Herpetology Notes, volume 14: 567-580 (2021) (published online on 21 March 2021)

Habitat use and activity of Bothrops bilineatus

smaragdinus Hoge, 1966 in the western Brazilian Amazon
(Serpentes: Viperidae)

Wirven Lima da Fonseca1, Radraque Rodrigues Correa1, Andesson de Souza Oliveira1,

Igor Soares de Oliveira2, and Paulo Sérgio Bernarde1,*

Abstract. We present information on the habitat use and activity of the pitviper Bothrops bilineatus smaragdinus in floodplain
forests of the western Brazilian Amazon. During a one-year period (April 2018–March 2019) we performed 816 h of time-
limited visual searches and recorded 35 individuals, with a further two recorded through occasional encounters, for a total of
37 snakes. Pitvipers were encountered at a frequency of one snake every 23 h (0.04 snakes/h). At night, snakes displayed sit-
and-wait hunting activity in 84.9% of recorded cases, while movement was observed in only 15.1%. Adult specimens mainly
positioned themselves on branches during hunting, whereas juveniles utilized leaves. Occurrence of snakes was significantly
correlated with the frequency of three frog species (Osteocephalus leprieuri, O. taurinus, Scinax ruber), which are part of
these vipers’ diet, and it was inversely correlated to rainfall and trail flooding. Bothrops b. smaragdinus was most frequently
encountered during the dry season, at the lowest recorded relative humidity, and less frequently during the rainy season. The
low encounter frequency may be associated with increased tree canopy humidity and vertical amphibian migration of the
Osteocephalus species, which occur at lower heights during the dry season.

Keywords. Arboreality, pitviper, sit-and-wait strategy, activity patterns, ecology

