A new species of Erodium L\'Her. (Geraniaceae) endemic to Australia

Share Embed


Descrição do Produto

Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2002 141 Original Article A NEW SOUTH AUSTRALIAN SPECIES OF ERODIUM M. L. ALARCÓN

Botanical Journal of the Linnean Society, 2003, 141, 243–250. With 14 figures

A new species of Erodium L’Hér. (Geraniaceae) endemic to Australia MARÍA LUISA ALARCÓN1*, JUAN JOSÉ ALDASORO1, CARLOS AEDO1 and CARMEN NAVARRO1 1Real

Jardín Botánico, Consejo Superior de Investigaciones Científicas, Plaza de Murillo 2, 28014 Madrid, Spain. 2Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense, 28040 Madrid, Spain Received May 2002; accepted for publication August 2002

Erodium janszii Alarcón, Aldasoro, Aedo & Navarro is described as a new species endemic to Southern Australia. Characters allowing discrimination between it and the following species, E. aureum, E. cygnorum, E. crinitum and E. carolinianum, are reported, with particular emphasis on the latter, which is the closest. A detailed illustrated description, as well as a distribution map, are also provided. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250.

ADDITIONAL KEYWORDS: awn indument – glandular hairs – mericarp bristles – morphology – papillae – sepal indument – taxonomy – variability.

INTRODUCTION The genus Erodium L’Hér. shows the highest diversity in the Mediterranean basin (aproximately 60 species), but a few endemic species occur in North America (1), South America (1), Central Asia (2) and Australia (5) (Knuth, 1912; Guittonneau, 1972). Carolin (1958) revised the taxonomy of Erodium in Australia, elaborating a key to differentiate endemic taxa. Later, Carolin (1970) described a new species, E. angustilobum. Aldasoro et al. (2000a) argued that E. cygnorum ssp. glandulosum should be treated as a different species, considering that the differences with E. cygnorum ssp. cygnorum (and with the other taxa) are significant, and proposed the name E. carolinianum. Previously undetected floral characters of Erodium were provided by Aldasoro, Aedo & Navarro (2000b). These include the presence of broad flat hairs in the upper petals of some species that reflect light and attract certain insects. The only endemic species showing this character is E. aureum. ET AL.

*Corresponding author. E-mail: [email protected]

Ten annual species are reported in Australia (Carolin, 1967, 1986), four of which are endemic: E. aureum Carolin, E. crinitum Carolin, E. cygnorum Nees in Lehm. and E. carolinianum Aldasoro et al. No recent studies are available on Australian species of Erodium. The present contribution is part of our current world taxonomic revision of the genus.

MATERIAL AND METHODS This paper is based on herbarium specimens, from the following herbaria: AD, CANB, COI, K, MA, MEL, MO, NSW, P, PERTH, W. A total of 320 specimens were examined. For scanning electron microscopy (SEM), mericarps were sectioned, glued on aluminium stubs, coated with 40–50 nm gold, and examined in a JEOL-TSM T330A scanning electron microscope at 20 kV. Flowers were taken from specimens, soaked in warm water with 1% NaOH or with 2–3 drops of liquid soap. After two hours they were transferred to water, treated by the critical point drying method (Cohen, 1974), and glued on aluminium stubs, coated with gold, and examined by SEM.

© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

243

244

M. L. ALARCÓN ET AL.

Figure 1. Erodium janszii (a,c–e, Donner 3266, NSW 459934; b, Corrik 7271, MEL 592017; f–k, Corrik 7271, NSW 461849; l–n, Kennedy s.n., MEL 282160). a,b, habit; c, leaf margin, sheaf; d, leaf margin, abaxial side; e, bracts; f, pedicel; g, flower; h, sepal; i, petal; j, staminal filament; k, staminode; l, mericarp; m, apical pit; n, seed.

© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

A NEW SOUTH AUSTRALIAN SPECIES OF ERODIUM Several quantitative characters were recorded and measured using a Brown and Acutee 599-571-3 digital caliper. To represent the variability of each descriptor within species, box-plots were prepared. These plots contained medians and percentiles and were calculated using the STATISTICA 5.1 package.

