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NOTES ON GEOGRAPHIC DISTRIBUTION Check List 14 (3): 509–518 
https://doi.org/10.15560/14.3.509
New records of genera and species of myxomycetes (Amoebozoa) 
from the Neotropics
Carlos Lado1, Arturo Estrada-Torres2, Carlos Rojas3, 4
1 Real Jardín Botánico (RJB, CSIC), Plaza de Murillo 2, 28014 Madrid, Spain. 2 Centro Tlaxcala de Biología de la Conducta, Universidad 
Autónoma de Tlaxcala, Km 1.5 carretera Tlaxcala-Puebla s/n, 90070, Mexico. 3 Engineering Research Institute, Universidad de Costa Rica, San 
Pedro de Montes de Oca, 11501-Costa Rica. 4 Experimental Interdisciplinary Station of Agroecological Models (FEIMA), University of Costa 
Rica, Turrialba, 30502-Costa Rica.
Corresponding author: Carlos Rojas, carlos.rojasalvarado@ucr.ac.cr
Abstract
During field surveys of myxomycetes in Central America, 2 previously unrecorded genera and 4 species, viz. Crate-
rium muscorum Ing, Dictydiaethalium dictyosporum Nann.-Bremek., Physarina echinocephala Höhn, and Stemonaria 
rufipes, were identified. Some of these are new for the Neotropics. These noteworthy range extensions of these species 
to the Mesoamerican biodiversity hotspot adds to our knowledge of rarely recorded myxomycetes worldwide. Images 
of the more relevant taxonomic characters are provided, and for some species, this is the first illustrations with macro 
and microphotographs and scanning electron microscopy images.
Keywords 
Biodiversity; geographical distribution; Mesoamerica; SEM.
Academic editor: Adalgisa Fernanda Cabral | Received 18 December 2017 | Accepted 5 April 2018 | Published 11 May 2018
Citation: Lado C, Estrada-Torres A, Rojas C (2018) New records of genera and species of myxomycetes (Amoebozoa) from the Neotropics. Check 
List 14 (3): 509–518. https://doi.org/10.15560/14.3.509
Introduction the limited availability of basic taxonomic information 
The myxomycetes have been documented in Central on some rarely documented species on the global level 
America for over 100 years (Lado and Wrigley de Bas- (Lado and Eliasson 2017), such task is highly significant 
anta 2008). In this geographical area, the implementation to strengthen the current catalog of data on myxomycetes.
of regional research agendas relating to myxomycetes has The latter is particularly relevant for a group of 
been limited by the centralization of biological research microorganisms which has not received much attention 
in Costa Rica and Panama. Despite this, a recent effort from scientists carrying out broader types of analyses. 
to normalize the investigation on myxomycetes in the Despite this fact, some efforts to address large-scale 
northern part of Central America has generated recent population assessments (e.g. Rojas et al. 2011, Lado et 
advances (Morales 2017). al. 2013, 2014), including assessing worldwide patterns 
Based on the scant historical efforts to investigate of distribution (Aguilar et al. 2014) and biogeography 
myxomycetes in Central America (see Rojas and Doss (e.g. Estrada-Torres et al. 2013, Dagamac et al. 2017), 
2013), it is not surprising to discover new records for have been carried out. These works have allowed a bet-
these microorganisms in such region. However, given ter understanding of myxomycete distribution across 
Copyright Lado et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted 
use, distribution, and reproduction in any medium, provided the original author and source are credited.
510 Check List 14 (3)
different ecosystems and geographical localities. How- with taxonomic comments relating to either the original 
ever, large informational gaps still exist for most rarely descriptions or other relevant observations on distribu-
recorded species. tion made by other authors. Forest classification follows 
In the present work, 4 undocumented species for Cen- Holdridge (1967). 
tral America are described, illustrated and commented 
upon. The objective of this effort is to provide the com-
munity of researchers in the field of microbial biology Results
with integrated taxonomic and ecological data on the The 4 species considered in the present work represent 
myxomycete species being considered. This is important new records for the Central American region and for 
since it provides new information for more accurate taxo- Costa Rica. Three of these species also represent new 
nomic identifications and higher resolution ecological records for the Neotropics. The collecting location of 
analyses. each specimen is presented in Figure 1. 
