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Expansion of the Candida tanzawaensis yeast clade: 16 novel Candida species from basidiocarp-feeding beetles

¹ Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, USA
² Department of Entomology, University of Georgia, Athens, GA 30602, USA

Correspondence
Sung-Oui Suh
ssuh{at}lsu.edu

	ABSTRACT

TOP ABSTRACT INTRODUCTION METHODS RESULTS AND DISCUSSION REFERENCES

 
A major clade of new yeast taxa from the digestive tract of basidiocarp-feeding beetles is recognized based on rRNA gene sequence analyses. Almost 30 % of 650 gut isolates formed a statistically well-supported clade that included Candida tanzawaensis. The yeasts in the clade were isolated from 11 families of beetles, of which Tenebrionidae and Erotylidae were most commonly sampled. Repeated isolation of certain yeasts from the same beetle species at different times and places indicated strong host associations. Sexual reproduction was never observed in the yeasts. Based on comparisons of small- and large-subunit rRNA gene sequences and morphological and physiological traits, the yeasts were placed in Candida ambrosiae and in 16 other undescribed taxa. In this report, the novel species in the genus Candida are described and their relationships with other taxa in the Saccharomycetes are discussed. The novel species and their type strains are as follows: Candida guaymorum (NRRL Y-27568^T=CBS 9823^T), Candida bokatorum (NRRL Y-27571^T=CBS 9824^T), Candida kunorum (NRRL Y-27580^T=CBS 9825^T), Candida terraborum (NRRL Y-27573^T=CBS 9826^T), Candida emberorum (NRRL Y-27606^T=CBS 9827^T), Candida wounanorum (NRRL Y-27574^T=CBS 9828^T), Candida yuchorum (NRRL Y-27569^T=CBS 9829^T), Candida chickasaworum (NRRL Y-27566^T=CBS 9830^T), Candida choctaworum (NRRL Y-27584^T=CBS 9831^T), Candida bolitotheri (NRRL Y-27587^T=CBS 9832^T), Candida atakaporum (NRRL Y-27570^T=CBS 9833^T), Candida panamericana (NRRL Y-27567^T=CBS 9834^T), Candida bribrorum (NRRL Y-27572^T=CBS 9835^T), Candida maxii (NRRL Y-27588^T=CBS 9836^T), Candida anneliseae (NRRL Y-27563^T=CBS 9837^T) and Candida taliae (NRRL Y-27589^T=CBS 9838^T).

Published online ahead of print on 28 June 2004 as DOI 10.1099/ijs.0.63246-0.

The GenBank/EMBL/DDBJ accession numbers for the sequences determined in this study are listed in Table 1 and Fig. 1.

	INTRODUCTION

TOP ABSTRACT INTRODUCTION METHODS RESULTS AND DISCUSSION REFERENCES

 
Yeasts and yeast-like endosymbionts have been reported from a variety of insects, including planthoppers, aphids and beetles (e.g. Nardon & Grenier, 1989; Noda & Omura, 1992; Suh et al., 2003, 2004). Although some endosymbionts from anobiid beetles and planthoppers show an affinity to certain filamentous ascomycetes (Jones & Blackwell, 1996; Noda et al., 1995; Noda & Kodama, 1996; Suh et al., 2001), the majority of the fungal endosymbionts have been identified as true yeasts (Ascomycetes: Saccharomycetes) (Jones et al., 1999). We were interested in sampling insects more broadly in order to determine if associations between additional fungi and insect hosts might be discovered. During a study of fungi from the digestive tract of basidiocarp-feeding beetles, about 650 yeasts were isolated from beetles in 26 families (Suh & Blackwell, 2004). Based on sequence comparisons of the D1/D2 loop of the large subunit (LSU) rRNA gene used for rapid yeast characterization, about 30 % of the yeast isolates formed a clade (CT clade) with Candida tanzawaensis, a yeast isolated from mosses in Japan (Nakase et al., 1988). Until the relatively recent report of six novel species (Candida ambrosiae, Candida canberraensis, Candida caryicola, Candida prunicola, Candida pyralidae and Candida xylopsoci), C. tanzawaensis had no known close relatives (Kurtzman, 2001). Currently, only one of the seven previously described species is known from multiple collections; in fact, C. tanzawaensis awaited description for 22 years in the vain hope that another isolate would be discovered (Nakase et al., 1988). Here, we describe 16 novel taxa, many with multiple isolates, from the gut of fungus-feeding beetles and compare the characteristics of all 23 members of the large CT clade, thereby recognizing unsuspected diversity in what appears to be a largely insect-associated clade.

