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Taxonomic evaluation of Cyrtandra (Gesneriaceae) from Sulawesi has resulted in the recognition of 39 species. Of these, four species are described as new here: Cyrtandra balgooyi H.J.Atkins & Karton., C. flavomaculata H.J.Atkins & Karton., C. longistamina H.J.Atkins & Karton. and C. parvicalyx H.J.Atkins & Karton. Eleven names are lectotypified and three names are placed in synonymy. Provisional conservation assessments suggest that seven of these species are Critically Endangered, six are Endangered, three are Vulnerable, two are Data Deficient and 21 are of Least Concern. Illustrations, distribution maps, and a key to the Cyrtandra species of Sulawesi are presented. Evaluasi taksonomi Cyrtandra (Gesneriaceae) dari Sulawesi menghasilkan pengenalan 39 jenis. Diantara jenis-jenis tersebut, empat jenis dideskripsikan sebagai jenis yang baru disini: Cyrtandra balgooyi H.J.Atkins & Karton., C. flavomaculata H.J.Atkins & Karton., C. longistamina H.J.Atkins & Karton. and C. parvicalyx H.J.Atkins & Karton. Sebelas nama telah dilektotipifikasikan dan tiga nama ditempatkan dalam sinonim. Penilaian konservasi sementara menyarankan bahwa tujuh dari jenisjenis ini dinyatakan Critically Endangered, enam dinyatakan Endangered, tiga dinyatakan Vulnerable, dua dinyatakan Data Deficient dan 21 sisanya dinyatakan Least Concern. Ilustrasi, peta distribusi dan kunci identifikasi untuk jenis-jenis Cyrtandra dari Sulawesi disajikan.

Tropical rainforest communities are often characterized by a small number of species-rich genera that contribute disproportionately to the alpha diversity in these habitats. In the Pacific Basin there are nearly 200 species of Cyrtandra, most of which are white-flowered woody shrubs that are single-island endemics. Within these island communities, multiple Cyrtandra species are commonly observed to occur sympatrically in wet forest understories, forming swarms of what appear to be ecologically similar taxa. The aim of this study was to determine if communities of these plants are randomly assembled with respect to phylogenetic relatedness and traits that are ecologically relevant. Using a combination of ten functional traits and a well-resolved species phylogeny, I examined community assembly within 34 species of Cyrtandra across three Pacific archipelagoes. Coexisting species were generally found to be more closely related and more phenotypically similar than would be expected by chance. This pattern was observed at both broad (island communities) and fine (site communities) spatial scales. The retention of phylogenetic signal in floral traits and the strong influence of these traits on the observed degree of phylogenetic clustering may indicate that pollinators act as a biotic filter for closely related species of Cyrtandra. In contrast, the absence of phylogenetic signal in most leaf traits, coupled with the lower contribution of these traits to niche clustering, suggests that environmental filtering along this trait axis is minimal in the observed communities. This study supports the theory that plant communities are not randomly assembled, and instead, that niche-based processes structure biodiversity at broad and fine spatial scales in diverse congeneric species assemblages.

Cyrtandra (Gesneriaceae), with over 800 species, is a mega-diverse genus which presents considerable taxonomic challenges due to its size. A well-sampled phylogeny of the genus across Southeast Asia has confirmed that all but one of the sections within Clarke’s 1883 genus-wide infrageneric classification are polyphyletic. It also shows that there are high levels of homoplasy in key morphological characters, although it is possible to use morphological characters to define clades in parts of the phylogenetic tree. There is some geographic structure in the phylogeny, but there is also evidence of dispersal between islands. A practical approach for tackling the taxonomy of Cyrtandra in the region, through phylogenetically informed taxonomic revisions of geographic areas, an approach which combines evidence from molecular, morphological and distribution data, is discussed. Completing our understanding of species diversity and delimitation in this genus will allow us to maximise the use of Cyrtandra as a tool for studying biogeography, speciation, diversification and conservation prioritisation in the rainforests of Southeast Asia.

Cyrtandra tempestii, a species endemic to the southeast coast of Taveuni, Fiji, is reevaluated in light of recent field collections and a thorough examination of herbarium specimens. A revised description of this species based on observations and morphological measurements from the field is presented here. Cyrtandra tuiwawai sp. nov. was formerly included under C. tempestii based on similarities in floral morphology and is described here as a new species endemic to the montane forests of Taveuni. Habitat preference, as well as morphological characters, distinguish the two species from one another. The addition of C. tuiwawai brings the total number of Cyrtandra species in Fiji to 42, and supports the need for continued fieldwork in the region.

As part of ongoing molecular phylogenetic work on the large Gesneriaceae genus Cyrtandra, new insights into the taxonomy and relationships of the Cyrtandra of Japan, Taiwan and Batan Island in the northern Philippines have emerged. Cyrtandra umbellifera is confirmed as a species with a distribution that includes both Taiwan and Batan Island. Cyrtandra yaeyamae is found to be distinct from the widespread C. cumingii, with a distribution that includes both the Ryukyu Islands in Japan and Batan Island.

Tropical rain forests are the most biologically diverse of terrestrial biomes. A few species-rich genera contribute disproportionately to this diversity and yet have often been neglected taxonomically because their sheer size makes them so challenging. Understanding the causes and mechanisms of these lineage diversifications is, however, central to understanding the evolution of plant diversity and for effectively planning its conservation. To realise the potential of these large genera, timecalibrated, phylogenetic studies are required to place the species in their evolutionary and biogeographic context, effectively plan future taxonomic work and to investigate the patterns and processes of their diversification. Cyrtandra is the largest genus in the Gesneriaceae with over 800 species of herbs, shrubs and occasionally trees, and is one of the fifty largest plant genera in the world. It is found from the Nicobar Islands in the Indian Ocean, throughout Malesia, in Taiwan and the southern Japanese islands, in northern Australia and east across Polynesia to Hawaii. It is found in forest, from sea level to 3000m. Taxonomic work is required throughout the distribution of Cyrtandra but primarily in its centres of diversity in Southeast Asia, on islands such as Sulawesi and Borneo. There is no functional infrageneric classification in the genus greatly increasing the difficulty of working with the group and hampering the planning of future taxonomic work. This study includes a taxonomic revision of Cyrtandra on the Indonesian island of Sulawesi which has resulted in the recognition of 40 species. Of these, five species are highlighted as new in this thesis: Cyrtandra longistamina H.J.Atkins & Karton. ined., Cyrtandra balgooyi H.J.Atkins & Karton. ined. Cyrtandra flavomaculata H.J.Atkins & Karton. ined., Cyrtandra parvicalyx H.J.Atkins & Karton. ined. and Cyrtandra sopuensis H.J.Atkins & Karton. ined. Eleven species are lectotypified and three names are placed in to synonymy. Provisional conservation assessments show that two of these species are Critically Endanagered, one is Endangered, thirteen are Vulnerable, one is Near Threatened and 22 are of Least concern. Generation of a well-sampled phylogeny of Cyrtandra across Southeast Asia and the Pacific and optimisation of eight macromorphological characters, as part of the current study, has allowed the development of a strategy for tackling this large genus taxonomically in the future in the absence of a functional infrageneric system. The poor diagnosability of the six major clades, the homoplasy of the macromorphological characters, combined with some degree of floristic exchange between islands, showed that a single approach, either based on geography, morphology or molecular data, would not be satisfactory for Cyrtandra and, similar to other large genera, an integrative method would be most successful. I assessed Clark et al. (2013)’s concept of a phylogenetically informed revision of a geographic area on the Cyrtandra of Sulawesi and Borneo and found it to be a promising approach. I now have a key to fourteen lineages on Borneo that can form the basis for taxonomic research on the 200 plus species on that island in the future. Molecular dating, ancestral range estimation and biogeographic stochastic mapping included here have allowed the examination of the geo-temporal patterns and evolution of southeast Asian Cyrtandra and to show that the great diversity of Cyrtandra seen in the Malesian region results from a recent radiation, with most speciation taking place in the last five million years. Borneo was recovered as the most likely ancestral range of the genus, with the current distribution of species resulting from a west to east migration across Malesia that corresponds with island emergence and mountain building. High levels of floristic exchange between the islands on the Sunda shelf and an important role for the Philippines as a stepping stone to Wallacea and New Guinea are uncovered. These patterns underlie much of the plant diversity in the region and form an emerging paradigm in Southeast Asian plant biogeography. Novel observations on speciation dynamics for southeast Asian Cyrtandra made here include (i) the absence of hybridisation, possiby due to the role of pre-zygotic barriers associated with high levels of floral divergence in sympatric populations in southeast Asia; (ii) a remarkably stable genome with no variation in chromosome numbers recorded across its distribution, including the Pacific. These two observations are notable as they are in contrast to other large genera such as Begonia, Inga and Solanum where hybridisation and ploidy changes are common. The dense phylogenetic sampling for interspecific and intraspecific diversity, combined with new trait and geographical datasets have provided much insight into the origin of the diversity of Cyrtandra. Future work can focus on the taxonomy of Cyrtandra on the large islands of Borneo and New Guinea and on the genomic architecture of species differences.

