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River rocks as sleeping perches for Norops oxylophus and Basiliscus plumifrons in the Talamanca Mountains, Costa Rica

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We report the first observation of a nocturnal perch site for both Norops oxylophus and Bassiliscus plumifrons on river rocks in Costa Rica.
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Acknowledgments.––We thank Rancho El Coyote for providing the accommodations and use of their facil-
ities during our stay. We also thank Abelino Cota and the students at Fausto’s Lab for field assistance, and Carla
Sette for her comments on this manuscript. The collecting permit number (SGPA/DGVS/08184/16) was issued
by the Dirección General de Vida Silvestre of México to Patricia Galina Tessaro. This project was supported by
CONACyT (1319).
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rafaeL a. Lara-resendiz1, Yaredh ramírez-enríquez2, isaí VaLLe-Jiménez2,
Jorge h. VaLdez-ViLLaViCenCio3, fausto r. méndez-de La Cruz4, and PatriCia gaLina-tessaro2
1Department of Ecology and Evolutionary Biology, Earth and Marine Sciences Building A316, University of California,
Santa Cruz, 95064 California, United States. E-mail: rafas.lara@gmail.com
2Centro de Investigaciones Biológicas del Noroeste, La Paz, Baja California Sur, Mexico.
E-mails: y.ramz02@hotmail.com, bioisai@gmail.com, and pgalina04@cibnor.mx
3Conservación de Fauna del Noroeste, Ensenada, Baja California, C.P. 22785, Mexico.
E-mail: j_h_valdez@yahoo.com.mx
4Laboratorio de Herpetología, Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de
México, A.P. 70515, C.P. 04510, Mexico. E-mail: faustor@ib.unam.mx
River rocks as sleeping perches for Norops oxylophus and Basiliscus plumifrons
in the Cordillera de Talamanca, Costa Rica
Lizards (Sauria: Squamata) are an extraordinarily diversified group of reptiles, with over 6,000 species worldwide
(Uetz et al., 2016). In Costa Rica, they represent 36% (86) of all the reptile species; 37 of these lizards are anoles
(Norops, Dactyloa: Dactyloidae), and three of the four species of basilisks (Basiliscus: Corytophanidae) occur in the
country (Sasa et al., 2010; Hedges and Conn, 2012; G. Köhler 2010, 2011; G. Köhler and Vargas, 2010; G. Köhler
et al., 2014; J. Köhler et al., 2012; J. J. Köhler et al., 2015).
Anoles are one of the most successful of all the lizard groups (Pianka and Vitt, 2003). Williams (1983) used
the term “ecomorph” to describe anoles with specific morphologies associated with distinctive microhabitats on dif-
ferent Caribbean islands (Pianka and Vitt, 2003). Based on that assessment, anoles have served as highly successful
models for studies on community ecology (Pianka and Vitt, 2003). In addition, anoles also display phenomenally
high levels of species richness, along with ecomorphological variation in mainland Central- and South America
(Pinto et al., 2008). Anoles avoid self-competition by using different parts of the habitat, and thus occupy a vast
array of habitats with predictable evolutionary trajectories (Losos, 2009; Crandella et al., 2014). Consequently,
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Other Contributions
this taxonomically diversified and highly ecologically divergent group of Neotropical lizards has been considered
to constitute an ideal model system for addressing biological questions (D’Cruze and Stafford, 2006). They also
have been conceptualized as recurrently evolving toward the occupancy of a distinct set of niches (Losos et al.,
2003), wherein each species is associated with a specific suite of morphological and ecological characteristics
(Walguarnery et al., 2012).
The Stream Anole (Norops oxylophus) is a medium-large species that occurs at low elevations on the Atlantic
versant from northeastern Honduras to eastern Costa Rica (McCranie and Köhler, 2015). Formerly, some authors
have considered N. lionotus as a synonym of N. oxylophus, but here we follow McCranie and Köhler (2015, and
references therein) in recognizing these taxa as separate species. This partially aquatic anole has been reported to
occur in streamside habitats in humid tropical or subtropical forest and gallery forest, where individuals often are
found at the water’s edge (Fitch 1970, 1973; Savage, 2002). Fitch (1973) found most individuals perching on rocks
in a stream or in the rocky riparian zone below high water mark during the day; some individuals, however, were
found in dense vegetation, but always within 1–2 m of the water’s edge. This species also has been reported to take
refuge among the tree roots on stream banks or under debris on the adjacent forest floor, and at night to sleep on the
streamside vegetation (McCranie and Köhler, 2015).
