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Nepenthes fractiflexa (Nepenthaceae), a new Bornean pitcher plant exhibiting concaulescent metatopy and a high degree of axillary bud activation
Abstract
Nepenthes fractiflexa is described as a new species from Sarawak and Kalimantan. The species is characterised by a number of unusual growth habits: plants readily produce aerial offshoots from their leaf axils, giving rise to a highly branched architecture of a complexity rarely seen in the genus; even those axillary buds not destined to develop further are activated and often elaborated into bract-like prophylls up to 5 cm long; and the inflorescence emerges close to the middle of the internode rather than from the leaf axil, as is typical of all other known species, leading us to make the first proposal for concaulescence in Nepenthes. These traits—coupled with the plant’s distinctive, long-decurrent petiolar wings and fractiflex climbing stems—distinguish it from its putative closest relative, N. mollis. The known distributions of N. fractiflexa and N. mollis are reviewed in light of recent discoveries. The apparent geographical disjunction between the two species is discussed in the context of edaphic and other ecological factors. Owing to its large range and remote, predominantly ridgetop habitat—but small number of recorded localities and individuals—the conservation status of N. fractiflexa is assessed as Near Threatened under the IUCN 3.1 criteria.
... Illegal logging has been identified as a threat to Nepenthes species from Cambodia (Mey et al., 2010), Indonesian Borneo (Giesen, 2000;Meijaard et al., 2005), and throughout the Philippines (Amoroso et al. 2009Gronemeyer et al., 2014;Cheek et al., 2015;Lagunday et al., 2017, Lagunday andAmoroso, 2019). The discovery of undescribed Nepenthes species and new populations of known species along the margins of logging roads and in areas made accessible by logging activitiesdfor N. dactylifera, N. ephippiata, N. fractiflexa, N. glandulifera, N. pitopangii, N. platychila and N. vogelii, among others (Lee 2004;Lee et al., 2009;Robinson et al., 2019a;Golos et al., 2020; A.T. Cross and A. Robinson pers. obs.)dhas led to an indisputable observation bias of Nepenthes in anthropogenic ruderal habitats, and occasional but not necessarily correct assertions that these plants might actually benefit from such disturbance (discussed in Golos et al., 2020). ...
... The discovery of undescribed Nepenthes species and new populations of known species along the margins of logging roads and in areas made accessible by logging activitiesdfor N. dactylifera, N. ephippiata, N. fractiflexa, N. glandulifera, N. pitopangii, N. platychila and N. vogelii, among others (Lee 2004;Lee et al., 2009;Robinson et al., 2019a;Golos et al., 2020; A.T. Cross and A. Robinson pers. obs.)dhas led to an indisputable observation bias of Nepenthes in anthropogenic ruderal habitats, and occasional but not necessarily correct assertions that these plants might actually benefit from such disturbance (discussed in Golos et al., 2020). However, these species are naturally often overwhelmingly epiphytic, and while many Nepenthes certainly proliferate in disturbed sites (typically in the company of resam fern Dicranopteris linearis, Gleicheniaceae), repeated visits to ruderal habitats that do not undergo periodic maintenance (slashing, mowing etc.) show that Nepenthes are often ephemeral and the succession of secondary growth eventually renders conditions unfavourable for their survival (A. ...
