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Department of Botany, National Museum of Nature and Science, Amakubo 4-1-1, Tsukuba, Ibaraki 305-0005, Japan
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONTAXONOMYLITERATURE CITED |
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To reevaluate two dictyostelid species, namely, Polysphondylium pallidum and P. album, 92 isolates of the P. pallidum complex from their type localities were examined based on mating relationships and morphological characteristics. In the mating tests three heterothallic mating groups were found among the isolates. They also were different morphologically from each other. These results suggested that they belonged to distinct taxa. By comparison of the three mating groups with the type specimens of P. pallidum and P. album, two of them were identified as P. pallidum and P. album. Based on the examined isolates P. pallidum and P. album were redescribed in detail.
Key words: dictyostelid cellular slime molds, mating, Polysphondylium album, Polysphondylium pallidum, species complex, taxonomy
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONTAXONOMYLITERATURE CITED |
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, 1989). In this complex P. pallidum Olive and its related species such as P. album Olive are included.
In 1901 P. pallidum originally was described based on the isolates from animal dungs collected in Liberia, West Africa, and Massachusetts, USA (Olive 1901). Thereafter this species was found in many areas of the world. According to Swanson et al (1999) P. pallidum is a cosmopolitan species.
P. album was described simultaneously with P. pallidum as the second species of white Polysphondylia based on an isolate from toad dung collected in Florida, USA (Olive 1901). This species was distinguished from P. pallidum mainly in having prostrate sorocarps, larger number of branches per whorl and larger sori. However Raper (1951) stated that the identity of P. album was questionable. Indeed it was reported that unpigmented Polysphondylia presented a continuous spectrum of sorocarp pattern and size and furthermore sorocarp patterns and dimensions of parts were influenced strongly by culture conditions such as temperature and humidity (Becker 1959, Raper 1984). Therefore Raper (1984) amended the description of P. pallidum based on many isolates and proposed that P. album was a synonym of P. pallidum.
On the other hand two syngens or mating groups of P. pallidum were discovered in eastern North America (Eisenberg and Francis 1977). Some presumed taxa that are morphologically similar to P. pallidum also were found (Hagiwara et al 1992, Hagiwara 1995, Hagiwara and Kawakami 2000). In addition we reported that at least two heterothallic mating groups (i.e. groups A and B) were present in seven representative strains of P. pallidum (Kawakami and Hagiwara 2002). These two groups were clearly distinguished morphologically. Group A had more numerous branches per whorl, round sorophore bases and ovoid tip cells. In contrast group B was characterized by fewer branches per whorl, clavate sorophore bases and subulate tip cells. It was shown by Schaap et al (2006) that the two groups were phylogenetically different from each other. These studies raise two questions: "What is a true P. pallidum?" and "Is it true that P. album is a synonym of P. pallidum?"
To reevaluate P. pallidum and P. album taxonomically we obtained isolates of the P. pallidum complex from their type localities, examined these isolates based on mating relationships and morphological characteristics and compared the isolates with their type specimens.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONTAXONOMYLITERATURE CITED |
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Isolation of the dictyostelids was performed according to the procedures described by Hagiwara (1989). A suspension (ca. 0.5 mL) of pregrown cells of Escherichia coli in sterile distilled water was spread on the surface of each hay infusion agar plate. Next small particles of soil samples or soil suspensions in sterile distilled water were spread on the surface of the plates. The plates were incubated at 20 C under 12:12 h light:dark conditions. After 1 wk the dictyostelids of the P. pallidum complex growing on the plates were isolated.
Mating test.— –
The mating tests were performed according to the procedures described by Kawakami and Hagiwara (1999). Spores of each pair of strains were inoculated into small colonies of E. coli on 0.1% lactose/proteose peptone agar plates (6 cm diam). For underwater cultures 5–6 mL of sterile Bonner’s salt solution (Bonner 1947) was added to each plate after the spores had germinated. Cultures were incubated at 25 C in the dark, and the presence or absence of macrocysts were examined after 3–5 wk. It is known that macrocysts are formed during sexual cycle (Raper 1984). Two pairs, namely PN500 and PN600 and CK8 and CK9, were used respectively as the mating tester strains of group A and group B.
