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Anolis caceresae HOFMANN & TOWNSEND, 2018

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Higher TaxaDactyloidae, Iguania, Sauria, Squamata (lizards)
Common NamesE: Berta’s Anole
Spanish: Pichete de Berta 
SynonymNorops caceresae HOFMANN & TOWNSEND 2018
Anolis crassulus — HAHN 1971: 111
Anolis crassulus — MEYER & WILSON 1971: 106
Anolis crassulus — MEYER & WILSON 1973: 16
Anolis crassulus — WILSON 1983: 125
Anolis crassulus — WILSON and TOWNSEND 2007: 135
Anolis crassulus — TOWNSEND & WILSON 2009: 64
Anolis crassulus — WILSON & JOHNSON 2010: 125 (in part)
Anolis crassulus — TOWNSEND & WILSON 2010: 474 (in part).
Norops crassulus — McCranie et al. 1992: 208
Norops crassulus — WILSON & MCCRANIE 1994: 148
Norops crassulus — WILSON & MCCRANIE 2002: 91
Norops crassulus — WILSON & MCCRANIE 2003a: 24 (in part)
Norops crassulus — WILSON & MCCRANIE 2003b: 37
Norops crassulus — KÖHLER & OBERMEIER 1998: 127 (in part)
Norops crassulus — KÖHLER 2008: 104 (in part)
Norops crassulus — SOLIS et al. 2014: 129 (in part)
Norops crassulus — TOWNSEND 2014: 239 (in part)
Norops crassulus — JOHNSON et al. 2015: 81 (in part)
Norops crassulus — MCCRANIE 2015: 368 (in part)
Norops crassulus — MCCRANIE & KÖHLER 2015: 49 (in part).
Norops crassulus (Honduras) — KÖHLER et al. 1999: 286
Anolis crassulus (Honduras) — HOFMANN & TOWNSEND 2017: 8 
DistributionSW Honduras (Comayagua, Intibucá, La Paz, Lempira), elevation 1,200–2,260 m

