Professional Documents
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Briología en el INECOL
Las hepáticas, los antocerotes y los musgos son comúnmente referidos como briofitas y tradicionalmente se
clasificaban como un solo grupo, División Bryophyta (Margulis y Schwartz 1988, Raven, Evert y Eichhorn 1986).
Los tres grupos de briofitas ciertamente comparten similitudes, pero hay evidencia en aumento de que juntos
no forman un grupo monofilético (Mishler y Churchill 1984, Bremer 1985, Kenrick y Crane 1991, Lewis et al
1997).
Análisis recientes son consistentes con la naturaleza parafilética de los tres grupos de briofitas y la colocación
de los musgos como hermanos de las plantas vasculares (Polysporagiomorpha, sensu Kenrick y Crane 1997).
Aun es incierto si las hepáticas o los antoceros son el linaje mas basal entre las plantas terrestres (Duff y Nickrent
1999). Algunos análisis filogenéticos han mostrado que los musgos están más relacionados con las plantas
vasculares que con las otros dos grupos de briofitas.
El esporofito no ramificado y con un solo esporangio de los musgos es comparable con los esporofitos
ramificados y con tres esporangios en Horneophython y Aglaophyton (Rhynia) major.
Otras características comparables entre los musgos y las plantas vasculares son los estomas, la forma de los
gametangios, especialmente los arquegonios en forma de botella, la cutícula, y las células conductoras internas
(Mishler and Churchill 1984, Mishler, Lewis et al 1994).
Las relaciones filogenéticas entre los tres linajes de briofitas permanece como una de las grandes preguntas sin
resolver en la biología evolutiva de las plantas (Goffinet 2000, Mishler et al 1994, Shaw y Renzaglia 2004).
REFERENCIAS
Bremer K. 1985. Summary of green plant phylogeny and classification. Cladistics 1: 369-385.
Duff R. J. y Nickrent D. L. 1999. Phylogenetic relationships of land plants using mitochondrial small- subunit
rDNA sequences. Am. J. Bot. 86(3): 372–386.
Garbary D. J., Renzaglia K. S. y Duckett J. G. 1993. The phylogeny of land plants: a cladistic analysis based on
male gametogenesis. Pl. Syst. Evol. 188: 237-269.
Goffinet B. 2000. Origin and phylogenetic relationships of the Bryophyta. En: Shaw A. J. y Goffinet B. (eds)
Bryophyte Biology. Cambridge University Press. 124-149 p.
Kenrick P. y Crane P. R. 1991. Water-conducting cells of early fossil land plants: Implications for the early
evolution of tracheophytes. Bot. Gaz. 152: 335-345.
Kenrick P. y Crane P. R. 1997. The origin and early diversification of land plants: a cladistic study. Smithsonian
Institution Press, Washington.
Lewis L. A., Mishler B. D. y Vilgalys R. 1997. Phylogenetic relationships of the liverworts (Hepaticae), a basal
embryophyte lineage, inferred from nucleotide sequence data of the chloroplast gene rbcL. Mol. Phyl. Evol.
7:377–393.
Margulis L. y Schwartz, V. 1988. Five Kingdoms: An illustrated guide to the phyla of life on earth. W. H. Freeman
y Company, New York.
Mishler B.D. y Churchill S.P. 1984. A cladistic approach to the phylogeny of bryophytes. Brittonia 36: 406-424.
Mishler B.D, Lewis L.A., Renzaglia K.S., Garbary D.J., Delwiche C.F., Zechman F.W., Kantz T. S. y Chapman R.
L. 1994. Phylogenetic relationships of the green algae and bryophytes. Ann. Miss. Bot. Gard. 81: 451-483.
Raven P. H., Evert R. F. y Eichhorn S. E. 1986. Biology of Plants. Worth Publishers Inc., NY. 775 pp.
Renzaglia KS, Schuette S, Duff RJ, Ligrone R, Shaw AJ, et al. (2007) Bryophyte phylogeny: Advancing the
molecular and morphological frontiers. The Bryologist: Vol. 110, No. 2 pp. 179–213.