Morphological observation and terminology.— –
Procedures of morphological observation followed Hagiwara (1989) and Kawakami and Hagiwara (2002). The subcultures of isolates were done with E. coli as food on nonnutrient agar plates to which a granular type of activated charcoal was added at 20 C under 12:12 h light:dark conditions.
Morphological terminology was defined according to those described by Hagiwara (1989). The terms "tip cell" and "whorl index" were defined by Kawakami and Hagiwara (2002); the tip cell represents the terminal cell of a sorophore tip, and the whorl index indicates the ratio of the number of whorls with four or more branches to the total number of whorls examined and is usually calculated using 40 whorls per isolate.
Type specimens of P. pallidum and P. album.— –
The voucher type specimens used for the original descriptions of P. pallidum and P. album have been preserved in the Farlow Reference Library and Herbarium of Cryptogamic Botany (FH), Harvard University, Cambridge, Massachusetts USA. They consist of three dry and three slide specimens of P. pallidum and one dry and two slide specimens of P. album (TABLE I). These specimens are judged as syntypes of each species based on their dates of preparation and/or origins.
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. B: number of branches per whorl; BB(max): width of the branch base at the thickest part (µm); BL: branch length (µm); BT: width of the branch tip at a level 50 µm below the top (µm); IL: internode length (µm); L/W: mean length/width ratio of spores per isolate; MD: mean size of 20 spores per isolate (µm); SB: width of the sorophore base at a level 100 µm above the bottom (µm); SB(max): width of the sorophore base at the thickest part (µm); ST: width of the sorophore tip 50 µm below the top (µm); TC: length of the tip cell (µm); TL: length of the terminal segment that is not lengthened (less than three times IL) (µm); and WI: whorl index. The measurements of dimensions are given as arithmetic ranges or means ± standard deviation, and the number of observations "n" is indicated in parentheses. |
RESULTS AND DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONTAXONOMYLITERATURE CITED |
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.) Fourteen isolates from Florida belonged to group A. Forty isolates from Sierra Leone and 24 from Florida belonged to group B. The other 14 isolates (eight from Sierra Leone and six from Florida) did not belong to either group A or group B. Among these three isolates from Sierra Leone belonged to another heterothallic mating group, namely AS21, which mated with both AS2 and AS23. This mating group was designated group C.
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). The three mating groups clearly were distinguished from one another by a combination of the number of branches per whorl, shape of sorophore base, shape and length of tip cell and spore size. The characteristics of each group are as follows. Group A had higher WI values, round to clavate sorophore bases and typically ovoid and shorter tip cells. On the other hand group B had lower WI values, clavate sorophore bases and typically subulate and longer tip cells. Two isolates (e.g. AS42 and AS50) had an exceptionally large number of branches per whorl (WI: 0.85 and 0.83 respectively). Group C had clavate sorophore bases, typically ovoid and shorter tip cells and smaller spores. In addition this group produced numerous microcysts on agar plates at 20 C under the usual conditions for sorocarp formation. Therefore these results suggested that the three groups represented distinct taxa.
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) probably belonged to any of the three groups; based on their morphological characteristics two isolates from Florida belonged to group A; four from Sierra Leone and four from Florida, to group B; and one from Sierra Leone, to group C. In this study the isolates certified for their mating groups were used for the following comparison of the type specimens of P. pallidum and P. album.
Morphological comparison of the type specimens of P. pallidum and P. album.— –
The original descriptions of P. pallidum and P. album are brief and limited to a few morphological characters (Olive 1901). Hence we examined the type specimens of both species based on several morphological characters (cf. the number of branches per whorl, shape of sorophore base and tip, branch length, etc).
Polysphondylium pallidum Olive #2 (FIGS. 1A–1C)
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), WI: 0.08 (n = 26).
