Riama columbiana ( Andersson, 1914 )

Sanchez-Pacheco, Santiago J., 2010, Lectotype Designation And Redescription Of The Gymnophthalmid Lizard Riama Columbiana (Andersson, 1914) With Notes On The Type Locality, Papéis Avulsos de Zoologia (São Paulo) 50 (2), pp. 31-41 : 32-38

Riama columbiana ( Andersson, 1914)

Figs. 1-4

Proctoporus columbianus Andersson, 1914:3-6 [Original description. Syntypes from “ Colombia ”, Collected by P. Nisser]

Riama columbiana (Andersson) : Doan & Castoe, 2005:409 [first use of combination]

Lectotype − NRM 1631 , an adult male collected approximately 1825-1832 by Pedro Nisser. Fig. 1 .

Type locality − Colombia, probably in Municipio de Sonsón, Departamento de Antioquia. The locality reported by Andersson was simply “ Colombia ”. Slightly more information is found on the jar label at NRM, which states “probably from the Rio Magdalena area”. The syntypes were collected by Pedro Nisser, probably between 1825 and 1832. He was a Swedish miner that arrived in Colombia in 1825; he married Maria Martínez de Nisser , a famous woman of the post-independence revolution in Colombia, for which there is a vast historical literature about her and her husband (e.g. Wassen, 1969 among others). Pedro Nisser lived in the south of Antioquia Department, Sonsón Municipality, where he occasionally collected specimens of fauna that were sent or taken to the NRM. Although he did not always live in Sonsón, his works in Colombia were at the southern area of Antioquia department, which, together with records of distribution of specimens of Riama columbiana recently collected, leads me to believe that the syntypes were collected in the area .

Paralectotypes − NRM 1633-34 and NRM 6168 , two females and a badly damaged specimen (respectively) collected with the lectotype.

Referred material (Numbers in parentheses correspond to the localities in the map of the Fig. 3 ) – ICN 11302 , a male collected on March 2007 by Luis Felipe Barrera at (1) finca La Cristalina, vereda La Cristalina, municipio de Neira, departamento de Caldas, Colombia, 2300 m , approximately 05°10’24.2”N, 75°26’53”W; ICN 11295-98 (a female, a juvenile male, a female and a male respectively), a series collected on 19 February 2008 by Taran Grant at (2) Bosques de la CHEC, predio La Mesa, vereda Montaño, municipio de Villa María, departamento de Caldas, Colombia, 2640 m , approximately 4°57.864’N, 75°25.967’W; ICN 11299-01 (a male, a juvenile female and a juvenile male respectively), a series collected on August- October 2006 by Susana Velásquez and José Cirel Lopez at (3) Bosques de la CHEC, predio La Mesa, vereda Montaño, municipio de Villa María, departamento de Caldas, Colombia, 2600 m , approximately 04°58’6”N, 75°26’11”W; MHNUC 0088 , a female collected on 6 May 2007 by Julian Andres Rojas at (4) vereda Montaño, municipio de Villa María, departamento de Caldas, Colombia, 2450 m ; IAvH-R 5194, a female collected by A. Pulido at (5) Parque Municipal Campo Alegre , municipio de Santa Rosa de Cabal , departamento de Risaralda, Colombia; IAvH-R 4941, a female collected at (6) Santuário de Fauna y Flora Otun Quimbaya, departamento de Risaralda, Colombia, approximately 4°43’N, 75°34’W; ICN 6479 , a female collected on 31 May 1981 by J.D. Lynch (7) between the haciendas El Brillante and San Julian, vereda San Julian, municipio de Calarcá, departamento de Quindío, Colombia, 2100 m .