	METHODS

TOP ABSTRACT INTRODUCTION METHODS RESULTS AND DISCUSSION REFERENCES

 
Yeast isolation and identification.
Host beetles were collected from several different localities in Vermont, the southeastern USA, and Barro Colorado Island, Panama. Vouchers were deposited at the University of Georgia Collection of Arthropods, Athens, GA. The beetles were taken to the laboratory and usually placed in Petri dishes for 1–3 days without food prior to dissection. Surface disinfection was by submersion in 95 % ethanol for 1–2 min. The alcohol wash was followed by a 0·7 % saline rinse; the rinse liquid was plated on acidified YM agar (Difco YM broth, 2 % plain agar, adjusted to pH 3·5 with HCl) as a negative control. The beetle gut was removed aseptically under a dissecting microscope and transferred to tubes containing 0·7 % saline. The gut segments were crushed in the saline solution with a pipette tip and streaked with a loop onto the surface of an acidified YM agar plate. Plates were incubated at 25 °C, and single colonies were streaked for purification. Cultures were maintained on YM agar or 2 % malt extract agar (Yarrow, 1998). The isolates were initially screened and grouped using the D1/D2 loop sequence of the LSU rRNA gene. Selected isolates from each LSU genotype have been deposited at the Agricultural Research Service Culture Collection (NRRL) and Centraalbureau voor Schimmelcultures (CBS); holotype specimens were deposited at NRRL as lyophilized cultures (Table 1). The morphological observations and metabolic tests comprising the yeast standard description were performed on at least one isolate from each LSU genotype, according to established methods (Yarrow, 1998; Barnett et al., 2000). Isolates within each clade were crossed in all combinations and observed on YM agar, 2 % malt agar and cornmeal agar at 17 °C for 6 weeks in order to detect ascospore development. Bright-field microscopic examination was the usual method of observation, but some cell preparations were examined more closely with fluorescence microscopy using 0·02 % Calcofluor white as an optical brightener to visualize yeast cell walls more clearly.

View larger version (83K): [in this window] [in a new window]   Fig. 1. Consensus of two most-parsimonious trees obtained from combined SSU and LSU rRNA gene sequence data. Candida rhagii was chosen as the outgroup taxon based on analyses including a broader range of taxa (not shown). GenBank/EMBL/DDBJ accession numbers after the names of yeast species are for SSU and LSU rRNA gene sequence, respectively. Tree length=514; consistency index=0·6381; homoplasy index=0·3619; retention index=0·8444; rescaled consistency index=0·5388. Numbers on tree branches indicate the percentages of bootstrap samplings derived from 1000 samples that supported the internal branches by 50 % or higher.

	RESULTS AND DISCUSSION

TOP ABSTRACT INTRODUCTION METHODS RESULTS AND DISCUSSION REFERENCES

Novel species in the CT clade
In addition to the D1/D2 region of the LSU rRNA gene sequences, complete sequences of the SSU rRNA gene (about 1750 bp) were determined for at least one isolate from each LSU genotype (see Table 1 for selected isolates), and these were combined with the LSU rRNA gene sequences in a dataset for better estimation of a phylogeny. A most-parsimonious tree was constructed from the combined dataset of SSU and LSU rRNA gene sequences by comparing the yeasts with taxa in Ascomycota and other yeasts alone (Fig. 1). The 39 gut yeasts we compared were representatives of the total 164 isolates in the clade (Fig. 1) with C. tanzawaensis and the other six previously described clade members. The clade was well supported statistically by 100 % bootstrap value (Fig. 1). Of the seven previously described taxa, C. xylopsoci, C. pyralidae, C. prunicola and C. caryicola were basal to the other yeasts in the CT clade. C. ambrosiae, C. tanzawaensis and C. canberraensis, however, were included among the beetle gut yeasts (Fig. 1). Within the CT clade, the beetle isolates were resolved into 17 subclades with sufficient divergence to warrant species-level recognition (Kurtzman & Robnett, 1998; Kurtzman, 2000; Suh & Blackwell, 2004) (Fig. 1).

As mentioned earlier, only one subclade (including Erot8, Niti14 and Tene7) had D1/D2 sequences identical to the previously described yeast C. ambrosiae, which we consider to be conspecific with these isolates. There were marked differences between C. ambrosiae and the other 16 subclades in the CT clade with 6 bp or more (usually more than 20 bp) difference between the subclades. The genetic variation among multiple isolates within subclades was lower (Table 1; Fig. 1). For example, yeasts designated Endo2 and Erot24 occurred in a common subclade and differed from each other by 3 bp of D1/D2 sequence. Also, there were minor D1/D2 sequence differences between common subclade members Tene28 and Tene30 or Erot20 and Tene19 (Table 1). Within all 17 subclades of the CT clade, however, the sequence variation of D1/D2 among subclade members was always less than 3 bp, within the range of species-level variation recognized in studies of other ascomycete yeast species (Kurtzman & Robnett, 1998; Table 1).