RESULTS ERODIUM JANSZII M.L. ALARCÓN, J.J. ALDASORO, C. NAVARRO & C. AEDO SP. NOV. (FIG. 1) A specie sat simili E. carolinianum Aldasoro et al. descripta hodie facile distinguitur foveola mericarpiorum pilis non glanduliferis praedita; distinguitur insuper mericarpiis pilis non glanduliferis undique multo frequentioribus atque, ut infra dicimus, diversis. We chose the epithet to honour Willem Jansz, the first European to arrive to Australia. Description: Annual plant, caulescent, with short roots. Stems 5–50 cm long, ascending, hairy, with patent glandular hairs 0.4–0.8 mm long. Leaves with 3–5 pairs of secondary nerves; 14–45 mm long and 0.9–42 mm wide; triangular-ovate, generally 3 lobate, crenate; the first leaves are usually less deeply divided, sometimes nearly entire; with glandular hairs 0.5–1.1 mm long; cauline leaves, more or less

245

similar to the basal ones; stipules 0.3–6 mm long, triangular, membranous, hairy, generally with glandular hairs 0.9–1.5 mm long in the margins and glandular hairs 0.4–0.7 mm long in the abaxial part, whitish or brownish. Inflorescences axillary, 2–5 flowered; bracts 2–5, 1.7–5 mm long and 1.2–2 mm wide, free, triangular, membranous, brownish, with glandular hairs 0.2– 1.3 mm long in the margins and in the abaxial part; pedicels hairy, then with eglandular or with glandular hairs 0.5–0.8 mm long. Sepals 4–8.7 mm long and 1.7–4 mm wide (7.5–15.6 mm long and 2.5–5 mm wide during fructification), with glandular hairs of 0.3– 1.2 mm long and eglandular hairs of 0.4–1 mm long, more or less patent; awn 0.7–1 mm long (0.9–1.1 mm long during the fructification). Petals 6.7–11 mm long and 3.4–5 mm wide, bluish-violet; claw ciliate. Stamen filaments 2.6–4.2 mm long and 0.8–1.1 mm wide, subulate, glabrous, green; anthers 0.8–1.2 mm long, dark brown; pollen 62–68 µm in diameter, striate or striate-reticulate; without supratectal processes, tricolpate; staminodes 1.2–3.2 mm long and 0.6–1.2 mm wide, glabrous. Nectaries 0.6 mm long, green. Gynoecium 4.3–5 mm long, hairy, with crowded eglandular hairs; stigmas 0.4–0.6 mm long. Fruit 45–90 mm long; mericarp 7.1–9.8 mm long, papillose, with crowded bristles of dissimilar size, some longer and others shorter, arising all from a ridge derived from papillae fusion in the mericarp surface; pit as long as wide

Figure 2. Distribution map in Australia of Erodium janszii (▲) and Erodium carolinianum (•) based on studied specimens. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

246

M. L. ALARCÓN ET AL.

(ratio pit length/pit width = 0.64–0.97), with crowded eglandular hairs and some sparse, long-stalked glands which are also at the start of the awn. Seed 3.6–4.8 mm long and 1–1.6 mm wide, longitudinal section elliptical. Type: South Australia. Eyre Peninsula, Gawler Ranges, Mount Ive, 32°15′S, 126°18′E, 28.ix.1969, N.N. Donner 3266 (NSW 459934 holotype; AD 97017255 isotype). Habitat: Open habitats on generally dry soils of extratropical Southern Australia (Carolin, 1958). Distribution: New South Western Australia (Fig. 2).

Wales,

Southern

and

TAXONOMIC DISCUSSION Five annual species of Erodium are endemic to Australia. All of them show a considerable variability in most features (i.e. plant size, leaf shape, indument, and flower size, etc.), but some were demonstrated as distinctive. These characters are mainly located on sepals and mericarps. The calyx indument shows interesting differences: in E. crinitum and E. cygnorum it is eglandular, while the rest of species show sepals with long glandular hairs. Erodium crinitum and E. cygnorum also have differences in the mericarp, E. crinitum has two ridges in the mericarp apex instead of a clear-cut pit (Fig. 3). Erodium cygnorum shows pits (most commonly eglandular) and 0–2(3) ridges in the mericarp (Fig. 4), and retrorse eglandular hairs on the sepals (Fig. 5).

