroidei, which includes 29 families that are spread over all continents except Antarctica (Sullivan et al., 2006). Due to their widespread distribution (Fig. 1), research focusing on the biogeography of siluriform fishes often indicates that South America may have served as the group’s center of origin (Briggs, 2005; Sullivan et al., 2006).

FIGURE 1| World map showing the distribution of Siluriformes (red areas) according to Nelson et al. (2016) and GBIF (2023), with illustrations of important families in distinct regions.

Loricariidae species, which rank third among the most biodiverse fish families, are known by several names, including armored catfishes, plecos, “cascudos”, “bodós”, and “acaris”. Together with Characidae and Cyprinidae, it corresponds for about 50% of all Ostariophysi species (Nelson et al., 2016). According to Fricke et al. (2023), 1,051 valid species are divided into six subfamilies: Delturinae, Hypoptopomatinae, Hypostominae, Lithogeninae, Loricariinae, and Rhinelepinae. Their distinct anatomy, which includes a suckermouth positioned ventral and dermal plates covering the body, allows them to be recognized despite their widespread distribution in the Neotropical region. Loricariids are iliophagous, with a vast diversity of feeding habitats ranging from wood to detritus and algae (Buck, Sazima, 1995; Lujan et al., 2017). They demonstrate a broad range of sizes, from less than 5 cm (e.g., Ribeiro et al., 2012) to approximately 1 m (Lujan et al., 2010). Given a combination of excessive reaping for the aquarium trade and the destruction of their habitats, some loricariids are threatened (Barros et al., 2023). Indeed, the high ornamental trade of plecos led to the development of the L-code (or L-number). This code was developed by aquarists, aiming to create a temporary “voucher number” for morphological variants identified by aquarium trade that lack a proper taxonomic identification or are difficult to classify only by morphological features (Stawikowski et al., 2004). Recently, 223 Loricariidae species have been assigned from their L-numbers to valid species name (Novák et al., 2022).

The description of fish species is often complex. While some species present a conservation of morphological features, body-color variation and polymorphisms are frequently observed in fish (Price et al., 2008). In this context, the use of complementary methods is fundamental to assessing all biodiversity, in especial cytogenetics (Stace, 2000), the field of genetics that studies chromosomes from their structure and molecular composition to their organization and variation among species. Cytogenetics is an important tool for discovering the Neotropical fish biodiversity, as exemplified by the outstanding number of new and/or cryptic species in addition to karyomorphs with absence of hybrid forms reported (e.g., Pazian et al., 2012; Ferreira et al., 2014; Oliveira et al., 2016; Ramirez et al., 2017; Rocha-Reis et al., 2018). Among loricariids, Artoni, Bertollo (2001) suggest that 2n = 54 might represent the basal diploid number for this family, with distinct trends in evolution among its subfamilies. Considering the continuous growing number of studies involving Loricariidae species, here we review for the first time the scientific literature that includes cytogenetic data for Loricariidae species. We discussed the trends in chromosomal evolution in this representative fish group and the main gaps and future directions for studies.

Material and methods

A detailed search was conducted on Google Scholar until December of 2023, using the keywords “Cytogenetics Loricariidae” and “Chromosomes Loricariidae”. During the screening of research papers, all the duplicate papers and cytogenetic papers without karyotype information were excluded from our compilation (for example Traldi et al., 2019, where only mapping of repetitive sequences was conducted, without addressing the diploid number — 2n and karyotype structure). We included all papers found published between 1968 and 2023 in journals in any language available. For more assurance on the quality of results, the data analyzed was double-checked by the authors: the resulted table comprising the cytogenetic features was checked by each author, comparing with the original papers to determine whether such articles would comply with the inclusion and exclusion criteria. We followed the classification of chromosomes by the centromeric position and arm ratios, proposed by Levan et al. (1964) and standardized for fish in Arai (2011) as metacentric (m), submetacentric (sm), subtelocentric (st) and acrocentric (a).

From each paper, we extracted cytogenetic and biogeographic data such as diploid number (2n), fundamental number (NF), karyotype formula (KF), sex chromosome system (SCS) and AgNOR/18S rDNA sites (simple or multiple). Additionally, locality and geographical coordinates of the populations investigated were also compiled, with the city name, state, and country indicated according to the coordinates provided in the papers. Species and family names were checked against Eschmeyer’s Catalog of Fishes (Fricke et al., 2023) and updated according to the valid names. Taxonomic abbreviations were maintained as present in the papers, following: “sp.” for specimens not fully identified, “prope” for specimens with resemblance to a described taxon but not identical, “aff.” for closely related to an identified taxon or with uncertainty in taxonomic identification, “cf.” for specimens similar to or comparable to a taxon but with doubts, “n.sp.” for proposed new species, and “L.” for locally variants of specimens. For the papers in which the geographical coordinates were available, we plotted those sample points into maps using QGIS 3.28.2.

Results

We compiled 125 papers that included cytogenetic data on Loricariidae species. These studies comprised a diversity of 234 species, here included those identified as “sp.”, prope, “aff.”, “cf.”, “n.sp.”, and “L” as distinct species, from 48 genus. Hypostominae was the subfamily most represented with 142 species karyotyped, followed by Loricariinae (54 spp.), Hypoptopomatinae (34 spp.), and Delturinae and Rhinelepinae with 2 spp. each, with absence of data for Lithogeninae. The Hypostomus genus was the most represented with 26 valid species karyotyped, in a total of 159 records when including all populational data and those identified as “sp.”, prope, “aff.”, “cf.”, “n. sp.”, and “L”. The geographical coordinates plotted into maps show a distribution of species cytogenetically investigated in Brazil, Argentina, and Paraguay, comprising six distinct river basins (Fig. 2). Although species from Ecuador and Venezuela were also compiled, those papers do not present geographical coordinates of the sample sites.

FIGURE 2| Partial maps of South America showing the distribution of Loricariidae species (red dots) investigated by cytogenetics that have the geographical coordinates published together. A. Map highlighting the geopolitical regions of South America, with emphasis on Brazil, Paraguay, and part of Argentina territory; B. Map highlighting the hydrographic basins of South America, with emphasis on Amazonas (green), Tocantins-Araguaia (yellow), Paraná/Paraguay (purple), São Francisco (cyan), Atlantic East and Atlantic South (red) River Basins.

Diploid number varied from 2n = 33 in males of Rineloricaria teffeana (Steindachner, 1879) (Marajó et al., 2022) to 2n = 96 in Hemipsilichthys sp. (Kavalco et al., 2004, 2005). The fundamental number ranged from 34 in R. teffeana (Marajó et al., 2022) to 142 in Hypostomus topavae (Godoy, 1969) (Kamei et al., 2017), and the simple distribution of Ag-NOR/18S rDNA was the most common with 269 records, against 110 records of multiple sites. Seven sex chromosomes systems were described for 31 Loricariidae species, the simple XX/XY, XX/X0, and ZZ/ZW, and the multiples X₁X₁X₂X₂/X₁X₂Y, XX/XY₁Y₂, ZZ/ZW₁W₂, and Z₁Z₁Z₂Z₂/Z₁Z₂W₁W₂. B chromosomes were found in five species, varying from 1B (e.g., de Souza et al., 2009) to 3B chromosomes (e.g., Porto et al., 2010). Complete results were compiled in Tab. 1.

TABLE 1 | Review of cytogenetic data by subfamily of Loricariidae published as articles. Geographical coordinates ending with asterisk indicates mismatched coordinates. Diploid number (2n), fundamental number (NF), karyotype formula (KF), sex chromosome system (SCS) are displayed.

Subfamily/Genus	Species	Locality	Latitude	Longitude	Basin	2n	NF	Ag-NOR – 18S	KF	SCS	Reference/ Observation
Loricariinae
Brochiloricaria	macrodon (Kner, 1853)					58♀♂	78		18m + 2sm + 38st/a		Michele et al. (1977). Reported as Loricaria macrodon
Farlowella	amazonum (Günther, 1864)	Agua Boa stream, Mundo Novo, MS, Brazil				58♀♂	110	Simple	6m + 38sm + 8st + 6a		Fernandes et al. (2012)
	amazonum	Dourado stream, Japorã, MS, Brazil	23°51’04.9"S	54°09’51.1"W	Iguatemi – Paraná	58♀♂	110	Simple	12m + 30sm + 10st + 6a		Fernandes et al. (2015)
	cf. amazonum	Paraná do Piloto, Barcelos, AM, Brazil	0°56’04.8"S	62°58’01.6"W	Negro – Amazonas	58♀♂	116	Simple	14m + 30sm + 14st		Marajó et al. (2018)
	schreitmuelleri Arnold, 1936	Jundiá stream, Manaus, AM, Brazil	2°19’43.8"S	60°04’40.4"W	Cuieiras – Amazonas	58♀♂	112	Simple	10m + 30sm + 14st + 4a		Marajó et al. (2018)
	hahni Meinken, 1937	Dourado stream, Mundo Novo, MS, Brazil	23°51’04.9"S	54°26’31.4"W	Paraná	58♀♂	110	Simple	12m + 30sm + 10st + 6a		Fernandes et al. (2021)
	hahni	Iguaçu River, Capanema, PR, Brazil	25°38’18.7"S	54°28’01.7"W	Paraná	58♀♂	112	Simple	12m + 20sm + 22st + 4a		Fernandes et al. (2021)
Harttia	carvalhoi Miranda Ribeiro, 1939	Ribeirão grande, Pindamonhangaba, SP, Brazil	22°46’03.0"S	45°26’07.1"W	Paraíba do Sul	52♀, 53♂		Simple	18m + 18sm + 8st + 8a or 17m + 18sm + 8st + 10a	XX/XY₁Y₂	Centofante et al. (2006)
	carvalhoi	Pindamonhangaba, SP, Brazil			Paraíba do Sul	52♀, 53♂		Simple	16m + 16sm + 12st + 8a ♀, 15m + 16sm + 12st + 10a ♂	XX/XY₁Y₂	Blanco et al. (2013, 2017); Deon et al. (2022a,b)
	carvalhoi					52♀♂		Simple	30m/sm + 20st/a		Alves et al. (2003). Reported as Harttia loricariformis, corrected in Centofante et al. (2006)
	dissidens Rapp Py-Daniel & Oliveira, 2001	Tambor stream, Rurópolis, PA, Brazil	4°05’37.8"S	55°00’30.2"W		54♀♂	92	Simple	20m + 26sm + 8a		Sassi et al. (2021, 2023a,b)
	duriventris Rapp Py-Daniel & Oliveira, 2001	Parauapebas River, Canaã dos Carajás, PA, Brazil	6°30’06.5"S	50°02’35.3"W	Tocantins-Araguaia	55♂, 56♀	96, 98	Simple	16m + 16sm + 16st + 8a ♀, 17m + 16sm + 16st + 6a ♂	X₁X₁X₂X₂/X₁X₂Y	Sassi et al. (2020, 2023a,b)
	gracilis Oyakawa, 1993	Machadinho stream, Santo Antonio do Pinhal, MG, Brazil	22°48’31.0"S	45°41’21.0"W	Sapucaí-Mirim	58♀♂		Simple	20m + 22sm + 8st + 8a		Blanco et al. (2017); Deon et al. (2022a,b)
	guianensis Rapp Py-Daniel & Oliveira, 2001	Paraíso stream, Alenquer, PA, Brazil	1°29’02.2"S	54°50’31.2"W		58♀♂	96	Simple	20m + 26sm + 2st + 10a		Sassi et al., 2021, 2023a,b
	intermontana Oliveira & Oyakawa, 2019	Piranga River, Carandaí, MG, Brazil	20°59’34.0"S	43°43’30.0"W		52♀, 53♂	90, 91		14m + 12sm + 12st + 14a ♀, 13m + 12sm + 13st + 15a ♂	XX/XY₁Y₂	Deon et al. (2020, 2022)
	kronei Miranda Ribeiro, 1908					58♀♂		Simple	40m/sm + 18st/a		Alves et al. (2003)
	kronei	Açungui River, Campo Largo, PR, Brazil			Ribeira	58♀♂	106	Simple	14m + 20sm + 14st + 10a		Blanco et al. (2013, 2017); Deon et al. (2022a,b)
	longipinna Langeani, Oyakawa & Montoya-Burgos, 2001	São Francisco River, Pirapora, MG, Brazil	17°21’22.3"S	44°56’59.5"W	São Francisco	58♀♂	102	Simple	16m + 12sm + 16st + 14a + 0-2Bs		Blanco et al. (2013, 2017); Deon et al. (2022a,b)
	loricariformis Steindachner, 1877	Paraitinga River, Silveiras, SP, Brazil	22°52’22.5"S	44°51’00.4"W	Paraíba do Sul	56♀♂	106		16m + 22sm + 10st + 8a		Kavalco et al. (2004, 2005); Blanco et al. (2017); Deon et al. (2022a,b)
	punctata Rapp Py-Daniel & Oliveira, 2001	Itiquira River, Formosa, GO, Brazil	15°19’25.0"S	47°25’26.0"W	Tocantins-Araguaia	57♂, 58♀	116	Simple	16m + 20sm + 12st + 10a ♀, 16m + 21sm + 12st + 8a ♂	X₁X₁X₂X₂/X₁X₂Y	Blanco et al. (2014, 2017); Deon et al. (2022); Sassi et al. (2021, 2023a)
	rondoni Oyakawa, Fichberg & Rapp Py-Daniel, 2018	13 de Maio River, Cachoeira da Serra, PA, Brazil	8°38’53.0"S	55°01’41.0"W	Xingu – Amazon	54♀♂	100	Simple	20m + 26sm + 4st + 4a	XX/XY	Sassi et al. (2020, 2023a,b)
	sp. 1	Macacos stream, Silveiras, SP, Brazil	22°40’43.0"S	44°51’25.0"W		56♀, 57♂	94		14m + 14sm + 10st + 18a ♀, 13m + 14sm + 10st + 20a ♂	XX/XY₁Y₂	Deon et al. (2020, 2022a,b)
	sp. 2	Barra Grande River, Prudentópolis, PR, Brazil	24°58’40.7"S	51°07’34.3"W		62♀♂	104		16m + 14sm + 12st + 20a		Deon et al. (2020, 2022a,b)
	sp. 3	Rio do Peixe, Cachoeira da Serra, PA, Brazil	8°39’20.7"S	55°09’24.1"W		54♀♂	96	Simple	16m + 18sm + 14st + 6a		Sassi et al. (2021, 2023a,b)
	torrenticola Oyakawa, 1993	Araras stream, Brazil			São Francisco	56♀♂	98	Simple	16m + 10sm + 16st + 14a		Blanco et al. (2012a, 2017); Deon et al. (2022a,b)
	villasboas Oyakawa, Fichberg & Rapp Py-Daniel, 2018	Curuá River, Cachoeira da Serra, PA, Brazil	8°44’09.0"S	54°57’46.0"W	Xingu – Amazon	55♂, 56♀	96, 98	Simple	18m + 24sm + 6st + 8a ♀, 19m + 24sm + 6st + 6a ♂	X₁X₁X₂X₂/X₁X₂Y	Sassi et al. (2020, 2023a,b)
Hemiodontichthys	sp. 1	São Francisco stream, Brazil			Purus	46♀♂			42m/sm + 4st/a		Carvalho et al. (2018)
	sp. 2	Iquiri stream, Brazil			Purus	58♀♂			28m/sm + 30st/a		Carvalho et al. (2018)
Loricaria	cataphracta Linnaeus, 1758	Córrego da Onça, Coxim, MS, Brazil			Paraguai	64♀♂	86	Simple and Multiple	12m + 8sm + 2st + 42a		Porto et al. (2014b)
	cataphracta					64	76		12m/sm + 52st/a		Fenocchio et al. (2003). Reported as Loricaria carinata
	cf. cataphracta	Iguaçu River, Brazil			Paraná	64♀♂		Simple	12m + 8sm + 2st + 42a		Takagui et al. (2020)
	simillima Regan, 1904	Paraná River, Brazil	27°06’22.5"S	55°31’20.0"W	Paraná	64♀♂	88	Simple	12m + 12sm + 40st/a		Benitez et al. (2017)
	simillima	Miranda River, Brazil			Paraguai	64♀♂		Simple	12m + 8sm + 2st + 42a		Takagui et al. (2020)
Loricariichthys	anus (Valenciennes, 1835)	Capivara stream, Laguna dos Patos system, RS, Brazil	30°17’34.0"S	51°19’21.2"W		54♀♂	82	Simple	10m + 18sm + 26a		Takagui et al. (2014)
	anus	Itapeva Lagoon, Tramandaí system, Brazil	29°32’11.9"S	49°55’47.9"W		54♀♂	80	Simple	10m + 16sm + 28a		Takagui et al. (2014)
	anus	Chimiray stream, Argentina	28°05’49.4"S	55°41’37.3"W	Uruguay	54♀♂	82	Simple	10m + 18sm + 26a		Takagui et al. (2014)
	anus	Iguaçu River, Brazil			Paraná	54♀♂		Simple	14m + 14sm + 26a		Takagui et al. (2020)
	maculatus (Bloch, 1794)				La Plata	56	78		22m/sm + 34st/a		Fenocchio et al. (2003)
	platymetopon Isbrücker & Nijssen, 1979	Jacutinga River, Brazil	23°13’25.3"S	50°58’47.4"W	Paranapanema	54♀♂	80	Simple	10m + 16sm + 28a		Takagui et al. (2014)
	platymetopon	Paranapanema River, Brazil	22°42’30.3"S	51°04’08.4"W		54♀♂	80	Simple	10m + 16sm + 28a		Takagui et al. (2014)
	platymetopon	Miranda River, Brazil				54♀♂		Simple	10m + 16sm + 28a		Takagui et al. (2020)
Pyxiloricaria	menezesi Isbrücker & Nijssen, 1984	Miranda River, Brazil			Paraná	68♀♂		Simple	8m + 6sm + 2st + 52a		Takagui et al. (2020)
Proloricaria	prolixa (Isbrücker & Nijssen, 1978)	Três Bocas stream, Brazil			Paraná	62♀♂		Simple	20m + 6sm + 36a		Takagui et al. (2020)
Rineloricaria	aequalicuspis Reis & Cardoso, 2001	Maquine River, RS, Brazil	29°39’10.4"S	50°12’31.8"W	Southeast Atlantic	68♀♂		Simple	68st/a		Venturelli et al. (2021)
	cadeae (Hensel, 1868)					66♀♂			2m + 64st/a		Alves et al. (2003)
	cadeae	Guaíba Lake, RS, Brazil				64♀♂		Simple	2m/sm + 62st/a		Maia et al. (2010)
	cadeae	Forquetinha River, RS, Brazil	29°24’22.4"S	52°03’19.2"W		64♀♂		Simple	64st/a		Venturelli et al. (2021)
	capitonia Ghazzi, 2008	Uruguai River, São Carlos, SC, Brazil			Uruguai	64♀♂	70	Simple	4m + 2sm + 58st/a		Primo et al. (2017)
	kronei (Miranda Ribeiro, 1911)	Itapocu River, Santa Catarina, Brazil				64♀♂			6m/sm + 58st/a		Alves et al. (2003)
	lanceolata (Günther, 1868)	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		48♀♂	54	Multiple	4m + 2st + 42a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		48♀♂	53	Multiple	3m + 2st + 43a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		48♀♂	55	Multiple	3m + 2sm + 2st + 41a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		47♀♂	54	Multiple	4m + 1sm + 2st + 40a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		47♀♂	53	Multiple	3m + 1sm + 2st + 41a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		47♀♂	52	Multiple	2m + 1sm + 2st + 42a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		46♀♂	54	Multiple	4m + 2sm + 2st + 38a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		46♀♂	53	Multiple	3m + 2sm + 2st + 39a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		46♀♂	52	Multiple	2m + 2sm + 2st + 40a		Porto et al. (2014a)
	lanceolata	Onça Stream, Coxim, MS, Brazil	18°32’20.0"S	54°33’43.0"W		45	53		4m + 2sm + 37a		Porto et al. (2014a)
	latirostris (Boulenger, 1900)	Laranjinha River, Ventania, PR, Brazil			Cinzas	46♀♂	60	Simple	10m + 4sm + 32st/a		Primo et al. (2017)
	latirostris	Barra Grande River, Prudentópolis, PR, Brazil			Ivaí	46♀♂	60	Simple	10m + 4sm + 32st/a		Primo et al. (2017)
	latirostris	Das Pedras River, Ventania, PR, Brazil	24°07’39.29"S	50°14’16.21"W		46♀♂	60	Simple	10m + 4sm + 32st/a		Glugoski et al. (2018)
	latirostris	Piumhi River, Piumhi, MG, Brazil	20°31’55"S	46°02’42"W		48♀♂	60	Simple	6m + 6sm + 36st/a		Glugoski et al. (2018)
	latirostris	Laranjinha River, Ventania, PR, Brazil	23°58’13"S	50°14’01"W		46♀♂	60	Simple	10m + 4sm + 32st/a		Glugoski et al. (2022)
	latirostris	Piumhi River, Piumhi, MG, Brazil	20°31’55"S	46°02’42"W		48♀♂	60	Simple	6m + 6sm + 36st/a		Glugoski et al. (2022)
	lima (Kner, 1853)	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	70♀♂	72	Simple	2st + 68a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	70♀♂	74	Simple	2m + 2st + 66a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	69♀♂	72	Simple	1m + 2st + 66a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	69♀♂	73	Simple	1m + 1sm + 2st + 65a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	68♀♂	72	Simple	2m + 2st + 64a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	68♀♂	74	Simple	1m + 1m + 2sm + 2st + 62a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	66♀♂	72	Simple	2m + 1m + 1m + 2st + 60a		Rosa et al. (2012)
	lima	Ribeira River, Areia stream, or Açungui River Ponta Grossa, PR, Brazil			Paraná	66♀♂	72	Simple	1m + 1m + 1m + 1st + st + 60a		Rosa et al. (2012)
	longicauda Reis, 1983	Cerquinha Lagoon, RS, Brazil	30°13’56.5"S	50°15’40.9"W	Southeast Atlantic	70♀♂		Simple	8m/sm + 62st/a		Venturelli et al. (2021)
	malabarbai Rodriguez & Reis, 2008	Forquetinha River, RS, Brazil	29°24’22.4"S	52°03’19.2"W	Southeast Atlantic	64♀♂		Simple	4m/sm + 60st/a		Venturelli et al. (2021)
	microlepidogaster (Regan, 1904)	Forquetinha River, RS, Brazil	29°24’22.4"S	52°03’19.2"W	Southeast Atlantic	68♀♂		Simple	68st/a		Venturelli et al. (2021)
	n. sp.	Betari River, São Paulo, Brazil				70♀♂			2sm + 68a		Alves et al. (2003)
	parva (Boulenger, 1895)					48♀♂					Hinegardner, Rosen (1972). Reported as Loricaria parva
	parva	Miranda River, Brazil			Paraguai	60♀♂		Simple	6m/sm + 54st/a		Takagui et al. (2020)
	pentamaculata Langeani & de Araujo, 1994	Tauá stream, Brazil			Paraná	56-59♀♂	64-65		8m/sm + 48st/a or 9m/sm + 47st/a + 0-3Bs		Porto et al., 2010
	pentamaculata	Jacuaca stream, tributary of Tibagi River, PR, Brazil				56♀♂			8m/sm + 48st/a		Maia et al. (2010)
	pentamaculata	Água do Oito stream, tributary of Tibagi River, PR, Brazil				56♀♂			8m/sm + 48st/a		Maia et al. (2010)
	pentamaculata	Tatupeba stream, Brazil			Paraná	56♀♂	64	Multiple	8m/sm + 48st/a		Porto et al. (2011)
	pentamaculata	Tauá stream, Brazil			Paraná	56♀♂	64-65	Simple	8m/sm + 48st/a or 9m/sm + 47st/a		Porto et al. (2011)
	pentamaculata	Keller River, Brazil			Paraná	56♀♂	64	Simple	8m/sm + 48st/a		Porto et al. (2011)
	pentamaculata	Barra Grande River, Prudentópolis, PR, Brazil			Ivaí	56♀♂	70	Simple	4m + 10sm + 42st/a		Primo et al. (2017)
	pentamaculata	Barra Grande River, Prudentópolis, PR, Brazil			Ivaí	54♀♂	64	Simple	6m + 4sm + 44st/a		Primo et al. (2018)
	pentamaculata	Juruba River, Apucarana, PR, Brazil			Tibagi	56♀♂	70	Simple	4m + 10sm + 42st/a		Primo et al. (2017)
	pentamaculata	Itiz stream, Brazil			Ivaí	56♀♂	64	Simple	8m/sm + 48st/a		Cius et al. (2022)
	pentamaculata karyomorph A	Barra Grande river, Prudentópolis, PR, Brazil	25°05’21.0"S	50°57’22.0"W	Ivaí	56♀♂	64	Simple			Glugoski et al. (2023)
	pentamaculata karyomorph B	Barra Grande river, Prudentópolis, PR, Brazil	25°05’21.0"S	50°57’22.0"W	Ivaí	55♀♂	64	Simple			Glugoski et al. (2023)
	pentamaculata karyomorph C	Barra Grande river, Prudentópolis, PR, Brazil	25°05’21.0"S	50°57’22.0"W	Ivaí	54♀♂	64	Simple			Glugoski et al. (2023)
	quadrensis Reis, 1983	Quadros lagoon, RS, Brazil	29°44’42.8"S	50°06’54.3"W	Southeast Atlantic	70♀♂		Simple	6m/sm + 64st/a		Venturelli et al. (2021)
	reisi Ghazzi, 2008	Chimiray stream, Brazil			Uruguai	60♀♂		Simple	60st/a		Takagui et al. (2020)
	stellata Ghazzi, 2008	Uruguai River, São Carlos, SC, Brazil			Uruguai	54♀♂	74	Simple	6m + 14sm + 34st/a		Primo et al. (2017)
	strigilata (Hensel, 1868)	Forquetinha River, RS, Brazil			Southeast Atlantic	68♀♂		Simple	6m/sm + 62st/a		Maia et al. (2010)
	teffeana (Steindachner, 1879)	Tefé River, Tefé, AM, Brazil	3°21’14.9"S	64°40’27.9"W	Amazon	33♂, 34♀	34	Simple	1sm + 32a	X₁X₁X₂X₂/X₁X₂Y	Marajó et al. (2022)
Spatuloricaria	sp.	Xingu River, Altamira, PA, Brazil	3°19’30.0"S	52°11’10.0"W	Xingu – Amazon	66♀♂	92	Simple	4m + 7sm + 6st + 16a		Ferreira et al. (2014)
	sp.	Caripetuba River, Abaetuba, PA, Brazil	1°37’23.5"S	48°55’33.0"W		66♀♂	82		6m + 10sm + 50a		Almeida et al. (2023)
Sturisoma	barbatum (Kner, 1853)	Miranda River, Brazil			Paraguai	66♀♂		Simple	20m + 18sm + 16st + 12a		Takagui et al. (2020)
	cf. nigrirostrum Fowler, 1940	Araguaia River, Barra do Garças, MT, Brazil			Tocantins-Araguaia	74♀♂			20m + 18sm + 36st/a		Artoni, Bertollo (2001)
Hypoptopomatinae
Corumbataia	cuestae Britski, 1997	Alambari stream, Botucatu, SP, Brazil				54♀♂	108	Simple	34m + 20sm		Cristina et al. (2005)
	cuestae	Lapa stream, SP, Brazil			Paraná	54♀♂		Simple	14m + 10sm + 3st/a		Camilo, Moreira Filho (2005)
	tocantinensis Britski, 1997	Vermelho River, Goiás, GO, Brazil				54♀♂	108	Simple	28m + 26sm		Cristina et al. (2005)
Hisonotus	depressicauda (Miranda Ribeiro, 1918)	Santo Inácio River, Bofete, SP, Brazil				54♀♂		Simple	14m + 18sm + 2st + 10a		Andreata et al. (1994). Reported as Microlepidogaster depressicauda
	leucofrenatus (Miranda Ribeiro, 1908)	Poço Grande stream, Juquiá, SP, Brazil				54-56♀♂		Simple	24m + 26sm + 4st + 2a	ZZ/ZW	Andreata et al. (1993). Reported as Microlepidogaster leucofrenatus
	leucofrenatus	Marumbi River, Morretes, PR, Brazil				54-56♀♂		Simple	22m + 24sm + 4st + 2a	ZZ/ZW (discarded by Andreata et al., 2010)	Andreata et al. (1993). Reported as Microlepidogaster leucofrenatus
	leucofrenatus	Cavalo stream, Jaraguá do Sul, SC, Brazil				54♀♂		Simple	22m + 24sm + 6st + 2a		Andreata et al. (2006)
	leucofrenatus	Marumbi River, Morretes, PR, Brazil				54-55♀♂		Simple	22m + 24sm + 4st + 2a		Andreata et al. (2010)
	leucofrenatus	Cavalo stream, Jaraguá do Sul, SC, Brazil				54♀♂		Simple	22m + 24sm + 6st + 2a		Andreata et al. (2010)
	nigricauda (Boulenger, 1891)	Guaíba River, Eldorado do Sul, RS, Brazil				54♀♂		Simple	26m + 20sm + 8st		Andreata et al. (2006)
	sp. A	Paraitinga River, Salesópolis, SP, Brazil				54♀♂		Simple	26m + 26sm + 2st		Andreata et al. (2006)
	sp. D	Grande stream, Pindamonhangaba, SP, Brazil				54♀♂		Simple	26m + 26sm + 2st		Andreata et al. (2006)
Hypoptopoma	inexspectatum (Holmberg, 1893)	Piraí River, Poconé, MT, Brazil				54♀♂	82	Simple	10m + 18sm + 8st + 18a		Cristina et al. (2005). Reported as Hypoptopoma guentheri

