The rediscovery of Pimelodella longipinnis (Borodin, 1927), an enigmatic Atlantic Rainforest catfish species from Southeastern Brazil (Siluriformes: Heptapteridae)

This article is a redescription of Pimelodella longipinnis, an enigmatic catfish previously known only from its holotype and with uncertain type locality. The species is redescribed based on recently collected materials from streams of the Mata Atlântica bioregion, in Santos municipality, São Paulo State, Brazil. Pimelodella longipinnis is assigned to a putatively monophyletic group, the Pimelodella leptosoma-group, diagnosed by the presence of a supraoccipital process not reaching the anterior nuchal plate, with a gap of ca. 20-25% of the supraoccipital process total length, and whose tip notably surpasses the midpoint of the complex vertebra in dorsal view. We also present a list of fish species described from a shipping sent to the American Museum of Natural History from the former Museu Paulista (now Museu de Zoologia da Universidade de São Paulo), of which P. longipinnis was part.


INTRODUCTION
Pimelodella Eigenmann, 1888 is a genus of Neotropical catfish family Heptapteridae distributed throughout cis-and trans-Andean drainages. With 82 species currently considered valid (Fricke et al., 2021), a conservative morphology and putatively wide distributions, the genus currently represents one of the most difficult bottlenecks for understanding the diversity of Neotropical catfishes. In addition, Pimelodella still lacks comprehensive taxonomic or phylogenetic studies, which result in great difficulty in diagnosing several individuals found in scientific collections (Slobodian et al., 2017), even those collected in well-sampled locations.
Currently, the genus Pimelodella can only be diagnosed from other Heptapteridae by a unique character combination. These include small to medium-size catfishes, usually between 12 and 30 cm of standard length [although some species, e.g., Pimelodella cristata (Müller & Troschel, 1849), may exceed 30 cm]; supraoccipital process long, usually reaching the anterior nuchal plate; anterior and posterior fontanels open, elongated, separated by an epiphyseal bar; eyes well-delimited by a free orbital rim, especially pronounced anteriorly and dorsally; pectoral fins with one unbranched, rigid and pungent spinous ray at leading edge, bearing usually both anterior denticulations and posterior serrae, followed by 7-9 (usually 8) branched rays; branchiostegal rays usually 6; caudal fin deeply forked; median caudal-fin rays not articulated to hypural plates; hypural 5 autogenous, but contacting the hypural plate; body generally with a dark midlateral stripe extending from the snout or just posterior to the opercle to the insertion of, or onto, median caudal-fin rays (Slobodian et al., 2017).
Pimelodella species are distributed throughout the Central and South America, occurring in all the major basins, including the Amazonas, Orinoco, Paraná-Paraguay, São Francisco, and Atlantic Coastal drainages, with the Amazon basin harboring its highest diversity (Slobodian, 2017). The Mata Atlântica and Upper Paraná bioregions (sensu Dagosta et al., 2020) are among the most well-sampled and intensively studied Neotropical areas, which is possibly influenced by their proximity to large urban centers (Agostinho et al., 2007;Dagosta et al., 2020). However, the closeness of these drainages to large metropolitan centers, such as São Paulo and Rio de Janeiro, has historically posed a series of risks to fishes inhabiting these areas. Currently, these taxa are among the most threatened due to habitat destruction, introduction of non-native species, pollution, among other anthropic deleterious activities (Agostinho et al., 2007;Silva et al., 2020).
Recent ichthyological expeditions to the South Atlantic coastal streams in the State of São Paulo yielded a distinctive species of Pimelodella, which revealed to fit the diagnosis of Pimelodella longipinnis. This species was not included in previous reviews of South Atlantic coastal stream fishes (e.g., Guazzelli, 1997;Menezes et al., 2007), being this the first work to offer a more precise distribution range to P. longipinnis, as well as a redescription of the species.

