Integrative taxonomy clarifies the armoured catfish Hypostomus pusarum (Starks) species complex (Siluriformes: Loricariidae) and reveals a new species in the drainages of Northeastern Brazil Author Lustosa-Costa, Silvia Yasmin Author Ramos, Telton Pedro Anselmo Author Zawadzki, Cláudio Henrique Author Jacobina, Uedson Pereira Author Lima, Sergio Maia Queiroz text Zoological Journal of the Linnean Society 2024 2024-07-04 201 3 1 30 http://dx.doi.org/10.1093/zoolinnean/zlae059 journal article 302777 10.1093/zoolinnean/zlae059 fea30d55-e582-4306-9c14-c79ddc8c8b4f 0024-4082 13738347 Hypostomus cari sp. nov. ( Fig. 12 ; Table 2 ) Zoobank registration: urn:lsid:zoobank.org:act:F2652781* DB20*4BA7*9C32*CE77D691E82A Hypostomus aff. pusarum . —Lustosa*Costa et al. 2022:10/26 (listed as non* type specimens); Ramos et al. 2014:5 ( Hypostomus sp. 1 /sp. 4 in the check*list); Lima et al. 2017 ( Hypostomus sp. 1 / sp. 4, checklist); Silva et al. 2015:5 ( Hypostomus sp. 1 , checklist); Lustosa*Costa et al. 2022:6 ( Hypostomus aff. pusarum , phylogeny of genus in the Maranhão * Piauí ecoregion); Silva et al. 2023:8 ( Hypostomus sp.1 /sp. 4, checklist). Holotype : NºP 25422, 127.3 mm CP, Parnaíba River , Alto Parnaíba , Maranhão , Brazil . 09°05 ʹ 21 ʹʹ S , 45°55 ʹ 32 ʹʹ W , 9 Agosto 2013 , Claudio H. Zawadzki and Tiago Debona. Paratypes : All from Brazil . Parnaíba River basin. MCP 50049 , 2 , 76.4–80.9 mm SL, collected with holotype. MZUSP 98641 , 28 , 15.3–72.9 mm SL, Pureza River , between Alto Parnaíba and Tasso Fragoso , Maranhão , 8°47 ʹ 10 ʹʹ S , 45°56 ʹ 42 ʹʹ W , 6 Apr 2001 , A. Oyakawa , V . Garuưi, and J.C. Nolasco. NUP 15946 , 23, 31.0–132.0 mm SL, collected with holotype . NUP 15956 , 1, 67.9 mm SL, Corrente River , Corrente, Piauí , 10°26 ʹ 29.4 ʹʹ S , 45°10 ʹ 23.2 ʹʹ W 12 Aug 2013 , C.H. Zawadzki and D. Baumgartner. UFPB 11075 , 1 , 90.5 mm SL, Pureza River , in the road MA*006, between Alto Parnaíba and Tarso Fragoso , Maranhão , 08°47 ʹ 10 ʹʹ S , 45°56 ʹ 41 ʹʹ W , 7 Feb 2009 , T . P.A. Ramos , P. Charvet , R . T . C. Ramos , G. Moro , and E. Franca. UFPB 11088 , 24 , 47.3–97.7 mm SL, Parnaíba River , Tasso Fragoso , Maranhão, 07°33 ʹ 24 ʹʹ S , 45°14 ʹ 58 ʹʹ W , 3 Apr 2005 , W. Severi and E. França . UFPB 11076 , 16 , 56.1–150.4 mm SL, Balsas River , Parnaíba basin, Sambaíba, Maranhão, 07°08 ʹ 32.3 ʹʹ S , 45°20 ʹ 39.3 ʹʹ W , 08 Feb 2009 , T . P.A. Ramos , P. Charvet , R . T . C. Ramos , G. Moro , and E. Franca. UFPB 8240 , 2 , 103.4 –107.0 mm SL, Parnaíba River , Alto Parnaíba , Maranhão , 09°06 ʹ 52 ʹʹ S , 45°55 ʹ 35 ʹʹ W , 01 Apr 2010 , T . Ramos, S . Ramos, and M. Silva. UFRN 5511 , 4, 97.0– 138.1 mm SL, Muquem Creek , Povoado Manga , Barão do Grajaú , Maranhão , 06°47 ʹ 05.6‘S, 43°15 ʹ 46.4 ʹʹ W , 10 Dec 2018 , L. Neto , S. Costa , T . Ramos , and Y. Ponce. UFRN 3090 , 29, 16.6–95.9 mm SL, Parnaíba River , Alto Parnaíba , Maranhão , 09°06 ʹ 54 ʹʹ S , 45°55 ʹ 37 ʹʹ W , 23 Jun 2014 , T . Ramos , S. Lima, L. Neto, M. Silva, L. Medeiros, and R. Paiva. UFRN 2744 , 5, 57.8– 102.5 mm SL, Corrente River , Corrente , Piauí , 10°23 ʹ 14 ʹʹ S , 45°15 ʹ 05 ʹʹ W , 18 Apr 2014 , T . Ramos, L . Neto, M . Germano, and L. Medeiros . UFPB 11077 , 1, 164.5 mm SL, Ingazeiro reser* voir, Paulistana, Piauí, 08°09 ʹ 49 ʹʹ S, 41°08 ʹ 13 ʹʹ W, 10 Dec 2010, T. Ramos and S. Ramos. UFPB 11101 , 3, 130.0–151.0 mm SL, Ingazeiro dam, Paulistana , Piauí , 08°20 ʹ 39 ʹʹ S , 41°08 ʹ 16 ʹʹ W , 19 May 2009 , T . Ramos. UFRN 5555 , 1, 120.6 mm SL, between the rivers Parnaíba and Canindé , Amarante , Piauí , 6°15 ʹ 06 ʹʹ S , 42°51 ʹ 03 ʹʹ W , 09 Dec 2018 , L. Neto , S. Costa , T . Ramos , and Y. Ponce. UFRN 5664 , 3, 85.7–100.8 mm SL, Guaribas River , Torrões village , Picos , Piauí , 07°09 ʹ 11 ʹʹ S , 41°30 ʹ 53 ʹʹ W , 31 Dec 2018 , L. Neto , S. Costa , º. Jacobina , and L. Calado. UFPB 11069 , 1, 145.1 mm SL, reservoir in the Paracati River , São Francisco do Piauí, Piauí , 08°13 ʹ 27 ʹʹ S , 41°24 ʹ 48 ʹʹ W , 09 Dec 2010 , T . Ramos and S. Ramos. UFPB 11052 , 2 , 55.1 * 101.2 mm SL, Lagoa