Patterns of cell elongation in the determination of the final shape in galls of Baccharopelma dracunculifoliae (Psyllidae) on Baccharis dracunculifolia DC (Asteraceae)

Cell redifferentiation, division, and elongation are recurrent processes, which occur during gall development, and are dependent on the cellulose microfibrils reorientation. We hypothesized that changes in the microfibrils orientation from non-galled tissues to galled ones occur and determine the final gall shape. This determination is caused by a new tissue zonation, its hyperplasia, and relative cell hypertrophy. The impact of the insect’s activity on these patterns of cell development was herein tested in Baccharopelma dracunculifoliae—Baccharis dracunculifolia system. In this system, the microfibrils are oriented perpendicularly to the longest cell axis in elongated cells and randomly in isodiametric ones, either in non-galled or in galled tissues. The isodiametric cells of the abaxial epidermis in non-galled tissues divided and elongated periclinally, forming the outer gall epidermis. The anticlinally elongated cells of the abaxial palisade layer and the isodiametric cells of the spongy parenchyma originated the gall outer cortex with hypertrophied and periclinally elongated cells. The anticlinally elongated cells of the adaxial palisade layer originated the inner cortex with hypertrophied and periclinally elongated cells in young and mature galls and isodiametric cells in senescent galls. The isodiametric cells of the adaxial epidermis elongated periclinally in the inner gall epidermis. The current investigation demonstrates the role of cellulose microfibril reorientation for gall development. Once many factors other than this reorientation act on gall development, it should be interesting to check the possible relationship of the new cell elongation patterns with the pectic composition of the cell walls.     

Morphogenesis of galls induced by Baccharopelma dracunculifoliae (Hemiptera: Psyllidae) on Baccharis dracunculifolia (Asteraceae) leaves.

The commonest insect gall on Baccharis dracunculifolia (Asteraceae) leaves is induced by Baccharopelma dracunculifoliae (Hemiptera, Psyllidae). The gall-inducing insect attacks young leaves in both the unfolded and the fully expanded stages. Four developmental phases were observed in this type of gall: 1) A folding phase, during which the leaf lamina folded upward alongside the midrib and the edges of the upper portion of the leaf approached each other, forming a longitudinal slit. A single chamber was formed on the adaxial surface of the leaf; 2) A swelling phase, in which the folded leaf tissues thickened and the edges of the leaf drew closer together, narrowing the slit. In this phase the gall matured, turning succulent, fusiform and pale green. The single nymphal chamber was lined with white wax and was able to house from one to several nymphs; 3) A dehiscence phase, characterized by the opening of the slit to release inducers; and 4) A senescence phase, when the gall turned dark and dry. The dermal system of the mature gall was composed of a single-layered epidermis. The mesophyll was swollen, and the swelling was due mainly to hyperplasia of the parenchyma. The vascular tissues along the midrib vein were conspicuous and the perivascular fibers resembled parenchymal cells. The hypertrophied secretory cavities contained low lipophylic content. This gall does not form nutritive tissue, but salivary sheaths left by the inducers were observed near the parenchyma, vascular bundles and secretory cavities. This study complements our current knowledge of gall biology and sheds further light on the plasticity of plant tissues stimulated by biotic factors.     

Floral-like destiny induced by a galling Cecidomyiidae on the axillary buds of Marcetia taxifolia (Melastomataceae)

The galls induced by Cecidomyiidae, Diptera, are very diverse, with conspicuous evidence of tissue manipulation by the galling herbivores. Bud galls, as those induced by an unidentified Cecidomyiidae species on Marcetia taxifolia, Melastomataceae, can be considered as one of the most complex type of prosoplasma galls. The gall-inducer manipulate the axillary meristem of the plant in a way that gall morphogenesis may present both vegetative and reproductive features of the host plant. Herein, we analyzed traces of determinate and indeterminate growth in the bud gall of M. taxifolia, looking for parallels between the features of the leaves and flowers, natural fates of the meristematic cells. The bud galls are induced by the cecidomyiid fly, and are formed by the connation of eight leaf primordia, a common process in ovary morphogenesis. The bud gall corresponds to a pistil-shaped gall morphotype, with anatomical features similar to those of an hypanthium and sepals. The gall mimics an ovary, which has protective barriers at the apex, and a nutritive tissue (with storage of lipids and proteins) or a placenta, respectively, at the basal portion. The redifferentiation of the promeristem into a nutritive tissue at the base of the gall confers a determinate destiny to the axillary bud. Comparatively, the gradients of cell expansion and of accumulation of primary metabolites also indicate that the gall and the ovary are convergent structures. Some constraints of the host plant cells, such as the absence of lignification, and the accumulation of polyphenols, lipids and terpenoids, are not altered and may confer chemical protection for plant tissues and the larva against oxidative stress.     

