Morphology and taxonomy of the microsporidium Liebermannia covasacrae n. sp. from the grasshopper Covasacris pallidinota (Orthoptera, Acrididae)

During a survey for grasshopper pathogens in Argentina in 2005-2006, individual Covasacris pallidinota from halophylous grasslands in Laprida, Buenos Aires province were found to be infected with a microsporidium. Infection was restricted to the salivary gland epithelial cells. The microsporidium produced ovocylindrical spores averaging 2.6 ± 0.28 × 1.4 ± 0.12 μm (range 2.2-3.4 × 1.1-1.7 μm), which resembled in size and shape the spores of Liebermannia patagonica and L. dichroplusae, two recently described species that also parasitize Argentine grasshoppers. The life cycle of the microsporidium included the formation of polynucleate, diplokaryotic, moniliform, merogonial plasmodia wrapped in flattened cisterns of the host endoplasmic reticulum (ER). Plasmodia divided to produce diplokaryotic cells. The latter underwent elongation, dissociation of diplokarya counterparts, vacuolization, dismantling of the host ER envelope, and deposition of electron-dense material outside the plasma membrane. The resultant binucleate sporogonial plasmodia divided into two uninucleate sporoblasts, which eventually transformed into spores. Uninucleate spores contained a lamellar polaroplast, embraced by an elongated polar sac, anchoring disc, 3-5 polar filament coils, and a cluster of anastomizing tubules (sporoblast trans-Golgi, posterosome) at the posterior end. Sequence similarity of the SSU rDNA of the newly discovered microsporidium (Genbank accession no. EU709818) to L. patagonica and L. dichroplusae was 99% and 97%, respectively, suggesting that the three species belong to one genus. All three species fell into one clade in SSU rDNA-based phylogenetic trees produced by neighbor joining, maximum parsimony, and maximum likelihood analyses with 100% statistical support. We assign the name Liebermannia covasacrae to this microsporidium. It can be easily differentiated from both congeners by host species, tissue tropism, type of sporogony, and several features of morphology. Comparison of the three Liebermannia spp. demonstrates that the nuclear phase (dikaryotic versus monokaryotic spores) and type of sporogony (polysporous versus disporous) may vary in closely related species.     

New case of long-term persistence of Paranosema locustae (Microsporidia) in melanopline grasshoppers (Orthoptera: Acrididae: Melanoplinae) of Argentina

We report an additional case of long-term persistence of Paranosema locustae in grasshoppers of Argentina. The pathogen was introduced from North America on rangeland at Loncopué, Neuquén province. Microsporidia were not detected in pre-introduction samples whereas infected grasshoppers were found 11 years after introduction. Affected grasshoppers were the melanoplines Dichroplus elongatus, Dichroplus maculipennis, and Scotussa lemniscata, some of them with high spore loads. The case highlights the ability of P. locustae to recycle in local grasshopper communities by parasitizing susceptible species other than the natural hosts.     

Mate guarding and male mate choice in the chameleon grasshopper <Emphasis Type="Italic">Kosciuscola tristis</Emphasis> (Orthoptera: Acrididae)

In the wild, male chameleon grasshoppers (Kosciuscola tristis) are frequently observed mounted on the back of females even when not in copula, and will fight off other usurping males. If this behaviour is mate guarding and reflects investment in male mate choice, then we expect males to preferably guard females based on reliable cues of quality. Cues for female quality likely include female size and egg development that together may indicate fecundity. We investigated male mate choice in the field expressed as mate-guarding preference, by comparing size and egg development in guarded and unguarded females. We found no difference between guarded and unguarded females in measures of fecundity or body size. The majority of females sampled did not contain any viable eggs. This finding suggests that male K. tristis indiscriminately guard females in a scramble mating system.     

Metaphase I orientation of Robertsonian trivalents in the water-hyacinth grasshopper, Cornops aquaticum (Acrididae, Orthoptera)

Trivalents resulting from polymorphic Robertsonian rearrangements must have a regular orientation in metaphase I if the polymorphisms are to be maintained. It has been argued that redistribution of proximal and interstitial chiasmata to more distal positions is necessary for a convergent orientation, the only one that produces viable gametes. Cornops aquaticum is a South-American grasshopper that lives and feeds on water-hyacinths, and has three polymorphic Robertsonian rearrangements in its southernmost distribution area in Central Argentina and Uruguay. The orientation of trivalents in metaphase I, the formation of abnormal spermatids and the frequency and position of chiasmata in the trivalents, was analysed in a polymorphic population of C. aquaticus. In this study we observed a correlation between the number of trivalents with the frequency of abnormal spermatids; additionally, the number of chiasmata, especially proximal and interstitial ones, was strongly correlated with the frequency of the linear orientation. Therefore we confirmed our previous assumption, based on other evidence, that the chiasmata redistribution in fusion carriers is essential to the maintenance of the polymorphisms.     