Isbrueckerichthys	duseni (Miranda Ribeiro, 1907)	Açungui River, Brazil			Ribeira	54♀♂	108	Simple	20m + 20sm + 14st		Ziemniczak et al. (2012)
	duseni	Betari River, Iporanga, SP, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
Kronichthys	lacerta (Nichols, 1919)	Marumbi River, Morretes, PR, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
	lacerta	Marumbi River, Morretes, PR, Brazil				54♀♂		Simple	20m + 20sm + 14st		Alves et al. (2012a)
	lacerta	Açungui River, Brazil			Ribeira	54♀♂	108	Simple	22m + 22sm + 10st		Ziemniczak et al. (2012)
	subteres Miranda Ribeiro, 1908	Betari River, Iporanga, SP, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
Microlepidogaster	sp. A	Alambari River, Botucatu, SP, Brazil				54♀♂		Multiple	30m + 20sm + 4st		Andreata et al. (1994)
	sp. B	Móia stream, Penápolis, SP, Brazil				54♀♂		Simple	22m + 28sm + 4st		Andreata et al. (1994)
Neoplecostomus	microps (Steindachner, 1877)	Paraitinga River, Brazil	22°52’22.5"S	44°51’41.0"W	Paraíba do Sul	54♀♂	108		24m + 20sm + 10st		Kavalco et al. (2004)
	microps	Paraitinga River, Brazil	22°52’22.5"S	44°51’41.0"W	Paraíba do Sul	54♀♂	108	Simple	24m + 20sm + 10st		Kavalco et al. (2005)
	microps	Grande stream, Campos do Jordão, SP, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
	microps	Paraíba do Sul River, Brazil				54♀♂		Simple	24m + 20sm + 10st		Alves et al. (2012a)
	paranensis Langeani, 1990	Sapateiro stream, Barbacena, MG, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
	paranensis	Hortelã stream, Botucatu, SP, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
	yapo Zawadzki, Pavanelli & Langeani, 2008	Verde River, PR, Brazil			Tibagi	54♀♂	108	Simple	18m + 20sm + 16st		Ziemniczak et al. (2012)
Otocinclus	aff. vestitus	Livramento River, Belém, PA, Brazil				72♀♂		Simple	22m + 12sm + 4st + 34a		Andreata et al. (1994)
	affinis Steindachner, 1877	Biguá River, Miracatu, SP, Brazil				54♀♂		Simple	46m + 8sm		Andreata et al. (1994)
	affinis	Bonito River, Niterói, RJ, Brazil				54♀♂		Simple	40m + 12sm + 12st		Andreata et al. (1994)
	flexilis Cope, 1894	Santo Antônio da Patrulha, RS, Brazil				54♀♂	108	Simple	36m + 18sm		Cristina et al. (2005)
	vittatus Regan, 1904	Taquari River, Coxim, MS, Brazil				54♀♂	108	Multiple	36m + 18sm		Cristina et al. (2005)
	vittatus	Cuiabá River, Santo Antônio do Leverger, MT, Brazil				54♀♂	76	Simple	12m + 10sm + 14st + 18a		Cristina et al. (2005)
Otothyris	juquiae Garavello, Britski & Schaefer, 1998	Rio Preto stream, Itanhaém, SP, Brazil				54♀♂	96	Simple	32m + 10sm + 12st		Cristina et al. (2005)
	travassosi Garavello, Britski & Schaefer, 1998	Ribeira da Terra Firme River, Canavieiras, BA, Brazil				54♀♂	96	Simple	26m + 16sm + 12st		Cristina et al. (2005)
Otothyropsis	cf. polyodon	Córrego Dourado, MT, Brazil	23°51’04.9"S	54°25’13.9"W		54♀♂	108	Simple	18m + 28sm + 8st		Fernandes et al. (2016)
Pareiorhaphis	splendens (Bizerril, 1995)	Marumbi River, Morretes, PR, Brazil				54♀♂			20m + 30sm + 4st		Alves et al. (2005). Reported as Hemipsilichthys splendens
	splendens	Marumbi River, Morretes, PR, Brazil				54♀♂		Simple	20m + 20sm + 14st		Alves et al. (2012a)
	splendens	São João River, Garuva, SC, Brazil				54♀♂			20m + 30sm + 4st		Alves et al. (2005). Reported as Hemipsilichthys splendens
	steindachneri (Miranda Ribeiro, 1918)	Cavalo stream, Jaraguá do Sul, SC, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005). Reported as Hemipsilichthys steindachneri
	vestigipinnis (Pereira & Reis, 1992)	Caveiras River, Painel, SC, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005). Reported as Hemipsilichthys vestigipinnis
Pareiorhina	brachyrhyncha Chamon, Aranda & Buckup, 2005	Ribeirão Grande creek, Pindamonhangaba, SP, Brazil			Paraíba do Sul	54♀♂		Simple	18m + 30sm + 6st		Centofante et al. (2011)
	rudolphi (Miranda Ribeiro, 1911)	Convento stream, Pindamonhangaba, SP, Brazil				54♀♂			26m + 16sm + 12st		Alves et al. (2005)
	rudolphi	Marimbondo creek, Visconde de Mauá, RJ, Brazil			Paraíba do Sul	54♀♂		Simple	18m + 32sm + 4st		Centofante et al. (2011)
Parotocinclus	maculicauda (Steindachner, 1877)	Açungui River, Brazil			Ribeira	54♀♂	108	Simple	20m + 20sm + 14st		Ziemniczak et al. (2012)
	maculicauda	Poço Grande stream, Juquiá, SP, Brazil				54♀♂		Simple	20m + 32sm + 2st		Andreata et al. (1994)
Pseudotocinclus	tietensis (Ihering, 1907)	Tietê River, Paranapiacaba, SP, Brazil				54♀♂		Simple	26m + 20sm + 6st	XX/XY	Andreata et al. (1992)
	n. sp.	Juquiá River, Juquitiba, SP, Brazil				54♀♂		Simple	22m + 24sm + 8st		Cristina et al. (2005)
Pseudotothyris	obtusa (Miranda Ribeiro, 1911)	Tributary of Itanhaém River, Itanhaém, SP, Brazil				54♀♂		Simple	26m + 18sm + 4st + 6a		Andreata et al. (1994)
Schizolecis	guentheri (Miranda Ribeiro, 1918)	Parati-Mirim stream, Parati, RJ, Brazil				54♀♂	102	Simple	30m + 18sm + 6a		Cristina et al. (2005)
	guentheri	Sítio do Meio stream, Mongaguá, SP, Brazil				54♀♂	102	Simple	30m + 18sm + 6a		Cristina et al. (2005)
	guentheri	Descoberto stream, Guaratuba, PR, Brazil				54♀♂	102	Simple	30m + 18sm + 6a		Cristina et al. (2005)
	guentheri	Garuva River, Garuva, SC, Brazil				54♀♂	102	Simple	30m + 18sm + 6a		Cristina et al. (2005)
Hypostominae
Ancistomus	spilomma (Cardoso & Lucinda, 2003)	Araguaia River, Pontal do Araguaia, MT, Brazil	15°50’15.0"S	51°58’43.0"W		52♀♂		Multiple	25m + 21sm + 6st ♀, 24m + 22sm + 6st ♂	ZZ/ZW	Oliveira et al. (2006). Reported as Hemiancistrus spilomma
	spinosissimus (Cardoso & Lucinda, 2003)	Araguaia River, Pontal do Araguaia, MT, Brazil	15°50’15.0"S	51°58’43.0"W		52♀♂		Simple	26m + 22sm + 4st		Oliveira et al. (2006). Reported as Hemiancistrus spinosissimus
	feldbergae	PA, Brazil				52♀♂		Simple	38m/sm + 14st		Pety et al. (2018). Reported as Peckoltia feldbergae
Ancistrus	abilhoai Bifi, Pavanelli & Zawadzki, 2009	Iguaçu River, União da Vitória, PR, Brazil	26°15’64”S*	51°06’28”W*		48♀♂	90	Simple	22m + 14sm + 6st + 6a		Ribeiro et al. (2015)
	abilhoai	Timbó River, Santa Cruz do Timbó, SC, Brazil	26°30’37.0"S	50°46’55.0"W		48♀♂	90	Simple	22m + 14sm + 6st + 6a		Ribeiro et al. (2015)
	aff. dolichopterus Kner, 1854	Aiapuá Lake, Piagaçu-Purus Sustainable Development Reserve, AM, Brazil	4°27’26.0"S	62°11’56.0"W		52♀♂	78, 79	Simple	16m + 8sm + 2st + 26a ♂, 16m + 9sm + 2st + 25a ♀	ZZ/ZW	Oliveira et al. (2007)
	aff. dolichopterus	Aiapuá Lake, Piagaçu-Purus Sustainable Development Reserve, AM, Brazil	4°27’26.0"S	62°11’56.0"W		52♀♂	78, 79	Simple	16m + 8sm + 2st + 26a ♂, 16m + 9sm + 2st + 25a ♀	ZZ/ZW	Favarato et al. (2016). Reported as Ancistrus sp. Balbina
	aguaboensis Fisch-Muller, Mazzoni & Weber, 2001	Ribeirão Bandeirinha, GO, Brazil			Tocantins-Araguaia	50♀♂	86	Simple	16m + 10sm + 4st + 20a		Glugoski et al. (2020)
	aguaboensis	Ribeirão Bandeirinha, GO, Brazil			Tocantins-Araguaia	50♀♂	86	Simple	16m + 10sm + 4st + 20a		Schott et al. (2022)
	cf. dubius Eigenmann & Eigenmann, 1889	Serra das Araras, Barra do Bugres, MT, Brazil				44♀♂		Simple	18m + 10sm + 16st/a	ZZ/ZW	Mariotto et al. (2004)
	cf. dubius	Coxipó River, Chapada dos Guimarães, MT, Brazil				42♀♂	84		24m + 10sm + 8st		Mariotto et al. (2006)
	cf. dubius	Pari creek, Cuiabá, MT, Brazil				42♀♂	84		24m + 10sm + 8st	XX/XY	Mariotto et al. (2006)
	cf. dubius	Flechas creek, Cáceres, MT, Brazil				42♀♂	84		24m + 10sm + 8st	XX/XY	Mariotto et al. (2006)
	cf. dubius	Fundo creek, Poconé, MT, Brazil				42♀♂	84		24m + 10sm + 8st	XX/XY	Mariotto et al. (2006)
	cf. dubius	Flechas stream, Brazil	15°58’07.0"S	57°19’07.0"W	Paraguai	42♀♂	84	Simple	24m + 10sm + 8st	XX/XY	Mariotto et al. (2011)
	cf. dubius	Fundo stream, Brazil	16°14’17.0"S	56°37’31.0"W	Paraguai	42♀♂	84	Simple	24m + 10sm + 8st	XX/XY	Mariotto et al. (2011)
	cf. dubius	Pari stream, Brazil	15°36’06.0"S	56°12’19.0"W	Paraguai	42♀♂	84	Simple	24m + 10sm + 8st	XX/XY	Mariotto et al. (2011)
	cf. multispinis (Regan, 1912)	Ribeirão Grande, SP, Brazil			Paraíba do Sul	52♀♂	84	Simple	16m + 10sm + 6st + 20a		Glugoski et al. (2020)
	cf. multispinis	Ribeirão Grande, SP, Brazil			Paraíba do Sul	52♀♂	84	Simple	16m + 10sm + 6st + 20a		Schott et al. (2022)
	cirrhosus (Valenciennes, 1836)	Maracapucú River, Abaetuba, PA, Brazil	1°45’29.2"S	48°56’57.0"W	Tocantins-Araguaia	34♀♂	68	Simple	20m + 14sm		Santos da Silva et al. (2023)
	cirrhosus	Ilha do Capim, Abaetuba, PA, Brazil	1°34’02.8"S	48°51’49.1"W	Tocantins-Araguaia	34♀♂	68	Simple	20m + 14sm		Santos da Silva et al. (2023). Reported as Ancistrus sp. 2
	cirrhosus	Maracapucú River, Abaetuba, PA, Brazil	1°45’29.2"S	48°56’57.0"W	Tocantins-Araguaia	34♀♂	68	Simple	20m + 14sm		Santos da Silva et al. (2022a). Reported as Ancistrus sp. 2
	cirrhosus	Ilha do Capim, Abaetuba, PA, Brazil	1°34’02.8"S	48°51’49.1"W	Tocantins-Araguaia	34♀♂	68	Simple	20m + 14sm		Santos da Silva et al. (2022a)
	claro Knaack, 1999	Coxipó River, Brazil	15°21’59.0"S	55°57’11.0"W	Paraguai	54♀♂	84	Simple	14m + 8sm + 8st + 24a		Mariotto et al. (2011)
	claro	Coxipó River, Chapada dos Guimarães, MT, Brazil	15°21’00.0"S	55°57’00.0"W	Paraguai	54♀♂	84	Simple	14m + 8sm + 8st + 24a		Mariotto et al. (2013)
	clementinae Rendahl, 1937	Río Palenque (Cantón Pasaje), Ecuador				52♂, 53♀	100, 101	Simple	48m/sm + 5a ♀, 48m/sm + 4st/a ♂	ZZ/ZW₁W₂	Nirchio et al. (2023)
	clementinae	Río La Moquillada (Cantón Las Lajas), Ecuador				52♂, 53♀	100, 101	Simple	48m/sm + 5a ♀, 48m/sm + 4st/a ♂	ZZ/ZW₁W₂	Nirchio et al. (2023)
	cuiabae Knaack, 1999	Arrombado, Poconé, MT, Brazil	16°21’21.0"S	56°27’55.0"W		34♀♂	68	Simple	20m + 8sm + 6st		Mariotto et al. (2009)
	cuiabae	Arrombado, Poconé, MT, Brazil	16°21’21.0"S	56°27’55.0"W		34♀♂	67	Multiple	19m + 8sm + 6st + 1a		Mariotto et al. (2009)
	cuiabae	Arrombado, Poconé, MT, Brazil	16°21’21.0"S	56°27’55.0"W		34♀♂	66	Simple	18m + 8sm + 6st + 2a		Mariotto et al. (2009)
	cuiabae	Arrombado bay, Brazil	16°21’21.0"S	56°27’55.0"W	Paraguai	34♀♂	68	Simple	20m + 8sm + 6st		Mariotto et al. (2011)
	dolichopterus Kner, 1854	Demeni River, Barcelos, AM, Brazil	0°25’19.0"S	62°54’42.0"W	Negro	52♀♂	79, 8	Simple	12m + 12sm + 4st + 24a ♂, 11m + 12sm + 4st + 25a ♀	Z₁Z₁Z₂Z₂/Z₁Z₂W₁W₂	Oliveira et al. (2008)
	dolichopterus	Demeni River, Barcelos, AM, Brazil	0°25’19.0"S	62°54’42.0"W	Negro	52♀♂	79, 8	Simple	12m + 12sm + 4st + 24a ♂, 11m + 12sm + 4st + 25a ♀	Z₁Z₁Z₂Z₂/Z₁Z₂W₁W₂	Favarato et al. (2016)
	dubius Eigenmann & Eigenmann, 1889	Barretinho stream, Presidente Figueiredo, AM, Brazil	1°58’20.0"S	59°29’48.0"W		38♀, 39♂	76, 78	Simple	27m + 10sm + 2st ♂, 26m + 10sm + 2st ♀	XX/XY₁Y₂	Oliveira et al. (2008)
	dubius	Barretinho stream, Presidente Figueiredo, AM, Brazil	1°58’20.0"S	59°29’48.0"W		38♀, 39♂	76, 78	Simple	27m + 10sm + 2st ♂, 26m + 10sm + 2st ♀	XX/XY₁Y₂	Favarato et al. (2016). Reported as Ancistrus sp. Barcelos
	maximus de Oliveira, Zuanon, Zawadzki & Rapp Py-Daniel, 2015	Igarapé Macoari, RR, Brazil	1°10’00.0"N	61°51’00.0"W		46♀♂	81, 82	Simple	18m + 11sm + 6st + 11a ♂, 18m + 12sm + 6st + 10a ♀	XX/XY	Oliveira et al. (2010)
	maximus	Igarapé Macoari, RR, Brazil	1°10’00.0"N	61°51’00.0"W		46♀♂	81, 82	Simple	18m + 11sm + 6st + 11a ♂, 18m + 12sm + 6st + 10a ♀	XX/XY	Favarato et al. (2016)
	multispinis (Regan, 1912)	Itapocu River, SC, Brazil				52♀♂		Simple	28m/sm + 24st/a		Alves et al. (2003)
	n. sp. 1	São Francisco River, AC, Brazil				38♀♂		Simple	30m/sm + 8st		Alves et al. (2003)
	n. sp. 1	Vermelho River, GO , Brazil				39♂, 40♀		Simple	33m + 6sm or 34m + 6sm	XX/X0	Alves et al. (2006)
	n. sp. 2	Betari River, SP, Brazil				52♀♂		Simple	30m/sm + 20st/a		Alves et al. (2003)
	n. sp. 2	Garuva River, SC, Brazil				52♀♂		Simple	10m + 16sm + 12st + 14a		Alves et al. (2006)
	ranunculus Muller, Rapp Py-Daniel & Zuanon, 1994	Xingu River, Altamira, PA, Brazil	3°15’21.0"S	52°12’45.0"W	Xingu	48♀♂	82	Simple	20m + 8sm + 6st + 14a ♂, 19m + 9sm + 6st + 14a ♀	ZZ/ZW	Oliveira et al. (2007)
	ranunculus	Xingu River, Altamira, PA, Brazil	3°15’21.0"S	52°12’45.0"W	Xingu	48♀♂	82	Simple	20m + 8sm + 6st + 14a ♂, 19m + 9sm + 6st + 14a ♀	ZZ/ZW	Favarato et al. (2016). Reported as Ancistrus sp. Piagaçu
	sp.	Criminoso stream, MS, Brazil	18°29’20.0"S	54°45’14.0"W	Paraguai	42♀♂	84	Simple	18m + 16sm + 8st		Prizon et al. (2016)
	sp.	Ivaí River, Brazil			Ivaí	50♀♂	88		20m + 12sm + 6st + 12a		Barros et al. (2017)
	sp.	Ivaí River, Brazil			Ivaí	50♀♂	88		20m + 12sm + 6st + 12a		Schott et al. (2022)
	sp. 01	Pipa stream, Serra de São Vicente, MT, Brazil	14°33’00.0"S	57°24’00.0"W	Paraguai	54♀♂	84	Simple	14m + 8sm + 8st + 24a		Mariotto et al. (2013)
	sp. 03	Pari stream, Cuiabá, MT, Brazil	15°36’00.0"S	56°12’00.0"W	Paraguai	54♀♂	84	Simple	14m + 8sm + 8st + 24a		Mariotto et al. (2013)
	sp. 04	Sepotuba River, Brazil	14°41’35.0"S	57°48’14.0"W	Paraguai	52♀♂	82	Simple	16m + 8sm + 6st + 22a		Mariotto et al. (2011)
	sp. 04	São José stream, Sepotuba River, Tangará da Serra, MT, Brazil	14°41’00.0"S	57°48’00.0"W	Paraguai	52♀♂	82	Simple	16m + 8sm + 6st + 22a		Mariotto et al. (2013)
	sp. 06	Matrixã River, Brazil	10°03’07.0"S	57°36’27.0"W	Amazon	50♀♂	86	Simple	18m + 10sm + 8st + 14a		Mariotto et al. (2011)
	sp. 06	Matrixã River, Nova Monte Verde, MT, Brazil	10°03’00.0"S	57°36’00.0"W	Amazon	50♀♂	86	Simple	18m + 10sm + 8st + 14a		Mariotto et al. (2013)
	sp. 08	Currupira River, Brazil	15°07’59.0"S	56°49’47.0"W	Paraguai	44♀♂	80	Simple	18m + 10sm + 8st + 8a	ZZ/ZW	Mariotto et al. (2011)
	sp. 1	Quianduba River, Abaetuba, PA, Brazil	1°45’18.2"S	49°00’38.8"W	Tocantins-Araguaia	38♀♂	72	Simple	20m + 14sm + 4st	XX/XY	Santos da Silva et al. (2022a)
	sp. 1	Quianduba River, Abaetuba, PA, Brazil	1°45’18.2"S	49°00’38.8"W	Tocantins-Araguaia	38♀♂	72	Simple	20m + 14sm + 4st	XX/XY	Santos da Silva et al. (2023)
	sp. 1	Maracapucú River, Abaetuba, PA, Brazil	1°45’29.2"S	48°56’57.0"W	Tocantins-Araguaia	38♀♂	72	Simple	20m + 14sm + 4st	XX/XY	Santos da Silva et al. (2023)
	sp. 1	Ilha do Capim, Abaetuba, PA, Brazil	1°34’02.8"S	48°51’49.1"W	Tocantins-Araguaia	38♀♂	72	Simple	20m + 14sm + 4st	XX/XY	Santos da Silva et al. (2023)
	sp. 13	Salgadinho stream, Brazil	14°40’14.0"S	52°21’50.0"W	Tocantins-Araguaia	40♀♂	80	Simple	26 m + 10sm + 4st		Mariotto et al. (2011)
	sp. 13	Salgadinho stream, Nova Xavantina, MT, Brazil	14°40’00.0"S	52°21’00.0"W	Tocantins-Araguaia	40♀♂	80	Simple	30m + 6sm + 4st		Mariotto et al. (2013)
	sp. Catalão	Catalão Lake, AM, Brazil	3°10’45.0"S	59°54’25.0"W	Amazonas	34♀♂		Simple	22m + 8sm + 4st	XX/XY	Favarato et al. (2016)
	sp. Dimona	Fazenda Dimona, Brazil	2°19’00.0"S	60°04’00.0"W		52♀♂	78	Simple	16m + 8sm + 2st + 26a		Oliveira et al. (2010)
	sp. L1	Mourão River, Brazil				50♀♂		Simple	12m + 18sm + 12st + 8a		Prizon et al. (2017)
	sp. L10	São Francisco Falso River, Brazil				50♀♂		Simple	10m + 18sm + 16st + 6a		Prizon et al. (2017)
	sp. L2	Stream 19, Brazil				50♀♂		Simple	12m + 18sm + 12st + 8a ♀, 11m + 18sm + 13st + 8a ♂	XX/XY	Prizon et al. (2017)
	sp. L3	Keller River, Brazil				50♀♂		Simple	12m + 18sm + 12st + 8a ♀, 11m + 18sm + 13st + 8a ♂	XX/XY	Prizon et al. (2017)
	sp. L6	São Francisco Verdadeiro River, Brazil				50♀♂		Simple	14m + 16sm + 14st + 6a		Prizon et al. (2017)
	sp. L8	Arroyo San Juan, Misiones, Posada, Argentina				50♀♂		Simple	10m + 14sm + 12st + 14a		Prizon et al. (2017)
	sp. L9	Ocoí River, Brazil			Paraná	50♀♂		Simple	10m + 18sm + 16st + 6a		Prizon et al. (2017)
	sp. Purus	Purus River, AM, Brazil	4°51’00.0"S	62°38’00.0"W		34♀♂	68	Simple	21m + 11sm + 2st ♂, 20m + 12sm + 2st ♀	XX/XY	Oliveira et al. (2010)
