Cibarial Pump Research Articles

Born suckers: the cibarial pump of Philaenus spumarius scales across ontogeny to ensure functional equivalence.

A select group of hemipterans within the suborder Auchenorrhyncha are the only animals that feed exclusively on xylem sap - a nutritionally poor liquid that exists under negative pressure within a plant's xylem vessels. To consume it, xylem-feeding bugs have evolved enlarged cibarial pumps capable of generating enormous negative pressures. A previous study examining the allometry of this feeding model suggested that small xylem feeders pay relatively higher energetic costs while feeding, favouring the evolution of larger-bodied species. However, this interspecific analysis only considered adult xylem-feeding insects and neglected the considerable intraspecific change in size that occurs across the insect's development. Here, we examine the changes in cibarial pump morphology and function that occur during the development of Philaenus spumarius, the common meadow spittlebug. We show that the cibarial pump scales largely as expected from isometry and that the maximum negative pressure is mass independent, indicating that size has no effect on the xylem-feeding capacity of juvenile spittlebugs. We conclude that a first instar nymph with a body mass 2% of the adult can still feed at the >1 MPa tension present in a plant's xylem vessels without a substantial energetic disadvantage.

The cibarial pump of the xylem-feeding froghopper Philaenus spumarius produces negative pressures exceeding 1 MPa.

The xylem sap of vascular plants is an unlikely source of nutrition, being both nutrient poor and held under tensions (negative pressures) that can exceed 1 MPa. But some insects feed on xylem sap exclusively, extracting copious quantities using a muscular cibarial pump. However, neither the strength of the insect's suction, nor the direct energetic cost of xylem ingestion, have ever been quantified. Philaenus spumarius froghoppers were used to address these gaps in our knowledge. Micro-CT scans of its cibarium and measurements of cibarial muscle sarcomere length revealed that P. spumarius can generate a maximum tension of 1.3 ± 0.2 MPa within its cibarium. The energetic cost of xylem extraction was quantified using respirometry to measure the metabolic rate (MR) of P. spumarius while they fed on hydroponically grown legumes, while xylem sap excretion rate and cibarial pumping frequency were simultaneously recorded. Increasing the plants' xylem tensions up to 1.1 MPa by exposing their roots to polyethylene glycol did not reduce the insects' rate of xylem excretion, but significantly increased both MR and pumping frequency. We conclude that P. spumarius can gain energy feeding on xylem sap containing previously reported energy densities and at xylem tensions up to their maximum suction capacity.

Life habits and evolutionary biology of new two-winged long-proboscid scorpionflies from mid-Cretaceous Myanmar amber

Long-proboscid scorpionflies are enigmatic, mid-Mesozoic insects associated with gymnosperm pollination. One major lineage, Aneuretopsychina, consists of four families plus two haustellate clades, Diptera and Siphonaptera. One clade, Pseudopolycentropodidae, from mid-Cretaceous Myanmar amber, contains Parapolycentropus. Here, we newly establish Dualula, assigned to Dualulidae, constituting the fifth lineage. Parapolycentropus and Dualula lineages are small, two-winged, with unique siphonate mouthparts for imbibing pollination drops. A cibarial pump provides siphonal food inflow; in Dualula, the siphon base surrounds a hypopharynx housing a small, valved pump constricted to a narrow salivary duct supplying outgoing enzymes for food fluidization. Indirect evidence links long-proboscid mouthpart structure with contemporaneous tubulate ovulate organs. Direct evidence of gymnospermous Cycadopites pollen is associated with one Parapolycentropus specimen. Parapolycentropus and Dualula exhibit hind-wing reduction that would precede haltere formation, likely caused by Ultrabithorax. Distinctive, male Aneuretopsychina genitalia are evident from specimens in copulo, supplemented by mixed-sex individuals of likely male mating swarms.

Studies on gas exchange in the meadow spittlebug, Philaenus spumarius: the metabolic cost of feeding on, and living in, xylem sap.

