Geleznowia calycina (J.Drumm. ex Harv.) Benth.

Anderson, Benjamin M., Binks, Rachel M., Byrne, Margaret, Crawford, Andrew D. & Shepherd, Kelly A., 2023, Using RADseq to resolve species boundaries in a morphologically complex group of yellow-flowered shrubs (Geleznowia, Rutaceae), Australian Systematic Botany 36 (4), pp. 277-311 : 295-298

Geleznowia calycina (J.Drumm. ex Harv.) Benth. , Fl. Austral. 1: 348 (30 May 1863)

Sanfordia calycina J.Drumm. ex Harv. in W. J.Hooker (ed.), Hooker’s J. Bot. Kew Gard. Misc. 7: 54 (1855); Eriostemon sanfordii F.Muell., Fragm. 1(5): 107 (Apr. 1859), nom. nov., as ‘ sandfordii ’.

Type citation: ‘ On sand plains to the east and west of the southern branch of the Hill River, and in similar situations to the south of the Irwin .’ Type: Western Australia, between Moore and Murchison Rivers, s. dat., J. Drummond 6: 83 (lecto, here designated: MEL 232716 ! [left-hand fragment]; isolecto: K 000717327 image!; PERTH 00967556 ! [ex Herbario Musei Britannici]; possible isolecto: TCD 0013332 image!) .

Geleznowia macrocarpa Benth., Fl. Austral. 1: 347 (30 May 1863). Type citation: ‘ W . Australia. Murchison river , Oldfield. ’ Type: W [estern] Aust [ralia], Murchison R [iver], s. dat., A. F. Oldfield s.n. (lecto, here designated: K 000717328 image!; isolecto: MEL 232721 !; MEL 232723 !) .

Erect single- or multiple-stemmed subshrub or shrub 0.3–0.75(1) m high; older branches light brown to light grey–brown and glabrous, younger branches yellowish-green with an indumentum of scattered to moderately dense, simple hairs up to 0.1 mm long. Leaves glaucous green to dark green, elliptic to obovate, 3.8–7.2 mm long, 2.1–4.7 mm wide, adaxial surface glabrous or rarely with scattered minute hairs to 0.05 mm long, abaxial surface glandular–verrucose and glabrous or rarely with scattered minute hairs up to 0.05 mm long. Flowers 5–7 (rarely 1–4), yellow, terminal inflorescences 15–22 mm long. Pedicel of central flower 2.8–3.7 mm long, with dense hairs 0.2–0.6 mm long. Bracts (7)10–14, pale yellow to yellow, usually tinged red post-pollination, elliptic to broadly obovate, 8.2–16.9 mm long, 5–10.3 mm wide; adaxial surface with scattered to moderately dense hairs 0.05–0.1 mm long; abaxial surface glandular–verrucose, glabrous or sometimes with scattered hairs 0.05–0.1 mm long. Bracteoles 8–12, usually paired below each flower except central flower, narrowly elliptic to obovate, 8.5–12.7 mm long, 2.5–6.7 mm wide, sometimes with an attenuate base, adaxial surface with scattered to moderately dense hairs 0.05–0.1 mm long; abaxial surface scattered to moderately dense hairs 0.05–0.3 mm long. Sepals pale yellow to yellow, elliptic to broadly elliptic, longer than petals, 9–12.3 mm long, 4.7–8 mm wide, glabrous or sometimes with hairs 0.05–0.1 mm long. Petals yellow, narrowly elliptic, cupped, coriaceous, 5.2–7.6 mm long, 2–3.7 mm wide, glabrous. Stamens 10; filaments 3–4.8 mm long, broadening at base up to 0.4–0.7 mm wide, glabrous; anthers oblong, 1.1–1.9 mm long, 0.3–0.7 mm wide. Carpels 5, free, with two ovules per carpel, total length 1.3–2 mm, total width 2–2.9 mm wide, verrucose, glabrous or with scattered to rarely moderately dense hairs up to 0.1 mm long. Style glabrous, 4–4.7 mm long, 0.4–0.5 mm wide; stigma obovoid, broader than style apex, 0.3–0.4 mm long, 0.4–0.5 mm wide. Fruit obovoid, 6–10 mm long, 8–9.8 mm wide. Seeds dark brown to black, 4.2–5.3 mm long, 2.7–3.5 mm wide; aril pale cream 2.3–3.3 mm long ( Fig. 10 ).