Introduction factors, including spatial and temporal temperature and

humidity distributions, all vary significantly from the
Although arboreal amphibians and reptiles form a
soil to the upper canopy layer (Madigosky and Vatnick,
very diverse component of the tropical environment
2000; Madigosky, 2004; Shaw, 2004). As it concerns
and present some remarkable ecological interactions
arboreal snakes, environmental features like foliage
with other species (e.g., Duellman, 1978; Shaw, 2004;
structure, microclimate, and prey type and availability,
Dial and Roughgarden, 2004; Wilson et al., 2007), those
as well as snake body size, weight, and shape, may
that inhabit tropical rainforest canopies are among the
determine the perch and forage heights that these
least studied vertebrates (e.g., Kays and Allison, 2001;
snakes are able to reach and explore (Lillywhite and
Guayasmin et al., 2006). In the Amazon, arboreal and
Henderson, 1993).
sub-arboreal snakes account for 19–35% of community
Information on the natural history of snakes is
species composition (e.g., Duellman, 1978; Martins and
paramount to uncovering their evolutionary biology and
Oliveira, 1998; Bernarde and Abe, 2006). Within this
ecology (Greene, 1997). However, vast forested areas
three-dimensional environment, biotic factors, including
have experienced rapid deterioration in the last several
plant structure, epiphytic plant availability, and
decades, especially in the Amazon, which makes obtaining
predator-prey and host-parasite dynamics, and abiotic
ecology and biodiversity knowledge a priority for future
management and conservation actions (Martins and
Oliveira, 1998). Arboreal snakes are sensitive to changes
in habitat and are more likely to be severely affected by
1
Laboratório de Herpetologia, Centro Multidisciplinar, Campus
deforestation, due to their adaptations and specializations
Floresta, Universidade Federal do Acre, Cruzeiro do Sul, Acre
69980-000, Brazil. to their specific habitat (Lillywhite and Henderson, 1993).
2
Laboratório de Etnociências, Ecologia e Conservação, Centro Furthermore, assessments of habitat use and activity of
de Educação e Letras, Campus Floresta, Universidade Federal venomous arboreal snakes may also contribute to the
do Acre, Cruzeiro do Sul, Acre 69980-000, Brazil. understanding of the circumstances in which snakebite
*
Corresponding author. E-mail: snakebernarde@hotmail.com occurs, providing valuable data for prevention guidelines
© 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. (Sazima, 1988; Oliveira and Martins, 2001).
568
Pitvipers of the genus Bothrops Wagler, 1824 occur
from Mexico to Argentina (Campbell and Lamar, 2004)
and currently comprise 45 species (Uetz et al., 2020).
These snakes inhabit several types of habitats, ranging
from open areas to dense forest and including habitat
disturbed by human activities (Campbell and Lamar,
2004). Available studies on the ecology of these snakes
are generally focused on their activity and habitat
use (e.g., Sazima, 1988; Oliveira and Martins, 2001;
Nogueira et al., 2003; Wasko and Sasa, 2012; Leão et
al., 2014), and mostly cover terrestrial species, with
relatively few studies of arboreal ones (e.g., Turci et
al., 2009; Andrade et al., 2010; Marques et al., 2012).
Additional studies on the ecology of arboreal pitvipers
are therefore not only desirable but essential to fulfil a
notorious knowledge gap concerning these animals.
Bothrops bilineatus (Wied-Neuwied, 1821) is considered
a complex of species comprising four clades (Dal Vechio et
al., 2018). The taxon for which we report our observations
herein corresponds to the “Western Amazonia” clade
(Dal Vechio et al., 2018), corresponding to the subspecies
B. b. smaragdinus Hoge, 1966. This arboreal snake
presents nocturnal habits and feeds on amphibians, small
mammals, lizards, bats, and other snakes (Dixon and
Soini, 1986; Martins et al., 2002; Campbell and Lamar,
2004; Turci et al., 2009; Fonseca et al., 2019a; Venegas
et al., 2019). Although B. b. smaragdinus is generally
considered uncommon in the wild (Bernarde et al.,
2011a), some reports of locations where it may be one of
the most frequent animals found during visual searches
in the natural environment are available (Silva et al.,
2020). There is evidence that this species may be the
second-ranked venomous snake concerning snakebites in
some regions (e.g., Letícia, Amazonas, Colombia; Shell,
Pastaza, Ecuador; Cruzeiro do Sul, Acre, Brazil; Haad,
1980/81; Smalligan et al., 2004; Mota-da-Silva et al.,
2019). Due to the high vulnerability of arboreal snakes
to habitat devastation and their medical interest, we here
present novel information on the habitat use and activity
patterns of B. b. smaragdinus in a floodplain forest located
in the western Brazilian Amazon.
Material and Methods
Study area. This study took place in the Lower Moa
River Forest (LMRF) in Cruzeiro do Sul Municipality,
Acre State, Brazil (7.3738°S, 72.4728°W; Fig. 1), a
site in the western Amazon. This region presents a hot
and humid tropical climate with an average annual
temperature of 24°C (Ribeiro, 1977), with the driest
months occurring from May–October. The LMRF is
Wirven Lima da Fonseca et al.
characterized as an “Open Alluvial Forest with Palm
Trees” (Acre, 2010) that is partially flooded seasonally
due to its proximity to the Moa River. The LMRF canopy
presents an average height of approximately 20 m, with
a few emerging tree species that can reach heights up
to 35 m. The canopy is dense, primarily composed of
the tree genera Brosimum (Moraceae), Enterolobium
(Fabaceae), and Virola (Myristicaceae) (Turci et al.,
2009). Large trees present tabular roots (“sapopembas”)
as an adaptation to the unstable LMRF soil and seasonal
floods (Turci et al., 2009). The understory displays
a high abundance of grasses (Poaceae) and spiny
palms (Astrocaryum, Arecaceae), and relatively few
herbaceous species, such as heliconias (Heliconiaceae),
arrowroot (Marantaceae), and wild gingers (Costaceae).
The diversity of spiny palms is relatively low, and only
a few species, including Euterpe precatoria, Iriartella
stenocarpa, and Bactris sp., have been recorded in the
area (Turci et al., 2009).
Data collection. Fieldwork was conducted by four
researchers during a one-year period, between April
2018 and March 2019. Three permanent trails were
systematically inspected, one of length 800 m and
two of 600-m length each (Fig. 1). The time-limited
visual search method (TLVS; Campbell and Christman,
1982) was applied, which consists of slowly walking
along a transect while searching for snakes visually
exposed in the environment. This method allows for
documentation of activity and habitat use patterns in the
field. A single person can search both sides of a trail,
but two people can optimize the method by searching
one side each. When this method is carefully performed,
two individuals can search approximately 400 m of a
trail in about 3 h. In this study, around 68 TLVS person-
hours were undertaken each month, totalling 816
TLVS person-hours, as follows: 3 h on Trail 1 (400 m
for each pair of researchers) and 4 h on Trails 2 and 3
(two researchers simultaneously on each track). Field
sampling was performed between 18:00–22:30 h. All
B. b. smaragdinus individuals recorded on Trails 2 and 3
were collected as voucher specimens (License SISBIO
12178), while individuals encountered on Trail 1 were
only observed and not collected. The following data
were recorded for each collected specimen: total length
(TL), sex, and gut contents. Voucher specimens have
been deposited at the Universidade Federal do Acre,
Campus Floresta Herpetological Collection (UFACF
4299–4317). Seven daytime visits were made in an
attempt to record the activity of specimens observed
during previous nights.
Habitat Use and Activity of Bothrops bilineatus smaragdinus 569

Figure 1. Location of the study area (red circle) and tracks sampled in the Lower Moa River Forest, Cruzeiro do Sul
Municipality, Acre State, Brazil. In the inset at lower left, Acre State is identified by the black colour. In the inset at upper right,
trail colours include blue (Trail 1), red (Trail 2), and yellow (Trail 3).

