Myxomycete species concepts, monotypic genera, the fossil record, and additional examples of good taxonomic practice

Harold W. Keller1 Sydney E. Everhart

Department of Biology, University of Central Missouri, Warrensburg MO 64093, USA 8 0

Conceptos de especies en mixomicetos, géneros monotípicos, el registro 0 2

fósil y ejemplos adicionales de una buena práctica taxonómica , 9 1 - 9

Resumen. Se destacan y amplían algunos de los principales elementos para una buena : 7 2

práctica taxonómica. Se revisan los conceptos de especie en los mixomicetos, a la vez que se A Í

discuten los géneros monotípicos, con ejemplos en Badhamiopsis ainoae, Protophysarum G O

phloiogenum y Trabrooksia applanata. Se sugiere que las secuencias de ADN resolverán el L O C

rango taxonómico al que los géneros monotípicos deben de asignarse en la clasificación de I M los mixomicetos. Se evalúa y discute por primera vez la evidencia fósil de mixomicetos E D

encontrada en ámbar. Perichaena brevifila, P. microspora, P. pedata y P. syncarpon habitan A

exclusivamente en la hojarasca y son un ejemplo como las diferencias ecológicas y los N A C patrones de estacionalidad basados en las observaciones de campo registradas en los datos I X

de las colecciones, pueden complementar las diferencias morfológicas para la separación de E M

las distintas especies. El futuro de la Sistemática de los mixomicetos requiere un cambio de la A T S taxonomía descriptiva a estudios de mayor profundidad basados en hipótesis para probar I V E

relaciones filogéneticas, patrones biogeográficos y restricciones de las especies a hábitats con R

características ecológicas especiales.

/

Palabras clave: Badhamiopsis, ecología, Perichaena, Protophysarum, estacionalidad,

o c

Trabrooksia. i x é M

n e

Abstract. This paper highlights and expands on some of the major points of good taxonomic a s e

practice. Myxomycete species concepts are reviewed, monotypic genera are discussed and r p m

critiqued, and case study examples are given for Badhamiopsis ainoae, Protophysarum I

. a phloiogenum, and Trabrooksia applanata. Monotypic genera are suggested for DNA í g o l

sequencing to resolve the correct taxonomic rank in myxomycete classification. Fossil o c i M

evidence of myxomycetes found in amber is evaluated and discussed for the first time. e d Perichaena species represented by P. brevifila, P. microspora, P. pedata, and P. syncarpon, are a n a

restricted to leaf litter habitats and serve as examples of fruiting seasonality patterns and c i x ecological differences based on detailed field observations recorded in collection data. This e M

a

additional ecological information can supplement morphological differences in distinct t s i v

species. The future of myxomycete systematics requires a shift from descriptive taxonomy to e R

8

in-depth studies using hypotheses that test phylogenetic relationships, biogeographical 0 0 2

patterns of distribution, and the restriction of species to habitats with special ecological

characteristics. © Key words: Badhamiopsis, ecology, Perichaena, Protophysarum, seasonality, Trabrooksia.

Recibido 23 de junio 2006; aceptado 20 de junio 2007. Received 23 June 2006; accepted 20 June 2007.

Autor para correspondencia: Harold W. Keller L

[email protected] or [email protected] A N I G I R O structure details of scanning electron microscopy (SEM). Many new myxomycete species have been Schnittler and Mitchell [31] suggested five criteria L A N Introduction I

Spore-to-spore cultivation and multiple collections were discovered by both amateurs and professionals since the last that authors of new species follow as a prelude to describing G I R

lacking, complicating accurate species descriptions, world monograph published in 1969 [23]. Monographic new species: 1. search literature published throughout the O The number of myxomycete species recognized by Martin especially when considering the variability and limited publications are necessary to assess the validity of new world for possible matching descriptions; 2. reference the and Alexopoulos [23] was roughly 422 compared to the most number of fruiting body and morphological characters. This species but have a long preparation time, requiring specimens new taxon with several specimens from more than one recent estimates of 600 by Nannenga-Bremekamp [27], and approach to taxonomy was followed in many countries, on loan from herbaria (although electronic databases speed locality; 3. create exact descriptions based on SEM 925 of Yamamoto [37]. Schnittler and Mitchell [31] cited resulting in a proliferation of species. this process, when available) and examination of specimens observations, color standardized with color charts, and spore- 1,012 subgeneric taxa of myxomycetes described as valid, The involvement of non-academic taxonomists from closely related species. In addition, accurate species to-spore cultures to ascertain constancy of morphological including 866 at the species level, and Lado [21] noted 900 working privately as individuals is possible because many descriptions are based on light microscopy and SEM quality characters; 4. compare morphologically similar species to legitimate names for accepted species. Lado's taxonomic myxomycete species produce fruiting bodies visible to the photographs that require more time. make sure characters deviate in more than one character, such database with 1,012 taxa could be subdivided into 446 taxa naked eye and materials needed for the collection and For example, to adequately monograph the genus as clustered versus free spores; 5. give details of the habitat estimated to be common (more than 20 collections and preservation of myxomycete fruiting bodies are readily Cribraria (which has more than 40 species that are often such as vegetation type, elevation, and localities. These d r o

reported from several localities) 258 to be rare (known from available. Furthermore, identification of myxomycetes can difficult to distinguish at the species level), it would probably criteria were previously discussed by Keller [16] in his c e r