Parnaguá, less margin, Parnaguá, Piauí , 10°14 ʹ 01 ʹʹ S , 44°40 ʹ 43 ʹʹ W , 23 Jun.06, W. Severi. UFPB 8009 , 2 , 124.3 – 144.2 mm SL, Parnaíba River , Benedito Leite, Piauí , 07°14 ʹ 43 ʹʹ S , 44°34 ʹ 16 ʹʹ W , 17 Nov 2009 , T . Ramos, M . Silva and G. Moro . Non-types: All from Brazil . Parnaíba River Basin, Maranhão : UFPB 11091 , 1 , 52.2–91.8 mm SL, Marcelino Creek , tributary of the Parnaíba River , Tasso Fragoso , 08°26 ʹ 47 ʹʹ S , 45°47 ʹ 09 ʹʹ W , 3 Apr 2005 , W. Severi and E. França. UFPB 11050 , 3 , 17.5– 77.5 mm SL Parnaíba River , Alto Parnaíba , 09°07 ʹ 51 ʹʹ S , 45°55 ʹ 45 ʹʹ W , 24 Apr 2011 , T . Ramos . UFPB 1051 , 13 , 56.1– 78.7 mm SL, Parnaíba River , Alto Parnaíba , 09°06 ʹ 52 ʹʹ S , 45°55 ʹ 35 ʹʹ W , 24 Apr 2011 , T . Ramos, S . Ramos , and M. Silva . UFPB 11087 , 10 , 38.3–67.4 mm SL, Marcelino Creek , in the road MA*006 / BR *330, next to Tasso Fragosso , 07°33 ʹ 24 ʹʹ S , 45°14 ʹ 58 ʹʹ W , 03 Apr 2005 , W. Severi and E. Franca. UFPB 11152 , 16 , 95.5 mm SL, Parnaíba River , Benedito Leite , 07°13 ʹ 30 ʹʹ S , 044°31 ʹ 56 ʹʹ W , 17 Sep 2009 , T . Ramos, M . Silva , and G. Moro . UFPB 11085 , 1 , 71.1 mm SL, Balsas River , Parnaíba basin, Benedito Leite , 07°14 ʹ 09 ʹʹ S , 44°34 ʹ 24 ʹʹ W , 01 Apr 2005 , W. Severi and E. França. UFPB 11064 , 5 , 21.9–46.9 mm SL, Parnaíba River , Loreto , 07°33 ʹ 02 ʹʹ S , 45°14 ʹ 10 ʹʹ W , 23 Apr 2011 , T . Ramos, S . Ramos , and M. Silva . UFPB 11058 , 7 , 68.8– 106.2 mm SL, Parnaíba River , Magalhães de Almeida , 3°18 ʹ 24 ʹʹ S , 42°05 ʹ 36 ʹʹ W , 13 Oct 2010 , T . Ramos . UFPB 11123 , 1 , 94.3 mm SL, Parnaíba River , Parnarama , 5°39 ʹ 14 ʹʹ S , 43°04 ʹ 57 ʹʹ W , 22 Sep 2009 , T . Ramos, M . Silva , and G. Moro . UFPB 11054 , 5 , 50.0– 87.5 mm SL, Parnaíba River , town of Beira * Rio , Buriti , 03°53 ʹ 22 ʹʹ S , 42°43 ʹ 25 ʹʹ W , 18 Apr 2011 , T . Ramos . UFPB 8243 , 1 , 64.8 mm SL, Parnaíba River , town of Manga , Barão de Grajaú , 06°47 ʹ 03 ʹʹ S , 43°15 ʹ 46 ʹʹ W , 05 Apr 2010 , T . Ramos . UFPB 11065 , 11 , 13.29–89.30 mm SL, Parnaíba River , São Francisco do Maranhão , 06°15 ʹ 02 ʹʹ S , 42°51 ʹ 19 ʹʹ W , 21 Apr 2011 , T . Ramos . UFPB 8244 , 1 , 47.8 mm SL, Parnaíba River , São Francisco do Maranhão , 6°15 ʹ 55 ʹʹ S , 42°51 ʹ 21 ʹʹ W , 06 Apr 2010 , T . Ramos . UFPB 11083 , 1 , 12.5–12.7 mm SL, Parnaíba River , São Francisco do Maranhão , 06°14 ʹ 47 ʹʹ S , 42°51 ʹ 24 ʹʹ W , 20 Apr 2011 , T . Ramos and S. Ramos . UFPB 11057 , 1 , 40 , 39.0– 64.5 mm SL, Parnaíba River , Benedito Leite , 7°13 ʹ 28 ʹʹ S , 44°31 ʹ 55 ʹʹ W , 22 Apr 2011 , T . Ramos. UFRN 5666 , 1, 65.9 mm SL, Muquem Creek , Manga Village , Barão de Grajaú , 06°45 ʹ 12.7 ʹʹ S , 43°16 ʹ 26.8 ʹʹ W , 10 Dec 2018 , L. Neto, S. Costa, T . Ramos , and Y.Ponce. Parnaíba River Basin , Piauí : UFPB 11157 , 1, 107.9 mm SL, Rio Grande reservoir, Rio Grande do Piauí , 07°46 ʹ 39.3 ʹʹ S , 43°08 ʹ 47.7 ʹʹ W , 07 Dec 2010 , T . Ramos and S. Ramos . UFPB 11137 , 1 , 67.4 mm SL, Jacobina reservoir, Jacobina do Piauí , 07°56 ʹ 03 ʹʹ S , 41°12 ʹ 29 ʹʹ W , 10 Dec 2010, T. Ramos and S. Ramos. UFPB 11110 , 3, 42.0–83.5 mm SL, São Francisco lagoon, Rio Grande do Piauí , 07°46 ʹ 39 ʹʹ S , 43°08 ʹ 47 ʹʹ W , 15 Dec 2010 , T . Ramos and S. Ramos. UFPB 11135 , 1, 119.9 mm SL, public market, ºnião, 04°35 ʹ 24 ʹʹ S, 42°52 ʹ 31 ʹʹ W, 24 Sep 2009, T. Ramos, M. Silva and G. Moro. UFPB 11105 , 30, 20.4–43.2 mm SL, tributary of the Gurgueia River, Bom Jesus, 09°00 ʹ 44 ʹʹ S , 44°18 ʹ 10 ʹʹ W , 22 Jun 2006, W. Severi. UFPB 11104 , 10, 20.9–37.5 mm SL, tributary of the Gurgueia River, Bom Jesus, 09°05 ʹ 26 ʹʹ S , 44°21 ʹ 46 ʹʹ W , 22 Jun 2006 , T . Ramos. UFPB 11096 , 1, 42.4 mm SL, Bonfim Creek, ºruçui, 07°15 ʹ 43 ʹʹ S , 44°31 ʹ 43 ʹʹ W , 13 Nov 2009 , T . Ramos, M. Silva, and G. Moro. UFPB 11146 , 1, 45.0 mm SL, Tributary of the Longá River, road from Barra, 03°57 ʹ 53 ʹʹ S , 42°16 ʹ 28 ʹʹ W , 17 Jun 2006 , W. Severi. UFPB 11147 , 3, 39.5– 47.7 mm SL, Cruz Creek, tributary of the Poty River, BR *316, Monsenhor Gil, 05°35 ʹ 00 ʹʹ S , 42°36 ʹ 53 ʹʹ W , 20 Jun 2006 , W. Severi. UFPB 11086, 3, 24.9–41.7 mm SL, Muquila Creek, PI 130, Amarante, 06°13 ʹ 09 ʹʹ S , 42°53 ʹ 30 ʹʹ W , 08 Apr 2005 , W. Severi. UFPB 11138, 3, 33.4–85.4 mm SL, Manda Creek, Figure 12. Hypostomus cari , new species , Holotype, NºP 25422, 127.3 mm SL, Parnaíba River, Alto Parnaiba, Maranhão, Brazil. Table 2. Morphometric data and counts of Hypostomus cari , new species. Holotype data are included in range. N range = 20. SL, Standard length. SD, Standard deviation.