Cyrtandra argentii Olivar, H.J.Atkins & Muellner sp. nov., endemic to the Philippines and named after George Argent, is herein described and illustrated. Collections associated with this new species are often confused with three other species, namely C. ferruginea Merr., C. villosissima Merr., and C. hirtigera H.J.Atkins & Cronk. Distinguishing characters including keys, updated descriptions, distribution maps, and photos of live specimens are provided to aid identification of the four species. Following the International Union for the Conservation of Nature (IUCN) criteria, C. argentii sp. nov. is considered to be Near Threatened (NT) due to its distribution in a zone susceptible to anthropogenic pressure and the lack of any formal protection.

Genus Cyrtandra J.R.Forst. & G.Forst. (Forster & Forster 1775) Key to the studied species 1. Mature leaves anisophyllous (i.e., smaller leaves less than half the length of the larger leaves in a pair)..................................................................................................................... C. ferruginea Merr. ��� Mature leaves subequal..................................................................................................................... 2 2. Indumentum white; inflorescences pendulous and pedunculate.................................................................................................................................... C. argentii Olivar, H.J. Atkins & Muellner sp. nov. ��� Indumentum ferruginous; inflorescences erect and subsessile......................................................... 3 3. Calyx divided almost to the base; corolla red.................................................... C. villosissima Merr. ��� Calyx fused for half or more of its length; corolla yellowish green or dull reddish orange........................................................................................................................ C. hirtigera H.J.Atkins & Cronk, Published as part of Olivar, Jay Edneil C., Atkins, Hannah J., Hauenschild, Frank & Muellner-Riehl, Alexandra N., 2020, Cyrtandra argentii, a new species of Cyrtandra (Gesneriaceae) from the Philippines, and a review of the C. villosissima group, pp. 1-15 in European Journal of Taxonomy 676 on page 4, DOI: 10.5852/ejt.2020.676, http://zenodo.org/record/3928370

Cyrtandra ferruginea Merr. (Merrill 1915) Figs 1, 3B, 4B Material examined Type PHILIPPINES ��� Luzon, Camarines, Mt Cauayan; 9 Dec. 1913; Phil. Pl. Ramos 1548 (syntypes: BM!, GH!, NY!, P!, US!). Additional material PHILIPPINES ��� Mt Isarog, Camarines Sur; Aug. 1915; Ramos 23554 (US00081328) ��� ibid.; 22 Mar. 1997; Mendum et al. 29182 (E00057041) ��� ibid.; 23 Mar. 1997; Argent et al. 20182 (L.3805694) ��� Mt Bulusan, Sorsogon; Dec. 1915; Elmer 16074 (L.2818244, US00081329, U.1341267, P03884333) ��� ibid.; 19 Jun. 1958; Sinclair 9624 (E00631523) ��� Mt Mayon, Albay; 15 Nov. 1991; Reynoso, Romero & Fuentes 3584 (E00316099) ��� Catanduanes; 11 Dec. 1917; Ramos 30288 (US00081330, P03884332) ��� Mt Malinao, Albay; 29 Oct. 1995; Reynoso, Sagcal & Fernando 21406 (L.3805666). Description An erect suffrutescent plant up to 1 m in height. Stems terete with ferruginous hairs all throughout. Leaves opposite, anisophyllous; petioles 4���7 cm long, densely hirsute; blades ca 20 �� 12 cm, oblong to oblong-elliptic slightly falcate, apex acute or slightly acuminate, base acute or rounded, pronouncedly asymmetrical, not decurrent, margins denticulate, 10 pairs of lateral veins, curving and uniting at the margin, densely hirsute on both sides; blades of smaller leaves of a pair 6.5���9�� 2���2.5 cm, resembling the major leaves in all other respects. Inflorescences cymose, axillary, erect, pedunculate, with 1���3 flowers; peduncle 2���3 cm long, densely hirsute; bracts green, ca 10 �� 1 mm, linear lanceolate, densely hirsute on both surfaces, persistent. Calyx tubular, pale green, 20���30 mm long, upper lobes ca 3 mm long, lower lobes ca 7 mm long, acuminate, densely hirsute externally, with glandular hairs internally at base of lobes. Corolla white, 50���65 mm long, funnel-shaped, upper lobes rounded, ca 8�� 9 mm, lower and lateral lobes rounded, ca 5�� 6 mm, densely hirsute, with glandular hairs internally, lobes slightly recurved. Stamens 2; filaments ca 10 mm long, attached ca 18 mm from base of corolla, sparsely covered with glandular hairs; anthers ca 1.5 mm long, thecae parallel, coherent at apices; staminodes 3, lateral staminodes ca 4 mm long, central staminode ca 1 mm long. Gynoecium ca 25 mm long overall; disc cupular with undulate margin, ca 2 mm long, glabrous; ovary 8���9 mm long, with glandular hairs throughout; style ca 12 mm long, with glandular hairs throughout; stigma bilobed, ca 1 mm across. Fruits lanceolate, green, hirsute, verrucose, 30���40 �� 6 mm; calyx persistent and entirely enclosing the fruit, style persistent. Distribution and habitat Cyrtandra ferruginea is found growing in damp forests at approximately 500���800 m a.s.l. and can be found on Catanduanes, Mt Isarog, Mt Mayon, Mt Malinao and Mt Bulusan (Fig. 1). Notes Cyrtandra ferruginea is morphologically most similar to C. argentii sp. nov., but can be separated by the following characters: ferruginous anisophyllous leaves, 1���3 flowered simple cymes, and hirsute corolla., Published as part of Olivar, Jay Edneil C., Atkins, Hannah J., Hauenschild, Frank & Muellner-Riehl, Alexandra N., 2020, Cyrtandra argentii, a new species of Cyrtandra (Gesneriaceae) from the Philippines, and a review of the C. villosissima group, pp. 1-15 in European Journal of Taxonomy 676 on pages 7-9, DOI: 10.5852/ejt.2020.676, http://zenodo.org/record/3928370, {"references":["Merrill E. D. 1915. New or noteworthy Philippine plants, XI. The Philippine Journal of Science. Section C, Botany 10: 1 - 84.","Pelser P. B., Barcelona J. F. & Nickrent D. L. 2011 onwards. Co's Digital Flora of the Philippines [online]. Available from www. philippineplants. org [accessed 2 May 2019]."]}