Carnivorous plants (CPs)—those possessing specific strategies to attract, capture and kill animal prey and obtain nutrition through the absorption of their biomass—are harbingers of anthropogenic degradation and destruction of ecosystems. CPs exhibit highly specialised and often very sensitive ecologies, being generally restricted to nutrient-impoverished habitats where carnivory offers a competitive advantage. As such, they are often the first species to disappear following habitat degradation, land use change, and alteration to natural ecological processes, and are at significant risk from processes such as eutrophication and weed invasion, and even poorly-understood impacts such as airborne nitrogen inputs. Many of the world’s 860 species of CPs are found in wetland habitats, which represent some of the most cleared and heavily degraded ecosystems on Earth. Global diversity hotspots for CPs are likewise located in some of the most heavily cleared and disturbed areas of the planet—southwestern Western Australia, Southeast Asia, Mediterranean Europe, central eastern Brazil, and the southeastern United States—placing their conservation at odds with human developmental interests. Many carnivorous plant species exhibit extreme range-restriction and are wholly localised to specific geological formations, microhabitats or elevations, with nowhere to move to in the face of environmental change such as a warming, drying climate. We provide the first systematic examination of the conservation status and threats to all CPs globally, compiling full or partial assessments of conservation status category for 860 species from 18 genera, and provide ten recommendations towards better conservation and management of this iconic group. A total of 69 species were assessed as Critically Endangered (8% of all species), 47 as Endangered (6%), 104 as Vulnerable (12%), and 23 as Near Threatened (3%). Slightly over 60% of CPs (521 species) were assessed as Least Concern. At least 89 species are known from only a single location based on current knowledge. Data on threatening processes were available for 790 species, with the most common threatening processes including Agriculture and Aquaculture (impacting 170 species), Natural Systems Modifications (168 species), Climate Change and Severe Weather (158 species), Energy Production and Mining (127 species), Human Intrusions and Disturbance (126 species), and Biological Resource Use (98 species). Almost a quarter of all species were impacted upon by three or more threatening processes. The most significant threats placing species at imminent risk of extinction include the continuing clearing of natural habitat for urban and agricultural development and the illegal collection of individuals from the wild for horticultural trade. The complex and specialised ecological requirements of CPs, together with the multifaceted threats they face, make conservation difficult and repatriation even to restored areas challenging. As the number of vulnerable, endangered and extinct carnivorous plant species continues to grow, despite significant conservation efforts in many regions and greater awareness of their ecological requirements, it is clear that a paradigm shift is required in our approach to the preservation of this unique group of plants in order to achieve long-term conservation successes.
... This genus of at least 160 accepted species is primarily distributed across the Malesian and Papuasian biogeographic regions, with centres of diversity in Sumatra, Borneo, and the Philippines (Danser 1928;Jebb & Cheek 1997;Clarke et al. 2018). In the last 20 years, the genus has received significant attention from botanists and enthusiasts alike, resulting in a series of significant discoveries that have led to the description of many taxa at species rank, as well as emended circumscriptions of existing species concepts (Akhriadi et al. 2004;Clarke et al. 2006;Mey et al. 2011;Cheek & Jebb 2013;Robinson et al. 2019;Bianchi et al. 2020;Golos et al. 2020). ...
A species of Nepenthes endemic to Peninsular Malaysia and distinctive for its extremely developed lid hairs is described and illustrated. Nepenthes berbulu is a highland species known only from a handful of peaks in the Titiwangsa Range. It is classified here within the recently defined N. macfarlanei group, for which a dichotomous key is provided. The description of this taxon brings the total number of Nepenthes species recognised from Peninsular Malaysia to eighteen.
... Tropical pitcher plants in the genus Nepenthes L. comprise over 160 described species occurring throughout Southeast Asia and a few outlying Paleotropical regions including Madagascar, the Indian subcontinent, New Caledonia, and Australia (McPherson, Robinson, & Fleischmann, 2009;Golos et al., 2020;Tamizi et al., 2020). These plants are characterized by their pitchers, which are modified leaves that evolved for the attraction, capture, and digestion of arthropod prey to fulfill the plants' nutritional requirements in the nitrogen-poor soils where they grow (Juniper, Robins, & Joel, 1989). ...
