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Full text of "Breviora"

MCZ 
LIBRARY 



I 13 2C09 

HARVARD 
UNIVERSITY 



B RE V I 





Nl Mseium oi v^onnparaitive /^oology 



US ISSN 0006-9698 



Cambridge, Mass. 



7 December 2008 



Number 514 



THE AMPHIBIANS AND REPTILES OF THE ESTACION BIOLOGICA JATUN 
SACHA IN THE LOWLAND RAINFOREST OF AMAZONIAN ECUADOR: 

A 20-YEAR RECORD 

Gregory O. Vigle 1 

Abstract. The amphibian and reptile fauna within a small area in a lowland rainforest fragment reserve in 
Amazonian Ecuador was intensively surveyed over an initial 2- year period via removal sampling (1986-88), 
coinciding with the construction of a road through the area and a subsequent surge of increased forest conversion 
and fragmentation in surrounding areas. A time-constrained transect sampling technique was employed to facilitate 
later, post-fragmentation visual-encounter monitoring to gather long-term data on species richness and composition 
(on a chronologically coarse scale). Between 1998 and 2005. an approximately equivalent quantity of sampling effort 
was accumulated in the same small area to compare with data from 1986-88. in an attempt to evaluate the efficacj of 
this reserve in conserving the herpetofaunal community documented more than 10 years earlier. 

Based on a total of 6.722 individual records obtained over 1.117 effort-hours of sampling divided into two primarj 
and five secondary sampling periods, herpetofaunal species richness was among the highest yet reported from a single 
locality in Amazonia, with 84 amphibian and 82 reptile species recorded from 1986 through 2007. A complete species 
list is provided, with commentary on noteworthy records. Of 73 amphibian and 46 reptile species recorded in 1986-88 
sampling. 68 amphibians (93%) and 40 reptiles (87%) were recorded again in sampling from 1998 through 2007: 1 1 
amphibian and 33 reptile species not recorded in 1986-88 were added from 1998 to 2007. Pairwise comparisons of 
species composition among all sampling periods using a similarity index revealed a > 50% similarity for both 
taxonomic groups across all periods: similarity of the two primary periods (1986-88 vs. 1998-2005) was 0.90 for 
amphibians and 0.70 for reptiles. 

The present study provides another single-site reference point for the mega-diverse herpetofauna of the upper 
Amazon basin, but is distinct in offering long-term data on species persistence; results from the 20-year span oi~ 
sampling suggest that this small reserve has. since 1986. successfully conserved this fauna. These results concur with 
previous long-term studies from the central Amazon basin in suggesting that relatively small reserves and other forest 
remnants with sufficient habitat diversitj may substantially contribute to the continued survival of species-rich 
Amazon rainforest herpetofaunal communities amidst widespread regional deforestation. 

Key words: Amazon: amphibians: biodiversity: conservation: Ecuador: Estacion Biologica Jatun Sacha; forest 
fragmentation: herpetofauna: monitoring: reptiles: reserve; species richness: tropical rainforest 



'School of Life Sciences, Arizona State University, Tempe, AZ 85287-1501, U.S.A.; e-mail: vigle@imap2. 
asu.edu. or Librarius« librarius.org 



© The President and Fellows o\' Harvard College 2008. 



BREVIORA 



No. 514 



Resumen. Los anfibios y reptiles que se encuentran en un area pequena dentro de una reserva fragmentada de 
bosque lluvioso en la Amazonia ecuatoriana han sido intensivamente investigados en un periodo inicial de dos afios 
mediante la recoleccion de muestras (1986-88). Esta actividad coincidio con la construction de un camino a traves 
del area y la subsiguiente oleada de conversion y fragmentation del bosque en las areas circundantes. Se empleo una 
tecnica de muestreo de transectos de tiempo restringido para facilitar luego el monitoreo de encuentro visual post- 
fragmentation y reunir datos de largo plazo sobre la riqueza y composicion de las especies (en una escala cronologica 
desigual). Entre 1998 y 2005 se acumulo un esfuerzo de investigation y muestreo equivalente en la misma area 
reducida para compararla con la information de 1986 a 1988, en un intento por evaluar la eficiencia de esta reserva 
en la conservation de la fauna herpetologica documentada mas de 10 ahos atras. 

Basada en un total de 6,722 registros individuales obtenidos durante 1.117 horas de recoleccion de muestras 
divididas en dos periodos principales y cinco periodos secundarios de muestreo, la diversidad herpetofaunistica 
estuvo entre las mas altas reportadas hasta el momento en una unica ubicacion en la Amazonia, con 84 especies de 
anfibios y 82 especies de reptiles registrados de 1986 a 2007. Se provee una lista completa de especies. con 
comentarios en los registros mas significativos. De las 73 especies de anfibios y 46 especies de reptiles registradas de 
1986 a 1988, 68 anfibios (93%) y 40 reptiles (87%) fueron registrados nuevamente en el muestreo de 1998 a 2007; 1 1 
especies de anfibios y 33 especies de reptiles no registrados entre 1986 y 1988 fueron anadidos en 1998 a 2007. 
Mediante un analisis de composicion de pareja entre todos los periodos de muestreo usando un indice de similitud 
revelaron un > 50% de similitud para ambos grupos taxonomicos entre periodos; la similitud de estos dos periodos 
principales (1986-88 vs. 1998-2005) fue 0.90 por anfibios y 0.70 por reptiles. 

El presente estudio provee otro punto de referenda de sitio unico para la herpetofauna megadiversa de la zona alta 
de la cuenca amazonica, pero es diferente ya que ofrece datos de largo plazo en la persistencia de especies; los 
resultados de un lapso de muestreo de 20 ahos sugieren que esta pequena reserva. desde 1986 ha conservado 
exitosamente esta fauna. Estos resultados coinciden con estudios previos de largo plaza de la cuenca central de la 
Amazonia en sugerir que las reservas relativamente pequenas y otros remanentes de bosque con suficiente diversidad 
de habitat pueden contribuir sustancialmente a la continua supervivencia de la alta diversidad de comunidades 
herpetofaunisticas de los bosques lluviosos amazonicos en medio de la extendida deforestation regional. 



INTRODUCTION 

The rainforests of the Amazon basin 
harbor the highest concentration of biodi- 
versity on Earth, yet they are being destroyed 
by human activities at an alarming pace; one 
prominent threat is deforestation and the 
fragmentation of previously continuous for- 
est into smaller island remnants of remaining 
habitat (Bierregaard et al, 2001; Lovejoy et 
al, 1984; Myers. 1984). 

Regionally, and on the basis of surveys from 
single localities, the lowland rainforests of the 
western Amazon basin of Ecuador, Peru, and 
Colombia are home to the highest concentra- 
tion of amphibian and reptile species richness 
known from any area on Earth (Campbell and 



Lamar, 2004; Dixon and Soini, 1986; Duell- 
man, 1999; Frost, 2007; Peters and Donoso- 
Barros, 1970; Peters and Orejas-Miranda, 
1970). Reports on single-site studies have been 
reviewed by Duellman (1988, 2005) and Duell- 
man and Thomas (1996) (also see Doan and 
Arizabal, 2002; Lynch, 2005). Within the 
immediate area of the Ecuadorian Amazon, 
published studies on the entire herpetofauna 
are available for several single or clustered 
sites, including Almendariz (1987), Cisneros- 
Heredia (2003), Duellman (1978), Duellman 
and Mendelson (1995), Izquierdo et al. (2000), 
and Lescure and Gasc (1986). 

From these studies, there has been an 
increasing awareness of the geographic and 
ecological heterogeneity of the Amazonian 



2008 



A MA/ON I OKI SI I RACiMFNT 1 II RIMTOl A I IN A 



herpetofauna, and the neea\ to survey addi- 
tional localities to better understand both 
local and regional distribution patterns of 
this mega-diverse but still poorb known 
fauna (Doan and Ari/abal. 2002; Duellman, 
2005: Lynch, 2005). 

The Estacion Biologica Jatun Sacha 
(EBJS) is a small remnant of lowland 
tropical wet forest located at the western 
edge of the Amazon Basin in eastern 
Ecuador, established as a reserve in 1985 
shortly before the construction of a road 
through the area. 

In 1986, coinciding with the onset of road 
construction, an intensive survey of the 
amphibian and reptile fauna was initiated. 
Although the first goal was to conduct a 
standard inventory via removal sampling, 
the ultimate fate of this new reserve as 
eventually becoming an isolated habitat 
island was recognized at the time, and a 
simple time-constrained transect sampling 
method was implemented to facilitate future 
long-term monitoring via visual-encounter 
surveys (VES) and to attempt to assess the 
long-term efficacy of the reserve in conserv- 
ing the herpetofauna. A general initial 
prediction was that species richness would 
eventually decline or otherwise exhibit obvi- 
ous changes in community composition. 

Between 1998 and 2005. a second phase of 
sampling was undertaken, employing the 
same methods over the same small area, 
but with most individual records gathered 
via VES efforts and few collected aside from 
road-kill specimens. The 1986-88 and 
1998-2005 periods are treated as equivalent 
"'primary'" units for comparative purposes, 
and each is further divided into equivalent 
shorter "secondary" periods on the basis of 
an arbitrary (and partially climate-based) 
division of total sampling effort (see Meth- 
ods). 

With the exception of Cusco Amazonico 
in Peru (Duellman. 1995. 2005). most previ- 



ous studies from single upper Amazonian 
sites were conducted over relatively short 
spans of time (5 years or less), and within 
largely undisturbed expanses of continuous 
forest. The present paper is distinct in 
presenting data on the long-term composi- 
tion of the herpetofauna] community within 
a forest fragment remnant; as such, it is 
offered as another single-site reference point 
for the upper Amazon, in the form of a 
traditional species checklist, and as a neces- 
sary preface to forthcoming accounts on 
species-level abundance and distribution 
derived from the same sampling efforts. 

This study was undertaken several years 
before widespread interest in reports of 
amphibian declines (Blaustein, 1994; Stuart 
et al , 2004); despite concerns over various 
"enigmatic'" declines with causal factors 
ranging from epidemic disease to global 
climate change, the effects of habitat change 
are understudied and perhaps underempha- 
sized for both amphibians and reptiles 
(Gardner et al, 2007), especially in the 
Neotropical lowlands, where long-term mon- 
itoring data remain very rare (Duellman, 
2005:49; Whitfield et a!.. 2007:8355). The 
EBJS provides an established lowland re- 
serve with a well-known history as a forest 
fragment, and is an appropriate site for 
monitoring the effects of habitat change on 
an exceptionally species-rich Neotropical 
herpetofauna. 

METHODS 

The following summary of methodology is 
divided into three sections, progressing from 
( 1 ) a detailed description of the physical 
characteristics, history, and climate of the 
EBJS; to (2) an account of the long-term 
division of sampling effort and major goals 
within the context of this history: and to (3) 
the description of specific sampling protocols 
employed in the field as well as the scope and 



BREVIORA 



No. 514 



deposition of specimens and records thus 
obtained. Online material will be available 
through the journal's website. 

