A revision of the genus Orthostichella (Neckeraceae)

 

Orthostichella C. Müll. is a genus of predominantly epiphytic, frequently pendulous mosses of tropical and subtropical American-African distribution. Schwägrichen (1816) described the oldest species now placed in Orthostichella (as Hypnum hexastichum Schwägr.), but the species generally accepted in the genus were first brought together by Müller (1850) under Neckera Hedw. sect. Pseudopilotrichum. C. Müll. subsection Orthostichella C. Müll. Müller (1879) began using his subsection Orthostichella at the generic level, however, he did not indicate this at the time, and because three names were placed in the genus, it is considerd invalid. Müller (1890) again used Orthostichella at the generic level, but once more this name is invalid because he described five species in the genus. In 1897 Müller yet again used Orthostichella at the generic level, but because he described only one new species the genus becomes validated under Article 42 (dealing with descriptio generico-specifica provisions) of the International Code of Botanical Nomenclature (Greuter 2000).

 

Orthostichella has long been associated with Pilotrichella (C. Müll.) Besch. The relationship between the two taxa began when Bescherelle (1872) elevated Müller’s (1850) Neckera Hedw. sect. Pseudopilotrichum. C. Müll subsect. Pilotrichella to generic rank. The newly established Pilotrichella was broadly conceived and included four sections: Orthostichella (C. Müll.) Besch., Eupilotrichella (C. Müll.) Besch., Papillaria (C. Müll.) Besch., and Meteoridium (C. Müll.) Besch. Papillaria had previously been removed from this group by Lorentz (1864). Jaeger & Sauerbeck (1877) refined Pilotrichella by dividing it into two unranked groupings: Eupilotrichella (including Bescherelle’s sect. Meteoridium) and Orthostichella. Section Meteoridium was removed from Pilotrichella by Brotherus (1906) who also positioned the genus (with sections Orthostichella and Eupilotrichella) in the tribe Meteorieae. This association of Orthostichella with Pilotrichella and the placement of Pilotrichella in the Meteoriaceae was followed by Fleischer (1908), Brotherus (1925), Bartram (1949), Florschütz (1964), Walther (1983), Vitt (1984), Spessard-Schueth (1994), Churchill & Linares (1995), Duarte-Bello (1997), and Magill & van Rooy (1998).

 

Orthostichella differs significantly from Pilotrichella in its smaller plant size, and in often having primary stolons as well as stipitate stems. It also differs from Pilotrichella in having spirally ranked leaves with sparsely developed alar cells, and often its leaves have a single or ddouble costae. Sporophytically Orthostichella differs from Pilotrichella in having shorter setae and smooth to papillose exostome teeth. Also the presence of large spores (to 64 μm) in Pilotrichella distinguishes the two taxa. In addition to Müller’s treatments (1879, 1890, 1897), Orthostichella has been recognized as a genus distinct from Pilotrichella by Buck (1994, 1994a), Buck & Goffinet (2000), Gradstein et al. (2001), Allen & Magill (2003), and Goffinet & Buck (2004).

 

The Meteoriaceae are usually placed in the Leucodontales (Fleischer 1908, Brotherus 1925, Walther 1983, Vitt 1984). The family, however, was transferred to the Hypnales by Buck (1994) and placed near the Brachytheciaceae in part because its exostome teeth are often horizontally striate at base and it lacks stolon-like primary stems that are tightly adherent to the substrate as well as greatly reduced stolon leaves. Buck (1994, 1994a) also reconsidered Pilotrichella and its systematic placement. As a result Orthostichella, Pilotrichella, Weymouthia Broth. and Squamidium (C. Müll.) Broth. were transferred to the Lembophyllaceae.

 

There are a number of tropical and subtropical pleurocarpous genera that grow pendulous in predominately epiphytic habitats. These genera present classification problems because they appear to represent several phylogenetic lines as judged by their very different peristomial forms, but they exhibit considerable convergence in their gametophytic features. Orthostichella is one of these problematic genera. It is difficult to decide if it belongs in the Meteoriaceae or Lembophyllaceae because its reduced peristome shows affinities to both families and it is hard to determine whether its gametophytic features are indicative of propinquity of descent or convergence. This situation is further complicated because the Meteoriaceae and Lembophyllaceae are so similar (e.g., compare the family descriptions of the Lembophyllaceae and Meteoriaceae in Buck & Goffinet 2000).

 

The leaves and stems of most genera placed in the Meteoriaceae often have parts that are intensely black. This odd feature is usually only noticed in passing, but it is so distinctive that when present one can immediately assign unknown specimens to the Meteoriaceae. Genera placed in the Lembophyllaceae can be green, yellowish red or brown, but they never exhibit this intense, at times shiny, black color. Furthermore, all of the genera now placed in the Meteoriaceae that can produce this intense black coloration also have single costae. On the basis of these two features it appears that Squamidium should be returned to the Meteoriaceae. Pilotrichella and Weymouthia which are ecostate or have short double costae and a green, yellowish red or brown coloration seem properly placed in the Lembophyllaceae. Orthostichella on the other hand seems better placed in the Neckeraceae by virture of its stolon/stipe morphology, poorly and variably developed leaf costae, and yellowish white neckeroid peristome. Within the Neckeraceae the genus is isolated by virtue of its non-complanate leaves that are often arranged in spiral rows.

 

Orthostichella has a complex morphology with creeping primary stems and erect or pendent secondary stems. Its stolons, stems, and branches are identical in morphological structure. It lacks a stem central strand, and its axillary hairs are usually reddish throughout. The secondary stems can be strongly stipitate or evenly foliate throughout. Remarkably, stipitate and evenly foliate secondary stems can sometimes be found on the same primary stem. Its branches often end in filiform attenuations or stolons. These different structures can be morphologically distinct, but more often they intergrade one into another or they can be abruptly transformed from one structure to another. As a rule the leaves on the branches, and sometimes the secondary stems are variously spirally ranked while those on the stolons and primary stems are unranked. But, this rule is often violated. Most leaves are cuspidate, have distinctive incurved, serrulate upper leaf margins, elongate, smooth, firm-walled leaf cells, and weakly developed, reddish yellow, firm-walled alar cells. The costa in O. versicolor is one of its oddest features. In some species most leaves are ecostate, however, here and there leaves can be found with short double, short single, or long single costae. In other species most leaves have a long single or long double costa but here and there ecostate leaves can be found. Gametophytic features of Orthostichella that show significant variation include: 1. relative plant size; 2. leaf shape; 3. leaf stance; 4. costa development; and 5. leaf apex shape.

 

The sporophytes of Orthostichella are rarely encountered but, they are uniform throughout the genus. The setae are elongate-flexuose and variously papillose roughened above. The genus has ovoid to short-cylindrical capsules, long-rostrate opercula, and mostly hairy, cucullate calyptrae. The Orthostichella peristome is diplolepideous and reduced with yellowish white exostomes and endostomes that are nearly the same length as the exostome teeth. The more or less linear exostome teeth are lightly horizontally striate on the dorsal (outer) surface at base. The endostome has a low basal membrane with filamentous, narrowly perforated segments, and cilia are usually absent. Even though the Orthostichella peristome is significantly reduced in form, the presence at the base of the exostome teeth of horizontal striae indicates the peristome is basically hypnoid.

 

The name Orthostichella combines the Greek orthos (straight) and Stichos (row) with the Latin substantival suffix -ella (diminutive). The name refers to the strong tendency for ranked leaves in the genus, although the branch leaves are often in spiral rather than straight rows.

 

Orthostichella C. Müll., Bull. Herb. Boissier 5: 204. 1897.

 

Type: O. filamentosula C. Müll. (Buck 1994).

 

Neckera sect. Pseudopilotrichu  C. Müll.,Syn. 2: 123. 1850. Pseudopilotrichum (C. Müll.) Buck & Allen in Buck, J. Hattori Bot. Lab. 75: 69. 1994. Lectotype: N. hexasticha Schwägr. C. Müll. (Buck 1994).

Neckera sect. Pseudopilotrichum subsect. Orthostichella C. Müll., Syn. 2: 123. 1850. Lectotype: N. hexasticha Schwägr. C. Müll. (Buck 1994).

 

Plants slender or medium-sized, dull, yellow-green, reddish yellow or brownish red, in dense or loose mats often with pendent strands. Stolons red, creeping; leaves widely spaced, reduced, erect, broadly ovate or ovate-triangular, abruptly long-pointed, ecostate or costa short and double. Primary stems occasionally stipitate, horizontal or erect, frequently and irregularly branched, at times with flagelliform tips, yellowish green, becoming dark red with age, in cross section with sclerodermis, enlarged cortical cells and no central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3–4 cells long, basal 1 cell quadrate to subquadrate, reddish brown, upper 2–3 cells broadly cylindrical, hyaline or reddish; rhizoids only on stolons red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Stipe leaves ovate below, piliferous above, auriculate, cordate, rounded at base or straight to the insertions. Stipe leaves ovate,; costae short and double; alar cells well developed. Secondary stem and branch apices turgid; stem leaves not seriately ranked, ovate, erect, variously clasping below, rounded to the base, abruptly narrowly acuminate, at times hair-pointed; branch leaves obscurely seriately ranked when dry, distinctly so when wet, leaves panduriform, broadly oblong, obovate, oblong-obovate, or broadly obovate, erect to erect-spreading, appressed and at times clasping below, abruptly flexed and spreading above, stem leaves often auriculate, apex abruptly acuminate, cuspidate to long-cuspidate; margins broadly incurved, sharply serrulate above, plane or broadly incurved, serrulate below, entire or weakly toothed on basal auriculation; costae short double, short single, long single, or absent; leaf cells long flexuose, smooth to slightly porose, outer basal cells shorter and broader, inner basal cells elongate, porose, alar cells subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 2–4 rows, yellow across the insertion. Asexual reproduction by deciduous leaves. Dioicous. Perigonia gemmate, lateral on secondary stems and branches, perigonial leaves ovate, acuminate, costa absent; paraphyses present, antheridia few. Perichaetia lateral, paraphyses and archegonia numerous; perichaetial leaves lanceolate, vaginula often densely hairy. Setae short, wavy, smooth below, roughened above, yellow. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, neck moderately developed; exothecial cells small, rounded in several rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate; annuli rudimentary, consisting of 2–3 rows of thin-walled enlarged cells; peristome yellowish white, exostome teeth linear, dorsal (outer) surface horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened. Calyptrae, cucullate, hairy or smooth.

 

Key to species of Orthostichella.

 

1. Branch leaves spreading from insertions.......................................................................................... 2

1. Branch leaves at base appressed and clasping stems....................................................................... 5

2. Branch leaves cuspidate; leaves consistently costate....................................... 4. O. pandurifolia

2. Branch leaves apiculate to mucronate....................................................................................... 3

3. All leaves with a long, single percurruct costa......................................................... 3. O. longinervis

3. Leaves mostly ecostate, sporatically costae, the costa short and double or single and ending at
midleaf................................................................... .............................................................. 4

4. Stem leaves hair-pointed ................................................................................. 7. O. welwitschii

4. Stem leaves cuspidate ....................................................................................... 6. O. versicolor

5. Stipe and secondary stem leaves rounded to the insertion; setae 1.0–1.5 mm long.............................

..................................................................................................................... 2. O. hexasticha

5. Stipe and secondary stem leaves auriculate; setae 3.5–4.0 mm long................................................ 4

6. Auriculate leaf bases serrate or dentate............................................................. 1. O. calomicra

6. Auriculate leaf bases entire ................................................................................... 5. O. roseana

 

 

1. Orthostichella calomicra (Broth.) Allen & Magill, comb. nov.

 

Pilotrichella calomicra Broth., Bol. Soc. Brot. 8: 180. 1890. Protologue: São Tomé and Príncipe. Ins. S. Tomé, ubi legit Fr. Quintas. Types: Afr. occ., ins. S. Thomé, leg. Fr. Quintas (H, holotype); Afr. occ. ins. S. Thomé, leg. Fr. Quintas (Comm. Broth.) (S, isotype); Ins. S. Thomé, leg. Fr. Quintas (S, isotype).

Pilotrichella isleana Besch. var. virescens Besch., Ann. Sci. Nat., Bot. sér. 6, 10: 267. 1880. Protologue. Réunion: Plaine des Cafres, G. De L’isle; Cilaos, hauts du Matarum, Valentin, 1876. Types. La Reunion. G.-de l’Isle [Herb. Mus. Paris] (H, lectotype designated here); La Reunion. M. g.-de l’Isle, com. Brotherus (S, 2 isolectotypes).

Pilotrichella holstii Broth., Bot. Jahrb. Syst. 20: 197. 1894. Protologue: Kenya/Tanzania. Usambara; ohne nähere Standorte (Holst n. 698). Types: Usambara, leg. C. Holst 698 (H, holotype); .Usambara, leg. C. Holst, com. Broth. (S, isotype).

Pilotrichella muelleri Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 33. 1895. Protologue: Cameroon. Habitat in Camerunia in ramulis arborum, ubi supra Beam pagum c. 1,600 metra supra mare m. Julio a. 1891 legi [Dusén]. Types: Africa occ. in monte Camerunensibus in ramulis arborum c. 1600 m. Julii 13 1891, P. Dusén 262 (S, holotype); Musci Africani in Camerunia a P. Dusén collecti. 262. Orthostichella Duseni C. M. In montibus Camerunensibus ad Bueam pagum c. 1600 metra supra mare in ramulis arborum die 13 m. Julii a. 1891 (H, NY, S [4 collections], isotypes).

Orthostichella duseni C. Müll., Kongl. Svenska Vetensk. Acad. Handl. 28(2): 33. 1895, invalid name listed in synonymy. Pilotrichella duseni (C. Müll.) Par., Index Bryol. Suppl. 270. 1900, illegitimate name includes type of species with priority. Based on Dusén Musci Camerunia n. 262 (H, S).