Holotype	Range	Mean	SD
Standard length	127.3	90.4–143.6	106.1	14.9
Percent of SL
Predorsal length	41.9	38.4–42.8	40.7	1.14
Head length	33.7	31.0–35.4	33.4	1.17
Cleithral width	31.9	27.8–33.7	31	1.28
Head depth	19.6	17.4–22.8	19.8	1.21
Interdorsal distance	19	12.42–19	14.9	1.45
Caudal peduncle length	32.9	28.2–34.1	31.2	1.44
Caudal peduncle depth	10.1	8.7–11.0	9.6	0.48
Dorsal*spine length	30.1	16.6–31.5	26.7	2.71
Pectoral*pelvic distance	24.2	20.4–26.3	23.2	1.34
Percent of head length
Cleithral width	94.6	87.1–95.9	91.4	3.1
Head depth	58.2	54.8–66.8	60.8	3.7
Snout length	65	59.2–66.3	63	1.53
Orbital diameter	16.5	11.8–17.6	15.2	1.48
Interobital width	39.5	34.4–44.0	38.7	2.22
Mandibular width	13.2	10.6–18.5	15	2.02
Other percents
Orbital diameter in snout length	25.5	18.8–28.7	24.2	2.58
Orbital diameter in interorbital length	41.9	27.2–49.8	39.6	5.52
Mandibular length in interobital length	55.7	32.2–55.7	41	5.99
Dorsal*spine length in predorsal length	71.7	55.7–78.0	66.4	5.54
First pectoral*fin ray length in predorsal length	80.8	63.6–87.4	75.2	5.5
Lower caudal*fin length in predorsal length	*	71.7–96.1	85.6	7.17
Adipose*fin length in caudal*peduncle depth	73.7	72.4–99.4	86.6	6.1
Caudal*peduncle depth in caudal*peduncle length	30.8	27.2–37.6	30.9	2.28
Interdorsal length in dorsal*fin base length	53.4	39.1–66.3	57.5	5.75
Lower lip width in lower lip length	28.9	23.4–35.5	28.4	3.07
Counts mode
Median plates series	25	25–26	26	0.3
Plates bordering supraoccipital	3	3–3	3	0
Predorsal plates	6	6–7	7	0.4
Dorsal plates below dorsal*fin bases	8	8–9	8	0.3
Plates between dorsal and adipose fins	6	6–7	7	0.4
Plates between adipose and caudal fins	4	3–4	4	0.5
Plates between anal and caudal fins	12	11–13	12	0.6
Premaxillary teeth	22	22–36	36	3.1
Dentary teeth	23	23–35	35	2.8

tributary of Poty River, Bandarrá, Beneditinos, 05°24 ʹ 15 ʹʹ S , 42°27 ʹ 20 ʹʹ W , 20 Jun 2006 , W. Severi. UFPB 11068 , 1, 75.3 mm SL, Negro Creek, Castelhanos, Palmeirais, 05°46 ʹ 12 ʹʹ S , 43°04 ʹ 23 ʹʹ W , 09 Apr 2005 , W. Severi. UFPB 11155 , 47, 29.2– 50.5 mm SL, Matões Creek, tributary of the Gurgueia River, Barra Verde, Bom Jesus, 9°14 ʹ 08 ʹʹ S , 44°27 ʹ 31 ʹʹ W , 22 Jun 2007 , W. Severi. UFPB 11136 , 5, 29.6–81.6 mm SL, Fundo Creek, in the road PI*130, Castelhanos, Palmeirais, 06°07 ʹ 37 ʹʹ S , 42°58 ʹ 39 ʹʹ W , 07 Apr 2005 , W. Severi. UFPB 11067 , 1, 33.1 mm SL, Passagem das Ovelhas Creek, Caxingo, 03°24 ʹ 25 ʹʹ S, 41°53 ʹ 49 ʹʹ W, 16 Jun 2006, W. Severi. UFPB 11141 , 2, 34.7–80.0 mm SL, Povo dos Homens Creek, tributary of the Longá River, Caxingo, 03°25 ʹ 12 ʹʹ S , 41°53 ʹ 46 ʹʹ W , 16 Jun 2006 , W. Severi. UFPB 11145 , 9, 27.3–33.8 mm SL, Berlenga River, tributaryofthePotiRiver, BR *316, PassagemFranca, 05°51 ʹ 31 ʹʹ S , 42°26 ʹ 07 ʹʹ W , 20 Jun 2006 , W. Severi. UFPB 11144 , 2, 66.7– 121.4 mm SL, Canindé River, Francisco Aires, 06°35 ʹ 12 ʹʹ S , 42°42 ʹ 16 ʹʹ W , 19 Nov 2009 , T . Ramos, M. Silva, and G. Moro. UFRN 2121 , 1, 66.8 mm SL, Canindé River, downstream of the dam Pedra Redonda, Conceição do Canindé, 07°58 ʹ 30 ʹʹ S , 41°59 ʹ 26 ʹʹ W , 18 Aug 2013 , A. Moraes, R . Paiva, L. Neto, M. Silva, T . Ramos, and S. Lima . UFPB 11059 , 5, 16.6–43.3 mm SL, Canindé River, Conceição do Canindé, 07°59 ʹ 49 ʹʹ S , 41°29 ʹ 59 ʹʹ W , 19 Jul 2010 , T . Ramos and S. Ramos. UFPB 11040 , 11, 43.2–84.2 mm SL, Canindé River, Oeiras, 06°59 ʹ 12 ʹʹ S , 42°05 ʹ 17 ʹʹ W , 21 Jun 2011 , W. Severi. UFPB 11045 , 5, 20.1– 45.0 mm SL, Canindé River, in the road BR 343, Amarante, 06°17 ʹ 41 ʹʹ S , 43°28 ʹ 11 ʹʹ W , 08 Apr 2005 , W. Severi. UFPB 11048 , 1, 32.9 