Cytological and histochemical gradients on two <Emphasis Type="BoldItalic">Copaifera langsdorffii</Emphasis> Desf. (Fabaceae)—Cecidomyiidae gall systems

Previous ultrastructural and histochemical analysis proposed patterns in the accumulation of substances in galls of Diptera: Cecidomyiidae in some plant species of the temperate region. Similar analyses were done to verify the conservativeness of these patterns in the Neotropical region, where a great number of species of Cecidomyiidae is responsible for a wide diversity of morphotypes. Two gall morphotypes induced by Cecidomyiidae in a unique host plant, Copaifera langsdorffii, were studied. The gradients of carbohydrates and the activity of invertases and acid phosphatases were similar, but the cytological gradients and distribution of proteins evidenced that the sites of the induction as well as the amount of neoformed tissues may be peculiar to each gall system. The production of lipids just in the secretory cavities either in the non-galled or galled tissues indicated a potentiality of the host plant which could not be manipulated by the galling insects. Further, the absence of nucleus in the nutritive tissue, an exclusive feature of the horn-shaped galls, indicates cell death attributed to the feeding habit of the galling herbivore.     

Morphometric analysis of young petiole galls on the narrow-leaf cottonwood, <Emphasis Type="Italic">Populus angustifolia</Emphasis>, by the sugarbeet root aphid, <Emphasis Type="Italic">Pemphigus betae</Emphasis>

An insect-induced gall is a highly specialized structure resulting from atypical development of plant tissue induced by a reaction to the presence and activity of an insect. The insect induces a differentiation of tissues with features and functions of an ectopic organ, providing nutrition and protection to the galling insect from natural enemies and environmental stresses. In this anatomical and cytological study, we characterized how the gall-inducing aphid Pemphigus betae reshapes the leaf morphology of the narrow-leaf cottonwood Populus angustifolia to form a leaf fold gall. Young galls displayed a bend on one side of the midvein toward the center of the leaf and back to create a fold on the abaxial side of the leaf. This fold was formed abaxially by periclinal and anticlinal divisions, effectively eliminating intercellular spaces from the spongy parenchyma. Galls at this stage exhibited both cell hypertrophy and tissue hyperplasia. Cells on the adaxial surface were more numerous and smaller than cells near the abaxial surface were, creating the large fold that surrounds the insect. Mesophyll cells exhibited some features typical of nutritive cells induced by other galling insects, including conspicuous nucleolus, reduced and fragmented vacuole, smaller and degraded chloroplasts, and dense cytoplasm compared to ungalled tissue. Even though aphids feed on the contents of phloem and do not directly consume the gall tissue, they induce changes in the plant vascular system, which lead to nutrient accumulation to support the growing aphid numbers in mature galls.     

EVOLUTION OF DIFFERENT GALL TYPES IN WILLOW-FEEDING SAWFLIES (HYMENOPTERA: TENTHREDINIDAE)

The sawflies that feed on the plant family Salicaceae can be divided into eight informal groups based on larval feeding habit or gall type: (1) species with free-living larvae; (2) leaf folders; (3) leaf blade gallers; (4) apical leaf gallers; (5) basal leaf gallers; (6) midrib and petiole gallers; (7) stem gallers; and (8) bud gallers. It has been proposed that the galling habit evolved from free-living larvae via leaf folders, and that the different gall types evolved gradually in the sequence mentioned above. Thus, the galling site would have “wandered” from the leaf margin toward the stem as a result of gradual changes in oviposition site preference. Allozyme data from eight informative loci were used to reconstruct the phylogeny of 18 representative sawfly species. The results suggest that indeed leaf folders seem to be a basal group; leaf blade gallers evolved independently of the other true gallers; apical and basal leaf gallers are not the ancestors of petiole and bud gallers, but they may share a common galling ancestor; bud gallers evolved from midrib/petiole gallers; and stem gallers are polyphyletic. The cause for the observed wandering of the galling site could be intraspecific competition due to a possible “nutrient shading effect” of galls situated closer to the host plant's main vascular system.     