 

Plants slender, dull, yellow-green, reddish yellow or brownish red, in dense mats. Stolons red, creeping; leaves widely spaced, reduced, erect, broadly ovate, abruptly long-pointed, ecostate or costa short and double, 0.6-0-8 mm long. Primary stems occasionally stipitate, horizontal or erect, frequently and irregularly branched, at times with flagelliform tips, to 5 cm long, yellowish green, becoming dark red with age, in cross section with sclerodermis of 3–5 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3–4 cells long, basal 1 cell quadrate to subquadrate, reddish brown, upper 2–3 cells broadly cylindrical, hyaline or reddish; rhizoids only on stolons red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Secondary stem and branch apices turgid; stem and branch leaves differentiated only in size, obscurely seriately ranked when dry, distinctly so when wet, leaves panduriform, broadly oblong, appressed and clasping below, broadly obovate, abruptly flexed and spreading above, stem leaves 1.0–1.5 mm long, strongly auriculate, apex apbruptly acuminate, long-cuspidate, cuspid 80–140 µm long; margins broadly incurved, sharply serrulate above, plane or broadly incurved, serrulate below, weakly toothed on basal auriculation; costae short double, short single, or absent; leaf cells long flexuose, smooth to slightly porose, 18–40 x 3–5 μm, outer basal cells shorter and broader, inner basal cells elongate, porose, alar cells subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 2–4 rows, yellow across the insertion. Dioicous. Setae short, wavy, 2–3 mm long, smooth below, roughened above, yellow. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.0–1.5 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate, 1.5 mm long; annuli rudimentary; peristome yellowish white, exostome teeth linear, 0.5–0.6 mm long, dorsal (outer) surface horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 17–26 μm long. Calyptrae 2 mm long, cucullate, hairy.

 

Etymology. The specific epithet calomicron combines the Greek Kalo- (rope) with micro (small) and refers to its distinctly serriately ranked leaves.

Distribution. West-Central and East Tropical Africa, Western Indian Ocean.

Illustrations. Dusén (1895, p. 33 a & b; 3f 1).

Ecology. On trees and shady rocks; 1200–1900 m.

 

Selected specimens examined

São Tomé and Príncipe. São Tomé: Quintas (H).

CAMEROON. Bueam (H, NY, S)

TANZANIA. Arusha: Crosby & Crosby 9119 (BM, FH, H, MO, NY, S, JE); Tanga: Pócs 8531I (MO).

MADAGASCAR. Antserana: Crosby & Crosby 7180 (H, MO, NY, S); Antananarivo: Crosby & Crosby 9281 (MO); Fianarantsoa: Crosby & Crosby 6873 (BM, JE, MO, NY); Toamasina: Magill et al. 9548 (B, FH, G, MO, NY).

REUNION. Arrondissement au Vent: Crosby & Crosby 8980 (B, BM, CAS, DUKE, E, F, FH, G, H, JE, MEXU, MO, NICH, NY, PC, S, UC, US).

 

Orthostichella calomicra is a small to medium-sized species with closely spaced, panduriform stem and branch leaves. The primary stems of O. calomicra are often branched close to the stolons, as a result the plants often form compact mats. The stem and branch leaves are appressed, clasping below and abruptly flexed and spreading above. The stem and branch leaves are strongly auriculate at base and the basal margins are nearly always weakly toothed. The costae of O. calomicra are consistently weak and the alar cells are poorly developed.

 

Morphologically O. calomicra is extremely close to O. roseana which differs in having a looser habit, entire basal leaf margins, longer setae, and smaller spores. In size and aspect it is nearly identical to O. hexasticha, and both species have the same leaf stance. Orthostichella hexasticha differs from O. calomicra in having stem leaves that are rounded to the insertion rather than strongly auriculate Orthostichella versicolor has forms that often have strongly auriculate stipe and, less often, stem leaves. Furthermore, O. versicolor and O. calomicra are more less similar in size. Orthostichella calomicra consistently differs from O. versicolor in having branch leaves that are appressed and clasping at base.

 

2. Orthostichella hexasticha   (Schwägr.) Buck, Bryologist 97: 435. 1994.

 

Hypnum hexastichum Schwägr., Sp. Musc. Frond. Suppl. 1(2): 210. 1816. Isothecium hexastichum (Schwägr.) Brid., Bryol. Univ. 2: 380. 1827. Pilotrichum hexasticum (Schwägr.) C. Müll., Bot. Zeitung (Berlin) 6: 767. 1848. Neckera hexasticha (Schwägr.) C. Müll., Syn. Musc. Frond. 2: 126. 1850. Meteorium hexastichum (Schwägr.) Mitt., J. Linn. Soc., Bot. 12: 432. 1869. Pilotrichella hexasticha (Schwägr.) Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 257. 1877. Protologue: Dominican Republic. In Domingo insula lectum communicavere cl. Thouin et Hornemann. Holotype: Hypnum hexastichum Domingo Thouin. Diund [?]. Hornemann. (G).

 

Pilotrichella tenella Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 261 1877, new name for Neckera tenella C. Müll., Bot. Zeitung (Berlin) 17: 238. 1859, not Neckera tenella Schwägr., 1827., illegitimate homonym. Protologue: Insula St. Domingo. A bryologo Tönder suecico Lund. Mohr accepti. Lectotype (designated here): Leskea n. sp. tenella ex S^t domingoa Mohr & Weber [Herb. Swartzii] (S).

 

Plants slender to medium-sized, dull, light green, yellow-green, reddish yellow or brownish green, in loose mats often with pendent strands. Primary stems red, creeping; leaves reduced, erect, ovate-triangular, 0.4–1.0 mm long. Secondary stems from primary stems stipitate, others horizontal, erect or pendent, frequently and irregularly branched, branches at times with flagelliform tips, to 4–10(–25) cm long, yellowish green, becoming dark red with age, in cross section with sclerodermis of 3–4 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 2–3 cells long, basal cell quadrate to subquadrate, reddish brown, upper 1–2 cells broadly cylindrical, hyaline (occasionally reddish); rhizoids on primary stems, at base of stipitate secondary stems, or on flagelliform branch tips, red, not or irregularly branched, from clusters of initials abaxial to the leaf insertion. Secondary stem and branch apices turgid; stem and branch leaves differentiated only in size, obscurely seriately ranked when dry, distinctly so when wet, leaves panduriform, broadly oblong, appressed, more or less clasping below, broadly obovate, abruptly flexed and spreading above, 0.5–1.2 mm long, at times somewhat auriculate, apex acuminate, long-cuspidate; margins broadly incurved, sharply serrulate above, plane or broadly incurved, serruate or entire below; costae short double, short single, or absent; leaf cells long flexuose, smooth to slightly porose, 16–37 x 3–4 μm, outer basal cells shorter and broader, inner basal cells elongate, porose, alar cells subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 4–6 rows, yellow across the insertion. Dioicous. Perigonia gemmate, lateral on secondary stems and branches, 0.8–1.0 mm long, perigonial leaves ovate, acuminate, costa absent; paraphyses present, antheridia few. Perichaetia lateral, 2.5 mm long, paraphyses and archegonia numerous; perichaetial leaves lanceolate 1.8–2.2 mm long, vaginula densely hairy. Setae short, wavy, 1.5–2.4 mm long, smooth, yellow. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.0–1.5 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate, 1.0 mm long; annulus not seen; peristome yellowish white, exostome teeth linear, to 0.5 mm long, dorsal (outer) surface lightly horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane low, segments filamentous, narrowly perforated, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 14–20 μm long. Calyptrae narrowly cucullate, with a few long hairs.

 

Etymology. The specific epithet hexasticha combines the Greek hexa- (six) and -sticha (in a row) in reference to its seriately ranked leaves.

Distribution. Caribbean (Cuba, Haiti, Dominican Republic).

Illustrations. Duarte-Bello (1997, Pl. 203); Buck (1998, Pl. 91 1–8).

 

Orthostichella hexasticha   A. Habit. B. Median leaf cells. C. Leaf apex. D. branch leaf. E. basal leaf cells. F. Stem leaf.

 

Orthostichella hexasticha   A. Calyptra. B. Sporophyte. C. Operculum. D. Exothecial cells showing stomata. E. Exothecial Cells. F. Axillary hairs. G. Stem cross-section.

 

Ecology. On rocks (laterite), tree trunks, branches, twigs, shrubs, lianas, and rotting logs in pine forests, cloud forests, and roadbanks; 700–2000.

 

Selected specimens examined. CUBA. Guantánamo: Gradstein, Bryophyta Neotropica 370 (MO); Gramma: Taylor 528 (NY); Sancti Spíritus: Buck 23441 (NY); Santiago de Cuba: Leon 11228 (FH, NY); Villa Clara: Pócs & Borhidi 9011/AH (MO). HAITI. Ouest: Imshaug 23071 (MO); Sud: Duncan 37 (MO); Sud’est: Buck 9395 (H, NY). DOMINICAN REPUBLIC. Azua: Mejía & Zanoni 8270 (MO, NY); Barahona: Abbott 1789 (B, BM, FH, NY); Independencia: Bolay 41 (MO); La Estrelleta: Reese 15410 (NY); La Vega: Zanoni et al. 39270 (NY); Pedernales: Steere 22830 (NY); San Rafael: Norris et al. 6804 (NY).

 

The most distinctive feature of O. hexasticha is its leaf shape. The strongly panduriform leaves are erect, clasping at base, then abruptly reflexed outward, and finally curved inward toward the stem. The upper portion of the leaves is strongly concave with broadly incurved, sharply serrulate upper margins that nearly meet below apex. The basal leaf margins are rounded to the insertion or somewhat auriculate, often broadly incurved, entire or serrulate at base. This leaf shape gives the plants an overall aspect that is helpful in recognizing the species. Unfortunately there are numerous collections (including the type of O. hexasticha) that consist of small plants that have this apect weakly developed (see e.g., Norris B6612 MO, NY). Furthermore, these collections have leaves that are distinctly ranked when dry. They must be examined with a compound microscope before they can be positively identified. In these collections of O. hexasticha the presence of panduriform leaves that are not or moderately auriculate at base will separate them from all other Neotropical members of the genus. The branches and stems of O. hexasticha frequently end in flagelliform tips.

 

Squamidium nigricans (Hook.) Broth. is similar in size and aspect to O. hexasticha. Plants of S. nigricans usually have some parts with an intense blackish color and this feature can be used to separate it from O. hexasticha which never has this color. In addition, S. nigricans never has flagelliform branch and stem tips. Microscopically, the presence in the leaves of a long, faint single costa, and greatly enlarged, strongly differentiated alar cells distinguished S. nigricans from O. hexasticha.

 

Orthostichella hexasticha is found in Cuba, Hispaniola, and Puerto Rico. The Puerto Rican collections of O. hexasticha represent the small form noted above. Buck (1998) considered these collections to be O. versicolor.

 

3. Orthostichella longinervis (Ren. & Card.) Allen & Magill, comb. nov.

 

Pilotrichella loninervis Ren. & Card., Bull. Soc. Roy. Bot. Beligique 32(2): 23. 1893. Orthostichopsis longinervis (Ren. & Card.) Broth., Nat. Pflanzenfam. 1(3): 805. 1906. Protologue: Madagascar: ad truncos; Diego Suarez (Chenagon); in silvis ditionis Antsianaka (fratres Perrot); inter Vinanintelo et Ikongo (D^r Besson).

 

Plants slender to medium-sized, dull, light green, yellow-green, reddish yellow or brownish green, in loose mats at times with pendent strands. Stolons yellow to red, creeping; leaves widely spaced, imbricate, narrowly ovate, abruptly cuspidate, to 1.5 mm long; costa single, long, extending into the leaf apex; alar cells moderately, dark-red, quadrate,. Primary stems at times strongly stipitate or foliose to the stolons, horizontal, erect or pendent, 2–12 cm long, frequently and irregularly branched, branches often with flagelliform tips, yellowish green, becoming dark red with age, in cross section with sclerodermis of 2–5 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3–4 cells long, basal 1–2 cells quadrate to subquadrate, reddish brown, upper 2–3 cells broadly cylindrical, hyaline or reddish; rhizoids consistently on stolons, also at base of stipitate secondary stems, red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Stipe leaves broadly ovate, abruptly cuspidate, rounded at base; costae long, single, reaching into the apex; alar cells moderately developed. Secondary stem and branch apices turgid; stem and branch leaves differentiated only in size, stem leaves not or obscurely seriately ranked when dry, branch leaves often ranked, dry or wet, leaves obovate, oblong-obovate, to panduriform, erect to erect-spreading, variously clasping below, 0.8–1.3 mm long, rounded to the insertion, apex shorlty cuspidate; margins broadly incurved and subentire to serrulate above, plane or broadly incurved, serrulate or entire below; costae consistenly long, single, usually subpercurrent, at times wavy, often with short lateral spurs; leaf cells long flexuose, smooth, 20–45 x 3–5, outer basal cells shorter and broader, inner basal cells near insertion elongate, often porose, alar cells red to red-yellow, subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 3–6 rows. Dioicous. Perigonia gemmate, lateral on secondary stems and branches, 1 mm long, perigonial leaves ovate, acuminate, costa absent; paraphyses present, antheridia numerous. Perichaetia lateral, 1.2-1.8 mm long, paraphyses and archegonia numerous; perichaetial leaves lanceolate, vaginula lightly hairy. Setae short, wavy, 2–3 mm long, smooth below, roughened-papillose above, yellow, becoming reddish with age. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.5 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; operculum and annulus not seen; peristome yellowish red, exostome teeth linear, 0.2–0.4 mm long, dorsal (outer) surface horizontally papillose-striate at base, papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 14–18 μm long. Calyptra not seen.

 

Etymology. The specific epithet longinervis combines the Latin longus- (long) with nervus (nerve) and refers to the long costa found in the leaf of the species.

Distribution. Western Indian Ocean.

Illustrations. Renauld and Cardot (1895–1905, Pl. 83 1).

Ecology. On tree branches; 800–1050 m.

 

Specimens examined.

MADAGASCAR. Antananarivo: Crosby & Crosby 5190 (MO); Antseranana: Magill et al. 9947 (MO); Toamasina: Magill et al. 9664 (BM, FH, H, MO, NY, S), Crosby & Crosby 9319 (MO).

