J OURNAL of the A DELAIDE B G OTANIC ARDENS AN OPEN ACCESS JOURNAL FOR AUSTRALIAN SYSTEMATIC BOTANY flora.sa.gov.au/jabg Published by the STATE HERBARIUM OF SOUTH AUSTRALIA on behalf of the BOARD OF THE BOTANIC GARDENS AND STATE HERBARIUM © Board of the Botanic Gardens and State Herbarium, Adelaide, South Australia © Department of Environment, Water and Natural Resources, Government of South Australia All rights reserved State Herbarium of South Australia PO Box 2732 Kent Town SA 5071 Australia J. Adelaide Bot. Gard. 25 (2012) 55–70 © 2012 Board of the Botanic Gardens & State Herbarium, Government of South Australia © 2012 Department of Environment, Water and Natural Resources, Govt of South Australia Notes on Hibbertia (Dilleniaceae) 7. H. hermanniifolia group (subgen. Hemistemma) from mainly temperate eastern Australia H.R. Toelken State Herbarium of South Australia, DENR Science Resource Centre, P.O. Box 2732, Kent Town, South Australia 5071 e-mail: [email protected] Abstract A taxonomic re-evaluation of the H. hermanniifolia group confirmed that it should be segregated from Hibbertia §Tomentosae Bentham (1863). The following taxa are recognised (newly described ones are in bold): H. acaulothrix Toelken, H. hermanniifolia DC. subsp. hermanniifolia and subsp. recondita Toelken, H. hexandra C.T.White, H. planifolia Toelken, H. reticulata Toelken, H. spathulata N.A.Wakef. subsp. spathulata and subsp. pleioclada Toelken. Desriptions, a key to the species and subspecies as well as illustrations are provided. The vestiture and its developments are reviewed. Keywords: Dilleniaceae, Hibbertia, nomenclature, revision, taxonomy, eastern Australia. Introduction have become available from a number of populations with usually disjunct localities of the characteristic Bentham (1863) included in his taxon Hibbertia ‘Inselberg’ distribution so often found in hibbertias §Tomentosae species of the H. melhanioides and H. along the Great Divide. The present treatment attempted tomentosa groups (Toelken 2010) as well as the H. to evaluate and classify these divergent and disjunct hermanniifolia group, which is here revised. All three populations, as many of them are now represented by together with other groups are now included in Hibbertia several collections to provide a wider range of their local subgen. Hemistemma (Touars) Horn. natural variation. However, this does not apply to three Hibbertia hermanniifolia was the first species of recent collections from the Northern Tableland of New this. It was described by Candolle (1817) based on South Wales which are here included as subspecies of a collection by G. Caley from Dovedale near Sydney, H. hermanniifolia and H. spathulata. They are well out but Bentham and subsequent botanists could not locate of hitherto known distribution ranges of the two species this second Dovedale, so that it was not recollected for and require more field work. 100 years. The second collection of it was only made in 1913 when E. Cheel and J.L. Boorman rediscovered it at Characteristics of the vestiture (cf. vestiture below) what is now Bents Basin along the lower Nepean River again proved a useful tool as in other groups of (cf. Notes under H. hermanniifolia). The species occurs Hibbertia (Toelken 1998, 2000, 2010). Bentham (1863) here only in one very restricted area and this is still the largely followed Candolle (1824) when he placed H. only locality from where typical H. hermanniifolia has hermanniifolia into his §Tomentosae, but Toelken been recorded. (2010) noted that it is distinguished by normal fascicled Approximately thirty years later C.T. White hairs subtended on the upper leaf surface by base cells, collected in southern Queensland and described H. while species of the §Tomentosae, consisting of the H. hexandra in 1942. He considered it to come “between tomentosa and H. melhanioides groups, have rosette-like H. hermanniaefolia and H. velutina R.Br.” (White fascicled hairs or scales. Horn (2005, 2009) had already 1942, p. 201), the latter is here considered to be only established this separation in his molecular analyses of distantly related to the complex as it is part of Hibbertia the genus and family. §Tomentosae, a taxon published by Bentham (1863) Characters without rank (cf. Toelken 2010, p.1). The first collection from Victoria of another species different from H. Leaves. The margins at the base of the leaf lamina continue hermanniifolia was described by Wakefield (1957) into ridges or wings on either side of the persistent as H. spathulata N.A.Wakef. As more collections leaf bases on the branches. The leaves therefore seem from Victoria followed the difference between these sessile, but, similar to the more or less terete petioles two species became less clear, but Willis (1973) in other parts of the subgen. Hemistemma (Thouars) distinguished H. hermanniifolia by its stalked fascicled Horn. They are also often variously elongated, so that hairs and stalked flowers. Since then more collections the petiole is usually ill defined although approximate 55 H.R. Toelken J. Adelaide Bot. Gard. 25 (2011) measurements for this section of the leaf are provided in the descriptions. Species of the H. hermanniifolia group do not develop a marked ridge or wing from the base of the central vein on the undersurface of the leaves, which characteristically continues on the leaf base and the branches of species of the H. tomentosa and H. melhanioides groups (Toelken 2010). No young seedlings have been recorded for this group, but juvenile plants about 10 cm tall, e.g. G.L.Stebbins A-63, NSW224829, without the basal leaves still had a few leaves which had two shallow subterminal teeth/lobes each with an obvious vein-end from a lateral vein more or less at the apex (Fig. 3I, 4B). Adult leaves including those of coppicing