Bisgaard's taxa

Taxon 1

All 11 isolates of this taxon originally reported by Bisgaard (20)* were demonstrated in mixed flora from the intestinal tract of apparently normal ducks. These isolates were later classified with Pasteurella sensu stricto based upon DNA:DNA hybridizations (27) and named Pasteurella anatis by Mutters et al. (Int. J. Syst. Bact. 1985, 35, 309-322). P. anatis has subsequently also been obtained from ducks suffering from respiratory disease (50).

Crossed immunoelectrophoresis applied to representative strains from 11 different Pasteurella spp. showed that P. anatis formed a separate cluster branching deeply with P. langaa showing less than 70% similarity with the core group of genus Pasteurella (Schmid et al. Zbl. Bakt. 1991, 275, 16-27). A new genus, Gallibacterium gen. nov., suggested to include the avian [P.] haemolytica – [A.] salpingitidis – [P.] anatis complex was reported by Christensen et al. (119), since these taxa formed a monophyletic unit with 16S rRNA similarities above 95%. Strains originally reported as [P.]anatis are now classified as G. anatis biovar anatis. Korczak et al. (Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399) demonstrated that rpoB gene sequence analysis was useful for separating Gallibacterium spp. from other taxa of Pasteurellaceae

Taxon 2 and 3 complex

Organisms associated with salpingitis and peritonitis in ducks were first reported as atypical

Actinobacillus lignieresii by Bisgaard (4). Comparative investigations of avian Pasteurellaceae including additional isolates of atypical Actinobacillus lignieresii obtained from ducks, geese and pigeons subsequently resulted in classification of these organisms as taxa 2 and 3 based upon differences in production of acid from (+)-L-arabinose and dulcitol (20).

A total of 35 strains isolated from the respiratory tract, liver, heart and spleen of pigeons and different species of Psittaciformes were reported by Beichel (1986). Nineteen of these were reinvestigated by Bisgaard et al. (90) and biotyped as previously described (50). The host spectrum was extended to include galliform birds like partridges and pheasants (50). The importance of taxon 2 and taxon 3 in salpingitis in web-footed birds was subsequently confirmed by Bisgaard (61).

On the basis of DNA-DNA hybridizations these taxa were shown to form a large distinct group which seems to represent a new genus with several species within the family Pasteurellaceae Pohl 1981 (27). However, full matrix DNA-DNA hybridizations were not carried out.

De Ley et al. (36) performed hybridizations between labelled rRNA from seven representative members of the family Pasteurellaceae to 53 strains of Pasteurellaceae and showed that strain HIM 730-3 (F420) of taxon 2/3 (according to Bisgaard et al. (51) and not taxon 2 as listed by De Ley et al ., (36) and strain NCTC 11412 of taxon 3 biovar 2 clustered with different nodes of the rRNA branches outlined, indicating major genetic diversity within taxon 2 and 3.

Different polyamine patterns were obtained with a duck isolate of taxon 2 biovar 1 (F150 T ) and a parakeet isolate of taxon 3 biovar 1 (F450 T ) according to Busse et al. (Int. J. Syst. Bact. 1997, 47, 698-708). Both isolates had profiles different from the newly established genera Gallibacterium (119) and

Avibacterium (138) of avian origin. Comparison of phena defined by protein profiling with species/groups previously established by DNA-DNA hybridization, carbohydrate profiling and biovar typing showed that the best correlation existed between DNA-DNA hybridization and biovar typing. A correlation between results obtained from DNA-DNA hybridizations and protein profiling was not observed. Protein profiling, however, indicated a connection between protein profiles and hosts of isolation (51). Host related bacterial lineages of the taxon 2 and 3 complex have subsequently been demonstrated by amplified fragment length polymorphism (AFLP) typing (158).

Phylogenetic analysis by 16S rRNA gene sequence comparison has shown that taxon 2 biovar 1 (NCTC 11414) and taxon 3 biovar 2 (CCUG 15565 T, wrongly referred to as CCUG 15563 in the article) cluster together forming subcluster 3D of Dewhirst together with strains subsequently classified as Gallibacterium and [ Pasteurella ] langaaensis (Dewhirst et al ., Zbl. Bakt. 1993, 279, 35-44). In a subsequent investigation including avian taxa, Bisgaard taxon 2 and taxon 3 formed a separate entity within the avian cluster (127). A single isolate from septicaemia in a budgerigar classified as trehalose positive taxon 3 biovar 1 only showed 94.5% 16S rRNA gene sequence similarity to biovar 2 of taxon 3 underlining the heterogeneity of this complex group.

