Neotropical Research Articles

Isospora pichororei n. sp. (Chromista: Apicomplexa: Eimeriidae) from rufous-capped spinetails Synallaxis ruficapilla Vieillot, 1819 (Passeriformes: Furnariidae: Synallaxiinae) in South America

Spinetails are a suboscine passerines of the genus Synallaxis Vieillot, 1818 which have great interest for ornithology, given the wide diversity of 37 species that are distributed throughout the Neotropical region. Despite this wide diversity and distribution, Synallaxis spp. have never been recorded as hosts of coccidian parasites. In this context, the current study describes a new species of Isospora Schneider, 1881 from rufous-capped spinetails Synallaxis ruficapilla Vieillot, 1819 captured in the Itatiaia National Park, which is a federal conservation unit in Southeastern Brazil. The oocysts of Isospora pichororei Genovez-Oliveira & Berto n. sp. are subspheroidal to ovoidal, measuring on average 25 by 21 μm. Micropyle is present, but discrete. Oocyst residuum absent, but one or two polar granules are present. Sporocysts are ellipsoidal with slightly pointed posterior end, measuring on average 17 by 10 μm. Stieda and sub-Stieda bodies are present. Sporocyst residuum is clustered among the vermiform sporozoites, which have striations, refractile bodies and nucleus. This morphology was different from the other Isospora spp. recorded in the host family Furnariidae. Molecular identification was targeted by the amplification and sequencing of a locus of the mitochondrial cytochrome c oxidase subunit 1 (cox1) gene. This sequence had the highest similarity of 99.5% with a sequence deposited for Isospora oliveirai Ortúzar-Ferreira & Berto, 2020, which is a coccidian species that parasitizes suboscine tityrids Schiffornis virescens (Lafresnaye, 1838), also in the Itatiaia National Park. Phylogenetic analysis grouped some species in subclades, including I. pichororei with I. oliveirai; however, it was inconclusive in an expectation of parasite-host coevolution. Finally, I. pichororei is established as new to science, being the first description from Synallaxinae and the third description from Furnariidae. Furthermore, this is the first Isospora sp. from the host family Furnariidae to have a molecular supplementation by sequencing a locus of the cox1 gene of the mitochondrial genome.

Description of juvenile instars of Scapheremaeus tonatiuh Palacios-Vargas, Ríos & Vázquez, 1998 (Acari, Oribatida, Cymbaeremaeidae)

The oribatid mite species Scapheremaeus tonatiuh (Oribatida, Cymbaeremaeidae) is recorded for the first time in the U.S.A. (initially described from Mexico). Its juvenile instars are described, based on materials collected from arboreal habitats in Florida, U.S.A. (Neotropical region). The larval instar is characterized by the presence of gastronotic setae с3, la, lm, lp long, lanceolate having setiform apex, and h1 very long, subflagellate; the nymphal instars are characterized by the presence of exuvial scalps, 12 pairs of gastronotic setae (da, dm, dp absent) of which setae с3, la, lm, lp, h1, h2 are long, phylliform but h3 is very long, subflagellate. The main morphological traits of the adult are summarized.

The Neotropical Orthonevra Macquart, 1829 (Diptera: Syrphidae)

The Neotropical species of the flower fly genus Orthonevra (Diptera: Syrphidae) are revised and 12 new species are described and illustrated: Orthonevra batman sp. nov., O. cthulhu sp. nov., O. gli sp. nov., O. gouletmasnerorum sp. nov., O. hypnotica sp. nov., O. minuosetulata sp. nov., O. nebulosa sp. nov., O. neiae sp. nov., O. roseae sp. nov., O. sternolobosa sp. nov., O. theta sp. nov., and O. thompsoni sp. nov. In addition, seven species are redescribed and illustrated: O. argentina (Brèthes, 1922), O. chilensis Thompson, 1999, O. flukei (Sedman, 1964), O. gewgaw (Hull, 1941), O. neotropica (Shannon, 1925), O. quadristriata (Shannon & Aubertin, 1933) and O. shannoni (Curran, 1926), and three Nearctic (one extending into the Neotropical region) species are diagnosed: O. bellula (Williston, 1882), O. nitida (Wiedemman, 1830) and O. nitidula (Curran, 1925) (for a total of 20 species with Neotropical distribution). Lectotypes are designated for two species, Chrysogater bellulus Williston and C. nitidus Wiedemann, and C. lugubris Jaennicke (1867) is removed from Orthonevra and left incertae sedis. A dichotomous and an interactive pictorial key to these species are also presented.