The Green Basilisk (Basiliscus plumifrons) is a large arboreal lizard (Taylor, 1956) with a distribution that
extends from Honduras to Panama (Savage, 2002). This species inhabits lowland moist and wet forests and premon-
tane slopes, mainly along stream courses but especially when the streams are located within mature forests (Savage,
2002). This lizard usually is associated with water, with notable tendencies for aquatic and arboreal habits (Savage,
2002). Basiliscus plumifrons has been observed perching on several substrate types, including among boulders at
the edge of a swift mountain stream (Fitch, 1973). It also has been reported as fairly abundant in trees along river-
banks in the area of Tortuguero, in northeastern Costa Rica (Hirth, 1963). The biology of B. plumifrons is similar to
that of B. basiliscus (Savage, 2002). Individuals of the latter species sleep on perches from near ground level to 20
m above ground level, and adults tend to favor perches near or above watercourses (Van Devender, 1983).
Parque Nacional Barbilla (9°56'31"N, 83°25'12"W) was established in 1982 as a biological reserve, and
obtained its current status as a national park in 1997 (SINAC, 2016). Its topography consists of rough terrain in
a mountainous region, its surface area is composed of 11,943 ha, and its location mostly is bordered by three riv-
ers (Barbilla, Moravia, and Dantas) that largely form the boundary between the provinces of Cartago and Limón.
Presently, this park is the last forested area (and floral and faunal refuge) in the central Atlantic area of the Cordillera
de Talamanca (SINAC, 2016). This area receives a high amount of rainfall, and also experiences a wide a variety of
climatic regimes on account its elevation, which ranges from 110 to 1,617 m (SINAC, 2016). The reptile fauna of
the park has not been well documented, however, but theoretically should contain a high number of species.
We conducted a survey of the reptiles and other vertebrates at Parque Nacional Barbilla from 17 to 28 April
of 2017. During this time we searched along three rivers (Barbilla, San Miguel, Dantas) at night for several hours.
On the evenings of the 26th and 27th of April we hiked along the Río Dantas, near the indigenous community
of Tsinikicha On 26 April 2017 at 2130 h, we encountered a male N. oxylophus perched on a rock in the river
(09°59'13.4"N, 83°28'11.9"W; elev. 399 m), which protruded ca. 40 cm above the water level; a similar rock was
positioned next to it (Fig. 1). The stream at that point was ca. 12 m wide, with the two rocks located in the middle of
the river where the current was strong. The surrounding vegetation was gallery forest, with mature forest all around.
The following night at ca. 2200 h, we observed a juvenile B. plumifrons perched on a boulder along the side of the
river, but which extended well into the river and protruded ca. 2 m above the water level. The lizard was perched
inside a crevice in the rock, near the middle of the boulder and facing the flowing water (Fig. 2). To our knowledge,
this is the first observation of a nocturnal perch site for both N. oxylophus and B. plumifrons on a substrate with
these characteristics.
We found little information on the nocturnal perch sites of these two relatively common lizards. Some of this
information was anecdotal, such as that provided by Guyer and Donnelly (2005) when they discussed the capture
of this species. These authors indicated that B. plumifrons slept on small trees or shrubs along streams and rivers.
Additionally, Leenders, (2001) reported having observed juveniles basking on large boulders in a forest stream
during the day, and that they avoided capture by diving into the water and clinging to the underside of rocks, where
they hid in air-pockets; he also noted that this anole retreats into rock crevices during periods of inactivity, at night,
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or when the air temperature is low. Nonetheless, Leenders (2001) did not specify that N. oxylophus sleeps on rocks
in streams. Van Devender (1983) noted that the closely related B. basiliscus minimizes threats by sleeping on veg-
etation overhanging water, and when disturbed will fall or jump into the water.
