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The Free Library > Science and Technology > Biological sciences > Acta Botanica Mexicana > October 1, 2016
The Free Library > Date > 2016 > October > 1 > Acta Botanica Mexicana
Cambios en la composicion de la comunidad fungica de biopeliculas sobre roca calcarea a traves de una cronosecuencia en Campeche, Mexico.
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Changes in fungal community composition of biofilms on limestone across a chronosequence in Campeche, Mexico
It is well known that rocks, either in natural geological settings or as part of monuments, are common habitats for a wide range of microorganisms (Scheerer et al., 2009; Miller et al., 2012). The colonization of lithic substrates by microbial communities is influenced by the properties of the substrate, such as porosity, surface roughness and mineralogical composition -bioreceptivity- (see review in Miller et al., 2012), in addition to climate and microclimatic conditions (Guillitte, 1995; Ortega-Morales et al., 1999; Gaylarde and Gaylarde, 2005; Barberousse et al., 2006). Furthermore, communities of microorganisms growing on lithic substrates, including fungi, may respond differentially to environmental conditions over time based on their ecophysiological requirements (Scheerer et al., 2009; Mihajlovski et al., 2014).
In the tropics and subtropics, rocks are capable of being colonized by microorganisms due to high levels of relative humidity and particular bioreceptivity of the limestone (Kumar and Kumar, 1999; Gomez-Cornelio et al., 2012). Gaylarde and Gaylarde (2005) found that the macro- and micro-environments of different geographical regions play an important role in the biomass and composition of the microorganism groups that compose biofilms. For example, microbial biomass in Latin America is dominated by cyanobacteria and fungi, while in Europe phototrophs, including algae and cyanobacteria, are the most common organisms. Furthermore, the development of biofilms on rocks represents an important stage in the primary succession of terrestrial ecosystems (Chertov et al., 2004; Gorbushina, 2007). In this process, fungi that form part of biofilms physically and chemically deteriorate rock, and thus actively participate in the formation of protosoil and minerals and also accelerate this process, enabling subsequent colonization of the substrate by mosses, lichens or plants (Gorbushina and Krumbein, 2000; Sterflinger, 2000; Gadd, 2007). Although molecular techniques are commonly used to study communities in the field of environmental microbiology, the traditional techniques of isolation and identification of fungi are of vital importance in order to phenotypically characterize fungi and to determine their role on epilithic substrates (Ruibal et al., 2005; Gleeson et al., 2010).
Fungal epilithic communities have been studied in a wide range of environments and for several lithotypes (Sterflinger and Krumbein, 1997; Sterflinger and Prillinger, 2001; Urzi et al., 2001; Gorbushina et al., 2002; Ruibal et al., 2005; Ruibal et al., 2009; Tang and Lian, 2012). However, most research has not considered the influence of time on the colonization patterns of fungal communities. One exception was the study of Lan et al. (2010), in which fungal communities of young and old biofilms on sandstone were found substantially different. Furthermore, the importance of filamentous fungi as rock colonizers and their ecological role in environments are not well-understood, especially in tropical and subtropical climates. More attention has been placed on the microcolonial fungi, meristematic fungi and yeasts of temperate climates (Sterflinger and Krumbein, 1997; Gorbushina et al., 2002; Chertov et al., 2004; Gorbushina et al., 2005; Ruibal et al., 2005; 2009; Sterflinger et al., 2012). Therefore, in order to expand our current understanding of the fungal community associated with limestone, we studied the culturable subset of fungi in biofilms exposed to similar environmental conditions and substratum properties. A chronosequence was considered by examining the biofilms of three walls constructed one, five and 10 years ago.
Study area and climate variables
The coastal city of Campeche, Mexico has a subtropical climate and an altitudinal range of 3-10 m. The studied biofilms were relatively categorized as young, middle-aged and old, corresponding to walls that were constructed with limestone rock fragments one, five and 10 years ago, respectively, according to historical documentation (Fig. 1). The colonization of buildings by fungi may initiate shortly after construction but the formation of biofilm usually takes several years (Barberousse et al., 2006; Gomez-Cornelio et al., 2012; Adamson et al., 2013). Hence, the monthly climate data were obtained from the local meteorological observatory in Campeche, in order to calculate the annual means of climate variables as well as the mean conditions corresponding to the number of years since walls were constructed and exposed to the environment. The considered climatic variables were: minimum and maximum temperature, minimum, mean and maximum relative humidity and mean rainfall (Table 1).
Limestone walls with comparable characteristics of exposure to the surrounding environment were chosen. All walls were composed of rock blocks and had similar substratum properties (or bioreceptivity), vertical surfaces and homogeneous coverage of biofilms. In addition, all walls were oriented towards the north where low solar irradiation and high relative humidity prevail in comparison to facades oriented towards other directions (Adamson et al., 2013; Ortega-Morales et al., 2013). Further criteria for selecting the walls included the absence of surrounding vegetation and low levels of human disturbance, such as painting or washing. Sites with automobile traffic or post-construction remodeling works were avoided. Samples were taken from above the height of one meter in order to avoid confounding factors such as potential microbial colonization due to splashing water. Hence, time elapsed since construction of the walls and establishment of biofilms was the main influential variable considered in the analysis of fungal community structure.
Biofilm sampling and fungal isolation
In the dry season of January 2014, we sampled biofilms on the three selected limestone surfaces. Mean maximum and minimum temperatures in January were 34 [degrees]C and 11.4 [degrees]C, respectively. Mean rainfall was 23 mm, and the mean relative humidity was 83%. On each wall, we scraped three surface areas of 3 x 3 cm to a maximum depth of 3 mm, using a sterile scalpel. Scraping was performed by the same person to avoid bias. The biomasses of the biofilms scraped from the wall were placed in sterile Petri dishes and transferred to the laboratory for processing.
Fungi were isolated by washing and filtration of particles technique (Bills et al., 2004). One gram of each scraped biofilm was placed in a washing apparatus with micro-sieves with pores of 250, 125, 100 and 75 [micro]m (MINI-SIEVE INSERT-ASTD, Bel-Art Products, Pequannock, New Jersey, USA), and was consequently washed and filtered for 10 min using bi-distilled water. This technique reduces the isolation of propagules from spores, favoring only the isolation of fungi attached to rock particles (Bills et al., 2004; Arias-Mota and Here dia-Abarca, 2014) that may present active bioweathering or serve a protective function on the surface of the limestone.
Particles trapped on the 75 [micro]m sieve were transferred to sterile filter paper and incubated for 24 h at 27 [degrees]C to remove excess water. In order to isolate the greatest number of species, we used two culture media: a copiotrophic medium composed of 2% malt extract, 2% agar and 0.2% CaC[O.sub.3] (MEAC) and an oligotrophic medium of 0.2% CaC[O.sub.3] and 2% agar (CCOA). Both media were adjusted to pH 7.7 and supplemented with chloramphenicol (200 mg [L.sup.-1]) to inhibit the growth of bacteria. Media were prepared with CaC[O.sub.3], since it is the main component of limestone (Burford et al., 2003). Under a stereomicroscope, 50 particles were transferred to 10 plates with MEAC (5 particles per plate); this procedure was repeated for the CCOA medium. Plates were incubated at 27 [degrees]C in darkness. After the fourth day, plates were inspected daily for a period of four weeks. All fungal colonies that emerged from the particles were purified in inclined tubes with MEAC.
Morphological identification of fungi
The fungal isolates were identified according to macroscopic characteristics, such as coloration, diameter, texture, pigmentation, margin appearance, zonality and production of exudates in the culture medium, in addition to the morphological characteristics of their reproductive and vegetative structures, including color, conidiogenesis, spore type and size. Fungal isolates that sporulated were identified using the taxonomic keys of Booth (1971), Ellis (1971; 1976), Sutton (1980), Pitt (2000), Klich (2002), Boerema et al. (2004), Domsch et al. (2007) and Seifert et al. (2011). The identity of the species with more than eight isolates was confirmed by performing genomic DNA extraction and sequencing the ITS region (data not shown).
Fungal colonies that did not sporulate were inoculated into the following culture media: cornmeal agar, oatmeal agar, potato-carrot agar, Czapek dox agar, potato dextrose agar and V8 agar. Plates were then subjected to cyclical periods of light/darkness (12/12 h) to promote sporulation (Bills et al., 2004) and incubated at 27 [degrees]C. Every fourth day for up to six weeks, plates were checked for signs of reproductive structures. Isolates that not produced spores were separated into morphotaxa, according to their macroscopic and microscopic morphology in the different culture media. All fungal isolates were conserved in malt extract broth supplemented with glycerol (20% [vol/vol]) at -80 [degrees]C; agar plugs with mycelium were conserved in sterile distilled water at room temperature.
The fungal communities of the sampled biofilms were analyzed according to species richness, defined as the number of different fungi species per biofilm, in addition to species abundance as the number of fungal isolates per identified species. The colonization frequency of the particles was determined as number of emerged fungal species (one or two per particle) from particles divided by the number of inoculated particles, multiplied by 100 in order to obtain the percentage of particles with adhered mycelium (Bills et al., 2004). In order to determine the substrates that have been reported for the fungi that were identified at the species level, we used the literature previously employed in the identification of fungi and performed a search in Summon system. Fungal diversity was calculated with the Simpsons (D') and Shannons (H') diversity indices, in addition to the Shannon (J') evenness index, which were performed in the EstimateS 9.1 software (Colwell, 2013). In order to determine the similarity and composition of the fungal species found in the three biofilms, the Jaccard index was calculated, and a Venn diagram was created.