The accumulated evidence outlined above underlines the uncertain taxonomic position of the taxon 2 and taxon 3 complex of avian origin and the lack of unambiguous diagnostic possibilities associated with these organisms which makes interpretation of studies on these organisms difficult and prevents progress in understanding of their epidemiology and pathogenesis.

For the same reason a subset of 23 strains representing existing biovars characterized by AFLP (158) were investigated by phylogenetic analysis of the 16S rRNA, rpoB , infB and recN gene sequences. Moreover, the recN gene sequences were used for estimation of whole genome similarity to allow comparison with previously published DNA-DNA hybridizations. Selected strains were additionally characterized by polyamine profiling. Clusters outlined by

Bojesen et al. (158) and shown not to belong to Gallibacterium sensu stricto were excluded from the investigation and these taxa will be published separately. These studies allowed the identification of five groups with unique properties allowing the suggestion of five new species of Gallibacterium: G. melopsittaci, G. trehalae, G. columbinum, G. salpingitigis and a not yet named species (181). Isolates of taxon 2 and 3 sharing the properties of Volucribacter are under investigation.

Taxon 4

Isolates classified as taxon 4 were originally obtained in mixed culture from the respiratory tract of apparently healthy chickens (20). These organismes shared the cultural and biochemical characters of bacteria previously reported as Pasteurella sp. by Clark & Godfrey (Av. Dis. 1960, 4, 280-290). These organisms were later classified with Pasteurella sensu stricto based upon DNA:DNA hybridizations (27) and named Pasteurella langaa by Mutters et al. (Int. J. Syst. Bact. 1985, 35, 309-322). Crossed immunoelectrophoresis applied to representative strains from 11 different Pasteurella spp. showed that [P.] langaa formed a separate cluster branching deeply with [P.] anatis showing less than 70% similarity with the core group of genus Pasteurella (Schmid et al. Zbl. Bakt. 1991, 275, 16-27). Based upon 16S rRNA similarities Dewhirst et al. (Zbl. Bakt. 1993, 279, 35-44) showed that the type strain of P. langaa branched deeply with [P.] anatis and the taxon 2 and 3 complex Phylogenetic relationships between taxa of Pasteurellaceae based upon maximum likelihood analysis of 16S rRNA gene sequences demonstrated the highest relationship between [P.] langaa and [P.] caballi (127) while Korczak et al. (Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399) showed that [P.] langaa was phylogenetically related with [A.] rossii. However, the phylogenetic tree based upon partial rpoB sequences showed that [P.] langaa formed a separate group together with [P.] caballi, [A.] rossii and [A.] porcinus. The final taxonomic position of [P.] langaa remains to be determined.

Taxon 5, 6 7 and 8

Organisms tentatively classified as taxon 5, 6, 7 and 8 were obtained from the posterior pharynx of healthy guinea pigs originating from two different colonies of well-managed and conventionally reared guinea pigs (22). The G + C content (mol %) of DNA from these taxa were 40.0-42.1%, 51.2%, 44.1% and 39.6%, respectively. DNA:DNA hybridizations did not allow classification with known taxa. rRNA cistron similarities of taxa belonging to Pasteurellaceae showed that taxon 7 remained ungrouped on the common root of [A.] actinomycetemcomitans, [H.] aphrophilus, H. influenzae, A. lignieresii and P. multocida (36). Comparison of 16S rRNA gene sequences showed that taxon 5 and 7 were unique while taxon 6 and 8 were related with taxon 10 and Actinobacillus, respectively (129). Similar results have been obtained for rpoB (Korczak et al. Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399). Additional isolates of taxon 6 and 8 from guinea pigs misclassified as Actinobacillus sp. have been reported by Bisgaard (20) and Boot & Bisgaard (58). These taxa remains to be classified and named.

Taxon 9

Trehalose negative isolates of Actinobacillus equuli were established as a separate group tentatively named taxon 9 in 1993 (50). These organisms had been obtained from the mucous membrane of oropharynx of apparently normal horses in addition to arthritis. Additional investigations have also associated these bacteria with cases of septicaemia in a horse and foal (70, 85). Subsequent investigations of 18 isolates showed that taxon 9 represents two novel species (115), A. arthritidis and Actinobacillus genomospecies 2, both of which have been obtained from as well diseased as normal horses. The closest 16S rRNA sequence based relationships for these species were A. ureae and A. hominis, respectively. According to the phylogenetic tree of the family Pasteurellaceae based on partial rpoB sequences A. lignieresii, A. pleuropneumoniae, A. arthritidis and A. genomospecies 1 and 2 form a separate cluster with a bootstrap value of 100 (Korczak et al. Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399).