branches of H. acaulothrix (e.g. S.Clarke s.n.) are not lobed or toothed as has been recorded for the H. tomentosa and H. melhanioides groups (Toelken 2010). However, the Fig. 1. H. acaulothrix. Adaxial leaf surface covered with erect fascicled leaves of H. reticulata, and rarely also in H. hexandra, hairs with 1-3 arms and subtended by a collar of ridged base cells. — exhibit short veinlets from the sinuate supermarginal J.D.Briggs 2080. — Leaf mounted on specimen holder and scanned uncoated using a JEOL Neoscope JCM-5000 Table Top SEM (Nikon, veins to the leaf margins (Fig. 4H), but in contrast to Tokyo) under low vacuum; accelerating voltage 15kV. those from juvenile leaves no obvious vein-ends were observed. These veinlets are also not continuations from surface, but the cover of sessile hairs below them was the lateral veins to the leaf margin (unlike in juvenile still more sparse than in adult leaves as also reported for leaves above), but seem to develop at irregular intervals, the H. melhanioides and H. tomentosa groups (Toelken sometimes even several times, from the supermarginal 2010). between any two connections with lateral veins. No On the adaxial surface of mature leaves at least one such veinlets were observed in any of the other species row of epidermal cells form a collar around the base of of this group even when the often dense tomentum was each hair. The walls of these cells become thickened and removed. often ridged (cf. Fig. 1). They stand out because they Vestiture. All species have predominantly fascicled hairs are often lighter than other green cells, and are similar and in H. reticulata these are overtopped by hooked to the base cells in the H. aspera group (Toelken 1998). simple hairs on proximal margins of the leaf lamina. Often they become very pronounced particularly when These hooked hairs are different to the occasional that leaf is very exposed and the hairs have worn off, so simple hairs on the upper leaf surface of, for instance that eventually some leaves especially of H. hexandra H. acaulothrix (Fig. 1) which usually have a similar are completely covered with such thickened cells. habit (including base cells, cf. below) to those of the However, the base cells of leaves of H. planifolia and H. surrounding fascicled hairs, which have arms varying spathulata are usually bulging similar to ‘goose-bumps’ in number (1−3) and size sometimes even on the same on human skin, but do not develop visibly thickened hair. Leaves of coppicing branches of the same species walls under the persistent dense hairs. They are even (S.Clarke s.n.) have mainly such simple hairs on the less well developed in H. reticulata. adaxial leaf surface, but even here they seem to merge Normally the base cells, which were only observed into few multiangulate fascicled hairs of similar size and on the upper leaf surface, are only slightly raised above habit and are therefore grouped together. the surrounding epidermis and form a bulging or beady These multiangulate fascicled hairs are, however, collar around each hair. On the undersurface of the never found to become rosette-like as in the very similar leaves the stalked hairs, although not unique to this species of the H. melhanioides and H. tomentosa groups, group (cf. H. alopecotis, Toelken 2010), are very well nor could the latter species be shown to have any developed in H. hermanniifolia. The stalk is made up multiangulate hairs, even in juvenile stages, although of cells similar to those of epidermis, which seem to this would have been the obvious derivation of the have formed a protrusion, as the basal tubercle of the rosette-like fascicled hairs (Toelken 2010). fascicled hair is found distally but partly hidden by a The hairs on the upper leaf surface of adult as well layer of epidermis-like cells. The cells of the stalks as juvenile plants, as much as they are known, are often develop thick walls similar to the base cells and become sparse and usually with very few arms. The leaves of filled with an amorphous brown substance, which is seedlings of typical H. hermanniifolia (G.L.Stebbins presumed to be tannins. This development of the stalk A-63, NSW224829) examined had already lost the from the epidermis is further supported by the occasional lowest about fifteen leaves, but still retained a few with presence of sessile fascicled hairs at the base or on the shallow subterminal lobes or rounded teeth. These leaves base of the stalk (Fig. 2) in the typical subspecies of had already a few hairs with short stalks on the abaxial H. hermanniifolia. The development of the stalks has, 56 J. Adelaide Bot. Gard. 25 (2011) Notes on Hibbertia 7 (Dilleniaceae) however, not been studied as no seedlings with very immature leaves could be obtained. Flowers. The terminal flowers are subtended by a bract that is always found immediately below the calyx as in the §Tomentosae but unlike the varying positions in the §Vestitae. The internode below the bract here referred to as peduncle (propodium; Conn 1995) is more or less elongated. In H. hermanniifolia subsp. hermanniifolia the internodes on the branches are usually elongate, so that one can more easily examine the flower buds, which are usually immediately overtopped by continued vegetative growth from the axil of the first leaf at the base of the peduncle. After at least two leaves another flower develops terminally and sympodial growth continues in a similar pattern to that described for H. superans (Toelken 2000, Fig. 11). As the branch thickens, the base of the slender peduncle is often pushed aside, so that it is usually not in a strictly leaf-opposed position, as it is commonly found in other Hibbertia species. In some dried specimens flowers may even seem axillary, but younger buds examined showed more clearly a terminal Fig. 