The evolution of sporophore in Reticulariaceae (Myxomycetes)
Recent molecular studies have clarified the phylogenetic relations within the family Reticulariaceae (Myxomycetes, Myxogastrea), allowing to reconstruct the ways of the morphological evolution within the family. We conclude, that the general tendency of the sporophore evolution in Reticulariaceae...
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Інститут ботаніки ім. М.Г. Холодного НАН України
2016
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irk-123456789-1783682021-02-19T01:28:30Z The evolution of sporophore in Reticulariaceae (Myxomycetes) Leontyev, D.V. Спорові рослини та гриби Recent molecular studies have clarified the phylogenetic relations within the family Reticulariaceae (Myxomycetes, Myxogastrea), allowing to reconstruct the ways of the morphological evolution within the family. We conclude, that the general tendency of the sporophore evolution in Reticulariaceae was the transformation of the stalked forms to the sessile ones, followed by the development of the metameric sporophores, called pseudoaethalia and aethalia, from the separate sporocarps. The common ancestor of the family was similar to Alwisia lloydiae having stalked sporocarps and tubular capillitium. At least four evolutionary lineages originated form this ancestor: 1) Alwisia → Tubifera, 2) Alwisia → Rigidotubula (and probably Siphoptychium), 3) Alwisia → (Lycogala) → Thecotubifera, 4) Alwisia → Lycogala → Reticularia. In each of these branches, the processes like a loss of stalks, transition from spherical to elongated sporothecae, transformation of solitary sporocarps to grouped ones, a loss of capillitium and formation of pseudocapillitium, have occurred independently. Therefore, the type of the fruiting body, and presence of capillitium or pseudocapillitium, cannot be considered as sufficient criteria for separating myxomycete genera. Молекулярно-генетичні дослідження прояснили філогенетичні відносини між таксонами родини Reticulariaceae (Myxomycetes, Myxogastrea), надали змогу реконструювати шляхи морфологічної еволюції цього таксона. Згідно з одержаними даними загальною тенденцією еволюції спорофорів у Reticulariaceae є трансформація стебельчастих форм у сидячі, яка супроводжувалася формуванням з вільних спорокарпів метамерних плодових тіл ‒ псевдоеталіїв та еталіїв. Спільний предок родини був подібний до сучасної Alwisia lloydiae і мав стебельчасті спорокарпи та трубчастий капіліцій. Від цього предка започатковані принаймні чотири філогенетичні лінії: Alwisia → Tubifera; Alwisia → Rigidotubula (та, ймовірно, Siphoptychium); Alwisia → (Lycogala) → Thecotubifera; Alwisia → Lycogala → Reticularia. У кожній із цих ліній незалежно відбувалися такі еволюційні процеси, як втрата ніжок, перехід від сферичних до циліндричних споротек і від одиночних до згрупованих спорокарпів, втрата капіліцію та формування псевдокапіліцію. Зважаючи на це, тип плодового тіла та наявність капіліцію або псевдокапіліцію не можуть розглядатися як вирішальні критерії для розмежування родів міксоміцетів. Молекулярно-генетические исследования прояснили филогенетические взаимоотношения между таксонами семейства Reticulariaceae (Myxomycetes, Myxogastrea), дав возможность реконструировать пути морфологической эволюции этого таксона. Согласно полученным данным, общей тенденцией в эволюции спорофора Reticulariaceae является трансформация стебельчатых форм в сидячие, сопровождавшаяся формированием из свободных спорокарпов метамерных плодовых тел – псевдоэталиев и эталиев. Общий предок семейства был близок к современной Alwisia lloydiae; он имел стебельчатые спорокарпы и трубчатый капиллиций. От этого предка берут начало, по крайней мере, четыре филогенетические линии: Alwisia → Tubifera; Alwisia → Rigidotubula (и, вероятно, Siphoptychium); Alwisia → (Lycogala) → Thecotubifera; Alwisia → Lycogala → Reticularia. В каждой из этих линий независимо происходили такие эволюционные процессы, как потеря ножек, переход от сферических споротек к цилиндрическим и от одиночных спорокарпов к сгруппированным, потеря капиллиция и формирование псевдокапиллиция. В связи с этим тип плодового тела, а также наличие капиллиция и псевдокапиллиция не могут рассматриваться в качестве решающих критериев для разграничения родов миксомицетов. 2016 Article The evolution of sporophore in Reticulariaceae (Myxomycetes) / D.V. Leontyev // Український ботанічний журнал. — 2016. — Т. 73, № 2. — С. 178-184. — Бібліогр.: 14 назв. — англ. 0372-4123 DOI: http://dx.doi.org/10.15407/ukrbotj73.02.178 http://dspace.nbuv.gov.ua/handle/123456789/178368 en Український ботанічний журнал Інститут ботаніки ім. М.Г. Холодного НАН України |
