Paper · 2022 · PeerJ

Need for split: integrative taxonomy reveals unnoticed diversity in the subaquatic species of Pseudohygrohypnum (Pylaisiaceae, Bryophyta)

Vladimir E. Fedosov; Anna V. Shkurko; Alina V. Fedorova; et al.

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Open access · cc-by

Abstract

We present an integrative molecular and morphological study of subaquatic representatives of the genus Pseudohygrohypnum (Pylaisiaceae, Bryophyta), supplemented by distribution modelling of the revealed phylogenetic lineages. Phylogenetic analyses of nuclear and plastid datasets combined with the assemble species by automatic partitioning (ASAP) algorithm revealed eight distinct species within the traditionally circumscribed P. eugyrium and P. subeugyrium . These species are therefore yet another example of seemingly widely distributed taxa that harbour molecularly well-differentiated lineages with narrower distribution ranges. Studied accessions that were previously assigned to P. eugyrium form three clearly allopatric lineages, associated with temperate regions of Europe, eastern North America and eastern Asia. Remarkably, accessions falling under the current morphological concept of P. subeugyrium were shown to be even more diverse, containing five phylogenetic lineages. Three of these lineages occur under harsh Asian continental climates from cool-temperate to Arctic regions, while the remaining two, referred to P. subeugyrium s.str. and P. purpurascens , have more oceanic North Atlantic and East Asian distributions. Niche identity and similarity tests suggested no similarity in the distributions of the phylogenetically related lineages but revealed the identity of two East Asian species and the similarity of two pairs of unrelated species. A morphological survey confirmed the distinctness of all eight phylogenetic lineages, requiring the description of five new species. Pseudohygrohypnum appalachianum and P. orientale are described for North American and East Asian plants of P. eugyrium s.l., while P. sibiricum , P. subarcticum and P. neglectum are described for the three continental, predominantly Asian lineages of P. subeugyrium s.l. Our results highlight the importance of nontropical Asia as a center of bryophyte diversity. Phylogenic dating suggests that the diversification of subaquatic Pseudohygrohypnum lineages appeared in late Miocene, while mesophilous species of the genus split before Miocene cooling, in climatic conditions close to those where the ancestor of Pseudohygrohypnum appeared. We speculate that radiation of the P. subeugyrium complex in temperate Asia might have been driven by progressive cooling, aridification, and increases in seasonality, temperature and humidity gradients. Our results parallel those of several integrative taxonomic studies of North Asian mosses, which have resulted in a number of newly revealed species. These include various endemics from continental areas of Asia suggesting that the so-called Rapoport’s rule of low diversity and wide distribution range in subpolar regions might not be applicable to bryophytes. Rather, the strong climatic oscillations in these regions may have served as a driving force of speciation and niche divergence.

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Claims 14

Observationobservedoriginal resultmeasured

Claim · model-written summary

Phylogenetic analyses revealed eight distinct species within P. eugyrium and P. subeugyrium.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
distinct speciese0
Value
eighte0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“revealed eight distinct species within the traditionally circumscribed P. eugyrium and P. subeugyrium”

✓ located verbatim in the source · abstract, characters 358–459 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Accessions previously assigned to P. eugyrium form three allopatric lineages associated with temperate Europe, eastern North America and eastern Asia.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
allopatric lineagese0
Value
threee0
Setting
temperate regions of Europee0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“Studied accessions that were previously assigned to P. eugyrium form three clearly allopatric lineages, associated with temperate regions of Europe, eastern North America and eastern Asia”

✓ located verbatim in the source · abstract, characters 636–823 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Accessions assigned to P. subeugyrium contain five phylogenetic lineages.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
phylogenetic lineagese0
Value
fivee0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“accessions falling under the current morphological concept of P. subeugyrium were shown to be even more diverse, containing five phylogenetic lineages”

✓ located verbatim in the source · abstract, characters 837–987 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Three of the P. subeugyrium lineages occur under harsh Asian continental climates from cool-temperate to Arctic regions.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
occur under harsh Asian continental climatese0
Setting
cool-temperate to Arctic regionse0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
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“Three of these lineages occur under harsh Asian continental climates from cool-temperate to Arctic regions”

✓ located verbatim in the source · abstract, characters 989–1095 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

