identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
B281A1F3C85D593A9770E2E91C0D6D7B.text	B281A1F3C85D593A9770E2E91C0D6D7B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Frullania chiapasensis Mamontov, K. Feldberg, Schaf. - Verw., Gradst. 2025	<div><p>Frullania chiapasensis Mamontov, K. Feldberg, Schäf.-Verw., Gradst. sp. nov.</p><p>Holotype.</p><p>Geoscientific Collection of the University of Göttingen, Germany (GZG), GZG.BST.22085; syninclusion  Parmotrema specimen 4.</p><p>Etymology.</p><p>The species is named after the location of the amber deposit in Chiapas, Mexico.</p><p>Age and stratigraphic level.</p><p>15‒23 Ma, Langhian – Aquitanian (early to middle Miocene), Simojovel, Chiapas, Mexico.</p><p>Diagnosis.</p><p>Incubously foliated liverwort with conduplicate-trilobed, entire-margined leaves; dorsal lobe obliquely ovate to elliptical, apex rounded to obtuse, lacking ocelli; ventral lobule  Frullania - type, saccate, adnate to lobe at a distance of ca. 0.8 of the stem width, parallel or somewhat converging to stem and at places leaning against it, clavate to obovate, not constricted above the postical opening, surface smooth, opening extending along the abaxial side of the lobe for ca. 0.32–0.50 of the lobule length, sickle-shaped; underleaves triangular to obovate, longer than wide, widest in the upper half, bilobed 0.2–0.3 × their length, upper half of underleaves armed with 2 short teeth on both sides.</p><p>Description.</p><p>Two gametophyte fragments. Intact shoot ca. 2.64 mm long, 0.33–0.35 mm wide with leaves, dark reddish brown (Figs 1 A, B, 2); with three broken, lateral branches [not clearly visible]. Stem only slightly darker in color than leaves, 35–45 µm in diameter, surface cells not visible. Lateral leaves incubous, alternate, contiguous to imbricate near apex, widely spreading, conduplicate-trilobed (Figs 1 A – C, 2 A). Dorsal lobe in dorsal aspect convex to nearly flat, partially with reflexed margin, obliquely ovate to elliptical (Figs 1 A – D, 2 A), 150–200 µm long × 120–150 µm wide, slightly longer than wide, length: width ratio ca. 1.2–1.3: 1; margin entire, apex rounded to obtuse, inner margin barely extending beyond the farther edge of the stem; lobe base not clearly visible. Lobe cells almost isodiametric or rectangular to 5–6 - angled (Fig. 1 D, F), 15–20 × 10–20 µm in the middle of the lobe, marginal cells somewhat smaller; cell walls slightly thickened, with indistinct trigones, no intermediate thickenings seen; with one central mammilla, ca. 5 µm in diameter. Ocelli not observed. Ventral lobules  Frullania - type, inflated, saccate, explanate lobules not observed; mostly + / - parallel with the stem, isolated lobules obliquely spreading at an angle of ca. 35 ° to the stem, or leaning against it; remotely inserted, distance between anterior base of lobule and edge of stem ca. 0.8 × the stem width; lobules clavate or obovate (Figs 1 C – E, 2), 110–140 µm long × 75–90 µm wide at the widest part, longer than wide, length: width ratio 1.5–1.6: 1; apex broadly rounded or angulate; gradually applanate towards the postical opening, not constricted above it, the portion adjoining the opening dorsiventrally compressed in comparison to the gibbous upper half, pinched together; opening very wide, extending along the abaxial side of the lobe for ca. 0.32–0.50 × the lobule length, sickle-shaped (Fig. 2), the apex of the opening with a discolored, gibbous, protuberant cell; the lobules almost symmetric, or somewhat asymmetric due to the sickle-shaped opening, widest in upper third, rarely near the middle; the valves (free margins of the opening) entire, parallel, equal in size; the lobule surface smooth, the margins of the opening crenulate-sinuate, especially near the apex (i. e., the upper end of the sinus), due to protuberant angles between the pairs of cells and strongly concave external walls of these cells. Styli indistinctly visible, triangular (Figs 1 D, E, 2), with expanded base from several cells and at least 6 cells high, with a short apex of 2 cells. Underleaves 1 per leaf pair, distant, transversely inserted, ca. 2.1–2.5 × the stem width, mostly longer than wide (Figs 1 C, E, 2), 110–140 µm long × 100–120 µm wide, triangular to obovate, flat, gradually broadening from a cuneate base; lower two thirds entire-margined, upper third armed on both sides with 1–2 short teeth of 1–3 cells; bilobed 0.2–0.3 × their length, sinus acute-angled, the sinus base broadly rounded; lobes widely triangular, obliquely directed to the stem axis, wider than long, with acute to obtuse apex, inner margins of lobes entire, slightly arcuate. Rhizoids possibly in fascicles arising from the upper part of the underleaves (Fig. 2 B). Asexual reproduction not observed. Sterile.