Tropical pitcher plants (Nepenthes) are carnivorous plants that trap and digest prey using highly modified fluid-filled leaves known as pitchers. Prey are digested by plant-secreted enzymes and pitcher symbionts. Pitchers exert control over abiotic properties of the digestive fluid such as pH levels that can influence its symbionts. Here we examine natural variation in pH and dissolved mineral concentrations in three sympatric Nepenthes species, assessing correlations between fluid properties and pitcher traits. We use addition experiments to investigate differences in protein digestion/absorption rates between species. Fluid pH and dissolved mineral levels both showed distinct patterns corresponding to pitcher developmental stages in N. gracilis and N. rafflesiana, whereas N. ampullaria differs from its congeners in exhibiting far less variation in fluid pH, as well as less clear evidence of protein depletion. This study further elucidates the properties of pitchers as habitats, revealing ways in which the host plant regulates that habitat.
... The Nepenthes flora of Kalimantan is poorly known compared to that of Malaysian and Bruneian Borneo, with relatively few modern records. Thus, the new discoveries that have emerged recently after expeditions to certain remote areas of Kalimantan (Robinson et al. 2019;Golos et al. 2020) are not surprising. ...
Nepenthes pudica, a new species from North Kalimantan, Indonesia, is described and illustrated. The species belongs to the N. hirsuta group (sensu Cheek and Jebb 1999) but exhibits some characters that are unique within the group or even within the genus. Above all, it produces underground, achlorophyllous shoots with well-developed, ventricose lower pitchers that form in soil cavities or directly in the soil. No lower pitchers are formed above ground. The main part of its prey are ants, besides other litter-and soil-inhabiting species of invertebrates. A number of infaunal species were found in both aerial and underground pitchers, mainly Diptera and nematodes. Nepenthes pudica is known only from a few neighbouring localities in the Mentarang Hulu district of North Kalimantan, where it grows on ridgetops at an elevation of 1100–1300 m. Its discovery underlines the natural richness of Borneo's rainforest and the necessity to preserve this important ecosystem with its enormous and still undiscovered biodiversity.
... Indonesia is considered as one of the most significant diversity hotspots for Nepenthes species, including 73 taxa of which 36 occur in Sumatra, 3 species in Java, 21 in Indonesian Borneo, 13 in Sulawesi, 6 in Maluku islands, and 13 in Indonesian New Guinea (Cheek & Jebb 2001, Akhriadi et al. 2009, Robinson et al. 2011, Cheek 2015, Cheek & Jebb 2016aCheek et al. 2018, Robinson et al. 2019, Golos et al. 2020). At the current state of knowledge, no record of Nepenthes there is in Lesser Sunda Islands (Cheek & Jebb 2001). ...
A new species of Nepenthes—Nepenthes putaiguneung—from the highlands of Sumatra (Indonesia) is described and
illustrated. This new species is morphologically similar with N. Singalana from which differs in having narrowly spathulate
leaves, more gracile upper pitchers with a finely ribbed peristome, lacking teeth on its inner edge, lid with a basal crest and
a short tooth-like triangular appendage near the apex on the underside (not lacking such appendages), the lid glands of lower
pitcher confined to the midline (not scattered evenly over the whole surface), and pedicel of lower male flowers usually with
minutely short bracteole. N. putaiguneung is also similar to N. mikei but differs in having narrowly spathulate leaves with
obtuse apex, lower pitcher with slender infundibulate shape below and cylindrical above, peristome of lower pitcher with
short and denser distinct ribs, lid of lower and upper pitcher with a prominent glandular crest at the base of midrib, several
distinct glands on the lid of lower pitcher that clustered along the midrib; and pedicel of lower male flowers that usually with
minutely short bracteole.
Compendium of new species of Carnivorous Plants published in 2020, first part: Genlisea, Drosera, Nepenthes.
VICTORIANO, M. 2021. A new species of Nepenthes (Nepenthaceae) and its natural hybrids from Aceh, Sumatra, Indonesia. Reinwardtia 20(1): 17–26. — A new species of Nepenthes (Nepenthaceae) from Aceh Province, Indonesia, Nepenthes longiptera Victoriano is herein described and illustrated. The species is unique among all other Nepenthes in Sumatra by the presence of wings on its upper pitchers. Comprehensive description, photographs, geographical distribution and preliminary IUCN conservation assessment are provided for the new species. Hybrids of this new taxon with other species are also reported in this paper.