The remaining two P. subeugyrium lineages have more oceanic North Atlantic and East Asian distributions.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
distributionse0
Setting
more oceanic North Atlantic and East Asiane0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“while the remaining two, referred to P. subeugyrium s.str. and P. purpurascens , have more oceanic North Atlantic and East Asian distributions”

✓ located verbatim in the source · abstract, characters 1097–1239 · show in text ↑
Observationobservedoriginal resultno effectmeasured

Claim · model-written summary

Niche identity and similarity tests suggested no similarity in the distributions of phylogenetically related lineages.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
no similarity in the distributionse0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“Niche identity and similarity tests suggested no similarity in the distributions of the phylogenetically related lineages but revealed the identity of two East Asian species and the similarity of two pairs of unrelated species”

✓ located verbatim in the source · abstract, characters 1241–1467 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Niche tests revealed the identity of two East Asian Pseudohygrohypnum species.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
identity of two East Asian speciese0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“Niche identity and similarity tests suggested no similarity in the distributions of the phylogenetically related lineages but revealed the identity of two East Asian species and the similarity of two pairs of unrelated species”

✓ located verbatim in the source · abstract, characters 1241–1467 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Niche tests revealed similarity of two pairs of unrelated Pseudohygrohypnum species.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
similarity of two pairs of unrelated speciese0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“Niche identity and similarity tests suggested no similarity in the distributions of the phylogenetically related lineages but revealed the identity of two East Asian species and the similarity of two pairs of unrelated species”

✓ located verbatim in the source · abstract, characters 1241–1467 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

A morphological survey confirmed the distinctness of all eight phylogenetic lineages.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
distinctness of all eight phylogenetic lineagese0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“A morphological survey confirmed the distinctness of all eight phylogenetic lineages, requiring the description of five new species”

✓ located verbatim in the source · abstract, characters 1469–1600 · show in text ↑
Observationobservedoriginal resultbasis n/a

Claim · model-written summary

Pseudohygrohypnum appalachianum and P. orientale are described as new species for North American and East Asian plants.

Taxon
Pseudohygrohypnum appalachianum and P. orientalePseudohygrohypnum appalachianumPseudohygrohypnum orientalee0
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are describede0

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e0

“Pseudohygrohypnum appalachianum and P. orientale are described for North American and East Asian plants of P. eugyrium s.l.”

✓ located verbatim in the source · abstract, characters 1602–1725 · show in text ↑
Observationobservedoriginal resultbasis n/a

Claim · model-written summary

P. sibiricum, P. subarcticum and P. neglectum are described as new species for three continental Asian lineages.

Outcome
are describede0

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e0

“P. sibiricum , P. subarcticum and P. neglectum are described for the three continental, predominantly Asian lineages of P. subeugyrium s.l.”

✓ located verbatim in the source · abstract, characters 1733–1872 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Phylogenetic dating suggests that diversification of subaquatic Pseudohygrohypnum lineages appeared in the late Miocene.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
the diversification of subaquatic Pseudohygrohypnum lineagese0
Qualifier
appeared in late Miocenee0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“Phylogenic dating suggests that the diversification of subaquatic Pseudohygrohypnum lineages appeared in late Miocene”

✓ located verbatim in the source · abstract, characters 1966–2083 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Mesophilous Pseudohygrohypnum species split before Miocene cooling.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
mesophilous species of the genus splite0
Qualifier
before Miocene coolinge0

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t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“while mesophilous species of the genus split before Miocene cooling, in climatic conditions close to those where the ancestor of Pseudohygrohypnum appeared”

✓ located verbatim in the source · abstract, characters 2085–2240 · show in text ↑
Mechanismhypothesisoriginal resultbasis n/a

Claim · model-written summary

Radiation of the P. subeugyrium complex in temperate Asia might have been driven by progressive cooling, aridification and increased seasonality and gradients.

Taxon
PseudohygrohypnumPseudohygrohypnumt0
Outcome
radiation of the P. subeugyrium complexe0
Qualifier
progressive cooling, aridification, and increases in seasonality, temperature and humidity gradientse0

Verbatim

t0

“subaquatic species of Pseudohygrohypnum”

✓ located verbatim in the source · title, characters 72–111 · show in text ↑
e0

“We speculate that radiation of the P. subeugyrium complex in temperate Asia might have been driven by progressive cooling, aridification, and increases in seasonality, temperature and humidity gradients”

✓ located verbatim in the source · abstract, characters 2242–2444 · show in text ↑