l i

2-20 collections and more than one locality), and 305 reported be learned quickly because fruiting body terminology is take five to ten years. To avoid others from publishing first, Plenary Address at ICSEM2 held at Madrid, Spain under the s s o f only from the type locality (one or a few collections) [31]. A relatively simple. Specimens can be examined using tools authors are prone to publish hastily, compromising good following topical headings: importance of ecological field e h t

figure by Schnittler and Mitchell [31] of the rate of species such as dissecting needles, handheld blowers, and hand lenses taxonomic practice. The pressure to publish quickly is related collections; importance of collecting; importance of type , a r e

descriptions beginning about 1965 shows a dramatically that are inexpensive to purchase or are handcrafted [34]. to the prestige factor associated with describing a new species collections; importance of spore-to-spore cultivation; living n e g

c

increasing number of described new taxa per year, throughout Slide preparations do not require complex chemicals, tap and not for work to clarify species concepts and solve cultures – a biological standard; importance of monographic i p y the world. water is sufficient. Moist chamber bark cultures are simple to taxonomic problems. works; importance of computerization of mycological t o n What are the reasons for this “explosion” of many prepare using materials available at commercial stores. Bark The myxomycete species problem and concepts as collections; importance of DNA sequencing techniques. It is o m

, s new species in the last 35 years? Certainly more cultures yield plasmodia, plasmodial tracks, and developing discussed by Clark [5] suggested that taxonomists become unfortunate that Schnittler and Mitchell [31] did not review, t p e c

myxomycologists were searching in more habitats, such as fruiting bodies easily observed at 10 to 100 times more familiar with the potential morphological variation due cite, or include these observations about good taxonomic n o c

8

the canopy of living trees [17] and different areas of the world magnification [19]. Myxomycetes, much like the macrofungi to geographical restriction of apomictic clonal lines and also practice [16]. s 0 e i 0 c 2 [32]. However, the increase seems to be present during the (mushrooms), pique the curiosity of amateur collectors that acquire a better understanding of population, developmental, The purpose of this paper will highlight and expand e

p , s

7 e

2 time when Nanennega-Bremekamp and many other non- eventually result in publication of new species without and reproductive biology. Genetic mating systems in on some of the major points of “good taxonomic practice” t

e A c Í y

G academic taxonomists were working privately as individuals, academic, professional taxonomists as collaborators. Indeed, Didymium iridis (Ditmar) Fr. and Didymium squamulosum previously noted with an additional caveat associated with m O o L myxomycete fruiting bodies often win photographic prizes at (Alb. & Schwein.) Fr. produce species complexes consisting habitats and seasonality patterns. Myxomycete species x O coupled with a trend for the “splitting” of genera and species, y C I M

.

M resulted in too many dubious species based on single or less the annual North American Mycological Society meeting of related sibling species that are potential morphospecies concepts will be reviewed, monotypic genera will be E

. E S

D , t

than four collections from a restricted locality. because of their striking beauty [19]. In comparison, related often geographically isolated. His final summation is worth a documented and critiqued, and case studies given to evaluate r A a N h

Unfortunately, many single collections only known from the groups of organisms, the dictyostelids and protostelids, direct quotation: “Therefore, it is suggested that taxonomists the past, present, and future application of methodologies that r A e C v I E X type locality had too few fruiting bodies and resulted in require laboratory culture methods and equipment not undertake the naming of new morphospecies with due care, will enhance our understanding of taxa in the Myxomycetes , . E M W . species descriptions based on limited material, inadequate for available to most amateurs so these groups were studied and that they base their descriptions on reasonable number of [18]. H A

T , r S I comparisons with other taxa. Furthermore, type specimens almost exclusively by professional, academic sporophores (as many as possible) collected from a number of e l l V e E

R were often retained in private herbaria and papers were myxomycologists. Thus, the taxonomic confusion and different areas (as widespread as possible), combined with a Monotypic Genera – human artifact or real? K published in regional and not international journals that number of synonyms for dictyostelids and protostelids were basic understanding of the population and developmental Monotypic genera are those genera that include only one undergo a rigorous merit review process. In most cases minimized, especially in view of the fact that culture biology of the myxomycetes”. This paper should be required species. Higher taxonomic ranks are human creations that species were illustrated with line drawings that lack the fine techniques are required for their isolation and observation. reading for all myxomycete taxonomists. give relative order to the species and represent hierarchies of 1 0 1 1 Table 1. List of monotypic genera

diverging characters [33]. Therefore, ranking taxa, including well preserved columella with capillitial attachments still L Martin and Alexopolous (1969) Erionema aureum Penz. A N I

