identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
B9438794FF92FFF582850A88D8AEBA04.text	B9438794FF92FFF582850A88D8AEBA04.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Anthoceros subtilis Stephani (1916: 1003	<div><p>Anthoceros subtilis Stephani (1916: 1003). Fig. 2, 3A–B.</p> <p>Type:— INDIA. South Canara, Mangalore, August 1907, R. Pfleiderer 6127 (holotype G-G00060882).</p> <p>For synonyms see Hasegawa (1998), Peng &amp; Zhu (2013) and Villarreal et al. (2015).</p> <p>Monoicous. Plants growing on soil in rosettes. Thallus light to dark green, up to 2.6 mm in length and 3.5 mm in width. Dorsal lamellae absent or scarce. Margins of thallus highly crenulate. Thallus up to 22 cells thick in solid regions (106– 599 μm), cavernous, schizogenous cavities 75–150 × 115–155 μm, 1–3 layers of schizogenous cavities in transverse section. Single chloroplast per cell with central pyrenoid. Rhizoids hyaline to brown; unbranched. Ventral spheric Nostoc colonies abundant. Antheridia scattered. Antheridial chamber with up to 15 antheridia per cavity, antheridia yellowish to dark brown, 102–125 μm μm in length, with four-tiered jacket cell arrangement. Archegonia scattered, with five neck cells covered by mucilage. Involucres erect, cylindrical, up to 1.5 mm long, with smooth surface or with very few lamellae, 3–6 cells thick in solid regions (28–159 μm), cavernous, schizogenous cavities 15–30 × 22–47 μm, one layer of schizogenous cavities in transverse section.</p> <p>Up to three sporophytes per plant. Sporophyte foot with a layer of palisade-like cells. Capsules erect, up to 10 mm long, opening by two longitudinal valves. Epidermal layer of capsule one cell thick in transverse section. Epidermal cells of capsule rectangular, elongated, 7.5–19 × 37–85 μm and thick-walled, stomata abundant (17 per mm 2), stomata 24–40 × 42–58 μm. Assimilative layer 4–5 cells (26.4–95.8 μm) thick in transverse section. Columella present within the sporophyte, extending to the tip, rigid and brown, 16 cells in cross section. Sporogenous layer one spore tetrad thick. Pseudoelaters short, thin-walled, light brown to dark brown at maturity, 42.8–81.6 μm long (65.1±26.9, n=22), 3.9–33.4 μm in width, and 1–5 cells at maturity. Spores light brown to brown at maturity, 26.6–42.1 μm (34.9±3.5, n=31) in equatorial diameter. Distal spore surface with numerous papillate to tuberculate projections, up to 3 μm in height and 2.2 μm in width, sometimes confluent at their bases. Proximal spore surface with a distinct trilete mark, 0.89–1.5 μm in width, irregularly covered with verrucae; triangular areas covered with numerous papillate to tuberculate projections, up to 2.8 μm in height and 2.5 μm in width, sometimes confluent at their bases. The entire spore surface is covered with minute papillae, up to 0.8 μm in height and 0.4 μm in width.</p> <p>Habitat: —Growing on the sides of roads and around coffee plantations in tropical montane cloud forest, with other hornworts such as A. telaganus Stephani (1916:1005) and Phaeoceros sp., ca. 1300 m.</p> <p>Distribution:— China, Central African Republic, India, Japan, Korean Peninsula, Nepal, Thailand, Vietnam (Asthana &amp; Srivastava 1991, Hasegawa 1998, Peng &amp; Zhu 2013, Villarreal et al. 2015, Choi et al. 2021) and Mexico.</p> <p>Affinities and differentiation:—Mexican populations of A. subtilis are characterized by small, brown spores (26.6–42.1 μm), with papillate to tuberculate projections covering both proximal and distal surfaces. Within Mexican taxa, A. aff sambesianus Stephani (1916: 996) shares with A. subtilis the small sized plants (Ibarra-Morales et al. 2015), up to 7 mm in diameter in A. aff sambesianus and up to 2.6 mm in length and 3.5 mm in width in A. subtilis, but can be differentiated by spore ornamentation pattern. Anthoceros aff sambesianus possess a low reticulated network with foramens and central sub-globose tubercles in triangular areas of proximal face (Fig. 3 C-D), while A. subtilis presents irregular tuberculate projections and minute papillae (Fig. 3 A-B).</p> <p>According to Peng &amp; Zhu (2013), A. subtilis resembles A. angustus Stephani (1916: 1001) in spore morphology, nevertheless, they can be differentiated by the presence of marginal spongy gemmae on the thallus margins of A. angustus, absent in A. subtilis; and by the size of the pseudoelaters, which are considerably longer in A. angustus (40–496 µm) and somewhat shorter in A. subtilis (42–199 µm). Anthoceros angustus is restricted to Asia (Peng &amp; Zhu 2013).