The larval head structures of Podagricomela shirahatai (Chûjô) (Chrysomelidae,Galerucinae, Alticini) and morphological effects of leaf mining

Head structures of the leaf mining larva of the chrysomelid species Podagricomela shirahatai are described and illustrated. Internal and external structures were reconstructed three dimensionally based on image stacks obtained with microcomputed tomography. The larval head is characterized by prognathism, a dorsoventrally compressed shape, a flattened maxillolabial complex, a completely reduced coronal suture, and the presence of a deep, V‐shaped posterior emargination of the head capsule. Internal structures are not distinctly affected by leaf mining. The cephalic features are compared with conditions found in surface feeding and root feeding alticine larvae and also with characters of chrysomeline larvae of Chrysomela populi Linnaeus. Possible correlations between modifications of the larval head and different feeding behaviors are discussed. Characters are also discussed with respect to possible phylogenetic implications. Some derived features are apparently due to phylogenetic constraints. Apomorphies characterizing alticine larvae with distinctly different life habits are the loss of M. frontoepharyngalis (M. 9), the origin of M. tentoriostipitalis (M. 18) from the head capsule, two insertions of M. tentoriopraementalis inferior (M. 29) and the reduction of stemmata. The study underlines that the anatomical study of chrysomeloid larvae is not only highly desirable in a phylogenetic context, but also crucial for understanding the evolution of different life strategies in this extremely successful group of Coleoptera. J. Morphol. 276:446–457, 2015. © 2014 Wiley Periodicals, Inc.     

The role of an epipharyngeal sensillum in the perception of feeding deterrents by Leptinotarsa decemlineata larvae

An epipharyngeal taste sensillum in Leptinotarsa decemlineata larvae was studied. Electron microscopy showed that the sensillum is innervated by five neurons. Electrophysiological experiments showed that one of these cells responds to water, a second to sucrose and a third to two feeding deterrents that were also effective in a behavioural test. Receptor cells sensitive to feeding deterrents were not previously reported for L.␣decemlineata larvae or adults. The response of the sucrose-sensitive cell was strongly inhibited by one of the two feeding deterrents and only slightly by the other feeding deterrent. The relationship between the behavioural and electrophysiological results is discussed in order to elucidate the neural code of feeding deterrents in L. decemlineata larvae. We conclude that probably both the response of the deterrent cell and peripheral interactions exerted by feeding deterrents on the sucrose-sensitive cell determine the potency of feeding deterrents. The present results provide a physiological basis for the hypothesis that the presence or absence of feeding deterrents in potential food plants is a decisive cue in food plant selection by L. decemlineata larvae. Accepted: 25 March 1998     

Identification of larvae of European species of Chironomus Meigen (Diptera: Chironomidae) by morphological characters

Abstract Variation in the morphology of larvae of twenty-six cytologically identified species of Chironomus Meigen from Europe is described. Characters studied include tubules on posterior abdominal segments, head pigmentation, mental size and teeth, mandibular and epipharyngeal teeth and antennae. A key to the species, based on these characters and ventromental plate features, is presented.     

Feeding ecology of a group of buffy‐headed marmosets (Callithrix flaviceps): fungi as a preferred resource

Mycophagy is a relatively rare behavior in primates and has only been recorded in five callitrichid species. Here, we present data on the feeding ecology of a free‐ranging group of Callithrix flaviceps, which was studied in the Augusto Ruschi Biological Reserve, Southeastern Brazil, in 2008. In contrast with other marmosets, which are typically gummivorous, the study group was predominantly mycophagous–insectivorous, with fungi corresponding to 64.8% of total feeding records, and gum (6.1%) and fruit (3.3%) together providing only a minor part of the diet. Prey corresponded to 25.8% of the group's diet. The fungi (Mycocitrus spp.) consumed by the marmosets were found attached to the stems of Merostachys bamboo. As the animal component of the group's diet was similar to that recorded in studies of other marmosets, we propose that fungi were exploited primarily as a substitute for plant material, in particular exudates. This highly mycophagous diet may be determined by two principal factors: (1) the abundance of fungi within the study area, and (2) the avoidance of bark gouging, for which C. flaviceps may be less specialized than most other marmosets. These conclusions are supported by comparisons with other marmoset groups, which indicate an ecological specialization for mycophagy in C. flaviceps, and that the species will resort to gummivory in habitats where fungi are scarce. Am. J. Primatol. 72:515–521, 2010. © 2010 Wiley‐Liss, Inc.     