mm SL, Contrato River, tributary of the Gurgueia River, Parnaíba basin, Contrato, 09°34 ʹ 45 ʹʹ S , 45°01 ʹ 45 ʹʹ W , 22 Jun 2006 , T . Ramos. UFRN 2722 , 46.0– 86.2 mm SL, 8, Corrente River, in the road BR 135, Corrente, 10°25 ʹ 30 ʹʹ S , 45°11 ʹ 47 ʹʹ W , 18 Apr 2014 , T . Ramos, L. Neto, M. Germano, and L. Medeiros. UFRN 1791 , 9, 87.7–155.7 mm SL, Guaribas River, Torrões, Picos, 07°09 ʹ 13 ʹʹ S , 41°30 ʹ 10 ʹʹ W , 23 Mar 2013 , S. Lima, T. Ramos, W. Berbel *Filho, L. Neto, S. Moraes and A. Moraes. UFRN 3754 , 3, 59.3–76.6 mm SL, Gurguéia River, in the road BR *135, São Gonçalo do Gurguéia, 10°01 ʹ 37 ʹʹ S , 45°18 ʹ 12 ʹʹ W , 19 Jun 2014 , S. Lima, T. Ramos, R. Paiva, M. Silva, M. Germano, Y. Ponce, L. Neto, and L. Medeiros. UFRN 2867 , 3, 38.4–57.5 mm SL, Gurguéia River, in the road BR 135, São Gonçalo do Gurguéia, 10°1 ʹ 37 ʹʹ S , 45°18 ʹ 12 ʹʹ W , 19 Jun 2014 , S. Lima, R. Paiva, M. Silva, and Y. Ponce. UFPB 11115 , 3, 45, 88.0– 63.4 mm SL, Gurgueia River, Cristino Castro, 08°48 ʹ 55 ʹʹ S , 44°13 ʹ 47 ʹʹ W , 22 Jun 2006 , W. Severi. UFPB 11107 , 7, 25, 74.0– 72.9 mm SL, Gurguéia River, Jurumenha, 07°04 ʹ 36 ʹʹ S , 43°30 ʹ 54 ʹʹ W , 07 Apr 2005 , W. Severi. UFPB 11112 , 20, 13.6–108.4 mm SL, Itaueira River, tributary of Parnaiba, Floriano, 06°48 ʹ 30 ʹʹ S , 42°57 ʹ 30 ʹʹ W , 07 Apr 2005 , W. Severi. UFPB 11060, 2, 39.0– 47.5 mm SL, Itaueira River, in the road BR *230, Floriano, 06°48 ʹ 43 ʹʹ S , 42°57 ʹ 30 ʹʹ W , 07 Apr 2005 , W. Severi. UFPB 11133 , 6, 41.4–94.4 mm SL, Longá River, Nossa Senhora de Nazaré, 04°40 ʹ 20 ʹʹ S , 42°13 ʹ 00 ʹʹ W , 23 Jun 2012 , T . Ramos. UFPB 11118 , 4, 38.1–47.9 mm SL, Parnaíba River, ºnião, 04°33 ʹ 46 ʹʹ S , 42°52 ʹ 03 ʹʹ W , 23 Sep 2009 , T . Ramos, M. Silva, and G. Moro. UFPB 11103 , 2, 40.4–64.5 mm SL, Parnaíba River, Amarante, 06°14 ʹ 36 ʹʹ S , 42°51 ʹ 24 ʹʹ W , 08 Apr 2005 , W. Severi. UFPB 8241 , 2, 19.9–43.4 mm SL, Parnaíba River, Ribeiro Gonçalves, 07°33 ʹ 24 ʹʹ S , 45°14 ʹ 58 ʹʹ W , 03 Apr 2010 , T . Ramos and S. Ramos. UFPB 11122 , 2, 73.0– 94.2 mm SL, Parnaíba River, Ribeiro Gonçalves, 07°33 ʹ 08 ʹʹ S , 45°14 ʹ 17 ʹʹ W , 10 Feb 2009 , T . Ramos. UFPB 8392 , 11, 17, 28.0– 47.1 mm SL, Parnaíba River, Santa Filomena, 07°08 ʹ 04 ʹʹ S , 45°55 ʹ 45 ʹʹ W , 02 Apr 2010 , T . Ramos. UFPB 9556 , 8, 58.1–89.3 mm SL, Parnaíba River, BR 321, Ribeiro Goncalves, 07°33 ʹ 07 ʹʹ S , 45°14 ʹ 13 ʹʹ W , 31 Mar 2005 , W. Severi. UFPB 8246 , 1, 17.64 mm SL, Parnaíba River, town Mangueira, Palmeirais, 05°39 ʹ 40 ʹʹ S , 43°05 ʹ 08 ʹʹ W , 07 Apr 2010 , T . Ramos and S. Ramos. UFPB 11156 , 7, 54.2– 70.6 mm SL, Piauí River, BR *020, São João do Piauí , 08°22 ʹ 00 ʹʹ S , 42°14 ʹ 11 ʹʹ W, 21 Jun 2006, W. Severi. UFPB 11151 , 1, 93.7 mm SL, Poti River, in the road PI*115, Juazeiro do Piauí, 5°11 ʹ 21 ʹʹ S, 41°42 ʹ 31 ʹʹ W , 19 Jun 2006 , W. Severi. UFPB 11153 , 1, 76.4 mm SL, Sambito River, Santa Cruz dos Milagres, 05°55 ʹ 40 ʹʹ S , 42°04 ʹ 48 ʹʹ W , 18 Nov 2009 , T . Ramos. UFPB 11139 , 3, 37.5– 74.9 mm SL, Sambito River, São Felix do Piauí , 05°55 ʹ 39 ʹʹ S , 42°04 ʹ 47 ʹʹ W , 12 Sep 2009 , T . Ramos, M. Silva, and G. Moro. UFPB 11079 , 5, 83, 44.0– 123.8 mm SL, Sambito River, Sítio Miguel Alves, Aroazes, 06°08 ʹ 27 ʹʹ S , 42°02 ʹ 40 ʹʹ W , 10 Sep 2009 , T . Ramos, M. Silva, and G. Moro. UFPB 11149 , 4, 42.44–77.1 mm SL, Sambito River, dowstream Mesa de Pedra dam, Valença, 06°14 ʹ 11 ʹʹ S , 42°04 ʹ 47 ʹʹ W , 09 Sep 2009 , T . Ramos, M. Silva, and G. Moro. UFRN 1238 , 3, 52.9–87.5 mm SL, Sambito River, Mesa de Pedra dam, Aroazes, 06°11 ʹ 32 ʹʹ S , 41°59 ʹ 35 ʹʹ W , 24 Mar 2013 , S. Lima, T. Ramos, W. Berbel *Filho, L. Neto, S. Moraes, and A. Moraes. UFRN 5633 , 1, 59.3 mm SL, Canindé River, Amarante, 06°17 ʹ 42.7 ʹʹ S , 42°48 ʹ 49.4 ʹʹ W , 08 Dec 2018 , L. Neto, S. Costa, T . Ramos, and Y. Ponce. UFRN 5652 , 10, 29.2–40.9, Guaribas River, Torrões village, Picos, 07°09 ʹ 11.8 ʹʹ S , 41°30 ʹ 53.7 ʹʹ W , 31 Dec 2018 , L. Neto, S. Costa, º. Jacobina, and L. Calado. UFRN 5655 , 3, 55.0– 56.2 mm SL, Pejuaba River, up* stream of the dam Piracuruca, Alto