Vulnerability assessment of natural heritage in the North-Eastern Ecuadorian Amazon using land use cover and nature protection status

This study evaluates the vulnerability of natural heritage in the Northeast Ecuadorian Amazon (NEA), from the perspective of intrinsic vulnerability. Therefore, two proxy indicators of vulnerability were established: (i) the ecological integrity vulnerability index (EIVI), indicating the potential loss of ecological integrity, assessed by protection status (2018); (ii) the biological diversity vulnerability index (BVI), deriving the potential loss of biological diversity, using land cover/land use (2014). Biodiversity values were derived with land cover-related biodiversity data, and (iii) and compared with spatial congruence/correlation between (i) and (ii). The EIVI was directly derived using protection status, which defines vulnerability levels by ranking them according to corresponding levels of anthropic development allowance. A robust relationship between land use classes and species richness was found for both endemic vascular plants in neotropical regions and multi-taxa in tropical forests worldwide, validating the BVI. Furthermore, the moderate spatial correlation between protection status and land cover (r = 0.52) may be partially explained by 3,060 km2 of unprotected primary forest. Overall, 42 % of the NEA have a high degree of congruence between potential biodiversity and protection status. In conclusion, this study provides insights into nature conservation efforts in areas with sparse biodiversity data using readily available geographic information.

Induction of Bone Malformations in Neotropical Fish (Megaleporinus Macrocephalus) Exposed to Pyriproxyfen During Initial Stage of Development

Objective: Megaleporinus macrocephalus, a native species from the Neotropical region, was used as a model to evaluate the effects of pyriproxyfen on larval development at a dose that could be found in the environment. Theoretical Framework: Pyriproxyfen is a compound used to control of the mosquito Aedes aegypti, vector of Neotropical diseases. In insects, pyriproxyfen acts as a juvenile hormone analogue, inhibiting embryogenesis and the development of characteristics of adults through the inhibition of ecdysone. Pyriproxyfen has demonstrated effects on the survival rate, behavior, and morphology during development of non-target organisms. Method: In this study Megaleporinus macrocephalus was exposed to a dose of 2.5μg/L of pyriproxyfen on the 8th day after hatching and the presence of bone malformations was evaluated. For the evaluation was applied technique of diaphanization and radiological imaging. Results and Discussion: Out of the total of the individuals, 43% had some type of bone malformation: 24% had scoliosis and kyphosis of the vertebral column, 9% had reduction of pectoral fins, 5% had malformation in the anal fin and 5% absented at least one of the pelvic fins. These data reinforce the action of pyriproxyfen as a teratogen, acting on larval development in reduced doses, which can be found in the environment. Our study warns that these molecules might interact in the mechanism of gene expression mediated by retinoic acid, important morphogen during embryogenesis and organogenesis. Research Implications: The practical and theoretical implications of this research are discussed, providing insights into how the results can be applied or influence practices in the field of teratology. These implications could encompass control of the vectors, conservation biology and the prevention of embryonic and fetal anomalies. Originality/Value: This study not only does it reinforce the urgency to study how these molecules interact but also suggests caution when using the compound as a controller for vectors until the true security is acknowledged.

Biological invasions negatively impact global protected areas

Protected areas underpin global biodiversity conservation and sustainability agendas. Biological invasions increasingly threaten the ecological functioning and long-term conservation value of protected areas, while a lack of information on impact impedes management decisions. We collated data from effects of biological invasions in protected areas to provide the first quantitative analysis of their global impacts. Based on 300 reported effects from 44 invasive species, we show that there are overall negative impacts from invasive species on both biotic and abiotic characteristics of protected areas globally. Impacts were pervasive across population, community, and ecosystem scales, and for the vast majority of invasive taxa with sufficient data. Negative impacts have been incurred around the world, with National Parks and World Heritage Sites in the Neartic and Neotropical regions the most studied. Notwithstanding context-dependencies and uneven research efforts, the recurrent negative impacts of invasive species indicate that current efforts are insufficient to curb current stressors and meet conservation and sustainability targets on land and in water. To address the risk of biological invasions in protected areas, it is imperative to prioritise fundamental research on ecological interactions, establish robust monitoring and prevention programs, and raise awareness through global initiatives.

