Braunia plicata (Mitt.) Jaeger,

Luna, Efraín De, 2022, Lectotypification, taxonomy and distribution of Braunia plicata (Bryopsida, Hedwigiaceae) in South America and Mexico, Phytotaxa 532 (1), pp. 101-106 : 102-106

Braunia plicata (Mitt.) Jaeger, Ber. St. Gall. Naturw. Ges. 1874-75: 171. (Ad. 2: 87). 1876. ( Figs. 1–4 , 15–25).

Basionym: Hedwigia plicata Mitt., J. Linn. Soc., Bot. 12: 406-407. 1869. Protologue. “Andes Quitensis, Jameson. In Bolivia, Bridges in herb. Hooker”.

Lectotype (designated here):— BOLIVIA. 1846, Bridges s. n. (Lectotype: BM! (herb. Hooker-BM) [bc.BM000960770!]; isolectotypes, BM! (herb. Wilson), NY (2)! [bc.NY00913932, bc.NY00913953]).

Braunia plicata var. canescens Card., Rev. Bryol. 38: 38. 1911. Protologue. “Etat de Hidalgo: Cuyamaloya, rochers (Pringle, 1908, nº 10627). Etat de Mexico: Toluca (Pringle, 1908, nº 15171)”.

Lectotype (designated here):— MEXICO. Toluca, Aug 4, 1908, Pringle 15171 (Lectotype P (herb. Cardot) [bc.PC0701219]; isolectotypes, NY! [bc.NY00792519]; S!, FH!) [proparte, the specimen at FH contains mostly plants of B. andrieuxi ]; MEXICO. Hidalgo, Cuyamaloya, Pringle 10627 (BM (2)! [bc. BM000960771, bc.BM000960772], E (2) [bc.E00108810, bc.E00246841], H! [bc. H3301459], JE (2)! [bc.JE04000337, bc.JE04000338], MO! [bc.MO-407560], NY! [bc.NY00792521], P(2) [bc.PC0147304, bc.PC0701217], S!, VT [bc.UVMVT031831], US!).

Nomenclatural note 1. The selection of a specimen collected by Bridges from Bolivia in 1846 as lectotype for B. plicata conforms strictly to the species concept in the original description ( Mitten 1869). The original description of Hedwigia plicata in Mitten (1869) reads: “Folia erecto-patentia, imbricata, latissime ovalia, concava, cuspide brevi interdum albescente terminata, quater plicata, margine late reflexo apice serrulato, … “. These leaf features constitute the core of the species concept of B. plicata , as recognized here. Only the plants in Bridges specimens display leaves widely ovate, leaf acumen hyaline, and leaf margin at apex serrulate to dentate ( Fig. 1 ). I have seen four specimens of the Bridges collection. There are two duplicates at BM (Bridges s. n. BM-herb. Hooker; Bridges s. n. BM-herb. Wilson) and two duplicates at NY (Bridges s. n. NY-herb. Mitten). These two specimens at NY consist of only few stems glued on a card (bc.NY00913932, bc.NY00913953). The specimen at BM-herb Hooker (bc.BM000960770) is selected here as lectotype ( Fig. 1 ), because the type citation makes clear reference to a particular specimen as follows: “In Bolivia, Bridges in herb. Hooker”.

Nomenclatural note 2. Mitten (1869) cited jointly in the protologue of B. plicata a specimen from Ecuador as “Andes quitenses, Jameson ”. This syntype is difficult to discern among several specimens collected by Jameson, due to incomplete collection data. Two duplicates at NY-herb. Mitten with herbarium labels “Andes Quitensis, Jameson s.n.”, no year, are very likely this syntype (bc.NY0001244170, bc.NY0001244171). Plants in these Jameson specimens do not display the features of B. plicata , such as obovate leaves, revolute leaf margins, hyaline leaf acumens, and dentate leaf apices. Instead, these Jameson syntypes have the features of B. cirrhifolia , namely, leaves are long ovate lanceolate, leaf margins reflexed at the base, leaf acumens are concolorous, and the leaf apex is entire ( Fig. 2 ). Also, the upper leaf cells are long and sinuous, just like in B. cirrhifolia ( Fig. 3 ). Additionally, I have seen the following Jameson specimens from Ecuador, which are also B. cirrhifolia (ordered chronologically): Quito, Jameson 135, 1845 (BM!-herb. Wilson, BM!-herb. Hooker, G!), Jameson s.n., 1846 (BM!- herb. Wilson, BM!-herb. Hooker, BM!-herb. Dixon), Jameson s.n., 1847 (BM!- herb. Dixon), Jameson 1, 1847 (BM!-herb. Hooker, BR!, MICH!, NY!), Jameson 25, 1847 (BM!-herb. Lyon), Jameson s.n., 1848 (G!), and Jameson s.n. 1851 (G!). Moreover, there are images online of a specimen at P (bc.PC0701220) labeled “ Jameson s.n. Quito, Mitten Herbarium, NYBG”. This specimen is very likely B. cirrhifolia since it consists of one very large stem (over 4 cm long), leaves with long concolorous acumens, and sporophytes with a very long seta (2 cm). In conclusion, all Jameson specimens I have seen actually correspond to B. cirrhifolia . Therefore, the Jameson syntype cited in the protologue cannot possibly be selected as lectotype for B. plicata . The identity of the syntype “Andes Quitensis, Jameson s.n., no year, NY!-herb. Mitten (bc.NY01244170, bc.NY01244171) is Braunia cirrhifolia .