</p><p>Discussion.</p><p>The entire conduplicate-trilobed leaves with an entire-margined dorsal lobe and  Frullania - type lobules (Figs 1 A – C, 2 A) as well as the bilobed underleaves (Figs 1 C, E, 2) allow for a reliable generic assignment.  Frullania is a large genus of porellalean leafy liverworts with a center of diversity in tropical and subtropical regions, where it often grows epiphytic in the canopy, but it is also a common element of temperate and arctic zones and can grow under much drier conditions than  Lejeuneaceae (e. g., Gradstein et al. 2001).</p><p>Frullania is the most diverse genus found in amber. Four species occur in Cretaceous Kachin amber (Feldberg et al. 2021 a, 2021 b), and 16 species have been described from Cenozoic European and African ambers (Bouju et al. 2021; Feldberg et al. 2021 a). To date, only three  Frullania fossils are known from New World deposits: a small fragment described at subgenus level from Dominican amber (Heinrichs and Schmidt 2010) and two fossils from Mexican amber, including  Frullania sp. (Juárez-Martínez et al. 2023) and the recently described  F. delgadilloi Juárez-Mart. &amp; Estrada-Ruiz, the latter representing the most completely preserved specimen (Juárez-Martínez et al. 2024). The Dominican fossil differs from  F. chiapasensis in possessing obliquely spreading lobules, small, acuminate styli, and more deeply bilobed, wider than long underleaves (Heinrichs and Schmidt 2010, figs 1–5).  Frullania sp. from Mexican amber is differentiated by its lobules which are not much longer than wide, constricted above the only slightly arched opening, and much smaller in relation to the lobes (Juárez-Martínez et al. 2023, fig. 3). Moreover, the leaf lobules in this specimen have a crook-shaped mouth similar to those in the subgenus  Meteoriopsis Spruce. By contrast, in  F. chiapasensis the lobule mouth is sickle-shaped and resembles those of numerous species in the subgenera Diastaloba s. l. and  Microfrullania (R. M. Schust.) R. M. Schust. (see also below). The underleaves of  Frullania sp. are not well visible and cannot be compared.  Frullania delgadilloi differs from  F. chiapasensis in lobule and underleaf shape. The lobules of  F. delgadilloi are significantly smaller in relation to the lobes and strongly oblique with the apices oriented outwards, resembling the Dominican  Frullania sp. in this aspect. Furthermore, the lobules of  F. delgadilloi are inserted further from the stem (ca. 1–2 times the stem width), while those of  F. chiapasensis are larger in relation to the lobes, less remote (ca. 0.8 times the stem width), and oriented mostly parallel to the stem. The lobes of  F. delgadilloi possess scattered larger cells, interpreted by the authors as ocelli, which seem to be absent from  F. chiapasensis . The underleaves in  F. delgadilloi are ovate, four or more times wider than the stem (according to our measurements of the shoots imaged in the description of  F. delgadilloi), bilobed to ca. 0.3 of their length, with the lobes narrowly acute and angular along the outer margins (Juárez-Martínez et al. 2024, fig. 3 A, F). In contrast, the underleaves of  F. chiapasensis are triangular to obovate, less than three times wider than the stem, and bilobed to 0.2–0.3 of their length, with 1–2 short but distinct teeth at the base of the obtuse to acute lobes. However, the species also resemble each other in several ways:  F. chiapasensis is characterized by a sickle-shaped lobule mouth that extends to ca. 0.32–0.50 of the lobule length and has crenulate margins as well as a protuberant cell at the apex. Juárez-Martínez et al. (2024) describe the opening as “ asymmetrical slightly crenulate with incision reaching about ca. 0.25 of lobule length ”; a protuberant cell is not mentioned. Furthermore, both species have large, foliose styli. We consider especially the differences in lobule shape and orientation to be significant enough to describe a second  Frullania species for Mexican Chiapas amber, though both species might be related at subgenus or section level.</p><p>Other liverwort inclusions similar to  F. chiapasensis are  F. baltica Grolle from Baltic and Bitterfeld amber and  F. schmalhausenii Mamontov et al. from Rovno amber. However, these species are not only older than  F. chiapasensis but can easily be differentiated based on their lobules, which are obliquely spreading, with the apices oriented away from the stem like in  F. delgadilloi, instead of lobules oriented mostly parallel to the stem. They can also be distinguished by their low abaxial lobule opening, which ends at less than 0.1 of the lobule length, while the lobe opening of  F. chiapasensis is strongly extended along the lateral lobe margin and hence sickle-shaped (Figs 1, 2; Grolle and Meister 2004, plate 5; Mamontov et al. 2019, figs 2, 3 b, c, g, j, k).