Genus Cyrtandra J.R.Forst. & G.Forst. (Forster & Forster 1775) Key to the studied species 1. Mature leaves anisophyllous (i.e., smaller leaves less than half the length of the larger leaves in a pair)..................................................................................................................... C. ferruginea Merr. – Mature leaves subequal..................................................................................................................... 2 2. Indumentum white; inflorescences pendulous and pedunculate.................................................................................................................................... C. argentii Olivar, H.J. Atkins & Muellner sp. nov. – Indumentum ferruginous; inflorescences erect and subsessile......................................................... 3 3. Calyx divided almost to the base; corolla red.................................................... C. villosissima Merr. – Calyx fused for half or more of its length; corolla yellowish green or dull reddish orange........................................................................................................................ C. hirtigera H.J.Atkins & Cronk

Cyrtandra hirtigera H.J.Atkins & Cronk (Atkins & Cronk 2001 ) Figs 1, 3C, 4C Material examined Type PHILIPPINES • Palawan, Cleopatra’s Needle; 29 Jan. 1998; Cronk et al. 25433 (holotype: PNH!; isotypes: E!, K!, L!). Additional material PHILIPPINES • San Vicente, Palawan; 4 Aug. 1988; Soejarto & Madulid 6353 (L.2818243) • Mt Beaufort, Palawan; 12 Mar. 1984; Ridsdale SMHI 23 (L.2818051) • Pagdanan Range, Palawan; 22 Apr. 1984; Podzorski SMHI 934 (L.2818049) • Mt Mantalingahan, Palawan; 5 May 1948; Edaño 122 (L.2818048) • ibid.; 4 Mar. 1992; Argent & Romero 9666 (L.3805814) • Malampaya Bay, Palawan; Oct. 1922; Merrill 11573 (US00081506) • Mt Capoas, Palawan; Apr. 1913; Merrill 9500 (US00081487) • Palawan; Apr. 1906; Foxworthy 581 (US00081485, P03899657) • Taytay, Palawan; 31 Jan. 1991; Stone 327 (L.2822680). Description An erect suffrutescent plant, up to 2–2.5 m in height. Stems terete, with crimson or white hairs throughout. Leaves opposite, subequal; petioles 5–8 cm long, densely hirsute; blades 22–30 × 15–18 cm, broadly elliptic, slightly falcate, apex acuminate, base cuneate, not decurrent, margins denticulate, 12–14 pairs of lateral veins, curving and uniting at the margins, densely hirsute on both sides. Inflorescences cymous, axillary, erect, subsessile, with 4–many flowers; peduncle 4–5 mm, densely hirsute; bracts green, ca 1 cm × 5 mm, lanceolate, densely hirsute on both surfaces, persistent; bracteoles up to 5 mm long, densely hirsute on both surfaces; pedicels ca 5 mm long, densely hirsute. Calyx tubular, red or green, ca 1–1.5 cm long, lobes ca 5 mm long, lower lobes ca 6 mm long, acute or narrowly acuminate, densely hirsute externally, with scattered glandular hairs internally and has prominent tufts of hairs at the base. Corolla dull reddish orange or yellowish green, ca 2 cm long, funnel-shaped, lobes slightly bilabiate or subequal, upper lobes, 1–1.5 × 2 mm, lateral lobes 1× 1.5 mm, lower lobes 2× 1.5 mm; glandular hairs externally and internally. Stamens 2; filaments ca 1–1.5 cm long, attached ca 12–13 mm from base of corolla, glabrous; anthers ca 2 mm long, thecae parallel, coherent at apices; staminodes 2, 5– 8 mm long. Gynoecium ca 20 mm long overall; disc cupular with undulate margin, ca 1.5 mm long, glabrous; ovary ca 4–5 mm long, glabrous; style ca 10 mm long, with eglandular hairs throughout; stigma bilobed, ca 2.5 mm across. Fruits ovoid, green, glabrous, ca 1 cm × 5 mm; inflorescence bracts persistent, calyx and style not persistent. Distribution and habitat Cyrtandra hirtigera is distributed throughout the island of Palawan and is usually found on slopes near gullies at 30–900 m a.s.l. Notes Atkins & Cronk (2001) described two varieties of this species, C. hirtigera var. hirtigera and C. hirtigera var. chlorina, distinguishable by color and shape of their calyces and corolla limbs. Cyrtandra hirtigera var. hirtigera has a crimson indumentum, red calyces with acute lobe apices, and reddish orange corollas with slightly bilabiate limbs. Cyrtandra hirtigera var. chlorina has pale indumentum, green calyces with acuminate lobe apices, and yellowish green corollas with subequal lobes.

Cyrtandra villosissima Merr. (Merrill 1906) Figs 1, 3D, 4D Material examined Type PHILIPPINES ��� Mindanao, Camp Keithley, Lake Lanao; Jan. 1906; Mrs Clemens 51 (holotype: AAH!; isotypes: F!, US!). Additional material PHILIPPINES ��� Mt Malindang, Misamis Occidental; 16 Mar. 2004; Opiso et al. 2088 (L.3794253) ��� ibid.; May 1993; Gaerlan et al. 10925 (K000184579, L.3794112) ��� Mt Hibok-Hibok, Camiguin; 1999; RBGE & PNH 48 (E00743749) ��� Lake Balunsasayao, Negros Oriental; 11 Sep. 1953; Britton 357 (L.2826671) ��� Cuernos Mountains, Negros Oriental; 13 May 1948; Eda��o 7393 (AAH00092001) ��� ibid.; Mar. 1908; Elmer 9511 (L.2826672, US00081486) ��� Mahilucot River, Bukidnon; Jul. 1920; Ramos & Eda��o 38649 (L.2826670, US00081488) ��� Mt Daho, Jolo; Sep. 1924; Ramos & Eda��o 43913 (P03899658). Description An erect suffrutescent plant up to 7m in height. Stems terete or slightly grooved,with ferruginous woolly hairs throughout. Leaves opposite, subequal; petioles 3���5 cm long, densely hirsute; blades 11���20�� 3.5���8 cm, oblong-ovate to ovate-lanceolate, slightly falcate, apex acuminate, base acute or acuminate, pronouncedly asymmetrical, not decurrent, margins denticulate, 12���14 pairs of lateral veins, curving and uniting at the margins, densely hirsute on both sides. Inflorescences cymous, axillary, erect, subsessile, with 4���many flowers; peduncle 4���5 mm, densely hirsute; bracts green, ca 1 cm, linear, densely hirsute on both surfaces, persistent; bracteoles up to 5 mm long, densely hirsute on both surfaces; pedicels ca 5 mm long, densely hirsute. Calyx tubular, pale green, lobes linear ca 1.5 cm �� 1 mm, densely hirsute. Corolla red, ca 18 mm long, funnel-shaped, lobes orbicular-ovate ca 4 mm long, densely hirsute. Stamens 2; filaments ca 2 mm long; anthers ca 2.5 mm long, thecae parallel, coherent at apices. Gynoecium ca 20 mm long overall, densely hirsute; disc cupular, glabrous; style densely hirsute. Fruits oblong, green, densely hirsute, ca 1 cm �� 4.5 mm; calyx and style persistent. Distribution and habitat Cyrtandra villosissima is distributed throughout the island of Mindanao and extends to the island of Negros in the Visayas and is usually found in well-shaded areas near ravines. Notes Cyrtandra villosissima is vegetatively similar to C. hirtigera. It is distinguishable by its red corolla and green calyces with distinctly linear lobes. Based on available distribution data, C. hirtigera appears to be restricted to the island of Palawan while C. villosissima can be found from Negros Island to the island of Mindanao., Published as part of Olivar, Jay Edneil C., Atkins, Hannah J., Hauenschild, Frank & Muellner-Riehl, Alexandra N., 2020, Cyrtandra argentii, a new species of Cyrtandra (Gesneriaceae) from the Philippines, and a review of the C. villosissima group, pp. 1-15 in European Journal of Taxonomy 676 on page 11, DOI: 10.5852/ejt.2020.676, http://zenodo.org/record/3928370, {"references":["Merrill E. D. 1906. New or noteworthy Philippine plants, V. Philippine Journal of Science 1 (Supplement): 169 - 246."]}

