Pathogenic hantaviruses, northeastern Argentina and eastern Paraguay.We describe the first, to our knowledge, cases of hantavirus pulmonary syndrome hantavirus pulmonary syndrome An often fatal RTI caused by a hantavirus; the first cluster occurred in the Four Corners region of Southwestern US Epidemiology Mean age 32, 61% ♀, 72% Native American Case definition Unexplained bilateral interstitial in northeastern Argentina and eastern Paraguay. Andes and Juquitiba (JUQ) viruses were characterized. JUQV was also confirmed in 5 Oligoryzomys nigripes reservoir species from Misiones. A novel Akodon-borne genetic hantavirus hantavirus, any of a genus (Hantavirus) of single-stranded RNA viruses that are carried by rodents and transmitted to humans when they inhale vapors from contaminated rodent urine, saliva, or feces. There are many strains of hantavirus. lineage was detected in 1 rodent from the Biologic Reserve of Limoy. ********** Members of the genus Hantavirus (family Bunyaviridae) are commonly transmitted to humans through rodents and may cause 2 severe human diseases: hemorrhagic fever with renal syndrome hemorrhagic fever with renal syndrome n. See epidemic hemorrhagic fever. and hantavirus pulmonary syndrome (HPS See Seer*HPS. ) (1). The number of recognized human cases and the number of distinct hantavirus genotypes identified have increased during recent years in Argentina and in the 3 southern HPS-endemic states of Brazil Brazil is divided into twenty-six estados ("states"; singular estado) and one district, the Distrito Federal ("Federal District") which contains the capital city, Brasília. (2,3). Six pathogenic Andes virus (ANDV) lineages that cause HPS have so far been found to circulate in Argentina in 3 HPS-endemic areas: Oran and Bermejo (BMJ BMJ n abbr (= British Medical Journal) → vom BMA herausgegebene Zeitschrift ) in the north; Lechiguanas (LEC (1) (LAN Emulation Client) A software driver that provides LAN emulation (LANE) in an ATM network. It resides in an ATM end station or in a computer system that provides the LAN to ATM conversion, often known as a LAN access device. See LANE. ), Hu39694, and AND Cent (Central) Plata in the central provinces; and ANDV in the South (4-7). Different hantavirus genetic lineages associated with HPS cases were reported in Brazil, such as Juquitiba virus (JUQV), Castelo dos Sonhos virus (CASV CASV Cooperstown All-Star Village CASV Clear Advanced Super View ), and Araraquara virus (ARAV ARAV Association of Reptilian and Amphibian Veterinarians ARAV Aravan Bat Virus ARAV Army Aviator ARAV Aegis Readiness Assessment Vehicles ) (8). Recently, pathogenic hantaviruses from Parana, southern Brazil, have been reported to belong to the same clade clade Cladus, subtype Genetics A branch of biological taxa or species that share features inherited from a common ancestor; a single phylogenetic group or line. See Inheritance, Species. as the Oligoryzomys nigripes-associated strains (9). In western Paraguay, cases were associated with Laguna Negra virus (LNV LNV Landbouw, Natuur en Voedselkwaliteit (Agriculture, nature and food quality Government ministry, Netherlands) LNV Londolovit, Papua New Guinea - Londolovit (Airport Code) ) from Calomys laucha (10). In northern Argentina and Bolivia, LNV was obtained from C. callosus (11,12). No cases have been reported in other areas of Paraguay, to our knowledge, until this study. We describe 3 HPS cases that occurred in northeastern Argentina, Misiones Province, which borders 2 other hantavirus-endemic countries, Brazil and Paraguay. We also describe what we believe to be the first case that occurred in eastern Paraguay and analyzed rodents captured in Misiones and characterized a novel genetic hantavirus lineage from the Biologic Reserve of Limoy in eastern Paraguay. The Study Three HPS cases were confirmed in Misiones Province, Argentina, in the following patients: a 14-year-old boy from Santa Ana in November 2003, a 28-year-old man from Leandro N. Alem in December 2003, and a 12year-old boy living in the Dos Arroyos locality of Alem city. In January 2005, HPS was confirmed in a 19-yearold girl living in Pirapo, a rural area of Itapua Department, Paraguay. Figure 1 shows the approximate geographic location of exposure sites for patients included in this study. Clinical manifestations in the 4 patients studied were