</p><p>Because Mexican amber is a rather young deposit and several extant species have been found in Dominican amber (Gradstein 1993; Kubilius et al. 2017; Feldberg et al. 2021 a), it is very important to compare the new fossil to the extant diversity. As one of the largest and taxonomically most complex genera of leafy liverworts,  Frullania contains more than 2,000 published names (von Konrat et al. 2010; Hentschel et al. 2015) and 675 currently accepted taxa (Söderström et al. 2016; Schäfer-Verwimp and Winter 2023). Many extant  Frullania species and subspecies are insufficiently known, and information about their morphology is only available in protologues published in the 19 th and early 20 th centuries. Recent studies have greatly expanded the knowledge of the complex taxonomy of the genus, and it seems that its diversity is even greater than anticipated, as new species are found regularly (e. g., Hentschel et al. 2009, 2015; Heinrichs et al. 2010; Ramaiya et al. 2010; von Konrat et al. 2010, 2012, 2013; Carter et al. 2017; Silva et al. 2017; Mamontov et al. 2020; Schäfer-Verwimp and Winter 2020, 2022, 2023). Consequently, the assessment of morphological differences between  F. chiapasensis and the extant species of the genus is difficult. Moreover, the morphological homoplasies within  Frullania challenge the subgeneric assignment of numerous extant non-sequenced species, and even more so that of fossils. Based on the combination of the characteristics of leaf lobes, lobules, and underleaves,  F. chiapasensis is similar to species of several subgenera: namely  Caulisequa, Diastaloba I, Diastaloba II, Diastaloba III,  Frullania, Mammillosae,  Meteoriopsis, and  Microfrullania . To compare them with  F. chiapasensis, we have analyzed the information (protologues, descriptions, and figures) for all species of the mentioned subgenera, which are accepted in Söderström et al. (2016). Some members of these subgenera are well distinguished from  F. chiapasensis by the presence of moniliate ocelli in leaf lobes or by the shape of the underleaves (if the underleaves are entire at apex or appendiculate at base, for example). However, many taxa can be distinguished from the newly described species only with difficulty. Therefore, we compare extant species similar to  F. chiapasensis in a morphology matrix based on 5 + 11 critical characters (see material &amp; methods and Appendix 1). Most species of subgenera  Frullania and  Meteoriopsis were excluded from consideration based on characters 1–5, while  F. chiapasensis + 144 species meeting these criteria have been chosen to map their approximate differences and similarities based on 11 additional characters.</p><p>Based on the character comparison, the extant species morphologically most similar to  F. chiapasensis is  F. simmondsii Steph., as described in Hattori (1979, fig. 42). According to Appendix 1, the species coincide with each other in all 11 characters. However, based on the description and illustration in Hattori (1979), the stylus of  F. simmondsii is similar to that in numerous species of subg. Diastaloba s. l. by leaving a narrow foliose strip extending along the keel, and thus being wider than long. In contrast, the stylus of  F. chiapasensis resembles the foliaceous styli found in numerous species of subg. Trachycolea by being longer than wide. Summarizing all differences between both species,  F. chiapasensis can be distinguished from  F. simmondsii as follows: (1) leaf lobes longer than wide in  F. chiapasensis vs. wider than long in  F. simmondsii; (2) stylus foliaceous, longer than wide vs. filiform, when flat narrowly foliaceous, wider than long; (3) stem underleaves mostly longer than wide, narrowly obcuneate at base, with the margins concave in the lower two-thirds vs. stem underleaves mostly wider than long, widely obcuneate at base, with the margins largely convex in lower two-thirds.</p><p>Besides  F. simmondsii, we have identified eight more species, each differing from  F. chiapasensis in one of the listed 11 characters (Appendix 1). The species are  F. colliculosa von Konrat et al.,  F. eplicata Steph.,  F. gabonensis Vanden Berghen,  F. hodgsoniae von Konrat et al.,  F. multilaceroides S. Hatt.,  F. scalaris S. Hatt.,  F. subtilissima (=  F. taxodiocola and  F. exilis Taylor) and  F. vaga Mitt. Among them, five species ( F. colliculosa,  F. gabonensis,  F. hodgsoniae,  F. multilaceroides,  F. subtilissima) differ from  F. chiapasensis in the lobule arrangement. The lobules in the mentioned five species are obliquely patent, spreading from the stem at angles of 25–55 ° or more, whereas in  F. chiapasensis the lobules are slightly patent or largely