| institution |
Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| collection |
DSpace DC |
| language |
English |
| topic |
Спорові рослини та гриби Спорові рослини та гриби |
| spellingShingle |
Спорові рослини та гриби Спорові рослини та гриби Leontyev, D.V. The evolution of sporophore in Reticulariaceae (Myxomycetes) Український ботанічний журнал |
| description |
Recent molecular studies have clarified the phylogenetic relations within the family Reticulariaceae
(Myxomycetes, Myxogastrea), allowing to reconstruct the ways of the morphological evolution within the family. We
conclude, that the general tendency of the sporophore evolution in Reticulariaceae was the transformation of the
stalked forms to the sessile ones, followed by the development of the metameric sporophores, called pseudoaethalia
and aethalia, from the separate sporocarps. The common ancestor of the family was similar to Alwisia lloydiae having
stalked sporocarps and tubular capillitium. At least four evolutionary lineages originated form this ancestor: 1) Alwisia → Tubifera, 2) Alwisia → Rigidotubula (and probably Siphoptychium), 3) Alwisia → (Lycogala) → Thecotubifera,
4) Alwisia → Lycogala → Reticularia. In each of these branches, the processes like a loss of stalks, transition from
spherical to elongated sporothecae, transformation of solitary sporocarps to grouped ones, a loss of capillitium and
formation of pseudocapillitium, have occurred independently. Therefore, the type of the fruiting body, and presence of
capillitium or pseudocapillitium, cannot be considered as sufficient criteria for separating myxomycete genera. |
| format |
Article |
| author |
Leontyev, D.V. |
| author_facet |
Leontyev, D.V. |
| author_sort |
Leontyev, D.V. |
| title |
The evolution of sporophore in Reticulariaceae (Myxomycetes) |
| title_short |
The evolution of sporophore in Reticulariaceae (Myxomycetes) |
| title_full |
The evolution of sporophore in Reticulariaceae (Myxomycetes) |
| title_fullStr |
The evolution of sporophore in Reticulariaceae (Myxomycetes) |
| title_full_unstemmed |
The evolution of sporophore in Reticulariaceae (Myxomycetes) |
| title_sort |
evolution of sporophore in reticulariaceae (myxomycetes) |
| publisher |
Інститут ботаніки ім. М.Г. Холодного НАН України |
| publishDate |
2016 |
| topic_facet |
Спорові рослини та гриби |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/178368 |
| citation_txt |
The evolution of sporophore in Reticulariaceae (Myxomycetes) / D.V. Leontyev // Український ботанічний журнал. — 2016. — Т. 73, № 2. — С. 178-184. — Бібліогр.: 14 назв. — англ. |
| series |
Український ботанічний журнал |
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AT leontyevdv theevolutionofsporophoreinreticulariaceaemyxomycetes AT leontyevdv evolutionofsporophoreinreticulariaceaemyxomycetes |
| first_indexed |
2025-07-15T16:50:22Z |
| last_indexed |
2025-07-15T16:50:22Z |
| _version_ |
1837732434571952128 |
| fulltext |
178 ISSN 0372-4123. Ukr. Bot. J., 2016, 73(2)
Myxogastrea, or Myxomycetes, are the group of ameboid
eukaryotes, able to produce macroscopic fruiting bodies
of complicate structure. Within the group, five orders
and about twelve families are traditionally recognized,
among which the family Reticulariaceae is characterized
by the absence of true capillitium and pseudoaethaliate
or aethaliate structure of the fruiting bodies (Nannenga-
Bremekamp, 1991; Neubert et al., 1993; Lado, Pando,
1997; Ing, 1999; Poulain et al., 2011).