I) Description of the study site: forest, 
history, and climate 

The EBJS is located in eastern Ecuador in 
Napo Province at elevations of 400 to 450 m 
(01°04'S, 77°37'W at main station complex; 
see Figs. 1-4): the reserve is owned and 
administered by Fundacion Jatun Sacha 
(FJS), an Ecuadorian conservation founda- 
tion. The EBJS was the first of several 
reserves established by FJS and originally 
(1985-88) consisted of 230 ha of preserved 
forest owned by three independent landown- 
ers. After an initial 150-ha expansion funded 
by the Grateful Dead music group in 1989 
(via their Rex Foundation), subsequent 
acquisitions of additional land have greatly 
increased the size of the reserve, and current 
total land holdings are 2,334 ha. 

The EBJS forest is classified as Tropical 
Wet Forest (Canadas, 1983; Holdridge, 
1967) and lies between the confluence of 
the Napo River to the north and smaller 
Arajuno River to the south (Figs. 1, 4). 
Forest within reserve boundaries consists of 
approximately 70% primary forest, with the 
remaining 30% composed of various succes- 
sional stages of secondary forest growth, 
including approximately 95 ha of open-for- 
mation riparian habitats (Fig. 3). 

Although the EBJS preserves a substantial 
tract of lowland rainforest, since 1985 the 
upper Napo region has undergone substan- 
tial changes resulting from deforestation and 
forest fragmentation because of increasing 
colonization and development. Most adja- 
cent bordering areas have been logged or 
affected by agricultural activities, although 
no precise survey data are available. Satellite 
images showing general overviews of forest 
cover at the EBJS and in adjacent areas 




Figure 1. Current land holdings of the EBJS reserve 
(gray shaded areas within dark borders); location in 
eastern Ecuador is indicated by arrow on inset map at 
top left. Star indicates EBJS main station complex at 
approximately 01°04'S, 77°37'W. The area located 
between land tracts A and B comprises the Central 
Study Area (CSA; see Fig. 2); tract A is cleared cattle 
pasture, whereas tract B is a mixture of agricultural 
fields and various secondary forest successional stages. 
Also see text and Figures 3. 4. Approximately to scale. 

(dating from 2000 and 2001) are presented in 
Figure 4 and online Figure 1 . 

Before the early 1980s, the region was 
sparsely populated and accessible only by 
trails or river transport, although forest 
conversion through small-scale logging and 
agricultural clearing had already affected the 
area for many years, especially along the 
banks of both major rivers. In March 1986, 
the construction of a road (from the west) 
reached the north-central area of the EBJS 
(Figs. 1-4), and ongoing construction now 
extends approximately 30 km further east. 



2008 



\\l \/()N I OKI SI I R \(,\|| \|||1 Ki-i |()| ,\l \ \ 




Figure 2. The Central Study Area (CSA) in the 
north-central region of the EBJS located between two 
deforested land tracts. A and B (also see Figs. 1. 4. and 
text). All sampling efforts as summarized in the text and 
online Table 1 were conducted on trail routes (irregular 
lines in central area) extending north and south from the 
EBJS main station complex (star) and north from the 
Librarius Study Center complex (solid circle), as well as 
along the margins of the road to a distance of 
approximately 300 m to the east and west of the main 
station. The gray-shaded area is shown in detail in 
Figure 3. Approximately to scale. 

The discovery of oil reserves in the region 
has accelerated the process of colonization in 
recent years, constituting an increased threat 
to the EBJS reserve in the form of additional 
r6ad traffic, and deforestation in nearby 
areas. In November 2003. a high-pressure 
crude oil pipeline was installed along the side 
of the road (buried to a depth of 2 m), and in 
early 2006. the road was widened by 4 m and 
paved with asphalt. Although the immediate 
area affected by the road itself within the 



reserve is relatively small, the larger regional 
effect o\' increased development is the pre- 
dominant threat, as the KB.IS has increas- 
ingly become an "island" reserve surrounded 
b\ fragmented forest. 

The present study reports primarily on 
sampling conducted in a small (approximate- 
ly 100 ha) central study area (CSA) border- 
ing the Napo River (Figs. 1 4); the CSA 
includes all major habitats represented in the 
area (primary forest, and various river- 
floodplain forests, swamps, and secondary 
forest; Fig. 3), in contrast to land acquisi- 
tions since 1990, which have focused almost 
entirely on additional primary forest to the 
south and west. This concentration of forest 
types provides the most appropriate site to 
sample the overall amphibian and reptile 
diversity, in that some species occupy only 
limited portions of the various habitats 
within the EBJS as a whole. 

The CSA is also the area most directly 
affected by the road (and forest clearing and 
fragmentation in adjacent areas) for the 
longest period of time. Two deforested tracts 
of land, in total area approximately equal in 
size to the CSA, border this area directly and 
entirely to both the east and west (see 
Figs. 1, 2, 4); both of these adjacent areas 
had already been partially cleared before 
1986, although the largest (to the west) had 
undergone 10 years of secondary growth 
before being cleared again in 1990. 

Thus, although the total area of preserved 
forest is today far larger by the acquisition of 
additional land, in keeping with the long- 
term goals of the study begun in 1986 (see 
below), subsequent sampling efforts have 
focused almost exclusively on this same small 
area. 

The Ecuadorian Amazon region receives 
one of the highest levels of rainfall within all 
of lowland Amazonia. In contrast to more 
easterly areas, however (exhibiting pro- 
nounced wet and dry seasons), climate at 



BREVIORA 



No. 514 




Figure 3. Map (bottom, approximately to scale), and diagrammatic cross section (top) showing relief and 
habitat distribution within the CSA (also see Figs. 1, 2. 4). All 1986-88 period sampling was conducted within this 
narrow plot and along the margins of the road to a distance of ~300 m to the east and west; approximately 70% of 
1998-2005 period sampling effort was also conducted within this same small area. Areas A to D form the floodplain 
of the Rio Napo, area E includes the road (R) and early successional growth and ephemeral ponds (EP) along the 
road margins, and area F is closed-canopy primary forest on steeply rolling hills bisected by small streams (S). Within 
the floodplain region, area A is an ephemeral fluvial swamp surrounded by crop trees and successional growth; area 
B was (until 1990) cleared and planted with corn every —6 months; area C is older secondary forest with many large 
trees; area D is a permanent (flowing) fluvial swamp with low Heliconia growth. Aside from increased secondary 
growth in areas A, B, and C (secondary forest in area B is now continuous with area C). this habitat distribution has 
remained largely unchanged since 1986. 



the EBJS is far less seasonal, and rainfall can 
be highly irregular within any given year. 
The yearly distribution of rainfall is relevant 
to the distribution of sampling effort for the 
purpose of comparing faunal records be- 
tween sampling periods (discussed below). 
Rainfall has been recorded at the EBJS since 
1986. although data are incomplete for some 
years; for one 10-year period with complete 
data for each month (1994-2003; see online 
Table 1 ), total rainfall per year averaged 
3,71 1 mm (ranging from 3,051 to 4,593 mm). 
This record also indicates that March- 
August are on average the wettest months, 
and September-February show the lowest 
average rainfall. Within any single year, 
however, this general pattern can vary 
greatly; for example, in 1996 January had 
the highest rainfall of any month (510 mm), 
and in 1997 rainfall for June and July (255 



and 210 mm, respectively) was lower than 
both November and December (335 mm in 
each month). 

II) Temporal distribution of sampling effort 
and major goals 

A Coarse-Scale Sequence from "Before" 
to "After." Collections and records of 
observations of the amphibian and reptile 
fauna of the EBJS include two major 
(primary) sampling periods, each including 
approximately equal sampling effort, the 
first from 15 March 1986 to 15 February 
1988 (536.0 hours total effort) and the 
second spanning 1 July 1998 through De- 
cember 2005 (581.0 hours). Each primary 
sampling period is arbitrarily subdivided into 
secondary sampling periods (discussed be- 
low). The two primary sampling periods 



2008 



AM A/ON I OR1 SI I K \ci\ll \I III RIM I ( )| \l \ \ 





Figure 4. Landsat 7 satellite image (30 m/pixel resolution), captured 9 September 2001. showing forest cover 
within the EBJS reserve and closely adjacent areas. Refer to Figures 1-3 for location of major landscape features 
relative to reserve borders; the two outlined rectangles approximately indicate deforested areas A and B to the east 
and west of the CSA. Also see online Figure 1 showing forest cover over a larger area. At right, adult Ranitomeya 
duellmani (Dendrobatidae). shown 1.5X natural size. 



reported here can be interpreted as "pre- 
fragmentation" (1986-88) and "postfrag- 
mentation" (1998-2005) data on the EBJS 
fauna, although as previously noted, forest 
conversion processes in the area had been 
ongoing even before 1985. 

An initial program of removal sampling 
from 1986 to 1988 was undertaken as a 
baseline inventory of the fauna to facilitate 
subsequent monitoring of the same area in 
later years (1998-2005) via VES efforts, with 
three major goals. The first goal was to 
generate an initial species list (and a record 
of museum specimens), while employing a 
sampling technique that could be accurately 
repeated in later years. 

A second goal was to accumulate equiv- 
alent amounts of sampling effort for each 
sampling period, with the use of results 
from each to monitor trends in the presence 
or absence (persistence) of the many species 



to assess the overall stability and similarity 
of the community over time (on a chrono- 
logically coarse scale). This is an ongoing 
process; hypothetically, each sampling pe- 
riod (and all periods combined) can provide 
a predictive "before" reference point to 
compare with subsequent equivalent sam- 
pling efforts "after,"'' as one measure of the 
efficacy (or lack thereof) of the EBJS 
reserve in preserving the herpetofauna at 
this increasingly isolated site (the third 
major goal, with potential larger scale 
relevance for conservation in the region as 
a whole). Sorenson's quotient of similarity 
(SQ = 2J[A + E\) is used to compare the 
percent similarity among all sampling 
periods (see Results and Table 2). This 
simple formula estimates similarity inde- 
pendent o'i the individual abundance of 
each species, where A is the number of 
species in sample 1, B is the number in 



BREVIORA 



No. 514 



Table 1. Distribution of numbers of amphibian and reptile species (top) and individual specimen records 

(bottom) for all sampling periods from the ebjs (also see text, appendix 1, and online table 1). species 

totals for the 1998-2005 primary period indicate the number of species recorded, followed by the number of 

additional species (+) recorded later from 2006 or 2007. far right column presents the total number of 
species from all sourc1 s (including "r" species) from 1986 through 2007, as listed in appendix 1. individual 
specimen totals indicate the number of individuals recorded in each period; numbers in parentheses indicate 

the individuals per effort-hour for each period. 









No. 


by Sampling 


Period 














Total 






Total 


Total: All Sources 




1986-1987 1987-1988 1986-1988 1998-2001 2002-2003 


2004-2005 


1998-2005* 


1986-2007* 


Species: 


















Amphibian: 


63 


63 


73 


63 


54 


61 


78(+l) 


84 


Reptile: 


33 


39 


46 


34 


31 


55 


66(+7) 


82 


Total: 


96 


102 


119 


98 


86 


116 


144(+8) 


166 


Individuals: 


















Amphibian: 


1.298 


1,329 


2,627 


1.407 


989 


831 


3,227 


5,854 




(4.87) 


(4.92) 


(4.90) 


(6.54) 


(5.47) 


(4.47) 


(5.55) 


(5.24) 


Reptile: 


152 


170 


322 


157 


155 


234 


546 


868 




(0.57) 


(0.62) 


(0.60) 


(0.73) 


(0.85) 


(1.26) 


(0.93: 


(0.77) 


Total: 


1,450 


1,499 


2,949 


1,564 


1.144 


1,065 


3.773 


6.722 


""Individual totals in the 


1998-2005 and 1986 


-2007 columns do not include recoi 


•ds from 2006 and 2007. 



sample 2, and J is the number of species 
common to both samples. These pairwise 
comparisons result in SQ values between 
and 1, indicating no similarity and 1 
indicating complete similarity (SQ X 100 = 
% similarity). 