During fieldwork, individuals were observed for 5
min in order to record their activity patterns according
to the following protocol: (1) encounter time; (2) perch
substrate; (3) height above ground (measured with a
common tape measure or laser distance measurer); (4)
activity (hunting, moving, or resting), as inferred from
the individual’s posture (see Oliveira and Martins, 2001;
Turci et al., 2009); and (5) direction of an individual’s
head in relation to the tree trunk when an individual
was hunting on branches. When encountered again,
individual snakes could be recognized by their natural
colour patterns on the head, body, and tail, associated
with individual size, through the use of pictures taken in
the field with a Nikon Coolpix P900 camera. This was
facilitated by several snakes that remained stationary, in
the same trees or branches, or close to the same spot where
they were previously observed (at a distance of up to 30
m). Bothrops b. smaragdinus individuals encountered
more than once during the same month were not counted
multiple times for the seasonality analyses. Individuals
with a total body length of ≤ 30 cm were considered
juveniles, as this length corresponds to the approximate
size of neonates (25.5–27 cm; Grego et al., 2012).
Collection of environmental variables. Frogs
considered potential prey for B. b. smaragdinus as
well as others seen in the vegetation during TLVS
were counted and identified (Bernarde et al., 2013).
Temperature and relative humidity data were recorded
at the beginning and the end of daily fieldwork using
an Akrom Kr811 thermo-hygrometer (Akrom Produtos
Eletrônicos, São Leopoldo, Brazil) positioned 1.5 m
aboveground, and the data were then averaged for
subsequent analyses. Rainfall data were obtained from
a meteorological station located 3.5 km from the study
area (Inmet, 2019). A common measuring tape or laser
measurer (for very high heights) were used to define the
height of the substrate on which the snakes were found,
the diameter of tree trunks at breast height (DBH) was
calculated from their circumference as obtained with
a measuring tape, and the diameters of branches were
measured with callipers when possible. The flooding
rate of each trail was obtained by considering the
percentage of flooded linear trail distance (i.e., if 100 m
of a 400-m trail were flooded, this percentage would be
given as 25%), and the average flood values per month
were used.
570 Wirven Lima da Fonseca et al.

Data analysis. The Spearman correlation test was used (below 90%) were observed during the dry season,
to evaluate the relationship between B. b. smaragdinus between June and October (Table 1). All trails presented
occurrences and predictor variables (prey, rainfall, high flood levels (40–100%) as lake levels increased
temperature, relative humidity, flooding), as primary in the LMRF, which occurred in the rainy season from
data were not normally distributed. This correlation test December–March (Table 1).
statistically determines the strength of a relationship Activity.­ Bothrops b. smaragdinus were mostly
between paired data. Results were considered significant observed performing hunting activities at night (n = 63,
at P < 0.05. 82.9%; Fig. 2A–C, E, G, H), followed by locomotion
activity (n = 13, 17.1%; Fig. 2D). Individuals performing
Results ambush hunting exhibited a caudal luring strategy on 44
occasions (Fig. 2C). During daytime encounters (n = 17),
Encounters. A total of 35 B. b. smaragdinus individuals
individuals were most often observed resting (n = 14;
was recorded on the three investigated trails during our
Fig. 2F), and only three individuals hunted in the early
surveys (ca. one individual per 23 h of searching, or
morning (Table 2).
0.04 snake/h; Fig. 2; Table 1; Appendices I, II). Eighteen
Correlation between biotic and abiotic factors.
individuals were collected as voucher specimens on
The occurrence of B. b. smaragdinus was positively
Trails 2 and 3 (11 males with TL 499–668 mm and mean
correlated with the frequency of three amphibian prey
TL = 588 mm; seven females with TL 316–758 mm and
species (Osteocephalus leprieurii, O. taurinus, Scinax
mean TL = 578 mm). The stomach of one specimen
ruber; R = 0.707; P = 0.01; n = 12). Conversely, the
contained a treefrog, Scinax ruber (Hylidae), and another
occurrence of B. b. smaragdinus was inversely correlated
a lizard, Gonatodes humeralis (Sphaerodactylidae).
with rainfall (R = –0.6430; P = 0.02; n = 12) and flooding
Bothrops b. smaragdinus was the most abundant snake
(R = –0.9325; P < 0.0001; n = 12). No correlations
recorded during the surveys, corresponding to 27% of
were observed between B. b. smaragdinus occurrence
total snake species found (n = 130 individuals of 21
and relative humidity (R = –0.4588; P = 0.1336; I =
species). On Trail 1, 17 B. b. smaragdinus were observed
12), temperature (R = 0.2699; P = 0.39; n = 12) and
on 58 occasions, including both encounters and repeated
the frequency of all amphibian species detected on the
encounters. In addition, one juvenile was occasionally
vegetation (R = –0.3364; P = 0.28; n = 12).
recorded on Trail 1 after completing a TLVS on Trail
Substrate use. In general, B. b. smaragdinus was
2, and another was collected off the trails. A total of 37
observed between 0.3 m and 18.0 m in height (mean =
individuals were recorded, and most presented a brown
6.4 m; n = 63 observations) when hunting on trees with
(59.5%) tail tip, while the rest (40.5%) exhibited a white
diameters varying between 1.9 cm to 45.1 cm (mean =
tail tip, independent of sex or age (Fig. 2A, B).
12.2 cm; n = 49) (Appendix I). Most individuals were
Seasonality. Forty-one of the encounters (75.9%)
found up to 5 m in height (n = 39, 51%; Fig. 3A).
occurred during the dry season, with no individuals
Individuals > 30 cm in total length were observed
encountered during rainy season from December–March
coiled up on branches during hunting activities (n = 56
(Table 1). Three juveniles (approximately 250 mm in total
observations; Fig. 2A–C, E), and only a single individual
length) were observed in November, at the beginning of
was found 0.3 m above the ground, as it was hunting on
the rainy season. On the other hand, frogs were present
a fallen palm leaf. Juveniles < 30 cm in length (n = 4)
on the vegetation during all months, exhibiting a slightly
were observed on leaves (n = 6 observations, including
higher abundance in the rainy season, when 53.8% of all
two repeated observations; Fig. 2G, H). Diameters of
observed frogs were encountered (Table 1; Appendix II).
branches used for hunting ranged from 5.2–12.3 mm
Osteocephalus leprieurii and O. taurinus were the
(mean = 8.4 mm; n = 28). Resting activity was recorded
most abundant anuran species detected throughout
14 times, and the individuals usually remained on the
year, corresponding to 50% of all amphibians recorded
same substrate used for hunting during the previous night
on the vegetation (Appendix II). Furthermore, these
(heights from 1.8–11.4 m; mean = 4.8 m; n = 9; Fig. 3B).
species were the most frequent during the dry season
Based on 57 observations, we calculated snake residence
(64.4% of encounters). The third most abundant
time on the same substrate (branch and tree). In most
amphibian was the toad Rhinella margaritifera (35%
cases (17), the snake remained on the same substrate
of encounters), which was seen more frequently during
only during the night of the first encounter, followed
the rainy season (70% of all records for this species;
by two days (9) and three days (6). In two occasions,
Appendix II). The lowest relative air humidity levels
Habitat Use and Activity of Bothrops bilineatus smaragdinus 571