First record of subgenus Synaldis Foerster (Hymenoptera, Braconidae, Alysiinae, Dinotrema Foerster) from Chile, with description of ten new species.

Synaldis is a taxon within the Aspilota group with a contentious taxonomic history, currently classified as a subgenus of the genus Dinotrema. Species of Synaldis were only documented in the Neotropical region in 2017, and until then, the Neotropical fauna of this subgenus was represented by five species from Brazil. In this study, Synaldis is reported for the first time in Chile, with the description and illustration of ten new species, namely: Dinotrema (Synaldis) acarinareolatumsp. nov., D. (S.) brunneumsp. nov., D. (S.) chilensesp. nov., D. (S.) daltonisp. nov., D. (S.) flavumsp. nov., D. (S.) latusdentertiumsp. nov., D. (S.) perisfelipoisp. nov., D. (S.) pilosicaudatumsp. nov., D. (S.) puyehuesp. nov., and D. (S.) veraesp. nov. The studied specimens were collected during expeditions to southern Chile, in the Valdivian temperate rainforest at Parque Nacional de Puyehue. This study also includes a dichotomous identification key for Neotropical species of Synaldis, as well as a discussion of the primary morphological characters used to distinguish species within the Neotropical and Nearctic regions.

When the predator becomes the prey: new records of intraguild predation among Central American and Caribbean arachnids (Arachnida: Amblypygi, Araneae, Scorpiones)

ABSTRACT Intraguild predation occurs when an organism preys on a competitor. This ecological interaction can have considerable consequences on the population dynamics of predators and indirectly affect their prey. In arachnids, it is a quite common phenomenon, but it has been poorly documented in the Neotropical region. Here, we present records of 16 species feeding on other 14 species of arachnids. The most common families that acted as predators were Buthidae (Scorpiones) and Salticidae (Araneae), while the most common prey were members of the families Araneidae and Ctenidae (Araneae). In our survey, Drake Bay, in Costa Rica, is overrepresented due to years of systematic observations by two of the coauthors.

The role of changing landscape in the dispersal of a soil‐feeding termite in Suriname and French Guiana

Abstract Neotropical forest ecosystems harbour significant biodiversity. To develop effective insect conservation practices, it is important to understand the factors that influence the diversity and population structure of the species. Dispersal, a key determinant of population structure, is well studied in termites nesting in wood, for which it can be influenced by wood transport, or in termites living in urban environments. However, understanding the dispersal of termites whose mobility remains unaffected by wood transport remains understudied. We investigated the dispersal of Embiratermes neotenicus, a soil‐feeding species with short dispersal distance, in the Neotropical region, where both intact and degraded forests exist. Using mitochondrial and nuclear data, we analysed genetic diversity, structure and factors contributing to population differentiation in Suriname and French Guiana at multiple scales for 70 colonies. The population in French Guiana is the ancestral population in the region that subsequently expanded. Significant genetic differentiation between populations was observed, with distinct patterns identified in Suriname and French Guiana. The Suriname population showed higher genetic diversity and no subpopulation differentiation, whereas the French Guiana population showed substructure into distinct genetic clusters. Analyses at the scale of all colonies suggest the influence of landscape features, such as the Maroni River, on genetic differentiation. At the local scale, genetic differentiation between colonies increases with forest alteration, even when this does not include major changes in forest cover. Our results highlight the sensitivity of soil‐feeding termite populations to habitat change. We argue that multi‐scale studies are needed for a comprehensive understanding of genetic patterns, especially for species with short dispersal distances.

