Two new species of semiaquatic Anolis (Squamata: Dactyloidae) from Costa Rica Chaves, Gerardo Ryan, Mason J. Bolaños, Federico Márquez, Cruz Köhler, Gunther Poe, Steven Zootaxa 2023 2023-07-25 5319 2 249 262 675PJ Chaves & Ryan & Bolaños & Márquez & Köhler & Poe, 2023 Chaves & Ryan & Bolaños & Márquez & Köhler & Poe 2023 [151,324,1542,1569] Squamata Dactyloidae Anolis Animalia 8 257 Chordata species riparius sp. nov.     Holotype. UCR 5579 ( Fig 6) collected on  7 September 2009by Douglas Robinsonand his students; an adult male from Pacuarito river, Pacuarito10 KmS-SW of Pérez Zeledón, 9.31390 N, - 83.77220 W,  880 masl, San José Province, Costa Rica.    Paratypes(all from Costa Rica: San José Province).  Adultmales: SMF 82183–82184(collected  9 October 2002by Axel Fläschendräger) from road between Platanillo and San Isidro, 9.31056 N; - 83.77500 W, ca. 850 masl;  SMF 92428(collected  15 February 2010by Gunther Köhler) from  1 kmN Cedral 9.35789 N, - 83.56094 W,  1425 masl; UCR 7893 (collected 24 August 1979by Douglas Robinson) from I.85 Km NE Alfombra, 9.3167 N, - 83.7722 W, ca. 930 masl; MCZ-R 18171 (collected 15 December 2007by Steve Poe and Mason J. Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón ( 9.31670 N, - 83.77220 W, 930 masl);  MSB 95201(collected on  10 January 2009by MasonJ Ryanand Ian Latella) from Alfombra, 12 KmS-SW of Pérez Zeledón( 9.31670 N, - 83.77220 W,  930 masl).   Adultfemales: SMF 82185–82186(collected  9 October 2002by Axel Fläschendräger) from road between Platanillo and San Isidro(ca.  850 masl);  SMF 83093–83094(collected  9 October 2002by Axel Fläschendräger) from near Platanillo(ca.  850 masl);  SMF 92429(collected  15 February 2010by Gunther Köhler) from  1 kmN Cedral( 9.35789 N, - 83,56094 W,  1425 masl); UCR 5580 (same data of the holotype); UCR 15959 (collected  3 March 2001by MasonJ Ryan) from Mollejones( 9.22930 N, - 83.64200 W,  800 masl); UCR 16162 (collected 2 July 2001by Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón ( 9.31670 N, - 83.77220 W, 930 masl);  MCZR-186143 (collected  15 December 2007by Steve Poeand MasonJ Ryan) from Alfombra, 12 KmS-SW of Pérez Zeledón( 9.31670 N, - 83.77220 W,  930 masl);  MSB 95200(collected  15 December 2007by Steve Poeand MasonJ Ryan) from Alfombra, 12 KmS-SW of Pérez Zeledón( 9.31670 N, - 83.77220 W,  930 masl); UCR 16638 (collected  15 March 2002by Franklin Aguilar) from Fila Piedras Blancas area( 9.50130 N, - 83.91280 W,  900 masl) in the Zona Protectora Los Santosin the Río Savegrebasin.   Diagnosis.  Anolis riparius  sp. nov.and  A. aquaticusare the only semiaquatic Middle American anoles that have a large orange-red dewlap with yellow in males ( Fig. 6B).  Anolis riparius  sp. nov.differs from  A. aquaticusin possessing smaller dorsal scales of the head and body (e.g., 15–19 scales across the snout between the second canthals in  A. riparius  sp. nov.versus 7–14 in  A. aquaticus, Fig. 5B), and is phylogenetically distinct (see below).  Anolis riparius  sp. nov.is distinguished from the other Middle American semiaquatic anoles as follows: from  A. robinsoni  sp. nov.by male dewlap color (chocolate brown with ill-defined brick red horizontal stripes in  A. robinsoni  sp. nov.); from  A. lionotusand  A. poecilopusby male dewlap color (solid yellow-orange in  A. lionotusand  A. poecilopus), lack of postcloacal scales in males (usually present in  A. lionotusand  A. poecilopus), and possession of zero to three enlarged rows of middorsal scales (a broadband of 10–24 enlarged middorsal scale rows in  A. poecilopusand  A. lionotus); from  A. barkeriby smaller size (maximum SVL 73 mmin  A. riparius  sp. nov., 91 mmin  A. barkeri), presence of distinctly expanded toe pads (ratio width of expanded pads / width of distal phalanx 1.8–2.2 in  A. riparius  sp. nov.versus 1.4–1.6 in  A. barkeri), and presence of a single row of middorsal caudal scales (double row in  A. barkeri).   