Additional goals were to assess population 
fluctuations by gathering time-series abun- 
dance (and ecological distribution) data for 
all species individually at the EBJS. The 
description and evaluation of species-level 
patterns of abundance and distribution are, 
however, beyond the scope of this paper, and 
Table 1 presents data solely on total num- 
bers of individuals in each major group 
(amphibians and reptiles) from each sam- 
pling period. Interpretation of this coarse- 
scale data is problematic, in that multiple 
variables influence abundance of the com- 
ponent species independently, especially 
among amphibians, with fluctuations in 
activity and apparent abundance often de- 
pendent on brief and irregular periods of 



high rainfall (but also see Summary and 
Discussion). 

Although the division of sampling effort 
presented in online Table 1 results in two 
primary periods approximately equal in total 
effort-hours, the second primary period 
spanned a far longer number of years. The 
arbitrary division of sampling effort as 
shown in Table 1 and online Table 1 is a 
compromise that uses the available sampling 
effort and climate data to provide the most 
equivalent periods for comparative purposes. 

The first primary sampling period is 
divided into two secondary periods (see 
online Table 1 ), providing two approximate- 
ly equal units (in total effort-hours); this 
division also approximately corresponds to 
each of two annual climate periods (see 
below). 

The second, much longer primary sam- 
pling period from 1998 to 2005 is divided 
into three secondary periods, varying greatly 
in the monthly distribution of effort-hours 



2008 



AMAZON 1 OKI SI 1 RAC.MI NI 111 Rl'l IOI UNA 



1~ VHl t 2. PaIRWISI COMPARISONS Ol \ll PRIMARY AND SECONDARE SAMPLING PERIODS FROM Mil EBJS FOR AMPHIBIANS 
AND REPTILES. Till COTA1 NUMBERS Ol SP1 CIES FROM EACH I'l RIOD \KI IMiliAl I I) IN m >i l )i \< I i\ I m COMMON CELL, 
\M> llll M MBERS OF SPECIES SHARED BY PERIODS ARI INDICATED Al mi UPPER RIGHT. SIMILARITY QI OTIENT (SQ) VALUES 

■\RI INDICAT1 I) IN ITALICS \l I OW] K l l l l 



Sampling Period 


1986 1987 


19S7 1988 


1998 2001 


2002 2003 


2004-2005 


1986-1988 


1998 2005 


Amphibians 
















1986 1987 


63 


53 


53 


47 


51 




59 


1987-1988 


.85 


63 


53 


45 


49 




60 


199S 2001 


.83 


.83 


64 


45 


48 


59 




2002 2003 


.80 


.77 


.76 


54 


51 


51 




20(14 2005 


.82 


.79 


.77 


.89 


61 


57 




1986-1988 


— 




.86 


.80 


.85 


73 


68 


1998-2005 


.84 


.85 


— 






.90 


7S 


Reptiles 
















1986-1987 


33 


26 


23 


17 


25 




30 


1987-1988 


0. 72 


39 


25 


19 


34 


— 


34 


1998-2001 


0.69 


0.68 


34 


18 


27 


27 




2002-2003 


0.53 


0.54 


0.55 


31 


24 


20 




2004-2005 


0.57 


0. 72 


0.61 


0.56 


55 


33 




1986-1988 


— 


— 


0.67 


0.52 


0.65 


46 


39 


1998-2005 


0.61 


0.65 


— 






0. 70 


66 



each year (online Table 1); this division, 
however, results in the most equal partition- 
ing of the total sampling effort from the 
1998-2005 period. 

Relative to annual climate variation, both 
the 1986-88 and 1998-2005 primary periods 
each include in total approximately equal 
proportions of sampling effort expended 
during the wetter months (March to August) 
and drier months (September to February) 
(see online Table 1). 

Ill) Field sampling protocol, museum and 
VES records, and taxonomy 

Most amphibian species at the EBJS are 
nocturnal; some reptiles are also nocturnal, 
whereas others may encountered resting at 
night, and sampling effort was almost 
exclusively concentrated on searches at 
night, with very little total effort (24 hours; 
2.1%) expended by day (these hours and 
records are lumped together with night 
sampling efforts). Thus, this study focused 
primarily upon the nocturnal herpetofauna 



(consisting mostly of amphibians) and those 
diurnal species of both amphibians and 
reptiles that typically rest at night within 
the areas searched. 

Sampling at night consisted of carefully 
searching the ground and vegetation, with 
spotlights, along the perimeters of established 
trails (transects) from ground level to a height 
of approximately 2 m aboveground and to a 
distance of approximately 2 m on either side 
of each trail. Although the actual lengths of 
trail segments searched per time unit of 
sampling effort were generally consistent 
(typically on the order of approximately 300 
linear meters per hour), the base unit used for 
comparisons between sampling periods is the 
person-effort-hour, defined as the hourly 
search effort expended by a single experienced 
person using the methods noted above. This 
was thus a form of time-constrained sampling 
technique (Campbell and Christman. 1982: 
Crump and Scott, 1994), applied to both 
removal and VES sampling efforts to facili- 
tate comparisons between sampling periods. 
Most nightly sampling periods ranged from 2 



II) 



BREVIORA 



No. 514 



to 5 hours in duration, and were conducted 
between 19:00 and 24:00. 

During the first primary period (1986 88), 
most specimens of most species encountered 
were collected and preserved, with 2,810 of 
the total 2,949 individual records represented 
by museum specimens. Collecting is now 
restricted at the EBJS, and records from the 
second primary period (1998 2005) are 
based almost entirely on VES efforts, includ- 
ing a total of 3,773 individuals. With only a 
few exceptions (see Appendix 1 and Results), 
the author personally identified all specimens 
(from all sampling periods) via direct obser- 
vation or examination of photographs (see 
below). 

Of the total 1,117 effort-hours of sam- 
pling, 635 hours (56.8%) were conducted 
personally by the author, whereas the re- 
maining 482 hours (43.2%) were conducted 
by persons who were trained on-site and 
supervised by the author, termed student 
collectors (SCs). Relative proportions of 
sampling effort by the author versus SCs 
were approximately equal for each primary 
period. Persons who assisted as SCs (and 
EBJS Park Guards), and their respective 
sampling effort contributions, are noted in 
the Acknowledgments. 

For all sampling before August 2003, the 
author was present on-site with all SCs and 
reviewed the results of each night's sampling 
efforts immediately: for the 1986^88 period, 
and from 1998 to 2001, all specimens encoun- 
tered by SCs were captured and then identi- 
fied (in 1986 88, most were preserved; in 
1998-2001, all were released within 48 hours). 

Beginning in March 2002, the physical 
capture of specimens by SCs was discontin- 
ued and digital cameras were used to 
photograph every amphibian and reptile 
specimen encountered; all other aspects of 
sampling methods remained the same, and 
identifications were determined solely from 
photos. 



From August 2003-December of 2005, a 
large portion of the total sampling effort 
(231.5 hours) was conducted by Park 
Guards employed by the EBJS. In a project 
funded by the Declining Amphibian Popu- 
lations Task Force (DAPTF), park guards 
were trained on-site in search techniques and 
the use of digital cameras. Park Guards then 
continued periodic sampling in the author's 
absence, and images of all individuals 
encountered were saved on compact disc 
(CD) for later review and identification 
(1,065 identified individuals). 

Although most of the 6,722 individuals 
from all sampling periods were recorded 
during regularly measured sampling efforts, 
a total of 376 records (5.6%) represent 
individuals that were randomly encountered 
outside the 1,117 hours summarized in on- 
line Table 1. Appendix 1 also includes all 
species known to the author from the EBJS 
since 1986 from all sources, including addi- 
tional records from 2006 and 2007, and a few 
important records from other investigators 
(see Appendix 1, Table 1, and Results). 

The majority (75%) of specimens collected 
during the 1986-88 removal sampling effort 
are deposited in the herpetological collec- 
tions of the Museum of Comparative Zool- 
ogy at Harvard University (MCZ); the 
remainder are housed in other collections 
(primarily in Ecuador). The following list of 
additional museum collections is presented in 
approximate decreasing order of the total 
number of included specimens at each 
museum (with acronyms): Museo Ecuator- 
iano de Ciencias Naturales, Quito (MECN); 
Museo de Zoologia, Pontifica Universidad 
Catolica del Ecuador, Quito (QCAZ); Amer- 
ican Museum of Natural History (AMNH); 
Florida Museum of Natural History (UF); 
United States National Museum (USNM); 
University of Kansas Museum of Natural 
History (KU). All specimens collected from 
1998 to 2007 are deposited at the QCAZ (all 



2008 



AMAZON FOREST FRAGMENT I II RIMTOFAUNA 



referenced by the author's "GV" field 
numbers in species accounts below). Dupli- 
eaie copies o\~ the photographic record from 
2003 to 2005 (with identifications and full 
data) are deposited at MCZ, MECN. and 
QCAZ and with the DAPTF. Online Ap- 
pendix 1 presents a complete list of all 
museum specimens; full data for MCZ 
specimens can be accessed on-line at www. 
mcz.harvard.edu. 

Since studies were initiated in 1986, the 
nomenclature for more than 70% of species 
o( amphibians (and some reptiles) known 
from the EBJS has been revised; for some 
species, both generic and family assignments 
have changed as a result of numerous recent 
phylogenetic investigations (see Frost, 2007). 
The present paper uses the most current 
published taxonomy, following the frequent- 
ly updated online references of the QCAZ of 
Coloma (2005-2007) for amphibians (which 
largely follows Frost, 2007), and Coloma et 
al (2000-2007) and Torres-Carvajal (2000- 
2007) for reptiles (excepting Iguanidae). 
These nomenclatural changes are likely to 
remain the subject of future debate (Frost, 
2007; Wiens, 2007, 2008); adoption of the 
most recently proposed binomials as em- 
ployed here is not necessarily intended as an 
endorsement of any of these arrangements, 
as much as an attempt to best present the 
most current and complete historical se- 
quence of taxonomic opinion. 

RESULTS 

Appendix 1 lists all species reported from 
the EBJS by all sources from 1986 to 2007, 
indicating the presence or absence of records 
in each sampling period; Table 1 summarizes 
data on numbers of species and individual 
records for all sampling periods. 

In total, 84 species of amphibians and 82 
species of reptiles have been recorded from 
the EBJS since March 1986. Total amphibian 



species include 79 anurans, 2 salamanders, 
and 3 eaeeilians. whereas reptiles include 4 
chelonians, 2 eroeodilians. 1 amphisbaenian, 
21 lizards, and 54 snakes. 

During the first primary period (1986 88), 
a total of 73 species of amphibians and 46 
reptiles were recorded, whereas the second 
primary period (1998 2005) included a total 
of 78 species of amphibians and 66 reptiles. 
These totals, and comparisons of sampling 
period totals presented below, all exclude 
three species included in Appendix 1 on the 
basis solely of reports by EBJS staff C'R" 
species; see Appendix 1 ). 

