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 Redescription of Hypostomus pusarum ( Starks, 1913 ) ( Figs 5 , 6 ; Table 1 ; Supporting Information, Table S3) Plecostomus pusarum : Starks 1913: 36 . Type locality: Ceará* Mirim River , Rio Grande do Norte, Brazil . Holotype : CAS *Sº 22225 . Paratype : CAS *Sº 22221 . Hypostomus pusarum : —Böhlke 1953:46 (catalogue of spe* cies); Ferraris 2007:259 (checklist); Isbrücker 1980:32 (catalogue of species); Burgess 1989:432 (atlas of marine and freshwater species); Isbrücker 2001:28 (genus nomenclature); Isbrücker 2002:19 (genus nomenclature); Britski and Garavello 2002:287 (comparative material); Reis et al. 2003:362 (check* list); Armbruster 2004:80 (literature compilation); Zanata et al. 2013:247 (comparative material); Zanata and Pitanga 2016:231 (comparative material); Zawadzki et al. 2017b:571 (compara* tive material); Ramos et al. 2017:8 (comparative material); Zawadzki et al. 2017a:564 (comparative material and picture of Hypostomus pusarum ); Da Costa et al. 2017 (checklist); Lima et al. 2017 (checklist); Lustosa*Costa et al. 2022 (comparative material). Figure 3. Genetic distances (Kimura 2 parameters) among species of the genus Hypostomus from the NCCD ecoregion, the Hypostomus aff. pusarum , and other species of the São Francisco and Parnaíba ecoregions. Basin abbreviations: APO (Apodi*Mossoro), PAR (Piranhas*Acu), PBN (Paraiba do Norte), and SAFR (Sao Francisco). Plecostomus jaguribensis (Supporting Information, Table S4, Fig. S1 ): Fowler 1915:264 , fig. 3 Type locality: Jaguaribe River in Barro Alto, Brazil . Holotype : ANSP 39930. Paratype : ANSP 39931. New synonym. Hypostomus jaguribensis : — Type catalogue: Böhlke 1984:123 ; Ferraris 2007:259 (checklist); Isbrücker 1980:32 (catalogue of species); Burgess 1989:432 (atlas of marine and freshwater species); Isbrücker 2001:28 (genus nomenclature); Isbrücker 2002:19 (genus nomenclature); Reis et al. 2003:362 (check* list); Armbruster 2004:80 (literature compilation); Zanata et al. 2013:247 (comparative material); Ramos et al. 2017:8 (com* parative material); Zawadzki et al. 2017a:571 (comparative material); Berbel*Filho et al. 2018 (checklist), Lima et al. 2017 (checklist). Plecostomus carvalhoi (Supporting Information, Table S5, Fig S2 ): Miranda Ribeiro 1937:54. Type locality: Granjeiro River, Crato, Ceará , Brazil . Lectotype : MNRJ 924A. Paralectotypes : MNRJ 924 (3). New synonym. Hypostomus carvalhoi : —Type catalogue: Miranda*Ribeiro 1955:401; Ferraris 2007:259 (checklist); Böhlke 1984:124 (catalogue of species); Miranda*Ribeiro 1953:401 ( lectotype designation); Zawadzki et al. 2017:12 (hypothesis that H. salgadae could be a junior synonym of H. carvalhoi ); Isbrücker 1980:21 (catalogue of species); Burgess 1989:431 (checklist); Isbrücker 2001:28 , 31 (genus nomenclature); Isbrücker 2002:19 (genus nomen* clature); Reis et al. 2003:362 (checklist); Armbruster 2004:80 (literature compilation); Zanata et al. 2013:247 (comparative material); Zawadzki et al. 2017:571 (comparative material); Zawadzki et al. 2017 and Zawadzki et al. 2019 (declare MNRJ 28870 (2) as syntypes ). Lima et al. 2017 , Berbel*Filho et al. 2018 (checklist). Figure 4. Maximum likelihood tree with phylogenetic relationships of the Hypostomus pusarum complex and Hypostomus aff. pusarum . Ŋe tree is associated with the occurrence of populations. On the right, a haplotype network for the COI molecular marker of the species of the H. pusarum complex and Hypostomus aff. pusarum . Each circle represents a haplotype and the size of the circles is proportional to the number of individuals that have the haplotype. Black dashes between circles indicate mutational steps. Abbreviations of the hydrographic basins: Ceará * Mirim (CEA), Jaguaribe (JAG), Granjeiro (GÞ), Choró (CHO), Trairí (TÞ), Salgado (SAL), São Francisco (SFR) and Parnaíba (PNB). Ecoregion abbreviations: north*east Caatinga and Coastal Drainage ( NCCD ), São Francisco (SAFR) and Parnaíba (PNBA). Plecostomus nudiventris (Supporting Information, Table S6, Fig. S3 ): Fowler 1941:147 , figs 44–46. Type locality: Choró River, Ceará , near Fortaleza, Brazil . Holotype : ANSP 69402. New synonym. Hypostomus nudiventris : — Type catalogue: Böhlke 1984:123 ; Ferraris 2007:259 (checklist); Isbrücker 1980:29 (catalogue of species); Burgess 1989:432 (checklist); Zanata et al. 2013:248 (comparative material); Zawadzki et al. 2017:12 (comparative material); Berbel*Filho et al. 2018 (checklist); Lima et al. 2017 (checklist). Ancistrus salgadae (Supporting Information, Fig. S4 ): Fowler 1941:154 , figs 56–58. Type locality: Salgado River, Icó, State of Ceará , Brazil , 6°25 ʹ S , 38°50 ʹ W . Holotype (single): ANSP 69440 (juvenile). New synonym. Hypostomus salgadae : — Type catalogue: Böhlke 1984:124 ; Ferraris 2007:300 (checklist); Isbrücker 1980:56 (catalogue of species); Burgess 1989:435 (checklist); Isbrücker 2002:12 (genus nomenclature); Isbrücker 2001:25 (genus nomencla* ture); Reis et al. 2003:318 (checklist); Ferraris 2007:300 (spe* cies inquirenda in Loricariidae ); Armbruster et al. 2015:99 (valid as Hypostomus salgadae ); Zawadzki et al. 2017:10 (pos* sibly a synonym of Hypostomus carvalhoi ); Berbel*Filho et al. 2018 (checklist); Lima et al. 2017 (checklist). New synonym. Plecostomus plecostomus papariae (Supporting Information, Table S7, Fig. S5 ): Fowler 1941:145 , figs 41–43. Type locality: Lago Papary, Rio Grande