Resource-mediated impact of spider predation risk on performance in the grasshopper Ageneotettix deorum (Orthoptera: Acrididae)

In response to increased exposure to predators when searching for food, many prey increase the frequency of antipredator behaviors, potentially reducing foraging rate and food intake. Such direct, nonlethal interactions between predators and prey resulting in reduced food intake can indirectly influence lifecycle development through effects on growth, developmental rate, and survival. We investigated the general hypothesis that individual performance of a herbivorous insect can be negatively affected when exposed to nonlethal predation risk, and that the response can be mediated by food quality. This hypothesis was tested using the common rangeland grasshopper Ageneotettix deorum with and without exposure to common wolf spider predators (Lycosidae, Schizocosa spp.) on both untreated natural and fertilized vegetation. All spiders were rendered temporarily incapable of direct feeding by restricting function of the chelicerae with beeswax. Detectable responses by grasshoppers to spiders indicate indirect consequences for lifecycle development. Grasshopper performance was measured as hind femur growth, duration of nymphal lifecycle stages, and survivorship in a caged field experiment conducted over 2 years. Grasshoppers developed faster and grew 3–5% larger when allowed to forage on fertilized vegetation in the absence of risk from a spider predator. Failure-time analysis illustrated enhanced survival probability in response to elevated food quality and the negative effects of grasshopper susceptibility to nonlethal predation risk. Performance on food of relatively low, ambient quality with no predation risk equaled that of grasshoppers caged with high quality vegetation in the presence of a modified spider. Increased resource quality can clearly moderate the negative life history responses caused by the behavioral modification of grasshoppers when exposed to spider predation risk, a compensatory response.     

Floral colour change in Pedicularis monbeigiana (Orobanchaceae)

We examined the effects of the retention of colour-changed flowers on long- and short-distance attractiveness of bumblebees and the likelihood of successive flower visits by bumblebees in Pedicularis monbeigiana. The lower lip changed colour with age from white to purple. Hand geitonogamous pollination significantly reduced seed production. No pollen limitation occurred in this species. Purple-phase flowers contributed minimally to pollinator attractiveness at long distance. The combination of less reproductive flowers with a lower amount of reward and floral colour change enabled plants to direct pollinators to reproductive, highly rewarding white flowers at close range. A high percentage of purple-phase flowers in an inflorescence was associated with a marked reduction in the frequency of successive flower visits to individual plants. We suggest floral colour change in P. monbeigiana may serve as a mechanism for enhancing inter-individual pollen transfer and reducing intra-individual pollen transfer.     

The complete mitochondrial genome of Gomphocerus tibetanus Uvarov, 1935 (Orthoptera: Acrididae: Gomphocerinae)

The complete nucleotide sequence of the mitochondrial genome (mitogenome) of Gomphocerus tibetanus Uvarov, 1935 (Orthoptera: Acrididae: Gomphocerinae) was determined. It is 15,571 bp in length and contains 74.8% A + T. All Gomphocerus tibetanus protein-coding sequences start with a typical ATN codon. The usual termination codons (TAA and TAG) were found from 13 PCGs except COI and COII which took incomplete codon T as termination codons. All tRNA genes could be folded into the typical cloverleaf secondary structure, except tRNA^Ser(AGN) lacking of dihydrouridine (D) arm. The sizes of the large and small ribosomal RNA genes are 1313 and 822 bp, respectively. The A + T content of the A + T-rich region is 82.3%. A preliminary analysis on characteristics of Gomphocerinae mitogenome was made by comparision among three Gomphocerinae mitogenomes and Locusta migratoria.     

The adaptive significance of ontogenetic colour change in a tropical python

Ontogenetic colour change is typically associated with changes in size, vulnerability or habitat, but assessment of its functional significance requires quantification of the colour signals from the receivers' perspective. The tropical python, Morelia viridis, is an ideal species to establish the functional significance of ontogenetic colour change. Neonates hatch either yellow or red and both the morphs change to green with age. Here, we show that colour change from red or yellow to green provides camouflage from visually oriented avian predators in the different habitats used by juveniles and adults. This reflects changes in foraging behaviour and vulnerability as individuals mature and provides a rare demonstration of the adaptive value of ontogenetic colour change.     