Analysis of fungal composition and diversity
In the mycological analysis 844 isolates were recovered, distributed in 73 genera and 202 species (Table 2). The identified species were grouped as follows: 149 Ascomycota, one Basidiomycota and 52 Mycelia sterilia. Hyphomicetous asexual species of Ascomycota (108 species) dominated, while Coelomycetous species represented 21% of the isolates (38 species). The genera with highest number of species (>4) and isolates (>17) in the fungal community were Aspergillus P. Micheli ex Haller, Cladosporium Link, Curvularia Boedijn, Fusarium Link, Microsphaeropsis Hohn., Myrothecium Tode, Nodulisporium Preuss, Paraconiothyrium Verkley and Phoma Sacc. (Table 2).
We found the largest number of isolates (322) in the middle-aged biofilm, followed by the old and young biofilms with 268 and 254 isolates, respectively. However, species richness and diversity were higher in the old biofilm and lower in the middle-aged biofilm (Table 3). The Shannon evenness index of the old biofilm generated a value close to 1, and in the middle-aged biofilm, a value of 0.47 (Table 3). The Jaccard's similarity index showed a low degree of similarity among the young, middle-aged and old fungal communities inhabiting the biofilms. The resulting index values were similar for the comparisons of young and middle-aged (0.18), young and old (0.16) and middle-aged and old (0.15) biofilms.
Overall, of the 202 species identified, 26% were isolated from inoculated particles in both culture media (52 species), while 36% (73 species) were found exclusively in the oligotrophic medium (CCOA) and 38% in the copiotrophic medium (MEAC). Additionally, in the analysis of colonization frequency of particles from all three biofilms, we observed a high number of particles with adhered mycelia (Table 3). The middle-aged biofilm showed the highest percentage (80%) of colonization; however, most mycelium that emerged from these particles belonged to the species Curvularia lunata (Wakker) Boedijn, Fusarium oxysporum Schltdl. and Fusarium redoleos Wollenw. (Table 2). The young and old biofilms presented a minor colonization frequency of particles (Table 3).
In the old biofilm, a high number of isolated species (35%) was specific to only one of the media, and a slightly lower proportion was present in both media (30 %). In young and middle-aged biofilms, 41% of species were indistinctly isolated from both media (Table 2). Isolates of Lasiodiplodia theobromae (Pat.) Griffon & Maubl. and Nigrospora oryzae (Berk. & Broome) Petch were obtained in the MEAC medium. A high percentage (>70%) of species from the genera Aspergillus, Penicillium Link and Trichoderma Pers. were also isolated. Meanwhile, in the CCOA medium the lithic species Friedmanniomyces simplex Selbmann, de Hoog, Mazzaglia, Friedmann & Onofri (6 isolates) was found as well as Stachybotrys Corda species and a large number of uncommon species (Table 2).
Of the 124 fungi identified at the species level, many were associated with numerous substrates, based on the literature review to determine with which substrates identified fungal species had been previously associated (Fig. 2). Thirty-one species were identified as cosmopolitan and belonged to the genera Cladosporium, Curvularia, Fusarium and Penicillium, including several common species, such as Aureobasidium pullulans (de Bary & Lowenthal) G. Arnaud and Geotrichum candidum Link (Domsch et al., 2007). Most of the identified species (81) have been reported in soil, mainly species of the genera Aspergillus, Microdochium Syd., Monodictys S. Hughes, Myrothecium, Phoma, Scolecobasidium E.V. Abbott and Trichoderma. Many species (78) were also associated with plants, including the genera Alternaria, Colletotrichum Corda, Microsphaeropsis, Monodictys, Myrothecium, Phoma and Sarocladium W. Gams & D. Hawksw. 44 of the identified species have been found in litter, corresponding to the genera of Myrothecium, Scolecobasidium and Stachybotrys. Forty species (genus Monodictys) have been associated with air, and 39 species with wood (genera Nodulisporium and Phlyctema Desm.). Finally, 40 and 31 species have been reported on rocks and in water, respectively.
Species dominance
With respect to species abundance, 61% of all fungal species were isolated only once, and 6% were isolated more than 10 times. The remaining fungi (33%) were isolated from 2 to 9 times. The encountered fungal community was mainly dominated by fungi that contain melanin at some or all of their reproductive stages (149 species). Only 53 species (26%) were found with hyaline structures without pigments (Table 2). In all biofilms, we found that species composition has an approximate ratio of 4:1 of melanized fungi to hyaline species.
In regard to the fungal communities, most co-existing groups of species may be associated with a particular biofilm, characterized by time of exposure of the substrate (limestone) to the environment; this is shown in the Venn diagram (Fig. 3). In the young biofilm, we found 49 exelusive species, including several from the genera Colletotrichum and Coleophoma Hohn.; the species Monodyctis paradoxa (Corda) S. Hughes, Myrothecium cinctum (Corda) Sacc., Phoma adianticola (E. Young) Boerema and Phoma paspali P.R. Johnst. were also prominent. In the middle-aged biofilm, only 32 exclusive species were found, although these were not frequent (> 2 isolates). The old biofilms had the highest number of exclusive species (82), among these species of the genera Aspergillus, Lasiodiplodia Ellis & Everh., Penicillium and Stachybotrys, in addition to Xylariales spp. and Hyphomycete 1.
The Venn diagram shows 16 species in the core group (three biofilms) of the fungal community (Fig. 3 and Table 2). Meanwhile, Cladosporium cladosporioides (Fresen.) G.A. de Vries, Curvularia clavata B.L. Jain, Nigrospora oryzae, Phoma eupyrena Sacc., Phoma herbarum Westend. and Westerdykella minutispora (P.N. Mathur) Gruyter, Aveskamp & Verkley were abundant in the young biofilm. In the middle-aged biofilm Curvularia lunata, Curvularia pallescens Boedijn, Friedmanniomyces simplex, Fusarium oxysporum, Fusarium redoleos and Scolecobasidium constrictum E.V. Abbott, and in the old biofilm Clypeopycnis sp., Curvularia australiensis (Tsuda & Ueyama) Manamgoda, L. Cai & K.D. Hyde and Pestalotiopsis maculans (Corda) Nag Raj were frequent (Fig. 4 and Table 2). Twenty-five species were present in at least two biofilms; the young and old biofilms had the highest number of shared species (12). Paraconiothyrium sp. was dominant and was isolated 19 times in the young biofilm, although its abundance diminished in the middle-aged biofilm. Cladosporium oxysporum Berk. & M.A. Curtis, Cladosporium sphaerospermum Penz. and Nodulisporium puniceum (Cooke & Ellis) Deighton were found at a higher frequency in the young biofilm in comparison to the old biofilm. Curvularia verruculosa Tandon & Bilgrami ex. M.B. Ellis was also more frequent in the middle-aged biofilm than the old biofilm. Meanwhile, the species Nodulisporium sylviforme Deighton and Paraphoma fimeti (Brunaud) Gruyter, Aveskamp & Verkley were more dominant in the old biofilm than in the young biofilm. Finally, Myrothecium roridum Tode was more frequent in the old biofilm than in the middle-aged.
The fungal communities of biofilms occurring on limestone walls, considering time elapsed since wall construction, presented different degrees of microbial colonization. In the 5- and 10-year-old samples, colonization was evident by the green and black biomass of the biofilms that were visibly observed (Fig. 1), which is consistent with the findings of Adamson et al. (2013). The phototrophs colonizing such substrates are mainly composed of filamentous cyanobacteria and cocoidal bacteria (Scheerer et al., 2009); these have been found on Mayan buildings in the Yucatan peninsula (Ortega-Morales et al., 1999; 2013). The metabolic products of these organisms provide nutritional support for the establishment of heterotrophic communities, including fungi, allowing for their colonization (De la Torre et al., 1991).
The fungal composition was dominated by species of the Ascomycota class, which are common in rock substrates; these Hyphomicetous asexual species are able to colonize rocks during the first year of exposure (Ruibal et al., 2008; Gleeson et al., 2010; Hallman et al., 2011; Gomez-Cornelio et al., 2012). In contrast, Coelomycetous species are known to colonize limestone in Mediterranean regions (Wollenzien et al., 1995). The identification of only one taxon belonging to Basidiomycota may indicate that this group of organisms is not common in biofilms developing on limestone, as confirmed by Tang and Lian (2012), who used culture-independent methods; however, species of this group may exist in sterile form. The genera Aspergillus, Cladosporium, Curvularia, Fusarium, Microsphaeropsis, Myrothecium, Nodulisporium, Paraconiothyrium and Phoma are common in subtropical environments and in this study (Table 2). For other regions, some of the most common, dominant genera that have been reported on epilithic substrates include Alternaria, Cladosporium, Fusarium, Penicillium, Phoma and Trichoderma (Wollenzien et al., 1995; Kumar and Kumar, 1999; Gorbushina and Krumbein, 2000; Urzi et al., 2001; Gorbushina et al., 2002). All these genera were also identified in this study, but only Cladosporium, Fusarium and Phoma were common and dominant. These differences may be dictated by environmental factors and the bioreceptivity of rocks.
Some isolates showed no reproductive structures (8%) and were identified as Mycelia sterilia (25% of morphospecies). However, few isolates belonged to these morphospecies, indicating that they are rare in the community (Table 2). The finding of sterile mycelia commonly occurs in microbiological studies of other substrates (Arias-Mota and Heredia-Abarca, 2014; Rocha et al., 2014; Khirilla et al., 2015), including limestone substrates (Gomez-Cornelio et al., 2012). These fungi may be classified according to morphotype based on their morphological characteristics (Paulus et al., 2003). The production of less complex structures and poorly elaborated reproduction systems may represent an adaptation strategy in order to conserve energy on certain substrates (Ruibal et al., 2005) due to lack of nutrients or water.
