Seasonal variation in larval host plants and consequences for Heliconius erato (Lepidoptera: Nymphalidae) adult body size

Abstract Adult body size, a key life history component, varies strongly within and between Heliconius erato phyllis (Lepidoptera: Nymphalidae) populations. In the present study, we determined whether seasonal variation in adult body size is temperature related and/or determined by seasonal changes of host plants (Passifloraceae) used by the larval stage. A population of H. erato phyllis located in a Eucalyptus plantation (Barba Negra Forest, Barra do Ribeiro County, Rio Grande do Sul State, Brazil) was sampled every 45 days from March 1997 to October 1998 to quantify seasonal variation in adult body size and use of larval host plants. In the laboratory, the effects of the following factors on adult body size were quantified: (i) host plant species (Passiflora misera or Passiflora suberosa); (ii) food quantity consumed by larvae (experimentally manipulated for each passion vine species); (iii) winter and summer temperatures (15 and 25°C, respectively); and (iv) the interaction between host plant species and temperature. Adults emerging during summer were larger than those emerging in other seasons. Female butterflies oviposited selectively on P. misera even when the dominant passion vine was P. suberosa. They only switched from using P. misera to P. suberosa during later autumn and winter, when P. misera vines were completely defoliated. The laboratory feeding trials with both passion vines showed a strong positive association between food quantity consumed by larvae and adult size. They also confirmed that adults are larger when their larvae are reared on P. misera than on P. suberosa. Temperature during larval development had no effect on H. erato phyllis adult size. Thus, seasonal variation of H. erato phyllis adult size in a given place is primarily determined by the availability and quality of host plant species used by the larval stage.     

Absence of Learning and Local Specialization on Host Plant Selection by <Emphasis Type="BoldItalic">Heliconius erato</Emphasis>

There is considerable interspecific variation in larval host plants (Passifloraceae) used among Heliconius erato (Lepidoptera: Nymphalidae) populations. This study evaluates the role of learning and the influence of interspecific variation in host plant attributes on such local specialization in H. erato host plant choices. Experiments were conducted under laboratory, insectary, and field conditions, with the two most widely used host plants in southern Brazil (Passiflora suberosa Linnaeus and Passiflora misera Humbold, Bonpland et Kunth). Larval feeding preference and induction were evaluated through choice tests for all instars. Oviposition was evaluated in relation to host plant preference, Hopkins host-selection principle, and conditioning time (for 3, 7, 11, or 15 days). Also, oviposition choice was tested regarding density, intemode length, and presence of intact terminal bud on P. suberosa and P. misera shoots. Both larvae and adults of H. erato phyllis showed preference for P. misera compared to P. suberosa, under all conditions. Larval feeding preference could not be induced for most instars. The Hopkins effect was not detected and oviposition choice could not be conditioned. Females alternated use of host plant species as a function of variation in either density or presence of terminal buds on shoots. Thus, our data indicate host plant preference in H. erato phyllis is not learned but innate. Therefore, we concluded that variation in local use of host plant by this butterfly in southern Brazil results from qualitative and quantitative variation of the passion vine species.     

Oviposition in <Emphasis Type="Italic">Heliconius erato</Emphasis> (Lepidoptera,Nymphalidae): how Essential Is Drumming Behavior for Host-Plant Selection?

Quantitative studies on the behavioral events involved in oviposition by phytophagous insects are scant. One of these events is drumming behavior, a rapid extension of forelegs after landing on a host, which remains largely under studied. In this study, quali- and quantitative analyses of Heliconius erato phyllis (Lepidoptera: Nymphalidae) oviposition events and drumming behavior in relation to both preferred (Passiflora misera and P. suberosa) and non-preferred (P. caerulea and P. alata) hosts were performed. For the first time, drumming behavior is described by images made during the butterfly oviposition process. The same set of females was assigned to both single- and multiple-choice tests under insectary conditions, and their behaviors toward each host were recorded on video. The resulting images were analyzed frame by frame. Both frequency and duration of behavioral events related to oviposition varied on preferred vs non-preferred hosts. On preferred hosts, behaviors that were associated with egg deposition (inspecting flight, drumming and abdomen touching) were pronounced; subsequently, oviposition occurred within a few seconds. On non-preferred host plants, behaviors that did not predict oviposition (flying and resting) predominated, and oviposition on these hosts was negligible. Drumming occurred on all plants and resulted in a faster decision-making process on high-quality hosts (prompt acceptance) as well as on lethal hosts (prompt rejection) compared to the host that confers poor larval performance (delayed rejection). Thus, drumming is crucial for decision-making related in host-plant selection by H. erato phyllis, and influences other behaviors involved in the oviposition process.     