</p> <p>Specimens examined:— MEXICO. Chiapas: <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-92.08139&amp;materialsCitation.latitude=15.0675" title="Search Plazi for locations around (long -92.08139/lat 15.0675)">Unión Juarez</a>, roadside, 1438 m, 15°04’03” N 92°04’53” W, 26 July 2019, Ariadna Ibarra-Morales 2019 -14b (FCME).</p> <p>Remarks: —Mexican populations of this species were always found growing with other bryophytes. In one of the most crowded patches, some rosettes of A. subtilis were formed by very small plants of less than 1 mm in capsule height and thallus diameter. Some also exhibited an unusual erect morphology of the thallus, similar to the thallus morphology that has been reported in A. erectus Kashyap (1915: 9) (Asthana &amp; Srivastava 1991). In these dwarf plants, the surface of the thallus was barely large enough for the base of the involucre to fit. This dwarf morphotype was probably a result of space and resource availability and may have also been affected by a fungal infection, since red hyphae were found within the gametophyte of some of these plants.</p> <p>Previous reports of this species from Asia (Asthana &amp; Srivastava 1991, Hasegawa, 1998; Peng &amp; Zhu 2013) document slightly larger thallus size (5–6 mm in diameter) and larger spores (35–45 µm) than those of Mexican plants (thallus up to 3.5 mm and spores of 26.6–42.1 μm), but spore morphology is very similar to that illustrated by Peng &amp; Zhu (2013) from China.</p></div> 	https://treatment.plazi.org/id/B9438794FF92FFF582850A88D8AEBA04	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	Ibarra-Morales, Ariadna	Ibarra-Morales, Ariadna (2021): Anthoceros subtilis and A. telaganus (Anthocerotaceae): two new records for America from the hornwort flora of Mexico. Phytotaxa 529 (1): 93-104, DOI: 10.11646/phytotaxa.529.1.7
B9438794FF97FFF68285097ADA50BFE0.text	B9438794FF97FFF68285097ADA50BFE0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Anthoceros telaganus Stephani (1916: 1005	<div><p>Anthoceros telaganus Stephani. Fig. 4, 5.</p> <p>Type:— Indonesia, Java, I’rov. Preanger, Telaga bodas, Schiffner #19 (holotype G-G00061328, isotype NY-NY00231473).</p> <p>Monoicous. Plants growing on soil in rosettes. Thallus dark green, growing up to 15 mm in length and 15 mm in width. Dorsal lamellae are scarce to abundant. Margins of thallus lobulate or irregularly crenulate. Thallus up to 15 cells (368–845 μm) thick in solid regions, cavernous, schizogenous cavities 113–286 × 159–481 μm, 1–2 layers of schizogenous cavities in transverse section. One chloroplast per cell with a central pyrenoid. Rhizoids hyaline and unbranched. Ventral spheric Nostoc colonies are abundant. Antheridia scattered. Antheridial chamber with up to 10 antheridia per cavity, yellowish to brown, 59–106 μm in length, and with four-tiered jacket cell arrangement. Archegonia not observed. Involucres erect and cylindrical, up to 5 mm long, surface smooth to abundantly lamellate, 7–9 cells (154–262 μm) thick in solid regions, cavernous, schizogenous cavities 32–86 × 45–154 μm, 1 (–2) layer of schizogenous cavities in transverse section.</p> <p>Up to seven sporophytes per gametophyte. Sporophyte foot with a layer of palisade-like cells. Capsules erect, up to 30 mm long, opening by two longitudinal valves. Epidermal layer of capsule one cell thick in transverse section. Epidermal cells of capsule rectangular, elongated, 7–14 × 57–121 μm and thick-walled, stomata abundant (10 per mm 2), 34–46 × 44–64 μm.Assimilative layer 3–5 cells (32.89–114.4 μm) thick in transverse section. Columella present within sporophyte, extending to the tip, rigid and brownish to blackish, 16 cells in cross section. Sporogenous layer one spore tetrad thick. Pseudoelaters elongate, thin-walled, dark brown at maturity, 39.8–104.9 μm (73.2±16.8, n=19) long, 3.8– 26.7 μm in width, 1–2 cells at maturity. Spores black at maturity, 32.7–53.21 μm (40.48±3.8, n=44) in diameter. Distal spore surface has numerous spinose-lamellate projections, up to 3.7 μm in height, frequently irregularly confluent at their bases, often tipped by a papilla of 0.53–2.98 μm in diameter. Proximal spore surface has a distinct trilete mark, 0.9–3.66 μm in width, irregularly covered with verrucae, a smooth strip along either side of trilete mark, up to 3.98 μm in width (2.89 μm in average), triangular areas covered with numerous spinose-lamellate projections, up to 3.8 μm in height, frequently irregularly confluent at their bases, tipped by a papilla of 0.64–3.25 μm in diameter.</p> <p>Habitat:—Growing on the sides of roads and around coffee plantations in tropical montane cloud forest, with other hornworts such as A. subtilis and Phaeoceros sp., 1200–1300 m.