We observed minor morphological differences among the isolates on YM and cornmeal agars at 5–7 days incubation and in YM broth after 5 days incubation at 25 °C, but morphological variation that consistently distinguished the yeasts within clades was not evident. One morphological feature common to all isolates was the lack of ascospore production. Physiological characters (Table 2), however, were more variable and these traits were useful to separate the other species of the CT clade. Below, we characterize and describe 16 novel species of beetle gut yeasts and compare them to the previously described yeasts of the CT clade (Fig. 1 and Table 2).

Description of Candida guaymorum Suh & Blackwell sp. nov.
Candida guaymorum (gu.ay.mo'rum. N.L. m. gen. guaymorum to commemorate the Guaymí, a group of indigenous people of Panama with detailed knowledge of the forest flora).

After 7 days growth in YM broth at 25 °C, cells are globose to ellipsoidal (1·25–6·25x1·25–7·5 µm), and occur singly, in pairs or in short chains (Fig. 2a). Pseudohyphae are present. After 7 days on YM agar at 25 °C, colonies are white to cream-coloured with pale-pinkish perimeter on some old colonies, smooth, shiny, flat and filamentous in margin. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae are present; septate hyphae are absent. Aerobic growth is white, shiny and smooth with filamentous margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

View larger version (101K): [in this window] [in a new window]   Fig. 2. Budding yeast cells, pseudohyphae and septate hyphae of the novel species. (a) C. guaymorum NRRL Y-27568T; (b) C. bokatorum NRRL Y-27571T; (c) C. kunorum NRRL Y-27580T; (d) C. terraborum NRRL Y-27573T; (e) C. emberorum NRRL Y-27606T; (f) C. wounanorum NRRL Y-27574T; (g) C. yuchorum NRRL Y-27569T; (h) C. chickasaworum NRRL Y-27566T; (i) C. choctaworum NRRL Y-27584T; (j) Candida bolitotheri NRRL Y-27587T; (k) C. atakaporum NRRL Y-27570T; (l) C. panamericana NRRL Y-27567T. (a–f, h–l) Seven days, half-strength cornmeal agar, 25 °C; (g) 5 days, YM broth, 25 °C. Bars, 5 µm.

Latin diagnosis of Candida bokatorum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ellipsoideae (2–6x3–6 µm), singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, hebes, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae fiunt; hyphae verae fiunt. Ascosporae non fiunt. Glucosum, maltosum (infirme, variabilitre), sucrosum (infirme, variabilitre), trehalosum et cellobiosum fermentantur. Galactosum, methyl -D-glucosidum, melibiosum, lactosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (variabilitre), D-xylosum, D-arabinosum, sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum, glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (lente, infirme), acidum succinicum, acidum citricum, ethanolum et propane-1,2-diolum (infirme, variabilitre). Non assimilantur L-sorbosum, D-ribosum, L-arabinosum, L-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, xylitolum, L-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, L-lysinum, cadaverinum et glucosaminum (infirme, variabilitre). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 40 °C. Variabilitre in medio 10 µg ml^–1 cycloheximido addito, non crescit in medio 100 µg ml^–1. Typus: NRRL Y-27571^T (=CBS 9824^T), designat stirpem typicum. Isolata a ile coleopterorum (Pselaphacus signatus; Erotylidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida bokatorum Suh & Blackwell sp. nov.
Candida bokatorum (bo.ka.to'rum. N.L. m. gen. bokatorum to commemorate the Bókatá, a group of indigenous people of Panama, linguistically related to the Guaymí).

After 7 days growth in YM broth at 25 °C, cells are globose to ellipsoidal (2–6x3–6 µm), mostly subglobose, and occur singly, in pairs or in short chains (Fig. 2b). Pseudohyphae may be present. After 7 days on YM agar at 25 °C, colonies are white to cream in colour, butyrous and smooth. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae are present. Septate hyphae may be present. Aerobic growth is white and smooth. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27571^T (=CBS 9824^T).

