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Bothrops sonene sp. nov. (MUSA 4350) from Pampas del Heath, Bahuaja-Sonene National Park, department of Madre de Dios, Peru.
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We describe a new species of pitviper of the genus Bothrops from the Peruvian Pampas del Heath, in the Bahuaja-Sonene National Park. Pampas del Heath is an area of seasonally flooded savannas and a northwestern extension of the Gran Chaco Boliviano-Paraguayo. The new species is easily distinguished from its congeners by the exclusive combination of...
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... province of Tambopata, department of Madre de Dios, Peru. An adult male, MUBI 14679, collected on August 14, 2015 by J.A. Ochoa, G. Sebastián & K. Huamani (12°57'47.47" S, 68°54'57.01" W; 210 m asl) (Figs. 5B, 5F, 6C). An immature male, MUSA 4350, collected on November 13, 2015 by R. Santa Cruz (12°57'2.47" S, 68°54'51.05" W; 209 m asl) (Figs. 4, 5C, 6A, D). An immature male, MUSM 33453, collected on between June 02-25, 1996 by J. Cadle (12°59'20.10" S, 68°55'48.45" W; 208 m asl) (Fig. ...
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Citations
... Species sampled for each genus were selected based on availability of specimens for morphological study. We followed Campbell and Lamar (2004), Carrasco et al. (2009Carrasco et al. ( , 2010Carrasco et al. ( , 2019 and The Reptile Database (Uetz et al., 2023) for taxonomic identification of voucher specimens. For a few species, we obtained morphological data from the literature (McCranie, 1988;Crother et al., 1992;Fuji and Peng, 2001;Marques et al., 2002;Campbell and Lamar, 2004;Guo and Zhao, 2006;Maduwage et al., 2009;Jadin et al., 2010Jadin et al., , 2011Barbo et al., 2016). ...
... Names for clades of species within Bothrops and Crotalus are those used in previous studies (Blair and S anchez-Ram ırez, 2016;Carbajal-M arquez et al., 2020;Carrasco et al., 2012;Fenwick et al., 2009;Hamdan et al., 2020;Murphy et al., 2002;Reyes-Velasco et al., 2013), with the exception of the Bothrops lanceolatus clade, which we considered is included within the Bothrops atrox clade. The definition of the Bothrops neuwiedi clade follows Carrasco et al. (2019). ...
... Atheris chlorechis ( Definition of characters follows Marx and Rabb (1965), Burger (1971), Marx et al. (1988), Pesantes (1989), Gutberlet Jr and Harvey (2002), Fenwick et al. (2009) and Carrasco et al. (2012Carrasco et al. ( , 2019. Some of the characters are newly proposed here. ...
Crotalines (pitvipers) in the Americas are distributed from southern Canada to southern Argentina, and are represented by 13 genera and 163 species that constitute a monophyletic group. Their phylogenetic relationships have been assessed mostly based on DNA sequences, while morphological data have scarcely been used for phylogenetic inquiry. We present a total‐evidence phylogeny of New World pitvipers, the most taxon/character comprehensive phylogeny to date. Our analysis includes all genera, morphological data from external morphology, cranial osteology and hemipenial morphology, and DNA sequences from mitochondrial and nuclear genes. We performed analyses with parsimony as an optimality criterion, using different schemes for character weighting. We evaluated the contribution of the different sources of characters to the phylogeny through analyses of reduced datasets and calculation of weighted homoplasy and retention indexes. We performed a morphological character analysis to identify synapomorphies for the main clades. In terms of biogeography, our results support a single colonization event of the Americas by pitvipers, and a cladogenetic event into a Neotropical clade and a North American/Neotropical clade. The results also shed light on the previously unstable position of some taxa, although they could not sufficiently resolve the position of Bothrops lojanus, which may lead to the paraphyly of either Bothrops or Bothrocophias. The morphological character analyses demonstrated that an important phylogenetic signal is contained in characters related to head scalation, the jaws and the dorsum of the skull, and allowed us to detect morphological convergences in external morphology associated with arboreality.