Figure 2. Shown are several Bothrops bilineatus smaragdinus specimens encountered during our surveys. Adults may
exhibit white (A) or brown (B) tail tips. Individuals performing different activities include (C) the use of caudal luring to
hunt, (D) locomotion, (E) and sit-and-wait hunting with a tongue flick. (F) The same individual in (E) is shown resting
during the day in the same tree. (G) and (H) are juveniles (< 30 cm in total length) hunting on leaves. Photos taken by
Wirven Lima da Fonseca (A–D, H) and Paulo Sérgio Bernarde (E–G).
572 Wirven Lima da Fonseca et al.

Table 1. Bothrops bilineatus smaragdinus encounters on trails during the Time Limited Visual Searches in the Lower Moa River
Forest, Cruzeiro do Sul Municipality, Acre State, Brazil, and the biotic and abiotic variables relevant to the present study. Reported
variables include encounters with the snake (Encounters); the general presence of frogs on the vegetation (Frogs); the presence of
the Osteocephalus leprieurii, O. taurinus, and Scinax ruber, the three frog species recorded as prey of B. b. smaragdinus (Prey);
the average temperature at night (Temperature, °C); the average relative humidity at night (Humidity, %); total monthly rainfall
(Rainfall, mm); and the linear percentage of trails flooded (Flood, %). Months are indicated by capital letters, beginning in April
2018 and ending in March 2019. Repeated encounters with the same individual during the same month were not taken into
account, only in different months.

2018 2019
Variable
A M J J A S O N D J F M
Encounters 9 1 5 9 9 12 5 4 0 0 0 0
Frogs 48 56 56 46 62 54 97 104 58 119 34 69
Prey 40 26 33 29 50 46 58 52 20 26 13 16
Temperature 24.8 25.1 25.5 26 25 25.4 26.8 26.2 25.7 23.8 23.5 24.4
Humidity 97.5 92.6 86.3 88 86 87.8 85.4 92.8 92.8 90.9 93.6 91.9
Rainfall 243.2 166 23.6 37.4 69.6 33.4 233.4 397.8 191.2 358.6 412.8 389.4
Flood 0 20 0 0 0 0 20 20 40 80 100 60

up to 15 and 17 days residence times were documented the diet of B. b. smaragdinus in this study (with prey
(Fig. 3B). Individuals moving on the vegetation were including only two frogs and one lizard), we confirmed
observed between 1.4 and 19.9 m in height (mean = 5.7 that adults of this snake species prey on ectothermic
m; n = 13; Appendix I). animals (e.g., Martins et al., 2002) and also often use
caudal luring as a strategy (Fonseca et al., 2019a).
Discussion Moreover, the specific prey found in the stomachs of
three specimens (Osteocephalus taurinus, Scinax ruber,
Bothrops b. smaragdinus was the most frequently
Gonatodes humeralis) represents only arboreal habits
encountered snake species during our surveys in the
(Dixon and Soini, 1986; Miranda et al., 2015).
LMRF (Silva et al., 2020). The encounter rate for the
Although males (11 individuals) were more frequently
species (0.04 snakes per hour) was higher than that
encountered than females (8), the overall number
recorded for B. atrox, which was 0.02 in the same study
of specimens collected is insufficient to adequately
area and 0.02 in the Adolfo Ducke Forest Reserve, in
discuss sex ratios. In general, males displayed a larger
Manaus (Oliveira and Martins, 2001; Turci et al., 2009).
average size (588 mm) than females (578), but the
The high abundance of B. b. smaragdinus in this locality
largest specimen was a female of length 758 mm, while
is intriguing since this species had not previously been
the largest male was 668 mm in length. At this same
recorded in species survey studies developed in terra
locality, Turci et al. (2009) recorded a similar pattern,
firme areas (e.g., Avila-Pires et al., 2009; Bernarde et al.,
of a 780 mm female and a 670 mm male specimen.
2011b; Fonseca et al., 2019b). This may indicate that B.
However, in Bothrops species, females tend to be larger
b. smaragdinus may be less frequent or more difficult to
than males (Almeida-Santos and Salomão, 2002; Silva
detect by visual searches in other vegetation structures.
et al., 2017; Bisneto and Kaefer, 2019). Thus, the pattern
Both male and female juvenile and adult B. b.
we detected may be due to insufficient sampling.
smaragdinus in this population exhibited tail tips
Unlike the general seasonality occurrence pattern
with a colour different from the rest of the body. Tail
observed for Amazonian snake species registered by
tips can be white or brown, regardless of age and sex
TLVS, where abundance is higher during the rainy season
(Fonseca et al., 2019a). Tail tip colour retention in adult
(Martins and Oliveira, 1998; Oliveira and Martins, 2001;
Bothrops species indicates that these individuals may
Bernarde and Abe, 2006), B. b. smaragdinus was more
feed on ectothermic prey and use their tail as a decoy
frequently encountered in the dry season (74.5% of the
and lure prey as a hunting strategy (Martins et al., 2002).
records), when relative humidity was lowest. On the
Although we obtained relatively little information on
other hand, frogs (potential prey) were registered on the
Habitat Use and Activity of Bothrops bilineatus smaragdinus 573
Table 2. Daytime Bothrops bilineatus smaragdinus observations on Trail 1. All observations were made in
2018.
No. of Humidity Temperature
Date Time of Day Activity
Individuals (%) (°C)
27 April 09:00–9:30 h 3 Hunting 90.3 23.1
15 June 09:40 h 1 Sleeping 90.7 23.9
7 July 14:40–15:45 h 6 Sleeping 74.2 27.7
20 July 09:05 h 1 Sleeping 88.6 22.8
18 September 14:20–15:15 h 4 Sleeping 56.5 33.1
17 October 17:35 h 1 Sleeping 82.0 29.0
8 November 17:20 h 1 Sleeping 89.7 25.8