The traditional understanding of the taxonomical
structure and discriminative features of Reticulariaceae
was reconsidered in our recent study, based on the
18S rDNA phylogeny (Leontyev et al., 2014a; 2014b;
2015). The family, with a few exceptions, appeared
to be a monophyletic taxon, subdivided to at least
six clusters, which corresponds to the genera Alwisia
Berk. & Broome, Lycogala Adans., Reticularia
Bull., Tubifera G.F. Gmel., Rigidotubula ad int. and
Thecotubifera ad int., among which Alwisia was re-
erected (Leontyev et al., 2014a; 2014b), and two last
names are to be published as new to science. One more
genus, the recently re-erected Siphoptychium Rostaf.,
was united with Reticulariaceae based on morphological
data (Leontyev, 2015).
The genera Reticularia and Tubifera, species
Alwisia bombarda Berk. & Broome, Lycogala
epidendrum (L.) Fr., Tubifera ferruginosa (Batsch) J.F.
Gmel., and T. dictyoderma Nann.-Bremek. &
Loer. in their traditional understanding appeared
to be polyphyletic. A revision of these taxa, based
on morphological and molecular data, allowed to
describe 9 new species: Alwisia lloydiae Leontyev,
S.L. Stephenson et Schnittler, A. morula G. Moreno,
Leontyev, D.W. Mitch., S.L. Stephenson, C. Rojas &
Schnittler, A. repens Leontyev, Schnittler, G. Moreno,
S.L. Stephenson, D.W. Mitchell & C. Rojas,
Tubifera applanata Leontyev & Fefelov, T. corymbosa
Leontyev, Schnittler, S.L. Stephenson & L.M. Walker,
T. dudkae (Leontyev & G. Moreno) Leontyev,
Moreno & Schnittler, T. magna Leontyev, Schnittler,
S.L. Stephenson & T. Kryvomaz, T. montana Leontyev,
Schnittler & S.L. Stephenson, T. pseudomicrosperma
Leontyev, Schnittler & S.L. Stephenson, and two
subspecies: T. ferruginosa subsp. ferruginosa Leontyev,
Schnittler & S.L. Stephenson and T. ferruginosa subsp.
acutissima Leontyev, Schnittler & S.L. Stephenson
(Leontyev et al., 2014a; 2014b; 2015). Two more species
(Rigidotubula reticulata ad int., R. violacea ad int.)
and one new combination (Thecotubifera dictyoderma
ad int., comb. nov. pro Tubifera dictyoderma Nann.-
Bremek. & Loer.) are prepared for the publication.
The species of Reticularia which have warty spores
with olive, golden-yellow or brown pigmentation © D.V. LEONTYEV, 2016
doi: 10.15407/ukrbotj73.02.178
D.V. LEONTYEV
Kharkiv State Zooveterinary Academy
1, Akademichna Str., Kharkiv, 62134, Ukraine
protista@mail.ru
THE EVOLUTION OF SPOROPHORE IN RETICULARIACEAE (MYXOMYCETES)
Leontyev D.V. The evolution of sporophore in Reticulariaceae (Myxomycetes). Ukr. Bot. J., 2016, 73(2): 178—184.
Kharkiv State Zooveterinary Academy
1, Akademichna Str., Kharkiv, 62134, Ukraine
protista@mail.ru
Abstract. Recent molecular studies have clarified the phylogenetic relations within the family Reticulariaceae
(Myxomycetes, Myxogastrea), allowing to reconstruct the ways of the morphological evolution within the family. We
conclude, that the general tendency of the sporophore evolution in Reticulariaceae was the transformation of the
stalked forms to the sessile ones, followed by the development of the metameric sporophores, called pseudoaethalia
and aethalia, from the separate sporocarps. The common ancestor of the family was similar to Alwisia lloydiae having
stalked sporocarps and tubular capillitium. At least four evolutionary lineages originated form this ancestor: 1) Alwisia →
Tubifera, 2) Alwisia → Rigidotubula (and probably Siphoptychium), 3) Alwisia → (Lycogala) → Thecotubifera,
4) Alwisia → Lycogala → Reticularia. In each of these branches, the processes like a loss of stalks, transition from
spherical to elongated sporothecae, transformation of solitary sporocarps to grouped ones, a loss of capillitium and
formation of pseudocapillitium, have occurred independently. Therefore, the type of the fruiting body, and presence of
capillitium or pseudocapillitium, cannot be considered as sufficient criteria for separating myxomycete genera.