Developmental anatomy and immunocytochemistry reveal the neo-ontogenesis of the leaf tissues of Psidium myrtoides (Myrtaceae) towards the globoid galls of Nothotrioza myrtoidis (Triozidae)

Key message

The temporal balance between hyperplasia and hypertrophy, and the new functions of different cell lineages led to cell transformations in a centrifugal gradient that determines the gall globoid shape.

Abstract

Plant galls develop by the redifferentiation of new cell types originated from those of the host plants, with new functional and structural designs related to the composition of cell walls and cell contents. Variations in cell wall composition have just started to be explored with the perspective of gall development, and are herein related to the histochemical gradients previously detected on Psidium myrtoides galls. Young and mature leaves of P. myrtoides and galls of Nothotrioza myrtoidis at different developmental stages were analysed using anatomical, cytometrical and immunocytochemical approaches. The gall parenchyma presents transformations in the size and shape of the cells in distinct tissue layers, and variations of pectin and protein domains in cell walls. The temporal balance between tissue hyperplasia and cell hypertrophy, and the new functions of different cell lineages led to cell transformations in a centrifugal gradient, which determines the globoid shape of the gall. The distribution of cell wall epitopes affected cell wall flexibility and rigidity, towards gall maturation. By senescence, it provided functional stability for the outer cortical parenchyma. The detection of the demethylesterified homogalacturonans (HGAs) denoted the activity of the pectin methylesterases (PMEs) during the senescent phase, and was a novel time-based detection linked to the increased rigidity of the cell walls, and to the gall opening. Current investigation firstly reports the influence of immunocytochemistry of plant cell walls over the development of leaf tissues, determining their neo-ontogenesis towards a new phenotype, i.e., the globoid gall morphotype.     

Leaf anatomy of the Pittosporaceae R. Br.

WILKINSON, H. P., 1992. Leaf anatomy of the Pittosporaceae R. Br. An anatomical study of the leaves of 58 species representing all nine genera has been made. The anatomical characters found to be of most use in distinguishing taxa are: in surface view-cuticular architecture as seen with the SEM, stomatal outline, occasionally stomatal density, presence/absence of hairs, hair type in adult leaves; in transverse section-petiole/midrib outline, midrib number of vascular bundles and number of secretory ducts, adaxial epidermis in 1/2 layers, dimensions of adaxial epidermal cells, thickness of outer wall of adaxial epidermis, occasionally chlorenchyma interrupted/not interrupted above the midrib vascular tissue in Pittosporum species; leaf margin.     

Synchronization of gallers with host plant phenology

In addition to various bottom-up effects, the synchronization of herbivores with their host plant phenology determines quality and quantity of food resources and affects the preference–performance linkage and abundance of herbivores. The synchronization has a more critical meaning for such short-lived galling insects as cecidomyiid adults and young aphid stem mothers than for other insects. This review, first, presents general information about gall midges and gall aphids, together with their life history patterns and some ecological attributes. Second, some important topics of galling insect–host plant relation are briefly reviewed. Then, synchronization patterns between gall midge emergence and host plant phenology are analyzed to discuss the adaptive strategies of gall midges and to show how the amount of available food resources is affected by the time lag in synchronization. The spatial distribution pattern and the preference–performance linkage of aphid stem mothers is also discussed in relation to synchronization. Received: October 2, 1998 / Accepted: July 3, 2000     

Leaf anatomy of Desmos and Dasymaschalon (Annonaceae) from China in relation to taxonomic significance

Leaf morphology in four species of Desmos and three species of Dasymaschalon was comparatively studied using scanning electron microscopy (SEM) together with epidermal maceration and paraffin methods. The results showed that there were some remarkable foliar anatomical differences between Desmos and Dasymaschalon. In leaves of Desmos, some of the adaxial epidermal cells were enlarged into globose cells each containing one large cluster crystal, while other epidermal cells were normal without any crystal, and in abaxial epidermis each cell contained one smaller cluster crystal. The leaf structure was typically bifacial, and the mesophyll cells were differentiated into palisade tissue and spongy tissue. Oil cells were distributed in the second layer of palisade and the whole spongy tissue, and the number of oil cells per mm leaf width ranged from 4 to 6. The vascular tissue in the midrib was separated into bundles by parenchyma cells. In leaves of Dasymaschalon, all the adaxial epidermal cells contained one cluster crystal, and the crystal size was similar to that of thecrystals in abaxial epidermal cells. The leaf structure was more or less isobilateral. Oil cells were distributed only in the spongy tissue between the two layers of the palisade, and the number of oil cells per mm leaf width ranged from 2 to 3. The vascular tissue in the midrib formed a continuous circle. It is clear that the anatomical differences between Desmos and Dasymaschalon are remarkable, supporting the treatment of Desmos and Dasymaschalon as two independent genera.     