MATERIAL AND METHODS
Measurements are point-to-point distances taken with a digital caliper under a dissecting scope, following Slobodian & Pastana (2018). Body measurements are presented as proportions of Standard Length (SL) and head parts are presented as proportions of Head Length (HL) (except for measurements of barbels, which are offered as proportions of SL). Meristics and descriptions of fin position follow Bockmann & Castro (2010). Vertebral counts include the Weberian complex elements counted as five, plus all free vertebrae and the compound caudal centrum (PU1+U1) counted as one (sensu Lundberg & Baskin, 1969). Number of specimens with each meristic value is presented in parentheses, followed by an asterisk indicating values for the holotype when a range is presented.
Clearing and staining protocol follow Taylor & Van Dyke (1985), with the prepared specimens indicated by "c&s". Osteological data was also obtained with aid of X-ray images of the holotype and five comparative specimens, which are indicated by "xr". Osteological and cephalic laterosensory canals terminology follows primarily Bockmann & Miquelarena (2008), with modifications proposed by Pastana et al. (2020). Nomenclature for pectoral-fin and dorsal-fin ornamentations follow Slobodian & Pastana (2018), with modifications of Kubicek et al. (2019). Fading coloration marks on holotype were observed with the aid of an ultraviolet light lantern (Netto-Ferreira & Luckenbill, 2017). Food preference was investigated using the stomach contents of six specimens, obtained by stomach dissection. In situ behavior was registered with a stationary camera Panasonic Lumix DMC-TS20 on three occasions, in four locations, during 15 minutes in the afternoon. Categorization of observed behavior follows Wilson & Roys (1994) and Morgan & Fine (2020).
Institutional abbreviations follow Sabaj (2020). Maps were produced using Google Earth Web and Quantum GIS 2.18, and edited in Adobe Photoshop CC 2019. Photos were edited in Adobe Photoshop CC 2019. Illustrations of pectoral-fin spines were produced with Adobe Illustrator CC 2019, based on photographs and direct observation. Specimens were preserved in 70% ethanol unless indicated as cleared and stained (c&s), which are preserved in glycerol. Comparative material of Pimelodella species is as presented in Slobodian et al. (2017), with additional specimens listed in the "Comparative material" section.

RESULTS
A thorough revision of Pimelodella specimens from the main scientific collections in Brazil and abroad by the first author revealed the occurrence of an enigmatic Pimelodella species occurring in three collection sites from Rio Quilombo and Rio Jurubatuba (Santos municipality, São Paulo State, Brazil). Such specimens were not congruent neither with Pimelodella species previously attributed to the Mata Atlântica in recent reviews of the region (e.g., Menezes et al., 2007) nor with those reported on the unpublished dissertation on Pimelodella species from coastal rivers in Southern and Southeastern Brazil (Guazzelli, 1997).
Upon comparison with the types of Pimelodella species from Mata Atlântica and Upper Paraná bioregions, the recently collected material was found to be undistinguishable from Pimelodella longipinnis (Borodin, 1927a). Morphometric data from specimens from the Rio Quilombo and Rio Jurubatuba, altogether with other morphological characteristics (see below), are strikingly congruent with data collected from the holotype of P. longipinnis (AMNH 8642) (Table 1). These fishes share relatively short maxillary barbels, reaching between verticals through pelvic-fin origin and anal-fin terminus (38.9-64.6% SL), dorsal-fin first branched ray somewhat short (10.6-22.5% SL), and adipose-fin length three to four times in SL (14.6-34.8% SL).
In addition, radiographs of the specimens examined from Rio Quilombo and Rio Jurubatuba reveal that their supraoccipital process do not reach the anterior prenuchal plate. Among the Pimelodella species from the Mata Atântica and Upper Paraná, this condition is shared only with P. itapicuruensis and P. longipinnis. Nevertheless, the type specimens of Pimelodella itapicuruensis (FMNH 57951,57953,57986,57987,and 57988) have maxillary barbels usually surpassing anal-fin terminus (65.3-87.5% SL), first dorsal-fin branched ray (the longest) 20.9-26.2% SL, and adipose-fin length roughly three times in SL (32.1-38.6% SL), all characteristics that do not match the specimens from Rio Quilombo and Rio Jurubatuba. In fact, these characteristics are highlighted by Eigenmann (1917) as being diagnostic of P. itapicuruensis in its original description (Eigenmann, 1917: p. 247-248, pl. XXXI, fig. 3).
Further evidence pointing that the material from from Rio Quilombo and Rio Jurubatuba represent Pimelodella longipinnis comes from its color pattern. When observed under ultraviolet light (following Netto-Ferreira & Luckenbill, 2017), the holotype of P. longipinnis revealed to have some fading coloration marks, despite no marks being presented in the original description (Borodin, 1927a). This fish has a poorly-defined, dark brown midlateral stripe extending from orbit to caudal fin origin, altogether with a poorly-defined darker region near dorsal fin base. These marks match perfectly the color pattern observed in the specimens from Rio Quilombo and Rio Jurubatuba.
In sum, based on the strikingly morphological similarities between the holotype of P. longipinnis and the recently collected specimens from Rio Quilombo and Rio Jurubatuba, we conclude these last belong to P. longipinnis. Therefore, this species is redescribed herein in light of a detailed examination of the holotype and additional 63 recently-collected specimens. An updated geographic range is provided for Pimelodella longipinnis, which now includes South Atlantic coastal streams of the São Paulo State. (Borodin, 1927) Figs. 1-4; Table 1 Synonymy Rhamdella longipinnis Borodin, 1927a: 6-7

