Abstract
A review of Rostraria in the Azores is given, and includes a key to the genus in the Azores and the description and illustration of a new species, R. azorica S. Hend., from the island of Santa Maria.
INTRODUCTION
Rostraria Trin. (=Lophochloa Rchb., see Brummitt, 1978) (Aveneae, Poaceae) is a genus of about ten species, distributed around the Mediterranean and Middle East in dry, disturbed habitats. It is sister to the genus Koeleria Pers. (see for example Soreng & Davis, 2000), but is distinguished from the latter by being annual rather than perennial and by having a more distinct awn (Clayton & Renvoize, 1986).
Rostraria cristata (L.) Tzvelev is the member of the genus most commonly encountered in the Macaronesian region. It was first recorded in the Azores in 1838 by Hochstetter & Guthnick (Seubert & Hochstetter, 1843: 10; Seubert, 1844: 20) from São Miguel, and has since been found on all the other Azorean islands with the exception of Pico and Flores (Fig. 1).
Location of the Azores and distribution of Rostraria azorica (m) and R. cristata (d).
Location of the Azores and distribution of Rostraria azorica (m) and R. cristata (d).
In their checklist of Macaronesian vascular plants, Eriksson, Hansen & Sunding (1979) (see also later editions as Hansen & Sunding, 1985, 1993) recognized Rostraria pumila (Desf.) Tzvelev (as Lophochloa pumila (Desf.) Bor.) from the Canary Islands and Porto Santo; a new record of the species from the Cape Verde Islands is also established here (see p. 130). Rostraria pumila is in general considered not to occur in the Azores. However, in his monograph of the genus, Domin (1907) alluded to the presence of R. pumila on the archipelago, as Koeleria pumila (Desf.) Domin, ‘in insulis Canariensibus et Azoricis’. (Latterly, Bor & Guest, 1968, also referred to the species in the Azores, but this reference is probably taken from Domin.) It is impossible to determine whether or not the record is in error as no Azorean specimens are cited, but because R. pumila is recognized as occurring elsewhere in Macaronesia, its presence in the Azores is at least phytogeographically feasible.
An unusual form of Rostraria cristata was noted by C. E. Hubbard in 1969 when he examined a specimen from Santa Maria collected by A. R. Pinto da Silva (Estação Agronómica Nacional, Oeiras, Portugal) and P. Dansereau (then Director of the New York Botanical Gardens) during a collecting expedition to the Azores in 1969 (Dansereau & da Silva 289). Hubbard wrote (11.vi.1969) ‘From the evidence available, I should regard your Azores grass as a new species of Lophochloa’ (see Silva & Silva, 1974: 84), to which Silva & Silva responded ‘I hope that in soon [sic] Hubbard will find time to publish it’. It appears that he did not.
Ten years later, Eriksson, Hansen & Sunding (1979) (and following editions as Hansen & Sunding, 1985, 1993) included the species Lophochloa azorica A.Hans. in their checklist. Cited as an endemic and found only on the island of Santa Maria, it seems that this indeed must be the same species as that recognized, but not formally described, by Hubbard. However, as there is also no record of Alfred Hansen ever publishing the species name, it must be considered a nomen nudum.
During a revision of the grasses of the Azores, examination of herbarium material studied by Hubbard (Dansereau & da Silva 289) and Hansen (12.vi.1974, Hansen 92), both held at the Royal Botanic Gardens, Kew (K), confirmed that both specimens belong to the genus Rostraria but are distinct from R. cristata; additional specimens and ecological information collected by HS have reinforced our convictions.
TAXONOMIC SUMMARY
ROSTRARIA AZORICA S. HEND. SP. NOV. (FIG. 2)
Rostraria azorica sp. nov.: (A) habit; (B) spikelet; (C) anther. Rostraria cristata: (D) habit; (E) spikelet; (F) anther. (A) Schäfer 526 (BM). (B,C) Schäfer 460 (BM). (D) Schäfer 472 (BM). (E,F) Schäfer 273 (BM).