Nomenclatural note 3. Cardot (1911) cited two specimens in the description of B. plicata var canescens: Pringle 15171 and Pringle 10627. I have seen several duplicates of both collections. The specimen Pringle 10627 should not be considered as lectotype since most duplicates consist of mixtures of different Braunia species (see Nomenclatural note 4). Only the selection of Pringle 15171 (P-herb. Cardot) as the lectotype preserves the identity of this taxon without confusion. The duplicates Pringle 15171 at NY! and S! are good isolectotypes. But the specimen of Pringle 15171 at FH contains mostly plants of B. andrieuxii , which I separated in an envelope with a note.

Nomenclatural note 4. The syntype specimen Pringle 10627 of B. plicata var canescens is problematic since most duplicates consist of mixtures of up to three species: B. plicata , B. andrieuxii , and B. secunda . Notably, Thériot (1926) had already warned that not all duplicates of Pringle 10627 were really B. plicata var canescens . Only seven duplicates of Pringle 10627 (BM!, H!, JE!, MO!, NY! [bc.00792521], S!, US!) are authentic B. plicata . But I have seen other eight duplicates, which are misidentified; five specimens (FH!, G!, NSW!, S!, B!) are B. andrieuxii , and the other three duplicates (MICH!, NY!, JE! [bc.JE04000338]) are B. secunda . Additionally, one of the duplicates of Pringle 10627 at P (bc.PC0147304) is certainly not genuine B. plicata . The images available at the MNHN web site show leaf features that correspond most likely to B. andrieuxi . The images online in the same web site for two other duplicates (bc.PC0701217, bc.PC0701218) show plants, with hyaline leaf apices, which are likely B. plicata .

Plants in dense mats; main stems sympodial, plagiotropic, 3–(4–5)– 7 cm long, closely branched; branches elongated, ending in obtuse, rosulate tips; flagelliform branches common; pseudoparaphylia foliose, with papillose cells. Leaves ovate or broadly ovate, 1.9–2.2 mm long, 0.8–1.1 mm wide, concave, imbricate, spreading when moist, strongly plicate; margins recurved to acumen; high base, widest point at or above middle of leaf; apex cuspidate, hyaline, margins sharply dentate; apical leaf cells homogeneous, elongated, narrow elliptic (5–7:1), 18–28 µm long, not sinuose, with marginal papillae; upper and median leaf cells 14–18 µm long, sub-quadrate to short elliptic (2:1), walls sinuose, papillae 2-3, low, on side walls, overarching lumen.

Autoicous. Archegonia medial on short vaginula (1.4–1.6 mm); perichaetial leaves narrow lanceolate, 2.4–2.9 mm long, less than twice the length of vegetative leaves, covering 1 mm of seta, acuminate, plicate, perichaetial paraphyses biseriate at base, immersed. Setae 7 (9–10) 12 mm long; capsule exserted, necks 0.3–0.4 mm, broad, gradually attenuated, urns 1.7–2.5 mm long, narrowly elliptic, mouth narrow, nearly 1/2 as wide as urn, capsule walls smooth, exothecial cells rectangular; stomates superficial; opercula base conical, short oblique rostrate. Calyptrae cucullate, large. Spores 29–31 µm in diameter.