Diagnosis
Pimelodella longipinnis differs from all its congeners, except P. bockmanni Slobodian & Pastana, 2018; P. itapicuruensis; P. leptosoma (Fowler, 1914); P. megalura Miranda Ribeiro, 1918;P. metae Eigenmann, 1917;P. montana Allen, 1942;P. peruensis Fowler, 1915, P. robinsoni, P. tapatapae Eigenmann, 1920 (Fowler, 1941) and P. yuncensis Steindachner, 1902, by having the supraoccipital process not reaching the anterior prenuchal plate (vs. supraoccipital process articulating with the anterior nuchal plate in all other Pimelodella species). From the remaining species, it differs from P. tapatapae by having the tip of the maxillary barbel reaching the region between verticals through pelvic-fin origin and anal-fin terminus (vs. reaching the caudal-fin origin). It differs from P. bockmanni, P. itapicuruensis, P. leptosoma, P. megalura, P. metae, and P. robinsoni by having dark coloration on lateral and dorsal regions of body (vs. body coloration paler, sometimes slightly darker anterodorsally). It differs from P. montana by the absence of a paired dorsolateral stripe extending from supraoccipital process to anterior one third of adipose-fin base (vs. a single dorsolateral stripe present). It is distinguished from P. peruensis and P. yuncensis by the dorsal fin presenting a hyaline stripe at the second fourth of dorsal-fin length, and maxillary barbel surpassing the vertical through pelvic-fin origin (vs. dorsal fin completely dark, without hyaline stripe; maxillary barbel very short, reaching the vertical up through half pectoral fin, but rarely surpassing the pelvic-fin origin).
When compared to Pimelodella species from the Mata Atlântica and Upper Paraná bioregions, Pimelodella longipinnis can be distinguished from P. avanhandavae, P. bahiana, P. boschmai, P. brasiliensis, P. eigenmanni, P. gracilis, P. harttii, P. ignobilis, P. kronei, P. lateristriga, P. meeki, P. pappenheimi, P. pectinifera, P. rudolphi, and P. transitoria by having the supraoccipital process not reaching the anterior prenuchal plate (vs. supraoccipital process articulating with the anterior nuchal plate in the mentioned Pimelodella species) and hypural 5 variably fused to hypurals 3+4 (vs. hypural 5 never fused to hypurals 3+4). Pimelodella longipinnis can also be distinguished from P. gracilis by the maxillary barbels reaching between verticals through pelvic-fin origin and anal-fin terminus (vs. maxillary barbels surpassing anal-fin terminus, usually reaching caudal-fin origin in P. gracilis) and by 41-42 total vertebrae (vs. 46 total vertebrae). It differs from P. avanhandavae, P. bahiana, P. brasiliensis, P. eigenmanni, P. harttii, P. ignobilis, P. kronei, P. lateristriga, P. pappenheimi, P. rudolphi, and P. transitoria by the dorsal-fin Table 1. Morphometric data for holotype and other 63 specimens of Pimelodella longipinnis. Abbreviations: Max = maximum; Min = minimum; n = number of specimens; SD = standard deviation; x = average. spine being approximately half to two-thirds of first dorsal-fin ray total length (vs. dorsal-fin spine roughly threefourths of first dorsal-fin ray total length). It also differs from P. avanhandavae, P. gracilis, and P. itapicuruensis by the short adipose fin, three to four times in SL (vs. two and half to three times in SL in P. avanhandavae and P. itapicuruensis; two to two and half times in SL in P. gracilis). Pimelodella longipinnis further differs from all Mata Atlântica and Upper Paraná species by having an overall dark body coloration, with purplish hue in life, with dark brown midlateral stripe narrow, extending from posterior portion of orbit to caudal-fin origin.