Figures 3–8. Scanning electron micrographs of mericarps and sepals indument. Fig. 3. Pit of E. crinitum (Barritt 407, MEL 1618245). Fig. 4. Pit of E. cygnorum (Maconochie 1823, NSW 459938). Fig. 5. Sepal of E. cygnorum (Seymon 12453, NSW 459936). Fig. 6. Sepal of E. janszii (Corrick 7271, NSW 461849). Fig. 7. Pit of E. aureum (Weber 3138, MEL 2059775). Fig. 8. Papilleae and short stalked glands in the pit of E. aureum (Weber 3138, MEL 2059775). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

A NEW SOUTH AUSTRALIAN SPECIES OF ERODIUM 0,55

Ratio first lobe width/leaf width

0,45

0,35

0,25

0,15

0,05 car (8)

jans (7)

cri (9)

aur (10)

cyg (16)

Non-Outlier Max Non-Outlier Min 75% 25% Median Outliers

Figure 9. Box plot of ratio first lobe width/total leaf width, indicating the differences and variability in Australian species of Erodium relating to shape and size of leaf lobes; the lobes are relatively narrower and more variable in width in E. cygnorum than in the other four endemic species. Numbers in brackets are the number of specimens measured. Car: Erodium carolinianum; jans: E. janszii; cri: E. crinitum; aur: E. aureum; cyg: E. cygnorum.

247

Carolin (1970) distinguished E. angustilobum from E. cygnorum on the basis of its narrow leaf lobes. However, E. cygnorum shows a large variability in leaf shape and size, which includes those with narrow lobes (data not shown). These forms have no other associated distinctive character, and consequently we have subsumed E. angustilobum under E. cygnorum. Erodium janszii can be distinguished from E. cygnorum using the ratio first lobe width/leaf width (Fig. 9). The three remaining species show glandular hairs on sepals (Fig. 6). Erodium aureum has broad flat hairs on petals, and hemispherical papillae (15–17 µm in diameter, Figs 7,8) and short stalked glandular hairs in the mericarp pit, with two short cells in the stalk (total length: 40–60 µm, Figs 7,8). Erodium carolinianum shows a different type of glandular hairs in the pit, with longer basal cells (total length: 65– 120 µm, Figs 10,11) also evident at the base of the awn, while E. janszii shows these glandular hairs only in the awn base (Fig. 13). It can also be distinguished by the differences in pit shape and indument. The pit is as long as wide in E. janszii, while it is wider than long in E. carolinianum (Fig. 14). Under the pit there is a furrow which is also clearly wider in E. carolinianum. In addition, E. janszii shows short applicate eglandular hairs 50–90 µm long all over the pit (Fig. 12), these hairs are absent in E. carolinianum. Other distinctive characters occur on the mericarp surface and the calyx: the bristles are considerably more crowded in E. janszii (sparse in E. carolinianum). Thus the mericarp wall is completely masked in the

KEY TO THE SPECIES OF ERODIUM IN AUSTRALIA 1. Leaves pinnatisect.............................................................................................................................................................. 2 2. Mericarps with short stalked glandular hairs in the pit .................................................................... E. moschatum 2. Mericarps without short stalked glandular hairs in the pit ............................................................... E. cicutarium 1. Leaves ± deeply partite...................................................................................................................................................... 3 3. Sepals without glandular hairs ........................................................................................................................................ 4 4. Sepals with two types of hairs: many shorter and a few longer, all of them retrorse .......................... E. crinitum 4. Sepals with only one type of hairs, short and retrorse .......................................................................... E. cygnorum 3. Sepals with glandular hairs .............................................................................................................................................. 5 5. Mericarps without a clear-cut pit, with 2–4 transversal ridges beside the awn insertion ....................... E. botrys 5. Mericarps with a clear-cut pit..................................................................................................................................... 6 6. Fruit shorter than 29 mm.................................................................................................................................................. 7 7. Pedicels and sepals with short stalked glandular hairs 0.3–0.4 mm long. Sepal mucro 0.2–0.4 mm long ............................................................................................................................ E. aureum 7. Pedicels and sepals with long stalked glandular hairs 0.7–1.1 mm long. Sepal mucro 0.7–1.1 mm long................................................................................................................................. E. malacoides 6. Fruit longer than 34 mm .................................................................................................................................................. 8 8. Mericarps without glandular hairs in the pit, usually with eglandular hairs ............................ E. brachycarpum 8. Mericarps with long stalked glandular hairs in the pit (60–120 µm long), without papillae................................ 9 9. Apical pit of mericarp with crowded eglandular hairs, and sometimes sparse glandular ones in the top, near the awn; mericarp with crowded bristles, covering completely the surface .................... E. janszii 9. Apical pit of mericarp with only sparse glandular hairs; mericarp with sparse bristles, leaving most of mericarp surface uncovered......................................................................................... E. carolinianum

© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

248

M. L. ALARCÓN ET AL.

Table 1. Characters measured in Erodium carolinianum and E. janszii E. carolinianum Calyx Sepal length during anthesis (mm) Sepal length during fructification (mm) Length of eglandular hairs (mm) Length of glandular hairs (mm) Mericarp: bristles Bristles density (bristles/mm2) Maximal bristle length (µm) Mericarp: awn Length of eglandular hairs in the awn insertion (µm) Length of glandular hairs in the awn insertion (µm) Mericarp: pit Pit length (µm) (distance from the ridge to the awn insertion) Pit width (µm) (measured perpendiculary to the ridge) Ratio pit length/pit width Hair types and abundance

first species, while it is visible in the second. The calyx shows shorter eglandular hairs in E. janszii than in E. carolinianum. Due to large variability in leaves and stems we were not able to find any distinctive characters relating to these organs. The most remarkable features are summarized in Table 1. The distribution areas of the five endemic species are largely similar (Fig. 2 and Carolin, 1958). Both E. cygnorum (2n = 60) and E. carolinianum (2n = 60) are hexaploid, E. crinitum (2n = 40) is tetraploid and E. aureum is diploid (2n = 20) (Carolin, 1958, 1970). The chromosome number of E. janszii is unknown.

MATERIAL

EXAMINED

E. janszii Alarcón et al. sp. nov. NEW SOUTH WALES: Wilcannia Darling River, 31°34′S, 143°23′E, 1886, Kennedy s.n. (MEL 281860); Far Western Plains, near mt. Robe 35 Km NNW of Broken Hill, 31°39′S, 141°21′E, 29.viii.1981, Corrick 7271 (MEL 592017, NSW 461849); Barrier Ranges, 31°25′S, 141°25′E, 1889, Wehl s.n. (MEL 281970); Wilcannia Darling River, 31°34′S, 143°23′E, 1885, Kennedy s.n. (MEL 282160). WESTERN AUSTRALIA: Fraser Range, 32°2′S, 122°46′E, 1874, Dempster s.n. (MEL 282210); Fraser Range, Cootaorina Creek, 32°2′S, 122°46′E, 4.x.1891, Holms s.n. (MEL 281971).

E. janszii

5.2–9.7 10–12 1–1.3 0.3–1.3

5.7–8.7 10.3–15.6 0.4–1 0.3–1.2

48–56 495–625

140–167 583–854

50–171 31.4–185

56–125 56–147

314–780 625–960 0.47–0.83 sparse glandular hairs

435–670 (780) 470–810 (1020) 0.64–0.97 sparse glandular and crowded eglandular hairs

E. carolinianum Aldasoro et al. NORTHERN TERRITORIES: Hale River, 23°16′S, 134°24′E, 9.ix.1957, Chippendale 2/1957 (PERTH 3304469); Ooraminna Rockhole, 24°5′S, 134°2′E, 21.viii.1962, Swinbourne 408 (CANB 225602); Mt. Pyroclast, 6 Km S of Gosse Bluff, 23°54′S, 132°18′E, 24.vi.1992, Fletcher 78 (MEL 2022133); Alice Springs, 23°32′S, 133°21′E, 8.viii.1968, Nelson 1721 (CANB 218563); Uluru, Ayers, Rock Montain Olga, 25°17′S, 130°44′E, 20.v.1988, Lazarides & Palmer 284 (CANB 385772); 5 m E New Crown, 25°41′S, 134°47′E, 14.vii.1968, Must 124 (CANB 313907). QUEENSLAND: 56 m WNW of Birdsville, 25°32′S, 139°1′E, 26.ix.1966, Boyland 310 (CANB 243103). SOUTHERN AUSTRALIA: Koonamore Station, 32°4′S, 139°23′E, 2.viii.1963, Crisp 259 (CANB 64415); Tinga-Tingana, 28°48′S, 140°10′E, 20.viii.1968, Carrick 1948 (PERTH 3305643); Bunyeroo Valley Lookout, 31°24′S, 138°35′E, 21.viii.1964, Phillips 134 (CANB 6591); Far NE, Cooper Creek, 31°5′S, 149°16′E, 12.viii.1968, Jackson 402 (AD); Region 5, Flinders Ranges, 30°45′S, 138°20′E, 12.vii.1989, Murfet 698 (AD); Gairdner Torrens, 32°17′S, 135°56′E, 30.viii.1989, Badman 3597 (MA 592447); Lake Frome downs, Black Oak Bore, 30°47′S, 139°44′E, 24.vii.1971, Weber 2088 (W); Northern Flinders Range, Mt. Aleck, 31°39′S, 138°27′E, 24.viii.1863, Kuchel 992 (P); Lake Frome downs, near Broughams, 30°47′S, 139°44′E, 6.viii.1971, Weber 2230 (MO); Kudna Rockhole, 31°7′S, 130°38′E, 19.ii.1967, Symon 4632 (CANB

© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

A NEW SOUTH AUSTRALIAN SPECIES OF ERODIUM

249

Figures 10–13. Scanning electron micrographs of mericarps. Fig. 10. Glands in the pit of Erodium carolinianum (Chippendale 9167, PERTH 3305627). Fig. 11. Top of the mericarp of E. carolinianum showing the pit (Chippendale 9167, PERTH 3305627). Fig. 12. Pit of E. janszii showing short applicate eglandular hairs (Corrik 7271, NSW 461849). Fig. 13. Top of the mericarp of E. janszii showing the pit and glandular hairs in the awn base (Donner 3266, NSW 459934).