Nepenthes diabolica, a new species of tropical pitcher plant from Central Sulawesi, is described and illustrated. It is a high-montane, terrestrial to epiphytic taxon, distinguished by its elaborately flanged peristome and rufous, woolly pitcher indumentum. N. diabolica is a member of section Tentaculatae, showing morphological affinities with N. hamata. Nevertheless, the two species can be distinguished by morphological characters as well as by ecology and geography. A diagnostic key to the species of section Tentaculatae is provided. Owing to its highly restricted range, small number of recorded individuals, and desirability among plant collectors, the conservation status of N. diabolica is assessed as Critically Endangered under the IUCN criteria. Finally, N. hamata, until now based on lost types and mixed paratypes, is lectotypified using recently uncovered original material.
Nepenthes maximoides sp. nov. (Sect. Alatae) is described and assessed as Critically Endangered (Possibly Extinct) from Luzon, Philippines and appears unrecorded in 110 years. The spectacular, large, narrowly funnel-shaped upper pitchers, lids with recurved basal and filiform apical appendages, unlike any other species in the Philippines, closely resemble those of N. maxima (Sect. Regiae) of Sulawesi-New Guinea, likely due to convergent evolution. Following recent phylogenomic analysis, sect. Alatae is divided into two, Sect. Alatae sensu stricto of Luzon to Sibuyan (including N. maximoides), and Sect. Micramphorae, expanded and recircumscribed to encompass those species of the southern Visayas, and Mindanao. A key is provided to the six species now recognised in the newly narrowly recircumscribed Sect. Alatae. The number of Nepenthes species recorded from Luzon has increased from two in 2001, to eight in 2020, all but one of which are endemic to that island, and four of which appear to be point endemics.
Nepenthes maximoides sp. nov. (Sect. Alatae) is described and assessed as Critically Endangered (Possibly Extinct) from Luzon, Philippines and appears unrecorded in 110 years. The spectacular, large, narrowly funnel-shaped upper pitchers, lids with recurved basal, and filiform apical appendages, unlike any other species in the Philippines, closely resemble those of N. maxima (Sect. Regiae) of Sulawesi-New Guinea, likely due to convergent evolution. Following recent phylogenomic analysis, sect. Alatae is divided into two, Sect. Alatae sensu stricto of Luzon to Sibuyan (including N. maximoides), and Sect. Micramphorae, expanded and recircumscribed to encompass those species of the southern Visayas, and Mindanao. A key is provided to the six species now recognised in the narrowly recircumscribed Sect. Alatae. The number of Nepenthes species recorded from Luzon has increased from two in 2001, to eight in 2020, all but one of which are endemic to that island, and four of which appear to be point endemics.
A report from a trip to Southern Thailand, with Nepenthes rosea described as a new species of the N. thorelii aggregate. Discovery, habitat and morphology of a unique taxon belonging to one of the most confusing groups within the genus.
A new species of Nepenthes L., N. appendiculata, from Sarawak, is described and illustrated. This new taxon is known only from the Hose Mountains and can easily be distinguished from all other Bornean members of the genus by its pitcher morphology, and particularly the unique glandular appendage present on the underside of the lids of its upper pitchers.
Two species previously known only from Kalimantan, N. campanulata Kurata and Nepenthes ephippiata Dans. are newly recorded for Sarawak. Nepenthes glandulfera Chi.C. Lee, a new species from the Hose Mountains in central Sarawak, is described and illustrated. Additional information and new locality records in Sarawak for N. hurrelliana Cheek & A. Lamb and N. vogelii Schuit. & de Vogel are provided.