Abstract. Revolutionary new concepts of bryophyte relationships have emerged from molecular phylogenetic
analyses conducted since the onset of the 21st century. For example, sequence data contradict the historical
notion that isophylly in leafy liverworts is plesiomorphic and that simple thalloid liverworts are monophyletic.
Also contrary to traditional views are the concepts that Leiosporoceros is genetically distinct from other hornworts
and that Oedipodium is sister to the peristomate mosses. Substantial increases in ultrastructural and anatomical
data likewise have provided new insights on interrelationships. Because of this recent deluge in evolutionary
studies on bryophytes, it is an opportune time to co-examine contemporary morphological knowledge and novel
molecular hypotheses. An understanding of bryophyte evolution and biology is essential to identify structural
innovations that accompanied early land colonization and to illuminate the evolution of more complicated body
plans in tracheophytes. In this review, we examine the progress that has been made since the 1999 International
Botanical Congress in clarifying the evolutionary history of the three groups of bryophytes. The state of our
knowledge on interrelationships is discussed, with poorly-known, genetically divergent taxa illustrated for each
group. Our review of bryophyte evolution includes a reëvaluation of the evolution of sperm cells, sporogenesis,
stomata, symbioses, conducting cells and chloroplast ultrastructure in hornworts. We explore the prospects for
future discoveries and advances with an emphasis on fundamental evolutionary problems that remain and the
challenges that must be met to resolve them.
Shaw J. y Renzaglia K. 2004. Phylogeny and diversification of Bryophytes. Am J. Bot. 91 (10): 1557-1581.
Abstract. The bryophytes comprise three phyla of embryophytes that are well established to occupy the first
nodes among extant lineages in the land-plant tree of life. The three bryophyte groups (hornworts, liverworts,
mosses) may not form a monophyletic clade, but they share life history features including dominant free-living
gametophytes and matrotrophic monosporangiate sporophytes. Because of their unique vegetative and
reproductive innovations and their critical position in embryophyte phylogeny, studies of bryophytes are crucial
to understanding the evolution of land plant morphology and genomes. This review focuses on phylogenetic
relationships within each of the three divisions of bryophytes and relates morphological diversity to new insights
about those relationships. Most previous work has been on the mosses, but progress on understanding the
phylogeny of hornworts and liverworts is advancing at a rapid pace. Multilocus multigenome studies have been
successful at resolving deep relationships within the mosses and liverworts, whereas single-gene analyses have
advanced understanding of hornwort evolution.
Phylogenetic relationships of the Thuidiaceae and the non-monophyly of the Thuidiaceae and the Leskeaceae
based on rbc L, rps 4 and the rps 4-trn S intergenic spacer
D García-Avila, E De Luna, AE Newton. 2009
The Bryologist 112 (1), 80-93
by Efrain De Luna
are licensed under a Creative Commons Attribution-Noncommercial-Share Alike 2.5 Mexico License.
BRIOLOGÍA
Briología en el INECOL
Los Musgos
Los musgos
Como linaje, los musgos son un grupo histórico crucial en el entendimiento de la transición de la vida a la
tierra y la vascularizacion. Los gametofitos retienen algunas características de las algas verdes ancestrales
(clorofila a y b, almidón, espermátidas con dos undulipodios). En cambio, los esporofitos despliegan
innovaciones clave para la vida fuera del agua como: estomas, un eje simple de células conductoras en un
esporofito no ramificado, y esporas liberadas al aire producidas en un esporangio sencillo apical.
El esporofito de los musgos con un solo esporangio es el nivel estructural mas simple entre todas las plantas
terrestres. El siguiente nivel estructural se encuentra en dos grupos fósiles: Horneophythopsida y Aglaophyton
(Rhynia) major, donde el esporofito es ramificado y produce varios esporangios apicales. El esporofito muestra
la organización estructural mas compleja en las traqueofitas.
Clasificación
Los musgos tanto fósiles como actuales consisten en por lo menos unas 10 000 especies. Entre los géneros
más conocidos se encuentran: Sphagnum, Andreaea, Polytrichum, Funaria, Bryum y Thuidium.