Latin diagnosis of Candida kunorum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae subglobosae aut fusiformes (2–5x3–6 µm), singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, hebes, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae non fiunt; hyphae verae non fiunt. Ascosporae non fiunt. Glucosum, galactosum (infirme), maltosum (infirme), trehalosum et lactosum (infirme) fermentantur. Sucrosum, methyl -D-glucosidum, melibiosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (lente), D-ribosum (infirme), D-xylosum (lente), D-arabinosum (infirme), sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum (lente), arbutinum, melezitosum, glycerolum, ribitolum, xylitolum (infirme), L-arabinitolum (infirme), D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum, acidum succinicum, acidum citricum (lente), ethanolum et propane-1,2-diolum (infirme). Non assimilantur L-sorbosum, L-arabinosum, L-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, butano-2,3-diolum, acidum quinicum et D-glucaratum. Assimilantur ethylaminum, L-lysinum, cadaverinum et glucosaminum (infirme). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 35 °C. Typus: NRRL Y-27580^T (=CBS 9825^T), designat stirpem typicum. Isolata a ile coleopterorum (Teichostethus testaceous; Nitidulidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida kunorum Suh & Blackwell sp. nov.
Candida kunorum (ku.no'rum. N.L. m. gen. kunorum to commemorate the Kuna, a group of indigenous people of Panama with a high degree of literacy and a desire for autonomy).

After 7 days growth in YM broth at 25 °C, cells are subglobose to fusiform (2–5x3–6 µm), mostly subglobose, and occur singly, in pairs or in short chains (Fig. 2c). After 7 days on YM agar at 25 °C, colonies are white, membranous and butyrous with smooth margin. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae are absent. Aerobic growth is white and smooth. No ascospores produced after 6 weeks at 17 °C from the single strain on YM agar or half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27580^T (=CBS 9825^T).

Latin diagnosis of Candida terraborum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ellipsoideae (4–6x5–7 µm), singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, hebes, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae fiunt; hyphae verae non fiunt. Ascosporae non fiunt. Glucosum (lente), galactosum (infirme), maltosum (lente), sucrosum (lente) et trehalosum (lente) fermentantur. Methyl -D-glucosidum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (infirme), D-xylosum (infirme), sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum, glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum (infirme), 2-keto-D-gluconatum, D-gluconatum (infirme), acidum succinicum, acidum citricum, ethanolum (lente) et propane-1,2-diolum (infirme). Non assimilantur D-sorbosum, D-ribosum, L-arabinosum, D-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, xylitolum, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum (infirme). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. 30 °C crescit neque 35 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27573^T (=CBS 9826^T), designat stirpem typicum. Isolata a ile coleopterorum (Iphiclus sedecimmaculatus; Erotylidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida terraborum Suh & Blackwell sp. nov.
Candida terraborum (ter.ra.bo'rum. N.L. m. gen. terraborum to commemorate the Térraba, a group of indigenous people of Panama, survivors of epidemics that once decimated their numbers).

After 7 days growth in YM broth at 25 °C, cells are globose to ellipsoidal (4–6x5–7 µm), mostly subglobose, and occur singly, in pairs or in short chains (Fig. 2d). Big and elongated cells are also observed. After 7 days on YM agar at 25 °C, colonies are off-white, butyrous and smooth. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae are present; septate hyphae are absent. Aerobic growth is white with fuzzy margin. No ascospores produced after 6 weeks at 17 °C from the single strain on YM agar or half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27573^T (=CBS 9826^T).

Latin diagnosis of Candida emberorum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ellipsoideae (1·25–5x2·5–6·25 µm), singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, hebes, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae fiunt; hyphae verae non fiunt. Ascosporae non fiunt. Glucosum, galactosum, trehalosum et cellobiosum (lente, infirme) fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (lente, infirme), D-xylosum, sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum (lente, infirme), glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (lente, infirme), acidum succinicum, acidum citricum et ethanolum. Non assimilantur D-sorbosum, D-ribosum, L-arabinosum, D-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, xylitolum, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, propane-1,2-diolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum, glucosaminum et D-tryptophanum (infirme). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum et imidazolum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 40 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27606^T (=CBS 9827^T), designat stirpem typicum. Isolata a ile coleopterorum (Triplax alvarengai; Erotylidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida emberorum Suh & Blackwell sp. nov.
Candida emberorum (em.be.ro'rum. N.L. m. gen. emberozrum to commemorate the Emberá, a group of indigenous people of Panama, knowledgeable in botanical lore).

After 7 days growth in YM broth at 25 °C, cells are globose to ellipsoidal (1·25–5x2·5–6·25 µm), mostly globose or subglobose, and occur singly, in pairs or in short chains (Fig. 2e). Pseudohyphae may be present. After 7 days on YM agar at 25 °C, colonies are white to cream-coloured with pale-pinkish perimeter, smooth and butyrous. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae are present; septate hyphae are absent. Aerobic growth is white, dull and smooth with filamentous margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27606^T (=CBS 9827^T).