Diagnosis – Riama columbiana possesses the following characteristics: (1) frontonasal distinctly longer to distinctly shorter than frontal (in adults usually shorter than frontal); (2) prefrontal scales absent; (3) nasoloreal suture usually complete (= loreal present), rarely incomplete; (4) supraoculars four, 2, 2+3, 2+4, or 2+3+ 4 in contact with ciliaries (one specimen has only the fourth in contact with ciliaries on the right side); (5) superciliary series incomplete (1, 2, 4, 1+1, 1+2, 2+1, 2+2, 3+1 or 3+2); (6) supralabial-subocular fusion absent; (7) postoculars usually three, rarely two; (8) postparietals 2-4, usually three; (9) supratympanic temporals three; (10) genials two or three, usually three; (11) dorsal scales rectangular, juxtaposed, with a low, rounded keel (smooth on the neck); (12) longitudinal dorsal scale rows in males 25-32, in females 23-33; (13) transverse dorsal scale rows in males 42-46, in females 41-49; (14) ventral scales smooth, in 22-23 transverse scale rows; (15) lateral scale rows 3-5, usually three; (16) femoral pores per hind limb in males 10 (the lectotype has nine on the left hind limb), in females 0-2, usually zero; scales between femoral pores in males 5-6, in females 4-6; (17) subdigital scales on Toe I 4-6; (18) limbs not overlapping when adpressed against body in adults; (19) anterior cloacal plate scales two and posterior five; (20) dorsum brown; dorsolateral stripes absent; lateral ocelli present in males, usually absent in females; ventral scales with dark, centrally positioned, linearly arranged spots.

Riama columbiana can be distinguished from all known Colombian congeners and R. simotera (from Colombian-Ecuadorian border) by the high number of transverse dorsal scale rows (41-49 vs. 33-39 in R. striata , 34-41 in R. laevis and 34-39 in R. simotera ) and the texture of the dorsal scales (keeled vs. strongly striated in R. striata , and smooth in R. laevis and R. simotera [at least anteriorly]). Additionally, it can be distinguished by the following traits (condition for R. columbiana in parentheses): R. striata : suture nasoloreal usually absent [= loreal absent] (usually complete [= loreal present]) and superciliary series complete (incomplete). R. laevis : superciliary series complete (incomplete). R. simotera : nasoloreal suture absent or incomplete [= loreal absent] (usually complete [= loreal present]), femoral pores in males 6 or 7 (9-10), in females 5-7 (0-2) and scales between femoral pores in males usually two (5-6).

All Ecuadorian and Peruvian Riama : R. balneator , R. cashcaensis , R. labionis , R. petrorum , R. unicolor and R. vespertina : nasoloreal suture absent [= loreal absent] (usually complete [= loreal present]); R. colomaromani : nasoloreal suture absent or incomplete [= loreal absent] (usually complete [= loreal present]); R. anatoloros : nasoloreal suture absent or incomplete, rarely complete (usually complete), superciliary series usually complete (incomplete); R. orcesi : nasoloreal suture usually absent [= loreal absent] (usually complete [= loreal present]), femoral pores per leg in males 10-14 (9-10); R. raneyi : supraoculars two or three, usually three (four); R. hyposticta : femoral pores per leg in males 4-6 (9-10); R. oculata : femoral pores per leg in males 6-7 (9-10); R. meleagris : femoral pores per leg in females nine (0-2); R. vieta : dorsal scales rugose (keeled); R. stigmatoral : transverse dorsal scale rows 36-41 (41-49), postoculars two (usually three), scales between femoral pores in males 0-2 (5-6); R. laudahnae : supraoculars three (four), supralabial-subocular fusion present (absent). All congeners from Venezuela and Trinidad and Tobago: R. rhodogaster : differentiated lateral scale rows absent (present), scales between femoral pores in males absent (5-6), in females one (4-6); R. inanis : transverse dorsal scale rows 29-33 (41-49), transverse ventral scale rows 14-18 (22-23) and femoral pores per hind limb in males 11-12 (9-10), in females 7-8 (0-2); R. luctuosa , R. shrevei and R. achlyens : limbs overlapping when adpressed against body in adults (not overlapping).

