Museum of Comparative Zoology Breviora 19

1 MCZ LIBRARY I 13 2C09 HARVARD UNIVERSITY RE V B I Nl Mseium oi v^onnparaitive /^oology US ISSN Cambridge, Mass 0006-9698 Number 514 December 2008 THE AMPHIBIANS AND REPTILES OF THE ESTACION BIOLOGICA JATUN SACHA IN THE LOWLAND RAINFOREST OF AMAZONIAN ECUADOR: A 20-YEAR RECORD Gregory O Vigle Abstract The amphibian and reptile fauna within a small Amazonian Ecuador was intensively surveyed over an initial area 2- in a lowland rainforest fragment reserve in 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 later, surrounding areas in A time-constrained transect sampling technique was employed to facilitate 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 this reserve in in the same small area to compare with data from 1986-88 in an attempt to evaluate the efficacj of 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: amphibian and 33 reptile species not recorded in 1986-88 were added from 1998 to 2007 Pairwise comparisons of species composition among taxonomic groups across amphibians and 0.70 for all all sampling periods using a similarity index revealed a > 50% similarity for both two primary periods (1986-88 vs 1998-2005) was 0.90 for periods: similarity of the reptiles The present study provides another Amazon basin, but is sampling suggest that single-site reference point for the distinct in offering long-term data this small reserve has since 1986 successfully previous long-term studies from the central remnants with Amazon sufficient habitat diversitj rainforest herpetofaunal Key words: on species mega-diverse herpetofauna of the upper from the 20-year span oi~ These results concur with small reserves and other forest persistence; results conserved this fauna Amazon basin in suggesting that relatively may substantially contribute to the continued survival of species-rich communities amidst widespread regional deforestation 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.; asu.edu or Librariusô librarius.org â The President and Fellows o\' Harvard College 2008 e-mail: vigle@imap2 BREVIORA Resumen Los anfibios y bosque lluvioso en mediante la reptiles que se encuentran en un area pequena dentro de una reserva fragmentada de Amazonia ecuatoriana han la No 514 del area y la subsiguiente oleada un periodo sido intensivamente investigados en recoleccion de muestras (1986-88) Esta actividad coincidio la de conversion y fragmentation del bosque en tecnica de muestreo de transectos de tiempo restringido para facilitar luego las areas el de dos afios inicial construction de un camino a traves circundantes Se empleo una monitoreo de encuentro visual post- fragmentation y reunir datos de largo plazo sobre la riqueza y composicion de las especies (en una escala cronologica 2005 se acumulo un esfuerzo de investigation y muestreo equivalente en la misma area desigual) Entre 1998 y reducida para compararla en la conservation de Basada en un total la information de 1986 a 1988, en un intento por evaluar la eficiencia de esta reserva fauna herpetologica documentada mas de 10 ahos atras la 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 comentarios en los registros mas significativos De en una unica ubicacion en 1986 a 2007 Se provee una anfibios y 82 especies de reptiles registrados de las la 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 especies de anfibios y 33 especies de reptiles Mediante un analisis revelaron un > 50% no registrados entre 1986 de composicion de pareja entre todos de similitud para Amazonia, 84 especies de completa de especies lista los el muestreo de 1998 a 2007; y 1988 fueron anadidos en 1998 a 2007 periodos de muestreo usando un indice de similitud 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 de la punto de referenda de sitio unico para la herpetofauna megadiversa de cuenca amazonica, pero es diferente ya que ofrece datos de largo plazo en la persistencia la zona alta de especies; los resultados de un lapso de muestreo de 20 ahos sugieren que esta pequena reserva desde 1986 conservado exitosamente esta fauna Estos resultados coinciden estudios previos de largo plaza de Amazonia en sugerir que las reservas la cuenca central de la relativamente pequenas y otros remanentes de bosque suficiente diversidad de habitat pueden contribuir sustancialmente a la continua supervivencia de herpetofaunisticas de los bosques lluviosos amazonicos en INTRODUCTION The rainforests of the Amazon basin harbor the highest concentration of biodion Earth, yet they are being destroyed by human activities at an alarming pace; one prominent threat is deforestation and the versity medio de la la alta diversidad de comunidades extendida deforestation regional Lamar, 2004; Dixon and Soini, 1986; Duellman, 1999; Frost, 2007; Peters and DonosoBarros, 1970; Peters and Orejas-Miranda, 1970) Reports on single-site studies have been reviewed by Duellman (1988, 2005) and Duellman and Thomas (1996) (also see