Taxon 10

Organisms tentatively classified with taxon 10 remains to be properly published. However, strains classified with this group have previously been included in other studies for comparison (127, 129, 133, 136). Taxon 10 produces gas from glucose, and has so far only been obtained from horses and horse bites in humans (Bisgaard, unpublished data). 16S rRNA gene sequence phylogeny demonstrated a close relationship with taxon 6 and [P.] aerogenes biovars 17 and 18 (136). These organisms remain to be classified and named to improve proper identification and general understanding on the importance of these bacteria.

Taxon 11

Evidence obtained to indicate that equine isolates of organisms previously reported as A. suis or haemolytic variants of A. equuli might constitute a separate taxon provisionally named taxon 11 was first published by Bisgaard et al. (25). Four biovars were initially reported, and selected DNA:DNA hybridizations did not support classification with A. suis. Additional biovars have been published, and several reports have associated these organisms with pathological lesions. However, other papers seem to point to a true predilection of taxon 11 for the respiratory tract of horses (50). Subsequent studies demonstrated that A. equuli and taxon 11 represent two different genotypes which differ with respect to disease pattern and epidemiology. For the same reasons two subspecies of A. equuli have been proposed: A. equuli subsp. e quuli (former A. equuli) and A. equuli subsp. haemolyticus (former taxon 11) (111).

RTX toxin activity (eqx) has been demonstrated in isolates of A. equuli subsp. haemolytica (Berthoud et al. Vet. Microbiol. 2002, 87, 159-174), and a stronger cytotoxicity towards horse-compared to pig lymphocytes was reported by Kuhnert et al. (Vet. Microbiol. 2003, 92, 161-167). So far, eqx has not been demonstrated in A. equuli subsp. equuli associated with sleepy foal disease and septicaemia in piglets.

Taxon 12

Several unclassified isolates of Pasteurellaceae were obtained during the investigation of the aerobic pharyngeal flora of healthy white laboratory mice originating from four different colonies. Nineteen isolates formed a group tentatively designated taxon 12. The G + C content of DNA and genome size of the reference strain were 46.9 mol % and 1.4 x 10 9 Dalton, respectively.

Taxon 12 was linked to Actinobacillus via the avian haemolytic Actinobacillus-like complex (taxon 26) by DNA:DNA hybridizations (27) and named A. muris (29). Subsequent studies excluded this taxon from Actinobacillus sensu stricto (133). Recent taxonomic investigations classified [A.] muris with the rodent cluster of Olsen et al. (Bergeys Manual of Systematic Bacteriology 2005 vol 2B p. 855). Phylogeny based on rpoB sequences, however, failed to group this taxon with other groups (Korczak et al. Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399). Further investigations are needed to clarify the taxonomy and significance of this taxon.

Taxon 13

Routine phenotypic characterization of P. multocida-like organisms obtained from pneumonic calf lungs often result in a fermentation pattern different from what is generally accepted for P.multocida sensu stricto. A group of indole-, mannitol- and sorbitol-negative isolates was tentatively designated taxon 13 (26). Selected DNA:DNA hybridizations classified ornithine positive taxon 13 isolates with P. multocida biovar 6 ( P. canis) while ornithine negative isolates were classified as V-factor independent isolates of [H.] avium. Additional DNA:DNA hybridizations demonstrated 80-92% DNA binding with P. canis for ornithine positive isolates of taxon 13 while ornithine negative isolates demonstrated 88% DNA binding with V-factor requiering isolates of [H.] avium biovar 1 classified as [P.] avium (now Avibacterium avium). Ornithine negative isolates of taxon 13 also demonstrated 69-81% DNA binding with P. stomatis (Mutters et al. Int. J. Syst. Bact. 1985, 35, 309-322). Additional information of the distribution and importance of these organisms have been published (39, 41,50).

Since routine identification of taxa belonging to Pasteurella rest on a single or a few phenotypic characters, and isolates aberrant for these characters have been reported more focus has been made on improvement of specific diagnostic genetic tools of importance for both the veterinary and medical professions. New diagnostic tools developed, however, questioned the correct classification of taxon 13 as discussed by Christensen et al. (131) who finally demonstrated that taxon 13 represents P. multocida sensu stricto, explaining positive reactions with probes (107) and in PCR- methods developed for detection of P. multocida(see 131 for references). Based upon MLST these organisms form a separate group (Unpublished data).

