Diversity patterns and chronobiology of hawkmoths (Lepidoptera,Sphingidae) in the Brazilian Amazon rainforest

As biodiversity loss rapidly increases through habitat degradation in the Amazon rainforest, the need to characterize and understand the species diversity becomes even more important. In this study we used empirical and published datasets to assess the diversity patterns and produce the first overview of the sphingid fauna in the Brazilian Amazon. We compared the diversity patterns in distinct areas in the biome by analyzing hawkmoth assemblages considering both species composition and abundance, and asked whether these communities are structured according to environmental factors. Additionally, we provide information of diel activity pattern of sphingids and evaluated the importance of time in sampling effort. We found that the Brazilian Amazon may harbor more than 80 % of the hawkmoth species that occur in Brazil and more than half of the species recorded in South America. Species composition and assemblage structure is determined by the quality of the habitat (disturbed or undisturbed vegetation), temperature and relative humidity. Finally, we show that the temporal activity of sphingids presents distinct patterns at different taxonomic levels, highlighting the importance of full night collections to better characterize the fauna. Our results show that habitat alteration can be an important factor affecting sphingid assemblages, illustrating the importance of Protected Areas in species maintenance.     

The neuroethology of sound production in tiger moths (Lepidoptera,Arctiidae)

When stimulated either acoustically or tactually, certain species of arctiid moths rhythmically emit trains of clicks from metathoracic tymbals. The purpose of the experiments presented here was to determine the location within the central nervous system (CNS) of the proposed tymbal central pattern generator (CPG) in Cycnia tenera. Motor neuron impulses that underlie tymbal activation were recorded extracellularly from the tymbal nerve while moths were subjected to selective severing of the suboesophageal, prothoracic, pterothoracic and abdominal ganglia connectives. Motor output evoked by either acoustic or tactile stimulation originates from a common CPG because tymbal nerve spikes in both cases are similar in amplitude, waveform and rhythmicity. Our results showed: (1) removal of the CNS posterior of the second abdominal neuromere had no effect, (2) removal of the head decreased the responsiveness of the animal to acoustic stimulation and, (3) severing the connectives between the prothoracic and pterothoracic ganglia abolished responses to acoustic stimuli and diminished responses to tactile stimuli. We conclude that although the minimal circuitry sufficient for activating the tymbals resides in the pterothoracic ganglion, the prothoracic and cephalic ganglia are required for the normal, and in particular, auditory-evoked operation of the tymbal CPG.Abbreviations ASR acoustic startle response - CNS central nervous system - CPG central pattern generator - dB peSPL decibel peak equivalent sound pressure level (rms re 20 Pa) - ISI inter-spike interval     

Museum archives revisited: Central Asiatic hawkmoths reveal exceptionally high late Pliocene species diversification (Lepidoptera,Sphingidae)

Three high‐elevation Hyles species of Central Asia have proven difficult to sample, and thus, only a limited number of specimens are available for study. Ancient DNA techniques were applied to sequence two mitochondrial genes from ‘historic’ museum specimens of Hyles gallii, Hyles renneri and Hyles salangensis to elucidate the phylogenetic relationships of these species. This approach enabled us to include the holotypes and paratypes. The status of H. salangensis as a species endemic to a mountain range north of Kabul in Afghanistan is confirmed by this study. It is most closely related to Hyles nicaea and H. gallii, and quite distant from the clade comprising the species from Hyles vespertilio to Hyles tithymali, despite this group and H. salangensis both completely lacking an arolium on the pretarsus. Our results show that the samples assigned to H. renneri and Hyles livornica tatsienluica are conspecific and so we reinstate Hyles tatsienluica stat. nov. as the valid name for this species and synonymize H. renneri syn. nov. with it. This study shows that the distribution range postulated for H. tatsienluica extends from Nepal well into the mountains of south‐western China. The distribution ranges of H. livornica and H. tatsienluica overlap. The study confirms the previously proposed synonymies of Hyles nepalensis, Hyles gallii intermedia and Hyles gallii tibetanica with H. gallii. Extensive species sampling (over 80% of Hyles species) in this study allowed additional analyses. The dated phylogeny reveals the global Hyles hawkmoth radiation to be much more recent than previously thought: it began in the Late Miocene and culminated in a Pleistocene burst of diversification in the Northern Hemisphere. Ancestral ranges of basal nodes were reconstructed as highly equivocal, but the Neotropics has the highest probability in the two oldest nodes. Although the origins of the Madagascan and Australian species also remain ambiguous, a large crown clade of fifteen species was reconstructed to have originated in the Palaearctic. The wide distribution ranges of the two migratory species, H. livornica and H. gallii, appear to blur any traces of the biogeographic origin of the clades containing these species. Specialization in larval host plant use onto particular plant families from the ancestral condition ‘polyphagous’ may have led to an increased rate of speciation and phylogenetic diversification in three subgroups of Hyles (the Hawaiian clade, the Hyles centralasiae group and the Hyles euphorbiae complex).     

