Chusquea anelythra , Nees 1835
Chusquea anelythra Nees (1835: 491). = Dendragrostis anelytra Nees ex Munro (1868: 63), nom. inval. Type (lectotype, designated here) :— BRAZIL. No locality or date (fl.), F . Sellow s.n., K! (H.965.88-7, fragm. ex B). Isolectotypes: BAA! (BAA00001764, fragm. ex B), BR! (BR6865504, fragm. ex B), E!...
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Zenodo
2021
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Online Access: | https://dx.doi.org/10.5281/zenodo.5817269 https://zenodo.org/record/5817269 |
Summary: | Chusquea anelythra Nees (1835: 491). = Dendragrostis anelytra Nees ex Munro (1868: 63), nom. inval. Type (lectotype, designated here) :— BRAZIL. No locality or date (fl.), F . Sellow s.n., K! (H.965.88-7, fragm. ex B). Isolectotypes: BAA! (BAA00001764, fragm. ex B), BR! (BR6865504, fragm. ex B), E! (E00373678, fragm. ex B), G! (G00099343, fragm. ex B), K! (H.965.88-8 and H.965.88-9, fragm. ex B), L! (catalogue number 908.86-1058, barcode L1233849, fragm. ex B), LE! (LE 01072717, fragm. ex. B), and P! (P00625465, fragm. ex B). Figs. 1, 2, and 3. Lectotypification comments : —In the protologue of Chusquea anelythra , Nees (1835) only cited “ In Brasilia. Sellow. Herb. Reg. Ber. ”, without any indication of a collection number for the Sellow specimen. However, as part of the Catalogue of New World Grasses, Clark (2000) indicated that the specimen “ F. Sellow 1248 ” was the holotype of this species, supposedly housed at B, in addition to four isotypes, two of them housed at LE (0556.01 and 3191.01) and the other two at US (catalogue numbers 2874614 and 557551, barcodes 00036597 and 00036596, respectively). According to Dr. L. Orlova, curator of the Herbarium of Higher Plants at LE (pers. comm.), there is a unique specimen currently deposited at LE (barcode 01072717) under the name Chusquea anelythra . Although we confirm that this specimen is a fragment from the type-specimen of C. anelythra , we are not sure about the correspondence between the current LE specimen and those two numbers cited by Clark (2000) from the LE collection. Thus, we have decided to consider only this recent specimen from LE cited here as an isolectotype of C. anelythra . We also analyzed the other putative type specimen housed at US (catalogue number 1021552, barcode 00009928). The printed label of this specimen shows that it originally was sent from the Berlin herbarium to the herbarium at the Bureau of Science in Manila, P. I., from which it ended up at US. The blue paper label with “Brasilia” and “Reliquiae d. Sello” (but no collection number) printed on it also has a Bureau of Science stamp on it. The specimen itself has a paper collecting tag with “1248” written on it, which was inferred by Clark (2000) to represent Sellow’s collection number. We confirm that this specimen represents C . anelythra and suggest that it may also correspond to duplicates of the original collection of C. anelythra . Moreover, as no collection number was mentioned in the protologue of this species (Nees 1835), we cannot exclude the possibility that all specimens labeled as “ Sellow 1248 ” or labeled as “ Sellow ” with a paper tag carrying the number “1248” actually correspond to duplicates of the original material. However, even if this is correct, the duplicate at US at most would be an isotype, and certainly not the holotype. Several fragments of the type material of this species derived from the collections housed at B exist, as indicated on the labels of these sheets. Some of these materials are incomplete and only consist of leaves, for example those deposited at BAA and US (barcode 00036597). According to Dr. R. Vogt, curator of the collection of the herbarium B (pers. comm.), neither the specimen “ Sellow ” nor the specimen “ Sellow 1248 ” are extant at this herbarium. Many specimens housed at B were destroyed during World War II (Merrill 1943), including the type collections