In this study, greater species richness was found in the old biofilm. However, in another study on sandstone, the highest number of species was reported for fresh biofilms in comparison to older biofilms (Lan et al., 2010). Meanwhile, in this study the Shannon evenness index showed that the species identified from isolates were almost equitable in the old biofilm. The lowest evenness was obtained in the middle-aged biofilm, probably due to the presence of several dominant species, such as Curvularia lunata and Fusarium redoleos (Table 3). We found higher diversity values in comparison to other studies on fungal communities colonizing distinct substrates of plant litter and endophytes (Collado et al., 2007; Reverchon et al., 2010). This diversity may be due to the establishment and accumulation of propagules on bare rock at a constant and rapid speed. For example, an increase of 9 x [10.sup.2] to 7.5 x [10.sup.5] colony-forming units was documented in only 11 weeks (Gorbushina and Krumbein, 2000). Under ideal environmental conditions, a high number of fungi could colonize and grow on limestone.
These results, in addition to those of a previous study that investigated the fungal communities on bare limestone (Gomez-Cornelio et al., 2012), suggest that the species richness of limestone in subtropical environments is high in comparison to other rock surfaces, that have been studied in Europe, for example, and in particular in the Mediterranean (De la Torre et al., 1991; Sterflinger and Prillinger, 2001; Gorbushina et al., 2002; Ruibal et al., 2005; Hallman et al., 2011). Although the intrinsic characteristics of rock, such as its mineral composition, porosity and roughness, have been reported to influence the colonization of microbial communities (Guillitte, 1995; Burford et al., 2003; Lan et al., 2010), the study of Tomaselli et al. (2000) did not find a relationship among existing organisms and the petrographic characteristics of rock. In this study, the high values of species richness and diversity may be attributed to favorable environmental factors in the subtropics (Table 1) and time elapsed since initial colonization (Gaylarde and Gaylarde, 2005; Mihajlovski et al., 2014).
Furthermore, the Jaccard's similarity index showed low values of similarity among fungal communities corresponding to different ages. This is notable in considering that the middle-aged and old biofilms were located less than 1 km from each other. These biofilms also had a similar chromatic aspect; therefore, one might expect a high degree of similarity. Meanwhile, the young biofilm was located at an approximate distance of 10.5 kilometers from the other biofilms and had a visible although incipient colonization of microorganisms. Microclimatic differences present at each site may contribute towards the formation of a unique and particular mycobiota (Mihajlovski et al., 2014). Our results differed from those reported for sandstone, in which no differences in eukaryotic composition were found between fresh and young biofilms (Lan et al., 2010). However, to the contrary, on a serpentine substrate fungal communities were found to have low similarity (Daghino et al., 2012). These findings highlight that the community composition and diversity of fungi are not always determined by substrate bioreceptivity.
Based on the fungi that emerged from the analyzed particles that were scraped from biofilms on limestone, we were able to evaluate the fungal network and find a high proportion of active mycelium. The use of an oligotrophic culture medium (CCOA) allowed for the characterization of the cultivable fungi, which represented a complete, diverse and functional community (Ruibal et al., 2005). Although many of these species require water to grow, such as species of the Stachybotrys genus (Jain et al., 2009), the products of the extracellular matrix and the retention of water in rock pores could allow for the growth of water-demanding species. A large variety of specialized fungi were observed in this study; these species could represent an autochthonous community specific to limestone, a substrate with limited nutrients. However, due to the intrinsic environmental conditions of the tropics, nutrients may be transported by air and deposited on the rock as dust (Kumar and Kumar, 1999; Gorbushina and Krumbein, 2000; Ruibal et al., 2009), thereby providing conditions for the colonization of specialized fungal species in the production of exopolymers allowing fungi to adhere to the surfaces; some fungi are also capable of biomineralizing limestone. Furthermore, many fungi are able to successfully establish on limestone since they precipitate calcium; these represent the main sink of toxic forms of calcium in the soil or other environments (Sterflinger, 2000). Hence, fungi are essential members of the microbial communities that develop in the biofilms of limestone.
The composition of fungal communities may also be influenced by surrounding substrates (Urzi et al., 2001; Hallman et al., 2011). As previously mentioned, we identified the substrates previously reported in the literature for the 124 fungi that were identified in our study at the species level. A high proportion of fungi may have originated from soil or plants, and to a lesser extent, from decomposing litter and/or wood. Also, these fungal species could come from other rocks or from spores suspended in air or water (Fig. 2). However, the establishment and development of fungi over a period of time may be determined by the interactions of species with their environmental conditions, such as relative humidity and temperature (Gorbushina and Krumbein, 2000; Gorbushina et al., 2005), in addition to the bioreceptivity of the substrate. The variety of fungi reported for other surrounding substrates confirms that a large quantity of propagules may potentially reach and colonize rock surfaces. Therefore, according to our findings, limestone surfaces could act as a reservoir of fungal species and function as a fungal source via the dispersion of species under ideal conditions.
Most species isolated from biofilms occurred only once or twice, and few species showed dominance. The isolation technique used in this study may promote the growth of rare and uncommon species; these results do concur with those found by other authors who used both taxonomic and molecular identification techniques (Collado et al., 2007; Ruibal et al., 2008; Gomez-Cornelio et al., 2012). Additionally, the abundance of dominant species of the three analyzed biofilms was variable and may be determined by time of exposure, similar to the variations observed in Lan et al. (2010) in the microorganism community of old and fresh biofilms.
The fungal community contained melanin at some or all of their reproductive stages (74% of the species); this concurs with the reports of other authors, in which the dominant fungi isolated from monuments also contained pigmentation (Sterflinger and Krumbein, 1997; Gorbushina et al., 2002; Lan et al., 2010). Pigmentation in fungi may have different functions or result from the environmental conditions experienced on the rock surface (Scheerer et al., 2009; Hallman et al., 2011). Therefore, melanized communities of fungi have been shown to occur at a high frequency on rock surfaces, although composition may differ from one community to another, as mentioned in the results.
Moreover, fungal propagules are capable of quickly colonizing rock surfaces. Gorbushina and Krumbein (2000) have suggested that fluctuations in environmental conditions, such as those that lead to deficits in nutrients and water on the rock surfaces, promote changes in the diversity of the fungal community. In our results the environmental conditions and the characteristics of the limestone substrate of the biofilms were similar; thus one would expect to find a relationship among the studied fungal communities. However, it may be necessary to study the roles of dominant species during succession of fungal communities. For example, it has been shown that the hyphae of common fungal species biomineralize the surface of the limestone (Burford et al., 2006) and thus lead to subsequent changes. In this study the composition of the fungal community is likely determined by time of exposure to the environment and species interactions, which may then facilitate or inhibit colonization by other species, leading to changes in the composition of the fungal chronosequence associated with limestone (Fryar, 2002). During this process, the functional properties of fungi on the limestone may also be potentially affected.
In this study, the fungal communities immersed in biofilms were different at each stage of development, defined by the time of exposure of the limestone substrate to the environment. Although the mineral composition of rock substrates has been found to have a certain degree of influence on the structure of fungal communities, in this study limestone samples with similar characteristics of bioreceptivity, such as rock color, roughness and porosity, were selected. Environmental conditions were also similar across sites in the city of Campeche, Mexico. Therefore, in addition to time of exposure, the differentiation in the community structure and diversity of fungi in this study may be determined by the interactions among the species of each biofilm; this should be confirmed by subsequent studies. A particular species composition was isolated in each biofilm corresponding to a different developmental stage, although a common pool of hyaline and melanized fungi appear to colonize rock with great success and may have specific functions on the rock substrate. In future studies, biotic factors, including interactions among bacterial, fungal and algal species, should be studied in order to determine their influence on the structure of the fungal community.
We are grateful to Julio C. Rojas Leon and Hugo Perales Rivera for their comments on an early version of the manuscript. This research was supported by institutional funding from El Colegio de la Frontera Sur and the Universidad Autonoma de Campeche. We extend our thanks to the Comision Nacional del Agua for the provision of meteorological data and to the Consejo Nacional de Ciencia y Tecnologia for the doctoral scholarship awarded to S.G.C. We thank two anonymous reviewers and the editor of the manuscript for suggested improvements; also thanks to Allison Marie Jermain for reviewing the English version of the manuscript.
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Sergio Gomez-Comelio (1,4), Otto Ortega-Morales (2), Alejandro Moron-Rios (1), Manuela Reyes-Estebanez (2), Susana de la Rosa-Garcia (3)