Key words: 18S rDNA, aethalium, capillitium, convergence, paralellism, phylogeny, pseudoaethalium, pseudocapilliti-
tum, spore ornamentation, sporocarp, sporotheca.
179ISSN 0372-4123. Укр. ботан. журн., 2016, 73(2)
Fig. 1. Phylogeny of the Reticulariaceae
based on 18S rDNA 5’-domain sequences.
The tree was constructed by Bayesian
Inference. ML bootstrap replicates above
50 and Bayesian posterior probabilities
above 0.5 are shown for each branch. A
hyphen indicates a conflicting topology.
The scale bar indicates the fraction
of substitutions per site. Asterisk: the
genotype displayed by the type specimen
of the respective taxon.
180 ISSN 0372-4123. Ukr. Bot. J., 2016, 73(2)
Last common ancestor
Alwisia lloydiae Alwisia bombarda
Confluence
of stalks
Lycogala flavofuscum
Reduction
of stalk
Reduction of hypothallus
Lycogala conicum, L. exiguum
Grouping of sporocarps
Reduction of capillitium
Alwisia morula
Stalk reduction
Alwisia repens
Pseudoaethalium formation
Lycogala epidendrum
Spongy hypothallus
Formation
of aethalium
Alwisia
Hypothallic stalk
Tubifera dimorphotheca
Formation of cylindrical
sporothecae
Tubifera corymbosa Tubifera dudkae
Tubifera ferruginosa
T.montana, T.magna
Cartilaginous peridium
Rigidotubula violacea,
R. reticulata
Siphoptychium
casparyi
Secondary
formation
of capillitium
Tubifera applanata
Formation of columella
Secondary
formation of
capillitium
Reduction
of spherical sporothecae
Tubifera microsperma,
T. pseudomicrosperma
Reduction of spherical
sporothecae
Formation of
cortex
Tubifera Reticularia
Siphoptychium?
Reticularia
splendens,
R. jurana
Reduction of capillitium
Thecotubifera
dictyoderma
?
Siphoptychium?
Lycogala
?
Reticularia lycoperdon
Reticularia lobata
Formation of
pseudocapillitium
Ancestral form
Reticularia intermedia
Branched pseudocapillitium
?
Pseudoaethalium formation
Ancestral form
Fig. 2. Evolution of the sporophore in Reticulariaceae. The genealogy of taxa corresponds to the current understanding
of the phylogeny of the group (see Fig. 1). Bars indicate the most important evolutionary changes. Dotted arrows show
the alternative evolutionary scenarios. Gray filling unites the members of the same genus.
181ISSN 0372-4123. Укр. ботан. журн., 2016, 73(2)
(R. liceoides, R. olivacea, R. aurea, R. rubiginosa,
R. simulans) on the basis of obtained data were excluded
from the family Reticulariaceae. The first two species
were removed to Cribrariaceae, with the simultaneous
re-erection of the genus Licaethalium Rostaf. (Leontyev
et al., 2015).
The current understanding of the phylogeny of
Reticulariaceae, based on these data, is shown in Fig. 1.
The results obtained have allowed to look at the
morphological diversity of Reticulariaceae from a new
point. In contrast to the traditional understanding, the
family appeared to include species with stalked and
sessile sporocarps (Alwisia lloydiae, A. repens, A. morula),
pseudoaethalia with the entire cortex (Thecotubifera
dictyoderma) and pseudoaethalia, composed of spherical
sporothecae (Tubifera dudkae). The true capillitium was
found to be rather common structure within the family,
being developed in Alwisia, Lycogala and Siphoptychium,
while the threads, described in sporocarps of Tubifera
(Nannenga-Bremekamp, 1991; Neubert et al., 1993),
appeared to be fungal hyphae, feeding with the spores,
but not a capillitium (Leontyev et al., 2015).
