CSIRO PUBLISHING Australian Systematic Botany, 2013, 26, 167–185 http://dx.doi.org/10.1071/SB12037 Karrabina gen. nov. (Cunoniaceae), for the Australian species previously placed in Geissois, and a synopsis of genera in the tribe Geissoieae H. C. F. Hopkins A,D, A. C. Rozefelds B and Y. Pillon C A c/o Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK. Queensland Museum, GPO Box 3300, South Brisbane, Qld 4101, Australia. Email: andrew.rozefelds@qm.qld.gov.au C Tropical Conservation Biology and Environmental Science program, University of Hawai‘i at Hilo, 200 West Kawili Street, Hilo, HI 96720, USA. Email: pillon@hawaii.edu D Corresponding author. Email: h.fortune-hopkins@kew.org B Abstract. Comparison of the four species groups comprising the tribe Geissoieae (Lamanonia in South America, Geissois sensu stricto from islands in the south-western Pacific, and Pseudoweinmannia and Geissois in Australia) showed differences in the stipules, inflorescences and flowers. Molecular phylogenetic analyses suggested that Geissois sensu lato is paraphyletic, with Australian Geissois being closer to Pseudoweinmannia than to New Caledonian Geissois. The name Karrabina is published to accommodate the Australian species formerly placed in Geissois, namely K. benthamiana Rozefelds & H.C.Hopkins and K. biagiana Rozefelds & H.C.Hopkins. Distinguishing characters for Karrabina are: four lateral stipules per node that vary from free to largely fused between and sometimes across the petioles; inflorescences with a variable number of metamers and racemes, with both median and lateral modules that always lack a median raceme; and pale flowers with five or six calyx lobes and shortly exserted stamens. Geissois s.s. differs in having: two intrapetiolar stipules per node; inflorescences consisting of few metamers and few racemes, with the modules being always lateral and possessing a median raceme; flowers with four red calyx lobes and prominently exserted stamens. The generic limits of Lamanonia and Pseudoweinmannia remain unchanged. Received 11 December 2012, accepted 22 April 2013, published online 18 October 2013 Introduction One of the well supported clades in Bradford and Barnes’s (2001) phylogeny of the Cunoniaceae comprised three genera, namely Lamanonia Vell. (five species from South America), Pseudoweinmannia Engl. (two species from eastern Australia) and Geissois Labill. s.l., which together form the tribe Geissoieae. As traditionally circumscribed, Geissois s.l. consists of two distinct groups of species; ~19 red-flowered taxa (Geissois s.s.) occur in various types of vegetation, including forest on montane islands in the south-western Pacific (Fiji, Vanuatu, New Caledonia, and Vanikoro in the Solomon Islands) and two creamish-flowered species, G. benthamiana F.Muell. and G. biagiana (F.Muell.) F.Muell., which grow in rainforest in eastern Australia (Bradford and Barnes 2001; Schimanski and Rozefelds 2002; Hopkins 2006). Overviews of the family by Dickison and Rutishauser (1990), Hufford and Dickison (1992) and Bradford and Barnes (2001), and morphological studies of the two groups within Geissois s.l. (Schimanski and Rozefelds 2002; Hopkins 2008), have all questioned the present circumscription of Geissois and suggested that relationships within the Geissoieae need clarification. Journal compilation
CSIRO 2013 In the present paper, we address the following questions: what are the phylogenetic relationships among the four species groups in the tribe Geissoieae; and what morphological characters define each of these groups? Because we conclude that each of the four groups should be recognised as a separate genus, we publish the new generic name Karrabina in the Taxonomy section, together with synopses for Geissois s.s., Lamanonia and Pseudoweinmannia. Except when discussing the historical background, we pre-empt our conclusions by referring to the Australian species previously placed in Geissois s.l. as Karrabina, to avoid repetition of the phrase ‘Geissois in Australia’. Historical background Schimanski and Rozefelds (2002) described the taxonomic history of the two Australian species of Geissois s.l. and the key points are summarised here, to show how views on relationships among the species groups in the Geissoieae have developed and to establish that, if excluded from Geissois, no other generic name is available for the Australian species traditionally placed under this name. www.publish.csiro.au/journals/asb 168 Australian Systematic Botany The name Geissois was originally published by de Labillardière (1824–1825), on the basis of a red-flowered plant from New Caledonia. Several other red-flowered species were described from this island during the 19th century (Brongniart and Gris 1862) as well as from Fiji (Gray 1856) and Vanuatu (Seemann 1865–1873) (see Hopkins 2006). In 1865, Mueller described two species from Australia, which he placed in Geissois, namely G. benthamiana and G. biagiana, and, subsequently, these two taxa have almost always been treated in the Australian literature as belonging in this genus (e.g. Francis and Chippendale 1970; Stanley and Ross 1983; Harden 1990; Hyland and Whiffin 1993; Schimanski and Rozefelds 2002; Boland et al. 2006). The only exceptions have been their occasional placement in a catch-all Weinmannia L. (e.g. von Mueller 1865–1866, 1867–1868; Bailey 1900) or its spelling variant Windmannia P.Browne (Kuntze 1891), sometimes with members of other Australian genera that are now also accepted as distinct from Weinmannia. Taxonomic overviews of the family before 1990, by A. Engler, J. Hutchinson and R. D. Hoogland, all included both Pacific and Australian species in a widely circumscribed Geissois that was distinct from both Lamanonia and Pseudoweinmannia. Engler (1928) placed Lamanonia (as Belangera Cambess.) and Geissois together in the Belangereae, characterising this tribe by an indefinite number of stamens (A¥), and he distinguished between the genera by the number of sepals (six in Lamanonia, four or five in Geissois). He also established the genus Pseudoweinmannia but placed it in the tribe Cunonieae, in which the number of stamens was twice the number of calyx lobes (A = 2K). Hutchinson (1967) also treated Geissois and Lamanonia as distinct although his generic descriptions did not differentiate between them. Both genera were said to have numerous stamens, but they were not adjacent in his key because Lamanonia had more than 20 stamens and, by implication, Geissois had fewer. Hoogland worked extensively on the Cunoniaceae and did much preparatory work for the regional floras of New Caledonia and Australia. His unpublished notes at the Muséum National d’Histoire Naturelle in Paris include floral descriptions for several species of Geissois and, together with his only publication on the genus (Hoogland 1984), they give no indication that he questioned the traditional broad circumscription of this genus. Subsequent publications have, however, drawn attention to the morphological heterogeneity of Geissois s.l. and suggested various possible relationships among the genera that form the Geissoieae. In a study of stipular development throughout the family, Dickison and Rutishauser (1990, fig. 53) suggested that the Australian species of Geissois (‘Geissois biagiana group’) were more closely related to Lamanonia than to the Pacific species (‘Geissois pruinosa group’). Their proposed phylogeny was: G. pruinosa group (G. biagiana group, Lamanonia), with Pseudoweinmannia on the same main branch of their cladogram (Dickison and Rutishauser 1990, fig. 52). They noted that Pseudoweinmannia shares the character of bifid stipules with the G. biagiana group, but is uniquely characterised by its wood, fruits and seeds. In their cladistic analysis of the Cunoniaceae, based on morphological data, Hufford and Dickison (1992) combined H. C. F. Hopkins et al. Geissois and Lamanonia into a single operational taxonomic unit (OTU), although they did not make any formal taxonomic changes. The synapomorphies they identified for Geissois– Lamanonia were apetalous flowers with a large number of stamens and digitately compound leaves, and they noted that segregating Lamanonia on the basis of its hexamerous perianth would leave Geissois without apomorphies. Pseudoweinmannia, because of its non-winged seeds, was separate and sister to Geissois–Lamanonia in their analysis, although they suggested that it might be nested within the latter group. On the basis of both DNA sequence data (rbcL, trnL–trnF) and morphology, Bradford and Barnes (2001) rejected the idea of uniting Geissois and Lamanonia and they placed Geissois, Lamanonia and Pseudoweinmannia together in a clade that they recognised taxonomically as the tribe Geissoieae. Although Geissois had no morphological synapomorphy, they indicated the following apomorphies for the two groups that they recognised within it (with characters unique in the family in italic script): ‘Geissois 1 (Pacific) – Stipules axillary; racemes usually cauliflorous; flowers red. Geissois 2 (Australia) – Foliaceous stipules overlapping across leaf axil; inflorescence module (IM) of multiple metamers; IM medial.’ The relationships indicated in their strict consensus tree (Bradford and Barnes 2001, fig. 4) were: Lamanonia (Pseudoweinmannia (Geissois 1, Geissois 2)); however, Lamanonia was not included in their molecular sampling. Bradford et al. (2004) again treated Geissois in a broad sense but noted its heterogeneity. Molecular phylogenetics To complement the dataset of Bradford and Barnes (2001), new rbcL and trnLc–f sequences were generated for the following three accessions: Geissois hirsuta Brongn. & Gris (Pillon et al. 33 (NOU), GenBank accessions for rbcL: JX236030; trnLc–f: JX236027), Lamanonia ternata Vell. (Lucas 309 (K), JX236032, JX236029) and Schizomeria serrata Hochr. (Chase 1223 (K), JX236031, JX236028). Our dataset included all genera currently recognised in the family Cunoniaceae with the exception of Aistopetalum Schltr. (unplaced to tribe) and Opocunonia Schltr. (Caldcluvieae), which are both endemic to New Guinea and for which it has therefore proved difficult to obtain material for analysis. Best-fit models of DNA substitution for both trnL and rbcL were GTR+g in both according to jModelTest analyses (Posada 2008). A maximum likelihood analysis was run in PHYML 3.0 (Guindon et al. 2010); branch support was assessed with 1000 bootstrap replicates. A Bayesian analysis was run in MrBayes 3.2 (Ronquist and Huelsenbeck 2003), on a partitioned dataset (unlinked), with one million generations and a tree sampled every 1000th generation and an initial burn-in of 100 000 generations. In both Bayesian and maximum-likelihood analyses, we recovered a well supported monophyletic tribe Geissoieae, with the following internal relationships: Lamanonia (Geissois (Karrabina, Pseudoweinmannia)), which received moderate Karrabina gen. nov. (Geissoieae: Cunoniaceae) Australian Systematic Botany bootstrap support and relatively high posterior probabilities (Fig. 1). Geissois s.l. appears therefore to be paraphyletic because the Australian species (Karrabina benthamiana) is closer to the Australian Pseudoweinmannia than it is to the New Caledonian species (Geissois hirsuta). These analyses also gave moderate support to the placement of Gillbeea F.Muell. as sister to the tribe Geissoieae. This genus of three species from Australia and New Guinea shares few morphological characters with the Geissoieae, except for its stipules; it has four free lateral stipules per node (Rozefelds and Pellow 2000; Hopkins and Hoogland 2002), as does Lamanonia (see below), and this character may thus be a synapomorphy for the clade. However, further evidence is needed before including Gillbeea in an expanded tribe Geissoieae, which is a morphologically coherent assemblage as currently circumscribed. Another study with different genes (the nuclear loci ncpGS and PHYC) and a more thorough sampling of Geissois (Pillon 2011) supported the monophyly of this group. G. hippocastanifolia Guillaumin and G. polyphylla Guillaumin (both from New Caledonia) formed a clade sister to the rest of the species from New Caledonia (including G. racemosa Labill., to which the type belongs, and the unusual G. hirsuta, see below) and the single endemic from Vanuatu (G. denhamii Seem.). Morphologically, G. hippocastanifolia and G. polyphylla have several distinctive characters; both have five or usually more leaflets per leaf and large coriaceous, oblong stipules that are flattened around the margins in bud, and in which the inner surface bears abundant small, curled, pale hairs and spherical red glands. Relationships among species in the second clade are ML 97 not clear. All have either three, five or a mixture of three and five leaflets per leaf, but never more. In general, species in this clade have smaller, less coriaceous stipules, but if they are large and leathery, they are cordate not oblong (G. lanceolata (Guillaumin) H.C.Hopkins, G. magnifica Baker f.), or exceptionally long and narrow (G. stipularis A.C.Sm.). Furthermore, the stipules in this clade are not flattened around the margins in bud and their inner surfaces are either glabrous or bear sparse straight hairs and no glands (except in G. hirsuta, where the stipules have indumentum similar to that in G. hippocastanifolia and G. polyphylla). It is likely that the taxa from Fiji and the Solomon Islands belong in this second clade. Geissois hirsuta is unusual because it has some apparently plesiomorphic characters (the leaflet margins in adult plants are toothed, not entire, and the inflorescence is composed of triads, not monads; see below for terminology) as well as other unique features (the pedicels articulate at the base, rather than above it; the capsules are ovoid with circular seeds, rather than ellipsoid with elliptic seeds). However, molecular data put this species in the group of taxa with either three, five or a mixture of three and five leaflets and not in an early divergent position, as its toothed leaflet margins might suggest. Comparative morphology Bradford and Barnes (2001, p. 373) defined the Geissoieae as follows (with synapomorphies for the tribe in italic script): ‘Trees and shrubs. Stipules lateral or axillary. Leaves decussate, palmately compound, stipels present. Inflorescence racemose; flowers maturing synchronously to acropetally. Flowers bisexual; Weinmannia madagascariensis 0.01 0.007 96 79 52 77 66 76 99 95 87 100 90 53 56 100 100 Caldcluvia paniculata Karrabina benthamiana Pseudoweinmannia lachnocarpa Geissois hirsuta 100 97 98 100 87 Lamanonia ternata Gillbeea adenopetala 56 96 Davidsonia pruriens Hooglandia ignambiensis Spiraeanthemum samoense Acrophyllum australe Eucryphia cordifolia Ackama paniculosa Spiraeopsis celebica Caldcluvia paniculata Karrabina benthamiana Pseudoweinmannia lachnocarpa Geissois hirsuta Lamanonia ternata Gillbeea adenopetala Anodopetalum biglandulosum Platylophus trifoliatus Ceratopetalum gummiferum Schizomeria serrata 0.0370 Bauera rubioides 97 90 Eucryphia cordifolia Ackama paniculosa Spiraeopsis celebica 96 95 Callicoma serratifolia Codia discolor Pullea cf. glabra Pullea cf. glabra Acrophyllum australe 66 Cunonia atrorubens Pancheria engleriana Vesselowskya rubifolia 100 Callicoma serratifolia Codia discolor 55 100 99 Vesselowskya rubifolia 67 Weinmannia madagascariensis Bayes Cunonia atrorubens Pancheria engleriana 75 169 Hooglandia ignambiensis 100 99 100 100 100 Anodopetalum biglandulosum Platylophus trifoliatus Ceratopetalum gummiferum Schizomeria serrata Bauera rubioides Davidsonia pruriens Spiraeanthemum samoense Fig. 1. Phylogenetic trees of the genera of Cunoniaceae based on rbcL and trnLc–f sequences, using maximum-likelihood (ML) (left) and Bayesian inference (right). Numbers above branches indicate bootstrap values above 50 (left) or posterior probabilities (right). 170 Australian Systematic Botany petals absent; androecium polystemonous; pollen dicolporate; ovary bicarpellate. Fruit capsular or indehiscent; seeds winged or not.’ In the following review, data for gross-morphological characters cover all, or almost all, of the species in each genus and are taken from our observations, largely based on herbarium specimens, supplemented by data from published sources, including those summarised in Bradford and Barnes (2001). Data on micromorphological characters are from published sources or were supplied by Richard Barnes (pers. comm.) for leaf cuticular features but are not necessarily based on all species in each group. Table 1 provides a summary of the characters discussed, Fig. 2 illustrates a member of each of the four genera, and Fig. 3 compares critical features of Geissois, Karrabina and Lamanonia. Leaves Cunoniaceae typically have opposite or whorled decussate leaves (alternate only in Davidsonia) that can be simple or compound. Leaflets and stipels All members of the Geissoieae have opposite, digitately (or palmately) compound leaves with caducous stipels at the base of the outermost lateral leaflets (Bradford and Barnes 2001), although the stipels are often lost in herbarium specimens. Digitately compound leaves with three leaflets could be described as trifoliolate, although this term also includes leaves in which the lateral leaflets are markedly different in shape from the terminal one, whereas in the Geissoieae, only slight differences in shape between lateral and terminal leaflets occur. In all four species groups, the leaves typically have either three, five or a mixture of three and five leaflets although, as mentioned above, two species of Geissois can have more (namely, five or seven in G. hippocastanifolia, seven or nine in G. polyphylla). Trifoliolate leaves are relatively widespread in the Cunoniaceae and digitately compound ones also occur in Vesselowskya Pamp. (tribe Cunonieae; Rozefelds et al. 2001) and some members of the Schizomeriae, but these genera lack stipels, and conversely, stipels occur in a few genera outside the Geissoieae (Bradford and Barnes 2001, appendix 3). Margins and venation Cunoniaceae are generally characterised by toothed leaf(let) margins, although species with entire margins occur throughout the family. Adult trees of Karrabina, Lamanonia and Pseudoweinmannia all have dentate leaflet margins and semicraspedodromous venation, as do the seedling leaves in Geissois (Fogliani et al. 2009); all species of Geissois except G. hirsuta have leaflets with entire margins and brochidodromous venation in the adult plants. Micromorphological characters Schimanski and Rozefelds (2002) gave a detailed account for Karrabina and some comparable data exist for Geissois and Lamanonia (R. Barnes, pers. comm.), whereas there are few data for Pseudoweinmannia. Karrabina resembles Geissois rather than Lamanonia, in having well developed areoles H. C. F. Hopkins et al. and strongly percurrent