Doan and fragmentation of previously continuous for- 2002; Lynch, 2005) Within the immediate area of the Ecuadorian Amazon, Arizabal, remnants of remaining published studies on the entire herpetofauna habitat (Bierregaard et al, 2001; Lovejoy et are available for several single or clustered al, 1984; Myers 1984) sites, est into smaller island Regionally, and on the basis of surveys from single localities, the lowland rainforests of the Amazon basin Colombia are home to western of Ecuador, Peru, and the highest concentra- amphibian and reptile species richness known from any area on Earth (Campbell and tion of including Almendariz (1987), Cisneros- Heredia (2003), Duellman (1978), Duellman and Mendelson (1995), Izquierdo and Lescure and Gasc (1986) From these studies, there et al (2000), has been an increasing awareness of the geographic ecological heterogeneity of the and Amazonian A MA/ON 2008 I OKI SI I RACiMFNT herpetofauna, and the neea\ to survey additional understand both better to localities II RIMTOl A I IN A ous studies from single upper Amazonian were conducted over relatively short sites and regional distribution patterns of known this mega-diverse but still poorb fauna (Doan and Ari/abal 2002; Duellman, spans of time 2005: Lynch, 2005) presenting data on the long-term composi- local The Estacion (EBJS) is tropical wet of edge forest community within western offered as another single-site reference point Basin in eastern fragment remnant; as such, forest upper Amazon, is it in the form of a 1985 traditional species checklist, and as a neces- before the construction of a road sary a ultimate fate of becoming eventually this an was recognized new reserve as habitat isolated 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 the time, at long-term efficacy of the reserve herpetofauna A in conserv- general prediction was that species richness initial would eventually decline or otherwise exhibit obvi- ous changes in the for preface 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 The 1986-88 and road-kill specimens forthcoming accounts on to and abundance species-level coinciding with the onset of road 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 in the tion of the herpetofauna] distribution derived from the same sampling efforts construction, ing distinct is reserve in the In 1986, island paper present at through the area the The forest a located Ecuador, established as shortly and within less), Sacha lowland remnant Amazon years or of Jatun Biologica small a (5 largely undisturbed expanses of continuous This study was undertaken several years widespread before interest of reports in amphibian declines (Blaustein, 1994; Stuart al et , 2004); despite concerns over various "enigmatic'" causal with declines from epidemic disease ranging to factors global climate change, the effects of habitat change are understudied sized for (Gardner both et and perhaps underemphaamphibians and reptiles al, especially 2007), in Neotropical lowlands, where long-term itoring the mon- data remain very rare (Duellman, 2005:49; Whitfield et EBJS provides an a! 2007:8355) established lowland The re- serve with a well-known history as a forest fragment, and monitoring the an is an appropriate effects exceptionally site for of habitat change on species-rich Neotropical herpetofauna METHODS 1998-2005 periods are treated as equivalent The following summary of methodology "'primary'" units for comparative purposes, divided into three sections, progressing from 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 Methods) in With the exception of Cusco Amazonico Peru (Duellman 1995 2005) most previ- ( ) a detailed characteristics, EBJS; to (2) description history, is of the physical and climate of the an account of the long-term and major goals division of sampling effort 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 Description of the study I) history, The EBJS Napo site: forest, and climate Ecuador in Province at elevations of 400 to 450 m (01°04'S, is located in eastern 77°37'W at main station complex; the reserve is owned and by Fundacion Jatun Sacha (FJS), an Ecuadorian conservation foundation The EBJS was the first of several reserves established by FJS and originally Figs 1-4): see administered (1985-88) consisted of 230 of preserved forest owned by After an ers three independent landown- initial 150-ha expansion funded by the Grateful Dead music group Rex their (via Foundation), 1989 in subsequent acquisitions of additional land have greatly increased the size of the reserve, and current total land holdings are 2,334 Figure 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 Current land holdings of the EBJS reserve (gray shaded areas within dark borders); location in eastern top Ecuador left (Figs tracts 4) Study Area (CSA; see pasture, whereas tract 70% primary forest, with the 30% composed of various succes- fields remaining Also see stages station at