herpetofauna of the santa cruz forest reserve in the

Introduction

The Amazon Basin is one of the most complex and biodiverse regions on the planet (Kauffmann-Zeh, 1999; Myers et al., 2000). The extent of this biodiversity is so pronounced that new species across different taxonomic groups are discovered each year within the basin (e.g., ferns: Cárdenas et al., 2016; insects: Perry et al., 2016; fish: Oberdorff et al., 2019). Much of this biodiversity is imperiled, however, because of anthropogenic activities (Barlow et al., 2016). Deforestation and habitat degradation as a result of increased mining and oil drilling (Finer et al., 2008; Espejo et al., 2018), agricultural practices (Glinskis and Gutiérrez-Vélez, 2019), and

cattle ranching (Veiga et al., 2002) are major ecological concerns. These industries require large swaths of disturbed landscapes, which negatively affects wildlife populations and, therefore, the existence of species and whole communities that live in the basin (Barlow et al., 2016). For example, oil palm plantations fragment habitats, which support less diverse communities of wildlife and contribute to polluted waterways (Vijay et al., 2016). To mitigate these effects, conservation areas such as national forests and ecological stations must be established and maintained throughout the Amazon because of the extensive endemism that occurs across different areas of the basin (da Silva et al., 2005; Peres, 2005; Freitas et al., 2017). In order to best conserve this rich biodiversity through the further establishment of conservation priorities, we must first understand community compositions within the different regions that make up the Amazon Basin.

Amphibians and reptiles make up a significant proportion of tetrapods in the Amazon and are, therefore, of significant conservation interest, although their presence can be obscured by their small size and/or cryptic behavioural patterns. Data on the biogeography, conservation status, and general life history of many

Herpetology Notes, volume 13: 753-767 (2020) (published online on 21 September 2020)

Herpetofauna of the Santa Cruz Forest Reserve in the Peruvian Amazon Basin

Matthew F. Metcalf1,*, Alexander Marsh1, Emerson Torres Pacaya2, Devon Graham2, and Charles W. Gunnels IV1

1 Department of Biological Sciences, Florida Gulf Coast University, 10501 FGCU Blvd S., Fort Myers, Florida 33965, USA.

2 Project Amazonas, 701 E. Commercial Blvd, #200, Fort Lauderdale, Florida 33334, USA.

* Corresponding author. E-mail: [email protected]

Abstract. The Amazon is the largest rainforest on the planet and is known for its biodiversity despite growing deforestation from agricultural development, urbanisation, wildfires, and unregulated mining. It is increasingly important to better understand regionalised biodiversity through fauna inventories because of these anthropogenic pressures. We conducted this study between May 2019 and August 2019 to identify herpetofauna biodiversity at the Santa Cruz Forest Reserve in Loreto, Peru. During the study period, we recorded amphibians and reptiles via visual encounter surveys and opportunistic encounters. We further supplemented these sightings by secondary data compiled from previous observations. In total, we recorded 200 species of herpetofauna at this reserve: 15 families of amphibian comprising 106 species, and 21 families of reptiles comprising 94 species. The visual encounter surveys adequately sampled the diversity of amphibians that would have been expected given this protocol and the timeframe of the study. In contrast, this procedure did not sample reptiles as well in terms of both species richness and diversity; more reptile species would be expected given additional sampling. The combined approach of this study, however, provided a robust inventory of the Santa Cruz Forest Reserve, which can be used to support future on-site studies and management plans throughout the region.

Keywords. Amphibians, Reptiles, Amazon, Peru, Inventory, Reserve

Matthew F. Metcalf et al.754

Amazonian herpetofauna are severely lacking, despite the fact that amphibians and reptiles are key indicators of environmental disruptions (Stuart et al., 2004). For example, these taxa have highlighted the effect of globally infectious diseases such as chytrid fungus (Skerratt et al., 2007; Russell et al., 2019) and the direct biodiversity loss in frogs (Scheele et al., 2019), as well as the indirect biodiversity collapse in tropical snakes (Zipkin et al., 2020). As industrialisation and agricultural activities advance throughout the Amazon, reductions and complete losses of amphibian and reptile populations are expected (Nobre et al., 2016). Consequently, biodiversity inventories are crucial to evaluate conservation status of herpetofauna communities (Oliveira et al., 2017; Shaney et al., 2017), identify regional endemism and new species (da Silva et al., 2005; Treurnicht et al., 2017; de Oliveira et al., 2020), and develop proactive conservation management strategies that mitigate the most destructive effects of deforestation (Groves et al., 2002; Guralnick et al., 2018). Herpetofauna surveys in the Amazon Basin are critical to create community baseline structures to assist future research and identify conservation management priorities (Fonseca et al., 2019).