	sp. Purus	Purus River, AM, Brazil	4°51’00.0"S	62°38’00.0"W		34♀♂	68	Simple	21m + 11sm + 2st ♂, 20m + 12sm + 2st ♀	XX/XY	Favarato et al. (2016). Reported as Ancistrus sp. Macoari
	sp. Trombetas	Trombetas River, PA, Brazil	1°27’00.0"S	56°21’00.0"W	Trombetas	38♀♂	73	Simple	22m + 8sm + 5st + 3a		Oliveira et al. (2010)
	sp. Vermelho	Demeni River, middle Negro River, AM, Brazil	0°23’00.0"S	62°50’00.0"W	Negro	42♀♂	78	Simple	26m + 6sm + 4st + 6a		Oliveira et al. (2010)
	taunayi Miranda Ribeiro, 1918	Cascalho stream, Mondaí, SC, Brazil			Uruguai	50♀♂		Simple	22m + 10sm + 10st + 8a	ZZ/ZW	Konerat et al. (2015)
	tombador Fisch-Muller, Cardoso, da Silva & Bertaco, 2005	Preto River, Diamantino, MT, Brazil	14°40’00.0"S	52°21’00.0"W	Amazon	50♀♂	84	Simple	14m + 12sm + 8st + 16a		Mariotto et al. (2013)
Aphanotorulus	emarginatus (Valenciennes, 1840)	Araguaia River, Barra do Garças, MT, Brazil				52♀♂		Simple	16m + 30sm + 6st		Artoni, Bertollo (2001). Reported as Hypostomus emarginatus
Araichthys	loro Zawadzki, Bifi & Mariotto, 2016	Papagaio River, Brazil	12°79′67”S*	58°39’02”W*	Amazon	54♀♂	108	Simple	36m + 12sm + 6st		Mariotto et al. (2019)
Baryancistrus	aff. niveatus	Xingu River, Altamira, PA, Brazil	3°12’48.0"S	52°12’41.7"W	Xingu – Amazon	52♀♂	104	Simple	16m + 32sm + 4st		Souza et al. (2004)
	xanthellus Rapp Py-Daniel, Zuanon & de Oliveira, 2011	Volta Grande do Xingu, Altamira, PA, Brazil	3°36’31.5"S	51°34’57.4"W	Xingu-Amazon	52♀♂	104	Simple	16m + 28sm + 8st		Medeiros et al. (2016)
	xanthellus	Volta Grande do Xingu, Altamira, PA, Brazil	3°23’28.2"S	51°44’29.3"W	Xingu-Amazon	52♀♂	104	Simple	16m + 28sm + 8st		Medeiros et al. (2016)
	xanthellus	Volta Grande do Xingu, Altamira, PA, Brazil	3°22’29.7"S	51°42’25.0"W	Xingu-Amazon	52♀♂	104	Simple	16m + 28sm + 8st		Medeiros et al. (2016)
	xanthellus	Volta Grande do Xingu, Altamira, PA, Brazil	3°35’38.6"S	51°49’36.0"W	Xingu-Amazon	52♀♂	104	Simple	16m + 28sm + 8st		Medeiros et al. (2016)
Corymbophanes	n. sp.	Chopotó River, Desterro de Melo, MG, Brazil				54♀♂			20m + 20sm + 14st		Alves et al. (2005)
Hemiancistrus	sp.	Araguaia River, Barra do Garças, MT, Brazil				52♀♂		Simple	20m + 20sm + 12st/a		Artoni, Bertollo (2001)
Hypancistrus	cf. debilittera Armbruster, Lujan & Taphorn, 2007	Uatumã River, AM, Brazil			Uatumã-Amazon	52♀♂	104	Multiple	34m/sm + 18st		Santos et al. (2023)
	cf. debilittera	Uatumã River, AM, Brazil			Uatumã-Amazon	52♀♂	105	Multiple	34m/sm + 18st		Silva et al. (2014)
	L066	Xingu River, PA, Brazil			Xingu-Amazon	52♀♂	104	Multiple	40m/sm + 12st/a		Cardoso et al. (2016)
	L333	Xingu River, PA, Brazil			Xingu-Amazon	52♀♂	104	Multiple	40m/sm + 12st/a		Cardoso et al. (2016)
	sp. "Pão"	Xingu River, Belo Monte hydroeletric plant	3°06’12.8"S	51°43’53.9"W		52♀♂	94		40m/sm + 12st/a		Almeida et al. (2023)
	sp. "Pão"	Altamira, PA				52♀♂	104		22m + 18sm + 12st		Santos et al. (2023)
	zebra Isbrücker & Nijssen, 1991	Xingu River, PA, Brazil			Xingu-Amazon	52♀♂	104	Multiple	34m/sm + 18st		Silva et al. (2014)
	zebra	Xingu River, PA, Brazil			Xingu-Amazon	52♀♂	104	Multiple	40m/sm + 12st/a		Cardoso et al. (2016)
	zebra	Xingu River, Belo Monte hydroeletric plant	3°06’12.8"S	51°43’53.9"W		52♀♂	94		40m/sm + 12st/a		Almeida et al. (2023)
	zebra	Altamira, PA				52♀♂	104		16m + 20sm + 16st		Silva et al. (2014)
Hypostomus	aff. agna (Miranda Ribeiro, 1907)	Cavalo stream, Jaraguá do Sul, SC, Brazil			Southern Brazil Coast	74♀♂		Multiple	8m + 10sm + 32st + 24a		Martinez et al. (2011)
	aff. ancistroides (Ihering, 1911)	São Miguel Arcanjo, SP, Brazil	23°54’14.3"S	47°55’46.8"W	Parapanema	66♀♂		Multiple	16m + 12sm + 12st + 26a ♀, 17m + 12sm + 12st + 25a ♂	XX/XY	Rocha-Reis et al. (2018)
	aff. ancistroides	Keller River, Brazil	23°38’25.9"S	51°51’32.8"W		68♀♂	119, 118	Multiple	16m + 13sm + 22st + 17a ♀, 16m + 12sm + 22st + 18a ♂	ZZ/ZW	Kamei et al. (2017)
	aff. ancistroides	Ponta Grossa, PR, Brazil			Tibagi	66♀♂	104	Multiple	12m + 16sm + 10st + 28a		Maurutto et al. (2012)
	aff. cochliodon Kner, 1854	Esparramo stream, Rondonópolis, MT, Brazil	16°30’19.6"S	54°40’31.6"W	Paraguai	64♀♂		Multiple	18m + 20sm + 26st/a		Becker et al. (2014)
	aff. cochliodon	Pitaluga stream, Rondonópolis, MT, Brazil	16°28’38.1"S	54°32’08.3"W	Paraguai	64♀♂		Multiple	18m + 20sm + 26st/a		Becker et al. (2014)
	aff. hermanni (Ihering, 1905)	Keller River, Brazil	23°38’25.9"S	51°51’32.8"W		72♀♂	124	Multiple	12m + 22sm + 18st + 20a		Kamei et al. (2017)
	aff. paulinus (Ihering, 1905)	Piquiri River, PR, Brazil				74♀♂		Simple	10m + 12sm + 20st + 32a		Bueno et al. (2013)
	aff. paulinus	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		76♀♂	102	Simple	8m + 18sm + 50st/a		Rubert et al. (2016)
	aff. tietensis (Ihering, 1905)	Pirapó River, Brazil	23°18’15.0"S	51°53’40.0"W	Paranapanema	76♀♂	114	Simple	8m + 6sm + 24st + 38a		Paula et al. (2022)
	aff. tietensis	Do Campo River, Brazil	24°04’41.0"S	52°26’12.0"W	Ivaí	76♀♂	114	Multiple	8m + 6sm + 24st + 38a		Paula et al. (2022)
	aff. topavae (Gody, 1969)	Mogi Guaçu River, Pirassununga, SP, Brazil				70♀♂	102	Multiple	18m + 14sm + 38st/a		Artoni, Bertollo (1996). Reported as Hypostomus sp. A
	aff. topavae	Mogi Guaçu River, Pirassununga, SP, Brazil				70♀♂	102	Multiple	18m + 14sm + 38st/a		Lorscheider et al. (2015)
	aff. unae (Steindachner, 1878)	Contas River, Brazil	13°51’51.0"S	40°04’54.0"W	Contas	76♀♂	104	Simple	12m + 16sm + 48st/a		Bitencourt et al. (2012)
	aff. unae	Preto do Costa River, Brazil	13°45’84”S*	39°56’47”W*	Contas	76♀♂	108	Simple	12m + 20sm + 44st/a		Bitencourt et al. (2012)
	aff. unae	Oricó River, Brazil	14°08’03.0"S	39°21’30.0"W	Contas	76♀♂	100	Simple	10m + 14sm + 52st/a		Bitencourt et al. (2012)
	aff. unae	Preto do Criciúma River, Brazil	13°55’45.0"S	39°57’57.0"W	Contas	76♀♂	106	Simple	10m + 20sm + 46st/a		Bitencourt et al. (2012)
	affinis (Steindachner, 1877)	Jacuí stream, Brazil	22°52’00.0"S	44°51’00.0"W	Paraíba do Sul	66♀♂	106		14m + 14sm + 12st + 12a		Kavalco et al. (2004)
	affinis	Jacuí stream, Brazil	22°52’00.0"S	44°51’00.0"W	Paraíba do Sul	66♀♂	106	Multiple	14m + 14sm + 12st + 12a		Kavalco et al. (2005)
	affinis	Jacuí creek, Cunha, SP, Brazil	23°02’25.9"S	44°56’02.7"W	Paraíba do Sul	66♀♂	104	Multiple	12m + 12sm + 14st + 28a		Brandão et al. (2018)
	affinis	Paraíba do Sul River, Itaocara, RJ, Brazil	21°39’41.1"S	42°04’28.3"W	Paraíba do Sul	66♀♂	104	Multiple	12m + 12sm + 14st + 28a		Brandão et al. (2018)
	albopunctatus (Regan, 1908)	Mogi Guaçu River, Pirassununga, SP, Brazil				74♀♂	104	Multiple	10m + 20sm + 44st/a		Artoni, Bertollo (1996)
	albopunctatus	Piquiri River, Brazil				74♀♂		Multiple	8m + 14sm + 16st + 36a		Bueno et al. (2013)
	albopunctatus	Piquiri River, Nova Laranjeiras, Brazil	24°56’54.0"S	52°35’49.0"W		74♀♂		Simple	8m + 14sm + 16st + 36a		Bueno et al. (2014)
	albopunctatus	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		74♀♂	104	Multiple	10m + 20sm + 44st/a		Rubert et al. (2016)
	albopunctatus	Mogi Guaçu River, Pirassununga, SP, Brazil				74♀♂	104	Multiple	10m + 20sm + 44st/a		Lorscheider et al. (2015)
	albopunctatus	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		74♀♂			10 + 18sm + 46st/a		Rubert et al. (2022)
	ancistroides (Ihering, 1911)					68♀♂	105M, 106F		10m + 27sm + 31st/a ♂, 10m + 28sm + 30st/a ♀		Michele et al. (1977)
	ancistroides	Rio Monjolinho, São Carlos, SP, Brazil				68♀♂	102	Multiple	16m + 18sm + 34st/a		Artoni, Bertollo (1996). Reported as Plecostomus strigaticeps
	ancistroides	Araquá River, Botucatu, SP, Brazil				68♀♂		Multiple	18m + 10sm + 12st + 28a		Alves et al. (2006)
	ancistroides				Paranapanema	68♀♂	104	Multiple	10m + 26sm + 32st/a		Rubert et al. (2011)
	ancistroides	Corumbataí River, SP, Brazil				68♀♂		Multiple	16m + 4sm + 16st + 32a		Alves et al. (2012b)
	ancistroides	Piquiri River, Formosa do Oeste, PR, Brazil				68♀♂			14m + 14sm + 8st + 32a		Bueno et al. (2012)
	ancistroides	Dourados stream, Mandaguari, PR, Brazil			Pirapó	68♀♂		Multiple	14m + 12sm + 18st + 24a		Endo et al. (2012)
	ancistroides	Maringá stream, Maringá, PR, Brazil			Pirapó	68♀♂		Multiple	16m + 12sm + 18st + 22a		Endo et al. (2012)
	ancistroides	Ximbaúva stream, Ourizona, PR, Brazil			Ivaí	68♀♂		Multiple	8m + 10sm + 18sm + 32a		Endo et al. (2012)
	ancistroides	Piquiri River, Brazil				68♀♂		Multiple	14m + 14sm + 8st + 32a		Bueno et al. (2013)
	ancistroides	Água Boa stream, Mundo Novo, MS, Brazil				68♀♂	116	Multiple	14m + 24sm + 10st + 20a		Fernandes et al. (2012)
	ancistroides	Dourado stream, Mundo Novo, MS, Brazil				68♀♂	116	Multiple	10m + 22sm + 16st + 20a		Fernandes et al. (2012)
	ancistroides	Dourado stream, Mundo Novo, MS, Brazil				68♀♂	120	Multiple	14m + 16sm + 22st + 16a		Traldi et al. (2013)
	ancistroides	Hortelã stream, Botucatu, SP, Brazil	22°56’28.9"S	48°35’03.2"W		68♀♂		Multiple	10m + 20sm + 10st + 28a		Pansonato-Alves et al. (2013)
	ancistroides	Piquiri River, Nova Laranjeiras, Brazil	24°56’54.0"S	52°35’49.0"W		68♀♂		Multiple	14m + 14sm + 8st + 32a		Bueno et al. (2014)
	ancistroides	Monjolinho River, São Carlos, SP, Brazil				68♀♂	102	Multiple	16m + 18sm + 34st/a		Lorscheider et al. (2015)
	cf. heraldoi Zawadzki, Weber & Pavanelli, 2008	Mogi Guaçu River, Pirassununga, SP, Brazil			Mogi Guaçu	72♀♂		Simple	6m + 6sm + 26st + 34a		Martinez et al. (2011)
	cf. plecostomus (Linnaeus, 1758)	Severo stream, Brazil	9°54’30.8"S	56°03’33.9"W	Amazon	68♀♂	120/ 121	Multiple	14m + 24sm + 14st + 16a ♂, 15m + 24sm + 14st + 15a ♀	ZZ/ZW	Oliveira et al. (2015)
	cf. topavae	Carrapato stream, Penápolis, SP, Brazil			Paraná	80♀♂		Multiple	6m + 8sm + 42st + 24a		Martinez et al. (2011)
	cf. wuchereri (Günther, 1864)	Mutum River, Jequié, BA, Brazil	13°43’18.0"S	39°51’20.0"W	Contas	76♀♂	104	Simple	10m + 18sm + 48st/a		Bitencourt et al. (2011b)
	cf. wuchereri	Una River, Valença, BA, Brazil	13°21’55.0"S	39°04’35.0"W	Recôncavo Sul	76♀♂	104	Simple	10m + 18sm + 48st/a		Bitencourt et al. (2011b)
	cochliodon Kner, 1854	Iguaçu River, Brazil				64♀♂		Simple	12m + 16sm + 16st + 20a		Bueno et al. (2014)
	cochliodon	Iguaçu River, Foz do Iguaçu, PR, Brazil	25°38’53.0"S	54°27’28.0"W		64♀♂		Simple	12m + 16sm + 16st + 20a		Rubert et al. (2016)
	cochliodon	Piraputanga River, Cáceres, MT, Brazil	16°03’33.0"S	57°40’33.0"W	Paraguai	64♀♂	100	Multiple	16m + 20sm + 28st/a		Rubert et al. (2016)
	commersoni Valenciennes, 1836	Iguaçu River, Brazil				68♀♂		Multiple	12m + 14sm + 14st + 28a		Bueno et al. (2013)
	commersoni	Lake of Ney Braga hydroeletric plant, Mangueririnha, PR, Brazil				68♀♂	100	Multiple	12m + 12sm + 8st + 36a		Maurutto et al. (2012)
	commersoni	Piquiri River, Nova Laranjeiras, Brazil	24°56’54.0"S	52°35’49.0"W		68♀♂		Multiple	12m + 14sm + 14st + 28a		Bueno et al. (2014)
	commersoni	Iguaçu River, Foz do Iguaçu, PR, Brazil	25°38’53.0"S	54°27’28.0"W		68♀♂		Multiple	12m + 14sm + 14st + 28a		Bueno et al. (2014)
	commersoni	Iguaçu River, PR, Brazil	26°15’01.1"S	51°06’10.7"W	Paraná	68♀♂	106	Multiple	12m + 12sm + 14st + 30a		Lorscheider et al. (2018)
	derbyi (Haseman, 1911)	Iguaçu River, Curitiba, PR, Brazil				66♀♂	82	Multiple	6m + 10sm + 20st + 30a		Maurutto et al. (2012)
	derbyi	Iguaçu River, PR, Brazil	26°15’01.1"S	51°06’10.7"W	Paraná	68♀♂	102	Simple	12m + 12sm + 10st + 34a		Lorscheider et al. (2018)
	faveolus Zawadzki, Birindelli & Lima, 2008	Taquaralzinho River, MT, Brazil				64♀♂		Simple	18m + 8sm + 22st + 16a		Bueno et al. (2013)
	faveolus	Taquaralzinho River, Barra do Garças, MT, Brazil	15°40’42.0"S	52°17’52.0"W		64♀♂		Simple	18m + 8sm + 22st + 16a		Bueno et al. (2014)
	goyazensis (Regan, 1908)	Vermelho River, GO, Brazil				72♀♂		Simple	10m + 16sm + 10st + 36a		Alves et al. (2006)
	hermanni (Ihering, 1905)	Piquiri River, PR, Brazil				72♀♂		Multiple	10m + 8sm + 32st + 22a		Bueno et al. (2013)
	hermanni	Piquiri River, Nova Laranjeiras, Brazil	24°56’54.0"S	52°35’49.0"W		72♀♂		Multiple	10m + 8sm + 32st + 22a		Bueno et al. (2014)
	hermanni	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		72♀♂	98	Simple	8m + 18sm + 46st/a		Rubert et al. (2016)
	hermanni	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		72♀♂			8m + 18sm + 46st/a		Rubert et al. (2022)
	iheringii (Regan, 1908)	da Lapa stream, Ipeúna, SP, Brazil				80♀♂	132	Multiple	8m + 16sm + 28st + 28a		Traldi et al. (2012)
	iheringii	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		80♀♂	102	Simple	8m + 14sm + 58st/a		Rubert et al. (2016)
	iheringii	Mogi Guaçu River, Pirassununga, SP, Brazil				76♀♂	114	Simple	8m + 30sm + 38st/a		Artoni, Bertollo (1996). Reported as Hypostomus aff. auroguttatus
	iheringii	Mogi Guaçu River, Pirassununga, SP, Brazil				76♀♂	114	Simple	8m + 30sm + 38st/a		Lorscheider et al. (2015)
	iheringii	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		80♀♂			8m + 14sm + 58st/a		Rubert et al. (2022)
	jaguar Zanata, Sardeiro & Zawadzki, 2013					76♀♂		Multiple	10m + 20sm + 46st/a		Anjos et al. (2020)
	macrops (Eigenmann & Eigenmann, 1888)					68♀♂	92		10m + 14sm + 44st/a		Michele et al. (1977)
	mutucae Knaack, 1999	Claro/Mutuca River, Chapada dos Guimarães, MT, Brazil	15°20’13.0"S	55°53’45.0"W	Paraguai	82♀♂	104	Simple	4m + 18sm + 60st/a		Rubert et al. (2016)
	mutucae	Claro River, Brazil	15°20’13.0"S	55°53’45.0"W		82♀♂		Simple	4m + 18sm + 60st/a		Rubert et al. (2022)
	myersi (Gosline, 1947)	Iguaçu River, PR, Brazil	26°15’01.1"S	51°06’10.7"W	Paraná	74♀♂	114	Simple	12m + 16sm + 12st + 34a		Lorscheider et al. (2018)
	nigromaculatus (Schubart, 1964)	Mogi Guaçu River, Cachoeira de Emas, Pirassununga, SP, Brazil				76♀♂	104	Multiple	8m + 20sm + 48st/a		Rubert et al. (2008)
	nigromaculatus	Três Bocas stream, Londrina, PR, Brazil				76♀♂	102	Multiple	6m + 20sm + 50st/a		Rubert et al. (2008)
	nigromaculatus	dos Apertados stream, Londrina, PR, Brazil				76♀♂	102	Multiple	6m + 20sm + 50st/a		Rubert et al. (2008)
	nigromaculatus	Lapa stream, Ipeúna, SP, Brazil				76♀♂	140	Simple	12m + 22sm + 30st + 12a		Traldi et al. (2013)
	nigromaculatus	Hortelã stream, Botucatu, SP, Brazil	22°56’28.9"S	48°35’03.2"W		76♀♂		Simple	8m + 18sm + 20st + 30a		Pansonato-Alves et al. (2013)
	nigromaculatus	Pirapitinga River, Caldas Novas, GO, Brazil	17°43’37.0"S	48°32’54.0"W		74♀♂	104	Simple	8m + 22sm + 44st/a		Rubert et al. (2016)
	paulinus (Ihering, 1905)					74♀♂	104		10m + 20sm + 44st/a		Michele et al. (1977). Reported as Plecostomus ancistroides
	paulinus	Três Bocas stream, PR, Brazil			Paranapanema	76♀♂	98	Simple	6m + 16sm + 54st/a		Rubert et al. (2011)
	paulinus	Apertados stream, PR, Brazil			Paranapanema	76♀♂	98	Simple	6m + 16sm + 54st/a		Rubert et al. (2011)
	paulinus					74♀♂		Simple	10m + 12sm + 20st + 32a		Bueno et al. (2014)
	paulinus	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		76♀♂	98	Simple	6m + 16sm + 54st/a		Rubert et al. (2016)
	paulinus	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	100	Multiple	10m + 18sm + 44st/a		Artoni, Bertollo (1996). Reported as Hypostomus sp. C
	paulinus	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	100	Multiple	10m + 18sm + 44st/a		Lorscheider et al., 2015
	paulinus	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	106	Multiple	14m + 20sm + 38st/a		Artoni, Bertollo (1996). Reported as Hypostomus sp. D2
	paulinus	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	106	Multiple	14m + 20sm + 38st/a		Lorscheider et al. (2015)