similar to those reported for ANDV infections: fever, myalgia myalgia /my·al·gia/ (mi-al´jah) muscular pain.myal´gic epidemic myalgia see under pleurodynia. my·al·gia n. , headache, and vomiting, soon followed by pulmonary edema Pulmonary Edema Definition Pulmonary edema is a condition in which fluid accumulates in the lungs, usually because the heart's left ventricle does not pump adequately. . Thrombocytopenia Thrombocytopenia Definition Thrombocytopenia is an abnormal drop in the number of blood cells involved in forming blood clots. These cells are called platelets. and hemoconcentration were reported, renal involvement was minimal, neither oliguria oliguria /ol·i·gu·ria/ (ol?i-gu´re-ah) diminished urine production and excretion in relation to fluid intake.oligu´ric ol·i·gu·ri·a n. Abnormally slight or infrequent urination. nor renal failure was observed in any case-patient, but all case-patients showed petechiae Petechiae Tiny purple or red spots on the skin associated with endocarditis, resulting from hemorrhages under the skin's surface. Mentioned in: Endocarditis, Hantavirus Infections, Hemorrhagic Fevers, Idiopathic Thrombocytopenic Purpura . The 4 HPS case-patients had immunoglobulin M (IgM) and IgG antibodies to ANDV N recombinant protein by ELISA ELISA (e-li´sah) Enzyme-Linked Immuno-Sorbent Assay; any enzyme immunoassay using an enzyme-labeled immunoreactant and an immunosorbent. ELISA n. (13) and all survived. These cases led us to investigate reservoirs for hantaviruses in Misiones by using Sherman live-capture traps (H.B. Sherman Traps, Tallahassee, FL, USA). A total of 59 rodents were trapped at 2 study sites in Misiones, where O. nigripes was the most frequently captured rodent (42 specimens), followed by Akodon montensis (11 specimens). The rodent species was identified by morphologic features, in particular, qualitative external and cranial cranial /cra·ni·al/ (-al) 1. pertaining to the cranium. 2. toward the head end of the body; a synonym of superior in humans and other bipeds. cra·ni·al adj. characteristics. Animals were tested for ANDV IgG antibodies (Table 1). The capture done in August 2005 in Santa Ana and Leandro N. Alem cities found 5 (11.9%) ANDV-positive O. nigripes rodents of 42 tested from this species. [FIGURE 1 OMITTED] Before we discovered the case in Itapua, we had conducted a serosurvey of rodents in 3 departments in eastern Paraguay (Table 1). Fifty-one rodent specimens were collected, of which 32 were sigmodontine rodents and 20 were Akodon cursor. Only 1 A. cursor rodent obtained from the Limoy Biologic Reserve, Paraguay, was seropositive seropositive /se·ro·pos·i·tive/ (-poz´i-tiv) showing positive results on serological examination; showing a high level of antibody. se·ro·pos·i·tive adj. . Because of the diversity of akodontine rodents, precise diagnosis, based solely on morphometric characteristics, is not always possible. To confirm the morphologic identification of the A. cursor rodent, we compared the mitochondrial mitochondrial pertaining to mitochondria. mitochondrial RNAs a unique set of tRNAs, mRNAs, rRNAs, transcribed from mitochondrial DNA by a mitochondrial-specific RNA polymerase, that account for about 4% of the total cell RNA that control region (fragment of 245 nt) with that of an A. montensis rodent used as a control. Mitochondrial DNA sequencing of A. cursor from Limoy found higher identity (94%) with an A. cursor from Paraguay (AF296264) than with A. montensis (90%). Positive voucher specimens were archived at the Museo Argentino de Ciencias Naturales and at the University of Buenos Aires To enter any of the available programmes of study in the university, students who have successfully completed high school must pass a first year common to all faculties. This first year is called "CBC", which stands for "Ciclo Básico Común" (Common Basic Cycle). . Amplification by reverse transcription-PCR was performed on the 4 human blood samples and on the lung tissues of the 6 seropositive rodents. Initially, a substantial portion of the nucleoprotein nucleoprotein Macromolecular complex consisting of a protein linked to a nucleic acid, either DNA or RNA. The proteins that combine with DNA are generally of characteristic types called histones and protamines. N coding region of the S segment (nt 50-954) and different fragments of the encoding region