Of 73 species of amphibians recorded in 
1986 88, only five (6.8%) were not recorded 
again from 1998 to 2007 {Hylomantis hulli, 
Hypsihoas geographicus, Nyctimantis rugi- 
ceps, Edalorhina perezi, and Strabomantis 
sulcatus), and 1 1 species that were not 
recorded in 1986-88 were added between 
1998 and 2007 (see Appendix 1 ). 

Of 46 reptiles recorded in 1986-88, six 
species (13.0%) were not recorded again in 
1998-2007 (C/wlonoiciis denticulate), Anolis 
nitens scypheus, Mabuya nigropunctata, Ur- 
acentron flaviceps, Liophis reginae semili- 
neata, and Ninia hudsoni); 33 reptile species 
were added in 1998-2007. Of the latter 33, 
three species (Anolis transversalis, Polychrus 
liogaster, and Corallus caninus) were record- 
ed by other investigators in 1989 but are 
treated as new additions relative to the 1986- 
88 sampling efforts (see Appendix 1, and 
species accounts below). 

In total, 93.2% of amphibian and 87.0% of 
reptile species recorded at the EBJS in initial 
1986-88 sampling were recorded at some 
point again between 1998 and 2007. 

One crude measure of similarity among 
sampling periods is the total number of 
species recorded in each period relative to 
the quantity of sampling effort, which can be 
expressed as simply the number of species 
per effort-hour (SPH). Total SPH continu- 



12 



B RE VI O R A 



No. 514 



ally decreases with increasing sampling effort 
as species accumulation approaches an as- 
ymptote; relevant comparisons can be made 
between the approximately equivalent five 
secondary and two primary periods. SPH 
values noted below exclude species recorded 
solely from 2006-07 (see Appendix 1 and 
Table 1). 

SPH values for amphibians for secondary 
periods are 0.24 (1986-87), 0.23 (1987-88), 
0.30 (1998-2001), 0.30 (2002-03), and 0.33 
(2004^05). Amphibian SPH for the 1986-88 
primary period was 0.14, and 0.13 for the 
1998-2005 primary period. SPH values for 
reptiles are 0.12 (1986-87), 0.14 (1987-88), 
0.16 (1998-2001), 0.17 (2002 03), and 0.29 
(2004-05) for secondary periods and 0.08 
(1986-88) and 0.13 (1998-2005) for the two 
primary periods. These data, while crude, 
indicate that SPH exhibited no obvious 
decreasing trend for amphibians (with values 
for both primary periods nearly equal), 
whereas reptile SPH values suggest an 
increasing trend over all comparable sam- 
pling periods between 1986 and 2005. 

Table 2 presents pairwise comparisons of 
all sampling periods using Sorenson's simi- 
larity quotient (SQ; see Methods), including 
the similarity of both primary periods to all 
secondary periods. Primary versus secondary 
periods, however, are not directly compara- 
ble, in that total sampling effort for each 
primary period is far greater than total effort 
for each secondary period. Similarity in the 
long-term persistence of species richness and 
composition is a central focus, and thus the 
chronologically most widely spaced periods 
are of special interest. Comparisons of 
similarity in species composition in Table 2 
exclude records from 2006 and 2007 (Ap- 
pendix 1 and Table 1). 

For amphibians, SQ values indicate a high 
percent similarity among all sampling peri- 
ods, ranging from 0.76 to 0.90. Similarity of 
each of the first two secondary periods to all 



three later equivalent periods ranges from 
0.77 to 0.83. The amphibian species SQ for 
the two primary periods in total is 0.90. 
Together, these values suggest a robust 
persistence in overall species composition of 
the EBJS amphibian community over an 
almost 20-year span of sampling. 

For reptiles, SQ values are lower and 
more variable than for amphibians, ranging 
from 0.53 to 0.72 for equivalent periods. 
Reptile SQ for the two primary periods is 
0.70, despite an almost equal total number 
of species; however, as summarized in 
Table 1, apparent abundance of reptiles as 
a group is far lower overall than for 
amphibians, with most species (especially 
snakes; > 65% of total reptile species) 
represented by far fewer individual and 
sampling-period records. 

The 0.70 similarity of the two primary 
periods nevertheless suggests a strong overall 
persistence in reptile species composition 
given that the majority of these persistent 
species were commonly encountered in reg- 
ular night sampling (whereas many others 
were known only from random encounters), 
and 27 of 66 species recorded in the 1998- 
2005 period (excluding 2006 and 2007 
records) were not recorded in the first 
primary period. 

SELECTED SPECIES ACCOUNTS 

Having summarized data on species com- 
position over all sampling periods, it is 
appropriate to review some especially note- 
worthy species records. These include species 
described on the basis of type material 
collected at the EBJS, species not yet 
described when first recorded from the EBJS 
but later described on the basis of material 
collected elsewhere, and records that possibly 
represent undescribed species or for which 
identification is uncertain. Also noted are 
contributions from other investigators and 



2008 



AMAZON FOREST FRAGMENT HERPETOFAUNA 



I 5 



informants, including published studies on 
various species conducted at the EBJS, and a 
number o\' range extensions. Full names of 
contributors (with abbreviations) are pre- 
sented initially, with abbreviations alone 
used for any subsequent citations. For some 
accounts in which taxonomie history is 
noted, older generic names are used, with 
current names (as listed in Appendix 1) 
indicated in parentheses. The following 
accounts are presented in the same taxonom- 
ie order as in Appendix 1 . 

Amphibians 

Anura: Bufonidae 

Ehinella "margaritifera" (Laurenti) 

Previously widely known under the specif- 
ic epithet "typhonius," toads currently 
grouped under the name "margaritifera" 
include multiple cryptic species (as many as 
a dozen or more) distributed throughout the 
lowland Amazon basin (Fouquet et al, 2007; 
Hass et al, 1995). Duellman and Mendelson 
(1995) described and illustrated three forms 
from Loreto, Peru; among preserved adults 
available from the EBJS (from 1986 88), 
apparently at least two very similar forms are 
represented. 

Anura: Dendrobatidae 

AUobates zaparo (Silverstone), 

Ameerega bilinguis (Jungfer), and 

Ameerega parvula (Boulenger) 

Collections of these three similar species 
were conducted at the EBJS and nearby 
areas from 2003 to 2005 by Catherine R. 
Darst (C.R.D.) pursuant to studies concern- 
ing diet, toxicity, and the evolution of 
aposematic coloration and predator avoid- 
ance among these and other dendrobatids 
(Darst et al., 2005, 2006; Darst and Cum- 
mings, 2006). As a result of these efforts, the 
distribution patterns of all three species at 



the EBJS are now better understood; A. 
zaparo and A. bilinguis co-occur extensively 
in primary forest within the CSA, whereas A. 
parvula apparently occurs only in primary 
forest areas to the south (C.R.D., personal 
communication). 

Hyloxalus sauli (Edwards) 

This species is included on the basis of a 
report by Morley Read (M.R.) of males 
heard calling within the CSA in January 
2005; he has extensively documented the 
vocalizations oi~ Amazonian anurans and is 
familiar with this species at other localities 
(see Read, 2000). In addition, two individu- 
als collected in the 1987-88 period were 
tentatively identified as H. sauli (and are 
included as such in Appendix 1); these 
specimens were however apparently mis- 
placed and never cataloged into the MCZ 
collections, and their identity could not be 
confirmed. 

Among Hyloxalus species known from 
Amazonian Ecuador, two additional species 
(H. bocagei and H. shuar) could be expected 
at the EBJS on the basis of their known 
distributions (Coloma, 1995, 2005-2007). 
The paucity of Hyloxalus records is puzzling 
given the presence of presumed ideal habitat 
at the EBJS; although unidentified juveniles 
have occasionally been encountered while 
active by day (most from areas to the south 
of the CSA), adults have remained remark- 
ably elusive, and no records of Hyloxalus 
have been obtained in night sampling. 

Anura: Hylidae 
Cruziohyla craspedopus ( Funkhouser) 

This species was initially recorded by M.R. 
from a single adult photographed near the 
EBJS main station in April 1998. A second 
adult was found by Paul S. Hamilton 
(P.S.H.) in June 2007, and a third by M.R. 



14 



BREVIORA 



No. 514 



in August 2007 (none collected, and all 
within the CSA). 

Hylomantis hulli (Duellman and Mendelson) 

Three specimens (MCZ A 111179-180, 
118541; see online Fig. 2) collected in 1987 
were initially identified as Phyllomedusa 
(Hylomantis) buckleyi, a similar species 
known from the eastern Andean slopes in 
Ecuador from 400 to 1,870 m elevation 
(Cannatella, 1980). Following the descrip- 
tion of Phyllomedusa (Hylomantis) hulli by 
Duellman and Mendelson (1995), all three 
EBJS specimens were re-examined and found 
to conform to H. hulli, all exhibiting the 
diagnostic coarsely granular dorsal skin with 
heel calcars absent (skin finely granular and 
prominent calcars present, in H. buckleyi). 
These specimens apparently represent both 
the westernmost locality, and highest eleva- 
tional records (approximately 420 m) yet 
reported for this species. 

Hypsiboas cinerascens (Boulenger) 

In January 2005, M.R. reported multiple 
males of Hypisiboas cinerascens calling from 
a swamp area near the Rio Napo (within the 
CSA); this is the only known EBJS record. 

Osteocephalus cabrerai (Cochran and Goin) 

A large series of specimens collected in the 
1986-88 period were initially identified as 
Osteocephalus buckleyi (see MCZ online 
catalog and online Appendix 1); Hyla 
(Osteocephalus) cabrerai was described by 
Cochran and Goin (1970:215) but then was 
relegated to the synonomy of O. buckleyi by 
Trueb and Duellman (1971). Duellman and 
Mendelson (1995) removed O. cabrerai from 
the synonomy of O. buckleyi on the basis of a 
single specimen collected in Loreto, Peru, 
and provided color photos of the Loreto 



specimen of O. cabrerai and another of an O. 
buckleyi from the same region. On the basis 
of numerous photos of living specimens from 
the EBJS, as well as examination of speci- 
mens in the MCZ, all represent O. cabrerai. 
Because these were not directly compared 
with other specimens of either species and 
the two are morphologically very similar, the 
entire series at the MCZ remains cataloged 
as O. buckleyi. 

Osteocephalus deridens ( Jungfer, et al. ) and 
Osteocephalus fuscifacies (Jungfer, et al.) 

Although Osteocephalus deridens and O. 
fuscifacies were collected in the 1986 88 
period, both were initially confused under 
the name O. leprieurii. Following descrip- 
tions by Jungfer et al. (2000) on the basis of 
holotypes collected at the EBJS, the speci- 
mens from the 1986 88 period were reexam- 
ined, revealing one adult male of each species 
(O. deridens: MCZ A 111191; O. fuscifacies: 
MCZ A 111189). A call tape of the O. 
deridens was also recorded postcapture in 
1988, further confirming its identity (online 
Audio File 1 and online Fig. 3). 

Osteocephalus deridens is represented by 
multiple records since 2001, but the single O. 
fuscifacies from 1987 remains the only record 
obtained in this study; the O. fuscifacies record 
in Appendix 1 includes only the holotype 
collected in 1997 (Jungfer et al, 2000). 