Rostraria azorica sp. nov.: (A) habit; (B) spikelet; (C) anther. Rostraria cristata: (D) habit; (E) spikelet; (F) anther. (A) Schäfer 526 (BM). (B,C) Schäfer 460 (BM). (D) Schäfer 472 (BM). (E,F) Schäfer 273 (BM).
Type: Azores, Santa Maria, on roadside near the airport, 12.vi.1974, A. Hansen 92 (holo. K!).
Note: Hansen's specimen from K is designated as holotype, as it is a more complete specimen than that of Dansereau & da Silva. It is also annotated with the unpublished epithet Lophochloa azorica A.Hans.
Lophochloa azorica A.Hans. nom . nud.
Diagnosis:R. cristatae (L.) Tzvelev similis, sed glumis longioribus, glumis inferioribus 2–3-nervibus, glumis superioribus 5-nervibus, aristis longioribus, antheris elongatis differt.
Description: Annual, caespitose, with sometimes geniculate culms 2.5–12(−18) cm tall, unbranched. leaf sheaths pubescent above and below, hairs to c. 1 mm long. ligule membranous, 0.5–1.5 mm long, obtuse, lacerate. blades linear, flat, 2–11 cm long, 1.5–3 mm wide, ± densely pubescent above and below. panicle lax, narrowly oblong, not lobed, 1.5–8 cm long, 3–9 mm wide, rachis scabrid/shortly pubescent. spikelets obovate, 5.5–9 mm long, 2–5 mm wide, with 3–8 florets; rachilla with short hairs 0.1–0.5 mm long, not produced, i.e. does not extend beyond uppermost floret. glumes unequal, acute, hyaline, usually glabrous, scabrid on the keel, shorter than the spikelet; the lower narrowly elliptic, 2–3-nerved, 3–6.5 mm long; the upper elliptic (4–)5-nerved, 4–7 mm long, sometimes apiculate to c. 0.4 mm. lemma elliptic, 5-nerved, 5–6 mm, acute, faintly 2-fid, smooth or papillose; awn inserted in the upper 2/3–4/5 of the lemma, straight or somewhat geniculate, antrorsely aculeolate; first lemma pubescent, the hairs up to c. 0.75 mm, awn (0–) 2.25–4 mm; second lemma ± glabrous, with awn 3.25–6 mm, longer than (or occasionally equal to) that of the first. palea 2-nerved, 2-keeled, 3/4 to almost equal to length of lemma, bidentate with setae 0.3–1 mm long, glabrous but scabrid on keels. stamens 3. anthers elongate (0.9–)1–1.6 mm long. caryopsis not seen. chromosome number unknown.
Colour photograph in Schäfer 2002: 219(3) (as Lophochloa azorica A.Hans.).
Distribution: Only known from the island of Santa Maria, Azores (Fig. 1).
Habitat and ecology:Rostraria azorica is scattered to locally common on coastal cliffs and sandy ground close to the sea, on walls, in vineyards and on steep roadside slopes, often with Gaudinia coarctata (Link) Dur. & Schinz., from sea level to around 180 m (Fig. 3). There is little apparent difference in ecology between R. azorica and R. cristata, and the two species can be found coexisting in some habitats.
Distribution and abundance of Rostraria azorica (d) and R. cristata (▪) on Santa Maria, the Azores. Size of symbol relates to a subjective measure of abundance, ranging from rare (smallest size) to common (largest size).
Distribution and abundance of Rostraria azorica (d) and R. cristata (▪) on Santa Maria, the Azores. Size of symbol relates to a subjective measure of abundance, ranging from rare (smallest size) to common (largest size).
Notes: The main morphological differences between R. cristata and R. azorica are outlined in Table 1 and Figure 2. Rostraria azorica is similar to R. cristata, but differs in its distinctly larger spikelets, glumes, lemmas and anthers, and by its longer awns. Despite this, R. azorica tends to be rather smaller in stature than R. cristata. Furthermore, the lower glume has 2–3 nerves as opposed to 1, and the upper glume has 5 nerves as opposed to 3; this disparity in venation is likely a consequence of the larger size of the glumes (T. Cope, pers. comm. 2002). The insertion of the awn on the lemma is somewhat lower in R. azorica than in R. cristata which has a subapical awn; also the leaf blades tend to be more densely and uniformly hairy in R. azorica than in R. cristata.