3. Relaciones filogenéticas.
La gran mayoría de especies de musgos constituyen tres grupos informales de Ordenes de la Clase Bryopsida:
1. Diphysciales, Funariales, Timmiales, Encalyptales,
2. Grimmiales, Dicranales, Pottiales, Fissidentales,
3. Orthotrichales, Splachnales, Hedwigiales, Bryales, Rhizogoniales, Hookeriales e Hypnales.
4. Lecturas adicionales.
REFERENCIAS
Crosby M. R., Magill R. E., Allen B. y He S. 2000. A checklist of the mosses. Missouri Botanical Garden, St.
Louis, Missouri, USA, website: www.mobot.org/MOBOT/tropicos/most/checklist.shtml.
Mishler B.D. y Churchill S.P. 1984. A cladistic approach to the phylogeny of bryophytes. Brittonia 36: 406-424.
Shaw J. y Renzaglia K. 2004. Phylogeny and diversification of Bryophytes. Am J. Bot. 91 (10): 1557-1581.
Galería de Bryophyta
BRIOLOGÍA
Briología en el INECOL
Las Hepáticas
Las Hepáticas
Este grupo de briofitas es muy diverso (más o menos 8 000 especies), pues incluye tanto las formas talosas
como las foliosas.
Algunos de los géneros más conocidos son: Marchantia, Plagiochila, Frullania y Metzgeria.
Marchantia chenopoda
Clasificación:
iii. La presencia de ácido lunulárico se ha interpretado como una sinapomorfia para las Marchantiophyta, pero
queda la duda si este compuesto también está presente en algunas algas verdes, las cuales no se han
estudiado (Kenrick y Crane 1997. p. 69).
Relaciones dentro del grupo.
Las relaciones filogenéticas entre los seis subgrupos de hepáticas son muy inciertas. Únicamente se ha podido
reconocer la relación de grupos hermanos entre Metzgeriales (talosas simples) y Jungermaniales (foliosas). Las
relaciones entre los otros cuatro órdenes y éste último grupo aún no están resueltas.
Marchantia chenopoda
REFERENCIAS
Kenrick P. y Crane P. R. 1997. The origin and early diversification of land plants: a cladistic study. Smithsonian
Institution Press. Washington D.C., USA.
Goremykin, V. V. & F. H. Hellwig. 2005. Evidence for the most basal split in land plants dividing bryophyte and
tracheophyte lineages. Plant Systematics and Evolution 254: 93-103.
Abstract The problem of relationships among the major basal living groups of land plants is long standing, yet
the uncertainty as to the phylogenetic affinity of these lines persists in the literature. Molecular and modern
cladistic studies of the phylogenetic relationships of the above groups resulted in a large number of conflicting
topologies. However, with the exception of the cladistic analyses of spermatogenesis, suggesting monophyly of
extant bryophytes, these studies agree the paraphyletic bryophyte grade is basal within the embryophyte tree.
Here we would like to present analyses on the basis of the concatenated datasets of nucleotide and amino-
acid sequences of 57 protein-coding genes common to 17 chloroplast genomes of land plants and a
charophyte alga Chaetosphaeridium globosum. Character-wise, these are the largest datasets currently
available to address the problem of basal relationships within embryophytes. Main lineages of bryophytes, i.e
liverworts, hornworts and mosses are represented in our alignments with a single taxon, whereas 14 taxa
represent the tracheophytes. With our data, phylogeny with liverwort basal appears to be and artifact related
to high and unequal A+T contents among the sequences analysed. Reducing this compositional bias and
applying methods developed to counter it, we recovered an alternative, strongly supported topology wherein
both bryophytes and tracheophytes are monophyletic. Within bryophytes, hornworts are basal and liverworts
are sister to mosses.
Lewis LA, BD. Mishler & R Vilgalys. 1997. Phylogenetic Relationships of the Liverworts (Hepaticae), a Basal
Embryophyte Lineage, Inferred from Nucleotide Sequence data of the Chloroplast Gene rbcL. Molecular
Phylogenetics and Evolution 7: 377-393.