... We generated a dataset of 52 quantitative and qualitative characters usually evaluated in taxonomic studies on Bothrops (e.g., Barbo et al., 2012Barbo et al., , 2016Carrasco et al., 2019). These characters were organized in 44 discrete, 5 continuous, and 3 categorical variables (full list of analyzed variables are presented in Table S3). ...
Brazilian continental islands represent a natural laboratory to study speciation driven by recent phenotypic and genotypic divergence. The Bothrops jararaca species group is distributed in the Brazilian Atlantic Forest and on most of the Brazilian continental islands. The group is currently composed of the mainland common lancehead (B. jararaca) and four insular species (B. alcatraz, B. insularis, B. otavioi, and B. sazimai). Here, we evaluate mitochondrial DNA and morphological diversity of the B. jararaca species group and aim to provide additional evidence to understand insularization processes on the Brazilian coast. Our results, interpreted together with a comprehensive review of geomorphological data, provide a new conceptual framework for understanding the colonization process of the Brazilian continental islands. This framework suggests a history of multiple rounds of periodic isolation and reconnection between insular populations and their mainland relatives throughout the last 420,000 years. Furthermore, although some insular populations may have speciated prior to the last glacial maximum, other species likely diverged within the last 11,000 years. Additionally, the repeated evolution of size and dietary shift in the B. jararaca species group suggests a remarkable case of convergent adaptation. Our study provides evidence that the Bothrops from Ilha da Moela (Brazilian state of São Paulo) represents an undescribed species, presenting a distinct phenotype, and an exclusive history of isolation and adaptation. We describe this unique lancehead as a new species and we suggest it should be listed as critically endangered based on its endemicity to a small island that is severely impacted by constant and longstanding human presence.
http://zoobank.org/urn:lsid:zoobank.org:pub:B1F65287-CEC4-4F9C-90D7-42314A497590
... Bothrops Wagler, 1824 is one of the most diverse genera of pitvipers (Viperidae: Crotalinae) (Campbell and Lamar 2004;Carrasco et al. 2012), currently composed of 46 recognized species (Uetz et al. 2021) that are distributed in Central America and primarily in South America (Campbell and Lamar 2004;Nogueira et al. 2019). From the earliest phylogenetic studies using molecular evidence (Werman 1992;Salomão et al. 1997Salomão et al. , 1999Parkinson 1999;W€ uster et al. 2002) to the recent analyses based on large taxon sampling (Alencar et al. 2016;Zaher et al. 2019;Hamdan et al. 2020) and integrative approaches (Fenwick et al. 2009;Carrasco et al. 2012Carrasco et al. , 2019, the genus has been regularly divided in the following six main species groups: Bothrops alternatus A.M.C. Duméril, Bibron and A.H.A. Duméril, 1854;Bothrops atrox (Linnaeus, 1758); Bothrops jararaca (Wied-Neuwied, 1824); Bothrops jararacussu Lacerda, 1884; Bothrops neuwiedi Wagler, 1824; and Bothrops taeniatus Wagler, 1824. Some few species with uncertain phylogenetic position (e.g., Bothrops pictus (Tschudi, 1845); Bothrops roedingeri Mertens, 1942;Bothrops barnetti Parker, 1938;Bothrops lojanus Parker, 1930;Bothrops andianus Amaral, 1923;and Bothrops venezuelensis Sandner-Montilla, 1952) have been considered incertae sedis (Fenwick et al. 2009;Carrasco et al. 2012), and other species demonstrating more basal divergences from recent phylogenies have been classified as belonging to the genus Bothrocophias Campbell, 2001 (e.g., Bothrocophias microphthalmus (Cope, 1875) and Bothrocophias hyoprora (Amaral, 1935)). ...
... Among these six assemblages of closely related species, the groups atrox, jararaca, jararacussu, and taeniatus are present within a distribution strictly connected to forested areas, whereas the distribution of the alternatus and neuwiedi groups are associated with open formations (e.g., Cerrado and Caatinga; Carrasco et al. 2019;Nogueira et al. 2019). The forest-dwelling groups occur throughout two of the principal forested areas of South America east of the Andes, the Brazilian Atlantic Forest (BAF) and the Amazon Rainforest (Ab'Sáber 1977). ...