parallel to the stem. Moreover, in  F. colliculosa,  F. gabonensis,  F. hodgsoniae, and  F. multilaceroides the underleaves of leading stems are usually slightly broader than long and broadest in the middle, whereas in  F. chiapasensis the underleaves are mostly longer than wide and broadest at the upper third. In the studied extant specimens of  F. subtilissima from Costa-Rica (Schäfer-Verwimp &amp; Holz SV / H- 0486 / A, Schäfer-Verwimp &amp; Holz SV / H- 0429 / Z, MHA), the width and length of leaf lobes exceed the length of leaf lobules ca. 2 and 2.8 times, respectively, thus the leaf lobes are ca. 8–10 times larger than lobules, while the lobe apex is often apiculate and the margins of underleaves entire or barely angulate. In contrast, in  F. chiapasensis the length and width of leaf lobes exceeds the length of leaf lobules ca. 1.4 and 1.6 times, respectively, thus the leaf lobes are only ca. 2–3 times larger than lobules, while the lobe apex is obtuse to rounded, and the margins of underleaves are armed with 1–2 teeth of 1–3 cells on both sides. Among the remaining three species,  F. eplicata differs from  F. chiapasensis by the low position of the lobule mouth apex that is similar to species of subgenera  Frullania,  Meteoriopsis, and Trachycolea. Moreover, well-developed underleaves in  F. eplicata are up to 3 times broader than the stem and wider than long (Vanden Bergen 1976, fig. 21 A). The second species,  F. scalaris, is very similar to  F. chiapasensis in the shape and size of leaf lobes and lobules. However, the leaf lobules in the former species are apically overlapping the stem, while the underleaves are described and illustrated as only slightly broader than the stem, entire margined, broadest in the middle, almost bilobed to 0.5 of their length, with triangular-lanceolate lobes (Hattori 1977, fig. 15). In contrast, in  F. chiapasensis the leaf lobules are mostly parallel to the stem, while the underleaves are 2.1–2.5 times broader than stem, toothed in the upper third, bilobed to 0.2–0.3 of their length, with widely triangular lobes. The last species,  F. vaga, is described to have only shallowly bilobed, entire margined underleaves (Mitten 1865), while other distinctions are not evident from the protologue.</p><p>The remaining 135 species in Appendix 1, apart from the 9 species discussed above, can be distinguished from  F. chiapasensis by two to seven characters. Therefore, the latter taxon can be considered as a separate species which essentially differs from all the described extant and fossil  Frullania species. The subgeneric assignment of  F. chiapasensis remains problematic because the species morphologically most similar to it are treated now as belonging to different subgenera, namely Diastaloba I and  Microfrullania (Söderström et al. 2016).</p></div>	https://treatment.plazi.org/id/B281A1F3C85D593A9770E2E91C0D6D7B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Feldberg, Kathrin;Kaasalainen, Ulla;Mamontov, Yuriy S.;Gradstein, S. Robbert;Schäfer-Verwimp, Alfons;Divakar, Pradeep K.;Schmidt, Alexander R.	Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K., Schmidt, Alexander R. (2025): Extending the fossil record of Miocene neotropical epiphyte communities. Fossil Record 28 (1): 79-102, DOI: 10.3897/fr.28.137758
BA36D971E7925D309496C443A93304A0.text	BA36D971E7925D309496C443A93304A0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parmotrema undetermined-1	<div><p>Parmotrema specimen 1</p><p>Specimen.</p><p>Stuttgart State Museum of Natural History, Germany (SMNS), SMNS -DO-4928-M; syninclusion unidentified leafy liverwort of the order  Porellales .</p><p>Age and stratigraphic level.</p><p>15‒20 Ma, Langhian – Burdigalian (early to middle Miocene), La Toca Formation, Dominican Republic.</p><p>Description.</p><p>Lichen fragment approximately 5.4 × 2.6 mm in diameter. Thallus foliose, lobate. Lobes robust, flat, and linear, 0.4–0.9 mm wide, lobe apices truncate (Fig. 7 B). Upper surface slightly uneven, brown, with prominent dark margins (Fig. 7 C). Medulla not visible. Lower cortex dark. Marginal cilia long (up to at least 0.8 mm), dark, thick and tapered (Fig. 7 B, C). Rhizines dark, shorter than cilia. Isidia abundant, laminal, finger-shaped, up to 150 µm long (Fig. 7 C). Apothecia, soredia, or pycnidia not present.</p><p>Discussion.</p><p>The general habit and long, thick, and tapered marginal cilia identify the specimen as  Parmotrema .  