Population trends of Finnish butterflies (Lepidoptera: Hesperioidea, Papilionoidea) in 1991–2000

The National Butterfly Recording Scheme in Finland (NAFI) is based on data collected using a uniform questionnaire for the whole country. During the first decade of the scheme (1991–2000), a total of 432 voluntary amateur and professional lepidopterists participated by providing data on 1.5 million individuals representing 94 indigenous and 11 non-resident species. Although the 10-year period is not long enough to provide extensive conclusions about changes in the fauna, changes in either the geographical distributions or population densities exhibited a downward trend for 15 species and an upward trend for 11 species. The decreased species were dominated by those inhabiting bogs and fens (4) and open sandy or rocky habitats (5), whereas the increased species were mainly typical of forest verges and clearings (8). The results indicated in particular the effects of drainage of peatlands and overgrowth of meadows. On the other hand, the distribution maps published in the national bulletin revealed northward expansion for seven species, which, together with some increasing migrants, may indicate the effects of possible climatic warming. By providing quantitative knowledge of possible changes in the distribution and abundance of butterflies, NAFI may be used to protect the Finnish butterfly fauna.     

Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae)

The historical biogeography of insects in South America is largely unknown, as dated phylogenies have not been available for most groups. We have studied the phylogenetic relationships and historical biogeography of a subtribe of butterflies, Phyciodina in the family Nymphalidae, based on one mitochondrial gene (COI) and two nuclear gene regions (EF-1alpha and wingless). The subtribe comprises 89 species mainly found in tropical South America, with a few species in North America and the Greater Antilles. We find that the enigmatic genus Antillea is sister to the rest of Phyciodina, and suggest that it should be included in the subtribe. Several genera are found to be polyphyletic or nested within another genus, and are proposed to be synonymised. These are Dagon, Castilia, Telenassa and Janatella, which we propose should be synonymised with Eresia. Brazilian "Ortilia" form an independent lineage and require a new genus name. The diversification of Phyciodina has probably taken place over the past about 34 MYA. The ancestral phyciodine colonised South America from North America through a possible landspan that connected the Greater Antilles to South America about 34MYA. A vicariance event left the ancestral Antillea on the Greater Antilles, while the ancestral 0e on South America colonised the Guyanan Shield and soon after the Brazilian Shield. We hypothesise that the Brazilian Shield was an important area for the diversification of Phyciodina. From there, the ancestor of Anthanassa, Eresia and Tegosa colonised NW South America, where especially Eresia diversified in concert with the rising of the Andes beginning about 20 MYA. Central America was colonised from NW South America about 15 MYA by the ancestors of Anthanassa and Phyciodes. Our study is the first to use a dated phylogeny to study the historical biogeography of a group of South American species of butterflies.     

Listening for males and bats: spectral processing in the hearing organ of <Emphasis Type="Italic">Neoconocephalus bivocatus</Emphasis> (Orthoptera: Tettigoniidae)

Tettigoniids use hearing for mate finding and the avoidance of predators (mainly bats). Using intracellular recordings, we studied the response properties of auditory receptor cells of Neoconocephalus bivocatus to different sound frequencies, with a special focus on the frequency ranges representative of male calls and bat cries. We found several response properties that may represent adaptations for hearing in both contexts. Receptor cells with characteristic frequencies close to the dominant frequency of the communication signal were more broadly tuned, thus extending their range of high sensitivity. This increases the number of cells responding to the dominant frequency of the male call at low signal amplitudes, which should improve long distance call localization. Many cells tuned to audio frequencies had intermediate thresholds for ultrasound. As a consequence, a large number of receptors should be recruited at intermediate amplitudes of bat cries. This collective response of many receptors may function to emphasize predator information in the sensory system, and correlates with the amplitude range at which ultrasound elicits evasive behavior in tettigoniids. We compare our results with spectral processing in crickets, and discuss that both groups evolved different adaptations for the perceptual tasks of mate and predator detection.     

The eyes of Macrosoma sp. (Lepidoptera: Hedyloidea): a nocturnal butterfly with superposition optics

The visual system of nocturnal Hedyloidea butterflies was investigated for the first time, using light and electron microscopy. This study was undertaken to determine whether hedylids possess the classic superposition eye design characteristic of most moths, or apposition eyes of true butterflies (Papilionoidea), and, to gain insights into the sensory ecology of the Hedyloidea. We show that Macrosoma heliconiaria possesses a superposition-type visual mechanism, characterized by long cylindrical crystalline cones, a lack of corneal processes, 8 constricted retinular sense cells, rhabdoms separated from the crystalline cones forming a translucent 'clear zone', and tight networks of trachea that form a tapetum proximal to the retina and which also surround the rhabdoms to form a tracheal sheath. Dark-adapted individuals of M. heliconiaria, M. conifera, and M. rubidinarea exhibited distal retinular pigment migration, forming an eye glow. Correspondingly, light-exposure induced pigment to migrate proximally, causing the eye glow to be replaced by a dark pseudopupil. Other characteristics of the visual system, including relative eye size, facet size, and external morphology of the optic lobes, are mostly 'moth like' and correlate with an active, nocturnal lifestyle. The results are discussed in relation to the evolution of lepidopteran eyes, and the sensory ecology of this poorly understood butterfly superfamily.     