do Norte , Brazil . Holotype : ANSP 69398. Paratypes : ANSP 69399 (1), 69400*01 (2). New synonym. Hypostomus papariae : — Type catalogue: Böhlke 1984:124 ; Ferraris 2007:300 (checklist); Isbrücker 1980:29 (catalogue of species); Burgess 1989:432 (checklist); Isbrücker 2001:28 , 31 (genus nomenclature); Isbrücker 2002:19 (genus nomen* clature); Reis et al. 2003:361 (checklist); Armbruster 2004:79 (literature compilation); Zanata et al. 2013:247 (comparative material); Zawadzki et al. 2017:12 (comparative material); Ramos et al. 2017:8 (comparative material); Zawadzki et al. 2017:571 (comparative material); Berbel*Filho et al. 2018 (checklist); Lima et al. 2017 (checklist); Medeiros et al. 2019 (checklist). Figure 5. On the less, holotype of Plecostomus pusarum : CAS*Sº 22225, 142.6 mm SL. Ceará*Mirim River, Ceará*Mirim, Rio Grande do Norte, Brazil. Photo: Department of Ichthyology, California Academy of Sciences. On the right, topotype of Hypostomus pusarum : ºFRN 1127, 130.1 mm SL. Ceará*Mirim River, Estrela do Norte farm, between Ceará*Mirim and Taipu, Rio Grande do Norte. Diagnosis Hypostomus pusarum is distinguished from H. alatus , H. albopunctatus , H. arecuta , 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 byhavingdarkspotsonyellowish*brownbody (vs.light spots on dark body); 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 , and H. yaku by having less than 54 teeth in dentary and premaxilla (vs. over 65 in dentary and 60 in premaxilla). Hypostomus pusarum is distin* guished from species of the H. cochliodon group ( sensu Zawadzki and Hollanda*Carvalho 2014) by having slender viliform bi* cuspid teeth (vs. robust spoon*shaped teeth), and having a dental angle between 90° and 120° (vs. dental angles of 80° to 90° de* grees). It differs from H. affinis , H. ancistroides , H. aspilogaster , H. boulengeri , H. careopinnatus , H. carinatus , H. commersoni , H. delimai , H. dlouhyi , H. formosae , H. hemiurus , H. hoplonites , H. krikati , H. micromaculatus , H. niceforoi , H. nigrolineatus , H. piratatu , H. plecostomus , H. scabriceps , H. spiniger , H. tapijara , H. watwata , H. velhochico , and H. wilsoni for having moderate or in* conspicuous keels (vs. developed and conspicuous keels); from H. lamberti by having less than 12 dark spots on the first plate on dorsal series, medium to large spots on head and anterior portion of trunk from (vs. extremely small dark spots on head and anterior portion of trunk and more than 13 dark spots on the first plate in the dorsal series); from H. nigropunctatus by having a single pre* dorsal plate posteriorly bordering supraocipital (vs. three); from H. subcarinatus by having dorsal*fin colour background greyish* brownin life (vs. blue). Hypostomus pusarum differs from H. jaguar by having the lower first caudal*fin ray longer than the upper ray (vs. caudal rays of almost the same length); from H. garmani , H. guajupia , H. hermanni , H. latirostris , H. lima , and H. macrops by having moderate or inconspicuous keels (vs. lacking keels). Hypostomus pusarum is distinguished from H. brevis , H. kuarup , H. melanephelis , H. mutucae , H. nigromacutlatus , H. paulinus , and H. velhomonge , by having the abdomen almost entirely covered by plates, except at pelvic*fin insertions (vs. abdomen without plates); from H. bolivianus and H. perdido , by having bicuspid teeth (vs. unicuspid teeth); from H. yaku for lacking strongly* developed odontodes along keels on lateral series of plates (vs. strongly*developed odontodes), from H. albopunctatus , H. commersoni , H. derbyi , H. formosae , H. heraldoi , H. paulinus , H. jaguar , H. leucophaeus , H. nigromaculatus , and H. perdido , by the dorsal fin surpassing the adipose fin when depressed (vs. dorsal fin usually not reaching adipose fin). Hypostomus pusarum differs from H. garmani , H. guajupia , H. hermanni , H. latirostris , and H. lima by having rounded spots larger than the diameter of the eye (vs. spots smaller than the diameter of the eye in H. pusarum ); differs from H. vaillanti by having dark spots on the body (vs. dark vermiculations over the body). Figure 6. Hypostomus pusarum photographed in life immediately asser capture. A, ºFPB 7886, 183.4 mm SL, Ceará*Mirim River basin, Ceará* Mirim, Rio Grande do Norte, Brazil. B, ºFRN 0607, 120.1 mm SL, ºmbuzeiro River, tributary of the Jaguaribe River, Aiuaba, Ceará, Brazil. Description Meristic and morphometric data in Table 1 . Head and snout rounded in dorsal view, slightly triangular in some specimens. Eye moderate in size, positioned dorsolaterally; dorsal margin of orbit slightly elevated, interorbital region straight. Greatest body width at cleithrum region, gradually tapering to caudal fin. Highest region of body at insertion of first unbranched dorsal*fin ray. Mouth ventral, oral disc rounded with papillae present throughout lower lip, papillae larger closest to mouth, distally decreasing in size; internal median buccal papilla large, conspicuous and with visible villi; with rough appearance. Lower lip not reaching transverse line between gill openings. Dentaries angled between 90° and 120° to each other.Teeth bicuspid,with cusp larger than cusp, curved towards buccal cavity, with distal portion darker (yellowish) than tooth base (whitish).Barbel usually smaller than eye diameter. Table 1. Morphometric data and counts of Hypostomus pusarum . SL, Standard length. N = 150, SD, standard deviation.