Spittlebugs (superfamily Cercopoidea) live within a mass of frothy, spittle-like foam that is produced as a by-product of their xylem-feeding habits. The wet spittle represents a unique respiratory environment for an insect, potentially acting either as a reserve of trapped oxygen (O2) or as a significant barrier to O2 diffusion from the surrounding atmosphere. Feeding on xylem sap under tension is also assumed to be energetically expensive, potentially placing further constraints on their gas exchange. To understand the respiratory strategies used by spittlebugs, this study measured the PO2 within the spittle of the meadow spittlebug, Philaenus spumarius, as well as the non-feeding metabolic rate (RMR) and respiratory quotient (RQ) of both nymphs and adults. The metabolic rate of nymphs feeding on xylem was also measured. In separate experiments, the ability of a nymph to obtain O2 from bubbles while submerged in foam was determined using a glass microscope slide coated in an O2-sensitive fluorophore. We determined that P. spumarius breathes atmospheric O2 by extending the tip of its abdomen outside of its spittle, rather than respiring the O2 trapped in air bubbles within the foam. However, spittlebugs can temporarily use these air bubbles to breathe when forcibly submerged. V̇O2 and V̇CO2 did not differ statistically within life stages, giving a RQ of 0.92 for nymphs and 0.95 for adults. Feeding on xylem was found to increase the nymphs' V̇CO2 by only 20% above their RMR. From this cost of feeding, cibarial pump pressures were estimated to be between -0.05 and -0.26 MPa.

Morphological fine tuning of the feeding apparatus to proboscis length in Hesperiidae (Lepidoptera).

The form and function of the hesperiid feeding apparatus was studied in detail. The butterflies in the family Hesperiidae are of particular interest because the longest proboscis ever recorded in Papilionoidea was found in the Neotropical genus Damas. We focused on the functional morphology by comparing proboscis morphology as well as size and composition of both the stipes pump and the cibarial suction pump in skippers with short and extremely long proboscis. Results revealed that all studied Hesperiidae have the same proboscis micromorphology and sensilla endowment regardless of the proboscis length. However, the numbers of internal muscles of the proboscis, the morphology of the stipes pump as well as the pumping organs for nectar uptake are related to the proboscis length. We conclude that the low number of tip sensilla compared to proboscis length is responsible for remarkably longer manipulation times of long-proboscid species during flower visits. The organs for proboscis movements and nectar uptake organs are well tuned to the respective proboscis length and are accordingly bigger in species with a proboscis that measures twice the body length.

AUSTIN HUMBERT, and JAMES JOY, Department of Biological Sciences, Marshall University, Huntington, WV, 25755. Sensory sensilla associated with the cibarial pump of Aedes vexans (Diptera: Culicidae).

Sensilla of the cibarium, beginning at the base of the labrum and extending proximally through the stomodaeum, were examined in both female and male Aedes vexans (Meigen) collected from Wayne Co., WV in June of 2016. Two types of sensilla were observed; (1) setiform, and (2) basiconic. Both forms of sensilla were aggregated in the distal region of the cibarium surrounding a hard palate, with the basicones being medially positioned and the setiforms lateral. Additionally, two pairs of basiconic sensilla were consistently observed in the proximal end (i.e., stomodaeum) of the cibarium in every individual examined.

Salt controls feeding decisions in a blood-sucking insect

Salts are necessary for maintaining homeostatic conditions within the body of all living organisms. Like with all essential nutrients, deficient or excessive ingestion of salts can result in adverse health effects. The taste system is a primary sensory modality that helps animals to make adequate feeding decisions in terms of salt consumption. In this work we show that sodium and potassium chloride salts modulate the feeding behavior of Rhodnius prolixus in a concentration-dependent manner. Feeding is only triggered by an optimal concentration of any of these salts (0.1–0.15M) and in presence of the phagostimulant ATP. Conversely, feeding solutions that do not contain salts or have a high-salt concentration (>0.3M) are not ingested by insects. Notably, we show that feeding decisions of insects cannot be explained as an osmotic effect, because they still feed over hyperosmotic solutions bearing the optimal salt concentration. Insects perceive optimal-salt, no-salt and high-salt solutions as different gustatory information, as revealed the electromyogram recordings of the cibarial pump. Moreover, because insects do a continuous gustatory monitoring of the incoming food during feeding, sudden changes beyond the optimal sodium concentration decrease and even inhibit feeding. The administration of amiloride, a sodium channel blocker, noticeably reduces the ingestion of the optimal sodium solution but not of the optimal potassium solution. Salt detection seems to occur at least through two salt receptors, one amiloride-sensitive and another amiloride-insensitive. Our results confirm the importance of the gustatory system in R. prolixus, showing the relevant role that salts play on their feeding decisions.