(1) El Colegio de la Frontera Sur, Av. Rancho poligono 2A, Parque Industrial Lerma, 24500 Campeche, Mexico.
(2) Universidad Autonoma de Campeche, Departamento de Microbiologia Ambiental y Biotecnologia, Avenida Agustin Melgar s/n, 24039 Campeche, Mexico.
(3) Universidad Juarez Autonoma de Tabasco, Division Academica de CienciasBiologicas, Carretera Villahermo-sa-Cardenas km 0.5 s/n, entronque a Bosques de Saloya, 86150 Villahermosa, Tabasco, Mexico.
(4) Author for correspondence: sgomez@ecosur.edu.mx
To cite as:
Gomez-Cornelio, S., O. Ortega-Morales, A. Moron-Rios, M. Reyes-Estebanez y S. de la Rosa-Garcia. 2016. Changes in fungal community composition of biofilms on limestone across a chronosequence in Campeche, Mexico. Acta Botanica Mexicana 117: 59-77.
Received: 28 de marzo de 2016.
Reviewed: 6 de julio de 2016.
Accepted: 2 de septiembre de 2016.
Table 1: Geographical location, color and climatic parameters of
biofilms developed on the surface of sampled limestone walls. (Values
are expressed as means [+ or -] 1SD).
Young biofilm
Geographical location                         19[degrees]49'30.7"N
90[degrees]32'51.2"W
Mean temperature ([degrees]C)   Minimum         16.5 [+ or -] 2.5
Maximum         37.1 [+ or -] 4.3
Mean relative humidity (%)      Minimum         43.8 [+ or -] 10.7
Mean            78.6 [+ or -] 5.6
Mean rainfall (mm)              Rainy season   224.9 [+ or -] 11.3
Dry season      20.4 [+ or -] 19
Degree of colonization (visual inspection)     Not observed
Middle-aged biofilm
Geographical location                          19[degrees]49'30.5"N
Mean relative humidity (%)      Minimum         39.9 [+ or -] 9.4
Mean              75 [+ or -] 5.8
Mean rainfall (mm)              Rainy season   192.6 [+ or -] 54
Degree of colonization (visual inspection)     Dark green
Old biofilm
90[degrees]33'14"W
Degree of colonization (visual inspection)     Black
Table 2: Epilithic fungal community in terms of abundance of species
isolated from biofilms on limestone at different stages of development
and color of their reproductive structures. (M: melanized and H:
hyaline).
(a) Copiotrophic medium (MEAC).
(b) Oligotrophic medium (CCOA).
* Mycelia sterilia with one or more isolates, which are added in the
Epilithic fungi                      Coloration   Young     Middle-
biofilm   aged
Sondariafimicola (Roberge ex         M                      2
Desm.) Ces. & De Not.
Xylariales sp. 1                     M
Coelomycetous asexual species of
Ascochyta carpathica (Allesch.)      M
Keissl.
Clypeopycnis sp.                     M            2         1
Coleophoma sp.                       M            7
Colletotrichum crassipes (Speg.)     H            1
Colletotrichum dematium (Pers.)      M            1
Colletotrichum gloeosporioides       H            1
(Penz.) Penz. & Sacc.
Coniothyrium multiporum (V.H.        M                      1
Pawar, P.N. Mathur &
Thirum.) Verkley & Gruyter         M
Cytospora polygoni-sieboldii Henn.
Lasiodiplodia theobromae (Pat.)      M
Griffon & Maubl.
Microsphaeropsis arundinis (S.       M                      1
Ahmad) B. Sutton
Microsphaeropsis sp. 1               M
Microsphaeropsis sp. 2               M                      2
Neosetophoma samararum (Desm.)       M
Gruyter, Aveskamp & Verkley
Paraconiothyrium sp.                 M            19        3
Paraphoma chrysanthemicola           M            1
(Hollos) Gruyter, Aveskamp &
Verkley
Paraphomafimeti (Brunaud) Gruyter,   M            2
Aveskamp & Verkley
Pestalotiopsis maculans (Corda)      M            5         3
Nag Raj
Peyronellaea aurea (Gruyter,         M            2
Noordel. & Boerema) Aveskamp,
Gruyter & Verkley
Peyronellaea gardeniae (S. Chandra   M            1
& Tandon) Aveskamp, Gruyter &
Phlyctema lappae (P. Karst.) Sacc.   M            1
Phoma adianticola (E. Young)         M            4
Boerema
Phoma crystallifera Gruyter,         M
Noordel. & Boerema
Phoma eupyrena Sacc.                 M            23        12
Phoma herbarum Westend.              M            4         1
Phoma heteroderae Sen Y. Chen,       M            2
D.W. Dicks. & Kimbr.
Phoma leveillei Boerema & G.J.       M
Phoma multirostrata (P.N. Mathur,    M                      1
S.K. Menon & Thirum.)
Dorenb. & Boerema
Phoma paspali P.R. Johnst.           M            5
Phoma pratorum P.R. Johnst. &        M                      1
Phoma proteae Crous                  M
Phoma putamina Speg.                 M            1
Phoma sp. 1                          M            1
Phoma tropica R. Schneid. &          M                      2
Phomopsis putator (Nitschke)         M
Traverso
Pleurophomopsis lignicola Petr.      M            1         1
Pyrenochaetopsis pratorum (Berk.     M            1
& M.A. Curtis) M.B. Ellis
Westerdykella minutispora (P.N.      M            4         1
Mathur) Gruyter, Aveskamp &
Hyphomicetous asexual species of
Acremoniella velutina (Fuckel)       M            1
Sacc.
Acremonium brachypenium W. Gams      H                      2
Acremonium fusidioides (Nicot) W.    H
Gams
Acremonium rutilum W. Gams           H
Acremonium sordidulum W. Gams &      H
D. Hawksw.
Agaricodochium sp.                   H            1
Alternaria longipes (Ellis &         M
Everh.) E.W. Mason
Alternaria tenuissima (Kunze)        M            1         1
Arxiella terrestris Papendorf        H
Aspergillus aculeatus Iizuka         M
Aspergillus alliaceus Thom &         H
Aspergillus awamori Nakaz.           M
Aspergillus foetidus Thom & Raper    M
Aspergillus fumigatus Fresen.        M
Aspergillus japonicus Saito          M
Aspergillus niger Tiegh.             M
Aureobasidium pullulans (de Bary     M
& Lowenthal) G. Arnaud
Badarisama sp.                       M                      1
Baudoinia sp.                        M
Calcarisporium sp.                   M            1
Capnobotryella antalyensis Sert &    M
Sterfl.
Chaetasbolisia falcata V.A.M.        M            1
Mill. & Bonar
Chromelosporium sp.                  H
Cladosporium cladosporioides         M            22        10
(Fresen.) G.A. de Vries
Cladosporium oxysporum Berk. &       M            10
M.A. Curtis
Cladosporium sphaerospermum Penz.    M            9
Cladosporium tenuissimum Cooke       M            2
Corynespora citricola M.B. Ellis     M            1
Corynesporella pinarensis R.F.       M
Curvularia australiensis (Tsuda &    M            1         1
Ueyama) Manamgoda, L.
Cai & K.D. Hyde Curvularia           M                      2
brachyspora Boedijn
Curvularia clavata B.L. Jain         M            18        3
Curvularia fallax Boedijn            M            1
Curvularia hawaiiensis (Bugnic.      M                      1
ex M.B. Ellis) Manamgoda,
L. Cai & K.D. Hyde Curvularia        M            10        105
lunata (Wakker) Boedijn
Curvularia pallescens Boedijn        M            6         13
Curvularia spicifera (Bainier)       M            1
Boedijn
Curvularia verruculosa Tandon &      M                      7
Bilgrami ex. M.B. Ellis
Curvularia sp.                       M
Echinocatena sp.                     M
Exochalara longissima (Grove) W.     M
Gams & Hol.-Jech.
Friedmanniomyces simplex Selbmann,   M            1         3
de Hoog, Mazzaglia,
Friedmann & Onofri
Fusarium camptoceras Wollenw. &      H            1
Reinking
Fusarium equiseti (Corda) Sacc.      H            1
Fusarium flocciferum Corda           H            1         1
Fusarium incarnatum (Desm.) Sacc.    H
Fusarium oxysporum Schltdl.          H            8         40
Fusarium redolens Wollenw.           H            14        41
Fusarium sacchari (E.J. Butler &     H            1
Hafiz Khan) W. Gams
Fusarium solani (Mart.) Sacc.        H            2
Fusarium subglutinans (Wollenw. &    H            1
Reinking) P.E. Nelson,
Toussoun & Marasas
Fusarium tabacinum (J.F.H. Beyma)    H                      1
W. Gams
Fusarium ventricosum Appel &         H            1
Wollenw.
Gabarnaudia sp.                      H
Geotrichum candidum Link             H
Gilmaniella subornata Morinaga,      M                      1
Minoura & Udagawa
Graphium penicillioides Corda        M
Hyphomycete 1                        M
Microdochium dimerum (Penz.) Arx     H                      1
Microdochium nivale (Fr.) Samuels    H            2
& I.C. Hallett
Monodictys fluctuata (Tandon &       M            1
Bilgrami) M.B. Ellis
Monodictys paradoxa (Corda) S.       M            3
Myrothecium cinctum (Corda) Sacc.    M            3
Myrothecium roridum Tode             M                      12
Myrothecium sp. 1                    M            1         3
Myrothecium sp. 2                    M                      1
Nalanthamala madreeya Subram.        H                      1
Nigrospora oryzae (Berk. & Broome)   M            3         1
Nodulisporium acervatum (Massee)     M
Nodulisporium ochraceum Preuss       M            1
Nodulisporium puniceum (Cooke &      M            3
Ellis) Deighton
Nodulisporium radians (Berk.)        M
Nodulisporium sp. 1                  M            1
Nodulisporium sp. 2                  M
Nodulisporium sylviforme Deighton    M            1
Nodulisporium thelenum (Sacc.) G.    M            1
Sm.
Ochroconis tshawytschae (Doty &      M
D.W. Slater) Kiril. &
Al-Achmed
Penicillium citreonigrum Dierckx     H
Penicillium dierckxii Biourge        H
Penicillium islandicum Sopp          H
Penicillium oxalicum Currie &        H
Periconia igniaria E.W. Mason &      M            1
M.B. Ellis
Periconiella mucunae M.B. Ellis      M
Prathoda longissima (Deighton &      M            1
MacGarvie) E.G.
Pseudohelicomyces albus Gamica &     H
E. Valenz.
Pseudopithomyces chartarum (Berk.    M            1
& M.A. Curtis) J.F. Li,
Ariyawansa & K.D. Hyde
Pseudoramichloridium brasilianum     M                      1
(Arzanlou & Crous)
Cheew. & Crous
Ramichloridium apiculatum (J.H.      M                      2
Mill., Giddens & A.A. Foster)
de Hoog
Sarocladium kiliense (Grutz)         H                      1
Summerb.
Sarocladium strictum (W. Gams)       H            1
Scolecobasidium constrictum E.V.     M            3         4
Sepedonium sp.                       H
Stachybotrys microspora (B.L.        M
Mathur & Sankhla) S.C. Jong
& E.E. Davis
Stachybotrys nephrospora Hansf.      M
Stachybotrys renispora P.C. Misra    M
Tolypocladium sp.                    H            1
Torula herbarum (Pers.) Link         M
Trichobotrys sp.                     M                      1
Trichocladium sp.                    M                      1
Trichoderma aggressivum Samuels &    H            1
Trichoderma harzianum Rifai          H                      2
Trichoderma longibrachiatum Rifai    H
Trichoderma ovalisporum Samuels      H
& Schroers
Trichoderma strigosum Bissett        H