Molecular identification of dipteran pests (Diptera: Sciaroidea) from shiitake mushroom

On shiitake farms, mycophagous maggots can cause serious damage by preventing formation of the fruiting body. Recently, these pests have significantly reduced shiitake production in Korea. However, larvae and female adults cannot be identified due to their lack of morphological characteristics. Therefore, farmers and applied entomologists are unable to determine which species is the primary cause of the shiitake damage. In this study, mycophagous flies (colonized larvae) were collected from damaged shiitake farms and subsequently identified by matching identified males with the cytochrome c oxidase subunit I (COI) sequences from the larvae. Divergences of the COI sequences among the species discriminated the clusters clearly, and the mycophagous pests were identified as Camptomyia corticalis and C. heterobia. Interestingly, these two species coexisted under the bark of shiitake oak bed logs.     

Juvenile hormone modulation of insect cold hardening: Ice-nucleating activity

Data are presented offering the first evidence for probable endocrine involvement in the control of cold hardening in Eurosta solidaginis. Juvenile hormone (JH) deprivation experiments in which the corpora allata were removed by head ligation resulted in a loss of supercooling (SC) capacity in larvae collected over 2 years. This loss of supercooling capacity is indicative of synthesis of organismal pools of ice-nucleating agents (INA). Larval sensitivity to JH removal (ligation) on SC is seasonally dependent. For example, in 1983, larvae were most sensitive in October, secondarily so in September, and relatively insensitive in December regardless of acclimation temperature. While in 1984, larvae acclimated to +5 °C were most sensitive in November, secondarily so in October, and relatively insensitive in December, and larvae acclimated to +15 °C were most sensitive in October and December and least so in November. Supercooling point elevations as great as 7 °C over controls were observed with maximal responses occurring within 1 day following ligation. In 1983, juvenile hormone replacement following ligation generally resulted in an expansion of supercooling capacity when compared to controls. As with ligation, the sensitivity to JH replacement on SC was seasonally dependent: September and October larvae being the most sensitive with December larvae being insensitive. Larvae collected in 1984 were given a greater dose of JH than those in the previous year and showed no significant change in SC over ligated-acetone controls. Hormone analog potentiation experiments in which unligated larvae collected in 1983 and acclimated to +5 °C were given methoprene resulted in depression of supercooling points for September and October larvae. JH titres appear to play an important role in the regulation of SC capacity in E. solidaginis larvae.     

Soil burrowing Muscina angustifrons (Diptera: Muscidae) larvae excrete spores capable of forming mycorrhizae underground

Muscina angustifrons (Diptera: Muscidae) is a mycophagous species that exploits a variety of fungi, including ectomycorrhizal fungi. Larvae of this species have been shown to feed on sporocarps (including spores), and full-grown larvae leave sporocarps and pupate 0–6?cm below the soil surface. In this study, we examined whether M. angustifrons larvae are capable of transporting ectomycorrhizal fungal spores and enhancing ectomycorrhiza growth on host-plant roots. Full-grown larvae were found to move horizontally 10–20?cm from their feeding sites and burrow underground. These wandering larvae retained ectomycorrhizal fungal spores in their intestines, which were excreted following relocation to underground pupation sites. Excreted spores retained germination and infection capacities to form ectomycorrhiza on host-plant roots. In the infection experiments, ectomycorrhizal fungal spores applied in the vicinity of underground host-plant roots were more effective in forming ectomycorrhiza than those applied to the ground surface, suggesting that belowground transportation of spores by M. angustifrons larvae could enhance ectomycorrhizal formation. These results suggested that M. angustifrons larvae act as a short-distance spore transporter of ectomycorrhizal fungi.     