the creation of monotypic genera, is highly subjective. The retaining a surface net intact, typical of an extant species of Arcyodes incarnata (Alb. & Schwein.) O. F. Cook Kelleromyxa fimicola (Dearn. & Bisby) Eliasson G I Barbeyella minutissima Meyl. Leocarpus fragilis (Dicks.) Rostaf. R problem is exacerbated by the practice of publishing new Stemonitis, possibly S. splendens. The preservation is O Brefeldia maxima (Fr.) Rostaf. Leptoderma iridescens G. Lister names in local, non-peer reviewed journals. Thus, new remarkable and there is no question that this is a species of Calomyxa metallica (Berk.) Nieuwl. Lindbladia tubulina Fr. descriptions and changes can be proposed without a literature Stemonitis and a myxomycete. Even the black (dark) color of Calonema aureum Morgan Listerella paradoxa E. Jahn. Cienkowskia reticulata* (Alb. & Schwein.) Rostaf. Minakatella longifila G. Lister review and assessment by experts in the field. The current the fruiting body structural parts (hypothallus, stalks, and Clastoderma debaryanum A. Blytt Mucilago crustacea F. H. Wigg. number of monotypic genera in the Myxomycetes was capillitial threads) are still attached to the columella with Cornuvia serpula (Wigand) Rostaf. Physarella oblonga (Berk. & M. A. Curtis) Morgan determined using three sources. In accordance with Martin branching patterns and a surface network intact as seen in Erionema aureum Penz. Protophysarum phloiogenum M. Blackw. & Alexop. Leocarpus fragilis (Dicks.) Rostaf. Prototrichia metallica (Berk.) Massee and Alexopolous [23], monotypic genera represent 17 of 53 optical section [6]. Lindbladia tubulina Fr. Trabrooksia applanata H. W. Keller genera or 32%, according to Keller and Braun [19] monotypic Arcyria sulcata Dörfelt & Schimdt is another Listerella paradoxa E. Jahn. Willkommlangea reticulata* (Alb. & Schwein.) Kuntze Minakatella longifila G. Lister genera represent 21 of 57 genera or 39%, and according to example of a myxomycete species from fossilized Baltic Lado NOMENMYX (2001) Mucilago crustacea F. H. Wigg. Arcyriatella congregata Hochg. & Gottsb. Lado [21] monotypic genera represent 14 of 59 genera or amber that was provided with a Latin diagnosis and described Physarella oblonga (Berk. & M. A. Curtis) Morgan d Barbeyella minutissiuma Meyl. r Prototrichia metallica (Berk.) Massee o 24%. Species which represent monotypic genera according as a new species. The paper title “The oldest fossil c Brefeldia maxima (Fr.) Rostaf. e r

Wilczekia evelinae Meyl. l Cornuvia serpula (Wigand) Rostaf. i to each source are shown in Table 1. myxogastroid slime mould” is based on Baltic amber from a s s

Keller and Braun (1999) Kelleromyxa fimicola (Dearn. & Bisby) Eliasson o f similar time-period (early Tertiary, Eocene) as the Stemonitis Arcyodes incarnata (Alb. & Schwein.) O. F. Cook Leocarpus fragilis (Dicks.) Rostaf. e h t

Fossil Record species. However, it is not clear which myxomycete species Arcyriatella congregata Hochg. & Gottsb. Lindbladia tubulina Fr. , a r

Badhamiopsis ainoae (Yamash.) T. E. Brooks & H. W.Keller Listerella paradoxa E. Jahn. e

Monotypic genera may be valid if it is the last remaining came first in the fossil record. The color illustrations (see [8] n Barbeyella minutissima Meyl. Minakatella longifila G. Lister e g

c

species from a group that has otherwise gone extinct, Figures 1-4) collectively show the stalk and capillitial threads Brefeldia maxima (Fr.) Rostaf. Mucilago crustacea F.H. Wigg. i p y however, this validation would require representative attached at the base to the calyculus. The color of the fruiting Calomyxa metallica (Berk.) Nieuwl. Physarella oblonga (Berk. & M. A. Curtis) Morgan t o

Cornuvia serpula (Wigand) Rostaf. Protophysarum phloiogenum M. Blackw & Alexop. n examples within the fossil record. Protozoa are known from body is not mentioned in the species description nor is the Dictydiaethalium plumbeum (Schumach.) Rostaf. o Prototrichia metallica (Berk.) Massee m

,

s the fossil record preserved in amber [10], however color evident in the amber. The capillitum appears as a coiled Willkommlangea reticulata* (Alb. & Schwein) Kuntze t p e c

myxomycete preservation is rare. Fossilized myxomycetes network of apparently elastic threads. Special light * Willkommlangea reticulata and Cienkowskia reticulata are now recognized as synonyms. n o c

8

are only known as preserved in amber, which is plant resin microscopic techniques (laser scanning-microscope) show s 0 e i 0 c 2 that hardened under the right conditions and over a long the ornamentation (cogs and rings) on the capillitial threads e

p

, surface or internal areas of the stalk and upper parts show no solid mass whereas myxomycete species usually have a s

7 e

2 period of time. Considering the habitats myxomycetes typical of Arcyria species [8]. The general habit appears t

cellular detail, however, the filaments are more likely fungal powdery spore mass. The spore mass in some myxomycete e A c Í y G occupied and their fragile structure, fossilization of fruiting similar to the extant myxomycete species Arcryia denudata in origin. Images (see [7] Figure 1, G) show the thickened species may stay within the spore case when the top portion m O o L (L.) Wettst., suggesting myxomycete fruiting bodies have x O bodies or spores in amber is the most likely and stage . To the margin is enrolled, unlike Protophysarum which has a undergoes circumscissile dehiscence and the lid falls away y C I M

.

M best of our knowledge, no myxomycologist has evaluated changed little in 35 to 40 million years. E

delicate spherical spore case that is thin and fragile without a from the peridium, as in the stalked species Licea operculata . E S

D , t

fossilized myxomycete taxa represented by fruiting bodies in Protophysarum balticum Dörfelt & Schmidt was

cuplike base. Myxomycete peridia and stalks are not cellular (Wingate) G.W. Martin. However, Protophysarum balticum r A a N h

the fossil record and this represents the most current review. described as a new myxomycete species in Baltic amber from r A

whereas lichens often have filaments or cells indicative of more closely resmbles the calicioid lichen, Chaenotheca e C v I E X Myxomycete fruiting bodies have been reported in the Tertiary period. A Latin diagnosis was not included, only , . E fungi. Dr. Thorsten Lumbsch, an expert lichenologist at the species, fossilized in amber and illustrated by Rikkinen [30] M W . Baltic amber [6, 8]. The first certain myxomycete in the fossil an English description. This specimen in amber clearly lacks H A Chicago Field Museum, reviewed the text and illustrations of (see mature ascoma in [30] Figures 1 and 2).