Latin diagnosis of Candida wounanorum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ovoidae (2–4x3–6 µm), plerumque subglobosae, singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, butyrosa. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae non fiunt; hyphae verae non fiunt. Ascosporae non fiunt. Glucosum (lente), galactosum (infirme) et trehalosum (lente) fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (lente), D-xylosum, sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum (infirme), glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, acidum succinicum (lente), acidum citricum et ethanolum. Non assimilantur D-sorbosum, D-ribosum, L-arabinosum, D-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, xylitolum, D-arabinitolum, galactitolum, inositolum, D-gluconatum, D-glucuronatum, DL-acidum lacticum, methanolum, propane-1,2-diolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum et cadaverinum. Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, glucosaminum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. In temperatura 30 °C crescit neque 35 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27574^T (=CBS 9828^T), designat stirpem typicum. Isolata a ile coleopterorum (Mycotretus dorsonotatus; Erotylidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida wounanorum Suh & Blackwell sp. nov.
Candida wounanorum (wou.na.no'rum. N.L. m. gen. wounanorum to commemorate the Wounan, a group of indigenous people of Panama known for their artistry in using plant materials; they inhabited the same region as the Emberá).

After 7 days growth in YM broth at 25 °C, cells are globose to oval (2–4x3–6 µm), mostly subglobose, and occur singly, in pairs or in short chains (Fig. 2f). After 7 days on YM agar at 25 °C, colonies are off-white with fuzzy white spots, butyrous and smooth. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae are absent. Aerobic growth is white, shiny, smooth and filamentous in margin. No ascospores produced after 6 weeks at 17 °C from the single strain on YM agar or half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27574^T (=CBS 9828^T).

Latin diagnosis of Candida yuchorum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ellipsoideae (2·5–7x2·5–7 µm), plerumque globosae, singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, teres, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae et hyphae verae fiunt. Ascosporae non fiunt. Glucosum, galactosum (lente), trehalosum et cellobiosum (lente) fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-xylosum, sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (lente), acidum succinicum et acidum citricum. Non assimilantur D-sorbosum, D-glucosaminum, D-ribosum, L-arabinosum, D-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, melezitosum, inulinum, amylum solubile, erythritolum, xylitolum, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, ethanolum, propane-1,2-diolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum, glucosaminum et D-tryptophanum (lente). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum et imidazolum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 40 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27569^T (=CBS 9829^T), designat stirpem typicum. Isolata a ile coleopterorum (Tritoma atriventris; Erotylidae), Athens, GA, USA, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida yuchorum Suh & Blackwell sp. nov.
Candida yuchorum (yu.cho'rum. N.L. m. gen. yuchorum to commemorate the Yuchi, native Americans of the southeastern USA; their public worship was tied to the corn harvest when food was shared).

After 7 days growth in YM broth at 25 °C, cells are globose to ellipsoidal (2·5–7x2·5–7 µm), mostly globose or subglobose, and occur singly, in pairs or in short chains (Fig. 2g). Pseudohyphae and septate hyphae may be present. After 7 days on YM agar at 25 °C, colonies are cream-coloured, smooth and shiny with filamentous margin. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae may be present. Aerobic growth is white and dull. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27569^T (=CBS 9829^T).

Latin diagnosis of Candida chickasaworum Suh & Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut fusiformes (2–6x2–8 µm), singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae et hyphae verae fiunt. Ascosporae non fiunt. Glucosum, galactosum et trehalosum fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (lente, infirme), D-xylosum, sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum (variabilitre), glycerolum, ribitolum, xylitolum (infirme, variabilitre), D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (variabilitre), acidum succinicum, acidum citricum, ethanolum et propane-1,2-diolum (infirme, variabilitre). Non assimilantur D-sorbosum, D-ribosum, L-arabinosum, D-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum (variabilitre). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est (variabilitre). In temperatura 30 °C crescit neque 35 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27566^T (=CBS 9830^T), designat stirpem typicum. Isolata a ile coleopterorum (Tritoma sp.; Erotylidae), Athens, GA, USA, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida chickasaworum Suh & Blackwell sp. nov.
Candida chickasaworum (chic.ka.sa.wo'rum. N.L. m. gen. chickasaworum to commemorate the Chickasaw, native Americans who in earlier times made use of many endemic and introduced southeastern USA plants).