2. H. hermanniifolia subsp. hermanniifolia. Stalked fascicled hairs with sessile fascicled hairs at or sometimes on the base of the stalk origin of the flowers. ×30. — I.R.Telford 7225. The position of the flower is not quite so obvious in most specimens of H. hermanniifolia subsp. recondita, The stamens are variable in size and some of them H. aculothrix, H. planifolia and H. spathulata partly are reduced to staminodes with more or less reduced, because of lateral displacement, but mainly as a result indehiscent anthers or anthers lacking (Fig. 3D, 4D). The of abbreviated internodes between the clustered leaves number of staminodes varies from one plant to another. on distal branches, so that it becomes impossible to There are often more staminodes, or all of them occur in determine the position of the flowers. the lateral bundles. The fertile stamens are also variable In H. hexandra, and to lesser degree also in H. in length and especially the central one (rarely few) of reticulata, flowers are also borne on elongated branches, the dorsal bundle is usually considerably longer than but here the flowers are found terminal on fascicled others. The filament of the central stamen, as well as the lateral branches toward the apex of the main branches. anthers of the other stamens of that bundle are usually Usually there are few such branches scattered along the leaning over the ovaries, so that the large anther is upper reaches of larger branches. However, on actively usually placed between the bases of the two erect styles. growing branches the flowers occur in distal groups of Other anthers of different sizes are normally tightly these fascicled branches, at first on short lateral branches arranged around this to form a broadening column with few leaves. Higher up these leaves decrease in downwards. The rotate flower with a central anther number and size acropetally to two caducous leaflets column is unusual in Hibbertia and resembles those of (up to 4 mm long) at the base of the peduncle in the Solanum flowers, except that two styles on either side axil of the uppermost leaves. These axillary flowers in of the anthers are at first erect, but become strongly such continuous inflorescences are born at successive recurved, so that the stigmas are directed dorsally and nodes unlike the terminal flowers, which develop at well away from the anthers. The filaments are usually intervals of several nodes/leaves, as also recorded for slightly longer in H. elongata and H. spathulata, and the §Tomentosae (cf. Toelken 2010, Fig. 12A). as a result the anthers in dried material are rarely close Stamens. Five more or less clear bundles of stamens together. A similar arrangement, though not as tightly are arranged around the ovaries in most flowers. These grouped, is also found in H. hexandra. Here, however, bundles seem to agree with the stamen trunks, which the styles remain more or less erect, as in some tropical divide up to provide vascular traces to the individual species of the H. melhanioides group, where the anthers stamens (Wilson 1965, Tucker & Bernhardt 2000), as are also often unequally long (e.g. H. bicarpellata, there are usually the same number of stamens in each Toelken 2010), but are not forming a central column. A bundle. Sometimes, particularly in flowers with more similar spacial arrangement of the anthers and styles as than 15 stamens, the dorsal two bundles (in relation to in the H. hermanniifolia complex is also often observed the bract) are more clustered, so that these bundles are in the H. stricta complex (Toelken 2010b) except here less easily distinguished. However, the few stamens of all the stamens are in one dorsal cluster. This spatial H. hexandra are also not always clearly grouped (Fig. arrangement seems to represent a distinct feature of the 4E). pollinating syndrome. 57 H.R. Toelken J. Adelaide Bot. Gard. 25 (2011) Ovary. In all species of the H. hermanniifolia group the leaves of H. reticulata); juvenile leaves broader, with a ovaries are densely hirsute with erect-spreading fascicled pair of rounded subterminal teeth/lobes but not repeated hairs. Similar to the species of the H. melhanioides in coppicing leaves, with subterminal and terminal group (Toelken 2010a) the style bases (Fig. 4F, 4J), vein-ends. Flowers terminal on more or less reduced particularly of H. hexandra and H. reticulata, are often short shoots, hence sometimes appearing axillary, ± to various degrees hairy. Other specimens of both the stalked, with narrowly ovoid to ellipsoidal buds; bracts species vary to glabrous. linear to linear-spathulate. Calyx with lobes subequal to dissimilar, acute to cuspidate, ± ridged distally, Seeds. The more or less lobed cup-shaped apex of the with multiangulate fascicled hairs outside and inside. arils varies only slightly in length, similar to those in the Petals entire to scarcely bilobed. Stamens 6−25 (–48), H. melhanioides and H. tomentosa groups. in groups around the ovaries, with varying numbers Taxonomy of staminodes; anthers linear-obloid, 1−3 longer ones. Pistils 2; ovaries each with 2 (−6) basal/ parietal ovules, Historically Hibbertia hermanniifolia was placed densely fascicled-tomentose; styles erect on either side by Candolle (1817, 1824) into a group of species of stamens, with style base ± hairy. distinguished by the stamens being arranged around the densely hairy ovaries. Bentham (1863) largely Notes. This distinctive group is restricted to temperate agreed and named the group Hibbertia §Tomentosae. areas (except H. reticulata) along mainly the eastern In that group Toelken (2010a) segregated the tropical escarpment of the Great Divide in Queensland, New species of the H. melhanioides and H. tomentosa groups South Wales and Victoria, where the present taxa seem to from the H. hermanniifolia group