... To estimate the phylogenetic position of the population from Pico do Jabre, we built a concatenated molecular matrix consisting of the two mitochondrial gene fragments cytochrome b (cytb) and NADH dehydrogenase subunit 4 (nd4). We included in our molecular dataset representatives of all groups of Bothrops, and the majority of our matrix was composed of sequences downloaded from GenBank (Supplementary Table S3), 1 derived from studies focused on the B. jararaca (Barbo et al. 2021a), B. neuwiedi (Machado et al. 2014;Carrasco et al. 2019), and B. jararacussu groups (Dal Vechio et al. 2021). We generated cytb and nd4 sequences for the individual IBSP 92904 from Pico do Jabre. ...
Here we describe a new species of Lancehead (Bothrops jabrensis sp. nov.) based on three individuals sampled from a previously unknown population from Pico do Jabre, an isolated and small Caatinga moist-forest enclave (CMFE) located in northeastern Brazil. Although this new species has an external morphology resembling those found in representatives of the Bothrops jararaca (Wied-Neuwied, 1824) species group, B. jabrensis can be diagnosed by a combination of meristic and color characters. Molecular phylogenetic analysis indicates the new species represents a unique and highly divergent lineage within Bothrops revealing the existence of a previously unknown phylogenetic lineage that has been evolving as an independent unit for more than 8 million years. Additionally, the estimated divergence time of this lineage conflicts with some proposed scenarios of historical processes associated with the evolution of CMFEs. Finally, the uniqueness of this species indicates its relevance for the maintenance of the phylogenetic diversity of Lanceheads in South America. Like other CMFEs, Pico do Jabre is consistently threatened by poaching, illegal fires, deforestation for agricultural purposes, and illegal logging. The restricted distribution of B. jabrensis, in a small and disturbed CMFE, strongly suggests that this species is critically endangered and is likely approaching extinction as a natural population.
... Bothrops bilineatus has been classified as belonging to the B. taeniatus species group by all recent studies that are focused on the systematic of Bothrops [e.g. (15,16)]. The species group is composed of six species of arboreal pit vipers that include B. bilineatus, B. chloromelas, B. oligolepis, B. medusa, B. pulchra, and B. taeniatus. ...
The two-striped forest-pitviper (Bothrops bilineatus) is an arboreal snake that is currently represented by two subspecies (B. b. bilineatus and B. b. smaragdinus) that comprise a species complex, and its distribution is in the Amazon and the Atlantic Forest. The rarity of encounters with this snake is reflected in the low occurrence of cases of snakebites throughout its geographic distribution and the resulting low number of published clinical reports. However, in some areas, B. bilineatus proves to be more frequent and causes envenomations in a greater proportion. Herein, we review the main aspects of the species complex B. bilineatus, including its biology, ecology, taxonomy, morphology, genetic and molecular studies, geographic distribution, conservation status, venom, pathophysiology and clinical aspects, and epidemiology. In addition, the different antivenoms available for the treatment of envenomations caused by B. bilineatus are presented along with suggestions for future studies that are needed for a better understanding of the snakebites caused by this snake.
... Bothrops species are distributed in both forest blocks and open areas over the Neotropics, making them an ideal model for studying biogeographical processes in the Neotropical region. The group counts with recently phylogenies available [57,58], and The Reptile Database [56] currently recognizes 45 species of lanceheads. Within these species, a group of 18 species stands out, as it is composed mainly by forest lanceheads, including Bothrops moojeni, which occurs in gallery forests throughout the Cerrado region [59], a geographic domain with savannah-like open vegetation. ...
... To perform biogeographic analyses described below, we used the recently published molecular dataset provided by Carrasco et al. [58] to generate a dated molecular phylogeny for our focal clade. In their study, Carrasco et al. [58] generated a total evidence phylogeny of the genus Bothrops, with the description of a new species. ...
... To perform biogeographic analyses described below, we used the recently published molecular dataset provided by Carrasco et al. [58] to generate a dated molecular phylogeny for our focal clade. In their study, Carrasco et al. [58] generated a total evidence phylogeny of the genus Bothrops, with the description of a new species. Their dataset comprised only mitochondrial genes (12s, 16s, cox1, cytb and nd4), with 412 sequences. ...