Methods Craterium muscorum Ing 
Trans. Brit. Mycol. Soc. 78(3): 443 (1982)
For the present study, 3 of the specimens were collected Figures 2–8
as part of research expeditions made by all the authors 
and 1 was isolated in a moist chamber culture. All collec- New record. Costa Rica, San José Province, Dota 
tions were made in Costa Rica between 2015 and 2017, County, in a patch of forest on km 77 of the Southern 
under the framework of Costa Rican Law 7788 (article 4 Interamerican Route (09.6154° N, 083.8192° W, ca 3000 
and transitory) and Resolution 5861-2005 from Univer- m), Arturo Estrada-Torres, 2 March 2016, on bryophytes, 
sity of Costa Rica. in evergreen lower montane wet forest, MA-Fungi 91170.
Field collections were made using the opportunistic Description. Sporocarps grouped or scattered, stalked, 
method described by Cannon and Sutton (2004). Fol- 1.6–2 mm in total height. Sporotheca subglobose, 0.6–
lowing this method, substrates where fruiting bodies 0.8 mm diameter, blackish (65. br. Black-62. d. gy. Br). 
of myxomycetes tend to develop were surveyed. When Hypothallus membranous, blackish. Stalk stout, erect, 
myxomycetes were discovered, they were extracted and cylindrical, attenuate toward the apex, 1–1.3 mm long, 
placed in small cardboard boxes for identification and dark brown to blackish (65. br. Black-62. d. gy. Br), filled 
storage. The specimen collected from a moist chamber with whitish, rhombic crystals of lime. Peridium single, 
appeared on ground litter as a substrate. For this method, membranous, slightly iridescent, dark grayish brown (62. 
explained by Stephenson and Stempen (1994), the d. gy. Br), blackish toward the base, slightly paler toward 
decomposing plant material was placed in a Petri dish the apex, orange brown (54. br O-55. s. Br) by transmitted 
previously lined with filter paper and water was added. light; dehiscence irregular, by the upper part of the sporo-
After 24 hours, excess water was discarded, and the moist theca, the basal part remaining like an irregular calyculus. 
chamber was kept under observation for 10 weeks. In a Columella concolour with the stalk, whitish in the upper 
similar manner, for the field collections, once the fruiting section, capitate form, reaching the middle of the sporo-
bodies were identified, they were extracted and placed in theca, filled with lime. Capillitium rigid, almost entirely 
small cardboard boxes for additional study. limy, fragile, merging with the columella, branched and 
For identification, all microscopic measurements, anastomosed, as a whitish tridimensional net, filled with 
observations and illustrations were made using a Nikon lime. Spores free, black in mass, dark brown (59. d. 
Eclipse 80i microscope with Nomarski interference con- Br-61. gy. Br) by transmitted light, subglobose, 12.5–15 
trast and a Leica M205 stereomicroscope provided with µm diameter, reticulated, sometimes with the reticulum 
a digital camera. The material was mounted directly in broken by Light Microscopy, simple reticulate type under 
Hoyer’s medium or polyvinyl alcohol (PVA). Three of the SEM, sculptured with muri of 1–1.3 µm height, making 
species considered in this study were also observed under a small number of meshes to a hemisphere. Plasmodium 
a scanning electron microscope to analyze ultrastructural not observed.
details, whereas the fourth one was not examined this 
way due to limited material. All the selected specimens Notes. Lister and G. Lister (1904), who originally de-
were treated with the critical point dried material tech- scribed this species as Badhamia rubiginosa var. globosa 
nique, and all the observations were made in the Real Lister & G. Lister, found it in the north of Wales at the 
Jardín Botánico, CSIC, using a Hitachi S-3000N scan- end of the 19th and beginning of the 20th century and 
ning electron microscope at 10–15 kV. Color notations mentioned that this species was “always occurring on 
are from the ISCC-NBS Color Name Charts Illustrated moss and ferns growing on wet rocks” in ravines. Martin 
with Centroid Colors (Kelly 1965). Nomenclature of the and Alexopoulos (1969) included it as a synonym of Bad-
species follows Lado (2005–2018). For the description hamia obovata (Peck) S.J. Smith (as var. globosa), but Ing 
of the spores using SEM, the terminology proposed by (1982) clarified the nomenclatural status of this species. 