mainly on the basis represent local relics judging by their often very disjunct rosette-like hairs to scales as opposed to fascicled hairs, restricted distribution. Widely separated populations of, respectively. for instance, the subspecies of H. hermanniifolia and H. spathulata indicate that some of these species once were H. hermanniifolia group more widespread. Shrubs with terete branches covered with raised leaf bases (not ridged or winged). Vestiture of multiangulate Affinities. Bentham (1863) placed H. hermanniifolia as fascicled hairs with their base often surrounded by base the first species into the Hibbertia §Tomentosae next to cells with thickened lateral walls, or at first only radial the H. §Vestitae and H. §Ochrolasiae and included them walls thickened. Leaves cuneately linear-triangular in his sect. Euhibbertia (=sect. Hibbertia), because the to obovate, with lamina gradually constricted into stamens in their flowers are arranged around the ovaries. indistinct petiole, truncate to cuspidate with scarcely Based on molecular evidence all of these three groups are recurved apex, sometimes emarginate, flat and with now separated from the species of H. subgen. Hibbertia slightly revolute margins, lateral and intramarginal veins and placed in the H. subgen. Hemistemma (Horn 2005, incompletely or not visible (except for mainly young 2009). Furthermore Horn (2009) demonstrated that Key to the species and subspecies based largely on floral characters 1. Lower surface of leaves without stalked fascicled hairs 2. Peduncle > 5 mm long 3. Stamens 32−48; upper leaf surface with distinct rugose-reticulate veins; northern Qld ................. H. reticulata 3: Stamens 6 (−12); upper leaf surface with only a distinct mainly central vein; Qld, NSW ............... H. hexandra 2: Peduncle ≤ 5 mm long 4. Undersurface of leaves not visible between revolute margins and central vein, which is ± obscured by dense vestiture; shrublets decumbent to prostrate; NSW ....................................... H. planifolia 4: Undersurface of leaves well exposed between revolute margins and obvious central vein; shrubs erect to spreading 5. Bracts 7.8−10.4 mm long; NSW ........................................................ H. acaulothrix 5: Bracts 3.4−5.4 mm long 6. Leaves with scarcely recurved margins; upper leaf surface puberulous to tuberculate or glabrescent; Qld, NSW ........................................................................ H. hexandra 6: Leaves with obviously revolute margins; upper leaf surface tomentose, rarely pubescent 7. Hairs on upper leaf surface with up to 8–10 (–14) arms; NSW ............... H. spathulata subsp. pleioclada 7: Hairs on upper leaf surface with 2–6 (–8) arms 8. Undersurface of leaves with some shortly stalked hairs; hairs on upper surface of leaves with 2 or 3 (–5) arms; NSW, Vic. ...................................... H. hermanniifolia subsp. recondita 8: Undersurface of leaves with sessile hairs; hairs on upper surface of leaves with 6–8 (–14) arms; Vic. ...................................................... H. spathulata subsp. spathulata 1: Lower surface of leaves with some stalked fascicled hairs 9. Stalk of fascicled hairs 0.35−0.7 mm long; peduncle (9.4–) 10−15 (–17.7) mm long; stamens and staminodes 18–28; NSW ........................................... H. hermanniifolia subsp. hermanniifolia 9: Stalk of fascicled hairs 0.1−0.3 mm long; peduncle (2–) 3−7 (–9) mm long; stamens and staminodes up to 15; NSW, Vic. ........................................................ H. hermanniifolia subsp. recondita 58 J. Adelaide Bot. Gard. 25 (2011) Notes on Hibbertia 7 (Dilleniaceae) the H. hermanniifolia group formed a separate clade H. melhanioides and H. tomentosa groups, while the and one of three sister clades is the H. aspera group. H. §Vestitae from temperate eastern Australia are part Toelken (2010a) divided the H. §Tomentosae into the of one of the other two clades separate from the H. H. melhanioides and H. tomentosa groups and here hermanniifolia and H. aspera clades (cf. above). Both distinguishes the H. hermanniifolia group mainly on the the H. §Ochrolasiae and H. §Vestitae have more strongly basis of their rosette-like hairs or scales as compared to revolute leaves and a more well developed central vein. multiangulate fascicled hairs respectively. Species of the H. hermanniifolia group are here The H. hermanniifolia and H. aspera groups, which alphabetically arranged in order to avoid renumbering had previously been widely separated, because of the them once the whole genus has been revised. different arrangement of the stamens, now demonstrate Hibbertia acaulothrix Toelken, sp. nov. one of several cases in Hibbertia where Horn (2009) A H. hermanniifolia floribus plus minusve sessilibus could show a switch from the original stamens around pilisque faciculatis sessilibus; a H. spathulata pilis the ovaries to stamens in one group to one side of the simplicibus foliorum paginis supernis bracteisque ovaries in another. Morphologically this affinity is longioribus calice differt. supported because, in contrast to the other two sister Typus: New South Wales, 10 km N Bemboka, clades, both groups have flat leaves with a scarcely 16.x.1986, J.D.Briggs 2080 (holo.: NSW; iso.: MEL; developed central vein and develop similar base cells CANB – n.v.). (cf. vestiture) around the tubercles of the fascicled hairs Hibbertia hermanniifolia auct. non DC.: G.J.Harden & J.Everett in G.J.Harden, Fl. New South Wales 1: 300 on the adaxial leaf surfaces. While most of the species (1990), pro parte. of the H. melhanioides and H. tomentosa groups have toothed/lobed leaves of juvenile and regenerating shoots Shrubs up to 1.1 m tall, with several stems, erect; (Toelken 2010a), the H. hermanniifolia group were here branches bluntly