</p> <p>Distribution:— Indonesia (Java and Sumatra) (Meijer 1957, Hasegawa 1998, Söderström et al. 2010) and Mexico.</p> <p>Affinities and differentiation:—Only one species known from Mexico, Anthoceros orizabensis (Stephani 1916: 965) Hässel de Menéndez (1990: 211) (SEM spore micrographs of this species can be found in Hässel de Menéndez (1990), plate 4, fig. 9 on page 212), resembles the spore ornamentation of A. telaganus, sharing the smooth strip along the trilete mark. Nevertheless, the smooth strip in A. orizabensis is incomplete, not reaching the equatorial girdle of the spores; moreover, the distal face of the spore has spines in A. orizabensis, while A. telaganus has a spinose-lamellate reticulated distal face.</p> <p>FIGURE 5. Scanning electron microscopy (SEM) micrographs of spores from Mexican populations of Anthoceros telaganus. A-B. From A. Ibarra-Morales 2019-11; C-D. From A. Ibarra-Morales 2019-20. A &amp; C distal surface, B &amp; D proximal surface.</p> <p>Other American species that share the smooth strip along the spore trilete mark of the proximal face are Anthoceros cavernosus Stephani (1916: 998), A. caucasicus Stephani (1923: 427) and A. fusiformis Austin (1875: 28) (Hässel de Menéndez 1990, Sérgio et al. 2020). Anthoceros fusiformis frequently presents a strap shaped thallus and lacks a pyrenoid in the chloroplasts (Hasegawa 1993), both characters differentiate it from A. telaganus. Regarding A. cavernosus and A. caucasicus, both species have spores with spinose outgrows reticulated at the base with ridges enclosing both lumina and pits on the distal face, and the proximal face with scattered spinose outgrowths, sometimes confluent at their bases (SEM spore micrographs of these species can be found in Hässel de Menéndez 1990 and Sérgio et al. 2020). In A. telaganus the spores present blunter papillate projections and an incomplete reticulum on the distal face, while the proximal face presents spinose-lamellate projections that are tipped by a papilla.</p> <p>In the location here reported, A. subtilis can be differentiated from A. telaganus by spore ornamentation, mature spore color and size, light brown to brown in the first species and black in the second, and smaller spores in A. subtilis. Plants not bearing sporophytes are difficult to differentiate. Thalli of Anthoceros subtilis are smaller and paler in color than those of A. telaganus. Anthoceros telaganus also presents an abundantly lamellate thallus morphotype that could be easily identified in this location.</p> <p>Specimens examined:— MEXICO. Chiapas: Unión Juarez, roadside, 1260 m, 15°03’35”N 92°04’44”W, 24 July 2019, Ariadna Ibarra-Morales 2019 -11 (FCME); 1438 m, 15°04’03”N 92°04’53”W, 26 July 2019, Ariadna Ibarra-Morales 2019-20 (FCME).</p> <p>Remarks: —Within the mature region of the sporophyte, colorless and shrunken (probably aborted) spores were observed amongst black mature regular-size spores in all plants examined of this species. The percentage of these inviable spores could not be estimated, but it was observed frequently that at least one spore of almost every tetrad displayed this condition and remain attached to a fully mature spore. This phenomenon was not observed in any other hornwort species growing in the same locations.</p> <p>Morphological variation within Mexican plants of this species was observed mainly in two characters: dorsal lamellae on the thallus and spore ornamentation. Dorsal lamellae on the thallus presented a variation range from nearly absent to abundant and plants expressing different ranges of this character could be found in the same population, but generally not close together in the same rosette. Regarding spore ornamentation, spores of northern populations (A. Ibarra -Morales #2019-20) were, in most of the individuals examined, more spinose and forming a more reticulated pattern (Fig. 5C–D) when compared to the ornamentation of southern populations (A. Ibarra -Morales #2019-11) (Fig. 5A–B); nevertheless, both spore morphologies were present in all collections.</p> <p>It is noteworthy that the type (Stephani 1916) and the collections here described, present similar ecological preferences, both being collected at or near coffee plantations. Meijer (1957) also states that this species grows on cultivated areas at 700–2000 m, and on earth walls and roadsides.</p></div> 	https://treatment.plazi.org/id/B9438794FF97FFF68285097ADA50BFE0	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	Ibarra-Morales, Ariadna	Ibarra-Morales, Ariadna (2021): Anthoceros subtilis and A. telaganus (Anthocerotaceae): two new records for America from the hornwort flora of Mexico. Phytotaxa 529 (1): 93-104, DOI: 10.11646/phytotaxa.529.1.7