Abstract. Sequence data from the chloroplast-encoded generbcL were obtained for 24 liverworts, a basal
group of embryophytes. Maximum likelihood and parsimony analyses of these data, along with data from other
major green plant lineages, confirm hypotheses based on morphological data, such as the paraphyly of
bryophytes, and the basal position of liverworts. Molecular data corroborate the deep separation between the
complex thalloid and leafy/simple thalloid liverworts implied by morphological data, but the monophyly of
liverworts could not be rejected. The effects of accounting for site-to-site rate heterogeneity in these data
were examined using maximum likelihood methods. Comparison of trees obtained with and without rate
heterogeneity showed that simply allowing for heterogeneity had a greater improvement on likelihood score
than optimization of transition/transversion bias. Incorporation of site-to-site rate heterogeneity in the larger
analysis, however, did not necessarily change which topology was favored. Properties ofrbcL sequences from
the two liverwort groups were compared. Significantly different substitution rates were found between
leafy/simple thalloid and complex thalloid liverwort taxa, with rates ofrbcL sequence evolution in leafy/simple
thalloid taxa being higher and more indicative of those of vascular plants, and with those of complex thalloid
taxa (such asMarchantia) being slower. Codon usage inrbcL in complex thalloid liverworts was biased toward
NNU and NNA, compared to the leafy/simple thalloid liverworts. Although base composition and relative
substitution rates differed between the two groups, no significant differences were detected within each of the
two groups of liverworts. The signal present in first and second codon sites versus third codon sites was
compared. While the third codon positions inrbcL across this taxon sampling are highly variable (with only 15
constant sites of 439), the trees obtained were in general agreement with trees from the entire data set and
with trees obtained from independent sources of data. The presence of signal in third codon positions across
greater than 400 MY of plant evolution means that definitions of saturation based on pair-wise comparisons of
sequences inadequately assess phylogenetic signal.
Wheeler JA. 2000. Molecular Phylogenetic Reconstructions of the Marchantioid Liverwort Radiation. The
Bryologist: Vol. 103, No. 2 pp. 314–333
Abstract Molecular phylogenies of the complex-thalloid liverworts (Marchantiales) were reconstructed using
independent nuclear and plastid data sets to explore relative age, relationships, and character evolution in this
ancient group. The sample includes 10 carpocephalate taxa and 24 acarpocephalate taxa (emphasizing Riccia)
within Marchantiales sensu stricto. In addition, Monoclea, Sphaerocarpos, Riella, three Metzgeriales
(Fossombronia, Pellia, and Blasia), the hornwort Anthoceros, four mosses, and outgroup Coleochaete are also
sampled. Two nucleotide sequence alignments were used 1) partial nuclear-encoded Large Subunit rDNA (LSU
rDNA) for all 48 taxa and 2) the plastid-encoded trnL-F region for the marchantioids and outgroup Blasia.
Alignment-ambiguous regions of each alignment were culled. A combined matrix consisting of concatenated
nuclear and plastid culled alignments was assembled for marchantioids and Blasia. The two alignments were
utilized in four analyses: 1) nuclear LSU rDNA for all taxa, 2) nuclear LSU rDNA for marchantioids + Blasia, 3)
plastid trnL-F region for marchantioids plus Blasia, and 4) combined nuclear and plastid data for marchantioids
plus Blasia. Selected pairwise comparisons reveal significant rate heterogeneity in the nuclear LSU rDNA data;
metzgerioid liverworts, hornworts and primitive mosses evolve significantly slower than other taxa relative to
the outgroup Coleochaete. The LSU rDNA genes of some marchantioid taxa and sampled bryalean mosses are
apparently evolving relatively fast. Rate heterogeneity is also documented within Marchantiales. Lunularia
positions as the most basal of sampled Marchantiopsida; Sphaerocarpales, Marchantia, and Corsinia represent
early diverging lines. A monophyletic Aytoniaceae, Cleveaceae, and Riccia are indicated. Topologies imply that
extant acarpocephalate taxa are derived from carpocephalate forms. Monoclea positions well within
Marchantiales sensu stricto. A well-supported long branch (Decay Index = 19) unites all sampled
Marchantiopsida and isolates this clade from other liverworts and bryophytes. This long branch may suggest
extensive extinction of proto- and eomarchantioid forms that led to modern taxa. A recurring theme in the
topologies presented here is the unresolved marchantioid polytomy that follows well-supported basal nodes. A
similar polytomy results from either independent data set and may correspond to a rapid radiation of
marchantioid forms (e.g., Aytoniaceae, Cleveaceae, Targionia, Monoclea, and riccioids) coincident with
extreme conditions and ecological reorganizations of the Permo-Triassic. The origin of Marchantiopsida
probably occurred long before; amidst, perhaps, a series of long-extinct Blasia-like ancestors that colonized
and innovated on any of various xeric surfaces (either cool or warm) that were available throughout
embryophyte history in the Paleozoic.