Rammeloo (1974, 1975) has been utilized. A descrip- The known distribution of the species according to 
tion of specimens of each species is provided, along Lister (1925) was limited to Great Britain, Ireland, Ger-
Lado et al. | Neotropical myxomycetes  511
Figure 1. Map of Costa Rica showing the collecting locations for 1) Craterium muscorum, 2) Dictydiaethalium dictyosporum, 3) Physarina 
echinocephala and 4) Stemonaria rufipes. Forests are seasonal on the Pacific slope (A: 2 and 4) and non-seasonal on the Caribbean slope (B: 
1 and 3).
many (Hosltein) and the Sandwich Islands in the South New records. Costa Rica, Alajuela Province, Atenas 
Pacific Ocean. According to the Global Biodiversity County, in a pasture about 2 km SW of Escobal 
Information Facility (GBIF) data portal, some specimens (09.9297°N, 084.4520°W, ca 330 m), Arturo Estrada-
also have been found in France, other parts of Germany Torres, 4 March 2016, on decayed wood, in a seasonal 
and Nepal. Stojanowska and Panek (2003) also reported premontane moist forest, MA-Fungi 91171.
it from Poland. The present record represents the first Description. Sporocarps densely packed as a pseudoa-
one for the American continent and the entire Neotropi- ethalium, at maturity simulating an aethalium, effused to 
cal region and widely broadens the known distribution pulvinate, depressed, 5–10 × 1.2–2.5 mm, olive brown 
of the species. Interestingly, this specimen was found (95. M. Ol Br-94. L. Ol Br). Sporocarps sessile, concres-
in the Cerro de la Muerte area, previously reported as a cent to coalescent, closely compacted in a palisade layer, 
region where myxomycete assemblages include species cylindrical. Hypothallus dark brown, protruding from the 
traditionally associated with temperate areas (Leontyev pseudoaethalium. Peridium single, membranous, slender 
et al. 2014, Rojas et al. 2015). The affinity of this species and evanescent at the base, persistent at the apex as plates 
for bryophytes and colder wet environments is supported of 75–150 µm diameter, slightly polyhedral to rounded, 
by the present study. dark grayish brown (62. d. gy. Br), dark yellow (88. d. 
This species is very similar to Craterium dictyospo- Y) by transmitted light; dehiscence apical by breaking 
rum (Rostaf.) H. Neubert, Nowotny & K. Baumann, but between the plates. Columella absent. Capillitium absent. 
the reticulation of the spores, which is strongly marked, Pseudocapillitium thread-like, olivaceous (94. l. Ol Br), 
almost complete (Figs 7, 8), and with a prominent edge fading to darker (95. m. Ol Br), hanging from the borders 
in C. muscorum, sets them apart. In addition, the lack of of the peridial plates, visible only after dehiscence, the 
lime in the peridium (Fig. 2), very evident in C. dictyos- threads 1–2 mm long, flat, 7–12 µm diameter, translu-
porum, is also an important difference. cent, yellowish (86. l. Y-90. gy. Y) by transmitted light, 
smooth but with a fringe of around 2 µm wide in one 
Dictydiaethalium dictyosporum Nann.-Bremek. side, sometimes undulate. Spores free, dark yellow (88. 
Proc. Kon. Ned. Akad. Wetensch., C. 69(3): 345 (1966) d. Y) in mass, pale yellow (89. p. Y-90. gy. Y) by trans-
Figures 9–17 mitted light, subglobose, 12.5–14 µm diameter, banded 
512 Check List 14 (3)
Figures 2–8. Craterium muscorum Ing (MA-Fungi 91170). 2. Stipitate sporocarp. 3. Sporocarp showing the columella. 4-5. Reticulate spores 
by LM. 6. Sporocarp under SEM. 7–8. Spores under SEM showing the ornamentation reticulate type, sculptured with muri. Scale bars 2–3 = 
0.5 mm; 4–5, 7–8 = 10 µm; 6 = 1 mm.