at 77°37'W The area located and B comprises the Central Fig 2); tract B is A is cleared cattle a mixture of agricultural and various secondary forest successional stages text and Figures Approximately to scale approximately sional EBJS main A Forest within reserve boundaries consists of 1, map complex indicated by arrow on inset approximately 01°04'S, between land Arajuno River to the south is Star indicates of secondary forest growth, including approximately 95 of open-for- (dating from 2000 and 2001) are presented in mation riparian habitats Figure and online Figure (Fig 3) Although the EBJS preserves a substantial tract upper tial of lowland rainforest, since 1985 the Napo region has undergone substan- changes resulting from deforestation and forest fragmentation because of increasing colonization and development Most adja- 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 area for many had already affected the years, especially along the March cent bordering areas have been logged or banks of both major affected by agricultural activities, although the construction of a road (from the west) no reached the north-central area of the precise survey data are available Satellite images showing general overviews of forest cover at the EBJS and in adjacent areas (Figs 1-4), rivers In 1986, EBJS and ongoing construction now extends approximately 30 km further east \/()N \\l 2008 I OKI SI I R \(,\|| \|||1 Ki-i reserve effect |()| ,\l \ \ relatively small, the larger regional is increased development o\' the pre- is dominant threat, as the KB.IS has increasingly become an "island" reserve surrounded fragmented b\ forest The present study reports primarily on sampling conducted in a small (approximate- (CSA) border- 100 ha) central study area ly ing Napo the River (Figs 4); CSA the major habitats represented in the area (primary forest, and various riverfloodplain forests, swamps, and secondary includes forest; all Fig 3), in contrast to land acquisi- 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 tions since 1990, portions limited EBJS within the CSA The of various the most also the area is habitats as a whole directly affected by the road (and forest clearing 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 and text) All sampling efforts as summarized in the text and were conducted on trail routes (irregular online Table lines in central area) extending north and south from the EBJS main station complex (star) and north from the Figure Librarius Study Center complex (solid circle), as well as along the margins of the road approximately 300 Figure m to the east The gray-shaded area station Approximately to to a distance of and west of the main shown in detail in is fragmentation in adjacent longest period of time of land, in total size to the entirely Figs 1, 2, Two both 4); the east and west partially cleared (see before 1986, although the largest (to the west) 10 the deforested tracts both of these adjacent areas had already been undergone and area directly and this years had of secondary growth before being cleared again scale for area approximately equal in CSA, border to areas) in 1990 Thus, although the total area of preserved The discovery of oil reserves in the region forest is today far larger by the acquisition of has accelerated the process of colonization in additional land, in keeping with the long- recent years, constituting an increased threat term goals of the study begun to the EBJS r6ad traffic, areas crude In oil reserve in the form of additional and deforestation in nearby November 2003 a high-pressure was installed along the side pipeline of the road (buried to a depth of m), and in was widened by m and paved with asphalt Although the immediate area affected by the road itself within the early 2006 the road below), subsequent in 1986 (see 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 however (exhibiting proeasterly areas, nounced wet and dry seasons), climate at BREVIORA Map No 514 and diagrammatic cross section (top) showing relief and 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 Figure (bottom, approximately to habitat distribution within the the EBJS is CSA far less seasonal, scale), (also see Figs 1, and can rainfall 4) and 210 mm, respectively) was lower than be highly irregular within any given year both November and December (335 The each month) yearly distribution of rainfall is relevant mm in to the distribution of sampling effort for the purpose of comparing faunal records between sampling periods (discussed below) EBJS since 1986 although data are incomplete for some Rainfall has been recorded at the years; for one 10-year period with complete data for each Table 3,71 This 1 ), month (1994-2003; total rainfall see online per year averaged II) Temporal distribution of sampling and major goals A to effort Coarse-Scale Sequence from "Before" "After." and Collections records of observations of the amphibian and