tertiary venation, but it shares with Lamanonia (and not Geissois s.s.) the character of T-pieces on the stomates. All three genera have hydathodes on the veins and areoles and unicellular hair bases that are centrally thickened at the base. Hairs Cunoniaceae typically have simple hairs though small paired hairs and rarely stellate trichomes have been recorded (respectively in Codia J.R.Forst. & G.Forst. and Spiraeopsis Miq.). The indumentum in the Geissoieae consists primarily of hairs that are small to large, straight to crispate and adpressed to erect, depending on the genus and species. Indumentum is present on the foliage in a few species of Geissois and Lamanonia, and on the inflorescence axes, calyx lobes, ovaries and fruits in at least some members of all four genera. In Geissois, minute, weak, curled hairs are commonly present in addition to larger trichomes. On the stipules, these minute hairs are either distributed over the adaxial surface (G. hippocastanifolia, G. hirsuta, G. polyphylla) or they form a yellow rim around the adaxial margin (most species); on the inner surface of the calyx lobes, they are frequently visible towards the margins, whereas in the centre of the lobes, they are typically concealed beneath longer, pale thick-walled hairs. The minute curled hairs resemble the thin-walled, often paired hairs that form a dense, pale, felty layer on the lower leaf surface in several species of Codia (tribe Codieae) (Carpenter and Buchanan 1993; Barnes and Hill 1999), whereas in Geissois, they have not been studied at high magnification and we do not know whether they are paired. In Lamanonia, similar hairs occur on the inner surface of the calyx lobes and beneath longer hairs on the fruits, whereas the dense indumentum on the lower surface of the leaflets in L. ulei (Engl.) L.B.Sm. is composed of larger hairs. Glands Small, subspherical trichomes on leaf surfaces and other parts have been recorded in genera scattered through the family and are referred to as ‘glands’. Dark red or sometimes yellow glands are abundant in three species of Geissois (G. hippocastanifolia, G. hirsuta, G. polyphylla) among the dense indumentum of minute curled hairs on the inner surface of the stipules, and are present occasionally on other structures, including young stems and leaves, pedicels and floral buds. They usually appear to be sessile when viewed at 40
, although they are clearly stalked in one collection (Franc 1812 (K, P), G. racemosa). Rather similar dark red glands occur on the leaves, inflorescence axes and pedicels in Lamanonia cuneata (Cambess.) Kuntze, but they are usually elliptic rather than circular in outline, and short-stalked globular or glandular hairs were recorded by R. D. Hoogland (unpubl. data) on the stipules and twigs respectively of the species now in Karrabina. No glands have been recorded for Pseudoweinmannia. Domatia Various types of domatia occur throughout the family (Hopkins and Hoogland 2002). In Geissois, small pouches, apparently without hairs when viewed at 40
, are generally Karrabina gen. nov. (Geissoieae: Cunoniaceae) Australian Systematic Botany 171 Table 1. Comparison of the genera in tribe Geissoieae In the body of the table, characters that differ significantly among genera are in bold. Data for all genera are based on personal observations, plus Dickison and Rutishauser (1990), Bradford and Barnes (2001) and other references cited in the text. Sources for individual genera include the following: for Geissois, Hopkins (2006, 2008) and Hopkins et al. (in press); for Karrabina, Bailey (1883, 1900), Boland et al. (2006), R. D. Hoogland (unpubl. data) and Schimanski and Rozefelds (2002); for Lamanonia, Leite (1983) and Zickel and Leitão Filho (1993); for Pseudoweinmannia, R. D. Hoogland (unpubl. data), Francis and Chippendale (1970) and Rozefelds and Pellow (2011); IMs, inflorescence modules Parameter Geissois Karrabina Lamanonia Pseudoweinmannia Number of species Distribution 19 New Caledonia, Fiji, Vanuatu, Solomon Islands (Vanikoro) 2 Eastern Australia 5 Brazil, Paraguay, Argentina 2 Eastern Australia 2 per node, intrapetiolar; ovate, oblong, to almost circular 4 per node, free-lateral to partially connate along one margin (interpetiolar) or both; foliose, ±circular Lateral primordia not connate or late connate, especially between petioles 4 per node, free-lateral Digitately compound, leaflets 3 Toothed Yes Semi-craspedodromous 4 per node, free-lateral; falcate to semicordate 2 per node, interpetiolar, derived from 4 lateral primorida; lanceolate, fugaceous Lateral primordia not connate Lateral primordia early connate between petioles ? Digitately compound, leaflets 3–5 Toothed YesA Semi-craspedodromous ? Digitately compound, leaflets 3 Toothed Yes Semi-craspedodromous Strongly percurrentA Random reticulateA Sometimes present (esp. G. balansae) Absent Often present Weakly percurrent to reticulateB Absent AbsentA Well developedA LignifiedA or parenchymatousB AbsentA YesA PresentA Well developedA LignifiedA PresentA ImperfectA ParenchymatousA ? ? ParenchymatousB AbsentA YesA RareA YesA ? ? YesA YesA YesA ? IMs usually of 2 metamers, with median raceme, either monads (most spp.) or triads (G. hirsuta); lateral: axillary (few spp.) or ramiflorous (most) IMs with variable number of metamers and racemes, sometimes median but without median raceme; often distal to leaves At or near base Racemes simple, in distal leaf axils At or near base IMs lateral or median, variable in complexity, including dyads, triads and tetrads, with or without median racemes, sometimes auxiliary At or near base 1 1 1 1 4 (4–)5–6(–7) 6 (4–)5–7(–8) Often spreading Cup-shaped to spreading Reflexed in old flowers Cup-shaped to spreading 8–30 ~16–32 ~25–60+ (9–)15–23(–30?) Foliage characters Stipules in adult foliage Developmental type for stipules Axillary primordial ridge (or lateral primordia early connate across leaf axils?) Stipules in seedlings Leaf form in adult plants 4 per node, free-lateral Digitately compound,leaflets 3, 5, 7, 9 Entire (except G. hirsuta) YesA Brochidodromous (semicraspedodromous in G. hirsuta) Strongly percurrentA Leaflet margin in adult Stipels Secondary venation Tertiary venation Domatia Micromorphology T-pieces on stomates Areoles Vein sheaths Terminal idioblasts Hydathodes on veins and areoles Unicellular hairbases with centrally thickened base Structure of inflorescence Articulation of pedicel Floral characters Number of perianth whorls Number of perianth lobes Perianth lobes at anthesis Number of stamens Usually near mid-point (at base in G. hirsuta) (continued next page) 172 Australian Systematic Botany H. C. F. Hopkins et al. Table 1. (continued ) Parameter Number of series of stamens Stamens in bud Form of disc Number of carpels and styles Number of ovules per locule Flower colour Length of stamens in relation to perianth Pollinator Fruits Seeds Geissois Karrabina Lamanonia Pseudoweinmannia 1 1 >1 1 Curled Annular, lateral to ovary 2(3) Straight Annular, adnate to ovary 2(3) Straight Annular, adnate to ovary 2 Straight Annular, lateral to ovary 2 20–36(–42) ~12–20(–22) ~16–20 4–6(8) Red, with disc often yellow Cream, green or yellowish Prominently exserted Exserted Cream to yellowish with orange disc Exserted Cream or whitish, with pink disc Exserted Birds 2-valved capsule, glabrous, or sparsely and coarsely hairy, or velvety; calyx caducous Probably insects 2-valved capsule with short-woolly indumentum; calyx caducous Insects 2-valved capsule with sparse to dense silkyvelvety indumentum; calyx caducous Numerous, flat, winged, with sculptured surface Numerous, flat, winged, with sculptured surface Numerous, flat, winged, with sculptured surface Probably insects Tardily dehiscent with placental proliferations; pericarp with long woolly indumentum; calyx persistent ~4, spherical, not winged A Source for micromorphological characters R. Barnes (pers. comm.). Source for micromorphological characters Dickison (1975a). B well developed in G. balansae Brongn. & Gris ex Guillaumin and smaller domatia occur in a few other taxa (G. denhamii (see Wheatley 1992), G. pentaphylla C.T.White, G. superba Gillespie and G. ternata). In Lamanonia, domatia are commonly present in most (all?) species, with tufts of hairs in expanding leaves, but without hairs in older foliage. Domatia have not been recorded from either Karrabina or Pseudoweinmannia. Stipules All members of Cunoniaceae have stipules, and these can be categorised as free-lateral (four per node when leaves opposite), interpetiolar (two per node if leaves opposite; most genera) or intrapetiolar (two per node and leaves opposite). The details of structure and development are different in each species group in the Geissoieae (Rutishauser and Dickison 1989; Dickison and Rutishauser 1990; Bradford and Barnes 2001; Schimanski and Rozefelds 2002) and homologies are indicated by Schimanski and Rozefelds (2002, fig. 33). The foliage of adult Lamanonia trees has four free lateral stipules per node, with no tendency for them to become connate during development (Figs 2A, 3L) (see also illustrations in Engler 1928; Leite 1983; Zickel and Leitão Filho 1993). Although this genus was not studied by Dickison and Rutishauser, it is reasonable to assume that the free lateral stipules develop from four lateral primordia. Fully grown stipules associated with mature leaves are falcate to semicordate, somewhat foliaceous, acute at the apex and fugacious to