 

Orthostichella longinervis is a medium-sized species that consistently has extremely well developed stipitate stems. There are some collections of O. panduraefolia with strongly stipitate stems, and while the feature occurs here and there among the other Orthostichella species, it is never as prominent as found in O. longinervis. The critical feature of O. longinervis, however, is its costal structure. The costa is consistently expressed, long, typically subpercurrent, strongly developed, and also often has short lateral spurs. This feature makes the species unique in the genus, which typically has an extremely variable costa: double, single, or absent on leaves from a single plant. Indeed, these two features of O. longinervis are so odd within the context of Orthostichella that they raise questions as to whether the species is properly placed there.

Athough the gametophytes of Orthostichella can be extremely variable the same can not be said for its sporophytes. Indeed, within all of its species, the features of the Orthostichella sporophyte are remarkable stable. In this regard it seems important that the O. longinervis sporophyte is also identical in form to that of the other species. Furthermore, in terms of stem morphology, leaf shape, margin stance and serration, areolation, and alar cell development there is nothing unusual in gametophytes of O. longinervis to indicate it is misplaced in Orthostichella.

Renauld and Cardot (1883) considered O. longinervis to closely approach the aspect of Porotrichum, but to differ, among other ways, in having concave, subcocheariform, imbricate branch leaves in distinct seriate rows. They also regarded the presence of a long, single costa in O. longinervis as evidence that it might only be a form of Pilotrichella subimbricata or P. chrysoneura, both of which are now placed in Orthostichopsis. This latter view of Orthostichella longinervis was adopted by Brotherus (1906) when he transferred the species into Orthostichopis. Although the costa form in the leaves of Orthostichella longinerivis is identical to that found in Orthostichopsis, the taxon is clearly misplaced within Orthostichopsis because it has foliose rather than filamentous pseudoparaphyllia. The presence of filamentous pseudoparaphyllia is a sine qua non feature not only of Orthostichopsis but also the Pterobryaceae.

Although Orthostichella longinervis differs significantly from Porotrichum in having concave, spirally ranked leaves, there are other genera in the Neckeraceae that lack complanate leaves. In particular Porotrichodendron has one one species (P. lindigii (Hampe) Buck) with tumid, evenly foliate leaves. Evidence against the placement of O. longinervis in Porotrichodendron can be found in its lack of a stem central strand, as well as the very close similarity of its sporophyte to those of other Orthostichella species. On balance, this species appears best placed in Orthostichella where it occupies a critical position in supporting the placement of the genus within the Neckeraceae.

 

4. Orthostichella pandurifolia (C. Müll.) Buck, Bryologist 97: 435. 1994.

 

Neckera panduraefoliaC, Müll., Bot. Zeitung (Berlin) 13: 767. 1855. Pilotrichella pandurifolia (C.Müller) Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 255. 1877. Protologue: South Africa. Promontor. bonae spei: Zeyher copiose sterilem arboream anno 1823 legit. Type: South Africa. Prom. b. spei: Zeyher 1823 (isotype NY).

 

Neckera decolorans Hampe in C. Müll., Linnaea 40: 265. 1876, invalid name listed in synonymy. Pilotrichella decolorans C.Müller in Kindberg, Enum. Bryin. Exot., suppl. 2 102 1891, invalid name, lacks a description. Based on: Insula Johana, Comoros, leg. Hildebrandt [Herb. Hampe] (BM).

Pilotrichella kuntzei C. Müll., Hedwigia 38: 127. 1889. Protologue: Habitatio. Prom. bonae spei, Kingwilliamstown, “im Perie-Walde” ad Polypodium Eckloni: Dr. Otto Kuntze le. et. mis. 1894, synomized by Magill & Rooy (1998).

Orthostichella capillicaulis C. Müll., Flora 73: 491. 1890, invalid name, genus invalid at time of publication. Pilotrichella capillicaulis C. Müll ex Kindb. Enum. Bryin. Exot., suppl. 2:102 1891, new name for Orthostichella capillicaulis C. Müll. Protologue: Tanzania. Africa or. trop., Leikipia in occidente montis Kenia, ad pedem der Aberdare-Kette: L. Höhnel Novembri 1887 in Exped. Telekiana. Type: Tanzania. Afr. or. Kenia, leg. L. Höhnel (H, isotype).

Orthostichella curvifrons C. Müll., Flora 73: 491. 1890, invalid name, genus invalid at time of publication. Pilotrichella curvifrons C. Müll ex Kindb., Enum. Bryin. Exot., suppl. 2: 102 1891. new name for Orthostichella curvifrons C. Müll., Protologue: Tanzania. Africa or. trop., Leikipia in regione meridionali montis Kenia, ad pedem der Aberdare-Kette: L. Höhnel Novembri 1887 in Exped. Telekiana. Type: Tanzania. Afr. or. Leikipia, leg. L. Höhnel (H, isotype).

Orthostichella tenellula C. Müll., Flora 73: 492. 1890, invalid name, genus invalid at time of publication. Pilotrichella tenellula C. Müll. ex Kindb., Enum. Bryin. Exot., Suppl. 2: 102. 1891, new new for Orthostichella tenellula C. Müll. Protologue: Patria. Africa or. trop., in sylva primaeva montis Kilima-Ndscharo: L. Höhnel in Exped. Telekiana 1887. Type: Tanzania. Afr. or., Kilima-Ndscharo, leg. Höhnel (H, isotype).

Pilotrichella cuspidata Broth., Bot. Jahrb. Syst. 24: 255. 1897. Protologue: Pondoland: in feuchtem Laubgebüsch an dem Unterholz an allen Zweigen und an den Bäumchen rankend und wie Bartmoos herabhängend nur in einem Gebüsch des Egorawaldes gefunden, daselbst aber eine grosse Strecke des Gebüsches damit behangen (Beyrich n. 38, Bachmann n. 6). Types: Pondoland, Bachmann n^o 6 (BM, lectotype, designated here); Pondoland, Beyrich 38 (BM, syntype).

Pilotrichella incurva Broth., Bot. Jahrb. Syst. 24: 256. 1897. Protologue: Angola: Huilla (Antunes). Type. Afr. occ., Huilla, leg. Antunes comm. Brotherus (S, isotype).

Pilotrichella stuhlmanni Broth., Bot. Jahrb. Syst. 24: 256. 1897. Protologue: Seengebiet: Bukoba (Stuhlmann), Mau, an Bäumen c 2100 m (Scott Elliot n. 73). Types: Bukoba, Stuhlmann (S, syntype); Bukoba, Stuhlmann 1890, (S, syntype).

Pilotrichella conferta Ren. & Card., Bull. Soc. Roy. Bot. Beligique 38(1): 24. 1899 [1900]. Protologue: Lesotho. Hab. Africa australis: Lessouto (Vernet, herb. Boissier). Type. Africa australis: Lessouto, Leg. Vernet, 1879 (PC, holotype; BM, S, isotypes).

Orthostichella leptopteridea C. Müll. in Par., Index Bryol. Suppl. 271. 1900, invalid name, lacks a description. Pilotrichella leptopteridea Par., Index Bryol. Suppl. 271. 1900, invalid name, lacks a description, Based on: Africa trop. Schoah, in viciniis urbis Ankober legit et in ventre Horpensis Museo. zool. Florentino misit 1887 (Stopfmaterial), cl. D. Traversi, Hb. E. Levier (S)

Pilotrichella percordata Broth in P. Vard., Rev. Bryol. 47: 53. 1920, invalid name, lacks a description. Based on: Kikouyou, British East Africa, 1910. A. G. Allan 315, ex herb. Rev. D. Lillie (H); . Kikouyou, British East Africa, 1910. A. G. Allan, ex herb. Rev. D. Lillie, [Herb. Potier de la Varde], (PC).

Pilotrichella cordata Brotherus in Potier de la Varde, Bull. Soc. Bot. France 71: 1057. 1924. Protologue: [Tanganyka ?] Kiswani (leg. R.P. Wetzler) Invalid name, lacks a description. Based on Kiswani, (Mambra) Tanganyka Territory, Juni 1924, coll: R. P. Wetzler, comm. R. P. Soul (PC); Usumbara, Kiswani, leg. R. P. Soul (FH, H).

 

Plants slender to medium-sized, dull, light green, yellow-green, reddish yellow or brownish green, in loose mats often with pendent strands. Stolons yellow to red, creeping; leaves widely spaced, reduced, erect, ovate-triangular, abruptly acuminate, to 1.6 mm long; ecostate or costa short and double; alar cells well developed, dark-red, quadrate,. Primary stems at times stipitate or foliose to the stolons, horizontal, erect or pendent, frequently and irregularly branched, branches often with flagelliform tips, 2–15 cm long, yellowish green, becoming dark red with age, in cross section with sclerodermis of 2–5 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3–4 cells long, basal 1–2 cells quadrate to subquadrate, reddish brown, upper 2–3 cells broadly cylindrical, hyaline or reddish; rhizoids consistently on stolons, also at base of stipitate secondary stems, red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Stipe leaves ovate below, piliferous above, auriculate to strongly rounded at base; costae short and double; alar cells well developed. Secondary stem and branch apices turgid; stem and branch leaves differentiated only in size, stem leaves not or obscurely seriately ranked when dry, branch leaves often distinctly seriately ranked, dry or wet, leaves obovate to oblong-obovate, erect to erect-spreading, variously clasping below, 1.0–2.0 mm long, at times somewhat auriculate, apex long cuspidate, branch leaf cuspid 140-200 µm long; margins broadly incurved and subentire to serrulate above, plane or broadly incurved, serrulate or entire below; costae usually short double or short single, occasionally absent; leaf cells long flexuose, smooth, upper cells 20–46 x 4–6, median cells 30–60 x 3–4 μm, outer basal cells shorter and broader, inner basal cells near insertion elongate, often porose, alar cells red to red-yellow, subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 4–6 rows, yellow across the insertion. Asexual reproduction by deciduous leaves. Dioicous. Perigonia gemmate, lateral on secondary stems and branches, 0.8–1.2 mm long, perigonial leaves ovate, acuminate, costa absent; paraphyses present, antheridia numerous. Perichaetia lateral, 1.4-2.0 mm long, paraphyses and archegonia numerous; perichaetial leaves lanceolate 1.4–2.0 mm long, vaginula densely hairy. Setae short, wavy, 1.5–3.5 mm long, smooth below, roughened-papillose above, yellow, becoming yellowish red with age. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.0–2.0 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate, 1.0–1.5 mm long; annuli rudimentary, consisting of 2–3 rows of thin-walled enlarged cells; peristome yellowish white, exostome teeth linear, 0.3–0.4 mm long, dorsal (outer) surface horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 14–20 μm long. Calyptrae, 2–3 mm long,narrowly cucullate, lightly hairy.

 

Etymology. The specific epithet panduraefolia combines the Latin panduratus (fiddle-shaped) with folium (leaf).

Distribution. Northeast, East, and South Tropical Africa, Southern Africa, Western Indian Ocean.

Illustrations. Magill and Rooy (1998, Fig. 160 1–11).

Ecology. On tree trunks and branches, also fairly common on rocks; 100–1800 m.

 

Selected specimens examined.

DJIBOUTI. leg. unknown, 1906 (NY, S).

KENYA. Central: Magill 13892 (BM, H, MO, NY, S).

TANZANIA. Arusha: Pocs & Pocs 6213 (B).

ANGOLA: Huilla Antunes (S).

LESOTHO Vernet (S).

SWAZILAND. Hhohho: Magill 3497 (MO).

SOUTH AFRICA. Eastern Cape: Perold 4142 (MO); Kwazula-Natal: Magill 5173 (H, MO); Limpopo: Magill 6543 (H, MO); Mpumalanga: Koekemoer 2219 (MO).

COMORO ISLANDS. Johana Island: Hildebrandt (BM).

 

Orthostichella pandurifolia has creeping primary stems from which the secondary stems can be erect or pendent. The secondary stems can be distinctly long-stipitate or evenly foliate throughout, with both types of foliate secondary stems sometimes found on the same primary stem. Its branches commonly end in slender attenuations. The branch leaves, and sometimes the secondary stems, of O. pandurifolia are typically occur in distinct spiral ranks while those on the stolons and primary stems are unranked. The leaves are long-cuspidate, have incurved, serrulate upper leaf margins, elongate, smooth, firm-walled leaf cells, and weakly developed, reddish yellow, firm-walled alar cells. Most leaves in O. pandurifolia have a single or double costae that sometimes reache well above midleaf, however, here and there ecostate leaves can be found. This costal variation is opposite that of O. versicolor in which most leaves are ecostate, and only occasionally do the leaves have short double, short single, or long single costae. Sporophytes are not often found in O. pandurifolia. The setae are short (1.5–3.5 mm long), flexuose, and roughened above. The operculum is long, obliquely rostrate, and the peristome is pale-yellow and greatly reduced. Distinctive features of the peristome include its narrow, mostly papillose exostome teeth that strongly cross-striate at the very base, and narrow endostomial segments that are perforated but not keeled. The calyptrae are cucullate and sparsely hairy and the vaginula are densely hairy.

 

There are two major forms of O. pandurifolia. At the southern end of its range, the species has very strongly ranked branch leaves, and plants with deciduous leaves are unknown. In this form of the species sporophytes are occasionally found. The second form of the species (nomenclaturally centered on Pilotrichella stuhlmanni Broth.) is found in the middle and northern parts of its range. The plants in this region have weakly ranked branch leaves, and are nearly always found with deciduous leaves. None of the plants from this region have been found with sporophytes. Orthostichella pandurifolia is primarily an epiphytic species. There are however many collections that have been collected on boulders, and unlike O. versicolor, the saxicolous condition appears to be a primary rather than a secondary habitat for O. pandurifolia.