of different plant families, Poaceae among them, so this is almost certainly what happened to the previously mentioned samples of C. anelythra . Given this, it is also impossible to verify if the material “ Sellow 1248 ” is part of the type collection of C. anelythra, so we exclude them from the proposed lectotypification. As stated by the International Code of Nomenclature for algae, fungi, and plants (ICN) (Turland et al. 2018), when the holotype or previously designated lectotype has been destroyed or lost, and there are duplicates of the original material, a substitute lectotype should be designated (Turland et al. 2018, Arts. 9.3 and 9.11). This is the case for Chusquea anelythra , in which the original specimen(s) collected by “ Sellow ” and designated by Nees (1835) as the type was (or were) destroyed, thus requiring the designation of a lectotype for this name. We found nine duplicates of the original material of this species (“ Sellow ”) housed at the herbaria BAA, BR, E, G, K, L, LE, and P, all ex B. We here designate one of the specimens housed at K (H.965.88-7, fragm. ex B) (Fig. 1) as the lectotype of Chusquea anelythra. Emended description : —Rhizomes unknown. Culms 3‒6 m tall, 0.6‒1.3 cm in diameter, more or less erect at the base then scandent or clambering towards the apex; internodes (11.2‒) 13.5‒16.1 (‒20.5) cm long, terete, solid, vinaceous or green, sometimes mottled, scabrous with a white waxy band (1‒2 cm long) just below the node, becoming glabrous on the upper half, otherwise completely glabrous. Culm leaves (7.9‒) 9.1‒10.7 cm long, not reaching the next node, the juncture of the sheath and blade abaxially a faint line or obscure; sheaths (6.2‒) 7.2‒8.4 cm long, from 3‒4 (‒4.7) times as long as the blade, persistent, margins ciliate on both or only one side, the overlapping one fused to the sheath at the base for 1.9‒2.2 cm, abaxially pubescent to hispid at the base with a concentration of trichomes in the central bud region, then scabrous towards the apex, adaxially glabrous, tessellate, waxy, shiny; summit extension absent; girdles 2.5‒5 mm, with a prominent skirt, brownish to dark brown, abaxially pubescent to hispid with spaced trichomes; outer ligule absent; inner ligule 1‒1.5 mm long, glabrous, erect, rigid or sometimes slightly membranous, white or yellowish, apex truncate and ciliate; blades 1.7‒2.5 cm long, broadly triangular, non-pseudopetiolate, erect, persistent, abaxially scabrous and densely waxy, adaxially glabrous at the base, becoming pubescent on the upper half then pilose at the apex, tessellate, the midrib not evident, the margins membranous and ciliate on both sides, rarely glabrous, the apex long setose, 1‒2.5 mm long. Nodes at mid-culm with a circular central bud subtended by 8‒10 smaller, subequal subsidiary buds in 1 row, in a constellate arrangement; central bud prophyll short, pilose with margins long ciliate; nodal line dipping slightly below the bud/branch complement; nodal region 5‒6.5 (‒8) mm long; supranodal ridge visible, slightly raised and prominent; nodal line raised and prominent. Branching infra-extravaginal, being initially infravaginal, often becoming also extravaginal as branches develop, leafy subsidiary branches 5‒11 (‒16) per node, 23‒50 cm long, often rebranching, geniculate at the first node, nodes prominent giving a knobby appearance. Foliage leaves 6‒13 (‒18) per complement; sheaths 2.8‒5.2 (‒6.3) cm long, scabrous to hispid on the upper half, straminous to brownish with green or vinaceous spots, keeled on the upper half, the margins long ciliate on only one side, summit extension asymmetrical, hispid to hirsute, on the overlapping margin (0.2‒) 0.45‒1.2 mm long, on the underlapping one 1.4‒3 (‒4.2) mm long and fused to the inner ligule; outer ligule asymmetrical, pubescent or scabrous, erect and rigid, apically ciliolate to long ciliate (trichomes 0.5‒4 mm long), the smaller side 0.3‒0.9 (‒1.3) mm