reptile fauna of EBJS the include two major mm (ranging from 3,051 to 4,593 (primary) sampling periods, each including record that approximately equal also indicates mm) 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 first 1988 from 15 sampling March 1986 (536.0 hours second spanning total July to effort, 15 effort) the February and the 1998 through De- cember 2005 (581.0 hours) Each primary sampling period is arbitrarily subdivided into secondary sampling periods (discussed below) The two primary sampling periods AM A/ON 2008 Figure within the Landsat EBJS satellite reserve and OR1 SI I K \ci\ll \I III RIM I ( \ \l )| \ image (30 m/pixel resolution), captured September 2001 showing closely adjacent areas Refer to Figures 1-3 for location of relative to reserve borders; the and west of the CSA Also I forest cover major landscape features two outlined rectangles approximately indicate deforested areas A and B to the east showing forest cover over a larger area At right, adult Ranitomeya see online Figure duellmani (Dendrobatidae) shown 1.5X natural size reported here can be interpreted as "pre- to assess the overall stability (1986-88) and "postfragmentation" (1998-2005) data on the EBJS of the community over time (on a chrono- fauna, although as previously noted, forest process; hypothetically, each sampling pe- conversion processes in the area had been riod (and ongoing even before 1985 a fragmentation" An program of removal sampling from 1986 to 1988 was undertaken as a initial baseline inventory of the fauna to facilitate subsequent monitoring of the same area in later years (1998-2005) via VES efforts, with logically coarse scale) This periods combined) can provide "before" reference point to compare with subsequent equivalent sampling efforts "after,"'' as one measure of the efficacy (or lack thereof) of the EBJS reserve in preserving the herpetofauna at this increasingly repeated in later years percent second goal was to accumulate equivalent 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 an ongoing is all major goal, A similarity predictive 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 three and isolated with (the third larger scale site potential relevance for conservation in the region as a whole) Sorenson's quotient of similarity (SQ = 2J[A + periods E\) is (see used to compare the among similarity Results all and Table sampling This 2) simple formula estimates similarity inde- pendent o'i the individual each species, where A species in sample 1, B is is abundance of the the number of number in BREVIORA No 514 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 Table recorded later from 2006 or 2007 far right column presents the total number of (including "r" species) from 1986 through 2007, as listed in appendix individual specimen totals indicate the number of individuals recorded in each period; numbers in parentheses indicate the individuals per effort-hour for each period additional species species (+) from all sourc1 s AMAZON OKI 2008 1~ VHl AND \M> t RAC.MI NI IOI 111 Rl'l UNA PaIRWISI COMPARISONS Ol \ll PRIMARY AND SECONDARE SAMPLING PERIODS FROM Mil EBJS FOR AMPHIBIANS \< NUMBERS Ol SP1 CIES FROM EACH i\ RIOD \KI IMiliAl I) IN m m COMMON CELL, MBERS OF SPECIES SHARED BY PERIODS ARI INDICATED Al mi UPPER RIGHT SIMILARITY QI OTIENT (SQ) VALUES REPTILES llll SI M Till COTA1 I'l \RI Sampling Period INDICAT1 I) IN ITALICS I \l I OW] K l l l l >i l )i I I BREVIORA II) to 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 museum by specimens Collecting is now EBJS, and records from the second primary period (1998 2005) are based almost entirely on VES efforts, includrestricted at the ing a total of 3,773 individuals few exceptions (see Appendix the author personally identified (from all With only a and Results), all specimens sampling periods) via direct obser- vation or examination of photographs (see below) Of 1,117 effort-hours of sam- the total 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 EBJS Park Guards), and their SCs (and respective sampling effort contributions, are noted in No 514 From August 2003-December large portion was conducted by Park Guards employed by the EBJS In a project funded by the Declining Amphibian Populations Task Force (DAPTF), park guards were trained on-site in search techniques and the use of digital cameras Park Guards then (231.5 hours) continued periodic sampling in the author's and absence, images (CD) for later Beginning in March 2002, the physical identification Although most of the 6,722 individuals from all sampling periods were recorded during regularly measured sampling efforts, a total of 376 records represent (5.6%) randomly encountered outside the 1,117 hours summarized in online Table Appendix also includes all species known to the author from the EBJS since 1986 from all sources, including additional records from 2006 and 2007, and a few important records from other investigators (see Appendix 1, Table 1, and Results) individuals that were The majority (75%) of specimens collected during the 1986-88 removal sampling effort are deposited in the herpetological collec- Museum of Comparative Zool- (primarily in Ecuador) and from 1998 to 2001, all specimens encountered by SCs were captured and then identified (in 1986 88, most were preserved; in 1998-2001, all were released within 48 hours) and review reviewed the results of each night's sampling 1986^88 period, individuals all (1,065 identified individuals) tions of the efforts immediately: for the of encountered were saved on compact disc Acknowledgments For all sampling before August 2003, the author was present on-site with all SCs and the of 2005, a of the total sampling effort ogy at Harvard University (MCZ); the remainder are housed in other collections additional museum The following collections is list of presented in approximate decreasing order of the total number of included specimens at each museum (with acronyms): Museo Ecuatoriano de Ciencias Naturales, Quito (MECN); Museo de Zoologia, Pontifica Universidad capture of specimens by SCs was discontin- Catolica del Ecuador, Quito and digital cameras were used to photograph every amphibian and reptile ican (QCAZ); Amer- sampling methods remained the same, and Museum of Natural History (AMNH); Florida Museum of Natural History (UF); United States National Museum (USNM); University of Kansas Museum of Natural were determined solely from History (KU) All specimens collected from ued specimen encountered; identifications photos all other aspects of 1998 to 2007 are deposited at the QCAZ (all BREVIORA IS an adult ently the reports are in a fluvial Serpentes: Colubridae Napo At raet us species the floodplain of the Rio swamp on (within crocodilus C No 514 CSA) in 1990 No additional known from the EBJS; however, Among material collected in 1986-88 (as VES some might survive in more remote areas of the Rio Arajuno on the south side of the well as additional reserve diagnostic characters of lepidosis conform to Atractus major Sauria: Iguanidae recorded by Laurie initially communication) (via and transversalis in May photos) of the Vitt (personal 1989; identification was is The single P shown in online Fig liogaster from 2006 was a (QCAZ: GV2000-0349) collected by EBJS Park Guard Milton G Orozco (M.G.O); this and previous EBJS records apparently represent the Ecuador, and a for this a possibly distinct cryptic and evaluation of additional specimens of Atractus individuals All sp have a 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 kill extension as species pending the collection W Lamar (W.W.L.) 8) road referred to tentatively dorsal pattern of alternating short and long 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 series consis- from all other EBJS specimens A major and are here treated later liogaster P determined by William were P liogaster J 173868-871; online form a tently varying Polychrus liogaster (Boulenger) Both A (MCZ R Fig 9); however, these Anolis transversalis (Dumeril) and records since 1998) are specimens of a small Atractus, which in very reports from first range substantial species, range approximately 1,000 extending km to the from the nearest reported records in bands extend unbroken dorsum In contrast, all A major exhibit a dull brown dorsal ground color with short dark brown bands and and bands, all laterally across the markings (with short pale tan broken middorsally with few (if any) bands laterally continuous across the dorsum Morphologically, there is a irregular outlines) that are striking difference in the size of the eye; the eyes of Atractus sp are distinctly smaller head size than are those of A major These differences are obvious if specimens are compared directly relative to the NW Additional Notes on Colubridae western Brazil (see Avila-Pires, 1995:131, 134) Several colubrid species and period records are included in Appendix on the basis of Serpentes: Boidae reports or identifications by other investigators, Corallus caninus (Linneaus) but otherwise require no further commentary The first record was an adult photographed by M.R in 1989 (date unknown; not collected); two additional adults were recorded in 2006 (all were found within the CSA) pavonina, Drepanoides anomalus, and Oxy- Records from rhopus formosus 1998 were Guards Willan G to 2005 of Dipsas obtained Poveda and by Park M.G.O.; identifications (from photos) for all of these were kindly determined by W.W.L ; AMAZON 2008 Pseustes basis o\~ a Jungfer in identified I OKI ST I RAOMI NT sulphureus is included on the s photograph taken by Karl-Heinz December 1997 and subsequently by W.W.L III RIM IOI AUNA being preserved and to assess the persistence the community under study; the success of any conservation effort is best measured by documenting community persistence (and o\' records and identifications for Drymobius rhombifer, Oxyrhopus