Fig. 1. An adult male Stream Anole (Norops oxylophus) found sleeping on a rock in the middle of the Río Dantas in Parque
Nacional Barbilla, Costa Rica. '© Bárbara I. Escobar-Anleu
Fig. 2. A young Green Basilisk (Basiliscus plumifrons) found sleeping on the groove of a boulder along the Río Dantas in
Parque Nacional Barbilla, Costa Rica. '© Bárbara I. Escobar-Anleu
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421
Vitt et al. (1995) reported that individuals of N. oxylophus were observed using a variety of microhabitats
along streams in southeastern Nicaragua, including logs, the stems of grasses, leaves above the ground, leaf litter on
the ground, vines, and open ground. Lizards often were found at night perched on leaves or vines, usually above or
near the water. Four of 32 (12.5%) anoles were found sleeping on the cliff face of a waterfall (Vitt et al., 1995), but
these authors did not specify the sleeping substrate for these four individuals. Norops oxylophus also can be found
on moss-covered tree trunks during the day (Guyer and Donnelly, 2005). Norops aquaticus, an ecologically similar
species, also has been observed perching and feeding on rocks, logs, and low vegetation during the day (Savage,
2002). Differences between diurnal and nocturnal perches may result from different selective pressures during day
and night (Singhal et al., 2007).
Anole sleeping perches have been described in differing detail for several species in several countries.
Branches and the leaves of ferns, herbs, grasses, bushes, shrubs, and trees, have been reported as sleeping perches
for these lizards: 23 juveniles and seven adults of N. uniformis (as A. uniformis) were found sleeping on leaves of 14
species of plants (Cabrera-Guzmán and Reynoso, 2010). Preferences by certain species for particular types of sleep-
ing sites, such as structurally unstable perches, high perches, or warm perches, also have been reported (Mohanty et
al., 2016). Sleeping perch selection could have thermoregulatory implications (Christian et al., 1984), or influence
the timing of display behaviors to conspecifics in the morning (Andrews, 1971).
The most frequent perch types used by Basiliscus galeritus in Colombia were branches (46.15%) and leaves
(38.46%), although some individuals were observed on vines, stems, and on the ground (Hernández-Córdoba et
al., 2012). Small juveniles are more capable of using their ability to run on water (Glasheen and McMahon, 1996),
and thus they can use low perches to escape. Larger individuals, however, use higher perches so they can find other
escape routes (Hernández-Córdoba et al., 2012). Several other lizard species in tropical areas have been reported
to sleep on vegetation (Pianka and Vitt, 2003). On the Andaman Islands of India, near the Myanmar coast, most
individuals of two species of the genus Coryphophylax have been reported to sleep on leaves (Mohanty et al., 2016).
The use of rocks as sleeping perches by tropical lizards, therefore, apparently is an uncommon behavior.
Acknowledgments.––We conducted these observations during the Integrated Project course by Icomvis,
Universidad Nacional (UNA), Costa Rica. We are grateful for the logistical and financial support received from
UNA and SINAC. We thank Luis Ruedas for kindly reviewing an early draft of the manuscript. Louis W. Porras
provided valuable comments and suggestions that improved the manuscript.
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José m. mora1 and BárBara i. esCoBar-anLeu1,2
1Instituto Internacional en Conservación y Manejo de Vida Silvestre (ICOMVIS), Universidad Nacional (UNA), Heredia,
Costa Rica. Emails: josemora07@gmail.com and bisa_e8a9@hotmail.com (JMM, Corresponding author)
2Escuela de Biología, Universidad de San Carlos de Guatemala, Edificio T-10, Ciudad Universitaria, Zona 12, Ciudad
de Guatemala, Guatemala.
... Opportunistic use of these aquatic sites for sleep could potentially confer hydration and/or thermoregulatory advantages, as well as provide safety from terrestrial predators (Rall, 1985;Wikramanayake & Green, 1989;Doody et al., 2014). Additionally, rapid escape from predators has been argued as a benefit of terrestrial sleep sites situated close to or overhanging large rivers or lakes ( Avila-Pires, 1995;Doody et al., 2014;Mora & Escobar-Anleu, 2017). ...
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Divergent evolution of hemipenial morphology between closely related forms appears to be a common phenom-enon in mainland anoles. We discuss possible evolutionary scenarios for this finding, based on different lines of evidence from our work on Anolis polylepis and A. osa from southern Costa Rica. Molecular analysis suggests a very recent separa-tion of these two forms and the occurrence of hybridisation in their contact zone. Evidence from crossbreeding experi-ments supports these assumptions. The results of an examination of female cloacal morphology indicate co-evolution of male and female genital morphologies and thereby contradict functional neutrality of hemipenial morphology.