Taxon 14

A phenotypically homogeneous group of avian Pasteurella-like organisms including two biovars were reported by Bisgaard & Mutters (28). DNA base composition of biovars 1 and 2 varied between 53.1 (biovar 1) and 52.0 mol % (biovar 2) while both biovars demonstrated a genome mass of 1.9 x 10 9 Dalton. Ninety-seven % DNA similarity was observed between biovars 1 and 2. DNA:DNA hybridizations, however, failed to classify these organisms, tentatively named taxon 14. DNA:rRNA hybridizations, classified taxon 14 at the common root of the seven rRNA cistrons outlined for Pasteurellaceae (36). These organisms also demonstrated a separate polyamine pattern (Busse et al. Int. J. Syst. Bact. 1997, 47, 698-708). 16S rRNA gene sequence phylogeny classified taxon 14 with taxon 32 and 40, and [P.] testudinis (127, 129, 133). Similar observations have been reported for rpoB (Korczak et al. Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399).

Taxon 14 has been obtained from upper respiratory tract lesions in ducks, turkeys, pigeons, a goose and a peafowl (28, 50) and blepharoconjunctivitis in turkeys (Bisgaard et al. WPSA, 4 th Int. Symp. on Turkey Prod. 2007 pp. 59-50). These organisms still remain to be classified and named , to improve proper identification and general understanding on the epidemiology and importance of these bacteria.

Taxon 15

Reports on [P.] haemolytica (M. haemolytica) in hosts other than ruminants were questioned by Bisgaard (24). Isolates from pigs were sufficiently different from bovine isolates to constitute a new taxon, tentatively designated taxon 15. Additional investigations including mol % G + C in DNA, genome mass, and DNA:DNA hybridizations confirmed that taxon 15 constitutes a separate group within the family Pasteurellaceae (33). Based upon quantitative evaluation of phenotypic data Angen et al. (74) outlined two biovars of taxon 15 and demonstrated that taxon 15 biovar 1 and biogroup 6 of [P.] haemolytica from ruminants could not be separated by phenotypic tests. The same was valid for taxon 15 biovar 2 and biogroup 7 of [P.] haemolytica from ruminants. Subsequent genotypic studies including ribotyping and multilocus enzyme electrophoresis (MLEE) showed that [P.] haemolytica biogroup 7 did not cluster with taxon 15, and with a few exceptions, porcine isolates formed a separate cluster by ribotyping while MLEE results did not allow a clear separation between bovine and porcine isolates (78). Based upon maximum-likelihood analysis of 16S rRNA gene sequences taxon 15 and [P.] haemolytica biogroup 6 formed a separate cluster branching into a porcine and bovine line. Strains representing these lines, however, demonstrated a DNA:DNA relationship on species level. For the same reason these organisms were classified with the same species, Mannheimia varigena (87). Taxon 36 demonstrated 88 % DNA binding with M. varigena confirming the high genetic affiliation between these groups demonstrated by ribotyping (78). rpoB based phylogeny also clustered porcine and bovine isolates of M. varigena together (Korczak et al. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399).

So far M. varigena has been isolated from pneumonia, mastitis and septicaemia as well as the oral cavity, rumen and intestines of cattle. Isolates of porcine origin have been associated with septicaemia, enteritis or pneumonia, but isolates from the upper respiratory tract have also been reported (50, 87).

Taxon 16

During characterization of Pasteurella-like organisms obtained from the oral cavity of dogs and cats Bisgaard & Mutters (31) identified a homogeneous group of bacteria which might represent urease negative P. dagmatis or indole positive [P.] gallinarum ( Av. gallinarum). Mol % G + C in DNA varied between 43.5 and 44.3 % while the genome mass varied between 1.6-1.8 x 10 9 Dalton. However, DNA:DNA hybridizations did not allow classification with known species of Pasteurellaceae. Although strains investigated included both Danish and a UK isolate the authors desisted from naming the new taxon tentatively called taxon 16 (31). A total of 30 isolates were reported by Muhairwa et al. (99). Two major clusters showing approximately 35 % similarity were observed by ribotyping, the closest neighbours being P. dagmatis and P. canis. rRNA:DNA hybridizations left this taxon ungrouped at the common root of the rRNA branches of P. multocida, A. lignieresii, H. Influenzae, [H.] aphrophilus and [A.] actinomycetemcomitans (36). The 16S rRNA relationship of taxon 16 has remained ambiguous (127, 129, 133). These organisms have also been obtained from bite wounds in humans inflicted by dogs (Frederiksen, personal communication). These organisms remain to be classified and named.