Description of lectotype − SVL 73.4 mm, tail broken; right side of head in nasal area partially damaged; translucent epidermal layer that covers dorsal surface of scales on head lost; rostral scale wider than long, higher than adjacent supralabials, in contact with frontonasal, nasals, and first supralabials posteriorly; frontonasal longer than wide, widest posteriorly, in contact with nasals and loreal laterally, anteriormost supraoculars posterolaterally, and frontal posteriorly, slightly longer (3.4 mm) than frontal (3.3 mm), not in contact with anteriormost superciliary posterolaterally; nasal longer than wide, nostril approximately in center of scale, in contact with first and second supralabial; loreal pentagonal, shortest suture with second supralabial, in contact with frontonasal dorsally, second supralabials ventrally, anteriormost superciliary posterodorsally and frenocular posteroventrally; prefrontals absent; frontal longer than wide, anterior suture nearly straight, lateral sutures slightly concave, posterior suture angular with point directed posteriorly, not in contact with anteriormost superciliary anterolaterally, in contact with first and second supraoculars laterally, frontoparietals posteriorly; frontoparietals pentagonal in contact with second and third supraoculars anterolaterally, parietals and interparietal posteriorly; supraoculars four, second in contact with ciliaries; superciliaries 1+2, anterior superciliary lies between loreal, first and second supraoculars, and anteriormost ciliaries; major palpebral disc divided, unpigmented; frenocular quadrangular, in contact with loreal anteriorly and second and third supralabials ventrally; circumorbital scales between posteriormost supraocular and frenocular six; postoculars three; interparietal hexagonal, longer than wide, distinctly widest and somewhat rounded posteriorly, in contact with parietals laterally, postparietals posteriorly; parietals in contact with third and fourth supraocular anterolaterally, temporal scales laterally, postparietals posteriorly; postparietals two, in broad contact; anterior temporals polygonal; supratympanic temporals three; supralabials six; infralabials five; mental wider than long, in contact with first infralabials and postmental posteriorly; postmental single, hexagonal, in contact with first and second infralabials; genials irregularly shaped; scales rows between genials and collar fold (along midventral line) 11; medialmost gular scale rows not distinctly enlarged; posteriormost gular row enfolded posteriorly, concealing some scale rows; lateral neck scales squarish or rounded.

Dorsal scales rectangular (most have lost the translucent epidermal layer), longer than wide, juxtaposed, in 42 transverse rows; some middorsal scales irregularly arranged; longitudinal dorsal scales rows at fifth transverse ventral scale row 17, at 10 th transverse ventral scale row 25, at 15 th transverse ventral scale row 25; lateral scale rows at fifth transverse ventral scale row 10/11, at 10 th transverse ventral scale row three, at 15 th transverse ventral scale row three; lateral scales on body near insertion of forelimb small to granular; ventrals smooth; complete transverse ventral scale rows 23; longitudinal ventral scale rows at midbody 11; anterior cloacal plate scales two; posterior cloacal plate scales five; tail broken at 29 th subcaudal; scales on tail rectangular and juxtaposed; dorsal, dorsolateral, and ventrolateral tail scales smooth; midventral subcaudals smooth, wider than adjacent scales, square.

Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, of varying sizes, juxtaposed; midbrachial anterodorsal scale at least twice as large as adjacent scales, overlapping adjacent distal scale; anteroventral, ventral, and posteroventral brachial scales roundish, juxtaposed; antebrachial scales polygonal, of various sizes; medial antebrachial small, rounded diamonds or ovals, subimbricate; dorsal manus scales polygonal, subimbricate; palmar scales small, oval; dorsal scales on fingers smooth, quadrangular, covering dorsal half of digit, overhanging subdigitals scales, 3/2 on I, four on II, six on III, six on IV, four on V; subdigital scales 3/4 on I, five on II, eight on III, nine on IV, 6/5 on V; anterior thigh scales polygonal, 2-3 times larger than adjacent scales, becoming smaller ventrally, proximally; anterodorsal thigh scales polygonal, subimbricate; posterior thigh scales small, oval to granular, in nearly regular vertical rows; femoral pores 9/10; scales between medialmost femoral pores six; anterior and anteromedial crus scales polygonal, subimbricate, mid-crus scale at least twice as larger as adjacent scales; lateral posterolateral, and posteromedial crus scales smaller than anterior scales, subimbricate; dorsal pes scales polygonal, subimbricate; scales on dorsal surface of digits single, quadrangular, smooth, overhanging subdigital scales, three on I, five on II, eight on III, 11/10 on IV, seven on V; subdigital scales single or double, five on I, 7/8 on II, 10/9 on III, 14 on IV, 10 on V.

Coloration of the types − In preservative (70% ethanol), see Andersson’s (1914:4-5) original description.

