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<mods:title id="4D20F98790E34DE121EDBD322997DFD4">An integrative taxonomic study of the genus Lethocolea (Marchantiophyta: Acrobolbaceae)</mods:title>
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<mods:namePart id="5E1B75502A622320400BB5BE6FD4CD09">Gradstein, S. Robbert</mods:namePart>
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<subSubSection id="4571335FBD7D439FD056FA04736A9449" type="nomenclature">
<paragraph id="EDBB4F3AB6C02B90E6E9A02EAC92D8FB">
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<taxonomicName id="5852F3442EC72617C1C7D84C6015AB3D" ID-CoL="5DPH" authority="Mitt." authorityName="Mitt." class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="BC28AA7BBDF5861C83C0BE7B40FBDADF" italics="true">Lethocolea</emphasis>
Mitt.
</taxonomicName>
(
<treatmentCitation id="9586219BBE447622CFCF551D45FC2D60" author="Hooker JD" page="751" year="1867">
<bibRefCitation id="455723A1161030F70E03E436793C016C" author="Hooker JD" refId="B26" refString="Hooker JD (1867) Handbook of the New Zealand Flora 2. Reeve &amp; Co., London, 393 - 798." year="1867">Hooker 1867: 751</bibRefCitation>
</treatmentCitation>
, 753) nom. cons.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="A2B49BA1DF41C5F3218E59343F9ED4EC" type="reference_group">
<paragraph id="9827AC1C6CEE736D3966C1D90ABF3804">
<treatmentCitationGroup id="FC2B3CF574DB72859CEBE18E28BCD298">
<taxonomicName id="E282114765CE08411F24713DAF168412" ID-CoL="9RL6R" authority="Taylor" authorityName="Taylor" class="Jungermanniopsida" family="Acrobolbaceae" genus="Podanthe" higherTaxonomySource="CoL,GBIF" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="7A45CB21E140BC8DBDA6B943BB55D95C" italics="true">Podanthe</emphasis>
<comment id="9CA371BE22342D312569355BBF94AB15">
Taylor (
<treatmentCitation id="8934EA756237AB9F844C7A3FA529914A" author="Taylor T" page="413" year="1846">
<bibRefCitation id="B0B51B9EE310927BC9FD6CD9C1BE298D" author="Taylor T" firstAuthor="Taylor" page="413" refId="B61" refString="Taylor T (1846) New Hepaticae. London Journal of Botany 5: 258284, 365417." year="1846">Taylor 1846: 413</bibRefCitation>
</treatmentCitation>
) Type species:
<taxonomicName id="F3D8FC2CE238F015C15D8A3E18024650" ID-CoL="9MZDF" authority="Taylor" authorityName="Taylor" class="Jungermanniopsida" family="Acrobolbaceae" genus="Podanthe" higherTaxonomySource="CoL,GBIF" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="squamata">
<emphasis id="A833F2761B9EBD738CE68D310235913B" italics="true">Podanthe squamata</emphasis>
Taylor
</taxonomicName>
(=
<taxonomicName id="B8CA50C336EA3795F775538BF02869BF" ID-CoL="6Q2MH" authority="(Taylor) G. A. M. Scott &amp; K. G. Beckm." authorityName="G. A. M. Scott &amp; K. G. Beckm." baseAuthorityName="Taylor" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="pansa">
<emphasis id="B63B2CB64EEF8F3E13DB883A7DB6B8AB" italics="true">Lethocolea pansa</emphasis>
(Taylor) G. A. M. Scott &amp; K. G. Beckm.
</taxonomicName>
).
</comment>
</taxonomicName>
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</paragraph>
</subSubSection>
<subSubSection id="SECID0E5JBG" type="type species">
<paragraph id="412C24696B5FC1184CDAEA2FAB1209C5">
<heading id="BF11D2CE2874497A8733926F943F6009" reason="title">Type species.</heading>
</paragraph>
<paragraph id="694981639A8CB17B3D60B52E22B7B62D">
<taxonomicName id="000FDF06E8EE4D0B92BC992ECA815ACE" authority="Mitt." authorityName="Mitt." class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="drummondii">
<emphasis id="BB2CC3093362A756D0889DD532AC9F0C" italics="true">Lethocolea drummondii</emphasis>
Mitt.