ridged to scarcely flanged from the found to develop toothed leaves only on seedlings and leaf base, tomentose or pubescent. Vestiture ± persistent, all leaves in the H. aspera group are entire (Toelken dense multiangulate fascicled hairs sometimes 1998). overtopped by larger broad-based ones on whole plant; The H. §Ochrolasiae from Western Australia form on branches dense, with erect to spreading multiangulate according to Horn (2009) part of a sister clade to the fascicled hairs (6−9 often unequal arms) overtopped by Key to species and subspecies based on vegetative characters As the revolute margins and central vein often bear different hairs they are here, for convenience, excluded in the use of ‘undersurface of leaves’, which are therefore not synonymous with the abaxial leaf surface (cf. Toelken 1998, 2000). References to the central vein always apply to the abaxial leaf surface unless specified, because here it is usually clearly visible. 1. Upper leaf surface with ± rugose-reticulate veins, with hooked simple hairs towards the proximal margins overtopping fascicled hairs; Qld .............................................................. H. reticulata 1: Upper leaf surface without veins or with incomplete vein network, without hooked hairs overtopping fascicled hairs 2. Stalked fascicled hairs present on branches and veins of abaxial surface of leaves .................. H. hermanniifolia 3. Stalk of fascicled hairs 0.35−0.7 mm long; peduncle (9.4–) 10−15 (–17.7) mm long; NSW .......... ............................................................. H. hermanniifolia subsp. hermanniifolia 3: Stalk of fascicled hairs 0.1−0.3 mm long; peduncle (2–) 3−7 (–9) mm long; NSW, Vic ............. .................................................................. H. hermanniifolia subsp. recondita 2: Stalked fascicled hairs absent on branches and leaves 4. Upper leaf surface sparsely hairy, becoming tuberculate or glabrescent; arms of hairs scarcely longer than basal tubercle; leaf lamina (4.6–) 6−18 (–22.8) mm broad; Qld, NSW ......................... H. hexandra 4: Upper leaf surface tomentose to hirsute; arms of hairs at least twice as long as basal tubercle; leaf lamina 1–5.4 (–8.3) mm broad 5. Upper leaf surface hirsute with arms of larger flaccid hairs 2−4 times longer than those on shorter hairs (particularly on flanks of cuneate base), fascicled hairs with 2 or 3 (–5 on margins) arms,not persistent; NSW .................................................................... H. acaulothrix 5: Upper leaf surface velvety to tomentose with arms of larger erect-spreading hairs scarcely longer than those of shorter ones, fascicled hairs with (3) 4−6 (–12 on margins) arms, persistent 6. Central vein ± obscured by dense vestiture; undersurface of leaves not visible between revolute margins and central vein, shrublets decumbent to prostrate; NSW ............................ H. planifolia 6: Central vein obvious; undersurface of leaves visible between revolute margins and central vein; shrublets erect to spreading .......................................................... H. spathulata 7. Hairs on upper leaf surface with up to 8–10 (–14) arms; NSW ............... H. spathulata subsp. pleioclada 7: Hairs on upper leaf surface 2–6 (–8) arms 8. Undersurface of leaves with some shortly stalked hairs; hairs on upper surface of leaves with 2 or 3 (–5) arms; NSW, Vic. ...................................... H. hermanniifolia subsp. recondita 8: Undersurface of leaves with sessile hairs; hairs on upper surface of leaves with 6–8 (–14) arms; Vic. ...................................................... H. spathulata subsp. spathulata 59 H.R. Toelken J. Adelaide Bot. Gard. 25 (2011) Fig. 3. A–D H. acaulothrix: A flowering branch ×1; B leaf from below ×3; C transverse section through mid-leaf ×9; D flower with calyx and corolla removed to show erect position of stigmas, 1 larger stamen and several staminodes ×10. E–G H. spathulata subsp. spathulata: E branch ×1; F leaf from below ×4; G transverse section through mid-leaf ×10. H–K H. hermanniifolia subsp. hermanniifolia: H flowering branch ×1; I juvenile leaf from above ×4; J transverse section through mid-leaf ×8; K seed ×10. L–N H. planifolia: L branch ×1; M leaf from below ×4; N transverse section through mid-leaf ×9. — A–D J.D.Briggs 2080; E–G N.G.Walsh 2209; H, J, K I.R.Telford 7225, I G.L.Stebbins A-63; L–N R.Coveney NSW 127188. or with scattered larger, often almost stalked broad-based to spreading multiangulate broad-based fascicled hairs ones (11−16 usually unequal arms); on leaves above (7−14 unequal to subequal, antrorse to radial arms) over moderately dense, spreading to antrorse, appearing dense fine narrow-based multiangulate fascicled hairs sessile (basal tubercle surrounded by base cells) (5−10 subequal arms, but not always distinguishable); on subequal multiangulate fascicled hairs (1−3 usually bracts above dense, with erect hairs (1−3 often unequal unequal arms or up to 5 on margin and often increasing arms), below dense, with erect multiangulate fascicled 2−4 times the length towards the flanks) wearing off hairs (6−9 subequal arms) or rarely with some broad- sooner or later; on leaves below scattered larger erect based ones (8−11 subequal arms) mainly on the central 60 J. Adelaide Bot. Gard. 25 (2011) Notes on Hibbertia 7 (Dilleniaceae) vein; on outer calyx lobes outside dense, with mainly Conservation status. In Wadbilliga National Park cactiform broad-based fascicled hairs or some erect “several thousand plants scattered over at least 3 ha.” multiangulate narrower-based fascicled hairs towards the were recorded by J.D.Briggs 2080. margins and the apex, inside ± dense antrorse narrow- Diagnostic features. The vestiture of simple or scarcely based fascicled hairs on upper third; on