Main differences between Rostraria azorica and R. cristata. All measurements are taken from specimens collected in the Azores
| Character | R. azorica | R. cristata |
|---|---|---|
| Culms | ||
| Length (cm) | 2.5–18 | 4.5–30 (−60) |
| Panicles | ||
| Shape | Narrowly cylindrical | Cylindrical/pyramidal |
| Lax/dense | Lax | Lax/dense |
| Lobed | No | Yes/no |
| Spikelets | ||
| Length (mm) | 5.5–8.75 | (2) 2.25–5 |
| No. florets | 3–8 | 3–6 |
| Lower glume | ||
| Length (mm) | 3–6.5 | 1.5–3.5 |
| No. nerves | 2–3 | 1 |
| Upper glume | ||
| Length (mm) | 4–7.5 | 2–4.2 |
| No. nerves | (4) 5 | 3 |
| Lemma | ||
| Length (mm) | 5–6 | 1.5–3.5 |
| Awn position on lemma | Upper 2/3–4/5 | Subapical |
| Awn length (mm), lemma 1 | 0–4 | 0.2–1.5 |
| Awn length (mm), lemma 2 | 3.25–6 | 0.75–2 |
| Anthers | ||
| Shape | Narrowly oblong | Suborbicular to oblong |
| Length (mm) | (0.9) 1–1.4 | 0.2–0.6 |
| Character | R. azorica | R. cristata |
|---|---|---|
| Culms | ||
| Length (cm) | 2.5–18 | 4.5–30 (−60) |
| Panicles | ||
| Shape | Narrowly cylindrical | Cylindrical/pyramidal |
| Lax/dense | Lax | Lax/dense |
| Lobed | No | Yes/no |
| Spikelets | ||
| Length (mm) | 5.5–8.75 | (2) 2.25–5 |
| No. florets | 3–8 | 3–6 |
| Lower glume | ||
| Length (mm) | 3–6.5 | 1.5–3.5 |
| No. nerves | 2–3 | 1 |
| Upper glume | ||
| Length (mm) | 4–7.5 | 2–4.2 |
| No. nerves | (4) 5 | 3 |
| Lemma | ||
| Length (mm) | 5–6 | 1.5–3.5 |
| Awn position on lemma | Upper 2/3–4/5 | Subapical |
| Awn length (mm), lemma 1 | 0–4 | 0.2–1.5 |
| Awn length (mm), lemma 2 | 3.25–6 | 0.75–2 |
| Anthers | ||
| Shape | Narrowly oblong | Suborbicular to oblong |
| Length (mm) | (0.9) 1–1.4 | 0.2–0.6 |
Main differences between Rostraria azorica and R. cristata. All measurements are taken from specimens collected in the Azores
| Character | R. azorica | R. cristata |
|---|---|---|
| Culms | ||
| Length (cm) | 2.5–18 | 4.5–30 (−60) |
| Panicles | ||
| Shape | Narrowly cylindrical | Cylindrical/pyramidal |
| Lax/dense | Lax | Lax/dense |
| Lobed | No | Yes/no |
| Spikelets | ||
| Length (mm) | 5.5–8.75 | (2) 2.25–5 |
| No. florets | 3–8 | 3–6 |
| Lower glume | ||
| Length (mm) | 3–6.5 | 1.5–3.5 |
| No. nerves | 2–3 | 1 |
| Upper glume | ||
| Length (mm) | 4–7.5 | 2–4.2 |
| No. nerves | (4) 5 | 3 |
| Lemma | ||
| Length (mm) | 5–6 | 1.5–3.5 |
| Awn position on lemma | Upper 2/3–4/5 | Subapical |
| Awn length (mm), lemma 1 | 0–4 | 0.2–1.5 |
| Awn length (mm), lemma 2 | 3.25–6 | 0.75–2 |
| Anthers | ||
| Shape | Narrowly oblong | Suborbicular to oblong |
| Length (mm) | (0.9) 1–1.4 | 0.2–0.6 |