Isolation and characterization of microsatellite markers from the sweet passion fruit (Passiflora alata Curtis: Passifloraceae)

Passiflora alata is one of the commercialized species of the passion vines, grown for its edible fruits. It is found in South America, mainly Paraguay, Argentina, Peru and Brazil. The development of a set of microsatellite markers was proposed to provide tools for analysing both genetic structure of wild populations and the reproductive system of the species, which are poorly understood. Seven markers were developed from a P. alata‐genomic library enriched for simple sequence repeats (SSR). Twelve individuals were genotyped, and the levels of the expected heterozygosity (H_E) did show that those markers could be used to develop P. alata genetic studies.     

The adult body size variation of Dryas iulia (Lepidoptera,Nymphalidae, Heliconiinae) in different populations is more influenced by temperature variation than by host‐plant availability during the seasons

The body size of insects is affected by environmental conditions during development and can present considerable intraspecific variations, which can be seen as an ultimate consequence/adaptation to environmental conditions. This paper evaluated whether the body size of the butterfly Dryas iulia from subtropical populations was influenced by changing climate conditions and food source availability during the seasons. The likely reasons behind body size variation were also investigated. First, field data on body size variation, host‐plant availability and climate fluctuation throughout the seasons were recorded. Then, the effects of host‐plant species and temperature on body size were analyzed by controlled experiments. Field data revealed that body size and host‐plant availability varied significantly through the seasons. Populations had the smallest body size during the spring and the biggest size during summer, whereas host‐plant availability was lower during winter and higher during spring. The controlled experiments revealed that both temperature and host‐plant had significant effect on the plasticity of body size. Larvae subjected to winter temperature treatment led to smaller butterflies when compared to immatures reared under summer temperature treatment, and larvae fed with Passiflora misera produced bigger adults when compared to larvae reared on Passiflora suberosa. The combination of data gathered in the field and in the laboratory suggests that seasonal body size variation in D. iulia is related mainly to differences in the temperatures to which larvae are subjected during development, while host‐plant shifts caused by differential availability of food through the seasons had slightly effects on the variation observed.     

Ultrastructure of the corona of scented and scentless flowers of Passiflora spp. (Passifloraceae)

To know thoroughly the structure and function of the corona of Passiflora, the anatomy and ultrastructure of two species were analysed in relation to the emission of odour perceivable by humans: a scented one, P. caerulea L., and a scentless one, P. suberosa L.Both species exhibited secretory tissue, whose cells were characterised by dense cytoplasm, numerous mitochondria and vacuoles. Evidence of granulocrine secretion was detected. Nevertheless, there were differences concerning some cytological structures: P. suberosa lacked smooth endoplasmic reticulum (sER) and starch but had large and many lipidic globules, while P. caerulea had few dictyosomes, scarce lipidic content, a greater proportion of sER/rough endoplasmic reticulum (rER) and amyloplasts. The cellular features of P. caerulea correspond with those of fragrance tissues. The secretion appearance and quantity were also different between both species: P. caerulea exhibited sparse drops on the cuticle in contrast to P. suberosa, which secrets a wax-like material. If this is the final product of the secretory process or just a vehicle that contributes to the emission of volatile compounds, as occurs in certain osmophores, needs further confirmation with chemical analysis.Results are discussed in the context of the pollination syndromes of each species and their florivores.     

Patterns of herbivory on Passiflora leaf tissues and species by generalized and specialized feeding insects

Abstract.