persistent. When they are large, their venation is campylodromous (e.g. Glaziou 17623 (K), L. grandistipularis (Taub.) Taub). Similar free lateral stipules occur in a few other genera of the Cunoniaceae (e.g. Gillbeea, Caldcluvia D.Don). The stipules of Karrabina are also lateral and free at inception (Dickison and Rutishauser 1990). In K. benthamiana, they remain free during growth and overlap across the leaf axil (Fig. 3G) and are comparable to those of Lamanonia grandistipularis. In K. biagiana, they are partially (i.e. late) connate down one margin, to become interpetiolar but deeply lobed, or they can be partially connate on both margins and then somewhat cup-shaped (Schimanski and Rozefelds 2002). Individual stipules are foliaceous and rounded when separate or kidney-shaped with deep indentations if partially fused down the interpetiolar margins; the venation is campylodromous but the apex is never acute (Fig. 3H). They are fugaceous to persistent in adult foliage. Geissois has two intrapetiolar (or axillary) stipules per node in adult plants (Fig. 3C, D) (Dickison and Rutishauser 1990; Bradford and Barnes 2001; Hopkins 2008), although seedlings have four free lateral stipules per node (Fogliani et al. 2009). Rarely, opposite stipules may be shortly connate at their base (e.g. in G. superba, illustrated in Smith 1952, and G. magnifica, illustrated in Hopkins 2008); however, this is a secondary modification. Each stipule develops from an intrapetiolar ridge (Dickison and Rutishauser 1990, fig. 2), and Schimanski and Rozefelds (2002) suggested that each ridge develops from two lateral primordia by early connation across the leaf axil. Fully grown stipules in Geissois are ovate, oblong, cordate or sometimes almost circular, with the base being cuneate, rounded or auriculate, and the apex obtuse to rounded or rarely bifid, and they are caducous to persistent. When the stipules are large, the venation is again campylodromous (e.g. G. lanceolata, G. magnifica). In Pseudoweinmannia, the fully grown stipules appear to be interpetiolar, but because they develop from four free lateral primordia by early connation and basal elongation of the common sheath area (Dickison and Rutishauser 1990), they were scored as lateral at inception by Bradford and Barnes (2001). Young stipules are bifid at the apex, although this Karrabina gen. nov. (Geissoieae: Cunoniaceae) Australian Systematic Botany A B C D Fig. 2. Representative species of the four genera in the tribe Geissoieae. A. Lamanonia speciosa. B. Geissois racemosa. C. Pseudoweinmannia lachnocarpa. D. Karrabina benthamiana. A is from de Saint-Hilaire et al. (1829–1833), originally published as Belangera speciosa; B is from de Labillardière (1824–1825); C is from Maiden (1917), originally published as Geissois lachnocarpa; D is from Maiden (1916), originally published as Geissois benthamii; note that the apparently terminal leaf is an error. 173 174 Australian Systematic Botany H. C. F. Hopkins et al. J A E K B F L G C D H Fig. 3. Comparison of the flowers, inflorescences and stipules in Geissois, Karrabina and Lamanonia. A. Flower, G. balansae. B. Shoot with three inflorescence modules, G. lanceolata. C. Stipules and apical bud, G. velutina. D. Stipules, G. lanceolata. E. Flower, K. benthamiana. F. Shoot with inflorescence, K. biagiana. G. Stipules, K. benthamiana; note one stipule is mis-shapen. H. Stipules, K. biagiana. J. Flower, L. ternata. K. Shoot with axillary racemes, L. ternata. L. Stipules, L. ternata. Drawn from McPherson 2348 (A), Jaffré 1015 (B), MacKee 34219 (C), MacKee 20378 (D), specimen with label for F. Mueller at P (E), specimen with label for F. Mueller at P (F, H), photograph in Boland et al. (2006) (G), Lourteig 2268 (J), Prance & Silva 59546 (K) and Glaziou 103 (L). Drawn by D. Storez. Karrabina gen. nov. (Geissoieae: Cunoniaceae) feature becomes inconspicuous during development and the mature stipules in the adult foliage are lanceolate, acute towards the apex and fugaceous. Stipules with bifid apices occur elsewhere in Cunoniaceae, especially in Codia and Callicoma Andrews (also Codieae) (Rutishauser and Dickison 1989), which suggests early interpetiolar connation between the primordia in these genera as well, although these authors considered that they had interpetiolar stipules, whereas Pseudoweinmannia was said to have non-interpetiolar ones. Inflorescences The Cunoniaceae show a range of inflorescence types, but all members of the Geissoieae have racemose inflorescences, with the flowers maturing synchronously or somewhat acropetally (Bradford and Barnes 2001). Elsewhere in the family, racemose inflorescences are found in three genera in the Cunonieae (Weinmannia, Cunonia L., Vesselowskya) and the details of their structure are taxonomically significant (e.g. Hoogland et al. 1997 for Cunonia; Bradford 1998 and Hopkins and Bradford 1998 for Weinmannia). Bradford (1998) described complex inflorescences in terms of repeating units, or inflorescence modules (IMs), each module consisting of one or more metamers, where a metamer comprises an internode, the node at its distal end, and any associated structures at that node such as buds, bracts and racemes. Although it is obvious to a casual observer that the inflorescences in the four species groups of the Geissoieae are different, describing these differences has proved difficult, even when using the modular approach of Bradford (1998). Of the four species groups, Lamanonia has the simplest inflorescences, with individual racemes being inserted singly in the axils of leaves towards the distal end of a shoot (Figs 2A, 3K). The lack of bracts or bract scars on the peduncle of each raceme indicates that each axillary structure is indeed a simple raceme, not an IM. The inflorescences of Geissois (Figs 2B, 3B) are more complex, although they have a more or less consistent structure (Hopkins 2008). In all species except G. hirsuta, they appear, superficially, to be composed of simple racemes, but each ‘raceme’ is in fact a monad, i.e. an IM, usually composed of two metamers, in which the peduncle bears two pairs of fugaceous bracts and a single median (terminal) raceme. G. hirsuta differs in that each IM is a triad in which, in addition to the median raceme, lateral racemes develop in the axils of the proximal pair of bract scars on the peduncle. In most species of Geissois, the IMs are ramiflorous, although in a few (G. hirsuta, G. superba, sometimes G. lanceolata and G. magnifica), they may be borne in leaf axils towards the shoot apex. Where IMs are in the axil of an extant leaf, usually only one monad or triad develops per axil, but where they are ramiflorous, they occur singly or in groups of up to 20 per half node. In the remaining two genera, the structure of the inflorescence is more plastic. In Pseudoweinmannia (Fig. 2C), racemes can be arranged in opposite pairs to form dyads that arise either in leaf axils or at leafless nodes immediately proximal to the distal leafbearing node of the shoot, or sometimes racemes form triads at leafless nodes. Inflorescences may also have auxiliary IMs (i.e. ones that develop from buds on either side of an axillary Australian Systematic Botany 175 bud or shoot), which generally consist of one metamer with a short basal segment bearing two, three or four racemes at its distal end. In Karrabina, the inflorescences frequently consist of racemes organised in opposite pairs at leafless nodes towards the distal end of a shoot (Figs 2D, 3F). In contrast to most species of Geissois, IMs in Karrabina are never ramiflorous nor in clusters at nodes. In Karrabina benthamiana, inflorescences or modules commonly consists of two metamers, with two successive pairs of racemes inserted on either a lateral leafless axis or a median leaf-bearing shoot, and IMs usually have a dormant apical bud between the distal pair of racemes. In Williams s.n. (NE 35786b), illustrated by Schimanski and Rozefelds (2002, fig. 24), and Bradford & Snow 848 (K), racemes are borne on lateral axes inserted in leaf axils at or near the distal end of a median shoot. In White 7533 (K), the apical bud between a single pair of opposite axillary racemes on a median shoot has continued vegetative growth beyond the racemes in the fruiting stage. In Karrabina biagiana, the inflorescences are usually more elaborate than in K. benthamiana, although Schimanski and Rozefelds (2002, fig. 23, based on Elick 44, QRS 87014) illustrated a relatively simple example. A more typical example is seen in Bradford & Snow 849 (K) (Fig. 4), in which the inflorescence has a median IM plus several lateral ones, and each IM is composed of one to three metamers. Most of the IMs have racemes inserted in opposite pairs at leafless nodes, towards the distal end of their axes, but one or two nodes of the axis, beyond the racemes, continue to develop and produce young leaves. Lateral IMs arise from nodes at the base of the median IM, either in the axils of fully developed leaves, or in the axils of reduced leaves, or from leafless nodes. Although most lateral IMs repeat the structure of the median IM, with the distal node(s) producing young leaves, dyads also occur in which a basal segment bears a pair of opposite lateral racemes and the apical bud within the IM is either aborted or dormant. Inflorescences in other specimens of K. biagiana are similar but with minor variations. Some homologies between inflorescence types can be inferred as follows. The