Ultrastructure of the mouthparts of adult sweetpotato whitefly,Bemisia tabaci Gennadius (Homoptera: Aleyrodidae)

The fine structure of the mouthparts of the whitefly, Bemisia tabaci Gennadius (Homoptera: Aleyrodidae), was examined by scanning and transmission electron microscopy. Adult whitefly mouthparts are similar to those of other homopterans, especially aphids, being composed of the labrum, the labium, and the stylets. The stylet bundle is the feeding organ of the whitefly and is composed of 2 mandibular stylets and 2 maxillary stylets. Mandibular stylets, which are located on the outer aspect of the stylet bundle, each contain 2 dendrites. The tips of the mandibular stylets are curved inward, and there are barb-like ridges on the lateral aspects, which probably function in piercing and cutting plant tissues and in anchoring the stylets in the tissues. The maxillary stylets are not innervated and are interlocked to form 2 separate compartments, the food canal and salivary canal. At the distal end of the interlocked maxillary stylets, there is a small depression, which may allow for mixing of the salivary canal and food canal components. Movement of the B. tabaci stylets during feeding is discussed in comparison with other homopterans.     

Fine structure of the mouthparts of Diostrombus politus and Proutista moesta (Hemiptera: Derbidae)

Insect mouthparts are important sensory and feeding structures, morphological studies of this organ can provide additional data for phylogenetic studies. The planthopper superfamily Fulgoroidea is among the dominant groups of phytophagous insects, however, the mouthparts ultrastructure of this superfamily remains unsatisfactorily studied. This study investigates the ultrastructure of the mouthparts of two species in the family Derbidae (Hemiptera: Fulgoroidea), Diostrombus politus Uhler and Proutista moesta (Westwood), using a scanning electron microscope. The results show that these two derbids are of a typical piercing-sucking type found in Hemiptera. They consist of a cone-shaped labrum, a three-segmented labium and a stylet fascicle with two interlocked maxillary stylets incompletely wrapped by two mandibular stylets. The arrangement of the sensilla on the labial tip differ slightly between the two derbid species, and the subapical labial sensilla are likely different among genera in the family Derbidae.     

Morphology of the mouthparts of the spittlebug Philagra albinotata Uhler (Hemiptera: Cercopoidea: Aphrophoridae)

Mouthparts associated with feeding behavior and feeding habits are important sensory and feeding structures in insects. To obtain a better understanding of feeding in Cercopoidea, the morphology of mouthparts of the spittlebug, Philagra albinotata Uhler was examined using scanning electron microscopy. The mouthparts of P. albinotata are of the typical piercing–sucking type found in Hemiptera, comprising a cone-shaped labrum, a tube-like, three-segmented labium with a deep groove on the anterior side, and a stylet fascicle consisting of two mandibular and two maxillary stylets. The mandibles consist of a dorsal smooth region and a ventral serrate region near the apical half of the external convex region, and bear five nodules or teeth on the dorsal external convex region on the distal extremity; these are regarded as unique features that distinguish spittlebugs from other groups of Hemiptera. The externally smooth maxillary stylets, interlocked to form a larger food canal and a smaller salivary canal, are asymmetrical only in the internal position of longitudinal carinae and grooves. One dendritic canal is found in each maxilla and one in each mandible. Two types of sensilla trichodea, three types of sensilla basiconica and groups of multi-peg structures occur in different locations on the labium, specifically the labial tip with two lateral lobes divided into anterior sensory fields with ten small peg sensilla arranged in a 5 + 4 + 1 pattern and one big peg sensillum, and posterior sensory fields with four sensilla trichodea. Compared with those of previously studied Auchenorrhyncha, the mouthparts of P. albinotata may be distinguished by the shape of the mandibles, the multi-peg structures and a tooth between the salivary canal and the food canal on the extreme end of the stylets. The mouthpart morphology is illustrated using scanning electron micrographs, and the taxonomic and putative functional significance of the different structures is briefly discussed.     