The emergence of the diagonal of open/dry vegetations, including Chaco, Cerrado and Caatinga, is suggested to have acted as a dispersal barrier for terrestrial organisms by fragmenting a single large forest that existed in South America into the present Atlantic and Amazon forests. Here we tested the hypothesis that the expansion of the South American diagonal of open/dry landscapes acted as a vicariant process for forest lanceheads of the genus Bothrops , by analyzing the temporal range dynamics of those snakes. We estimated ancestral geographic ranges of the focal lancehead clade and its sister clade using a Bayesian dated phylogeny and the BioGeoBEARS package. We compared nine Maximum Likelihood models to infer ancestral range probabilities and their related biogeographic processes. The best fitting models (DECTS and DIVALIKETS) recovered the ancestor of our focal clade in the Amazon biogeographic region of northwestern South America. Vicariant processes in two different subclades resulted in disjunct geographic distributions in the Amazon and the Atlantic Forest. Dispersal processes must have occurred mostly within the Amazon and the Atlantic Forest and not between them. Our results suggest the fragmentation of a single ancient large forest into the Atlantic and Amazon forests acting as a driver of vicariant processes for the snake lineage studied, highlighting the importance of the diagonal of open/dry landscapes in shaping distribution patterns of terrestrial biota in South America.
... Because of the representativeness of the viperid group in the Americas and the relatively well-known phylogenetic relationships for certain genera, they are of great importance as ecological and phylogeographical models (Castoe et al., 2009;Alencar et al., 2018;Jadin et al., 2019). And, although New World pitvipers have been described since the beginning of modern taxonomy (e.g., Crotalus horridus Linneaus 1758), many new taxa continue to be uncovered (Jadin et al., 2012;Townsend et al., 2013;Grü nwald et al., 2015;Doan et al., 2016;Carrasco et al., 2019;Timms et al., 2019) suggesting greater species diversity than is currently recognized. ...
The jumping pitvipers of the genus Metlapilcoatlus are generally associated with montane environments and are widely distributed from central and southern Mexico to Panama. In this study, we analyzed the phylogenetic position and compared the morphology of a population from the Sierra Madre Oriental, within the Sierra Gorda Biosphere Reserve, to other species of jumping pitvipers. We implemented two mtDNA gene fragments (cyt b and ND4) of 50 specimens of the five Metlapilcoatlus species for phylogenetic reconstruction using Bayesian inference and maximum likelihood. We used a linear discriminant analysis and comparative statistics of external and hemipenial characters to assess the morphological differences among Metlapilcoatlus lineages. Our analyses support a strong genetic and morphological distinction of Metlapilcoatlus nummifer populations established to the north of the Trans-Mexican Volcanic Belt (Hidalgo, Northern Veracruz, and Querétaro) from other Metlapilcoatlus taxa. Based on this evidence we describe the populations from the north as a new species.
... Venomous snakes in the genus Bothrops Wagler 1824 (family Viperidae) have been the focus of several phylogenetic and diversification studies over recent years (Wüster et al. 2008;Fenwick et al. 2009;Carrasco et al. 2012;Alencar et al. 2016), with an equally active taxonomy revelling unrecognised diversity at the species level (Ferrarezzi and Freire 2001;Marques et al. 2002;Da Silva and Rodrigues 2008;Folleco-Fernández 2010;Barbo et al. 2012Barbo et al. , 2016Carrasco et al. 2019;Timms et al. 2019). Despite these efforts, conflicting topologies, taxonomic uncertainty, and underestimation of species diversity persist (Machado et al. 2014;Dal Vechio et al. 2018. ...
... otavioi andB. sazimai, Barbo et al. 2012, 2016;Carrasco et al. 2019;B. monsignifer Timms et al. 2019) and phylogenetic studies failing to fully resolve relationships within the neuwiedi, atrox, taeniatus and jararacussu species groups (Machado et al. 2014;Saldarriaga-Córdoba et al. 2017;Dal Vechio et al. 2018. ...