| Character | R. azorica | R. cristata |
|---|---|---|
| Culms | ||
| Length (cm) | 2.5–18 | 4.5–30 (−60) |
| Panicles | ||
| Shape | Narrowly cylindrical | Cylindrical/pyramidal |
| Lax/dense | Lax | Lax/dense |
| Lobed | No | Yes/no |
| Spikelets | ||
| Length (mm) | 5.5–8.75 | (2) 2.25–5 |
| No. florets | 3–8 | 3–6 |
| Lower glume | ||
| Length (mm) | 3–6.5 | 1.5–3.5 |
| No. nerves | 2–3 | 1 |
| Upper glume | ||
| Length (mm) | 4–7.5 | 2–4.2 |
| No. nerves | (4) 5 | 3 |
| Lemma | ||
| Length (mm) | 5–6 | 1.5–3.5 |
| Awn position on lemma | Upper 2/3–4/5 | Subapical |
| Awn length (mm), lemma 1 | 0–4 | 0.2–1.5 |
| Awn length (mm), lemma 2 | 3.25–6 | 0.75–2 |
| Anthers | ||
| Shape | Narrowly oblong | Suborbicular to oblong |
| Length (mm) | (0.9) 1–1.4 | 0.2–0.6 |
Specimens examined: AZORES. Santa Maria: Praia, roadside above beach, c. 2 m, UTM 9169, 2.iv.2001, Schäfer 15 (AZU, BM); S. airport, roadside, c. 100 m, UTM 9363, 3.iv.2001, Schäfer 39 (AZU); east side of airport runway, earthy path, c. 90 m, UTM 9263, 7.iv.2001, Schäfer 123 (AZU, BM); W. Ponta do Carpinteiro, earthy path, c. 70 m, UTM 9562, 8.iv.2001, Schäfer 168 (AZU); Fonte da Grimanesa, steep dry bank in east-facing vineyards, c. 100 m, UTM 8976, 24.iv.2001, Schäfer 460 (AZU, BM); Lagoinhas, vineyards north-east of Lagoinhas, c. 30 m, UTM 9770, 30.iv.2001, Schäfer 526 (AZU, BM); Maia, in the paved vineyards, 27.v.1969, P.Dansereau & A.R. Pinto da Silva 289 (K); on roadside near the airport, 12.vi.1974, A.Hansen 92 (K).
ROSTRARIA CRISTATA (FIG . 2 )
Rostraria cristata (L.) Tzvelev Novosti Sist .Vyssh. Rast. 7: 47 (1970 publ. 1971).
Basionym: Festuca cristata L., Sp. pl. 76 (1753). Lectotype: Portugal, 92.24 (LINN), designated by Sherif & Siddiqi in F. Libya 145: 167, 1988.
Homotypic synonyms:Koeleria cristata (L.) Bertol. Amoen. Ital. 67 (1819) non Pers. Lophochloa cristata (L) Hyl. Nord. Kärlväxtfl. 1: 283 (1953); Bot. Not. 1953: 355. Koeleria phleoides (Vill.) Pers. Syn. Pl. 1:97 (1805). Lophochloa phleoides (Vill.) Rchb. Fl. Germ. Excurs. 42 (1830). Koeleria phleoides var. typica Domin Bibl. Bot. 65: 257 (1906) nom. inval. Koeleria phleoides var. azorensis Domin Bibl. Bot. 65: 268 (1906) synon. nov.
Distribution: Southern Europe, North Africa, Macaronesia (Azores, Madeira, Canary Islands and Cape Verde Islands), and from south-west Asia to north-west India. Introduced in southern Africa, Australia and America.
In the Azores, it is known from the islands of Santa Maria, São Miguel, Terceira, Graciosa, São Jorge, Faial and Corvo (Fig. 1).