of the M segment: G1 glycoprotein glycoprotein (glī'kōprō`tēn), organic compound composed of both a protein and a carbohydrate joined together in covalent chemical linkage. (nt 41-443), G1-G2 glycoprotein (nt 1,728 1,976), and G2 glycoprotein (nt 2,715-2,941) were amplified and subsequently sequenced. Comparison of the viral 905-nt N fragment sequence from case-patient 1 showed the highest degree of identity, [approximately equal to] 90%, with LEC (Table 2). The strains from case-patients 2 and 3 showed little genetic variation between them and were [approximately equal to] 95% identical to ARAV from Parana city, Brazil (12), in the same fragment. Comparison of a G1-G2 fragment available for JUQV strain (nt 1,867-1,976) with that from case-patient 2 showed a 93.6% identity. The G2 fragment from case-patient 3 was 95.6% identical to that of JUQV. Thus, these results suggest that the strains from case-patients 2 and 3 are JUQV, although they demonstrate that the strain called Araucaria araucaria Any pinelike coniferous plant (see conifer) of the genus Araucaria (family Araucariaceae). Found in South America, the Phoenix Islands, and Australia, araucaria trees are magnificent evergreens, with whorled branches and stiff, flattened, pointed leaves. would also be JUQV. Sequences from the 5 positive O. nigripes from Misiones showed little variability between them and the N fragments were 98.3% identical to those in strains from casepatients 2 and 3. The case-patient from Itapua, Paraguay, showed the greatest nucleotide identity (93.5%) with BMJ lineage, in the N fragment; identity at the amino acid level was 99.3%. Another different strain was obtained from the A. cursor rodent captured in Limoy Reserve; this was the most distinct strain. The highest degree of identity exhibited was [approximately equal to] 77% (Table 2). The G1 fragment from this strain was compared with the closest related hantavirus and showed 67% identity with strains isolated from O. nigripes, case-patients 2 and 3, and LNV. The N-encoding sequences were further subjected to phylogenetic phy·lo·ge·net·ic adj. 1. Of or relating to phylogeny or phylogenetics. 2. Relating to or based on evolutionary development or history. analysis. All virus sequences from Misiones and the Itapua case form a monophyletic monophyletic /mono·phy·let·ic/ (mon?o-fi-let´ik) descended from a common ancestor or stem cell. mon·o·phy·let·ic adj. 1. Descended or derived from one original stock or source. group together with ANDV lineages, nonpathogenic Pergamino (PRG PRG Parti Radical de Gauche (French: Left Radical Party) PRG Purge PRG Programming Research Group (Oxford University) PRG Preliminary Remediation Goal PRG People's Revolutionary Government ) and Maciel (Figure 2), and CASV and ARAV (done in a 643-nt parsimonious par·si·mo·ni·ous adj. Excessively sparing or frugal. par  si·mo tree, data not shown). The other South American clade was
formed with Rio Mearim virus and Anajatuba virus from Brazil, together
with LNV, HTN HTN HypertensionHTN High Blood Pressure HTN Hierarchical Task Network HTN Hughes Television Network HTN Hospitality Training Network (Sydney, Australia) HTN Histotechnology (program of study) virus 007, and Rio Mamore virus from Paraguay, Peru, and Bolivia, respectively, with a moderate support of 63%. Sequences from case-patients 2 and 3 from Misiones grouped together with JUQV (Araucaria) to form a separate clade, since they were from the most divergent strains. The sequence from case-patient 1 grouped with LEC lineage from Central Argentina. Different alignment parameters and phylogenetic methods produced the same results in trees with similar topology; however, bootstrap See boot. (operating system, compiler) bootstrap - To load and initialise the operating system on a computer. Normally abbreviated to "boot". From the curious expression "to pull oneself up by one's bootstraps", one of the legendary feats of Baron von Munchhausen. supports were moderate or low level for some lineages. The sequence from the A. cursor rodent was a quite genetically distinguishable virus lineage, separated and apart from Choclo and Maporal viruses. [FIGURE 2 OMITTED] Conclusions HPS is an emerging disease in South America, and investigations strengthen the belief that the disease is underestimated. Despite being surrounded by HPS-endemic countries, Misiones