T , r S I record was a species assigned to Stemonitis splendens Rostaf. the features of Photophysarum phloiogenum M. Blackw. & e l l V this paper [7]. He indicated that the specimen is not a In addition to fruiting bodies, other myxomycete life e E R in Baltic amber from the Tertiary, Eocene approximately 35 to Alexop. based on the following morphological characters: myxomycete but a lichen in the calicioid group sometimes cycle stages have been preserved in amber, such as the K 40 million years ago. Images (see [6] Figure 1-7, Plate 15) the stalk is much larger and thicker, the basal part of the spore referred to as “stubble lichens”. Stalked species of Licea plasmodium of a physaraceous species [35]. Even so, there clearly show stalked sporangia with hypothallus and case is persistent, somewhat turbinate, with a thickened wall, sometimes resemble calicioid lichens (Keller, pers. obs.), are problems with the description of the fossilized columella. Further images (see [6] Figure 6 and 7) represent a sometimes only a cuplike base remains. Apparently the however the spores of Protophysarum balticum appear as a plasmodium. The objects inside the amber are a continuous 3 2 1 1 Table 2. Seasonal occurrence of Perichaena species

section that is described as the veins of a phaneroplasmodium Protophysarum was the extinct sister species to L Months A N I

and vesicles that are described as part of a plasmodium that Protophysarum phloiogenum, according to some species J F M A M J J A S O N D Total G I

Perichaena brevifila 1 1 2 3 6 13 R has budded off and sclerotized. Every part of the plasmodium concepts, the genus would no longer be considered O P. microspora 3 3 3 9 appears as though it is a network of bubbles that Waggoner monotypic and would be a good example where an extant P. pedata 2 4 3 1 10 P. syncarpon 1 1 8 8 4 1 1 24 and Poinar [35] describe as spherules, and suggest is the result species represents a monotypic genus. of initial plasmodial sclerotization. Inside each spherule, “specimen found on upper layer of decaying leaves under full recognition of four separate species. Details about the habitat Waggoner and Poinar also state that there are typically 0 to 6 Additional Methods of Good Taxonomic Practice cover of a bush in an urban landscape” rather than “found on may explain why there is a seasonal difference in occurrence spherical nuclei that are 8 to 24 µm in diameter and classify leaves.” Furthermore, the collection date of fruiting bodies of the species and why there are a few collections that are the plasmodium in the Physarales. However, the Physarales Phylogenetic Analysis should always be recorded. A good example of the use of outliers. If the species of Perichaena require similar average is an order in which the nuclei are characteristically small, for The description of a new species that represents a monotypic detailed habitat description and seasonality of fruiting to temperatures for fruiting, the occurrence of P. brevifila at the example, the nuclei of Physarum polycephalum Schwein. are genus is compelling when experts agree the specimens differ support classification is in the monograph of Perichaena in a bottom of the decaying leaf litter may explain the difference in well documented [11] as globose to elliptical in shape and 2.5 from all other genera by more than one character. When a d r

thesis by Keller [14]. The thesis recorded the seasonal seasonality. It takes longer for average temperatures at the o to 7 µm in diameter. The “nuclei” described by Waggoner and genus is based on only one character it is open to questionable c e r

l

occurrence of Perichaena species from shaded, decaying bottom of a leaf litter to reach the same temperature at the top i

Poinar are outside of the size range characteristic of the interpretation. The authors suggest the use of DNA analysis s s o

leaves or straw stacks. Detailed notes about the habitat of leaf litter. The months where only one collection is f Physarales and outside of the size range expected for most as additional evidence to support monotypic genera [9]. e h t

showed that P. brevifila T. E. Brooks & H. W. Keller was recorded may also be explained by the habitat. Leaf cover myxomycetes. The images are unconvincing, especially the Good candidates for genetic analysis are: Arcyriatella , a r

found near the bottom of leaf litter and other species were may provide enough protection for the fruiting bodies to be e

scattered bubbles and questionable “nuclei” that lack any congregata Hochg. & Gottsb., Erionema aureum Penz., n e g

found near the top of leaf litter, and P. microspora Penz. & preserved for months, which would explain the collection of c

organization or internal details suggesting a plasmodium or Kelleromyxa fimicola (Dearn. & Bisby) Eliasson, Leocarpus i p

Lister was collected only from Florida and Louisiana and is P. brevifila in January and March, and the collection of P. y sclerotium. It is doubtful that the objects inside the amber are fragilis (Dicks.) Rostaf., Lindbladia tubulina Fr., Listerella t o n

therefore considered a Southeastern species in the United syncarpon in January and P. pedata in November. o actually a myxomycete plasmodium based on the description paradoxa E. Jahn., Minakatella longifila G. Lister, m

,

States of America (U.S.A.) The assumption is that collectors regularly and s by Waggoner and Poinar and the amorphous properties of a Physarella oblonga (Berk. & M. A. Curtis) Morgan, t p e