Fine structure and sensory apparatus of the mouthparts of the pear psyllid,Cacopsylla chinensis (Yang et Li) (Hemiptera: Psyllidae)

The pear psyllid, Cacopsylla chinensis (Yang et Li) (Hemiptera: Psyllidae), is one of the most significant economic pests of pear in China, causing direct damage through feeding by the highly specialized piercing–sucking mouthparts. The ultrastructural morphology and sensory apparatus of the mouthparts of the adult were examined using scanning and transmission electron microscopy. The piercing–sucking mouthparts of C. chinensis are composed of a three-segmented labium with a deep groove in the anterior side, a stylet fascicle consisting of two mandibular and two maxillary stylets, and a pyramid-shaped labrum. Proximal to the labium, the stylet fascicle forms a large loop within a membranous crumena. Mandibles, with more than ten teeth on the external convex region, can be seen on the distal extremity. Smooth maxillary stylets are interlocked to form a larger food canal and a smaller salivary canal. One dendritic canal housing 2 dendrites is also found in each mandible. Two types of sensilla trichodea, four types of sensilla basiconica, single as well as groups of sensilla campaniformia, and oval flattened sensilla occur in different locations on the labium, whereas a kind of sensilla basiconica is at the junction of the labrum and anteclypeus. Sensilla trichodea and sensilla campaniformia, always present with denticles, are present on the middle labial segment. Three types of sensilla basiconica, two types of sensilla trichodea and two oval flattened sensilla are located on the distal labial segment. The mouthpart morphology and abundance of sensilla located on the labium in C. chinensis are illustrated, along with a brief discussion of their taxonomic and putative functional significance.     

Morphological description of the mouthparts of the Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae)

Scanning (SEM) and transmission (TEM) electron microscopy were used to elucidate the morphology of the rostrum, as well as the mandibular and maxillary stylets of the psyllid Diaphorina citri, vector of phloem-inhabiting bacteria associated with citrus huanglongbing (HLB) disease. D. citri has a cone-shaped rostrum that extends behind the pair of prothoracic coxae. The stylet bundle comprises a pair of mandibular (Md) and maxillary (Mx) stylets with a mean length of 513.3 μm; when retracted, their proximal portions form a loop and are stored in the crumena (Cr). Serial cross-sections of the rostrum revealed that the mandibles are always projected in front of the maxillary stylets. The two maxillary stylets form the food and salivary canals, with diameters of 0.9 μm and 0.4 μm respectively. These two canals merge at the end of the stylets forming a common duct with a length of 4.3 μm and a mean diameter of 0.9 μm. The acrostyle, a distinct anatomical structure present in the common duct of aphid maxillary stylets, was not observed by TEM in the ultrathin cross-sections of the common duct (CD) of D. citri. This study provides new information on D. citri mouthparts that may help to understand the feeding behaviour of this important vector of HLB-associated bacteria.     

Pronymphs,hatching, and proboscis assembly in leafhoppers and froghoppers (Hemiptera,Cicadellidae and Aphrophoridae)

Pharate 1st instar nymphs enclosed in the embryonic cuticle, referred to as pronymphs, were studied in a froghopper Aphrophora pectoralis Mats. (Aphrophoridae) and the leafhoppers Oncopsis flavicollis (L.), Populicerus populi (L.), Alebra wahlbergi (Boh.), Igutettix oculatus (Lindb.), and Scenergates viridis (Vilb.) (Cicadellidae). The species vary in the relative length of the pronymphal antennae and details of sculpturing of the cephalic region. No egg bursting structures were observed, except small denticles on the crown region of S. viridis pronymphs. Rudimentary mandibular and maxillary stylets of a pronymph are external, short, tubular appendages containing tips of the corresponding nymphal stylets, whose more basal parts develop inside of the head. Casting off of the embryonic cuticle results in the nymphal stylets being passively pulled out and assuming a close-set parallel orientation. Once the sheaths of unsclerotized cuticle secreted by the peripodial epithelium and enveloping each developing stylet have been cast off with the exuviae, the bare stylets become squeezed and interlocked into a functional bundle. The roles of the maxillary plates, clypeus, labrum, and labium in the stylet bundle assembly are discussed. The process repeats after each molt.     