inner calyx lobes branched fascicled hairs on the upper leaf surface is outside ± dense band of cactiform fascicled hairs along similar to that of H. hermanniifolia, but H. acaulothrix the centre becoming smaller laterally and with some is distinguished by the absence of stalked fascicled hairs fascicled cilia on the broad membranous margin, inside and flowers on elongated peduncles. Similarly the latter glabrous. Leaves without axillary tufts of hairs; petiole is distinguished from H. spathulata by the presence of 0−0.3 mm long, indistinct; lamina cuneately oblong- simple hairs on the upper leaf surface, and bracts are oblanceolate, (4.6–) 6−15 (–22.3) × (1.6–) 2.5−4.5 (–5.4) longer than the calyx. mm, usually truncate-emarginate rarely rounded and with scarcely recurved vein-end, gradually constricted Variation. Although the species is at present known into petiole, flat, entire, above distinctly grooved along only from a few well separated localities, the different the central vein and puberulous or sometimes pubescent, populations show little variation. Leaves of young fast- becoming tuberculate, below with central vein broader growing branches have usually predominantly simple and often raised above revolute margins or rarely with hairs on the adaxial leaf surface and these and at least some lateral veins visible on upper half, velutinous, the larger hairs of the abaxial surface are more or less slightly discolourous; juvenile leaves (coppicing) larger antrorse. Since a similar trend was found in both H. but similar to normal leaves, entire, with finer and larger hermanniifolia and H. spathulata the absence of stalked hairs. Flowers single, terminal, sessile or almost so, often fascicled hairs in this species is significant. on lateral shoots, with flower bud ellipsoidal; penduncle Etymology. The epithet “acaulo-thrix”, Greek, “stalkless- ± absent; bracts linear-oblanceolate to linear-spathulate, hair” (noun in apposition) refers to the sessile hairs of 7.8−10.4 × 1.9−2.4 mm, usually longer than outer this species in comparison to those of the very similar calyx, pointed, ridged, below pubescent to tomentose or H. hermanniifolia. velutinous, above tomentose. Calyx with lobes slightly unequal but forming a range; outer calyx lobes (3/2) Specimens examined linear-lanceolate to -oblanceolate, (5.4–) 6−6.8 (–7.2) new South wAleS: S.Clarke s.n., Water Board land off × 2.5−2.8 mm, scarcely longer than inner ones, pointed Long Nose Point Road, 27.iii.2001 (WOLL); M.D.Crisp 2235 & I.R.Telford, 3.3 km E Mt Coricudgy, 27.x.1976 to acute, ridged, outside stubble-like to velutinous along (CANB); P.Gilmour WO33, Brogo River near Yankees Creek, margins, inside puberulous to pubescent on the upper Wadbilliga National Park, 1.iv.1983 (CANB); D.J.McGillivray third and the margins; inner calyx lobes (2/3) oblong- 1605, between Kekeelbon Mts and Mt Coricudgy, 13.ii.1966 ovate, (4.2–) 4.5−5.5 (–6.3) × 3.1−3.8 (–4.1) mm, acute (NSW); D.Tilley WOLL 2113, Wattle Ridge, Hilltop, iii.82 to cuspidate, scarcely ridged, stubble-like to pubescent (WOLL); T. & S.Whaite 3275, Kekeelbon Mts, 1.ix.1969 on a central band and with fascicled cilia on broad (NSW). membranous margin, inside glabrous. Petals obovate, (7.4–) 7.8−9.5 (–10.2) × (4.3–) 5.0−7.0 (–8.8) mm long, Hibbertia hermanniifolia DC. scarcely bilobed. Stamens 9−13 (plus some staminodes), Regn. Veg. Syst. Nat. 1: 431 (1817), as “hermanniaefolia”; subequal except for 1 (–3) longer ones, in ± 5 bundles Prodr. 1: 75 (1824), as “hermanniaefolia”; Benth., Fl. Austral. 1: 30 (1863), as “hermanniaefolia”; F.Muell., around the ovaries; filaments narrow throughout, 1.3−1.8 Syst. Cens. 1: 2 (1882), as “hermannifolia”; C.Moore, mm long, slightly connate basally; anthers narrowly Census Pl. New South Wales 1 (1884), as “hermannifolia”; obloid, rarely obloid-ovoid, 1.1−1.3 mm long, 1 (–3) C.Moore & Betche, Handb. Fl. New South Wales 10 longer ones (1.8−1.9 mm long) in the centre, abruptly (1893), as “hermannifolia”; W.A.Dixon, Pl. New South constricted above and below. Pistils 2; ovaries obovoid, Wales 29 (1906), as “hermannifolia”; N.C.W.Beadle et each with 2 (–4) ovules above one another, hirsute, al., Vasc. Pl. Sydney edn 1: 194 (1963); J.H.Willis, Handb. style attached to dorsal apex of ovary then ± erect such Pl. Victoria 2: 386 (1973); G.J.Harden & J.Everett in G.J.Harden, Fl. New South Wales 1: 300 (1990); Toelken that stigmas are lateral to the upper part of the longer in N.G.Walsh & Entwisle, Fl. Victoria 3: 305 (1996). — ± incurved anther(s). Fruiting peduncle not elongating. Type citation: “in montibus Novae Hollandiae. Caley. (v. Seeds not seen. Flowering: usually October−April. Fig. s. sp. in h. [dried specimen seen in herb.] Lambert)”. Type: 1, 3A–D. “m Lambert” (holo.: G-DC). — see Notes below. Distribution and ecology. Recorded associated with Shrubs 0.8−1.5 (–3) m tall, with several stems, sedimentary rocks from very different woodlands, spreading; branches bluntly ridged to flanged from the such as with Eucalyptus sieberi L.A.S.Johnson or E. leaf base, hispid to pubescent. Vestiture ± persistent, multicaulis Blakeley or associated with Allocasuarina usually dense multiangulate fascicled hairs over whole littoralis (Salisb.) L.A.S.Johnson, Corymbia gummifera plant but overtopped by few to many stalked ones (Gaertn.) K.D.Hill & L.A.S.Johnson and Leptospermum mainly on branches, leaves below and outside of outer trinervium (Sm.) Joy Thomps.; found in few widely calyx lobes; on branches dense, with erect to spreading separated localities in New South Wales (CT, CC, SC). multiangulate narrow-based fascicled hairs (7−10 often 61 H.R. Toelken J. Adelaide Bot. Gard. 25 (2011) unequal arms) of different sizes overtopped by more or upper half; inner calyx lobes (2/3) ovate, 5.1−6.4 × less spreading stalked