Parmotrema is an extant genus with approximately 300 species mainly distributed in tropical and subtropical regions, especially in the Pacific Islands and South America (Thell et al. 2012). It is part of the parmelioid crown group of the  Parmeliaceae (Pizarro et al. 2018) where apothecial and conidial characters, growth form, cortical and medullar chemistry, and presence, absence and / or type of cell-wall polysaccharides, marginal cilia, rhizines, and surface features, like epicortex and pseudocyphellae, have been used to separate genera (Crespo et al. 2011). Like in many parmelioid genera, molecular phylogenetics have repeatedly shaped the generic boundaries of  Parmotrema (e. g., Blanco et al. 2005; Divakar et al. 2005, 2017; Crespo et al. 2011). As the species level taxonomy of  Parmotrema mainly relies on chemical and other characters not observable in the fossil inclusions, comparisons of fossils to extant species are often ineffectual. However, the genus currently includes several infrageneric groups that, at some point, have been acknowledged as separate genera based on morphology, anatomy, and chemistry but which were found to be nested within  Parmotrema in molecular phylogenetic analyses (Elix 1993; Blanco et al. 2005; Divakar et al. 2005, 2017; Crespo et al. 2011).  Parmotrema specimen 1 with the smooth upper surface, dark lower surface, and very long, branched marginal cilia resembles the  Parmotrema s. str. group within the genus  Parmotrema . The group is, for example, characterized by broad lobes, broad naked marginal zone on the lower surface, marginal cilia, and the lack of maculae on the upper surface (Elix 1993). The extant  Parmotrema s. str. group contains more than 250 species distributed especially in the tropical regions of the world (Elix 1993).</p></div>	https://treatment.plazi.org/id/BA36D971E7925D309496C443A93304A0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Feldberg, Kathrin;Kaasalainen, Ulla;Mamontov, Yuriy S.;Gradstein, S. Robbert;Schäfer-Verwimp, Alfons;Divakar, Pradeep K.;Schmidt, Alexander R.	Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K., Schmidt, Alexander R. (2025): Extending the fossil record of Miocene neotropical epiphyte communities. Fossil Record 28 (1): 79-102, DOI: 10.3897/fr.28.137758
CB108EEC9EB15CA981CFD92C76327C05.text	CB108EEC9EB15CA981CFD92C76327C05.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parmotrema undetermined-2	<div><p>Parmotrema specimen 2</p><p>Specimen.</p><p>Stuttgart State Museum of Natural History, Germany (SMNS), SMNS -DO-4929-M.</p><p>Age and stratigraphic level.</p><p>15‒20 Ma, Langhian – Burdigalian (early to middle Miocene), La Toca Formation, Dominican Republic.</p><p>Description.</p><p>Three lichen fragments situated very closely together, most probably originating from the same thallus (Fig. 7 E); whole inclusion approximately 5.0 × 2.4 mm in diameter. Thallus foliose, lobate. Lobes flat, broad to linear, 0.4–1.0 mm wide, lobe apices truncate to rounded (Fig. 7 E). Upper surface smooth and brown, with dark margins (Fig. 7 D). Marginal cilia dark, branched, and long (up to 0.8 mm) (Fig. 7 E). Medulla not visible. Lower cortex dark. Rhizines dark. Pycnidia abundant on the upper surface (Fig. 7 D). Apothecia, isidia, or soredia not present.</p><p>Discussion.</p><p>The general habit, very long, branched marginal cilia, and dark lower surface identify the specimen as  Parmotrema . Of the infrageneric groups, the dark lower surface without rhizines reaching the margins could represent, for example, the  Parmotrema s. str. group.</p></div>	https://treatment.plazi.org/id/CB108EEC9EB15CA981CFD92C76327C05	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Feldberg, Kathrin;Kaasalainen, Ulla;Mamontov, Yuriy S.;Gradstein, S. Robbert;Schäfer-Verwimp, Alfons;Divakar, Pradeep K.;Schmidt, Alexander R.	Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K., Schmidt, Alexander R. (2025): Extending the fossil record of Miocene neotropical epiphyte communities. Fossil Record 28 (1): 79-102, DOI: 10.3897/fr.28.137758
FB66CEF003875E48827155D844A39F21.text	FB66CEF003875E48827155D844A39F21.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parmotrema undetermined-3	<div><p>Parmotrema specimen 3</p><p>Specimen.</p><p>American Museum of Natural History, New York, AMNH DR-14-294.</p><p>Age and stratigraphic level.</p><p>15‒20 Ma, Langhian – Burdigalian (early to middle Miocene), La Toca Formation, Dominican Republic.</p><p>Description.</p><p>Lichen fragment approximately 1.7 × 1.3 mm in diameter. Thallus foliose, lobate. Lobes flat, linear, approximately 0.5 mm wide and 40 µm thick, lobe apices truncate to rounded. Upper surface slightly uneven with effigurate maculae, with prominent dark margins (Fig. 8 A). Marginal cilia long (up to 0.9 mm), branching, tapering (Fig. 8 A). Medulla pale (Fig. 8 C). Lower cortex dark (Fig. 8 B, C). Rhizines dark, mainly simple (Fig. 8 B). Apothecia, isidia, soredia, or pycnidia not present.</p><p>Discussion.