petola digitalis, 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 expertise; a shortage of Atractus resources often renders such efforts difficult Initial Chironius occipitoalbus, exoletus, Oxyrhopus melanogenys, Siphlophis compresand Umbrivaga pygmaea (from June 2005) and D.F.D reported Oxybelis fulgidus from June 2007 sus, over fluctuations) long periods time of 1994; Teder et al, 2007) (Blaustein et al, Unfortunately, poorly known, yet speciesrich, rainforest ecosystems require ongoing sampling effort combined with taxonomic and limited in human and scope (Bierregaard Crump, 2003) The techniques employed utilized minimal resources, in material et al., 1997; study this including one principal investigator (the author), persons who were Serpentes: Viperidae Lachesis muta (Linnaeus) Both G.A.S and Park Guard Gabriel 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 Tapuy (G.T.) have observed individual collected in the area, although the exact locality is long-term data on the persistence of species richness and composition Although tropical herpetofaunas have recently been undertaken (Doan, 2003; Funk many respects, et al., 2003), employed must be both consistent and sustainable over time if an is accurate assessment of the DISCUSSION be attained (Teder In studies evaluating sampling methods for monitoring whatever method unknown SUMMARY AND 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 EBJS forms the a microcosm relevant to a number of important Amazon Assuming this community is to methods employed in et al., 2007) that the study gathered accurate data on long- rainforest con- term species persistence as a measure of servation, including questions concerning the conservation efficacy, results reported here issues pertaining to appropriate identify minimum reserve size required to and conserve viable communi- biotic ties, the long-term effects of continuing development and fragmentation on surviving habitat islands, and the need to inventory and monitor all component organisms these questions Bierregaard et ( al , 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, to address monitoring over the subsequent 10 years (1998-2007) revealed that 92% of amphibian 997, 200 and reptile species initially recorded in 1986 Laurance et al 1997; Lewin 1984; Simberloff and Abele 1982; Turner, 1996; Turner and some point later during 1998 2007 The number of species Corlett, 1996) added in 1998 2007 for both amphibians (11) and reptiles (33) far exceeds the number The need monitor is to first essential inventory to and later determine what is 88 were also present at of 1986-88 species lacking subsequent rec- BREWORA 20 No 514 ords from 1998-2007 (thus there were very almost entirely unstudied; additional species few possible "ghosts"; see Dayton may The 1998) for partially attributed to a vehicular increased effort is combination of (1) in and traffic the collecting in particular period increased road-kill resulting specimens, (2) the use of digital cameras to document randomly encountered individuals, and (3) contributions from other investigators New species records for both groups now continue to accumulate; snakes although dominate, a pre- amphibian large (Rhaebo glaberrimus) was added within the CSA as recently as June 2007 Pairwise among better comparisons for amphibians sampling periods (far equivalent sampled than represented) all larity in species composition (from 0.77 to whereas similarity for 0.90), but exceeds 0.50 for is records reptiles, in total reveal a high percent simi- reptiles lower is equivalent pairs, and all Crude 0.70 for the two primary periods measures of both species (SPH) and individuals per sampling effort-hour for both major among sampling periods, but none suggest any pronounced declines Interpretation of group-level abundance data is problematic; given the large sample sizes represented, however, lumped data on even coarse scale provides (and baselines ranges of variation) that could be used to predict patterns and monitor long-term expected of abundance at the EBJS over future equivalent sampling periods Perhaps the most remarkable among the Neotropical and Ecuador nity, but the concentration of 161 recorded from the CSA is not commuspecies within a sampled area of only 100 directly bordering two large tracts 80% of deforested land (Figs 1^4) More than of total sampling effort was conducted along precisely the same trails site is already For amphibians, only Tipu- Santa tini EBJS highest ever recorded for any single (both Cecilia NE to the located in 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 as a result of colonization leaving the and EBJS second only many years oil extraction, to Leticia currently extant at EBJS the sites Reptile species richness greater is than reported for (Cisneros-Heredia, Tiputini and among Tiputini in amphibian species richness 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 amphibians in