Province had no reported HPS cases until 2003. Furthermore, HPS cases have not been documented in eastern Paraguay. Two pathogenic hantaviruses that cause HPS have so far been proven to circulate in Misiones: LEC and JUQV. We confirmed O. nigripes as the reservoir species associated with 2 JUQV cases in Misiones. In eastern Paraguay, Itapua Department, BMJ lineage produced HPS. We have already reported a BMJ case in Bolivia (7). LEC was characterized originally from an O. flavescens mouse trapped in the Rio de la Plata River La Plata River River, eastern central Puerto Rico. It flows about 45 mi (70 km) northwest and north, to empty into the Atlantic Ocean. Part of it is dammed to create a lake which provides hydroelectric power. , an area where several HPS cases had occurred (6). Species of Akodon are found throughout South America. To our knowledge, no Akodon-borne hantavirus has been reported to be associated with cases in South America. A. azarae is the most abundant sigmodontine species widely distributed in rural and peridomestic habits of central Argentina. In Buenos Aires Province Buenos Aires Province (IPA: [ˈbwenos ˈaiɾes], Spanish: Provincia de Buenos Aires) is the wealthiest and most populated province of Argentina. , PRG was characterized in A. azarae populations. Despite the absence of reported HPS cases associated with this species in the studied area, a high seroprevalence seroprevalence Immunology The proportion of a population that is seropositive–ie, has been exposed to a particular pathogen or immunogen; the seropositivity of a population is calculated as the number of individuals who produce a particular antibody divided , [approximately equal to] 10%, has been detected (14). We characterized a distinct Akodon-borne hantavirus at the Biologic Reserve of Paraguay, although we did not investigate whether this virus can produce illness. A prior study analyzed a collection of sigmodontine rodents from the major biomes of Paraguay where 1 A. montensis and 2 O. nigripes were positive for viral RNA RNA: see nucleic acid. RNA in full ribonucleic acid One of the two main types of nucleic acid (the other being DNA), which functions in cellular protein synthesis in all living cells and replaces DNA as the carrier of genetic (15). Precise identification of source populations in the reservoir and collection of quantitative data on their relative contribution to hantavirus transmission will be essential for disease control in the 3-country frontier. Acknowledgments We acknowledge the invaluable contributions of physicians and epidemiologists in data collection and continued support, which made this research possible; we particularly thank Sandra Grenon. We are also grateful to Pablo Teta and Ulf Dreschler for technical rodent field work and to Guillermo Cancros for administrative support. This research was supported in part by grant nos. DPC/ CD/078/03 and DPC/CD/133/06 from the Pan American Health Organization The Pan American Health Organization (PAHO) is an international public health agency with 100 years of experience in working to improve health and living standards of the countries of the Americas. It serves as the specialized organization for health of the Inter-American System. . References (1.) Childs JE, Ksiazek TG, Spiropoulou CF, Krebs JW, Morzunov S, Maupin GO, et al. Serologic se·rol·o·gy n. pl. se·rol·o·gies 1. The science that deals with the properties and reactions of serums, especially blood serum. 2. and genetic identification of Peromyseus maniculatus as the primary rodent reservoir for a new hantavirus in the southwestern United States. J Infect Dis. 1994; 169:1271-80. (2.) Raboni SM, Rubio G, De Borba L, Zeferino A, Skraba I, Goldenberg S, et al. Clinical survey of hantavirus in southern Brazil and the development of specific molecular diagnosis tools. Am J Trop Med Hyg. 2005;72:8004. (3.) Suzuki A, Bisordi I, Levis S, Garcia J, Pereira LE, Souza RP, et al. Identifying rodent hantavirus reservoirs, Brazil. Emerg Infect Dis. 2004;10:2127-34. (4.) Padula P J, Colavecchia SB, Martinez VP, Gonzalez Della Valle MO, Edelstein A, Miguel SD, et al. Genetic diversity, distribution, and serological serological pertaining to or emanating from serology. serological test one involving examination of blood serum usually for antibody. features of hantavirus infection in five countries in South America. J Clin Microbiol. 