The Peruvian Amazon is one of the most biologically diverse regions within Amazonia (Rodríguez and Knell, 2004; Finer et al., 2008). Peru has among the highest number of newly discovered amphibian species globally (Catenazzi and Bustamante, 2007; Wake and Vredenburg, 2008; Lehr et al., 2017; Santa-Cruz et al.,

2019). In addition, the Peruvian Amazon is expected to have among the world’s most diverse species richness of amphibian and reptile communities (Vasconcelos et al., 2012; Roll et al., 2017). This is also a rapidly developing area. For example, the human population in the province of Loreto has grown by 28.6% from 1993 to 2017 and the urban core of Iquitos, an otherwise isolated city in the eastern Peruvian Amazon, has grown by 26.4% from 2000 to 2015 (INEI, 2020). To add to a growing list of herpetofaunal inventories throughout Loreto (see Table 1), this study describes the composition of herpetofauna in the Santa Cruz Forest Reserve in Loreto, Peru. These valuable insights into amphibian and reptile community structures will provide baseline data for future studies, as well as detail the ecological and geographic importance of the Santa Cruz Forest Reserve.

Materials and Methods

Study site.—The Santa Cruz Forest Reserve (-3.5214ºS, -73.1803ºW; 115.82 m a.s.l.) is owned and operated by Project Amazonas, which purchased the first parcel in 2008 (Fig. 1). Additional plots of primary and secondary forest and agricultural lands were gradually incorporated to create 232 hectares of contiguous protected area. The reserve is located approximately 22 km northeast of the city of Iquitos. Bordering the reserve is an agricultural property used for cattle ranching and yuca (cassava) and plantain farming. There are also three adjacent community reserves which increase the forest continuity outside of the Santa Cruz

Table 1. Herpetofauna observed in selected rapid biological inventories from nearby regions within Loreto, Peru, including observations from the Santa Cruz Forest Reserve. LOCATION Ns An Ca Gy Ch Cr Lz Am Sn SOURCE

Ampiyacu, Apayacu, Yaguas, Medio Putumayo

104 62 1 1 3 3 19 - 15 Rodríguez and Knell (2004)

Sierra del Divisor 109 67 1 – 2 1 17 – 21 Barbosa de Souza and Rivera (2006)

Mastés 109 74 – – 2 2 18 – 13 Gordo et al. (2006)

Nanay-Mazán-Arabela 93 53 1 – 2 2 23 – 12 Catenazzi and Bustamante (2007)

Cuyabeno-Güeppí 107 59 – – 3 3 22 1 19 Yánez-Muñoz and Venegas (2008)

Maijuna 108 63 1 2 1 1 22 – 18 von May and Venegas (2010)

Yaguas-Cotuhé 128 73 1 1 4 3 21 – 25 von May and Mueses-Cisneros (2011)

Ere-Campuya-Algodón 128 68 – – 4 2 22 1 31 Venegas and Gagliardi-Urrautia (2013)

Medio Putumayo-Algodón 142 89 1 – 4 3 20 – 25 Chávez and Mueses-Cisneros (2016)

Santa Cruz Forest Reserve 200 102 1 3 3 2 34 1 54 This study

Abbreviations: Ns – number of species; An – Anuran; Ca – Caudata; Gy – Gymnophiona; Ch – Chelonian; Cr – Crocodylian; Lz – Lizard; Am – Amphisbaenian; Sn – Snake.

Table 1. Herpetofauna observed in selected rapid biological inventories from nearby regions within Loreto, Peru, including observations from the Santa Cruz Forest Reserve.

Forest Reserve boundaries. Much of the reserve is primary forest (~60%) and the rest is secondary forest, which is being actively restored through the planting of native primary forest vegetation.

Sampling.—Sampling of herpetofauna occurred between May 2019 and August 2019 using visual encounter surveys (VES) along nine different 250-meter transects throughout the reserve. We conducted VES similarly to methods used by Doan (2003), which did not include time constraints. Observers walked at a reasonable pace but used as much time as necessary to sufficiently sample each transect. VES transects served to sample amphibians and reptiles found on the ground and within the lower canopy. We then conducted transects during multiple time periods to best capture herpetofauna across varying activity patterns. We aggregated diurnal surveys into four time slots: 06:00 – 08:00 h, 08:00 – 10:00 h, 10:00 – 12:00 h, and 12:00 – 14:00 h. Similarly, we conducted nocturnal surveys across four time slots: 18:00 – 20:00 h, 20:00 – 22:00 h, 22:00 – 24:00 h, and 24:00 – 02:00 h.