long, truncate or lightly concave, the bigger one (0.7‒) 0.9‒1.9 mm long, lobed; inner ligule asymmetrical, membranous, pubescent to scabrous, ciliolate at the apex, toward the overlapping margin 0.6‒1.6 mm long and free from the summit extension, on the underlapping one 2‒3.4 (‒4.3) mm long and fused to the summit extension; pseudopetioles 1‒4.9 mm long, thick, distinct, abaxially pubescent to scabrous with a tuft of trichomes at the base that extends into the blade, adaxially hispid-pilose to scabrous with deposition of wax, pulvinus dark brown; blades (6.5‒) 7‒18.3 (‒20) cm long, 0.7‒1.3 cm wide, L: W 8‒19.4 (‒23), lanceolate to linear-lanceolate, non-tessellate, abaxially slightly scabrous with a tuft of trichomes at the base, adaxially pilose-scabrous at the base, becoming glabrous towards the apex, otherwise scabrous, base subcordate to asymmetrical, one side attenuate and the other one subcordate to rounded-truncate, midrib slightly excentric, abaxially prominent throughout the blade, except at the apex, adaxially prominent only on the lower half of the blade, both margins scabrous or one scabrous and the other smooth, apex acute to long setose, 0.95‒3.8 (‒6.1) mm long. Synflorescences (7.0‒) 9.4‒15.2 cm long, paniculate, open, pyramidal, with the lower branches reflexed, subtended by 2 spatheate bracts, the first (lower) one with the sheath 6.1‒9.4 cm long, papyraceous, somewhat overlapping, blade 2‒4.2 (‒7.6) cm long, brownish green, the second (upper) one with the sheath 8.1‒12.9 cm long, slightly membranous, expanded, blade 2.5‒6.7 (‒16) cm long, stramineous, rarely green; rachis triquetrous, densely pilose; branches and pedicels angular, densely pilose, all subtended by a scar or rim or occasionally a scale-like subtending bract 0.8‒1.3 mm long, the primary branches pulvinate, 1‒5 lowermost branches erect when young and divergent to reflexed at maturity, the lowermost one 5.8‒8.3 cm long, the second 7‒8.7 cm long, the third 5.9‒7.8 cm long, the fourth 4.4‒5 (‒7.1) cm long, and the fifth 3.2‒3.6 (‒5.9) cm long, the sixth and higher order branches and pedicels erect to appressed to the primary branches; pedicels 1‒4.5 mm long. Spikelets 6.6‒9 mm long, 1.2‒1.6 mm wide, globose to slightly dorsally compressed; glumes I and II scale-like, obtuse; glume I 0.1‒0.3 mm long, <1/30 the spikelet length, slightly pubescent at the base, becoming glabrous towards the apex; glume II 0.2‒0.45 mm long, <1/25 the spikelet length, glabrous; glumes III and IV navicular for the upper half, abaxially glabrous at the base then pubescent to pilose towards the apex, 3-nerved; glume III (3.6‒) 4.2‒5.6 (‒6) mm long, ca. 2/3 the spikelet length, awned (the awn 0.3‒1.3 mm long); glume IV (4.5‒) 5‒7 mm long, 3/4 to 4/5 the spikelet length, awned [the awn 0.2‒0.7 (‒1.2) mm long]; lemma (6‒) 6.5‒7.6 (‒8), somewhat navicular, apex setose, awned (the awn 0.3‒1.2 mm long), abaxially glabrous at the base, becoming pubescent with a tuft of trichomes at the apex, 7- nerved; palea 6.1‒7.7 mm long, overlapping the lemma, 2-apiculate (the apicule 0.1‒0.5 mm long), 4-nerved, 2-keeled, strongly sulcate only for the upper half, the sulcus scabrid, abaxially pubescent then pilose towards the apex. Lodicules 3, obovate to broadly obovate, glabrous, hyaline, apically long ciliate; anterior pair 1‒1.8 mm long, the posterior one 0.9‒1.4 mm long. Stamens 3; anthers 2.5‒4.3 mm long, brown. Caryopsis not seen. Taxonomic comments: —Within the Chusquea meyeriana informal group, C. anelythra , C. anelytroides , and C. attenuata share some vegetative characters (Table 1), such as culm leaves not reaching the next node, sheath and blade not very well-differentiated, and the blade short and deltate to broadly triangular, in addition to foliage leaves with the outer ligule asymmetrical (U-shaped or less usually asymmetrical in C. attenuata ) (Figs. 2 and 3). Reproductively, these species can be recognized by the presence of 2‒4 