Development of feeding structures in larval fish with different life histories: winter flounder and Atlantic cod

The size at which feeding structures developed and shifts in head proportions occurred, differed between Atlantic cod Gadus morhua and winter flounder Pseudopleuronectes americanus. The sequence and timing of the development of feeding structures may not be dependent on size, but may occur because they are necessary to meet specific requirements offish larvae feeding in the plankton. In early larval stages development of feeding structures was similar in number and type and was necessary for first-feeding in both species. In later stages, significant differences between species occurred in the timing of the development of feeding structures. In cod differentiation of new structures and changes in head proportions occurred at about two-thirds of the way through larval life, which coincided with an increase in growth. In flounder changes in feeding morphology did not occur during the symmetrical larval stage, but occurred only after metamorphosis to the asymmetrical demersal juvenile stage. Differences between cod and flounder in the size at which feeding structures develop may reflect life history adaptations expressed in the duration of the pelagic larval stage, as well as differences in juvenile habitat and feeding ecology.     

Precibarial and cibarial chemosensilla in the whitefly,Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae)

The internal anatomy of the anterior alimentary canal of the whitefly, Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae) B-biotype, was examined by light, scanning, and transmission electron microscopy to elucidate the location and number of precibarial and cibarial gustatory sensilla. Elucidation of the epipharyngeal organ complex within the precibarium revealed 10 precibarial sensilla located proximal to where the paired maxillary stylets diverge on their retraction. The sensory organ complex within the cibarium consists of 8 sensilla, 6 on the epipharyngeal sclerite with 2 found within the hypopharyngeal sclerite. Fine structure investigation revealed the individual neurons to terminate at sensillar pores, which allow direct contact with passing fluids, thus supporting a chemosensory function. Ultrastructure of the neurons is similar to that of precibarial and cibarial gustatory chemosensilla found in other piercing-sucking insects. Their importance to whitefly feeding is discussed.     

EVOLUTIONARY LOSS OF LARVAL FEEDING: DEVELOPMENT,FORM AND FUNCTION IN A FACULTATIVELY FEEDING LARVA,BRISASTER LATIFRONS

Species with large eggs and nonfeeding larvae have evolved many times from ancestors with smaller eggs and feeding larvae in numerous groups of aquatic invertebrates and amphibians. This change in reproductive allocation and larval form is often accompanied by dramatic changes in development. Little is known of this transformation because the intermediate form (a facultatively feeding larva) is rare. Knowledge of facultatively feeding larvae may help explain the conditions under which nonfeeding larvae evolve. Two hypotheses concerning the evolutionary loss of larval feeding are as follows: (1) large eggs evolve before modifications in larval development, and (2) the intermediate form (facultatively feeding larva) is evolutionarily short-lived. I show that larvae of a heart urchin, Brisaster latifrons, are capable of feeding but do not require food to complete larval development. Food for larvae appears to have little effect on larval growth and development. The development, form, and suspension feeding mechanism of these larvae are similar to those of obligate-feeding larvae of other echinoids. Feeding rates of Brisaster larvae are similar to cooccurring, obligate-feeding echinoid larvae but are low relative to the large size of Brisaster larvae. The comparison shows that in Brisaster large egg size, independence from larval food, and relatively low feeding rate have evolved before the heterochronies and modified developmental mechanisms common in nonfeeding echinoid larvae. If it is general, the result suggests that hypotheses concerning the origin of nonfeeding larval development should be based on ecological factors that affect natural selection for large eggs, rather than on the evolution of heterochronies and developmental novelties in particular clades. I also discuss alternative hypotheses concerning the evolutionary persistence of facultative larval feeding as a reproductive strategy. These hypotheses could be tested against a phylogenetic hypothesis.     

Influence of watering and trenching ponderosa pine on a pine sawfly

Summary Neodiprion autumnalis (Smith) larvae were caged for two successive years on root-trenched, watered, and untreated ponderosa pine (Pinus ponderosa Doug. ex Laws.) to determine effects of host moisture stress on larval feeding. Levels of moisture stress (as measured by the Scholander pressure chamber) differed significantly among treatment levels during 1984 and 1985 larval feeding periods. Differences in larval feeding success were not detected in 1984. In 1985, however, larvae on trenched (stressed) trees clipped and rejected more foliage, consumed more needles, had lower pupal weights, lower survival, and a longer feeding period than larvae on watered or untreated trees. Frass production did not differ among treatment levels. The length of the feeding period was shorter for larvae on watered trees than for larvae on untreated trees, but other measures of feeding success did not significantly differ between watered and untreated trees.     

Larval biology of butterflyfishes (Pisces,Chaetodontidae): what do we really know?