Description
Morphometric data in Table 1. Body depressed, depth at dorsal-fin origin five to seven times in SL and compressed, body width at dorsal-fin origin seven to ten times in SL (Fig. 1). Greatest body depth at dorsal-fin origin. Dorsal body profile convex from snout to origin of dorsal fin, slightly concave from that point to origin of adipose fin, convex along base of adipose fin, and concave along caudal peduncle. Ventral profile of body slightly convex from snout to branchiostegal membrane, then again convex in separate arc between branchiostegal membrane and pectoral-fin, convex between pectoral end pelvic fins, slightly convex from pelvic to anal fin, and concave from that point to caudal-fin origin.
Head moderately deep, head depth (at base of supraoccipital process) ½ to ⅔ of head length. Mouth subterminal. Eye elliptical, its longest diameter four to seven times in head. Bony interorbital distance slightly lesser than eye diameter. Barbels thin, slightly compressed, and elliptical in cross-section. Maxillary barbel when parallel to main body axis reaching to vertical through area between pelvic-fin origin and anal-fin terminus. Outer mental barbel when parallel to main body axis, reaching to verticals through area between origin and distal third of adpressed pectoral-fin. Inner mental barbel, when parallel to main body axis, reaching to verticals through area between posterior ventral limit of branchiostegal membrane and pectoral-fin origin. Supraoccipital process narrow, roughly rectangular, contacting the dorsal lamina of Weberian complex only at anterior tip. Branchiostegal rays 6 (49). Pseudotympanum large, oval, dorsal to posterior process of cleithrum and posteriorly reaching 6 th (2) or 7 th (2*) vertebra. Posterior process of cleithrum triangular, narrow, with straight dorsal margin. Anus and urogenital papilla adjacent. Urogenital papilla short, tubular and somewhat triangular. Anus roughly positioned at vertical through first third of adpressed pelvic fin; urogenital papilla near vertical through second third of adpressed pelvic fin.

Coloration in alcohol
Background body coloration mostly brown, ranging from dark brown dorsally to light brown or yellowish on ventral region of head and body (Fig. 3). Overall head coloration medium brown, with visible dark grey pigment along posterior fontanel, corresponding to a sheet of pigment covering the brain at this region. Dorsal region of maxillary barbel covered with dark brown to gray chromatophores. Outer and inner mental barbels hyaline. A poorly-defined, narrow, dark brown to dark gray midlateral stripe extending from orbit to caudal fin origin, where it broadens, and continues at median caudal-fin rays (Fig. 3). Poorly-defined darker region near dorsal fin base, varying from dark brown to dark gray coloration. Poorly-defined dark brown to dark gray col-   Slobodian, V. et al.: Rediscovery of Pimelodella longipinnis Pap. Avulsos Zool., 2021;v.61: e20216173 6/13 oration at lateroventral region of body, posterior to vertical through cleithrum process and reaching pelvic-fin origin, fading dorsally near midlateral stripe. Additional dark grayish area at pseudotympanum region, exhibiting denser concentration of chromatophores, merging with midlateral stripe. Dorsal fin dark gray, with hyaline stripe near fin base, starting roughly from the first branchedray. Pectoral, pelvic, anal and caudal fins with scattered dark brown chromatophores along rays, and membrane between rays mostly hyaline.

Coloration in life
Background body coloration dark grey to purple (Fig. 4). Some specimens are slightly lighter, presenting a medium grey background body coloration. Ventral region of head and body cream colored, except at pectoral and pelvic-fin insertions, and near mental barbels insertions, that present scattered grey chromatophores. All dark brown/gray marks observed in alcohol specimens are also observable in life: midlateral stripe (and darker area at the pseudotympanum), along posterior fontanel, near dorsal-fin base, between cleithrum process and pelvic-fin origin, and along dorsal fin (except by the hyaline stripe). Dark grayish area at pseudotympanum slightly lighter in life than in ethanol-preserved specimens.