</taxonomicName>
, nom. illeg. (=
<taxonomicName id="2B891166AE4056488EE19914C97BA043" authority="(Taylor) G. A. M. Scott &amp; K. G. Beckm." authorityName="G. A. M. Scott &amp; K. G. Beckm." baseAuthorityName="Taylor" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="pansa">
<emphasis id="59C0539722E0FB4573CCFBA9BA09075E" italics="true">L. pansa</emphasis>
(Taylor) G. A. M. Scott &amp; K. G. Beckm.
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0E1KBG" type="description">
<paragraph id="F319B085885736A6A0F886A313CD2666">
<heading id="BB57C94444B21983C7A5ADBD2BC5EF8F" reason="title">Description.</heading>
</paragraph>
<paragraph id="FF50476E88B2756B23950DFFC96AD134">
<underline id="F934EE479C9018F3351423C5D8340ADD">Plants</underline>
dioicous, rarely paroicous (
<taxonomicName id="7472C05658B43F9C0DF62E7DC831B2C9" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="naruto-toganensis">
<emphasis id="0153BF65E8B97B57688431E49AFC096F" italics="true">L. naruto-toganensis</emphasis>
</taxonomicName>
), prostrate, pale green to yellowish-green to olive-green to brownish, sometimes reddish or purplish, little-branched, branching ventral-intercalary or (rarely) lateral-intercalary, ventral branches stoloniform.
<underline id="A106F3DDB4D146614BFC495FF5F2CA6E">Stems</underline>
rather fleshy, usually fragile and made up of thin-walled cells, sometimes rigid and with slightly thickened epidermal walls, epidermis cells similar in size to medullary cells, rarely slightly smaller, ventral region of stem sometimes with fungal hyphae inside the cells.
<underline id="963A97D75B13281938BAA766DCA526FC">Rhizoids</underline>
scattered to slightly fascicled, hyaline to light brown to pink, rarely purplish (
<taxonomicName id="5963FE111A5A031795EABD150B983648" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="naruto-toganensis">
<emphasis id="D6EC53473A7D62660D22D00EBDD5E434" italics="true">L. naruto-toganensis</emphasis>
</taxonomicName>
), sometimes with fungal hyphae.
<underline id="219C6BF45112663B1D8208C54B401A52">Leaves</underline>
succubous, alternate, imbricate, lamina flat to convex, or concave with convex margins, undivided, suborbicular to ovate to ovate-oblong to lingulate, insertion line reaching dorsal midline of stem, apex rounded to truncate to emarginate, margins entire, sometimes with a border of thick-walled cells, dorsal leaf base not or shortly decurrent, ventral leaf base not decurrent, lower ventral half of the leaf lamina with an area of enlarged, hyaline cells with little or no chlorophyll.
<underline id="30718CE0515B08AEF01F73952B762F6A">Leaf cells</underline>
isodiametric to elongate, fully thin-walled or with very small to medium-sized trigones, trigones usually hyaline, rarely reddish, cuticle coarsely papillose or smooth; oil bodies large, 13 per cell, usually ellipsoid, dark brown to greyish-brown (exceptionally colourless), sometimes persistent in dried material.
<underline id="E85A6FC33A8F442D6620DE9DED809446">Underleaves</underline>
absent.
<underline id="24188BAEDAA4D932305FC98D412314A5">Androecia</underline>
terminal or intercalary on main shoots, bracts saccate (not saccate in
<taxonomicName id="261330D74DD2830829C202FE58D17113" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="naruto-toganensis">
<emphasis id="EC887246A3C0EAD7035E68BDE6183A63" italics="true">L. naruto-toganensis</emphasis>
</taxonomicName>
), antheridia 1 ( 6) per bract, antheridial stalk short, up to 7 cells long, irregularly biseriate.