The phylogenetic relationships of Morgan's Sphinx, Xanthopan morganii (Walker), the tribe Acherontiini, and allied long-tongued hawkmoths (Lepidoptera: Sphingidae, Sphinginae)

A cladistic analysis is presented of the hawkmoths of the tribe Acherontiini, Morgan's Sphinx (Xanthopan morganii (Walker)), and related genera. The study aims to test the monophyly of tribe Acherontiini; the hypothesis that all taxa with extremely long probosces (some Acherontiini, Meganoton rubescens, Neococytius, Xanthopan) form a monophyletic group, or at least fall within a single reasonably compact clade; and, within this group, to determine whether Xanthopan is more closely related to Acherontiini or to Cocytius and Neococytius. The data set comprises 109 characters derived from adult and immature stage morphology, biology and behaviour. These data were analysed using equal weighting, successive approximations character weighting (SACW) and implied weighting. All weighting schemes agreed on the monophyly of Acherontiini and of a group of genera comprising Amphimoea, Cocytius and Neococytius (the Cocytius group). Several other generic and suprageneric clades were also consistently recovered. However, those hawkmoths with extremely long probosces were never recovered as a monophyletic group. The relationships of Xanthopan were also ambiguous. Equal weighting and SACW placed Xanthopan + Meganoton rubescens (Butler) as sister to the Cocytius group, while implied weighting placed Xanthopan as sister to Acherontiini. This latter relationship is based primarily on shared possession of a pilifer/palp hearing organ. Further analyses suggested the two components of this organ were not biologically independent. Downweighting this feature accordingly resulted in all weighting schemes converging on the topology found by equal weighting. Exclusion of the incomplete subset of immature stage data had no effect under implied weighting but equal weighting and SACW now recovered a Neotropical clade comprising Manduca and the Cocytius group, while Xanthopan was placed with M. rubescens and Panogena. Downweighting the pilifer/palp hearing organ under implied weighting again caused convergence with the equal weighting/SACW results. Thus, the relationships of Xanthopan remain equivocal and further data, particularly from the immature stages, will be required to elucidate its phylogenetic position further. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 2002, 135 , 471–527.     

Measuring range sizes of South-East Asian hawkmoths (Lepidoptera: Sphingidae): effects of scale, resolution and phylogeny

Aim  The effects of resolution and spatial extent on range measures were explored in estimates of the geographic distribution of tropical hawkmoths. Furthermore, data were tested for phylogenetic autocorrelation.
Location  South-East Asia.
Methods  Various range measures, such as geographic information system (GIS)-supported range estimates, minimum convex polygons, latitudinal and longitudinal extents, and their products, were derived from original distribution records and compared to each other. A taxonomic classification of the species was used to analyse phylogenetic effects on range sizes.
Results  Range size measures exhibit a strongly right-skewed frequency distribution with many geographically restricted species and few widespread taxa. Rankings from GIS-supported, comprehensive range size estimates do not deviate greatly from more crude measurements of lower resolution. Comprehensive ranges and ranges within South-East Asia are correlated strongly, but already at this rather large scale the ranking of species changes considerably. Other measures of occupancy with an increasingly more localized consideration of 'range' show decreasing strengths of correlation. We found a weak, but significant, autocorrelation in range area data: related groups of species have ranges of similar size.
Main conclusions  Spatial resolution did not affect range ranking greatly in our data. However, macroecological studies based only on parts of species' ranges must be viewed critically, particularly if their extent is small compared to comprehensive ranges. Phylogenetic non-independence of range size data must be considered in comparative analyses.     

Contribution of the maxillary muscles to proboscis movement in hawkmoths (Lepidoptera: Sphingidae)--an electrophysiological study

The role of the maxillary muscles in the uncoiling and coiling movements of hawkmoths (Sphingidae) has been examined by electromyogram recordings, combined with video analysis. The maxillary muscles of adult Lepidoptera can be divided into two groups, galeal and stipital muscles. The galea contains two basal muscles and two series of oblique longitudinal muscles, which run through the entire length of the galea. Three muscles insert on the stipes, taking their origin on the tentorium and on parts of the cranium and gena, respectively. Proboscis extension is initiated by an elevation of the galea base caused by the basal galeal muscles. The actual uncoiling of the proboscis spiral is accompanied by rapid compressions of the stipites which are caused by two of the stipital muscles. The study provides strong support for the hypothesis that uncoiling is brought about by an increase of hemolymph pressure by the stipites forcing hemolymph into the galeae. Recoiling is caused by the contraction of both sets of oblique longitudinal galeal muscles supported by elasticity of the galea cuticle. Finally, the remaining stipital muscle pulls down the galea base which brings the coiled proboscis back to its resting position where it is held in the U-shaped groove of the labium without further muscle activity.     