Note. Sorocarps were significantly tangled (FIG. 1B). #3
Sorocarps with 2–4 branches per whorl, WI: 0.6 (n = 6).
Note. Sorocarps were significantly tangled. #4
Sorocarp with 2 branches per whorl, WI: 0.0 (n = 1).
Note. Sorocarps were significantly tangled. 2-6-97 (FIGS. 1D–1F)
Approximately 17 sorocarps were mounted.
Sorocarps with 1–3 nodes (n = 13), with 2–4 branches per whorl, WI: 0.13 (n = 8). Sorophore bases clavate, 12–26 µm in diameter at the thickest part (n = 5) (FIG. 1D); tips acuminate, simple, 2.5–4.5 µm in diameter at a level 50 µm below the top (n = 6); tip cells subulate-like or spathulate-like; terminal segments 462–1100 µm in length (n = 3); internode segments 471–660 µm in length (n = 4). Branches 90–400 µm in length (n = 15); bases clavate, 4.5–19.5 µm at the thickest part (n = 16); tips acuminate, simple, 2.5–6.5 µm at a level 50 µm below the top (n = 11). Spores oblong to elliptical, usually 1.6–2.3 times longer than their width, mostly 5.2–6.5(–7.6) x 2.6–4.1 µm (n = 44) (FIG. 1F). Microcyst-like structures globose, 3.2–7.8 µm in diameter (n = 44).
Note. Tip cells were not so clearly observed (FIG. 1E).
2-24-97 (FIG. 1G)
Approximately 20 sorocarps were mounted.
Sorocarps with 1–3 nodes (n = 10), with 2–3 (mostly 2) branches per whorl, WI: 0.0 (n = 12). Sorophore bases clavate, 12–25 µm in diameter at the thickest part (n = 7); tips acuminate, simple, 3–4.5 µm in diameter at a level 50 µm below the top (n = 3); tip cells subulate-like or spathulate-like; terminal segments 260–440 µm in length (n = 2); internode segments 326–568 µm in length (n = 8). Branches 79–266 µm in length (n = 19); bases clavate, 5.5–17.5 µm at the thickest part (n = 20); tips acuminate, simple, 3–5 µm at a level 50 µm below the top (n = 4). Spores oblong to elliptical (FIG. 1G), usually 1.7–2.4 times longer than their width, mostly 5–7.3 x 2.4–3.8 µm (n = 44). Microcyst-like structures globose, 4.9–8.4 µm in diameter (n = 44).
Note. Tip cells were not so clearly observed.
3-10-01
Two sorocarps were mounted.
Sorocarps with 5 nodes (n = 1), with 2–5 branches per whorl, WI: 0.6 (n = 5), or irregular branches. Sorophore tips acuminate, simple, 3.5 µm in diameter at a level 50 µm below the top (n = 2); tip cells subulate-like; internode segments 576–735 µm in length (n = 4). Branches 235–306 µm in length (n = 5); tips acuminate, simple, 4.0–4.5 µm at a level 50 µm below the top (n = 4). Spores oblong to elliptical, usually 1.5–2.3 times longer than their width, mostly 5.2–6.9 x 2.3–4.1 µm (n = 44).
Note. Sorophore bases were not detected. Tip cells, terminal segments, and branch bases were obscure. Microcyst-like structures were not observed.
Polysphondylium album Olive #1 (FIGS. 2A–2C)
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), WI: 1.0 (n = 9).
Note. Sorocarps were significantly tangled (FIG. 2B). 10/2/’97 (FIGS. 2D and 2E)
One sorocarp was mounted.
Sorocarp with 4 nodes (n = 1), with 2–5 branches per whorl, WI: 0.33 (n = 6). Internode segments 317–524 µm in length (n = 5). Branches 101–224 (–282) µm in length (n = 11); bases clavate, 9–15 µm at the thickest part (n = 5); tips acuminate, simple, 7.5 µm at a level 50 µm below the top (n = 1).
Note. Sorophore base and tip were obscure. Spores were clustered (FIG. 2e).