New knowledge about the phylogeny of the
group, together with a deeper understanding of its
morphological variability, gives us a chance to understand
how the structure of these organisms was changed in the
process of evolution. In a response to this possibility,
we composed a concept which describes regularities of
the evolutionary changes of the sporophore structure in
Reticulariaceae (Fig. 2).
According to the 18S rDNA phylogeny (Fig. 1),
the basis of the evolutionary tree of the Reticulariaceae
is formed by the genus Alwisia. This correlates with
its morphological features, among which the type
of sporophore is of greatest importance. It is proven
that the primary type of myxomycetes sporophore is
the stalked sporocarp: such fructifications are typical
for all sister groups of myxomycetes (Dictyodtelia,
Ceratiomyxales, Protostelia s. l.), and also for the
Echinosteliales, the order which is believed to be closest
to the last common ancestor of myxomycetes (Fiore-
Donno et al., 2005; 2008; 2012). Within Lucisporidia
(the bright-spored myxomycetes), the basal position is
occupied by the Cribrariaceae, most of which also have
stalked sporocarps. Finally, this is the only type of the
fruiting body known in all the orders of Myxogastrea.
Therefore, the presence of this structure makes Alwisia
the best candidate for the role of the last common
ancestor of Reticulariaceae.
The next plesiomorphic character of Alwisia is the
presence of a well-developed capillitium, resembling
that of another bright-spored myxomycetes,
Dianemataceae and Trichiaeceae. This similarity
supports a hypothesis that the presence of capillitium
is the basal characteristic for the Reticularaceae and for
the Lucisporodia in general.
Among species of Alwisia, two are characterized
by well developed, free stalks, and another two
(A. bombarda, A. lloydiae) – by the presence of
capillitium (A. morula, A. lloydiae). There is only one
known species, which unites both plesiomorphies lost
by most of Reticulariaceae, the A. lloydiae, which we
consider as the closest to the last common ancestor
of the family. It is noteworthy that this species occurs
in the Australian continent and in Tasmania, the well
known refuges of a relict biota.
From the basal form, related with A. lloydiae, at
least three evolutionary lineages have originated. First
of them is represented by the vector A. lloydiae →
A. morula → A. repens, in which the sporothecae lose
the capillitium (A. morula) and their stalks become
procumbent (A. repens). We suppose that this lineage
has continued in the genera Tubifera, Rigidotubula,
and probably Siphoptychium. Simultaneously to the
loss of capillitium and stalks, the spore dissemination
type changed from the active way, with the help of
capillitium, to the passive one, using rain drops and/or
insects.
Another vector is formed by species which did not
lose the capillitium but grouped their sporothecae in a
common confluent stalk (A. bombarda). Sporothecae
became thinner, as it happens in all Reticulariaceae with
tightly grouped sporothecae (see below).
Finally, the third lineage joins Alwisia with
the phylogenetic cluster, which unites Lycogala,
Thecotubifera and Reticularia. The consequence of
branching of this contemporary genera from Alwisia-like
ancestor is not yet clear; however, taking into account
the attribution of sporophores of Lycogala to the free
sporocarps (Leontyev et al., 2014c), this genus appears
to be the most «archaic» representative of the group.
This is noteworthy that among Lycogala, Thecotubifera
and Reticularia, the only first genus has the tubular
capillitium, close in its morphology, ultrastructure and
ontogenesis to that of Alwisia.
The lineage which binds A. lloydiae with the genus
Tubifera, is divided to two subclades that we call «small-
spored» and «large-spored» (Leontyev et al., 2015).
First of them includes the species of Tubifera with the
182 ISSN 0372-4123. Ukr. Bot. J., 2016, 73(2)
spores less than 6.5 μm diam., mostly annulate and non-
iridescent peridium, flattened apices of sporothecae
and a dull pigmentation of the young sporophores.