Mouthpart morphology and stylet penetration of host plants by the glassy-winged sharpshooter, Homalodisca coagulata, (Homoptera: Cicadellidae)

The ultrastructural morphology of the mouthparts of the glassy-winged sharpshooter, Homalodisca coagulata, and method of plant penetration was examined using light microscopy, scanning electron microscopy, and transmission electron microscopy methods. The gross morphology of the labrum, labium, and stylet fascicle was consistent with what has been described for other plant-sucking homopterans. The ultrastructural examination of the mouthparts revealed unique details that have previously gone unreported. Several types of sensilla-like structures having the form of pegs and multi-lobed objects were identified on the outer surfaces of the labrum and within the labial groove. Dendritic canals terminated in an extensive network of smaller canals at the distal tip of the maxillary stylets below a series of surface denticles suggesting that this area may have a sensory function associated with locating xylem elements of host plants. Examination of salivary sheath pathways established that 65% of the plant penetrations by this insect terminated in the xylem vessels of the host plant. Probing by the insect was largely intracellular and terminal branching of a single probe site was common. Plant surface feeding sites varied with the stage of development which correlates with the depth of the xylem vessels and the length of the maxillary stylets of the various instars.     

Mouthparts and stylet penetration of the lac insect Kerria lacca (Kerr) (Hemiptera:Tachardiidae)

Hitherto less known aspects on mouthpart morphology and penetration mechanism of the lac insect Kerria lacca have been explored. Unique details of the mouthparts, i.e. morphology of labium and stylets and salivary sheath have been brought out. The gross morphology of the mouthparts though resembled other plant sucking homopterans; a two-segmented labium with symmetrically distributed six pairs of contact-chemoreceptors on its surface was distinct; the mandibular stylets had serrations on its extreme apical region, while the maxillary stylets had their external surface smooth with parallel longitudinal grooves on their inner surface. Formation of flanges, salivary sheath and penetration pathway observed along with probing and penetration of the stylets intracellularly up to the phloem cells, as illustrated herein, are the addition to the existing knowledge on the structural details of the mouthparts and the feeding behavior thereupon.     

Correlation of stylet activities by the glassy-winged sharpshooter, Homalodisca coagulata (Say), with electrical penetration graph (EPG) waveforms

Glassy-winged sharpshooter, Homalodisca coagulata (Say), is an efficient vector of Xylella fastidiosa (Xf), the causal bacterium of Pierce's disease, and leaf scorch in almond and oleander. Acquisition and inoculation of Xf occur sometime during the process of stylet penetration into the plant. That process is most rigorously studied via electrical penetration graph (EPG) monitoring of insect feeding. This study provides part of the crucial biological meanings that define the waveforms of each new insect species recorded by EPG. By synchronizing AC EPG waveforms with high-magnification video of H. coagulata stylet penetration in artifical diet, we correlated stylet activities with three previously described EPG pathway waveforms, A1, B1 and B2, as well as one ingestion waveform, C. Waveform A1 occured at the beginning of stylet penetration. This waveform was correlated with salivary sheath trunk formation, repetitive stylet movements involving retraction of both maxillary stylets and one mandibular stylet, extension of the stylet fascicle, and the fluttering-like movements of the maxillary stylet tips. Waveform B1 was ubitquious, interspersed throughout the other waveforms. B1 sub-type B1w was correlated with salivation followed by maxillary tip fluttering. This tip fluttering also occurred before and during B1 sub-type B1s, but was not directly correlated with either the occurrence or frequency of this waveform. Waveform B2 was correlated with sawing-like maxillary stylet movements, which usually occurred during salivary sheath branching. Waveform C was correlated with ingestion. Fluid outflow was also observed as a mechanism to clear the maxillary tips from debris during waveform C. This detailed understanding of stylet penetration behaviors of H. coagulata is an important step toward identifying the instant of bacterial inoculation which, in turn, will be applied to studies of disease epidemiology and development of host plant resistance.     

Oviposition behaviour in Lygus rugulipennis: a morpho-functional study

Before oviposition, a Lygus rugulipennis (Poppius) (Heteroptera: Miridae) female explores the substrate with her mouthparts, performing ‘probing’ behaviour which eventually leads to a choice of oviposition site. In this study, morphological and behavioural observations were carried out in order to define the mechanisms through which female bugs recognise suitable oviposition sites. The morphological study was conducted using electron‐microscopy techniques, while the behavioural experiments aimed at the temporary (using ZnSO₄) or permanent (i.e., cauterization of the rostrum and ablation of both stylets and labium) inactivation of sensory structures located in the labium tip and in the stylets, which are probably involved in oviposition site selection. The tip of the labium of L. rugulipennis females is characterised by the presence of 11–12 uniporous gustatory sensilla which are innervated by 3–6 sensory neurones. One aporous mechanoreceptor is located ventrally on both areas of the labium tip. Other aporous mechanosensilla are found more proximally on the labium. ZnSO₄ labium tip treatment did not affect oviposition site selection, while stylet amputations, as well as rostrum cauterisations, resulted in almost complete oviposition failure. Labium tip‐ablated females oviposited similarly to control females. These results suggest that the sensory neurones associated with the stylets are involved in the location of the oviposition site. Uniporous gustatory sensilla on the labium tip are not involved in the final steps of the oviposition behaviour. However, they seem to play a role in assessing the suitability of the substrate as a food source.     