Phyllomedusa coelestis (Cope) 

This species (online Fig. 4) is known from 
only two records at the EBJS (both adults, 
not collected), the first found by M.R. in 
January 2005, and the second by the author 
in October 2005; both records were from 
primary forest within the CSA. All diagnos- 
tic characters of the latter fully conform to 
the redescription and illustration provided 
by Duellman and Mendelson (1995). These 



2008 



\M \/()N I OKI SI 1 RACMINI III Rl'l l()l \l \ \ 



15 



apparently represent only the second and 
third records known from Ecuador, extend- 
ing the range approximately 200 km west 
from Yasuni. and are the highest elevational 
records tor this speeies (420 m). 

Scinax funereus (Cope) 

Although Scinax funereus was not initially 
recorded in this study until July 2003, 
Duellman and Wiens (1993) reported a single 
speeimen from the EBJS colleeted by Luis 
Coloma in May 1990. 

Anura: Mierohylidae 
Microhylidae incerta sedis 

This species is included on the basis of a 
single specimen (QCAZ: GV2000-0329) col- 
lected by day in June 2001 on the south side 
of the EBJS in primary forest near the Rio 
Arajuno. Although resembling Chiasmocleis 
ventrimaculata in dorsal and ventral colora- 
tion, at 32.0 mm snout-vent length (SVL), it 
is much larger (SVL 20 24 mm in C 
ventrimaculata) and apparently conforms to 
no species yet reported from Amazonian 
Ecuador. The dorsum was dull grayish- 
brown (body and limbs) with a thin pale 
middorsal line, the venter (throat, body, and 
limbs) mottled cream and pale brown; skin 
smooth (dorsally and ventrally), dorsal 
occipital fold absent; fingers and toes lacking 
webbing; tympanum and annulus not evi- 
dent; eyes very small, iris brown. 

Anura: Strabomantidae 

PristinuuUis aureolineatus 

(Guayasamin et al. ) 

The first record of P. aureolineatus from 
the EBJS is a single adult (MCZ A 109239) 
collected in May 1986 and initially identified 
as Eleutherodactylus ( Prist inumt is) lacrimo- 
sus, with two additional individuals (not 



collected) recorded in the 1998 2005 sam- 
pling period. Following description by 
Guayasamin et al. (2006), in which P. 
aureolinatus was assigned to the lacrimosus 
assemblage on the basis of type material 
from widely scattered localities in northeast- 
ern Ecuador and Peru, the three EBJS 
records were recognized as conforming to 
P. aureolineatus. These apparently represent 
both the westernmost locality and the highest 
elevational records (approximately 400 
425 m) yet reported for this species. 

Pristimantis librarius (Flores and Vigle) 

Flores and Vigle (1994) described 
Eleutherodactylus (Pristimantis) librarius on 
the basis of a series of 40 specimens, all 
collected at the EBJS during the initial 1986 
88 sampling period. This species was appar- 
ently known only from the EBJS until Elmer 
(2004) reported P. librarius from several 
localities throughout lowland eastern Ecua- 
dor. 

Pristimantis ockendeni (Boulenger) 

Kathryn R. Elmer made extensive collec- 
tions of Eleutherodactylus {Pristimantis) ock- 
endeni at the EBJS and throughout eastern 
Ecuador from 2003 to 2005 pursuant to 
studies on population genetic structure and 
diversity (Elmer et al, 2006, 2007a, 2007b). 
As a result of these studies, Elmer and 
Cannatella (2008) recently identified and 
described three cryptic species in eastern 
Ecuador (Pristimantis achuar, P. altamnis, 
and P. kichwarum), all of which had previ- 
ously been confused under the name E. 
ockendeni; they further speculate that 
Eleutherodactylus ( Pristimantis) ockendeni 
{sensu stricto) is most likely a name that 
has been misapplied to Ecuadorian popula- 
tions for many years and suggest that this 
species probably does not occur in Ecuador. 



16 



BREVIORA 



No. 514 



Among these three new species, both P. 
altamnis and P. kichwarum are reported from 
the EBJS. Presumably, records reported here 
include specimens of both new species (among 
material collected in 1986-88, as well as VES 
records from 1998 to 2005), although none of 
the many existing specimens (or photographic 
records) have yet been examined by the 
author (or Elmer and Cannatella, 2008) in 
comparison to these very recent descriptions, 
and all are thus included here under the single 
prior name "ockendeni." 

Pristimantis species I 

Pristimantis sp. I was initially recorded in 
the 1986-88 sampling period on the basis of 
a 32.0-mm SVL gravid adult female (MCZ A 
129304; online Fig. 5) and seven adult males 
ranging from 20.0 to 22.0 mm SVL. Al- 
though uncommon at the EBJS, additional 
records were obtained between 1998 and 
2005. Aside from its larger size, in both 
morphology and color Pristimantis sp. I is 
remarkably similar to P. martiae (males 
13.2-16.8 mm SVL, females 18.3 23.0 mm 
SVL; Lynch, 1974), a far more common 
species at the EBJS. Both species share a 
range of characters, including the lack of an 
external tympanum (although a partial 
tympanic annulus is faintly evident in some 
specimens of both species), similar snout 
shape, skin texture, dark supratympanic and 
canthal stripes, and a bronze iris with a 
broad dark-reddish horizontal median 
streak. Remarkably, both also share the 
same polymorphic dorsal coloration, with 
two discrete morphs exhibited in both 
species. In both species, some individuals of 
both sexes are pale brown to gray dorsally 
with 3 or 4 darker chevron-shaped transverse 
bands, whereas others have a lineate pattern 
of longitudinal pale to dark brown stripes of 
varying widths from the tip of the snout to 
the vent. Lynch (1974) noted a 3:1 ratio in 



chevromlineate morphs among specimens of 
P. martiae cited in the original description; at 
the EBJS, a nearly identical ratio was evident 
among 61 P. martiae collected in 1986-88. 
For Pristimantis sp. I, the lineate morph was 
more common, in an approximately opposite 
proportion. 

Duellman and Mendelson (1995) de- 
scribed Eleutherodactylus (Pristimantis) de- 
ltas from Loreto, Peru (approximately 
210 km SE of the EBJS) on the basis of a 
single adult female. This species bears a close 
resemblance to Pristimantis sp. I, sharing a 
nearly identical dorsal pattern of pale and 
darker brown stripes, and dark supratym- 
panic and canthal stripes; at 30.9 mm SVL, 
the holotype of P. delius also falls precisely 
within the observed size range for adult 
female Pristimantis sp. I at the EBJS. The 
two differ however in that P. delius exhibits a 
distinct tympanic membrane and annulus 
and lacks vomerine odontophores (tympanic 
membrane not evident, but with faintly 
evident annulus, and vomerine odonto- 
phores present in all preserved specimens of 
Pristimantis sp. I). Despite these differences, 
Pristimantis sp. I might be conspecific with 
P. delius; specimens from the EBJS have not 
yet been compared with the holotype of P. 
delius, and final determination must await 
direct comparisons. 

Pristimantis species II 

This species is included on the basis of a 
single individual (sex undetermined) found, 
but not collected, within the CSA in June 
1999 (online Fig. 6). Because this individual 
cannot be referred to any described species, a 
brief description is provided. 

SVL 21.5 mm; limbs short and robust in 
proportion to body, hands and feet lack 
webbing, first finger shorter than second, 
finger and toe pads expanded and rounded. 
Prominent supratympanic fold present, but 



2008 



AM A/ON I OKI SI I RAOMI Nl III KIM loiAUNA 



17 



tympanic membrane and annulus not evi- 
dent. Snout rounded dorsally and in profile. 
Skin o\' entire dorsum (head, body, and 
limbs) thick and glandular in appearance, 
covered with low. rounded tubercles; tuber- 
eles coalesce dorsolateral^ to form faintly 
evident, broken dorsolateral ridges. Entire 
dorsum uniform grayish-brown at night, but 
dark brown (almost black) by day. with 
widely scattered metallic-gold Hecks 0.5- 
2.0 mm in diameter (most obvious under 
magnification); faintly evident darker trans- 
verse bands dorsally on arms and legs. Entire 
venter brown except for pale gray groin area 
with a bluish tinge; anterior, posterior, and 
other concealed surfaces of legs unmarked 
pale gray. Iris silver with fine black reticula- 
tions dorsally, denser black reticulations 
ventrally, a broad rusty-red median stripe, 
and a very narrow red pupil ring. 



forest trail within the CSA in June L999. 
Because this individual (not collected) was 
clearly distinct from the other two species 
known from the EB.IS. but could not be 
identified by direct examination of the 
specimen in the field, a brief description 
follows (also see online Eig. 7). 

Total length approximately 360 mm; mid- 
body diameter 1112 mm; dorsum deep blue 
fading to paler blue ventrolaterally; midven- 
trally also deep blue, slightly darker than 
dorsum. Head elongate, tentacles thin and 
white to cream; eyes not evident externally 
aside from very small pin-point dermal 
depressions. Number of costal grooves ap- 
proximately 1 10. 

Reptiles 

Chelonia: Chelidae 

Mesoclemmys gibba (Schweigger) 



Caudata: Plethodontidae 

Bolitoglossa equatoriana (Brame and Wake) 

and Bolitoglossa peruviana (Boulenger) 

Cisneros-Heredia (2006) reported several 
new geographic records for B. equatoriana 
and cited one specimen collected at the EBJS 
in 1999 as representing the southernmost 
record for this species, which is known only 
from east-central Ecuador; the EBJS is 
apparently also the easternmost and highest 
elevation locality. 

Both B. equatoriana and B. peruviana were 
among the most ubiquitous and abundant 
amphibians in all forest habitats at the EBJS 
throughout all sampling periods, each repre- 
sented by more than 300 individual records 
Since 1986. 



DeForce et al. (2004) conducted mark- 
recapture studies on Mesoelenmivs gibba at a 
small pond within the CSA in 2001 but were 
unable to estimate population size because of 
extreme "trap-happy" behavior. 

Chelonia: Podocnemididae 
Podocnemis unifilis (Troschel) 

This species is included on the basis of a 
photograph of several adults basking on a 
log in the Rio Arajuno on the south side of 
the EBJS (taken by a visitor; date unknown). 
The presence of P. unifilis along the Rio 
Arajuno bordering the EBJS is also pre- 
sumed on the basis of observations of this 
species from the Rio Arajuno near its 
confluence with the Rio Napo. 



Gymnophiona: Caeciliidae 
Caeciliidae incerta sedis 



Crocodylia: Crocodylidae 
Caiman e. crocodilus (Linnaeus) 



This species is included on the basis of a EBJS Founder and Director, G. Alejandro 

single individual found at night on a primary Suarez (G.A.S.) observed what was appar- 



IS 



BREVIORA 



No. 514 



ently an adult C. crocodilus in a fluvial 
swamp on the floodplain of the Rio Napo 
(within the CSA) in 1990. No additional 
reports are known from the EBJS; however, 
some might survive in more remote areas of 
the Rio Arajuno on the south side of the 
reserve. 

Sauria: Iguanidae 

Anolis transversalis (Dumeril) and 

Polychrus liogaster (Boulenger) 

Both A. transversalis and P. liogaster were 
initially recorded by Laurie J. Vitt (personal 
communication) in May 1989; identification 
(via photos) of the P. liogaster was later 
determined by William W. Lamar (W.W.L.). 
Although excluded from 1986-88 sampling 
period totals, both species were recorded 
again in the 1998-2005 period (and in 2006). 
The former was recorded in June 2005 (by 
P.S.H.) and August 2006 by Alexis Harrison 
(one adult each), and the latter from July 
2003 and September 2006 (also one adult 
each; the 2003 record is shown in online Fig. 
8). 