1,100

1,000

Ratio pit width / length

0,900

0,800

0,700

0,600

0,500

0,400 E. carolinianum (9)

E. janszii (11) Non-Outlier Max Non-Outlier Min 75% 25%

188405); c. 48 km NE of Frome Downs Homestead, 31°3′S, 140°11′E, 25.vii.1971, Whibley 3485 (CANB 326394); 20 km SW of Chambers Gorge, 30°58′S, 139°16′E, 24.ix.1973, Sihhes & Ollerenshaw 901 (CANB 62748); Huckittan Homestead, 22°35′S, 135°35′E, 21.vii.1970, Latz 679 (CANB 35881). WESTERN AUSTRALIA: Burt Plain, 23°7′S, 133°17′E, 26.vii.1962, Chippendale 9167 (PERTH 3305627); Reid, 30°49′S, 128°25′E, 1.ix.1962, Aplin 1676 (PERTH 305562); Zraza Range, 6.ix.1962, Aplin 1783 (PERTH 3303748); Mte Eveline, 26°9′S, 127°11′E, 22.viii.1962, George 3835 (PERTH 3305589); Mte. Fanny, 28°53′S, 114°57′E, 8.vii.1963, George 4825 (PERTH 3305554); Mte. Olga, 31°42′S, 119°34′E, 12.vii.1963, George 4987, PERTH 3305619; Loongana SA, 164 km N of this place, 29°28′S, 127°4′E, 20.ix.1984, Downing 949 (PERTH 3304930); Mte Aloysius, 26°0′S, 128°35′E, 21.vii.1963, George 5234 (PERTH 3305570); 35 m N of Mundrabilla, 30°15′S, 127°31′E, 19.viii.1974, Mitchell 47 (PERTH 3304256).

Median

Figure 14. Box plot of the pit dimensions in Erodium janszii and E. carolinianum. Numbers in brackets are the number of specimens measured.

ACKNOWLEDGEMENTS We thank M. Laínz for preparing the Latin diagnosis. This work was supported by the Flora iberica project

© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

250

M. L. ALARCÓN ET AL.

number PB96-0849, and the Erodium project number REN2000-0818.

REFERENCES Aldasoro JJ, Aedo C, Navarro C, Sáez LI. 2000a. A new name in Erodium L’Hér. ex Aiton (Geraniaceae). Anales del Jardín Botánico de Madrid 57 (2): 407. Aldasoro JJ, Aedo C, Navarro C. 2000b. Insect attracting structures on Erodium petals (Geraniaceae). Plant Biology 2: 471–481. Carolin RC. 1958. The species of the genus Erodium L’Hér. endemic to Australia. Proceedings of the Linnean Society of New South Wales 33 (2): 92–100. Carolin RC. 1967. Erodium L’Hér. Contributions from the New South Wales National Herbarium 102: 12–18.

Carolin RC. 1970. A new species of Erodium L’Hér. from Australia. Proceedings of the Linnean Society of New South Wales 94 (3): 212–213. Carolin RC. 1986. Erodium L’Hér. ex Aiton. In: Jessop JP, Toelken HR, eds Flora of South Australia, 4th edn, vol. 2. Adelaide, Australia: D.J. Woolman, Government Printer, 717–719. Cohen AL. 1974. Critical point drying. In: Hayat, MA, ed. Principles and techniques of scanning electron microscopy, vol. 1: 44–112. New York: Van Nostrand Reinhold. Guittonneau GG. 1972. Contribution à l’étude biosystématique du genre Erodium L’Hér. dans le bassin méditérranéen occidental. Boissiera 20: 1–154. Knuth R. 1912. Erodium L’Hér. In: Engler A, ed. Das Pflanzenreich IV.129 (Heft 53) Leipzig: Engelmann, 221–290, 583– 587.

© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 243–250

Lihat lebih banyak...

Comentários

Copyright © 2017 DADOSPDF Inc.