We conducted transects across habitat types found at the reserve and adjacent pasture lands with three replicate sampling transects for each habitat type (primary forest, secondary forest, and pasture lands), resulting in nine unique transects. Over the entire research period, we sampled each transect once per diurnal and nocturnal time slot so that every replicate transect was sampled across all four diurnal and all four nocturnal time slots.

Our data include species collected by opportunistic encounters (OE) during the timeframe of this study in 2019. This sampling type represents any species found during the overall stay at Santa Cruz (May to August 2019) but outside of the organised transects. For example, we captured one Drymarchon corais while hiking an unused trail. We recorded this observation as OE because researchers captured the snake outside of a designated VES transect but within the boundaries of the Santa Cruz Forest Reserve during the timeframe of this study. Regardless of sampling method, we attempted to capture all encountered animals and then photograph the individual to confirm species identification. We safely released all specimens at or near the point of capture. In order to supplement our data, we included secondary data (SD) contributed by Project Amazonas. Secondary data included the observation of herpetofauna species - whether they were visual, through photographs, or voucher specimens - accumulated and vetted by Project Amazonas staff and/or other researchers since acquiring the property in 2008.

Data analysis.—We ran statistical analyses in R (R Core Team, 2017) using the vegan package (Oksanen et al., 2007) to create a species accumulation curve of all amphibians and reptiles found during visual encounter

Herpetofauna of the Santa Cruz Forest Reserve in the Peruvian Amazon Basin 755

Figure 1. Top map shows the position of Santa Cruz Forest Reserve relative to Iquitos and surrounding communities along the Amazon River Basin in Peru. Bottom map shows the borders of the Santa Cruz Forest Reserve (outlined in white) with adjacent agricultural pasture (outlined in yellow).

surveys. We also extrapolated species richness and diversity for animals sampled during the VES because observed species frequently underestimates real species richness (Hughes et al., 2001). We measured species richness and diversity indices in terms of HILL numbers (Hill, 1973), which provides multiple benefits in context of this study (Chao et al., 2014). HILL numbers show expected doubling property when combining independent species assemblages. In addition, they are defined in terms of expected numbers of species, providing a clear and biologically relevant unit for evaluation and comparison (Chao et al., 2014). We extrapolated estimates of species richness and diversity for the first three HILL numbers based on separate evaluations of all collected amphibian and reptiles using the iNEXT package (Hsieh et al., 2016). We used this approach to estimate species richness, the Shannon diversity index, and the Simpson diversity index, which were represented by q = 0, q = 1, q = 2 respectively.

Results

We surveyed herpetofauna throughout the Santa Cruz Forest Reserve from May to August 2019. Over this period, we conducted 72 VES transects, totalling 170.3 person-hours of sampling effort. Nocturnal surveys required more time (x̄ = 65.89 min per 250-m transect) than diurnal surveys (x̄ = 32.69 min per 250-m transect). We recorded additional species opportunistically during the study timeline and from previous records collected by researchers and staff at Santa Crus Forest Reserve since 2008.

We found 102 herpetofauna species during the duration of this study. Of the 571 amphibians we sampled across the 72 VES transects, all were anurans except for one caudata (Bolitoglossa altamazonica). By comparison, we observed only 110 reptiles during the same VES transects. All of these reptiles were lizards, except for five snake species. During the VES, we found Bothrops atrox, Chironius scurrulus, and Imantodes cenchoa on one occasion each, and Micrurus surinamensis and Siphlophis compressus on two occasions each. Amphibians found during VES and OE observations accounted for 53% of species (54 species) across 11 families (Fig. 2, Table 2) and reptiles made up 47% of species (48 species) within 17 families (Fig. 3, Table 3). For amphibians, we observed 41.5% of species exclusively during VES transects and 24.5% exclusively during opportunistic encounters. Overall, we found 75.5% of amphibian species during VES transects and 56.6% in opportunistic encounters. Reptiles, however,

Figure 2. Sample of amphibians found at the Santa Cruz Forest Reserve. (A) Rhinella dapsilis, (B) Vitreorana ritae, (C) Ranitomeya ventrimaculata, (D) Dendropsophus triangulum, (E) Boana punctata, (F) Osteocephalus mutabor, (G) Lithodytes lineatus, (H) Chiasmocleis bassleri, (I) Phyllomedusa bicolor, (J) Pipa snethlageae. Photos by Alexander Marsh.


trended differently. We recorded only one species (Chironius scurrulus; 2%) exclusively in VES transects. During opportunistic encounters, we observed 51% of the reptile species exclusively. Overall, we documented 45% of reptiles across both methods.