Habitat. Commonly grows on trees, but also colonizes andesitic rocks in exposed habitats and within forests at middle to high elevations (1500–2700 m).

Distribution. Bolivia, Peru, Mexico.

Specimens examined:— BOLIVIA. Bridges s.n., 1846 (BM!-herb. Hooker, BM!-herb. Wilson, NY!-herb. Hooker, NY!-herb. Wilson); La Paz: Saavedra, NW of Chuma, Lewis 79-947 (F!); Cochabamba: Iscaypata, cerca de Quillacollo, De Luna 2094 (DUKE!, LPB!); in market at Cochabamba, Hermann 25280 (NY!); Santa Cruz: In der Cordillere von Santa Cruz, 1600m, Herzog 3482 (JE!, S!); Comarapa und Abra de Catalina, Herzog 3703 (JE!, S!).

— PERU. Cusco: Calca, Pisac, Marin 622 (F!); Laguna Urpicancha, near Huacarpay, Inohue 828 (JE!).

— MEXICO. Aguascalientes: San José de Gracia, La Congoja, Villalobos 8 (MEXU! NY!); Hidalgo: Cuyamaloya, Pringle 10627 (BM!, H!, JE!, MO!, NY!, S!, US!); ridge above Real del Monte, near Pachuca, Sharp 797 (MICH!, US!); S of San Juan del Rio, Rauh & Dull 122 (H!); Estado de México: Toluca, Pringle 15171 (FH! proparte, NY!, S!); Cerro Gordo, Cárdenas 2200 (DUKE!); San Martín de las Pirámides, Cerro Gordo, Allen 12943 (MO!); Nuevo León: 3 mi NW of Picacho Onofre, Nesom &Wells 34 (DUKE!); Cerro del Viejo, 15 mi W Dulces Nombres, Meyer & Rogers 149 (G!, NY!); Querétaro: cerro El Pinguical, cerca de Pinal de Amoles, Ocampo 620 (XAL); Tlaxcala: Mt. Tlacuapanga, Sharp 433 (MICH!); cerca de Villareal, al NE de Huamantla, De Luna 2646 (XAL!), 2718 (XAL!), 2720 (XAL!).

Discussion. This is strictly the species concept from that of Mitten (1869). Morphological characters distinguishing B. plicata are the leaves imbricate, strongly plicate, and concave ( Fig. 1C ). The leaf apex is cuspidate, hyaline, and dentate ( Figs. 1D , 4A–C, 4I–J ). Shoots are often dark-green to brownish, especially darker green along the leaf plications, which contrasts sharply with the hyaline leaf acumens ( Fig. 1A–C ). Braunia plicata is mostly montane Andean. Among the South American species with hyaline leaf acumens, the dentate apex and the narrowly elongated apical cells ( Figs. 1A , 4H ) separate B. plicata from B. incana , B. canescens and B. subincana . The following key is proposed to help in distinguishing B. plicata from the other three species with hyaline apices.

Figures

FIGURE 1. Lectotype of Braunia plicata. A) Specimen Bridges s. n., 1846, at BM (herb. Hooker). B) Detail of lectotype, showing one capsule. C) Detail of leaves with hyaline apex. D) Leaf apex hyaline and dentate. (Photographs by E. De Luna).

FIGURE 2. Jameson syntype of Braunia plicata. A–B) Vegetative leaves in the Jameson syntype. C) Perichaetial leaf in the Jameson syntype. D–E) Vegetative leaves characteristic of Braunia cirrhifolia. F. Perichaetial leaf of B. cirrhifolia.A–C from Jameson s.n. (syntype of B. plicata, NY-herb. Mitten). D–F from Jameson 1 (BM), type for B. cirrhifolia.

FIGURE 3. Jameson syntype of Braunia plicata. A–B) Leaf cells in the apex and upper leaf, in the Jameson syntype. C–D) Leaf cells in the apex and upper leaf, characteristic of B. cirrhifolia. A–B from Jameson s.n. (syntype of B. plicata, NY-herb. Mitten). C–D from Jameson 1 (BM), type for B. cirrhifolia.

FIGURE 4. Gametophytic and sporophytic features of Braunia plicata. A–C, I–J) Vegetative leaves. D–E) Perichaetial leaves. F, K) Capsules. G) Upper leaf cells. H) Apical leaf cells.A–H from lectotype Bridges s.n. (BM-herb-Hooker). I–K from Herzog 3703 (JE).