Synopsis Relatively little is known of the pelagic portion of the life history of butterflyfishes. Eggs are small (<1 mm), pelagic and hatch in less than 30 hours. Most species pass through a so-called tholichthys larval interval characterized by elaborate, distinctive head spination:Coradion larvae have different head spination. While older chaetodontid larvae can be identified by adult characters, young (preflexion) larvae generally cannot now be identified below family. In tropical plankton studies chaetodontid larvae averaged <0.1% of larvae captured, and occurred in 13% of samples. This rarity is a major hindrance to further work, but is not unexpected in view of adult abundance. Larvae of a few taxa are most abundant in shelf waters, but larvae of many chaetodontid taxa seem to be most abundant in oceanic waters. In either case, waters near reefs have the fewest chaetodontid larvae. Offshore maxima of larvae appear to exist a few kilometers seaward of Great Barrier Reef ribbon reefs. Chaetodontid larvae may prefer the upper portion of the water column. Both size and age at settlement vary widely within the family and the large genusChaetodon, and the latter varies widely within species. Average size at settlement is less than 20 mm and age is less than 40 days. No correlation was found between size and age at settlement. Behaviour and feeding of chaetodontid larvae are essentially unstudied. Chaetodontid larvae seem to be least abundant in winter. The implications of these conclusions are discussed and some suggestions for further research are made. In all areas more work is needed.     

Chemoreceptors in the preoral cavity of the tobacco hornworm, Manduca sexta, and their possible function in feeding behaviour

Feeding preference tests showed that antenna- and maxillectomized larvae of the tobacco hornworm do not lose the ability to discriminate between a host plant (tomato) and a non-host plant (canna). When the labrum was also removed, no difference in acceptability of the two plants could be found. This suggested that chemoreceptors existed on the labrum. Scanning electronmicroscope pictures of the epipharynx revealed one pair of possible chemoreceptive structures. Using electrophysiological methods, the gustatory function of these sensilla was established and some properties were studied. Activity of not more than three different sensory cells of each epipharyngeal sensillum was observed. One cell was found to be sensitive to various distasteful compounds. Saps of tomato and canna do not stimulate particular sensory cells; however, the behavioural responses towards these plants are totally different. It was suggested that these sense organs relay information on the chemical composition of the plants in certain patterns of activity. The feeding tests on different diets showed that maxillectomy reduced the feeding stimulant effect of sucrose, but not completely. No activity of a specific sugar receptor in the epipharyngeal sensilla was observed electrophysiologically. However, the response of two cells to salt was inhibited in the presence of sugar.The epipharyngeal chemoreceptors function as a second checkpoint after the maxillary chemoreceptors for feeding deterrent compounds. It is more likely that the actual food intake in maxillectomized larvae is controlled by the presence of feeding deterrents than by the presence of feeding stimulants.

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Zusammenfassung Wie Wirtswahlversuche zeigten, verlieren Larven des Tabakhornwurms nach Antennen- und Maxillenentfernung nicht die Fähigkeit zwischen einer Wirtspflanze (Tomate) und einer Nichtwirtspflanze (Canna) zu unterscheiden. Wenn jedoch auch das Labrum entfernt wurde, konnte kein Unterschied mehr in der Bevorzugung einer der beiden Pflanzen gefunden werden. Dies läßt Chemorezeptoren im Labrum vermuten. Raster-elektronenmikroskopische Bilder des Epipharynx ließen ein Paar von Rezeptoren mit möglicherweise chemorezeptiven Strukturen erkennen. Mit elektrophysiologischen Methoden wurde die Geschmacksfunktion dieser Sensillen nachgewiesen und es wurden einige Eigenschaften untersucht. Dabei wurde die Aktivität von nicht mehr als drei verschiedenen Sinneszellen in jedem Epipharynxsensillum festgestellt. Eine Zelle erwies sich als empfindlich für verschiedene, geschmacklich unangenehme Verbindungen. Saft von Tomaten und Canna stimulierten keine besonderen Sinneszellen; dennoch sind die Verhaltensreaktionen gegenüber diesen Pflanzen völlig verschieden. Es wird vermutet, daß dieses Sinnesorgan Information über die chemische Zusammensetzung der Pflanzen in bestimmten Aktivitätsmustern übermittelt. Die Fraßtests an verschiedenen Diäten zeigten, daß Maxillenektomie die fraßstimulierende Wirkung von Saccharose vermindert, allerdings nicht vollständig. Elektrophysiologisch wurde keine Aktivität eines spezifischen epipharyngalen Zuckerrezeptors in den epipharyngalen Sensillen festgestellt. Jedoch wurde die Reaktion von zwei Zellen auf Salz in Gegenwart von Zucker gehemmt.Die epipharyngalen Chemorezeptoren arbeiten als eine zweite Kontrollstelle nach den maxillaren Chemorezeptoren für fraßabschreckende Verbindungen. Es ist wahrscheinlich, daß die tatsächliche Nahrungsaufnahme bei Larven nach Maxillenentfernung eher durch die Gegenwart von Fraßdeterrentien gesteuert wird als durch die Gegenwart von Fraßstimulantien.

