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Xylose Medium, and Furthermore Tested for Amylolytic and Lipolytic Activity (After PERSOONIA Published by the Rijksherbarium, Leiden Volume 13, Part 1, pp. 97-105 (1986) Notes on Zygorhynchus species M.A.A. Schipper Centraalbureau voor Schimmelcultures, Baarn* An account is given of species of Zygorhynchus maintained in the CBS collec- to tion, with emphasis on zygospore formation. A key species is added. difficult isolates of of all CBS Two to identify Zygorhynchus spp., prompted a study for the the strains in the genus comparison with original diagnoses and with descrip- tions given in the monograph of the genus by Hesseltine & al. (1959). This study, with of emphasis on zygospore formation, led to a re-evaluation some key characteristics used by Hesseltine & al. (1959). All strains maintained in the CBS collection sub Zygorhynchus californiensis, Z. ex- ponens var. exponens, Z. exponens var. smithii, Z. heterogamus, Z. japonicus, Z. macro- and Z. examined. carpus, Z. moelleri, Z. psychrophilus, sp. were METHODS 3°C Cultures were grown on beerwort agar at temperatures from 15 to 36°C, at 20° from increments; Z. psychrophilus, which does not grow at or above C, was grown 15°C. 5—20°C. 0 to For similar reasons, Z. japonicus was grown at Following the of methodology Hesseltine & al. (1959), cultures were also grown on D-glucose and D- xylose medium, and furthermore tested for amylolytic and lipolytic activity (after Hankin & Anagnostakis, 1975). 4% 2% Media: beerwort agar containing resp. sugar; D-glucose and D-xylose medium, 20 7H 0 0.25 2.0 KH2P04 containing D-glucose or D-xylose g, MgS0 L-asparagin 4 2 g, g, 0.5 thiamine-HCl 0.25 trace elements solution 15 1 de-ionized g, g, 1 ml, agar g in 1 malt malt extract 10 extract 4 V8 water; yeast agar: g, yeast g, glucose 4 g per litre; juice agar (200 ml V8 per litre);oatmeal agar: extract of 30 g flakes per litre. Zygorhynchus Vuillemin(1903) Type species: Zygorhynchus heterogamus (Vuill.) Vuill. (1903) =Mucor heterogamus Vuill. (1886). Vuillemin(1903) distinguished Zygorhynchus from Mucor on differencesof the zygo- spore-apparatus. the both and the In Zygorhynchus suspensors are unequal, in size shape, smaller one straight and short, the larger long, curving upwards and widening towards a pyriform shape. The most conspicuous feature in Zygorhynchus is the occurrence of the com- Address: P.O. Box 273, 3740 AG Baarn, The Netherlands. 97 98 PERSOONI A Vol. 13, Part 1, 1986 Fig. 1. Zygosporic stages. — a. Zygorhynchus. — b. Mucor. (Schematic.) with the aerial plete zygospore apparatus, extremely unequal suspensors, on same hypha. In the closely related genus Mucor, suspensors, varying from equal to rather un- equal, always originate from different hyphae, both in heterothallic and homothallic species (Fig. 1). In some species of Zygorhynchus however, some be , zygospores may formed following the Mucor pattern. Green (1927) found that in Z. moelleri Vuill. branch 'sometimes the gametangia were produced on different zygophores and that one might produce two gametangia which might fuse with others on independent zygo- phores'. Hesseltine & al. (1959) described and discussed the mixture of Zygorhynchus and where both Mucor patterns in Z. exponens Burgeff, patterns are quite common. observed the The same situation was in present study in CBS 154.69, Z. japonicus Ko- minami. Zygorhynchus exponens Burgeff—Figs. 2, 4e in Abh. 4: 34. 1924. Zygorhynchus exponens Burgeff Bot. The three strains of Z. exponens maintained in the CBS collection were all derived ' from denoted ' isolate of the the type strain. The strain niger by Burgeff, a single-spore different from the others. type strain, is (now) not 24° colonies of all three showed when On beerwort agar at C grey and white patches, 27-30°C and formed both the grown at they were yellow grey. Zygospores were in Zygorhynchus and Mucor fashion, the latter prevailing. — of forest Material examined. CBS 141.20,type strain Zygorhynchus exponens, ex soil, 403.58 1492 = of Geisenheim, Germany, 11. Burgeff = CBS = NRRL Blakeslee C-972, subculture — isolate from 141.20. CBS 508.48, strain ‘niger’, single spore CBS 141.20,H. Burgeff. Hesseltine considered formation this intermediate & al. (1959) zygospore in species between Z. moelleriand homothallicMucor species. SCHIPPKR: Notes on Zygorhynchus 99 Fig. 2. Zygosporic stages. — a. Mucor plumbeus, CBS 213.75 (+) x 848.73 B (—). — b. Zygo- CBS 141.20. — c. Z. CBS 508.48. — d. Z. CBS 403.58 rhynchus exponens, exponens, exponens, — Z. CBS 404.58. e. exponens var. smithii, 100 P E R S O O N I A Vol. 13, Part 1, 1986 The of Z. colonies of with patchy colouring exponens is reminiscent strains an un- balanced sexual state e.g. azygosporic descendantsof normally heterothallicspecies. Mor- phologically, Z. exponens is most closely related to Mucor plumbeus: like M. plum- beus, Zygorhynchus exponens has short racemose branches with small sporangia; break- conical columellae ing sporangial wall; (some) with a short apical projection; globose, verrucose sporangiospores; brown zygospores