Cyrtandra hirtigera H.J.Atkins & Cronk (Atkins & Cronk 2001 ) Figs 1, 3C, 4C Material examined Type PHILIPPINES ��� Palawan, Cleopatra���s Needle; 29 Jan. 1998; Cronk et al. 25433 (holotype: PNH!; isotypes: E!, K!, L!). Additional material PHILIPPINES ��� San Vicente, Palawan; 4 Aug. 1988; Soejarto & Madulid 6353 (L.2818243) ��� Mt Beaufort, Palawan; 12 Mar. 1984; Ridsdale SMHI 23 (L.2818051) ��� Pagdanan Range, Palawan; 22 Apr. 1984; Podzorski SMHI 934 (L.2818049) ��� Mt Mantalingahan, Palawan; 5 May 1948; Eda��o 122 (L.2818048) ��� ibid.; 4 Mar. 1992; Argent & Romero 9666 (L.3805814) ��� Malampaya Bay, Palawan; Oct. 1922; Merrill 11573 (US00081506) ��� Mt Capoas, Palawan; Apr. 1913; Merrill 9500 (US00081487) ��� Palawan; Apr. 1906; Foxworthy 581 (US00081485, P03899657) ��� Taytay, Palawan; 31 Jan. 1991; Stone 327 (L.2822680). Description An erect suffrutescent plant, up to 2���2.5 m in height. Stems terete, with crimson or white hairs throughout. Leaves opposite, subequal; petioles 5���8 cm long, densely hirsute; blades 22���30 �� 15���18 cm, broadly elliptic, slightly falcate, apex acuminate, base cuneate, not decurrent, margins denticulate, 12���14 pairs of lateral veins, curving and uniting at the margins, densely hirsute on both sides. Inflorescences cymous, axillary, erect, subsessile, with 4���many flowers; peduncle 4���5 mm, densely hirsute; bracts green, ca 1 cm �� 5 mm, lanceolate, densely hirsute on both surfaces, persistent; bracteoles up to 5 mm long, densely hirsute on both surfaces; pedicels ca 5 mm long, densely hirsute. Calyx tubular, red or green, ca 1���1.5 cm long, lobes ca 5 mm long, lower lobes ca 6 mm long, acute or narrowly acuminate, densely hirsute externally, with scattered glandular hairs internally and has prominent tufts of hairs at the base. Corolla dull reddish orange or yellowish green, ca 2 cm long, funnel-shaped, lobes slightly bilabiate or subequal, upper lobes, 1���1.5 �� 2 mm, lateral lobes 1�� 1.5 mm, lower lobes 2�� 1.5 mm; glandular hairs externally and internally. Stamens 2; filaments ca 1���1.5 cm long, attached ca 12���13 mm from base of corolla, glabrous; anthers ca 2 mm long, thecae parallel, coherent at apices; staminodes 2, 5��� 8 mm long. Gynoecium ca 20 mm long overall; disc cupular with undulate margin, ca 1.5 mm long, glabrous; ovary ca 4���5 mm long, glabrous; style ca 10 mm long, with eglandular hairs throughout; stigma bilobed, ca 2.5 mm across. Fruits ovoid, green, glabrous, ca 1 cm �� 5 mm; inflorescence bracts persistent, calyx and style not persistent. Distribution and habitat Cyrtandra hirtigera is distributed throughout the island of Palawan and is usually found on slopes near gullies at 30���900 m a.s.l. Notes Atkins & Cronk (2001) described two varieties of this species, C. hirtigera var. hirtigera and C. hirtigera var. chlorina, distinguishable by color and shape of their calyces and corolla limbs. Cyrtandra hirtigera var. hirtigera has a crimson indumentum, red calyces with acute lobe apices, and reddish orange corollas with slightly bilabiate limbs. Cyrtandra hirtigera var. chlorina has pale indumentum, green calyces with acuminate lobe apices, and yellowish green corollas with subequal lobes., Published as part of Olivar, Jay Edneil C., Atkins, Hannah J., Hauenschild, Frank & Muellner-Riehl, Alexandra N., 2020, Cyrtandra argentii, a new species of Cyrtandra (Gesneriaceae) from the Philippines, and a review of the C. villosissima group, pp. 1-15 in European Journal of Taxonomy 676 on page 10, DOI: 10.5852/ejt.2020.676, http://zenodo.org/record/3928370, {"references":["Atkins H. [J.] & Cronk Q. C. B. 2001. The genus Cyrtandra (Gesneriaceae) in Palawan, Philippines. Edinburgh Journal of Botany 58 (3): 443 - 458. https: // doi. org / 10.1017 / s 0960428601000762"]}

The islands of Southeast Asia comprise one of the most geologically and biogeographically complex areas in the world and are a centre of exceptional floristic diversity, harbouring 45,000 species of flowering plants. Cyrtandra, with over 800 species of herbs and shrubs, is the largest genus in the family Gesneriaceae and is one of the most emblematic and species-rich genera of the Malesian rainforest understorey. The high number of species and tendency to narrow endemism make Cyrtandra an ideal genus for examining biogeographic patterns. We sampled 128 Cyrtandra taxa from key localities across Southeast Asia to evaluate the geo-temporal patterns and evolutionary dynamics of this clade. One nuclear and four chloroplast regions were used for phylogenetic reconstruction, molecular dating, and ancestral range estimation. Results from the dating analysis suggest that the great diversity of Cyrtandra seen in the Malesian region results from a recent radiation, with most speciation taking place in the last five million years. Borneo was recovered as the most likely ancestral range of the genus, with the current distribution of species resulting from a west to east migration across Malesia that corresponds with island emergence and mountain building. Lastly, our investigation into the biogeographic history of the genus indicates high levels of floristic exchange between the islands on the Sunda shelf and the important role of the Philippines as a stepping stone to Wallacea and New Guinea. These patterns underlie much of the plant diversity in the region and form an emerging paradigm in Southeast Asian plant biogeography.

Study on the used of medicinal plants by Marind people who lived at Wasur National Park, Merauke was conducted by using descriptive methods which include observation, interview, documentation, literature review techniques, as well as emic and ethic approaches. During the study there were 46 species which belong to 26 families plants found as medicinal plants used by Marind people to cure about 30 diseases. Among 46 species, there were 7 potential medicinal plant species which known and used by most Marind people. The seven potential medicinal plant species were Ipomoea triloba L., Bauhinia sp., Pittosporum sp., Kingiodendron platycarpum Bent., Sophora tool mentosa L. Cyrtandra sp., dan Tinopspora disstiflora L. Part of plant used vary from leaf, root, bark, fruit and other part of plant. Compare to other part of plant, leaf was the most common used as traditional medicinal plant. Key words : traditional medicinal plant, Marind people, Wasur National Park

Eleven new species of Cyrtandra (Gesneriaceae) from Sulawesi are described and illustrated: C. albiflora Karton. & H.J.Atkins, C. boliohutensis Karton. & H.J.Atkins, C. gambutensis Karton. & H.J.Atkins, C. hekensis Karton. & H.J.Atkins, C. hendrianii Karton. & H.J.Atkins, C. hispidula Karton. & H.J.Atkins, C. kinhoii Karton. & H.J.Atkins, C. multinervis Karton. & R.Bone, C. nitida Karton. & H.J.Atkins, C. rantemarioensis Karton. & R.Bone and C. rubribracteata Karton. & H.J.Atkins. Illustrations, maps and preliminary conservation assessments are provided for all the species.