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The electrophysiological part of this study was carried out by G. de B. during a stay at the Dept. of Biology, Princeton University.     

Feeding resumption, morphological changes and mortality during starvation in Japanese flounder larvae

Japanese flounder Paralichthys olivaceus larvae established first feeding 3 days after hatching (DAH) at c . 17° C. Non-fed fish reached irreversible starvation at age 5 DAH. Non-fed fish showed similar feeding rate and feeding intensity as the fed fish when they were provided with prey before 5 DAH, after which the starved larvae did not feed even when prey became available. None of the six morphological measurements examined (total length, body height, eye height, head height, gut height and myotome height) showed significant differences between the non-fed and fed larvae until 5 DAH. Normal development continued only in the fed group, and the non-fed larvae showed reverse growth or body collapse after 5 DAH. Owing to the shrinkage and collapse at the top of head due to starvation, head height could be a sensitive indicator of starvation in Japanese flounder larvae. In the fed treatments, high mortality occurred from first feeding (3 DAH) to irreversible starvation (5 DAH), accounting for about two-thirds to three-quarters of the overall mortality (46–52%) throughout the experiments. This mortality was not prey density or larval density dependent. Mortality during the same period in the non-fed larvae accounted for about a third of the overall mortality (100%).     

Re‐establishment of biting mouthparts in desert‐living dung beetles (Scarabaeidae: Scarabaeinae) feeding on plant litter—Old structures reacquired or new ones evolved?

Evolution of mouthparts in adult dung beetles (Scarabaeidae: Scarabaeinae) for eating moist, fresh dung was linked with a loss of the ability to chew. However, the desert‐living genus Pachysoma, probably evolved from a wet‐dung feeding, Scarabaeus‐like ancestor, has switched to a diet of dry fecal pellets (of rodents or small ruminants) and plant litter that requires re‐establishment of chewing. Indeed, gut contents of a litter‐feeding Pachysoma species indicate efficient food comminution. Based on scanning electron microscopy, cutting and grinding mouthpart structures in six Pachysoma species, of two lineages and with different food preferences, are described and compared with homologous structures in wet‐dung feeding Scarabaeus species. In Pachysoma, cutting and breaking of large food items is probably performed by a clypeal scraper, a prominent epipharyngeal tooth and large maxillary galeal hooks. Further comminution is achieved by a large, grinding area evolved on the mandibular molae. Interspecific differences and the probable function and evolution of these structures are discussed. Particularly, the unique tools for cutting/breaking are completely novel structures and not results of some reacquisition of normal biting mouthparts. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Condition and size of damselflies: a field study of food limitation

Summary Based on evidence from field manipulations, several authors have recently suggested that interference competition among larval odonates reduces individual growth rates and biomass by reducing foraging rates. This study was designed to test the effects of food shortage on condition (relative mass per unit head width) of larval Ischnura verticalis (Odonata: Coenagrionidae) under laboratory conditions and to use these results to estimate the degree of food shortage of larvae under naturally occurring field conditions. In the laboratory, there were marked differences in condition of larvae fed diets ranging from ad libitum feeding with worms to ad libitum feeding with Daphnia 1 day out of every 8. Condition of larvae collected from May through October from 17 different sites in southern Ontario indicated that, for most of the year, larvae had conditions similar to those fed ad libitum with Daphnia in the laboratory. There was no evidence that larval condition was related to population density. Condition of larvae in most sites during July was similar to that of larvae fed poor diets in the laboratory. It is unlikely that the low conditions were due to competition as there were no correlations with density across sites and population densities during July were at their lowest. Adult head widths showed a seasonal decline from mid June to the end of the flight season. There was no evidence that head widths were related to population density although there was some evidence that head widths of males were positively related to larval condition. My results do not support the hypothesis that competition is important in affecting foraging rates and subsequent development of larvae. Contrasts between my results and other studies may stem from difficulties with the interpretation of field experiments, that densities in my study may have been low due to fish predation, and/or that I. verticalis larvae are slow moving relative to other larvae and thus less likely to interact.     