Secondary data from previous observations added 52 species of amphibians to the Santa Cruz Forest Reserve’s list, which includes one new order (Gymnophiona) and four new families. Secondary data also added 46 reptile species to the list which includes one new order (Amphisbaenia) and three new families. The VES and OE data from this study added 16 new species of amphibians and nine new species of reptiles that had not been recorded in the Santa Cruz Forest Reserve previously.

The upward trajectory of the species accumulation curve suggests that the herpetofauna community that could be found during the 72 VES transects was more diverse than the 102 observed species (Fig. 4A). This trend in the species accumulation curve seemed to be a result of reptile sampling. The 72 transects appeared to be deficient in identifying the species richness of reptiles that would be found with VES transects (Fig. 4B). The 110 individually observed reptiles represented 24 species. However, the iNEXT extrapolation of species richness (q = 0) suggests that an additional 13 species would be found (37.4 species ± 26.87–86.34 95% CI) after sampling at least 700 animals. In contrast, visual encounter surveys appeared to be a stronger measure for recording the species richness and diversity of amphibians (Fig. 4B). The iNEXT extrapolation curves suggest that amphibian diversity would approach its asymptote only after the number of collected animals doubled from the 571 amphibians across the 42 species that we found during VES transects. However, this extra sampling would be expected to discover only two additional amphibian species (44.4 species ± 42.46–55.09 95% CI). Observed measures of amphibian and reptile diversity (Shannon index, q = 1 and Simpson index, q = 2) were closer to the expected asymptotes based on the number of collected animals.

Discussion

Of the 200 species observed at the Santa Cruz Forest Reserve, seven species are listed by the IUCN Red List as either data deficient or vulnerable. Five amphibians are listed as data deficient (Allobates conspicuus, Pristimantis delius, Ranitomeya amazonica, Ranitomeya variabilis, and Vitreorana ritae) and two reptiles (turtles) listed as vulnerable (Chelonoidis denticulata and

Figure 3. Sample of reptiles found at the Santa Cruz Forest Reserve. (A) Paleosuchus trigonatus, (B) Anolis transversalis, (C) Copeoglossum nigropunctatum, (D) Plica plica, (E) Epicrates cenchria, (F) Atractus snethlageae, (G) Siphlophis compressus, (H) Micrurus suranimensis, (I) Lachesis muta, (J) Podocnemis unifilis. Photos by Alexander Marsh.


Podocnemis unifilis). However, the true conservation status of many of these species remains unclear as most are either unstudied or very poorly studied. In addition, taxonomic determinations remain an ever-evolving and prevalent issue for many Amazonian amphibians and reptiles. As cryptic species continue to be recognised (e.g., Elmer and Cannatella, 2008; Sturaro et al., 2017), it is expected the species richness in this biological hotspot will increase.

The VES was the most rigorous methodology used in this study, although this method sampled only those animals that would make use of the ground and/or forest understory. Other microhabitats, such as the mid- and upper-canopy and aquatic environments, could not be sampled with this procedure, which explains why we encountered fewer species with VES than with the opportunistic encounters and secondary data. Given the sampling pressure utilised in this study, VES transects were extremely successful in sampling the amphibian community. However, the reptile community curves suggest more species would be observed with additional sampling. The ability of the VES to sample a relatively complete amphibian community likely reflects that frogs were much more numerous than the entire class of reptiles. The addition of opportunistic encounters and secondary data produced a more robust collection of species found in the reserve, yet these procedures lack methodological rigor due to varying factors, such as observer expertise and variable sampling intensities. We encourage continued floral and faunal inventories in this area, especially when supplemented by additional and complimentary sampling methods. For example, studies have shown micro-environmental characteristics may influence niche distributions in groups (Menin et al., 2007) such as Pristimantis (López-Rojas et al., 2015). Future studies are planned to incorporate these new methods into identifying a more complete herpetofauna community at this location.

As predicted for the Peruvian Amazon (Vasconcelos et al., 2012; Roll et al., 2017), the Santa Cruz Forest Reserve showed a high degree of herpetological biodiversity despite its small size. Several rapid biological inventories have recently documented herpetofauna communities throughout the Loreto region surrounding the Santa Cruz Forest Reserve (Table 1). This study’s approach detected a comparably high species richness in reference to these inventories, despite our study involving only one location, a significantly smaller study area, and lacking multiple capture methods. The Santa Cruz Forest Reserve is centrally located between many of these rapid inventories and may serve as an important

habitat corridor between sites. We did, however, observe very few snakes during VES transects, which may be a result of their secretive habits and movement patterns in addition to transects being conducted over a short timeframe. Further research should reveal a more concrete understanding of snake assemblages in this area.