reticulated, the meshes small, about 1 µm across and southwest Pacific). The material is conserved in the BM 
bands about 1 µm high by LM, densely pilate to simple herbarium and no additional specimens have been pub-
reticulate type under SEM, sometimes the reticulum is lished except for a record reported by Keller (1973) for 
broken. Plasmodium not observed. the Florida Everglades (USA), identified by Nannenga-
Notes. This species was described by Nannenga- Bremekamp; a collection made by H. Seraoui (5 October 
Bremekamp (1966) on wood collected during 1914 in 2007, HS 3379) on decayed wood in New Caledonia 
New Caledonia (in the subregion of Melanesia, in the (again), deposited in Marianne Meyer´s collection under 
Lado et al. | Neotropical myxomycetes  513
Figures 9–17. Dictydiaethalium dictyosporum Nann.-Bremek. (MA-Fungi 91171). 9. Sporocarps densely packed as a pseudoaethalium. 10–11. 
Sporocarps showing the remaining peridium and the pseudocapillitium thread-like. 12. Plate of the peridium with pseudocapillitium under 
LM. 13. Flat and undulated pseudocapillitial thread, tape-like by LM. 14–15. Plate of the peridium and pseudocapillitium under SEM. 16–17. 
Spores under SEM. Scale bars 9 = 2 mm; 10 = 1 mm; 11 = 0.5 mm; 12 = 50 µm; 13 = 10 µm; 14–15 = 100 µm: 16–17 = 10 µm.
number MM30415 (Poulain et al. 2011); and a report record presented herein is the fifth globally, and the first 
from Ranade et al. (2012) for India. In the BR herbarium, one for Central America and the Neotropics. It is interest-
according to the GBIF data portal (https://www.gbif.org/ ing to note that all reports have been made in tropical/
occurrence/665667634), there is a specimen from Florida subtropical areas with strong seasonality. The record 
with an identical collecting date and location as Keller´s from Florida and one from New Caledonia were collected 
material and probably corresponds to this specimen. The in October during the fall and the record from Costa Rica 
514 Check List 14 (3)
Figures 18–25. Physarina echinocephala Höhn. (MA-Fungi 91172). 18. Sporocarp showing the typical lime projections of the sporotheca, 
radiating from the peridial membrane. 19. Open sporocarps showing the columella. 20. Open sporotheca showing the peridium with 
brownish areola at the contact with the white spiculate projections.  21–22. Capillitial threads and spores by LM. 23-24. Spores under LM. 25 
Limy spiculate projections of the peridium by LM. Scale bars 18–20 = 0.5 mm; 21–22 = 5 µm; 23–25= 10 µm.
in March during the spring. In all locations at that time of Description. Sporocarps grouped, stalked, 0.6–1 mm 
the year, the temperature is warm (around 26 °C) and the in total height. Sporotheca whitish, limy, subglobose, 
precipitation oscillates around 200 liters per square meter 0.4–0.6 mm diameter, with spiculate lime projections. 
for the respective monthly period. Stalk conical, 0.3–0.5 mm long, whitish, colorless to pale 
The distinctive character of this species is the spore orange yellow (73. P. OY) by transmitted light, filled with 
ornamentation, composed by fused verrucae or little lime crystals. Peridium membranous, grayish, with many 
pilae, giving a reticulate pattern, sometimes broken, as whitish, lime projections radiate from the membrane, 
observed by SEM (Figs 16, 17). The flat threads of the areolate by transmitted light, colorless but brownish at 
pseudocapillitium (Figs 12, 13), tape-like and undulated, the contact with the white spiculate projections, the pro-
also contribute to the characterization of the species. In jections 25–75 µm long, in form of cones or pila, filled 
the specimen examined in the present study, the spore size with lime granules; dehiscence irregular. Columella 
is larger than the one reported by Nannenga-Bremekamp representing a projection of the stalk into the sporotheca, 
in the original description (10–12 µm diameter). The subglobose to dome-shaped, light yellowish brown (76. 
same was observed by Keller for the Florida specimen. l. y. Br), occupying a third of the sporotheca, limy. Cap-
illitium filiform, limeless, brown (58. m. Br-59. d. Br), 
Physarina echinocephala Höhn. branched, with few anastomoses. Spores free, blackish in 
Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl. mass, light brown (57. l. Br) by transmitted light, subglo-
118:432 (1909) bose to slightly polygonal, 9–10 µm diameter, minutely 
Figures 18–25 warted. Plasmodium not observed.