inflorescence of Lamanonia, with simple racemes in successive pairs of opposite leaf axils, could be transformed into the type of inflorescence in Karrabina (with sets of dyads), by the suppression of the leaves at the raceme-bearing nodes accompanied by the shortening of the basal segments on the lateral raceme-bearing axes. However, the relationship between inflorescences in Geissois and Lamanonia is less straight forward. The axillary monads of Geissois are not homologous to the simple racemes of Lamanonia because each monad is a modified shoot system composed of two metamers. The homologies for Pseudoweinmannia are complicated by the plasticity of the inflorescences and the development of IMs from auxiliary buds. Those parts of the inflorescence with opposite pairs of racemes and nodes where leaves have been suppressed, are homologous to inflorescences in Karrabina benthamiana. Inflorescences in Pseudoweinmannia, that have either several axillary racemes per node or consist of axillary triads, resemble the inflorescences of Geissois, although in Pseudoweinmannia, each IM comprises only one metamer, not two. 176 Australian Systematic Botany H. C. F. Hopkins et al. H D E F C G A B Fig. 4. Karrabina biagiana. A. Shoot with leaves and inflorescence. B. Stipule. C. Stamen. D. Flowers post anthesis, with anthers fallen. E. Gynoecium and disc attached to pedicel. F. Transverse section through ovary. G. Longitudinal section through ovary. H. Dehisced fruit. Drawn from Bradford & Snow 849 (A, B), Gray 1235 (C–G) and Dockrill 34 (H). Drawn by Patricia K. R. Davies. Karrabina gen. nov. (Geissoieae: Cunoniaceae) Australian Systematic Botany Flowers All members of Cunoniaceae have a polysymmetric perianth and androecium, and usually the ovary is superior, with the number of styles (strictly stylodia) being equal to the number of carpels. The floral ground plan for the family, using the notation of Prenner et al. (2010), appears to be: B K4–6* C4–6* A4–6$4–6* G(3–5)* Vx¥. Flowers in the Geissoieae are bisexual with a single perianth whorl, usually referred to as a calyx, which is valvate in bud. The stamens are numerous (A > 2K), with bithecal anthers incised at the apex and lacking extensions of the connective. The anthers are dorsifixed, introrse and versatile in all four species groups. The disc is annular and the superior gynoecium has 2 (or rarely 3) carpels fused at the level of the ovary and surmounted by 2(3) styles with terminal stigmas. Elsewhere in the family, apetalous flowers are relatively common, occurring in approximately eight other genera, and polystemony also occurs in Eucryphia Cav. and Bauera Banks ex Andrews (Dickison 1975b, 1978; Bradford and Barnes 2001). Pedicels and bracts In all four genera, each pedicel is subtended by a small fugaceous or caducous bract that is typically linear to narrowly triangular or ovate and somewhat carinate. These bracts are usually 1–2 mm long in Karrabina, Pseudoweinmannia and most species of Geissois, up to ~5 mm in Lamanonia, and up to 7 mm in G. hirsuta. Some variation occurs in the position of the articulation of the pedicel. In Pseudoweinmannia, Karrabina and Lamanonia, the pedicels articulate at or shortly above the base (<1 mm); pedicels that articulate just above the base often have a marked change in the density of the indumentum at this point. In Geissois, the articulation is near the mid-point in most species, just above 177 the base in G. polyphylla and G. hippocastanifolia, and at the base in G. hirsuta. Perianth Geissois has a four-merous perianth and only very rarely have five calyx lobes been recorded. Different sources give slightly different numbers of perianth parts for the other genera; however, the commonest numbers are five or six for Karrabina, six for Lamanonia and five to seven for Pseudoweinmannia. Androecium The members of Geissoieae all show complex polyandry, and in all four groups, the number of stamens is irregular rather than being a multiple of the number of calyx lobes (Table 1). In Geissois, Karrabina and Pseudoweinmannia, the number of stamens is small relative to Lamanonia (never more than 32) and, at anthesis, the filaments appear to be inserted in a single series between the calyx lobes and the disc, with the filaments all being similar in length within any flower. In G. pruinosa Brongn. & Gris, Matthews and Endress (2002) noted that the median (episepalous) stamen is initiated first and the subsequent initiation occurs laterally and progressively extends to the sepals margins. Large buds of K. biagiana and K. benthamiana clearly show their stamens in groups (Schimanski and Rozefelds 2002, figs 25, 27) and in Pseudoweinmannia lachnocarpa (F.Muell.) Engl. (Rozefelds and Pellow 2011, fig. 4E), the median ones in each group, opposite the midpoint of a calyx lobe, have the longest filaments, which suggests that they were initiated first. Lamanonia generally has more numerous stamens (~25–60; Table 2) that appear in mature flowers to be inserted in one series and partially and irregularly in a second and often a third series; the filament bases are crowded, with a tendency for Table 2. Number of stamens per flower in Lamanonia species, on the basis of literature sources and herbarium specimens Herbarium specimens were identified as follows: Z&L_1, listed in Zickel and Leitão Filho (1993); Z&L_2, listed under both L. brasiliensis and L. ternata in Zickel and Leitão Filho (1993); Zickel, det. slip on specimen; Leite, listed in Leite (1983); HCFH, determined by Hopkins Specimen or literature reference Species Specimen determined by Zickel and Leitão Filho (1993) Prance & Silva 59074 Ratter et al. 3524 Ratter et al. 3768 brasiliensis brasiliensis brasiliensis brasiliensis? Z&L_1 Type number Z&L_2 Zickel and Leitão Filho (1993) Glaziou 8247 chabertii chabertii Zickel and Leitão Filho (1993) Dusen 10889 Hatschbach 11986 Widgren s.n. Plant material Herbarium 40 49 39 46 Flower Flower Flower P K K Type number Numerous 44 Flower K cuneata cuneata cuneata cuneata Z&L_1 Zickel HCFH Numerous 56, 58, 60 52 55, 59 Flower Flower Flower K, P K P Zickel and Leitão Filho (1993) Glaziou 17623 grandistipularis grandistipularis Type number Bud, flower K, P Leite (1983) Zickel and Leitão Filho (1993) Blanchet 3374 Glaziou 2495 Hatschbach 23180 Hatschbach 28368 Prance & Silva 59546 ternata ternata ternata ternata ternata ternata ternata Buds, flowers Flower Flower Flower Flower P P K K P Z&L_1 Z&L_1 Leite Zickel Z&L_1 Number of stamens per flower 30 41, 44 35–40 25–30 26, 28, 28, 30, 31 30 47 46 59 178 Australian Systematic Botany adjacent ones to be shortly connate at the base. The stamens in large buds are in groups or fascicles, with the median ones in each group being longer than those on either side (Leite 1983, fig. 3A, B), but again this is less evident at anthesis. Although the direction of stamen development has not been studied in detail in any of these groups, the available evidence strongly suggests that all genera exhibit centrifugal androecial development. The floral formulae in the generic descriptions show the androecia in all four genera as being polysymmetrical with an indefinite number of stamens (A¥*) at anthesis. The formula in bud is also given for Karrabina, Lamanonia and Pseudoweinmannia. The filaments of Karrabina (Schimanski and Rozefelds 2002), Lamanonia (Leite 1983, fig. 3A, B) and Pseudoweinmannia (Rozefelds and Pellow 2011) are more or less straight in bud, whereas in Geissois they are irregularly coiled (Endress and Stumpf 1991; A. C. Rozefelds, pers. obs. 2002). This may be because the filaments are comparatively longer at anthesis in Geissois than in the other genera. Disc It is not clear to what extent differences in the form of the disc are taxonomically significant within the tribe. In Geissois, the disc is lateral to the base of the ovary and it is quite large and fleshy, often being yellow in otherwise reddish flowers, and it secretes large volumes of nectar, which correlates with the adaptation to bird-pollination in this genus. In contrast, the disc in Karrabina, Pseudoweinmannia and Lamanonia is adnate to the base of the ovary so it lies vertically beneath the upper part of the ovary (Zickel and Leitão Filho 1993; Schimanski and Rozefelds 2002, figs 28, 30; Rozefelds and Pellow 2011). In Karrabina, the disc appears to be relatively inconspicuous, compared to Geissois, and we have no information on nectar secretion. In Lamanonia, it is inconspicuous in drawings and rehydrated flowers, whereas in fresh material, it forms an orange ring that contrasts with the rest of the flower, which is cream to yellow, and it is nectariferous (Leite 1983; Zickel and Leitão Filho 1993). In Pseudoweinmannia lachnocarpa it is red to red-brown in colour in an otherwise whitish flower, and it is distinctly lobed (Rozefelds and Pellow H. C. F. Hopkins et al. 2011; see photo by B. Jago in Australian Tropical Rainforest Plants 2010). Ovary The outer surface of the ovaries in all four genera is densely hairy, except in a few species of Geissois. All genera exhibit axile placentation and each of the two locules per ovary in Karrabina, Geissois and Lamanonia has numerous (~10–36) ovules arranged in two rows, whereas Pseudoweinmannia has 2–6 ovules per locule. Schimanski and Rozefelds (2002) illustrated the arrangement of ovules in Karrabina and stated that they were arranged in two rows per carpel, but they erroneously gave the number of ovules