Isolation and characterization of a native strain of the entomopathogenic fungus Beauveria bassiana for the control of the palm weevil Dynamis borassi (Coleoptera: Curculionidae) in the neotropics

This study aimed to isolate and characterize a native strain of Beauveria bassiana, coded as Bv065, showcasing its potential as a biological control agent targeting the palm weevil Dynamis borassi. Originating from a naturally infected D. borassi specimen collected in southwestern Colombia, the fungus underwent molecular identification and was identified as B. bassiana, exhibiting high sequence similarity with known reference strains. The physiological characterization revealed that Bv065 thrived within a temperature range of 25 to 30 °C and a pH range of 6 to 9. Moreover, the key carbon sources that allow optimal growth of the strain were identified through metabolic profiling, including sucrose, D-mannose, and γ-amino-butyric acid. These findings offer strategic insights for scalability and formulation methodologies. Additionally, enzymatic analyses unveiled robust protease activity within Bv065, crucial for catalysing insect cuticle degradation and facilitating host penetration, thus accentuating its entomopathogenic potential. Subsequent evaluations exposed Bv065’s pathogenicity against D. borassi, causing significant mortality within nine days of exposure, albeit exhibiting limited effectiveness against Rhynchophorus palmarum. This study underscores the importance of understanding optimal growth conditions and metabolic preferences of B. bassiana strains for developing effective biopesticides. The findings suggest Bv065 as a promising candidate for integrated pest management strategies in neotropical regions, particularly for controlling palm weevil infestations in coconut and peach palm cultivation. Future research avenues include refining mass production methodologies, formulating novel delivery systems, and conducting comprehensive field efficacy trials to unlock the full potential of Bv065 in fostering sustainable pest management practices. Overall, this study contributes to the growing body of knowledge on entomopathogenic fungi and their pivotal role in biological control, offering nuanced perspectives on eco-friendly alternatives to conventional insecticidal interventions.

Revisiting the biogeographical regionalization of the Pacific Lowlands biogeographical province using bird distributional data

Abstract The individual tracks of 47 bird species endemic to the Pacific Lowlands biogeographical province, which belongs to the Mesoamerican dominion of the Neotropical region, were used to revise its biogeographical regionalization. As a result, the six biogeographical districts previously recognized are classified in two subprovinces: the Coastal Lowlands of Nayarit-Sinaloa subprovince, stat. nov., which includes the Sinaloan, Tres Marías Islands, Revillagigedo Islands, and Guerreran districts; and the Coastal Lowlands of Southwestern Mexico subprovince, stat. nov., which includes the Tehuanan and Tapachultecan districts. Bird species supporting the biogeographical subprovinces and districts recognized are listed. The recognition of the subprovinces encompassing the districts represents an improvement of this biogeographical regionalization.

An updated checklist of Endomychidae and Anamorphidae (Coleoptera: Coccinelloidea) from El Salvador, with additional new records from the Neotropical region

The checklist of the species of fungus beetle families Endomychidae and Anamorphidae from El Salvador is updated. Only six species of Endomychidae and one Anamorphidae have been previously recorded from the country, with specific localities reported only for two of them. The study of specimens collected in El Salvador over 60 years ago by Jan and Bohumila Bechyné and held in the Royal Belgian Institute of Natural Sciences, Brussels, allowed us to have a better idea on the diversity and distribution of these beetles in the country. Eight species of Endomychidae and one of Anamorphidae arerep orted here, including three first records for El Salvador: Anidrytus guatemalae Arrow, Epipocus bifidus Gerstaecker andStenotarsus ovalis Arrow. Additionally, first records for species of Endomychidae from countries of the Neotropical region are provided: Anidrytus batesi Strohecker from Peru, A. ovatulus Gerstaecker from Guyana, Ephebus hirtulus Stroheckerfrom Paraguay, Epopterus vacuus Gerstaecker from Costa Rica and Acinaces unicolor Tomaszewska from Peru.

Evolution of fossoriality in microteiid lizards

Abstract Morphology is among the most important traits influencing the interaction of individual animals with their environments. Fossoriality reflects this functional association between morphology and the use of subterranean habitats and their associated environmental characteristics. Lizards in the families Gymnophthalmidae and Alopoglossidae are model organisms to examine the interplay between morphology and fossoriality because great morphological diversity exists among species, including varying degrees of body elongation and limb reduction, and they have a wide geographical distribution in the Neotropical region. We analysed the morphology of 101 microteiid species and created an index to evaluate their degree of fossoriality. From this index, we traced the evolution of fossoriality in these lizards and assessed its primary environmental correlates. We found that fossoriality evolved independently in several lineages, mainly associated with high temperature and low precipitation, characteristic of more arid and sandy environments.