Ecological notes
In both localities, P. longipinnis was collected in clear water, shallow streams, near sandy and rocky bottoms, with scattered leaf litter (Fig. 6), and also at water column. Collecting tools involved electric fishing and dipnet. Specimens were found in schools with five to 20 specimens, hiding at rocky crevices. Gut contents revealed an omnivore diet, with Trichoptera (80%), shrimps (15%), tadpoles (3%), and unidentified plant material (2%).
Subaquatic observations (Supplementary Material S1) revealed conspecific agonistic behaviors between P. longipinnis individuals. The observed behaviors are according to the following categories described for the channel catfish Ictalurus punctatus (Ictaluridae) (Wilson & Roys, 1994): head thrust (abrupt swing of head, laterally, against other fish), head wag (series of head thrusts), tail beat (tail thrusts as a means of displacement another fish), and tail thrust (powerful lateral swing of the tail). Furthermore, a pectoral brush (pectoral-fin brush along the side of the opponent, either parallel or antiparallel), behavior described for the blue catfish Ictalurus furcatus (Ictaluridae) (Morgan & Fine, 2020) was also observed. Described behaviors are probably related to territory dispute (Wilson & Roys, 1994) and can be observed in the video provided in the Supplementary material.

Conservation status
Pimelodella longipinnis is so far known only from Southeastern Mata Atlântica streams at the estuarine region of Santos municipality. Both Rio Quilombo and Rio Jurubatuba have stretches protected by a Brazilian park, the Parque Estadual da Serra do Mar (PESM) (Fig. 5), which is the largest Protected Area of Mata Atlântica rainforest, with 332 hectares, covering 25 municipalities from São Paulo and Rio de Janeiro States (Fundação Florestal, 2021). Despite collections inside PESM have not been authorized, we believe that P. longipinnis distribution might include protected areas inside the PESM. Therefore, P. longipinnis is categorized herein as Least Concern (LC) according to the International Union for Conservation of Nature criteria (IUCN, 2012).
A complete revision of Pimelodella species by Slobodian and de Pinna (in prep.) in underway, as well  as a phylogenetic study of the relationships of its species (Slobodian et al., in prep.). In this present context of limited taxonomic and phylogenetic knowledge about Pimelodella, the recognition of putatively monophyletic subsets within the genus has proven to be a useful approach to describe new species (cf., Slobodian et al., 2017;Slobodian & Pastana, 2018). Despite most Pimelodella species present a long supraoccipital process (i.e., reaching the anterior nuchal plate), some species present a slightly shorter supraoccipital process that never reaches the nuchal plate. Slobodian & Pastana (2018) presented a list of eleven Pimelodella species bearing this condition (the Pimelodella leptosoma-group), which includes P. longipinnis. These species exhibit a gap between the tip of the supraoccipital process and the anterior nuchal plate, corresponding to ca. 20-25% of the supraoccipital process total length, whose tip notably surpasses the midpoint of the complex vertebra in dorsal view (Slobodian & Pastana, 2018). This mentioned gap is significantly shorter than that one found in Rhamdella species (e.g., Bockmann & Miquelarena, 2008: fig. 7), whose tip of the supraoccipital process does not reach the midpoint of the complex vertebra in dorsal view. Also, the distalmost portion of the supraoccipital process of Pimelodella leptosoma-group species is embedded in skin and connective tissue, what makes its length accurately determined only through dissection or with x-ray images. Those might be the reasons that made Bockmann & Miquelarena (2008: p. 46) suggest that P. longipinnis has a "sharp long supraoccipital process that contacts the predorsal plate", altogether the authors affirmation that further discrimination of different states for this feature (their Character 8) was not pertinent to that study, focused on Rhamdella species.
Therefore, despite we disagree with Bockmann & Miquelarena's (2008) assumption that P. longipinnis supraoccipital process "contacts the predorsal plate", we agree the condition found in P. longipinnis is different from those found in Rhamdella species. Additional discussion on such matters can be found in Slobodian & Pastana (2018). It is worth mentioning that a recently published work on Heptapteridae relationships using molecular data (Silva et al., 2021) recovered Pimelodella as a polyphyletic genus of the subfamily Rhamdiinae. However, the authors did not include species of Rhamdella neither Pimelodella leptosoma-group in their analysis. Therefore, the evolution of the supraoccipital process in members of Rhamdiinae still awaits investigation under a phylogenetic approach.