vegetation during the entire year at the study area, peaking that the assessed B. b. smaragdinus population migrates
during the dry season, although with lower uniformity vertically and seasonally throughout the year, perhaps in
and diversity compared to the rainy season (Miranda et correlation with the availability of Osteocephalus prey.
al., 2015). In addition, amphibian species belonging to the Regardless of the abundance of frogs in the lower forest
genus Osteocephalus, recorded as the stomach content of strata (i.e., < 2 m in height above ground), the third most
B. b. smaragdinus (Dixon and Soini, 1986), were more abundant species was Rhinella margaritifera, which
abundant during the dry season, and the B. b. smaragdinus presents diurnal activity on litter and uses the vegetation as
frequency was correlated with Osteocephalus encounters a resting spot, remaining still (Miranda et al., 2015). Thus,
throughout the year. The two frog species belonging to this amphibian is not expected to be part (or an important
this genus were most frequently observed at heights in the
trees above 3.5 m and up to 12 m (Miranda et al., 2015),
and they are known to be common canopy inhabitants Figure 3. (A) Range of documented heights at which
(Doan, 2004; Guayasamin et al., 2006). Bothrops bilineatus smaragdinus were observed during
The increased encounter rate with these amphibians nocturnal hunting activity (n = 63). (B) Residence time of B.
during the driest months is likely due to decreased b. smaragdinus in the same location during consecutive days
canopy humidity during this period in tropical (n = 56).
forests (Freiberg, 1997; Madigosky and Vatnick,
2000; Madigosky, 2004). As a consequence of the
dry conditions higher in the canopy, these species
descend to lower strata and become detectable during
visual searches (Doan, 2004). Low humidity is also an
important factor that can influence the activity of some
snake species (Lillywhite and Henderson, 1993; Daltry
et al., 1998; Moore and Gillingham, 2006; Subach
et al., 2009). In the present study, this relationship
seems to be indirect, through prey availability (here:
Osteocephalus treefrogs), since no correlation between
B. b. smaragdinus frequency and humidity levels was
observed. Frogs and squamate reptiles represent two
canopy ecology extremes, as the former are generally
intolerant of desiccation while the latter are tolerant
(Shaw, 2004). In addition, snake prey location and
distribution are significant factors in the evolution of
snake habitat selection (Reinert, 1993). Thus, spatial
and temporal prey availability may be determinant for
the occurrence of certain snake species (e.g., Chandler
and Tolson, 1990; Lillywhite and Henderson, 1993;
Bernarde et al., 2000; Madsen et al., 2006). It is likely
574
portion) of the B. b. smaragdinus diet, since these snakes
preferably hunt more active prey (Turci et al., 2009;
Fonseca et al., 2019a; Venegas et al., 2019). Most frogs in
the study area that use vegetation occur at heights < 2.1 m
(Miranda et al., 2015), whereas B. b. smaragdinus usually
hunts in higher strata. This reinforces the hypothesis of
lower prey availability during the rainy season. However,
other factors, such as competition or predation, should
also be considered as potential determinants for B. b.
smaragdinus habitat selection.
Another hypothesis may explain the low frequency
of B. b. smaragdinus in lower strata during the rainy
season, which is the snakes’ attempt to minimize
competition vertically. Another viperid, B. atrox, is also
present in this forest, and it is the second-most abundant
snake species (Silva et al., 2020). This generalist snake
may be a potential B. b. smaragdinus competitor, since
it is mostly active during the rainy season (Oliveira
and Martins, 2001; Turci et al., 2009), feeds on prey
in common with B. b. smaragdinus (frogs including
Osteocephalus, small lizards, small mammals), and
hunts on the vegetation at heights of up to 1.5 m
(Martins and Oliveira, 1998; Bisneto and Kaefer, 2019).
Furthermore, B. atrox is also ophiophagous, including
cannibalism (Martins and Oliveira, 1998; Bernarde and
Abe, 2010; Rodrigues et al., 2016; Bisneto and Kaefer,
2019), and it is abundant during the rainy season in this
forest (Turci et al., 2009). Thus, it may also represent a
potential B. b. smaragdinus predator. Compared to B.
b. smaragdinus, B. atrox reaches a larger size (over 1.5
m; Martins and Oliveira, 1998; Oliveira and Martins,
2001). In addition, B. atrox juveniles were observed at
a height of 1.5 m, and two adults were seen during this
study hunting at heights of 2.5 and 4.2 m during the rainy
season, which reinforces the potential predator status of
B. atrox on B. b. smaragdinus. Despite the record of a
pitviper (Porthidium lansbergii) as a prey of B. asper
(Roldan et al., 2011), a species belonging to the B. atrox
species group, no records of B. atrox preying on other
viperid species are available, suggesting that further
studies are required in this regard. Prey availability
may determine the mobility range of arboreal snakes,
and competition and/or other factors may lead to dietary
changes or to optional terrestrially of strictly arboreal
species (Lillywhite and Henderson, 1993).
One adult B. b. smaragdinus specimen was observed
hunting at only 0.3 m from the ground during the
rainy season. Turci et al. (2009) also observed an adult
female in the same situation in the rainy season and,