<underline id="C91FC72D511C3D10533B928803F5D5BC">Gynoecia</underline>
terminal on main shoots, female bracts in 24 pairs, bracts slightly larger than vegetative leaves (except for the inner ones), standing upwards or spreading outwards; perianth absent; archegonia ca 2030 per gynoecium.
<underline id="04EC05288B915698A52B5095C781FFCE">Marsupia</underline>
pendent, shortly to elongate cylindrical, up to 310 ( 20) mm long, green and with smooth surface when young, brown and with usually hairy surface when mature, marsupial canal narrow, lined by few or numerous large, elongate, papilliform, mucilaginous cells; archegonia, calyptra and developing sporophyte carried to the base of the marsupium, archegonia with a short neck.
<underline id="D439B51830BA8360CF4DF7236FAD3F3C">Setae</underline>
whitish, up to
<quantity id="D00A710C7C5B3ABC80C8A8EBBF322260" metricMagnitude="-2" metricUnit="m" metricValue="1.2" unit="cm" value="1.2">1.2 cm</quantity>
long after elongation, massive, made up of ca 70 rows of cells (Fig.
<figureCitation id="1114D410AC02F2EAE3E98E8A2491BE78" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 D</figureCitation>
), with distinct trigones.
<underline id="730E6DA29B97FC86C3167977301ADF85">Capsules</underline>
cylindrical, scarcely wider than the seta, tip acute to apiculate (Fig.
<figureCitation id="6F7B7B110CADC3A54F3E61EFEA358DD3" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 C</figureCitation>
), capsule dehiscence complete or incomplete, splitting to the base into 4 or in only 23 valves with two adjacent valves remaining partially connate, capsule wall 24 - stratose (Fig.
<figureCitation id="74515F7CA2B4A334E4C61009B7109C7A" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 F</figureCitation>
), cell walls of the outer layer yellowish to reddish-brown, with small, orange to reddish-brown nodular thickenings evenly spaced along the outer longitudinal wall and turning into conspicuous thickening bands on radial walls, cell walls of the inner layers hyaline, without thickenings (seen in
<taxonomicName id="374EBA7A6356BE72191526042B42157C" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="congesta">
<emphasis id="50129A027C23125085E76AE7E121AE9D" italics="true">L. congesta</emphasis>
</taxonomicName>
only); elaters with 12 spirals (seen in
<taxonomicName id="CC14FFEA3E2021DD6E2F26B7884ECD20" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="congesta">
<emphasis id="E2CDEF9BB91FE431D51422EC67EAF3FD" italics="true">L. congesta</emphasis>
</taxonomicName>
only), narrowly fusiform, 130200 × ca 10 µm, surface finely punctate; spores isodiametrical, ca 25 µm in diameter, yellowish to brown, surface finely papillose.
<underline id="51D31404ECE066AEBA59BD86E659DE36">Asexual reproduction</underline>
(subgen.