Holotype	Range	Mean	SD
Standard length	142.6	56.8–206.2	131.2	28.8
Per cent of SL
Predorsal length	41.6	36.0–46.2	39.9	1.4
Head length	34.3	29.9–39.2	32.8	1.49
Cleithral width	34	22.4–36.1	31.5	1.42
Head depth	21.1	9.95–22.5	19.5	1.34
Interdorsal distance	17.2	11.1–20.4	15.2	1.84
Caudal peduncle length	26.9	26.2–37.0	30.4	1.51
Caudal peduncle depth	10.5	9.02–17.72	10.1	0.47
Dorsal*spine length	34.6	25.4–35.84	31.1	2.1
Pectoral*pelvic distance	25.1	20.2–39.0	23.5	1.93
Per cent of head length
Cleithral width	99.1	90.4–99.8	96.41	2.5
Head depth	61.6	55–66.5	59.7	2.5
Snout length	63.0	57.0–67.2	62.7	1.78
Orbital diameter	14.2	10.2–17.6	13.7	1.49
Interobital width	41.6	36.6–46.2	40.3	1.98
Mandibular width	15.9	10.8–20.5	15.4	1.96
Other per cents
Orbital diameter in snout length	22.0	15.9–30.9	21.8	2.69
Orbital diameter in interorbital length	34.3	25.5–44.0	34	4.01
Mandibular length in interobital length	38.2	27.1–48.0	38.6	5.5
Dorsal*spine length in predorsal length	83.3	65.0–90.8	79.2	5.70
First pectoral*fin ray length in predorsal length	82.6	66.5–92.5	80.1	4.61
Lower caudal*fin length in predorsal length	104.2	82.3–110.7	95.5	7.00
Adipose*fin length in caudal*peduncle depth	66.6	70.0–99.7	83.9	6.95
Caudal*peduncle depth in caudal*peduncle length	39	27.2–39.6	33.2	2.18
Interdorsal length in dorsal*fin base length	60.0	47.2–81.3	58.8	6.34
Lower lip width in lower lip length	32.1	20.0–42.6	29.1	3.27
Counts mode
Median plates series	26	25–27	26	0.35
Plates bordering supraoccipital	3	1–3	3	0.27
Predorsal plates	1	3	3	0
Dorsal plates below dorsal*fin bases	8	8–9	8	0.11
Plates between dorsal and adipose fins	7	5–8	6	0.57
Plates between adipose and caudal fins	4	2–5	4	0.51
Plates between anal and caudal fins	12	11–13	12	0.42
Premaxillary teeth	36	23–54	36	6.39
Dentary teeth	35	18–49	35	6.35

Dorsal and lateral surfaces of body covered with plates, ex* cept for small bare portion at tip of snout in some specimens, and across dorsal*fin base. Dorsal profile straight from snout to interorbital region, slightly concave from orbit to insertion of first dorsal*fin ray. Ventral profile straight from tip of snout to anal*fin, belly covered by minute plates in specimens over 90 mm SL, except region of insertion of pelvic and pectoral fins, portion below lower lip, and urogenital region; ventral re* gion with naked areas in some specimens smaller than 90 mm SL. Supraoccipital bone with small elevation forming incon* of spicuous keel on head. Lateral region of trunk with five rows slightly keeled plates and small odontodes, lateral plate keels visible, more easily observed in first row of plates close to dorsal fin. Pre*anal plate absent. Dorsal fin II, 7; extending beyond base of adipose fin when depressed, reaching base of adipose fin in some individuals. Second dorsal*fin ray slightly curved. Pectoral fin I, 6; first un* branched pectoral*fin ray slightly curved, widening towards to distal portion of ray; when depressed reaching half length of first pelvic*fin ray; odontodes of first unbranched pectoral*fin ray en* larged distally.Adipose fin developed and inserted five to seven plates asser posterior insertion of dorsal fin. Pelvic fin I, 5; first pelvic*fin ray tapered at base and distal portion, widened on medial portion; when depressed first ray reaching half anal*fin length. First unbranched adipose*fin ray compressed and slightly curved backwards. Caudal fin I, 14, I; dorsal lobe smaller than ventral lobe; unbranched. Colour in alcohol: Overall ground colour of dorsal and ventral re* gions of head, trunk and fins greyish*brown ( Fig. 5 ). First rays of pectoral and pelvic fins yellowish on distal portion. Spots, close to each other on dorsal portion of head. Spots more widely spaced and less conspicuous towards porterior portion of body. In some specimens, spots longitudinally arranged from plate where lateral line begins to end of caudal peduncle. Inter*radial region of pectoral, pelvic and anal fins with a single row of darker spots than on back in most specimens. Dorsal fin with one row of transverse spots on interradial area, darker in individuals up to 70 mm SL; usually one row of inconspicuous spots in individ* uals from 70 mm to 120 mm SL, and two rows of conspicuous spots in adult individuals (larger than 150 mm SL) ( Fig. 7 ). Paưern of spots ontogenetically variable. Ventral region light yellow, lighter than dorsal portions of body; ventral dark spots also lighter than others that cover dorsal surface of body. Region of abdomen usually with small spots (smaller than diameter of pupil), random and close to each other in most individuals. In some specimens, the spots are larger than the diameter of the pupil and more conspicuous ( Fig. 8 ). In some examples, dark