</p><p>The general habit and long, branching and tapering marginal cilia identify the specimen as  Parmotrema . The effigurate maculae on the upper surface and tapered marginal cilia of  Parmotrema specimen 3 resemble the former genus  Canomaculina (Elix 1993) which has been synonymized with  Parmotrema (Blanco et al. 2005) . The  Canomaculina group is characterized by tapered cilia, effigurate-maculate upper surface of the lobes, and simple and furcate to branched rhizines (Elix 1993). The  Canomaculina group contains more than ten extant species and has a center of distribution in South America (Elix 1993; Blanco et al. 2005).</p></div>	https://treatment.plazi.org/id/FB66CEF003875E48827155D844A39F21	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Feldberg, Kathrin;Kaasalainen, Ulla;Mamontov, Yuriy S.;Gradstein, S. Robbert;Schäfer-Verwimp, Alfons;Divakar, Pradeep K.;Schmidt, Alexander R.	Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K., Schmidt, Alexander R. (2025): Extending the fossil record of Miocene neotropical epiphyte communities. Fossil Record 28 (1): 79-102, DOI: 10.3897/fr.28.137758
6716A1229A05503BB070BBC997A034A5.text	6716A1229A05503BB070BBC997A034A5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parmotrema undetermined-4	<div><p>Parmotrema specimen 4</p><p>Specimen.</p><p>Geoscientific Collection of the University of Göttingen, Germany (GZG), GZG.BST.22085; syninclusion  Frullania chiapasensis .</p><p>Age and stratigraphic level.</p><p>15‒23 Ma, Langhian – Aquitanian (early to middle Miocene), Simojovel, Chiapas, Mexico.</p><p>Description.</p><p>Lichen fragment approximately 2.3 × 4.5 mm in diameter. Thallus foliose, lobate. Lobes flat and linear, 0.5-1.0 mm wide, lobe apices truncate to slightly rounded (Fig. 8 D). Upper surface smooth, brown (Fig. 8 D). Marginal cilia dark, mainly simple, up to 0.5 mm long (Fig. 8 D). Medulla not visible. Lower cortex concolorous with the upper surface (Fig. 8 E). Rhizines dark, abundantly branched (Fig. 8 E). Apothecia, isidia, soredia, or pycnidia not present.</p><p>Discussion.</p><p>The general habit and long, tapering marginal cilia identify the specimen as  Parmotrema . The branching rhizines and long, tapered marginal cilia resemble the  Canomaculina group. However, unlike the  Canomaculina group, the fossil inclusion seems to possess a smooth upper surface of the lobes that visibly lack maculae.</p></div>	https://treatment.plazi.org/id/6716A1229A05503BB070BBC997A034A5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Feldberg, Kathrin;Kaasalainen, Ulla;Mamontov, Yuriy S.;Gradstein, S. Robbert;Schäfer-Verwimp, Alfons;Divakar, Pradeep K.;Schmidt, Alexander R.	Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K., Schmidt, Alexander R. (2025): Extending the fossil record of Miocene neotropical epiphyte communities. Fossil Record 28 (1): 79-102, DOI: 10.3897/fr.28.137758
B8AF380D312455309F95494016EBD03E.text	B8AF380D312455309F95494016EBD03E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Thysananthus patrickmuelleri K. Feldberg, Gradst., Schaf. - Verw., Mamontov 2025	<div><p>Thysananthus patrickmuelleri K. Feldberg, Gradst., Schäf.-Verw., Mamontov sp. nov.</p><p>Holotype.</p><p>Geoscientific Collection of the University of Göttingen, Germany (GZG), GZG.BST.22086.</p><p>Additional specimens examined.</p><p>Patrick Müller Amber Collection (Zweibrücken, Germany) MEX 70, MEX 71.</p><p>Etymology.</p><p>The specific epithet honors the amber collector Patrick Müller who has generously supported our research by providing numerous amber fossils for study, including all known specimens of this species.</p><p>Age and stratigraphic level.</p><p>15‒23 Ma, Langhian – Aquitanian (early to middle Miocene), Simojovel, Chiapas, Mexico.</p><p>Diagnosis.</p><p>Incubously foliated liverwort with a ventral merophyte at least five cells wide; flagelliform branches present on lower parts of the shoots; lateral leaves conduplicate-bilobed; dorsal lobe convex, asymmetrically oblong-ovate with a rounded to obtuse apex, median lobe cells mostly elongate with distinctly cordate trigones; ventral lobule  Lejeunea - type, ovate-rectangular with one hooked apical tooth and a truncate apex; underleaves suborbicular to rounded quadrate to obovate with a rounded to emarginate, recurved apex and entire, mostly revolute margins.</p><p>Description.