Cyrtandra (Gesneriaceae) is among the largest genera of flowering plants in the remote oceanic islands of the Pacific, with an estimated 175 species distributed across an area that extends from the Solomon Islands, east to the Marquesas Islands, and north to the Hawaiian Islands. The vast majority of species are single-island endemics that inhabit upland rainforests. Although previous molecular phylogenetic studies greatly advanced our understanding of the diversification of Pacific Cyrtandra, a number of uncertainties remain regarding phylogenetic relationships, divergence times, and biogeographic patterns within this large and widely dispersed group. In the present study, five loci (ITS, ETS, Cyrt1, psbA-trnH, and rpl32-trnL) were amplified and sequenced for phylogenetic reconstruction of 121 Cyrtandra taxa. Maximum likelihood and Bayesian inference confirmed that C. taviunensis from Fiji is sister to the remaining members of the Pacific clade. Dating analyses and ancestral area estimation indicates that the Pacific clade of Cyrtandra originated in Fiji during the Miocene ca. 9mya. All major crown lineages within the Pacific clade appeared < 5mya, coincident with the emergence of numerous Pacific islands and a subsequent increase in available habitat. The biogeographic history of Cyrtandra in the Pacific has been shaped by extinction, dispersal distance, and founder events. Biogeographic stochastic mapping analyses suggest that cladogenesis within Pacific Cyrtandra involved a combination of narrow (within-area) sympatry and founder events. A mean of 24 founder events was recovered between Pacific archipelagos, while a mean of 10 founder events was recovered within the Hawaiian archipelago.

Cyrtandra (Gesneriaceae) is a genus of flowering plants with over 800 species distributed throughout Southeast Asia and the Pacific Islands. On the Hawaiian Islands, 60 named species and over 89 putative hybrids exist, most of which are identified on the basis of morphology. Despite many previous studies on the Hawaiian Cyrtandra lineage, questions regarding the reconciliation of morphology and genetics remain, many of which can be attributed to the relatively young age and evidence of hybridization between species. We utilized targeted enrichment, high‐throughput sequencing, and modern phylogenomics tools to test 31 Hawaiian Cyrtandra samples (22 species, two putative hybrids, four species with two samples each, one species with four samples) and two outgroups for species relationships and hybridization in the presence of incomplete lineage sorting (ILS). Both concatenated and species‐tree methods were used to reconstruct species relationships, and network analyses were conducted to test for hybridization. We expected to see high levels of ILS and putative hybrids intermediate to their parent species. Phylogenies reconstructed from the concatenated and species‐tree methods were highly incongruent, most likely due to high levels of incomplete lineage sorting. Network analyses inferred gene flow within this lineage, but not always between taxa that we expected. Multiple hybridizations were inferred, but many were on deeper branches of the island lineages suggesting a long history of hybridization. We demonstrated the utility of high‐throughput sequencing and a phylogenomic approach using 569 loci to understanding species relationships and gene flow in the presence of ILS.

Two new species of Cyrtandra (Gesneriaceae) are described; one apparently endemic to Yapen Island, Tanah Papua, the other present in Tanah Papua and in Papua New Guinea: both species are described and illustrated, and preliminary conservation assessments given. Notes on the identity of other known collections of Cyrtandra from Yapen are provided.

Summary Understanding of the role of environmental change in the decline of endangered species is critical to designing scale-appropriate restoration plans. For locally endemic rare plants on the brink of extinction, frugivory can drastically reduce local recruitment by dispersing seeds away from geographically isolated populations. Dispersal of seeds away from isolated populations can ultimately lead to population decline. For localized endemic plants, fine-scale changes in microhabitat can further limit population persistence. Evaluating the individual and combined impact of frugivores and microhabitat heterogeneity on the short-term (i.e. transient) and long-term (i.e. asymptotic) dynamics of plants will provide insight into the drivers of species rarity. In this study, we used four years of demographic data to develop matrix projection models for a long-lived shrub, Cyrtandra dentata (H. St. John & Storey) (Gesneriaceae), which is endemic to the island of O'ahu in Hawai'i. Furthermore, we evaluated the individual and combined influence of a non-native frugivorous bird, Leiothrix lutea, and microhabitat heterogeneity on the short-term and long-term C. dentata population dynamics. Frugivory by L. lutea decreased the short-term and long-term population growth rates. However, under the current level of frugivory at the field site the C. dentata population was projected to persist over time. Conversely, the removal of optimum microhabitat for seedling establishment (i.e. rocky gulch walls and boulders in the gulch bottom) reduced the short-term and long-term population growth rates from growing to declining. Survival of mature C. dentata plants had the greatest influence on long-term population dynamics, followed by the growth of seedlings and immature plants. The importance of mature plant survival was even greater when we simulated the combined effect of frugivory and the loss of optimal microhabitat, relative to population dynamics based on field conditions. In the short-term (10 years), however, earlier life stages had the greatest influence on population growth rate. Synthesis and applications. This study emphasizes how important it is to decouple rare plant management strategies in the short versus long-term in order to prioritize restoration actions, particularly when faced with multiple stressors not all of which can be feasibly managed. From an applied conservation perspective, our findings also illustrate that the life stage that, if improved by management, would have the greatest influence on population dynamics is dependent on the timeframe of interest and initial conditions of the population. This article is protected by copyright. All rights reserved.

International audience; Aim The diversity and composition of species pools within oceanic archipelagos is determined by a combination of colonization, abiotic tolerance, in situ diversification, biotic interactions and extinction. The signature of biogeographic events and evolutionary processes, however, may be masked by recent coalescence and hybridization between closely related species. We used the species-rich plant genus Cyrtandra (Gesneriaceae) to investigate the roles of colonization and hybridization in shaping community assemblages on the geologically young Hawai'i Island. Location Hawaiian Islands. Taxon Cyrtandra (Gesneriaceae). Methods We sampled 29 Cyrtandra taxa and putative hybrids across the main Hawaiian Islands and generated single nucleotide polymorphisms (SNPs) from nine single-copy nuclear genes. Maximum likelihood and Bayesian inference were used to reconstruct phylogenetic relationships, divergence times were estimated using secondary calibrations and island ages, and ancestral area estimation was done using likelihood models. Lastly, we used a Bayesian population assignment test and principal components analysis to infer population genetic structure. Results Cyrtandra colonization of the main Hawaiian Islands appears to have followed the progression rule, whereby the oldest high Hawaiian Islands (Kaua'i and O'ahu) were colonized first, followed by colonization of the younger islands as habitat became available. Hawai'i Island was colonized four times, with two dispersal events from O'ahu and two from Maui Nui. The different colonization events gave rise to significantly uneven numbers of species, and hybridization among the incipient lineages was detected in the form of intermediate genotypes. Main conclusions Our investigation into community assembly in a species-rich plant genus on a geologically young oceanic island revealed a history of multiple colonizations and hybridization among colonizing lineages. The rapid diversification (3.5 species/My) of one of four Cyrtandra lineages on Hawai'i Island may be the result of hybridization between genetically diverse lineages that stem from independent colonization events. Multiple colonization events followed by a merging of lineages may be particularly common during early-stage community assembly on islands and, through the generation of genetic variation, may be especially important for species diversification.

Cyrtandra Forster & Forster (1776: 5) is the largest genus in the family Gesneriaceae, with over 800 species distributed throughout Southeast Asia and the Pacific (Atkins et al. 2013). Members of the genus are important understory elements in primary forests, with varying habits (Burtt 2001). The Philippines is considered a center of diversity for the genus, with approximately 105–150 spp. (Atkins et al. 2013, Johnson et al. 2017).