Veronaea musae M.B. Ellis            M                      1
Verruconis verruculosa (R.Y. Roy,    M                      1
R.S. Dwivedi & R.R. Mishra)
Samerp. & de Hoog
Geotrichopsis sp.                    H
Mycelia sterilia
Mycelia sterilia (Morphotaxon 01)    H            1         1
Mycelia sterilia (Morphotaxon 02)    M            2
Mycelia sterilia (Morphotaxon        M            1
06-10 *)
Mycelia sterilia (Morphotaxon 16)    H            1
Mycelia sterilia (Morphotaxon        M                      3
Mycelia sterilia (Morphotaxon 21)    M                      2
Mycelia sterilia (Morphotaxon        H                      1
Mycelia sterilia (Morphotaxon 27)    H
Mycelia sterilia (Morphotaxon 28)    M
Mycelia sterilia (Morphotaxon        M
Mycelia sterilia (Morphotaxon        H
Epilithic fungi                      Old       Total
Sondariafimicola (Roberge ex         1         3 (a,b)
Xylariales sp. 1                     1         1 (b)
Xylariales sp. 2                     1         1 (a)
Ascochyta carpathica (Allesch.)      2         2 (a)
Clypeopycnis sp.                     3         6 (a,b)
Coleophoma sp.                                 7 (a,b)
Colletotrichum crassipes (Speg.)               1 (a)
Colletotrichum dematium (Pers.)                1 (a)
Colletotrichum gloeosporioides                 1 (b)
Coniothyrium multiporum (V.H.                  1 (b)
Thirum.) Verkley & Gruyter           1         1 (a)
Cytospora polygoni-sieboldii
Henn.
Lasiodiplodia theobromae (Pat.)      7         7 (a)
Microsphaeropsis arundinis (S.       2         3 (a,b)
Microsphaeropsis sp. 1               1         1 (b)
Microsphaeropsis sp. 2                         2 (a,b)
Neosetophoma samararum (Desm.)       1         1 (a)
Paraconiothyrium sp.                           22 (a,b)
Paraphoma chrysanthemicola                     1 (b)
Paraphomafimeti (Brunaud) Gruyter,   3         5 (a,b)
Pestalotiopsis maculans (Corda)      10        18 (a,b)
Peyronellaea aurea (Gruyter,                   2 (a,b)
Peyronellaea gardeniae (S. Chandra             1 (b)
Phlyctema lappae (P. Karst.) Sacc.             1 (a)
Phoma adianticola (E. Young)                   4 (a)
Phoma crystallifera Gruyter,         1         1 (b)
Phoma eupyrena Sacc.                 15        50 (a,b)
Phoma herbarum Westend.              2         7 (a,b)
Phoma heteroderae Sen Y. Chen,                 2 (a,b)
Phoma leveillei Boerema & G.J.       1         1 (b)
Phoma multirostrata (P.N. Mathur,    1         2 (b)
Phoma paspali P.R. Johnst.                     5 (a)
Phoma pratorum P.R. Johnst. &                  1 (a)
Phoma proteae Crous                  1         1 (b)
Phoma putamina Speg.                 2         3 (a,b)
Phoma sp. 1                                    1 (a)
Phoma sp. 2                                    2 (a,b)
Phoma tropica R. Schneid. &                    2 (a,b)
Phomopsis putator (Nitschke)         1         1 (a)
Pleurophomopsis lignicola Petr.                2 (a)
Pyrenochaetopsis pratorum (Berk.               1 (b)
Westerdykella minutispora (P.N.      1         6 (a,b)
Acremoniella velutina (Fuckel)                 1 (a)
Acremonium brachypenium W. Gams                2 (b)
Acremonium fusidioides (Nicot) W.    1         1 (b)
Acremonium rutilum W. Gams           2         2 (b)
Acremonium sordidulum W. Gams &      1         1 (a)
Agaricodochium sp.                             1 (a)
Alternaria longipes (Ellis &         1         1 (a)
Alternaria tenuissima (Kunze)                  2 (a,b)
Arxiella terrestris Papendorf        1         1 (b)
Aspergillus aculeatus Iizuka         2         2 (a)
Aspergillus alliaceus Thom &         1         1 (a)
Aspergillus awamori Nakaz.           2         2 (a)
Aspergillus foetidus Thom & Raper    2         2 (a)
Aspergillus fumigatus Fresen.        2         2 (a)
Aspergillus japonicus Saito          1         1 (a)
Aspergillus niger Tiegh.             7         7 (a)
Aureobasidium pullulans (de Bary     1         1 (a)
Badarisama sp.                                 1 (a)
Baudoinia sp.                        1         1 (b)
Calcarisporium sp.                             1 (a)
Capnobotryella antalyensis Sert &    1         1 (a)
Chaetasbolisia falcata V.A.M.                  1 (b)
Chromelosporium sp.                  1         1 (a)
Cladosporium cladosporioides         8         40 (a,b)
Cladosporium oxysporum Berk. &       8         18 (a,b)
Cladosporium sphaerospermum Penz.    2         11 (a-b)
Cladosporium tenuissimum Cooke       2         4 (a,b)
Corynespora citricola M.B. Ellis               1 (a)
Corynesporella pinarensis R.F.       1         1 (b)
Curvularia australiensis (Tsuda &    5         7 (a,b)
Cai & K.D. Hyde Curvularia                     2 (a)
Curvularia clavata B.L. Jain         1         22 (a,b)
Curvularia fallax Boedijn                      1 (b)
Curvularia hawaiiensis (Bugnic.                1 (b)
L. Cai & K.D. Hyde Curvularia        26        141 (a,b)
Curvularia pallescens Boedijn        11        30 (a,b)
Curvularia spicifera (Bainier)                 1 (a)
Curvularia verruculosa Tandon &      1         8 (a,b)
Curvularia sp.                       2         2 (a,b)
Echinocatena sp.                     1         1 (b)
Exochalara longissima (Grove) W.     2         2 (a)
Friedmanniomyces simplex Selbmann,   2         6 (b)
Fusarium camptoceras Wollenw. &                1 (b)
Fusarium equiseti (Corda) Sacc.                1 (a)
Fusarium flocciferum Corda                     2 (a,b)
Fusarium incarnatum (Desm.) Sacc.    1         1 (b)
Fusarium oxysporum Schltdl.          4         52 (a,b)
Fusarium redolens Wollenw.           9         64 (a,b)
Fusarium sacchari (E.J. Butler &               1 (a)
Fusarium solani (Mart.) Sacc.        2         4 (a,b)
Fusarium subglutinans (Wollenw. &              1 (b)
Fusarium tabacinum (J.F.H. Beyma)              1 (b)
Fusarium ventricosum Appel &                   1 (a)
Gabarnaudia sp.                      1         1 (b)
Geotrichum candidum Link             2         2 (a,b)
Gilmaniella subornata Morinaga,                1 (a)
Graphium penicillioides Corda        2         2 (a,b)
Hyphomycete 1                        11        11 (a,b)
Microdochium dimerum (Penz.) Arx     2         3 (b)
Microdochium nivale (Fr.) Samuels              2 (b)
Monodictys fluctuata (Tandon &                 1 (b)
Monodictys paradoxa (Corda) S.                 3 (a,b)
Myrothecium cinctum (Corda) Sacc.              3 (a)
Myrothecium roridum Tode             16        28 (a,b)
Myrothecium sp. 1                              4 (a,b)
Myrothecium sp. 2                              1 (a)
Nalanthamala madreeya Subram.                  1 (b)
Nigrospora oryzae (Berk. & Broome)   1         5 (a)
Nodulisporium acervatum (Massee)     1         1 (b)
Nodulisporium ochraceum Preuss                 1 (a)
Nodulisporium puniceum (Cooke &      2         5 (a,b)
Nodulisporium radians (Berk.)        1         1 (b)
Nodulisporium sp. 1                            1 (b)
Nodulisporium sp. 2                  1         1 (a)
Nodulisporium sylviforme Deighton    4         5 (a,b)
Nodulisporium thelenum (Sacc.) G.    1         2 (a,b)
Ochroconis tshawytschae (Doty &      1         1 (b)
Penicillium citreonigrum Dierckx     1         1 (a)
Penicillium dierckxii Biourge        1         1 (b)
Penicillium islandicum Sopp          1         1 (a)
Penicillium oxalicum Currie &        1         1 (a)
Periconia igniaria E.W. Mason &                1 (a)
Periconiella mucunae M.B. Ellis      1         1 (a)
Prathoda longissima (Deighton &                1 (b)
Pseudohelicomyces albus Gamica &     2         2 (a,b)
Pseudopithomyces chartarum (Berk.              1 (a)
Pseudoramichloridium brasilianum               1 (a)
Ramichloridium apiculatum (J.H.                2 (b)
Sarocladium kiliense (Grutz)                   1 (a)
Sarocladium strictum (W. Gams)                 1 (b)
Scolecobasidium constrictum E.V.     1         8 (a,b)
Sepedonium sp.                       1         1 (b)
Stachybotrys microspora (B.L.        1         1 (b)
Stachybotrys nephrospora Hansf.      1         1 (b)
Stachybotrys renispora P.C. Misra    1         1 (b)
Tolypocladium sp.                              1 (b)
Torula herbarum (Pers.) Link         1         1 (a)
Trichobotrys sp.                               1 (a)
Trichocladium sp.                              1 (b)
Trichoderma aggressivum Samuels &    1         2 (a)
Trichoderma harzianum Rifai          1         3 (a,b)
Trichoderma longibrachiatum Rifai    2         2 (a)
Trichoderma ovalisporum Samuels      1         1 (a)
Trichoderma strigosum Bissett        1         1 (a)
Veronaea musae M.B. Ellis                      1 (a)
Verruconis verruculosa (R.Y. Roy,              1 (a)
Geotrichopsis sp.                    1         1 (a)
Mycelia sterilia (Morphotaxon 01)    1         3 (b)
Mycelia sterilia (Morphotaxon 02)              2 (a,b)
Mycelia sterilia (Morphotaxon 03)              2 (b)
Mycelia sterilia (Morphotaxon 05)    1         2 (a,b)
Mycelia sterilia (Morphotaxon                  5 (a)
Mycelia sterilia (Morphotaxon                  5 (b)
Mycelia sterilia (Morphotaxon                  6 (a,b)
Mycelia sterilia (Morphotaxon        1         6 (a)
Mycelia sterilia (Morphotaxon        1         11 (b)
Mycelia sterilia (Morphotaxon        1         2 (b)
Table 3: Abundance, richness, diversity and evenness of the fungal
epilithic community colonizing biofilms on limestone at different
stages of development.
State of      Abundance    Richness     Simpson      Shannon
biofilms      (number of   (number of   Diversity    Diversity
isolates)    species)     index (D')   index (H')
Young         254          83           25.8         3.7
Middle-aged   322          59           6.8          2.7
Old           268          117          37.2         4.1
State of      Shannon      Particles
biofilms      evenness     colonization
index (J')   (%)
Young         0.67         73
Middle-aged   0.47         86
Old           0.75         75
Figure 2: Frequency of substrate types reported for the fungal
species isolated from biofilms according to the literature.
Plant-associated   78
Litter             44
Lithic             40
Air                40
Wood               39
Water              31
Soil               81
Note: Table made from pie graph.
COPYRIGHT 2016 Instituto de Ecologia, A.C.
Title Annotation:
Gomez-Cornelio, Sergio; Ortega-Morales, Otto; Moron-Rios, Alejandro; Reyes-Estebanez, Manuela; de la
Acta Botanica Mexicana
Germinacion y frecuencia de esporofitos de dos especies simpatricas de Blechnum (Blechnaceae).
Nuevos registros del genero Scutellinia (Pyronemataceae, Pezizales) para la micobiota mexicana.
Prosa escolar
Aspectos sociales