Distribution and habitat

Widespread through the northern sandplains from west of Gillingarra to south of Shark Bay ( Fig. 8 ) in the Avon Wheatbelt, Geraldton Sandplains, Swan Coastal Plain and Yalgoo bioregions (Department of Agriculture, Water and the Environment 2020). This species grows on flats, gentle slopes, or steeper lateritic slopes, in yellow, white or brown sand over limestone or with laterite. G. calycina is a disturbance opportunist and plants can be found growing in scrapes on the edge of gravel pits or road verges. Often found growing in low heath or shrubland associated with Allocasuarina , Acacia , Banksia , Hibbertia , Conospermum and Adenanthos .

Phenology

Early flowering species with flowers at anthesis from May to early August, and fruits forming in late August to September. The yellow petaloid bracts usually become distinctly tinged with red post-pollination ( Fig. 10 c ).

Conservation status

Reasonably widespread and not considered to be under threat at this time.

Typification

In April 1854, William Harvey met James Drummond in Perth and they discussed ‘several new and curious genera which he had recently discovered in the newly opened country 300 miles [~ 480 km] to the Northwards… One of the Rue family [ Rutaceae ] is a very curious & beautiful plant & quite a new type in the order’ ( Harvey 1854 a, p. 110). Drummond had briefly described this new species, without providing a name, as ‘a stiff upright-growing shrub, about 2 ft [~ 0.61 m] high; the flowers are borne in corymbs from nine inches to a foot in diameter; they are not very conspicuous, but are accompanied by numerous large bracts of a golden-yellow colour, which render this one of the most showy of our native plants; it appears on sand-plains to the east and west of the southern branch of the Hill River, and in other similar situations to the south of the Irwin River’ ( Drummond 1853, p. 122). Harvey offered to describe this species by using the name Sanfordia calycina proposed by Drummond ( Harvey 1854 b, 1855); however, it is unclear what specimens he may have examined to refine his description, which was published while he was still overseas prior to his return to Trinity College in Dublin in 1856. Harvey may have examined material with Drummond. Alternatively, Harvey could have used material sent to Melbourne while visiting in 1854, because he received other specimens from Western Australia during that time ( Harvey 1854 c). Of the several available syntypes for this name, Wilson (2013) treated the Drummond 83 specimen at the Royal Botanic Gardens, Kew ( K 000717327) as the ‘holotype’ and listed three sheets as ‘isotypes’ ( MEL 232716; PERTH 00967556; TCD 0013332). The sheet at Kew ( K 000717327) with a blue ‘83’ label has a ‘Herbarium Hookerianum 1867’ stamp and a note ‘between Moore and Murchison Rivers, W. Australia, J. Drummond 1853 ’, suggesting that it was received at Kew in 1853 before Harvey met Drummond, so he is unlikely to have seen this material. Similarly, the PERTH 00967556 specimen that was previously lodged at BM is dated 1854, again presumably when it was received there, and is also unlikely to have been seen by Harvey. Therefore, the mounted specimen on the left-hand side of the MEL 232716 sheet with a Drummond ‘83’ tag attached and a corresponding grey Mueller label on the right with ‘ Geleznowia verrucosa Turcz. var calycina, W.A. J.Dr. 83’ in his handwriting is selected here as the lectotype of G. calycina , whereas the packet of fragments on the right is isolectotype material of G. verrucosa (see typification under that name). The specimen of G. calycina at the Trinity College Herbarium ( TCD 0013332) was collected by Drummond and is not numbered, and the locality is given only as the Swan River Colony. The specimen was not annotated by Harvey, and the name under Geleznowia verrucosa is ‘ Sanfordia floribunda JDr’ not S. calycina as Drummond had proposed, so it is unclear whether this material was used by Harvey to form the protologue and, subsequently, lodged at TCD. Paul Wilson treated this specimen as a ‘probable isotype’ (in sched. 30 June 1993); however, this cannot be confirmed, so it is treated as a possible isolectotype here. A specimen at the Muséum National d’Histoire Naturelle ( P 06615735 image!), with a blue Mueller label and identified in his hand as Geleznowia verrucosa var. calycina , is another J. Drummond collection but lacks a collection number, date or locality, other than ‘ W. A.’. This specimen along with several other Drummond collections at MEL without collection dates or locality information are therefore excluded from consideration as type material of Sanfordia calycina .