The single P. liogaster from 2006 was a 
road kill (QCAZ: GV2000-0349) collected by 
EBJS Park Guard Milton G. Orozco 
(M.G.O); this and previous EBJS records 
apparently represent the first reports from 
Ecuador, and a very substantial range 
extension for this species, extending the 
range approximately 1,000 km to the NW 
from the nearest reported records in western 
Brazil (see Avila-Pires, 1995:131, 134). 

Serpentes: Boidae 
Corallus caninus (Linneaus) 

The first record was an adult photo- 
graphed by M.R. in 1989 (date unknown; 
not collected); two additional adults were 
recorded in 2006 (all were found within the 
CSA). 



Serpentes: Colubridae 
At raet us species 

Among material collected in 1986-88 (as 
well as additional VES records since 1998) 
are specimens of a small Atractus, which in 
diagnostic characters of lepidosis conform to 
Atractus major (MCZ R 173868-871; online 
Fig. 9); however, these form a series consis- 
tently varying from all other EBJS specimens 
referred to A. major and are here treated 
tentatively as a possibly distinct cryptic 
species pending the collection and evaluation 
of additional specimens. 

All individuals of Atractus sp. have a 
dorsal pattern of alternating short and long 
dark red bands, with each band separated by 
a shorter black band; the short red bands are 
paler and brighter in shade than the long 
bands, and all bands extend unbroken 
laterally across the dorsum. In contrast, all 
A. major exhibit a dull brown dorsal ground 
color with short dark brown bands and 
irregular markings (with short pale tan 
outlines) that are broken middorsally with 
few (if any) bands laterally continuous across 
the dorsum. Morphologically, there is a 
striking difference in the size of the eye; the 
eyes of Atractus sp. are distinctly smaller 
relative to head size than are those of A. 
major. These differences are obvious if 
specimens are compared directly. 

Additional Notes on Colubridae 

Several colubrid species and period records 
are included in Appendix 1 on the basis of 
reports or identifications by other investigators, 
but otherwise require no further commentary. 

Records from 1998 to 2005 of Dipsas 
pavonina, Drepanoides anomalus, and Oxy- 
rhopus formosus were obtained by Park 
Guards Willan G. Poveda and M.G.O.; 
identifications (from photos) for all of these 
were kindly determined by W.W.L. 



2008 



AMAZON I OKI ST I RAOMI NT III RIM IOI AUNA 



Pseustes s. sulphureus is included on the 
basis o\~ a photograph taken by Karl-Heinz 
Jungfer in December 1997 and subsequently 
identified by W.W.L. 

Initial records and identifications for 
Drymobius rhombifer, Oxyrhopus petola dig- 
italis, and Siphlophis cervinus were reported 
by Dale F. DeNardo (D.F.D.), Martin J. 
Fouquette, and P.S.H. from June 2005. They 
also reported all 2004 05 period records for 
Atractus occipitoalbus, Chironius exoletus, 
Oxyrhopus melanogenys, Siphlophis compres- 
sus, and Umbrivaga pygmaea (from June 
2005). and D.F.D. reported Oxybelis fulgidus 
from June 2007. 

Serpentes: Viperidae 
Lachesis muta (Linnaeus) 

Both G.A.S. and Park Guard Gabriel 
Tapuy (G.T.) have observed L. muta within 
the general upper Napo area, and G.A.S 
observed at least one within the CSA. A 
large skin owned by G.T. came from an 
individual collected in the area, although the 
exact locality is unknown. 

SUMMARY AND DISCUSSION 

In many respects, the EBJS forms a 
microcosm relevant to a number of important 
issues pertaining to Amazon rainforest con- 
servation, including questions concerning the 
appropriate minimum reserve size required to 
identify and conserve viable biotic communi- 
ties, the long-term effects of continuing 
development and fragmentation on surviving 
habitat islands, and the need to inventory and 
monitor all component organisms to address 
these questions ( Bierregaard et al. , 1 997, 200 1 ; 
Laurance et al. . 1997; Lewin. 1984; Simberloff 
and Abele. 1982; Turner, 1996; Turner and 
Corlett, 1996). 

The need to first inventory and later 
monitor is essential to determine what is 



being preserved and to assess the persistence 
o\' the community under study; the success of 
any conservation effort is best measured by 
documenting community persistence (and 
fluctuations) over long periods of time 
(Blaustein et al, 1994; Teder et al, 2007). 
Unfortunately, poorly known, yet species- 
rich, rainforest ecosystems require ongoing 
sampling effort combined with taxonomic 
expertise; a shortage of human and material 
resources often renders such efforts difficult 
and limited in scope (Bierregaard et al., 1997; 
Crump, 2003). 

The techniques employed in this study 
utilized minimal resources, including one 
principal investigator (the author), persons 
who were trained in field sampling methods 
(SCs and EBJS Park Guards), and (most 
recently) the use of new imaging technology 
(digital cameras) combined to gather simple 
long-term data on the persistence of species 
richness and composition. Although studies 
evaluating sampling methods for monitoring 
tropical herpetofaunas have recently been 
undertaken (Doan, 2003; Funk et al., 2003), 
whatever method is employed must be both 
consistent and sustainable over time if an 
accurate assessment of the community is to 
be attained (Teder et al., 2007). 

Assuming that the methods employed in 
this study gathered accurate data on long- 
term species persistence as a measure of 
conservation efficacy, results reported here 
suggest that the EBJS has, since 1986, very 
successfully conserved the resident herpeto- 
faunal community. After almost 10 years of 
isolation in an increasingly fragmented area, 
monitoring over the subsequent 10 years 
(1998-2007) revealed that 92% of amphibian 
and reptile species initially recorded in 1986 
88 were also present at some point later 
during 1998 2007. The number of species 
added in 1998 2007 for both amphibians 
(11) and reptiles (33) far exceeds the number 
of 1986-88 species lacking subsequent rec- 



20 



BREWORA 



No. 514 



ords from 1998-2007 (thus there were very 
few possible "ghosts"; see Dayton et al, 
1998). The increase in reptile species records 
for the 2004-05 period in particular is 
partially attributed to a combination of (1) 
increased vehicular traffic and increased 
effort in collecting the resulting road-kill 
specimens, (2) the use of digital cameras to 
document randomly encountered individu- 
als, and (3) contributions from other inves- 
tigators. New species records for both groups 
continue to accumulate; snakes now pre- 
dominate, although a large amphibian 
(Rhaebo glaberrimus) was added within the 
CSA as recently as June 2007. 

Pairwise comparisons for amphibians 
among equivalent sampling periods (far 
better sampled than reptiles, in total records 
represented) all reveal a high percent simi- 
larity in species composition (from 0.77 to 
0.90), whereas similarity for reptiles is lower 
but exceeds 0.50 for all equivalent pairs, and 
is 0.70 for the two primary periods. Crude 
measures of both species (SPH) and individ- 
uals per sampling effort-hour for both major 
groups vary among sampling periods, but 
none suggest any pronounced declines. In- 
terpretation of group-level abundance data is 
problematic; given the large sample sizes 
represented, however, lumped data on even 
this coarse scale provides baselines (and 
ranges of variation) that could be used to 
predict and monitor expected long-term 
patterns of abundance at the EBJS over 
future equivalent sampling periods. 

Perhaps the most remarkable result is not 
merely the robust persistence of the commu- 
nity, but the concentration of 161 species 
recorded from the CSA within a sampled area 
of only 100 ha directly bordering two large 
tracts of deforested land (Figs 1^4). More than 
80% of total sampling effort was conducted 
along precisely the same trails within the CSA 
over the entire 20-year span of sampling. Most 
of the interior regions of the reserve remain 



almost entirely unstudied; additional species 
may be expected from these areas. 

Including all records to date, the herpeto- 
faunal species richness of the EBJS is already 
among the highest ever recorded for any single 
Neotropical site. For amphibians, only Tipu- 
tini and Santa Cecilia (both located in 
Ecuador to the NE of the EBJS; Cisneros- 
Heredia, 2003; Duellman, 1978) and forests to 
the north of Leticia, Colombia (Lynch, 2005) 
have recorded more amphibian species (also 
see Doan and Arizabal, 2002; Duellman, 2005; 
Duellman and Thomas, 1996). Santa Cecilia 
has been completely deforested for many years 
as a result of colonization and oil extraction, 
leaving the EBJS second only to Leticia and 
Tiputini in amphibian species richness among 
currently extant sites. Reptile species richness 
at the EBJS is greater than reported for 
Tiputini (Cisneros-Heredia, 2003) and is 
exceeded only by Santa Cecilia within the 
upper Amazon region (Duellman, 1978) and 
Cusco Amazonico and Explorers Inn in the 
Tambopata region of southeastern Peru 
(Doan and Arizabal, 2002; Duellman, 2005). 

Only a few studies have addressed the 
long-term effects of forest fragmentation on 
anuran amphibians in lowland Amazon 
rainforest (none from the more species rich 
upper Amazon). Tocher et al. (1997) exam- 
ined pre- and post-isolation species richness 
among small (1-100 ha) fragments of prima- 
ry forest over a 10-year period in north- 
central Brazil (40 species). Contrary to their 
initial predictions, few species were lost from 
fragments following isolation, and fragments 
of all sizes showed increases in species 
richness compared with equal-sized plots 
within larger contiguous primary forest areas 
(partially attributed to an influx of distur- 
bance-tolerant species into fragments). Gas- 
con et al. (1999) also studied the persistence 
of anurans in fragments and surrounding 
disturbed matrix habitat in the central 
Amazon over spans of up to 19 years and 



:oos 



AMAZON FORI ST FRAGMENT HERPETOFA1 N \ 



21 



found that few species were lost, overall 
species richness increased, and matrix-toler- 
ant species (including many primary forest 
species) remained stable or increased in 
fragments. These investigators concluded 
that the Amazonian anuran community was 
less affected by fragmentation than other 
vertebrates and that many anuran species 
can better utilize and migrate through 
disturbed habitat, but they cautioned that 
further monitoring would be essential to 
discern more long-term patterns. 

In an earlier study from the same area, 
Zimmerman and Bierregaard (1986) com- 
pared species-area relation data derived from 
island biogeography theory with data on 
ecological factors in an attempt to test which 
approach was a better predictor of the 
minimum area required to preserve the 
anuran fauna (39 species). They concluded 
that on the basis solely of species-area data, 
only about 500 ha would be required to 
preserve this fauna, but habitat heterogeneity 
and, especially, the distribution of breeding 
sites were far better predictors of the 
minimum area needed to ensure community 
survival. Without sufficient breeding sites, 
even larger areas would not be adequate; 
with abundant breeding sites and heteroge- 
neity, an even smaller area might suffice. 

Results from this study provide another 
long-term reference point for an exception- 
ally rich Amazonian herpetofauna, suggest- 
ing that some relatively small reserves could 
be a viable conservation strategy for these 
taxa in response to forest fragmentation; the 
high concentration and persistence of species 
within the small but ecologically heteroge- 
neous (and extensively riparian) CSA is 
especially noteworthy in light of previous 
studies on Amazonian anurans. 