Removal of historical taxonomic bias and its impact on biogeographic analyses: a case study of Neotropical tardigrade fauna

Abstract Large-scale databases are crucial for macroecology research, yet not entirely bias-free. Studying the biogeography of metazoan microfauna utilizing such databases is challenging, and added biases can further hinder them. Incidence data of tardigrade species from Central and South America are geographically biased, comprising mostly species erroneously considered cosmopolitan. The impact of these biases on macroecological studies is still unknown. This paper evaluates how biased distribution records affect understanding of non-marine tardigrade distribution patterns in the Neotropical region. Using two datasets (including and excluding dubious records of allegedly cosmopolitan species), we assess how well biogeographic regions are sampled, estimate tardigrade species’ richness, and analyse whether including dubious records can alter our comprehension of their macroecology. We demonstrate how biogeographic regions are still not fully sampled, and that including unreliable records influences how many species remain to be discovered. Since records of ‘false cosmopolitan species’ represent most records, their removal increases uncertainty while yielding distribution patterns more likely to be accurate. Disregarding untrustworthy distribution records of ‘false cosmopolitan species’ is the first step to better understanding tardigrade macroecology in the Neotropics, and probably worldwide. However, additional, preferably systematic sampling is required before we can infer general tardigrade biodiversity patterns in under-sampled regions.

Breeding records of 325 bird species from Peru over 21 years based on citizen science data

AbstractBreeding is a key component of the life history of birds. Documenting diverse aspects of their breeding biology provides valuable natural history data that can be used to address questions of evolutionary, ecological, or conservation significance. Despite its importance, this information is scarce for most Neotropical species. We compiled 1180 breeding records corresponding to 325 bird species for the period 2000–2021, through an in‐depth review of photo or video records from Peru, sourced from Macaulay Library and iNaturalist platforms. We determined the geographical distribution of the breeding records, described patterns of activity and nesting behavior (i.e., host plant preference and clutch size), and explored the timing of breeding. Our study represents one of the few efforts to provide nationwide data on breeding birds in the Neotropical region. We also offer broad taxonomic coverage for further studies interested in testing hypotheses on the variation of reproductive traits and timing at both local and regional scales. The complete data set for this abstract published in the Data Article section of the journal is available in electronic format in MetaCat in JaLTER at http://db.cger.nies.go.jp/JaLTER/metacat/metacat/ERDP-2024-07.1/jalter-en.

Integrative taxonomy clarifies the armoured catfish Hypostomus pusarum (Starks) species complex (Siluriformes: Loricariidae) and reveals a new species in the drainages of Northeastern Brazil

Abstract Hypostomus is the most species-rich genus within the family Loricariidae and is widely distributed throughout the Neotropical region. Nonetheless, the diversity and distribution of these species have still large knowledge gaps. This scenario is more significant in some regions, such as the northeast of Brazil. In this region, the first species of the genus, H. pusarum, was described in the Northeast Caatinga and Costal Drainages ecoregion. Six congeners were subsequently described in the same ecoregion, all sharing the same colour pattern making them difficult to distinguish. All of them are collectively referred to as the H. pusarum complex. The present work seeks to clarify the diversity that constitutes the H. pusarum complex through an integrative study using molecular and morphological data. The results indicate that H. carvalhoi, H. jaguribensis, H. nudiventris, H. papariae, and H. salgadae are all junior synonyms of H. pusarum. However, one of the morphotypes that occurs in the Parnaíba River is a new species that differs from the others by the absence of a developed medial buccal papilla and the presence of a pre-anal plate. The data provided here highlight the importance of integrative taxonomy for assessing diversity in complex and diverse groups in the Neotropics.

First record of Acarophenacidae (Acari: Heterostigmata) from Mexico, with description of a new species of Paracarophenax associated with Cerambycidae (Insecta: Coleoptera)

Paracarophenax (Acari: Heterostigmata: Acarophenacidae) comprises eight distinct species of egg parasitoids associated mainly with various beetles, distributed across different global regions. In this study, a new species of Paracarophenax was found associated with cerambycid beetles´ collection samples deposited in the Colección de Insectos de la Universidad de Morelos, Morelos, Mexico. The aim of this paper is to describe the ninth species of the Paracarophenax, and provide an updated taxonomic key of the genus. The new species can be separated from others congeners by the presence of one spiniform seta on genu II, the seta l’ on femur I slightly spatulate, one seta on femur III, setae e slightly longer than f, and the distance between setae e and f slightly longer or equal to the distance between setae h<sub>1</sub> and h<sub>2</sub>. This study reports the family Acarophenacidae for the first time from Mexico, and the genus Paracarophenax from the Neotropical Region.

