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Deoxyribonucleic Acid Relatedness Between Serogroups and Serovars in the Family Leptospiraceae with Proposals for Seven New Leptospira Species PAUL0 H

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Deoxyribonucleic Acid Relatedness Between Serogroups and Serovars in the Family Leptospiraceae with Proposals for Seven New Leptospira Species PAUL0 H INTERNATIONAL JOURNAL OF SYSTEMATIC BACTERIOLOGY, OCt. 1987, p. 407415 Vol. 37, No. 4 0020-7713/87/040407-09$02.00/0 Copyright 0 1987, International Union of Microbiological Societies Deoxyribonucleic Acid Relatedness between Serogroups and Serovars in the Family Leptospiraceae with Proposals for Seven New Leptospira Species PAUL0 H. YASUDA,' ARNOLD G. STEIGERWALT,* KATHERINE R. SULZER,3 ARNOLD F. KAUFMANN,4* FAYE ROGERS,3 AND DON J. BRENNER2 Department of Microbiology, Instituto de Ciencias Biomedicas da Universidade de Sao Paulo, Sao Paulo, Brazil 05508, and Molecular Biology Laboratory,2 Leptospirosis Lab~ratory,~and Bacterial Zoonoses A~tivity,~Division of Bacterial Diseases, Center for Infectious Diseases, Centers for Disease Control, Atlanta, Georgia 30333 Deoxyribonucleic acid hybridization (hydroxyapatite method, 55 and 70°C) was used to characterize 38 serovars from 22 named serogroups of Leptospira interrogans and Leptospira bijlexa, 6 serovars from 4 new unnamed serogroups of Leptospira interrogans, and single serovars of the proposed species Leptospira parva and Leptonema illini. Deoxyribonucleic acid relatedness confirmed the validity of Leptospira parva and Leptonema illini. The well-accepted species Leptospira interrogans and Leptospira bijlexa, as currently defined, were extremely heterogeneous. Relatedness results revealed at least five new species among the parasitic serovars formerly included in Leptospira interrogans and two new species among the saprophytic serovars formerly included in Leptospira bijlexa. Serogrouping did not equate with species identification, as serovars from several different subserogroups belonged to different species. The new species named in this paper are Leptospira, noguchii, Leptospira weilii, Leptospira santarosai, Leptospira borgpetersenii, Leptospira meyeri, Leptospira wolbachii, and Leptospira inadai. Leptospiraceae is a family in the Spirochaetales whose Similar to salmonellae, the serovars (serotypes) of lepto- type genus is Leptospira (7, 12, 14). The three Leptospira spires are named, and antigenically related serovars are species are Leptospira interrogans, which contains a large organized into serogroups. Leptospira parva contains a number of serogroups whose strains are parasitic or patho- single serovar (13), and Leptonema illini contains two genic for humans and animals (14); Leptospira bijlexa, which serovars (1, 12). At least 19 serogroups that contain more contains a large number of serogroups whose strains are than 170 serovars are known for Leptospira interrogans, and primarily found in fresh surface waters and moist soil and are 38 serogroups with 65 serovars have been described for the rarely isolated from humans or animals (14); and Leptospira less-studied species Leptospira bijlexa (14). parva, which was isolated from tap water, is not pathogenic In previous DNA hybridization studies (2, lo), six DNA for hamsters, and is biochemically intermediate between L. relatedness groups were identified from 15 serovars. Among interrogans and L. bijlexa (13). Leptonema, with the single the pathogenic serovars, Leptospira interrogans serovars species Leptonema illini, was proposed as a second genus in australis, bataviae, icterohaemorrhagiae, kabura, muen- the Leptospiraceae (12). Leptonema illini deoxyribonucleic chen, and pomona were in one relatedness group, and acid (DNA) has a guanine plus cytosine (G+C) content of 51 serovars javanica, tarassovi, and perhaps celledoni were in to 53 mol% (12), compared with 47.4 mol% for Leptospira a second relatedness group. Among the saprophytic sero- parva (13), 36 to 39 mol% for Leptospira bijlexa (14), and 35 vars, Leptospira bijlexa serovars andamana, patoc, and to 41 mol% for Leptospira interrogans (14). Leptonema saopaulo were in one relatedness group, serovar codice was possesses cytoplasmic tubules which are absent in in a second, serovar ranarum was in a third, and Leptonema Leptospira, and the structure of the basal complex on its illini was in a fourth. flagella resembles that of gram-positive bacteria, whereas These studies indicated that leptospires contain a great this structure in Leptospira resembles that of gram-negative deal of heterogeneity. In this study we determined the DNA bacteria (12). Leptonema strains were isolated from the relatedness of a much larger sample of leptospires in an urine of a healthy bull, a turtle, and water. They are not attempt to improve the classification of this complex and pathogenic for animals and are biochemically similar to important group of bacteria. Leptospira bijexa except for their