Evidence for long-distance, chemical gall induction by an insect

Abstract We report that a chemical stimulus from a herbivore, a galling insect, changes plant morphology and physiology to benefit the herbivore. Previous studies could not determine whether insect galls are induced by mechanical or chemical stimuli because feeding and oviposition both occurred at the site of gall formation. We report that the mouthparts of a spruce‐galling insect, Adelges cooleyi, were inserted in stem phloem cells far from induced galls, that tissues between mouthparts and galls appeared normal, and that the ability to initiate galls was inversely correlated with distance from buds (potential gall sites). Thus the effects of chemical stimuli were unambiguously separated from any mechanical influence of probing stylets or ovipositors. Our results strongly suggest that galls were induced by a chemical stimulus transported to buds via vascular tissue and that its efficacy was dose‐dependent.     

中国假鹰爪属和皂帽花属植物叶的形态结构及其分类学意义

利用扫描电镜技术、叶片离析法和石蜡切片法研究了假鹰爪属Desmos 4种植物和皂帽花属Dasy-maschalon 3种植物叶片的形态结构。结果表明：假鹰爪属植物叶片近轴面表皮具大型球状含晶簇细胞和不含晶簇的表皮细胞两种类型,远轴面表皮细胞均具一较小的晶簇;叶肉组织明显分化为栅栏组织细胞和海绵组织细胞,油细胞分布于第2层的栅栏组织和海绵组织内,单位毫米叶宽油细胞数为4～6个;主脉维管组织被薄壁细胞分隔成束状。皂帽花属植物叶片近轴面表皮细胞形状相同,均具一晶簇,远轴面表皮细胞的晶簇和近轴面表皮细胞的晶簇相似;靠近上、下表皮的叶肉组织均分化为栅栏组织细胞,在两层栅栏组织细胞之间分化为一至几层海绵组织细胞,油细胞分布于海绵组织内,单位毫米叶宽油细胞数为2～3个;主脉维管组织形成连续的环状。由此可见两属叶的结构具有明显的差异,因而支持假鹰爪属和皂帽花属为两个独立属的观点。     

Plant stress and larval performance of a dipterous gall former

According to the plant vigour hypothesis, galling insects should respond positively and perform better on vigorous plants or plant parts, the opposite of the predictions of the plant stress hypothesis. I carried out field experiments to analyse the effects of sustained abiotic stress on the interactions between the common reed (Phragmites australis) and a gall-forming fly (Lipara lucens). The reed shoot diameter (a measure of plant vigour) is strongly affected by environmental conditions, where dry and/or nutrient-poor habitats produce thinner (stressed) shoots. L. lucens gall density is negatively correlated with shoot diameter. In a survival experiment with a wide range of shoot diameters, larval mortality was also highly correlated with shoot quality. Gall formation was higher on thinner, stressed shoots. An analysis of the gall tissues revealed that galls induced by L. lucens contain a high amount of a nutrient-rich feeding tissue. The impact of L. lucens is higher on thinner shoots. The results of this study showed that L. lucens performs better on stressed hosts, which contradicts the plant vigour hypothesis for galling insects. The low nutrient availability in the stressed shoots can be compensated by the production of galls with a nutrient-rich feeding tissue.     

Taxonomic significance of leaf anatomy of Aniselytron (Poaceae) as an evidence to support its generic validity against Calamagrostis s. l.