Alegre, 04°00 ʹ 15.6 ʹʹ S , 41°26 ʹ 53.4 ʹʹ W , 06 Dec 2018 , L. Neto, S. Costa, T . Ramos, and Y. Ponce. UFRN 5657 , 3, 61.7–64.0 mm SL, Caldeirão Creek, Floriano, 06°39 ʹ 26 ʹʹ S , 42°53 ʹ 44 ʹʹ W , 09 Dec 2018 , L. Neto, S. Costa, T . Ramos, and Y. Ponce. UFRN 5552 , 2, 74.2–82.7 mm SL, Caldeirão Creek, Floriano, Brasil , 06°39 ʹ 26.7 ʹʹ S , 42°53 ʹ 44.6 ʹʹ W , 09 Dec 2018 , L. Neto, S. Costa, T . Ramos, and Y. Ponce. UFRN 5668 , 5, 47.2–111.70 mm SL, Mulato Creek, Amarante, 06°14 ʹ 09.0 ʹʹ S , 42°51 ʹ 29.7 ʹʹ W , 09 Dec 2018 , L. Neto, S. Costa, T . Ramos, and Y.Ponce. UFRN 5513 , 5, 42.2–90.0 mm SL, Pejuaba River, upstream of the dam Piracuruca, Alto Alegre, 04°00 ʹ 15 ʹʹ S , 41°26 ʹ 53 ʹʹ W , 06 Dec 2018 , L. Neto, S. Costa, T . Ramos, and Y. Ponce. UFRN 3043 , 2, 56.4–111.7 mm SL, Cachoeira do Pintado, Água Quente River, Barreiras do Piauí , 10°11 ʹ 28.9 ʹʹ S , 45°51 ʹ 12.9 ʹʹ W , 20 Jun 2014 , T . Ramos, S. Lima, L. Neto, M. Silva, Y. Ponce, and R. Paiva. UFRN 2220 , 4, 32.7– 46.5 mm SL, Piauí River, downstream of the Jenipapo dam, Jenipapo, 08°27 ʹ 13 ʹʹ S , 42°09 ʹ 53 ʹʹ W , 18 Aug 2013 , A. Moraes, R . Paiva, L. Neto, M. Silva, T . Ramos, and S. Lima . UFRN 1492 , 4, 31.3–59.4 mm SL, Guaribas River, village Torrões, Picos, 07°09 ʹ 13 ʹʹ S , 41°30 ʹ 10 ʹʹ W , 23 Mar 2013 , S. Lima, T. Ramos, W. Berbel *Filho, L. Neto, S. Moraes, and A. Moraes. Parnaíba River Basin, Ceará : UFPB 11150 , 1, 75.6 mm SL, Cabaças Creek, tributary of Poti, Ibiapaba, 05°04 ʹ 49 ʹʹ S , 40°52 ʹ 01 ʹʹ W , 22 Jul 2009 , T . Ramos, M. Silva, and G. Moro. UFRN 5078 , 8, 46.1– 101.2 mm SL, Poty River, tablelands of Ibiapaba, Cratéus, 05°03 ʹ 27 ʹʹ S , 40°50 ʹ 13 ʹʹ W , 30 Jun 2018 , L. Neto, S. Costa, and T . Ramos. Diagnosis Hypostomus cari differs from H. affinis , H. ancistroides , H. argus , H. aspilogaster , H. borellii , H. boulengeri , H. careopinnatus , H. carinatus , H. crassicauda , H. derbyi , H. dlouhyi , H. formosae , H. hoplonites , H. krikati , H. lamberti , H. laplatae , H. melanephelis , H. micromaculatus , H. niceforoi , H. pantherinus , H. piratatu , H. plecostomus , H. robinii , H. spiniger , H. subcarinatus , H. tapijara , H. vaillanti , H. watwata , and H. wilsoni by lacking or having incospicuous keels (vs. strong and conspicuous keels). It dif* fers from H. alatus , H. albopunctatus, H. arecuta, H. bimbai, H. delimai , H. faveolus , H. þancisci , H. krishnamurtii , H. luteus , H. luteomaculatus , H.margaritifer , H. meleagris , H. microstomus , H. myersi , H. multidens , H. regani , H. roseopunctatus , H. sertanejo , H. strigaticeps , H. tietensis , and H. variipictus by having dark spots on body and fins (vs. lighter spots); from H. corantijni , H. crassicauda , H. gymnorhinchus , H. hemiurus , H. micromaculatus , H. nematopterus , and H. paucimaculatus by having roundish dark spots (vs. dark spots horizontally elongate—see: Weber et al . 2012 : figs 8, 9). Ŋe new species is distinguished from H. bolivianus , H. denticulatus , H. þeirei , H. isbrueckeri , H. johnii , H. kuarup , H. latirostris , H. macrops , H. mutucae , H. paulinus , H. renestoi , H. ternetzi , H. unae , H. velhomonge , and H. yaku by having smaller number of teeth on both premaxillary and den* tary, rarely more than 50 (vs. higher number of teeth on both pre* maxillary and dentary, more than 60), and from H. denticulatus , H. peckoltoides , H. velhochico , and H. yaku by having pectoral fins lacking hypertrophied odontodes, even in adult specimens (vs. strongly*developed odontodes). It differs from H. heraldoi for having the first pectoral*fin ray clearly longer than the first pelvic*fin ray (vs. first pectoral*fin ray equal to or shorter than the first pelvic*fin ray). Hypostomus cari differs from H. laplatae by having from 25 to 27 plates in lateral serie (vs. 31); from the H. cochliodon group ( sensu Zawadzki et al . 