After 7 days growth in YM broth at 25 °C, cells are globose, subglobose, to fusiform (2–6x2–8 µm), and occur singly, in pairs or in short chains (Fig. 2h). Pseudohyphae and septate hyphae may be present. After 7 days on YM agar at 25 °C, colonies are white to cream-coloured, butyrous and smooth with mycelial edge. After 10 days Dalmau plate culture on corn meal agar at 25 °C, elongated pseudohyphae and septate hyphae may be present. Aerobic growth is white to cream-coloured with mycelial margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27566^T (=CBS 9830^T).

Latin diagnosis of Candida choctaworum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ovoidae (2–6x2–8 µm), plerumque globosae et subglobosae, singulae vel binae. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, butyrosa, teres. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae et hyphae verae may fiunt. Ascosporae non fiunt. Glucosum, galactosum (variabilitre) et trehalosum fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-sorbosum, D-glucosaminum, D-ribosum (variabilitre), D-xylosum, L-arabinosum (infirme, variabilitre), D-arabinosum, trehalosum, cellobiosum, salicinum, arbutinum, glycerolum, erythritolum, ribitolum, xylitolum (variabilitre), D-arabinitolum (variabilitre), D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (variabilitre), acidum succinicum, acidum citricum, ethanolum et propane-1,2-diolum (infirme, variabilitre). Non assimilantur D-rhamnosum, sucrosum, maltosum, methyl -D-glucosidum, melibiosum, lactosum, raffinosum, melezitosum, inulinum, amylum solubile, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum (variabilitre). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 40 °C. Crescit in medio 100 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27584^T (=CBS 9831^T), designat stirpem typicum. Isolata a ile coleopterorum (Neomida bicornis; Tenebrionidae), Baton Rouge, LA, USA, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida choctaworum Suh & Blackwell sp. nov.
Candida choctaworum (choc.ta.wo'rum. N.L. m. gen. choctaworum to commemorate the Choctaw, native Americans warriors and code-talkers, many of whom were removed from their native lands in the southeastern USA).

After 7 days growth in YM broth at 25 °C, cells are globose to oval (2–6x2–8 µm), and occur singly, in pairs or in short chains (Fig. 2i). Pseudohyphae may be present. After 7 days on YM agar at 25 °C, colonies are white to cream-coloured, smooth and butyrous with slightly wrinkled edge. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae may be present. Aerobic growth is white, shiny, smooth and slightly fuzzy. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27584^T (=CBS 9831^T).

Latin diagnosis of Candida bolitotheri Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut subglobosae (2–6x2–6 µm), singulae vel binae. Pseudohyphae fiunt. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, teres, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae fiunt; hyphae verae fiunt. Ascosporae non fiunt. Glucosum, galactosum et trehalosum fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-sorbosum (variabilitre), D-glucosaminum, D-ribosum (variabilitre), D-xylosum, trehalosum, cellobiosum, salicinum, arbutinum, glycerolum, erythritolum, ribitolum, xylitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (lente, infirme), acidum succinicum, acidum citricum et ethanolum. Non assimilantur L-arabinosum, D-arabinosum, D-rhamnosum, sucrosum, maltosum, methyl -D-glucosidum, melibiosum, lactosum, raffinosum, melezitosum, inulinum, amylum solubile, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, propane-1,2-diolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum. Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 45 °C. Variabilitre in medio 10 µg ml^–1 cycloheximido addito, non crescit in medio 100 µg ml^–1. Typus: NRRL Y-27587^T (=CBS 9832^T), designat stirpem typicum. Isolata a ile coleopterorum (Bolitotherus cornutus; Tenebrionidae), Athens, GA, USA, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida bolitotheri Suh & Blackwell sp. nov.
Candida bolitotheri (bo.li.to.the'ri. N.L. n. gen. bolitotheri is named for the coleopteran host, Bolitotherus cornutus, occurring in Ganoderma basidiocarps from Vermont to Louisiana, USA).

After 7 days growth in YM broth at 25 °C, cells are globose to subglobose (2–6x2–6 µm), mostly globose, and occur singly or in short chains (Fig. 2j). Pseudohyphae may be present. After 7 days on YM agar at 25 °C, colonies are white to cream-coloured with very pale-pinkish from centre to edge, and smooth to slightly wrinkled centre with mycelial edge. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae are present. Septate hyphae may be present. Aerobic growth is white, shiny and smooth with filamentous margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27587^T (=CBS 9832^T).