Of the available syntypes for Geleznowia macrocarpa, Wilson treated the Kew specimen from Hooker’s Herbarium ( K 000717328) as the ‘holotype’. This specimen is here designated as the lectotype for Geleznowia macrocarpa . This sheet is an Oldfield collection from the Murchison River with a ‘Herbarium Hookerianum 1867’ stamp and was therefore available to Bentham. Similarly, the MEL 232723 specimen is an Oldfield collection from the same locality and is initialled with Bentham’s characteristic ‘B’, so it was also seen by him. The fragment on the MEL 232721 specimen also has a note in Bentham’s hand, indicating that the material is from the specimen sent to Hooker and as such is treated here as an isolectotype, even though there is no information to suggest it was an Oldfield collection. Overall, the quality of the type material is very poor, consisting of bare branchlets with a few loose leaves and flowers with the subtending bracts having fallen off, thus the total number of flowers or bracts within each inflorescence cannot be confirmed. Distinguishing features mentioned by Bentham (1863) in his protologue include narrow sepals and the ‘[c]occi (not yet fully ripe) more than twice as long as broad’. There are a few specimens at PERTH that have sepals narrower than typical (3.2–4.2 mm wide cf. 4.5–8.5(9.6) mm wide) and an ovary that is longer than broad (e.g. C. A. Gardner 7744 PERTH 0968617), corresponding to the flowers observed on the type. These specimens are intact and are currently identified as G. calycina because of the size and number of flowers and bracts within each inflorescence. Therefore, G. macrocarpa is currently retained herein as a synonym of G. calycina . It should be noted that within this ‘narrow sepal’ group there are a few specimens that have quite dense hairs and two specimens where the fruits also have simple hairs and unusual finger-like protrusions extending from the verrucose glands at the apex (e.g. A. M. Ashby 2172 PERTH 00969206; R. Bates 4052 PERTH 04274695).

Notes

This species can be distinguished from other Geleznowia by the following character combination: a low subshrub 0.3–0.75 mm high with dark green leaves; usually with 5–7 (sometimes 1–4) flowers per inflorescence, surrounded by (7)10–14 pale yellow to yellow petaloid bracts (often tinged red post-pollination), 8.2–16.9 mm long, 5–10.3 mm wide, abaxial surface glabrous or sometimes with scattered hairs 0.05–0.1 mm long; 8–12 bracteoles; sepals 9–12.3 mm long, 4.7–8 mm wide; and a broad stigma 0.4–0.5 mm wide. Geleznowia calycina most closely resembles G. uberiflora but can be distinguished from it by its maximum flower number of 5–7 flowers per inflorescence (cf. usually 1–3 flowers), with more bracts ((7)10–14 cf. (5)6–8) and more bracteoles (8–12 cf. 0–6). G. calycina is also an early flowering species with flowers appearing as early as May and fruits forming in late August to September (cf. peak flowering in August–September in G. uberiflora ). Moreover, the bracts and sepals in G. calycina usually turn red post-pollination ( Fig. 10 c ), which readily distinguishes this species when it is found growing with the later-flowering G. uberiflora ( Fig. 10 e ).

Geleznowia calycina is also somewhat morphologically similar to G. narcissoides but is readily distinguished by its fewer flowers, with 5–7 flowers per inflorescence (cf. (5) 7–10 flowers), greenish-yellow to yellow petaloid bracts (cf. pale lemon yellow), and bracts, bracteoles and sepals being glabrous or with an indumentum of scattered to moderately dense hairs up to 0.1 mm long (cf. an indumentum of moderately dense to dense hairs up to 1.2 mm long). G. calycina also flowers earlier than G. narcissoides , often showing the distinctive red blush to bracts post-pollination when co-occurring with G. calycina when its flowers are in bud or reaching anthesis in spring.

The type nominated by Broadhurst for her unpublished G. verrucosa subsp. formosa ms R. V. Smith 66/370 ( PERTH 0967580) (in sched.) falls within Wilson’s (2013) broad concept of G. verrucosa , but examination of this sheet confirms that it is a match for G. calycina . Other specimens determined by Broadhurst as G. verrucosa subsp. formosa ms represent various taxa, for example L. Broadhurst 11 ( PERTH 05645298) is G. narcissoides , whereas L. Broadhurst 3 ( PERTH 05496624) and L. Broadhurst 18 ( PERTH 05599016) are G. amabilis .