New record. Costa Rica, Cartago Province, Turrialba Notes. The most prominent difference that exists for the 
County, in ravine next to a creek, about 0.6 km SE of La collection checked herein and descriptions of the species 
Esperanza de Atirro (09.8003° N, 083.6439°W, ca 750 in the literature, is the color of the sporocarps. Höhnel 
m), Carlos Lado, 18 October 2017, on dead leaves, in an (1909) described it as brown chocolate and Lister (1925) 
evergreen premontane wet forest, MA-Fungi 91172. as pale pink or flesh colored. However, Alexopoulos and 
Lado et al. | Neotropical myxomycetes  515
Blackwell (1968), using a collection obtained from a remains as a collar or little calyculus, membranous, trans-
culture from Durango (Mexico), described it as “brown lucent, colorless by transmitted light, densely granulated. 
when still moist, but turning grayish-white upon drying Columella concolorous with the stalk, reaching the apex 
because of the lime which encrust them”. The color of the of the sporotheca, attenuate toward the apex, spreading as 
present collection, whitish, matches the description made a membranous layer just at the apex. Capillitium filiform, 
by the latter authors for mature forms. Also, the spores branched and with only a few anastomoses in the interior, 
observed in the present collection have the same size netted, isodiametric, dark grayish brown (62.d. gy. Br) 
(8.5–10.5 µm diameter) described by them. The darker by transmitted light, sparingly joined to the columella 
peridium (as areoles) in the point of insertion of the peg- along its whole length, the primary branches sometimes 
like protuberances (Figs 20, 25) is characteristic, but not with little membranous expansions at the connection of 
previously mentioned in the descriptions of this species. the columella, net with some membranous expansions 
Also, the light color and tenuous ornamentation of the at the nodes, the secondary branches in the lower part 
spores observed by transmitted light are remarkable of mostly united at the periphery, but not forming a surface 
this species (Figs 23, 24). The capillitium observed in this net, with the same lumen that the internal net, sometimes 
collection is longer and lacks the membranaceous expan- with spines, the apical secondary branches ending free. 
sions found by Alexopoulos and Blackwell (1968) in the Spores clustered, 3–7 spores in each cluster, strongly 
specimen from Mexico. coherent, dark brown to blackish (62. d. gy. Br-65. br 
According to Alexopoulos and Blackwell (1968) and black) in mass, grayish brown (60. l. gy. Br-61. gy. Br) 
Martin and Alexopoulos (1969), this species is known by transmitted light, subglobose, slightly depressed at 
only from Java, Thailand, and Mexico. Ndiritu and de external pole, 9–10 µm diam, warted to spinulose, with 
Hann (2014) also reported it for the African Continent slightly more dense ornamentation on the outside of each 
(Rwanda and Burundi). Its presence in the Turrialba cluster by LM, baculate to slightly pilate under SEM. 
region of Costa Rica seems to coincide more with the Plasmodium not observed.
first 2 locations due to climatic similarities. At least in 
the mid- to southern section of Thailand and in the island Notes. This is the second record of this species world-
of Java, premontane moist forests resemble structurally wide, as it was previously known only from Japan as 
those in the Neotropics. This species may have eluded reported by Nannenga-Bremekamp et al. (1984). Accord-
collection in other surveys in Central America due to poor ing to the GBIF data portal (GBIF 2016), there are 3 
surveys in ravines or riparian forests or simply because specimens recorded, 1 in the BR herbarium (the type 
fruitings were not present when surveys were carried collection) and 2 more, in the TNS herbarium, also from 
out. For example, it is interesting to note that the pres- Japan. This is the first specimen of the species outside 
ent record was found already colonized by microfungi Asia, and thus the first record for the Americas and the 
in a dry-period window after constant rainfall during the entire Neotropics. 