Coloration in life Fig. 2 − Based on field notes of Taran Grant for ICN 11295-98: Dorsum and flanks dark brown; flanks with tiny white spots surrounded by black blotches to form diffuse ocelli in some specimens. Ventral scales centrally dark brown, peripherally white or pale orange. Iris orange. Smallest individual ( ICN 11296, SVL 40.29 mm) is brightest orange ventrally, appears to be lost ontogenetically. Smallest specimen also has pattern of dark brown and black (more black on flanks).

Variation − SVL of largest male 77.3 mm, largest female 71.6 mm. The paralectotypes and referred specimens are similar to the lectotype with the following noteworthy exceptions: type series and ICN 11300 are the only specimens with frontonasal scale that is not in contact with anteriormost superciliary posterolaterally; frontonasal scale in contact with anteriormost superciliary posterolaterally in 11 specimens ( ICN 11295-99 , ICN 11301-02 , ICN 6479 , IAvH 4941, IAvH 5194, and MHNUC 0088 ); nasoloreal suture incomplete in ICN 11298 and incomplete/ complete in ICN 6479 ; loreal separated from supralabials by frenocular and nasal in ICN 6479 ; in two specimens ( ICN 11297 and MHNUC 0088 ) the anterior suture of the frontal scale is concave ; ICN 11295 and ICN 11298 have frontoparietals in contact only with third supraocular anterolaterally, ICN 11300 and ICN 11297 have this condition on the right side only.

Supraoculars in contact with ciliaries, the second ( NRM 6168, NRM 1634, MHNUC 0088, and on the left side of ICN 11295) or the second and the fourth ( ICN 11298, NRM 1633, ICN 11295 and ICN 11297 on the left side, ICN 11296 and ICN 11301 on the right side) or the second and the third ( ICN 11299 and ICN 11302; NRM 1633 and ICN 11300 on the right side) or the second, the third and the fourth ( ICN 6479, ICN 11296, ICN 11300 and ICN 11301 on the left side). ICN 11297 has the fourth supraocular in contact with ciliaries on the right side). Superciliary series incomplete (one in ICN 6479; two on the left side of ICN 11296 and ICN 11300; four on the right side of ICN 11297; 1+ 1 in NRM 1633, ICN 11299 and ICN 11302, and on the left side of NRM 6168; 1+ 2 in NRM 1634, on the right side of NRM 6168, and on the left side of MHNUC 0088, ICN 11295 and ICN 11301; 2+1 on the left side of ICN 11297, and on the right side of ICN 11300; 2+2 on the right side of MHNUC 0088, ICN 11295 and ICN 11296; 3+ 1 in ICN 11298; 3+2 on the right side of ICN 11301).

Femoral pore number is the most sexually dimorphic trait, with males having 9-10 on each leg, whereas females have from 0-2. Most of the females without femoral pores; however, the female MHNUC 0088 (femoral pores 2/1-1) has one proximal femoral pore and one distal femoral pore on the right leg (there is a hiatus), on the left leg there are two proximal femoral pores. The female ICN 11300 has one distal femoral pore per leg.

NRM 1634 and NRM 6168 have five supralabial scales on the right side; ICN 11302 , IAvH 4941 and IAvH 5194 have seven supralabials and ICN 11298 has four infralabials; postoculars two in IAvH 5194; circumorbital scales between posteriormost supraocular and frenocular seven in ICN 11298 ; type series are the only specimens with two postparietals, the referred specimens (except ICN 11297) have three postparietals; ICN 11297 has four. Paralectotypes and ICN 11299 , ICN 11302 and IAvH 5194 have two genials, and MHNUC 0088 , ICN 6479 , ICN 11295-98 , ICN 11300-01 and IAvH 4941 have three genials. In ICN 11299 the genial count is two because a small median scales occludes midventral contact of a third pair of large scales on the chin.

Longitudinal dorsal scale rows at fifth transverse ventral scale row 15 in NRM 1633, at 10 th transverse ventral scale row 23 in NRM 1633, at 15 th transverse ventral scale row 23 in NRM 1633 and 20 in NRM 1634; transverse dorsal scale rows 43 in NRM 1634; transverse ventral scale rows 22 in NRM 1633 and NRM 1634; lateral scale rows at 10 th transverse ventral scale row five in NRM 1633 and three on the left side of NRM 1634, at 15 th transverse ventral scale row five in NRM 1633 and six in NRM 1634. Subdigital scales on toe I four in NRM 1633.

