Abstract
Seed morphology of Hydrangeaceae was explored to describe the parameters of diversity, define characters and character states, and hypothesize monophyletic groups in the family on the basis of derived features. Eleven seed characters were defined that delimit four basic aspects of morphology, including (1) shape, (2) primary sculpture, (3) secondary sculpture, and (4) appendages. Character states could not be delimited readily in some aspects of diversity, including transectional shape and length of seeds. Despite continuous variation in seed length, when mapped on previous cladograms these data supported the hypothesis that seeds less than 1.0 mm long were plesiomorphic and that seed size had increased separately in different subclades. Seed size increased in association with wing loss in Fendlera, Fendlerella, and Whipplea. A shift from apical, interstylar dehiscence of fruits that produce a very limited gap for seed emergence in most members of the hydrangea clade to fruit wall fragmentation may have permitted an increase of seed size in Decumaria, Pileostegia, and Schizophragma. The data help to support hypotheses of monophyly for Deutzia and Philadelphus, although relevant character states appear to display parallelisms. The data also help to demonstrate the paraphyly of Hydrangea as traditionally circumscribed. A derived urceolate seed form allies Broussaisia and Dichroa with Hydrangea hirta and Hydrangea luteovenosa. These data indicate that fossil seeds recognized as Dichroa from the Middle Oligocene of Europe may be indistinguishable from those of certain Hydrangea species. Hydrangea heteromalla and Hydrangea paniculata share derived long, spindle-form seeds with a flange at the micropylar pole with Decumaria and Schizophragma, and these taxa may form a monophyletic group that includes Pileostegia. The tropical species of Hydrangea section Cornidia share a derived rarity of anticlinal crosswalls in the seed body. Current phylogenetic hypotheses indicate that the micropylar cowls that Cardiandra and Deinanthe share with Deutzia are evolutionary parallelisms. A derived funicular appendage supports prior hypotheses of monophyly for Fendlerella and Whipplea. These two genera may be most closely related to Fendlera, with which they share seed coat cells with convex long axes.
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Claims 15
Claim · model-written summary
Eleven seed characters were defined across four morphological aspects.
- Taxon
- HydrangeaceaeHydrangeaceaet0
- Outcome
- Eleven seed characterse0
- Value
- Elevene0
Verbatim
e0“Eleven seed characters were defined that delimit four basic aspects of morphology, including (1) shape, (2) primary sculpture, (3) secondary sculpture, and (4) appendages.”
t0“Hydrangeaceae”
Claim · model-written summary
Some character states could not be readily delimited, including seed transectional shape and length.
- Taxon
- HydrangeaceaeHydrangeaceaet0
- Organ / stage
- seedse0
- Outcome
- Character states could not be delimited readilye0
Verbatim
t0“Hydrangeaceae”
e0“Character states could not be delimited readily in some aspects of diversity, including transectional shape and length of seeds.”
Claim · model-written summary
Seed length data supported seeds under 1.0 mm as ancestral and independent seed size increases in subclades.
- Taxon
- HydrangeaceaeHydrangeaceaet0
- Outcome
- seeds less than 1.0 mm long were plesiomorphice0
- Value
- less than 1.0 mme0
Verbatim
t0“Hydrangeaceae”
e0“Despite continuous variation in seed length, when mapped on previous cladograms these data supported the hypothesis that seeds less than 1.0 mm long were plesiomorphic and that seed size had increased separately in different subclades.”
Claim · model-written summary
Seed size increased in association with wing loss in Fendlera, Fendlerella and Whipplea.
- Taxon
- Fendlera, Fendlerella, and Whipplea
Fendlera
FendlerellaWhippleae0
- Organ / stage
- winge0
- Outcome
- Seed sizee0
- Direction
- increasede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e0“Seed size increased in association with wing loss in Fendlera, Fendlerella, and Whipplea.”
Claim · model-written summary
Fruit wall fragmentation may have permitted larger seeds in Decumaria, Pileostegia and Schizophragma.
- Taxon
- Decumaria, Pileostegia, and SchizophragmaHydrangea sect. DecumariaHydrangea sect. PileostegiaSchizophragmae0
- Outcome
- increase of seed sizee0
- Qualifier
- fruit wall fragmentatione0
Verbatim
e0“A shift from apical, interstylar dehiscence of fruits that produce a very limited gap for seed emergence in most members of the hydrangea clade to fruit wall fragmentation may have permitted an increase of seed size in Decumaria, Pileostegia, and Schizophragma.”