Pathobiology of amber disease,caused by Serratia Spp., in the New Zealand grass grub,Costelytra zealandica

Amber disease in the New Zealand grass grub (Costelytra zealandica) is caused by some strains of Serratia entomophila or Serratia proteamaculans (Enterobacteriaceae). When treated with pathogenic isolates, larvae ceased feeding within 48 h, developed an amber coloration after 72 h, and entered a long chronic phase without feeding. An acute dose of 2-4 x 10(4) pathogenic bacteria was sufficient to produce disease in 50% of treated larvae. Time to death was directly related to temperature. At 15 degrees C, infected larvae remained in a chronic, nonfeeding state for more than 4 months prior to death. Nonpathogenic isolates, lacking the disease-causing plasmid (pADAP), had no effect on either feeding or disease. Twenty-four hours after ingestion, bacteria were found predominantly in the hindgut and growth occurred primarily within the fermentation chamber and in the head section of the larvae. Nonpathogenic strains did not multiply in treated larvae. Treatment of diseased larvae with antibiotic eliminated Serratia cells from the insects but did not result in restoration of feeding or the dark gut characteristic of the healthy larva.     

Biology of Campoletis chlorideae (Uchida) (Hym., Ichneumonidae) developing in Bt-treated, Bt-resistant Helicoverpa armigera (Hübner) (Lep., Noctuidae) larvae

Abstract:  Life history parameters in two generations of endoparasitoid Campoletis chlorideae (Uchida) were examined using Bacillus thuringiensis (Bt)-resistant Helicoverpa armigera (Hübner) larvae feeding on B. thuringiensis toxin Cry1Ac. In the laboratory, Bt toxin was fed to Bt-resistant host larvae continuously in case of Bt treatment and only before or after the host larvae were parasitized in Bt–P and P–Bt treatments, respectively. C. chlorideae pupae developed faster in Bt treatment than non-Bt treatment. The shortened pupal stage duration was mainly because of the feeding of host larvae on Bt-diet before being parasitized. Body length of adult male C. chlorideae developed inside Bt-treated Bt-resistant (Bt–Bt) H. armigera larvae significantly decreased, especially in host larvae feeding on Bt-diet after being parasitized. However, survival, pupal mortality and adult longevity of C. chlorideae were almost unaffected in Bt-resistant H. armigera larvae feeding on Bt-toxin. Furthermore, Bt-treated host larvae had the same effect on the F₁ progeny of C. chlorideae as the previous generation, and there was no significant difference between generations. This experiment suggests that there is very limited effect on the life history parameters in two generations of C. chlorideae parasitizing Bt–Bt H. armigera larvae. But both generations of C. chlorideae are affected when Bt-resistant H. armigera larvae fed on Bt toxin for different durations.     

Larval morphology enhances phylogenetic reconstruction in Cetoniidae (Coleoptera: Scarabaeoidea) and allows the interpretation of the evolution of larval feeding habits

Abstract.  The Cetoniidae, the showiest of scarabs, comprises some 3900 species in 515 genera, distributed worldwide except for subpolar areas and some offshore New Zealand islands. Parsimony analysis of 76 larval and adult characters and 42 terminal taxa supports the monophyly of Cetoniidae (sensu Krikken, 1984 ), but not of the traditionally considered subfamilies Cetoniinae and Trichiinae (sensu Krikken, 1984 ). In the study taxon, larval characters are shown to be more informative than those of adults for deeper phylogeny. The evolution of some larval characters (head and legs) in relation to feeding habits is discussed on the basis of phylogenetic analysis. The results show an evolutionary shift from wood associations to a higher ecological plasticity that allows the larvae to feed on wide sources of organic matter (including compost, dung, gopher burrows, packrat middens, ant debris piles, etc.).