Using a time-calibrated phylogenetic hypothesis including 768 Gesneriaceae species (out of [Formula: see text]3300 species) and more than 29,000 aligned bases from 26 gene regions, we test Gesneriaceae for diversification rate shifts and the possible proximal drivers of these shifts: geographic distributions, growth forms, and pollination syndromes. Bayesian Analysis of Macroevolutionary Mixtures analyses found five significant rate shifts in Beslerieae, core Nematanthus, core Columneinae, core Streptocarpus, and Pacific Cyrtandra These rate shifts correspond with shifts in diversification rates, as inferred by Binary State Speciation and Extinction Model and Geographic State Speciation and Extinction model, associated with hummingbird pollination, epiphytism, unifoliate growth, and geographic area. Our results suggest that diversification processes are extremely variable across Gesneriaceae clades with different combinations of characters influencing diversification rates in different clades. Diversification patterns between New and Old World lineages show dramatic differences, suggesting that the processes of diversification in Gesneriaceae are very different in these two geographic regions.

PREMISE OF THE STUDY Recent reviews of reproductive isolation (RI) in plants propose that boundaries between closely related species are maintained predominantly through prezygotic mechanisms. However, few experimental studies have explored how boundaries are maintained in long-lived species. Hawaiian Cyrtandra presents an intriguing challenge to our understanding of RI, as it comprises 60 shrub or small tree species that are almost exclusively restricted to wet forests, where sympatry of multiple species is common. METHODS We assessed the relative strengths of pre- and postzygotic barriers among four species of Cyrtandra occurring at the extremes of the main Hawaiian Island's natural island-age gradient, Kaua'i (4.7 Myr) and Hawai'i Island (0.6 Myr), to contrast the strengths and stages of reproductive isolation among species at different stages of divergence. KEY RESULTS A combination of F1 seed germination, F1 seedling survival, and F1 seedling growth isolated (61-91%) three of the species from sympatric relatives. In contrast, the fourth species was isolated (59%) from its sympatric relative through phenological differences alone. Significant postzygotic barriers in between-island crosses were also observed in one species. CONCLUSIONS Results suggest that boundaries between sympatric Cyrtandra species in Hawaii are maintained predominantly through postzygotic barriers. Observations from between-island crosses indicate that postzygotic barriers can arise in allopatry, which may be important in the initial divergence of populations. Future studies of RI in Cyrtandra should include a broader range of species to determine if postzygotic isolating barriers are foremost in the maintenance of species boundaries in this large genus.

A study on structure and association of ground cover plants commnunity was conducted in Cikiniki Resort, Gunung Halimun Salak National Park, West Java. The aimed of this study was to determine the structure and degree of association of ground cover plant that dominates with others ground cover plants. Plot observation using purposive sampling method. The sampling location was divided into two observation sites near the water source and away from the water source. We developed 20 plots with measure on 2 x 2 m2. The data obtained are frequency, density, dominance, Importance Value Index (INP), Distribution Index and Diversity Index Type. The relationship between abiotic factor and vegetation association was analyzed using Chi-Square analysis. The results showed that the dominant type was Schismatoglottis calyptrata (Roxb.) Zoll. & Moritzi (INP: 41.67%), Homalomena cordata Schott (INP: 16.87%), Cyrtandra populifolia Miq. (INP: 12.27%), Psychotria viridiflora Reinw. Ex Blume (INP: 11.79%) and Amischotolype mollissima (Blume) Hassk. (INP: 10,77%). Based on the calculation of chi-square (X2) associations S.calyptrata with other dominant species have low values of 0.26 (H. cordata), 0.25 (C. populifolia), 0.275 (P. viridiflora) and has no association with A. mollissima.

During fieldwork in Fiji, four new species of Cyrtandra (Gesneriaceae) were discovered and are described herein: C. gregoryi M.A.Johnson, sp. nov., C. hispida M.A.Johnson, sp. nov., C. longifructosa M.A.Johnson, sp. nov., and C. waisaliensis M.A.Johnson, sp. nov. The addition of four new species brings the current number of Fijian Cyrtandra to 41 endemic species. Two of the four species are known from only a single locality, and all of the new species are likely endangered or critically endangered. Continued fieldwork in the islands of Fiji is warranted in order to better understand current species distributions and population demographics of Cyrtandra in this species-rich and still poorly explored region of the South Pacific.

During the preparation of the Vascular Flora of the Marquesas Islands three new species of Cyrtandra (Gesneriaceae) have come to light and are described herein: C. uapouensis W. L. Wagner & Lorence, C. uahukaensis W. L. Wagner & Lorence, and C. kenwoodii W. L. Wagner & A. J. Wagner. Amended descriptions of the eight previously described Marquesan species are also provided as well as a key to the species. With the description of these the new species Cyrtandra in the Marquesas Islands consists of 11 species, six of which have been included in recent molecular phylogenetic studies of Pacific Cyrtandra, and appear to have arisen from one original introduction. If the other five species are members of this Marquesas clade then Cyrtandra would represent the largest lineage of Marquesas vascular plants. Psychotria is largest genus in the Marquesas Islands with 13 species, but is thought to consist of three separate lineages.

Four new species of Cyrtandra (Gesneriaceae) from the Latimojong Mountains, South Sulawesi (Sulawesi Selatan) are described and illustrated: C. floccosa R.Bone & H.J.Atkins, C. kjellbergii R.Bone & H.J.Atkins, C. purpureofucata R.Bone & H.J.Atkins, and C. spectabilis R.Bone & H.J.Atkins. Conservation recommendations are made for the four species.

Resolving species relationships within recent radiations requires analysis at the interface of phylogenetics and population genetics, where coalescence and hybridization may confound our understanding of relationships. We developed 18 new primer pairs for nuclear loci in Cyrtandra (Gesneriaceae), one of the largest plant radiations in the Pacific Islands, and tested the concordance of 14 loci in establishing the phylogenetic relationships of a small number of Hawaiian species. Four genes yielded tree topologies conflicting with the primary concordance tree, suggesting plastid capture and horizontal transfer via hybridization. Combining all concordant genes yielded a tree with stronger support and a different topology from the total-evidence tree. We conclude that a small number of genes may be insufficient for accurate reconstruction of the phylogenetic relationships among closely related species. Further, the combination of genes for phylogenetic analysis without preliminary concordance tests can yield an erroneous tree topology. It seems that the number of genes needed for phylogenetic analysis of closely related species is significantly greater than the small numbers commonly used, which fail to isolate coalescence, introgression and hybridization.

During the preparation of the Vascular Flora of the Marquesas Islands three new species of Coprosma (Rubiaceae, tribe Anthospermeae) have come to light and are described herein: Coprosma fatuhivaensis W. L. Wagner & Lorence, Coprosma meyeri W. L. Wagner & Lorence, and Coprosma temetiuensis W. L. Wagner & Lorence. Descriptions, illustrations, conservation status, and specimen citations are provided. Amended descriptions of three previously described Marquesan Coprosma species are also provided as well as a key to the species, four of which fall into the Critically Endangered (CR) and two into the Endangered (EN) category. With the description of these the new species, Coprosma becomes the sixth largest lineage in the Marquesas Islands with six species after Psychotria (one lineage which has 9 spp.), Cyrtandra (8 spp.), Bidens (8 spp.), Melicope (7 spp.), and Ixora (7 spp.).

Based on a considerably enlarged sampling, a phylogenetic analysis of the largest group of didymocarpoid Gesneriaceae, the “advanced Asiatic and Malesian genera”, was performed, covering all but 3 of the 60 genera presently recognised in this group (20 of these, mostly from China, are monotypic). The results suggest that no fewer than 17 out of the 57 genera examined are poly- (or rarely para-)phyletic. Highly polyphyletic are Briggsia, Chirita, Henckelia and Raphiocarpus. Only a dozen of the non-monotypic genera (including the three species-richest genera, Cyrtandra, Aeschynanthus and Agalmyla) are confirmed as monophyletic entities, though some exhibit considerable genetic variation. For eight genera, no statement can be made, as only one (of two or several) species was included in the analysis. For a dozen of the (particularly Chinese) monotypic genera a close relationship (or possible congenerity) with other genera was found. In China, only Allostigma, Cathayanthe, Conandron and Metapetrocosmea seem to have no strong affinities to other genera, indicating that they represent phylogenetically isolated lineages or represent remnants of previously larger and earlier diversified groups. The present study forms the foundation for targeted molecular, morphological and phytogeographic studies of the polyphyletic and monotypic genera and particular of clades of genera with interrelations uncovered here for the first time.