<taxonomicName id="7DA01DA8CC0258D6E66E5E6497D5402C" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="4DA9AB3E20BF0E2822F82A22BC6F3588" italics="true">Lethocolea</emphasis>
</taxonomicName>
) via large, disciform (sometimes angular-shaped) gemmae produced near the shoot apex on the dorsal stem surface or on the bases of the dorsal leaf margins, gemmae green to light brown,
<quantity id="FC53C0A40426D24287FAF61847E229E7" metricMagnitude="-3" metricUnit="m" metricValue="3.0" metricValueMax="5.0" metricValueMin="1.0" unit="mm" value="3.0" valueMax="5.0" valueMin="1.0">15 mm</quantity>
in diameter, 420 cells across, biconvex, 56 cells thick in the middle and 12 cells thick at the margin, sometimes with a 1 - celled stalk, with or without transparent, unistratose wing; gemmalings sprouting from the centre or the margin of the gemmae.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EEOBG" type="distribution">
<paragraph id="8A5C239137544B0E9E5CB3D76A0C228D">
<heading id="DFE655A37E663F52C0924D330FA3A883" reason="title">Distribution.</heading>
</paragraph>
<paragraph id="F045FC6AB6CAAC8A569CCE269773A8BD">
Widely distributed in Mediterranean and temperate regions of the Southern Hemisphere and in tropical mountains, extending northwards to
<collectingCountry id="A255057AAF71EFB8B098D1040E391CF6" name="Mexico">Mexico</collectingCountry>
,
<collectingCountry id="DB7D46CA78EBCD8C0F04EA91CD303626" name="India">India</collectingCountry>
, and
<collectingCountry id="C8B43732C09DBB4F104274777385A1F2" name="Japan">Japan</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EJOBG" type="habitat">
<paragraph id="024F19DB7EB1FC9FBAD03FE64FB96F7A">
<heading id="F9D4B504F78B5BE5DB7FF78C6E46B3E8" reason="title">Habitat.</heading>
</paragraph>
<paragraph id="18B89E5CC7BD37484093FED707F7E2DE">On sandy, loamy, or peaty soils subject to desiccation, in shaded and exposed sites, along trails, on earth walls and on soil over rock, sometimes in swampy, wet habitats, or floating on water, becoming deeply reddish-maroon. The plants sometimes grow partly buried in the substrate and may become brownish, somewhat purplish or carmine when growing in exposed sites.</paragraph>
</subSubSection>
<subSubSection id="SECID0EOOBG" type="notes">
<paragraph id="ED65567A95E29DF24357DC953D88A14D">
<heading id="21D854B47253F489B86801D2CEDE312C" reason="title">Notes.</heading>
</paragraph>
<paragraph id="4C3BB0AFCC96D5501B7E894CA7800B68">
<taxonomicName id="5068EA9E0EA761CE04C2A1CE3B758E75" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="E19B3EB7D650D35DFE9EBB200720FE50" italics="true">Lethocolea</emphasis>
</taxonomicName>
is readily recognized by: 1) creeping, little-branched leafy shoots with rather fragile stems made up of thin-walled cells, without or with one (or more) ventral stolons; 2) rhizoids scattered, hyaline to light brown to pink (exceptionally purplish); 3) leaves succubous, insertion line reaching dorsal midline of stem, alternate, imbricate, undivided, suborbicular to ovate-oblong to lingulate, with a broadly rounded apex, entire margins and an area of enlarged hyaline cells in the lower ventral half of the leaf; 4) leaf cells with or without trigones, trigones hyaline, cuticle coarsely papillose or smooth, oil bodies 13 per cell, large, finely granular, greyish-brown to dark brown; 5) underleaves absent; 6) gametoecia on main shoots; 7) sporophyte enclosed by a pendent marsupium at the shoot apex (perianth absent); 8) calyptra present, free, carried down the marsupial canal to the foot of the marsupium together with the unfertilized archegonia; 9) capsule cylindrical, tip acute to apiculate; 10) asexual reproduction by multicellular, disciform gemmae (in subgen.