spots are arranged linearly in the ventral region of the caudal peduncle ( Fig. 8B ). Colour in life: Colour paưern of living specimens similar to pre* served, except for darker brownish background and more con* spicuous spots ( Fig. 6 ). Iris operculum clearly visible. Some young individuals, when removed from the natural environment, present four trans* verse spots along length of dorsum, first close to insertion of first dorsal*fin ray, second at end of dorsal fin, third at begin* ning of adipose fin, and fourth on caudal peduncle (Supporting Information, Fig. S6 ). Some specimens with large dark spots on dorsum, while others with darker colours only on tips of caudal and dorsal fins ( Fig. 9 ). Seoual dimorphism: No sexual dimorphism was observed. Distribution and habitat: Hypostomus pusarum has been re* corded throughout the NCCD ecoregion, in the river basins of Paraíba do Meio ( Alagoas ), Acaraú, Choró, Cocó, Coreaú, Curu, Jaguaribe, Mundaú, Salgado ( Ceará ), Curimataú, Gramame, Mamanguape, Piranhas*Açu, Paraíba do Norte ( Paraíba ); Ipojuca ( Pernambuco ), Apodi*Mossoró, Catu, Ceará *Mirim, Doce, Piranhas*Açu, Potengi, Trairí, ( Rio Grande do Norte ). Specimens were also collected in the north tributaries of the São Francisco River ecoregion, in the Pajeú, Moxotó and Brígida River sub*basins ( Pernambuco ) and in the Parnaíba ecoregion, in the Poti River ( Ceará ) and Sambito River ( Piauí ). In the Parnaíba and São Francisco basins, the species was only found in Caatinga areas, while in the NCCD , a few sites are in Atlantic Forest coastal basins ( Fig. 10 ). Ecological notes and habitat: Hypostomus pusarum was found in environments with substrate composed of rocks and small pebbles, moderate current, and high clear water, as is the case of stretches where the species was collected in the Piranhas* Açu River. However, the species is also found in lentic environ* ments, with muddy substrate and high turbidity, such as dam environments. In the Ceará *Mirim River, type locality of the species ( Fig. 11 ), the environment is characterized as a perennial stretch of rapids, with depth ranging from 1.0 to 2.0 m, presence of mar* ginal vegetation, and rocky and sandy sediments. At this locality H. pusarum coexists with other species such as: Astyanao aff. bimaculatus (Linnaeus, 1758) , Cheirodon jaguaribensis Fowler, 1941 , Compsura heterura Eigenmann, 1915 , Hemigrammus marginatus Ellis, 1911 , H. rodwayi Durbin, 1909 , Serrapinnus heterodon (Eigenmann, 1915) , S. piaba (Lütken, 1875) , S. potiguar Jerep and Malabarba, 2014 from the family Characidae , Characidium bimaculatum Fowler, 1941 ( Crenuchidae ), Hoplias malabaricus (Bloch, 1794) ( Erythrinidae ), Leporinus piau Fowler, 1941 ( Anostomidae ), Steindachnerina notonota (Miranda*Ribeiro, 1937) ( Curimatidae ), Cichlasoma orientale Kullander 1983 , Saoatilia brasiliensis (Bloch, 1792) ( Cichlidae ), Eleotris pisonis (Gmelin, 1789) ( Eleotridae ), Poecilia vivipara Bloch and Schneider, 1801 ( Poecilidae ), and Synbranchus aff. marmoratus Bloch, 1795 ( Synbranchidae ). In the Jaguaribe and Salgado Rivers, it occurs in syntopy with H. sertanejo (Zawadzki et al ., 2017) . Individuals of the species were also collected in lentic envir* onments, with muddy sediment; some had macrophytes on the banks, while others did not. In the Argemiro de Figueiredo res* ervoir ( Paraíba ), for example, the sediment is muddy, with no macrophytes on its margin, in addition to intense anthropogenic impacts and low dissolved oxygen. In this ecosystem H. pusarum coexists with Apareiodon davisi Fowler, 1941 ( Parodontidae ), Prochilodus brevis Steindachnner, 1875 ( Prochilodontidae ), Triportheus signatus (Garman, 1890) ( Triportheidae ), H. malabaricus (Erytrinidae), Plagioscion squamosissimus (Heckel, 1840) ( Sciaenidae ), S. brasiliensis , Geophagus brasiliensis (Quoy and Gaimard, 1824) and Oreochromis niloticus (Linnaeus, 1758) ( Cichlidae ). Conservation status: Hypostomus pusarum was collected throughout the NCCD , in the states of Ceará , Rio Grande do Norte , Paraíba , Pernambuco and Alagoas , in the São Francisco ecoregion in the state of Pernambuco , and in the Parnaíba eco* region in the states of Ceará and Piauí , therefore, has a wide dis* tribution and is assessed as ‘Least Concern’ (LC) according to the International ºnion for Conservation of Nature (IºCN) categories and criteria (IºCN 2024). However, its geographic distribution, as well as the synonyms herein proposed should be updated. It is worth mentioning that the species was found in the Aiuaba Ecological Station, a protected area in the Jaguaribe River basin in Ceará ( Ávila et al . 