Habitat and ecology:Rostraria cristata ranges from rare (Faial) to very common (Santa Maria) and is found on walls, among cobblestones, on gravel paths, on dry sandy ground, on coastal cliffs and on cultivated ground. Rostraria cristata is more common than R. azorica both in respect to its distribution on Santa Maria (Fig. 3) and across all the islands (Fig. 1). See also notes on habitat and ecology for R. azorica.
Notes: A variety of Rostraria cristata from São Miguel was recognized by Domin (1907), [Koeleria phleoides] var. azorensis, based on a specimen held at the Copenhagen Botanical Museum (C), collected by Thomas Carew Hunt in 1845 [‘Botanical Society of London no. 292’]. Domin suggested that it may even have been worthy of recognition at subspecies level [‘Varietas egregia… . Forsan K. phleiodes subspecies nova’]. He distinguished the variety on the basis of the smaller spikelets (2–2.75 mm), the upper glume which is equal or subequal to (as opposed to shorter than) the florets, the lemma-like glumes in terms of their thin texture, and the golden-coloured panicle. Inspection of Domin's specimen at C and other material of R. cristata from the Azores confirms that some inflorescences not only have small spikelets but also glumes and lemmas which are smaller than typical R. cristata. However, in the authors' opinion, the variation is neither distinct nor consistent enough to warrant varietal status and is here synonymised with Rostraria cristata (L.) Tzvelev.
Synonymy: As there have been a large number of nomenclatural changes within Rostraria cristata (L.) Tzvelev, only a selective list is given above, including those names that have been used in the Macaronesian literature. For a more comprehensive list of synonymies up to 1907 see Domin (1907: 256).
Selected specimens examined: Azores. sine loc, on walls, paths, etc., vi.1838, H.J. Guthnick s.n. (BM). Faial: Horta, Monte da Guia, 100 m, 5.viii.1972, B. Gonçalves 4506 (BM); Horta, on roadside near station, 8.vii.1973, Hansen 245 (C); Cruz do Bravo, Caminho dos Trevos, on earthy path with Papaver dubium & Phalaris canariensis, c. 130 m, UTM 6755, 10.vii.1999, Schäfer 728 (AZU). Graciosa: Santa Cruz, 10 m, 25.vii.1972, B. Gonçalves 4361 (BM). São Miguel: sine loc, 1845, T.C.Hunt s.n. (BM); sine loc, 1845, T.C. Hunt s.n. (K); sine loc, 1857, H.Drouet s.n. (BM); sine loc, on walls, paths, etc., v.1838, C. Hochstetter 132 (Herb. R.J. Shuttleworth 1877) (BM); sine loc, on top of roadside wall, 1971, W. Brooke 11501 (BM); Ponta Delgada, 50 m, 16.v.1971, B. Gonçalves 2519 (BM); Ponta Delgada, waste place at airport, 28.vi.1971, Hansen 20 (C); Ponta Delgada, waste ground by piscine S.Pedro, 10.vi.1981[?], E.C. Wallace s.n. (RNG). Santa Maria: Vila do Porto, at harbour, 28.viii.1970, Hansen s.n. (C); Valverde, 140 m, 23.iv.1972, B. Gonçalves 3803 (BM); area between airport hotel and town, 11.vi.1974, Hansen 36 (C); south Anjos picnic place east of village; gravel area of parking place, UTM 9664, 2.iv.2001, Schäfer 6 (AZU); Praia de S. Lourenço, parking place – gravel area near the beach, c. 3 m, UTM 9573, 2.iv.2001, Schäfer 20 (AZU); Praia de S. Lourenço, parking place – gravel area near the beach, c. 3 m, UTM 9573, 2.iv.2001, Schäfer 21 (AZU); airport area, c. 90 m, UTM 9363, 3.iv.2001, Schäfer 40 (AZU); east of airport runway, c 90 m, UTM 9163, 7.iv.2001, Schäfer 119 (AZU, BM); Vila do Porto, old factory east of village, waste ground, UTM 9165, 7.iv.2001, Schäfer 138 (AZU); E. Ponta do Carpinteiro, c. 70 m, gravel path in pastures, UTM 9562, 8.iv.2001, Schäfer 160 (AZU); Almagreira, roadside south of village, c. 180 m, UTM 9268, 10.iv.2001, Schäfer 201 (AZU); Figueiral E. Vila do Porto, c. 40 m, pastures on the cliffs, UTM 9066, 15.iv.2001, Schäfer 270 (AZU, BM); Figueiral E. Vila do Porto, c. 40 m, coastal cliffs, UTM 9066, 15.iv.2001, Schäfer 273 (AZU, BM); Maia, road to Calheta, by roadside in east-facing vineyard slope, c. 200 m, UTM 8976, 24.iv.2001, Schäfer 457 (AZU); vineyard, east-facing rocky bank, 150 m, UTM 8976, 24.iv.2001, Schäfer 472 (AZU, BM). Terceria: Angra, roadside, 2.vi.1972, Hansen 154a (C).