 

Orthostichella pandurifolia has sometimes been confused with Squamidium brasiliense (Hornsch.) Broth because they both are pendent mosses with with weaklyn developed costae. Squamidium brasiliense differs from Orthostichella pandurifolia in having julaceous stems, consistently long, single costae, and exceptionally well-developed alar cells. Orthostichopsis species differ from Orthostichella pandurifolia in having filamentous pseudoparaphyllia, long, strongly developed, single costae, and cylindrical capsules.

 

Orthostichidium pentastichum (Brid.) Allen and O. involutifolium (Mitt.) Broth. are similar to Orthostichella pandurifolia in having ecostate leaves with broadly incurved upper leaf margins and weakly developed alar cells. Plants of O. pentasticum are more slender and less branched than those of Orthostichella pandurifolia, while those of O. involutifolium tend to be a little bit larger than those of Orthostichella pandurifolia, and its leaves are more stiffly erect as well as more strongly ranked and they have filamentous pseudoparaphyllia. Furthermore, both Orthostichidium species have filamentous pseudoparaphyllia, and O. involutifolium often has clusters of uniseriate gemmae around their branch primordia. Orthostichella pandurifolia never has asexual gemmae, and the branch primordia are surrounded by scale leaves.

 

5. Orthostichella roseana Allen & Magill, species nova.

 

Type: Ecuador. Vicinity of Huigra, mostly on the Hacienda de Licay, October 1918, J. N. Rose 2400 (holotype NY; isotypes B, FH, MO, NY, S).

 

Species haec a O. hexasticha inter alia foliis caulium auriculatis setaeque longioribus differt.

 

Plants slender, dull, light green, yellow-green, reddish yellow or brownish green, in loose mats often with pendent strands. Stolons red, creeping; leaves widely spaced, reduced, erect, broadly ovate, abruptly pointed, ecostate, 0.6-0-8 mm long. Primary stems stipitate, horizontal, erect or pendent, frequently and irregularly branched, branches frequently and irregularly branched, at times with flagelliform tips, to 10 cm long, yellowish green, becoming dark red with age, in cross section with sclerodermis of 3–5 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3–4 cells long, basal 1–2 cells quadrate to subquadrate, reddish brown, upper 2–3 cells broadly cylindrical, hyaline or reddish; rhizoids only on stolons and at the base of stipitate stems, red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Secondary stem and branch apices turgid; stem and branch leaves differentiated only in size, obscurely seriately ranked when dry, distinctly so when wet, leaves panduriform, broadly oblong, appressed and clasping below, broadly obovate, abruptly flexed and spreading above, stem leaves 1.0–1.2 mm long, strongly auriculate, apex acuminate, long-cuspidate; margins broadly incurved, sharply serrulate above, plane or broadly incurved, serrulate or entire below; costae short double, short single, or absent; leaf cells long flexuose, smooth to slightly porose, 18–50 x 3–6 μm, outer basal cells shorter and broader, inner basal cells elongate, porose, alar cells subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 2–4 rows, yellow across the insertion. Dioicous. Perigonia gemmate, lateral on secondary stems 1.0 mm long, perigonial leaves ovate, acuminate, costa absent; paraphyses present, antheridia numerous. Perichaetia lateral, 2.0–2.5 mm long, paraphyses and archegonia numerous; perichaetial leaves lanceolate 1.2–2.0 mm long, vaginula hairy. Setae short, wavy, 3.5–4.0 mm long, smooth below, roughened above, yellow. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.0–2.0 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate, 1.0–1.5 mm long; annulus not seen; peristome yellowish white, exostome teeth linear, to 0.5 mm long, dorsal (outer) surface horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 10–14 μm long. Immature calyptrae, 2 mm long, cucullate, lightly hairy.

 

Etymology. The specific epithet roseana honors the collector of the type specimen Joseph Nelson Rose (1862–1928) curator at the U.S. National Herbarium. Rose collected plants in the Central and Southeastern United States, Mexico, and South America. Stafleu & Cowen (1983).

Distribution. Western South America.

Ecology. Pendent from trees; 850–2500 m.

 

Orthostechella roseana A. Habit. B. Branch leaf apex. C. Stipe leaf. D. Leaf margin. E. Leaf apex. F. Stem cross-section. G-I. Branch leaves. J. Basal leaf cells. K-O. leaves.

 

Specimens examined. ECUADOR. Azuay: Steyermark 52753 (FH, L, MO, NY); Cañar: Steere & Balslev 25952 (NY); Chimborazo: Rose 24000 (B, FH, MO, NY, S), Pallatanga, Spruce 1218 (NY); Loja: Laegaard 18475B (MO); Pichincha: sin. coll. (FH, S). PERU. Cajamarca: Soukup 3908 (FH).

 

Orthostichella roseana is a small species with well spaced, panduriform stem and branch leaves. The stem and branch leaves are also appressed, clasping below and abruptly flexed and spreading above. The stem leaves are strongly auriculate at base. Stolons are difficult to find in the few known collections of O. roseana, however, the species does appear to be stipitate.

 

Morphologically O. roseana occupies an intermediate position between O. hexasticha and O. versicolor. In size and aspect it is nearly identical to O. hexasticha, and both species have the same leaf stance. Orthostichella hexasticha differs from O. roseana in having stem leaves that are rounded to the insertion rather than strongly auriculate, and shorter (1.5–2.4 mm) setae. The sporophytes of O. roseana are identical to those of O. versicolor. Furthermore, O. versicolor has two forms (the rigida-form and pachygaster-form) that often have strongly auriculate stipe and, less often, stem leaves. Most collections of O. versicolor are larger in size than O. roseana, but there are several forms of O. versicolor that are as small or smaller than O. roseana. Orthostichella roseana consistently differs from O. versicolor in having well spaced, panduriform branch leaves that are appressed and clasping at base.

 

Orthostichella roseana is presently found in the Adean regions of Ecuador and northern Peru. There is an especially confusing form of O. versicolor from Bolivia that has the aspect of O. roseana. It differs, however, in being somewhat larger in size and in having branch leaves that are more or less spreading from the insertions.

 

6. Orthostichella versicolor (C. Müll.) Allen & Buck, Mem. New York Bot. Gard. 76(3): 140. 2003.

 

Neckera versicolor C. Müll., Syn. Musc. Frond. 2(1): 127. 1850. Orthostichella versicolor (C. Müll.) Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn. ser. 3, 2: 277. 1870, combination invalid. Pilotrichella versicolor (C. Müll.) Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 258.1877. Protologue: Brasil. Brasilia, Rio Janeiro: Beyrich, Beske. Lectotype (designated here): Brasilia, Serra d’Estrella, Beyrich (BM); isolectotype: Serra d’Estrella et Nov. Friburg, ad radices arborum, Beyrich (BM).

 

Neckera viridis C. Müll., Syn. Musc. Frond. 2(1): 125. 1850. Pilotrichella viridis (C. Müll.) Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 258. 1877. Orthostichella viridis (C. Müll.) C. Müll., Linnaea 42: 493. 1879, combination invalid. Protologue: Venezuela. Venezuela, Caracas: Moritz. Coll. I. Thorreyana Hamburgensis No. 38. Lectotype (designated here): Carcaras, leg. Moritz No. 38 (BM); isolectotypes: Caracas, Moritz 38 (L); Venezuela, Caracas, Moritz, Müller in Hb. (NY).

Neckera rigida C. Müll., Syn. Musc. Frond. 2(1): 126. 1850. Pilotrichella rigida (C. Müll.) Besch., Mém. Soc. Sci. Nat. Cherbourg 16: 222. 1872. Protologue: Mexico. Mexico: Miquel, Deppe.Lectotype (designated here): Mexiko. Deppe (S); isolectotype: Mexico, Miquel (BM).

Neckera pachygaster C. Müll., Syn. Musc. Frond. 2(1): 126. 1850. Pilotrichella pachygaster (C. Müll.) Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 256. 1877, syn nov. Protologue: Venezuela. Venezuela, in montibus nivosis prov. Meridae: Moritz. Lectotype (designated here): Caracas. leg. Moritz 171. Neckera pachygaster CM., Columbien (Merida 1844/45) Moritz (BM); isolectotypes: Columbien, Merida, 1844/45, leg. Moritz 171 (H), Merida, Moritz 171 (L).

Neckera pachygaster C. Müll.var. gracilis C. Müll., Bot. Zeitung (Berlin) 15: 582 1857. Pilotrichella pachygaster var. gracilis (C.Müller) Paris, Index Bryol. 948 1897. Protologue: Colombia. Nova Granada, S. Martha, Mina, 4000 ped. alta: Funck et Schlim in Coll. Linden No. 1034. Lectotype: Nova Granada, S. Martha, Mina, 4000 ped., leg. Funck & Schlim (BM). Probably isolectotype: Voyage de L. Schlim. N^lle Grenade, prov. de Santa Marta. Hauteur 4000 fts. Minca, 1846 à 1852 fl. en Mai, N^o 1043 (L).

Pilotrichella mexicana Schimp. ex Besch., Mém. Soc. Sci. Nat. Cherbourg 16: 223. 1872. Protologue: Mexico. Mejico (F. Müller, Schimper comm.). Holotype: Herbarium Mexicanum, A°. 1853. Fred. Müller. Comm. H. Schlumber. [Herb. Bescherelle] (BM); isotype: Orizaba, lg. F. Müller (BM).

Pilotrichella pulchella W. P. Schimper ex Bescherelle, Mém. Soc. Sci. Nat. Cherbourg 16: 222 1872. Protologue: Mexico. Orizaba (F. Müller, Lorentz mihi comm.). Holotype: Mexico. Orizaba, 1853, legit F. Müller [Herb. Bescherelle] (BM); isotype: Mexico, Orizaba, leg. Müller 1853 (H).

Neckera ampullacea Hampe ex C. Müll., Linnaea 40: 264. 1876. Pilotrichella ampullacea (C. Müll.) Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 256. 1877. Protologue: Comoro Islands. Comoro – insula Johanna, ubi primum legit Prof. Peters Berol. 1843; inter alios muscos misit J. M. Hildebrandt 1875. Lecotype (designated here): Africa: Comoro – insula Johaña. leg. Hildebrandt, 1875 (H).

Pilotrichella caldensis Ångstr., Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 33(4): 34 1876. Protologue: Brazil. Regnell n. 3 a misit. Type. Brasilia, Rio de Janeiro, leg. A. F. Regnell 3a (Holotype S, isotypes H, S).

Pilotrichella pachygastrella C.Müller ex Ångstr., Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 33(4): 33. 1876. Protologue: Brazil Regnell No 3b et Henschen. Lectotype (designated here): Brasilia, A. F. Regnell n^o 3^b (S); isolectotype: Caldas, Regnell n^o 3^b (H, S). Syntype: Caldas Brasiliae, S. Henschen (BM, S).

Neckera cyathipoma C. Müll., Linnaea 42: 414. 1879. Pilotrichella cyathipoma (C. Müll.) Kindb., Enum. Bryin. Exot. 28. 1888. Protologue: Argentina. Argentinia subtropica, Rio seco prope Sn. Andrés, cum N. Avellandae saepius commixta, fertilis 19 Septbr. 1873. — Cuesta de Buyuyu, 15 Majo 1873, sterilis; in alia cuesta Boliviae sterilis; Jujui; Cuesta de Sn. Diego, 12. Junio 1873 fertilis; Cuesta de Sn. Rosa; ubique rarius fructificans. Quoque in Cuesta colorada inter Sn. Luis et Amareta Boliviae,14 Junio 1873. Types: Argentinia subtropica, Rio seco prope St. Andrés, cum N. Avellandae saepius commixta, 19 Septmb. 1873, P. G. Lorentz (lectotype designated here JE); Argentinia subtropica, Rio seco prope San. Andrés, cum N. Avellandae saepius commixta, 19 Sept. 1873, leg. P. G. Lorentz (isolectotype H); Argentinia subtropica. Cuesta de Buyuyu. P. G. Lorentz (syntype FH, H).

Pilotrichella inflatifolia C. Müll., Flora 69: 282. 1886. Protologue: São Tomé. Bom Successo, 1050–1100 met. alta, ad arbores, [Adolf Moller]. Types: V. afrika, S. Thomé: Moller, ded. C. Müller 1889 (S, lectotype designated here); Ins. S^t. Thome (1100 m. alt). Legit A. Moller (S, isotypes).

Pilotrichella leptoclada C.Müll., Flora 69: 282 1886. Protologue: São Tomé. Encostas do Pico de S. Thomé, alt. 1500–2100 met., ad arbores, [Adolf Moller]. Types: Africa, S. Thomé: Moller, ded. C. Müller 1889, (S, lectotype designated here); Vestafrica, S. Thomé, Moller 12, com. N. C. Kindberg, ex hb. C. M. 1889, (H, isolectotype); Afr. occ. ins. S. Thomé, 1885, leg. A. Moller (NY, isolectotype).

Pilotrichella guineensis Ångstr. ex C. Müll., Flora 69: 282. 1886. Protologue: ex Guinea. Type: Guinea, ad Afzelius (S, lectotype designated here).

Neckera moenkemeyeri C. Müll., Flora 69: 516. 1886. Pilotrichella moenkemeyeri (C. Müll.) Kindb. Enum. Bryin. Exot. 102. 1891. Pilotrichella maenkemeyeri Kindb. ex Par., Index Bryol. 952. 1897, orthographic variant. Protologue. Nigeria. Africa occid. tropica, territorium fluminis Niger, Old-Calabar, ad arborum truncos, 10 Novbr. 1883: W. Mönkemeyer. Types : Nigeria. Niger-gebiet, Alt-Calabar ad arborum truncos. Fl. Afr. trop. occ. 11/10 1884, leg. W. Mönkemeyer (JE, lectotype designated here, H, S, isolectotypes); Nigeria. Niger fl. Alt Calabar, ad arbor. truncos, 1884, W. Mönkemeyer (JE, isolectotype), syn. nov.

Pilotrichella obovata Kiaer in Wright, J. Bot. 26: 266. 1888, invalid name lacks a description. Based on: Madagascar, Mt. Ankaratra (Borgen 28). Musci Madagascarienses, Herb. Kiaer. In montibus Ankaratra 1877–1879 legit M. Borgen No. 28 (L).