Role of Phote-HrTH (Phormia terraenovae hypertrehalosemic hormone) in modulating the supercontractile muscles of the crop of adult Phormia regina Meigen

Phote-HrTH (Phormia terraenovae hypertrehalosemic hormone) has been demonstrated in the Diptera to be involved in flight metabolism, reproduction, and diapause. Each of these events needs the hormone’s action and requirement for carbohydrates is the common denominator. In Diptera, carbohydrates are taken up during feeding by action of the cibarial pump and are then stored in the crop. Using adult Phormia regina, both a bioassay and electrophysiological recordings show that Phote-HrTH slows down or inhibits the crop lobe muscles (P5) and, at the same time, stimulates the muscles of the pump 4 (P4) involved in pushing fluids out of the crop and up into the midgut for digestion. The EC50 for P4 was in the nanomolar range while the IC50 for P5 was 1.4–75.1pM. The effect of Phote-HrTH on P4/5 suggests that the peptide is important in coordinating the two pumps, which are involved in moving carbohydrates up into the midgut for digestion. The adult crop organ is an essential storage organ for carbohydrates and now should be considered an important structure capable of delivering nutrients to the midgut for digestion.

Drinking with an Unsealed Tube: Fluid Uptake Along the Butterfly Proboscis

Abstract Most adult Lepidoptera depend on a proboscis for fluid uptake. Although the proboscis has been regarded as a sealed tube with fluid uptake restricted to the distal end, recent evidence indicates that it is permeable along its entire length in at least some species. We, therefore, tested the effectiveness of the seal during feeding in four species of butterflies. Feeding rates in monarchs (Danaus plexippus L.), painted ladies (Vanessa cardui L.), and tiger swallowtails (Papilio glaucus L.) did not differ significantly when the proboscises were straightened and fully, versus partially, submersed in 1 or 15% sucrose solutions. To explore these results, we tested fluid uptake along the nearly transparent proboscises of buckeye butterflies (Junonia coenia Hübner) by applying colored droplets of water to the legular seam between the paired galeae. Colored fluid appeared in the food canal of straightened and naturally flexed proboscises within 10 s, regardless of whether the chemosensilla were stimulated with sugar. Statistically significant entry of fluid, however, occurred ≈30 s after droplets were applied and only if the proboscis was naturally flexed and stimulated with sucrose. The results suggest that fluid uptake along the length of the proboscis is influenced by changes in legular spacing when the butterfly naturally bends the proboscis and on activation of the cibarial pump when chemosensilla are stimulated with sugar.

Experimental analysis of the liquid-feeding mechanism of the butterfly Pieris rapae

The butterfly Pieirs rapae drinks liquid using a long proboscis. A high pressure gradient is induced in the proboscis when cibarial pump muscles contract. However, liquid feeding through the long proboscis poses a disadvantage of high flow resistance. Hence, butterflies may possess special features to compensate for this disadvantage and succeed in foraging. The main objective of this study is to analyze the liquid-feeding mechanism of butterflies. The systaltic motion of the cibarial pump organ was visualized using the synchrotron X-ray imaging technique. In addition, an ellipsoidal pump model was established based on synchrotron X-ray micro-computed tomography. To determine the relationship between the cyclic variation of the pump volume and the liquid-feeding flow, velocity fields of the intake flow at the tip of the proboscis were measured using micro-particle image velocimetry. Reynolds and Womersley numbers of liquid-feeding flow in the proboscis were ~1.40 and 0.129, respectively. The liquid-feeding flow could be characterized as a quasi-steady state laminar flow. Considering these results, we analyzed the dimensions of the feeding apparatus on the basis of minimum energy consumption during the liquid-feeding process. The relationship between the proboscis and the cibarial pump was determined when minimum energy consumption occurs. As a result, the volume of the cibarial pump is proportional to the cube of the radius of the proboscis. It seems that the liquid-feeding system of butterflies and other long-proboscid insects follow the cube relationship. The present results provide insights into the feeding strategies of liquid-feeding butterflies.