broad-based ones (8−16 (–25) 3.6−4.2 mm, acute to obtuse, without ridge, tomentose usually unequal arms) often near nodes; on leaves above to almost stubble-like towards the margins and with sometimes wearing off but retaining basal tubercle fascicled cilia along the membranous margins, inside and ring of base cells, moderately dense, with few to glabrous to puberulous towards the apex. Petals oblong- many erect simple to erect or spreading, often antrorse oblanceolate, rarely obovate, 6.8−9.4 × 4.7−6.8 mm, multiangulate narrow-based fascicled hairs (2, 3 often scarcely bilobed. Stamens 9−24 (plus few staminodes) unequal arms, rarely up to 5 arms on the Northern unequal, in 5 bundles around the ovary; filaments Tableland, New South Wales) of different sizes, filiform, 1.1−1.5 mm long, scarcely basally connate; becoming larger (up to 12 arms) towards the flanks of anthers narrowly obloid, 1.2−1.5 mm long, at least the recurved margins; on leaves below dense, with fine central one longer (1.8−2.1 mm), abruptly constricted spreading multiangulate narrow-based fascicled hairs of above and below, straight. Pistils 2; ovaries obovoid, different sizes (7−10 subequal arms) overtopped by few each with 2−4 ovules in pairs above one another, hirsute, to many stalked broad-based ones (11−16 (–23) usually with style attached to dorsal apex then more or less unequal arms) mainly restricted to veins and the revolute erect positioning stigmas laterally to the upper part of margins; on bracts above moderately dense, with the longer anther. Fruiting peduncle slightly elongated, erect multiangulate narrow-based fascicled hairs (3−5 ± recurved. Seeds obovoid to almost spherical, 2.5−2.9 subequal arms), below like leaves dense, with spreading × 2.9−3.25 mm, shiny-black; aril with fleshy base multiangulate narrow-based fascicled hairs of different expanding into ± 3 or 4 lobed cup-like membrane sizes overtopped by few larger stalked broad-based clasping the lower half of seed. fascicled hairs; on outer calyx lobes outside ± dense, Diagnostic features. Pedunculate flowers and the with spreading, often short-branched multiangulate, uniquely stalked fascicled hairs with their commonly often broad-based fascicled hairs of different sizes brown stalks, particularly on the undersurface of leaves, overtopped by ± stalked usually broad-based fascicled are distinctive of this species. Some of the latter hairs hairs scattered over the surface but becoming shorter were always present, although they were found to towards the margins, inside with ± dense antrorse grade into normal fascicled hairs with broad and thin narrow-based multiangulate fascicled hairs on often tubercles. The discolourous leaves of this species are more than the upper half; on inner calyx lobes outside ± usually accentuated by the sparse tomentum of simple dense, with spreading but short-branched multiangulate, and fascicled hairs (with few arms) on the upper leaf often broad-based fascicled hairs of different sizes surface, which is distinctive of the two subspecies. overtopped by scattered larger broad-based ones mainly The truncate leaves with a similar fascicled tomentum along the centre and with narrow-based fascicled cilia of vegetative material of some Asterolasia species on the membranous margins, inside glabrous except for (Rutaceae) have at times been confused with specimens a few antrorse fascicled hairs towards the apex. Leaves of Hibbertia hermanniifolia. without axillary tufts of hairs; petiole 0.2−1.4 mm long, indistinct; lamina obovate to oblanceolate, 3.3−30.2 Variation. The plants from the type locality, Bents Basin × 1.8−10.2 mm, usually ± truncate and with short on the Nepean River, have particularly large leaves and recurved apex, usually entire, flat to ± folded lengthwise hairs. They become smaller southwards in southern or grooved along the central vein and pubescent to New South Wales and in Victorian plants often appear sparsely velutinous, below with narrow central vein similar in size to those of H. spathulata. Throughout the rarely raised above revolute margins and rarely with distribution of the species the stalk of stalked hairs is usually incomplete lateral veins visible towards the more or less intensely brown-coloured, while the short apex, and hirsute to velutinous, discolourous; juvenile base of sessile hairs is without colour. The same applies leaves obtriangular, with three subequal terminal teeth, to less obviously stalked rosette-like fascicled hairs in H. flat, covered at least at first with fine sessile fascicled alopecota from the Northern Territory and Duretto and hairs, each with few arms on the upper surface and with Ladiges (1997) also report “often heavily pigmented” more on the undersurface. Flowers single, terminal or stalks of species of Boronia. often not leaf-opposed but beside petiole, with ovoid Similarly in the northern specimens the flowers are to ellipsoidal buds; peduncle thread-like but firm, (3–) usually overtopped by axillary vegetative growth so that 4−12 (–17.7) mm long, terete; bracts spathulate to they appear to be axillary, because they are commonly linear-oblanceolate, 3.3−5.5 × 0.8−2.9 mm, longer than pushed sideways. These flowers are rarely clearly leaf- half the outer calyx, leaf-like with revolute margins and opposed, nor do they show rudiments of axillary short central ridge, sparsely velutinous above and ± hirsute shoots as it is observed in the tropical species. The below. Calyx with lobes slightly unequal but forming specimen (G.L.Stebbins A-63, NSW224829) is the only a range; outer calyx lobes (3/2) linear-lanceolate to one that has a number of terminal flowers but then the lanceolate, 5.1−7.4 × 1.8−3.1 mm, slightly longer than peduncles are typically longer than 9 mm. inner ones, ridged, outside hirsute rarely velutinous, The number of stamens and staminodes of a flower inside pubescent with few antrorse fascicled hairs on were found to vary from one population to another. 