• 1 The feeding patterns of specialized (Heliconius butterflies) and generalized (the grasshopper, Osmilia flavolineata) herbivores on leaf age classes and species of neotropical vines in the genus Passiflora are compared by offering them leaf discs in choice experiments.
• 2 Heliconius larvae selected young meristematic leaves over medium aged and mature leaves of Passiflora auriculata on which to feed.
• 3 Adults of O. flavolineata consumed more young leaves of P. pittieri from an array of different ages of leaf tissue, but showed no preference for young over mature leaves of P. tetrastylus or P. data.
• 4 In choice tests conducted with medium aged leaves of four species of Passiflora, O. flavolineata selected P. tetrastylus over P. pittieri and P. vitifolia, which were consumed more than P. alata.
• 5 P. alata, which was avoided by O. flavolineata in feeding trials, was the only abundant Passiflora in neotropical habitats where generalized herbivores (grasshoppers and katydids) were common.
• 6 Our results suggest that Passiflora species in forest habitats where some generalized herbivores are rare have fewer chemical defences, but retain physical defences (tough leaves and hooked trichomes) rendering them unpalatable to specialized herbivores like the larvae of Heliconius butterflies.

     

Diurnal activity patterns and feeding ecology of the endemic geladas (Theropithecus gelada) in the Simien Mountains National Park,Ethiopia

Diurnal activity patterns and feeding ecology of geladas (Theropithecus gelada) in Chenek, Simien Mountains National Park, were studied from January 2010 to April 2010. Data were gathered for 7 days per month starting from early in the morning to dusk. The study site (1805 ha) was divided into five blocks and a total of 397 individuals were counted using simultaneous census method. One band with 21 individuals was selected as a study group and scan sampling was used to study their diurnal activity patterns and feeding ecology. Geladas dedicated significantly more time to foraging than to other diurnal activities. They spent the majority of feeding time foraging on grass, and allocated little time to feeding on leaves, shoots, and flowers. Geladas of all age classes devoted the least time to resting, and spent similar amount of time moving. The study band was observed to interact with people, and to steal food from tourists and local people. Raising local people's awareness of geladas ecology and behavior might be a necessary component in the species' successful management and conservation.     

Composition of the floral nectar of different subgenera of Argentinian Passiflora species

The composition of the floral nectar sugars and amino acids of four species of Passiflora (P. foetida, P. caerulea, P. suberosa, and P. misera) included in different infrageneric taxa and with distinct pollination mechanisms has been studied. The effect of weather and floral age on nectar volume, existence, and total and relative amounts of the various compounds was explored. The proportion of sugars was rather constant within a given species whereas the composition, number, and total quantity of amino acids showed great intraspecific and intra-plant variability; these nectar properties were independent of floral stage or meteorological conditions. Species belonging to the same subgenus displayed equivalent sugar ratios and similar total amount of amino acids, so these characteristics might be conservative in the genus. For all species, the amino acid concentration surpassed known values for their respective pollination syndromes, viz. bee and wasp-pollinated flowers. No relationship emerged between pollinators with different glossa length and nectars with distinct sugar ratios. Rather, nectar chemical composition seems to reflect taxonomic relationships.     

Feeding ecology and activity pattern of black-fronted titi monkeys (<Emphasis Type="Italic">Callicebus nigrifrons</Emphasis>) in a semideciduous tropical forest of southern Brazil

Most aspects of the ecology and behavior of Callicebus nigrifrons are still unknown. The information available about this species is mainly based on a few studies that also focused on other Callicebus. We examined the feeding behavior and activity pattern of a free-ranging pair of C. nigrifrons between March and November 2007 in an area of semideciduous tropical forest of southeastern Brazil. The study site is located at the southern limit of the Tropical Zone and is characterized by pronounced seasonality. As observed for other Callicebus monkeys, fruits were the most consumed food resource, accounting for 53% of the diet, which was complemented mainly by leaves (16%) but also by invertebrates and flowers (10% of each). A great variety of plant families (28) and species (62) were included in the diet. The titis spent 35% of their time feeding, distributing the remaining time between resting (30%) and traveling (24%). Data presented here indicate that C. nigrifrons prefer high-quality food items (fruit pulp), adding low-quality food items (such as leaves) as the availability of the higher-quality foods decreases. The amount of time spent traveling and resting did not change between seasons, but the time invested in feeding increased during the lean period. The activity pattern was not related to fruit availability, but in months with lower temperatures, monkeys spent more time feeding. We suggest that the feeding ecology and activity pattern of C. nigrifrons reflect adaptations related to annual fluctuations in food availability and temperature, respectively.     