Another subclade consists of the species with the spores
larger than 6.5 μm, mostly iridescent peridium, covered
with wavy folds, the convex apices of sporothecae and
bright pigmentation of immature sporophores. These
two subclades within Tubifera must have a common
ancestor. It probably had spherical sporothecae, found
in its evolutionary predecessor, A. repens. However, in
contrast with the last one, common ancestor of Tubifera
should have a spongy hypothallus, which is absent in all
Alwisia, but well developed in both subclades of Tubifera.
The structure, which meets these requirements, occurs
in T. dudkae which belongs to the «large-spored» clade.
All other species of the «large-spored» clade
(T. ferruginosa, T. magna, T. montana) have cylindrical
sporothecae, the presence of which seems to be the usual
result of the accretion of sporothecae in Reticulariaceae.
Independently, the transformation of spherical
sporothecae to cylindrical ones took place in the
«small-spored» clade. However, in this case we know
contemporary species which retained both types of
sporothecae: T. corymbosa, in which all cylindrical and
spherical sporothecae are seated on spongy hypothallus,
and T. dimorphotheca, in which spherical sporothecae
are situated on the lateral surface of the hypothallic
stalk, while cylindrical ones occupy its top. The
reduction of spherical sporothecae in T. dimorphotheca
could produce the forms similar to T. microsperma and
T. pseudomicrosperma.
The formation of the hypothallic stalk, characteristic
to T. dimorphotheca and T. microsperma, correlates
with the decreasing of the size of pseudoaethalium.
This suggests that the recovery of the stalks, completely
lost in the previous stages of Tubifera evolution, was
caused by the partial return to the strategy of the stalked
sporocarp, because the distribution using the rain drops
and insects is effective only in large fruiting bodies
(Schnittler, 2002).
The point of divergence, which led to the formation
of T. applanata within the «small-spored» clade,
remains unclear. This species is the only member of the
subcluster, which have large, sessile pseudoaethalia with
closely accreted prismatic sporothecae. Its origin from
an ancestor, common with T. corymbosa, looks the most
«parsimonic» scenario: it needs only the loss of spherical
sporothecae, while any other hypotheses claim for the
reduction of hypothallic stalk. Molecular data support
the origination of T. applanata and T. corymbosa from
the common ancestor.
Sporophores of the genera Rigidotubula and
Siphoptychium represent a further development of
tubiferoid pseudoaethalia with prismatic sporothecae,
which have got the cartilaginous peridium and
columella. With the appearance of these structures, the
Rigidotubula can be easily derived from T. applanata.
However, the molecular data do not support this simple
scenario and show that Rigidotubula has branched
from the trunk of Tubifera even before the last one was
subdivided on «small-» and «large-spored» clades, and
therefore, before Tubifera formed pseudoaethalia with
cylindrical sporothecae. It means that Rigidotubula
should has gone all the way from the heap of spherical
sporothecae to the pseudoaethalium, composed of
prismatic ones, independently from Tubifera. This is a
prominent example of parallel evolution.
The absence of molecular data about the phylogenetic
relations of the genus Siphoptychium does not allow to
determine its position in our evolutionary scheme.
However, the complex of morphological features, like
a prismatic sporothecae, cartilaginous peridium and
well developed columella, puts this genus close to
Rigidotubula. On the other hand, it should be noted
that the most important feature of Siphoptychium is
the branching of columella and the presence of the
perforations at the places where columella merges the
peridium. Both of these features bring Siphoptychium
close to Alwisia (the last one does not have columella,
but its capillitial threads have a similar structure
and ornamentation with the branches of columella
in Siphoptychium). Therefore, Siphoptychium may
represent a separate lineage of pseudoaethaliate forms
that comes directly from Alwisia-like ancestor.