Galería de Marchantiophyta
BRIOLOGÍA
Briología en el INECOL
Los Antoceros
Este es un grupo pequeño de más o menos 100 especies, clasificados entre 5 o 9 géneros.
Algunos grupos de géneros se han clasificado al nivel de familia, aunque Notothylas se ha propuesto a nivel de
orden.
ii. Pirenoides divididos por una membrana en cuerpos pirenoidales. En las algas el pirenoide no está dividido.
v. Simbiontes de Nostoc.
viii. Ultraestructura del espermatozoide, bilateralmente simétrica con los cuerpos basales similares y de
inserción adyacente (Fig. 22, Carothers y Duckett 1979. Bryophyte Systematics p. 439).
ix. División inicial del cigoto vertical.
x. Pseudoeláteres en el esporangio.
xi. Columnela. Hay una columna central de tejido estéril en el esporofito en los antoceros y en los musgos.
Esta estrucctura satisface el criterio de similitud y de conjunción pero no la prueba de congruencia. Esta
sinapomorfia por tanto es homoplásica en esos dos grupos de briofitas.
El grupo Anthocerophyta, contiene alrededor de 100-150 especies escasamente descritas. Los conceptos de
las interrelaciones de los antoceros basado en la morfología, y los esquemas de clasificación resultante,
muestran que no hay consenso en los niveles de orden, familia y género (Hyvönen y Piipo 1993, Shaw y
Renzaglia 2004).
Stech,M., D. Quandt & F. Wolfgang. 2003. Molecular circumscription of the hornworts (Anthocerotophyta)
based on the chloroplast DNA trnL–trnF region. Journal of Plant Research 116:389-398.
Abstract In phylogenetic trees generated from partial trnLUAA intron sequences, the hornworts (represented
by nine species from the genera Anthoceros, Dendroceros, Megaceros, Notothylas and Phaeoceros) are
resolved as a monophyletic group and are separated from the clades of mosses, liverworts and tracheophytes.
A secondary structure of the trnLUAA intron of Anthoceros agrestis is presented, displaying the arrangement of
the stem-loop regions P1–P9. Compensatory base-pair changes (coevolutionary sites) are detected in regions
P4/5 and P9 within the hornwort sequences. The original homology of the most variable region, P8, cannot be
detected anymore due to the extremely fast divergent evolution of this segment in the major land plant
groups. Similarly, a high sequence divergence occurs in the trnL–trnF intergenic spacer. Apart from
synapomorphic substitutions in the trnLUAA intron, the hornworts are characterised by a large P6 region
consisting of many repetitive elements. The molecular data therefore support the hornworts as representing
an independent land plant lineage (Anthocerotophyta). Although relationships between hornworts and the
other land plant groups remain unresolved in the trnLUAA intron trees, it is rather unlikely that bryophytes are
monophyletic in their traditional circumscription, i.e. comprising hornworts, mosses and liverworts.
Keywords Anthocerotophyta - Chloroplast–DNA relationship - Group I intron secondary structure - Molecular
evolution - trnLUAA intron - trnLUAA–trnFGAA intergenic spacer
Galería de Anthocerophyta
BRIOLOGÍA
Briología en el INECOL
La clasificación que aquí se presenta deriva de análisis cladísticos recientes basados en datos morfológicos y
moleculares. Varios estudios filogenéticos han cubierto la diversidad de los musgos al nivel de ordenes y se
basan en la investigación de los últimos años en la que se incrementaron las unidades de muestreo y los
genes secuenciados. Los primeros estudios obviamente incorporaron pocos representantes de todos los
musgos (Hedderson et al 1996, Newton et al 2000). Otros estudios han incrementado la cobertura, de manera
que al menos unos representantes de todos los ordenes ya han sido incluidos en algún análisis cladístico.