烟粉虱天敌日本刀角瓢虫的捕食行为

研究了日本刀角瓢虫雌成虫和幼虫捕食粉虱时的各种行为时间分配.结果表明,日本刀角瓢虫成虫的捕食行为可分为爬行、取食、清洁、静息、整翅和排泄6个部分.饥饿后的日本刀角瓢虫幼虫捕食烟粉虱卵时,幼虫各龄期间的取食和爬行时间无显著差异,对烟粉虱卵的处置时间随瓢虫幼虫虫龄的增大而缩短.日本刀角瓢虫雌成虫捕食烟粉虱卵时,饥饿后的雌成虫用于取食的时间显著长于非饥饿的雌成虫,而用于清洁、静息和爬行的时间显著短于非饥饿的雌成虫;饥饿后的瓢虫雌成虫对卵的处置时间显著长于非饥饿的雌成虫.饥饿后的日本刀角瓢虫雌成虫捕食烟粉虱若虫时,瓢虫雌成虫的取食时间随烟粉虱若虫虫龄的增大而增加,清洁和静息时间随烟粉虱若虫虫龄的增大而缩短,对若虫的处置时间随烟粉虱若虫虫龄的增大而增加.     

Passiflora tacanensis, a new species of Passiflora subgenus Decaloba supersection Cieca from Mexico

Passiflora tacanensis, a new species from southern Mexico, is described and illustrated. It was discovered during the course of a taxonomic revision ofPassiflora subgenusDecaloba supersectionCieca and is assigned to that group of apetalous passion flowers.Passiflora tacanensis is known only from one fruiting herbarium specimen that was collected in southern Mexico on Volcán Tacaná in a high mesophytic forest. It is distinguished from its closest relatives,P. coriacea, P. sexocellata, P. mcvaughiana, andP. xiikzodz, by its foliose stipules, leaves that lack laminar nectaries but possess petiolar nectaries, and wide seeds with a chalazal beak that is not well developed.     

A new Brazilian Passiflora leafminer: Spinivalva gaucha,gen. n., sp. n. (Lepidoptera,Gracillariidae, Gracillariinae), the first gracillariid without a sap-feeding instar

Male, female, pupa, larva and egg of a new genus and species of Gracillariidae (Gracillariinae), Spinivalva gaucha Moreira and Vargas from southern Brazil are described and illustrated with the aid of optical and scanning electron microscopy. A preliminary analysis of mitochondrial DNA sequences including members of related lineages is also provided. The immature stages are associated with Passiflora actinia, Passiflora misera and Passiflora suberosa (Passifloraceae), and build mines on the adaxial leaf surface. Initially the mines are serpentine in shape, but later in larval ontogeny become a blotch type. Although the larvae are hypermetamorphic as in other Gracillariidae, there is no sap-feeding instar in Spinivalva gaucha; the larva feeds on the palisade parenchyma, thus producing granular frass during all instars. Pupation occurs outside the mine; prior to pupating, the larva excretes numerous bubbles that are placed in rows on the lateral margins of the cocoon external surface. This is the second genus of gracillariid moth described for the Atlantic Rain Forest, and the second gracillariid species known to be associated with Passifloraceae.     

Cytogenetics of the genus Parasarcophaga (Sarcophagidae: Diptera)

Somatic chromosome complements of five sympatric species of the genus Parasarcophaga, viz. P. misera, P. albiceps, P. argyrostoma, P. ruficornis and P. knabi, are described. All the species have five pairs of meta/submetacentric autosomes and an XX/XY sex chromosome pair which is highly variable in size and shape. In P. misera and P. albiceps they are tiny dots while in P. knabi the metacentric X and Y chromosomes constitute almost one third of the genome. In P. ruficornis and P. argyrostoma they are telocentric chromosomes of moderate size. A comparative study of the C-banding patterns of P. ruficornis, P. knabi, P. argyrostoma and P. misera shows that autosomes of the former three species possess characteristic C-bands in pericentric regions while in P. misera they are absent. The heterochromatic sex chromosomes are C-band positive in all the species. However, with the exception of the tiny sex chromosomes of P. misera, the X chromosomes of other species show shorter or longer regions which stain rather lightly. These C-banded areas correspond to the heterochromatic areas revealed in orcein stained preparations. The evolutionary implications of these results are discussed.     