Cyrtandra sect. Dissimiles C. B. Cl., sect. Radicicaules Kraenzlin and sect. Radiciflorae Kraenzlin are lectotypified. Two varieties are raised to specific rank: C. dispar var. glabriflora B. C. Stone to C. stonei (Malay Peninsula), C. decurrens var. polyneura C. B. Cl. to C. polyneura (Sulawesi). Nineteen new species are described (11 from Sarawak, 1 from Sarawak and Kalimantan, 7 from Sabah—6 of them from Mt Kinabalu). Annotations are provided on 6 previously known species: C. calycina Benth. (New Guinea), C. debilis Kraenzlin (Kalimantan), C. farinosa C. B. Cl. (Sarawak), C. ligulifera C. B. Cl. (New Guinea), C. rubropicta Kraenzlin (Kalimantan and Sarawak), and C. russa C. B. Cl. (Sarawak).

The genus Cyrtandra is the largest in the Gesneriaceae family and is one of the most widely dispersed plant genera in southeast Asia and the Pacific. Species of Cyrtandra are morphologically diverse but characters are often homoplastic causing considerable difficulty in defining monophyletic classification units. In this study, we used molecular phylogenetic analysis of 88 taxa representing approximately 70 species to construct a well-resolved evolutionary hypothesis for Cyrtandra. Diversification rates analysis and ancestral range analysis were also conducted to infer timing of major lineage divergences and geographic origin of these lineages, principally among Pacific species. Using these data, we compared existing classification schemes to better understand the applicability of current taxonomy. Divergence time estimates support a diversification of the Pacific clade at approximately 20 MYBP. Although the origin of the Pacific lineage remains unresolved, ancestral range reconstruction analysis supports Fiji as the most likely "first-step" into the Pacific with subsequent dispersals to Hawai'i, and other archipelagos. A greater Fiji-Samoa region, corresponding with Takhtajan's Fijian Region, is implicated as a major Pacific region interface and possibly a center of origin for expansion of Cyrtandra throughout the Pacific. Among South Pacific taxa sampled, several supported clades in our evolutionary hypothesis are characterized by distinct morphological traits possibly warranting sectional rankings. Relationships among Hawaiian taxa are less resolved and the distributions of species within this clade do not consistently correspond to existing sectional rankings. More detailed, population-level research is needed to clarify these relationships. We argue that future sectional classifications should correspond with monophyletic lineages and that species-level relationships should be more closely studied within these lineages.

Island systems have long been useful models for understanding lineage diversification in a geographic context, especially pertaining to the importance of dispersal in the origin of new clades. Here we use a well-resolved phylogeny of the flowering plant genus Cyrtandra (Gesneriaceae) from the Pacific Islands to compare four methods of inferring ancestral geographic ranges in islands: two developed for character-state reconstruction that allow only single-island ranges and do not explicitly associate speciation with range evolution (Fitch parsimony [FP; parsimony-based] and stochastic mapping [SM; likelihood-based]) and two methods developed specifically for ancestral range reconstruction, in which widespread ranges (spanning islands) are integral to inferences about speciation scenarios (dispersal-vicariance analysis [DIVA; parsimony-based] and dispersal-extinction-cladogenesis [DEC; likelihood-based]). The methods yield conflicting results, which we interpret in light of their respective assumptions. FP exhibits the least power to unequivocally reconstruct ranges, likely due to a combination of having flat (uninformative) transition costs and not using branch length information. SM reconstructions generally agree with a prior hypothesis about dispersal-driven speciation across the Pacific, despite the conceptual mismatch between its character-based model and this mode of range evolution. In contrast with narrow extant ranges for species of Cyrtandra, DIVA reconstructs broad ancestral ranges at many nodes. DIVA results also conflict with geological information on island ages; we attribute these conflicts to the parsimony criterion not considering branch lengths or time, as well as vicariance being the sole means of divergence for widespread ancestors. DEC analyses incorporated geological information on island ages and allowed prior hypotheses about range size and dispersal rates to be evaluated in a likelihood framework and gave more nuanced inferences about range evolution and the geography of speciation than other methods tested. However, ancestral ranges at several nodes could not be conclusively resolved, due possibly to uncertainty in the phylogeny or the relative complexity of the underlying model. Of the methods tested, SM and DEC both converge on plausible hypotheses for area range histories in Cyrtandra, due in part to the consideration of branch lengths and/or timing of events. We suggest that DEC model-based methods for ancestral range inference could be improved by adopting a Bayesian SM approach, in which stochastic sampling of complete geographic histories could be integrated over alternative phylogenetic topologies. Likelihood-based estimates of ancestral ranges for Cyrtandra suggest a major dispersal route into the Pacific through the islands of Fiji and Samoa, motivating future biogeographic investigation of this poorly known region.

Nineteen miscellaneous species of Bornean Cyrtandra are dealt with. Cyrtandra atrichoides, C. congestiflora, C. crockerella, C. dulitiana, C. kanae, C. libauensis, C. plicata, C. vaginata and C. disparoides subsp. inconspicua are newly described. Descriptions and discussion are provided for C. erythrotricha and C. poulsenii, originally published with diagnoses only. Cyrtandra axillaris, C. longicarpa and C. microcarpa are also described, while C. borneensis, C. dajakorum, C. glomeruliflora, C. latens and C. prolata are reduced to synonymy.

Six new species of Cyrtandra from Borneo (C. apaensis, C. burleyana, C. filisecta, C. subsphaerocarpa, C. tibangensis, C. toreniiflora) are described. Cyrtandra albibracteata is re-described in order to correct gross errors in the original paper. The published diagnosis validating C. paragibbsiae from Brunei is supplemented by a full description.

Two groups of species are dealt with: first, Cyrtandra schizostyla together with three new species, all notable for the extraordinary elaboration of their anthers; second, C. velutina and C. diplotricha together with four new species, notable for their remarkably slender, soft-walled fruits. All species are fully described and identification keys provided.

Cyrtandra comprises at least 600 species distributed throughout Malesia, where it is known for many local endemics and in Polynesia and Micronesia, where it is present on most island groups, and is among the most successfully dispersing genera of the Pacific. To ascertain the origin of the oceanic Pacific island species of Cyrtandra, we sequenced the internal transcribed spacers of nuclear ribosomal DNA of samples from throughout its geographical range. Because all oceanic Pacific island species form a well-supported clade, these species apparently result from a single initial colonization into the Pacific, possibly by a species from the eastern rim of SE Asia via a NW-to-SE stepping stone migration. Hawaiian species form a monophyletic group, probably as a result of a single colonization. The Pacific island clade of Cyrtandra dispersed across huge distances, in contrast to the apparent localization of the SE Asian clades. Although highly vagile, the Pacific clade is restricted to oceanic islands. Individual species are often endemic to a single island, characteristic of the "supertramp" life form sensu Diamond (1974, Science 184: 803-806). The evolution of fleshy fruit within Cyrtandra provided an adaptation for colonization throughout the oceanic Pacific via bird dispersal from a single common ancestor.

Section Dissimiles C.B. Clarke is reviewed. Eleven species are known, one of which is newly described here (C. fulvisericea), and one species is reduced to synonymy (C. producta = C. trisepala). The section is centred on Borneo, with two outlying species in Sumatra and one in Peninsular Malaysia. Section Dissimiles is defined by a unique combination of characters, the most distinctive of these are: a zygomorphic calyx (the upper three lobes are fused to form one but the tips remain free, and the lower two lobes are completely separated to the base); anisophyllous leaf pairs, and a small (generally less than 2 cm long), predominantly white, somewhat translucent corolla.