Tropical forests remain at the forefront of biodiversity conservation concerns due to increasing anthropogenic threats (de Solar et al., 2016). The rainforests in the western Amazon, and the Peruvian Amazon Basin in particular, represent one of the largest remaining

Figure 4. Species diversity for amphibians and reptiles found during visual encounter surveys conducted in the Santa Cruz Forest Reserve. (A) Species accumulation curve (± 95% CI) for all herpetofauna observed during the consecutive visual encounter survey transects. (B) Sample interpolation (solid lines) and extrapolation (dashed lines) curves (± 95% CI) based on the number of sampled amphibians and reptiles, respectively. Three measures of diversity are illustrated in the panels: 0 = Species richness, 1 = Shannon index of biodiversity, and 2 = Simpson index of biodiversity.


Table 2. Amphibians recorded at the Santa Cruz Forest Reserve in Loreto, Peru. Record Types: VES = Visual Encounter Survey; OE = Opportunistic Encounter; SD = Secondary Data. IUCN Red List rankings: DD = Data Deficient, LC = Least Concern, NL = Not Listed, VU = Vulnerable. * indicates species only found in adjacent pasture land surveys.

Table 2.

SPECIES VES OE SD IUCN ANURA Aromobatidae

Allobates conspicuus (Morales, 2002) X X X DD

Allobates femoralis (Boulenger, 1884) X X X LC

Allobates marchesianus (Melin, 1941) X LC

Bufonidae

Rhaebo glaberrimus (Günther, 1869) X LC

Rhaebo guttatus (Schneider, 1799) X LC

Rhinella dapsilis (Myers and Carvalho, 1945) X X X LC

Rhinella margaritifera (Laurenti, 1768) X X X LC

Rhinella marina (Linnaeus, 1758) X X X LC

Centrolenidae

Vitreorana ritae (Lutz, 1952)* X DD

Ceratophryidae

Ceratophrys cornuta (Linnaeus, 1758) X LC

Craugastoridae

Niceforonia nigrovittata (Andersson, 1945) X LC

Noblella myrmecoides (Lynch, 1976) X LC

Oreobates quixensis Jiménez de la Espada, 1872 X X X LC

Pristimantis academicus Lehr et al., 2010 X NL

Pristimantis acuminatus (Schreve, 1935) X LC

Pristimantis altamazonicus (Barbour and Dunn, 1921) X X LC

Pristimantis carvalhoi (Lutz, 1952) X X LC

Pristimantis conspicillatus (Günther, 1858) X LC

Pristimantis croceoinguinis (Lynch, 1968) X X X LC

Pristimantis delius (Duellman and Mendelson, 1995) X DD

Pristimantis diadematus (Jiménez de la Espada, 1872) X LC

Pristimantis lanthanites (Lynch, 1975) X LC

Pristimantis malkini (Lynch, 1980) X LC

Pristimantis martiae (Lynch, 1974) X X X LC

Pristimantis ockendeni (Boulenger, 1912) X X X LC

Pristimantis orcus (Lehr et al. 2009) X LC

Pristimantis padiali (Moravec et al., 2010) X LC

Pristimantis peruvianus (Melin, 1941) X LC

Pristimantis variabilis (Lynch, 1968) X X LC

Pristimantis ventrimarmoratus (Boulenger, 1912) X LC

Strabomantis sulcatus (Cope, 1874) X X LC

Dendrobatidae

Ameerega hahneli (Boulenger, 1884) X X X LC

Ameerega trivittata (Spix, 1824) X LC


SPECIES VES OE SD IUCN Ranitomeya amazonica (Schulte, 1999) X DD Ranitomeya ventrimaculata (Shreve, 1935) X X LC