Developmental specialization and geographic structure of host plant use in a polyphagous grasshopper, Schistocerca emarginata (=lineata) (Orthoptera: Acrididae)

Host plant use and availability were determined in early nymphal and adult-stage Schistocerca emarginata (=lineata) (Orthoptera: Acrididae) populations at six localities in Texas, USA. Early instar nymphal populations were feeding almost exclusively on either Ptelea trifoliata (Rutaceae) or Rubus trivialis (Rosaceae). This study represents the first demonstration of a geographic structure of host plant specificity in a polyphagous grasshopper. Recognizing this geographic structure required investigations of both developmental and geographical variation in host plant use. Nymphal diet breadths were significantly less than adult diet breadths at four of six localities and smaller overall when pooled nymphal and adult diet breadths were compared among sites. Neither restricted nymphal mobility nor host plant availability accounted for the observed differences in host plant use between developmental stages and among localities. Evidence suggests that the differences in host use among populations are due to host-plant-associated genetic differentiation. Received: 4 June 1998 / Accepted: 10 March 1999     

Development of the subsoephageal body cells and the pericardiac cells during embryogenesis with diapause in Locusta migratoria (Linnaeus 1758) (Orthoptera: Acrididae)

During Locusta migratoria embryogenesis, the yolk is progressively degraded and the resulting metabolites are released in the haemolymph. We researched the organs possibly involved in the uptake of haemolymphatic proteins. Among organs originated from mesoderm, the SOB (suboesophageal bodies) situated in the embryonic head are remarkable by a very early acquisition of differentiated cytological characters, while most other cells of the embryo are undifferentiated. The SOB quite disappear before hatching. Just before katatrepsis stage, the other organs derived from mesoderm begin to differentiate, including the PC (pericardial cells) which take over from the SOB. These cells, situated in thorax and abdomen, are developed during the dorsal close of embryo. The development and the ultrastructural changes of the SOB cells and of the PC were studied during an embryogenesis with diapause. The morphology of embryos which enter diapause is comparable with that of a continuous development at the beginning of katatrepsis. However, the cells of SOB and PC cells suffer from remarkable changes not only physiologically but cytologically. At the beginning of diapause, the proteosynthetic activity practically disappears in the SOB cells and the lysis areas appear. Nevertheless, the exchanges between these cells and the haemolymph still remain important. For the period of cold, which is necessary to the resumption of development, the aspect of the SOB cells changes and in particular the areas of lysis become less wide. When the embryo reopens its development, the SOB cells show a proteosynthetic activity and the areas of lysis disappear. The changes of the SOB cells and of the PC cells are regularized during the resumption of the development: the SOB cells which had again taken a normal activity start to regress from the stage VII on, while the PC cells take over.     

Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)

Many gomphocerine grasshoppers communicate acoustically: a male's calling song is answered by a female which is approached phonotactically by the male. Signals and recognition mechanisms were investigated in Chorthippus biguttulus with regard to the cues which allow sex discrimination. (1) The stridulatory files on the hindfemur of both sexes are homologous in that they are derived from the same row of bristles, but convergent with respect to the “pegs”. In males the pegs are derived from the bristles, and in females from the wall of the bristle's cup. (2) Male and female songs are generated by similar, probably homologous motor programs, but differ in the duration, intensity, “gappyness” of syllables, risetime of pulses, and the frequency spectra. The hindleg co-ordination during stridulation and the resulting temporal song patterns are less variable in males than in females. (3) For both sexes, recognition of a mate's signal depends on species-specific syllable structure. For males it is essential that the female syllables consist of distinct short pulses, whereas females reject “gappy” syllables. Males strongly prefer “ramped” pulses, females respond to syllables irrespective of steeply or slowly rising ramps. Males react only to the low-frequency component, whereas females prefer spectra containing both, low and high frequency components. Accepted: 20 November 1996     

The roles of colour change and behaviour in the hygrothermal balance of a littoral isopod,Ligia oceanica

Summary Ligia oceanica can change its colour using melanophores, the animal's reflectance varying between about 2 and 10%. Darker individuals heat up more quickly, and to higher body temperatures, than do pale ones. Colour change shows an underlying circadian rhythm, though the pattern of this rhythm varies with temperature, humidity, background and time of year. In general the rhythm is such as to ensure maximum camouflage at the critical dusk period, but in some conditions hygrothermal needs are overriding and the animals are paler (to stay cool) or darker (to warm up). In addition, animals show short term colour modification; when transferred to differing backgrounds and temperatures their colours initially reflect background matching, but after 30–45 min are modified into thermally appropriate shades, dark at 5° C and pale at 20° C. Field-caught specimens showed body temperatures that varied with colouration, and modification of colour in relation to thermal needs, particularly by being paler than expected when forced into the open by daytime high tides, and darker than expected when active prior to dusk. Animals invariably selected dark backgrounds in choice chambers. However, choice of humidity depended on previous experience; saturated air was normally preferred, but warm animals chose drier air (to allow evaporative cooling) unless also water-stressed. They also tended to disperse to facilitate cooling, whereas aggregation increased with increasing RH and with decreasing temperature. The interactions of colour changes, behavioural choices, and activity patterns in controlling the hygrothermal belance of Ligia in the intertidal environment are discussed in the light of these results.     