when assessing the snake’s stomach content, found a
Wirven Lima da Fonseca et al.
rodent. All observations of this species in the LMRF
indicate strict arboreality while hunting (Turci et al.,
2009; Fonseca et al., 2019a), and it may hunt near
ground level only rarely or opportunistically. Although
competition and predation may have an effect on this B.
b. smaragdinus population, prey availability and habitat
structure are considered to be among the most important
factors that influence seasonal snake movements (e.g.,
Reinert, 1993; Gregory et al., 2001; Heard et al., 2004;
Hirai, 2004). During the rainy season, with the resulting
higher humidity, large areas of this forest become
flooded, and Osteocephalus encounters probably
decrease due to vertical migration towards upper strata
(e.g., Doan, 2004; Guayasamin et al., 2006). Thus, the
most plausible hypothesis is that B. b. smaragdinus is
more active in the upper strata during the rainy season
compared to the dry season, which reduces the potential
for detection by active search methods from the ground.
As reported by Turci et al. (2009) and Fonseca et al.
(2019a), B. b. smaragdinus is preferably a nocturnal
snake that applies an ambush strategy to hunt and often
uses caudal luring to decoy prey. The use of their tail
as bait is paramount for snakes that feed on active prey
(Fonseca et al., 2019a). For Osteocephalus, which is
a nocturnal treefrog group that commonly jumps onto
several types of substrate (trunks, branches, leaves,
palm leaves, vines, bushes) at different heights (Miranda
et al., 2015), this tactic may be efficient in the three-
dimensional environment where B. b. smaragdinus
generally occurs (Fonseca et al., 2019a). However, these
snakes were observed active up to nearly 20 m in height
above ground level, higher than previously recorded
(2 m: Duellman, 1978; 7 m: Turci et al., 2009; 18 m:
Fonseca et al., 2019a). In the Neotropics, most arboreal
snake species are active below 5 m in height, and mostly
near ground level (e.g., Lillywhite and Henderson,
1993; Martins and Oliveira, 1998). Our observations are
among the highest records for Amazonian snake species,
and they are the highest recorded for B. b. smaragdinus.
Bothrops b. smaragdinus used the same substrate to
rest during the day and remained in the same place for
up to 17 days. Its cryptic colouring (green) and its low
mobility probably contribute to avoiding or minimizing
predation pressure, especially by visually-oriented
diurnal predators, such as birds of prey, which are among
the main Neotropical forest predators (Lillywhite and
Henderson, 1993; Martins et al., 2008). Furthermore,
the use of vegetation for hunting and resting may
contribute to reduce the risk of predation by terrestrial
predators (e.g., Martins, 1993).
Habitat Use and Activity of Bothrops bilineatus smaragdinus
Juveniles and adults of the same species can use
substrate in different ways (e.g., Reinert, 1993; Oliveira
and Martins, 2001; Shine et al., 2002; Wilson et al.,
2007), which may be associated with morphology
(body size or colour), prey availability, and predation.
We recorded four juveniles (TL < 30 cm) hunting
by ambush on tree leaves. On the other hand, large
amphibians (e.g., Osteocephalus) were not observed
on small leaves (Miranda et al., 2015), which cannot
support their weight. However, juvenile Osteocephalus,
small hylids, and Pristimantis species were observed
on smaller leaves (Miranda et al., 2015), and these
represent suitable prey for B. b. smaragdinus juveniles.
Adult B. b. smaragdinus were mostly recorded hunting
on tree branches, but not on leaves (Turci et al., 2009;
Fonseca et al., 2019a; this study). This probably reflects
individual body size and weight, which prevents larger
individuals access to certain microhabitats, which
therefore become inaccessible to adults (e.g., Plummer,
1981; Henderson, 1993; Shine et al., 2002).
Conclusions
Although B. atrox is usually the most abundant viperid
in Amazon forests, B. b. smaragdinus may be the most
frequent snake species in certain environments (such as
floodplain forest) during visual night searches. Bothrops
b. smaragdinus generally hunts by ambushing its prey in
the forest canopy up to nearly 20 m above ground, and
it uses caudal luring as its main behavioural strategy to
attract prey. In a reflection of body size and the availability
of suitably-sized prey, adult B. b. smaragdinus preferably
use branches to hunt, whereas juveniles prefer leaves.
Bothrops b. smaragdinus was more frequently
observed during the driest period, when canopy humidity
is reduced and potential frog prey (including treefrogs of
the genus Osteocephalus) migrates vertically towards
the lower stratum. Conversely, during the rainy season
when flooding in the study area increased local relative
humidity, B. b. smaragdinus activity may shift to the
upper forest strata, where prey availability is higher, and
which makes the detection of these snakes more difficult
during visual night searches.
Acknowledgements. The authors would like to thank the
Coordination of Superior Level Staff Improvement (CAPES) –
Financial Code 001, the Brazilian National Council for Scientific
and the Technological Development (CNPq) for financial support
(PIBIC/UFAC), to authors RRC and ASO, and the Instituto Chico
Mendes de Conservação da Biodiversidade (ICMBio) for the
license permit (SISBIO/12.178).
575
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578 Wirven Lima da Fonseca et al.