On recognition of sound communication signals in bush crickets (Orthoptera,Tettigoniidae)

Two hypotheses have been proposed to explain the mechanisms of calling signal recognition in orthopterans: the filtration and resonance ones. To test these hypotheses, conspecific male calling songs and their models with modified temporal parameters were presented to females of bush crickets in ethological experiments. The models with a double pulse rate evoked positive phonotaxis of females while phase shift significantly complicated the recognition process. These data fit the resonance hypothesis.     

Phylogeny of the death's head hawkmoths, Acherontia[Laspeyres], and related genera (Lepidoptera: Sphingidae: Sphinginae: Acherontiini)

Abstract. Adult death's head hawkmoths (Acherontia species) have a unique feeding biology as cleptoparasites of honeybees, stealing honey from the combs, rather than imbibing nectar from flowers. The moths have a range of features, both morphological and behavioural, that enable them to successfully enter, feed and escape from the colonies. These adaptations may vary among the three Acherontia species and allow them each to target different species of honeybee. A cladistic analysis is presented of the hawkmoths of tribe Acherontiini. The study aims to resolve the relationships of the genera and species of Acherontiini, with a particular focus on the three species of Acherontia. The dataset comprises sixty‐five characters derived from adult, larval and pupal morphology, and larva host‐plant biology. These data are analysed using equal weighting and implied weighting. Acherontiini and each constituent genus are recovered as monophyletic. However, within Coelonia, there is ambiguity in that the sister‐species relationships C. brevis+C. fulvinotata and C. fulvinotata + C. solani are equally parsimonious under both weighting schemes. Furthermore, under equal weighting Agrius is placed equally parsimoniously as the sister group of either Acherontia + Coelonia or Callosphingia. Under implied weighting, however, only the latter relationship is most parsimonious (fit). Within Acherontia, A. atropos and A. styx are always recovered as sister species to the exclusion of A. lachesis. The results of the phylogenetic analysis provide an objective basis for future studies of the unique cleptoparasitic association of these moths.     

Intraspecific trait variability and community assembly in hawkmoths (Lepidoptera: Sphingidae) across an elevational gradient in the eastern Himalayas,India

1. We investigated some aspects of hawkmoth community assembly at 13 elevations along a 200‐ to 2770‐m transect in the eastern Himalayas, a little studied biodiversity hot spot of global importance. We measured the morphological traits of body mass, wing loading, and wing aspect ratio of 3,301 free‐ranging individuals of 76 species without having to collect or even constrain them. We used these trait measurements and T‐statistic metrics to assess the strength of intracommunity (“internal") and extra‐community (“external”) filters which determine the composition of communities vis‐a‐vis the regional pool of species.
2. The trait distribution of constituent species turned out to be nonrandom subsets of the community‐trait distribution, providing strong evidence for internal filtering in all elevational communities. The external filter metric was more ambiguous. However, the elevational dependence of many metrics including that of the internal filter provided evidence for external (i.e., environmental) filtering. On average, a species occupied as much as 50%–75% of the total community‐trait space, yet the T‐statistic metric for internal filter was sufficiently sensitive to detect a strong nonrandom structure in the trait distribution.
3. We suggest that the change in T‐statistic metrics along the environmental gradient may provide more clues to the process of community assembly than previously envisaged. A large, smoothly varying and well‐sampled environmental span would make it easier to discern them. Developing T‐statistics for combined analysis of multiple traits will perhaps provide a more accurate picture of internal/filtering and niche complementarity. Moths are a hyperdiverse taxon and a very important component of many ecosystems. Our technique for accurately measuring body and wing dimensions of free‐ranging moths can generate trait database for a large number of individuals in a time‐ and resource‐efficient manner for a variety of community assembly studies using this important taxon.