Type locality: near Río Agua Negra, 14.459°N, 88.385°W, 1,940 m above sea level, Reserva Biológica Opalaca, Departamento de Intibucá, Honduras (map in Fig. 6 in HOFMANN & TOWNSEND 2018)  
TypesHolotype: CM 161315 (field number JHT 3914), Figures 3 and 4, an adult male, collected on 28 May 2015 by Thomas J. Firneno Jr., Michael W. Itgen, Josiah H. Townsend, and Kayla D. Weinfurther.
Paratypes: (n = 22; 12 males, 10 females); CM 161303–161305, 161308, MVZ:Herp:286121–286122, KU 348836, KU 348838 [JHT 3739, 3741], (field num- bers JHT 3738–39, 3741–43, 3764, 3767, 3781), adult males, and CM 161301–161302, 161306–161307, MVZ:Herp:286123–286124, KU 348835, KU 348837 [JHT 3737, 3740] (field numbers JHT 3735–37, 3740, 3766, 3768–70), adult females, from near El Rodeo, 14.441°N, 88.145°W, 2100 m above sea level, Refugio de Vida Silvestre Mixcure, Departamento de Intibucá, Honduras, collected 23 May 2015 by Thomas J. Firneno Jr., Michael W. Itgen, Josiah H. Townsend, and Kayla D. Weinfurther; CM 161310, 161312–161314 (field numbers JHT 3887, 3906, 3910–11), adult males, CM 161309 (field number JHT 3822), an adult female, and CM 161311 (field number JHT 3888), a subadult female, with the same col- lection data as the holotype. 
DiagnosisDiagnosis.—A medium-sized species (maximum record- ed snout-vent length 57 mm in males: KU 348838, CM 161310, CM 161315; 54 mm in females: KU 348837, CM 161307) of beta anole in the genus Norops endemic to southwestern Honduras. Norops caceresae can be dif- ferentiated from all other anoles in Honduras, except those of the N. crassulus species subgroup, by the combination of having enlarged middorsal scales, strongly keeled ven- tral scales, two or fewer scales separating the supraorbital semicircles, four to seven loreal scale rows, suboculars and supralabials in contact, heterogeneous lateral body scales, enlarged postcloacal scales in males, and an orange-to-red male dewlap (in life). Norops caceresae can be diagnosed from the other species of the Norops crassulus species subgroup, except N. crassulus sensu stricto, as follows (known distributions in parentheses): from N. amplisqua- mosus McCranie et al., 1993 (northwestern Cortés, Hon- duras), N. heteropholidotus (Mertens, 1952) (northwestern El Salvador, southeastern Guatemala, and southwestern Honduras), N. muralla (Köhler et al., 1999) (northwestern Olancho, Honduras), N. sminthus (Francisco Morazan and southern Comayagua, Honduras), and N. wermuthi Köhler and Obermeier, 1998 (northern Nicaragua and extreme southeastern Honduras) by its strongly keeled ventral scales (versus smooth to weakly keeled); from N. anisol- epis (Chiapas, Mexico) by its larger size (maximum SVL 57 mm in males, 54 mm in females; versus 47 mm in male, 48 mm in female N. anisolepis) and its orange-to-red male dewlap (versus bright pink male dewlap in N. anisolepis); from N. haguei (Stuart, 1942) (Alta Verapaz, Guatemala) by its larger dorsal scales (34–43 between level of axilla and groin, approximately equal to or slightly smaller in size than ventrals; versus 41–57, dorsals much smaller than ventrals in N. haguei); from N. morazani (northern Francisco Morazán, Honduras) by having a hemipenis with an undivided asulcate processus (versus divided asulcate processus in N. morazani), and having 6–9 scales separating the nasals (2–4 in N. morazani); and from N. rubribarbaris Köhler et al., 1998 (Santa Bárbara, Hondu- ras) by having a TL/SVL 1.82–2.02 in males, 1.60–2.00 in females (versus 2.16–2.54 in males, 2.19–2.21 in females of N. rubribarbaris), and having 12–20 rows of enlarged dorsal scales (versus 8–11 in N. rubribarbaris).
Norops caceresae is most similar in external mor- phology to N. crassulus sensu stricto (highlands of cen- tral Guatemala, southwestern El Salvador, and Chiapas, Mexico). Norops caceresae is distinguished from all other populations assigned to N. crassulus by the following: (1) a longer head relative to width: HL/HW 1.49–1.70 (1.58 ± 0.07) in males, 1.50–1.70 (1.58 ± 0.07) in females (versus 1.32–1.65 [1.46 ± 0.06] in males, 1.32–1.56 [1.46 ± 0.07] in females of N. crassulus); (2) a shorter tail relative to snout–vent length: complete TL/SVL 1.79–2.02 (1.91 ± 0.01) in males, 1.60–2.00 (1.87 ± 0.14) in females (ver- sus 1.62–2.37 [2.20 ± 0.24] in males, 1.54–2.44 [2.07 ± 0.24] in females of N. crassulus) (3) a longer shank rela- tive to snout-vent length: ShL/SVL: 0.24–0.29 (0.26 ± 0.01) in males, 0.22–0.27 (0.25 ± 0.02) in females (versus 0.22–0.27 [0.24 ± 0.01] in males, 0.21–0.25 [0.23 ± 0.01] in females of N. crassulus); (4) a longer shank relative to head length: ShL/HL: 0.84–1.13 (0.94 ± 0.07) in males, 0.79–0.94 (0.92 ± 0.06) in females (versus 0.74–0.95 [0.83 ± 0.12] in males, 0.74–0.97 [0.84 ± 0.05] in females of N. crassulus); (5) a slightly higher average number of lamel- lae on Phalanges II-IV of the 4th toe: 25.7 ± 1.8 in males, 24.9 ± 1.3 in females (versus 23.3 ± 1.9 in males, 22.5 ± 2.0 in females of N. crassulus). 
CommentHabitat: Norops caceresae inhabits both intact and disturbed Mixed Transitional Cloud Forest, Broad-leaf Cloud Forest, and Mixed Cloud Forest between 1,200 and 2,260 m elevation. This diurnal species appears most abundant near forest edges and along streams and is locally abundant in the matrix of traditional agriculture and forest patches that typifies much of the Lenca Highlands (Fig. 6). At night, N. caceresae can be encountered asleep on low vegetation, typically >1.5 m above the ground.

Sympatry: N. heteropholidotus around a large stream through relatively intact Mixed Cloud Forest in the Cordillera de Opalaca in western Intibucá, Honduras, and with N. heteropholidotus and N. laeviventris (Wiegmann, 1834) in and around small Mixed Cloud Forest fragments at Cerro San Pedro La Loma in eastern Intibucá.

Group: member of the crassulus clade of Norops. 
EtymologyThe eponym caceresae is a noun in the genitive case, and is given in honor of Berta Isabel Cáceres Flores of La Esperanza, Departamento de Intibucá, Honduras (approximately 25 km from the type locality). Berta Cáceres was a community leader and environmental activist who cofounded the Consejo Cívico de Organizaciones Populares e Indígenas de Honduras (COPINH: Civic Council of Popular and Indigenous Organizations of Honduras), and, in the face of threats against her and her family and the murders of friends and colleagues, led grassroots efforts to unite communities against environmentally destructive actions and the privatization of native lands in Honduras. She was awarded the Goldman Prize for Conservation in 2015, and continued her work organizing and fighting for indigenous rights and environmental justice until she was assassinated in her home in La Esperanza on 3 March 2016. The authors named this species as a small gesture to honor Berta and to raise awareness for COPINH and their work, while continuing to draw attention to the plight of indigenous and environmental activists in Honduras, dozens of whom have been murdered over the past decade. 
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