A comparative study of leaf anatomy on Aniselytron Merr. and Calamagrostis Adans. s. l. was conducted to review the systematic status of Aniselytron Merr. Calamagrostis s. l. exhibits wide variation in many features, but basic leaf structures of the genus remain constant: absence of a midrib-keel; median and large vascular bundles are central, with double sheaths, accompanied by girders both adaxially and abaxially; prickles have a bulbous base and are not sunken; the abaxial epidermal cells are short and wide and relatively thick-walled. Aniselytron differs from Calamagrostis s. l. in: midrib-keel is present, consisting of a large central bundle with small ones on either side; all vascular bundles are abaxially situated, with abaxial girders only, parenchyma takes the place of the adaxial sclerenchyma; the bases of the prickles are sunken and are not bulbous; the abaxial epidermal cells are tall and thin-walled. These distinct anatomical features, in combination with the differences in spikelet structure and habitat, suggest that Aniselytron should be generically separated from and not merged with Calamagrostis s. l. Due to the adaxial parenchyma in the midrib which has never been found in Pooideae, Aniselytron might have a relationship with some other subfamily.     

泽米科植物羽片脉序和解剖学及其系统学意义

研究了苏铁目泽米科Zamiaceae 2亚科的所有4族(Stevenson系统, 1992)共10种代表植物的羽片脉序及解剖学特征,结果显示泽米科羽片脉序为二歧分叉的平行脉,无中脉。小刺双子铁Dioon spinulosum、大头非洲铁Encephalartos friderici-guilielmii和摩尔大泽米Macrozamia moorei等的平行脉末端以不同的形式互相连接,而鳞木铁Lepidozamia peroffskyana、粗壮角果铁Ceratozamia mexicana var. robusta、竹叶角果铁C. hildae、佛州泽米Zamia　floridana、柔叶泽米Z. debilis、鳞秕泽米Z. furfuracea和短尖泽米Z. muricata等的平行脉末端不连接而直达叶缘,其中鳞木铁、粗壮角果铁和竹叶角果铁的脉达叶缘后逐渐消失。羽片的横切面结构通常由表皮、下皮厚壁细胞和叶肉组成,表皮层包括上、下表皮各一层,叶肉可能同时分化出近上表面的栅栏组织和近下表面的栅栏组织,或仅有近上表面的栅栏组织分化,或无栅栏组织分化而完全为海绵组织。然而,泽米科没有典型的海绵组织和传输组织分化。小刺双子铁、大头非洲铁、鳞叶木铁和摩尔大泽米的羽片具有粘液道而无工字厚壁组织,在小刺双子铁中粘液道与维管束对生,在另3种中则与维管束轮生;但粗壮角果铁、竹叶角果铁、佛州泽米、柔叶泽米、鳞秕泽米和短尖泽米的羽片则具有工字厚壁组织而没有粘液道,其中粗壮角果铁和竹叶角果铁的羽片工字厚壁组织仅与上表皮相连,而佛州泽米、柔叶泽米、鳞秕泽米和短尖泽米的羽片工字厚壁组织与上、下表皮都相连。羽片脉序和解剖学特征支持Stevenson将泽米铁科分为两亚科的观点。     

杨树叶薄层培养中不定芽形态发生的细胞组织学研究

将杂种杨树(Populus nigra var.betulifolia×P.trichocarpe)NE299叶主脉用振动切片机横切成400μm或800μm的薄切片,培养在附加0.2mg/L BA和0.01mg/L NAA的木本植物培养基上。培养后,位于主脉维管束两侧中上部的维管束鞘薄壁细胞首先启动分裂。几乎同时,与其邻接的一些栅栏组织细胞也分裂,并很快形成胚性分生细胞团。主脉的愈伤组织主要由维管束鞘薄壁细胞,以及与其邻接的一些栅栏组织细胞和韧皮部的薄壁细胞分裂而来。不定芽通常发生在愈伤组织的周边区,也可以起源于维管组织结节(vascular nodules)周围的形成层状细胞。侧脉的维管束鞘细胞分裂活动很强,可不经愈伤组织直接长成不定芽。杨树叶主脉处的维管束鞘薄壁细胞在与叶肉组织相邻接的细胞中,通常含有少量较小的叶绿体,而位于背腹面的细胞中含有贮藏的淀粉。对形态发生的特定部位及其细胞进行了讨论。     

Why do the galls induced by Calophya duvauae Scott on Schinus polygamus (Cav.) Cabrera (Anacardiaceae) change colors?