Intravital microscopy and image analysis of Rhodnius prolixus (Hemiptera: Reduviidae) hematophagy: The challenge of blood intake from mouse skin

Hematophagous insects transmit many of the most dangerous parasitic diseases. The transmission usually occurs during hematophagy or just after as this is when the vector and the host are in contact. The contact time is determined by the feeding performance of the insect in each host. In triatomines, feeding performance interferes with both their life cycle and the vectorial competence to transmit the hemoflagellate Trypanosoma cruzi. Triatomine bugs are vessel feeders, obtaining their blood meals directly from the vessels (venules or arterioles) of their vertebrate hosts. The host blood intake rate is not constant during the feeding, and the sucking frequency of triatomines tends to be higher and to contain fewer interruptions in pigeons than in mice. To identify the difficulties encountered by triatomine bugs in obtaining blood meals from mouse skin, we used intravital microscopy techniques associated with electromyograms of the cibarial pump. To monitor the vibration of the cannulated vessels and the blood flow through the head of the insect during the engorgement phase, we introduced a novel method for image analysis. The mean number of vessels used during a Rhodnius prolixus blood meal was 3.4±1.2, and the insects fed more in venules (63%) than in arterioles (37%). An important increase in vascular permeability was observed throughout the feeding. Platelet aggregation, rolling and leukocyte adherence were analyzed on the venular endothelium, showing remarkable increases for some time following the R. prolixus feeding. The reduction in sucking frequency that was observed during insect feeding was likely due to the increased cibarial pump filling time. The monitoring of the vessel wall pulsation also permitted the registration of regurgitation-like movements during blood pumping, with these movements being recorded mostly during the second half of the feeding. The evaluation of blood flow through the head of the insect suggested that the regurgitation-like movements were not true regurgitations and were caused by abrupt difficulties in the function of the cibarial pump. The role of the platelet plugs and the changes in blood viscosity at the R. prolixus feeding site are discussed. The method introduced in the present study to analyze the images brings new insights into the interaction between hematophagous vectors and their hosts, reinforcing the importance of insect saliva throughout the feeding process.

Regulatory mechanisms and the role of calcium and potassium channels controlling supercontractile crop muscles in adult Phormia regina

Bioassays and electrophysiological recordings were conducted in the adult blowfly Phormia regina to provide new insights into the regulatory mechanisms governing the crop filling and emptying processes of the supercontractile crop muscles.The cibarial pump drives ingestion. Simultaneous multisite extracellular recordings show that crop lobe (P5) distension during ingestion of a 4.7 μl sugar meal does not require muscle activity by any of the other pumps of the system. Conversely, pumping of fluids toward the anterior of the crop system during crop emptying is brought about by active muscle contraction, in the form of a highly coordinated peristaltic wave starting from P5 and progressively propagating to P6, P4 and P3 pumps, with P5 contracting with a frequency about 3.4 times higher than the other pumps. The crop contraction rate is also modulated by hemolymph-borne factors such as sugars, through ligand recognition at a presumptive receptor site rather than by an osmotic effect, as assessed by both behavioural and electrophysiological experiments. In this respect, sugars of equal osmolarity produce different effects, glucose being inhibitory and mannose ineffective for crop muscles, while trehalose enhances crop activity.Finally, voltage and current clamp experiments show that the muscle action potentials (mAPs) at the P4 pump are sustained by a serotonin-sensitive calcium conductance. Serotonin enhances calcium entry into the muscle cells and this could lead, as an indirect modulatory effect, to activation of a Ca2+-activated K+ conductance (IK(Ca)), which sustains the following mAP repolarization phase in such a way that further mAPs can be generated early and the frequency consequently increased.

Immunohistological localization of serotonin in the CNS and feeding system of the stable fly STOMOXYS CALCITRANS L. (Diptera: Muscidae)

Serotonin, or 5-hydroxytryptamine (5-HT), plays critical roles as a neurotransmitter and neuromodulator that control or modulate many behaviors in insects, such as feeding. Neurons immunoreactive (IR) to 5-HT were detected in the central nervous system (CNS) of the larval and adult stages of the stable fly, Stomoxys calcitrans, using an immunohistological technique. The location and pattern of the 5-HT IR neurons are described and compared for these two different developmental stages. Anatomical features of the fly feeding system were analyzed in third instar larvae and adult flies using a combination of histological and immunohistological techniques. In third instar larvae, the cibarial dilator muscles were observed within the cibarial pump skeleton and innervated by 5-HT IR neurons in nerves arising from the brain. There were four pairs of nerves arising from the frontal surface of the larval brain that innervate the cibarial pump muscles, pharynx, and muscles controlling the mouth hooks. A strong serotoninergic innervation of the anterior stomatogastric system was observed, which suggests 5-HT may play a role in the coordination of different phases of food ingestion by larvae. Similarly, many 5-HT IR neurons were found in both the brain and the thoracico-abdominal ganglia in the adult, some of which innervate the cibarial pump dilator muscles and the stomatogastric muscles. This is tnhe first report describing neuromuscular structures of the stable fly feeding system. The results reported here suggest 5-HT may play a critical role in feeding behaviors of stable fly larvae and adults.