Taxon 17

Organisms received as A. equuli or unclassified Pasteurellaceae have been provisionally named taxon 17 (50). This group, however, differs from A. equuli in D(+) xylose, D(-) mannitol, β-glucuronidase and ONPX. A mouse isolate (P434) received from W. Frederiksen deviating from taxon 17 in indole, trehalose and a -fucosidase demonstrated 44 % DNA binding wit P. dagmatis NCTC 11617 (Ryll: Untersuchungen zur genetischen Klassification und praktischen Identifizierung der Pasteurellaceae der Nagetiere. Thesis, Justus-Liebing-Universität, Giessen, 1989). Phylogenetic investigations as determined by comparison of 16S rRNA gene sequences showed that [P.] pneumotropica NCTC 8141 T (Jawetz type) made up a separate cluster together with [A.] muris, [H.] influenza-murium, taxon 17 and taxon 22 (127). rpoB based phylogeny confirmed the close relationship between taxon 17 and P. pneumotropica type Jawetz (Korczak et al. Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399). This group is further characterized by a deletion of five bases through E. coli rrnB pos. 204-208 compared with other members of Pasteurellaceae (Unpublished data). The significance of taxon 17 associated with mice and rats remains to be investigated.

Taxon 18

A total of 26 isolates obtained from the rumen of sheep and classified as A. lignieresii were originally received from Dr. J.E.Phillips. Further investigations showed that these isolates might represent non-haemolytic and maltose and dextrine negative [P.] haemolytica and were provisionally named taxon 18 (30). Based upon differences in D (-) sorbitol and L (+) arabinose four different biovars were outlined. An additional biovar was outlined by Angen et al. (74) who also demonstrated that these organisms clustered together with the exception of biovar 5. However, biogroup 8 D of [P.] haemolytica and biovar 2 of taxon 20 also clustered with taxon 18. Although forming a separate cluster biovar 5 of taxon 18 branched deeply with biogroup 9 of [P.] haemolytica. Ribotyping confirmed the relationship between taxon 18 biovars 1, 3 and 4 and [P.]haemolytica biogroup 8 D while other isolates of taxon 18 biovars 3 clustered separately. By MLEE a single isolate of taxon 18 biovar 1 clustered deeply with selected isolates of [P.] haemolytica biogroups 1, 8 D and 10 (78). The phylogenetic relationships of the [P.] haemolytica-complex as revealed by maximum-likelihood analysis of 16S rRNA gene sequences resulted in five major groups of which two strains of taxon 18 biovars 1 and 3 (HPA 81 and 92 T) formed a separate group with HPA 98 ([ P.] haemolytica biogroup 8 D) and UT 26 (atypical biogroup 1 of [P.] haemolytica).

Two strains of taxon 18 biovars 1 and 3 (HPA 92 T and 109) demonstrated 92 % DNA binding and clustered with HPA 98 at 88 % DNA binding. UT 26 finally showed 83 % DNA binding with HPA 98 and 84 % binding with UT 27 (biogroup 10 of [P.] haemolytica). For the same reasons these organisms were named Mannheimia ruminalis (87). Species of Mannheimia also form a separate phylogenetic group by analysis of rpoB(Korczak et al. Int. J. Syst. Evol. Micriobiol. 2004, 54, 1393-1399).

Analysis of the leukotoxin genotype by Southern blot and the corresponding β-haemolytic phenotype on sheep blood agar plates revealed that both characters were present only in strains of biogroup 1 (UT26), biogroup 190 (HPA95, HPA114, and UT27), and taxon 18 biovar 2 (HPA113, HPA90, and UT38), whereas strains of biogroup 8D and taxon 18 biovars 1, 3, and 4 have lost the leukotoxin operon.