Ranitomeya variabilis (Zimmermann and Zimmermann, 1988) X DD

Hemiphractidae

Hemiphractus helioi (Sheil and Mendelson, 2001) X LC

Hemiphractus proboscideus (Jiménez de la Espada, 1870) X LC

Hylidae

Boana boans (Linnaeus, 1758) X X LC

Boana calcarata (Troschel in Schomburgk, 1848) X X LC

Boana cinerascens (Spix, 1824) X LC

Boana fasciata (Günther, 1859) X LC

Boana geographicus (Spix, 1824) X LC

Boana lanciformis (Cope, 1871) X X X LC

Boana microderma (Pyburn, 1977) X LC

Boana nympha (Faivovich et al. 2006) X X LC

Boana punctata (Schneider, 1799) X X LC

Dendropsophus brevifrons (Duellman and Crump, 1974) X LC

Dendropsophus koechlini (Duellman and Trueb, 1989) X LC

Dendropsophus leucophyllatus (Beireis, 1783) X LC

Dendropsophus marmoratus (Laurenti, 1768) X LC

Dendropsophus minutus (Peters, 1872) X LC

Dendropsophus miyatai (Vigle and Goberdhan-Vigle, 1990)* X LC

Dendropsophus parviceps (Boulenger, 1882) X LC

Dendropsophus rhodopeplus (Günther, 1858) X LC

Dendropsophus rossalleni (Goin, 1959) X LC

Dendropsophus sarayacuensis (Shreve, 1935) X LC

Dendropsophus triangulum (Günther, 1869)* X X X LC

Nyctimantis rugiceps (Boulenger, 1882) X LC

Osteocephalus buckleyi (Boulenger, 1882) X X LC

Osteocephalus cabrerai (Cochran and Goin, 1970) X X LC

Osteocephalus deridens (Jungfer et al., 2000) X LC

Osteocephalus leprieurii (Duméril and Bibron, 1841) X LC

Osteocephalus mutabor (Jungfer and Hödl, 2002) X X LC

Osteocephalus planiceps (Cope, 1874) X X X LC

Osteocephalus taurinus Steindachner, 1862 X X X LC

Osteocephalus yasuni (Ron and Pramuk, 1999) X LC

Scarthyla goinorum (Bokermann, 1962) X LC

Scinax cruentommus (Duellman, 1972) X LC

Scinax funereus (Cope, 1874) X LC

Scinax garbei (Miranda-Ribeiro, 1926) X LC

Scinax ruber (Laurenti, 1768) X LC

Sphaenorhynchus dorisae (Goin, 1957) X LC

Table 2. Continued.Table 2.





Bufonidae






Centrolenidae


Ceratophryidae


Craugastoridae






















Dendrobatidae




Table 2. Continued.

SPECIES VES OE SD IUCN Trachycephalus coriaceus (Peters, 1867) X LC Trachycephalus cunauaru (Gordo et al., 2013) X NL

Trachycephalus resinifictrix (Goeldi, 1907) X X LC

Trachycephalus typhonius (Linnaeus, 1758) X LC

Leptodactylidae

Adenomera andreae (Müller, 1923) X X X LC

Adenomera hylaedactyla (Cope, 1868) X X LC

Engystomops petersi (Jiménez de la Espada, 1872) X LC

Hydrolaetare schmidti (Cochran and Goin, 1959)* X LC

Leptodactylus bolivianus (Boulenger, 1898) X LC

Leptodactylus diedrus (Heyer, 1994) X X LC

Leptodactylus discodactylus (Boulenger, 1884) X LC

Leptodactylus leptodactyloides (Andersson, 1945) X LC

Leptodactylus pentadactylus (Laurenti, 1768) X X LC

Leptodactylus rhodonotus (Günther, 1869) X LC

Leptodactylus stenodema (Jiménez de la Espada, 1875) X LC

Leptodactylus wagneri (Peters, 1862) X X X LC

Lithodytes lineatus (Schneider, 1799) X X X LC

Pseudopaludicola ceratophyes (Rivero and Serna, 1985) X LC

Microhylidae

Chiasmocleis bassleri (Dunn, 1949) X X X LC

Chiasmocleis tridactyla (Duellman and Mendelson, 1995) X X X LC

Ctenophryne geayi (Mocquard, 1904) X LC

Hamptophryne boliviana (Parker, 1927) X X LC

Synapturanus rabus (Pyburn, 1977) X LC

Phyllomedusidae

Cruziohyla craspedopus (Funkhouser, 1957) X LC

Phyllomedusa bicolor (Boddaert, 1772) X X LC

Phyllomedusa tarsius (Cope, 1868) X X LC

Phyllomedusa vaillantii (Cope, 1868) X LC

Pipidae

Pipa pipa (Linnaeus, 1758) X X LC

Pipa snethlageae (Müller, 1914) X LC

CAUDATA

Plethodontidae

Bolitoglossa altamazonica (Cope, 1874) X X LC

GYMNOPHIONA

Caeciliidae

Siphonops annulatus (Mikan, 1822) X LC

Microcaecilia albiceps (Boulenger, 1882) X LC

Typhlonectidae

Typhlonectes compressicauda (Duméril and Bibron, 1841) X LC

Table 2.





Bufonidae






Centrolenidae


Ceratophryidae


Craugastoridae






















Dendrobatidae




Table 3. Reptiles recorded at the Santa Cruz Forest Reserve in Loreto, Peru. Record Types: VES = Visual Encounter Survey; OE = Opportunistic Encounter; SD = Secondary Data. IUCN Red List rankings: DD = Data Deficient, LC = Least Concern, NL = Not Listed, VU = Vulnerable. * indicates species only found in adjacent pasture land surveys.