Influence of the intestinal anticoagulant in the feeding performance of triatomine bugs (Hemiptera; Reduviidae)

Triatomines are haematophagous insects in all post-embryonic life stages. They are vectors of Trypanosoma cruzi, the causative agent of Chagas disease. Their vectorial ability is influenced by their feeding performance, which varies greatly amongst species. Recent work showed that inhibition of the coagulation process in the anterior midgut (crop) environment considerably influences the blood meal size. In this work, we performed a comparative study of the level of anticoagulant activity in the saliva and crop contents of three triatomine species – Triatoma infestans, Triatoma brasiliensis and Rhodnius prolixus – and correlated this with their feeding performance on live hosts. Moreover, the feeding parameters on a large diameter vessel influenced by the crop anticoagulants were evaluated in detail. The anticoagulant activity was significantly higher in the crop contents than in salivary glands, varying from 1.6-fold higher for R. prolixus to 70-fold higher for T. brasiliensis. Amongst the species, T. brasiliensis had the lowest crop anticoagulant activity, the lowest concentration of thrombin inhibitor, and took the longest to feed. Triatoma brasiliensis nymphs that had their intestinal anticoagulant (brasiliensin) knocked down by RNA interference had the lowest capacity to maintain cibarial pump frequency at higher levels throughout the feeding process and consequently a lower ingestion rate (mg/min), even when fed under favourable conditions (large diameter vessel). However, the feeding difficulty for brasiliensin knockdown T. brasiliensis nymphs was reversed by treating the host mice with heparin (a potent systemic anticoagulant) before blood feeding. The results indicate that crop anticoagulant activity influences modulation of the blood-pumping frequency to the intestine and significantly affects the feeding efficiency of triatomine spp. on live hosts.

Feeding Performance ofTriatoma brasiliensis(Hemiptera: Reduviidae) on Habitual Hosts:Thrichomys laurentius(Rodentia: Echimyidae) and Humans

Triatoma brasiliensis is an important vector of Trypanosoma cruzi in the semiarid zones of Northeastern Brazil where it is commonly found in human dwellings, peridomiciliary areas, and the wild environments. To better understand features that may be influencing the interaction of T. brasiliensis with its hosts, in the present work we compared the feeding performance of sylvatic and domestic populations of T. brasiliensis on humans and on Thrichomys laurentius, a wild rodent found in the natural ecotopes of T. brasiliensis. The analyses were performed using the electronic monitoring of the cibarial pump. The blood sources considerably affected the feeding parameters. Insects fed on T. laurentius had lower ingestion rate, gained less weight, had longer interruption time, remained less time in contact with the host, and had the contractions of the pump in a lower frequency during the feeding process in comparison to insects fed on humans. These results indicate that humans are better blood sources for T. brasiliensis than T. laurentius. The fact that there was no difference between the feeding performance of sylvatic and domestic insects suggests that feeding is not a barrier for the domiciliation. Together, these findings suggest that shelter and faster blood meals are qualities of human dwellings that facilitate the domiciliation of T. brasiliensis.

Feeding behavior of Triatoma vitticeps (Reduviidae: Triatominae) in the state of Minas Gerais, Brazil

The objective of this study was to evaluate the feeding behavior of Triatoma vitticeps through the identification of its food sources and the characterization of the blood ingestion process. In addition, we aimed to verify if the saliva of this vector interferes with the perception of the host during the feedings by creating a nervous impulse. Here, we demonstrated that the T. vitticeps saliva reduces, gradually and irreversibly, the amplitude of the compound action potential of the nervous fibre, which helps decrease the perception of the insect by the host. The precipitin reaction demonstrated the feeding eclecticism of this vector, with the identification of eight food sources - most of them found simultaneously in the same insect. The analysis of the electrical signals produced by the cibarial pump during meals demonstrated that the best feeding performance of T. vitticeps nymphs that fed on pigeons is mainly due to the higher contraction frequency of the pump. The longer contact period with the host to obtain a complete meal compared with other triatominae species of the same instar could favor the occurrence of multiple blood sources in T. vitticeps under natural conditions, as it was evidenced by the precipitin test.