2017 ). Common names: Bodó, Cascudo, Chupa*pedra, Limpa*vidro. Eoamined material: All from Brazil . Ceará *Mirim River– Rio Grande do Norte State : CAS 122225, 1, 142.6 mm SL, holo* type. CAS 122221, 4 ( 4 specimens measured from 26 inside the lot), 94.4–141.7 (48.6–141.7) mm SL, paratypes . NºP 14685, 10, 64.7–180.3 mm SL, Ceará *Mirim River, Jardim de Angicos, 05°39 ʹ 16.8 ʹʹ S , 35°58 ʹ 02.7 ʹʹ W . ºFRN 185, 15, 51.8* 133.9 mm SL, topotypes . ºFRN 842, 4, 65.9* 121.4 mm SL, Ceará *Mirim River, Ceará *Mirim, 05°38 ʹ 07 ʹʹ S , 35°24 ʹ 24 ʹʹ W , topotypes . ºFRN 563, 3, 56.6–70.7 mm SL, downstream of the ºsina São Francisco, Ceará *Mirim, 5°38 ʹ 7 ʹʹ S , 35°24 ʹ 24 ʹʹ W , topotypes . ºFRN 1063, 7, 76.0– 117.7 mm SL, Ceará *Mirim River, Ceará*Mirim, topotypes. ºFRN 1125, 5, 56.7–70.6 mm SL, Ceará *Mirim River, between Ceará *Mirim and Taipu, 5°37 ʹ 34.4 ʹʹ S , 35°32 ʹ 28.7 ʹʹ W . ºFRN 1127, 6, 66.2–130.4 mm SL, Ceará *Mirim River, between Ceará *Mirim and Taipu, 05°37 ʹ S , 35°33 ʹ W . ºFRN 119, 2, 120.1– 160.4 mm SL, Ceará *Mirim River. ºFPB 7701, 25, 98.2–177.6 mm SL, Ceará *Mirim River, Jardim de Angicos, 05°39 ʹ 17 ʹʹ S, 35°58 ʹ 05 ʹʹ W. ºFRN 1126, 2, 85.8–88.5 mm SL, Ceará *Mirim River, between Ceará *Mirim and Taipu, 05°37 ʹ S , 35°33 ʹ W . ºFRN 118, 2, 60.5–103.5 mm SL, Ceará *Mirim River. ºFRN 5654, 5, 36,97–74,20 mm SL, Ceará * Mirim River, 05°38 ʹ 07.4 ʹʹ S , 35°24 ʹ 24.8 ʹʹ W . Figure 7. Ontogenetic development of interradial spots on the dorsal fin of Hypostomus pusarum (A) ºFRN 0255, 62.6 mm SL, Seridó River, upstream of Ilha de Santana, Caicó, Rio Grande do Norte, Brazil, (B) ºFRN 1127, 116.9 mm SL, Ceará*Mirim River, between Ceará*Mirim and Taipu, Rio Grande do Norte, Brazil, and (C) ºFPB 7701, 170.4 mm SL, Ceará*Mirim River, Jardim de Angicos, Rio Grande do Norte, Brazil. Figure 8. Intraspecific variation of Hypostomus pusarum . A, lighter, smaller, and closely spaced spots in ventral region, 123.2 mm SL. B, darker, larger, and more widely spaced spots, 117.4 mm SL. ºFRN 608, ºmbuzeiro River, Jaguaribe River basin, São Nicolau, Ceará, Brazil. Figure 9. Intraspecific variation between individuals of Hypostomus pusarum collected in the same locality. A, dark spots on dorsum, 78.5 mm SL. B, no dark spots on body, spots only on tips of dorsal and caudal fins, 70.1 mm SL. ºFRN 5643, Lagoa de Remígio Dam, Remígio, Paraíba, Brazil. Jaguaribe River— Ceará State : ºFRN 0607, 4, 189.30– 202.15 mm SL, ºmbuzeiro River dam, at Ecological station of Aiuaba, Aiuaba, 06°36 ʹ 06.8 ʹʹ S , 40°07 ʹ 28.6 ʹʹ W . ºFPB 9222, 1, 113.6 mm SL, Oiticica reservoir, village Lagoa da Mata, Itatira, 04°40 ʹ 45 ʹʹ S , 39°41 ʹ 38 ʹʹ W . ºFRN 1256, 1, 17.4 mm SL, Jaguaribe River, Saboeiro, 06°32 ʹ 31 ʹʹ S , 39°54 ʹ 37 ʹʹ W . ºFRN 607, 4, 186.9– 198.5 mm SL, ºmbuzeiro River, tributary of the Jaguaribe River, Aiuaba, 06°36 ʹ 06 ʹʹ S , 40°07 ʹ 28 ʹʹ W . ºFRN 608, 8, 104.9– 183.8 mm SL, ºmbuzeiro River, downstream the dam of São Nicolau, São Nicolau, 06°40 ʹ 48 ʹʹ S , 40°04 ʹ 49 ʹʹ W . ºFPB 9226, 1, 142.6 mm SL, Bom Sucesso stream, Jaguaribe basin, Piquet Carneiro, 05°45 ʹ 57 ʹʹ S , 39°25 ʹ 54 ʹʹ W . ºFPB 7699, 1, 101.9 mm SL, Santana reservoir, Jaguaribe, 05°48 ʹ 29 ʹʹ S , 038°39 ʹ 09 ʹʹ W . ºFRN 1811, 3, 94.8–104.0 mm SL, Cruzado stream, Lavras da Mangabeira, 06°47 ʹ 08 ʹʹ S , 38°58 ʹ 34 ʹʹ W .ºFRN 1843, 1, 180.1 mm SL, ºmbuzeiro River, downstream the dam São Nicolau, São Nicolau, 06°40 ʹ 48 ʹʹ S , 40°04 ʹ 49 ʹʹ W . ºFRN 609, 2, 130.0– 134.0 mm SL, ºmbuzeiro River, downstream the dam São Nicolau, São Nicolau, 06°40 ʹ 48 ʹʹ S , 40°04 ʹ 49 ʹʹ W . Salgado River— Ceará State : ºFRN 5076, 4, 79.3–124.3 mm SL, Salgado River, Icó, 06°24 ʹ 53.8 ʹʹ S , 38°52 ʹ 23 ʹʹ W , topotypes . ºFPB 7700, 6, 81.0– 100.7 mm SL, Salgado River, Icó, 06°24 ʹ 28 ʹʹ S , 38°52 ʹ 06 ʹʹ W . ºFPB 9225, 16, 50.3–86.3 mm SL, Machado stream, tributary of the Salgado River, Lavras de Mangabeira, 06°42 ʹ 25 ʹʹ S , 39°01 ʹ 40 ʹʹ W . ºFRN 1843, 48, 17.3–120.3, Salgado River, Icó, 06°24 ʹ 53 ʹʹ S , 38°52 ʹ 23 ʹʹ W , topotypes . ºFRN 5061, 4, 58,00–78,30 mm SL, Salgado River, 06°24 ʹ 53.8 ʹʹ S , 38°52 ʹ 23 ʹʹ W . ºFRN 1813, 1, 75.31 mm SL, Salgado River, Icó, 06°24 ʹ 53.8 ʹʹ S , 38°52 ʹ 23 ʹʹ W . Granjeiro River— Ceará State : ºFRN 2017, 109, 100–124.35 mm SL, Granjeiro River, Crato, 07°15 ʹ 18.7 ʹʹ S , 39°25 ʹ 58.9 ʹʹ W , topotypes , Granjeiro River, Crato, 07°15 ʹ 18 ʹʹ S , 39°25 ʹ 58 ʹʹ W , topotypes . ºFRN 2189, 1, 54.4 mm SL, Granjeiro River, Crato, 07°15 ʹ 18 ʹʹ S , 39°25 ʹ 58 ʹʹ W , topotypes . Choró River— Ceará State : ºFRN 4888, 2, 129.6–146,0 mm SL, Choró River, Chorozinho, 04°18 ʹ 13.5 ʹʹ S , 38°29 ʹ 55.4 ʹʹ W , topotypes . ºFRN 2901, 22, 48.6–106.4 mm SL, Choró River, Chorozinho, 04°18 ʹ 13 ʹʹ S , 38°29 ʹ 55 ʹʹ W , topotypes . ºFPB 7697, 3, 78.3–118.4 mm SL, Chorozinho, Choró