Anterior alimentary canal of the pear Psylla,Psylla pyricola foerster (Homoptera,Psyllidae)

Scanning and light microscopy investigations of the anterior alimentary canal of the pear psylla, Psylla pyricola Foerster (Homoptera: Psyllidae), revealed the morphology of the labium and stylets, as well as the presence of sensory structures and a valve in the precibarium. The labium consists of three telescoping segments with an internal labial groove, which surrounds and supports the stylet bundle. Also a part of the labial groove is the internal labial clamp. The stylet bundle is comprised of paired styliform mandibles and maxillae, which interlock to form the food and salivary canals. The stylet bundle proximal to the labium forms a large loop within a membranous crumena. When fully retracted the coiled stylets are under tension. Stylet extension generates increasing tension so that when retracted the stylets readily recoil within the crumena. Penetration of leaf tissues by the stylet bundle is dependent on the interaction between stylet muscles, opening and closing of the labial clamp, the barbed stylet tips, and the ventral position of the labium. Proximal to the crumena the paired stylets separate and diverge at the entrance of the precibarium, which is formed by the interlocking of the epi-and hypopharynges. There are 18 sensory structures in the precibarium, as well as a precibarial valve. These structures appear to be homologous to similar structures observed previously in aphids and leafhoppers. The morphology and the location of the precibarial sensilla suggest that, like the precibarial sensory organs of aphids and leafhoppers, they are gustatory and probably mediate acceptance or rejection of plant fluids, thus playing a major role in locating tissues for feeding.     

Asian citrus psyllid stylet morphology and applicability to the model for inter-instar stylet replacement in the potato psyllid

In Hemiptera, presumptive stylets for each consecutive postembryonic instar are manufactured prior to ecdysis to replace the ecdysial stylets discarded with the exuviae. With the discovery that the bacterium “Candidatus” Liberibacter solanacearum accesses the tissues involved in the stylet replacement process of the potato psyllid, a hypothesis was formed that the bacterium could adhere to the stylets of freshly emerged instars and hence gain access to the host plant when feeding is resumed. Although unproven, it was imperative that a model for stylet replacement be built. Stylet morphology and the stylet replacement process of the Asian citrus psyllid (ACP), vector of “C.” L. asiaticus, causal pathogen of citrus greening disease, are comparable to the potato psyllid model system. Morphology consists of a basal terminus with its tab-shaped auricle, a base, shaft, and an apical terminus. Each of the four auricles act as a platform for the replacement apparatus, which is compacted into a tight aggregate of cells, the ‘end-cap’. As modeled, on apolysis of larval instar hypodermis, the aggregate ‘deconstructs’ and expands into a snail shell-shaped tube, the ‘atrium’, that houses the presumptive stylet as it is synthesized. Completed stylets then despool from the atrium and are fitted into their functional positions as the next instar emerges from its exuviae.     

Functional analyses in the hemipteran Oncopeltus fasciatus reveal conserved and derived aspects of appendage patterning in insects

The conservation of expression of appendage patterning genes, particularly Distal-less, has been shown in a wide taxonomic sampling of animals. However, the functional significance of this expression has been tested in only a few organisms. Here we report functional analyses of orthologues of the genes Distal-less, dachshund, and homothorax in the appendages of the milkweed bug Oncopeltus fasciatus (Hemiptera). This hemimetabolous insect has typical legs but highly derived mouthparts. Distal-less, dachshund, and homothorax are conserved in their individual expression patterns and functions in the legs of Oncopeltus, but their functions in other appendages are in some cases divergent. We find that specification of antennal identity does not require wild-type Distal-less activity in Oncopeltus as it does in Drosophila. Additionally, the mouthparts of Oncopeltus show novel patterns of gene expression and function, relative to other insects. Expression of Distal-less in the maxillary stylets of Oncopeltus does not seem necessary for proper development of this appendage, while dachshund and homothorax are crucial for formation of the mandibular and maxillary stylets. These data are used to evaluate hypotheses for the evolution of hemipteran mouthparts and the evolution of developmental mechanisms in insect appendages in general.     