as 5–10 per carpel (Schimanski and Rozefelds 2002, pp. 229, 232), whereas this is the number per row; elsewhere in their paper, the number of ovules or seeds per carpel is described by Schimanski and Rozefelds as 10–25 (and see Table 1). Floral biology and pollination Karrabina, Lamanonia and Pseudoweinmannia are all likely to be insect-pollinated, although we have found little published information. The flowers of Karrabina species are creamish according to Schimanski and Rozefelds (2002), the calyx lobes and stamens being the principal showy parts. A photo of K. biagiana (by B. Jago in Australian Tropical Rainforest Plants 2010) shows this species to have creamish flowers and the field notes of Bradford & Snow 848 record the flowers of K. benthamiana as pale green with pink styles. However, Bailey (1883, 1900) described the calyx and filaments in both species as yellow. We have no information on floral scent or potential pollinators but the small, actinomorphic flowers suggests visitation by small insects, including perhaps flies and beetles. In Lamanonia, the flowers are white, cream or yellowish and very fragrant, and beetles and thrips have been reported as floral visitors (Table 3). Pseudoweinmannia also has pale, sweetly fragrant flowers (calyx pale green, filaments and ovary cream, anthers yellow and disc red, according to the field notes of Bradford & Snow 858 (K), Table 3. Field data on floral characters and floral visitors for Lamanonia species All herbarium specimens mentioned are at K Lamanonia species Floral characters L. brasiliensis Tepals and androecium cream; ovary orange at base, pale green above Calyx and filaments white Flowers white Flowers yellow Flowers cream Sepals creamish-green, stamens white, anthers cream Petals (sic) yellow, stamens dull yellow, anthers reddish, styles white Flowers white Nectariferous disc orange L. chabertii L. cuneata L. grandistipularis L. ternata Flowers yellowish cream with a band of pale orange hairs round the base of the ovary Floral visitors Source of data Field notes of Ratter et al. 3524 Insects present Field notes of Prance & Silva 59074 Field notes of Pinto et al. FUEL 17456 Photo by C. Zickel Field notes of Hatschbach 11986 Field notes of Pirani et al. CFCR 12724 Field notes of Ganev 261 Pelidnota purpurea Burn. (Scarabaeidae) Flowers full of thrips (Thysanoptera) Field notes of Gentry & Zardini 49512 Leite (1983) Field notes of Ratter et al. 6455 Karrabina gen. nov. (Geissoieae: Cunoniaceae) and although we have no data on floral visitors, again the small, actinomorphic flowers are likely to be visited by small insects such as flies and beetles. In contrast, the predominantly red flowers of Geissois appear to be largely ornithophilous. The calyx lobes and prominently exserted filaments are red, the disc is often yellow and the ovary is red, yellow or green with exserted red styles; nectar is usually abundant and the flowers have little or no odour. Floral visitors include various species of nectar-feeding birds in New Caledonia (Meliphagidae and Psittacidae, Barré et al. 2010), Fiji and Vanuatu, as well as geckos, several types of insects and, occasionally, fruit-bats (observations and other references summarised in Hopkins et al., in press). The widespread ramiflory of this genus is no doubt related to ornithophily, although the significance of differences among species (e.g. racemes lax and pendent v. stiff and projecting) is not known. Fruits, seeds and dispersal Most members of the Geissoieae have dry, dehiscent capsules and this type of fruit is common throughout the family, whereas indehiscent fruits, whether dry or fleshy, are considered derived (Dickison 1989). The two-valved capsules of Karrabina, Lamanonia and Geissois dehisce from the apex. The style-bases are often persistent, although the perianth and androecium are usually not. After dehiscence, each valve has a tendency to split for a short distance at its apex; this is most pronounced in Lamanonia, giving the capsule a ‘four-pointed’ appearance, and this character is less strongly expressed in Geissois and Karrabina. The coriaceous central part of each valve has parchment-like extensions of the endocarp down either side and the valves have a tendency to remain partially joined in their lower halves by the placentas. Sometimes, especially in Karrabina, this occurs in a manner similar to that seen in Caldcluvia (Godley 1983, fig. 2) and sometimes in Geissois, Lamanonia and Karrabina, the endocarp eventually splits from the remnants of the placentas to form a free central column. The seeds in all three genera are small, flat, winged, without hairs and wind-dispersed. Within a single capsule, some variation usually occurs in the shape of the seeds. Seed surfaces are minutely sculptured (Dickison 1984; Smith 1985). Externally, the ovoid–ellipsoid capsules of Lamanonia and Karrabina are strikingly similar. In Lamanonia, the valves are sparsely to densely silky-velutinous with fine, slightly wavy, longish pale to brown hairs covering an under-layer of minute trichomes. The seeds are elliptic, with the embryo near the base at one side and the wing is asymmetrically developed at the apex and on one side (Dickison 1984, fig. 39; Zickel and Leitão Filho 1993). In Karrabina, the indumentum on the capsules is similar but more matted and the seeds are also similar (based on Dockrill 34 (K) for K. biagiana, and Longman s.n. (K) for K. benthamiana). Within Geissois, G. hirsuta has ellipsoid to broadly ovoid capsules with sparse, coarse hairs, whereas the remaining species have narrowly cylindrical, sometimes curved capsules that can be velutinous, coarsely hairy or glabrous. Except in G. hirsuta, the seeds are quite similar to those of Lamanonia but the wing is typically well developed at the apex, narrow at the Australian Systematic Botany 179 sides and sometimes slightly developed at the base (Dickison 1984, figs 37, 38), so that the embryo is more centrally positioned. The seeds of G. hirsuta are unique in the genus, being almost circular, with the embryo in the centre of an annular wing (ORSTOM 1986). Pseudoweinmannia has tardily dehiscent fruits (Floyd 1989; Harden 1990), resembling small hairy balls in which the styles, calyx and androecium (stamen filaments) are persistent (Fig. 2C) and is probably also anemochorous. Each fruit is broadly ovoid with abundant long woolly hairs and the locules are largely filled by placental proliferations between the seeds (Dickison 1984, fig. 21). The seeds are few (up to four per fruit) and each has a small, aril-like outgrowth on the testa which Dickison (1984) suggested might be an elaiosome, but we know of no current ecological or field studies to support his suggestion. Rozefelds and Pellow (2011) concluded that the thin-walled parenchmatous cells were derived from the placental proliferations around the point of attachment of the seed, and are therefore not an elaisosome. Despite a superficial similarity to the fruits of Codia, which develop from an inferior ovary, the fruits of Pseudoweinmannia are unique in the family. They appear markedly different from those of the other genera in the Geissoieae, but their underlying structure is similar because they are also derived from a superior bicarpellary ovary with two styles, as in the other three genera (Rozefelds and Pellow 2011). Discussion We confirmed the observation of Bradford and Barnes (2001) that the tribe Geissoieae contains four distinct groups. Both the molecular and morphological data showed that the two Australian species traditionally placed in Geissois are distinct from Geissois s.s. Publication of a new genus, Karrabina, to accommodate these two species, increases to eight the number of endemic genera of Cunoniaceae in Australia, the others being Acrophyllum Benth., Anodopetalum A.Cunn. ex Endl., Bauera, Callicoma, Davidsonia F.Muell., Pseudoweinmannia and Vesselowskya, now that Aphanopetalum Endl. has been excluded (Dickison et al. 1994), and the total number of genera in the family represented in Australia remains at 15. Dickison and Rutishauser (1990) suggested that the Australian species of Geissois, now Karrabina, had similarities to Lamanonia and these are now shown to include features of the leaves, the number of perianth parts and their colour, and the shape of the capsules and their indumentum, with the stipules and inflorescences showing clear homologies. However, these characters appear to be plesiomorphic for the tribe and the closest relative of Karrabina is Pseudoweinmannia, although these two genera have few or no synapomorphies (see Table 1), perhaps because the latter has many derived features. Geissois also has many characters that are unique in the tribe and that appear to be derived, including intrapetiolar stipules, red flowers with four perianth lobes, prominently exserted stamens and a large disc (all species), and entire leaflet margins, ramiflory and narrowly cylindrical capsules (most species). All of the genera of Cunoniaceae recognised by Bradford et al. (2004) were described before 1929, with the exceptions of Acsmithia Hoogl., which was published by Hoogland (1979) 180 Australian Systematic Botany and has now been recombined with Spiraeanthemum A.Gray (Pillon et al. 2009), and the recently rediscovered, rare Hooglandia McPherson & Lowry from New Caledonia (McPherson and Lowry 2004). This raises the question of why it has taken so long for the Australian species now