Based on comparisons of the holotype of Rhamdella longipinnis with recently collected material from coastal streams of Southeastern Brazil, we could redescribe this species. Despite P. longipinnis locality being uncertain in its description, referred to as "São Paulo prov?", the presence of this species in Mata Atlântica streams is congruent with other species from the same shipping sent to the American Museum of Natural History from the former Museu Paulista (now Museu de Zoologia da Universidade de São Paulo), later described by Nichols (1919a, b) and Borodin (1927a, b, c). Among the materials sent to AMNH are individuals of Kronichthys lacerta (Nichols, 1919b) and Chasmocranus truncatorostris Borodin, 1927a, from the Mata Atlântica bioregion in São Paulo and Santa Catarina States, respectively; Hypostomus brevis (Nichols, 1919b) and Caecorhamdella brasiliensis Borodin 1927b (junior-synonym of Pimelodella kronei), generically described from "São Paulo", but now known to occur in the Upper Paraná ecoregion (Y, Figueiredo, pers. comm.) and Mata Atlântica bioregion (Ribeira de Iguape basin), respectively; besides several other species from Upper Paraná, São Francisco and Paraguay ecoregions. A summary of the species described from the shipping sent to AMNH from the former Museu Paulista are presented in Table 2.
As demonstrated in Table 2, the species described by Nichols (1919a, b) and Borodin (1927a, b, c) are from several collection events, most of them in São Paulo State, in Upper Paraná and Mata Atlântica locations. Whenever specific localities are presented for the types, collections occurred between 1898 and 1910 by several collectors, such as W. Ehrhardt, E. Garbe, R. von Ihering, H. Luederwaldt and A. Oliveira (Nichols, 1919a, b;Borodin, 1927a, b, c). However, generically defined localities are not accompanied from collectors' identification, what makes impossible the further investigation of the localities, trying to exactly pinpoint the collection sites.
Furthermore, among Pimelodella species from Mata Atlântica and Upper Paraná (the two bioregions that occur in São Paulo State, the uncertain type locality of P. longipinnis), only two species have the supraoccipital process almost reaching the anterior nuchal plate: P. itapicuruensis and P. longipinnis, and our recently collected material was identified as P. longipinnis due to the striking similarity with the holotype of this species. On the other hand, P. itapicuruensis, described from Rio Itapicuru, Bahia State, has longer maxillary barbels (65.3-87.5% SL), reaching between verticals through anal-fin origin and terminus; pectoral-fin spine bearing at posterior margin 5-7 small, shallow, retrorse serrae along basal half (Slobodian et al., 2017: fig. 6B); and a dark brown midlateral stripe running from snout to median caudal-fin rays. Pimelodella longipinnis differs from P. itapicuruensis by having shorter maxillary barbels (38.9-64.6% SL), reaching to area between verticals through pelvic-fin origin and anal-fin terminus; posterior margin of pectoral-fin spine with 8-10 straight to retrorse serrae along its basal two-thirds, larger than the ones found in P. itapicuruensis (Fig. 2); and dark grey midlateral stripe running from region posterior to orbit to caudal-fin origin, where it broadens, and continuing to median caudal-fin rays extension (Figs. 3, 4) Pimelodella longipinnis can be promptly distinguished from all these by the shorter supraoccipital process and dark gray to brown background coloration (vs. supraoccipital process reaching prenuchal plate and light gray to yellowish background coloration). A thorough revision of Pimelodella material available in Ichthyological collections with collected material from Southeastern Mata Atlântica returned only the aforementioned materials identifiable as P. longipinnis, occurring exclusively in coastal streams of São Paulo State, Brazil. No other Pimelodella species was found syntopically with P. longipinnis. These findings lead us to believe that P. longipinnis has a restricted distribution, occurring in streams of Santos municipality region. Furthermore, this work brings awareness to a poorly-known species, that occurs in one of the most densely populated regions of Brazil, and which was omitted from other review publications about the region (e.g., Menezes et al., 2007), helping in filling an important gap on Neotropical fish taxonomy.

COMPARATIVE MATERIAL
In addition to the material listed in Slobodian et al. (2017), the following material was compared in this study:  Table 2. Summary of species described by Nichols and Borodin based on shipping sent to the American Museum of Natural History from the former Museu Paulista (now Museu de Zoologia da Universidade de São Paulo). Species are followed by type locality (with bioregion indication whenever available, following Dagosta et al., 2020) and other taxonomic observations.

Described as
Type locality (and corresponding bioregion sensu Dagosta et al., 2020) Authority Current status Transferred to current status by Observations