Latin diagnosis of Candida atakaporum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae subglobosae aut cylindratae (3–5x5–9 µm), singulae vel binae. Pseudohyphae fiunt. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae et hyphae verae fiunt. Ascosporae non fiunt. Glucosum et trehalosum (lente) fermentantur. Galactosum, maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (lente), D-xylosum, D-arabinosum (infirme), sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum, amylum solubile (infirme), glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum (infirme), DL-acidum lacticum (lente), acidum succinicum (lente), acidum citricum (lente), ethanolum et propane-1,2-diolum (infirme). Non assimilantur D-sorbosum, D-ribosum, L-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, erythritolum, xylitolum, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, methanolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum, glucosaminum (infirme) et D-tryptophanum (lente). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum et imidazolum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 40 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27570^T (=CBS 9833^T), designat stirpem typicum. Isolata a ile coleopterorum (Triplax festiva; Erotylidae), Baton Rouge, LA, USA, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida atakaporum Suh & Blackwell sp. nov.
Candida atakaporum (a.ta.ka.po'rum. N.L. m. gen. atakaporum commemorates the Atakapa, native Americans of the northwestern coast of the Gulf of Mexico and their unique language).

After 7 days growth in YM broth at 25 °C, cells are subglobose, ellipsoidal or cylindrical (3–5x5–9 µm), and occur singly, in pairs or in short chains (Fig. 2k). Pseudohyphae and septate hyphae may be present. After 7 days on YM agar at 25 °C, colonies are off-white with membranous margin and butyrous with small filamentous spots. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae are present. Aerobic growth is off-white and fuzzy. No ascospores produced after 6 weeks at 17 °C from the single strain on YM agar or half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27570^T (=CBS 9833^T).

Latin diagnosis of Candida panamericana Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ovoidae (3–6·25x3·75–7 µm), plerumque subglobosae, singulae vel binae. Pseudohyphae fiunt. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, teres, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae fiunt; hyphae verae fiunt. Ascosporae non fiunt. Glucosum, galactosum (lente), trehalosum et cellobiosum fermentantur. Maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum, D-xylosum, sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum, glycerolum, ribitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum, acidum succinicum et acidum citricum. Non assimilantur D-sorbosum, D-ribosum, L-arabinosum, D-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, erythritolum, xylitolum, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, ethanolum, propane-1,2-diolum, butano-2,3-diolum, acidum quinicum et D-glucaratum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum (variabilitre). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Vitaminae externae ad crescentiam necessaria non sunt. Augmentum non fiunt in temperatura 40 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27567^T (=CBS 9834^T), designat stirpem typicum. Isolata a ile coleopterorum (Mycotretus interstitialis; Erotylidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida panamericana Suh & Blackwell sp. nov.
Candida panamericana (pan.a.mer.i.can'a. N.L. f. adj. panamericana to call attention to its broad distribution spanning the regions from Louisiana to Panama).

After 7 days growth in YM broth at 25 °C, cells are globose to oval (3–6·25x3·75–7 µm), mostly subglobose, and occur singly, in pairs or in short chains (Fig. 2l). Some cells form clusters. Pseudohyphae are present. After 7 days on YM agar at 25 °C, colonies are white to off-white in colour, smooth, glistening and flat with filamentous margin. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae are present. Septate hyphae may be present. Aerobic growth is white, shiny and smooth with filamentous margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

The type strain is NRRL Y-27567^T (=CBS 9834^T).

Latin diagnosis of Candida bribrorum Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut ellipsoideae (3–8x3–8 µm), plerumque subglobosae, singulae vel binae. Pseudohyphae fiunt. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, butyrosa, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae et hyphae verae fiunt. Ascosporae non fiunt. Glucosum, galactosum (variabilitre), maltosum (infirme, variabilitre) et trehalosum fermentantur. Methyl -D-glucosidum, sucrosum, melibiosum, lactosum, cellobiosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-sorbosum (variabilitre), D-glucosaminum, D-ribosum (variabilitre), D-xylosum, L-arabinosum (variabilitre), D-arabinosum (variabilitre), sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum, glycerolum, erythritolum, ribitolum, xylitolum, D-glucitolum, D-mannitolum, gluconolactonum, 2-keto-D-gluconatum, D-gluconatum, acidum succinicum, acidum citricum, ethanolum et propane-1,2-diolum (variabilitre). Non assimilantur D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum (variabilitre). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. Augmentum non fiunt in temperatura 40 °C. Infirme crescit in medio 10 µg ml^–1 cycloheximido addito, variabilitre in 100 µg ml^–1. Typus: NRRL Y-27572^T (=CBS 9835^T), designat stirpem typicum. Isolata a ile coleopterorum (Pselaphacus sp.; Erotylidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida bribrorum Suh & Blackwell sp. nov.
Candida bribrorum (bri.bro'rum. N.L. m. gen. bribrorum to commemorate the Bribri, a small group of indigenous people of northern Panama, who apply ecological principles and recognize symbiotic relationships).