The Pollination of Mid Mesozoic Seed Plants and the Early History of Long-proboscid Insects1,2,3

Abstract A defining event for Mesozoic plant-pollinator interactions is the angiosperm radiation, which extended the reach of pollinating insects during the Early Cretaceous in a brief interval of geologic time. Recent evidence indicates that events beginning in the Early Permian and increasing during the Middle Jurassic provided repeated opportunities for insect feeding on pollen, pollination drops, and reproductive tissues of extinct gymnosperm lineages. Pollination was an associated development. Studies on the detailed mouthpart structure of several fluid-feeding insect lineages indicate targeting of certain tubular features of gymnosperm ovulate organs that previously were considered anomalous and difficult to interpret. One mouthpart type, the long-proboscid condition, consists of elongate, tubular (siphonate) proboscises that accessed surface fluids powered by a cibarial pump, often assisted with a distal proboscis sponging organ. These proboscises were received in ovulate organs through often intri...

Sharpshooter X Wave: Correlation of an Electrical Penetration Graph Waveform with Xylem Penetration Supports a Hypothesized Mechanism forXylella fastidiosaInoculation

AbstractElectrical penetration graph (EPG) monitoring is the most rigorous means of observation and quantification of feeding by piercing-sucking arthropods. Previous EPG studies with aphids and leafhoppers have demonstrated that the X wave identifies when the stylets of these phloem fluid-ingesting insects make contact with their preferred plant vascular cell, phloem sieve elements. This article presents the first direct evidence of an X wave identifying ingestion from a xylem tracheary element by a xylem fluid-ingesting type of leafhopper Homalodisca liturata Ball (Hemiptera: Cicadellidae: Cicadellinae), whose waveforms are nearly identical to those of the glassy-winged sharpshooter, Homalodisca vitripennis (Germar). We document consistent association of the sharpshooter X wave with salivary sheath termini in xylem, especially lignified secondary xylem cells, and absence of the X wave in the rare instance of ingestion from a nonxylem cell. The sharpshooter X wave is a complex, multicomponent waveform, composed of X wave-specific variants of waveform subtypes B1w (representing salivation), B1s (representing precibarial valve movement for tasting), types C1 (a new waveform type that may represent egestion) and C2 (a new designation for the waveform type representing ingestion/cibarial pumping). It is proposed that the sharpshooter X wave represents a blended suite of behaviors that function to 1) physically seal stylet tips into the cell via sheath salivation, 2) repeatedly taste then eject (egest) chemical constituents of the cell to determine acceptability, and 3) mechanically test the strength of the stylet seal via trial cibarial pumping (ingestion). It is further hypothesized that the X wave represents vector behaviors that control inoculation of the Pierce's disease bacterium, Xylella fastidiosa. The ingestion-(salivation and egestion) hypothesis is stated for the mechanism of transmission of X. fastidiosa.

The feeding process of Cimex lectularius (Linnaeus 1758) and Cimex hemipterus (Fabricius 1803) on different bloodmeal sources

The bedbugs Cimex lectularius and Cimex hemipterus are obligate hematophages in all their nymphal instars as well as in the adult stage. The efficiency with which the insects obtain blood from their hosts is directly related to their population dynamics. In the present study we compared the feeding process and salivary content in individuals of these two species when fed on different blood sources or host sites, using a cibarial pump electromyogram. Females ingested more blood than males but needed longer contact time with the host to complete the meal. The bedbug C. lectularius was more efficient than C. hemipterus in obtaining blood from mice and pigeons. With regard to the feeding site on mice, it was easier for the insects to obtain blood from the skin of the belly than that of the back. Individuals of C. hemipterus were able to maintain the cibarial pump functioning at higher frequencies for longer periods when fed on pigeons treated with anticoagulant. Although saliva from C. lectularius contained more hemeproteins and showed more anti-clotting activity its total protein content was similar to that of C. hemipterus. Overall, C. lectularius obtains a bloodmeal more efficiently from its hosts, which may have enabled this species to reach higher levels of infestation than C. hemipterus.

Telmatogeton murrayi sp. n. from Iceland and T. japonicus Tokunaga from Madeira (Diptera: Chironomidae)

Male and female imagines of Telmatogeton murrayi sp. n. from Iceland are described and male and female imagines of T. japonicus Tokunaga from Madeira redescribed. Telmatogeton murrayi differs from T. japonicus by its much larger size, higher chaetotaxy, in details of the cibarial pump, and in having the posterior widening of the female spermathecal duct nearly as wide as the anterior widening. The number of setae on the pedicel indicates that the European T. japonicus not may be conspecific with the Pacific T. japonicus. The effects of environmental induced size variation on morphological differences and meristic data are discussed.