Claim · model-written summary
Seed data support monophyly hypotheses for Deutzia and Philadelphus, with apparent parallel character states.
- Taxon
- Deutzia and Philadelphus
Deutzia
Philadelphuse0 - Outcome
- hypotheses of monophylye0
- Qualifier
- character states appear to display parallelismse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e0“The data help to support hypotheses of monophyly for Deutzia and Philadelphus, although relevant character states appear to display parallelisms.”
Claim · model-written summary
Seed data indicate traditional Hydrangea is paraphyletic.
- Outcome
- paraphylye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The data also help to demonstrate the paraphyly of Hydrangea as traditionally circumscribed.”
Claim · model-written summary
A derived urceolate seed form allies Broussaisia and Dichroa with H. hirta and H. luteovenosa.
- Taxon
- Broussaisia and DichroaHydrangea sect. BroussaisiaHydrangea sect. Dichroae0
- Outcome
- derived urceolate seed forme0
- Comparator
- Hydrangea hirta and Hydrangea luteovenosae0
Verbatim
e0“A derived urceolate seed form allies Broussaisia and Dichroa with Hydrangea hirta and Hydrangea luteovenosa.”
Claim · model-written summary
Middle Oligocene European fossil seeds assigned to Dichroa may be indistinguishable from some Hydrangea seeds.
- Taxon
- DichroaHydrangea sect. Dichroae0
- Outcome
- may be indistinguishablee0
- Comparator
- certain Hydrangea speciese0
- Setting
- Europee0
Verbatim
e0“These data indicate that fossil seeds recognized as Dichroa from the Middle Oligocene of Europe may be indistinguishable from those of certain Hydrangea species.”
Claim · model-written summary
H. heteromalla and H. paniculata share derived spindle-form seeds with Decumaria and Schizophragma.
- Taxon
- Hydrangea heteromalla and Hydrangea paniculata
Hydrangea paniculata
Hydrangea heteromallae0 - Outcome
- derived long, spindle-form seeds with a flange at the micropylar polee0
- Comparator
- Decumaria and Schizophragmae0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Hydrangea heteromalla and Hydrangea paniculata share derived long, spindle-form seeds with a flange at the micropylar pole with Decumaria and Schizophragma, and these taxa may form a monophyletic group that includes Pileostegia.”
Claim · model-written summary
The spindle-seeded taxa may form a monophyletic group including Pileostegia.
- Taxon
- HydrangeaceaeHydrangeaceaet0
- Outcome
- monophyletic group that includes Pileostegiae0
Verbatim
e0“these taxa may form a monophyletic group that includes Pileostegia”
t0“Hydrangeaceae”
Claim · model-written summary
Tropical Hydrangea section Cornidia has a derived rarity of anticlinal crosswalls in the seed body.
Verbatim
e0“The tropical species of Hydrangea section Cornidia share a derived rarity of anticlinal crosswalls in the seed body.”
Claim · model-written summary
Micropylar cowls shared by Cardiandra, Deinanthe and Deutzia are interpreted as evolutionary parallelisms.
- Taxon
- Cardiandra and DeinantheHydrangea sect. CardiandraHydrangea sect. Deinanthee0
- Outcome
- evolutionary parallelismse0
- Comparator
- Deutziae0
Verbatim
e0“Current phylogenetic hypotheses indicate that the micropylar cowls that Cardiandra and Deinanthe share with Deutzia are evolutionary parallelisms.”
Claim · model-written summary
A derived funicular appendage supports monophyly of Fendlerella and Whipplea.
- Taxon
- Fendlerella and Whipplea
FendlerellaWhippleae0
- Organ / stage
- funicular appendagee0
- Outcome
- hypotheses of monophylye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“A derived funicular appendage supports prior hypotheses of monophyly for Fendlerella and Whipplea.”
Claim · model-written summary
Fendlerella and Whipplea may be closest to Fendlera and share convex long-axis seed coat cells.
- Taxon
- Fendlerella and Whipplea
FendlerellaWhippleae1
- Outcome
- most closely related to Fendlerae0
- Qualifier
- seed coat cells with convex long axese0
- Comparator
- Fendlerae0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These two genera may be most closely related to Fendlera, with which they share seed coat cells with convex long axes.”
e1“A derived funicular appendage supports prior hypotheses of monophyly for Fendlerella and Whipplea.”