ability to grow in Trypticase soy broth (1, 12). In addition to pathogenicity, Leptospira interrogans differs from Leptospira bijlexa by its MATERIALS AND METHODS inability to grow in the presence of 8-azaguanine and its inability to grow at 13°C (14-17). Two biogroups within Bacterial strains. A total of 45 leptospire serovars from the Leptospira interrogans differ in their ability to grow in the three previously named species of Leptospira and 1 serovar presence of 2,6-diaminopurine (14). Leptospira parva does of Leptonema were studied. These organisms represent 18 not grow in the presence of either of the above chemicals and named and 4 unnamed serogroups of Leptospira inter- does grow at 13°C (13). In general, the phenotypic charac- rogans, 4 serogroups of Leptospira bijlexa, Leptospira teristics that reportedly differentiate the leptospires have parva, and Leptonema illini. Some of the serogroups are been based on evaluations of a limited number of strains. subdivided into subserogroups based on antigenic related- ness patterns for convenience in serotyping. Serovars rep- resenting these subserogroups were included in the study. * Corresponding author. Only the reference type strains for the serovars in the study 407 408 YASUDA ET AL. INT. J. SYST.BACTERIOL. TABLE 1. Strains of Leptospiraceae used in DNA relatedness studies Serovar Serogroup Subserogroup Strain Species assignment icterohaemorrhagiae Icterohaemorrhagiae Icterohaemorrhagiae RGAT (= ATCC 43642T) Lep tospira in terrogans copenhageni M20 Leptospira interrogans smith i Smithi Smith Leptospira interrogans sarmin Sarmin Sarmin Leptosp ira weilii canicola Canicola Canicola Hond Utrecht IV Lep tospira in t e rrog a n s schueflneri Schueffneri Vleermuis 90 Leptospira interrogans ballum Ballum Mus 127 Leptospiva borgpetersenii javanica Javanica Veldrat Batavia 46T (= ATCC 43292T) Leptospira borgpetersenii celledoni Celledoni CelledoniT (= ATCC 43285T) Lebtosbiru weiiii pomona Pomona Pomona Leptospira interrogans australis Aus tralis Australis Ballico Leptospira interrogans jalna Jalna Jalna Leptospira interrogans autumnalis Autumnalis Autumnalis Akiyami A Leptospira interrogans fortbragg Fortbragg Fort Bragg Lep tospira noguc hii louisiana Louisiana LSU 1945 Leptospira noguchii pyrogenes Pyrogenes Pyrogenes Salinem Leptospira interrogans zanoni Zanoni Zanoni Leptospira interrogans grippotyphosa Grippot yphosa Moskva V Lep tospira in t errogans djasiman Djasiman Djasiman Leptospira interrogans cynopteri Cynopteri 3522 C Pending study“ panama Panama CZ 214KT (= ATCC 43288T) Lept ospira nog uchii hebdomadis Hebdomadis Hebdomadis Hebdomadis Leptospira interrogans borincana Borincana HS 622 Leptospira santarosai mini Mini Sari Leptospira borgpetersenii sejroe Sejroe Sejroe M 84 Leptospira borgpetersenii WO& Wolffi 3705 Leptospira interrogans saxkoebing Saxkoebing Mus 24 Leptospira interrogans bataviae Bataviae Van Tienen Leptospira interrogans tarassovi Tarassovi Tarassovi Perepelicin Leptospira borgpetersenii bakeri Bakeri LT 79 Leptospira santarosai atlantae Atlantae LT 81 Leptospira santarosai navet Navet TRVL 109873 Leptospira santarosai shermani Shermani LT 821T (= ATCC 43286T) Leptospira santarosai peru New unnamed 1 MWlO Leptospira santarosai banana1 Aa14 Leptospira santarosai wuwain MW6 Leptospira santarosai “AGC” New unnamed 2 AGC Leptospira interrogans “84-011370” New unnamed 3 84-011370 Leptospira noguchii lyme New unnamed 4 loT (= ATCC 43289T) Leptospiva inadai ranarum Ranarum Iowa City FrogT (= ATCC 43287T) Leptospira meyeri codice Codice CDCT (= ATCC 43284T) Leptospira wolbachii semaranga Semaranga Veldrat Semarang 173 Leptospira meyeri patoc Patoc IT (= ATCC 23582T) Leptospira bifexa andaman Andamana CH 11 Lep tospira bif exa parva NA~ HT (= NCTC 11395T) Leptospiru parva illini NA 3055T (= ATCC 43287T) Leptonema illini ‘ Study of more serovars is needed for reliable species identification on the basis of DNA relatedness. NA, Not applicable. were used (Table 1). In the remainder of this paper we refer DNA methods. A11 strains were grown at 30°C in polysor- to leptospires by serovar name only (e.g., smithi instead of bate albumin medium (23). Cells were harvested by centrif- Leptospira interrogans serovar smithi) unless specifically ugation during the late logarithmic or stationary phase of noted otherwise. The term leptospires refers to members of growth. Unlabeled DNA was isolated and purified as previ- the genus Leptospira as well as to Leptonema illini. Strains ously described (3). DNA was labeled in vitro with 32P(11). were maintained in a polysorbate albumin medium (PLM-5; DNA relatedness was determined by the hydroxyapatite Armour Pharmaceutical Co., Kankakee, Ill.). The identity of method (3), except that, because of the low G+C contents of each strain was verified by our leptospirosis laboratory (a the DNAs of strains, hybridization reactions were done at an Food and Agricultural Organization/World Health Organiza-
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