Taxon 19

During investigations of porcine Pasteurellaceae (24) a separate group of isolates was outlined and tentatively named taxon 19. These isolates originated from septic infections in piglets and abortion in sows. However, only NCTC 10699 was included in the publication as a reference strain. Six out of 10 isolates investigated represented original isolates from Dr. N.S. Mair after whom these organisms subsequently were named (Sneath & Stevens Int. J. Syst. Bact. 1990, 40, 148-153).

rRNA:DNA hybridizations affiliated [P.] mairii at the common root of the rRNA brances of P. multocida, A. lignieresii, H. influenza and [H.] aphrophilus (36). An emended description of [P.] mairi was reported by Christensen et al. (136). Phylogenetic analysis based on 16S rRNA gene sequence comparison showed that [P.] aerogenes sensu stricto, [P.] mairii sensu stricto and [A.] seminis formed a monophyletic group representing a new genus candidate within the family Pasteurellaceae. Similar observations had previously been observed for rpoB by Korczak et al. (Int. J. Syst. Evol. Microbiol. 2004, 54, 1393-1399).

RTX toxins are predominantly observed in taxa of Pasteurellaceae and often associated with pathogenic representatives. The pax of [P.] aerogenes has been demonstrated in isolates associated with abortion in pigs, but not in those from other clinal conditions (Kuhnert et al. Infect. Immun. 2000, 68, 6-12). The pax gene has subsequently been demonstrated in all isolates of [P.] mairii investigated (155).

Taxon 20

Organisms classified with [P.] haemolytica were reported from 44 out of 81 European hares suffering from respiratory tract lesions. Similar organisms were obtained from mammary and uterine infections and from 10 out of 42 apparently asymptomatic carriers investigated (Louzis Rec. Méd. Vét. 1984, 160, 581-584). Phenotypically related organisms provisionally named taxon 20 were subsequently reported from cases of purulent bronchopneumonia and from conjunctivitis in European brown hares (40).

Two biovars of taxon 20 have been reported. Quantitative evaluation of phenotypic data clustered biovar 1 with [P.] granulomatis while biovar 2 clustered with taxon 18 (74). Ribotyping, however, clustered biovars 1 and 2 of taxon 20 together. MLEE finally classified these organisms with [ P.] granulomatis (78). Taxonomic investigations of the [Pasteurella] haemolytica-complex as evaluated by 16S rRNA sequencing and DNA:DNA hybridizations classified both isolates from ruminants and leprine species with Mannheimia granulomatis nov. rev. (87). A novel lineage of M. granulomatis has subsequently been reported from lesions in roe deer (163).

Taxon 21

These organisms have been isolated from rats and mice (50) and represent melibiose and raffinose positive [P.] pneumotropica type Jawets.

Taxon 22

Thes taxon is associated with rats and chickens (50).

Taxon 23 and 24

Organisms similar to Krause et al. (J. Appl. Bact. 1989, 67, 171-175) (50).

Taxon 25

Guinea pig isolates (50) representing a new taxon within Mannheimia

Taxon 26

Isolates from birds (20, 27, 50).

Taxon 27

Isolates from mice and rats (50).

Taxon 28

Originally received as atypical P. multocida and subsequently classified as O. rhinotracheale (Van Empel & Hafez, 1999) (53).

Taxon 29 and 30

Two taxa from rabbits not yet published.

Taxon 31

Non V –factor requiring Av. paragallinarum (46) (138).

Taxon 32

Hawk isolates (50) (127).

Taxon 33

Isolates from psittacine birds (90) classified as Volucribacter (132).

Taxon 34

Isolates from birds (127).

Taxon 35

Misclassified P. multocida from dogs not yet published.

Taxon 36

These organisms were classified with M. varigena (78, 87).

Taxon 37

Isolates from psittacine birds (90) classified as Psittacella sp. (170).

Taxon 38

Isolates from dogs not yet formally published.

Taxon 39

Bovine isolates classified with Mannheimia (104).

Taxon 40

Gull isolates (127).

Taxon 41

Isolates from rodents not yet published (Rat A/1a) (31).

Taxon 42

Porcine isolates of [P.] caballi (151).

Taxon 43

Melibiose negative and L (+) arabinose positive A. equuli. These organisms might represent Mannheimia isolates from horses not yet published.

Taxon 44

Psittacine isolates not yet published (B96/3 and B96/4 etc.).

Taxon 45 and 46

New species from large cats obtained from bite wounds in humans (140).

Taxon 47

Rat isolataes (Rat 44 sv) (118).

Taxon 48

(Ac 150 + P. pn 420). New murine taxa (200).

Taxon 49

V-factor requiring isolates from pheasants and partriges.

Taxon 50

Chicken isolates classified as Av. endocarditidis (164).

Taxon 51 and 52

(Ac 150 + P. pn 420). New murine taxa (200).

Taxon 53, 54 and 55

New avian taxa.