<taxonomicName id="E00ECA18B92A70F82605FD27B6832480" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="324AF4EAA5B1C79ADCAD1BB35BBBE0DA" italics="true">Lethocolea</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="73843DAB0868D43A50B18B4538CF6B64">
By its undivided succubous leaves, lack of underleaves, and rather large, finely granular oil bodies, one or two per cell,
<taxonomicName id="C7D3184DFAA38CD43BEF4D34CEA0014E" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="35ACCFAE603E5B3C1A37851BFB9B6D42" italics="true">Lethocolea</emphasis>
</taxonomicName>
has been confused with
<taxonomicName id="EC9884B6A51991502D790F02FF7AA0DF" class="Jungermanniopsida" family="Jackiellaceae" genus="Jackiella" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="6AF4AC820A0D5AC9323D3D420B33C23D" italics="true">Jackiella</emphasis>
</taxonomicName>
(
<taxonomicName id="814F74B52715E2096DE24EAE0241A9DA" class="Jungermanniopsida" family="Jackiellaceae" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="family">Jackiellaceae</taxonomicName>
),
<taxonomicName id="301A1A5282B936F6067DC613F8F746F0" class="Jungermanniopsida" family="Cephaloziaceae" genus="Odontoschisma" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="E60068C2FEEA2000F6CD3DEDD61E0AE4" italics="true">Odontoschisma</emphasis>
</taxonomicName>
(
<taxonomicName id="350CB9E094111C6AEC655CA58EF1F308" class="Jungermanniopsida" family="Cephaloziaceae" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="family">Cephaloziaceae</taxonomicName>
), and
<taxonomicName id="CD1257D637510D77EDFD6A3B1D524299" class="Jungermanniopsida" family="Solenostomataceae" genus="Solenostoma" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="D93DE6925EF71DDFAB467AA17BD540D5" italics="true">Solenostoma</emphasis>
</taxonomicName>
(
<taxonomicName id="F462509566F2FF21B4732F3C336270F5" class="Jungermanniopsida" family="Solenostomataceae" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="family">Solenostomataceae</taxonomicName>
). The latter three genera are distinguished from
<taxonomicName id="5009EDC7849F02E1CA6ECA6BE43C3198" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="0A8A6A49BC3D5AC2976510E485155C04" italics="true">Lethocolea</emphasis>
</taxonomicName>
by the absence of an area of enlarged-hyaline cells in the lower ventral half of the leaf and by the smooth or finely papillose cuticle covered by minute papillae much smaller than in
<taxonomicName id="B1DDF8097102332F2BCE46B76441C279" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="BCA4FF49C37D26FE0BA2AF45D1424D54" italics="true">Lethocolea</emphasis>
</taxonomicName>
.
<taxonomicName id="6B36E73426C1691A5A2D7F8BB5B06117" class="Jungermanniopsida" family="Jackiellaceae" genus="Jackiella" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="7B0571461073357354CA629239F080F4" italics="true">Jackiella</emphasis>
</taxonomicName>
and
<taxonomicName id="C003118011B4AFC7C5A76656119E4A50" class="Jungermanniopsida" family="Cephaloziaceae" genus="Odontoschisma" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="7051189303B4B2CC354A32878182018B" italics="true">Odontoschisma</emphasis>
</taxonomicName>
, moreover, have dorsal leaf insertions usually not reaching the dorsal midline of stem, gametangia on short ventral branches, and asexual reproduction by small, 12 - celled gemmae. The sporophyte is enclosed by a perianth in
<taxonomicName id="4713FF3AB400827D75D9FD77B759B2ED" class="Jungermanniopsida" family="Cephaloziaceae" genus="Odontoschisma" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="D66FC18AAB5C235D14BF5CED7E27AE55" italics="true">Odontoschisma</emphasis>
</taxonomicName>
and
<taxonomicName id="F0F034E433D82C15DA3DEAA35F9E6966" class="Jungermanniopsida" family="Solenostomataceae" genus="Solenostoma" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="88903EC1027F2D9D1AF9B2F338E289A9" italics="true">Solenostoma</emphasis>
</taxonomicName>
, and by a marsupium in
<taxonomicName id="BF2677C8418924F01D762BA68AD5EF7E" class="Jungermanniopsida" family="Jackiellaceae" genus="Jackiella" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="5178FB6FCA9BEC201C06B2521F6E62C1" italics="true">Jackiella</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="FF8C5FA04F7CA0E9B9A139A2DEFCDE9E">
Based on the results of the morphological and molecular study, four
<taxonomicName id="EF725AA7CA065054DE7E77CD77F12A66">
<emphasis id="5D51FE85B14E14E85FED6CC1B9800885" italics="true">Lethocolea</emphasis>
species
</taxonomicName>
are recognized in two subgenera, subgen.
<taxonomicName id="4CB8A6B0A8DEDFF762825F4511CE644F" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="5FD67995F3CFA47FB3B61E3358FFA170" italics="true">Lethocolea</emphasis>
</taxonomicName>
and subgen.