lowland Amazon from the more species rich upper Amazon) Tocher et al (1997) examined pre- and post-isolation species richness among small (1-100 ha) fragments of prima- anuran rainforest (none ry forest over a 10-year period in north- central Brazil (40 species) Contrary to their few species were lost from fragments following isolation, and fragments initial predictions, result merely the robust persistence of the records to date, the herpeto- all faunal species richness of the groups vary this be expected from these areas Including increase in reptile species records 2004-05 the al, et within the CSA of all showed increases in species compared with equal-sized plots sizes richness within larger contiguous primary forest areas (partially attributed to an influx of distur- bance-tolerant species into fragments) Gas- et al (1999) also studied the persistence of anurans over the entire 20-year span of sampling Most disturbed of the interior regions of the reserve remain Amazon in fragments and surrounding matrix habitat in the central over spans of up to 19 years and AMAZON :oos found few species were that species richness increased, remained These fragments that the less overall lost, forest increased or in concluded investigators Amazonian anuran community was affected by fragmentation vertebrates and can stable FRAGMENT HERPETOFA1 and matrix-toler- many primary ant species (including species) FORI ST better many anuran that and utilize than other migrate species through is of millions of years of biological evolution the in Amazonian rainforest This small reserve might provide a model for future conservation efforts in the but to region, repeat an admonition expressed in virtually address the studies on fragmented ecosystems, time and continued monitoring will be necessary to many questions that yet remain an earlier study from the same area, Zimmerman and ecological factors in an attempt to test which approach was minimum area predictor better a required of the preserve to the anuran fauna (39 species) They concluded that on the basis solely of species-area data, only about 500 would be required to preserve this fauna, but habitat heterogeneity and, especially, the distribution of breeding were minimum far better is dedicated in equal measure two persons, without whom the study would never have been initiated or completed 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 expansion of the EBJS reserve to the present day the can almost entirely be credited to the vision and dedication of EBJS Founder and Director G Alejandro Suarez, whose assistance of breeding sufficient This paper to community predictors area needed to ensure Without survival ACKNOWLEDGMENTS Bierregaard (1986) com- pared species-area relation data derived from island biogeography theory with data on sites, even larger areas would not be adequate; with with abundant breeding indispensable for neity, However, merely a brief instant within the far larger context all sites 21 although 20 years might represent a substantial time span from a human perspective, this monitoring would be essential to discern more long-term patterns In \ impacted landscape increasingly disturbed habitat, but they cautioned that further N sites and heteroge- an even smaller area might suffice Results from this study provide another long-term reference point for an exceptionally rich ing that Amazonian herpetofauna, suggestsome relatively small reserves could all on-site aspects of this study has been more than 20 years Although space does not permit the acknowledgement 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 ,, (with contributed effort- be a viable conservation strategy for these Collectors, taxa in response to forest fragmentation; the hours high concentration and persistence of species 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 within the small but ecologically heteroge- neous (and especially studies extensively noteworthy riparian) in light CSA is of previous on Amazonian anurans Contrary to some in 1986, the initial predictions EBJS herpetofauna has not suffered a catastrophic decline richness, despite more than 20 made evidently in species years amidst an total in parentheses): Brock Dolman School for Field Studies students with 1986-88 field work and During 1998 2007 Guards contributed field all (75.0), assisted logistics work, EBJS Park essential sampling effort BREVIORA 22 knowledge, and camaraderie; these included Milton Orozco (99.5 effort-hours), Willan Poveda (97.0), Gabriel Tapuy persons noted Elibar Jimenez (19.5), and (15.5) in In addition to all accounts of species, various No 514 with Assistance museum identifications, and loans and depositions of specimens use, since 1986 have been generously shared by the J Cadle, L Coloma, G Hanken, R Heyer, F Irish, J Rosado, V Wallach, and the late E Alberch, P late Flores, J additional contributions in the field from Losos, 1998 to 2007 are