2000;38:3029-35. (5.) Levis S, Morzunov S, Rowe JE, Enria DA, Pini N, Calderon G. Genetic diversity and epidemiology of hantaviruses in Argentina. J Infect Dis. 1998;177:529-38. (6.) Martinez VP, Colavecchia S, Garcia Alay M, Suzuki B, Trincheri A, Busto S, et al. Hantavirus pulmonary syndrome in Buenos Aires Province [in Spanish]. Medicina (B Aires). 2001;61:147-56. (7.) Padula P, Della Valle MG, Alai MG, Cortada P, Villagra M, Gianella A. Andes virus and first case report of Bermejo virus causing fatal pulmonary syndrome. Emerg Infect Dis. 2002;8:437-9. (8.) Johnson AM, de Souza LT, Ferreira IB, Pereira LE, Ksiazek TG, Rollin PE, et al. Genetic investigation of novel hantaviruses causing fatal HPS in Brazil. J Med Virol. 1999;59:527-35. (9.) Raboni SM, Probst CM, Bordignon J, Zeferino A, dos Santos CN. Hantaviruses in Central South America: phylogenetic analysis of the S segment from HPS cases in Parana, Brazil. J Med Virol. 2005;76:553-62. (10.) Johnson AM, Bowen MD, Ksiazek TG, Williams RJ, Bryan RT, Mills JN, et al. Laguna Negra virus associated with HPS in western Paraguay and Bolivia. Virology virology, study of viruses and their role in disease. Many viruses, such as animal RNA viruses and viruses that infect bacteria, or bacteriophages, have become useful laboratory tools in genetic studies and in work on the cellular metabolic control of gene expression . 1997;238:115-27. (11.) Levis S, Garcia J, Pini N, Calderon G, Ramirez J, Bravo D, et al. Hantavirus pulmonary syndrome in northwestern Argentina: circulation of Laguna Negra virus associated with Calomys callosus. Am J Trop Med Hyg. 2004;71:658-63. (12.) Carroll DS, Mills JN, Montgomery JM, Bausch DG, Blair PJ, Burans JP, et al. Hantavirus pulmonary syndrome in Central Bolivia: relationships between reservoir hosts, habitats, and viral genotypes. Am J Trop Med Hyg. 2005;72:42-6. (13.) Padula PJ, Rossi CM, Della Valle MO, Martinez PV, Colavecchia SB, Edelstein A, et al. Development and evaluation of a solid-phase enzyme immunoassay Immunoassay An assay that quantifies antigen or antibody by immunochemical means. The antigen can be a relatively simple substance such as a drug, or a complex one such as a protein or a virus. based on Andes hantavirus recombinant nucleoprotein. J Med Microbiol. 2000;49:149-55. (14.) Suarez OV, Cueto GR, Cavia R, Gomez Villafane IE, Bilenca DN, Edelstein A, et al. Prevalence of infection with hantavirus in rodent populations of central Argentina. Mere Inst Oswaldo Cruz. 2003;98:727-32. (15.) Chu YK, Owen RD, Gonzalez LM, Jonsson CB. The complex ecology of hantavirus in Paraguay. Am J Trop Med Hyg. 2003;69:263-8. Address for correspondence: Paula Padula, Laboratorio de Hantavirus, Departamento de Virologla, INEI-ANLIS "Dr. Carlos G. Malbran," Av. Velez Sarsfield 563-C1282AFS A distributed file system for large, widely dispersed Unix and Windows networks from Transarc Corporation, now part of IBM. It is noted for its ease of administration and expandability and stems from Carnegie-Mellon's Andrew File System. AFS - Andrew File System , Buenos Aires, Argentina; email: ppadula@gmail.com Dr Padula is a virologist virologist microbiologist specializing in virology. at the Instituto Nacional de Enfermedades Infecciosas-ANLIS "Dr C. G. Malbran" and the National Cientific and Technologic Investigations Council (CONICET CONICET Consejo Nacional de Investigaciones Científicas Y Técnicas (National Council for Science and Technology, Argentina) ), Argentina. Her research interests include hantaviruses and zoonotic diseases. Paula Padula,* Valeria P. Martinez,* Carla Bellomo,* Silvina Maidana,* Jorge San Juan, ([dagger]) Paulina Tagliaferri, ([dagger]) Severino Bargardi, ([section]) Cynthia Vazquez, ([paragraph]) Norma Colucci,([paragraph]) Julio Estevez, (#) and Maria Almiron** * Instituto Nacional de Enfermedades Infecciosas ANLIS "Dr. Carlos. G. Malbran," Buenos Aires, Argentina; ([dagger]) Hospital de Enfermedades Infecciosas "Francisco J. Muniz," Buenos Aires, Argentina; ([double dagger]) Hospital de Pediatria de Posadas, Misiones, Argentina; ([section]) Universidad Nacional de Misiones Faculties
Table 1. Sigmodontine rodents captured and surveyed for antibodies
to Andes hantavirus and viral RNA, northeast Argentina and
eastern Paraguay *
Location Period Species No.