Development of the mouthparts in embryos of Haplothrips verbasci (Osborn) (Insecta,Thysanoptera, Phlaeothripidae)

The asymmetric “punch and suck” mouthparts of larval Haplothrips verbasci develop from paired appendages in the late, post-anatrepsis embryo similar to those of other insects. Later, the labrum flexes ventrally over the stomodaeum, the right mandibular appendage degenerates, the maxillary appendages divide into inner (lacinial) and outer (stipital) lobes, and the hypopharynx arises from the venters of the mandibular and maxillary segments. All cephalic segments consolidate anteriorly prior to katatrepsis, their appendages flex ventrally, and the labial appendages fuse medially to form the labium and the primordia of the salivary glands and valve. The left mandible and the lacinial lobes of the maxillae invaginate into the head during and after katatrepsis to form the mandibular and maxillary stylet-secreting organs and these later deposit the cuticle of their respective stylets. Cuticle of the mandibular lever is deposited by labral cells at the apex of the mandibular sheath during and after hatching. That of each maxillary lever is secreted simultaneously into the lumen of a ventrally-directed diverticulum developing from stipital cells at the apex of each maxillary sheath. Shortly after katatrepsis, the maxillary and labial palpi originate respectively from cells in the outer wall of each stipital lobe and at the apex of the labium. Muscles of the mouthparts arise after katatrepsis from cephalic mesoderm and are fully-differentiated before cuticle of the mandibular and maxillary levers has been deposited. Gnathal morphogenesis in embryos of H. verbasci resembles that occurring in bug embryos and provides additional evidence that Thysanoptera and Hemiptera evolved from a common psocopteroid stem species having small, paired, biting and chewing mandibles and well developed lacinial stylets.     

Electrical penetration graphic waveforms in relation to the actual positions of the stylet tips of Nilaparvata lugens in rice tissue

The stylet penetration behavior of Nilaparvata lugens (Hemiptera: Delphacidae) in rice plants (Oryza sativa) was evaluated through the use of an electrical penetration graph (EPG). To accomplish this, we classified the EPG signals into seven different waveforms, np, N1, N2, N3, N4-a, N4-b, and N5, according to their shapes, amplitudes, and frequencies. The N4-b waveform was always preceded by N3 and N4-a, in that order. Continuous honeydew excretion only occurred during the N4-b period, and the honeydew deposited on a filter paper containing ninhydrin reagent during the N4-b period was stained violet. The tips of the stylets that were severed in the N3, N4-a, and N4-b periods were in the phloem region of rice. Moreover, the flow of plant sap after stylectomy only occurred during the N4-b period. Finally, sucrose was the only carbohydrate component identified when HPLC analysis of the plant sap was conducted. On the other hand, honeydew excretion hardly occurred during the N5 period and the tips of the stylets that were severed during the N5 period were located in the xylem region of rice. Based on the location of the stylets and honeydew excretion, the EPG waveforms for the stylet penetration behaviors of N. lugens were assigned to the following groups; np: non-penetration of stylets, N1: penetration initiation, N2: salivation and stylet movement, N3: an extracellular activity near the phloem region, N4-a: an intracellular activity in phloem region, N4-b: phloem sap ingestion, and N5: activity in the xylem region.     

The “acrostyle”: A newly described anatomical structure in aphid stylets

The recent demonstration that a plant virus could be retained on protein receptors located exclusively in a small area inside the common duct at the tip of aphid maxillary stylets indicated the possible existence of a distinct anatomical structure at this level. Since no distinct feature within the common duct of any aphid species has ever been reported in the literature, we first carefully re-examined the distal extremity of the maxillary stylets of Acyrthosiphon pisum using transmission- and scanning-electron microscopy. Here, we describe an area of the cuticle surface displaying a different structure that is limited to a “band” paving the bottom of the common duct in each opposing maxillary stylet. This band starts at the very distal extremity, adopts a “comma-like” shape as it continues up towards the salivary canal, reducing in width and disappearing before actually reaching it. Investigations on several aphid species led to the conclusion that this anatomical feature—which we have tentatively named the “acrostyle”—is highly conserved among aphids. We then produced an antibody recognizing a consensus peptide located in the middle of the RR-2 motif of cuticular proteins from A. pisum and showed that this motif is accessible specifically within the acrostyle, indicating a higher concentration of cuticular proteins. While it is clear that at least some viruses can use the acrostyle to interact with their aphid vectors to ensure plant-to-plant transmission, the role of this new “organ” in aphid biology is unknown and calls for further investigation in the near future.     