credited to N Batcha, Williams Assistance with obtaining collecting Goberdhan-Vigle, P Grefa, C James, S S Konningsor, R Sawby, W Schaedla, M Shindell, and students in Arizona State J in Matheus, F Sarmiento, Figueroa, S University tropical biology courses held at cias Naturales, terio useful advice and discussion through- funded by the thanks D DeNardo, M Douglas, M Fou- Museum W Lamar, D Oakey, D Pearson, B 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 Sullivan, Subsequent to completion of the present additional six species records and the Minis- tially and the E Williams late E of Comparative Zoology, Harvard 1998-2007 University; field work was par- supported by grants from the Arizona Graduate Research Develand the Declining Amphibian Task Force (DAPTF; with State University opment Office Populations special thanks to NOTES ADDED manuscript, INEFAN and de Agricultura y Ganaderia del Ecuador The 1986 88 phase of this study was out the second phase of sampling, the author quette, J The Museo Ecuatoriano de Cien- R Ulloa, the EBJS For Ecuador was provid- and research permits ed by IN Tim Halliday) PROOF m (600 forest within the EBJS south of the CSA) station, 2008, by Milton G in July from the EBJS have been reported; with the Orozco Although determination was made by exception of one record (dating from 1988), the author (and William were confirmed by the author from photographs (none collected) These are presented below in the same order the photograph, this record identifications all Appendix as AMPHIBIA: cagei One Dendrobatidae: Hyloxalus bo- adult was reported from an area km NW of the EBJS station, in July of 2008, by Ross J Maynard and Nathan A Shepard REPTILIA: Aniliidae: Anilius scytale A was recorded on the south side of the reserve (near the Rio Arajuno), km SSE of the EBJS station, in July 2008 by Ross J Maynard and Nathan A Shepard single juvenile Colubridae: Atractus torquatus This species tentatively included based on a photograph of one individual taken at night in primary is would apparently represent a substantial westward range extension and the first additional material W Lamar) based on is report from Ecuador; needed for confirmation Colubridae: Liophis cobella Reported by Bill Montgomery based on in July a photograph taken 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 was recorded by day near km SW One adult a stream in primary of the EBJS station forest, 3.5 March 2008, by Milton G Orozco in AMAZON 2008 I OKI ST I R \(,MI NI IIRPITOI Al INA 23 and distribution of records, 1986 2007.* Amphibians and ebjs since march 1986 over mm sampling periods, \is including all species reported from al sources through 2007 (also see text and Tables 1, 2) Presence or absi nce within v h swum lng period (and other columns) is indicated by + or - symbols.! Appendix reptiles I List oi species recorded from till i 24 BREVIORA Appendix Continued Presence in Sampling Period No 514 2008 AMAZON FOREST FRAGMENT HERPETOFAUNA \iti ndix I < 'ontinued 25 26 BREVIORA Appendix Continued No 514 zoos \\1 \/(>\ OKI SI Ai-i'i I K \(,M1 ndix M III Continued RIM |()| \( \ \ 27 BREVIORA 28 LITERATURE CITED Almendariz, A 1987 Contribution herpetofauna centroriental Ecuatoriana Politecnica: Biologia Revista de Information Tecnicola Natio- Cientifica, VII: 77-133 Escuela Politecnica nal Quito Ecuador Avila-Pires, C T Amazonia of Lizards 1995 S Brazilian Squamata) Zoologische Ver- handelingen, 299: 1-706 F W J SlTES, Jr., J Lynam, R K Didham M Andersen, C Gascon, M D Tocher A P Smith, V M Viana, T E Lovejov K E Sieving, E A Kramer, C Restrepo, and C Moritz 1997 Key priorities for the study of fragmented tropical ecosystems, pp 515-525 Laurance, W In: & F and fragmented Press, Chicago and London, 616 pp C Gascon, T E Lovejoy, and R Mesquita 2001 Lessons from Amazonia: The Ecology and Conservation of a Fragmented Forest New Haven u Range new extensions, provincial records from Ecuador, and natural history Avail- 042-06 Cochran D M., and C Goin J Frogs of 1970 Colombia United States National Museum Bulle- 288: 1-655 tin Coloma A 1995 Ecuadorian frogs of the genus L Colostethus (Anura: Dendrobatidae) The Univerof Museum Kansas Natural of History Miscellaneous Publication, 87: 1-75 Coloma, 2005-2007 Anfibios de Ecuador L A (Ed.) 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Amazon lowlands of of the Museum of Na- of The University of Kansas, 31: 1-22 2005 Discovery of the richest frog fauna in the world — an exploration of the forests to the north of Leticia Revista... subsequent 10 years (199 8-2007) revealed that 92% of amphibian 997, 200 and reptile species initially recorded in 198 6 Laurance et al 199 7; Lewin 198 4; Simberloff and Abele 198 2; Turner, 199 6; Turner... methods remained the same, and Museum of Natural History (AMNH); Florida Museum of Natural History (UF); United States National Museum (USNM); University of Kansas Museum of Natural were determined