Collections were made between 1930 and 1977 from equally collected during each month and from the same c

myxomycete plasmodium that would make the capture of a Protophysarum phloiogenum, Prototrichia metallica (Berk.) n o c

various locations in the U.S.A.: Perichaena brevifila from habitats. In general, myxomycete fruiting in the U.S.A. is 8

plasmodium by resin extremely unlikely. Arguments for the Massee, and Trabrooksia applanata H. W. Keller (Table 1). A s 0 e i 0 Georgia, Kansas, and Virginia, P. microspora from Florida, most diverse and abundant in June, July, and August. c 2 description of fossilized plasmodium would have been more good example of the use of phylogenetic analysis is e

p , s

7 Georgia, and Louisiana (collection data supplemented from Exceptions to this generality are known from California e

2 convincing had the authors provided an image of a partially represented by the genus Schenella. Schenella simplex T. t

e A c

Í labels provided by BPI), P. pedata (Lister & G. Lister) Lister where the rainy months are in January and February. Indeed, y

G sclerotized plasmodium in vitro compared to that in the Macbr. was analyzed using molecular DNA sequencing and m O o L ex E. Jahn from Illinois, Florida, and Kansas, and P. those months correspond to the months where myxomycete amber, provided reasoning for the difference in expected x O was found to be synonymous with the fungal gasteromycete y C I M

syncarpon T. E. Brooks from Iowa and Kansas. The fruiting is most diverse and abundant, lending support to the .

M versus observed nuclei size, and consulted the expertise of a (puffball) Pyrenogaster [9]. E

. E S

collection dates of the four Perichaena species that only seasonality of myxomycetes based on precipitation and D , t

professional myxomycologist. r A a

N occurred on either shaded, decaying leaves or straw stacks temperatures. Clearly, in the case of Perichaena species, h

The validation of monotypic genera in the Ecology and Seasonal Fruiting r A e C v I were compared (Table 2). The occurrence of P. brevifila was notes about the ecology and seasonal occurrrence of fruiting E X myxomycetes using the fossil record is highly problematic The ecology and seasonal occurrence should be recorded , . E M

mostly (>1 observation) in September, October, and bodies in the field add support to deliniation of species within W . considering the improbability of fossilization and difficulty in regularly when studying myxomycetes, since little is known H A

T , r

S November, whereas the species P. microspora and P. pedata the genus. The same could be true for other genera but more I assigning specimens to the correct genus. Fossilized about the phenology of myxomycetes. Although these are not e l l V e E occurred mostly in June, July, and August, and P. syncarpon observations of seasonal occurrence and ecology must be R specimens of both Stemonitis splendens and Arcyria sulcata characters used to classify myxomycetes, they may be K occurred mostly in July, August, and September (Table 2). recorded before that conclusion can be made. appear to be valid, however, it is doubtful that Protophysarum supporting characters by which to group or split Seasonality of fruiting alone is not a defining characteristic of balticum is even a myxomycete and more likely to be a morphologically similar specimens. Therefore, collections Perichaena species. However, these data add support to the calicioid lichen. However, if the fossilized species of should always thoroughly describe the habitat in detail, 5 4 1 1 16 REVISTA MEXICANA DE MICOLOGÍA 27, 2008 quotation identification.” The proved induced not classifying absence culture, compared form T “ T and considered myxomycete that spore description smaller Y recognized by 26, Populus Juniperus occurs and than was his genus T T Discussion T rabr rabr rabr rabrooksia amam., rabr collection also the 29]. first genus Alexopoulos 45 falls ooksia ooksia ooksia size of in ooksia on collections stable absence spores nor character 1980 balsamifera of the In

collected

Didymium because living was A vir in when a is within lime this of 1983 does

countries tested in synonym H. W applanata, applanata giniana new the keyed of applanata to Selected species. in authors in the described (8-9 This slime honor trees corticolous combined compared not of protologue M. numerous . Martin is variety key in 1969 are for Keller a lime µm)

commentary mention here, not L. L. 1962 L. stur such of mold of size known T to presence ,

Monotypic , The

Farr ravis , was a and quite Great Podocarpus T world the gisii , et numerous publications type , based is monotypic . by at by is T as applanata. with to al range in . collections myxomycetes an Ulmus genera variety updated [15]. described least highly E.

the

Acer Y T applanata [24] from 1 certain and any Britain, Hagelst, . 1-13 amamoto inherent Brooks monograph on E. the of iridescent is did for

negundo only other Brooks. not Genera seven sp. It elemental a but provided other suggestive µm micr macr and genus single has as not purposes Ireland,

T noting as and species unlike from yet, with cited rabr genus. or var states assign ospora not [36] revised traits ophylla a [15]. peridium

to environmentally L. case . Since collection. new but [24], ooksia here been micr various commemorate calcium. by , which the and with Fraxinus in of preclude of many of T

so [13, 19, study Whether Keller

rabr monotypic the This following: the should ospora as then a (Th.) the specimen including Japan. grown

far has limeless slightly a and U.S.A. Martin it ooksia states. genus direct taxon it other more been

Sw [15] was The His sp., has 25,

the the be Y or

in It . . ,

character presence specimens a plasmodiocarps crystals, stellate, aggregated the no ef carbonate light to T distinctly as flattened T Fructifications Morphological ( ( 173-176, detailed species distinguish that nomenclatural Alexopoulos D species. examination w one as T T b rabr rabrooksia e i rabr rabr e a d iridescent sessile r r e y synonym correct peridium r fervescence inclearlactophenol;

m a on there ooksia ooksia ooksia n b i u t a have

species the NOMENMYX is m color

to s noncalcareous in T or Didymium as sporangia. h e

dif rabr name d a irregularly of these with are species internal s v is

seen applanata p. of of absence surface, t ferent & e been is u o p. determination are thin, D.

