A peer-reviewed open-access journal Zookeys 569: 105-121 (2016) Reena #ZooKeys http:/ / ZOO keys -pen soft. net Launched to accelerate biodiversity research A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae) Jonathan W. Armbruster', Nathan K. Lujan? | Department of Biological Sciences, 101 Life Sciences Building, Auburn University, AL 36849, USA 2 Department of Biological Science, University of Toronto Scarborough, Toronto, ON MICIA4, Canada Corresponding author: Jonathan W. Armbruster (armbrjw@auburn.edu) Academiceditor: VV. Bogutskaya | Received 18 September 2015 | Accepted 12 December2015 | Published 26 February 2016 http://zoobank.ore/D8631FD8-6EF2-48F F-9656-75AEO0D0736F Citation: Armbruster JW, Lujan NK (2016) A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae). ZooKeys 569: 105-121. doi: 10.3897/zookeys.569.6630 Abstract A new species of the suckermouth armored catfish genus Peckoltia is described from the lower Ventuari River, a tributary of the upper Orinoco River in Amazonas State, Venezuela. Specimens of this species were formerly included in the wide-ranging Amazonian species P vittata, but a recent molecular phylog- eny found Orinoco individuals to be distantly related to Amazon Basin individuals spanning the range of P vittata syntypes. Detailed morphological examination confirmed distinctiveness of Orinoco specimens, and found them to be diagnosable from true P vittata by having generally greater than 25 teeth (vs. less), spots on the nape (vs. nape lacking spots), the upper lip with two to three black bar-shaped markings in a line like a moustache (vs. lips generally with a hyaline wash), and by the snout having a medial black line disconnected from the moustache markings (vs. medial snout stripe connected to a bar just above the lip). Peckoltia wernekei displays remarkable genetic similarity to its sister species, P /ujani, but differs mor- phologically by having dentary tooth rows meet at an angle less than 90° (vs. >90°), by having large faint blotches on the abdomen (vs. abdomen with no blotches), by a smaller internares width (21.2—26.6% vs. 28.5—46.5% of interorbital width), and a larger dorsal spine (148.1-178.6% vs. 80.1-134.5% of abdominal length). Keywords Ancistrini, Hypostominae, Molecular Phylogeny, Morphology, Peckoltia, Systematics, Taxonomy Copyright Jonathan W. Armbruster, Nathan K. Lujan. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 106 Jonathan W. Armbruster & Nathan K. Lujan / ZooKeys 569: 105-121 (2016) Introduction Peckoltia Miranda Ribeiro, 1912 is a genus of suckermouth armored catfishes (Loricariidae) with 18 currently described species (Armbruster et al. 2015). Armbruster (2004; 2008) had restricted Peckoltia to those species of Ancistrini that had dentary tooth rows meeting at an angle of 90° or less and that lacked the synapomorphies of similar genera like Hypancistrus and Panagolus (the latter then the Panaque dentex group); however, this arbitrary definition was not supported by the molecular analysis of Lujan et al. (2015), so Armbruster et al. (2015) recognized an expanded Peckoltia that currently lacks a morphological diagnosis, but is strongly monophyletic based on molecular evidence (Lujan et al. 2015). Armbruster (2008) had recognized a wide-ranging Peckoltia vittata (Steindachner, 1881) that included specimens from the Maranh4o to the Madeira and upper Ori- noco, but suggested that this putative range of P. vittata included multiple species. The syntype series of Peckoltia vittata itself contains specimens spanning more than 600 km of the main channel of the Amazon River, from the Xingu River to the Madeira River, making it difficult to know where one might reliably find the true P. vittata. Speci- mens from the Xingu, Madeira and Orinoco that were morphologically consistent with Peckoltia vittata were found to be polyphyletic in Lujan et al. (2015) and Lujan et al. (in review, Fig. 1). In addition, P. greedoi was recognized as distinct from P. vittata by Armbruster et al. (2015; specimens of P. greedoi were originally included among the specimens of P. vittata examined by Armbruster 2008). Lujan et al. (2015) found the putative specimens of Peckoltia vittata from the Ori- noco to be strongly supported as sister to another Orinoco species, P. /ujani Armbrust- er, Werneke