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Become greatly intelligent and know the ultimate.
Sa no jushasva dravinena medhe.
And so goddess of wealth, give us wealth and intellect.
For they wear the garlands of lotuses.
* Svaha is the wife of fire God. We give offerings to her and she gives it to fire, who gives it to devas.
Sa maam medha suprathika jushtaam.
Let it come to me with love and favour me.


text:
"""Tests for the Independent distribution."""

from __future__ import absolute_import
from __future__ import division
from __future__ import print_function

import importlib
import numpy as np

from tensorflow.contrib.distributions.python.ops import independent as independent_lib
from tensorflow.contrib.distributions.python.ops import mvn_diag as mvn_diag_lib
from tensorflow.contrib.distributions.python.ops import test_util
from tensorflow.python.ops import math_ops
from tensorflow.python.ops.distributions import normal as normal_lib
from tensorflow.python.platform import test
from tensorflow.python.platform import tf_logging

def try_import(name):  # pylint: disable=invalid-name
  module = None
  try:
    module = importlib.import_module(name)
  except ImportError as e:
    tf_logging.warning("Could not import %s: %s" % (name, str(e)))
  return module

stats = try_import("scipy.stats")

class ProductDistributionTest(
    test_util.VectorDistributionTestHelpers, test.TestCase):

  def testSampleAndLogProbUnivariate(self):
    loc = np.float32([-1., 1])
    scale = np.float32([0.1, 0.5])
    with self.test_session() as sess:
      ind = independent_lib.Independent(
          distribution=normal_lib.Normal(loc=loc, scale=scale),
          reduce_batch_ndims=1)

      x = ind.sample([4, 5])
      log_prob_x = ind.log_prob(x)
      x_, actual_log_prob_x = sess.run([x, log_prob_x])

      self.assertEqual([], ind.batch_shape)
      self.assertEqual([2], ind.event_shape)
      self.assertEqual([4, 5, 2], x.shape)
      self.assertEqual([4, 5], log_prob_x.shape)

      expected_log_prob_x = stats.norm(loc, scale).logpdf(x_).sum(-1)
      self.assertAllClose(expected_log_prob_x, actual_log_prob_x,
                          rtol=1e-5, atol=0.)

  def testSampleAndLogProbMultivariate(self):
    loc = np.float32([[-1., 1], [1, -1]])
    scale = np.float32([1., 0.5])
    with self.test_session() as sess:
      ind = independent_lib.Independent(
          distribution=mvn_diag_lib.MultivariateNormalDiag(
              loc=loc,
              scale_identity_multiplier=scale),
          reduce_batch_ndims=1)

      x = ind.sample([4, 5])
      log_prob_x = ind.log_prob(x)
      x_, actual_log_prob_x = sess.run([x, log_prob_x])

      self.assertEqual([], ind.batch_shape)
      self.assertEqual([2, 2], ind.event_shape)
      self.assertEqual([4, 5, 2, 2], x.shape)
      self.assertEqual([4, 5], log_prob_x.shape)

      expected_log_prob_x = stats.norm(loc, scale[:, None]).logpdf(
          x_).sum(-1).sum(-1)
      self.assertAllClose(expected_log_prob_x, actual_log_prob_x,
                          rtol=1e-6, atol=0.)

  def testSampleConsistentStats(self):
    loc = np.float32([[-1., 1], [1, -1]])
    scale = np.float32([1., 0.5])
    n_samp = 1e4
    with self.test_session() as sess:
      ind = independent_lib.Independent(
          distribution=mvn_diag_lib.MultivariateNormalDiag(
              loc=loc,
              scale_identity_multiplier=scale),
          reduce_batch_ndims=1)

      x = ind.sample(int(n_samp), seed=42)
      sample_mean = math_ops.reduce_mean(x, axis=0)
      sample_var = math_ops.reduce_mean(
          math_ops.squared_difference(x, sample_mean), axis=0)
      sample_std = math_ops.sqrt(sample_var)
      sample_entropy = -math_ops.reduce_mean(ind.log_prob(x), axis=0)

      [
          sample_mean_, sample_var_, sample_std_, sample_entropy_,
          actual_mean_, actual_var_, actual_std_, actual_entropy_,
          actual_mode_,
      ] = sess.run([
          sample_mean, sample_var, sample_std, sample_entropy,
          ind.mean(), ind.variance(), ind.stddev(), ind.entropy(), ind.mode(),
      ])

      self.assertAllClose(sample_mean_, actual_mean_, rtol=0.02, atol=0.)
      self.assertAllClose(sample_var_, actual_var_, rtol=0.04, atol=0.)
      self.assertAllClose(sample_std_, actual_std_, rtol=0.02, atol=0.)
      self.assertAllClose(sample_entropy_, actual_entropy_, rtol=0.01, atol=0.)
      self.assertAllClose(loc, actual_mode_, rtol=1e-6, atol=0.)

if __name__ == "__main__":
  test.main()

redpajama_set_name: RedPajamaGithub


text:
export function formatNumber(value) {
  return Number(value).toLocaleString();
}


text:
Die Gipfel-Glasschnecke (Eucobresia pegorarii) ist eine "Halbnacktschnecke" aus der Familie der Glasschnecken (Vitrinidae), die zu den Landlungenschnecken (Stylommatophora) gerechnet wird. Die Tiere können sich nicht mehr in das kleine Gehäuse zurückziehen.

Merkmale
Das rechtsgewundene, kleine Gehäuse ist ohrförmig, mit einem sehr flachen Gewinde. In der Seitenansicht ist es noch zu sehen. Es besitzt 2,5 langsam anwachsende Windungen und misst 4,3 bis 6 mm im Durchmesser, und 2,5 mm in der Höhe. Die Oberseite der Windungen ist sehr flach gewölbt, die Naht ist kaum eingetieft. Unten sind die Windungen stark gewölbt. Die Mündungsbreite beträgt 4,2 mm, die Mündungshöhe 3 mm. Die letzte Windung ist etwa breiter als das Gewinde. Ein Nabel ist nicht ausgebildet. Der Mündungsrand ist gerade und läuft dünn aus. Die stark querelliptische Mündung steht sehr schräg zur Windungsachse. Am unteren Rand der Mündung ist ein mäßig breiter Hautsaum vorhanden. Der Außenrand der Mündung ist schwach geschweift.

Die Schale ist sehr dünn und zerbrechlich. Das Embryonalgehäuse besteht aus 1¾ Windungen. Es ist milchig trüb. Der Teleoconch ist gelblich hornfarben und durchscheinend. Die Oberfläche ist schwach radial gerunzelt und stark glänzend.

Das ausgestreckte Tier wird bis etwa 18 mm lang. Der Weichkörper und der Mantel ist hellgrau bis etwas dunkler grau. Der Mantellappen ist schwarz und bedeckt nicht den Apex des Gehäuses. Die Begrenzung des schlitzförmigen Atemloches ist oben vorne fingerförmig vorgezogen.
Vom Oberrand dieses Schlitzes führt eine Mantelrinne zum Rücken (hierdurch auch
von Phenacolimax glacialis unterschieden).

Im zwittrigen Geschlechtsapparat mündet der Zwittergang (Ductus hermaphroditicus) basal in die Eiweißdrüse. Der Eisamenleiter (Spermovidukt) ist länglich-gestreckt. Der Samenleiter (Vas deferens) ist kurz und dringt in der unteren Hälfte in die Penishülle ein. Der Penis ist dick, mäßig lang und gekrümmt; er ist von einer zusätzlichen Gewebehülle (Penishülle) umgeben. Der Samenleiter dringt dann apikal in den Penis ein. Der Apex ist gerundet, der Penisretraktormuskel setzt apikal an. Im Innern des Penis ist ein Schwellkörper vorhanden. Im weiblichen Teil sind freier Eileiter (Ovidukt) und Vagina annähernd gleich lang, der Eileiter kann aber auch länger sein. Die Vagina ist aber wesentlich dicker. Die Spermathek besitzt einen sehr kurzen Stiel und eine vergleichsweise sehr große Blase, die sogar länger ist als der Stiel. Allerdings variiert die Länge des Stieles etwas. Die Blase erreicht den unteren Teil des Eisamenleiters. Eileiter und Spermathek münden in einer halbkugeligen Papille im oberen Teil der Vagina. Vagina und Penis münden zusammen in ein vergleichsweise langes Atrium.