After 7 days growth in YM broth at 25 °C, cells are globose to ellipsoidal (3–8x3–8 µm), mostly subglobose, and occur singly, in pairs or in short chains (Fig. 3a). Pseudohyphae are present. After 7 days on YM agar at 25 °C, colonies are white to cream in colour, butyrous and smooth surface with slightly fuzzy spots. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae are present. Aerobic growth is white to cream-coloured with fuzzy margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on half-strength cornmeal agar. See Table 2 for a summary of physiological and other characteristics.

View larger version (127K): [in this window] [in a new window]   Fig. 3. Budding yeast cells, pseudohyphae and septate hyphae of the novel species. (a) C. bribrorum NRRL Y-27572T; (b, c) C. maxii NRRL Y-27588T; (d) yeast cells of C. anneliseae NRRL Y-27563T; (e) C. taliae NRRL Y-27589T. (a–c) Seven days, YM broth, 25 °C; (d, e) 7 days, half-strength cornmeal agar, 25 °C. Bars, 5 µm.

Latin diagnosis of Candida maxii Suh et Blackwell sp. nov.
In medio liquido dextrosum et peptonum et extractum levidinis continente post 7 dies ad 25 °C cellulae vegetativae globosae aut subglobosae (2–6·25x3·75–6·25 µm), singulae vel binae. Pseudohyphae et hyphae verae fiunt. Cultura in agaro extramalti et faecis continente post 7 dies ad 25 °C, albida, margine ciliata. In agaro farina Zeae maydis confecto post 10 dies ad 25 °C, pseudohyphae et hyphae verae fiunt. Ascosporae non fiunt. Glucosum (lente), trehalosum (infirme) et cellobiosum (infirme) fermentantur. Galactosum, maltosum, methyl -D-glucosidum, sucrosum, melibiosum, lactosum, melezitosum, raffinosum, inulinum, amylum solubile et D-xylosum non fermentantur. Assimilantur glucosum, galactosum, D-glucosaminum (lente), D-ribosum, D-arabinosum (infirme), sucrosum, maltosum, trehalosum, methyl -D-glucosidum, cellobiosum, salicinum, arbutinum, melezitosum, glycerolum (lente), erythritolum, ribitolum, xylitolum (infirme), D-glucitolum, D-mannitolum, gluconolactonum (lente), 2-keto-D-gluconatum, D-gluconatum, acidum succinicum, acidum citricum et ethanolum. Non assimilantur D-sorbosum, D-xylosum, L-arabinosum, D-rhamnosum, melibiosum, lactosum, raffinosum, inulinum, amylum solubile, D-arabinitolum, galactitolum, inositolum, D-glucuronatum, DL-acidum lacticum, methanolum, propane-1,2-diolum, butano-2,3-diolum, acidum quinicum, D-glucaratum et D-galactonatum. Assimilantur ethylaminum, D-lysinum, cadaverinum et glucosaminum (infirme). Non assimilantur kali nitratum, sodii nitritum, creatinum, creatininum, imidazolum et D-tryptophanum. Amylum non formatur. Biotinum externum ad crescentiam necessarium est. 30 °C crescit neque 35 °C. Non crescit in medio 10 µg ml^–1 cycloheximido addito. Typus: NRRL Y-27588^T (=CBS 9836^T), designat stirpem typicum. Isolata a ile coleopterorum (Tenebrionidae), Barro Colorado Island, Panama, depositata in Collectione Culturarum (NRRL), Peoria, IL, USA.

Description of Candida maxii Suh & Blackwell sp. nov.
Candida maxii (max.i'i. N.L. m. gen. maxii for Max Vallone, whose grandparents generously support the Mycological Society of America).

After 7 days growth in YM broth at 25 °C, cells are globose to subglobose (2·5–6·25x3·75–6·25 µm). Pseudohyphae and septate hyphae are present (Fig. 3b, c). After 7 days on YM agar at 25 °C, colonies are white in colour, ridged, glistening and flat with filamentous margin. After 10 days Dalmau plate culture on corn meal agar at 25 °C, pseudohyphae and septate hyphae are present. Aerobic growth is white, shiny and smooth with filamentous margin. No ascospores produced after 6 weeks at 17 °C from individual strains on YM agar or strains crossed in all combinations on