River, 04°19 ʹ 22.2 ʹʹ S , 38°30 ʹ 31.7 ʹʹ W . ºFPB 9223, 9, 61.8–74.5 mm SL, Choró River, Itapiúna. Trairí River— Rio Grande do Norte State : ANSP 69398, 1, 94.3 mm SL, holotype . ANSP 69399, 1, 99.1 mm SL, paratype , collected along with the holotype , lago Papari. ºFPB 7693, 32, 44.5–59.3 mm SL, Ariri River, Nísea Floresta, 06°06 ʹ 34 ʹʹ S , 35°12 ʹ 09 ʹʹ W . ºFRN 2399, 2, 14.8* 19.1 mm SL, Trairí River, na BR * 101 in the São José de Mipibú, São José de Mipibú. NºP 14684, 10, 54.6–104.4 mm SL, Ariri River, Nísia Floresta. ºFRN 2304, 26, 98.9–183.5 mm SL, downstream Açude Novo, Santa Cruz, 06°13 ʹ 35.4 ʹʹ S , 36°01 ʹ 38.3 ʹʹ W . Piranhas-Açu River— Rio Grande do Norte State : ºFRN 0311, 31, 14.0– 44.5 mm SL, Piranhas*Açu River, Jardim de Piranhas, 06°22 ʹ 40.3 ʹʹ S , 37°21 ʹ 19.3 ʹʹ W . ºFRN 1378, 5, 73.80– 112.6 mm SL, Espinharas River reservoir, Serra Negra do Norte, 06°40 ʹ 11.7 ʹʹ S , 37°23 ʹ 13.5 ʹʹ W . ºFRN 1731, 5, 62.9* 88.4 mm SL, Piranhas*Açu River, Jardim de Piranhas, 06°22 ʹ 40 ʹʹ S , 37°21 ʹ 19 ʹʹ W. ºFRN 585, 3, 62.6–85.7 mm SL, Piranhas*Açu River, Jardim de Piranhas, 06°22 ʹ 40 ʹʹ S, 37°21 ʹ 19 ʹʹ W. ºFPB 9684, 1, 153.0 mm SL, Cipó stream, Sitio Cipó, São João do Sabugi, 06°38 ʹ 39 ʹʹ S , 37°10 ʹ 30 ʹʹ W . NºP 4795, 11, 140.0–207.0 mm SL, Acauã River, tributary of the Piranhas*Açu River. NºP 14683, 2, 103.1–135.0 mm SL, Piranhas*Açu River. Piranhas-Açu River— Paraíba State : ºFRN 1384, 4, 50.0– 101.7 mm SL, Piranhas River, village of Piranhas Velho, São José do Piranhas, 07°06 ʹ 18 ʹʹ S , 38°29 ʹ 28 ʹʹ W . ºFRN 1867, 2, 101.6– 91.6 mm SL, Pereiro River, tributary of Piranhas*Açu River, Bonito de Santa Fé, 07°16 ʹ 12 ʹʹ S , 38°30 ʹ 34 ʹʹ W . ºFRN 584, 26, 73.0– 150.1 mm SL, Piranhas*Açu River, São Bento, 06°28 ʹ 48 ʹʹ S , 37°26 ʹ 44 ʹʹ W . ºFPB 2919, 4, 34.6–45.4 mm SL, Espinharas River, tributary of Piranhas*Açu River, Açude Jatobá, Patos. NºP 14683, 2, 103.1– 135.1 mm SL, Piranhas*Açu, Aparecida, 06°48 ʹ 42.8 ʹʹ S , 38°4 ʹ 24.6 ʹʹ W . Paraíba do Norte River— Paraíba State : ºFRN 0730, 10, 126.4– 162.4 mm SL, Paraíba do Norte River, down* stream of the Epitácio Pessoa dam, Boqueirão, 07°29 ʹ 11.9 ʹʹ S 36°07 ʹ 24.5 ʹʹ W . ºFPB 6053, 2, 142.0–172.0 mm SL, Soledade reservoir, 7°04 ʹ 54 ʹʹ S , 36°22 ʹ 10 ʹʹ W . ºFPB 6054, 5, 97.7– 144.6 mm SL, Soledade reservoir, Soledade, 07°04 ʹ 54 ʹʹ S , 36°22 ʹ 10 ʹʹ W . ºFPB 6026, 2, 92.19–94.3 mm SL, Taperoá reser* voir, Taperoá River, tributary of Paraíba do Norte River, Taperoá, 07°13 ʹ 44 ʹʹ S , 36°50 ʹ 09 ʹʹ W . ºFPB 3661, 1, 144.9 mm SL, Avelós River, Taperoá River tributary of the Paraíba do Norte River, São João do Cariri. ºFRN 729, 1, 106.1 mm SL, Paraíba do Norte River, downstream of the Epitácio Pessoa reservoir, Boqueirão, 07°29 ʹ 11 ʹʹ S , 36°7 ʹ 24 ʹʹ W . ºFRN 434, 3, 53–60.5 mm SL, Paraíba do Norte River, in the BR 104, Barra de Santana, 07°31 ʹ 45 ʹʹ S , 35°59 ʹ 53 ʹʹ W . ºFRN 4148, 36, 73.0– 138.2 mm SL, Paraíba do Norte River, BR *104, Barra de Santana, 07°31 ʹ 45 ʹʹ S , 35°59 ʹ 55 ʹʹ W . ºFRN 475, 4, 22.8*52.9, Paraíba do Norte River, Cruz do Espírito Santo , 7°8 ʹ 57 ʹʹ S , 35°7 ʹ 17 ʹʹ W . ºFRN 1861, 2, 123.1– 142.6, Piancó River, Ibiara, 07°19 ʹ 15 ʹʹ S , 38°08 ʹ 57 ʹʹ W . ApodiMossoró River— Rio Grande do Norte State : ºFRN 1818, 6, 88.2–140.4 mm SL, Apodi*Mossoró River, downstream of the Santa Cruz dam, Apodi, 05°39 ʹ 29.5 ʹʹ S , 37°46 ʹ 17.5 ʹʹ W . ºFRN 1803, 2, 106.0– 129.1 mm SL, Apodi*Mossoró River, down* stream of the Santa Cruz reservoir, Apodi, 05°39 ʹ 29 ʹʹ S , 37°46 ʹ 17 ʹʹ W . ºFRN 3511, 2, 38.5–41.5 mm SL, Apodi*Mossoró River, São Francisco do Oeste, 05°59 ʹ 54 ʹʹ S , 38°09 ʹ 43 ʹʹ W . ºFRN 3490, 1, 40.7 mm SL, Apodi*Mossoró River, José da Penha, 06°22 ʹ 14 ʹʹ S , 38°17 ʹ 28 ʹʹ W . ºFRN 1635, 3, 60.6–67.6 mm SL, Apodi*Mossoró River, downstream Santa Cruz reservoir, Apodi, 05°39 ʹ 29 ʹʹ S , 37°46 ʹ 17 ʹʹ W . ºFRN 3444, 4, 78.8–90.6 mm SL, Apodi*Mossoró River, Mossoró, 05°17 ʹ 21 ʹʹ S , 37°23 ʹ 08 ʹʹ W . NºP 6177, 2, 92–152 mm SL, Apodi*Mossoró River, Mossoro, 05°13 ʹ 37.8 ʹʹ S , 37°21 ʹ 48.8 ʹʹ W . Gramame River— Paraíba State : ºFPB 7329, 2, 99.9–177.8 mm SL, Gramame River, Vascecada, Pedras de Fogo, 07°19 ʹ 31 ʹʹ S , 34°59 ʹ 32 ʹʹ W . ºFRN 7328, 2, 158.5– 172.1 mm SL, Gramame River, Vascecada, Pedras de Fogo, 07°19 ʹ 31 ʹʹ S , 34°59 ʹ 32 ʹʹ W . ºFPB 6702, 3, 116.7– 142.9 mm SL, Gramame River, João Pessoa. ºFPB 7328, 2, 158.5– 172.1 mm SL, Gramame River, Vascecada, Pedras de Fogo, 07°19 ʹ 31 ʹʹ S , 34°59 ʹ 32 ʹʹ W . Sambito River— Piauí State : ºFRN 11160, 2, 123.3– 149.2 mm SL, Sambito River, Sítio Miguel Alves, Aroazes, 06°08 ʹ 27.6 ʹʹ S , 042°02 ʹ 40.6 ʹʹ W . Poti River— Ceará State : ºFPB 9265, 8, 74.1–150.0 mm SL, Poti River, Crateús, 05°03 ʹ 27 ʹʹ S , 40°50 