more ooksia 152, r Didymium n of holotype descriptions g numerous Farr comparison a for

a points with i two

396-401); similar s stur s D. thin i c c compared Didymium membranous, every of i , spore

a a in stur

grayish stur rarely l n Color ef SEM taxa. of Gen. gisii applanata [ c Didymium n 2 a fused calcareous protruding the i specimens 1 gisii r n in gisii specimens calcium e species ] a nomenclatural stur g

mass, o

morphological

(see shape, should u

Myxomycetes: and brownish is a by Additional

two s by l

gisii plasmodiocarps n when of p. i

stur that morphologically premature t d e Brooks p [15]

peridial r , lacking 185-189); [21]. is e Didymium a Keller transparent . gisii t as form, taxa. r carbonate are t h i crust u or be listed d e were

lightly or Figures r in

i e u

prematurely

because p.

sometimes The s m expected other collections [4] Didymium stur o and specimens s 149-150). The structural characteristics in . of u and o treatment

as not r u names ( c characters view 71 N by sprinkled r size, T 7-10) crystals e a c species with sturgisii is rabr o and

e peridium seen. n synonym Braun of s [24]. Keller i along c

s a of usually a

since less compiled ooksia a reflected free a l and variable dried calcium c n M of of the to give silvery a d of Many either r

a

often e each with with both with This gisii r [19] [15] and that fact and o t n the the are i u or as of o in n p. s t ,

examined 81 harvested dead 1933, U.S.A., October Ntra. C. bark examined: Brooks are size into anastomosing, l may threads unattached, and shaped trabeculae all of anastomosing, The to light vertically two D. often or Physarales. a rg 78 Didymium. the in stur Lado, of needed e taxa. narrow attached capillitial and of be microscope 74 r moist yield aspen the

BPI Sra. peridium gisii attachments . Juniperus [4] mostly p unassociated Iowa, ornamentation Specimens The l T 1980, specimens MA-Fungi a and aligned

817869 rabr of

21 and s noncalcareous Enebral, Spain,E.Guadal [19] chamber (calcareous attachments m Sp and bark, When D. to

capillitial o cited

threads The August ec and with ooksia East Keller with d . and stur the collected giving i and im o ; oxycedrus (see moist c capillitial the Colorado, fructifications by en a to violaceous gisii. base below Okoboji, the cultures examined on examined. r Figures 17073

with and p has Keller s rarely the violaceous s [15] 1979, rise in system , bark [4, extremities pillars)

of chamber or simple, Braun upper forms, both m the by to

; 19]. to Figures sometimes L., branch, a [15]. ajara, system E. 9, T these Gilpin the of y on ra collected trabeculae C. branching, colors.

of 1 species Guadalajara, 23 is peridial

[19]. 1 br The In however Juniperus b often base subparallel, , prematurely decaying color with e D. Lado, distinctly oo environmental T March wetted

contrast, amajon-Guadalajara, on is County hyaline 4, lack spores ks s stur

Additional i a of

The m SEM) 5, ia seen but with with constant wall by p the

calcium and gisii , MA-Fungi 6 1980, slender l none 21 e ap capillitial additional , wood, appear as thurifera , in Chapman, Perigo and dif

plasmodiocarp. ends D. [12], T attached tubular pl in broad dry July were seen b amajon-Erata many ferent an were r stur collected some character a attenuating , collections conditions in carbonate, n at truncated, extending capillitial similar 4 c N 9, gisii with a the cited h

August funnel- species threads system seen 17177

e L., Slope, SEMs above in of 1979, d we field

BPI has the the the 22 by o re in in in in r .

Physarum appears scanning calcareous globose commentary Brooks T on livingtreesin pillars the When like calcareous, detach plasmodiocarp hyaline, crystalline), vertical, enclosing tubular recognition plasmodiocarps o dif lactophenol is clearlya non-calcareous chambers, proximity living corticolous T is mostsimilar Badhamiopsis Badhamiopsis rabr rab f

fers species. a pillars r

ooksia ook the from c trees [20] from are invaginations a to unbranched T non-calcareous with sia p . electron wo spores dense on i especially remain be l capillitium physaraceous myxomycete,however However bifurcate

of . and myxomycetes l thin,

. all indicative However i attenuating the t Specimens the additional and app

i a ainoae a petaloid ainoae a [20]. Badhamiopsis in appearanceand other fruiting Keller l and separate stalked

deposits same are the fieldandin bottom lan hyaline, attached n vines micrographs e The capillitial , genera removed, ata pillars, threads on t

, under , (Y w from and a the piece based of bodies o taxa Juniperus amash.) . dehiscence monotypic species plasmodiocarps, threads r as genus. examined were and of k where calcium Braun SEMs restricted

like . and certain

the in short, more assigned of The break white on T fail nucleata the system sometimes this h bark icicles upper T non-calcareous, e of The

attached moist chamber [19]. the attempt the lack

more easilyconfused . to Physaraceae or vir two carbonate. environmental E. and is calcareous slender Badhamia p genus produce in exposing r Latin giniana less to delicate a Brooks to e capillitium to were

the peridium,

the d the distinctive H. Badhamiopsis the o tax the when cited in m spike-like, to to high , case field calcareous appear diagnosis. Z. a i the upper often n bark bubbles . the illustrate & a

Badhamiopsis Badhamiopsis are hyaline, Li t granules or in a broken study resolution e , thevertical basis H. and , sometimes l base oriented its y consists

branching, Keller and conditions is possibly similar surface feature peridium. bifurcate, W

com in in absence stalked, e . usually in for ff Keller spike- of

moist open, merit close what clear Four u non- mon with (not s the the e of as of of to to d a