Description of the holotype( Figure 6). Adult male as indicated by dewlap and everted hemipenes SVL 73.3 mm; HL 20.1 mm; HL/SVL 0.27; HW 10.7 mm; EOH 1.64 mm; IL 1.45 mm; IL/EOH.88; SLS-I 0.1; AGL 31.26 mm; FL 18.34 mm; T4L 15.95 mm; T4W 1.54 mm; TL 135.00 mm, tail complete; TL/SVL 1.84; TH 5.994 mmand TW 3.42 mm. 18 scales across the snout between second canthals; two scales separate the nasal opening from rostral scale; ten postrostral scales between supralabials; 12 supralabial scales from rostral scale to level of middle of eye; four scales separating supraorbital semicircles; 2 rows of scales separate subocular scales from supralabials; one slightly elongate superciliary scale; nine scales separating interparietal scale from supraorbital semicircles; four enlarged scales in supraocular disk; eight loreal scales in column just anterior to eye; 12 postmental scales posteriorly in contact with mental scale between infralabials; 15 scales in the loreal region; 13 rows of single scales in the dewlap (21–48 scales per row, rows somewhat irregular); seven enlarge middorsal scale row; 16 dorsal scales in 5% of SVL; 12 ventral scales in 5% of SVL; 220 scales around midbody; 15 expanded lamellae under fourth toe; 23 small middorsal caudal scales before the first large caudal scale ( Fig. 6D). Preoccipital, enlarged infralabial, tail crest, middorsal, postcloacal scales absent; supraocular disc presents with small scales. Frontal region of head concave; scales in supraocular disc unicarinate; rostral scale with very weak cleft and overlaps mental scale; ear opening vertically ovoid; dorsal edge of ear lacks ornamentation; mental scale completely divided posteriorly and with its posterior edge convex; dewlap extends well on to chest with anterior insertion at level of anterior portion of the eye and without protruding scales at distal edge; dorsal scales strongly keeled and uniform in size; ventral scales in diagonal rows keeled and larger than dorsal scales; lateral scales homogeneous; tail is compressed and triangular in cross-section with the base taller than wide; middorsal caudal scales in single row and its size is less than three times than the adjacent scales ( Fig. 6D); the longest toe of adpressed limb reaches anterior to eye and, supradigital scales keeled and multicarinate.  Color in Life(based on field notes and color photographs; Fig. 6A and 6B). Dewlap yellow with orange-red horizontal streaks. Dorsal ground color olive to chestnut brown with transverse olive-green bars across dorsum down flanks extending to tail. A dirty cream-colored line extends posteriorly from the labial scales down the side of the body. Scattered small greenish spots are present on the dorsum. The dorsum and tail are marked with olive-green transverse bars. The limbs are marked with pale green to yellowish transverse bars with small punctuations.  Variation. Size of reproductive males (69.6 ± 5.3, 65.8–73.3 mm) was larger than females (58.0 ± 6.0, 49.1–65.1 mm). Table 1shows variation in the lepidosis characters of  Anolis riparius  sp. nov.. Individuals exhibited little variation in coloration. Although some individuals were dark, the amount and distribution of the light bands and dots were similar among all specimens.  Hemipenis morphology.The completely everted hemipenis of SMF 92428 ( Fig. 7B) is a stout bilobed organ. The sulcus spermaticus is bordered by well-developed sulcal lips and opens into two broad concave areas, one on each lobe. A small asulcate ridge is present. The apex is strongly calyculate, truncus with transverse folds.   Etymology. The specific epithet is an adjective taken directly from Latin (  riparius= of stream banks) about the riparian habitat of this species.   Distribution.  Anolis riparius  sp. nov.is known from streams in the riparian gallery forests of the Pacuar, Guabo, and Savegre river basins ( Fig. 1) from 100 to 1450 masl. This species inhabits lowland and premontane tropical wet forest life zones.   