and Tan, and part of a clade with another undescribed species known as Peckoltia sp. n. Meta L147. In this study, we re-examine specimens identified as P. vittata from the upper Orinoco and describe them as a new species clearly distinguish- able from Amazon Basin P. vittata. In addition, we provide a key to all of the current species of Peckoltia. Methods Methods follow Armbruster (2003) with the addition of counts of mid-dorsal and mid-ventral plates (the number of plates in these series from the head to caudal fin and excluding the last, triangular plate, which is beyond the hypural). Institutional abbreviations are as in Sabaj Pérez (2014). Names of skeletal characteristics are as in Schaefer (1987) and Geerinckx et al. (2007) and of plate rows as in Schaefer (1997). Full morphometric dataset is presented in Suppl. material 1, locality information for species described below is presented in Suppl. material 2. A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae) 107 1% compta (Tapajos) _ ~_P braueri (Branco) P.n.sp. Madeira L210 —/35/— P. vittata (Xingu) U 0.98/96/77, ; oseas~ P. lineola (Ventuari) 2 P. aff. vittata (Madeira) ~ ee ——— P. sabaji (Amazon/Orinoco) 2 osseanr Pe. furcata (Upper Amazon) iy a Sanne P. relictum (Upper Amazon) P. pankimpuju (Maranon) osaee-[ P. lujani (Orinoco) P. wernekei (Ventuari) 0.99/100/86 0.98/63/— P. N.Sp. Meta L147 Panagolus Posie Scobinancistrus Ancistomus ‘Panagolus’ koko 0.05 Figure |. Phylogenetic relationships within the Peckol/tia Clade (sensu Lujan et al. 2015), from Lujan et al. (in review). Results based on analysis of a 4293 base pair alignment consisting of two mitochondrial (16S, Cyt b) and three nuclear loci (RAG1, RAG2, MyH6). Node support values given in order as Bayes- ian posterior probability, maximum likelihood bootstrap and maximum parsimony bootstrap. Taxonomy Peckoltia wernekei Armbruster & Lujan, sp. n. http://zoobank.org/3488FBE3-34F0-4F5B-94D2-G0BFA849D945 Figs 3—4 and 5b, Table 1 Peckoltia aff. vittata (Orinoco) Lujan et al., 2015 [molecular phylogeny] Type locality. Ventuari River drainage, Amazonas State, Venezuela, South America Holotype. AUM 54314, 104.6 mm SL, VENEZUELA, Amazonas State, Ventu- ari River drainage, Marujeta Creek, 159 km E of San Fernando de Atabapo, 04.2948°, -066.2889°, N.K. Lujan, M. Sabaj Pérez, D.C. Werneke, T. Carvalho, V. Meza-Var- gas, 02 April 2010. Paratypes. All specimens Venezuela, Amazonas State, Ventuari River drainage: AUM 39248, 1, 63.2 mm SL, Ventuari River at beach at village of Moriche, 116 km NE of Macuruco, 169 km NE of San Fernando de Atabapo, 04.7503°, -066.3549°, D.C. Werneke, N.K. Lujan, M.H. Sabaj, L.S. de Souza, 7 April 2004. AUM 39313, 13, 2 cs, 56.8-87.3 mm SL, Manapiare River, 14.5 km NW of San Juan de Manapi- 108 Jonathan W. Armbruster & Nathan K. Lujan / ZooKeys 569: 105-121 (2016) Table 1. Selected morphometrics of Peckoltia wernekei. Numbers in parentheses refer to landmark numbers in Armbruster (2003). Holotype SD Min | Max SL, mm (1-20) 104.6 104.6 %SL Predorsal Length (1-10) 38.8 ak Ares 45.2 Head Length (1-7) 36.5 | 3a" S50 | Rio | ara 37.5 Head—dorsal Length (7—10) 6.7 23 79 Cleithral Width (8—9) 26.4 29 oe B07 Head-pectoral Length (1-12) 25.8 23 27, : : 2889 Thorax Length (12-13) 299 24.8 Pectoral-spine Length (12-29) Ble 23 32.0 34.7 Abdominal Length (13-14) 22.0 23 20.3 22.0 Pelvic-spine Length (13-30) era) 23 252 f ; 29.0 Postanal Length (14-15) 39.0 39.0 Anal-fin spine Length (14-31) 18.1 Lal Dorsal—pectoral Distance (10-12) 24.5 23 26.8 297 Dorsal spine Length (10-11) 32.6 vP) By Dae Dorsal-pelvic Distance (10-13) 24.9 we) 22.3 : : 24.9 Dorsal-fin base Length (10-16) 29.3 28.9 30.9 Dorsal-adipose Distance (16-17) 15.3 17.4 Adipose-spine Length (17-18) 9.4 25 10.7 12.0 Adipose-upper caudal Distance (17-19) 16.2 25 al 19.3 Caudal-peduncle Depth (15-19) 10.8 23 Lahey : 13.2 Adipose-lower caudal Distance (15-17) 22.9 23.0 24.8 Adipose-anal Distance (14-17) 20.8 23 20.3 21.0 Dorsal-anal Distance (14—16) 14.9 23 15.0 16.3 Pelvic-dorsal Distance (13—16) 27.5 23 2235 27D % Head Length es Es ee | Head-eye Length (5-7) 31.4 41.3 Orbit Diameter (4—5) 18.9 23 21.9 Don, Snout Length (1-4) DOT 23 56.3 Sh) Internares Width (2-3) 11.2 oh 1.3, : : 12:3 Interorbital Width (5-6) 43.0 49.6 Head Depth (7-12) 59.6 73.5 Mouth Length (1—24) 41.9 23 41.9 46.1 Mouth Width (2122) 42.4 ee ees 46.0 Barbel Length (22-23) 16.9 23 A ; ; 22.9 Dentary Tooth Cup Length (25—26) 11.3 13.8 Premaxillary Tooth Cup Length (27-28) 10.6 3 | a20 | 15s | 92, 15.4 are, 05.4286°, -066.1362°, N.K. Lujan, M.H. Sabaj, L.S. de Souza, D.C. Werneke, 12 April 2004. AUM 39839, 1, 31.5 mm SL, Manapiare River, 10 km NW of San Juan de Manapiare, 05.3868°, -066.1159°, N.K. Lujan, L.S. de Souza, D.C. Werneke, A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae) 109 ee ee a ¢ jg Ae Figure 2. Mouth of Peckoltia multispinis, holotype, NUW 8952. Photo by M.H. Sabaj and K. Luckenbill. ail / ee eS ae ¥ ; ee M.H. Sabaj, 14 April 2004. MCNG 56680, 13, 52.6-80.2 mm SL, same data as AUM 39313. Diagnosis. Peckoltia wernekei can be separated from all other Peckoltia by having a broken black line of pigment on the upper jaw (vs. solid line of pigment along snout edge or snout uniformly colored or mottled). Peckoltia wernekei can be further sepa- rated from P. vittata by generally having 25 or more teeth in at least one dentary or one premaxilla (vs. generally 24 or fewer; one specimen of P. wernekei had both upper and lower jaws with <25 teeth/ramus), by having a largely naked abdomen (abdomen with a few plates below pectoral girdle, between pelvic fins and along sides of abdomen; vs. most of ventral surface from the throat to the anus with small plates), and by having large, faint blotches on the abdomen (vs. abdomen uniform). Peckoltia wernekei can be further separated from upper Orinoco congeners as follows: from P. brevis and P. lineola by lacking short lines and spots on the head (vs. lines and spots present), from P. brevis, P. caenosa and P. lineola by having a largely naked abdomen (vs. abdomen fully plated), and by generally having 25 or more teeth per jaw ramus (vs. 22 or fewer in P. brevis, 21 or fewer in P. caenosa, and 19 or fewer in P. lineola); from P. lujani by having the dentaries meet at an angle less than 90° (vs. >90°), by having large, faint 110 Jonathan W. Armbruster & Nathan K. Lujan / ZooKeys 569: 105-121 (2016) Figure 3. Holotype of Peckoltia wernekei sp. n., AUM 54314, 104.6 mm SL, dorsal, lateral, and ventral views. Scale = 1 cm. Photos by J.W. Armbruster. blotches on the abdomen (vs. abdomen with no blotches), by a smaller internares width to interorbital width ratio (21.2—26.6% vs. 28.5—46.5%), and a larger dorsal spine to abdominal length ratio (148.1-178.6% vs. 80.1-134.5%); and from P. sabaji by having bands in the dorsal and caudal fins (vs. spots) and prominent dorsal saddles on the body (vs. large spots). Description. Morphometrics in Table 1. Counts and measurements based on 23 specimens. Small to medium-sized loricariids, largest specimen examined 104.6 mm SL. Body stout, but slightly narrower than P. vittata. Head gently sloped to su- praoccipital. Supraoccipital with tall, rounded crest. Supraoccipital crest raised slightly above nuchal region. Nuchal region rises slightly to nuchal plate. Dorsal slope decreas- ing in straight line to insertion of dorsal procurrent caudal-fin rays then ascending to A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae) 111 Figure 4. Paratype of Peckoltia wernekei sp. n., AUM 39313, 73.5 mm SL dorsal, lateral, and ventral views. Scale = 1 cm. Photos by J.W. Armbruster. caudal fin. Body depth greatest at anteriormost insertion of dorsal fin. Ventral profile flat to caudal fin. Caudal peduncle trapezoidal in cross section with dorsal surface flat- tened. Body widest at insertion of pectoral fins, narrowest at insertion of caudal fin. Snout rounded. Eye moderately sized (orbit diameter 18.9 + 1.0% of head length), dorsal rim of orbit forming tall crest that continues forward to area just anterior of nares as low, rounded ridge. Iris operculum present. Interorbital space with slight, rounded, median hump that is contiguous with ridge of parieto-supraoccipital. Parieto-supraoccipital pointed posteriorly with posterior point raised above nuchal region in small crest. Infraorbitals, frontal, nasal, compound pterotic and parieto-supraoccipital supporting odontodes. Preopercle generally supporting a single column of odontodes that gener- hi2 Jonathan W. Armbruster & Nathan K. Lujan / ZooKeys 569: 105-121 (2016) Figure 5. Live photos of A Peckoltia