Caaguazu, Paraguay Spring 2000 Akodon sp. 1
Calomys callosus 1
Caaguazu, Paraguay Summer 2001 Holochilus 4
brasiliensis
C. callosus 1
Central Paraguay Summer 2001 Oryzomys buccinatus 2
Reserve Limoy, Winter 2001 A. cursor 7
Paraguay C. callosus 2
Caaguazu, Paraguay Spring 2001 A. cursor 8
A. nigrita 1
Reserve Limoy, Summer 2002 A. cursor 5
Paraguay
Leandro N. Alem, Autumn 2004 Oligoryzomys sp. 9
Misiones, Argentina A. cursor 3
O. nigripes 1
O. flavescens 1
H. brasiliensis 1
Santa Ana Misiones, Winter 2005 O. nigripes 28
Argentina A. montensis 8
O. flavescens 1
Leandro N. Alem, Winter 2005 O. nigripes 14
Misiones, Argentina A. montensis 3
O. flavescens 3
C.laucha 2
Location Sero- RT-PCR-
positive positive
rodents, rodents
no (%)
Caaguazu, Paraguay 0
0
Caaguazu, Paraguay 0
0
Central Paraguay 0
Reserve Limoy, 1 (14.3) 1
Paraguay 0 0
Caaguazu, Paraguay 0
0
Reserve Limoy, 0
Paraguay
Leandro N. Alem, 0
Misiones, Argentina 0
0
0
0
Santa Ana Misiones, 3 (10.7) 3
Argentina
Leandro N. Alem, 2 (14.3) 2
Misiones, Argentina 0
0
0
* RT, reverse transcription.
Table 2. Comparison of the nucleotide (first) and amino acid
(second) sequences of the 905-nucleotide region of the N gene ORF
among human and rodent hantavirus strains from South America with
viruses from Misiones, Argentina, and eastern Paraguay *
Case 1 Cases Case Akodon
2 ,3 Itapua cursor
Limoy
Case 1 82.2 91.2 74.1
Cases 2, 3 94.7 84.2 75.8
Case Itapua 99.3 94.0 74.0
Akodon cursor 86.1 85.4 85.7
Limoy
Oligoryzomys 94.3 99.7 93.7 85.0
nigripes
LEC 99.7 94.3 99.0 85.7
BMJ 100.0 94.7 99.3 86.0
AND 95.7 94.7 95.0 85.7
JUQ 94.7 100.0 94.0 85.4
LN 86.7 87.4 86.0 85.0
ANAJ 87.7 89.0 87.4 88.0
SN 85.4 84.1 85.0 84.0
O.
nigripes
LEC BMJ AND
Case 1 82.1 96.5 92.3 83.8
Cases 2, 3 98.5 81.9 83.1 82.3
Case Itapua 84.0 90.8 93.5 83.5
Akodon cursor 75.7 74.8 74.7 75.7
Limoy
Oligoryzomys 82.0 82.4 82.1
nigripes
LEC 94.0 91.7 84.8
BMJ 94.3 99.7 84.3
AND 94.3 95.4 95.7
JUQ 99.7 94.4 94.7 94.7
LN 87.0 86.4 86.7 87.7
ANAJ 88.7 87.4 87.7 89.0
SN 84.0 85.1 85.4 83.7
JUQ LN ANAJ SN
Case 1 82.5 77.6 80.1 76.4
Cases 2, 3 95.1 80.1 80.0 76.7
Case Itapua 84.9 78.7 81.0 75.9
Akodon cursor 75.9 74.7 76.9 73.9
Limoy
Oligoryzomys 95.1 79.8 80.0 76.2
nigripes
LEC 83.0 78.8 80.2 76.1
BMJ 83.5 79.0 79.9 76.8
AND 83.4 78.8 79.2 75.7
JUQ 80.6 80.7 76.4
LN 87.4 80.2 75.4
ANAJ 89.0 91.4 75.5
SN 84.1 82.4 82.4
* ORF, open reading frame; LEC, Lechiguanas; BMJ, Bermejo; AND,
Andes; JUQ, Juquitiba, LN, Laguna Negra; ANAJ, Anajatuba, SN, Sin
Nombre. An expanded online version of Table 2 is available from
www.cdc.gov/EID/content/13/8/1211-T2.htm
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