ʹ 13 ʹʹ W . ºFPB 7443, 5, 42.2– 102.0mmSL,PotiRiver,Ibiapaba, 05°02 ʹ 43.9 ʹʹ S , 040°55 ʹ 19.8 ʹʹ W . ºFPB 11566, 10, 122.6– 172.8 mm SL, Poti River, Crateús. São Francisco River— Pernambuco State : ºFRN 0725, 3, 73.1– 96.5 mm SL, Serrinha stream, downstream Serrinha dam, Serra Talhada, 08°12 ʹ 41.4 ʹʹ S , 38°32 ʹ 04.2 ʹʹ W . ºFRN 0727, 10, 89.6– 134.7 mm SL, Pajeú River, downstream of the Jazigo dam, Serra Talhada, 08°00 ʹ 01.9 ʹʹ S , 38°14 ʹ 41.2 ʹʹ W . NºP 19224, 4, 139.4– 179.9 mm SL, Capibaribe River, Recife, 08°01 ʹ 37.9 ʹʹ S , 34°56 ʹ 38.0 ʹʹ W . NºP 4920, 6, 112.2– 186.2 mm SL, Acauã River (Marechal Dutra dam), Acari, 06°26 ʹ 11 ʹʹ S , 36°38 ʹ 28 ʹʹ W . NºP 13974, 2, 197.5– 221.7 mm SL, Itaparica reservoir, Petrolandia, 08°59 ʹ 24.0 ʹʹ S , 38°13 ʹ 50.0 ʹʹ W . NºP 5250, 5, 143.0–175.0 mm SL, Lagoon of Piató, Açu. NºP 4796, 7, 157.5– 181.7 mm SL, Acauã River (Marechal Dutra dam), Açu, 06°26 ʹ 11 ʹʹ S , 36°38 ʹ 28.0 ʹʹ W . NºP 13973, 1, 188.0 mm SL, Itaparica reservoir, Belém de São Francisco, 08°47 ʹ 36.7 ʹʹ S , 38°57 ʹ 14.68 ʹʹ W . NºP 4795, 11, 135.4– 205.7 mm SL, Acauã River (Marechal Dutra dam), Acari, 06°26 ʹ 11 ʹʹ S 36°38 ʹ 28.00 ʹʹ W . Ipojuca River— Pernambuco State : ºFPB 11373, 1, tributary of the Ipojuca River, Escada, 08°22 ʹ 45.8 ʹʹ S , 35°17 ʹ 24.7 ʹʹ S . Paraíba do Meio River—Alagoas State : ºFRN 3889, 17, Paraíba do Meio River, upstream of Quebrângulo, Quebrângulo, 09°18 ʹ 23.1 ʹʹ S, 36°28 ʹ 48.4 ʹʹ W . ºFRN 3867, 3, Paraíba do Meio River, Quebrângulo, 09°18 ʹ 45.4 ʹʹ S , 36°28 ʹ 04.7 ʹʹ W . Figure 10. Distribution of Hypostomus pusarum in the three ecoregions: Northeastern Caatinga and Coastal Drainages (NCCD), São Francisco (SAFR) and Parnaíba (PNBA). Figure 11. Type locality of Hypostomus pusarum , Ceará*Mirim River, Ceará*Mirim, Rio Grande do Norte, Brazil. Remarks In the original description of Hypostomus pusarum , the only species compared was H. plecostomus (Linnaeus, 1758) from Suriname ( Starks 1913 ). It worth noting that important taxo* nomic information on Hypostomus are the dentary and pre* maxillary teeth counts, which were not originally indicated. In its description, Starks (1913) analysed individuals from 150 to 200 mm SL collected in the Ceará*Mirim River, while the pre* sent redescription was based on a wide sampling, including the type series and material from several basins of the NCCD , Parnaíba, and São Francisco River basins, ranging from 90 to 207 mm SL, covering ontogenetic and population variation. Fowler (1915) described H. jaguribensis from the Jaguaribe River in Ceará State and compared it to H. unae (Steindachner, 1878) and H. wuchereri (Günther, 1864) from Bahia , H. aurogutatus Kner, 1854 from the Paraíba do Sul Basin, H. ancistroides (Ihering, 1911) from the Piracicaba River, H. albopunctatus (Regan, 1908) and H. variipictus (Ihering, 1911) from the Pardo River in São Paulo , but did not mention H. pusarum . Fowler (1915) pointed out morphological and meristic data in the type series of H. jaguribensis similar to those reported by Starks (1913) for H. pusarum , such as the number of plates in the median series (25 and 25–26), predorsal plates (three), plated belly, lower lobe of the caudal fin larger than the upper lobe, and the same colour paưern in both species. Fowler (1915) also described that in H. jaguribensis the spots on the inter*radial membrane of the dorsal fin appear in a single vertical row, but this may occasionally be irregular. Our results showed that in the material from the type locality of H. jaguribensis , individuals larger than 170 mm SL also have two transverse rows, as indicated in H. pusarum ( Starks, 1913 ) . Ŋis characteristic can be considered ontogenetic variation in which individuals up to 70 mm SL have only a single row of transverse spots on the inter*radial membrane, and from approximately 120 mm the spots become less conspicuous, and in some larger individuals they even separate into two rows of spots ( Fig. 7 ). Ŋis ontogenetic paưern has already been reported in the devel* opment of Isorineloricaria acuarius Ray and Armbruster, 2016 (Ray and Armbruster 2016). Ŋis fact may explain differences in the spots on the dorsal fin found by the authors who described the species of Hypostomus from Northeastern Brazil at the begin* ning of the 20th century, since few individuals and of different sizes were used. Alípio de Miranda Ribeiro (1937) described several species of vertebrates, among them, Hypostomus carvalhoi from the Granjeiro River, also in the Jaguaribe Basin in Ceará , based on four specimens and presented a short description. Ŋe meristic data of H. carvalhoi overlaps with those of the species previously described by Starks (1913) and Fowler (1915) , as well as the