... We examined 41 specimens of Bothrops brazili spanning the taxon's entire geographic distribution (Appendix A). We complemented this examination with data available in the literature (Amaral 1923;Hoge 1953Hoge , 1966Hoge & Lancini 1962;Roze 1966;Hoge et al. 1972;Cunha & Nascimento 1975, 1978, 1993Gasc & Rodrigues 1980;Chippaux 1986;Dixon & Soini 1986; Campbel & Lamar 1989, 2004 Ferrarrezzi & Freire 2001;Duellman 2005;Da Silva & Rodrigues, 2008;Carrasco et al. 2019;Timms et al. 2019). ...
Recent genetic studies have found unclear species boundaries and evidence of undescribed diversity in the poorly studied jararacussu species group within Bothrops. In this contribution, we investigate phenotypic and genetic diversity in the Amazonian snake Bothrops brazili to test previous assertions of unrecognised species diversity within this taxon. Our phylogenetic results and inferences of independently evolving lineages based on molecular data recover two divergent clades within B. brazili, one restricted to areas north and another to areas south of the Amazon River. Phylogenetic relationships between these lineages and other species in the jararacussu species group reveal B. brazili to be paraphyletic, with the northern clade inferred as sister to a clade composed of Atlantic Forest taxa (B. jararacussu, B. muriciensis, B. pirajai). External morphology (number of ventral and subcaudal scales) and colouration patterns (lateral trapezoidal marks) consistently separate the two lineages of B. brazili. We therefore recognise and describe the northern lineage as a new species of Bothrops, improving our knowledge of species diversity within a medically important clade of venomous South American snakes.
... 5,7 With about 50 species of extant pitvipers (http://www.reptiledatabase.org), 11 some of them described recently, [12][13][14] Bothrops (subfamily Crotalinae of Viperidae) represents the most successful South American pitviper radiation. Bothrops species inhabit a wide spectrum of ecoregions throughout tropical Latin America, from sea level to 3000 m of altitude, from northeastern Mexico to Argentina and some parts of the lower Caribbean islands. ...
We report a structural and functional proteomics characterization of the venoms of the two subspecies (bilineatus and smaragdinus) of the South American palm pitviper B. bilineatus from the Brazilian state of Rondônia and B. b. smaragdinus from Perú. These poorly known arboreal and mostly nocturnal generalist predators are widely distributed in lowland rain forest throughout the entire Amazon region, where they represent an important cause of snakebites. The three B. bilineatus spp venom sampled exhibit overall conserved proteomic profiles comprising components belonging to 11 venom protein classes, with PIII (34-40% of the total venom proteins) and PI (8-18%) SVMPs and their endogenous tripeptide inhibitors (SVMPi, 8-10%); bradykinin-potentiating-like peptides (BBPs, 10.7-15%); snake venom serine proteinases (SVSP, 5.5-14%); C-type lectin-like proteins (CTL, 3-10%); phospholipases A2 (PLA2, 2.8-7.6%); cysteine-rich secretory proteins (CRISP, 0.9-2.8%); L-amino acid oxidases (LAO, 0.9-5%) representing the major components of their common venom proteomes. Comparative analysis of the venom proteomes of the two geographic variants of B. b. smaragdinus with that of B. b. bilineatus revealed that the two Brazilian taxa share identical molecules between themselves but not with Peruvian B. b. smaragdinus, suggesting hybridization between the geographically close, possibly sympatric, Porto Velho (RO, BR) B. b. smaragdinus and B. b. bilineatus parental populations. The toxin arsenal of the South American palm pitvipers may account for the in vitro recorded collagenolytic, caseinolytic, PLA2, L-amino acid oxidase, thrombin-like and factor X-activating activities, and the clinical features of South American palm pitviper envenomings, i.e. local and progressively ascending pain, shock and loss of consciousness, spontaneous bleeding and profound coagulopathy. The remarkable crossreactivity of the Brazilian pentabothropic SAB antivenom toward the heterologous B. b. bilineatus venom suggests that the paraspecific antigenic determinants should had been already present in the venom of the last common ancestor of the Bothrops "jararaca" and "taeniatus" clades, an event that has been dated about 8.5 Mya in the mid-late Miocene epoch of the Cenozoic era.
... Lancehead snakes (genus Bothrops) are an excellent study system in which to investigate factors that influence reproduction. The genus contains 45 species widely distributed over a range of climatically distinct ecoregions in Central and South America (Carrasco et al., 2019;Uetz et al., 2019). Moreover, Bothrops species vary greatly in body sizes, body shapes, and macrohabitat use (Martins et al., 2001), and their phylogenetic relationships are reasonably known (Alencar et al., 2016), thus facilitating the interpretation of the evolution of reproductive patterns. ...