which are (often) borne between equal or inequal suspensors originating from different hyphae and zygospore ornamentation of similar patterns (Schipper & al., 1975). Occasionally sporangia and zygospores are formed on the same hypha. Sporangia, of sporangiospores and zygospores Mucor plumbeus and Zygorhynchus exponens are of about the same size. The differences between the two species are the shape of the of the of majority columellae and occurrence zygospores borne between suspensors from originating one hypha. Zygorhynchus exponens in the Zygorhynchus stage showed only moderately en- rather variable Unusual the characters larged suspensors, in shape (Fig. 2). though may the has be stable. After of the seem, species proved to sixty years repeated culturing, type strain still fits the author's drawings, and so does the 1948-single spore isolate. of of Dr. W. Gauger, University Nebraska, kindly germinated zygospores CBS 141.20. Germination was unusually fast: after only 8 days the first germination was observed. Thirteen viable germinations were obtained: two germ sporangia and eleven mycelial germinations. Both single germ spore isolates (five from one germ sporangium) and hyphal tip isolates (nine, taken at random) produced colonies identical to the parental one. of features CBS 404.58, the type strain Z. exponens var. smithii, shows similar and is regarded as synonymous. Material examined. — CBS strain of smithii 404.58, type Zygorhynchus exponens var. Hesseltine & al. = NRRL 2572 = 1MI 79624 = IFO 6665, ex soil, U.K. Zygorhynchus japonicus Kominami — Fig. 3 Zygorhynchus japonicus Kominami in Mykol. Zentbl. 5: 3. ('1914') 1915. Kominami's strain is no longer known to exist. In 1968 a similar fungus was isolated by A. A. Milko (Pidoplichko & Milko, 1971) which was available for study: CBS 154.69 (= VKM F-1382, ex forest soil, Chesnovizkaja Region, USSR). In this strain, a Mucor found zygosporic condition was to occur alongside the Zygorhynchus condition. however CBS 154.69 did not grow at 25°C, good growth and sporulation occurred between 5 and 20°C. on 4% beerwort beerwort It was grown agar, 2% agar, maltyeast oatmeal at 20°C. agar, agar and V8 juice agar, In each colony the Mucor pattern of formation to The zygospore seemed prevail. typical ‘japonicus’ pattern, as described and occurred mixed with the the by Kominami, was rare, ‘Mucor’ pattern on same hyphae; indisputable azygospores were observed (Fig. 3). Pairs of zygospore-bearing aerial from the substratum distinct cultures. hyphae arising separately were in young Colonies derived from isolated zygospores were similar. SCHIPPFR: Notes on Zygorhynchus 101 — Fig. 3. Zygosporic stages in Zygorhynchus japonicus, CBS 154.69. a. ‘Typical’ pattern. — b. Prevailingpattern. — c. Azygospores. Zygorhynchus japonicus does not closely resemble any known Mucor species. There are some similarities with Mucor bacilliformis Hesseltine (1954). One of these is the of attraction of several aerial hyphae by a single one opposite sex resulting in an arach- undisturbed colonies. noid appearance in young Zygorhynchus heterogamus (Vuill.) Vuill.& Z. macrocarpus Ling — Fig. 4c Zygorhynchus heterogamus (Vuill.) Vuill. in Bull, trimest. Soc. mycol. Fr. 19: 116. 1903. in Revue Bot. 42: 7. 1930. Zygorhynchus macrocarpus Ling gen. 102 PERSOONIA Vol. 13, Part 1, 1986 Of both species some type material is still in existence: The holotype material of Z. heterogamus = Mucor heterogamus Vuill. consists of slides prepared by Vuilleminin 1886. (Hesseltine & al. (1959) studied and described these slides). CBS 215.27 = NRRL2663 = type strain of Z. macrocarpus, ex soil of peatery, France. The first description of Z. heterogamus, complemented with Vuillemin's drawings, fea- and the observations of the original slides by Hesseltine & al. gave the following tures: sporangia av. 50—60 jjni diam.; columellae globose (on narrow base); sporangio- spores globose to short ellipsoidal, 3—5.5 x 2.1—4 /urn; zygospores 45—150, mostly over 100 ptm diam, the larger suspensors subglobose. These features differ slightly from those given by Hesseltine & al. (1959) as a com- posite after five isolates, identified with Z. heterogamus and cultured on SMA viz. sporangia mostly up to 35 (15—55) /am diam.; columellae applanate to almost globose; sporangiospores irregular short oval, a few globose, 2—4 x 4—6.5(5 x 8) pm; zygo- spores 30—70 pm diam.; the larger suspensors abruptly inflated, globose. Material examined. CBS 215.27 = NRRL 2663 = type strain of Zygorhynchus macrocarpus = = Ling, ex soil of a peatery, France. — CBS 455.58 NRRL 1616 Harvard University Collection no. 202. — CBS 338.74, ex sediment in a drain pipe, Sweden. — CBS 580.83, ex sandy soil, potato- field, Haren, The Netherlands (1981). — CBS 594.83, ex soil, Paramo Cruz Verde, ca. 3000 m alt., Cundinamarca,Columbia, H. Valencia] isol. W. Gams. of fits The type strain Z. macrocarpus the above description of Z. heterogamus. Ling Young (1930) noted the similarity of zygospores in both species, but omitted to in- dicate differences.
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