	paulinus	Piracicaba River, Piracicaba, SP, Brazil	22°43’07.0"S	47°39’19.0"W		76♀♂			6m + 16sm + 54st/a		Rubert et al. (2022)
	plecostomus (Linnaeus, 1758)					54♀♂					Hinegardner, Rosen (1972)
	plecostomus					54♀♂			24m/sm + 12st + 18a		Muramoto et al. (1968). Reported as Plecostomus paulinus
	prope iheringii	Corumbataí River, SP, Brazil				74♀♂		Multiple	10m + 14sm + 20st + 30a		Alves et al. (2012b)
	prope paulinus	Corumbataí River, SP, Brazil				76♀♂		Simple	6m + 18sm + 12st + 40a		Alves et al. (2012b)
	prope paulinus	Corumbataí River, SP, Brazil				76♀♂		Simple	6m + 18sm + 12st + 40a		Alves et al. (2012b)
	prope plecostomus	Orinoco River, Bolivar, Venezuela				68♀♂		Simple	12m + 16sm + 12st + 28a		Alves et al. (2012b)
	prope unae	Contas River, Brazil	13°51’51.0"S	40°04’54.0"W	Contas	76♀♂	104	Multiple	12m + 16sm + 48st/a		Bitencourt et al. (2011a)
	prope unae	Contas River, Brazil	13°51’51.0"S	40°04’54.0"W	Contas	76♀♂	108	Multiple	12m + 20sm + 44st/a		Bitencourt et al. (2011a)
	prope unae	Oricó River, Brazil	14°08’03.0"S	39°21’30.0"W	Contas	76♀♂	100	Multiple	10m + 14sm + 52st/a		Bitencourt et al. (2011a)
	prope unae	Preto do Criciúma River, Brazil			Contas	76♀♂	106	Multiple	10m + 20sm + 46st/a		Bitencourt et al. (2011a)
	regani (Ihering, 1905)					72♀♂			10m + 20sm + 42st/a		Artoni, Bertollo (1996)
	regani	Araquá River, Botucatu, SP, Brazil				72♀♂		Multiple	12m + 18sm + 26st + 16a		Alves et al. (2006)
	regani	Jacutinga River, Brazil			Paranapanema	72♀♂	100	Multiple	10m + 18sm + 44st/a		Rubert et al. (2011)
	regani	Piumhi River, Piumhi, MG, Brazil	20°20’31.0"S	45°59’03.4"W	São Francisco	72♀♂	116	Simple	8m + 16sm + 20st + 28a		Mendes-Neto et al. (2011)
	regani	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂		Simple	6m + 6sm + 32st + 28a		Martinez et al. (2011)
	regani	Ivaí River, Floresta, PR, Brazil			Ivaí	72♀♂		Multiple	12m + 14sm + 26st + 20a		Endo et al. (2012)
	regani	Piquiri River, Brazil				72♀♂		Multiple	12m + 8sm + 10st + 42a		Bueno et al. (2013)
	regani	Piquiri River, Nova Laranjeiras, Brazil	24°56’54.0"S	52°35’49.0"W		72♀♂		Multiple	12m + 8sm + 10st + 42a		Bueno et al. (2014)
	regani	Mogi Guaçu River, Pirassununga, SP, Brazil	21°55’37.7"S	47°22’02.6"W		72♀♂	100	Simple	10m + 20sm + 42st/a		Rubert et al. (2016)
	regani	Pirapitinga River, Caldas Novas, GO, Brazil	17°43’37.0"S	48°32’54.0"W		72♀♂	106	SImple	12m + 22sm + 38st/a		Rubert et al. (2016)
	regani	Onça stream, Coxim, MS, Brazil	18°32’18.0"S	54°33’43.0"W	Paraguai	72♀♂		Simple	12m + 14sm + 18st + 28a		Ferreira et al. (2019)
	regani	Onça stream, Coxim, MS, Brazil	18°32’18.0"S	54°33’43.0"W	Paraguai	72♀♂		Simple	13m + 14sm + 17st + 18a		Ferreira et al. (2019)
	soniae Hollanda Carvalho & Weber, 2005	Alta Floresta, MT, Brazil	9°54’30.8"S	56°03’33.9"W	Teles Pires	64♀♂	112	Multiple	12m + 22sm + 14st + 16a	XX/XY	Oliveira et al. (2019)
	soniae	Alta Floresta, MT, Brazil	9°53’50.5"S	56°03’39.5"W	Teles Pires	64♀♂	112	Multiple	12m + 22sm + 14st + 16a	XX/XY	Oliveira et al. (2019)
	soniae	Alta Floresta, MT, Brazil	9°53’30.5"S	56°04’18.8"W	Teles Pires	64♀♂	112	Multiple	12m + 22sm + 14st + 16a	XX/XY	Oliveira et al. (2019)
	sp.	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	108	Multiple	10m + 26sm + 36st/a		Artoni, Bertollo (1996). Reported as Hypostomus sp. D1
	sp.	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	108	Multiple	10m + 26sm + 36st/a		Lorscheider et al. (2015)
	sp.	Fundo stream, Barra do Garças, MT, Brazil			Tocantins-Araguaia	64♀♂		Simple	14m + 24sm + 26st/a ♂, 15m + 24sm + 25st/a ♀	ZZ/ZW	Artoni et al. (1998)
	sp.	Mogi Guaçu River, Pirassununga, SP, Brazil			Mogi Guaçu	68♀♂		Multiple	6m + 6sm + 32st + 24a		Martinez et al. (2011)
	sp.	Esparramo stream, Rondonópolis, MT, Brazil	16°30’00.0"S	54°40’00.0"W	Paraguay	74♀♂		Multiple	12m + 20sm + 42st/a		Becker et al. (2014)
	sp. 2 Rio Perdido	Perdido River, Planalto da Bodoquena, MS, Brazil	21°17’09.0"S	56°41’46.0"W		84♀♂	106	Simple	6m + 16sm + 62st/a		Cereali et al. (2008)
	sp. 3 Córrego Salobrinha	Salobra River, Planalto da Bodoquena, MS, Brazil	20°41’34.0"S	56°44’25.0"W		82♀♂	100-102		6m + 12sm + 64st/a + 1-2aBs		Cereali et al. (2008)
	sp. 3 Córrego Salobrinha	Salobrinha stream, Planalto da Bodoquena, MS, Brazil	20°41’07.0"S	56°46’44.0"W		82♀♂	100-102		6m + 12sm + 64st/a + 1-2aBs		Cereali et al. (2008)
	sp. B	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂		Simple	12m + 18sm + 42st/a		Artoni et al. (1999a)
	sp. E	Mogi Guaçu River, Pirassununga, SP, Brazil				80♀♂		Multiple	8m + 16sm + 56st/a		Artoni et al. (1999a)
	sp. F	São Francisco River, Brazil				75, 76♀♂			10m + 16sm + 50st/a or 10m + 17sm + 48st/a		Artoni et al. (1999a)
	sp. Xingu-1	Xingu River, Altamira, PA, Brazil	3°12’48.0"S	52°12’41.7"W	Xingu – Amazon	64♀♂		Simple	32m/sm + 32st/a		Milhomem et al. (2010)
	sp. Xingu-2	Xingu River, Altamira, PA, Brazil	3°12’48.0"S	52°12’41.7"W	Xingu – Amazon	66♀♂		Simple	32m/sm + 34st/a		Milhomem et al. (2010)
	sp. Xingu-3	Xingu River, Altamira, PA, Brazil	3°12’48.0"S	52°12’41.7"W	Xingu – Amazon	65♀♂		Simple	38m/sm + 26st/a + 1B		Milhomem et al. (2010)
	spiniger (Hensel, 1870)	Forquetinha River, Forquetinha, RS, Brazil	29°21’43.5’’S	52° 07’39.6’’W		66♀♂		Multiple	10m + 16sm + 14st + 26a		Takagui et al. (2023)
	spiniger	Quadros Lagoon, RS, Brazil	29°21’43.5’’S	52° 07’39.6’’W		66♀♂		Multiple	10m + 16sm + 14st + 26a		Takagui et al. (2023)
	spiniger	Forquetinha River, Forquetinha, RS, Brazil	29°21’43.5"S	52°07’39.6"W		66♀♂	92	Multiple	10m + 16sm + 40st/a		Rubert et al. (2016). Reported as Hypostomus commersoni, corrected in Takagui et al. (2023)
	strigaticeps (Regan, 1908)					74♀♂	86		8m + 4sm + 62st/a		Michele et al. (1977). Reported as Plecostomus macrops
	strigaticeps				Paranapanema	72♀♂	98	Multiple	10m + 16sm + 46st/a		Rubert et al. (2011)
	strigaticeps	Corumbataí River, SP, Brazil				74♀♂		Multiple	10m + 14sm + 14st + 36a		Alves et al. (2012b)
	strigaticeps	Ivaí River, Floresta, PR, Brazil				72♀♂		Multiple	10m + 14sm + 18st + 30a		Endo et al. (2012)
	strigaticeps	Piquiri River, Brazil				72♀♂		Multiple	12m + 12sm + 18st + 30a		Bueno et al. (2013)
	strigaticeps	Água Boa stream, Mundo Novo, MS, Brazil				72♀♂	122	Multiple	12m + 18sm + 20st + 22a		Fernandes et al. (2012)
	strigaticeps	Hortelã stream, Botucatu, SP, Brazil	22°56’28.9"S	48°35’03.2"W		72♀♂		Multiple	10m + 18sm + 18st + 26a		Pansonato-Alves et al. (2013)
	strigaticeps	Piquiri River, Nova Laranjeiras, Brazil	24°56’54.0"S	52°35’49.0"W		72♀♂		Multiple	12m + 12sm + 18st + 30a		Bueno et al. (2014)
	strigaticeps	Atlântico stream, PR, Brazil	23°18’13.0"S	52°01’51.0"W	Paraná	72♀♂		Multiple	12m + 12sm + 18st + 30a		Baumgärtner et al. (2014)
	strigaticeps	Piquiri River, PR, Brazil	24°56’54.0"S	52°35’49.0"W	Paraná	72♀♂		Multiple	12m + 12sm + 18st + 30a		Baumgärtner et al. (2014)
	strigaticeps	Paraná River, PR, Brazil	24°48’43.0"S	54°19’16.0"W	Paraná	72♀♂		Multiple	12m + 12sm + 18st + 30a		Baumgärtner et al. (2014)
	strigaticeps	Iguassu River, PR, Brazil	25°39’02.0"S	54°27’25.0"W	Paraná	72♀♂		Multiple	12m + 12sm + 18st + 30a		Baumgärtner et al. (2014)
	strigaticeps	Mogi Guaçu River, Pirassununga, SP, Brazil	21°55’37.7"S	47°22’02.6"W		72♀♂	98	Simple	10m + 16sm + 46st/a		Rubert et al. (2016)
	strigaticeps	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	102	Simple	12m + 18sm + 42st/a		Lorscheider et al. (2015)
	strigaticeps	Mogi Guaçu River, Pirassununga, SP, Brazil				72♀♂	102	Simple	12m + 18sm + 42st/a		Artoni, Bertollo (1996). Reported as Hypostomus sp. B
	tapijara Oyakawa, Akama & Zanata, 2005	Ribeira de Iguape River, Registro, SP, Brazil				66♀♂	118	Multiple	14m + 24sm + 14st + 14a		Traldi et al. (2013)
	tietensis (Ihering, 1905)	Piraí River, Brazil	23°22’22.0"S	47°22’13.0"W	Paraná	72♀♂	108	Simple	8m + 8sm + 20st + 36a		Anjos et al. (2020)
	tietensis	Piraí River, Brazil	23°22’22.0"S	47°22’13.0"W	Paraná	72♀♂	109	Simple	8m + 8sm + 20st + 36a		Paula et al. (2022)
	topavae (Godoy, 1969)	Mogi Guaçu River, Pirassununga, SP, Brazil				80♀♂	104	Multiple	8m + 16sm + 56st/a		Artoni, Bertollo (1996). Reported as Hypostomus sp. E
	topavae	Mogi Guaçu River, Pirassununga, SP, Brazil				80♀♂	104	Multiple	8m + 16sm + 56st/a		Lorscheider et al. (2015)
	topavae	Piquiri River, Nova Laranjeiras, PR, Brazil				80♀♂			14m + 10sm + 26st + 30a		Bueno et al. (2012)
	topavae	Piquiri River, PR, Brazil				80♀♂		Simple	14m + 10sm + 26st + 30a		Bueno et al. (2013)
	topavae	Piquiri River, Nova Laranjeiras, PR, Brazil	24°56’54.0"S	52°35’49.0"W	Paraná	80♀♂		Simple	14m + 10sm + 26st + 30a		Bueno et al. (2014)
	topavae	Keller River, Brazil	23°38’25.9"S	51°51’32.8"W		80♀♂	142	Multiple	14m + 30sm + 18st + 18a		Kamei et al. (2017)
	unae (Steindachner, 1878)					76♀♂			10m + 20sm + 46st/a		Anjos et al. (2020)
Lasiancistrus	schomburgkii (Günther, 1864)	Massangana River, Brazil	9°80′48″S*	63°08′90″W*	Amazon	54♀♂	108	Simple	28m + 16sm + 10st		Mariotto et al. (2019)
	sp.	Cachoeira River, Brazil	14°64′66″S*	52°35′50″W*	Tocantins-Araguaia	54♀♂	108	Simple	28m + 16sm + 10st		Mariotto et al. (2019)
Megalancistrus	sp.	Cuiabá River, Brazil	15°62′90″S*	56°08′70″W*	Paraguai	52♀♂	104	Simple	28m + 16sm + 8st		Mariotto et al. (2019)
	parananus (Peters, 1881)	Piquiri River, Nova Laranjeiras, PR, Brazil				52♀♂		Simple	18m + 24sm + 10st		Bueno et al. (2018)
Panaqolus	sp.	Camarapi River, Portel, PA, Brazil				52♀♂		Simple	24m + 18sm + 10st/a		Ayres-Alves et al. (2017)
	tankei Cramer & Sousa, 2016	Xingu River, Brazil				52♀♂	104	Simple	6m + 38sm + 8st		Ferreira et al. (2021)
Panaque	armbrusteri Lujan, Hidalgo & Stewart, 2010	Xingu River, Altamira, PA, Brazil				52♀♂		Simple	26m + 20sm + 6st/a		Ayres-Alves et al. (2017)
	armbrusteri	Gorgulho da Rita, Altamira, PA, Brazil				52♀♂		Simple	26m + 20sm + 6st/a		Ayres-Alves et al. (2017)
	cf. nigrolineatus	Araguaia River, Barra do Garças, MT, Brazil				52♀♂		Simple	26m + 20sm + 6st/a		Artoni, Bertollo (2001)
Peckoltia	cavatica Armbruster & Werneke, 2005	PA, Brazil				52♀♂		Simple	38m/sm + 14st		Pety et al. (2018)
	multispinis (Holly, 1929)	PA, Brazil				52♀♂		Simple	28m/sm + 24st		Pety et al. (2018)
	oligospila (Günther, 1864)	PA, Brazil				52♀♂		Multiple	38m/sm + 14st		Pety et al. (2018)
	sabaji Armbruster, 2003	PA, Brazil				52♀♂		Multiple	38m/sm + 14st		Pety et al. (2018)
	sp. 1 Jari river	Jari River, Monte Dourado, PA, Brazil	03°18’14.9"N	52°03’29.3"W		52♀♂	102	Multiple	44m/sm + 6st + 2a + 1B		Souza et al. (2009)
	sp. 2 Jari river	Jari River, Monte Dourado, PA, Brazil	03°18’14.9"N	52°03’29.3"W		52♀♂	102		32m/sm + 18st + 2a		Souza et al. (2009)
	sp. 3 Jarumã	Abaetuba, PA, Brazil	1°42’41.9"S	48°51’45.9"W		52♀♂		Simple	46m/sm + 6st		Silva et al. (2021)
	sp. 4 Caripetuba	Abaetuba, PA, Brazil	1°37’23.5"S	48°55’33.0"W		52♀♂		Multiple	40m/sm + 12st		Silva et al. (2021)
	vittata (Steindachner, 1881)	Xingu River, Altamira, PA, Brazil	03°12’4"N	52°12’41.7"W		52♀♂	102	Simple	16m + 20sm + 14st + 2a		Souza et al. (2009)
	vittata	PA, Brazil				52♀♂		Simple	32m/sm + 18st + 2a		Pety et al. (2018)
Pseudacanthicus	leopardus (Fowler, 1914)	Xingu River, Brazil			Xingu – Amazon	52♀♂	104	Simple	18m + 34sm		Santos da Silva et al. (2022b)
	sp.	Xingu River, Brazil			Xingu – Amazon	52♀♂	104	Simple	18m + 34sm		Santos da Silva et al. (2022b)
	spinosus (Castelnau, 1855)	Tocantins River, Brazil			Tocantins-Araguaia	52♀♂	104	Simple	18m + 34sm		Santos da Silva et al. (2022b)
Pterygoplichthys	ambrosettii (Holmberg, 1893)	Preto River, Mirassolândia, SP, Brazil				52♀♂		Simple	16m + 24sm + 8st + 4a		Artoni et al. (1999b). Reported as Liposarcus anisitsi
	ambrosettii	Tietê River, Botucatu, SP, Brazil				52♀♂		Simple	28m + 12sm + 8st + 4a		Alves et al. (2006). Reported as Liposarcus anisitsi
	ambrosettii	Miranda River, MS, Brazil				52♀♂		Simple	8m + 14sm + 14st + 16a		Alves et al. (2006). Reported as Liposarcus anisitsi
	ambrosettii	Agua Boa stream, Brazil	23°50’16.7"S	54°20’55.5"W		52♀♂	100	Simple	14m + 26sm + 8st + 4a		Fernandes et al. (2015). Reported as Pterygoplichthys anisitsi
	ambrosettii	Paraná River, Guaíra, PR, Brazil				52♀♂		Simple	16m + 24sm + 8st + 4a		Bueno et al., 2018
	chrysostiktos (Birindelli, Zanata & Lima, 2007)					52♀♂		Simple	22m + 20sm + 10st		Anjos et al. (2020). Reported as Hypostomus chrysostiktos
	gibbiceps (Kner, 1854)	Orinoco River, Venezuela				52♀♂		Simple	20m + 24sm + 8st		Alves et al. (2006). Reported as Glyptoperichthys gibbiceps
	joselimaianus (Weber, 1991)	Quatro Bocas Lake, Araguaiana, MT, Brazil	15°23’20.1"S	51°42’45.9"W		52♀♂		Simple	28m + 16sm + 8st/a		Oliveira et al. (2006)
	multiradiatus (Hancock, 1828)	Orinoco River, Venezuela				52♀♂		Simple	22m + 18sm + 12st		Alves et al. (2006). Reported as Liposarcus multiradiatus
	multiradiatus	Orinoco River, Venezuela				52♀♂		Simple	22m + 18sm + 12st		Alves et al. (2012a). Reported as Liposarcus multiradiatus
	pardalis (Castelnau, 1855)	Porto dos Milagres, Amazonas River, Santarém, PA, Brazil				52♀♂		Simple	18m + 18sm + 8st + 8a		Guimarães et al. (2023)
	pardalis	Arapixuna, Amazonas River, Santarém, PA, Brazil				52♀♂		Simple	18m + 18sm + 8st + 8a		Guimarães et al. (2023)
	pardalis	Grande Lake, Amazonas River, Santarém, PA, Brazil				52♀♂		Simple	18m + 18sm + 8st + 8a		Guimarães et al. (2023)
	pardalis	Pacoval Lake, Amazonas River, Santarém, PA, Brazil				52♀♂		Simple	18m + 18sm + 8st + 8a		Guimarães et al. (2023)
Scobinancistrus	aureatus Burgess, 1994	Xingu River, Brazil			Xingu-Amazon	52♀♂		Simple	22m + 20sm + 10st		Cardoso et al. (2013)
	aureatus	Xingu River, Brazil			Xingu-Amazon	52♀♂		Simple	22m + 20sm + 10st		Ayres-Alves et al. (2017)
	aureatus	Xingu River, Belo Monte hydroeletric plant, Brazil	3°06’12.8"S	51°43’53.9"W		52♀♂			24m + 18sm + 10st		Almeida et al. (2023)
	aureatus	Xingu River, Gorgulho da Rita, Brazil	3°20’06.2"S	52°10’32.9"W		52♀♂			24m + 18sm + 10st		Almeida et al. (2023)
	pariolispos Isbrücker & Nijssen, 1989	Xingu River, Brazil			Xingu-Amazon	52♀♂		Simple	24m + 18sm + 10st		Cardoso et al. (2013)
	pariolispos	Xingu River, Brazil			Xingu-Amazon	52♀♂		Simple	24m + 18sm + 10st		Ayres-Alves et al. (2017)
	pariolispos	Xingu River, Belo Monte hydroeletric plant, Brazil	3°06’12.8"S	51°43’53.9"W		52♀♂			22m + 20sm + 10st		Almeida et al. (2023)
Transancistrus	santarosensis (Tan & Armbruster, 2012)	Río Dos Bocas, Río Palenque, Ecuador				54♀♂	106	Multiple	25m + 27sm + 2a ♀, 26m + 26sm + 2a ♂	ZZ/ZW	Tursellino et al. (2023)
Delturinae
Hemipsilichthys	sp.	Paraitinga River, Brazil	22°52’22.5"S	44°51’00.4"W	Paraíba do Sul	96♀♂	120		16m + 8sm + 72a		Kavalco et al. (2004). Reported as Upsilodus sp.
	sp.	Paraitinga River, Brazil	22°52’22.5"S	44°51’00.4"W	Paraíba do Sul	96♀♂	120	Simple	16m + 8sm + 72a		Kavalco et al. (2005)
	n. sp.	Patos River, Lapa, PR, Brazil				54♀♂			20m + 30sm + 4st		Alves et al. (2005)
Rhinelepinae
Pogonopoma	wertheimeri (Steindachner, 1867)	Mucurí River, Taquarinha, BA, Brazil				54♀♂		Multiple	20m + 30sm + 4st		Artoni, Bertollo (2001)
Rhinelepis	aspera Spix & Agassiz, 1829	Paraná River, Guaíra, PR, Brazil				54♀♂		Simple	20m + 26sm + 8st		Artoni, Bertollo (2001). Reported as Rhinelepis aspera
	aspera	Paraná River, Porto Rico, PR, Brazil				54♀♂		Simple	24m + 18sm + 12st		Endo et al. (2012)
	aspera	Paraná River, Santa Helena, PR, Brazil				54♀♂		Simple	20m + 26sm + 8st		Bueno et al. (2018)