Predicting the potential distribution and coexistence of Chagas disease vectors in the Americas.

Predicting the potential distribution and coexistence of suitable geographic areas for Chagas disease vectors in the Americas is a crucial task for understanding the eco-epidemiological dynamics of this disease. The potential distribution and coexistence of 3 species-Rhodnius prolixus (Hemiptera: Reduviidae), Cavernicola pilosa (Hemiptera: Reduviidae), and Rhodnius pictipes (Hemiptera: Reduviidae) were modeled. Presence records were obtained and environmental variables were selected based on correlation analysis, Jackknife analysis and knowledge of the biology and natural history of the species. The MaxEnt algorithm included in the kuenm package of R software was used for modeling the potential distribution, and various scenarios of the BAM diagram (Biotic, Abiotic, and Movement variables) were evaluated. The variables contributing to the final models were different for each species. Rhodnius pictipes showed a potential distribution in South America, particularly in Brazil, Bolivia, Peru, Colombia, Venezuela, Guyana, and Suriname. Areas with environmentally suitable conditions for R. prolixus were located in southern Brazil, Peru, Colombia, southern Mexico, Guatemala, El Salvador, and Honduras, whereas for C. pilosa they were in southeastern Brazil, southeastern Central America, Peru, Ecuador, Colombia, Venezuela, Guyana, Suriname, and French Guiana. Co-occurrence analysis revealed distinct patterns in the neotropical region, with some areas indicating the potential distribution of 1 or more species. In Brazil, occurrence and co-occurrence areas were concentrated in the northwest and southeast regions. Overall, this study provides valuable information on the potential distribution and coexistence of vectors, which can inform targeted vector control strategies and contribute to global efforts in combating Chagas disease.

Large Protected Areas Safeguard Mammalian Functional Diversity in Human-Modified Landscapes

Habitat loss and fragmentation are pervasive processes driving the disappearance of populations and species in the Neotropical region. Since species loss may translate into functional loss, assessing changes in the composition of assemblages’ functional traits might improve our understanding of the ecological roles played by species and ecosystem functioning. Here, we investigate how landscape structure and composition impact the functional diversity of terrestrial mammals in 18 forest patches composing eight protected areas in Southern Brazil. We used functional diversity (FD) based on dietary, physical, and behavioral traits and species vulnerability to extinction. We determined which landscape variables (patch size, proportions of forest and sugarcane, and patch isolation) most influenced mammal FD values by using a both-direction stepwise model selection from a linear global model. Finally, we evaluated the role of trophic guilds in explaining the variation in the FD values using a Principal Component Analysis. Between 2012 and 2017, using camera traps, we recorded 26 native medium- and large-sized mammals throughout the protected areas, of which 6 are regionally threatened, and 5 domestic/exotic species. Richness among the forest patches varied from 4 to 24 species (9.05 ± 5.83), while the FD values varied from 1.29 to 6.59 (2.62 ± 1.51). FD variation was best explained by patch size, which exhibited a strong positive correlation (adjusted R2 = 0.55, slope = 0.67, p < 0.001). Insectivores and frugivores presented the highest correlation with patch size, explaining most of the variation in the FD values. Our findings strengthen the paramount role of large protected areas in maintaining mammal diversity and their ecological functions in human-modified landscapes.

Revalidation of the termitophilous genus Ceratoxenus Mann and description of two new species from the Neotropical region (Coleoptera: Staphylinidae: Aleocharinae: Corotocini)

Termitogastrina is a subtribe of termitophilous rove beetles distributed mostly across Neotropical and Afrotropical regions. In 1923, William M. Mann established the genera Ceratoxenus and Xenopelta from material collected by Alfred E. Emerson in Guyana. Nine years later, Emerson himself synonymized the genera, in what seems to be a result from the misidentification of specimens. In this work, we revalidate the genus Ceratoxenus, provide its redescription and a taxonomical key for the species, along with a comparative diagnosis with the genus Xenopelta. Additionally, two new species of Ceratoxenus are described: C. akerata sp. nov. and C. inflatus sp. nov., respectively, from Panama and Guyana. This paper also marks a new record for the genus Ceratoxenus for Panama.