     

Effect of sound signals on spontaneous activity in grasshopper interneurons (Orthoptera,Tettigonidae)

According to one of hypotheses proposed for explaining mechanisms of sound signal recognition in insects, their CNS contains a group of rhythmically active neurons that function as a reference standard for comparison with perceived acoustic information. To check this hypothesis, the spontaneous neuronal activity and its changes in perception of conspecific and heterospecific signals (CS and HS) were analyzed in the CNS of two sympatric grasshopper species Tettigonia cantans and Metrioptera roeselii. The activity of individual neurons was assayed in fixed and freely moving insects. The results of the experiment have shown that in the thoracic part of the CNS there is a group of rhythmically active neurons that do not directly respond to sound signal but readjust their impulses under effect of its action. On presentation of CS the following reactions were observed: attenuation or enhancement of impulses; stabilization or destabilization of rhythm; regular increase or decrease in interspike intervals; phasic readjustments leading to synchronization of impulses with sound stimuli (pulses). No similar alterations were usually produced by HS; still, if they did appear, they were less pronounced or of opposite direction. These data indicate that the grasshopper auditory system affects markedly the rhythmically active neurons, their reaction depending considerably on temporal organization of sound signals. Selectivity of these reactions allows us to suggest that the rhythmically active neurons are directly related to the neuronal networks providing the sound signal recognition.Translated from Zhurnal Evolyutsionnoi Biokhimii i Fiziologii, Vol. 40, No. 6, 2004, pp. 531–538.Original Russian Text Copyright © 2004 by Zhantiev, Korsunovskaya, Chukanov.To the 100-Anniversary of A. K. VoskresenskayaThis revised version was published online in April 2005 with a corrected cover date.     

Effects of l-canavanine on arginine catabolism in manduca sexta (sphingidae: lepidoptera)

• 1.1. Hemolymph ornithine concentrations in tobacco horn worm larvae fed a 2.5 mM l-canavanine plus 25 mM l-arginine-supplemented artificial diet (CAAM) were higher than those in larvae fed diets supplemented with 2.5 mM canavanine (CAV), 25 mM arginine (ARG), or controls (CON).
• 2.2. Ornithine concentrations in CAV-treated larvae were significantly greater than the control or ARG treatment, but less than the CAAM treatment during the latter part of the wandering larval stage and during the pharate pupal stage.
• 3.3. Urea concentrations were greater during the active feeding stage with the CAAM- and ARG-treated larvae having significantly higher levels than control or CAV-treated larvae.
• 4.4. Urea concentrations in all treatments never exceeded 36.5% of the ornithine concentration.
• 5.5. Canavanine concentrations were higher in CAV-treated larvae than in CAAM-treated larvae.
• 6.6. During active feeding, arginine concentrations for all treatments were similar, but were lower in CAV- and CAAM-treated larvae during the pharate pupal stage.

     

The phylogenetic relationships of Chalcosiinae (Lepidoptera, Zygaenoidea, Zygaenidae)

The chalcosiine zygaenid moths constitute one of the most striking groups within the lower-ditrysian Lepidoptera, with highly diverse mimetic patterns, chemical defence systems, scent organs, copulatory mechanisms, hostplant utilization and diapause biology, plus a very disjunctive biogeographical pattern. In this paper we focus on the genus-level phylogenetics of this subfamily. A cladistic study was performed using 414 morphological and biochemical characters obtained from 411 species belonging to 186 species-groups of 73 genera plus 21 outgroups. Phylogenetic analysis using maximum parsimony leads to the following conclusions: (1) neither the current concept of Zygaenidae nor that of Chalcosiinae is monophyletic; (2) the previously proposed sister-group relationship of Zygaeninae + Chalcosiinae is rejected in favour of the relationship (Zygaeninae + ((Callizygaeninae + Cleoda ) + ( Heteropan + Chalcosiinae))); (3) except for the monobasic Aglaopini, none of the tribes sensu Alberti (1954 ) is monophyletic; (4) chalcosiine synapomorphies include structures of the chemical defence system, scent organs of adults and of the apodemal system of the male genitalia. A paired metathoracic androconial organ and a series of abdominal tergal corematal organs have been discovered, both being new to Lepidoptera. Due to highly homoplastic patterns in copulatory structures and wings that demonstrate significant sexual dimorphism, polymorphism and mimicry, 17 of the 69 'true' chalcosiine genera ( c . 25%) are shown to be either paraphyletic or polyphyletic. The present classification is therefore very misleading. Reductions of various parts of the male genitalia in some groups are accompanied by morphological and functional replacement involving the 8th abdominal segment. A prominent but convergent lock and key mechanism is revealed.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 161–341.     