The Lycogala inherited from Alwisia its tubular
capillitium, but not the stalk. Its sessile sporocarps
are the most probable predecessors of the aethalia
known in Reticularia. The stage, intermediate between
grouped sporocarps and aethalium, is known within this
lineage only in Thecotubifera, where the fruiting body
consists of prismatic sporothecae which form together
an entire, indivisible cortex. This allows to imagine a
simple evolutionary scenario, such as: Lycogala (sessile
sporocarps) → Thecotubifera (pseudoaethalia) →
Reticularia (aethalia). However, at least one reason
does not allow to support this hypothesis. Peculiarities
of the spores in Th. dictyoderma, such as very large
meshes with high borders, separate this species from all
another Reticulariaceae, while in Reticularia the spore
ornamentation is typical for the family (small meshes with
relatively low borders). It is unlikely that the ancestors of
183ISSN 0372-4123. Укр. ботан. журн., 2016, 73(2)
Reticularia once lost a typical ornamentation, and then
developed it again. Moreover, the species of Reticularia
with well-developed pseudocapillitium (R. splendens,
R. jurana) show the remnants of confluence of spherical,
bag-shaped or vermicular sporothecae, but not straight
and prismatic ones, like those in Thecotubifera. Thus,
much preferable is the scenario of independent origin of
Reticularia and Thecotubifera from a common ancestor,
that had a rudimentary pseudoaethalia (such a form is
yet unknown) or sporocarps (Lycogala). A link between
Lycogala and Reticularia can represent R. lobata, since
the latter, according to our assumption, has not aethalia,
but spherical sporocarps, that form the primitive
pseudoaethalia. This corresponds to the characters of
pseudoaethaliate ancestor of aethaliate Reticularia,
because, as noted above, it probably did not have
cylindrical sporothecae. The structure of sporophores
in R. lobata is very close to L. epidendrum and differs
from the latter only by lack of tubular capillitium, the
loss of which took place here independently from the
vector Alwisia → Tubifera.
The abovementioned discussion allows to make
several conclusions:
1. The general tendency of the morphological
evolution in Reticulariaceae is the transformation
of the stalked forms to the sessile ones, followed
by the formation of pseudoaethalia and aethalia.
Simultaneously, the spore dissemination type was
changed from the active way with the help of capillitium
to the passive one, through rain and insects.
2. The common ancestor of the family had stalked
sporocarps with well-developed tubular capillitium and
was similar in structure to Alwisia lloydiae. This ancestral
form produced at least four evolutionary lineages: 1)
Alwisia → Tubifera, 2) Alwisia → Rigidotubula (and
Siphoptychium, if it does not form a separate branch),
3) Alwisia → (Lycogala) → Thecotubifera, 4) Alwisia →
Lycogala → Reticularia lobata → aethalial species of
Reticularia.
3. The evolution of the fruiting bodies in
Reticulariaceae is characterized by the parallelism.
Processes like a loss of stalks, transition from spherical
to elongated sporothecae, transformation of sporocarps
to pseudoaethalia and aethalia, loss of capillitium
and formation of pseudocapillitium have occurred
repeatedly. Therefore, the morphological similarity
of the fruiting bodies does not necessarily indicate the
affinity of species. A type of the fruiting body cannot be
considered as a sufficient criterion for the separation of
the genera in myxomycetes.
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Леонтьєв Д.В. Еволюція спорофора у Reticulariaceae
(Myxomycetes). – Укр. ботан. журн. – 2016. –73(2): 178—184.
Харківська державна зооветеринарна академія, кафедра
біотехнології
вул. Академічна, 1, Мала Данилівка, Харків, 62341,
Україна
Молекулярно-генетичні дослідження прояснили філоге-
нетичні відносини між таксонами родини Reticulariaceae
(Myxomycetes, Myxogastrea), надали змогу реконструюва-
ти шляхи морфологічної еволюції цього таксона. Згідно
з одержаними даними загальною тенденцією еволюції
спорофорів у Reticulariaceae є трансформація стебельчас-
тих форм у сидячі, яка супроводжувалася формуванням з
вільних спорокарпів метамерних плодових тіл ‒ псевдо-
еталіїв та еталіїв. Спільний предок родини був подібний
до сучасної Alwisia lloydiae і мав стебельчасті спорокарпи
та трубчастий капіліцій. Від цього предка започатковані
принаймні чотири філогенетичні лінії: Alwisia → Tubif-
era; Alwisia → Rigidotubula (та, ймовірно, Siphoptychium);
Alwisia → (Lycogala) → Thecotubifera; Alwisia → Lycogala
→ Reticularia. У кожній із цих ліній незалежно відбували-
ся такі еволюційні процеси, як втрата ніжок, перехід від
сферичних до циліндричних споротек і від одиночних до
згрупованих спорокарпів, втрата капіліцію та формуван-
ня псевдокапіліцію. Зважаючи на це, тип плодового тіла
та наявність капіліцію або псевдокапіліцію не можуть
розглядатися як вирішальні критерії для розмежування
родів міксоміцетів.