<taxonomicName id="B0B037DB447339CCBD380322068B8E43" class="Jungermanniopsida" family="Acrobolbaceae" genus="Symphyomitra" higherTaxonomySource="CoL,GBIF" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="EDFAF6ABB58065B16D19F85F0E0E0689" italics="true">Symphyomitra</emphasis>
</taxonomicName>
. The two subgenera are distinguished by the presence or absence of disciform gemmae, the length and shape of the marsupium, and the incidence of papilliform cells lining the marsupial canal (
<bibRefCitation id="F1093AAD64FF73FF5A05DD9C30F055EF" author="Grolle" firstAuthor="Grolle" journalOrPublisher="Journal of the Hattori Botanical Laboratory" pagination="547-551" refId="B23" refString="Grolle R (1972) Miscellanea hepaticologica 121130. Journal of the Hattori Botanical Laboratory 36: 547551." title="Miscellanea hepaticologica 121-130." volume="36" year="1972">Grolle 1972</bibRefCitation>
). The two groups are fully supported in the molecular analysis (Fig.
<figureCitation id="E160E0BE826EB6E8F046A54A36085D71" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Maximum likelihood tree resulting from rps 4 sequences in the liverwort genus Lethocolea. Numbers below the branches correspond to bootstrap support." figureDoi="10.5091/plecevo.126936.figure2" httpUri="https://binary.pensoft.net/fig/1150547">2</figureCitation>
).
<bibRefCitation id="4E6A9CEDDC624E097FB97A771AEA4785" author="Schuster RM" firstAuthor="Schuster" journalOrPublisher="Nova Hedwigia Beiheft" pagination="1-721" refId="B48" refString="Schuster RM (2021) Austral Hepaticae III. Nova Hedwigia Beiheft 120: 1721." title="Austral Hepaticae III." volume="120" year="2021">Schuster (2021)</bibRefCitation>
treated the two subgenera as sections, but in view of the strong molecular support for the two groups we do not accept Schusters classification.
</paragraph>
<paragraph id="44FAC7E524317A661F145097C450B1DD">
Sporophytes are rare in
<taxonomicName id="950586368CE66A9666EC22534A80D34B" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="8F294620ADDF94181B6A1579D312E8CB" italics="true">Lethocolea</emphasis>
</taxonomicName>
; the above description is based on two sporophytic collections of
<taxonomicName id="2855BD81494BB909DD84C8FB841C0069" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="congesta">
<emphasis id="C9CF94DB0F0188950FF552E37C8A88AA" italics="true">L. congesta</emphasis>
</taxonomicName>
, with additions from
<bibRefCitation id="0D7B105876F4FF63C99690E628E7D1D3" author="Spruce R" firstAuthor="Spruce" journalOrPublisher="Transactions and Proceedings of the Botanical Society of Edinburgh" pagination="309-588" refId="B54" refString="Spruce R (1885) Hepaticae amazonicae et andinae. II. Transactions and Proceedings of the Botanical Society of Edinburgh 15: 309588." title="Hepaticae amazonicae et andinae. II." volume="15" year="1885">Spruce (1885)</bibRefCitation>
and
<bibRefCitation id="77EA1FFEE117013164702201CA96AC4A" author="Schuster RM" firstAuthor="Schuster" journalOrPublisher="Nova Hedwigia Beiheft" pagination="1-721" refId="B48" refString="Schuster RM (2021) Austral Hepaticae III. Nova Hedwigia Beiheft 120: 1721." title="Austral Hepaticae III." volume="120" year="2021">Schuster (2021)</bibRefCitation>
. The seta of
<taxonomicName id="BFDE240D03AD2EEA1D591985D87210B1" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="849ACC203145ACFC6D01BE3A3DB23662" italics="true">Lethocolea</emphasis>
</taxonomicName>
is massive and the capsule is cylindrical with an apiculate tip (Fig.