Discussion

The origin of the chromosomal diversity of Loricariidae is attributed to both ecological and molecular factors. Small and isolated populations, in addition to the low vagility, are the main ecological characteristics that have allowed the fixation of chromosomal rearrangements in Loricariidae, as suggested for Farlowella (Marajó et al., 2018), Harttia (Sassi et al., 2023), and Rineloricaria (Rosa et al., 2012). The diploid number 2n = 54 is often considered the ancestral diploid number for the family, since it is observed in most Loricariidae subfamilies and present in other Siluriformes (Artoni, Bertollo, 2001). However, there is no consensus on the matter particularly considering the new karyotypic description of basal taxa within subfamilies. Takagui et al. (2020) in a review of Loricariinae karyotypes argue that although predominant, 2n = 54 should not be considered the basal diploid number for the family because multiple divergences in the microstructure of karyotypes within the same 2n are recurrently seen throughout the family. Notably, the 2n range in Loricarioidei suggest that other numbers rather than 2n = 54 can be considered the plesiomorphic ones, as Astroblepidae presents 2n = 52–54, Scoloplacidae 2n = 50, Callichthyidae 2n = 40–134, and Trichomycteridae 2n = 32–62 (reviewed by Conde-Saldaña et al., 2018). Beyond that, the Diplomystidae (Siluroidei) presents 2n = 56 chromosomes (Campos et al., 1997; Oliveira, Gosztonyi, 2000), which might also suggest that as the ancestral 2n.

We plotted the 2n range per subfamily into the main molecular phylogenetic reconstructions of Loricariidae (Fig. 3) and 2n = 54 is the most widespread, being conserved in Rhinelepinae and present in all other subfamilies. Regardless matter whether 2n = 54 or 2n = 56 is the ancestral diploid number, it is noteworthy that chromosomal evolution in Loricariidae is complex. Considering the lower number of karyotyped species when compared to the valid richness, 234 spp. with cytogenetic data against 1,051 valid species (Fricke et al., 2023), it is difficult to establish the evolutionary pathways that led to the observed variability. Notably, while the stability is restricted to 2n in some genera, with high divergence in karyotype structure, as the 2n = 58 in Farlowella, 2n = 54 in Corumbataia, and 2n = 52 in Pterygoplichthys, stable 2n and karyotype is also observed, as the 2n = 52 (26m+20sm+6st/a) in two species of Panaque. Our compilation reveals that processes of ascending and descending dysploidy (i.e., the increase or decrease of 2n while preserving the genomic content) were frequent in most subfamilies but Rhinelepinae, which may exhibit the constant 2n = 54 as a symplesiomorphic trait.

FIGURE 3| Phylogenetic relationships of Loricariidae with cytogenetic data herein compiled in red. Molecular phylogenies modified from: A. Roxo et al. (2014); B. Lujan et al. (2015); C. Pereira, Reis (2017); D. Roxo et al. (2019).

Despite the 2n conservation in Rhinelepinae, variations in the karyotype structure are observed within and between species. Populations of Rhinelepis aspera Spix & Agassiz, 1829 from the same river differ in the karyotype formula (Artoni, Bertollo, 2001; Endo et al., 2012), suggesting that pericentric inversions played an important role in the chromosomal evolution of the species. Such mechanism is not restricted to Rhinelepinae but also observed in other loricariids, such as Ancistrus (Mariotto et al., 2009), Loricariichthys (Takagui et al., 2014), and Rineloricaria (Rosa et al., 2012; Primo et al., 2018). In addition, the multiple Ag-NOR observed in Pogonopoma wertheimeri (Steindachner, 1867) (Artoni, Bertollo, 2001) when compared to the simple distribution in other Rhinelepinae species suggest that other mechanisms are also important for the chromosomal evolution in the group. Notably, numeric and structural polymorphisms are frequently observed in loricariids, for example in the multiple karyomorphs of Rineloricaria pentamaculata Langeani & de Araujo, 1994 (Glugoski et al., 2023), and the presence of B chromosomes in Harttia longipinna Langeani, Oyakawa & Montoya-Burgos, 2001(Blanco et al., 2012). This chromosomal diversity was probably generated by a combination of rearrangements that include Robertsonian fusions and fissions, paracentric and pericentric inversions, and translocations (Artoni, Bertollo, 2001; Kavalco et al., 2004; Ziemniczak et al., 2012).