Orthostichella profusicaulis C. Müll., Flora 73: 493. 1890., invalid name. genus invalid. Pilotrichella profusicaulis (C. Müll.) Par., Index Bryol. 948. 1897, combination invalid. Protologue: Patria Africa or. trop. Ugueno, in regione meridionali montis Kilima-Ndscharo, 1500 m altitudinis: Dr. Hans Meyer 1889. Based on: Africa, Ugueno, Kilimandscharo, 1500 m, 1889, Hans Meyer, com. C. Müller 1890 (S).

Pilotrichella grimaldii Ren. & Card., Bull. Soc. Roy. Belgique 30(2): 192. 1891 [1892]. Protologue: Madagascar: Diego Suarez (Chenagon). Types: Madagascar, circa Diego Suarz, leg. Chenagon, Musci Mascareno-Madagascarienses N^o 39, Herb. F. Renauld, (S, isotype); Diego Suarez – Madagascar, leg. Cap. Chenagon (H, isotype); Madagascar: Diego Suarez, leg. Chenagon 1890, Herb. F. Renauld, (S, isotype).

Pilotrichella subpachygastrella Broth., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 21 Afd. 3(3): 46. 1895. Protologue. Brazil. Prov. S. Paulo, ad arbores montis Serra de Caracol (Mosén n. 428). E. Brasilia sine loco designato sub n. 24 misit amicissimus J. Cardot. Lecotype (designated here): Brasilia, prov. S. Paulo, ad arbores montis “Serra de Caracol” 15/8/1873, Hj. Mosén (H); isolectotype: Brazil. Prov. S. Paulo ad arbores montis Serra de Caracol. 18¹⁵/₈73. H. Mosén 428 (S).

Pilotrichella communis C. Müll. ex Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 28. 1895. Orthostichella communis (Dusén) C. Müll. in Paris, Index Bryol. 943. 1897, invalid name cited in synonymy. Protologue: Cameroon. Habitat in Camerunia, ubi ad Isangille (Oran) pagum m. Martio a. 1892 c. fr. parum evolutis legi [Dusén]. Type: Africa occ. Camerunia. Isangille pagum in ramis arboreus. Martii 1892, P. Dusén 651 (S, holotype). Musci Africani in Camerunia a P. Dusén collecti 651. Ad Isangille pagum in ramulis arborum die 23 m. Martii a 1892 (S, NY, isotypes), syn. nov.

Pilotrichella gracilicaulis C. Müll. ex Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 30. 1895. Orthostichella gracilicaulis (Dusén) C. Müll. in Paris, Index Bryol. 943. 1897, invalid name cited in synonymy. Protologue: Cameroon. Habitat in Camerunia in ramulis arborum, ubi ad Bomanam pagum c. 670 metra supra mare m. Julio a. 1892 c.fr. et inter Bibundi pagum et Bomanam pagum c. 300 metra supra mare m. Decembri a. 1890 c.fr. et Januario a 1892 c.fr. legi [Dusén]. Types. Musci Africani in Camerunia a P. Dusén collect. In montibus Camerunensibus prope Bomanam pagum c. 600 [sic] metra supra mare in truncis arborum die 19 m. Julio a. 1892, Dusén 164 (S, lectotype designated here). Musci Africani in Camerunia a P. Dusén collecti. Ad Bomanam pagum in ramulis. 19 July 1892 (S, syntype). Musci Africani in Camerunia a P. Dusén collect. In montibus Camerunensibus prope Bomanam pagum c. 620 [sic] metra supra mare in ramulis arborum m. Julio a. 1892, Dusén 164 (S, syntype). Musci Africani in Camerunia a P. Dusén collecti, Bibundi 5 January 1892 (S, syntype).

Pilotrichella turgidellacea C. Müll. ex Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 31. 1895. Orthostichella turgidellacea (C. Müll. ex Dusén) Buck, Bryologist 97: 435. 1994. Protologue: Cameroon. Habitat in Camerunia in ramis ramulisque arborum , ubi ad Bomanam pagum c. 670 metra supra mare m. Decembri a 1890 et supra Etome pagum c. 800 metra supra mare m. Januario a. 1892 legi [Dusén]. Types. Musci Africani in Camerunia a P. Dusén collecti. In montibus Camerunensibus ad Bomanam pagum c. 670 metra supra mare in truncis arobrum die 18 m. Decembris a. 1890, [Dusén] 131, P Duséns mossherbarium (S, lectotype designated here, S isolectotype); V. Afrika, Kamrum, Bomana, 18/12/90, P. Dusén (S, isolectotype); Africa occ., Camerunia prope Etome pagum in ramulis, Januario a. 1892, P. Dusén (S, syntype).

Orthostichella filamentosula C. Müll., Bull. Herb. Boissier 5: 204. 1897. Pilotrichella filamentosula (C. Müll.) Par., Index Bryol. Suppl. 271. 1900. Protologue: Guatemala. Sine loco speciali quam linteum involucris serviens (Packmaterial) in Museo zoologico Florentino 1892: Hb. Levier. Types: e viciniis urbis Guatemala. rudem accepit Mus. Zoolog. Florent. quam involucrum animalium, 1892, legit indigenus (lectotype designated here BM); e viciniis urbis Guatemala. rudem accepit cum pellibus animalium Mus. Zool. Florentinum, 1892, collector ignotus (isolectotype NY, H).

Pilotrichella subimbricata (Hampe) Jaeg., var. flageyi Ren. & Card. In Ren., Prodr. Fl. Bryol. Madagascar 196. 1898. Orthostichopsis subimbricata (Hampe) Broth. var. flageyi (Ren. & Card.) Card., Hist.Phys. Madagascar, Mousses 345. 1915. Protologue. Madagascar: Diego Suarez, Chenagon, 1890; Andrangoloaka, Sikora, 1892. Type: Madagascar, Diego Suarez, leg. Chenangon (S, lectotype, designated here), syn. nov.

Pilotrichella subpanduraefolia Par., Index Bryol. Suppl. 272. 1900, new name for Pilotrichella panduraefolia C. Müll. ex Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 32. 1895, illegitimate homonym not Pilotrichella panduraefolia (C. Müll.) Jaeg, 1877. Protologue: Habitat in Camerunia in truncis ramulisque arborum, ubi in montibus Rumpi ad Tokko pagum c. 900 metra supra mare m. Aprili a. 1892 et in montibus Camerunensibus c. 300 metra supra mare m. Januario a. 1892 legi [Dusén]. Types. Musci Africani in Camerunia a P. Dusén collecti. In montibus “Rumpi” ad Tokko pagum c. 900 metra supra mare in truncis arborum die 16 m. Aprilis a. 1892 [Dusén] 859 (S, lectotype; L, S isolectotypes); In montibus Camerunensibus c. 300 metra supra Bibundi pagum, in ramulis arborum m. Januario a. 1892, P. Dusén (S, syntypes).

Orthostichella microcarpa C. Müll., Hedwigia 40: 86. 1901. Pilotrichella microcarpa (C.Müll.) Broth., Nat. Pflanzenfam. I(3): 811 1906. Protologue: Brazil. Brasilia, Sa. Catharina, Serra Geral, in declivibus Serrae ad ramos arborum., Febr. 1890: E. Ule, Coll. No. 866. Type: Brasilia, S. Catharina, Serra Geral, auf Baum zweigen am abhang der Serra do Oratorio, 2/1890, leg. E. Ule 866 (lectotype designated here H).

Orthostichella mucronatula C. Müll., Hedwigia 40: 87. 1901. Pilotrichella mucronatula (C. Müll.) Broth., Nat. Pflanzenfam.  I(3): 811 1906. Protologue: Brazil. Brasilia, Sa. Catharina, Tubarão, in sylva ad ramulos arborum prope Conconhaz, Sept. 1889; E. Ule, Coll. No. 778; ad flumen Laranjeiras superius prope Orleans in Sierra Geral, Sept. 1889; idem, Coll. No. 779; Nova Venezia, ad truncos arborum sylvestrium, Julio 1891: idem, Coll 1169. Lectotype designated here: Brasilia, prov. S. Catharina, Tubarão, auf Baumzweigen in Wäldehen bei Conconhaz, Sept. 1889, leg E. Ule 778 (H); syntypes: An Bäumen am oberen Larenjeiras bei Orleans, Estado de Sta. Catharina: September 1889, E. Ule 779 (JE); S^ta Catharina, in arboribus ad flumen Laranjeiras prope Oreans in Serra Geral, leg. E. Ule 779 (JE); Brasilia, prov. S. Catarina, Orleans auf Baumen am oberen Laranjeiras, Sept. 1889, leg E. Ule 779 (H); An Baumstämmen im Walde bei Nova Venezia, Estado de Sta. Catharina: Juli 1891, E. Ule 1169 (BM, S); Brasilia, prov. S. Catarina, Nova Venezia, am Baumstämmen im Walde. Juli 1891, leg E. Ule 1169 (H).

Pilotrichella subpachygastrella var. minor Broth. in Bauer, Verh. Zool.-Bot. Ges. Wien 55: 579. 1905, invalid name, lacks a description. Based on: Brazil. Canóas, auf Baumrinde in Wädern [Brazil, Porto Alegre], Ed. M. Reineck & Jos. Czermak (B, BM, FH, JE, S).

Pilotrichella attenuata Broth., Denkschr. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl. 88: 739 1913. Protologue: Tanzania. Deutsch-Osafrika: West -Usambara, Wald unterhalb Mazumbei, an Bäumen, zirka 1200 m ü. M., 18./VIII. 1909, [Brunnthaler]. Holotype: West-Usambara, Wald unter Mazumbi, an Bäumen, 1200 m, 18/8 1909, leg. J. Brunnthaler.

Pilotrichella angustifolia Herz., Biblioth. Bot. 87: 112. 1916, syn nov.

Protologue: Bolivia. Ad Baumästen in der Talschlucht bei Tablas, ca. 1800 m, [Herzog] 4666. Type: An Baumästen in der Talschlucht bei Tablas, ca. 1800 m., Mai 1911, leg. Th. Herzog 4666 (holotype JE; isotypes H, L, S).

Pilotrichella cyathipoma var. laxiretis Herz., Biblioth. Bot. 87: 112. 1916, syn. nov.

Protologue: An Bäumen im Wald des Randgebirges bei Yacuiba, ca. 500 m., [Herzog] No. 2626. Type: Bolivia. Im Wald des Randgebirges bei Yacuiba, ca. 500 m., Oktober 1910, Th. Herzog No. 2626 (holotype JE, isotypes H, L, S).

Pilotrichella allionii Broth., Rev. Bryol. 47: 14. 1920 [1921], syn nov.

Protologue: Ecuador. Prov. de l'Oriente. Gualaquiza; in arboribus ad ripam fl. Bomboiza prope ejus confluentiam cum fl. Gualaquiza; 800–900 m [M. Allioni]. Gualaquiza; in silva El Salado; 950 m [M. Allioni]. Types: Ecuador - Oriente - V. Bomboiza. Ad arborum ramos in silva “El Salado” fregnens sed permisita Gualaquiza m 950, 25 Aug. 1910, M. Allioni 497 (H, lectotype designated here). Ecuador., prov. Azuay. Gualaquiza: ad ripam flum. Bomboiza prope ejus confluentiam cum fl. Gualaquiza, in arboribus, Jul. 1909, legit Rev. M. Allioni (H, syntype); V. Boimbizo, Ad truncos in sylva El .Salado, 950 m, 25 Viii 1910, M. Allioni (PC, isolectotype).

Pilotrichella rigens Broth., Denkschr. Kaiserl. Akad. Wiss., Math.- Naturwiss. Kl. 83: 309 1926, illegitimate homonym. Not Pilotrichella rigens Card., 1910. Protologue: Brazil. São Paulo. Prope Raiz da Serra; ad arbores; 20–50 m s. m. ([Schiffner] 695 p.p.). Prope Salto Grande do Rio Paranapanema; ad arbores; ca. 500 m s. m. ([Schiffner] 1716, 1718). In ripa sinistra fluminis Paranapanema ad cataractas “Salto Grande”; ad arbores ([Schiffner] 1795). Prope “Fazenda Bella Vista” in districtu urbis S.Cruz ad flumen Rio Pardo; ad ramulos; ca. 500 m s. m. ([Schiffner] 1937). In itinere Cerqueira Cesar–Frazenda “Bella Vista”; ad arbores ad flumen Rio Turvo; 500 m s. m. ([Schiffner] 1183). Apud cataractas “Salto dos Treis Ranjos” prope urbem Cerqueira Cesar; ad arbores; ca. 500 m s. m. ([Schiffner] 493). In silvaticis inter Faxina et Apiahy prope Lagoas ([Schiffner] 97). Apiahy, Catos Altas da Ribeira (Puiggari 1986). Ad viam inter Apiahy et Yporanga (Puiggari 48). Syntypes: Prov. Paraná: In ripa sinistra fluminis Paranapanema ad cataractas “Salto Grande”, ad arbores, 27.VII.1901, lgt. V. Schiffner 1795 (BM, H); São Paulo. Prope “Fazenda Bella Vista” in districtu urbis S.Cruz ad flumen Rio Pardo, ca. 500 m s. m., ad ramulos, 19.VII.1901, lgt. V. Schiffner 1937 (BM, H, S); São Paulo. Apud cataractas “Salto dos Treis Ranjos” prope urbem Cerqueira–Cesar. ca. 500 m s. m., ad arbores, 21.VII.1901, lgt. V. Schiffner 493 (BM, H, S); São Paulo. Prope Salto Grande do Rio Paranapanema. ca 500 m.s.m., ad arbores, 25.VII.1901, lgt. V. Schiffner 1716 (BM, H); São Paulo. Prope Salto Grande do Rio Paranapanema; ad arbores; ca. 500 m s. m. 25. VII. 1901, lgt. Schiffner 1718 (BM, H, S); São Paulo. In itinere Cerqueira Cesar–Frazenda “Bella Vista”. Ad flumen Rio Turvo, ad arbores, 500 m s. m., 22 VI 1901, lgt. V. Schiffner 1183, (BM, H, S); São Paulo, Apiahy, chemin d’ Yporanga, 7/1880, leg. J. J. Puiggari 48 (H); Brasilis, São Paulo, Apiahy, Catos Altas da Ribeira, 1/9/1881, leg. J. J. Puiggari 1986 (H); São Paulo. In silvaticis inter Faxina et Apiahy. ca. 800 m.s.m. pr. Lagoas, 23.VIII.1901, lgt.. V. Schiffner 97 (H); Brazilia. Prov. São Paulo. Prope Raiz da Serra; 20–50 m.s.m., ad arbores; 4.VI.1901, lgt. V. Schiffner 695 (BM, H).