Appendix I. Vegetation use by Bothrops bilineatus smaragdinus recorded during nocturnal limited visual
searches in the Lower Moa River Forest, Cruzeiro do Sul Municipality, Acre State, Brazil in 2018. The
use of a palm leaf, as opposed to a large leaf of a regular tree, is indicated by the substrate term “Palm.”
Instances of caudal luring are indicated by an asterisk (*). Tree trunk size is indicated as diameter at breast
height (DBH).

Branch
Time Height DBH
Individual Date Trail Substrate Diameter Activity
(h) (m) (cm)
(mm)
I 4 April 19:50 1.4 2 Branch 1.9 6.2 Moving
II 4 April 20:18 10.0 2 Branch 7.9 7.1 Hunting
III 4 April 21:45 8.6 2 Branch 7.5 10.6 Hunting*
IV 19 April 21:32 4.5 3 Branch 5.5 9.5 Hunting*
V 19 April 22:02 8.5 3 Branch 12 7.1 Hunting
VI 25 April 18:15 1.3 1 Branch 2.9 8.4 Hunting*
VI 26 April 18:15 1.3 1 Branch 2.9 8.4 Hunting
VII 26 April 18:50 2.5 1 Branch 5.5 9.1 Moving
VIII 26 April 19:20 0.3 1 Palm - 8.2 Hunting*
IX 26 April 21:23 6.7 1 Branch 5.6 9.2 Hunting*
X 18 May 19:10 10.6 1 Branch 7.4 8.9 Hunting*
XI 14 June 18:00 3.3 1 Palm 4.2 12.0 Moving
XII 14 June 18:00 18.0 1 Branch 39.4 - Hunting*
XIII 14 June 18:15 4.8 1 Leaf 17.6 7.4 Hunting*
VII 14 June 19:00 6.8 1 Branch 10.8 6.9 Hunting*
XIV 28 June 20:24 0.8 2 Branch 2.3 7.0 Hunting*
XV 1 July 19:50 5.7 2 Branch 11.3 5.2 Hunting*
XIII 4 July 18:20 2.0 1 Branch 7.1 7.8 Moving
VII 4 July 19:00 2.3 1 Branch 8.9 9.9 Hunting*
XVI 4 July 20:45 7.0 1 Branch 15.4 - Moving
XVII 4 July 20:50 6.7 1 Branch 13.6 8.8 Hunting*
XVIII 4 July 20:46 2.6 1 Branch 11.1 9.7 Hunting
VII 5 July 18:40 2.3 1 Branch 8.9 9.9 Hunting*
XVI 5 July 20:30 6.5 1 Branch 11.6 - Moving
XVIII 5 July 20:45 2.6 1 Branch 11.1 9.7 Hunting
IX 5 July 20:50 11.4 1 Branch 17.1 - Hunting*
XVII 5 July 20:50 6.7 1 Branch 13.6 8.8 Hunting*
XIX 6 July 18:31 7.3 1 Branch 31.7 - Hunting*
VII 6 July 18:50 2.3 1 Branch 8.9 9.9 Hunting*
XVI 6 July 20:30 8.9 1 Branch 11.5 - Hunting*
XVII 6 July 20:50 6.7 1 Branch 13.6 8.8 Hunting*
XVIII 6 July 20:45 2.6 1 Branch 11.1 9.7 Hunting*
IX 6 July 20:53 11.4 1 Branch 17.1 - Hunting*
XX 8 July 19:26 6.7 2 Branch 6.8 - Hunting*
XIII 22 August 18:15 5.9 1 Leaf 10.3 - Hunting*
XIX 22 August 18:40 7.3 1 Branch 14.2 - Hunting*
XVIII 22 August 20:40 8.3 1 Branch 16.3 - Hunting*
Habitat Use and Activity of Bothrops bilineatus smaragdinus 579
Appendix I. cont.