Ontogeny of territoriality in the desert clickerLigurotettix coquilletti (Orthoptera: Acrididae)

Field observations and experiments showed that settlement prior to reproductive maturation strongly influenced territorial success in the desert clicker(Ligurotettix coquilletti), an acridid grasshopper in which males defend individual host plants (Larrea tridentata bushes) as mating arenas. High tendencies to move and reposition among the host plants were displayed in two distinct episodes. The first occurred during early nymphal development, and it resulted in many individuals settling on high-qualityLarrea bushes. Prior work showed thatL. coquilletti that fed on foliage from these specific bushes exhibited higher relative growth rates and therefore were expected to eclose sooner. Early-eclosing males enjoyed a priority advantage in defending high-quality bushes as mating territories, and they consequently encountered more females and mated more frequently. Despite the distribution of most insects on high-quality bushes at the time of eclosion, though, a second movement episode occurred shortly thereafter. This reshuffling contrasts markedly with the site fidelity of mature adult males, most of which settle on mating territories, and it may function as a means of examining an enlarged sample of potential sites. The above results imply that territorial success of maleL. coquilletti does not result from retaining sites defended by the parents. This point is also supported by the finding that females do not oviposit particularly close to theLarrea bushes in which they reside regardless of their quality; most egg pods are deposited in bare soil midway between the bushes. Nonetheless, certain oviposition sites may be conducive to earlier hatch, and this can lead to earlier eclosion and ultimately to defense of a valuable territory. Therefore, the parental generation, through selective oviposition, may yet influence the success of their male offspring.     

Foraging patterns and dietary mixing in the field by the generalist grasshopperBrachystola magna (Orthoptera: Acrididae)

The foraging behavior of a cryptic romaleine grasshopper was observed in its natural habitat in southern Arizona. Individual polyphagy, dietary mixing, and the pattern of feeding on different substrates were monitored. Individuals were found to be extreme generalists with surprising levels of feeding on other insects. Differences in the foraging behavior between grasshopper species are discussed in relation to the habitat, the season, and the different defensive strategies.     

Consumption rates and the evolution of diet-induced plasticity in the head morphology of Melanoplus femurrubrum (Orthoptera: Acrididae)

Summary Phenotypic plasticity may be an ecologically important evolutionary response to natural selection in multiple environments. I have determined the effect of diet-induced developmental plasticity in the head size of grasshoppers (Melanoplus femurrubrum) onfeeding performance on two types of plants. Full-sib families were divided and raised on either red clover, Trifolium repens, or rye grass, Lolium perenne. In three different stages of ontogeny, grasshoppers raised on rye grass had significantly larger heads, relative to body size, than full-sibs raised on clover. A principal components analysis indicated that two to five relative head size characters covaried as a block in their plastic response to the feeding environment. Regressions of adjusted consumption rates (mg/sec) against relative head size revealed that larger head sizes, induced by the rye grass diet, enhanced consumption rates of rye grass, but not clover. Unexpectedly, a similar positive association was observed between head size and consumption rate for grasshoppers raised on clover when they were feeding on clover. These results support the inference that grasshoppers exhibit adaptive phenotypic plasticity. However, the unexpected influence of head size on consumption rates of clover indicates that the functional relationship between head morphology and feeding performance is complex and that variation in this relationship among plant environments is not sufficient to explain the evolution of diet-induced phenotypic plasticity.     

Transmission and perception of acoustic signals in the desert clicker,Ligurotettix coquilletti (Orthoptera: Acrididae)

Males of the desert clicker, Ligurotettix coquilletti (Acrididae: Orthoptera) defend a femalerequired resource, the creosote bush Larrea tridentata, in desert habitats of the southwestern United States. Males signal acoustically to each other as well as to searching females. The call is produced by tegminal/femoral stridulation where one or both legs are used for sound production. Sound pressure levels, measured laterally, are influenced by the intervening tegmen between the stridulating leg and the microphone. Differences in measured sound pressure levels between sides can vary up to 7 dB. When clicks are produced multiply,these multiple clicks may be 4 dB louder than single clicks. We examine the structure of the call and the effective broadcast area of single males by monitoring acoustic ascending neurons of the ventral nerve cord in the neck. By taking the neurophysiological preparation into the field, we were able to map the broadcast area of isolated males and also of males calling within aggregations. The distance over which the signal of isolated males could be detected was 8–14 m, whereas neural representation of the calls of males within aggregation were detectable within 4–6 m. The sound spectrum of the song, although having a major lower-frequency component around 10 kHz, has extensive power in the ultrasonic range. The tuning characteristics of the ascending auditory neuron matched the overall structure of the male call. The importance of the acoustic cue, as compared to visual cues, is discussed in relation to female attraction.