Table 3.

SPECIES VES OE SD IUCN AMPHISBAENIA Amphisbaenidae

Amphisbaena alba (Linnaeus, 1758) X LC

CROCODYLIA

Alligatoridae

Caiman crocodilus (Linnaeus, 1758) X LC

Paleosuchus trigonatus Schneider, 1801 X X LC

SQUAMATA: SAURA

Alopoglossidae

Alopoglossus angulatus (Linnaeus, 1758) X X LC

Alopoglossus atriventris (Duellman, 1973) X X X LC

Alopoglossus buckleyi (O’Shaughnessy, 1881) X X X LC

Ptychoglossus brevifrontalis (Boulenger, 1912) X LC

Dactyloidae

Anolis bombiceps Cope, 1875 X X X NL

Anolis fuscoauratus D’Orbigny, 1837 X X X NL

Anolis ortonii Cope, 1868 X X X NL

Anolis punctatus Daudin, 1802 X LC

Anolis trachyderma Cope, 1876 X X X NL

Anolis transversalis Duméril, 1851 X X X NL

Gekkonidae

Hemidactylus mabouia Moreau de Jonnés, 1818 X NL

Gymnophthalmidae

Arthrosaura reticulata (O’Shaughnessy, 1881) X X X LC

Cercosaura argulus (Peters, 1862) X X X LC

Cercosaura ocellata (Wagler, 1830) X LC

Cercosaura oshaughnessyi (Boulenger, 1885) X X NL

Iphisa elegans Gray, 1851 X LC

Loxopholis parietalis (Cope, 1886) X LC

Potamites ecpleopus (Cope, 1875) X LC

Hoplocercidae

Enyalioides laticeps (Guichenot, 1855) X X X LC

Enyalioides microlepis (O’Shaughnessy, 1881) X LC

Mabuyidae

Copeoglossum nigropunctatum (Spix, 1825) X X X LC

Phyllodactylidae

Thecadactylus solimoensis (Bergmann and Russell, 2007) X X LC

Polychrotidae

Polychrus marmoratus (Linnaeus, 1758) X LC

Sphaerodactylidae

Gonatodes concinnatus (O’Shaughnessy, 1881) X X X LC

Table 2.





Bufonidae






Centrolenidae


Ceratophryidae


Craugastoridae






















Dendrobatidae




Table 3. Continued.

SPECIES VES OE SD IUCN Gonatodes humeralis (Guichenot, 1855) X X X LC Pseudogonatodes guianensis Parker, 1935 X X X LC

Teiidae

Ameiva ameiva (Linnaeus, 1758) X X LC

Kentropyx altamazonica Cope, 1876 X LC

Kentropyx pelviceps Cope, 1868 X X X LC

Tupinambis teguixin (Linnaeus, 1758) X

Tropiduridae

Plica plica (Linnaeus, 1758) X X NL

Plica umbra (Linnaeus, 1758) X X LC

Stenocercus fimbriatus (Avila-Pires, 1995) X X LC

Uracentron flaviceps (Guichenot, 1855) X LC

SQUAMATA: SERPENTES

Boidae

Boa constrictor Linnaeus, 1758 X LC

Corallus caninus (Linnaeus, 1758) X LC

Corallus hortulanus (Linnaeus, 1758) X X LC

Epicrates cenchria (Linnaeus, 1758) X X NL

Eunectes murinus (Linnaeus, 1758) X NL

Colubridae

Chironius carinatus (Linnaeus, 1758) X X NL

Chironius exoletus (Linnaeus, 1758) X LC

Chironius fuscus (Linnaeus, 1758) X LC

Chironius multiventris (Schmidt and Walker, 1943) X LC

Chironius scurrulus (Wagler, 1824) X LC

Dendrophidion dendrophis (Schlegel, 1837) X LC

Drymarchon corais (Boie, 1827) X X LC

Drymoluber dichrous (Peters, 1863) X X LC

Leptophis ahaetulla (Linnaeus, 1758) X LC

Mastigodryas boddaerti (Sentzen, 1796) X LC

Oxybelis fulgidus (Daudin, 1803) X LC

Phrynonax poecilonotus (Günther, 1858) X LC

Spilotes pullatus (Linnaeus, 1758) X X LC

Spilotes sulphureus (Wagler, 1824) X LC

Dipsadidae

Atractus collaris (Peracca, 1897) X LC

Atractus major (Boulenger ,1894) X LC

Atractus snethlageae (da Cunha and do Nascimento, 1983) X LC

Clelia celia (Daudin, 1803) X X LC

Dipsas catesbyi (Sentzen, 1796) X X LC

Dipsas indica (Laurenti, 1768) X LC

Dipsas pavonina (Schlegel, 1837) X LC

Table 2.