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Based on differences in coloration, morphometrics, and scalation, I recognize seven species of anoles related to Anolis pentaprion in Central America: Anolis beckeri (southeastern Mexico to northern Nicaragua), A. cristifer (Pacific lowlands of Chiapas to central Guatemala), A. fungosus (Caribbean premontane zone from central Costa Rica to western Panama), A. pentaprion (Caribbean lowlands of southern Nicaragua to northwestern Colombia, also on Pacific versant in central and eastern Panama), A. salvini (Pacific premontane and lower montane zones from central Costa Rica to western Panama), A. utilensis (restricted to Isla de Utila, Honduras), and a new species (from the Pacific lowlands of Costa Rica and western Panama) described herein. The new species differs from A. pentaprion by having a red male dewlap with few large, widely spaced gorgetal scales scattered across the dewlap (pink with purple pigment between rows of gorgetals, and with regular rows of numerous small, narrowly spaced gorgetal scales in A. pentaprion). Furthermore, I resurrect the name A. beckeri Boulenger for the pentaprion-like populations of anoles distributed from southeastern Mexico to northern Nicaragua. Anolis beckeri differs from A. pentaprion most obviously in the dorsal tail scalation (a serrated caudal crest present in A. pentaprion versus no such crest in A. beckeri) and dewlap scalation (gorgetal rows with 17–25 narrowly spaced scales in A. pentaprion versus gorgetal rows with 4–10 widely spaced scales in A. beckeri). Finally, SMF 83608 is proposed as the neotype for Anolis pentaprion. I provide standardized descriptions of A. beckeri, A. pentaprion, and the new species described herein.
Article
I describe a new species of Anolis related to Anolis (or Norops) altae from Volcán Tenorio, Costa Rica. The new species differs from all other species in the A. altae complex by having the ventral surfaces of body and limbs distinctly reticulated with dark brown pigment and a male dewlap that is dark red with brown blotches, as well as in several morphometric and pholidotic characteristics. It further differs from its geographically nearest congener of this complex, A. monteverde, by having a bilobate hemipenis (unilobate in A. monteverde).
Article
Adaptive radiation, which results when a single ancestral species gives rise to many descendants, each adapted to a different part of the environment, is possibly the single most important source of biological diversity in the living world. One of the best-studied examples involves Caribbean Anolis lizards. With about 400 species, Anolis has played an important role in the development of ecological theory and has become a model system exemplifying the integration of ecological, evolutionary, and behavioral studies to understand evolutionary diversification. This major work, written by one of the best-known investigators of Anolis, reviews and synthesizes an immense literature. Jonathan B. Losos illustrates how different scientific approaches to the questions of adaptation and diversification can be integrated and examines evolutionary and ecological questions of interest to a broad range of biologists.
Article
In this review, we provide thorough descriptions of the 39 named species of the family Dactyloidae (commonly referred to as anoles) we recognize as occurring in Honduras. We follow two recent phylogenetic analyses suggesting eight genera of anoles are recognizable, two of which occur in Honduras (Anolis and Norops). Each species account contains a synonymy, statements on its geographic distribution, description, diagnosis/similar species statements, a list of illustrations, a remarks section, natural history comments, an etymology section, and a list of specimens examined; for a few species, a list of other records is included. The synonymy for each species includes the original description of the species with the original proposed combination, the type specimen(s), and direct quotes of the species' type locality. Each synonymy also includes the first use of the currently used scientific name and all known references to Honduran specimens and/or localities. Distribution maps with the known Honduran localities plotted are included for each species (with one exception), and other records are included on a few of those maps. Color photographs of an adult of each species are included as well as color photographs of the male dewlap of all species. Following the species accounts are two dichotomous identification keys (English and Spanish versions) to help the reader identify any specimen in hand. Following the identification keys is a section on species group relationships of each species known from the country. A distribution section that contains the distribution of each anole species in Honduras by department, forest formation, elevation, physiographic region, and ecophysiographic area; broad patterns of geographic distribution; historical units; and a discussion of Honduras as a distributional endpoint. A section on conservation discusses vulnerability gauges, IUCN Red List categories, and a section detailing each species' occurrence in Honduran protected areas.