placed in Karrabina to be recognised as distinct. Several factors have contributed. Although casual inspection suggests Karrabina is rather different from Geissois, its distinguishing characters have not proved easy to describe. The inflorescence structure is complex and variable in Karrabina and descriptions of the stipules for Karrabina and Geissois appear to overlap (in both, they are sometimes fused both between and across the leaf axils), although, in fact, each has stipules that are unique in the family. Few taxonomists have critically compared these two genera, in part because Cunoniaceae is a small, generally little known family and Geissois and Karrabina are allopatric, the latter having only two species, both with restricted distributions. The classical morphological and anatomical studies of the Cunoniaceae, which owe much to the work of Dickison (e.g. Dickison 1975a, 1975b, 1978, 1984, 1989 and others), sought primarily to document and compare the characters of genera rather than to question generic limits, and, until Bradford and Barnes (2001), no-one since Engler (1928) had looked in detail at generic delimitation over the entire family in a single publication. The parallel morphological and molecular studies presented here have helped redefine generic limits in the Geissoieae, as has been done previously for the tribe Spiraeanthemeae (Pillon et al. 2009). Further work is still needed elsewhere in the Cunoniaceae, such as in the Caldcluvieae, where some genera may not be monophyletic (Y. Pillon, pers. obs.). H. C. F. Hopkins et al. Taxonomy In the generic descriptions, features unique to each genus are in italic. Geographical distributions are shown in Fig. 5. 1. Karrabina gen. nov., Rozefelds & H.C.Hopkins Distinguished from Geissois by having: stipules four per node, free-lateral and overlapping across leaf axil or partially connate down one or both margins (rather than stipules two per node and intrapetiolar in Geissois); inflorescences with several simple racemes organised into complex lateral and sometimes median, leafless modules, each module consisting of an indefinite number of metamers with no median raceme (rather than inflorescences with one or three median racemes organised into simple, lateral, leafless modules, each usually comprising two metamers in Geissois); and creamish flowers commonly with five or six calyx lobes (rather than reddish flowers with four calyx lobes in Geissois). Type: Karrabina benthamiana (F.Muell.) Rozefelds & H.C. Hopkins. Key references: Schimanski and Rozefelds (2002), whose generic description for Geissois refers to Karrabina; Boland et al. (2006) under the name Geissois. Trees. Leaves opposite and decussate, petiolate, digitately 3foliolate; leaflets dentate, with semi-craspedodromous venation, domatia absent; stipels at distal end of petiole caducous. Stipules 4 per node, large, foliaceous, free-lateral and overlapping across leaf axil, or partially connate down one margin (interpetiolar) or both margins and then often notched, often persistent. Inflorescence racemose, usually distal to fully developed leaves, consisting of several simple racemes organised into complex lateral and sometimes median leafless IMs, IMs 100°E 110°E 120°E 130°E 140°E 150°E 160°E 170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W 100°W 90°W 80°W 70°W 60°W 50°W 40°W 30°N 30°N 20°N 20°N 10°N 10°N 0° 0° 10°S 10°S 20°S 20°S 30°S 30°S 40°S 40°S 50°S 50°S 60°S 60°S 100°E 110°E 120°E 130°E 140°E 150°E 160°E 170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W 100°W 90°W 80°W 70°W 60°W 50°W 40°W Fig. 5. Distributions of the genera in the tribe Geissoieae. A. Karrabina and Pseudoweinmannia; both genera occur in both of the regions indicated. B. Geissois. C. Lamanonia. Karrabina gen. nov. (Geissoieae: Cunoniaceae) consisting of an indefinite number of metamers, with no median raceme. Flowers bisexual, apetalous, cream to yellowish; pedicels articulating at or near base; calyx lobes 5 or 6, triangular, valvate in bud, scarcely spreading at anthesis; stamens numerous (~16–32), filaments in a single series, exserted; anthers bithecal, not apiculate; disc adnate to base of ovary, ribbed, with indentations corresponding to bases of filaments; gynoecium superior, ovoid–conical, densely woolly, bicarpellate and bilocular, with 2 styles; ovules 16–28(–32) per locule, in 2 rows. Floral formula: B K5–6* C0 A¥* at anthesis (A5–6x* in bud) G(2)* Vx¥. Fruits capsular, ovoid, dehiscing from the apex into 2 valves, with remnants of styles persisting, valves remaining partly connected by the placentae; exocarp velvety to woolly; seeds numerous, flat, distally winged. Etymology and pronunciation ‘Garrabin’ was referred to as the Aboriginal name for Geissois benthamiana by Dixon et al. (1990), but further research and comments from linguists suggest that the spelling ‘Karabin(y)’ is more appropriate. The scientific name Karrabina is therefore derived from the name given by Gresty (1946–1947), ‘Karabin(y)’, which is used for these trees in the Yugambeh Language Region that extends from the Logan River to the Tweed River in south-eastern Queensland (P. O’Connor, pers. comm. 2012). This language region overlaps a significant proportion of the range of K. benthamiana. ‘Karabin(y)’ is the earliest Aboriginal name found for the tree (M. Sharpe, pers. comm. 2011; S. Haworth, pers. comm. 2011), and Karrabina is one of the very few generic names for Australian plants that is based on the Aboriginal name of the plant. The initial consonant of Karrabina sounds like a ‘K’, which is consistent with Yugambeh language word usage (P. O’Connor, pers. comm. 2012); the ‘r’ should be pronounced as a ‘rr’, hence the spelling of the name, and the ‘n’ in the name should be pronounced as ‘ny’ as in canyon. Vernacular name The common name in English for Karrabina species (‘Red Car(r)abeen’) is obviously derived from the Aboriginal language word and this name has also been used for Pseudoweinmannia lachnocarpa (Australian Tropical Rainforest Plants 2010). ‘Carabeen’, in combination with other descriptive terms (e.g. Blush Carabeen, Grey Carabeen, White Carabeen) also applies to trees in some other genera, such as Aceratium, Elaeocarpus and Sloanea (Elaeocarpaceae) (Maiden 1917; Australian Tropical Rainforest Plants 2010) and has various spellings. New combinations Karrabina benthamiana (F.Muell.) Rozefelds & H.C.Hopkins Geissois benthamiana F.Muell., Fragmenta phytogeographiae Australiae 5, 16 (1865); Geissois benthamii F.Muell., loc. cit. 5, 180 (1866) ‘benthami’; Weinmannia benthamii F.Muell. loc. cit. 6, 188 (1868) ‘benthami’; Windmannia benthamii (F.Muell.) Kuntze, Revisio Generum Plantarum 1, 228 (1891); ‘benthami’; Weinmannia benthamii var. typica Domin, Beitrage zur Flora und Pflanzengeographie Australiens 708 & Biblotheca Botanica 89, Australian Systematic Botany 181 154 (1926) [nom. inval.]. Type: Cloud’s Creek, Clarence River, Beckler s.n., s.dat. (holotype: MEL104460). Weinmannia benthamii var. microcarpa Domin, Beitrage zur Flora und Pflanzengeographie Australiens, 708 & Bibliotheca Botanica 89, 154 (1926). Type: not designated (Queensland: Regenwälder der Tamborine Mts). Distribution: south-eastern Queensland and north-eastern New South Wales. Karrabina biagiana (F.Muell.) Rozefelds & H.C.Hopkins Weinmannia biagiana F.Muell., Fragmenta Phytographiae Australiae 5, 16 (1865); Geissois biagiana (F.Muell.) F.Muell., loc. cit. 5, 180 (1866); Windmannia biagiana (F.Muell.) Kuntze, Revisio Generum Plantarum 1, 228 (1891). Type: Seaview Range, Rockingham’s Bay, [Dallachy s.n.?], 3 December 1864 (holotype: MEL104551; isotypes: K000739679, P). Distribution: north-eastern Queensland. Notes The two species of Karrabina, both from rainforest in eastern Australia, differ in relatively minor characters of the leaves, stipules and inflorescences. Schimanski and Rozefelds (2002) provided a detailed generic description, a key and species descriptions, plus illustrations, distribution maps and a discussion of nomenclature. 2. Geissois Labill., Sertum austro-caledonicum 2, 50 (1825) Type: Geissois racemosa Labill., [New Caledonia, Balade], Labillardière s.n. [1793] [Lectotype: FI, Herb. Webb. 060970 (buds, fl. & fr.); isolectotypes: BM000600408; FI, Herb. Webb. 060971 p.p. (excl. fragm. with persistent stipules); K000739688; P (st.) (Hopkins 2006)]. Key references: Smith (1952, 1985) (Fiji); Hopkins (2008) (morphology); Pillon (2011); Hopkins et al. (in press). Trees. Leaves opposite and decussate, petiolate, digitately compound; leaflets 3, 5, 7 or 9, entire with brochidodromous venation (most species) or dentate with semi-craspedodromous venation (G. hirsuta only), domatia sometimes present; stipels at distal end of petiole caducous. Stipules 2 per node, intrapetiolar, persistent or caducous. Inflorescences racemose, IMs usually of 2 metamers with a median raceme, racemes single (monad) in most species, or in 3s (triad) (G. hirsuta only), IMs lateral and mostly ramiflorous (axillary in G. hirsuta and G. superba, and mixture of ramiflorous and axillary in G. lanceolata and G. magnifica). Flowers bisexual, apetalous, red; pedicels articulating near midpoint (most species) or base; calyx lobes 4, triangular, valvate in bud, often spreading at anthesis; stamens (7–) 11–22 (–30?), filaments in a single series, far exserted; anthers bithecal, not apiculate; disc well developed, lateral to ovary, incised; ovary superior, ovoid-conical, glabrous