El linaje mas basal es el de la Clase Sphagnopsida (1). Este grupo es hermano del clado grande del resto de
los musgos, dentro del cual la clase Andreaeopsida (3) es posiblemente el linaje basal.
A este nivel la posición y categoría taxonómica de Takakiopsida son inciertos. Unos análisis cladísticos colocan
a Takakia como grupo hermano de Andreaeopsida, pero otros estudios encuentran que es hermano de
Sphagnopsida. En cualquier caso, su reconocimiento taxonómico puede considerarse tentativamente al nivel
de clase. Takakiopsida (2) entonces incrementa el numero de clases a cinco.
El grupo de los musgos con peristoma incluye tres linajes principales: Polytrichopsida (4) con peristomas
nematodontos. El segundo grupo es de musgos con peristomas artrodontos que contiene a Tetraphidopsida. El
tercero es Bryopsida (5). Sin embargo, en la clasificación presentada por Shaw y Goffinet (2000) excluyen
Tetraphidopsida y colocan el orden Tetraphidales dentro de Polytrichopsida.
Varios factores han hecho posible esta nueva visión en las relaciones filogeneticas dentro de los musgos.
Nuevos datos empíricos se han acumulado de la morfología, anatomía, ontogenia, ultraestructura y secuencias
del DNA. Aunque la mayoría de los estudios recientes se basan en datos moleculares, el elemento mas
importante del progreso reciente ha sido el uso de métodos cladísticos para la interpretación de esos tipos de
evidencia. Un marco formal de las relaciones de los musgos esta disponible ahora que reemplaza las
estimaciones intuitivas de relaciones y clasificación fundamentadas mayormente en conceptos de similitud
general y escenarios evolutivos ad hoc.
Sphagnum (Sphagnopsida)
Clase 2 Takakiopsida
Orden 3 Takakiales
Clase 3 Andreaeopsida
Orden 4 Andreaeales
Orden 5 Andreaeobryales
Clase 4 Polytrichopsida
Orden 6 Polytrichales
Orden 7 Tetraphidales
Polytrichum (Polytrichopsida)
Clase 5 Bryopsida
Subclase 1 Diphysciidae
Orden 8 Diphysciales
Subclase 2 Funariidae
Orden 9 Funariales
Orden 10 Timmiales
Orden 11 Encalyptales
Subclase 3 Dicraniidae
Orden 12 Grimmiales
Orden 13 Seligeriales
Grimmia (Bryopsida, Dicraniidae)
Orden 14 Archidiales
Orden 15 Pottiales
Orden 16 Dicranales
Subclase 4 Bryidae
Orden 17
Orthotrichales
Orden 18 Splachnales
Orden 19 Hedwigiales
Orden 20 Bryales
Orden 23 Hookeriales
Orden 24 Hypnales
REFERENCIAS
Buck W. R., Cox C. J., Shaw A. J. y Goffinet B. 2004. Ordinal relationships of pleurocarpous mosses, with
special emphasis on the Hookeriales. Syst. Biod. 2 (2): 121–145.
Hedderson T. A., Chapman R. L. y Rootes W. L. 1996. Phylogenetic relatioships of Bryophytes inferred from
nuclear encoded rRNA gene sequences. Pl. Syst. Evol. 200:213-224.
Newton A. E., Cox C. J., Duckett J. G., Wheeler J., Goffinet B., Hedderson T. A. y Mishler B. D. 2000.
Evolution of the major moss lineages: Phylogenetic analyses based on multiple gene sequences and
morphology. The Bryologist 103:187-211.
Shaw A.J. y Goffinet B. (eds). 2000. Bryophyte Biology. Cambridge University Press, New York.
Galería de Bryophyta
Briofitas de México / Bryophytes of Mexico -
Hypopterygium tamariscinum (Hedw.) Brid.