7/4/’98
One sorocarp and two pseudoplasmodia were mounted.
Sorocarp with 3 nodes (n = 1), with 2–3 branches per whorl, WI: 0.0 (n = 3). Internode segments 260–422 µm in length (n = 3). Branches 112 µm in length (n = 1). Microcyst-like structures globose, 3.5–4.7 (–8.2) µm in diameter (n = 22).
Note. Sorophore base and tip were obscure. Branch bases and tips were also obscure. Spores were clustered.
In the type specimens of P. pallidum Specimen #2 was designated lectotype by Hagiwara (1989). Specimen #4 probably is derived from a culture related to Specimen #2 because its locality and substrate are the same as those of Specimen #2. For the same reason it is considered that Specimens 2-6-97 and 2-24-97 are derived from Specimen #2. On the other hand it is possible that Specimen 3-10-01 does not belong to P. pallidum because it has many irregular branches near its sorophore base and relatively larger number of branches per whorl than those of other specimens.
The spores of P. pallidum originally were described as "occasionally spherical, about 7–8 µ in diameter" (Olive 1901). In Specimens 2-6-97 and 2-24-97 the spherical materials indeed were observed and were in agreement in terms of size (3.2–8.4 µm diam) with the spherical spores described by Olive (1901) (FIG. 1G). However these materials were also in agreement in terms of size with the microcysts of group A or group B (See TAXONOMY). Therefore we confirmed that these materials are probably not spores but microcysts. In fact in our experience we have noted that microcysts often are attached to sorophore bases. We considered that Olive might have picked up sorophores mingled with microcysts while preparing slide specimens during his study.
In the type specimens of P. album, Specimen #1 was designated lectotype of P. album in this study because it is considered that Specimens 10/2/’97 and 7/4/’98 are derived from Specimen #1 based on their locality and substrate.
Olive (1901) stated that P. album sorocarps "are rather constantly weak at the base, so that the fructifications lie close to the substratum in a characteristic fashion." Such fructifications can be regarded as prostrate sorocarps. Indeed in our study the sorocarps of Specimen #1 appeared to be prostrate (FIG. 2B, C). On the other hand Becker (1959) reported that glass-topped cultures of P. pallidum had more numerous prostrate sorocarps than clay-topped cultures. In addition in our experience the formation of such a prostrate sorocarp depends on culture conditions and/or strains of the P. pallidum complex. Therefore we consider that this characteristic fashion is not restricted to P. album.
Olive (1901) stated that "the sori of P. album are larger" than those of P. pallidum. However it is known that the sorus size varies according to culture conditions, particularly humidity (Hagiwara 1989). In addition the sorus size was reported to show a significant difference even between the strains belonging to the same mating group (Kawakami and Hagiwara 2002). Therefore it is difficult to distinguish P. album from P. pallidum based on the sorus size.
Olive (1901) also reported that "the sori of P. album are usually more numerous in a whorl" than those of P. pallidum. This implies that P. album has more numerous branches per whorl than P. pallidum. This description was confirmed by the comparison of both lectotypes of P. pallidum and P. album; the WI value of P. album was obviously higher than that of P. pallidum (TABLE III). In the original descriptions (Olive 1901) the spore size (4–5.6 x 2.5–3 µm) of P. album is somewhat smaller than that (5–6.5 x 2.5–3 µm) of P. pallidum. However because spores of Specimens 10/2/’97 and 7/4/’98 of P. album were densely clustered and mostly in a slanted position in the slides (FIG. 2E) it is possible that the spore size of P. album was reported to be smaller than the actual size. Therefore the difference in spore length between P. pallidum and P. album should not be overestimated.
The morphological comparison of both lectotype specimens of P. pallidum and P. album revealed that P. album is characterized by a comparatively large number of branches per whorl and is clearly distinguishable from P. pallidum by high WI value (TABLE III). Therefore we consider that P. album is not a synonym of P. pallidum and is a distinct species.