lujani (uncataloged), photograph by N.K. Lujan, and B P. wernekei sp. n. AUM 39313 (was used as a live photo of P. vittata in Armbruster 2008). Photograph by M.H. Sabay Pérez. ally decrease in number with increasing body size; largest specimen with some odon- todes located posterodorsally on preopercle and two individuals without preopercular odontodes. Opercle with one to three rows of odontodes with numbers of rows and numbers of odontodes generally decreasing with increasing body size; largest indi- vidual without odontodes. Lips covered with short, wide papillae. Lower lip wide, reaching just to or slightly short of pectoral girdle; upper lip narrow. Edge of lower lip smooth. Maxillary barbel only barbel present, reaching about two-thirds of distance to gill opening from base of barbel. Median plates 24—26 (mode 24). Plates unkeeled, but first four or five plates of mid-ventral series bent to form slight ridge. Five caudal peduncle plate rows. Plates on all dorsolateral surfaces of body. Throat naked. Abdomen mostly naked except for a line one to three platelets wide along and slightly posterior to anterior margin of pectoral girdle, few uneven rows of platelets ventral to ventral plate series, patch A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae) 113 of platelets below posterior section of pelvic girdle; number of platelets on abdomen increases with body size. Evertible cheek plates supporting hypertrophied odontodes evertible perpendicular to head. Cheek odontodes 17—40 (mode 33). Longest evertible cheek odontode almost reaching vertical through posterior edge of pectoral-fin spine. Hypertrophied cheek odontodes relatively weak. Odontodes slightly longer than aver- age body odontodes present along dorsal-, adipose-, pelvic-, caudal-, and pectoral-fin spines; larger individuals with hypertrophied odontodes at tip of pectoral spine. Dorsal fin ii,7; dorsal spinelet V-shaped, dorsal-fin locking mechanism present, last ray of dorsal fin not reaching or just reaching preadipose plate when adpressed. Adipose fin with single preadipose plate and moderately long spine. Caudal fin i,14,i; caudal fin forked, ventral lobe longer than dorsal lobe; dorsal and ventral procurrent caudal-fin rays five. Pectoral fin i,6; pectoral-fin spine reaching just posterior to pelvic fin when adpressed ventral to pelvic fin. Pelvic fin i,5; pelvic-fin spine extending one to two plates posterior to anal fin when adpressed. Anal fin i,4; unbranched anal-fin ray slightly shorter than first branched ray. Teeth bicuspid with lateral lobe one-half to three-quarters length of medial lobe and lateral cusp half width of medial cusp. Eighteen to 32 left dentary teeth (mode 31; 1 of 23 with less than 25); 22-35 left premaxillary teeth (mode 32; 1 of 23 with less than 25); all specimens with at least one jaw ramus having 25 or more teeth. Color. Base color reddish brown. Head and nape mottled dark brown with dis- tinct, medial dark line along mesethmoid and slightly less distinct lines from lateral portion of naris to lip mark. Parieto-supraoccipital crest darker than surrounding ar- eas. Lip with dark mark consisting of two or three ovoid dashes of brown, lateral portions continue as line to lateral portions of nares (Fig. 6). Dark portion of lips separated from mesethmoid line by distinctly lighter C-shaped region (Fig. 6), which may continue less distinctly between mesethmoid line and lines lateral to nares. Lips may also have other spots. Head colors less distinct in larger specimens. Body with four distinct, oblique bars, first below anterior of dorsal fin, second below posterior end of dorsal fin and anterior part of interdorsal space, third beginning at preadipose plate to about posterior edge of adipose spine, and fourth at end of caudal peduncle; first and second bars fade into a dark wash ventrally connecting the two bars; third and fourth bars continuing around caudal peduncle; bars connected at median plate series. Secondary bars sometimes present between any two primary bars, but generally not as dark, darker ventrally than dorsally; posterior secondary bars darker than anterior; sec- ondary bars may connect across caudal peduncle. Pectoral-fin spine dark brown with alternating, similarly-sized dark and light spots, spots continuing as bands on fin; two