Variation and repeatability of ultrasonic sexual advertisement signals inAchroia grisella (Lepidoptera: Pyralidae)

MaleAchroia grisella (Lepidoptera: Pyralidae) produce ultrasonic advertisement signals for pair formation. The signals are generated by wing-fanning and consist of pairs of approximately 100-μs pulses of 70- to 130-kHz sound. In a previous study, playback experiments showed that femaleA. grisella preferred signals whose pulses were louder, longer, and delivered at a higher rate and included longer silent (asynchrony) intervals within pairs. To understand the variation and repeatability of these signal characters, acoustic signals of 18A. grisella males were recorded twice daily on the first 4 days and the seventh day after adult eclosion. ANOVA and MANOVA revealed significant among-individual and among-age variation for all signal characters. Peak amplitude, average amplitude, rate, and pulse length peaked 2–3 days after eclosion and then decreased. Asynchrony interval, however, continued to increase as males aged. Repeatabilities of peak amplitude, average amplitude, rate, and asynchrony interval were all relatively high (>0.5).     

Wallace’s line revisited: has vicariance or dispersal shaped the distribution of Malesian hawkmoths (Lepidoptera: Sphingidae)?

Large-scale biogeographical patterns of Sphingidae in the Malesian archipelago are investigated, using Geographic Information System-supported estimates of species ranges determined from a compilation of specimen-label databases. Distribution maps for all species and checklists for 114 islands are presented at http://www.sphingidae-sea.biozentrum.uni-wuerzburg.de . Phenetic analyses of island faunas reveal 'textbook'-like patterns of biogeographical associations in the region, with Wallace's line emerging as the main faunal discontinuity in the region. Further analyses, using partial Mantel statistics, indicate that historical features of geography (such as land bridges formed during periods of lower sea levels) are an important determinant of faunal similarity, but recent dispersal can also explain a significant portion of present day hawkmoth distributions. Faunal turnover in relation to distance between islands is steeper for Smerinthinae than for other subfamilies, indicating different dispersal abilities among higher taxonomic groups. These differences are discussed in the light of life-history differences between the subfamilies. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 455–468.     

The hemocytes of Hyalophora cecropia (Lepidoptera)

The hemocytes of selected stages of Hyalophora cecropia from first instar larvae to four-day-old adults were examined and compared with those of Samia cynthia and Antheraea polyphemus. Five classes and two subclasses of hemocytes are described in these moths: (1) prohemocytes, (2) plasmatocytes (of several morphological types), (3) spherule cells, (4) adipohemocytes (two subclasses), and (5) oenocytoids. All types except oenocytoids and subclass II adipohemocytes, are found in all stages examined. Mitotic figures were common among prohemocytes of most stages, but were seen only rarely among plasmatocytes and adipohemocytes, and were not seen among spherule cells or oenocytoids. Prohemocytes and plasmatocytes often contain lipid but rarely PAS positive material. Spherules of spherule cells are PAS positive, as are occasional cytoplasmic inclusions of oenocytoids. Adipohemocytes of both subclasses contain lipid and PAS positive materials in all stages examined. Adipohemocytes and plasmatocytes proved to be most active in phagocytizing ink. Relationships between hemocytes of these and other insects, and some possible functions of hemocytes are discussed.     

The Disphragis notabilis (Schaus) species-group in Costa Rica (Lepidoptera,Notodontidae)

The four described taxa in the Disphragis notabilis (Schaus) species-group are reviewed, including the types and their dissected genitalia. Disphragis hemicera (Schaus), stat. rev., is elevated to species rank, D. normula (Dognin) is retained as a synonym of D. notabilis, D. sobolis Miller is confirmed as distinct from D. hemicera, and D. bifurcata sp. n., is newly described. Both D. hemicera and D. bifurcata occur in Costa Rica. The known ranges of the other species are outlined. Defining characters of each species are presented and a key to species is provided. Unusual variation in the genitalia is noted.     

The adhesive devices in larvae of Lepidoptera (Insecta, Pterygota)

Adhesion to smooth surfaces by means of thin fluid lipid film was studied on living larvae of 71 species of Lepidoptera by a simple ”light reflection method”. The method made it possible to localize exactly the sites of adhesion and to estimate roughly the film thickness, within a certain range. Furthermore, it revealed the general presence of mobile lipid on the entire insect surface. The observations on living larvae were complemented by comparative structural studies of the adhesive parts with light and scanning electron microscopes on preserved specimens of 161 species. Specialized adhesive devices were found in great diversity on larval legs and prolegs, especially in larvae living in the open air on their food plants. Two main surface types of adhesive cuticle were found: (1) cuticle with a flexible smooth surface and (2) cuticle with very numerous small projections (microtrichia) with spatulate and recurved apices. Both the functional implications of the adhesive cuticular structure and the role of the adhesive fluid as well as the evolution of the adhesive devices are discussed. The adhesive effect is due to ”capillary” or meniscus forces. Accepted: 29 July 1999