Although rare in most fish species, being observed in only about 5% of Teleostei species (Arai, 2011; Sember et al., 2021), our compilation shows that Loricariidae species carry seven out of the nine known sex chromosome systems observed among fishes. Ancistrus was demonstrated to harbor the largest diversity of sex chromosomes with six of the seven recognized systems for the family distributed in 18 species, corresponding to about 23% of the valid species (Neuhaus et al., 2023). The simple XX/XY was the most predominant in the genus, recorded in A. maximus de Oliveira, Zuanon, Zawadzki & Rapp Py-Daniel, 2015 (Oliveira et al., 2010; Favarato et al., 2016), Ancistrus cf. dubius (Mariotto et al., 2011), Ancistrus sp. 1 Quianduba River (Silva et al., 2022, 2023), Ancistrus sp. 1 Maracapucú River (Santos da Silva et al., 2023), Ancistrus sp. 1 Ilha do Capim (Santos da Silva et al., 2023), Ancistrus sp. Catalão (Favarato et al., 2016), Ancistrus sp. L2 (Prizon et al., 2017), Ancistrus sp. L3 (Prizon et al., 2017), and Ancistrus sp. Purus (Oliveira et al., 2010; Favarato et al., 2016). Additionally, two multiple sex chromosome systems that were not observed in any other Loricariidaeare recorded in Ancistrus: ZZ/ZW₁W₂ in A. clementinae Rendahl, 1937 (Nirchio et al., 2023), and Z₁Z₁Z₂Z₂/Z₁Z₂W₁W₂ (Oliveira et al., 2008; Favarato et al., 2016). On other hand, Harttia harbor the highest number of multiple sex chromosome systems when compared to the number of valid species, with six occurrences representing about 25% of valid species (compiled in Sassi et al., 2021): XX/XY₁Y₂ in H. carvalhoi Miranda Ribeiro, 1939, H. intermontana Oliveira & Oyakawa, 2019, and Harttia sp. 1 (Centofante et al., 2006; Deon et al., 2020); X₁X₁X₂X₂/X₁X₂Y in H. duriventris Rapp Py-Daniel & Oliveira, 2001, H. punctata Rapp Py-Daniel & Oliveira, 2001, and H. villasboas Oyakawa, Fichberg & Rapp Py-Daniel, 2018 (Blanco et al., 2014; Sassi et al., 2020), in addition to putative simple XX/XY in H. rondoni Oyakawa, Fichberg & Rapp Py-Daniel, 2018 and H. torrenticola Oyakawa, 1993 (Deon et al., 2020; Sassi et al., 2020). The Loricariidae diversity of sex chromosomes was originated by rearrangements that include translocations (Blanco et al., 2014), centric fissions (Sassi et al., 2023); centric fusions (Centofante et al., 2006), and pericentric inversions (Artoni et al., 1998), including the combination of distinct rearrangements especially in the origin of multiple sex chromosome systems (Oliveira et al., 2008; Deon et al., 2022). Sex chromosomes in Loricariidae seems to have independent origins, but further research is required to explore the genomic content of those sex chromosomes and its origin. Indeed, there is a recognized lack of information regarding the effects of environmental cues and molecular/gene mechanisms in sex determination of Neotropical fishes (Fernandino, Hattori, 2019).

Most Loricariidae species present a single 18S rDNA/Ag-NOR site, which is also considered a plesiomorphic character in the group (Artoni, Bertollo, 2001; Kavalco et al., 2004), and the standard distribution in most vertebrates (Sochorová et al., 2018). Notably, such region in loricariids is involved in several chromosomal rearrangements, including the origin and differentiation of sex chromosomes, being considered evolutionary breakpoint regions in Ancistrus, Harttia, and Rineloricaria (Glugoski et al., 2018; Deon et al., 2022). Size heteromorphism in the Ag-NOR site is also common, probably because of unequal crossing-over between homologs (Takagui et al., 2020). According to our review, some genus conserved the simple Ag-NOR locus in all analyzed species to date (here included only those genera with more than one species karyotyped), namely as Baryancistrus, Corumbataia, Farlowella, Harttia, Hisonotus, Lasiancistrus, Loricaria, Loricariichthys, Neoplecostomus, Panaqolus, Panaque, Pareiorhina, Pseudacanthicus, Pterygoplichthys, and Scobinancistrus. On other hand, the multiple distribution of Ag-NOR seems to be conserved only in Hypancistrus, while other genus as Ancistrus, Hypostomus, and Rineloricaria present both simple and multiple distributions on chromosomes.

Although distributed throughout the Neotropical region, there is a predominance of cytogenetic studies in Loricariidae species from Brazil (Fig. 2). Few studies were conducted in other countries that includes Argentina, Ecuador, Paraguay, and Venezuela. Notably, those in Argentina and Paraguay were mostly restricted to the frontier region with Brazil. When accounting the Brazilian territory, is also notable that some regions are poorly represented in cytogenetic studies, especially the northeast in which at least nine states have not been included in the cytogenetic samplings. In addition, the Guianas Shield and Western Amazon regions are largely recognized as neglected regions in biogeographical and evolutionary studies (Cassemiro et al., 2023), also with little or absent cytogenetic information for Loricariidae. Despite the regional sampling gap problem, Loricariidae diversity is still insufficiently represented by cytogenetic studies. Our compilation recorded 234 species assessed by cytogenetic studies, that in comparison to the 1,051 valid species (Fricke et al., 2023), represents 22.26% of the family species richness. The diversity of genus assessed by cytogenetic studies was the highest in Hypoptopomatinae (42.1%), followed by Rhinelepinae (28.5%), Hypostominae (27.2%), Loricariinae (24.4%), Delturinae (20%), and Lithogeninae (0%). Besides Lithogeninae that do not have any species karyotyped to date, the subfamily Delturinae has only one genus and two unidentified species karyotyped: Hemipsilichthys sp. Paraitinga River (Kavalco et al., 2004, 2005), and Hemipsilichthys n. sp. Patos River (Alves et al., 2005). We suggest that further cytogenetic studies focus on expand the sampling in the northeast Brazil, the Western Amazon, the Guianas Shield, and other Neotropical countries, in addition to evaluate a more representative portion of the diversity.

Acknowledgments

The authors are grateful to all cytogeneticists who contributed to the increase of the knowledge about Loricariidae cytogenetics. We also thank to the Fundação de Amparo à Pesquisa do Estado de São Paulo (FMCS 2020/02681–9, 2022/04261–2, and 2023/08116–0; MBC AR-23/00955–2), to Conselho Nacional de Desenvolvimento Científico e Tecnológico – CNPQ (MBC 302928/2021–9), and Instituto Nacional de Ciência e Tecnologia da Biodiversidade e uso Sustentável de Peixes Neotropicais – INCT Peixes (405706/2022–7).

References

Almeida BRR, Souza LF, Alves TA, Cardoso AL, Oliveira JA, Ribas TFA et al. Chromosomal organization of multigene families and meiotic analysis in species of Loricariidae (Siluriformes) from Brazilian Amazon, with description of a new cytotype for genus Spatuloricaria. Biol Open. 2023; 12(11):bio060029. https://doi.org/10.1242/bio.060029

Alves AL, Borba RS, Pozzobon APB, Oliveira C, Nirchio M, Granado A et al. Localization of 18S ribosomal genes in suckermouth armoured catfishes Loricariidae (Teleostei, Siluriformes) with discussion on the Ag-NOR evolution. Comp Cytogen. 2012a; 6(3):315–21. https://doi.org/10.3897/CompCytogen.v6i3.2667

Alves AL, Borba RS, Oliveira C, Nirchio M, Granado A, Foresti F. Karyotypic diversity and evolutionary trends in the Neotropical catfish genus Hypostomus Lacépède, 1803 (Teleostei, Siluriformes, Loricariidae). Comp Cytogen. 2012b; 6(4):443–52. https://doi.org/10.3897/CompCytogen.v6i4.4028

Alves AL, Oliveira C, Foresti F. Karyotype variability in eight species of the subfamilies Loricariinae and Ancistrinae (Teleostei, Siluriformes, Loricariidae). Caryologia. 2003; 56(1):57–63. https://doi.org/10.1080/00087114.2003.10589308

Alves AL, Oliveira C, Foresti F. Comparative cytogenetic analysis of eleven species of subfamilies Neoplecostominae and Hypostominae (Siluriformes: Loricariidae). Genetica. 2005; 124:127–36. https://doi.org/10.1007/s10709-004-7561-4

Alves AL, Oliveira C, Nirchio M, Granado A, Foresti F. Karyotypic relationships among the tribes of Hypostominae (Siluriformes:Loricariidae) with description of XO sex chromosome system in a Neotropical fish species. Genetica. 2006; 128:1–09. https://doi.org/10.1007/s10709-005-0715-1

Andreata AA, Almeida-Toledo LF, Oliveira C, Toledo Filho SA. Chromosome Studies in Hypoptopomatinae (Pisces, Siluriformes, Loricariidae): I. XX/XY Sex Chromosome Heteromorphism in Pseudotocinclus tietensis. Cytologia. 1992; 57(3):369–72. https://doi.org/10.1508/cytologia.57.369

Andreata AA, Almeida-Toledo LF, Oliveira C, Toledo Filho SA. Chromosome studies in Hypoptopomatinae (Pisces, Siluriformes, Loricariidae) II. ZZ/ZW sex-chromosome system, B chromosomes, and constitutive heterochromatin differentiation in Microlepidogaster leucofrenatus. Cytogenet Genome Res. 1993; 63(4):215–20. https://doi.org/10.1159/000133538

Andreata AA, Almeida-Toledo LF, Oliveira C, Toledo Filho AS. Cytogenetic studies on the subfamily Hypoptopomatinae (Pisces, Siluriformes, Loricariidae). III. Analysis of seven species. Caryologia. 1994; 47(1):27–37. https://doi.org/10.1080/00087114.1994.10797279

Andreata AA, Ferreira DC, Foresti F, Oliveira C. Molecular cytogenetic study of heterochromatin in Hisonotus leucofrenatus (Teleostei, Loricariidae, Hypoptopomatinae). Hereditas. 2010; 147(1):10–17. https://doi.org/10.1111/j.1601-5223.2009.2149.x

Andreata AA, Oliveira C, Foresti F. Karyological characterization of four Neotropical fish species of the genus Hisonotus (Teleostei, Loricariidae, Hypoptopomatinae) from distinct Brazilian river basins. Genet Mol Biol. 2006; 29(1):62–66. https://doi.org/10.1590/S1415-47572006000100013

Anjos MS, Bitencourt JA, Nunes LA, Sarmento-Soares LM, Carvalho DC, Armbruster JW et al. Species delimitation based on integrative approach suggests reallocation of genus in Hypostomini catfish (Siluriformes, Loricariidae). Hydrobiologia. 2020; 847:563–78. https://doi.org/10.1007/s10750-019-04121-z

Arai R. Fish karyotypes a check list. Springer: Tokyo, Japan; 2011.

Arratia G, Kapoor BG, Chardon M, Diogo R. Catfishes Vol. 1 & 2. Enfield, NH: Science Publishers; 2003.

Artoni RF, Bertollo LAC. Cytogenetic studies on Hypostominae (Pisces, Siluriformes, Loricariidae). Considerations on karyotype evolution in the genus Hypostomus. Caryologia. 1996; 49(1):81–90. https://doi.org/10.1080/00087114.1996.10797353

Artoni RF, Bertollo LAC. Nature and distribution of constitutive heterochromatin in fishes, genus Hypostomus (Loricariidae). Genetica. 1999; 106:209–14. https://doi.org/10.1023/A:1003957719178

Artoni RF, Bertollo LAC. Trends in the karyotype evolution of Loricariidae fish (Siluriformes). Hereditas. 2001; 134(3):201–10. https://doi.org/10.1111/j.1601-5223.2001.00201.x

Artoni RF, Venere PC, Bertollo LAC. A heteromorphic ZZ/ZW sex chromosome system in fish, genus Hypostomus (Loricariidae). Cytologia. 1998; 63(4):421–25. https://doi.org/10.1508/cytologia.63.421

Ayres-Alves T, Cardoso AL, Nagamachi CY, Sousa LM, Pieczarka JC, Noronha RCR. Karyotypic evolution and chromosomal organization of repetitive DNA sequences in species of Panaque, Panaqolus, and Scobinancistrus (Siluriformes and Loricariidae) from the Amazon basin. Zebrafish. 2017; 14(3):251–60. https://doi.org/10.1089/zeb.2016.1373

Barros AV, Wolski MAV, Nogaroto V, Almeida MC, Moreira-Filho O, Vicari MR. Fragile sites, dysfunctional telomere and chromosome fusions: what is 5S rDNA role? Gene. 2017; 608:20–27. https://doi.org/10.1016/j.gene.2017.01.013

Barros FJT, Rodrigues ELC, Moura MCS, Torres RA, Paula EA, Sousa LM. Weight-length and length-length relationships of the endangered zebra pleco Hypancistrus zebra (Siluriformes, Loricariidae) from the Xingu River, Amazon, Brazil. J Appl Ichthyol. 2023; 5158180. https://doi.org/10.1155/2023/5158180

Baumgärtner L, Paiz LM, Zawadzki CH, Margarido VP, Castro ALBP. Heterochromatin polymorphism and physical mapping of 5S and 18S ribosomal DNA in four populations of Hypostomus strigaticeps (Regan, 1907) from the Paraná River basin, Brazil: evolutionary and environmental correlation. Zebrafish. 2014; 11(5):479–87. https://doi.org/10.1089/zeb.2014.1028

Becker QMC, Castro RJ, Silva AM, Vizzotto PC. Cytogenetic characterization of two species of Hypostomus (Siluriformes, Loricariidae) from tributaries of the Vermelho river, upper Paraguay river basin. Biodiversidade. 2014; 13(1):1–13.

Benitez MF, Pastori MC, Garrido GG, Takagui F, Giuliano L, Fenocchio AS. First cytogenetic characterization of Loricaria simillima (Loricariidae, Siluriformes) from Paraná River (Argentina) with emphasis in cytotaxonomy of Loricaria. Caryologi. 2017; 70(1):29–33. https://doi.org/10.1080/00087114.2016.1258158

Bitencourt JA, Affonso PRAM, Giuliano-Caetano L, Dias AL. Heterochromatin heterogeneity in Hypostomus prope unae (Steindachner, 1878) (Siluriformes, Loricariidae) from Northeastern Brazil. Comp Cytogen. 2011a; 5(4):329–44. https://doi.org/10.3897/CompCytogen.v5i4.1149

Bitencourt JA, Affonso PRAM, Giuliano-Caetano L, Dias AL. Identification of distinct evolutionary units in allopatric populations of Hypostomus cf. wuchereri Günther, 1864 (Siluriformes: Loricariidae): karyotypic evidence. Neotrop Ichthyol. 2011b; 9(2):317–24. https://doi.org/10.1590/S1679-62252011000200008

Bitencourt JA, Affonso PRAM, Giuliano-Caetano L, Carneiro PLS, Dias AL. Population divergence and peculiar karyoevolutionary trends in the loricariid fish Hypostomus aff. unae from northeastern Brazil. Genet Mol Res. 2012; 11(2):933–43. http://doi.org/10.4238/2012.April.13.1

Blanco DR, Vicari MR, Artoni RF, Traldi JB, Moreira-Filho O. Chromosomal characterization of armored catfish Harttia longipinna (Siluriformes, Loricariidae): first report of B chromosomes in the genus. Zool Sci. 2012; 29(9):604–09. http://doi.org/10.2108/zsj.29.604

Blanco DR, Vicari MR, Lui RL, Bertollo LAC, Traldi JB, Moreira-Filho O. The role of the Robertsonian rearrangements in the origin of the XX/XY₁Y₂ sex chromosome system and in the chromosomal differentiation in Harttia species (Siluriformes, Loricariidae). Rev Fish Biol Fish. 2013; 23(1):127–34. http://doi.org/10.1007/s11160-012-9283-5

Blanco DR, Vicari MR, Lui RL, Artoni RF, Almeida MC, Traldi JB et al. Origin of the X₁X₁X₂X₂/X₁X₂Y sex chromosome system of Harttia punctata (Siluriformes, Loricariidae) inferred from chromosome painting and FISH with ribosomal DNA markers. Genetica. 2014; 142(2):119–26. http://doi.org/10.1007/s10709-014-9759-4

Blanco DR, Vicari MR, Lui RL, Traldi JB, Bueno V, Martinez JF et al. Karyotype diversity and evolutionary trends in armored catfish species of the genus Harttia (Siluriformes: Loricariidae). Zebrafish. 2017; 14(2):169–76. http://doi.org/10.1089/zeb.2016.1377

Brandão KO, Rocha-Reis DA, Garcia C, Pazza R, Almeida-Toledo LF, Kavalco KF. Studies in two allopatric populations of Hypostomus affinis (Steindachner, 1877): the role of mapping the ribosomal genes to understand the chromosome evolution of the group. Comp Cytogen. 2018; 12(1):1–12. http://doi.org/10.3897/CompCytogen.v12i1.22052

Briggs JC. The biogeography of Otophysan fishes (Ostariophysi: Otophysi): a new appraisal. J Biogeography. 2005; 32(2):287–94. https://doi.org/10.1111/j.1365-2699.2004.01170.x

Buck S, Sazima I. An assemblage of mailed catfishes (Loricariidae) in southeastern Brazil: distribution, activity and feeding. Ichthyol Explor Freshw. 1995; 6(4):325–32.

Bueno V, Zawadzki CH, Margarido VP. Trends in chromosome evolution in the genus Hypostomus Lacépède, 1803 (Osteichthyes, Loricariidae): a new perspective about the correlation between diploid number and chromosomes types. Rev Fish Biol Fish. 2012; 22:241–50. http://doi.org/10.1007/s11160-011-9215-9

Bueno V, Venere PC, Zawadzki CH, Margarido VP. Karyotypic diversification in Hypostomus Lacépède, 1803 (Siluriformes, Loricariidae): biogeographical and phylogenetic perspectives. Rev Fish Biol Fish. 2013; 23(1):103–12. http://doi.org/10.1007/s11160-012-9280-8

Bueno V, Venere PC, Konerat JT, Zawadzki CH, Vicari MR, Margarido VP. Physical mapping of the 5S and 18S rDNA in ten species of Hypostomus Lacépède 1803 (Siluriformes: Loricariidae): evolutionary tendencies in the genus. Sci World J. 2014; 943825. http://doi.org/10.1155/2014/943825

Bueno V, Konerat JT, Zawadzki CH, Venere PC, Blanco DR, Margarido VP. Divergent chromosome evolution in Hypostominae tribes (Siluriformes: Loricariidae): correlation of chromosomal data with morphological and molecular phylogenies. Zebrafish. 2018; 15(5):492–503. http://doi.org/10.1089/zeb.2018.1612

Camilo FM, Moreira-Filho O. Karyotypic description of Corumbataia cuestae (Pisces, Loricariidae, Hypoptopomatinae). Cytologia. 2005; 70(1):47–51. https://doi.org/10.1508/cytologia.70.47

Campos H, Arratia G, Cuevas C. Karyotypes of the most primitive catfishes (Teleostei: Siluriformas: Diplomystidae). J Zool System Evol Res. 1997; 35:113–19. https://doi.org/10.1111/j.1439-0469.1997.tb00412.x

Cardoso AL, Carvalho HLS, Benathar TCM, Serrão SMG, Nagamachi CY, Pieczarka JC et al. Integrated cytogenetic and mitochondrial DNA analyses indicate that two different phenotypes of Hypancistrus (L066 and L333) belong to the same species. Zebrafish. 2016; 13(3):209–16. https://doi.org/10.1089/zeb.2015.1213

Cardoso AL, Sales KAH, Nagamachi CY, Pieczarka JC, Noronha RCR. Comparative cytogenetics of two species of genus Scobinancistrus (Siluriformes, Loricariidae, Ancistrini) from the Xingu River, Brazil. Comp Cytogen. 2013; 7(1):43–51. https://doi.org/10.3897/CompCytogen.v7i1.4128

Carvalho ML, Silva GJC, Melo S, et al. The non-monotypic status of the Neotropical fish genus Hemiodontichthys (Siluriformes, Loricariidae) evidenced by genetic approaches. Mitochondrial DNA Part A. 2018; 29(8):1224–30. https://doi.org/10.1080/24701394.2018.1431230

Cassemiro FAS, Albert JS, Antonelli A, Graham CH. Landscape dynamics and diversification of the megadiverse South American freshwater fish fauna. PNAS. 2023, 120(2):e2211974120. https://doi.org/10.1073/pnas.2211974120

Centofante L, Bertollo LAC, Moreira-Filho O. Cytogenetic characterization and description of an XX/XY₁Y₂ sex chromosome system in catfish Harttia carvalhoi (Siluriformes, Loricariidae). Cytogen Genome Res. 2006; 112(3–4):320–24. https://doi.org/10.1159/000089887