For the following comparison with the three mating groups, the lectotypes of P. pallidum and P. album and specimens derived from them were used (e.g. Specimens #2, #4, 2-6-97 and 2-24-97 of P. pallidum; Specimens #1, 10/2/’97 and 7/4/’98 of P. album).
Morphological comparison of the three mating groups with the type specimens of P. pallidum and P. album.— –
We examined whether the morphological characteristics of the three mating groups were present in the type specimens of P. pallidum and P. album. (The results are shown in TABLE III.) Group B was identical to the type specimens of P. pallidum in terms of B, WI, shape of sorophore base and tip, and SB(max). In addition other morphological data of group B considerably overlapped with those of the type specimens. On the other hand, group A was clearly different from the type specimens of P. pallidum in having high WI values and round to clavate sorophore bases. Group C also was distinctly different from the type specimens of P. pallidum in having high WI values and also did not considerably overlap with them in terms of BB(max).
The morphological data of the type specimens of P. album are poor (TABLE III). Nevertheless the WI values of group A were closest to that of the lectotype of P. album. In addition group A was identical to the type specimens of P. album in terms of BB(max) and also considerably overlapped them in terms of IL and BL. On the other hand group B did not considerably overlap with the type specimens of P. album in terms of IL. Similarly group C did not considerably overlap the type specimens of P. album in terms of BB(max). Therefore group A was most similar to the type specimens of P. album.
The morphological comparison of groups A, B and C with the type specimens of P. pallidum and P. album suggested that group A is identical to P. album, group B is identical to P. pallidum and group C is not identical to either P. album or P. pallidum. Thus in the present study we redescribe the morphological characteristics of P. pallidum and P. album in detail based on the examined isolates. It is necessary to examine whether group C is a new taxon by morphological comparison with other known species of the P. pallidum complex.
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONTAXONOMYLITERATURE CITED |
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, 4
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When cultured at 20 C on nonnutrient agar with Escherichia coli, sorocarps usually solitary (FIG. 3C) or sometimes clustered, with 1–8(–10) nodes, with 2–5 (–7) branches per whorl (FIG. 3D–E), phototropic, often prostrate. Sorophores colorless, delicate, sinuous, sometimes with collars, 1.9–9 mm long, gradually tapering from base to tip; bases clavate (FIG. 4A) or sometimes acuminate, 11.5–37.5 µm diam at the thickest part; tips acuminate (FIG. 4B) or sometimes piliform, simple, 1.5–7 µm diam at 50 µm below the top; tip cells typically subulate, or sometimes navicular, aciculate, slightly spathulate, cylindrical, or filiform (FIG. 4B), 12.5–50(–67.5) µm long; terminal segments 339–1135(–1540) µm long; internode segments 427–959 µm long. Branches delicate, 110–348(–405) µm long, often with collars (FIG. 4D), gradually tapering from base to tip; bases clavate (FIG. 4C), 5–20(–24) µm diam at the thickest part; tips acuminate (FIG. 4D), simple, 2–5(–6) µm diam at 50 µm below the top. Sori white, globose; terminal sori 27–174(–237) µm diam; lateral sori 16–106(–147) µm diam. Spores hyaline, oblong to elliptical, usually 1.5–2 times longer than their width, mostly 5.6–7.6 x 3.2–4.6(–4.9) µm, with unconsolidated polar granules and often with irregular granules (FIG. 3F). Pseudo-plasmodia radial or fan-shaped (FIG. 3A), centralized, mostly 335–3183 µm radius, and then often doughnut-shaped after aggregation (FIG. 3B), not migrating without sorophore formation. Microcysts globose to subglobose (FIG. 3G), 3.1–7.1 µm diam. Macrocysts globose to subglobose (FIG. 3H), mostly 17.5–35.8 µm diam except outermost fibrillar cell wall.