Activities and feeding behavior of free-ranging pregnant baboons

The activities and feeding behavior of pregnant baboons, Papio cynocephalus,were studied in two free-ranging groups in Amboseli National Park, Kenya. Both the environmental conditions and the age of their fetuses influenced females’ activity budgets and feeding behavior. Females fed more and rested less during the dry season than during the wet season, and most females spent progressively more time feeding and less time grooming than expected (based upon the amount of recent rainfall) as their fetuses matured. During the wet season, females also devoted significantly greater proportions of feeding time to consumption of grass blades, leaves, and flowers and significantly smaller proportions of their feeding time to feeding upon seeds, corms, and fruit. As their fetuses became older, the proportion of time spent feeding upon seeds increased, and the proportion of time spent feeding upon grass blades and fruit declined.     

Foraging strategies of yakushima macaques (Macaca fuscata yakui )

I investigated the activity budget and diet of Yakushima macaques (Macaca fuscata yakui,)in warm temperate broad- leaved forest of Yakushima, Japan. Both time spent feeding and time spent moving varied considerably between half- months. However, total time spent in active behaviors— feeding time plus moving time— was stable. The composition of the diet also showed considerable variation between half- months. The macaques fed mainly on fruits, seeds,mature leaves, fallen seeds, flowers, and young leaves, each of which accounted for more than 30% of feeding time in at least 1 half- month. They also ate insects and fungi, but each of them comprised ≤ 25 and ≤ 8% of feeding time in any half- month, respectively. Time spent feeding on mature leaves, young leaves, flowers, or fallen seeds is positively correlated with total time feeding and is negatively correlated with time moving. In contrast, time feeding on fruits, seeds, insects or fungi is negatively correlated with time feeding and is positively correlated with time moving. Foraging on foods that have a low energy content, a high density, and a relatively even distribution— mature leaves— or that need much manipulation to be processed— flowers and fallen seeds— increased feeding time, while foraging on foods for which monkeys must search intensively in the forest— fruits, seeds, insects, and fungi— led to increased moving time. I examined foraging strategies of Yakushima macaques in terms of moving costs and the quality of food items. Regarding time feeding on fruits, which have more energy and may need less manipulation than other foods, as a benefit, and moving time as a cost, they seemed to employ a strategy that balanced the costs and benefits of foraging.     

Karyology and cytotaxonomy of the genus Passiflora L. (Passifloraceae)

 The chromosomes of 31 species of Passiflora, distributed throughout the subgenera Astrophea, Calopathanthus, Distephana, Dysosmia, Passiflora, Plectostemma and Tacsonia were analysed. Three different karyotypes were observed: 2n = 12, 24, 36; 2n = 18, 72 and 2n = 20. The karyotype of these species was almost always constituted of metacentric and submetacentric chromosomes with variable karyotype symmetry. In the group with x = 6, represented by the subgenus Plectostemma, six diploid species with 2n = 12, one tetraploid with 2n = 24 (P. suberosa) and an intraspecific polyploid with 2n = 12, 36 (P. misera) were analysed. P. pentagona (subgenus Astrophea) may also be included in this karyological group since it presents 2n = 24 and may be of polyploid origin, with x = 6. The interphase nuclei in this group were areticulate, except those of P. morifolia and P. pentagona with semi-reticulate characteristics. Two small terminal heterochromatic blocks, positive for chromomycin A₃, were identified in the largest chromosome pair of P. capsularis and P. rubra, species very closely related, while P. tricuspis displayed four chromosomes with proximal blocks. In the group with x = 9, represented mainly by subgenus Passiflora, 20 species with 2n = 18 and one with 2n = 72 were studied. They presented chromosomes larger than those species with x = 6 and interphase nuclei of semi-reticulate type, except for P. mixta with areticulate nuclei. Four terminal CMA⁺ blocks were observed in P. edulis, six blocks in P. caerulea and P. racemosa, while five blocks were observed in the single P. amethystina plant analysed. P. foetida (subgenus Dysosmia), the only species with 2n = 20, exhibited six chromosomes with CMA⁺ blocks and interphase nuclei of the areticulate type. The meiotic analysis of representatives of the three groups (P. foetida, P. suberosa, P. cincinnata and P. racemosa) always presented regular pairing and regular chromosome segregation, except in P. jilekii where a tetravalent was observed. The analysis of the chromosome variation within the genus and the family suggests that the base number of Passiflora may be x₁ = 6 or x₁ = 12, whereas x₂ = 9 is only an important secondary base number. Received April 11, 2000 Accepted October 5, 2000     