Nepenthes chaniana, a new species from montane habitats in Sabah and Sarawak, is described. This species was previously considered as belonging to N.pilosa Dans., but is shown to have significant morphological differences from that species. The relationships between N. chaniana, N. pilosa and another closely related species, N. glandulifera, are discussed.
Nepenthes platychila Chi.C. Lee, a new species from the Hose Mountains in central Sarawak, Borneo, is described and illustrated.
An expedition to the highest peak of the Kemul Massif located a number of different Nepenthes taxa, including the type populations of N. fusca and N. mollis, neither of which have been observed in situ since they were first collected in 1925. Studies of the type form of N. fusca show that two further species should be considered, one largely restricted to Sabah’s Crocker range—here reinstated as N. zakriana—the second a southerly distributed species from Brunei, Sarawak and Kalimantan, named here as N. dactylifera. Analysis of the type population of N. mollis indicates that N. hurrelliana is a heterotypic synonym of N. mollis, and reveals an additional undescribed Nepenthes species from Sarawak and Kalimantan with a widespread but highly localised range.
Nepenthes vogelii Schuit. & de Vogel is described as a new species from Sarawak. It is compared with N. fusca Danser, from which it differs e.g. in the much smaller pitchers, of which the lid lacks appendages.
Nepenthes hurrelliana is described and illustrated as a new species from Sabah, Sarawak and Brunei in Borneo. Its relationships are discussed, a key to related species is presented and a conservation assessment is made.
The carnivorous nature of Nepenthes its ecology architecture morphology and the taxonomy of the New Guinea species are discussed. The unique architecture of N. ampullaria is described for the first time. A taxonomic summary with descriptions and drawings of all eleven species known to occur in New Guinea is presented (N. ampullaria, N. insignis, N. klossii, N. maxima, N. mirabilis, N. neoguineensis, N. paniculata, N. papuana, N. treubiana, N. lamii (as N. vieillardii) and one undescribed species (N. danseri). For each species the synonymy in New Guinea only is given and one undescribed species is outlined. All collections from New Guinea seen by the author are listed and assigned to species, and distribution maps are given. A collector index is also provided.
A skeletal world revision of the genus is presented to accompany a family account for Flora Malesiana. 82 species are recognised, of which 74 occur in the Malesiana region. Six species are described as new, one species is raised from infraspecific status, and five species are restored from synonymy. Many names are typified for the first time. Three widespread, or locally abundant hybrids are also included. Full descriptions are given for new (6) or recircumscribed (7) species, and emended descriptions of species are given where necessary (9). Critical notes are given for all the species. Little known and excluded species are discussed. An index to all published species names and an index of exsiccatae is given.
Two new paniculate species of Nepenthes, N. halmahera and N. weda, both allied to N. danseri Jebb & Cheek, are described respectively from lowland and lower montane forest on ultramafic substrate. Nepenthes weda appears to be unique in the genus due to the adaxial tepal surfaces which in the distal portion are hairy and lack nectar glands. This species is also unique among paniculate members of the genus in its forward-facing, concave subapical lid appendage. Both species are currently only known from the Weda Bay Nickel Project concession area in Halmahera, Indonesia, and are assessed as Critically Endangered using the 2012 IUCN standard. Two character state s are formally described and named for the first time in Nepenthes: nanophyll rosettes and multiseriate fringed pitcher wings. Stage-related heteromorphy in lid appendages is documented for the first time in the genus. Keys to the species of Nepenthes of Halmahera, and to the paniculate species of SE Asia are presented.
Background
The Palaeotropical pitcher plant genus Nepenthes (Nepenthaceae) is characterized by specialized nutrient sequestration strategies, narrow endemism, and a patchy distribution in which vicariance is believed to have played a fundamental role.
Scope
Using recent studies of the effects of climate, soil type and vicariance, we review patterns of diversity and endemism in Nepenthes. First we consider how climate influences the geographical range of the genus and diversity of prey trapping mechanisms. Second, we examine edaphic influences, specifically the relationship between limestone and ultramafic soils and the obligate edaphic endemic Nepenthes that inhabit them. Third, we examine the role of vicariance, with regards to the patchy distribution of suitable habitats throughout Southeast Asia, and the passive dispersal mechanism of Nepenthes seeds.