B: 3.2 ± 0.9 (n = 1040); WI: 0.0–0.5(–0.85); SB: 17.4 ± 4.8 (n = 125); SB(max): 22.9 ± 5.5 (n = 116); ST: 4.1 ± 1.1 (n = 124); TC: 29.2 ± 9.4 (n = 230); IL: 695 ± 107 (n = 184); TL: 787 ± 256 (n = 69); BB(max): 10.3 ± 3.3 (n = 142); BT: 3.2 ± 0.8 (n = 134); BL: 216 ± 49 (n = 214); MD: 6.1–7.0 x 3.4–4.0(–4.4); L/W: 1.5–1.9.
Habitat. – In forest soil; on ass dung.
Isolates examined. – AS18, 32, 33, 35, 42, 43, 44, 45, 47, 50, 51, 52, 53, 54, 56, 61, 62, 63, 64, 65, 68, 69, 70, 71, 73, 75, 76, 77, 81, 83, 85, 86, 88, 90, 91, 94, 96, 97, 98 and 99, from soils, evergreen forests and fields, Freetown, Sierra Leone, Nov 1995; FL5, 6, 20, 21, 25-2, 26, 27, 53, 54, 55, 56, 57, 58, 59 and 60, from soils, Pinus and Quercus forests, Eustis, and FL41-1, 41-3, 42, 45-3, 46, 48, and 49 from soils, deciduous forest, Eustis, and FL66, from soil, artificial grove, Orlando, Florida, USA, Oct 1996; CK8 (=TNS-C-98) and CK9, from soils, evergreen forest, Tokyo University Forest, Kiyosumi, Chiba Pref., Japan, Jun 1972. Five isolates, AS42, AS68, AS99, FL48, and FL55 were deposited in American Type Culture Collection (ATCC), Manassas, Virginia, USA.
World distribution. – Africa: Liberia, Sierra Leone. America: USA (Florida, Massachusetts), Canada. Asia: Japan, Nepal, Seychelles, Taiwan, Papua New Guinea.
Note. –
Hagiwara (1989) described that the sorocarps of this species were "not phototropic." In this study however it became clear that they were phototropic.
Polysphondylium album Olive, Proc. Amer. Acad. Arts and Sci. 37:342, 1901. FIGS. 5, 6
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When cultured at 20 C on nonnutrient agar with Escherichia coli, sorocarps usually solitary (FIG. 5C) or sometimes clustered, with 1–12 nodes, with 2–7(–12) branches per whorl (FIG. 5D–E), without or rarely with lengthened terminal segments, phototropic, often prostrate. Sorophores colorless, delicate, sinuous, sometimes with collars, 1–13 mm long, gradually tapering from base to tip; bases round to clavate (FIG. 6A) or rarely conical, 14.5–48(–74) µm diam at the thickest part; tips acuminate or nearly obtuse if lengthened (FIG. 6B), simple, 3–10 µm diam at 50 µm below the top; tip cells typically ovoid, or sometimes navicular or oblong to elliptical, 4–20.5(–25) µm long, or nearly cylindrical if terminal segments lengthened (FIG. 6B), 20–35 µm long; terminal segments 198–981(–4664) µm long; internode segments (187–)273–840(–1096) µm long. Branches delicate, 108–326 µm long, often with collars (FIG. 6D), gradually tapering from base to tip; bases clavate (FIG. 6C), 5–15(–27.5) µm diam at the thickest part; tips acuminate (FIG. 6D), simple, 2.5–6(–7.5) µm diam at 50 µm below the top. Sori white, globose; terminal sori 56–168(–201) µm diam; lateral sori 25–102 µm diam. Spores hyaline, oblong to elliptical, usually 1.5–2.2 times longer than their width, mostly 5.7–7.5(–8.4) x 3.2–4.3 µm, with unconsolidated polar granules (FIG. 5F). Pseudoplasmodia radial or fan-shaped (FIG. 5A), centralized, mostly 670–1843 µm radius and then sometimes doughnut-shaped after aggregation (FIG. 5B), not migrating without sorophore formation. Microcysts globose to subglobose (FIG. 5G), 3.1–8.7 µm diam. Macrocysts globose to subglobose (FIG. 5H), mostly 19.8–52.1 µm diam except outermost fibrillar cell wall.