Bufonidae






Centrolenidae


Ceratophryidae


Craugastoridae






















Dendrobatidae




wilderness areas, and boasts ample species richness (Rabosky et al., 2019). While habitat loss and fragmentation continue to occur throughout Amazonia,

there is growing scientific and social demand to mitigate negative effects on biodiversity and ecosystem services (Haila, 2002). In particular, preserved lands

Table 3. Continued.Table 2.





Bufonidae






Centrolenidae


Ceratophryidae


Craugastoridae






















Dendrobatidae



SPECIES VES OE SD IUCN Drepanoides anomalus (Jan, 1863) X LC Erythrolamprus reginae (Linnaeus, 1758) X X LC

Erythrolamprus typhlus (Linnaeus, 1758) X LC

Helicops angulatus (Linnaeus, 1758) X X LC

Helicops polylepis (Günther, 1861) X LC

Hydrops martii (Wagler, 1824) X LC

Imantodes cenchoa (Linnaeus, 1758) X X X LC

Imantodes lentiferus (Cope, 1894) X LC

Oxyrhopus formosus (Wied-Neuwied, 1820) X NL

Oxyrhopus melanogenys (Tschudi, 1845) X NL

Oxyrhopus petolarius (Linnaeus, 1758) X X LC

Oxyrhopus vanidicus (Lynch, 2009) X LC

Philodryas argentea (Daudin, 1803) X LC

Pseudoboa coronata (Schneider, 1801) X X LC

Siphlophis cervinus (Laurenti, 1768) X LC

Siphlophis compressus (Daudin, 1803) X X X LC

Taeniophallus brevirostris (Peters, 1863) X LC

Xenodon rabdocephalus (Wied-Neuwied, 1824) X LC

Elapidae

Micrurus hemprichii (Jan, 1858) X LC

Micrurus filiformis (Günther, 1859) X LC

Micrurus langsdorfii (Wagler, 1824) X LC

Micrurus lemniscatus (Linnaeus, 1758) X LC

Micrurus spixii (Wagler, 1824) X LC

Micrurus surinamensis (Cuvier, 1817) X X X LC

Viperidae

Bothrops atrox (Linnaeus, 1758) X X X NL

Bothrops bilineatus (Wied-Neuwied, 1821) X NL

Bothrops taeniatus (Wagler, 1824) X LC

Lachesis muta (Linnaeus, 1766) X X NL

TESTUDINES

Chelidae

Platemys platycephala (Schneider, 1792) X X NL

Podocnemididae

Podocnemis unifilis (Troschel, 1848) X X VU

Testudinidae

Chelonoidis denticulata (Linnaeus, 1766) X VU


and primary forests are increasingly important for protecting biodiversity throughout the tropics (Gibson et al., 2011). Protected reserves, sustainable use areas, and indigenous lands serve as essential biodiversity sanctuaries and economic mixed-use areas (Miranda et al., 2016; Karin et al., 2018; Kauano et al., 2020), yet may not be sufficient for species that fill a unique and narrow niche (e.g. freshwater turtles; Fagundes et al., 2016).

Conservation is trans-disciplinary and is, therefore, influenced by social and geo-political dynamics (Chamberlain et al., 2012; Benitez-Capistros et al., 2016). As the Santa Cruz Forest Reserve has grown since its recent inception, Project Amazonas has proactively maintained positive interactions with local communities through education, healthcare, research, and conservation. This multidisciplinary approach is of particular importance as the region continues to develop and human communities expand from urban centres into forested areas. The high species richness found at the Santa Cruz Forest Reserve highlights this successful approach and affirms the reserve’s vital role as a protected conservation unit in close proximity to Iquitos. Research such as this can now be used to inform conservation and wildlife management practices within the reserve and the surrounding areas.

Acknowledgments. Research was conducted under FGCU IACUC #1819-05. This project would not have been possible without the financial and overall support from the offices of FGCUScholars (Undergraduate Research) and the FGCU Honors College. We are extremely grateful to Project Amazonas for their hospitality and to everyone involved with the fantastic crew at the Santa Cruz Forest Reserve. We also want to thank the FGCU Animal Behavior Research Group for their discussions and edits to this and other manuscripts as well as Prairie Gunnels for assistance in the field. Finally, we want to extend our gratitude to the late Dr. John Herman and his family. John, without your guidance, mentorship, and friendship, we would not be the biologists we are today. We are humbled to continue your legacy for conserving biodiversity, especially snakes, throughout the world.

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Accepted by Fabrício Oda


herpetofauna of the santa cruz forest reserve in the

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