ICN 11302 was collected using pitfall traps in the interior of a gallery forest.

Remarks – Andersson (1914:3) described the texture of the dorsal scales of Riama columbiana as smooth, Distribution Fig. 3 − Riama columbiana occurs at moderate elevations (between 2100 m and 2640 m) on the western slope of the Cordillera Central, Colombian Andes, in the departments of Caldas, Risaralda and Quindío, and probably Antioquia (see comments on the type locality).

Habitat and ecology Fig. 4 – ICN 11299-01 were found buried, alone in an open area of pasture surrounded by secondary forest and reforested areas. ICN 11295-98 were also found buried (in topsoil 6 to 10 inches underground), one of them concealed beneath a rock. MHNUC 0088 was found under a log on the forest fragment edge delimited by pastures. which was repeated by all subsequent workers. Nevertheless, careful examination of the type series and more recently collected material shows that the dorsal scales are actually keeled. I believe Andersson’s (1914) erroneous description may be due to the lectotype having lost the translucent epidermal layer of the scales, giving them a smooth appearance. Further, in this species and other keeled species of Riama , keels are clearly visible only if the ethanol is permitted to evaporate from the scales and the angle of light is adjusted.

Andersson’s (1914) incorrect characterization of Riama columbiana has had important consequences in the Riama systematics. Kizirian & Coloma (1991), Kizirian (1995, 1996) and Doan & Schargel (2003) understandably considered the specimens KU 169946-48, from Tenerife, Valle del Cauca, Colombia, to be R. columbiana , and they derived species diagnoses from those specimens. Similarly, Doan (2003) and Doan & Castoe (2005) used these same specimens to represent R. columbiana in her phylogenetic analysis of Proctoporus sensu lato, and to allocate this species within Riama respectively. However, I reviewed the Tenerife’s series and these specimens certainly represent a different species. This is further supported by the difference in the number of femoral pores in males: 3-5 versus 9-10 in R. columbiana , the condition of the supralabial-subocular fusion: present versus absent in R. columbiana , and the number of transverse dorsal scale rows: 37-38 versus 41-49 in R. columbiana among others traits. Likewise, Castoe et al. (2004) did not analyze sequences of R. columbiana in their molecularbased phylogeny of gymnophthalmid lizards; as such, the phylogenetic placement of R. columbiana remains unknown and therefore the species treated here is allocated within Riama because its characteristics agree with those of the genus as provided by Doan & Castoe (2005). The placement of this species within Riama may be assessed in a future phylogenetic analysis.

Uzzell (1958) considered the specimens AMNH 38821 and AMNH 38822 from Ecuador as closely resemble species to Riama columbiana and, as it was said by Kizirian (1996), he put emphasis on the differences between these specimens and R. columbiana . Peters (1967) and Peters & Donoso-Barros (1970) mistakenly reported R. columbiana in Ecuador, due to a misunderstanding on Uzzell’s (1958) comments. Kizirian (1996) included them in the referred material of R. anataloros original description. Finally, Presch (1978) described the hemipenes of KU 133518 under the name Proctoporus columbianus ; however, as was stated by Kizirian & Coloma (1991), this specimen corresponds to Riama striata .

Figures

FIGURE 1: A) Dorsal and B) ventral view of the lectotype (NRM 1631 [SVL: 73.4 mm]) of Riama columbiana. Photos: Santiago J. Sánchez-Pacheco.

FIGURE 2: Riama columbiana in life.Two females (A, B: ICN 11295 [SVL: 69.2 mm] and C, D: ICN 11297 [SVL: 70.5 mm]), a subadult male (E: ICN 11298 [SVL: 57.4 mm]), and a juvenile male (F: ICN 11296 [SVL: 41.1 mm]). Photos: Taran Grant.

FIGURE 3: Distribution of Riama columbiana. Map with collection localities (black circles) for R. columbiana. Numbers correspond to the localities referenced in Referred material. Triangle = Approximated type locality, in Municipio de Sonsón, Departamento de Antioquia.

FIGURE 4: Forests of the CHEC on the western slopes of the Cordillera Central of the Colombian Andes in Caldas Department. ICN 11295-01 were found in grassland adjacent to the cloud forest. Photo: Susana Velásquez.