Centofante L, Vicari MR, Artoni RF, Buckup PA, Bertollo LAC, Moreira-Filho O. Chromosome analyses in species of Pareiorhina (Siluriformes, Loricariidae) and cytotaxonomic considerations on the group. Nucleus. 2011; 54(2):65–70. https://doi.org/10.1007/s13237-011-0035-z

Cereali SS, Pomini E, Rosa R, Zawadzki CH, Froehlich O, Giuliano-Caetano L. Karyotype description of two species of Hypostomus (Siluriformes, Loricariidae) of the Planalto da Bodoquena, Brazil. Genet Mol Res. 2008; 7(3):583–91. https://doi.org/10.4238/vol7-3gmr404

Cius A, Lorscheider CA, Barbosa LM, Zawadzki CH, Borin-Carvalho LA, Porto FE et al. Contributions of species Rineloricaria pentamaculata (Loricariidae:Loricariinae) in a karyoevolutionary context. Caryologia. 2022; 75(3):39–46. https://doi.org/10.36253/caryologia-1611

Cristina FD, Chiachio MC, Takako AK, Andreata AA, Foresti F, Oliveira C. Comparative cytogenetics of nine species of Hypoptopomatinae (Teleostei: Siluriformes: Loricariidae): the importance of structural rearrangements in chromosome evolution. Caryologia. 2005; 58(4):387–95. https://doi.org/10.1080/00087114.2005.10589478

Deon GA, Glugoski L, Vicari MR, Nogaroto V, Sassi FMC, Cioffi MB et al. Highly rearranged karyotypes and multiple sex chromosome systems in armored catfishes from the genus Harttia (Teleostei, Siluriformes). Genes. 2020; 11(11):1366. https://doi.org/10.3390/genes11111366

Deon GA, Glugoski L, Sassi FMC, Hatanaka T, Nogaroto V, Bertollo LAC et al. Chromosomal rearrangements and origin of the multiple XX/XY₁Y₂ sex chromosome system in Harttia Species (Siluriformes: Loricariidae). Front Genet. 2022a; 13:877522. https://doi.org/10.3389/fgene.2022.877522

Deon GA, Glugoski L, Hatanaka T, Sassi FMC, Nogaroto V, Bertollo LAC et al. Evolutionary breakpoint regions and chromosomal remodeling in Harttia (Siluriformes: Loricariidae) species diversification. Genet Mol Biol. 2022b; 45(2):e20210170. https://doi.org/10.1590/1678-4685-GMB-2021-0170

Endo KS, Martinez ERM, Zawadzki CH, Paiva LRS, Júlio Junior HF. Karyotype description of possible new species of the Hypostomus ancistroides complex (Teleostei: Loricariidae) and other Hypostominae. Acta Scient Biol Sci. 2012; 34(2):181–89. https://doi.org/10.4025/actascibiolsci.v34i2.9318

Evers H, Pinnegar JK, Taylor MI. Where are they all from? – sources and sustainability in the ornamental freshwater fish trade. J Fish Biol. 2019; 94(6):909–16. https://doi.org/10.1111/jfb.13930

Favarato RM, Silva M, Oliveira RR, Artoni RF, Feldberg E, Matoso DA. Cytogenetic diversity and the evolutionary dynamics of rDNA genes and telomeric sequences in the Ancistrus genus (Loricariidae: Ancistrini). Zebrafish. 2016; 13(2):103–11. https://doi.org/10.1089/zeb.2015.1140

Fenocchio AS, Pastori MC, Roncati HÁ, Moreira-Filho O, Bertollo LAC. A cytogenetic survey of the fish fauna from Argentina. Caryologia. 2003; 56(2):197–204. https://doi.org/10.1080/00087114.2003.10589325

Fernandes CA, Alves DS, Guterres ZR, Martins-Santos IC. Cytogenetic analysis of two locariid species (Teleostei, Siluriformes) from Iguatemi River (Paraná River drainage) in Brazil. Comp Cytogenet. 2015; 9(1):67–78. https://doi.org/10.3897/CompCytogen.v9i1.8804

Fernandes CA, Bailly D, Sant’Ana DRA, Alves DS. First cytogenetic record for a species of Otothyropsis Ribeiro, Carvalho & Melo, 2005 (Loricariidae, Hypoptopomatinae). Neotrop Ichthyol. 2016, 14(1):e150123. https://doi.org/10.1590/1982-0224-20150123

Fernandes CA, Damásio JF, Martins-Santos IC. Cytogenetic studies in species of family Loricariidae (Osteichthyes, Siluriformes) from Iguatemi river basin, Brazil. First cytogenetic report in Farlowella amazonum (Günther, 1864). Caryologia. 2012; 65(4):276–80. https://doi.org/10.1080/00087114.2012.752913

Fernandes CA, Paiz LM, Piscor D, Gavazzoni M, Carvalho LAB, Portela-Castro ALB et al. Chromosomal diversity in two allopatric populations of Farlowella hahni Meinken 1937 (Teleostei: Siluriformes): cytogenetics and cytochrome b analyses. Zebrafish. 2021; 18(1):66–72. https://doi.org/10.1089/zeb.2020.1966

Fernandino JI, Hattori RS. Sex determination in Neotropical fish: Implications ranging from aquaculture technology to ecological assessment. Gen Comp Endocrinol. 2019; 273:172–83. https://doi.org/10.1016/j.ygcen.2018.07.002

Ferreira AMV, Viana PF, Zuanon J,Ezaz T, Cioffi MB, Takagui FH et al. Cytogenetic analysis of Panaqolus tankei Cramer & Sousa, 2016 (Siluriformes, Loricariidae), an ornamental fish endemic to Xingu River, Brazil. Cytogenet Genome Res. 2021; 161(3–4):187–94. https://doi.org/10.1159/000514061

Ferreira GEB, Barbosa LM, Prizon-Nakajima AC, Paiva S, Vieira MMR, Gallo RB et al. Constitutive heterochromatin heteromorphism in the Neotropical armored catfish Hypostomus regani (Ihering, 1905) (Loricariidae, Hypostominae) from the Paraguay River basin (Mato Grosso do Sul, Brazil). Comp Cytogenet. 2019; 13(1):27–39. https://doi.org/10.3897/CompCytogen.v13i1.30134

Ferreira RO, Pereira AL, Nagamachi CY, Pieczarka JC, Sousa LM, Noronha RCR. Caracterização citogenética de uma espécie de Spatuloricaria (Siluriformes, Loricariidae) do rio Xingu, (Pará, Amazônia, Brasil). Biota Amazôn. 2014; 4(1):30–36. http://dx.doi.org/10.18561/2179-5746/biotaamazonia.v4n1p30-36

Fink S, Fink W. Interrelationships of the ostariophysan fishes (Teleostei). ZJLS. 1981; 72:297–353.

Fink S, Fink W. Interrelationships of ostariophysan fishes (Teleostei). In: Stiassny MLJ, Parenti LR, Johnson GD, editors. Interrelationships of fishes. New York: Academic Press; 1996. p.209–45.

Fricke R, Eschmeyer WN, Van der Laan R. Eschmeyer’s catalog of fishes: genera, species, references [Internet]. San Francisco: California Academy of Science; 2023. Available from: https://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp

Glugoski L, Giuliano-Caetano L, Moreira-Filho O, Vicari MR, Nogaroto V. Co-located hAT transposable element and 5S rDNA in an interstitial telomeric sequence suggest the formation of Robertsonian fusion in armored catfish. Gene. 2018; 650:49–54. https://doi.org/10.1016/j.gene.2018.01.099

Glugoski L, Deon G, Schott S, Vicari MR, Nogaroto V, Moreira-Filho O. Comparative cytogenetic analyses in Ancistrus species (Siluriformes: Loricariidae). Neotrop Ichthyol. 2020; 18(2):e200013. https://doi.org/10.1590/1982-0224-2020-0013

Glugoski L, Nogaroto V, Deon GA, Azambuja M, Moreira-Filho O, Vicari MR. Enriched tandem repeats in chromosomal fusion points of Rineloricaria latirostris (Boulenger, 1900) (Siluriformes: Loricariidae). Genome. 2022; 65(9):479–89. https://dx.doi.org/10.1139/gen-2022-0043

Glugoski L, Deon GA, Nogaroto V, Moreira-Filho O, Vicari MR. Robertsonian fusion site in Rineloricaria pentamaculata (Siluriformes: Loricariidae): involvement of 5S rDNA and satellite sequences. Cytogenet Genome Res. 2023; 162(11–12):657–64. https://doi.org/10.1159/000530636

Guimarães AS, Maciel LAM, Souza MFB, Rodrigues LRR. Karyotypic and molecular analysis of Pterygoplichthys pardalis (Castelnau 1855) from the lower Amazon River. Animals. 2023; 13(9):1533. https://doi.org/10.3390/ani13091533

Hinegardner R, Rosen DE. Cellular DNA content and the evolution of teleostean fishes. The American Naturalist. 1972; 106(951):621–44. https://doi.org/10.1086/282801

Kamei MCSL, Baumgärtner L, Paiva S, Zawadzki CH, Martins-Santos IC, Portela-Castro ALB. Chromosomal diversity of three species of Hypostomus Lacépède, 1803 (Siluriformes, Loricariidae), from the Paraná River basin, Brazil: a species complex in Hypostomus ancistroides Reinforced by a ZZ/ZW sex chromosome system. Zebrafish. 2017; 14(4):357–63. https://doi.org/10.1089/zeb.2017.1429

Kavalco KF, Pazza R, Bertollo LAC, Moreira-Filho O. Heterochromatin characterization of four fish species of the family Loricariidae (Siluriformes). Hereditas. 2004; 141(3):237–42. https://doi.org/10.1111/j.1601-5223.2004.01850.x

Kavalco KF, Pazza R, Bertollo LAC, Moreira-Filho O. Karyotypic diversity and evolution of Loricariidae (Pisces, Siluriformes). Heredity. 2005; 94(2):180–86. https://doi.org/10.1038/sj.hdy.6800595

Konerat JT, Bueno V, Margarido VP, Portela-Castro ALB, Martins-Santos IC. Diversity of sex chromosome systems in Ancistrini (Loricariidae, Hypostominae): ZZ/ZW in Ancistrus taunayi Miranda Ribeiro, 1918. Cytogenet Genome Res. 2015; 146(4):306–10. https://doi.org/10.1159/000441431

Levan A, Fredga K, Sandberg AA. Nomenclature for centromeric position on chromosomes. Hereditas. 1964; 52:201–20.

Lévêque C, Oberdorff T, Paugy D, Stassny MLJ, Tedesco PA. Global diversity of fish (Pisces) in freshwater. Hydrobiologia. 2008; 595:545–67. https://doi.org/10.1007/s10750-007-9034-0

Lorscheider CA, Oliveira JIN, Dulz TA,Nogaroto V, Martins-Santos IC, Vicari MR. Comparative cytogenetics among three sympatric Hypostomus species (Siluriformes: Loricariidae): an evolutionary analysis in a high endemic region. Braz Arch Biol Technol. 2018; 61:e18180417. http://dx.doi.org/10.1590/1678-4324-2018180417

Lorscheider CA, Zawadzki CH, Vicari MR, Martins-Santos IC, Artoni RF. Karyotypic diversity of the armoured catfish genus Hypostomus (Siluriformes: Loricariidae) in the context of its occurrence and distribution. J Fish Biol. 2015; 87(4):1099–110. https://doi.org/10.1111/jfb.12762

Lujan NK, Armbruster JW, Lovejoy NR, López-Fernández H. Multilocus molecular phylogeny of the suckermouth armored catfishes (Siluriformes: Loricariidae) with a focus on subfamily Hypostominae. Mol Phylogenet Evol. 2015; 82:269–88. https://doi.org/10.1016/j.ympev.2014.08.020

Lujan NK, Cramer CA, Covain R, Fisch-Muller S, López-Fernández H. Multilocus molecular phylogeny of the ornamental wood-eating catfishes (Siluriformes, Loricariidae, Panaqolus and Panaque) reveals undescribed diversity and parapatric clades. Mol Phylogenet Evol. 2017; 109:321–36. https://doi.org/10.1016/j.ympev.2016.12.040

Lujan NK, Hidalgo M, Stewart DJ. Revision of Panaque (Panaque), with descriptions of three new species from the Amazon Basin (Siluriformes, Loricariidae). Copeia. 2010; 4(2010):676–704. https://doi.org/10.1643/CI-09-185

Maia TPA, Giuliano-Caetano L, Rodriguez MS, Rubert M, Takagui FH, Dias AL. Chromosomal banding in three species of the genus Rineloricaria (Siluriformes, Loricariidae, Loricariinae). Ichthyol Res. 2010; 57(2):209–13. https://doi.org/10.1007/s10228-009-0145-7

Marajó L, Viana PF, Ferreira M, Rapp Py-Daniel LH, Feldberg E. Cytogenetics of two Farlowella species (Loricariidae: Loricariinae): implications on the taxonomic status of the species. Netrop Ichthyol. 2018; 16(4):e180029. https://doi.org/10.1590/1982-0224-20180029

Marajó L, Viana PF, Ferreira AMV, Rapp Py-Daniel LH, Cioffi MB, Sember A et al. Chromosomal rearrangements and the first indication of an ♀X₁X₁X₂X₂/♂X₁X₂Y sex chromosome system in Rineloricaria fishes (Teleostei: Siluriformes). J Fish Biol. 2022; 102(2):443–54. https://doi.org/10.1111/jfb.15275

Mariotto S, Artoni RF, Miyazawa CS. Occurrence of sexual chromosome, of the type ZZ/ZW, in Ancistrus cf. dubius (Loricariidae, Ancistrinae) of the Paraguay River Basin, Mato Grosso, Brazil. Caryologia. 2004; 57(4):327–31. https://doi.org/10.1080/00087114.2004.10589413

Mariotto S, Centofante L, Miyazawa CS, Bertollo LAC, Moreira-Filho O. Chromosome polymorphism in Ancistrus cuiabae Knaack, 1999 (Siluriformes: Loricariidae: Ancistrini). Netrop Ichthyol. 2009; 7(4):595–600. https://doi.org/10.1590/S1679-62252009000400006

Mariotto S, Centofante L, Moreira-Filho O. Diversity and chromosomal evolution in the genus Ancistrus Kner, 1854 (Loricariidae: Ancistrini) from three hydrographic basins of Mato Grosso State, Brazil. Netrop Ichthyol. 2013; 11(1):125–31. https://doi.org/10.1590/S1679-62252013000100015

Mariotto S, Centofante L, Venere PC, Ferreira DC, Artoni RF. New comparative cytogenetic data on three genera of armored catfishes of Ancistrini (Loricariidae: Hypostominae). Cytogenet Genome Res. 2019; 159(4):208–14. https://doi.org/10.1159/000504723

Mariotto S, Centofante L, Vicari MR, Artoni RF, Moreira-Filho O. Chromosomal diversification in ribosomal DNA sites in Ancistrus Kner, 1854 (Loricariidae, Ancistrini) from three hydrographic basins of Mato Grosso, Brazil. Comp Cytogenet. 2011; 5(4):289–300. https://doi.org/10.3897/CompCytogen.v5i4.1757

Mariotto S, Miyazawa CS. Ancistrus cf. dubius (Siluriformes, Ancistrinae), a complex of species. 1. Chromosomic characterization of four populations and occurence of sexual chromosomes of type XX/XY, in the Pantanal basin of Mato Grosso, Brazil. Caryologia. 2006, 59(4):299–304. https://doi.org/10.1080/00087114.2006.10797929

Martinez ERM, Zawadzki CH, Foresti F, Oliveira C. Cytogenetic analysis of five Hypostomus species (Siluriformes, Loricariidae). Genet Mol Biol. 2011; 34(4):562–68. https://doi.org/10.1590/S1415-47572011005000038

Maurutto FAM, Manvailer LFS, Sczepanski TS, Cestari MM, Artoni RF. Cytogenetic characterization of three allopatric species of Hypostomus Lacépède (1803) (Teleostei, Loricariidae). Caryologia. 2012; 65(4):340–46. https://doi.org/10.1080/00087114.2012.760882

Medeiros LA, Ginani EG, Sousa LM, Rapp Py-Daniel LH, Feldberg E. Cytogenetic analysis of Baryancistrus xanthellus (Siluriformes: Loricariidae: Ancistrini), an ornamental fish endemic to the Xingu River, Brazil. Neotrop Ichthyol. 2016; 14(2):e150108. https://doi.org/10.1590/1982-0224-20150108

Mendes-Neto EO, Vicari MR, Artoni RF, Moreira-Filho O. Description of karyotype in Hypostomus regani (Ihering, 1905) (Teleostei, Loricariidae) from the Piumhi river in Brazil with comments on karyotype variation found in Hypostomus. Comp Cytogenet. 2011; 5(2):133–42. https://doi.org/10.3897/CompCytogen.v5i2.964

Michele JL, Takahashi CS, Ferrari I. Karyotypic study of some species of the family Loricariidae. Cytologia. 1977; 42:539–46. https://doi.org/10.1508/cytologia.42.539

Milhomem SSR, Castro RR, Nagamachi CY, Souza A, Feldberg E, Pieczarka J. Different cytotypes in fishes of the genus Hypostomus Lcépède, 1803, (Siluriformes: Loricariidae) from Xingu river (Amazon region, Brazil). Comp Cytogenet. 2010; 4(1):45–54. https://doi.org/10.3897/compcytogen.v4i1.31

Muramoto J, Ohno S, Atkin NB. On the diploid state of the fish Order Ostariophysi. Chromosoma. 1968; 24:59–66. https://doi.org/10.1007/BF00329607

Nelson JS, Grande TC, Wilson MV. Fishes of the world. Hoboken, New Jersey: John Wiley & Sons; 2016.