Conclusions
Climate is the principal determinant of variation in pitcher functional traits and in perhumid environments, may drive the evolution of alternative nutrient sequestration strategies. Although little is known about the ecophysiological relationships between soil type and obligate edaphic Nepenthes, ultramafic and limestone substrates may strongly influence vegetation physiognomy, creating a diversity of environmental niches that are exploited by specialized Nepenthes species. Finally, the complex geology and geography of the Malay Archipelago drives diversification through vicariance.
In this paper, the authors deal with the morphological features of 10 abnormal branches of honeysuckle. The studied forms are of two different types. The first is illustrated by five specimens collected on a single hybrid bush called Lonicera nigra × orientalis. All of these show unusual union between restricted portions of different leaves. The arrangement of the fused parts permits one to infer that gamophylly, in this example, results indubitably from a real ontogenetic (postgenital) union. Concaulescence is the second kind of anomaly, shown in the samples from various cultivars belonging to the Lonicera tatarica complex. The occasional occurrence of accessory serial buds between the concrescent axes suggests that ontogenetic fusion could give rise also to such structures. In some particular instances, however, it is possible that unusual intercalary growth is combined with the first process.
Background and Aims
The architecture of a plant depends on the nature and relative arrangement of each of its parts; it is, at any given time, the expression of an equilibrium between endogenous growth processes and exogenous constraints exerted by the environment. The aim of architectural analysis is, by means of observation and sometimes experimentation, to identify and understand these endogenous processes and to separate them from the plasticity of their expression resulting from external influences.
Scope
Using the identification of several morphological criteria and considering the plant as a whole, from germination to death, architectural analysis is essentially a detailed, multilevel, comprehensive and dynamic approach to plant development. Despite their recent origin, architectural concepts and analysis methods provide a powerful tool for studying plant form and ontogeny. Completed by precise morphological observations and appropriated quantitative methods of analysis, recent researches in this field have greatly increased our understanding of plant structure and development and have led to the establishment of a real conceptual and methodological framework for plant form and structure analysis and representation. This paper is a summarized update of current knowledge on plant architecture and morphology; its implication and possible role in various aspects of modern plant biology is also discussed.
The species Nepenthes mikei and Nepenthes xiphioides (Nepenthaceae) from Gunung Pangulubau, North Sumatra, Indonesia are described and illustrated
A previously unrecognized species of Nepenthes (Nepenthaceae) from Puncak Angasan, Aceh Province, Sumatra, Indonesia is described and illustrated. The original description of N. mike is reinstated.
Nepenthaceae is one of the largest carnivorous plant families and features ecological and morphological adaptations indicating an impressive adaptive radiation. However, investigation of evolutionary and taxonomic questions is hindered by poor phylogenetic understanding, with previous molecular studies based on limited loci and taxa. We use high-throughput sequencing with a target-capture methodology based on a 353-loci, probe set to recover sequences for 197 samples, representing 151 described or putative Nepenthes species. Phylogenetic analyses were performed using supermatrix and maximum quartet species tree approaches. Our analyses confirm five Western outlier taxa, followed by N. danseri, as successively sister to the remainder of the group. We also find mostly consistent recovery of two major Southeast Asian clades. The first contains common or widespread lowland species plus a Wallacean-New Guinean clade. Within the second clade, sects. Insignes and Tentaculatae are well supported, while geographically defined clades representing Sumatra, Indochina, Peninsular Malaysia, Palawan, Mindanao and Borneo are also consistently recovered. However, we find considerable conflicting signal at the site and locus level, and often unstable backbone relationships. A handful of Bornean taxa are inconsistently placed and require further investigation. We make further suggestions for a modified infra-generic classification of genus Nepenthes.