One of the galling herbivores associated to the superhost Schinus polygamus (Cav.) Cabrera (Anacardiaceae) is Calophya duvauae Scott (Hemiptera: Calophyidae). The galls are located on the adaxial surface of leaves and vary from red to green. The levels of their pigments were herein investigated in relation to age. Samples were collected between June 2008 and March 2009, in a population of S. polygamus at Canguçu municipality, Rio Grande do Sul, Brazil. Galls were separated by color, measured, dissected, and the instar of the inducer was determined. The levels of photosynthetic (chlorophyll a and b, total chlorophylls, and carotenoids) and protective pigments (anthocyanins) were also evaluated. Red galls were more numerous than the green ones, and the induction should occur preferentially on young leaves, but may also occur on mature leaves. Immature stages of C. duvauae were observed in all samples throughout the year, characterizing its life cycle as multivoltine. There was a significant correlation between the instar of the inducer and the size of the gall (r = 0.675, p < 0.001), with larger galls corresponding to more advanced instars. The ratio between red and green galls varied over the samples, with the highest (96%) and lowest (38%) frequency of red galls observed in September, and December 2008, respectively. The average amounts of chlorophyll b and total chlorophylls were 70% lower in the gall tissues when compared to non-galled portions of galled leaves. There is a notable reduction in the contents of all pigments in the galls when compared to non-galled leaves, especially for total anthocyanins in green galls. The red galls presumably had the constant stimuli of healthy gall inducers, whereas in green galls they died prematurely, due to the interference of parasitoids and inquilines. The alterations in gall structure and color were related to the gradual decreasing in galling stimuli.     

Leaf gall polymorphism and molecular phylogeny of a new Bruggmanniella species (Diptera: Cecidomyiidae: Asphondyliini) associated with Litsea acuminata (Lauraceae) in Taiwan,with ecological comparisons and a species description

Two types of cecidomyiid leaf galls, cup‐shaped and umbrella‐shaped, occur on Litsea acuminata (Lauraceae) in Taiwan. Based on the concept of gall shapes as “extended phenotypes” of gall inducers, these two types could be induced by different gall midge species. However, galls with intermediate shapes between the two types were recently discovered, which implies that possible genetic exchanges occur between the gall inducers of both types. To clarify the taxonomic status of gall midges responsible for the two types of galls on L. acuminata, we undertook taxonomic, molecular phylogenetic and ecological studies. Our findings show that the two gall types are induced by the same Bruggmanniella species and the species is new to science. We describe the species forming this range of galls as Bruggmanniella litseae sp. n. , and compare their geographical distribution, galling position and morphometry. Based on our results, a possible evolutionary scenario of B. litseae sp. n. is discussed.     

The redifferentiation of nutritive cells in galls induced by Lepidoptera on Tibouchina pulchra (Cham.) Cogn. reveals predefined patterns of plant development

Insect galls may present nutritive tissues with distinct cytological features related to the order of the gall inducer. Galling Lepidoptera larvae chew plant cells and induce the redifferentiation of parenchymatic cells into nutritive ones. The nutritive cells in the galls induced by a microlepidoptera on the leaves of Tibouchina pulchra (Cham.) Cogn. (Melastomataceae) are organelle-rich, with developed Golgi apparatus, endoplasmic reticulum, ribosomes, polyribosomes, mitochondria, plastids, and one great central or several fragmented vacuoles. The nonobservance of the nuclei in the nutritive cells deserves special attention, and confers a similarity between the nutritive cells and the vascular conductive ones. The great amount of rough endoplasmic reticulum, ribosomes, polyribosomes, and mitochondria is indicative of the high metabolic status of these cells. They are vascular cambium-like, with high protein synthesis and lipid storage. The proteins are essential to enzymatic metabolism, and secondarily, to larvae nutrition, similarly to the lipid droplets which confer energetic profile to these nutritive cells. The living enucleated cells receive mRNA from their neighbor ones, which may support the high metabolic profile of endoplasmic reticulum and ribosomes observed in galls. Thus, the nutritive cells are stimulated by the galling larvae activity, generating a new cell type, whose redifferentiation includes a mix of intrinsic and common plant pathways.     

Cycad collicles (“leaflet nodules”)

The presence of small knobs/nodules found on the adaxial leaflet surface of some Encephalartos species was investigated in 134 cycad species of nine genera. The knobs, referred to as collicles, were found to be the result of connecting mucilage canal ‘bridges’ over vascular bundles, connecting mucilage canals situated between adjacent parallel vascular bundles. Collicles were found on some Encephalartos species only and not always on all individuals of the same species. The possible taxonomic value of these collicles is discussed.