Nirchio M, Oliveira C, Cioffi MB, Sassi FMC, Valdiviezo J, Paim FG et al. Occurrence of sex chromosomes in fish of the genus Ancistrus with a new description of multiple sex chromosomes in the Ecuadorian endemic Ancistrus clementinae (Loricariidae). Genes. 2023; 14:306. https://doi.org/10.3390/genes14020306

Novák J, Hofmann J, Hohl D, Magalhães ALB, Patoka J. Enigmatic armoured catfishes (Siluriformes: Callichthyidae and Loricariidae) in ornamental aquaculture: a new insight into Neotropical fish diversity. Aquaculture. 2022; 547:737460. https://doi.org/10.1016/j.aquaculture.2021.737460

Novák J, Kalous L, Patoka J. Modern ornamental aquaculture in Europe: early history of freshwater fish imports. Rev Aquaculture. 2020; 12(4):2042–60. https://doi.org/10.1111/raq.12421

Oliveira C, Gosztonyi AE. A cytogenetic study of Diplotnystes mesembrinus (Teleostei, Siluriformes, Diplomystidae) with a discussion of chromosome evolution in siluriforms. Caryologia. 2000; 53(1):31–37. https://doi.org/10.1080/00087114.2000.10589178

Oliveira LC, Ribeiro MO, Costa GM, Zawadzki CH, Prizon-Nakajima AC, Borin-Carvalho LA et al. Cytogenetic characterization of Hypostomus soniae Hollanda-Carvalho & Weber, 2004 from the Teles Pires River, southern Amazon basin: evidence of an early stage of an XX/XY sex chromosome system. Comp Cytogenet. 2019; 13(4):411–22. https://doi.org/10.3897/CompCytogen.v13i4.36205

Oliveira LC, Ribeiro MO, Dutra ES, Zawadzki CH, Portela-Castro ALB, Martins-Santos IC. Karyotype structure of Hypostomus cf. plecostomus (Linnaeus, 1758) from Tapajós River basin, Southern Amazon: occurrence of sex chromosomes (ZZ/ZW) and their evolutionary implications. Genet Mol Res. 2015; 14(2):6625–34. http://dx.doi.org/10.4238/2015.June.18.5

Oliveira RR, Feldberg E, Anjos MB, Zuanon J. Mechanisms of chromosomal evolution and its possible relation to natural history characteristics in Ancistrus catfishes (Siluriformes: Loricariidae). J Fish Biol. 2009; 75(9):2209–25. https://doi.org/10.1111/j.1095-8649.2009.02450.x

Oliveira RR, Feldberg E, Anjos MB, Zuanon J. Karyotype characterization and ZZ/ZW sex chromosome heteromorphism in two species of the catfish genus Ancistrus Kner, 1854 (Siluriformes: Loricariidae) from the Amazon basin. Neotrop Ichthyol. 2007; 5(3):301–06. https://doi.org/10.1590/S1679-62252007000300010

Oliveira RR, Feldberg E, Anjos MB, Zuanon J. Occurrence of multiple sexual chromosomes (XX/XY₁Y₂and Z₁Z₁Z₂Z₂/Z₁Z₂W₁W₂) in catfishes of the genus Ancistrus (Siluriformes: Loricariidae) from the Amazon basin. Genetica. 2008; 134(2):243–49. https://doi.org/10.1007/s10709-007-9231-9

Oliveira RR, Souza IL, Venere PC. Karyotype description of three species of Loricariidae (Siluriformes) and occurrence of the ZZ/ZW sexual system in Hemiancistrus spilomma Cardoso & Lucinda, 2003. Neotrop Ichthyol. 2006; 4(1):93–97. https://doi.org/10.1590/S1679-62252006000100010

Pansonato-Alves JC, Serrano EA, Utsunomia R, Scacchetti P, Oliveira C, Foresti F. Mapping five repetitive DNA classes in sympatric species of Hypostomus (Teleostei: Siluriformes: Loricariidae): analysis of chromosomal variability. Rev Fish Biol Fish. 2013; 23:477–89. https://doi.org/10.1007/s11160-013-9303-0

Paula GBN, Gavazzoni M, Zawadzki CH, Fernandes CA, Portela_castro ALB, Lui RL et al. Identification of cryptic species in allopatric populations of Hypostomus tietensis (Siluriformes: Loricariidae) through cytogenetics analyses. Neotrop Ichthyol. 2022; 20(2):e210158. https://doi.org/10.1590/1982-0224-2021-0158

Pazian MF, Pereira LHG, Shimabukuru-Dias CK, Oliveira C, Foresti F. Cytogenetic and molecular markers reveal the complexity of the genus Piabina Reinhardt, 1867 (Characiformes: Characidae). Neotrop Ichthyol. 2012; 10(2):329–40. https://doi.org/10.1590/S1679-62252012005000015

Pereira EH, Reis RE. Morphology-based phylogeny of the suckermouth armored catfishes, with emphasis on the Neoplecostominae (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2017; 4264(1):1–104. https://doi.org/10.11646/zootaxa.4264.1.1

Pety AM, Cardoso AL, Nagamachi CY, Pieczarka JC, Sousa LM, Rodrigues RC. In situ localization of ribosomal sites in Peckoltia and Ancistomus (Loricariidae: Hypostominae) from the Amazon basin. Zebrafish. 2018; 15(3):263–69. https://doi.org/10.1089/zeb.2017.1523

Porto FE, Portela-Castro ALB, Martins-Santos IC. Possible origins of B chromosomes in Rineloricaria pentamaculata (Loricariidae, Siluriformes) from the Paraná River basin. Genet Mol Res. 2010; 9(3):1654–59. https://doi.org/10.4238/vol9-3gmr859

Porto FE, Portela-Castro ALB, Martins-Santos IC. Chromosome polymorphism in Rineloricaria pentamaculata (Loricariidae, Siluriformes) of the Paraná River basin. Ichthyol Res. 2011; 58:225–31. https://doi.org/10.1007/s10228-011-0215-5

Porto FE, Vieira MMR, Barbosa LM, Borin-Carvalho LA, Vicari MR, Portela-Castro ALB et al. Chromosomal polymorphism in Rineloricaria lanceolata Günther, 1868 (Loricariidae: Loricariinae) of the Paraguay basin (Mato Grosso do Sul, Brazil): evidence of fusions and their consequences in the population. Zebrafish. 2014a; 11(4):318–24. https://doi.org/10.1089/zeb.2014.0996

Porto FE, Gindri BS, Vieira MMR, Borin LA, Portela-Castro ALB,Martins-Santos IC. Polymorphisms of the nucleolus organizing regions in Loricaria cataphracta (Siluriformes, Loricariidae) of the upper Paraguay River basin indicate an association with transposable elements. Genet Mol Res. 2014b; 13(1):1627–34. http://dx.doi.org/10.4238/2014.March.12.15

Price AC, Weadick CJ, Shim J, Rodd FH. Pigments, patterns, and fish behavior. Zebrafish. 2008; 5(4):297–307. https://doi.org/10.1089/zeb.2008.0551

Primo CC, Glugoski L, Almeida MC, Zawadzki CH, Moreira-Filho O, Vicari MR et al. Mechanisms of chromosomal diversification in species of Rineloricaria (Actinopterygii: Siluriformes: Loricariidae). Zebrafish. 2017; 14(2):161–68. https://doi.org/10.1089/zeb.2016.1386

Primo CC, Glugoski L, Vicari MR, Nogaroto V. Chromosome mapping and molecular characterization of the Tc1/Mariner element in Rineloricaria (Siluriformes: Loricariidae). Braz Arch Biol Technol. 2018; 61:e18170623. http://dx.doi.org/10.1590/1678-4324-2018170623

Prizon AC, Borin-Carvalho LA, Bruschi DP, Ribeiro MO, Barbosa LM, Ferreira GEB et al. Cytogenetic data on Ancistrus sp. (Siluriformes, Loricariidae) of the Paraguay River basin (MS) sheds light on intrageneric karyotype diversification. Comp Cytogenet. 2016; 10(4):625–36. https://doi.org/10.3897/CompCytogen.v10i4.8532

Prizon AC, Bruschi DP, Borin-Carvalho LA, Cius A, Barbosa LM, Ruiz HB et al. Hidden diversity in the populations of the armored catfish Ancistrus Kner, 1854 (Loricariidae, Hypostominae) from the Paraná River basin revealed by molecular and cytogenetic data. Front Genet. 2017; 8:185. https://doi.org/10.3389/fgene.2017.00185

Ramirez JL, Birindelli JL, Carvalho DC, Affonso PRAM, Venere PC, Ortega H et al. Revealing hidden diversity of the underestimated Neotropical ichthyofauna: DNA barcoding in the recently described genus Megaleporinus (Characiformes:Anostomidae). Front Genet. 2017; 8:149. https://doi.org/10.3389/fgene.2017.00149

Ribeiro AC, Lima FCT, Pereira EHL. A new genus and species of a minute suckermouth armored catfish (Siluriformes: Loricariidae) from the rio Tocantins drainage, Central Brazil: the smallest known loricariid catfish. Copeia. 2012; 2012(4):637–47. https://doi.org/10.1643/CI-11-137

Ribeiro MO, Noleto RB, Lorscheider CA, Porto FE, Prizon AC, Zawadzki CH et al. Cytogenetic description of Ancistrus abilhoai (Siluriformes: Loricariidae) from Iguaçu River basin, southern Brazil. Genet Mol Res. 2015; 14(2):4051–57. http://dx.doi.org/10.4238/2015.April.27.20

Rocha-Reis D, Brandão KO, Almeida-Toledo LF, Pazza R, Kavalco KF. The persevering cytotaxonomy: discovery of a unique XX/XY sex chromosome system in catfishes suggests the existence of a new, endemic and rare species. Cytogenet Gen Res. 2018; 156(1):45–55. https://doi.org/10.1159/000492959

Rosa KO, Ziemniczak K, Barros AV, Nogaroto V, Almeida M, Cestari MM et al. Numeric and structural chromosome polymorphism in Rineloricaria lima (Siluriformes: Loricariidae): fusion points carrying 5S rDNA or telomere sequence vestiges. Rev Fish Biol Fish. 2012; 22(3):739–49. https://doi.org/10.1007/s11160-011-9250-6

Roxo FF, Albert JS, Silva GS, Zawadzki CH, Foresti F, Oliveira C. Molecular phylogeny and biogeographic history of the armored Neotropical catfish subfamilies Hypoptopomatinae, Neoplecostominae and Otothyrinae (Siluriformes: Loricariidae). PLoS ONE. 2014; 9(8):e105564. https://doi.org/10.1371/journal.pone.0105564

Roxo FF, Ochoa LE, Sabaj MH, Lujan NK, Covain R, Silva GSC et al. Phylogenomic reappraisal of the Neotropical catfish family Loricariidae (Teleostei: Siluriformes) using ultraconserved elements. Mol Phylogenet Evol. 2019; 135:148–65. https://doi.org/10.1016/j.ympev.2019.02.017

Rubert M, Rosa R, Jerep FC, Bertollo LAC, Giuliano-Caetano L. Cytogenetic characterization of four species of the genus Hypostomus Lacépède, 1803 (Siluriformes, Loricariidae) with comments on its chromosomal diversity. Comp Cytogenet. 2011; 5(5):397–410. https://doi.org/10.3897/CompCytogen.v5i5.1589

Rubert M, Rosa R, Zawadzki CH, Mariotto S, Moreira-Filho O, Giuliano-Caetano L. Chromosome mapping of 18S ribosomal RNA genes in eleven Hypostomus species (Siluriformes, Loricariidae): diversity analysis of the sites. Zebrafish. 2016; 13(4):360–68. https://doi.org/10.1089/zeb.2016.1279

Rubert M, Takagui FH, Santos KF, Pompeo LRS, Rosa R, Zawadzki CH et al. Topotype-based chromosomal diversity among five species of freshwater armored catfishes in the Hypostomus auroguttatus supergroup (Actinopterygii: Siluriformes). Zool Sci. 2022; 39(5):446–52. https://doi.org/10.2108/zs210103

Rubert M, Zawadzki CH, Giuliano-Caetano L. Cytogenetic characterization of Hypostomus nigromaculatus (Siluriformes, Loricariidae). Neotrop Ichthyol. 2008, 6(1):93–100. https://doi.org/10.1590/S1679-62252008000100011

Saitoh K, Miya M, Inoue JG, Ishiguro NB, Nishida M. Mitochondrial genomics of ostariophysan fishes: perspectives on phylogeny and biogeography. J Mol Evol. 2003; 56(4):464–72. https://doi.org/10.1007/s00239-002-2417-y

Santos CEV, Almeida BRR, Tavares FS, Frade LSF, Cardoso AL, Sá ALA et al. Chromosomal mapping of the histone multigene family and U2 snRNA in Hypancistrus species (Siluriformes, Loricariidae) from the Brazilian Amazon. Zebrafish. 2023; 20(1):28–36. https://doi.org/10.1089/zeb.2022.0030

Santos da Silva K, Souza ACP, Pety AM, Noronha RCR, Vicari MR, Pieczarka JC et al. Comparative cytogenetics analysis among Peckoltia species (Siluriformes, Loricariidae): insights on karyotype evolution and biogeography in the Amazon region. Front Genet. 2021; 12:779464. https://doi.org/10.3389/fgene.2021.779464

Santos da Silva K, Glugoski L, Vicari MR, Souza ACP, Noronha RCR, Pieczarka JC et al. Chromosomal diversification in Ancistrus species (Siluriformes: Loricariidae) inferred from repetitive sequence analysis. Front Genet. 2022a; 13:838462. https://doi.org/10.3389/fgene.2022.838462

Santos da Silva K, Souza ACP, Rodrigues LRR, Pieczarka JC, Nagamachi CY. Chromosomal diversification in Pseudacanthicus species (Loricariidae, Hypostominae) revealed by comparative mapping of repetitive sequences. Animals. 2022b; 12:2612. https://doi.org/10.3390/ani12192612

Santos da Silva K, Glugoski L, Vicari MR, Souza ACP, Akama A, Pieczarka JC et al. Mechanisms of karyotypic diversification in Ancistrus (Siluriformes, Loricariidae): inferences from repetitive sequence analysis. Int J Mol Sci. 2023; 24(18):14159. https://doi.org/10.3390/ijms241814159

Sassi FMC, Deon GA, Moreira-Filho O, Vicari MR, Bertollo LAC, Liehr T et al. Multiple sex chromosomes and evolutionary relationships in Amazonian catfishes: the outstanding model of the genus Harttia (Siluriformes: Loricariidae). Genes. 2020; 11(10):1179. https://doi.org/10.3390/genes11101179

Sassi FMC, Moreira-Filho O, Deon GA, Sember A, Bertollo LAC, Liehr T et al. Adding new pieces to the puzzle of karyotype evolution in Harttia (Siluriformes, Loricariidae): investigation of Amazonian species. Biology. 2021; 10(9):922. https://doi.org/10.3390/biology10090922

Sassi FMC, Sember A, Deon GA, Liehr T, Padutsch N, Oyakawa OT et al. Homeology of sex chromosomes in Amazonian Harttia armored catfishes supports the X‑fission hypothesis for the X₁X₂Y sex chromosome system origin. Sci Rep. 2023; 13(1):15756. https://doi.org/10.1038/s41598-023-42617-w

Schott SCQ, Glugoski L, Azambuja M, Moreira-Filho O, Vicari MR, Nogaroto V. Comparative cytogenetic and sequence analysis of u small nuclear RNA genes in three Ancistrus species (Siluriformes: Loricariidae). Zebrafish. 2022; 19(5):200–09. https://doi.org/10.1089/zeb.2022.0040

Sember A, Nguyen P, Perez MF, Altmanová M, Petr R, Cioffi MB. Multiple sex chromosomes in teleost fishes from a cytogenetic perspective: state of the art and future challenges. Philos Trans Royal Soc London B Biol Sci. 2021; 376(1833):20200098. https://doi.org/10.1098/rstb.2020.0098

Silva KS, Souza ACP, Pety AM, Noronha RCR, Vicari MR, Pieczarka JC et al. Comparative cytogenetics analysis among Peckoltia species (Siluriformes, Loricariidae): insights on karyotype evolution and biogeography in the Amazon region. Front Genet. 2021; 12:779464. http://dx.doi.org/10.3389/fgene.2021.779464

Silva M, Ribeiro ED, Matoso DA, Sousa LM, Hrbek T, Rapp Py-Daniel L et al. Chromosomal polymorphism in two species of Hypancistrus (Siluriformes: Loricariidae): an integrative approach for understanding their biodiversity. Genetica. 2014; 142(2):127–39. https://doi.org/10.1007/s10709-014-9760-y

Sochorová J, Garcia S, Gálvez F, Symnonová R, Kovarík A. Evolutionary trends in animal ribosomal DNA loci: introduction to a new online database. Chromosoma. 2018; 127:141–50. https://doi.org/10.1007/s00412-017-0651-8

Souza ACP, Nagamachi CY, Milhomem SSR, Feldberg E, Pieczarka JC. Cytogenetic analysis in catfish species of the genus Peckoltia Miranda Ribeiro, 1912 (Teleostei: Siluriformes: Loricariidae). Comp Cytogenet. 2009; 3(2):103–09. https://doi.org/10.3897/compcytogen.v3i2.17

Souza ACP, Nascimento AL, Carvalho Jr. JR, Barros RMS, Feldberg E, Nagamachi CY et al. Karyotypic analysis of Baryancistrus aff. niveatus (Ancistrinae, Loricariidae) by C-banding, AgNOR, CMA3, DAPI and FISH. Caryologia. 2004; 57(3):219–23. https://doi.org/10.1080/00087114.2004.10589396

Stace CA. Cytology and cytogenetics as a fundamental taxonomic resource for the 20th and 21st centuries. Taxon. 2000; 49:451–77. https://doi.org/10.2307/1224344

Stawikowski R, Werner A, Seidel I. Fünfzehn jahre l-nummern/fifteen years of l-numbers. Die Aquarien- und Terrarienzeitschrift Special L-Nummern/L-Numbers, 4–5. Verlag Eugen Ulmer, Stuttgart, Germany; 2004.

Sullivan JP, Lundber JG, Hardman M. A phylogenetic analysis of the major groups of catfishes (Teleostei: Siluriformes) using rag1 and rag2 nuclear gene sequences. Mol Phylogenet Evol. 2006; 41(3):636–62. https://doi.org/10.1016/j.ympev.2006.05.044

Takagui FH, Baumgärtner L, Venturelli NB, Paiz LM, Viana P, Dionísio JF et al. Unrevealing the karyotypic evolution and cytotaxonomy of armored catfishes (Loricariinae) with emphasis in Sturisoma, Loricariichthys, Loricaria, Proloricaria, Pyxiloricaria, and Rineloricaria. Zebrafish. 2020; 17(5):319–32. https://doi.org/10.1089/zeb.2020.1893

Takagui FH, Rubert M, Dionisio JF, Baumgärtner L, Cardoso YP, Jerep FC et al. Cytogenetic markers reinforce the redescription of the armored pleco Hypostomus spiniger (Loricariidae – Hypostominae), an endemic species in the Uruguay River basin and Patos Lagoon system. Braz Arch Biol Technol. 2023; 66:e23220154. https://doi.org/10.1590/1678-4324-2023220154

Takagui FH, Venturelli NB, Dias AL, Swarça AC, Vicari MR, Fenocchio AS et al. The Importance of pericentric inversions in the karyotypic diversification of the species Loricariichthys anus and Loricariichthys platymetopon. Zebrafish. 2014; 11(4):300–05. https://doi.org/10.1089/zeb.2014.0985

Traldi JB, Blanco DR, Vicari MR, Martinez JF, Lui RL, Barros AV et al. Chromosomal diversity in Hypostomus (Siluriformes, Loricariidae) with emphasis on physical mapping of 18S and 5S rDNA sites. Genet Mol Res. 2013; 12(1):463–71. http://dx.doi.org/10.4238/2013.February.8.11

Traldi JB, Lui RL, Martinez JF, Vicari MR, Nogaroto V, Moreira-Filho O et al. Chromosomal distribution of the retroelements Rex1, Rex3 and Rex6 in species of the genus Harttia and Hypostomus (Siluriformes: Loricariidae). Neotrop Ichthyol. 2019; 17(2):e190010. https://doi.org/10.1590/1982-0224-20190010

Traldi JB, Vicari MR, Blanco DR, Martinez JF, Artoni RF, Moreira-Filho O. First karyotype description of Hypostomus iheringii (Regan, 1908): a case of heterochromatic polymorphism. Comp Cytogenet. 2012; 6(2):115–25. https://doi.org/10.3897/CompCytogen.v6i2.2595

Tursellino MN, Cioffi MB, Sassi FMC, Deon GA, Oliveira C, Kuranaka M et al. Integrating genomic and chromosomal data: a cytogenetic study of Transancistrus santarosensis (Loricariidae: Hypostominae) with characterization of a ZZ/ZW sex chromosome system. Genes. 2023; 14:1662. https://doi.org/10.3390/genes14091662

Venturelli NB, Takagui FH, Pompeo LRS, Rodriguez MS, Rosa R, Giuliano-Caetano L. Cytogenetic markers to understand chromosome diversification and conflicting taxonomic issues in Rineloricaria (Loricariidae: Loricariinae) from Rio Grande do Sul coastal drainages. Biologia. 2021; 76:2561–72. https://doi.org/10.1007/s11756-021-00748-3

Ziemniczak K, Barros AV, Rosa KO, Nogaroto V, Almeida MC, Cestari MM et al. Comparative cytogenetics of Loricariidae (Actinopterygii: Siluriformes): emphasis in Neoplecostominae and Hypoptopomatinae. Italian J Zool. 2012; 79(4):492–501. https://doi.org/10.1080/11250003.2012.676677

Authors

Francisco de Menezes Cavalcante Sassi¹, Marcelo de Bello Cioffi¹ and Orlando Moreira-Filho¹

^[1] Laboratório de Citogenética de Peixes, Departamento de Genética e Evolução, Universidade Federal de São Carlos, Rodovia Washington Luis, km 235, 13565-905 São Carlos, SP, Brazil. (FMCS) francisco.sassi@hotmail.com, (MBC) mbcioffi@ufscar.br (corresponding author), (OMF) omfilho@ufscar.br.

Authors’ Contribution

Francisco de Menezes Cavalcante Sassi: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Writing-original draft, Writing-review and editing.

Marcelo de Bello Cioffi: Conceptualization, Funding acquisition, Project administration, Supervision, Writing-original draft, Writing-review and editing.

Orlando Moreira-Filho: Conceptualization, Data curation, Investigation, Methodology, Supervision, Validation, Writing-review and editing.

Ethical Statement

Not applicable.

Competing Interests

The author declares no competing interests.

How to cite this article

Sassi FMC, Cioffi MB, Moreira-Filho O. A state-of-art review of Loricariidae (Ostariophysi: Siluriformes) cytogenetics. Neotrop Ichthyol. 2024; 22(4):e240050. https://doi.org/10.1590/1982-0224-2024-0050

Copyright

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

Diversity and Distributions Published by SBI

Accepted June 6, 2024

Submitted October 28, 2024 by Claudio Oliveira

Epub January 10, 2025

Abstract​

Introduction​

Material and methods

Results​

Discussion​

Acknowledgments​

References​