Ключові слова: 18S рДНК, еталій, капіліцій, паралелізм,
філогенія, псевдоеталій, псевдокапіліцій, орнаментація
спор, спорокарп, споротека.
Леонтьев Д.В. Эволюция спорофора у Reticulariaceae
(Myxomycetes). – Укр. ботан. журн. – 2016. –73(2): 178—184.
Харьковская государственная зооветеринарная
академия, кафедра биотехнологии
ул. Академическая, 1, Малая Даниловка, Харьков,
62341, Украина
Молекулярно-генетические исследования прояснили
филогенетические взаимоотношения между таксонами
семейства Reticulariaceae (Myxomycetes, Myxogastrea), дав
возможность реконструировать пути морфологической
эволюции этого таксона. Согласно полученным данным,
общей тенденцией в эволюции спорофора Reticulariaceae
является трансформация стебельчатых форм в сидячие,
сопровождавшаяся формированием из свободных спо-
рокарпов метамерных плодовых тел – псевдоэталиев и
эталиев. Общий предок семейства был близок к совре-
менной Alwisia lloydiae; он имел стебельчатые спорокар-
пы и трубчатый капиллиций. От этого предка берут на-
чало, по крайней мере, четыре филогенетические линии:
Alwisia → Tubifera; Alwisia → Rigidotubula (и, вероятно,
Siphoptychium); Alwisia → (Lycogala) → Thecotubifera; Al-
wisia → Lycogala → Reticularia. В каждой из этих линий
независимо происходили такие эволюционные процес-
сы, как потеря ножек, переход от сферических споро-
тек к цилиндрическим и от одиночных спорокарпов к
сгруппированным, потеря капиллиция и формирование
псевдокапиллиция. В связи с этим тип плодового тела, а
также наличие капиллиция и псевдокапиллиция не мо-
гут рассматриваться в качестве решающих критериев для
разграничения родов миксомицетов.
Ключевые слова: 18S рДНК, эталий, капиллиций,
параллелизм, филогения, псевдоэталий,
псевдокапиллиций, орнаментация спор, спорокарп,
споротека.
Savchenko K.G., Wasser S.P., Heluta V.P., Nevo E. Smut fungi of Israel (Biodiversity of Cyanoprocaryotes, Algae
and Fungi of Israel) / Institute of Evolution and Faculty of Natural Sciences, University of Haifa, Israel. – Koeltz
Scientific Books, 2015. – 160 p.
The book covers smut fungi (Ustilaginomycetes p.p., Exobasidiomycetes p.p., and Microbotryales) of Israel. A total
of 73 species in 15 genera are described. Three genera (Macalpinomyces, Melanustilospora and Schizonella) are new
records for Israel. The book is divided into two main parts: General Part providing data regarding environmental
conditions of Israel, morphology of smut fungi, materials and methods, historical studies, as well as the analysis of
species composition of smut fungi in Israel; Special Part (taxonomic part) providing information regarding detailed
macro- and micromorphological descriptions, distribution in Israel, general distribution, and notes.
Видання охоплює сажкові гриби (Ustilaginomycetes p.p., Exobasidiomycetes p.p. і Microbotryales) Ізраїлю.
Описано 73 види з 15 родів, три роди (Macalpinomyces, Melanustilospora і Schizonella) є новими для
Ізраїлю. Книга складається з двох основних частин. Загальна містить відомості про стан навколишнього
середовища країни, морфологію сажкових грибів, аналіз їх видового складу в Ізраїлі, матеріали та методи,
історію досліджень цих грибів. У спеціальній (таксономічній) частині монографії наводиться детальна
інформація про макро- та мікроморфологію видів сажкових грибів, їхнє поширення в Ізраїлі та світі,
подано також примітки.
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