to five dark bands on fin with number increasing with body size. Pelvic fin as pectoral but with two to four dark bands. Dorsal fin as pectoral but with dark bands distinctly wider than light bands, three or four dark bands. Anal and caudal fins as pectoral, but with light bands wider than dark bands (one to three dark bands in anal and three to five bands in caudal). Adipose spine with dorsal section of third dark bar covering base in all specimens, with some also having dark spot at tip of spine; in largest individual (holotype) basal and distal spots combine along posterior margin of spine, with ante- Jonathan W. Armbruster & Nathan K. Lujan / ZooKeys 569: 105-121 (2016) 114 Figure 6. Schematic of the mouth of Peckoltia wernekei based on the specimen in Figure 3. Dentaries (D) form less than a 90° angle with one another whereas premaxillae (PM) are relatively straight. A broad, broken band is formed from ovate blotches (OB) on the upper lip, separated from a line that runs along the mesethmoid (ML) by a C- shaped light space (C). braueri P @P caenosa cavatica lineola lujani P. P AP pti | Aiveg ; | eile { ae sabaji i . % t - wernekei P ie Figure 7. Distribution of Peckoltia around the western Guiana Shield. “T” indicates type locality of P. wernekei sp. n. A new species of Peckoltia from the Upper Orinoco (Siluriformes, Loricariidae) 115 rior edge having light space between spots. Abdomen mostly light, but with four to six large, faint blotches laterally and occasionally with one or two median faint blotches. Sexual dimorphism. None observed. Distribution. Known only from the Ventuari River, a right-bank tributary of the upper Orinoco River in Amazonas State, Venezuela (Fig. 7). Etymology. Patronym honoring David C. Werneke, Collection Manager of Fish- es at the Auburn University Museum, for his diligence, camaraderie and humor during three expeditions to the upper Orinoco Basin and for his long service as a Collection Manager at Auburn University. Key to the species of Peckoltia (after Armbruster (2008) and the present paper) 1 co | \9 Eyes reduced (orbit diameter <10% of HL); long caudal-fin filaments; lack- ing pigment or color reduced, light tan with very slightly darker saddles, mottled black and white or completely black............ Peckoltia pankimpuju Eyes normal (orbit diameter >13% of HL); caudal fin lacking filaments; color normal, with dark brown saddles, spots, or lines on a tan background)....... 2 Dentaries meeting at an angle less than 90° to just slightly greater .............. 3 Dentariésuncctineat-an angle ereater than ae. 0, dsesadsenssenenesepebioesene 17 Posterior margin of lower lip with finely branched fimbriate papillae (Fig. 2)..... ee renee arrest occ Seren Peckoltia multispinis Posterior margin of lower lip with simple fimbriae or smooth... 4 Distinct round spots present on head and/or sides of body ........eceeeeeeeeee 5 Color pattern on head consisting of large blotches, saddles or lines, lacking distinct-round spots*on head Or Sidlesi2c.5 ters scenes Soest tance 12 Some spots on the parieto-supraoccipital and/or compound pterotic combin- HYG VLO GM OGMAM TIES en tur.oteT matesd ie sieds Ve ontras icecbees yiouinece Pes AEs Duet cea t AW ots tase teat 6 All spots distinct, none combining to form lines... eee eeeeseeseeeeeeeeeeeees 7 Lines on head most prominent on compound pterotic, not radiating from a central point on the parieto-supraoccipital; lines on head approximately same width.or wider than ‘pupil ...7. 4h. Peckoltia lineola Lines on the head most prominent on the parieto-supraoccipital where they radiate from a central point; lines on head narrower than pupil... Lieeeecateeceesute even eA Ue ibe kisuliir mv OPE sal ONE A bode seadicebobei Peckoltia vermiculata ARES) dear vceteawane alos epee vase eres Riel tusgeecst yd. Gsltaag teach cis ne ea eg csp svena vane sarong iss 8 Spots present on caudal fin separate, not forming bands... eee 10 Watetaltbodyastiriaees withespots alate erste gcee setae at Peckoltia otali Wateralibody surtaces waitdlorsal saddles 9. sess. Seecsnc tee, eeee de lestachs Aeonatt 9 Spots on the dorsal fin; no spots on the abdomen; caudal fin lunate with up- per caudal-fin spine longer than lower spine (usually the tail is broken and this-chatacte mise @Dsetvable) tiie o..tssSoare