morphological and colour characters, such as dark spots more concentrated on the head, weakly keeled plates, thorax and ab* domen covered by small plates, and length of the depressed dorsal fin reaching to the adipose fin. Also in 1937, the naturalist Rodolpho von Ihering sent ma* terial collected by him in the drainages of the states of Piauí, Ceará, Rio Grande do Norte, Paraíba, and Pernambuco to the Philadelphia Academy of Natural Sciences. Ŋis material was analysed by Fowler (1941) who described several species, but neglected some important data, such as the type localities and coloration of the species. Among the armoured catfishes de* scribed by Fowler of the H. pusarum complex are H. papariae , H. nudiventris , and H. salgadae . Hypostomus nudiventris from the Choró River near Fortaleza and H. salgadae from the Salgado River (Jaguaribe Basin) in Icó, both from Ceará , have the same number of predorsal plates, number of teeth (10–12), and ab* domen without plates. In addition, diagnoses that could separate the two species were not provided, probably because, Fowler de* scribed, these two species in different genera. Ŋese species were described based only on the holotypes that present reduced sizes, H. salgadae with 29 mm , and H. nudiventris with 82 mm , in total length ( Fowler 1941 ). Many species of Hypostomus that have a plated belly only de* velop plates asser reaching a certain size, such as 80 mm SL in H. velhochico and 110 mm SL in H. faveolus Zawadzki et al ., 2008 (Zawadzki et al. 2008, Zawadzki et al. 2017, Lustosa*Costa et al. 2022). Based on our analyses, we could observe that some speci* mens of H. pusarum possess a completely plated belly (with the exception of the insertion area of the pectoral and pelvic fins, and urogenital papilla) only asser reaching 90 mm SL, which explains the fact that Fowler (1941) did not describe plates for the holo* types of H. nudiventris and H. salgadae , once as the specimens had not yet developed the ventral plates. In the description of H. salgadae , in addition to the plateless abdomen, Fowler mentioned a ‘wide frontal marginal area of the head’, which probably led him to describe it as Ancistrus . Fowler (1941:155) , also described the presence of four spots on the body of the holotype as ‘four dark blotches on body, first below front of dorsal, second below hind part of dorsal, third from adi* pose fin and last at caudal base’, treating that as a diagnostic fea* ture. However, based on our analysis of juveniles of H. pusarum collected in the Ceará *Mirim River and kept in an aquarium, the same spot paưern described by Fowler (1941) in H. salgadae was verified (Supporting Information, Fig. S6 ). Ŋis coloration can also be observed in some fixed individuals and has already been indicated as originated by stress in H. plecostomus , H. boulengeri (Eigenmann and Kennedy, 1903) , H. ancistroides , and H. faveolus (Zawadzki et al. 2008, 2010). Hypostomus papariae , was also described by Fowler (1941) from the Papary Lake, in Rio Grande do Norte State , and the type series is composed of three lots ( holotype and two of paratypes ). However, one lot of paratypes was collected in the Choró River (type locality of H. nudiventris ), without any comparison with this species in the description of H. papariae . Ŋe author then as* sumed that there were two different species in the Choró River, one with plates on the belly ( H. papariae ) and one without plates ( H. nudiventris ). However, H. papariae shows the same paưern of double spots on dorsal fin, and dorsal fin reaching to the adipose fin, which is observed in the illustration of Fowler (1941) . As in other descriptions of species from north*eastern Brazil , Fowler (1915 , 1941 ) did not cite Starks (1913) , and it is possible that he was not aware of it. Some characters of H. papariae , such as the two dark spots on the adipose fin, and rows of dark spots on pectoral, anal, pelvic, and caudal fins were not observed in the drawing made by Fowler (1941), or in topotypes, but, more conspicuous round spots were observed on the fins of some specimens collected in several basins. In recent collections in the Papary Lake, fresh* water fishes, like Hypostomus , were not found ( Medeiros et al. 2019 ). Ŋis lake is in the coastal portion of the Trairí Basin, sug* gesting that the type locality of Lake Papary in Fowler (1941) is a mistake ( Medeiros et al. 2019 ). Zawadzki et al. (2017) have proposed the synonymy of H. salgadae to H. carvalhoi . Now, through the analyses carried out in the present study, we concluded that both are junior synonyms of H. pusarum , the first described species of the complex, as well as H. jaguribensis , H. papariae , and H. nudiventris .