Selected specimens examined

WESTERN AUSTRALIA. 93 miles [~ 150 km] N of Geraldton on North West Coastal Highway , [60 km N of Binnu], 30 July 1967, A. M. Ashby 2172 ( PERTH 00969206 ); Kalbarri National Park , 30 Aug. 1984, R. Bates 4052 ( PERTH 04274695 ); 1.4 km from turnoff along Hawks Head Road , 31 May 1996, L . Broadhurst 13 ( PERTH 05496667 ); corner West Binnu Road and turnoff to Hutt River Province , 18 Oct. 1996, L. Broadhurst 17 ( PERTH 05599032 ); Yerina Springs Road, 8.2 km S of West Ogilvie Road , 3 Oct. 2004, A. Crawford ADC 590 ( PERTH 07118279 ); eastern verge of the Midlands Road ~ 4.6 km N of the Buntine–Marchagee Road ( N of Marchagee ), 7 Sep. 2007, A. Crawford ADC 1344 ( PERTH 07752318 ); 2.2 km E of the turnoff to the Loop and Z Bend gorges, Kalbarri NP on Kalbarri–Ajana Road, E of Kalbarri , 3 Oct. 2010, A. Crawford ADC 1386 ( PERTH 07828748 , NSW); Gravel Reserve E of White Gums Nature Reserve, 9 Sep. 2008, A. Crawford ADC 1839 ( PERTH 08202095 ); Strawberry–Walkaway Road, 28.7 miles (~ 46 km) S of Walkaway, 1976, Hj. Eichler 22002 ( CANB, NSW, PERTH 08280614 ); near Eradu , 9 Oct. 1945, C. A. Gardner 7744 ( PERTH 00968617 ); Strawberry–Walkaway Road, ~ 46 km (~ 28.7 miles) S of Walkaway , 28 Sep. 1976, R. W. Johnson 3366 ( AD, PERTH 00970808 ); 9.6 km W of the Hawkes Head Lookout Road on the Ajana–Kalbarri Road, E of Kalbarri , 1 Sep. 2021, K. A. Shepherd & B. M . Anderson KS 1848 ( PERTH 09508120 ); 10.7 km from Natta Road on Tomkins Road, E of Brand Highway , 22 Sep. 2009, K. A. Shepherd & J . Wege KS 1290 ( PERTH 08159882 ); 12.6 km N from the Port Gregory Road on Yerina Springs Road , SSE of Kalbarri, 23 Sep. 2009, K. A. Shepherd & J . Wege KS 1300 ( PERTH 08159998 ); 5.8 km S of Eneabba Drive on the Brand Highway , E side of the road, 19 Aug. 2020, K. A. Shepherd & C. F . Wilkins KS 1721 ( AD, BRI, CANB, K, MEL, NSW, NY, PERTH 09508066 ); 100 m N on Wells Road on the North West Coastal Highway on W side of road, N of Geraldton , 20 Aug. 2020, K. A. Shepherd & C. F . Wilkins KS 1722 ( PERTH 09508074 ); 8 km N on Yerina Springs Road from Port Gregory Road, N of Northampton , 20 Aug. 2020, K. A. Shepherd & C. F . Wilkins KS 1723 ( BM, CANB, MEL, NSW, P, PERTH 09508082 ); 100 m S of Ogilvie West Road on Yerina Spring Road , NW of Northampton, 20 Aug. 2020, K. A. Shepherd & C. F . Wilkins KS 1726 ( CANB, DNA, HO, MEL, MSB, NSW, PERTH 09508104 ); 3.2 km W on Binnu West Road from Yerina Spring Road, N of Northampton , 20 Aug. 2020, K. A. Shepherd & C. F . Wilkins KS 1729 ( CANB, MEL, NSW, PERTH 09508090 ); 9.6 km W of the Hawkes Head Lookout Road on the Ajana–Kalbarri Road, E of Kalbarri , 22 Aug. 2020, K. A. Shepherd & C. F . Wilkins KS 1739 ( AD, BM, CANB, MEL, NSW, PERTH 09508112 ) .

Figures

Fig. 10. Geleznowia calycina: (a) single-stemmed habit; (b) flowering heads showing the large pale yellow bracts subtending the inflorescence (some tinged red at the apex) with whorls of grey–green leaves below; (c) dense flowering head post-pollination, highlighting the red bracts and sepals; (d) open flower showing the broad pale yellow sepals with bracts below, bright yellow petals and obovoid stigma at the apex of the style; (e) mixed population of G. calycina (left) with a multiple-stemmed habit and clusters of flowers tinged red, compared with G. uberiflora (right) with single pale yellow terminal flowers. Vouchers: K.A. Shepherd & B.M. Anderson KS 1848 (a); K.A. Shepherd & C.F.Wilkins KS 1723 (b); K.A. Shepherd & J.A. Wege KS1300 (c); A.D. Crawford ADC 1344 (d); A.D. Crawford ADC 1840 (right) and ADC 1839 (left) (e). Photos: K. A. Shepherd (a–c, e); A. D. Crawford (d).

Fig. 8. Distributions of all seven Geleznowia species in Western Australia. The base map is sourced from GADM (ver. 3.6, see https://gadm.org), with grey borders showing IBRA regions and subregions (Department of Agriculture, Water and the Environment 2020).