2014) by having slender villiform bicuspid teeth (vs. robust spoon*shaped teeth), and having dentary angle between 90° and 115° (vs. dentary angles of 80° to 90°). It differs from H. pusarum by having medial buccal papilla inconspicuous or pointed and smooth (vs. medial buccal papilla well*developed, papillose and rough); by having a short dorsal fin, not reaching the origin of the adipose fin; by having pectoral fins with narrow distal end (vs. pectoral fins with wider distal end in adults, see Fig. 6 ); and for having pre*anal plate (vs. pre*anal plate absent). Description Meristic and morphometric data in Table 2 . Head and snout rounded in dorsal view, eye relatively large in size, positioned dorsolaterally, dorsal margin of orbit slightly elevated; dorsal region of head completely covered by dermal plates and odontodes. Predorsal plate with small elevation that forms in* conspicuous keel. Mouth ventral, oral disc rounded, lips with pa* pillae; on lower lip, papillae larger closest to mouth, decreasing in size further from oral cavity; internal buccal papilla absent in most specimens, not hypertrophied when present. Lower lip not reaching transverse line between gill openings. Dentary angle between 90° and 115°. Teeth villiform and bicuspid, with lateral cusp larger than medial cusp, curved towards oral cavity. Maxillary barbel slightly smaller than pupil. Dorsal and lateral surface of body covered with plates, except for dorsal*fin base. Tip of snout almost completely covered by plates. Dorsal profile straight from tip of snout to interorbital region, low keel on supraoccipital process; head broad, slightly depressed. Ŋree straight predorsal plates. Abdomen and thorax covered with plates, except area of insertion of pectoral and pelvic fins. Pre*anal plate present. Ventral region of head covered with small plates, except for below lower lip. Ventral region of body straight from snout to insertion of pelvic fins. Body widest at cleithrum, progressively narowing to end of caudal peduncle. Body covered by five lateral series of plates: dorsal series with inconspicuous keel from its origin to adipose fin; mid*dorsal series without keel; median series with complete lateral line and without keel; lateral line with 25 to 26 plates; midventral series bent to fourth or fissh plate. Dorsal fin II, 7; when depressed reaching origin of adi* pose fin. First dorsal*fin ray slightly curved. Pectoral fin I, 6; first unbranched pectoral*fin ray long and depressed, reaching one*third of first unbranched pelvic*fin ray, with poorly devel* oped odontodes on distal region; width of first pectoral*fin ray increasing slightly on distal region in some adult individuals. Adipose fin developed, inserted from six to seven plates asser posterior insertion of dorsal fin. Pelvic fin I, 5; slightly curved inwards, first ray unbranched with narrow proximal and distal regions and widened and flaưened centre, with small odontodes. Anal fin I, 4; first anal*fin ray with slightly concave distal edge, extending to seventh plate asser its origin. Caudal fin I, 14, I; ven* tral lobe slightly longer than dorsal, and same size in some spe* cimens. Colour in alcohol: Overall ground colour of dorsal and ventral regions of body, head and fins yellowish*brown. First rays of pectoral and pelvic fins yellowish at distal end. Small, dark, con* spicuous spots, close together in head region. Towards posterior portion of body spots more widely spaced, larger (larger than the diameter of the pupil). In most specimens, spots arranged longi* tudinal from plate in which lateral line begins to end of caudal peduncle. Inter*radial area of pectoral, pelvic and caudal fins, with darker spots, forming transverse bands. Bands five to eight on pectoral and pelvic fins, and five to 10 on caudal fin, with same colour as spots in back. Ŋe dorsal fin with row of transverse conspicuous spots in interradial membrane. Ventral region of body lighter than dorsal (light brown), also showing large and conspicuous spots. Some specimens with small and widely spaced spots only in ventral region. Colour in life: Colour paưern of live specimens resembling that of fixed specimens, except that light