17 Keller, H.W., Everhart, S.E. Myxomycete species concepts, monotypic genera, the fossil record ORIGINAL Botánico, CSIC, Madrid, Spain, Abstract Volume: 23-37.

show detailed fine structure of the fruiting body. National Monument, Arizona. the National Science Foundation, Biodiversity and Survey 17. Keller, H.W., 2004. Tree canopy biodiversity and student research L A

experiences in Great Smoky Mountains National Park. N I

Furthermore, the stalked habit, the branching capillitium, and Program, DEB #0343447, National Geographic Committee Systematics and geography of plants 74: 47-65. G I

18. Keller, H.W., 2005. What is a myxomycete species? In: Galindo, G., R larger spore size (15-18 µm in diameter) are characters that for Research and Exploration #7272-02, and, Discover Life in M.G. Santiago- Martínez, A. Montoya Esquivel, A. Estrada- O Conclusions Torres (eds.), Fifth International Congress on systematics and would exclude this taxon from the genus Badhamiopsis [22]. America Award #2002-17. We thank Carrie M. Harrington for ecology of Myxomycetes. Abstracts of oral and poster A single collection of Badhamiopsis cavifera Nann.-Brem. & the English to Spanish translation. presentations. Universidad Autónoma de Tlaxcala. 45. 19. Keller, H.W., K.L. Braun, 1999. Myxomycetes of Ohio: their systematics, Y. Yamamoto on moss from a living tree has a capillitium Phylogenetic relationships based on molecular biology and use in teaching. Ohio Biological Survey Bulletin New Series Vol. 13, Number 2, 1-182. typical of a Diderma or Didymium. Two sessile, pulvinate methodologies will highlight the taxonomy of Myxomycetes 20. Keller, H.W., T.E. Brooks, 1976. Corticolous Myxomycetes IV: Badhamiopsis, a new genus for Badhamia ainoae, Mycologia 68: st sporangia are illustrated that apparently represent the basis for in the 21 Century. The criteria for species concepts will References 834-841. 21. Lado, C., 2001. NOMENMYX. A nomenclatural taxabase of the description since no other fruiting bodies are mentioned in surely involve DNA analysis that focus on population Myxomycetes. Cuadernos de Trabajo Flora Micological Iberica 16: 1-221. the text. Without the holotype available to examine and such diversity and the complexities of understanding population 1. Blackwell, M., 1974. A study of sporophore development in the myxomycete Protophysarum phloiogenum. Archives for 22. Li Hui-zhong. 1989. Myxomycetes from China IV: Taxonomic study in a scanty specimen it is difficult to assess if this represents a differentiation. Species concepts should include more data Microbiology 99: 331-344. the Myxomycetes of Shennongjia. In: Fungi and Lichens of 2. Blackwell, M., C.J. Alexopoulos, 1975. Taxonomic Studies in the Shennongjia, Mycological and Lichenological Expedition to new taxon. However, the line drawings clearly indicate that than just morphological differences with additional Myxomycetes. IV. Protophysarum phloiogenum, a new genus Shenongjia, Academia Sinica. World Publishing Corp, Beijing. d and species of Physaraceae. Mycologia 67: 32-37. China. Pp. 1-78d. r o

the habit, the color and calcareous peridium, the obvious information from biogeographical patterns, habitat analysis 23. Martin, G.W., C.J. Alexopoulos, 1969. The Myxomycetes. University of c

3. Blackwell, M., R.L. Gilbertson, 1984. Distribution and sporulation e r

Iowa Press, Iowa. phenology of Myxomycetes in the Sonoran Desert of Arizona. l i

didymiaceous capillitial threads, exclude this taxon from the based on field observations over time, and seasonality Microbial Ecology 10: 369-377. 24. Martin, G.W, C.J. Alexopoulos, M.L. Farr, 1983. The genera of s s o

4. Brooks, T.E., 1967. A study of corticolous Myxomycetes. Ph.D. Myxomycetes. University of Iowa Press, Iowa. f genus Badhamiopsis [28]. patterns in circumscribed geographical areas. Selected Dissertation, University of Kansas, Lawrence, United States of 25. Mitchell, D.W., 2004. A key to the corticolous Myxomycota, Systematics e h t

America. monotypic genera represent sources of DNA that will and geography of plants 74: 261-285. , a 5. Clark, J., 2000. The species problem in the Myxomycetes. Stapfia 73: 39- 26. Mitchell, D.W., 2005. Myxomycetes, CD-ROM of illustrated keys and r e

Protophysarum phloiogenum, a monotypic case study eludicate taxonomic ranks (orders, families, and genera) and a 53. inventory of all world species. CD for MS Windows. DiKey, n e

6. Domke, W., 1952. Der erste sichere Fund eines Myxomycetes im dichotomous keys to various groups of organsisms. Inventory g

and synoptic keys of all legitimately described myxomycete taxa. c Protophysarum phloiogenum is an example of a good more correct assignment of problematic taxa such as Baltischen Berstein (Stemonitis splendens Rost. Fa. Succini fa. i

This is a private publication by the author and is regularly p

nov. Foss.). Mitteilungen aus dem Geologischen Staatsinstitut in y monotypic genus [2]. At first glance, the specimen appeared Protophysarum and Trabrooksia. Monotypic genera may be updated. t Hamburg 21: 154-161. o n