<figureCitation id="D7966DFB85DB7EC95620C57022D69AFF" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 C, D</figureCitation>
), like in other members of
<taxonomicName id="AC8F9FF3CFB55C019D82157AAE7D4660" authority="(Schuster 2021)" baseAuthorityName="Schuster" baseAuthorityYear="2021" class="Jungermanniopsida" family="Acrobolbaceae" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="family">
Acrobolbaceae (
<bibRefCitation id="6537AD773D28711EE5FEA222F4B50B90" author="Schuster" firstAuthor="Schuster" journalOrPublisher="Nova Hedwigia Beiheft" pagination="1-721" refId="B48" refString="Schuster RM (2021) Austral Hepaticae III. Nova Hedwigia Beiheft 120: 1721." title="Austral Hepaticae III." volume="120" year="2021">Schuster 2021</bibRefCitation>
)
</taxonomicName>
. The capsule of
<taxonomicName id="719BBA4681E57A9F2CA4DCB60BAEC918" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="congesta">
<emphasis id="8EEAAEE291671AF1664E6C837ECFDC3E" italics="true">L. congesta</emphasis>
</taxonomicName>
is unusual, however, in opening to the base into 24 valves (with two adjacent valves sometimes remaining partially connate) and in having a thin, mostly 2 - stratose wall, with limited 34 - stratose areas (Fig.
<figureCitation id="CE8CB31630DCAEE091B2DC55FF75A027" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 F</figureCitation>
) and no thickenings on the walls of the inner cell layers (Fig.
<figureCitation id="7A6136B2A67B12E5C7423C3E45419277" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 G, I</figureCitation>
) (capsule opening to the base into 4 valves, wall 410 - stratose and with thickenings in all cell layers in other
<taxonomicName id="22DD25C0DB896D9A5C950CB84F6D4154" baseAuthorityName="Schuster" baseAuthorityYear="2021" class="Jungermanniopsida" family="Acrobolbaceae" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="family">Acrobolbaceae</taxonomicName>
). Moreover, the elaters of
<taxonomicName id="DE700F8C97FD2FFC114DB86E3FC20E2F" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="species" species="congesta">
<emphasis id="76A981B197F5D79EF18A68C0BF60C6A3" italics="true">L. congesta</emphasis>
</taxonomicName>
possess 12 spirals (Fig.
<figureCitation id="B0B83DA2D1D0B6A2E56AC7C7F031B95E" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Lethocolea congesta. A, E. Female shoot with near-mature marsupium. B. Elater. C. Shoot apex with mature marsupium, female bracts and mature sporophyte before dehiscence, showing apiculate capsule. D. Cross section of seta. F. Cross sections of capsule. G. Outer cells of capsule wall, frontal view. H. Outer cells of capsule wall, lateral view. I. Inner cells of capsule wall, frontal view. J. Spores. K. Male bract with antheridium. L. Male plant with androecia. M. Cross section of stem. A from Onraedt 70. R. 4539; B J from Halle 139; K L from Larrain 45507; M from Holz 209. Drawn by Anna Luiza Ilkiu-Borges." figureDoi="10.5091/plecevo.126936.figure9" httpUri="https://binary.pensoft.net/fig/1150554">9 B</figureCitation>
) (elaters bispiral in
<taxonomicName id="B589497B7FD16E80E02DF9149673DA54" baseAuthorityName="Schuster" baseAuthorityYear="2021" class="Jungermanniopsida" family="Acrobolbaceae" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="family">Acrobolbaceae</taxonomicName>
). Data on capsule wall thickenings and elaters are still lacking for the other species of
<taxonomicName id="2C24054CFE089B8D9443C3A5AB893CB7" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="D284A335B52B68F100F89E2D87322084" italics="true">Lethocolea</emphasis>
</taxonomicName>
. The sporophyte of
<taxonomicName id="0EC1D0E9B9CAA5582EA861633FC59C58" class="Jungermanniopsida" family="Acrobolbaceae" genus="Lethocolea" kingdom="Plantae" order="Jungermanniales" phylum="Marchantiophyta" rank="genus">
<emphasis id="BA89812F936A4EC008D12204E1079F51" italics="true">Lethocolea</emphasis>
</taxonomicName>
clearly needs more study.
</paragraph>
</subSubSection>
</treatment>
</document>