previous weeks, suggesting that humidity may be a factor Its distinctive characters are the clustered spores (Figs 
accounting for its fruiting. 30, 31, 34, 36), a capillitium consisting of threads of 
uniform diameter, even in the primary branches, mostly 
Stemonaria rufipes Nann.-Bremek. & Y. Yamam. united at the periphery in the lower part, but not forming 
In Nannenga-Bremekamp, Yamamoto & Sharma, Proc. a surface net, and the collar remaining at the base of the 
Kon. Ned. Akad. Wetensch., C. 87(4): 456 (1984). sporotheca (Figs 26–29, 32, 33). The material checked in 
Figures 26–36 this study differs slightly from the description made by 
Nannenga-Bremekamp et al. (1984) in the smaller spore 
New record. Costa Rica, San José Province, Acosta clusters (3–7 spores per cluster versus 10–20 spores in 
County, next to the road along route 301, about 1.5 km the original description) and the non-piriform, ovoid 
NE of Caspirola (09.6882° N, 084.2736° W, ca 500 m.), or conical shape of the spores, but subglobose or even 
Carlos Rojas, 20 January 2015, from a moist chamber cul- somewhat flat on the outer side. The peridium of the 
ture prepared with ground litter, in a seasonal premontane Costa Rican specimen is ornamented with dense papil-
moist forest, deposited in the Myxogastrid Biorepository lae by transmitted light, a feature not mentioned in the 
at the Engineering Research Institute of University of original description. 
Costa Rica, Ro-6476. The specimen of the present study was found using 
Description. Sporocarps aggregated to grouped, stalked, the moist chamber technique, in a similar manner to the 
erect, 1.3–2.5 mm in total height. Sporotheca cylindrical, original description. However, ground litter was used 
with rounded apex and base, grayish brown to black- instead of bark. The area where the plant material was 
ish (62.d. gy. Br-65. br Black), 0.3–0.4 mm diameter. collected shows seasonality and it is under the influence 
Hypothallus membranous, confluent, common to several of occasional tropical storms (with strong precipitation), 
sporocarps. Stalk cylindrical, slightly attenuate toward given its short linear distance to the coast. Due to the 
the apex, 0.3–0.5 mm long, blackish (65. br black), hol- rugged topography and sparse population, the area is not 
low, opaque above, slightly pale at the base. Peridium heavily influenced by human activities and thus patches 
evanescent except at the base of the sporotheca where it of forest patches are mostly intact.
516 Check List 14 (3)
Figures 26–36. Stemonaria rufipes Nann.-Bremek. & Y. Yamam. (Ro-6476). 26. Sporocarp. 27. Detail of the apex of the sporotheca showing the 
columella and capillitium. 28. Base of the stalk. 29. Remain of the peridium around the stalk. 30–31. Clusters of spores under LM. 32. Detail 
of the apex of the sporotheca showing the capillitial threads and the membranous expansion of the columella under SEM. 33. Capillitium 
sparingly joined to the columella along its whole length, the primary branches sometimes with little membranous expansions at the connec-
tion with the columella, the secondary branches united at the periphery, but not forming a surface net under SEM. 34–36. Spores clustered, 
three to five spores to a cluster by SEM. Scale bars 26 = 1 mm; 27–28 = 100 µm; 29 = 50 µm; 30–31 = 10 µm; 32–33 = 100 µm; 34–36 = 10 µm.
Discussion albeit hard to document, host many interesting myxo-mycetes for the development of taxonomic studies. Even 
The new records presented here widely broaden the pre- though molecular techniques may provide important 
viously reported distributions of these 4 species. Their information about the identity of species making up a 
presence in the Mesoamerican Biodiversity Hotspot community, the complete characterization of species still 
provides evidence to demonstrate that tropical areas, requires morphological examination and careful descrip-
Lado et al. | Neotropical myxomycetes  517
tion of the material under examination. References
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was funded by the Spanish Government through the Lado C, Wrigley Basanta D de (2008) A review of Neotropical myxo-
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particularly for field trips, was obtained by University Diversity 59: 3–32. https://doi.org/10.1007/s13225-012-0159-8.C  
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