B: 4.5 ± 1.4 (n = 560); WI: 0.68–0.98; SB: 18.0 ± 4.1 (n = 123); SB(max): 30.1 ± 8.9 (n = 125); ST: 5.7 ± 1.6 (n = 103); TC: 11.2 ± 3.8 (n = 144); IL: 561 ± 111 (n = 281); TL: 551 ± 277 (n = 107); BB(max): 9.3 ± 3.1 (n = 170); BT: 4.3 ± 1.0 (n = 184); BL: 208 ± 36 (n = 386); MD: 6.3–7.1(–7.5) x 3.5–3.9; L/W: 1.7–2.
Habitat. – In forest soil; on toad dung.
Isolates examined. – FL1, 2, 3, 8, 11, 12, 13, 15, 19, 22, 24, 39 and 40 from soils, Pinus and Quercus forests, Eustis, and FL45-2, from soil, deciduous forest, Eustis, Florida, USA, Oct 1996; PN500 and PN600, from soils, Delaware, USA, 1977; V-1, from soil, Virginia, USA, Oct 1937; WS543, from soil, Wisconsin, USA, Sep 1969. Three isolates, FL2, FL11, and FL39 were deposited in ATCC.
World distribution. – America: USA (Delaware, Florida, Virginia, and Wisconsin).
Note. – Two strains, FL11 and FL45-2 sometimes produced the sorocarps with lengthened terminal segments.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: shkawak{at}yahoo.co.jp
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONTAXONOMYLITERATURE CITED |
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Eisenberg RM, Francis D. 1977. The breeding system of Polysphondylium pallidum, a cellular slime mold. J Protozool 24:182–183.[Medline]
Hagiwara H. 1982. Altitudinal distribution of dictyostelid cellular slime molds in the Gosainkund region of Nepal. In: Otani Y., ed. Reports on the cryptogamic study in Nepal. Tokyo: National Science Museum. p 105–117.
———. 1989. The taxonomic study of Japanese dictyostelid cellular slime molds. Tokyo: National Science Museum. 131 p.
———. 1995. Dictyostelids from the northern part of Ibaraki Prefecture, Central Japan (In Japanese, with English summary). Mem Nat Sci Mus, Tokyo 28:65–71.
———, Chiu-Yuan C, Zeng-Yung Y. 1992. Dictyostelid cellular slime molds of Taiwan. Bull Nat Sci Mus, Tokyo, B 18:39–52.
———, Kawakami S. 2000. Dictyostelids from the Fukiage Gardens of the Imperial Palace, Tokyo (In Japanese, with English summary). Mem Nat Sci Mus, Tokyo 34: 389–393.
Kawakami S, Hagiwara H. 1999. Macrocyst formation in three dictyostelid species, Dictyostelium monochasioides, Polysphondylium candidum, and P. pseudocandidum. Mycoscience 40:359–361.[CrossRef]
———, ———. 2002. Two mating groups of Polysphondylium pallidum, a dictyostelid cellular slime mold. Mycoscience 43:453–457.[CrossRef]
Olive EW. 1901. A preliminary enumeration of the Sorophoreae. Proc Am Acad Arts Sci 34:333–344.
Raper KB. 1951. Isolation, cultivation, and conservation of simple slime molds. Quart Rev Bio 26:169–190.[CrossRef]
———. 1984. The dictyostelids. Princeton, New Jersey: Princeton Univ. Press. 453 p.
Schaap P, Winckler T, Nelson M, Alvarez-Curto E, Elgie B, Hagiwara H, Cavender J, Milano-Curto A, Rozen DE, Dingermann T, Mutzel R, Baldauf SL. 2006. Molecular phylogeny and evolution of morphology in the social amoebas. Science 314:661–663.
Swanson AR, Vadell EM, Cavender JC. 1999. Global distribution of forest soil dictyostelids. J Biogeography 26:133–148.[CrossRef]
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