</p><p>Gametophyte fragments yellowish to dark reddish brown, 2.7–5.04 mm long, main shoots 0.52–1.16 mm wide with leaves (Figs 4 A – C, 5 A – C, 6 A – E). Branching irregular,  Lejeunea - type, primary branches either similar to main shoot, or flagelliform and microphyllous (Figs 4 B, D, 6 C, D); main shoot-like branches 1.26–4.8 mm long, 0.41–1.4 mm wide with leaves; flagelliform branches 0.2–0.24 mm wide; one secondary microphyllous branch becoming main shoot-like on upper part. Stem dark reddish brown to blackish, 40–70 µm in diameter; cortical cells possibly thick-walled and elongated [mostly obscured by underleaves], ventral merophyte ca. 5 cells wide; stem on large branches 40–60 µm in diameter, on flagelliform branches ca. 30 µm in diameter. Lateral leaves incubous, densely imbricate, alternate, widely spreading to erect-spreading or erect-appressed (especially on upper shoot parts), conduplicate-bilobed with large dorsal lobe and a  Lejeunea - type ventral lobule folded against the lobe (Figs 4 C, E, 5 C, 6 E), insertion line J-shaped. Dorsal lobe convex, asymmetrically oblong-ovate, on upper shoot parts falcate, 480–770 µm long × 220–300 µm wide in the middle, length: width ratio 1.4–3.5: 1, margin entire, postical margin nearly plane or slightly incurved for up to 0.7 × the lobe length, exterior to keel first weakly then abruptly arched towards apex, apex rounded to obtuse, antical margin regularly arched, lobes overlapping stem 0.5–2 × the stem width beyond the farther edge of the stem, dorsal base not visible; lobes on larger branches very similar to main shoot, short ovate to oblong-ovate, slightly falcate to straight, 230–840 µm long × 160–430 µm wide in the middle, length: width ratio 1.4–2.1: 1; lobes on flagelliform branches erect-spreading to erect, 150–160 µm long × 70–110 µm wide in the middle, length: width ratio 1.5–2.1: 1. Lobe cells rectangular to hexagonal (Figs 4 F, 5 D, E, 6 E), elongated at the lobe base and in the middle, slightly shorter to (sub) isodiametric near apex, up to 3 × as long as wide [often collapsed and deformed], basal cells 20–35 µm long × 7.5–20 µm wide, median cells 15–35 µm long × 7.5–17.5 µm wide, apical cells 7.5–20 µm long × 7.5–15 µm wide; cell walls thin, trigones cordate, not coalesced, few intermediate thickenings present. Ventral lobule ovate-rectangular, lower part inflated, free antical margin not inflexed (Figs 4 E, 5 C, D, 6 E), 0.3–0.4 × the length of the lobe, 200–310 µm long × 80–200 µm wide in the middle, length: width ratio 1.6–2.9: 1, free antical margin nearly straight to slightly arched, free lateral margin truncate and mostly forming an angle of ca. 90 ° with the postical lobe margin, apex with a single, often hooked tooth (Figs 4 E, 5 D), 10–15 µm (1–2 cells) long × 10–15 µm (1–2 cells) wide at base, hyaline papilla not seen, keel curved, leaves slightly emarginate at end of keel; no appendages at the base of the lobule or the keel seen; lobules on larger branches 0.3–0.5 × as long as lobes, 110–310 µm long × 70–160 µm wide in the middle, length: width ratio 1.3–2.4: 1; lobules on flagelliform branches 0.5–0.7 × as long as lobe, 90–110 µm long × 60–80 µm wide in the middle, length: width ratio 1.3–1.8: 1. Underleaves imbricate to contiguous, symmetrical, suborbicular to rounded quadrate to obovate, generally wider than long to as long as wide (Figs 4 C, E, 5 C, 6 E), 3–6 × wider than the stem, 150–310 µm long × 160–390 µm wide, length: width ratio 0.6–1: 1; slightly squarrose, concave with recurved margins to occasionally nearly plane, apex recurved, rounded to truncate to emarginate, margins entire, insertion line arched, base auriculate, not adnate to lateral leaf; underleaves on larger branches more often nearly plane, on lower parts slightly longer than wide, 160–280 µm long × 130–360 µm wide, length: width ratio 0.7–1.2: 1, apex rounded, often not recurved; underleaves on flagelliform branches not clearly visible, possibly ovate. Rhizoids in bundles near underleaf insertion, bundles up to 230 µm long, rhizoids ca. 10 µm in diameter with truncate and ampliate tips. Sterile.</p><p>Discussion.</p><p>The conduplicate-bilobed leaves with  Lejeunea - type lobules and the presence of underleaves clearly identify this specimen as a member of  Lejeuneaceae (Figs 4 C, E, 5 A – C, 6 E). The robust size, the undivided underleaves, and at least five cells wide ventral merophyte allow an assignment to subfamily  Ptychanthoideae (Figs 4 A, C, 5 A – C, 6 A, B, E). Although all specimens are sterile, the ovate, apically rounded dorsal lobes, the elongate median lobe cells with distinct cordate trigones, as well as the entire-margined underleaves, the probably thick-walled stem cells, and the presence of flagelliform branches on lower shoot parts are indicative of the genus  Thysananthus (Figs 4, 5, 6 A – E).