spatheate bracts subtending the synflorescences, glumes III and IV not reaching the full spikelet length, and the palea 2-apiculate (2-awned or less usually 2-apiculate in C. attenuata ) (Table 1, Fig. 2). ......continued on the next page These three species are similar during the initial stage of their development, which can lead to mistakes in the identification of specimens with juvenile culms. As the culms and branching develop, the three species become easier to differentiate, producing a set of morphological characters that can be useful to recognize them. Both Chusquea anelythra and C. anelytroides develop larger culms compared to C. attenuata , which is smaller and more delicate (Table 1). Despite these morphological similarities, C. anelythra can be differentiated vegetatively from C. anelytroides by the presence of the circular central bud ( vs. somewhat triangular in C. anelytroides ), culm leaf sheath shorter [(6.2‒) 7.2‒8.4 cm vs. 9.5‒10 cm long], foliage leaves with narrower blades (0.7‒1.3 cm vs. 1.4‒2.9 cm wide), and the inner ligule asymmetrical and reduced [<4.3 mm long on both sides vs. attenuate and 7‒20 (‒30) mm long], as well as the sheath summit extension that is (0.2‒) 0.45‒1.2 mm long on the overlapping margin and 1.4‒3 (‒4.2) mm long on the underlapping one ( vs. 2.0‒4.6 and 3‒7 mm, respectively) (Table 1; Figs. 2 and 3). Reproductively, C. anelythra differs from C. anelytroides by the presence of two spatheate bracts subtending the synflorescences ( vs. four), spikelets 6.6‒9 mm long ( vs. 9.5‒12 mm), relatively longer glumes III and IV, reaching ca. 2/3 and 3/4‒4/5 the spikelet length, respectively ( vs. ca. 1/3 and ca. 1/2), and shorter lemmas [(6‒) 6.5‒7.6 (‒8) mm long vs. 8‒10 mm] and paleas (6.1‒7.7 mm long vs. 8.4‒10 mm) (Table 1). Both C. anelythra and C. attenuata exhibit similarities in the length of their spikelets and included structures (such as glumes, lemma and palea) (Table 1), besides having a circular central bud, which is also shared with C. meyeriana (Clark 2001, Vidal et al. unpubl. data). Although infravaginal branching is reported as characteristic for all species of the Chusqua meyeriana informal group (Figs. 2 and 3), it is not very well described in the literature (Clark 2004, Fisher et al. 2014, Vidal et al. 2018, Andrade et al. 2019). Thus, after our extensive examination of all species of the C. meyeriana group, we here propose to describe this branching pattern in C. anelythra as infra-extravaginal, in which the branches emerge initially infravaginally, but often also become extravaginal as they develop. McClure & Smith (1967) gave the distribution of Chusquea anelythra as extending from Rio de Janeiro to Santa Catarina, based only on specimens collected in the latter state. We analyzed all specimens cited by these authors ( Reitz & Klein 6131 , 6453 , 6830, 7280 and 10462 ) and conclude that they do not in fact represent this species, although they do belong to the C . meyeriana informal group. Reitz & Klein 6830 and 7280 represent C. meyeriana or something closely related to it. The other three collections match other specimens from the state of Paraná, and although this entity resembles C . anelythra in many respects, it differs in key aspects of foliage leaf morphology and synflorescence structure and appears to be closer to C . wilkesii Munro (1868: 63). We therefore exclude all of these specimens from our concept of C . anelythra , and confirm that this species does not occur in southern Brazil. Judziewicz et al. (1999) considered this species as restricted to southeastern Brazil, occurring in the states of Espírito Santo, Minas Gerais, Rio de Janeiro, and São Paulo, although Clark (2001) did not confirm the occurrence of this species in São Paulo. Clark et al. (2020) also did not exclude the possibility that C. anelythra could occur in the states of São Paulo, Paraná, and Santa Catarina. Etymology: —According to McClure & Smith (1967), the specific epithet “ anelythra ” has a Greek origin and means an = “without” and elytra = “coverage or scale”, referring to the very reduced lower pair of glumes of this species. Habitat and distribution : — Chusquea anelythra is endemic to Brazil, occurring in montane forests and Atlantic forest fragments of Rio de Janeiro, Minas Gerais, Espírito Santo, and Bahia, along rivers and streams at 860‒1450 m.a.s.l. (Table 1, Figs. 3 and 4). During field trips by our research group, we recently found a large population of Chusquea anelythra ( Andrade et al. 180, 258, 269 ) at the Pedra Lascada ridge, in the municipality of Coaraci in southern Bahia, as indicated by Andrade (2019). We also found another collection of this species from Bahia housed at HUEFS, collected in the Peito-de-Moça ridge in the municipality of Arataca ( Amorim et al. 5367 ). These collections represent the first record of this species in Bahia. Conservation status : —The estimated area of occupancy (AOO) of Chusquea anelythra was 80 km ², whereas the extent of occurrence (EOO) was 158,204 km ². Following the IUCN Red List categories and criteria (IUCN Standards and Petitions Committee 2019), this species should be considered Vulnerable (VU) [B2ab(iii)], as it is restricted to montane forests and Atlantic forest fragments, with an AOO estimated to be less than 2,000 km ², and a number of locations equal to 10. Phenology: —During our herbarium survey, we identified only seven flowering collections of Chusquea anelythra : four of them from Rio de Janeiro, in November 1890 ( Glaziou 18634 ), November 1986 ( Martinelli et al. 11954 and 11959 ), October 1987 ( Martinelli et al. 12253 ), and May 1989 (spikelets very mature to disarticulating, Martinelli et al. 13435 ); only one from Minas Gerais in March 2002 (spikelets very mature to disarticulating, Santos-Gonçalves & Élcio 458 ); and two from Espírito Santo in December 2017 ( Pianissola 295 ) and January 2019 ( Pianissola 209 ). All specimens analyzed from Bahia were in the vegetative stage, but vegetative characters were enough to confirm the identification. The information available does not allow us to make accurate inferences about the flowering cycle of Chusquea anelythra , so we cannot discard the possibility of simply sporadic or asynchronous flowering events, as indicated for other Brazilian Chusquea species such as C. fruticosa Pianissola, Santos-Gonçalves & Clark in Pianissola et al. (2018: 239‒241), C. gouveiensis Vidal & Clark in Vidal et al. (2018: 78-83), C. imbricata Pianissola, Clark & Santos-Gonçalves in Pianissola et al. (2018: 241‒245), and C. tenella (Attigala et al. 2017, Pianissola et al. 2018, Vidal et al. 2018). However, if these were all gregarious flowering events, a minimum 15- to 16-year cycle, with flowering extending for up to three years, may be inferred. Additional specimens examined : — BRAZIL. No locality, no date, (fl.), Sellow 1248 (US). Bahia: Mun. Arataca, Serra do Peito-de-Moça, complexo Serra das Lontras, estrada que liga Arataca a Una, ramal ca. 22,4 km de Arataca com entrada no assentamento Santo Antônio, RPPN “Caminho das Pedras”, trilha de acesso ao Peito-de-Moça, após residência do Senhor “Mormaço”, 15°15’48”S, 39&de : Published as part of Vidal, Kaio Vinicius De A., Dorneles Welker, Cassiano A., Silva, Alaísma Pereira Da, Santos-Gonçalves, Ana Paula, Clark, Lynn G. & Oliveira, Reyjane P., 2021, Revisiting the circumscription of Chusquea anelythra (Poaceae-Bambusoideae- Bambuseae): lectotypification, redescription, geographic distribution, and conservation status, pp. 71-85 in Phytotaxa 529 (1) on pages 73-82, DOI: 10.11646/phytotaxa.529.1.5, http://zenodo.org/record/5814320 : {"references": ["Nees, C. G. D. (1835) Bambuseae brasilienses. Linnaea 9: 461 - 494.", "Munro, W. (1868) A monograph of the Bambusaceae, including descriptions of all the species. 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