Reproductive traits vary widely within and among snake species, and are influenced by a range of factors. However, additional studies are needed on several species, especially on tropical snake faunas, to fully understand the patterns of reproductive variation in snakes. Here, we characterized the reproductive biology of B. jararacussu from southeastern and southern Brazil. We combined macroscopic and microscopic examinations of the reproductive system of museum specimens with observations of free-ranging snakes to characterize size at sexual maturity, sexual size dimorphism, reproductive output, and male and female reproductive cycles. We compared our data with published literature and discuss the factors that may play a role in shaping the reproductive patterns in the species and the genus. Bothrops jararacussu shares several characteristics with its congeners such as autumn mating season, obligatory sperm storage in the female reproductive tract, seasonal timing of parturition (summer-autumn), female-biased SSD, maturity at larger body sizes in females, and a positive relationship between body size and litter size. These characteristics seem phylogenetically conserved in Bothrops. On the other hand, B. jararacussu exhibits some unique characteristics such as a high degree of SSD (one of the highest values recorded in snakes), a large female body size, and a large litter and offspring size, which are among the largest recorded in the genus. Moreover, larger females reproduce more frequently than smaller conspecifics. These characteristics may be collectively interpreted as the result of a strong selection for increased fecundity. Other peculiarities of the species include an asynchrony between spermiogenesis (summer-autumn) and the peak of SSK hypertrophy (autumn to spring) and a prolonged production of SSK granules. Because SSK hypertrophy and mating are androgen-dependent in snakes, the prolonged SSK hypertrophy suggests that male B. jararacussu may prolong their potential to mate (compared with its congeners), which may increase their reproductive success. Our results and previous literature collectively suggest that, in Bothrops, the evolution of SSD is driven by fecundity selection, variation in reproductive output is influenced by variation in female body size, and the timing of spermiogenesis is influenced by other factors in addition to temperature. We also suggest that male Bothrops have undergone multiple evolutionary shifts in the timing of spermiogenesis.
... Phylogenies based on mtDNA ( Alencar et al., 2016;Arteaga et al., 2016; the present study) recovered the species basal to or rooted within Bothrocophias, regarding Bothrops paraphyletic. A recent phylogenetic study combining molecular and morphological evidence recovered B. lojanus in a position that led Bothrocophias to be paraphyletic ( Carrasco et al., 2019). Hence, although undoubtedly B. lojanus is a basal member of the Bothrops + Bothrocophias radiation, it may still be considered an "unstable" taxon whose different positions regarding both genera will likely lead to some taxonomic rearrangement. ...
Based on the literature, we had predicted that the diversification within the Neotropical snake genus Bothrops occurred along a latitudinal gradient from north to south, with diversification into unoccupied niches through ecological opportunity, not correlated with geoclimatic events. Using a dated phylogeny and estimating likelihoods of ancestral states at cladogenesis events, we reconstructed ancestral areas and assessed major events of the diversification of Bothrops clades, and we also discuss systematic implications for this group. Based on the phylogeny we produced, B. lojanus was not considered as part of the genus Bothrops since the results recovered this species nested within the Bothrocophias clade. We infer that the diversification of the Miocene Bothrops pictus and Bothrops alternatus clades may be related to the uplift of the western slopes of the Andes and the Argentinian Patagonian Andes, respectively. The Pliocene Bothrops taeniatus and Bothrops osbornei clades may be related to the uplift of the eastern and northern Andes, respectively. The Plio‐Pleistocene Bothrops neuwiedi clade may be related to the habitat transitions from a warmer and forested environment to a cooler and open landscape; the Bothrops jararaca (i.e. island endemic species) and Bothrops lanceolatus clades to over‐water dispersal with island speciation; and Bothrops atrox clade to the appearance of the Panamanian land bridge. We found that a multitemporal and multidirectional history of diversification may be correlated with geoclimatic and dispersalist events. We argue that the vacant niche hypothesis by itself does not explain Bothrops diversification.