yellow body colour more intense, and spots on body darker and more conspicuous ( Fig. 13A ). Iris operculum visible, with light beige colour, forming notch in centre of dark iris, not visible in most fixed individuals. Two melanistic individuals (ºFRN 3043), with dark brown body colour and black spots ( Fig. 13B ). Distribution: Hypostomus cari is widely distributed in the Parnaíba ecoregion, is known in the upper, middle, and lower portions of the Parnaíba River basin ( Fig. 14 ). Ecological notes and habitat: Ŋe type locality of Hypostomus cari is the Parnaíba River in Povoado de Manga, Barão de Grajaú, Maranhão , Brazil . Ŋis locality is characterized by rapids, with rocky substrate and high oxygen flow ( Fig. 15 ). At that point, the species coexists with other species such as: Astyanao lacustris (Lütken, 1875) , Knodus victoriae (Steindachner, 1907) , Moenkhausia dichroura (Kner, 1858) , Tetragonopterus argenteus Cuvier, 1816 ( Characidae ), Hypostomus johnii , H. vaillanti , H. velhomonge , Loricaria parnahybae Steindachner, 1907 ( Loricariidae ), Prochilodus lacustris Steindachner , 1907 ( Prochilodontidae ), Caenotropus labyrinthicus (Kner, 1858) ( Chilodontidae ), Curimata macrops Eigenmann and Eigenmann, 1889, Steindachnerina notonota ( Curimatidae ), Geophagus parnaibae Staeck and Schindler, 2006 , Saoatilia brasiliensis ( Cichlidae ), Hassar affinis (Steindachner, 1881) ( Doradidae ), Hemiodus parnaguae Eigenmann and Henn, 1916 ( Hemiodontidae ), H. malabaricus (Erithrinidae), Leporinus piau ( Anostomidae ), Pellona flavipinnis (Valenciennes, 1837) ( Pristigasteridae ), Pimelodella parnahybae Fowler, 1941 ( Pimelodidae ), and Pterengraulis atherinoides (Linnaeus, 1766) ( Engraulidae ). Ŋe species has a wide distribution with records in the upper, middle, and lower portions of the Parnaíba River basin, both in the Caatinga and Cerrado, being the most widely distributed species of Hypostomus in the ecoregion (Lustosa* Costa et al. 2022). Conservation status: Ŋe new species has a wide geographic dis* tribution, collected throughout the Parnaíba River basin, in the states of Ceará , Maranhão and Piauí , in the upper, middle, and lower reaches of the basin, even occurring in preserved areas near the Nascentes do Rio Parnaíba National Park ( Silva et al. 2023 ). Ŋerefore, based on currently available data, and using the criteria of the IºCN Subcommiưee on Norms and Petitions (IºCN 2024), it is suggested that H. cari be classified as ‘Least Concern’ (LC). Type locality: Parnaíba River, Alto Parnaíba, Maranhão , Brasil . 09°05 ʹ 21 ʹʹ S 45°55 ʹ 32 ʹʹ W ( Fig. 15 ). Etymology: Ŋe specific epithet cari refers to one of the popular names that riverside fishermen from the Parnaíba River and other regions refer to the armoured catfishes of the Loricariidae family, such as those of the genus Hypostomus . Comparative material: Brazil : Hypostomus johnii : ºFRN 1341, 10, 10.4–64.6 mm SL, Sambito River, downstream Mesa de Pedra Dam, Aroazes, Piauí . ºFRN 1233, 32, 13.9–93.5 mm SL, Sambito River, downstream the Mesa de Pedra Dam, Aroazes, Piauí . ºFRN 2866, 3, 15.1–59.5 mm SL, Gurguéia River, under bridge on road BR 135, São Gonçalo do Gurguéia, Piauí . ºFRN 2708, 2, 35.4–59.7 mm SL, Tají River, Corrente, Piauí . ºFRN 3024, 5, 38.6–100.6 mm SL, ºruçuí*Vermelho River, Prata Village, Barreiras do Piauí , Piauí . ºFRN 2891, 8, 62.4–83.1 mm SL, Gurguéia River headwaters, São Gonçalo do Gurguéia, Piauí . ºFRN 1800, 6, 83.4–116.8 mm SL, Guaribas River, in Torrões Village, Picos, Piauí . ºFPB 3537, 9, 34.9–86.1 mm SL, Jacaraí River, under bridge on road PI*111, Piracuruca, Piauí . ºFPB 9535, 18, 57.3–94.5 mm SL, Longá River, Nossa Senhora de Nazaré, Piauí . ºFPB 11078, 6, 43.7– 67.5 mm SL, Gurguéia River, Jurumenha, Piauí . ºFPB 9267, 5, 92.4–129.1 mm SL, Poti River, Crateús, Ceará . ºFPB 9536, 12, 22.8–86.1 mm SL, Longá River, Cachoeira do ºrubu eco* logical park, Batalha, Piauí . ºFPB 9533, 1, 121.9 mm SL, São Nicolau River, Santa Cruz dos Milagres, Piauí . Hypostomus