Ähnliche Arten
Die Gipfel-Glasschnecke unterscheidet sich von der Ohrförmigen Glasschnecke (Eucobresia diaphana) und der Alm-Glasschnecke (Eucobresia nivalis) durch den kleineren Mantellappen, der den Apex des Gehäuses nicht bedeckt. Die Gipfel-Glasschnecke lässt sich von der Gletscher-Glasschnecke (Eucobresia glacialis) quasi nur anatomisch sicher unterscheiden. Bei der Gipfel-Glasschnecke besitzt das Gehäuse einen etwas schmaleren Hautsaum am unteren Rand der Mündung. Der Außenrand der Mündung ist bei der Gipfel-Glasschnecke schwach geschweift, bei der Gipfel-Glasschnecke gerundet. Die Unterschiede verschwimmen aber im Einzelfall. Die Gletscher-Glasschnecke unterscheidet sich durch den deutlich dickeren, konisch zulaufenden Penis und durch die stark verdickte Vagina.

Geographische Verbreitung und Lebensraum
Das Verbreitungsgebiet ist auf die Alpen beschränkt; Schweiz (Wallis, Graubünden), Deutschland (Bayerische Alpen), Österreichisch, Italien und Slowenien.

Die Tiere leben auf feuchten Bergwiesen und Rasenbändern an Felswänden, unter Steinen und Geröllhalden, überwiegend oberhalb der Baumgrenze, in der Schweiz zwischen 2000 und 3000 m, in Österreich zwischen 1240 und 3140 m über Meereshöhe.

Lebensweise
Lothar Forcart fand im August 1943 auf dem Riffelberg sowohl juvenile wie auch adulte Tiere. Daraus ist zu schließen, dass die Art einen mehrjährigen Entwicklungszyklus hat.

Taxonomie
Das Taxon wurde 1884 von Carlo Pollonera in der Kombination Vitrina pegorarii erstmals beschrieben. Es wird heute einheitlich in der Gattung Eucobresia Baker, 1929 geführt.

Gefährdung
Die Art ist in Bayern stark gefährdet. Auch nach der Einschätzung der IUCN ist die Art auf das Gesamtverbreitungsgebiet betrachtet stark gefährdet.

Literatur
 Michael P. Kerney, R. A. D. Cameron, Jürgen H. Jungbluth: Die Landschnecken Nord- und Mitteleuropas. 384 S., Paul Parey, Hamburg & Berlin 1983, ISBN 3-490-17918-8, S. 155.

Einzelnachweise

Glasschnecken

redpajama_set_name: RedPajamaWikipedia


text:
HomeMilitary AviationHere's what jumping from a C-130 with the 82nd Airborne Division looks like
Here's what jumping from a C-130 with the 82nd Airborne Division looks like
December 27, 2014 David Cenciotti Military Aviation 6
Paratrooper Point Of View
Ever wondered what it looks like to jump from a U.S. Air Force C-130 with the 82nd Airborne Division?
The following stunning footage, posted by the 82nd Airborne Division, provides a hint.
Based at Fort Bragg, North Carolina, the 82nd Airborne Division is a U.S. Army airborne infantry division specializing in parachute assault operations into denied areas.
Post by 82nd Airborne Division.
H/T to John Gresham for the heads-up
Airborne Assault (how it looks like if you are not wearing Night Vision Goggles)
Awesome, surreal footage: C-17 heavy cargo pallets airdrop through the NVG imagery
Pentagon confirms drone crash in Somalia. But it doesn't say it was an Austrian made PSYOPs-capable Camcopter
Night vision footage of U.S. Air Force C-17 dropping U.S. Army personnel onto a Drop Zone
This is what taking part to a U.S. Special Forces raid on board a 160th SOAR MH-60 Black Hawk looks like
This video taken 26 years ago today shows why low level airdrop can be extremely dangerous
[Photo] U.S. Army CH-47s flying over ocean of snow in Afghanistan
parachute launch
[Photo] French Navy Rafale flying below sea level over lake in Djibouti
Here's all we know about the Indonesian Airbus 320 disappeared over Java Sea
Meet the U.S. Air Force Reserve's First Female F-35A Lightning II Pilot
September 8, 2018 Tom Demerly F-35, Military Aviation 0
USAF Reserve Col. Regina Sabric Is Also Commander of 419th Fighter Wing. Colonel Regina Sabric, callsign "Torch", of Tobyhanna, Pennsylvania has become the first female Lockheed Martin F-35A Lightning II Joint Strike Fighter pilot in […]
[Video] All kinds of planes recovering at Nellis Air Force Base after Red Flag mission
February 19, 2014 David Cenciotti Military Aviation 5
Landing pattern can become quite busy at Nellis when aircraft all types start recovery at the end of their Red Flag missions. In order of appearance, the video shows the following aircraft returning to Nellis […]
Two B-52 strategic bombers have just arrived in Qatar to start pounding ISIS
April 9, 2016 David Cenciotti Iraq, Military Aviation, Syria, Troubled Areas, War on ISIS 1
The "Buff" has arrived at Al Udeid airbase. Two U.S. Air Force B-52 Stratofortress aircraft from Barksdale Air Force Base, Louisiana, have arrived at Al Udeid Air Base, Qatar, on Apr. 9, 2016. The aircraft, […]