Contrary to some initial predictions made 
in 1986, the EBJS herpetofauna has evidently 
not suffered a catastrophic decline in species 
richness, despite more than 20 years amidst an 



increasingly impacted landscape. However, 
although 20 years might represent a substan- 
tial time span from a human perspective, this 
is merely a brief instant within the far larger 
context of millions of years of biological 
evolution in the Amazonian rainforest. This 
small reserve might provide a model for future 
conservation efforts in the region, but to 
repeat an admonition expressed in virtually 
all studies on fragmented ecosystems, time 
and continued monitoring will be necessary to 
address the many questions that yet remain. 

ACKNOWLEDGMENTS 

This paper is dedicated in equal measure 
to two persons, without whom the study 
would never have been initiated or complet- 
ed. First, the guidance and material support 
provided by the late Ernest E. Williams 
formed the foundation for the 1986-88 field 
work and all research efforts that followed. 
Second, the continued survival and expan- 
sion of the EBJS reserve to the present day 
can almost entirely be credited to the vision 
and dedication of EBJS Founder and Direc- 
tor G. Alejandro Suarez, whose assistance 
with all on-site aspects of this study has been 
indispensable for more than 20 years. 

Although space does not permit the ac- 
knowledgement of all persons who have 
assisted in varying ways, it is essential to note 
many substantial contributions. For the 1986 
88 period, the following served as "Student 
Collectors, ,, (with total contributed effort- 
hours in parentheses): Brock Dolman (75.0), 
Deborah Goberdhan (67.5), Judith Futerfas 
(14.0), and George Root (14.0). The late J. 
Anderson, D. Beauchamp, B. Bochan, the late 
A. Gentry, K. Gretter, M. McColm, the late J. 
Miller, D. Neill, C. Nieukirk, W. Wilbert, and 
School for Field Studies students all assisted 
with 1986-88 field work and logistics. 

During 1998 2007 field work, EBJS Park 
Guards contributed essential sampling effort. 



22 



BREVIORA 



No. 514 



knowledge, and camaraderie; these included 
Milton Orozco (99.5 effort-hours), Willan 
Poveda (97.0), Elibar Jimenez (19.5), and 
Gabriel Tapuy (15.5). In addition to all 
persons noted in accounts of species, various 
additional contributions in the field from 
1998 to 2007 are credited to N. Batcha, S. 
Goberdhan-Vigle, P. Grefa, C. James, S. 
Konningsor, R. Sawby, W. Schaedla, M. 
Shindell, and students in Arizona State 
University tropical biology courses held at 
the EBJS. 

For useful advice and discussion through- 
out the second phase of sampling, the author 
thanks D. DeNardo, M. Douglas, M. Fou- 
quette, W. Lamar, D. Oakey, D. Pearson, B. 
Sullivan, and T. Swanson. Translation of the 
abstract was kindly provided by G. Cadena 
and J. Logback. Special thanks to W.E. 
Duellman, W.W. Lamar, J.B. Losos, and 
J.D. Lynch for critical reviews of the ms. 



Assistance with identifications, museum 
use, and loans and depositions of specimens 
since 1986 have been generously shared by the 
late P. Alberch, J. Cadle, L. Coloma, G. 
Flores, J. Hanken, R. Heyer, F. Irish, J. 
Losos, J. Rosado, V. Wallach, and the late E. 
Williams. Assistance with obtaining collecting 
and research permits in Ecuador was provid- 
ed by S. Figueroa, J. Matheus, F. Sarmiento, 
R. Ulloa, The Museo Ecuatoriano de Cien- 
cias Naturales, and INEFAN and the Minis- 
terio de Agricultura y Ganaderia del Ecuador. 

The 1986 88 phase of this study was 
funded by the late E. E. Williams and the 
Museum of Comparative Zoology, Harvard 
University; 1998-2007 field work was par- 
tially supported by grants from the Arizona 
State University Graduate Research Devel- 
opment Office and the Declining Amphibian 
Populations Task Force (DAPTF; with 
special thanks to Tim Halliday). 



NOTES ADDED IN PROOF 



Subsequent to completion of the present 
manuscript, six additional species records 
from the EBJS have been reported; with the 
exception of one record (dating from 1988), 
all identifications were confirmed by the 
author from photographs (none collected). 
These are presented below in the same order 
as Appendix 1 . 

AMPHIBIA: Dendrobatidae: Hyloxalus bo- 
cagei. One adult was reported from an area 
1 .5 km NW of the EBJS station, in July of 2008, 
by Ross J. Maynard and Nathan A. Shepard. 

REPTILIA: Aniliidae: Anilius scytale. A 
single juvenile was recorded on the south side 
of the reserve (near the Rio Arajuno), 4 km 
SSE of the EBJS station, in July 2008 by Ross 
J. Maynard and Nathan A. Shepard. 

Colubridae: Atr actus torquatus. This species 
is tentatively included based on a photograph 
of one individual taken at night in primary 



forest (600 m south of the EBJS station, 
within the CSA) in July 2008, by Milton G. 
Orozco. Although determination was made by 
the author (and William W. Lamar) based on 
the photograph, this record would apparently 
represent a substantial westward range exten- 
sion and the first report from Ecuador; 
additional material is needed for confirmation. 

Colubridae: Liophis cobella. Reported by 
Bill Montgomery based on a photograph taken 
in July 1988 and later determined by William 
W. Lamar (probably from within the CSA). 

Colubridae: Oxybelis aeneus. One adult was 
recorded by day from a secondary forest area 
2.5 km SW of the EBJS station in August 
2008, by Milton G. Orozco. 

Viperidae: Bothriopsis taeniata. One adult 
was recorded by day near a stream in primary 
forest, 3.5 km SW of the EBJS station in 
March 2008, by Milton G. Orozco. 



2008 



AMAZON I OKI ST I R \(,MI NI 1 IIRPITOI Al INA 



23 



Appendix I. List oi species and distribution of records, 1986 2007.* Amphibians and 
reptiles recorded from till ebjs since march 1986 over mm sampling periods, \is<> 

including all species reported from al 1 sources through 2007 (also see text and 
Tables 1, 2). Presence or absi nce within i v h swum lng period (and other columns) is 

indicated by + or - symbols.! 









Presence in Sampling 


Period 










1986 


1987 


1998- 


2002- 


2004 




Species 


DFR 


1987 


1988 


2001 


2003 


2005 


CSA 


MS 


AMPHIBIANS 


















Anura: Bufonidae 


















Rhaebo glaberrimus 


6/07 


- 


- 


- 


- 


+* 


+ 


- 


Rhinella testae 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


R. "margaritifera"* 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


R. marina 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Anura: Centrolenidae 


















Cochranella midas 


8/87 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Hyalinobatrachium munozorum 


8/87 


- 


+ 


+ 


- 


- 


+ 


+ 


Anura: Dendrobatidae 


















Allobates zaparo* 


3/86 


+ 


- 


+ 


+ 


+ 


+ 


+ 


Ameerega bilinguis* 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


A. hakneli 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


A. parvula* 


8/87 


+ 


- 


- 


+ 


+ 


- 


+ 


Hyloxalus sauli* 


11/87 


- 


+ 


- 


- 


+ 


+ 


+ 


Ranitomeya duellmani 


7/87 


+ 


+ 


+ 


- 


+ 


+ 


+ 


R. ventrimaculata 


6/86 


+ 


+ 


- 


- 


+ 


+ 


+ 


Anura: Hylidae 


















" Hyla" alboguttata genus incerta 


















sedis 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


C 'ritziohyla craspedopus* 


4/98 


- 


- 


+* 


- 


+* 


+ 


- 


Dendropsophus bifurcus 


5/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


D. brevifrons 


4/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


D. marmoratus 


4/86 


+ 


+ 


- 


+ 


+ 


+ 


+ 


D. minimis 


6/99 


- 


- 


+ 


- 


- 


+ 


+ 


D. parviceps 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


D. rhodopeplus 


5/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


D. sarayacuensis 


8/87 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


D. triangulum 


7/86 


+ 


+ 


- 


+ 


- 


+ 


+ 


Hylomantis hulli* 


5/87 


+ 


+ 


- 


- 


- 


+ 


+ 


Hypsiboas boons 


3/86 


+ 


- 


- 


+ 


+ 


+ 


+ 


H. calcaratus 


4/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


H. cinerascens* 


1/05 


- 


- 


- 


- 


+ 


+ 


- 


H. fasciatus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


i H. geographiats 


8/87 


+ 


- 


- 


- 


- 


+ 


+ 


H. lanciformis 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


H. punctatus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Nyctimantis rugiceps 


8/87 


- 


+ 


- 


— 


— 


+ 


+ 


Osteocephalus cabrerai* 


6/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


O. deridens* 


2/88 


- 


+ 


+ 


+ 


+ 


+ 


+ 


O. fuscifacies* 


8/87 


- 


+ 


+* 


- 


- 


+ 


+ 


O. planiceps 


6/87 


+ 


+ 


+ 


+ 


+ 


+ 


+ 



24 



BREVIORA 



No. 514 



Appendix 1. Continued. 



Presence in Sampling Period 







1986- 


1987- 


1998- 


2002- 


2004- 






Species 


DFR 


1987 


1988 


2001 


2003 


2005 


CSA 


MS 


0. verruciger 


8/87 


+ 


- 


+ 


- 


+ 


+ 


+ 


Phrynohyas venulosa 


4/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Phyllomedusa coelestis* 


1/05 


- 


- 


- 


- 


+ 


+ 


- 


P. larsius 


8/87 


- 


+ 


- 


- 


+ 


+ 


- 


P. tomopterna 


4/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. vail Ian ti 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Scinax cruentommas 


5/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


S. fiinereus* 


5/90 


- 


- 


- 


+ 


+ 


+ 


+ 


S. garbei 


8/87 


- 


+ 


+ 


+ 


+ 


+ 


+ 


S. ruber 


5/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Anura: Leiuperidae 


















Edalorhina perezi 


4/86 


+ 


+ 


- 


- 


- 


+ 


+ 


Engystomops petersi 


9/87 


- 


+ 


- 


+ 


+ 


+ 


+ 


Anura: Leptodactylidae 


















Leptodactylus andreae 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


L. lineatus 


5/86 


+ 


- 


+ 


+ 


+ 


+ 


+ 


L. mystaceus 


5/86 


+ 


+ 


- 


+ 


+ 


+ 


+ 


L. pentadactylus 


10/87 


- 


+ 


+ 


- 


- 


+ 


- 


L. rhodomystax 


6/03 


- 


- 


- 


+ 


+ 


+ 


- 


E. wagneri 


5/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Anura: Microhylidae 


















Chiasmocleis bassleri 


5/86 


+ 


- 


+ 


+ 


+ 


+ 


+ 


Syncope antenori 


3/86 


+ 


+ 


+ 


- 


- 


+ 


+ 


Microhylidae incerta sedis* 


6/01 


- 


- 


+ 


- 


- 


- 


+ 


Anura: Ranidae 


















Lithobates palniipes 


6/86 


+ 


+ 


+ 


- 


- 


+ 


+ 


Anura: Strabomantidae 


















Isodactylus nigrovittatus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Oreobales quixensis 


5/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Pristimantis acuminatus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. altamazonieus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. aweolineatus* 


5/86 


+ 


- 


+ 


- 


+ 


+ 


+ 


P. conspicillatus 


6/86 


+ 


- 


+ 


+ 


+ 


+ 


+ 


P. croceoinguinis 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. diadematus 


6/87 


- 


+ 


+ 


+ 


+ 


+ 


+ 


P. lacrimosus 


3/86 


+ 


+ 


+ 


- 


- 


+ 


+ 


P. lanlhanites 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. librarius* 


3/86 


+ 


+ 


+ 


- 


- 


+ 


+ 


P. malkini 


6/87 


+ 


+ 


+ 


+ 


+* 


+ 


+ 


P. martiae 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. ockendeni* 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. paululus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


P. pseudoacuminatus 


4/86 


+ 


+ 


+ 


- 


- 


+ 


+ 


P. quaquaversus 


6/86 


+ 


+ 


+ 


- 


- 


+ 


+ 


P. variabilis 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Pristimantis sp. I* 


3/86 


+ 


+ 


+ 


- 


+ 


+ 


+ 


Pristimantis sp. II* 


6/99 


- 


- 


+ 


- 


- 


+ 


- 


Strabomantis sulcatus 


5/86 


+ 




- 


- 


- 


+ 


+ 



2008 AMAZON FOREST FRAGMENT HERPETOFAUNA 25 

\iti ndix I < 'ontinued. 