Ecology. Little is known about the ecology and life history of  Anolis riparius  sp. nov.but it seems to share lifehistory traits with  A. aquaticusand  A. robinsoni  sp. nov.. What follows is based mainly on field observations by MJR from Alfombra and additional unnamed streams of the Rio Guabo watershed. This species may be common in small order and headwater streams but absent from some other streams within a watershed. During the day this species may be found perched on rocks and boulders along streams and can jump into the water or crawl into interstitial cracks between rocks and boulders when pursued. At night these lizards are found sleeping on moss mats, fern fronds, and other large leafy riparian vegetation, especially near the splash zone around boulders. We have also found juveniles and adults sleeping on vertical rock walls and boulders in and next to waterfalls. We have observed these lizards sleeping from 0.5 to 2.0 meters above the ground. At Alfombra on 21 December 2007MJR and SP found a congregation of seven gravid females on the same fern-covered boulder. One meter from the sleeping females was a cluster of 6 loosely aggregated eggs that were stuck to the moss on the same large boulder.  Morphological comparisons of the two new species and  Anolis aquaticus.  Anolis aquaticusis the smallest species in this complex. It has fewer scales for all traits except for the number of scales between the suboculars and supralabials ( Table 1 ). Anolis aquaticushas relatively larger dorsal scales, whereas  A. robinsoni  sp. nov.and  A. riparius  sp. nov.have small, granular dorsal scales that result in a higher number of total of scales around the body (more than 160, whereas  A. aquaticushas 160 or fewer). The number of scales between the second canthals is close to half in number in  A. aquaticusrelative to the other species ( Table 1, Fig. 4B, 5B, 6B), indicating the larger size of the scales in  A. aquaticus. The number of small middorsal caudal scale varies between 0 to 10 in  A. aquaticus( Fig. 5C) while the other species has more than 20 small middorsal caudal scales ( Fig. 4D and 4D). However, the most striking feature separating one of the three species of this group is the color of the male dewlap. The red and yellow dewlap of  A. aquaticusand  A. riparius  sp. nov.( Fig. 5Aand 6B) contrasts with the dark brown dewlap of  A. robinsoni  sp. nov.( Fig. 4B). We did not find any difference in hemipenes morphology among  A. aquaticus, A. riparius  sp. nov., and  A. robinsoni  sp. nov.( Fig. 7). 4164459315 2009-09-07 Costa Rica Perez Zeledon 880 9.3139 Douglas Robinson 1 -83.7722 Km 8 257 1 San Jose holotype 4164459304 2002-10-09 SMF Axel Flaschendrager Costa Rica 9.31056 Adult -83.775 8 257 SMF 82183-82184 1 San Jose paratype 4164459309 2010-02-15 SMF Gunther Kohler Costa Rica 1425 9.35789 1 km N Cedral 1 -83.56094 8 257 SMF 92428 1 San Jose paratype 4164459305 2009-01-10 MSB Costa Rica 930 9.3167 1 -83.7722 8 257 MSB 95201 1 San Jose paratype 4164459310 2002-10-09 SMF Axel Flaschendrager Costa Rica 850 Adult 8 257 SMF 82185-82186 1 San Jose paratype 4164459318 [225,1371,2010,2037] 2002-10-09 SMF Axel Flaschendrager Costa Rica 850 Platanillo 8 257 SMF 83093-83094 1 San Jose paratype 4164459303 2010-02-15 SMF Gunther Kohler Costa Rica 1425 9 258 9.35789 1 km N Cedral 1 -83.56094 8 257 SMF 92429 1 San Jose paratype 4164459308 2001-03-03 SMF Costa Rica 800 9.2293 1 km N Cedral 1 -83.642 9 258 1 San Jose holotype 4164459302 2007-12-15 MCZ Costa Rica 930 9.3167 1 -83.7722 9 258 1 San Jose paratype 4164459319 2007-12-15 MSB Costa Rica 930 9.3167 1 -83.7722 9 258 MSB 95200 1 San Jose paratype 4164459314 2002-03-15 Franklin Aguilar Costa Rica 900 9.5013 Fila Piedras Blancas area 1 -83.9128 9 258 1 San Jose paratype 4164459316 [415,1047,403,429] Zona Protectora Los Santos Costa Rica Rio Savegre 9 258 1 San Jose paratype