ROSTRARIA PUMILA
Rostraria pumila (Desf.) Tzvelev Novosti Sist. Vyssh. Rast., 7: 48 (1970 publ. 1971).
Basionym: Avena pumila Desf. Fl . Atl. 1: 103 (1798).
Type: Morocco. Broussonet 296 (1806) (lecto. LE).
Homotypic synonyms:Trisetum pumilum (Desf.) Kunth Rev. Gram. 1: 102 (1829). Koeleria pumila (Desf.) Domin, Bibl. Bot 65: 288 (1906). Trisetaria pumila (Desf.) Maire Bull. Soc. Hist. Nat. Afr. Nord 33: 93 (1942). Lophochloa pumila (Desf.) Bor. Grass. Burm. Ceyl. Ind. Pak. 445 (1960).
Heterotypic synonyms:Koeleria sinaica Boiss., Diagn. Plant. Orient. nov., 13: 53 (1853).
Distribution: Southern Europe, North Africa, Macaronesia (Canaries, Madeira, Cape Verde Islands (new record) and the Azores), and from the south-west of Asia to north-west India. Introduced in southern Africa and southern Australia.
There are no details of where the species occurred in the Azores, or whether it is still present on the islands.
Habitat and ecology: Unknown.
Notes:Rostraria pumila is quite distinct from the other two species, with subequal rather than unequal glumes, both of which are 3-nerved, and the lower of which is usually shortly and densely woolly. Unlike R. cristata and R. azorica, the rachilla is produced (i.e. it extends beyound the uppermost floret), and is densely villous.
Selected specimens examined: Cape Verde Islands. Santo Antão. Agua das Caldeiras, c. 1300–1500 m 24.xi.1980. W. Lobin 2208 (BM).
ACKNOWLEDGEMENTS
This paper is dedicated to Charles Edward Hubbard whose initial observations on Rostraria prompted this work. SH is grateful to the directors and staff of the herbaria cited above for the use of their collections, in particular Tom Cope at RBG Kew who provided valuable assistance and advice. Many thanks are due also to Lucy Smith for producing the plate and to Bill Baker, Richard Bateman and Bob Press for comments on earlier drafts of the manuscript. HS is grateful to the German National Merit Foundation for funding the field work in the Azores.
KEY TOROSTRARIAOF THE AZORES (ADAPTED FROM COPE, 1994)
- 1
Glumes unequal, the lower 1–3-nerved, shorter and narrower than the 3–5-nerved upper, thinly hairy or glabrous; rachilla not produced, subglabrous or with hairs up to 0.5 mm long............................................2
- 1.′
Glumes subequal, 3-nerved, the lower often shortly and densely woolly, the upper rarely so; rachilla produced, densely villous with hairs c. 1 mm long..............................................R. pumila (Desf.) Tzvelev
- 2
Lower glume 2–3-nerved, upper glume 4–5-nerved; anthers usually at least 1 mm, elongate.......R. azorica S.Hend.
- 2.′
Lower glume 1-nerved, upper glume 3-nerved; anthers 0.6 mm or less, oval............................R. cristata (L.) Tzvelev
REFERENCES