62 J. Adelaide Bot. Gard. 25 (2011) Notes on Hibbertia 7 (Dilleniaceae) Notes. In the protologue Candolle (1817) cites the type long. Flowers usually overtopped by axillary vegetative as “in montibus Novae Hollandiae” and being a Caley growth displacing the peduncle to a lateral position; specimen in Herb. Lambert, but none of this detail is peduncles (9.4–) 10–15 (–17.7) mm long; bracts found on the holotype specimen. A sheet (BM 834625) oblanceolate-spathulate, 1.9–2.9 mm broad. Stamens and of this species contains four branches, one marked “in staminodes 18–28. Flowering: September–February. Fig. the Mountains − Caley” and a second “Dove Dale Oct 2, 3H–K. 30-1807” in a different later handwriting (Bentham?). Distribution and ecology. Grows on sandy soils or rocky There is no indication whether they belong to the same slopes associated with sandstone cliffs in sclerophyll collection or two different ones and which specimen eucalypt forest of mainly Corymbia eximia (Schauer) then belongs to which collection, except that all four K.D.Hill & L.A.S.Johnson and Eucalyptus pilularis Sm. specimens are so similar that it seems likely that they in New South Wales (CC). are from the same plant of the only population this, the typical subspecies is known from. It could also not be Conservation status. Locally common in conserved established whether or not the Caley specimen/s in the area. British Museum are duplicates, i.e. possible isotypes Variation. Judging by a number of specimens collected particularly as Vallance et al. (2001) stated “Caley at this restricted population it would seem that this visited and revisited the place over the years.” subspecies shows little variation. Even juvenile plants The R.Brown specimen in Kew Herbarium was noted are remarkably similar to adult plants (see description). to have been received in 1880 (after flora treatment of Bentham, 1863) and is accompanied by a collector’s Specimens examined label but without a number added subsequently by J.J. new South wAleS (all from Bents Basin): D.Benson Bennett. This was found typical of that batch judging et al. NSW 167541, 20.ii.1984 (NSW); R.Brown [G.Caley] s.n., Dove Dale, 30.x.1807 (BM, K); E.Cheel & by other specimens. It is also annotated “Dove Dale Oct J.L.Boorman NSW 33652, 10.ix.1913 (BM, BRI, CANB, 30-1807” and thus unlikely to be collected by Brown. K, MEL, NSW); J.Pulley 897 & I.R.Telford, 29.ix.1971 Vallance et al. (2001) suggested he and G.Caley could (CANB); G.L.Stebbins A-63, 20.xi.1974 (CANB, NSW); only have visited Dovedale on 23.x.1803, but no such I.R.Telford 7225, 27.x.1978 (CANB). specimen could be located. A note concerning the exact locality of this Dovedale, Hibbertia hermanniifolia subsp. recondita Toelken, which Bentham was unable to resolve, was found in the subsp. nov. cover of the species in the National Herbarium of New A subspecie typica pedunculis brevibus usque ad 9 mm South Wales (NSW): longis, pilis brevibus foliisque parvis differt. “The type of this species was collected by Caley at Typus: Victoria, Mt Elizabeth 2, R.D.Hoogland 11911, 26.xi.1970 (holo.: AD98040099; iso.: BRI, MEL, ‘Dovedale’ (cf. Benth. [1863]) or, as cited by DC., ‘in NSW; A, B, BM, CANB, E, G, HBG, K, L, NE, OKLA, montibus Novae-Hollandiae’. This Dovedale (see label PERTH, UC – n.v.). on specimen of leaf from Nat. Herb. Vict. Mel.) is at Hibbertia hermanniifolia auctt. non DC.: Willis, Handb. Pl. Bent’s Basin, Nepean River. There was also another early Victoria 2: 386 (1973), pro parte; G.J.Harden & J.Everett property named ‘Dovedale’ on the Bellinger River (see in G.J.Harden, Fl. New South Wales 1: 300 (1990), pro Balliere’s N.S.W. Gazetteer & Road Guide compiled by parte; Toelken in N.G.Walsh & Entwisle, Fl. Victoria 3: R.P.Whilworth 1866) & and this must have lead Moore 305 (1996), pro parte. & Betche [1893 and Dixon 1906] to cite ‘Clarence River Leaves oblanceolate to obovate, (3.3–) 4–8.5 (–13.5) District’ in the Fl. N.S.W. Caley’s collections were from × (1.8–) 2.5–4 (–5.6) mm; stalk of fascicled hairs 0.1–0.3 the Central Coast & too early for a N. Coast area. N.C.F. mm long. Flowers not overtopped by axillary vegetative [Neridah C. Ford] 6/1957”. growth, or if displaced by some growth then peduncle The species consists of many, often disjunct leaf-opposed; peduncles (2–) 3–7 (–9) mm long; bracts populations which are here grouped into two subspecies. linear-oblanceolate, 0.9–1.4 mm broad. Stamens and The subsp. hermanniifolia is, however, restricted to a staminodes up to 15. Flowering: August–February. small area at Bents Basin. Distribution and ecology. Grows on dry slopes or on Key to subspecies skeletal soils in rock outcrops, often on granite, in shrub 1. Stalk of fascicled hairs 0.35−0.7 mm long; peduncle land to sclerophyll forest in southern New South Wales (9.4–) 10−15 (–17.7) mm long; combined stamens and (NT, SC) and north-eastern Victoria (EHL, EG). staminodes 18–28; NSW ....... subsp. hermanniifolia 1: Stalk of fascicled hairs 0.1−0.3 mm long; peduncle (2–) Conservation status. Locally common on Mt Poole, 3−7 (–9) mm long; combined stamens and staminodes up New South Wales (J.D.Briggs & D.E.Albrecht 1957A) to 15; NSW, Vic. .................. subsp. recondita and Mt Elizabeth 2, Victoria (F.E.Davies 640). Variation. Specimens from widely separated localities Hibbertia hermanniifolia subsp. hermanniifolia. on the Northern Tablelands are remarkably similar to the Leaves broadly obovate, (5.6–) 7–18 (–30.2) × (2.2–) southern populations of this subspecies. Even the number 30–75 (–10.2) mm; stalk of fascicled hairs 0.35–0.7 mm of stamens, the most critical character that separates 63