1,a: One chloroplast per cell without a central pyrenoid, proximal face of spore with a central wart
in each of the triangular areas ................ > NO! A. tristanianus
2, a:. Dehiscent capsules >10 mm long, distal spore surface with spines forming a differentiated
reticulum ....................3 No!
2, b: Dehiscent capsules ≤10 mm long, distal spore surface with spiny tubercles not forming a
differentiated reticulum -------> A. sambesianus!
- One chloroplast per cell with a central pyrenoid, proximal face of spore without a central
wart.........................2
2. Dehiscent capsules >10 mm long, distal spore surface with spines forming a differentiated
reticulum ....................3
- Dehiscent capsules ≤10 mm long, distal spore surface with spiny tubercles not forming a
differentiated reticulum
......................................................................................................................................... A.
sambesianus
4. Thalli with dorsal lamellae scarce or absent, proximal spore surface not foveolate
............................ A. hispidus
- Thalli with dorsal lamellae regular to abundant, proximal spore surface foveolate or incompletely
foveolate ........ 5
5. Proximal spore surface with subglobose tubercles, distal surface spinose not foveolate
........................ A. scariosus
- Proximal spore surface without subglobose tubercles, distal surface spinose with a foveolate
reticulum .... A. punctatus
BRIOLOGÍA
Briología en el INECOL
- en el suelo
- sobre rocas y piedras al lado de los caminos
- sobre la base de arbustos y arboles
- en la corteza de los troncos
- arriba en las ramas de arboles
División Bryophyta
Clase Polytrichopsida
Clase Bryopsida
Subclase Dicraniidae
Orden Pottiales
Orden Dicranales
Subclase Bryidae
Orden Orthotrichales
Orden Bryales
Orden Rhizogoniales
Orden Hookeriales
Orden Hypnales
Clase Polytrichopsida
Orden Polytrichales,
Polytrichaceae
Atrichum muelleri, 4389, 4403
Atrichum (Polytrichaceae)
Clase Bryopsida
Subclase Dicraniidae
Orden Pottiales
Pottiaceae Leptodontium viticulosoides var sulphureum 4394
Streptopogon concavifolium 4305
Calymperaceae Syrrhopodon parasiticus 4412
Syrrhopodon prolifer var scaber 4571
Orden Dicranales
Campylopus
Dicranum 4402
Dicranaceae Fissidens polypodiodes 4393
Leucobryum albidum 4395
Fissidentaceae
Leucobryaceae
Dicranum (Dicranaceae)
Fissidens (Fissidentaceae)
Leucobryum (Leucobryaceae)
Subclase Bryidae
Macrocoma (Orthotrichaceae)
Orden Bryales
Bryaceae
Bryum densifolium 4362, 4388
Mniaceae
Mnium 4310
Bryum (Bryaceae)
Orden Rhizogoniales
Rhizogonium (Rhizogoniaceae)
Racopilum (Racopilaceae)
Orden Hookeriales
Hookeriaceae
Hookeria acutifolia 4379
Hypopterigiaceae
Hypopterygium tamariscinum 4251
Leucomiaceae
Rhyncostegiopsis flexuosa 4400
Hookeria (Hookeriaceae)
Pilotrichaceae
Rhyncostegiopsis tunguraguana 4568
Hypopterygium (Hypopterygiaceae)
Orden Hypnales
Schoenobryum 4577
Schoenobryum concavifoluim 4306, 4566
Schoenobryum gardneri, LH B48448,
Neckera 4253
Papillaria (Meteoriaceae)
Neckera urnigera 4572, LH B48426,
Thuidium (Thuidiaceae)
Briófitas
Características generales
También son capaces de resistir temperaturas extremas: crecen tanto sobre rocas
del nivel de nieve permanente del Ártico y Antártida, como en lugares donde la
roca alcanza 70º al sol.
Requieren menor intensidad lumínica que las restantes plantas, por lo que son los
habitantes del interior de las cuevas, viviendo con solo 0,1 % de intensidad
lumínica.
Toleran un amplio rango de pH: los Sphagnum de las turberas viven a pH 3 - 4, los musgos
de la toba caliza están a pH 7- 8,5.