Presence 


in Sampling 


Pei iod 










1 986 


1987 


1998 


2002 


2004 




Species 


DFR 


1987 


1988 


2001 


2003 


2005 


CSA 


MS 


Caudata: Plethodontidae 


















Bolitoglossa equatoriana* 


! 86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


B. peruviana* 


J/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Gymnophiona: Caeciliidae 


















Oscaecilia bassleri 


6/03 








+ 




+ 


- 


Siphonops annulatus 


6/86 


+ 


+ 


+ 




+ 


+ 


+ 


Caeciliidae sp. incerta sedis* 


6/99 






+ 


- 


- 


+ 


- 


REPTILES 


















Chelonia: Chelidae 


















Mesoclemmys gibba* 


1/01 


- 




+ 


- 


+ 


+ 




Platemys platycephala 


7/04 


- 








+ 


+ 




Chelonia: Podocnemididae 


















Podocnemis unifilis* 


R 


- 


- 


- 


- 


- 


- 


- 


Chelonia: Testudinidae 


















Chelonoidis denticulata 


12/87 


- 


+ 




- 


- 


+ 


- 


Crocodylia: Crocodylidae 


















Caiman c. crocodilus* 


R 


- 


- 


- 


- 


- 


+ 


- 


Paleosuchus trigonal us 


7/98 


- 




+ 


+ 


+* 


+ 


- 


Amphisbaenia: Amphisbaenidae 


















Imphisbaena fuliginosa bassleri 


3/06 


- 


- 


- 


- 


+* 


+ 




Sauria: Gekkonidae 


















Gonatodes concinnatus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Pseudogonatodes guianensis 


3/86 


+ 


+ 




+ 


+ 


+ 


+ 


Thecadactylus solimoensis 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 




Sauria: Gymnopthalmidae 


















Alopoglossus angulatus 


8/06 










+* 






Buchia t. trisanale 


9/87 


- 


+ 






+ 


+ 


+ 


Cercosaura argulus 


6/86 


+ 


+ 


+ 




+ 


+ 


+ 


Leposoma parietale 


3/86 


+ 


+ 




+ 


+ 


+ 


+ 


Potamites strangulatus 


10/05 










+ 


+ 


+ 


Sauria: Iguanidae 


















Anolis /. fuscoauratus 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


A. nitens scypheus 


6/86 


+ 










+ 


+ 


A. ortonii 


7/86 


+ 


+ 


+ 




+ 


+ 


+ 


A. punctatus boulengeri 


8/87 




+ 


+ 


+ 


+ 


+ 


+ 


A. trachyderma 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


A. transversalis* 


5/89 










+ 


+ 




Enyalioides laticeps 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Plica umbra ochrocollaris 


3/86 


+ 


+ 


+ 




+ 


+ 


+ 


Polychrus liogaster* 


5/89 








+ 


+* 


+ 


+ 


Uracentron flaviceps 


1/88 




+ 








+ 




Sauria: Scincidae 


















Mabuya nigropunctata 


9/87 




+ 








+ 


+ 


Sauria: Teiidae 


















Kentropyx pelviceps 


3/S6 


+ 


+ 


+ 




+ 


+ 


+ 


Tupinambis teguixin 


4/86 


+ 


+ 


+ 




+ 


+ 




Serpentes: Boidae 


















Boa c. constrictor 


10/05 










+ 


+ 





26 



BREVIORA 



No. 514 



Appendix 1. Continued. 









Presence in Sampling 


Period 










1986 


1987- 


1998- 


2002- 


2004- 




Species 


DFR 


1987 


1988 


2001 


2003 


2005 


CSA 


MS 


Coral/us caninus* 


?/89 


- 


_ 


_ 


_ 


+* 


+ 




C. hortulanus 


6/87 


+ 


+ 


+ 


+ 


+ 


+ 


_ 


Epicrates c. cenchria 


8/87 


+ 


+ 


- 


+ 


+ 


+ 


+ 


Serpentes: Colubridae 


















Atractus elaps 


11/04 


- 


- 


- 


- 


+ 


+ 


_ 


A. gaigeae 


6/03 


- 


- 


- 


+ 


+ 


+ 


+ 


A. major 


3/86 


+ 


+ 


+ 


+ 


+* 


+ 


+ 


A. occipitoalbus* 


9/87 


- 


+ 


- 


- 


+ 


+ 


+ 


Atractus sp.* 


8/87 


+ 


+ 


+ 


- 


+ 


+ 


+ 


Chironius exoletus* 


7/03 


- 


- 


- 


+ 


+ 


+ 


_ 


C. f. fuscus 


8/87 


+ 


- 


+ 


- 


+* 


+ 


+ 


C. in. multiventris 


6/87 


+ 


+ 


- 


_ 


+ 


+ 


+ 


C. scurrulus 


11/04 


- 


- 


_ 


_ 


+ 


+ 


+ 


Clelia clelia 


8/87 


+ 


- 


+ 


_ 


+ 


+ 


+ 


Dendrophidion dendrophis 


9/87 


- 


+ 


+ 


+ 


+ 


+ 


+ 


Dipsas catesbyi 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


D. indica ecuadorensis 


3/86 


+ 


+ 


+ 


- 


+ 


+ 


+ 


D. pavonina* 


10/03 


- 


- 


- 


+ 


_ 


+ 


_ 


Drepanoides anomalus* 


7/87 


+ 


- 


_ 


_ 


+ 


+ 


+ 


Drymobins rhombifer* 


6/05 


- 


- 


- 


_ 


+ 


+ 


_ 


Drymoluber dichrous 


4/86 


+ 


- 


- 


_ 


+ 


+ 


+ 


Helicops angulatus 


6/87 


+ 


+ 


+ 


_ 


+ 


+ 


+ 


Imantodes cenchoa 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


I. leu tif eras 


3/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Leptodeira a. annulata 


3/86 


+ 


+ 


+ 


+ 


_ 


+ 


+ 


Leptophis ahaetulla nigromarginatus 


10/87 


- 


+ 


- 


- 


+ 


+ 


+ 


Liophis reginae semilineata 


7/87 


+ 


- 


- 


_ 


_ 


+ 


+ 


L. typhlus 


7/03 


- 


- 


+ 


_ 


+* 


+ 


+ 


Ninia hudsoni 


8/87 


+ 


+ 


_ 


_ 


_ 


+ 


+ 


Oxybelis fulgidus* 


7/03 


- 


_ 


_ 


+ 


+* 


+ 


+ 


Oxyrhopus formosus* 


9/05 


- 


- 


- 


_ 


+ 


+ 


+ 


0. melanogenys* 


6/03 


- 


- 


- 


+ 


+ 


+ 


+ 


0. petola digitalis* 


6/05 


- 


- 


- 


_ 


+ 


+ 


_ 


Pseudoboa coronata 


6/00 


- 


- 


+ 


_ 


+ 


+ 


+ 


Pseustes poecilonotus polylepis 


8/87 


- 


+ 


_ 


_ 


+* 


+ 


+ 


P. s. sulphureus* 


12/97 


- 


- 


+* 


_ 


_ 


+ 


_ 


Siphlophis cervinus* 


6/05 


- 


- 


- 


- 


+ 


+ 


_ 


S. compressus* 


7/03 


- 


- 


- 


+ 


+ 


+ 


_ 


Spilotes p. pullatus 


11/87 


- 


+ 


+ 


_ 


+ 


+ 


_ 


Taeniophallus brevirostris 


9/06 


- 


_ 


_ 


_ 


+* 


+ 


+ 


Taut ilia melanocephala 


11/05 


- 


- 


_ 


_ 


+ 


+ 


_ 


Umbrivaga pygmaea* 


7/03 


- 


_ 


_ 


+ 


+ 


+ 


+ 


Xenodon r. rabdocephalus 


4/06 


- 


- 


- 


_ 


+* 


+ 


+ 


X. sever its 


9/87 


- 


+ 


_ 


+ 


_ 


+ 


_ 


Xenoxybelis argenteus 


7/86 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Serpentes: Elapidae 


















Leptomicrurus narducii melanotics 


7/87 


+ 


_ 


_ 


+ 


+ 


+ 


+ 


Micrurus hemprichii ortoni 


6/99 


- 


- 


+ 


+ 


+ 


+ 





zoos 



\\1 \/(>\ 1 OKI SI I K \(,M1 M III RIM |()| \( \ \ 



27 



Ai-i'i ndix 1. Continued. 









Pi 


esence 


m Sampling 


Period 










I9S6 




1987 


1998 


2002- 


2004 




Species 


DFR 


1987 




1 988 


2001 


2003 


2005 


CSA 


MS 


\f lemniscatus helleri 


9/87 






+ 


+ 


+ 


+ 


+ 


+ 


M. ornatissimus 


9/87 






+ 






+ 


+ 


+ 


M. surinamensis 


6/00 








+ 


+ 


+ 


+ 




Serpentes: Viperidae 




















Bothriopsis bilineata smaragdina 


4/04 


- 










+ 


+ 




Bothrocophias hyoprora 


9/07 


- 




- 




- 


+* 




- 


Bothrops atrox 


3/86 


+ 




+ 


+ 


- 


+ 


+ 


+ 


Lachesis in. muta* 


R 














+ 


- 



*Asterisks by species names indicate that additional information is presented in the Results section of the text. 
Asterisks after + symbols for the 1998-2000 period denote species recorded earlier by other investigators (December 
1997 and April 1998); these are lumped with 1998-2001 species totals in Tables 1 and 2. Asterisks after + symbols for 
the 2004-05 period indicate that the record came instead from 2006 or 2007; these are not included in 2004-2005 or 
1998-2005 species totals in Tables 1 and 2. 

tDFR = Date of first record (month/year); an ""R" indicates that the species is included on the basis solely of 
reports by EBJS staff (dates unknown). CSA = Species presence/absence within the Central Study Area. MS = 
Species is represented (+) by at least one museum specimen obtained from sampling efforts reported here; see online 
Appendix 1. 



28 



BREVIORA 



No. 514 



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