to hairy, bicarpellate and bilocular, with 2 styles; ovules ~14–36 per locule, in 2 rows. Floral formula: B K4* C0 A¥* at anthesis, G(2)* Vx¥. Fruits capsular, cylindrical or ovoid (G. hirsuta only), dehiscing from the apex into 2 valves, often remaining united towards the base, eventually detaching from placentae which then form a free central column; exocarp glabrous, velutinous, or sparsely to densely coarse-hairy; seeds 182 Australian Systematic Botany numerous, flat, elliptic or round (G. hirsuta only) in outline, winged. Nineteen species from montane islands of the south-western Pacific: 13 in New Caledonia, four in Fiji, one in Vanuatu and one in the Solomon Islands (Hopkins 2006, 2007; Hopkins and Pillon 2011), in various forest types, including rainforest, riverine forest and open forest, and sometimes in maquis (scrubby vegetation especially on ultramafic soils). Notes Approximately two thirds of the species occur in New Caledonia (Hopkins et al., in press), and of these, approximately half grow on ultramafic substrates, suggesting that this local radiation is related, at least in part, to an ability to adapt to the unusual conditions found on such soils. Several species are hyperaccumulators of nickel (Jaffré et al. 1979). Outside New Caledonia, Geissois denhamii (Vanuatu) and G. ternata (Fiji) are markedly similar in their overall morphology and show similar variation in leaf size and in the density of indumentum on their stipules and ovaries. However, they can be distinguished by their calyx lobes (densely hairy on the adaxial surface in G. denhamii but almost glabrous in G. ternata). The distinction of G. stipularis (Fiji) from G. ternata requires clarification. Geissois superba from Viti Levu (Fiji) is somewhat isolated morphologically among the species with either three, five or a mixture of three and five leaflets. Its inflorescences are densely flowered, pendulous (not projecting) and often axillary, the opposite stipules can be adpressed at their base to form a cupshaped structure that retains water (phytotelm), and the leaves are often large. G. hirsuta also has pendulous, axillary inflorescences and large leaves, but the overall resemblance between these two taxa is not marked. Geissois pentaphylla is confined to the small islands of Vanikoro in Temotu Province, Solomon Islands, and is most similar to G. denhamii. Vanikoro belongs to the Santa Cruz group, which is geographically closer to the northern islands of Vanuatu than to any of the large members of the Solomon Islands. The biogeographical affinities between Vanikoro and Vanuatu are also demonstrated by their shared occurrence of Agathis Salisb., which is absent from the remainder of the Solomon Islands (Whitmore 1969). 3. Lamanonia Vell., Florae fluminensis, 228 (1829) Type: Lamanonia ternata Vell., Florae fluminensis Icones 5, tab. 104 (1831). Synonyms: Belangera Cambess. in Saint-Hilaire et al., Flora brasiliae meridionalis 2, 204 (1830). Polystemon D.Don, Edinburgh New Philosophical Journal 9, 95 (1830). Key references: Leite (1983); Zickel and Leitão Filho (1993). Trees or rarely shrubs. Leaves opposite and decussate, petiolate, digitately compound; leaflets 3(–4–)5, dentate with semi-craspedodromous venation, domatia often present; stipels at distal end of petiole caducous. Stipules 4 per node, lateral and free, often caducous. Inflorescence of simple racemes inserted in axils of distal leaves. Flowers bisexual, apetalous, white to yellow; pedicels articulating at or near base; calyx lobes 6, narrowly triangular, valvate in bud, spreading to reflexed at H. C. F. Hopkins et al. anthesis; stamens ~25–60, in >1 series, exserted; anthers bithecal, not apiculate; disc adnate to base of ovary, ribbed, rather inconspicuous in dried material; ovary superior, ovoid, tomentose, bicarpellate and bilocular, with 2 styles; ovules ~16–20 per locule, in 2 rows. Floral formula: B K6* C0 A¥* at anthesis [A6x* in bud] G(2)* Vx¥. Fruits capsular, ovoid, dehiscing from apex into 2 valves that often remain united towards the base by the placentae; exocarp sparsely to densely silky–velutinous; seeds numerous, flat, distally winged. Five species were recognised by Zickel and Leitão Filho (1993) from central, eastern and southern Brazil, Paraguay and northern Argentina, in various wooded habitats: L. brasiliensis Zickel & Leitão, L. chabertii (Pamp.) L.S.Sm., L. cuneata, L. grandistipularis, and L. ternata (which has numerous synonyms including L. glabra (Cambess.) Kuntze, L. speciosa (Cambess.) L.B.Sm., L. tomentosa (Cambess.) Kuntze, and L. ulei). Notes The description and illustration in Flora Fluminensis (Vellozo 1829, 1831) are difficult to interpret at species level because this is a closely knit genus and differences among species are generally small. Zickel and Leitão Filho (1993) indicated that although the flowers are relatively uniform, minor differences occur in the leaves, stipules and fruits that are useful in distinguishing among taxa. The density of the indumentum on the leaves varies within the widespread and abundant Lamanonia ternata, and Zickel and Leitão Filho (1993) included L. ulei from mountains in the State of Rio de Janeiro within this taxon. However, the indumentum on the lower surface of the leaflets in L. ulei is dense and pale and obscures the intervenium. This differs from the indumentum in other material of L. ternata, apparently without any intermediate forms, indicating that L. ulei is worthy of specific recognition (specimens examined: Brazil, Prov. Rio de Janeiro, Nova Friburgo, Pedra do Conico, 1300 m, January 1898, Ule Herb. Bras. 4551 [given as 4581 in the protologue] [holotype: B – image!]; State of Rio de Janeiro, Município de Santa Maria Madalena: Santa Maria Madalena, Pedra Dubois, 21580 S, 42010 W, 900–1195 m, 22 February 1983, Plowman & de Lima 12870 [K]). Fossil leaves from early Tertiary sediments in North Dakota were referred to Lamanonia by Hickey (1977). The fossil laminae are similar to individual leaflets of Lamanonia because some are slightly asymmetric, but semi-craspedodromous venation and toothed margins occur in genera in both the Cunoniaceae and Elaeocarpaceae, and because Hickey (1977) provided no unambiguous synapormophy, it is not possible to refer these leaves, with certainty, to an extant genus or family. 4. Pseudoweinmannia Engl., Naturlichen Pflanzenfamilien ed. 2, 18a, 249 (published 1928, dated 1930) Lectotype: Pseudoweinmannia lachnocarpa (F.Muell.) Engl., New South Wales, Tweed, [W.] Guilfoyle s.n., s.dat. (MEL104457) (Rozefelds and Pellow 2011). Key reference: Rozefelds and Pellow (2011). Trees. Leaves opposite and decussate, petiolate, digitately 3foliolate, leaflets dentate with semi-craspedodromous venation, domatia absent; stipels at distal end of petiole caducous. Stipules Karrabina gen. nov. (Geissoieae: Cunoniaceae) 2 per node and bifid when young (developing from 4 primordia), interpetiolar, lanceolate when fully grown, fugaceous. Inflorescence racemose, racemes single in opposite pairs, at leafy or leafless nodes, or in IMs (dyads, triads, tetrads), sometimes arising from auxiliary buds. Flowers bisexual, apetalous, whitish; pedicels articulating at or near base; calyx lobes (4–)5–7(–8), elliptic, valvate in bud, somewhat spreading at anthesis; stamens 12–25, in a single series, filaments flattened at base, shortly exserted; anthers bithecal, not apiculate; disc lateral to ovary, minutely incised, dark; ovary superior, ovoid, tomentose, bicarpellate and bilocular, with 2 styles; ovules 4–6 (–8) per locule. Floral formula: B K5–7* C0 A¥* at anthesis [A5–7x* in bud], G(2)* Vx8–12. Fruits indehiscent, almost spherical, exocarp densely covered with long hairs, calyx and styles persistent, placental proliferations giving appearance of 4 locules; seeds up to 4 per fruit, rounded pyramidal, not winged. Two species from rainforest in eastern Australia: P. lachnocarpa from south-eastern Queensland and northeastern New South Wales; P. apetala (Bailey) Engl. from north-eastern Queensland. Notes Both taxa in Pseudoweinmannia were formerly placed in Weinmannia (von Mueller 1872–1874; Bailey 1893) and P. lachnocarpa was included in Geissois by Maiden (1917); however, since the name Pseudoweinmannia was published by Engler in 1928, its position as a distinct genus has not been questioned in the Australian literature (e.g. Francis and Chippendale 1970; Harden 1990; Hyland and Whiffin 1993). Acknowledgements Helen Hopkins thanks the Royal Botanic Gardens, Kew, and the Muséum National d’Histoire Naturelle, Paris, for herbarium and library facilities; Richard Barnes for unpublished information on micromorphology; Jason Bradford for discussions on the Geissoieae, especially the structure of the inflorescence, and for material of Cunoniaceae from Australia; Peter Endress for help with the terminology of androecia; Carmen Zickel for a photograph of Lamanonia cuneata; Steve Bachman, Paul Little and John Stone for technical help with the illustrations; and Raj Saxena for advice on copyright. Andrew Rozefelds thanks Patricia O’Connor (Yugambeh Museum) for providing advice on the pronunciation and orthography of the name Karrabina; Margaret Sharpe and Sylvia Haworth advised on Aboriginal word usage in southern Queensland and on the most appropriate name and spelling for the new genus; BRI and K kindly arranged the loan of comparative material for study. 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