 

Nomenclatural notes

1. The protologue of Neckera moenkemeyeri C. Müll gives the year of the type collection as 1883 while the types in H, JE, and S give the year of collection as 1884. According to Vegter (1976), Mönkemeyer collected in southern Nigeria and Fernando Po between 1884–1885.

 

2. There is a specimen of in L that appears to be an isolectotype of Neckera pachygaster var. gracilis C. Müll. The collection bears the orginal label from the “Voyage de L. Schlim”, but differs from the protologue in two respects: its collection number is No. 1043 rather than No. 1034, and the town is given as Minca rather than Mina. The L collection is considerably larger than the BM collection, and the plants in both collections are identical.

 

3. None of the potential type collections of P. gracilicaulis in S exactly fit the protologue information for that species. There is one collection, marked “TYPUS” that contains a label in Dusén’s handwriting which is also marked in red “sp. nov!”, but this collection was made on 21 January 1891 at 700 meters. The only specimen clearly marked as from the Dusén herbarium is chosen here as the lectotype.

 

Plants slender to medium-sized, dull, light green, yellow-green, reddish yellow or brownish green, in loose mats often with pendent strands. Stolons yellow to red, creeping; leaves widely spaced, reduced, erect, ovate-triangular, abruptly acuminate, ecostate, 1.0–1.4 mm long. Primary stems at times stipitate or foliose to the stolons, horizontal, erect or pendent, frequently and irregularly branched, branches at times with flagelliform tips, to 4–10(–40) cm long, yellowish green, becoming dark red with age, in cross section with sclerodermis of 3–5 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3–4 cells long, basal 1–2 cells quadrate to subquadrate, reddish brown, upper 2–3 cells broadly cylindrical, hyaline or reddish; rhizoids consistently on stolons, also at base of stipitate secondary stems, or on flagelliform branch tips, red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Stipe leaves ovate, cordate, rounded at base or straight to the insertions. Secondary stem and branch apices turgid; stem and branch leaves differentiated only in size, stem leaves not or obscurely seriately ranked when dry, branch leaves often distinctly seriately ranked, dry or wet, leaves obovate to oblong-obovate, erect to erect-spreading, variously clasping below, 1.0–1.6 mm long, at times somewhat auriculate, apex cuspidate; margins broadly incurved and sharply serrulate above, plane or broadly incurved, serrulate or entire below; costae mostly absent, occasionally short double or short single; leaf cells long flexuose, smooth, upper cells 20–30 x 4–6, median cells 30–60 x 3–4 μm, outer basal cells shorter and broader, inner basal cells near insertion elongate, often porose, alar cells red to red-yellow, subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 4–6 rows, yellow across the insertion. Asexual reproduction by deciduous leaves common. Dioicous. Perigonia gemmate, lateral on secondary stems and branches, 0.8–1.2 mm long, perigonial leaves ovate, acuminate, costa absent; paraphyses present, antheridia numerous. Perichaetia lateral, 1.4-2.0 mm long, paraphyses and archegonia numerous; perichaetial leaves lanceolate 1.4–2.0 mm long, vaginula moderately hairy. Setae short, wavy, 2.5–7.0 mm long, smooth below, roughened above, yellow, becoming yellowish red with age. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.0–2.0 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate, 1.0–1.5 mm long; annuli rudimentary, consisting of 2–3 rows of thin-walled enlarged cells; peristome yellowish white, exostome teeth linear, to 0.5 mm long, dorsal (outer) surface lightly horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 14–20 μm long. Calyptrae, 2–3 mm long,narrowly cucullate, smooth to lightly hairy.

 

Etymology. The specific epithet vesicolor is a Latin adjective meaning “variously colored.” It probably refers to the fact that plants of O. veriscolor occur in various combinations of green, yellow, red, and brown.

Distribution. Mexico; Central America; Caribbean; Western, Northern, and Southern South America, Brazil; West and East Tropical Africa; Western Indian Ocean.

Illustrations. Herzog (1916, Fig. 41 a–b); Bartram (1949, Fig. 118 A–C); Florschütz (1964, Fig. 100); Churchill (1994, Fig. 27 C); Sharp et al.(1994, Fig. 537); Churchill (1995, Fig. 124 a–g); Buck (1998, Pl. 91 9–17); Gradstein et al. (2001, Fig. 156 A–I); Buck (2003, Fig. 126).

 

Orthostichella versicolor A. Branch with sporpohyte. B. Stem leaf. C. Branch leaf apex. D-F & K-L. Branch leaves. G. Median leaf cells. H. Peristome teeth and segment. I. Sporophyte with operculum. J. Basal leaf cells. M. Stem cross-section.

 

Ecology. On the trunks, branches, and twigs of trees, vines, shrubs, logs, leaves, and tree fern fronds, occasionally on moist rocks along rivers and boulders along roadsides and dry slopes; 50–2600 m.

 

Selected specimens examined.

MEXICO. Baja California Sur: Brandegee (NY); Chiapas: Laughlin 540 (MO); Colima [Socorro Island]: Felger 15853 (MO); Hidalgo: Magill 2317 (MO); Jalisco: Mexia 1338a (FH, MO, NY); Morelos: Pringle 15670 (FH, US); Oaxaco: Richards et al. 3801b (MO, NY, US); Puebla: Sharp 1000 (MO, NY); San Luis Potosí: Sharp 5888 (MO, NY); Sinaloa: H. S. Gentry 7283 (FH, NY); Tabasco: Ruiz 1459a (H); Tamaulipas: Manuel 333 (MO); Veracruz: Pringle 15147 (FH, MO, NY, US).

BELIZE. Cayo: Allen 18428 (BRM, MO); Toledo: Allen 19057 (BRM, MO). GUATEMALA. Alta Verapaz: Standley 91732 (FH, NY, S); Chimaltenango: Wenzel & Mitchell s. n. (MO); Chiquimula: Steyermark 31558 (FH, NY); Guatemala: Standley 80668 (FH, NY); Huehuetenango: Steyermark 49439 (FH); Izabal: Steyermark 39991 (FH, MO); Jalapa: Steyermark 32533 (FH, S); Petén: Lundell 2830 (NY, S); Quezaltenango: Standley 86702 (FH, NY, US, S); Quiché: Aguilar 915 (FH, S); Sacatepéquez: Standley 88951 (FH, NY, S); San Marcos: Croat 40907 (MO, NY); Suchitepéquez: Steyermark 46666 (FH, MO, US). EL SALVADOR. Ahuachapán: Padilla s.n. (NY). HONDURAS. Atlántida: Allen 17314 (MO, TEFH); Copán: Allen 17781 (MO); Cortés: Allen 14354 (MO, NY); Franciso Morazán: Allen 12389B (MO); Lempira: Allen 12148 (MO); Olancho: Allen 12777B (MO); Santa Bárbara: Crosby 2793 (MO); Yoro: Allen 13664 (MO). NICARAGUA. Estelí: Moreno 19249 (H, MO, NY, US, S); Granada: Almedo 1459B (MO); Jinotega: Stevens 16695 (MO, NIC); Madriz: Stevens 16137 (MO, NIC); Matagalpa: Stevens 22193 (MO); Zelaya: Schramm (FH, NY, US). COSTA RICA. Alajuela: Crosby & Crosby 5754 (BM, CR, G, H, NY, US); Cartago: Liesner & Judziewicz 14486 (MO, NY); Guanacaste: Crosby 3851 (MO, S); Hererdia: Crosby & Crosby 5780 (BM, CR, F, FH, G, H, JE, NY, S, US); Limón: Steere CR-57 (NY); Puntarenas: Lyon 511 (CR, MO); San José: Crosby & Crosby 5746 (BM, CR, F, FH, G, H, JE, MICH, NY, PC, S, US). PANAMA. Bocas Del Toro: Allen 5844 (MO); Canal Area: Bailey & Bailey 511 (NY); Chiriquí: Croat 15925 (MO); Coclé: Witherspoon 8771 (MO); Colón: Crosby 10354 (MO); Darién: Allen 8718 (MO); Panamá: Crosby 4489 (MO); Veraguas: Knapp & Sytsma 2543 (MO).

CUBA. Gramma: Ekman 5390 (FH); Pinar del Río: León 12731 (NY); Sancti Spíritus: Clement 11 (NY); Santiago de Cuba: Hamilton (NY). PUERTO RICO. Aguadilla: Stevens 1781 (NY); Arecibo: Britton & Marble 1177 (MO, NY, S); Ponce: Britton & Marble 974 (NY). DOMINICA. Elliot 639 (BM, FH).

COLOMBIA. Antioquia: Callejas, Roldán & Científico 10182 (NY); Boyacá: Amórtegui 357 (FH); Caquetá: Churchill & Betancur 16896 (H, MO, NY); Cauca: Churchill & Franco 16583 (H, MO, NY); Cundinamarca: Apollinaire (B, BM, FH, JE, L, MO, NY, S); Magdalena: Mägdefrau 1159 (B, MO); Nariño: Churchill & Rengifo 17599 (H, MO, NY, S); Risaralda: Betancur et al. 3110 (NY); Santander: Churchill, Franco & Parra 18760 (NY); Valle: Hartman 217 (MO); Vaupes: Schultes 11740 (FH). ECUADOR. Carchi: Steere 8871 (NY); Imbabura: Steere 9262 (NY); Morona-Santiago: Holm-Nielsen et al. 4632 (MO, S); Napo: Holm-Nielsen & Jeppesen 760 (MO); Pastaza: Holm-Nielsen & Jeppesen 406 (MO); Pichincha: Fransén 87 (NY, S); Tungurahua: Spruce 1197 (BM, NY); Zamora-Chinchipe: Ortega 544a (MO). PERU. Ayacucho: Cook & Gilbert 919 (NY); Cuzco: Cook & Gilbert 919 (BM); Junin: Gradstein, Bryophyta Neotropica 67 (B, H, L, MO, NY, S); San Martin: Frahm et al. 1362 (B, H, NY); Tumbes: Simpson BR-137 (MO). BOLIVIA. Chuquisaca: (Prov. Sud Cinti) Serrano 1318 (MO), (Prov. Calvo) Churchill 20693 (MO); Cochabamba: (Prov. Chapare) Churchill et al. 19857 (MO); La Paz (Prov. Inquisivi) Lewis 89-1018 (MO); Santa Cruz: (Prov. Caballero) Churchill et al. 21914 (MO), (Prov. Chiquitos) Lewis 85-1298 (US), (Prov. Florida) Churchill & Vásquez 21777 (MO), (Prov. Ibanez) Fuentes s.n. (MO), (Prov. Ichilo) Nee 38863 (MO, NY), (Prov. Vallegrande) Vargas 690 (MO); Tarija: (Prov. Arce) Lewis 84-2192 (US), (Prov. O’Connor) Lewis 79-532 (MO).

PARAGUAY. Alto Paraná: Zardini & Guerrero 39949 (MO); Caazapá: Zardini 7850B (MO, NY); Canendiyú: Zardini &bFranco 55007A (H, JE, MO, NY, US); Paraguarí: Hahn 746 (H, MO, NY, US); San Pedro: Zardini & Guerrero 42771 (MO). ARGENTINA. Jujuy: Hosseus 27/VI/33 (JE); Misiones: Hamilton s.n. (B, BM); Salta: Price et al. 1580 (MO); Tucumán: Buck 25968 (NY). URUGUAY. Montevideo: sin. coll. (NY).

VENEZUELA. Anzoátequi: Steyermark 61131 (FH); Aragua: Nee & Whalen 16883 (B, NY); Barinas: Nee & Whalen 17084 (NY); Falcon: Griffin & Wingfield PV-1648 (MO); Lara: Griffin & López 336 (B, MO); Mérida: Fendler 114 (NY); Monagas: Pursell 8960 (MO); Portuguesa: Steyermark et al. 126699 (MO); Táchira: Liesner & González 10923A (MO); Trujillo: Liesner et al 13029 (MO) SURINAME. Commewijne: Schallert (NY); Sipaliwini: Allen 25508 (B, BM, FH, H, JE, MO, NY, U, US). FRENCH GUIANA. St. Laurent-du-Maroni: Buck 25580 (NY).

BRAZIL. Federal District: Heringer 8303A (NY); Minas Gerais: Mexia 5358a (FH, MO); Paraná: Dusen 8495 (NY); Rio de Janeiro: Gradstein Bryophya Neotropica 218 (B, H, MO, NY, S); Rio Grande do Sul: Wasum et al. 2160 (MO, NY); Santa Catarina: Frahm 1652 (MO); São Paulo: Visnadi & Vital 515 (MO).

NIGERIA. Cross River: Mönkemeyer (JE).

GHANA. Capt. Benton (BM).

KENYA. Rift Valley: Bishop Harmington (BM); Kaibibich, Cherangani Hills, 1°12’N, 35°17 E, Friis 2772 (MO).

MALAWAI: Southern: Chapman & Chapman 9471 (MO).

TANZANIA. Arusha: Rojkowski 369 (B).

ZAIRE: Kivu: Hendrickx 3891 (MO).

ZIMBABWE. Inyanga: Müller 2646 (MO); Nyanga: Clarke 90 (MO).

COMORO ISLANDS. Anjouan Island: Magill & Pócs 11241 (B, BM, F, G, H, MO, NY, PC S, US); Grande Comore Island: Magill & Pócs 10924 (BM, L, FH, G, H, JE, MO, NY, S, US); Mayotte Island: Magill & Pócs 11620 (JE, MO, NY); Mohéli Island: Magill & Pócs 11729 (MO).