redpajama_set_name: RedPajamaCommonCrawl


text:
Fulanisation and Islamisation: Another salvo from Obasanjo
By OLUWOLE IGE On May 30, 2019
Deputy Editor, LEON USIGBE, looks at the various positions stirred by former President Olusegun Obasanjo's latest salvo on his perceived agenda to Fulanise and Islamise the African Continent.
President Olusegun Obasanjo was in his elements again when he recently raised alarm on what he observed as an ongoing move to Fulanise and Islamise Nigeria and Africa. The former president, true to his character, did not hold back, vocalising what many would have been reluctant to express about the rampaging Fulani herdsmen and Boko Haram insurgents and their perceived hidden agenda. Obasanjo alleged a grand plot by certain elements to foist a Fulani and Islamic agenda on Nigeria, other West African countries and the whole of Africa. He therefore advocated a well thought out global action against terrorism and other organised crimes.
He said: "It is no longer an issue of lack of education and lack of employment for our youths in Nigeria which it began as. It is now West African Fulanisation, African Islamisation and global organised crimes of human trafficking, money laundering, drug trafficking, gun trafficking, illegal mining and regime change."
Makinde warns NURTW on violence
He spoke on the topic "Mobilising Nigeria's Human and Natural Resources for National Development and Stability," at the second session of the Synod of the Church of Nigeria, Anglican Communion, held at the Cathedral of Saint Paul's Anglican Church, Oleh in Isoko South Local Government Area of Delta State.
The former military Head of State said the Federal Government has allowed the attacks by Boko Haram and herdsmen to fester by treating the matter with kid gloves. Thus, he stressed the need for President Muhamamdu Buhari to take the issue seriously, rally local and global stakeholders to deal with the killer herdsmen and Boko Haram once and for all "without favouritism or cuddling."
The two-term former president said: "Both Boko Haram and herdsmen's acts of violence were not treated as they should at the beginning. They have both incubated and developed beyond what Nigeria can handle alone. They are now combined and internationalised with ISIS in control.
"Yet, we could have dealt with both earlier and nip them in the bud, but Boko Haram boys were seen as rascals not requiring serious attention in administering holistic measures of stick and carrot. And when we woke up to the reality, it was turned to industry for all and sundry to supply materials and equipment that were already outdated and that were not fit for active military purpose.
"Soldiers were poorly trained for the unusual mission, poorly equipped, poorly motivated, poorly led and made to engage in propaganda rather than achieving results. Intelligence was poor and governments embarked on games of denials while paying ransoms which strengthened the insurgents and yet governments denied payments of ransoms. Today, the insecurity issue has gone beyond the wit and capacity of Nigerian government or even West African governments."
Continuing, the elder statesman admonished: "Government must appreciate where we are, summon each group that should make contributions one by one and subsequently collectively seek the way forward for all hands on deck and with the holistic approach of stick and carrot. There should be no sacred cow.
"Some of the groups that I will suggest to be contacted are: traditional rulers, past service chiefs (no matter how competent or incompetent they have been and how much they have contributed to the mess we are in), past heads of paramilitary organisations, private sector and civil society.
"Others are community leaders particularly in the most affected areas-present and past governors, present and past local government leaders, religious leaders, past Heads of States, past intelligence chiefs, past heads of Civil Service and relevant current and retired diplomats, members of the opposition and nay groups that may be deemed relevant."
Obasanjo found immediate support in the Southern and Middle Belt Leaders Forum (SMBLF) which warned the Federal Government not treat his alert on the fulanisation and Islamisation with levity.
Speaking on their behalf, Yinka Odumakin, who is the national publicity secretary, Afenifere, said Nigeria cannot afford to take Obasanjo's admonition with levity, stating his reasons: "Every line of what President Obasanjo said, we study it and we must have a response for it. Otherwise we are losing the country fast, because when we get to a stage where Fulani herdsmen are being elevated to the level of Afenifere and Ohanaeze, it is said that when short men begin to cast long shadows, the sun is about to set. Let the sun not set for Nigeria."
Odumaking said Obasanjo is not a frivolous man, as such, his statement cannot be dismissed with the wave of the hand. He added: "They have been talking about some people trying to destabilise Nigeria. You cannot say Obasanjo wants to destabilise Nigeria; Nigeria is a pet project for him. For him to come out and make those kinds of statements, and it is not as if he does not know what he is saying.
"Don't forget that the original Boko Haram was a response to Obasanjo's reforms in the country. What is called Sharia in many northern states today was a reaction against Obasanjo but even in all those years, Obasanjo did not make those statements. He managed these situations in the interest of national stability.
"I don't know whether you have seen the latest global terrorism index. After Iraq and Afghanistan, Nigeria is the third most terrorised country in the world today and in the global terrorism index of 2015, two of the first four most violent terrorist groups are in Nigeria- Boko Haram and Fulani herdsmen, which the government is now equating with Afenifere and Ohanaeze Ndigbo and Garba Shehu (presidential spokesman) is telling us we must respect Fulani herdsmen.
"About eight months ago, Wall Street Journal said that to release the Dapchi girls, the Federal Government paid Boko Haram Three Million Euros. Today, Boko Haram has become deadlier. When the Fulani herdsmen wanted N100 billion recently, the Federal Government was denying but the herdsmen came out to say 'yes, it is true', they are asking for the money because they have been promised the money since the previous regime.
"So, I think now with President Obasanjo raising a bar on this discourse, the Federal Government should talk with Nigerians. The president cannot just be saying kidnapping has become a new business. Who are the people behind the new business? What did we do to them? What is going on? It is a new line of business, so should we live with it?"
President Buhari during the NTA interview programme on Monday, May 27, 2019
Odumakin's position was echoed by the leader of Afenifere, Chief Ayo Adebanjo, who was however happy that after seeing things differently for years, Obasanjo seemed to have seen the light. He expressed the belief that the only way to resolve the national quagmire is to restructure the country along the line the late Premier of the old Western Region, Chief Obafemi Awolowo, advocated before the 1960 Independence.
The Afenifere leader observed that apart from the terrorists trying to Islamise the country, herdsmen have also infiltrated every part of the country and are wreaking havoc as they go along. Something must be done before it is too late, he advised. To this end, he noted that the leadership of the country must wake up to the reality of the situation by ensuring the nation is restructured immediately.
The Movement for Actualisation of the Sovereign State of Biafra (MASSOB) has also jumped to Obasanjo's side, maintaining that the former president has only exposed the secret of the current administration to destroy the middle belt region and the southern part of Nigeria. MASSOB's position was conveyed to the public through Samuel Edeson who pointed out that Obasanjo was courageous in voicing his stance on the issue of fulanisation and Islamisation.
"MASSOB commends General Obasanjo for being courageous and eloquent in revealing this unwanted truth which the present government doesn't want to hear. It will be recalled that MASSOB has earlier exposed this Islamisation agenda in pursuit, projection and actualisation of the grand command of Uthman Dan Fodio which is to Islamise/fulanise this geographical entity called Nigeria and destroy the entire Southern Nigeria but nobody believed us.
"MASSOB wishes to make it clearer again that there is no difference between Boko Haram and Fulani herdsmen. The terrorist herdsmen were sent to go and destroy the South and Middle Belt while the Boko Haram will carry their acts of terrorism in the North-East. That is why President Buhari can never declare Fulani herdsmen, a terrorist movement or even condemn the killing of the innocent citizens by these foot soldiers of Islam called Fulani herdsmen," MASSOB stated.
Former aviation minister, Chief Femi Fani-Kayode, hailed Obasanjo for being the only politically active Southern and Middle Belt leader that understands the dangers lying ahead in the country.
Fani-Kayode stated: "Obasanjo remains the only politically active Southern and Middle Belt leader that understands what is really going on, the grave danger that we are in and what lies ahead. He is deeply courageous and his latest contribution that there is an agenda to fulanise Nigeria and Islamise West Africa says it all.
"Obasanjo needs say no more. History and posterity will be kind to him for speaking the bitter truth and saying what almost every other Southern and Middle Belt politician and leader knows to be true but is too timid and too scared to say publicly."
Not everyone is pleased with Obasanjo's statement though. For instance, former Jigawa State governor, Sule Lamido, who served in Obasanjo's civilian administration as Minister of Foreign Affairs, believes the former president's disappointment with serving presidents should not be allowed to turn him to a religious and ethnic bigot.
"If it were said at a non-religious venue to a non-religious audience, may be it might have been more tolerable. Please, sir, don't let your disappointment with sitting presidents turn you into a bigot. You must not abandon the national stage. The cracks along the various divides in our national cohesion are already turning into huge gorges," Lamido saidd in a statement.
The Arewa Consultative Forum (ACF) too warned against giving ethnic and religious colouration to criminal activities. Its General Secretary, Anthony Sani, argued that doing so will only embolden the criminals to continue with their evil deeds.
"Former President Obasanjo may have his facts for making such comments. But for me, I do not want us to give ethnic and religious colouration to the criminal activities of some people. Else we provide them with platforms upon which to stand and commit crimes, knowing it is almost impossible to prosecute religion and ethnicity," he said.
He pointed out that there are moderate Muslims who he said are in the majority and opposed to religious fanatics. According to him, "such moderate Muslims need to be enlisted in the campaign against Islamic terrorists whose aims are not for piety but political, albeit attired in the garb of religious jihadists. If we offend the sensitivity of such moderate Muslims and push them to the side of the fanatics, we would be playing to the gallery by swallowing the bait put by the fanatics and that is what they want."
But Obasanjo is not one to shy away from firing shots and courting controversy. This is in line with his determination to remain consistent in speaking out whenever and wherever he sees evil, no matter whose ox is gored. In July 2018, seeing the direction of the country, he accused President Buhari of being confused and incompetent to deal with national issues.
"Nigeria, in recent times, has not been so divided along religious and regional lines as at today with inexcusable killings and devastating poverty and with government's wringing hands and apparently incompetent to stem the tide except giving one unrighteous and unacceptable justification after the other. The obvious indication is that the government is seemingly confused and has got to the end of its tether and the nation is being left divisively and perilously to drift," he declared at the time.
In January this year, he alleged that Buhari had plunged Nigeria back to the late General Sani Abacha era when state institutions were used to fight perceived enemies of the late Head of State.
He said in an open letter, titled Points for Concern and Action: "Today, another Abacha Era is here. The security institutions are being misused to fight all critics and opponents of Buhari and to derail our fledgling democracy. EFCC, Police and Code of Conduct Tribunal are also being equally misused to deal with those Buhari sees as enemies for criticising him or as those who may not do his bidding in manipulating election results.
"Criticism, choice and being different are inherent trade mark of democracy. If democracy is derailed or aborted, anarchy and authoritarianism will automatically follow. Today, as in the day of Abacha, Nigerians must rise up and do what they did in the time of Abacha: churches and mosques prayed; international community stood by us Nigerians. I was a beneficiary and my life was saved. Well-meaning Nigerians took appropriate actions and made sacrifices, some supreme, some less than supreme but God had the final say and He took the ultimate action."
The Federal Government has since taken exception to Obasanjo's latest outburst and has demanded that the former president withdraw the claim on the alleged fulanisation and Islamisation. Minister of Information and Culture, Alhaji Lai Mohammed, in a written response, argued that such "indiscreet, deeply offensive and patently divisive comments are far below the status of an elder statesman. It is particularly tragic that a man who fought to keep Nigeria one is the same one seeking to exploit the country's fault lines to divide it in the twilight of his life."
While observing that Boko Haram and the Islamic State's West African Province (ISWAP) are terrorist organisations which care little about ethnicity or religion when perpetrating their senseless killings and destruction, the government's spokesman declared: "Since the Boko Haram crisis, which has been simmering under the watch of Obasanjo, boiled over in 2009, the terrorist organisation has killed more Muslims than adherents of any other religion.
"The terrorist group blown up more mosques than any other houses of worship and it is not known to have spared any victim on the basis of their ethnicity. It is therefore absurd to say that Boko Haram and its ISWAP variant have as their goal the 'fulanisation and Islamisation' of Nigeria, West Africa or Africa."
In an apparent attempt to convince doubting Thomases that the president does not brook them, Mohammed recalled Buhari's characterisation of Boko Haram as "a mindless, godless group who are as far away from Islam as one can think of." The minister noted therefore that Obasanjo's comments were "as insensitive and mischievous as they are as offensive and divisive in a multi-ethnic and multi-religious country like Nigeria."
He wondered whether "there is no limit to how far the former President will go in throwing poisonous darts at his perceived political enemies." He further said Obasanjo's prescriptions for ending the Boko Haram/ISWAP crisis, which include seeking assistance outside the shores of Nigeria, are coming several years late as he noted that Buhari had done that and more since assuming office.
But having sent his message in his usual blunt, no-holds barred manner, Obasanjo has once again retreated, perhaps, to assess and assimilate its impact on government and stakeholders before returning at a future date.
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So as you all know. I'm on my fitness journey. Being or getting fit is all about trial and error. ( as you make errors don't beat yourself up about it. It's not a race). With that being said, I have made some adjustments throughout my journey thus far. Soooo.. Let's get to it.
I don't drink juice or anything really sugary anymore. I have a soda every once in a while, but that's very rare. I notice so much that if I drink juice or anything sugary I wake up with very bad headaches. So I've opted now for just water. Mostly flavored water or water that is cold. It quenches my thirst. ( No Sprite).
I eat most of my carbs in the morning. I do this to fuel my energy for the day (which is the whole point of carbs). After that I'm working out and eat like a lot of vegetables and fruit throughout the rest of the day. Alot of carbs are not needed at night because this is the time where I wind down anyways. I don't need the energy at the end of the day.
I do not check the scale every week anymore. I'm now doing it every month. It takes about 3 weeks to form a habit. The number on the scale is just a number. I'm focusing on living a healthy lifestyle and being less concerned with my a number on the scale. What I see in the mirror is all the motivation I need. ( remember muscle weighs more than fat).
CRAVINGS!!! There are some nights that I stay up super late. Staying up late can sometimes cause you to become hungry ( but not really). So to help calm my mindset of going out for fast food or getting something unhealthy, I keep cereal in the kitchen or a healthy granola bar. Afterwards, my craving that I have is pretty much gone.
I do not go grocery shopping when I'm hungry. I will be tempted to pick out something so unhealthy. I always have to eat something healthy and satisfying before I go grocery shopping if I'm hungry. ( It's a lifesaver and it saves me money). I love me some oreos at night with milk!! My old favorite late night snack. (shhhhh) Don't judge me.
This kinda goes along with #1, but I refrigerate my water. I have been honest with myself and realized that probably the reason it was so hard for me to drink water is because it was not cold. Now, before I go to bed, I will get my water jug and fill it up with water and in the morning I will have cold water to drink throughout the day. So I have no excuse.