Fallback Foods of Red Leaf Monkeys (Presbytis rubicunda) in Danum Valley, Borneo

Animals in Southeast Asia must cope with long periods of fruit scarcity of unpredictable duration between irregular mast fruiting events. Long-term data are necessary to examine the effect of mast fruiting on diet, and particularly on the selection of fallback foods during periods of fruit scarcity. No such data is available for colobine monkeys, which may consume substantial amounts of fruits and seeds when available. We studied the diet of red leaf monkeys (Presbytis rubicunda, Colobinae) in Danum Valley, Sabah, northern Borneo, using 25 mo of behavioral observation, phenology and vegetation surveys, and chemical analysis to compare leaves eaten with nonfood leaves. The monkeys spent 46% of their feeding time on young leaves, 38% on seeds, 12% on whole fruits, 2.0% on flowers, 1.0% on bark, and 1.2% on pith. They spent more time feeding on seeds and whole fruit when fruit availability was high and fed on young leaves of Spatholobus macropterus (liana, Leguminosae) as fallback foods. This species was by far the most important food, constituting 27.9% of the total feeding time, and the feeding time on this species negatively correlated with fruit availability. Consumed leaves contained more protein than nonconsumed leaves, and variation in time spent feeding on different leaves was explained by their abundance. These results suggest that red leaf monkeys show essentially the same response to the supra-annual increase in fruit availability as sympatric monogastric primates, increasing their seed and whole-fruit consumption. However, they depended more on young leaves, in particular Spatholobus macropterus, as fallback foods during fruit-scarce periods than did gibbons or orangutans. Their selection of fallback food appeared to be due to both nutrition and abundance.     

Behaviour of late-instar gypsy moth larvae in high and low density populations

ABSTRACT. 1. Using scaffolding and night-vision equipment, we observed fifth and sixth instars of the gypsy moth, Lymantria dispar (L.), on Quercus velutina Lam. in the field.
2. In low-density populations, larvae fed at night and spent the day resting in sheltered sites away from the canopy. In high-density populations, larvae remained in the canopy throughout the day and night, and the amount of feeding during daylight hours increased with population density.
3. Larvae at all population densities used a similar sequence of behaviours and sampled a number of leaves when selecting feeding sites, but larvae in high-density populations switched feeding sites more frequently and fed continuously for shorter periods.
4. Larvae seldom interfered with each other's feeding in any of the populations.
5. When fifth instars were collected from the field and held for 24 h in an electronic feeding monitor, they maintained feeding rhythms that were characteristic of their source populations. Larvae spent more time crawling and less time feeding when offered foliage from high-density rather than low-density populations.     

Walking,feeding and intraspecific interaction of larvae of Metaseiulus occidentalis,Typhlodromus pyri,Neoseiulus fallacis and Amblyseius andersoni held with and without eggs of Tetranychus urticae

Larvae of Metaseiulus occidentalis (Nesbitt), Typhlodromus pyri Scheuten, Neoseiulus fallacis (Garman) and Amblyseius andersoni Chant exhibited different activity levels when held on apple leaf or on tile arenas and given or not given eggs of Tetranychus urticae Koch and water (tiles only). M. occidentalis larvae held without prey exhibited high levels of walking (includes searching) during 24 hours of evaluation, whereas M. occidentalis larvae held with prey fed quickly and then became less active. Fed larvae of M. occidentalis were less active on leaves than tile. Larvae of T. pyri on leaves had a very low frequency of walking, almost never fed and quickly assumed a resting position during development. While much less active than M. occidentalis, fed and unfed T. pyri larvae walked more on tiles than leaves before resting. Larvae of N. fallacis and A. andersoni fed at low rates and were similarly active on tiles and leaves. Free water increased walking by M. occidentalis, A. andersoni and T. pyri on tile but not N. fallacis. M. occidentalis larvae interacted 5–7 times more often than larvae of the other three species. Cannibalism or scavenging was rarely seen and then only for M. occidentalis larvae. Larvae of all four mites walked, fed and interacted much more in the first 12 hours than the second 12 hours of tests, except unfed M. occidentalis. Unfed M. occidentalis larvae did not molt to protonymphs but unfed larvae of the other three species did. Unfed and fed protonymphs of all four species walked more at 4 hours after molting than larvae at 12–24 hours. Unfed and fed protonymphs of T. pyri or A. andersoni had similar walking frequencies, but unfed protonymphs of N. fallacis were more active than fed ones. Trends in larval activities are discussed relative to the life history of each species.