MADAGASCAR. Antseranana: Magill et al. 9961 (H, MO, NY); Antananarivo: Dorr & Rakotozafy 2806 (MO); Fianarantsoa: Crosby & Crosby 8642 (MO); Toamasina: Dorr 3141 (BM, FH, JE, MO, S).

 

Orthostichella veriscolor has a complex morphology with creeping primary stems and erect or pendent (some collections reported to hang 1.0–1.5 meters long) secondary stems. The secondary stems can be strongly stipitate or evenly foliate throughout. Remarkably, stipitate and evenly foliate secondary stems can sometimes be found on the same primary stem. Its branches commonly end in filiform attenuations or stolons. These different structures can be morphologically distinct, but more often they intergrade one into another or they can be abruptly transformed from one structure to another. As a rule the leaves on the branches, and sometimes the secondary stems, of O. versicolor are variously spirally ranked while those on the stolons and primary stems are unranked. But, this rule is often violated. Most leaves are cuspidate, have distinctive incurved, serrulate upper leaf margins, elongate, smooth, firm-walled leaf cells, and weakly developed, reddish yellow, firm-walled alar cells. The costa in O. versicolor is one of its oddest features. Most leaves are ecostate, however, here and there leaves can be found with short double, short single, or long single costae. Sporophytes are rarely encountered in this dioicous moss. The setae are 2–6 mm long, flexuose, and roughened above. The operculum is long, obliquely rostrate, and the peristome is pale-yellow and greatly reduced. Distinctive features of the peristome include its narrow, mostly papillose exostome teeth that weakly cross-striate at the very base, and narrow endostomial segments that are perforated but not keeled. The calyptrae are cucullate and either naked or sparsely hairy.

 

There are three major forms of O. veriscolor in the Neotropics. The common form of the species in the northern part of its range is nomenclaturally centered on Neckera rigida C. Müll. (Mexico) and Neckera pachgaster C. Müll. (Venezuela). In this form the plants are often turgid, distinctly stipitate, and the leaves are weakly ranked. The secondary stem leaves are strongly auriculate and deciduous branch leaves are not generally encountered. Throughout the southern part of its range the common form of the species has more slender plants that may or may not be stipitate. The secondary stem leaves are not or weakly auriculate, the branch leaves are distinctly ranked, and deciduous branch leaves are frequently encountered. The non-stipitate plants of this southern form are nomenclaturally centered on P. veriscolor C. Müll. (Brazil), while the stipitate plants are nomenclaturally centered on P. rigens Broth. (Brazil). Throughout the range of the species there are even more slender expressions, that lack stipitate secondary stems, have more or less unranked, non-auriculate stem leaves, and non-deciduous branch leaves. The oldest name at the species level for this expression is Pilotrichella pulchella Schimper ex Bescherelle (Mexico), but it had earlier been described as Neckera pachygaster var. gracilis C. Müll. (Venezuela).

 

Orthostichella versicolor is primarily an epiphytic species. It is occasionally collected on boulders, but field observations on plants in this habitat point to epiphytic plants that have been able to maintain growth after falling from the canopy. Collections taken from twigs tend to have plants small with weakly ranked leave, while those from tree trunks often consist of big plants with strongly ranked leaves. In some large collections that have been carefully collected and so show all parts of the population every one of the morphological expressions of the species can be found.

 

This species has generally been known as Pilotrichella pentasticha (Brid.) Wijk & Marg. or Orthostichella pentasticha (Brid.) Buck every since Robinson (1967) chose to use the oldest specific epithet (Hypnum pentastichum Brid.) for this species complex. Hypnum pentastichum was based on a specimen of unknown origin sent to Bridel by Philibert Commerson (1726–1773). This specimen is not in B, however, Marshall Crosby recently located the specimen at PC. The PC collection was made by Commerson on Mauritius in 1769 or 1770, and unfortunately it is not an Orthostichella but rather belongs to the genus Orthostichidium (Orthostichium pentastichum (Brid.) Allen, comb. nov. Basionym: Hypnum pentastichum Brid., Muscol. Recent. 2(2): 100. 1801). The second oldest name for this species complex is Hypnum imbricatum (P. Beauv.) which is based on a M. Bory-St. Vincent collection from Réunion. I have not been able to locate the type of this name. It is possible that it too is an Orthostichidium or even the same as O. pentastichum.

 

Orthostichella versicolor can be confused in the field with Squamidium nigricans (Hook.) Broth., Orthostichopsis tenuis (Jaeger) Broth., Orthostichopsis praetermissa Buck) or Orthostichidium quadrangulare (Schwaegr.) Allen because they all are pendent mosses with distinctly ranked leaves. Squamidium nigricans, Orthostichopsis tenuis, and O. praetermissa differ from Orthostichella versicolor in having a long single costae. Squamidium nigricans also has well-developed alar cells and often a black coloration, while the two Orthostichopsis species have filamentous pseudoparaphyllia, densely hairy calyptrae and vaginula, and cylindrical capsules. Orthostichopsis praetermissa further differs from Orthostichella versicolor in having hair-pointed leaves. Orthostichella versicolor can often be distinguished from these three species with a hand lens beause of its branches that frequently end in filiform attenuations and have deciduous leaves.

 

Orthostichidium quadrangulare is especially close to Orthostichella versicolor since both species have ecostate leaves with broadly incurved upper leaf margins and weakly developed alar cells. Plants of O. quadrangulare tend to be a little bit larger than those of Orthostichella versicolor, and its leaves are more stiffly erect as well as more strongly ranked. However, the only certain way to separate Orthostichidium quadrangulare from Orthostichella versicolor is to examine their branch primordia. In Orthostichidium quadrangulare the primordia often give rise to clusters of uniseriate gemmae and they are surrounded by filamentous pseudoparaphyllia. Orthostichella versicolor never has asexual gemmae, and the branch primordia are surrounded by scale leaves.

 

7. Orthostichella welwitschii  (Duby) Allen & Magill, comb. nov.

 

Neckera welwitschii Duby, Mém. Soc. Phys. Genève 21: 428. 1872. Meteorium welwitschii (Duby) Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 243. 1877. Pilotrichella welwitschii (Duby) Gepp in Hiern., Cat. Afr. Pl. 2(2): 292. 1901. Protologue: Angola. Frequenter in ramulis extremis variarum arborum pendulam in sylvis primitivis 2300–2400 ped. alt. montis Campolungo prov. Golungo-alto regni Angolensis reperiit cel. Welwitsch. Type. Welwitsch Iter Angolense, Dist. Golungo; 2400 feet, (S, isotype); Musci Angolensis N° 145. Distr. Gol°-alto, ad 2800 ped., ad truncos vituntos Coffee arab. indigense, in monte Campolungo freq. sine fr. Febr. 1855, leg. Welw. (H, isotype); Angola: distr. Golungo-Alto. Leg. Welwitsch, 1855. n° 145 (PC, isotype).

 

Pilotrichella sordidoviridis C. Müll. ex Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 27. 1895., Protologue: Habitat in Camerunia in ramulis arborum, ubi ad Bibundi emporium m. Julio a. 1892 c. fr. legi [Dusén]. Types: Africa occ., Camerunia ad Bibundi emporium in ramulis, Julio 1892, P. Dusén [P. Duséns mosssherbarium] (S, lectotype); Musci Africani in Camerunia. Ad Bibundi emporium in ramulis, 21/7 1892, P. Dusén (FH, S, isotypes); Musci Africani in Camerunia. Ad Bubundi pagum in truncis arborum m. Julio a. 1892, P. Dusén 829 (S, isotype).

Pilotrichella dewrevrei Ren. & Card., Bull. Soc. Roy. Bot. Beligique 38(2): 72. 1899. Pilotrichella sordido-veridis Dusén var. dewrevrei (Ren. & Card.) Ren. & Card., Bull. Soc. Roy. Bot. Beligique 38(1): 25. 1900. Protologue: Congo sine loco (Alfr. Dewèvre). Type: Congo belge. Leg. DeWèvre [Herb. J. Cardot] (PC, holotype).

Pilotrichella pilifolia Dixon, Smithsonian Misc. Collect. 69(8): 8. 1918. Prtologue. Uganda. Epiphytic; pendent in forest, Mabira, near Mubango, alt. 4,000 ft, July, 1916, Dümmer 2961; c. fr. Type: On tree trunks, pendent. Mabira Forest, Mubango, 4000 ft, Mardh 20, Drummer 2961 (FH, isotype).

 

 

Plants slender to medium-sized, dull, light green, yellow-green, reddish yellow or brownish green, in loose mats often with pendent strands. Stolons creeping, leaves well spaced. Primary stems foliose, horizontal or pendent, frequently and irregularly branched, branches often with flagelliform tips, 2–15 cm long, yellowish green, becoming dark red with age, in cross section with sclerodermis of 2–5 small, thick-walled, reddish orange cells, cortex cells enlarged, firm-walled, pale yellow, red with age, central strand absent; paraphyllia absent, pseudoparaphyllia absent, scale leaves present; axillary hairs 3 cells long, basal 1 cell quadrate to subquadrate, upper 2 cells broadly cylindrical, all cells reddish; rhizoids on stolons and on primary stems, red, not or irregularly branched, from clusters of initials abaxial to the leaf insertions. Primary stem and branch apices turgid; stem leaves not seriately ranked, ovate, erect, variously clasping below, 1.4–2.0 mm long, rounded to the base, abruptly narrowly acuminate, to hair-pointed; margins broadly incurved and subentire to serrulate above, plane or broadly incurved, serrulate or entire below; costae absent or very shortly single or double or short single; leaf cells long flexuose, smooth, upper cells 30–44 x 4–5, median cells 40–60 x 3–4 μm, outer basal cells shorter and broader, inner basal cells near insertion elongate, often porose, alar cells, subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 4–6 rows, yellow across the insertion. Branch leaves often distinctly seriately ranked, dry or wet, leaves obovate to oblong-obovate, often panduriform, erect to erect-spreading, variously clasping below, 0.9–1.5 mm long, rounded to the insertion, apex apiculate to short cuspidate, cuspid 40-80 µm long; margins broadly incurved and subentire to serrulate above, plane or broadly incurved, serrulate or entire below; costae very short, double, single, occasionally absent; leaf cells long flexuose, smooth, upper cells 20–40 x 4–5, median cells 30–60 x 3–4 μm, outer basal cells shorter and broader, inner basal cells near insertion elongate, often porose, alar cells red to red-yellow, subquadrate to short-rectangular extending up the margins in 2–4 rows and across the insertion in 4–6 rows, yellow across the insertion. Dioicous. Perigonia not seen. Perichaetia lateral, unfertilized perichaetia 1.0–1.5 mm long, paraphyses and archegonia numerous; fertilized perichaetial 2.0–2.5 mm long, leaves lanceolate 2–3 mm long, vaginula hairy. Setae wavy, 3–5 mm long, smooth below, roughened-papillose above, yellow, becoming yellowish red with age. Capsules exserted, erect to somewhat inclined, ovoid to short-cylindrical, 1.0–1.5 mm long, neck moderately developed; exothecial cells small, rounded in 2–3 rows below the mouth, subquadrate, irregularly short-rectangular, or oblate, firm-walled below, cells in neck smaller than the median cells; stomata superficial on neck; opercula obliquely long-rostrate, 1.0 mm long; annuli rudimentary, consisting of 2–3 rows of thin-walled enlarged cells; peristome yellowish white, exostome teeth linear, 0.3–0.4 mm long, dorsal (outer) surface lightly horizontally striate at base, smooth or finely papillose above, trabeculae weakly developed on both sides, endostome nearly as long as exostome, basal membrane very low, segments filamentous, narrowly perforated, not keeled, smooth to papillose, cilia rudimentary or absent. Spores round, lightly roughened, 18–24 μm long. Calyptra not seen.

 

Etymology. The specific epithet welwitschii is given in honor of its collector, Friedrich Martin Josef Welwitsch (25 Feb 1806–20 Oct 1872), an Austrian botanist. Welwitsch collected plants in Angola (1853–1861) for the Portuguese government, and during that time also collected in other parts of Southwest Africa (Stafleu & Cowan 1988, Frahm & Eggers 2001, includes portrait).

 

Illustrations: Dusén (1895, .p. 27 a--c, Pl. 2, fig. 19, Pl. 4 fig. 3)

Habitat: On tree trunks and branches; 852–1219 m.

Distribution. West, West Central, East, and South Tropical Africa.

Selected specimens examined.

ANGOLA. Dist. Golungo: Welwitsch (S).

CAMEROON. Bubundi, Dusén 829 (S).

IVORY COAST. In silva Akrèdiou, inter Thiassale et Dabou, A. Jolly (B).

TANZANIA. Stromgebiet des oberen Ruhudje, Landschaft [District] Lupembe, nordlich des Flusses, Schlieben 726 (B).

ZAIRE. Dewèvre (PC).

 

Orthostichella welwitschii is a small to medium-sized species with extremely long hair-pointed stem leaves. The species appears to lack stipitate primary stems, but this may be an artifact due to the few collections from which the species is known. The stem leaves of P. welwitschii are more or less similar in form to those of O. pandurifolia, however, they are not as consistently costate and its branch leaves are identical to those of O. versicolor. In terms of overall morphology O. welwitschii appears most similar to O. versicolor. Unfortunately the pendent stems of O. versicolor often have hair-pointed leaves and this is likely cause considerable difficulty in separating the two species. The stem leaves of O. welwitschii, however, consistently have extremely long hair-points and often the hair-points are kinked. This feature is especially pronounced in the stem buds. In most collections of O. versicolor short-pointed stem leaves can be found. There are included in this treatment of O. welwitwschii a number of small slender collections that consistent entirely of pendent stems and branches. These collections have hair-pointed stem leaves, but they may actually represent pendent forms of O. versicolor.

 

Excluded taxa.