Section Dissimiles C.B. Clarke is reviewed. Eleven species are known, one of which is newly described here (C. fulvisericea), and one species is reduced to synonymy (C. producta = C. trisepala). The section is centred on Borneo, with two outlying species in Sumatra and one in Peninsular Malaysia. Section Dissimiles is defined by a unique combination of characters, the most distinctive of these are: a zygomorphic calyx (the upper three lobes are fused to form one but the tips remain free, and the lower two lobes are completely separated to the base); anisophyllous leaf pairs, and a small (generally less than 2 cm long), predominantly white, somewhat translucent corolla.

Seven new species of Cyrtandra (Gesneriaceae) from Sulawesi are described and illustrated: C. gorontaloensis, C. fasciata, C. luteiflora, C. serratifolia, C. purpurea, C. tenuicarpa and C. roseiflora.

At a regional scale, the high species numbers (gamma diversity) of tropical forests have been explained by either a gradual accumulation of species through time (museum hypothesis) or, by contrast, rapid recent speciation in large genera. However, the origins of local rain forest diversity (alpha diversity) have been given little attention. Cyrtandra (Gesneriaceae), an understorey genus in the highly species-rich Indo-Malayan rain forest, has considerable capacity for producing local endemics, making it particularly suitable for studying diversity on a local scale. We sampled Cyrtandra species from one community on Mount Kerinci, Sumatra, and phylogenetic analyses of ITS sequences suggest that this community is an assembly of three distinct phyletic lineages: (1) a group of herbaceous or subshrub plants of Bornean affinity, (2) one member of a group of widespread shrubs forming Cyrtandra section Dissimiles and (3) a second group of shrubs. The evolutionary origin of this community is therefore not a result of rapid and recent speciation: it is assembled from species resulting from a gradual accumulation of diversity through time (museum hypothesis), although one lineage shows evidence of more recent, continuing speciation than the other two. The community includes two distantly related, apparently endemic species, but there is no evidence for a local adaptive radiation. The protection of representative species from each lineage would allow the conservation of genetic diversity. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81, 49–62.

Collections of Cyrtandra from the island of Palawan in the Philippines are reviewed. Twelve species are present, of which only five species are already described. A further three species and one variety are described here: C. cleopatrae H.J. Atkins & Cronk sp. nov., C. pulgarensis [Elmer ex] H.J. Atkins & Cronk sp. nov., C. hirtigera H.J. Atkins & Cronk sp. nov., and C. hirtigera var. chlorina H.J. Atkins & Cronk var. nov. Further material of the remaining four taxa (spp. B, C, D, E) will be needed before these can be formally named, although descriptions are provided here. All species are endemic except C. elatostemoides Elmer and C. hypochrysoides Kränzl., which also occur in Borneo and Luzon respectively. The pattern of narrow endemism shown by the cyrtandras of Palawan is commented on briefly.

Pollen of 34 accessions (23 species, one putative hybrid) of South Pacific and Hawaiian Cyrtandra was investigated by SEM. Most pollen grains are isopolar, 3-colpate, spheroidal in equatorial view, and circular in polar view. Only C. kamooloaensis (Hawaii) exhibits suboblate pollen grains. Average equatorial diameters range from 9 to 16mum. In general, colpi are long and tapering. Colpus membranes show granular elements which decrease in size from the equatorial to the polar regions. Most taxa exhibit a microreticulate mesocolpial exine pattern, in some cases with a tendency to reticulate or fossulate pattern. Lumina size varies within the mesocolpial region of single grains, average lumina diameter is either similar throughout grains or decreases towards the colpial borders or the apocolpial region. C. kamooloaensis exhibits the smallest lumina reported for Hawaiian Cyrtandra so far. In some species, distinct free-standing columellae are present. Except for a fine granular ornamentation of shallow groove...

A biogeographic and phylogenetic study of Cyrtandra (Gesneriaceae) in the Sundaland region (Borneo and Peninsular Malaysia) and the Philippines using nuclear ribosomal (ITS) DNA sequence data reveals a major division between the Cyrtandra floras of Sundaland and the Philippines. Palawan, the most westerly of the Philippine islands, emerges as an area of mixing between these two. The Bornean element in the Cyrtandra flora of Palawan (two species in our sample) appears to result from recent (i.e. Pleistocene) dispersal from Borneo. The remaining seven species sampled from Palawan are most closely related to those from elsewhere in the Philippines. However, the Palawan clade is sister to the other Philippine taxa, suggesting an ancient (possibly Pliocene) vicariance event. Huxley's line—a zoogeographic boundary placing Palawan and Borneo together—receives some support from this study as there is evidence of recent dispersal of Bornean flora into Palawan. However, in terms of more ancient biogeographic patterning of the region, Palawan has stronger links with the other Philippine islands.

The PCR-amplified atpB/rbcL spacer region of ten Cyrtandra species and representatives of the genera Didymocarpus, Didissandra, and Monophyllaea was sequenced and used for phylogenetic analyses. In Cyrtandra 83 variable positions were found over a length of 890bp. The data suggest that the Samoan species included in our study originated by at least two independent colonization events, while for the Hawaiian species the situation is not clear yet. With respect to generic relationships, Monophyllaea shows considerable divergence (9.9%) from the remaining taxa and thus is used as outgroup. Didymocarpus is most closely related to Cyrtandra, their genetic divergence (2.2%) being only slightly higher than the average distance within Cyrtandra (1.9%). The promoter regions for the atpB and rbcL genes, which have been described from Rubiaceae, were identified and found to be located at homologous positions. Of the two atpB promoters found in Rubiaceae only the distal one appears functional, the proximal promoter has been eroded by mutations and is disrupted in Didissandra by a 14bp insertion.

Interspecific hybridization among Hawaiian species ofCyrtandra (Gesneriaceae) was investigated using randomly amplified polymorphic DNA (RAPD) markers. Thirty-three different primers were used to investigate interspecific hybridization for 17 different putative hybrids based on morphological intermediacy and sympatry with putative parental species. RAPD data provided evidence for the hybrid origin of all putative hybrid taxa examined in this analysis. However, the patterns in the hybrid taxa were not found to be completely additive of the patterns found in the parental species. Markers missing in the hybrid taxa can be attributed to polymorphism in the populations of the parental species and the dominant nature of inheritance for RAPD markers. Unique markers found within hybrid taxa require further explanation but do not necessarily indicate that the taxa are not of hybrid origin. The implications suggest that these interspecific hybridization events had, and continue to have, an effect on the adaptive radiation and conservation biology ofCyrtandra.

There is not a ‘fragment’ in all nature, for every relative fragment of one thing is a full harmonious unit in itself. John Muir, 1867 ( A Thousand-Mile Walk to the Gulf , 1916) When we try to pick out anything by itself, we find it hitched to everything else in the Universe. John Muir, 1869 ( My First Summer in the Sierra , 1911) Plant diversity around the world A complete treatment of vegetation around the world would be impossible in a whole book let alone a single chapter. Instead, we concentrate on plants that inhabit different environmental extremes. We bring into focus the biotic relations of plants and, in addition, we consider some aspects of plant evolution in relation to the Earth's history and climate change, by looking at plants of islands. The greatest omission this chapter is an account of the forests of the world. Every botanist should visit a tropical rainforest at least once. No vegetation formation on Earth can compare to tropical rainforest in its staggering wealth of life forms, its diversity of species. It is the ‘Ultima Thule’ of the botanical world, after which everything else falls into perspective. … I measured my insignificance and climate change, by looking at plants of islands. against the quiet majesty of the trees. All botanists should be humble. From trampling weeds and cutting lawns they should go where they are lost in the immense structure of the forest. […]