velhomonge : MZºSP 5068, 3, 23.5–98.7 mm SL, Grajaú, Grajaú River, Department of Zoology Tour. MZºSP 98591, 1, 81.2 mm SL, Balsas, Balsas River, Parnaíba River basin. ºFRN 5514, 63.7–83.03 mm SL, Muquem stream, Manga, Barão do Grajaú, Maranhão . ºFRN 3092, 23, 14.5–84.5 mm SL, Parnaíba River, Alto Parnaíba, Maranhão . Hypostomus vaillanti : ºFRN 5648, 3, 167.5–240.2 mm SL, Parnaíba River, Manga Village, Barão do Grajaú, Maranhão. NMW 44273, 1, 130.5 mm SL, syntype . NMW 44277, 1, 168.8 mm SL, syn* type, Preto River (tributary of Grande River, São Francisco River) at Ibipetuba. NMW 44276, 3, 75.8–119.4 mm SL, syntype . MCZ 7842, 2, 121.4– 140.8 mm SL. MCZ 7817, 1, 113.4 mm SL, Poti River. MCZ 7833, 1, 192.0 mm SL, San Gonçalo. Hypostomus chrysostiktos : ºFBA 2787, 3, 160.2– 175.4 mm SL, Bahia, Iaçu, Paraguaçu River. Hypostomus þancisci : MSL 14038, 1, 180.0 mm SL, Minas Gerais, Três Marias reservoir. NºP 9940, 6, 111.0– 187.1 mm SL. NºP 9945, 2, 148.6– 150.7 mm SL, Minas Gerais, das Velhas River. ºFPB 3021, 4, 99.7–115.6 mm SL, Bahia, Santana, Corrente River. Hypostomus garmani : ºFPB 3018, 1, 89.0 mm SL, Bahia, Santana, Corrente River. Hypostomus jaguar : NºP 4448, 2, 126.8– 152.9 mm SL, Bahia, Itaberabá, Paraguaçu River. ºFBA 6501, 8, 119.2– 175.7 mm SL, Itaberabá, Paraguaçu River, 10 km from Vila Vicente. Hypostomus cf. margaritifer : ºFPB 3019, 1, 97.1 mm SL. ºFPB 3041, 1, 97.1 mm SL, Bahia, Corrente River, São Francisco River basin. Hypostomus unae : MSL 41334, 3, 55.2–120.8 mm SL, Bahia, Panelinha River, tributary of Pardo River. MSL 41473, 10, 80.2–126.5 mm SL, Bahia, Preto do Costa River, tributary of the Contas River; NºP 9811, 5, 78.9– 53.7 mm SL, Bahia, das Pedras River, de Contas River basin. NºP 9814, 81.5–102.7 mm SL, Bahia, Oricó River, de Contas River basin. Hypostomus brevicauda : BMNH 1864, 1, 19.1–17. 2, 189.0* 196.1 mm SL, syntypes . MSL 36709, 3, 52.7* 125.4 mm SL, Bahia, Camacã, Traíra creek. ºFBA 0066, 1, 133.92 mm SL, Bahia, Xique*Xique, Suacica River, São Francisco River basin. ºFPB 4541, 15, 140.5– 189.2 mm SL, Bahia, ºna, uma River, Brasília, 8 km SE of São José. ºFPB 4563, 1, 166.9 mm SL, Bahia, ºna, Aliança River. Hypostomus mutacae : MSL 28669, 67.7 mm SL, holotype , Mato Grosso, Mutuca River, tributary of Paraguai River. NºP 6641, 13, 52.4–109.2 mm SL. NºP 6642, 4, 62.1–98.1 mm SL, Mato Grosso, Claro River, tributary of Paraguai River. Hypostomus alatus : ºFPB 3020, 1, 122.2 mm SL. ºFPB 7459, 1, 121.1 mm SL, Bahia, Santana, Corrente River. ºFRN 5509, 1, 88.7 mm SL Riacho do Poção, Juazeiro, Bahia. NºP 9119, 1, 110.1 mm SL, Minas Gerais, Curimataí River. NºP 9829, 5, 139.0– 77.4 mm SL, Minas Gerais, das Velhas River. NºP 9837, 4, 124.4– 217.6 mm SL, Minas Gerais, Cipó River. Hypostomus lima : BMNH 1876.1.10 , 2, 72.9–86.1 mm SL, syntypes , Minas Gerais, Lagoa Santa. NºP5717, 4, 56.1–126.0 mm SL, Minas Gerais, Ribeirão dos Patos. NºP 5721, 2, 47.5–72.8 mm SL, Minas Gerais, Minhocas stream. NºP 9827, 18, 81.5* 181.5 mm SL, Minas Gerais, São Miguel. Hypostomus paulinus : NºP 5344, 1, 69.0 mm SL, São Paulo, Passa Cinco River, tribu* tary of Piracicaba River. NºP 5379, 2, 84.1–91.7 mm SL, São Paulo, Passa Cinco River, tributary of the Piracicaba River. NºP 5722, 1, 90.5 mm SL, São Paulo, Passa Cinco River, tribu* tary of the Piracicaba River. NºP 6411, 17, 46.5–103.28 mm SL; NºP 6413, 13, 45.9–98.9 mm SL, São Paulo, Piracicaba River, tributary of the Tietê River. Hypostomus multidens : NºP 5340, 1, 157.0 mm SL, São Paulo, Piraju, Paranapanema River, Chavantes reservoir. Hypostomus regani : BMNH 1905.6.7.3, 174.2 mm SL, holotype , São Paulo, Piracicaba River. Hypostomus strigaticeps : BMNH 1907.7.6.1012, syn* types, 3, 75.7–160.0 mm SL, São Paulo, Piracicaba River, Tietê River basin. NºP 4017, 2, 72.8–100.0 mm SL, São Paulo, Ipuã, Ipanema River, Tietê River basin. NºP 4538, 11, 82.0–140 mm SL, São Paulo , Corumbataí River, tributary of the Tietê River. Hypostomus yaku : NºP 15348, 6, 29.8–58.1 mm SL, Goiás , Quente River, Paranaíba River basin.