</p><p>Thysananthus is a large genus with 30 extant species and several fossil taxa (Gradstein 1993; Sukkharak and Gradstein 2014, 2017; Sukkharak 2015; Wang et al. 2016; Feldberg et al. 2021 a). It includes the former genus  Mastigolejeunea which was ranked at subgenus level based on molecular phylogenetic analyses and morphology (Sukkharak and Gradstein 2017). The oldest fossil of the genus is  T. contortus (Göpp. &amp; Berendt) Sukkharak &amp; Gradst. from Paleogene Baltic and Bitterfeld amber. This species has predominantly  Frullania - type branches, which become as vigorous as the main axis, and rarely  Lejeunea - type branches, as well as four lobule teeth (Grolle and Meister 2004, plates 14, 15, as  Mastigolejeunea contorta (Göpp. &amp; Berendt) Gradst. &amp; Grolle; Sukkharak and Gradstein 2017). More fossil species of  Thysananthus have been found in tropical Miocene ambers. The genus is known from Dominican amber with the extant species  Thysananthus auriculatus (Wilson &amp; Hook.) Sukkharak &amp; Gradst. (Gradstein 1993, as  Mastigolejeunea auriculata (Wilson &amp; Hook.) Steph.; Sukkharak and Gradstein 2017) as well as the extinct species  T. bidentulus (Gradst.) Sukkharak &amp; Gradst. (Gradstein 1993, as  Mastigolejeunea bidentula Gradst.; Sukkharak and Gradstein 2017) and  T. weiweianus N. - N. Yu &amp; Gradst. (Yu et al. 2020).  Thysananthus auriculatus can be differentiated from  T. patrickmuelleri by the less elongate leaf lobes, the more strongly incurved postical leaf margin, the oblique apical free margin of the lobule which continues into the postical margin of the lobe, and the either small or absent apical lobule tooth (Gradstein 1993, fig. 7; Sukkharak and Gradstein 2014, figs 4, 5).  Thysananthus bidentulus is more similar to  T. patrickmuelleri in its general habit, but differs in having oblong lobules with oblique apices, which continue into the postical margin of the lobe, and the presence of two one-celled lobule teeth (Gradstein 1993, fig. 8).  Thysananthus weiweianus differs from the new fossil species in having larger, often confluent trigones, a very small or absent lobule tooth, and more elongate underleaves with plane lateral margins (Yu et al. 2020, figs 1, 2). Another amber deposit that includes  Thysananthus is Miocene Ethiopian Shewa amber (Bouju et al. 2021, fig. 3).  Thysananthus aethiopicus Bouju et al. differs from the new Mexican fossil in having lobes with more strongly incurved postical margins and occasionally apiculate apices, as well as lobules with a very coarse apical tooth and sometimes a second tooth.</p><p>A specimen similar to  T. patrickmuelleri in its general habit has already been found in Mexican amber but was only identified as belonging to  Ptychanthoideae and possibly to  Mastigolejeunea (Fig. 6 F; Scheben et al. 2014). Many important diagnostic characters are not visible in this specimen, especially the walls of the lobe cells, which are crucial for correctly identifying the genus, and the structure of the lobules.</p><p>Compared to the extant species of  Thysananthus, the new fossil is very similar to the neotropical  T. plicatiflorus (Spruce) Sukkharak &amp; Gradst., a species widely distributed in the rainforests of northern South America (Gradstein 1994, 2021; Sukkharak and Gradstein 2014, fig. 21, as  Mastigolejeunea plicatiflora (Spruce) Steph.). Both species share elongate, ovate-oblong leaves with rounded apices and a plane postical margin, lobules with a single short tooth and a truncate apical free margin that terminates at the end of the keel, as well as auriculate underleaves. The two species differ markedly, however, in the orientation of the underleaves, which are very flat and have a plane apex in  T. plicatiflorus while being somewhat squarrose with a distinctly recurved apex in  T. patrickmuelleri (apex only occasionally plane on branch leaves). Another quite similar species is the Australasian  T. ligulatus (Lehm. &amp; Lindenb.) Sukkharak &amp; Gradst. (Sukkharak and Gradstein 2014, figs 18, 19, as  Mastigolejeunea ligulata (Lehm. &amp; Lindenb.) Schiffn.), which also has rather elongated lobes with rounded apices, plane margins, and auriculate underleaves. However, it can be differentiated by the lobules being smaller in relation to the lobes (0.2–0.3 times the lobe length vs. 0.3–0.4) and underleaves that are often longer than wide, spathulate, and not overlapping the stem as much as those of  T. patrickmuelleri (3–4 times wider than the stem vs. 3–6). Since the new fossil shows significant morphological differences to all known extinct and extant species of  Thysananthus, it is described as a new species.</p></div>	https://treatment.plazi.org/id/B8AF380D312455309F95494016EBD03E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Feldberg, Kathrin;Kaasalainen, Ulla;Mamontov, Yuriy S.;Gradstein, S. Robbert;Schäfer-Verwimp, Alfons;Divakar, Pradeep K.;Schmidt, Alexander R.	Feldberg, Kathrin, Kaasalainen, Ulla, Mamontov, Yuriy S., Gradstein, S. Robbert, Schäfer-Verwimp, Alfons, Divakar, Pradeep K., Schmidt, Alexander R. (2025): Extending the fossil record of Miocene neotropical epiphyte communities. Fossil Record 28 (1): 79-102, DOI: 10.3897/fr.28.137758