Many new questions on the regulation of shoot branching have been raised in recent years, prompting a review and reassessment of the role of each signal involved. Sugars and their signaling networks have been attributed a major role in the early events of axillary bud outgrowth, whereas cytokinin appears to play a critical role in the modulation of this process in response to the environment. Perception of the recently discovered hormone strigolactone is now quite well understood, while the downstream targets remain largely unknown. Recent literature has highlighted that auxin export from a bud is important for its subsequent growth.
Nepenthes is a genus of 130-160 species, almost half of which were described after 2001. The recent, rapid increase in species descriptions has been driven by application of a less rigorous species concept by botanists, taxonomic inflation, and discoveries of new taxa during explorations of remote parts of Southeast Asia. Many recently published species descriptions of Nepenthes are based entirely upon qualitative morphological information and are not supported by adequate research. Accordingly, the status of many Nepenthes taxa is contested. Evolution within the genus is not well understood, because nuclear and maternally inherited plastid genomes cannot resolve relationships between many species, particularly those that evolved recently through introgression or reticulate evolution. Improvement in our understanding of the systematics and evolution of Nepenthes requires the adoption of ‘best practice’ collection and preservation methods, and the application of quantitative analytical methods for morphological, genetic, and ecological information.
This chapter focuses on the geology of Offshore Brunei and Sabah. The Baram Delta of this region continues north-eastwards from the West Baram Line and extends outwards from the coast as far as the prominent linear deep, known as the NW Borneo Trough. Across the Morris Fault, the offshore area is subdivided into the Inboard Belt, the Outboard Belt, and the Thrust Sheet province. The Inboard Belt is subdivided into a southern and northern section by the Kinabalu Culmination. The Southern Inboard Belt is characterized by NNE-SSW trending anticlines with steep flanks and strongly faulted crests (Sabah Ridges), spaced at 5–20 km apart. The Northern Inboard Belt lies north of the Kinabalu Culmination and is characterized by an intersecting network of N-, E- and SE-trending compressional ridges and synclines. They terminate against the NE-trending Emerald Fault Zone to the west and the Crocker fold-belt to the east. The Outboard Belt occurs west of the northern Inboard Belt , is structurally complex, and contains both extensional and compressional structural features. The Thrust Sheet province is bounded to the NW and SW by steep thrust faults, and to the SE by normal faults that separate it from the Outboard Belt. The Thrust Sheet is interpreted as an allochthonous overthrust mass that borders the NW Borneo Trough;it could be a nappe of Rajang Group rocks that resulted from gravity sliding away from the uplifting Western Cordillera.
Résumé
Étude macroscopique de formes présentant des axes apparemment soudés chea plusieurs espèces de Dipsacus et Scabrosa. Il semble exister une relation entre la nature de l'aberration et le genre considéré. Chez Dipsacus il s'agit en général d'une vraie fasciaion; chez Scabiosa il y a, au contraire, généralement concaulescence. La structure de formes fasciées de Dipsacus s'explique bien par une augmentation du nombre des héices foliaites. Il n'en est pas de même dans les axes concaulescents de Scabiosa.
A dichotomous key and morphological descriptions and photographs of three species of pitcher plants recorded from Sabah were given. Nepenthes curtisii ssp. zakriana Adam and Wilcock is elevated to species level that is Nepenthes zakriana (Adam and Wilcock) Adam and Hafiza. A new species, Nepenthes naquiyuddinii Adam and Hafiza is described.
The idea that apically derived auxin inhibits shoot branching by inhibiting the activity of axillary buds was first proposed 70 years ago, but it soon became clear that its mechanism of action was complex and indirect. Recent advances in the study of axillary bud development and of auxin signal transduction are allowing a better understanding of the role of auxin in controlling shoot branching. These studies have identified a new role for auxin early in bud development as well as some of the second messengers involved in mediating the branch-inhibiting effects of auxin.
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