Branch
Time Height DBH
Individual Date Trail Substrate Diameter Activity
(h) (m) (cm)
(mm)
XXI 24 August 20:15 4.2 2 Branch 5.5 7.8 Hunting
XXII 24 August 21:23 3.2 2 Branch 5.8 6.9 Hunting
XXIII 24 August 21:40 3.9 2 Branch 7.7 11.2 Moving
XIII 27 August 18:15 5.9 1 Leaf 10.3 - Hunting*
XIX 27 August 18:30 13.5 1 Branch 45.1 - Hunting
XVIII 27 August 20:40 8.3 1 Branch 16.3 - Hunting
XIX 28 August 18:32 9.3 1 Branch 23.2 - Hunting*
XXIV 28 August 20:02 3.5 1 Branch 2.3 6.8 Moving
XVIII 28 August 20:30 8.3 1 Branch 16.3 - Hunting
XXV 29 August 18:15 7.5 3 Branch 6.7 - Hunting
XXVI 29 August 21:43 10.5 2 Branch 13.2 - Hunting
XI 12 September 18:10 1.4 1 Branch 9.4 8.7 Moving
XXIV 12 September 19:31 1.7 1 Branch 3.1 9.6 Hunting
XVI 12 September 20:35 1.8 1 Branch 9.3 10.4 Hunting*
XVII 12 September 20:55 3.4 1 Branch 10.9 12.1 Hunting*
XI 17 September 18:08 13.3 1 Branch 42 - Hunting*
XXVII 17 September 18:40 4.0 1 Branch 11.2 8.1 Moving
XXVIII 17 September 19:09 2.4 1 Branch 7.7 6.7 Moving
XXIV 17 September 19:45 4.8 1 Branch 4.4 - Hunting
XVI 17 September 20:30 1.8 1 Branch 9.3 10.4 Hunting
XVII 17 September 20:50 4.2 1 Branch 9.9 12.3 Hunting
XIX 20 September 19:07 5.9 1 Branch 12.3 - Hunting
XVI 20 September 20:30 1.8 1 Branch 9.3 10.4 Hunting
IX 20 September 20:55 4.8 1 Branch 9.2 - Hunting
XXIX 24 September 20:28 5.5 2 Branch 12.3 - Hunting
XXX 24 September 20:45 8.2 2 Branch 15.5 - Hunting*
XXXI 24 September 21:05 2.5 3 Branch 11.4 5.2 Hunting
XXXII 25 September 20:24 2.4 2 Branch 5.4 6.2 Hunting
XI 1 October 18:15 14.1 1 Branch 8.4 - Hunting
XIII 1 October 18:20 4.5 1 Leaf 14.5 - Hunting
XIX 1 October 18:30 1.6 1 Branch 4.3 7.6 Hunting*
XI 9 October 18:15 12.4 1 Branch 23.5 - Moving
XI 15 October 18:15 14.2 1 Branch 14.8 - Hunting*
XIX 15 October 18:35 12.4 1 Branch 16.8 - Hunting
XVI 15 October 18:35 1.8 1 Branch 12.6 12.2 Hunting*
XXXIII 17 October 21:30 3.5 2 Branch 6.8 6.7 Hunting*
XII 2 November 18:10 19.9 1 Branch 31.1 - Moving
XXXIV 6 November 18:30 2.3 1 Leaf 3.7 6.2 Hunting*
XXXV 12 November 21:00 3.3 1 Leaf 4.3 6.4 Hunting*
580 Wirven Lima da Fonseca et al.

Appendix II. Numbers of frogs observed along trails in the Lower Moa River Forest, Brazil, during our
surveys from April 2018–March 2019.

2018 2019
Taxon Total
A M J J A S O N D J F M

Bufonidae
Rhinella margaritifera 2 24 12 12 6 3 29 44 29 72 16 35 284

Craugastoridae
Pristimantis fenestratus 1 1
Pristimantis ockendeni 1 1
Pristimantis reichlei 1 1 1 1 1 1 6
Pristimantis sp. 1 1 2

Hylidae
Boana calcarata 2 2 8 2 14
Boana cinerascens 1 2 1 1 3 8
Boana fasciata 1 3 1 1 2 8
Boana geographica 3 3 8 2 16
Boana lanciformis 1 1
Boana punctata 1 1
Dendropsophus brevifrons 2 2
Dendropsophus marmoratus 1 1
Dendropsophus parviceps 2 8 10
Dendropsophus sp. 2 1 3 5 11
Osteocephalus leprieurii 38 17 18 13 27 33 34 20 8 10 6 10 234
Osteocephalus taurinus 2 9 14 16 23 13 22 32 12 16 5 6 170
Scinax ruber 1 2 2 5
Sphaenorhynchus dorisae 1 1
Trachycephalus typhonius 1 1

Phyllomedusidae
Cruziohyla craspedopus 1 1
Phyllomedusa bicolor 4 5 6 2 1 3 1 2 24
Phyllomedusa vaillantii 1 1
Total 48 56 56 46 62 54 97 104 58 119 34 69 803

Accepted by Mirco Solé