27. Nannenga-Bremekamp, N.E., 1991. A guide to temperate Myxomycetes, o to be a Lamproderma within the Stemonitaceae, however, the last representatives of taxa gone extinct but the paucity of 7. Dörfelt, H., A.R. Schmidt, 2006. An archaic slime mould in Baltic amber.

an English translation by A. Feest and Y. Burggraaf of De m

Palaeontology 49: 1013-1017. ,

Nederlandse Myxomyceten. Biopress Ltd., Bristol, England. s four distinct characters contradicted that placement. First, the a fossil record leaves many unanswered questions. Good 8. Dörfelt, H., A.R. Schmidt, P. Ullmann, J.Wunderlich, 2003. The oldest t 28. Nannenga-Bremekamp, N.E, Y. Yamamoto. 1988. Badhamiopsis p

fossil myxogastroid slime mould. Mycological Research 107: e 123-126. cavifera (Myxomycetes), a new species from Japan. Journal of c plasmodium was a phaneroplasmodium not an taxonomic practice and monographic studies using modern n

Japanese Botany 63: 28-30. o

9. Estrada-Torres, A., T.W. Gaither, D.L. Miller, C. Lado, H.W. Keller, 2005. c

8 aphanoplasmodium; second, sporangial development is methodologies are important because they lay the foundation The myxomycete genus Schenella: morphological and DNA 29. Neubert, H., W. Nowotny, K. Baumann, 1996. Die Myxomyceten s 0 e Deutschlands und des angtenzenden Alpenraumes unter i

0 sequence evidence for synonymy with the gasteromycete genus c 2 subhypothallic not epihypothallic; third, calcium carbonate for sound scientific research and allow the application of Pyrenogaster. Mycologia 97: 139-149. besonderer Beruchsichtigung Osterreichs, Band. 2. Physarales. e

p ,

Karlheinz Baumann: Gomaringen. s

7 10. Frederick, L., 1990. Phylum plasmodial slime molds, class Myxomycota. e

2 was found associated with the peridium and capillitium; and knowledge across a broad spectrum. The future of 30. Rikkinen, J. 2003. Calicioid lichens from European Tertiary amber. t In: Margulis, L., J.O. Corliss, M. Melkonian, and D.J. Chapman e A

Mycologia 95: 1032-1036. c

Í (eds.), Handbook of Protoctista. Jones and Bartlett Publishers, y G finally, the spores germinated via splitting open rather than myxomycete systematics requires a shift from descriptive Boston. Pp. 467-483.

31. Schnittler, M., D.W. Mitchell, 2000. Species diversity in Myxomycetes m O o L based on the morphological species concept – a critical through a pore. The description is a good example of patience 11. Gray, W.D., C.J. Alexopoulos, 1968. Biology of the Myxomycetes. New x O taxonomy to in-depth studies using hypotheses that test comparison. Stapfia 73: 55-62. y C York: Ronald Press. I M

.

M in taxonomic descriptions as the specimens were cultured phylogenetic relationships, biogeographical patterns of 12. Hagelstein, R., 1944. The Mycetozoa of North America based upon the 32. Stephenson, S.L, H. Stempen, 1994. Myxomycetes: a Handbook of Slime E

Molds. Timber Press, Portland, Oregon. . E

specimens in the Herbarium of the New York Botanical Garden. S

D , t spore-to-spore over a period of seven years, with variation distribution, and the restriction of species to habitats with Mineola, New York. 33. Stuessy, T.F., 1990. Plant Taxonomy. Columbia University Press, New r A

York, New York. a

N 13. Ing, B., 1999. The Myxomycetes of Britain and Ireland: an identification h

noted only in spore size. Furthermore, additional work on the special ecological characteristics. Only when r A handbook. Richmond Publishing Company, Slough, England. 34. Sundberg, W.J., H.W. Keller, 1996. Myxomycetes: some tools and tips on e C v I collection, care, and use of specimens. Inoculum 47 (4): 12-14. E

X 14. Keller, H.W., 1971. The genus Perichaena (Myxomycetes): a taxonomic species has been published [1, 3] which lends further support myxomycologists collaborate will we reveal answers to the , . E and cultural study. Ph.D. Dissertation, University of Iowa, Iowa. 35. Waggoner, B.M., G.O. Poinar, Jr., 1992. A fossil myxomycete M W . for the species, including analysis of sporophore development question: “What is a myxomycete species?” [18]. 15. Keller, H.W., 1980. Corticolous Myxomycetes VIII: Trabrooksia, a new plasmodium from Eocene-Oligocene amber of the Dominican H A

T genus, 72: 395-403. Republic. Journal of Protozoology 39: 639-642. , r S I and identification of the species from additional locations. e 36. Yamamoto, Y., 1994. Some new and rare records of Myxomycetes from l l V 16. Keller, H.W., 1996. Invited plenary address, Biosystematics of e E Myxomycetes: a futuristic view. In: Lado, C. and J.C. Hernandez Japan. Hikobia 11: 523-530. R Interestingly, the initial collections were made from the bark K (eds.), Second International Congress on the systematics and 37. Yamamoto, Y., 1998. The myxomycete biota of Japan. Shinjuku-ku, of living Ulmus americana L. in Colorado and later Acknowledgments ecology of Myxomycetes (ICSEM2), organized by Real Jardín Tokyo, Toyo Shorin Publishing. collections were made from the pith of dead saguaro, Carnegiea gigantea (Engelm.) Britt. & Rose, in Saguaro This research was financially supported in part by grants from 9 8 1 1