Sensory receptors associated with the stylets and cibarium of the rice brown planthopper,Nilapavarta lugens

Summary The stylets of Nilapavarta lugens consist of two maxillae that interlock to form separate food and salivary ducts partially surrounded by two mandibles. The ultrastructure of the sensory innervation of the stylets is described. Each maxilla possesses five neurones which extend to the tip of the stylet. The mandibles also contain five neurones, four of which are paired. The paired neurones comprise a shorter dendrite extending part of the way along the stylet and a longer one extending to the tip. The possible functions of these neurones are discussed. Gustatory receptors are located in the small passageway leading from the food duct to the cibarium. The receptors are in two distinct groups on the epipharyngeal side and one group on the hypopharyngeal side of the food canal. Two to five neurones innervate each receptor which connects to the food canal via a small pore.     

Assessing the validity of stylets as ageing tools in Octopus pallidus

A method to age octopus was determined through the quantification of growth increments within the stylet structure (highly reduced internal shells). To validate the periodicity of the increments, stylets were analysed from known-age laboratory-raised Octopus pallidus. The animals, which ranged from 3 to 8 months old, were exposed either to a simulated natural or to constant temperature regime. Transverse stylet sections were embedded in thermo-plastic cement and ground until a thin section was achieved. A pre-hatch region and first post-hatch increment was identified in the stylet microstructure. The number of increments, across all ages and temperature treatments, was in remarkably close agreement to age (number of days), clearly demonstrating that stylet increments are deposited daily throughout the lifecycle of O. pallidus. Morphometric analyses of the stylet indicated that increments were laid down regularly during stylet growth and that stylets grew in concert with body size, further supporting daily periodicity. This study successfully validates daily increments in stylets and thus demonstrate a method to age octopus, therefore, potentially having critical implications for future octopus research and the effective management of stocks worldwide.     

Observations on penetration of Barley leaves by the aphidRhopalosiphum maidis (Fitch)

Summary Penetration of leaves of barley,Hordeum vulgare L., by the corn leaf aphid,Rhopalosiphum maidis (Fitch), was studied with light, phase, and electron microscopes. Penetration of epidermis and mesophyll was largely intercellular, that of vascular bundles or veins largely intracellular. Like other aphids,R. maidis secretes a salivary sheath which surrounds the stylets. When mesophyll cells and parenchymatous elements of the veins were penetrated by stylets, their protoplasts were pushed to one side by intruding sheath material; hence, the protoplasts were not punctured by the stylets, although sometimes the plasmalemma of penetrated cells was ruptured by sheath material. The salivary sheaths ended more or less abruptly outside the walls of sieve elements being fed upon, the maxillary stylets projecting beyond the sheaths and into the sieve elements. Before penetrating a functional sieve element the aphid apparently flushes its stylets in order to clear them for ingestion of food. Salivary and food canals merge near the tips of the maxillary stylets to form a single canal, which ends short of the tips.This research was supported by the U.S. National Science Foundation (GB-8330).     

An investigation of preferential feeding habit in fourAsclepiadaceae by the Aphid,Aphis nerii B. de F.

Summary Penetration of the stems ofAraujia sericofera, Asclepias curassavica, Cynanchum ellipticum andSarcostemma viminale by stylets of the aphidAphis nerii was studied with light and differential interference contrast microscopes. Of a total of 118 stylets and 446 stylet tracks observed in cross-sections of stems of the fourAsclepiadaceae, 97 stylets and 372 stylet tracks terminated within the internal primary phloem. Of the remainder, 15 stylets and 74 tracks terminated within the external primary phloem. 22 stylets and 179 of the stylet tracks penetrated the external phloem on the way to the internal phloem. Of these, only four stylets and 32 of the stylet tracks showed signs of attempted probes of the external phloem. It is suggested thatAphis nerii may obtain not only its essential food requirements but also cardiac glycosides as a basis for chemical aposematism.