 

Neckera trichophoroides Hampe, Linnaea 38: 668. 1874. Pilotrichella trichophoroides (Hampe) Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 259. 1877. Meteorium trichophoroides (Hampe) Mitt., Trans. Roy. Soc. Victoria 19: 82. 1882. Barbella trichophoroides (Hampe) Broth., Nat. Pflanzenfam. 1(3): 284. 1906. Protologue: Lord Howe’s Island, Summit of Mount Gower. Type: Lord How’e Island, Summmit Mount Gower (S, isotype).

 

Fleischer (1908) made this a synonym of Barbella enervis (Mitt.) Fleisch. This opinion was confirmed by Streimann (1991) who further considered B. enervis a synonym of Barbella cubensis (Mitt.) Broth. Barbella cubensis is now a synonym of Barbellopsis trichophora (Mont.) Buck (Streimann 1993, Buck 1998). Wijk et al. (1964) attribute the name Neckera trichophoroides to Hampe ex C. Müll., however there is no indication in the protologue that Carl Müller had anything to do with the species.

 

Pilotrichella biformis Hampe var. congesta Ren. & Card., Bull. Soc. Roy. Bot. Beligique 41(1): 76. 1905. Squamidium biforme (Hampe) Broth. var. congesta (Ren. & Card.) Een, Lindbergia 3: 217. 1976. Protologue: La Réunion: S^t Philippe (rev. Rodriguez); Madagascar: Ampériféry (rev. Campenon). Types: Rodriguez (PC lectotype); Campenon (H, PC syntypes).

 

Allen and Crosby (1986) synonmized this taxon with Squamidium brasiliense (Hornsch.) Broth.

 

Pilotrichella capilliramea C. Müll. in  Par., Index Bryol. Suppl. 270. 1900, invalid name, lacks a description.

 

No material of this taxon was examined. It is restricted to the Philippines and is not likely to be an Orthostichella. The last checklist of Philippine mosses (Tan & Iwatsuki 1991) makes no mention of this taxon.

 

Pilotrichella chlorothrix C. Müll., Flora 73: 490. 1890. Squamidium chlorothrix (C. Müll.) Broth., Nat. Pflanzenfam. 1(3): 809. 1906. Protologue: Africa or. trop. in sylva primaeva montis Kilima-Ndscharo, 1500–3000 m: Dr. Hans Meyer 1889. Type: Africa or. trop. in sylva primaeva montis Kilima-Ndscharo, 1500–3000 m: Dr. Hans Meyer 1889 (H).

 

Allen and Crosby (1986) synonmized this taxon with Squamidium brasiliense (Hornsch.) Broth.

 

Orthostichopsis chrysoneura (Hampe ex C. Müll.) Broth. Nat. Pflanzenfam. 1(3): 805. 1906. Neckera chrysoneura Hampe ex C. Müll., Linnaea 40: 263. 1876. Pilotrichella chrysoneura (Hampe ex C. Müll.) Jaeg., Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 256. 1877. Protologue: Comoro Islands. Patria. Comoro-insula Johanna, unde ad filices arborescentes involvendas misit J. M. Hildebrandt 1875. Types. Africa orient: Comor-insula Johanna, ad arbores, leg. Hildebrandt 1875 (S, isotype); Archipelagus Comorensis, insula Johanna: J. M. Hilderandt, 1875 (S, isotype).

 

Renauld and Cardot (1915) considered Orthostichopsis chrysoneura a local race of the Madagascan O. subimbricata (Hampe) Broth). Argent (1973b) also placed O. chrysoneura into the synonymy of Orthostichopsis subimbricata, although he did not see the type of O. subimbricata. Orthostichopsis chrysoneura is gametophytically extremely close to the southern South American O. tenuis (Jaeg.) Broth. The two differ only in seta length: O. tenuis capsules exserted; O. subimbricata capsules immersed.

 

Pilotrichella densiramea Broth., Bot. Jahrb. Syst. 20: 197. 1894. Squamidium densirameum (Broth.) Broth., Nat. Pflanzenfam. 1(3): 809. 1906. Squamidiuim biforme (Hampe) Broth., var. densirameum (Broth.) Bizot, Acta Bot. Acad. Sci. Hung. 28: 33. 1982. Protologue: Tanzania. Usambara: Handeï, trockener Hochwald, an Baumrinde (Holst n. 9056) und Lugulua Wald, auf alten Bäumen (n. 2629a). Types: Usambara: Handeï, trockener Hochwald, an Baumrinde Holst n. 9056 (S, syntype); Lugulua Wald, auf alten Bäumen Holst 2629a, H, lectotype).

 

Allen and Crosby (1986) synonomized this taxon with Squamidium brasiliense (Hornsch.) Broth.

 

Pilotrichella dubia C. Müll. in Paris, Index Bryol. 944. 1897, invalid name lacks a description. Protologue: Pac.: Nov. Guinea. (M. Yule).

 

This taxon is excluded from Orthostichella based on its occurrence in New Guinea.

 

Pilotrichella eustichia Schimp. in Kindb., Enum. Bryin. Exot. Suppl. 2: 102. 1891, invalid name, lacks a description. No specimen was cited along with this name.

 

There is a collection in H from the Bescherelle herbarium named Pilotrichella eustichia Sch. The specimen, collected in Mauritius (Danty n. 14), is Orthostichidium pentastichum.

 

Pilotrichella kraussei Lor., Moosstudien 164. 1864. Protologue: Chile. Prope Valdiviam leg.Dr. Krausse. Isotype: Chile. Ad arborum ramos in silvis densis udis. 0–2000’ prope Correl, puerto de Valdivia, leg. Dr. H. Krausse (NY).

 

This taxon is synonymous with Weymouthia cochlearifolia (Schwägr.) Dix.

 

Pilotrichella lepto-erioclada C. Müll. in  Par., Index Bryol. 946. 1896, invalid name, lacks a description. Based on: Musci Africani in Camerunia a P. Dusén collecti. Ad N’dian emporium in ramulis arborum die 26 m. Martii a. 1892, Dusén 800 (S).

 

The plants in this collection belong to the Sematophyllaceae.

 

Pilotrichella percapillaris C. Müll. in Par., Index Bryol. 948. 1897, invalid name, lacks a description. Based on V. Afr., Kamerun, Jumbo 1080 m, 17/4/92, P. Dusén (S).

 

The plants in this collection are Rhacopilopsis trinitensis (C. Müll.) Britt. ex Dix.

 

Hildebrantiella phleoides (Desvaux ex Brid) Allen & Magill, comb. nov. Hypnum phleoides Desvaux ex Brid., Bryol. Univ. 2: 431. 1827. Pilotrichella phleoides (Desvaux ex Bridel) Jaeger, Ber. Thäigk. St. Gallischen Naturwiss. Ges. 1875–76: 261. 1877. Protologue: In insula Bourbonis [Reunion] in arborum cortice habitat. Type: Neckera phleoides mihi (Hypnum Desv.) Ins. Bourbon, hb. Brid. (JE, isotype).

 

Hildebrantiella pachyclada Besch., Ann. Sci. Nat., Bot. sér. 6, 10: 264. 1880, syn. nov.

 

 

Pilotrichella recurvula C. Müll., Hedwigia 41: 129. 1902. Protologue: Australia. Habitatio. Queensland, Kev Range, Ballendea, 1889; F. M. Bailey. Hb. Brotheri 1890.

 

Streimann (1991) examined the type of this species and determined it was identical to Calyptothecium recurvulum (Broth.) Broth..

 

Pilotrichella rehmanni C. Müll. in Geh., Rev. Bryol. 5: 70. 1878, invalid name, lacks a description.

 

This taxon is synonymous with Squamidium brasiliense (Hornsch.) Broth.

 

Pilotrichella serricola C.Müll. in Par., Index Bryol. 949 1897, invalid name, lacks a description. Based on E. Ule, Bryotheca brasiliensis n. 67 (BM).

 

This taxon is synonymous with Squamidium brasiliense (Hornsch.) Broth.

 

Pilotrichella subbiformis Ren. & Card., Bull. Soc. Roy. Bot. Beligique 39(2): 110. 1900. Erythrodontium subbiformis (Ren. & Card.) Broth., Nat. Pflanzenfam. 1(3): 888. 1907. Protologue: Congo. Rég. V: Kisantu (J. Gillet, 1900, n^o 712). Type. Congo belge: Kisantu, leg. Gillet, Herb. F. Renauld (S, isotype).

 

The very small isotype of this taxon at S belongs to Erythrodontium.

 

Pilotrichella subsemitorta Broth. in Naveau, Rev. Bryol., n.s. 1: 38. 1928, invalid name, lacks a description.

 

According to Theriót (1929) this is an error for Papillaria subsemitorta Broth. ex Ther.

 

Nomina nuda not seen.

 

Pilotrichella ankasinae C. Müller in Kindberg, Enum. Bryin. Exot., Suppl. 2 102 1891. Protologue: nothing. Invalid name, lacks a description.

 

Pilotrichella chrysoneura var. viridis Renauld & Cardot, Rev. Bot. Bull. Mens. 9: 221 1891. Protologue: [Madagascar]. Invalid name, lacks a description

 

Pilotrichella cuspidata Broth., var. gracilescens Tosco & Piovano, Allionia 4: 124. 1956. Protologue: Kenya (Central Prov.) – Massailand, foresta (Leg. G. Balbo, 6 genn. 1908); Tuthu, Monte Gasongori (Leg. G. Balbo, 7 lug. 1908). Invalid name, holotype not indicate.

 

Pilotrichella platyclada Par., Index Bryol. 948. 1897, invalid name lacks a description. Orthostichella platyclada C. Müll. in Par., Index Bryol. 948. 1897, invalid name lacks a description. Based on: Musci Africani in Camerunia a P. Dusén collecti. 846.

 

Pilotrichella porotrichoides C. Müller in Paris, Index Bryol. Suppl. 271 1900, invalid name, lacks a description. Based on: Sikora Pl. Madag. n. 14234

 

Pilotrichella stracheyana C. Müll. in Kern, Rev. Bryol. 19: 41. 1892, invalid name lacks a description. Protologue: As stuffing in a Satyr pheasant (Ceriornis satyra).

 

Types not seen.

 

Pilotrichella debilinervis Ren. & Card., Bull. Soc. Roy. Bot. Beligique 34(2): 70. Protologue: Hab. Bourbon: Salazie, in sylva “de Belouze” dicta (Chauvet; hb. de Poli).

 

Pilotrichella delicatula Brotherus in Mildbraed, Wiss. Erg. Deut. Zentr.-Afr. Exped., Bot. 2: 160 1910. Protologue: Vulkan-Gebiet [Central Africa]: Bambusmischwald von Bugoie, ca. 2100 m ü. M[ildbraed]. (steril im Sept. 1907–n. 2034). Illegitimate homonym, not Pilotrichella delicatula (Besch.) Schimp., (1897).

 

Hypnum imbricatum Palisot de Beauvois, Prodr. Aethéogam. 65. 1805. Pilotrichella imbricata (Palisot de Beauvois) Bescherelle, Mem. Soc. Sci. Nat. Cherbourg 16: 222 1872. Protologue: l'Ile de Bourbon [Reunion] Bory-St.-Vincent

 

Pilotrichella imbricatula C. Müller ex Geheeh, Abh. Naturwiss. Vereine Bremen 7: 209 1881. Protologue: [Madagascar] Wald von Ambatondrazaka, [Brogen or Borchgrewink] 6 Decbr. 1877. [= P. isleana Besch fid. Card. in Grand., Hist Madag. 39: 350, 1915.]

 

Pilotrichella imbricatula var. exserta Thériot, Recueil Publ. Soc. Havraise Etudes Diverses 1929: 116 1930. Protologue: [Madagascar] Massif de l'Andringitra. (leg. H. Humbert).

 

Pilotrichella imbricatula var. nervosa Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique 33(2): 126 1895. Pilotrichella isleana var. nervosa (Renauld & Cardot) Cardot in Grandidier Hist. Phys. Madagascar, Mousses 39: 351 1915. Protologue: Madagascar: inter Savondronina et Ranomafana (D^r Besson). — (Renauld, Musci Mascareno-Madagascarienses Exsiccati, n°.228).

 

Pilotrichella latiramea C.Müller ex Dusén, Kongl. Svenska Vetensk. Acad. Handl. 28(2): 29. 1895. Protologue: Habitat in Camerunia in ramulis arborum, ubi ad Ekundu Ndene emporium m. Julio a. 1892 legi [Dusén]

 

Neckera pseudo-imbricata C. Müll., Bot. Zeitung (Berlin) 20: 381. 1862. Pilotrichella pseudoimbricata (C. Müller) Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1875–76: 255 1877. Protologue: Comoro Islands. Insula Anjoana ad canal. de Mozambique Dr. Peters 1843.

 

Neckera pseudo-imbricata C. Müll. var. pallescens C. Müll., Linnaea 40: 265 1876. Pilotrichella pseudoimbricata var. pallescens (C. Müller) Paris, Index Bryol. 948 1897. No specimen cited in protologue.

 

Neckera pseudo-imbricata C. Müll. var. rufescens C. Müll., Linnaea 40: 265 1876. Pilotrichella pseudoimbricata var. rufescens (C. Müller) Paris, Index Bryol. 948 1897. No specimen cited in protologue.

 

Pilotrichella ragazzii Brizi, Annuario Reale Ist. Bot. Roma 5: 79. 1893. Prtologue: [Ethiopia]. Comunissima sul tronco degli alberi nella foresta di Fekeriè-Ghemb. 20 Maggio 1885. — Let-Marefià. 21 Marzo 1885. [V. Ragazzi].

 

Pilotrichella sericea C. Müll. ex Kindb., Enum. Bryin. Exot., Suppl. 2: 102. 1891, new new for Orthostichella sericea C. Müll., Flora 73: 492. 1890, invalid name due to an invalid generic name. Protologue: Patria. Africa or. trop., Leikipia in regione occidentalis montis Kilima-Ndscharo, ad pedem der Aberdare-Kette: L. Höhnel in Exped. Telekiana 1887.

 

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