The pineal gland of nocturnal mammals

Summary In the pineal gland of the pipistrelle bat two different populations of pinealocytes and glial cells were observed electron microscopically. The pinealocytes of populations I and II differ in their content of metabolically active cell organelles. In the pinealocytes of population I, granular vesicles originating from the Golgi apparatus were found in the perikaryon and especially in the endings of the pinealocyte processes. Granular vesicles appeared to be more numerous in hibernating nulliparous females. The pinealocytes of population II are characterized by the presence of small cytoplasmic vacuoles, probably originating from cisternae of the granular endoplasmic reticulum and containing flocculent material of moderate electron density. The classification of the pinealocytes belonging to population II is discussed.This collaboration was initiated with the aid of an SRC European short visit grant to P.A.R.The study was supported by the Foundation for Medical Research, the Netherlands (FUNGO, 13-35-33)     

Rainfall and the interaction of microclimate with larval resources in the population dynamics of checkerspot butterflies (Euphydryas editha) inhabiting serpentine grassland

Summary The interaction of host plant phenology and microclimatic heterogeneity was examined to determine its role in the population dynamics of checkerspot butterflies, Euphydryas editha, inhabiting serpentine grassland in California's outer Coast Range.Within the 2–3 hectares inhabited by a population of E. editha (Jasper Ridge Area H), microclimatic differences resulting from topographic heterogeneity largely determine the temporal and spatial pattern of senescence of the larval host plants, Plantago erecta and Orthocarpus densiflorus. Survival of larvae from hatching to diapause is extremely low as a result of unpredictable variation in the timing of larval development relative to the timing of host plant senescence, both of which are mediated by microclimatic patterns. During this study, population H declined to near extinction as a result of two consecutive years of record rainfall that apparently disrupted the tenuous temporal relationship between larval development and plant senescence. Retarded development of post-diapause larvae led to a late and extended flight season and delayed egg production; this in turn resulted in massive mortality of pre-diapause larvae due to starvation because host plant senescence occurred before larvae became large enough to enter diapause. Adult population size the following spring was the smallest in 25 years of study. This work emphasizes the importance of microclimatic heterogeneity for understanding population-level processes in small ectothermic animals and underlines the potential importance of such heterogeneity in the establishment of reserves designed to protect such animals     

Frequency modulated sound pattern analysis in the lesser bulldog bat: the role of interactions between adjacent frequency elements of complex sounds

A stereotyped approach phase vocalization response of Noctilio albiventris to artificial echoes simulating a virtual approaching object was used to assess the ability of the bat to analyze and extract distance information from the artificial echoes. The performance of the bats depended on the temporal pattern of frequency change of the continuously sweeping frequency modulated (FM) component of the signals. When the bats were presented with a CF/FM signal containing a time-reversed upward FM sweep, they responded with approach phase behavior at a performance level that was significantly below that seen with a CF/FM signal containing a naturally structured downward FM sweep. When the FM sweep was divided into a series of brief pure tone steps, the extent to which the bats showed a difference in their capability to process upward versus downward FM sweeps depended on the difference in frequency between the pure tone steps. The bats effectively processed downward but not upward FM sweeps when the difference in frequency between pure tone frequency elements of the FM sweeps was from about 100–200 Hz, but they effectually processed both downward and upward FM sweeps when the tonal elements composing the FM sweeps were separated by more than about 200 Hz. This suggests that the ability of the bats to effectively process downward but not upward FM sweeps is based on local interactions between adjacent frequency elements of the complex sounds.Abbreviations CF constant frequency - FM frequency modulated     

山西省蝶类多样性与地带分布

结合自然地理与植被条件对山西蝶类多样性分布的现状进行研究,结果表明,其特点是：不同林区差异性大,呈南北递减趋势,受小气候环境影响较大,资源呈片段化分布,耕作区种类多样性丧失严重。根据已知的216种蝶类在山西15个林区的分布情况,采用聚类平均法(UPGMA)对这15个林区的蝶类多样性进行聚类分析,在相似性系数为0.85时,可将山西林区划分为5个蝶类分布地带。按照地理位置和自然景观的不同,分别称之为：Ⅰ.南端山地蝴蝶区;Ⅱ.南部盆地蝴蝶区;Ⅲ.中南部山地丘陵蝴蝶区;Ⅳ.中部高山蝴蝶区;Ⅴ.西北部丘陵蝴蝶区。在关键区系分析中,得知南端山地蝴蝶区和中部高山蝴蝶区是山西蝶类多样性分布的重要地区,对这两个区系加以保护,即可保护山西已知蝴蝶种类的97.7%。     

Host specificity among Maculinea butterflies in Myrmica ant nests

Summary Ecological studies have been made of all 5 European species of Maculinea. These confirm that M. nausithous and M. rebeli live underground in Myrmica ant nests for 10 months of the year, as has long been known for the other 3 species. The main discovery was that each Maculinea species depends on a single, and different, host species of Myrmica. This specificity contradicts previous papers and scientific reviews of the relationship between Maculinea and ants. Therefore, early records are re-examined and 3 reasons are given to explain why most are misleading when applied to wild populations. Dependence on a single, rather than any, species of Myrmica explains why Maculinea populations exist in only a small minority of biotopes where their foodplants and Myrmica ants abound. It also explains the puzzling disappearance of Maculinea populations from apparently suitable sites. The discovery that M. alcon and M. rebeli depend on separate species of Myrmica that are not even closely related strengthens the argument that these butterflies are good species.     

Hox genes are essential for the development of eyespots in Bicyclus anynana butterflies

The eyespot patterns found on the wings of nymphalid butterflies are novel traits that originated first in hindwings and subsequently in forewings, suggesting that eyespot development might be dependent on Hox genes. Hindwings differ from forewings in the expression of Ultrabithorax (Ubx), but the function of this Hox gene in eyespot development as well as that of another Hox gene Antennapedia (Antp), expressed specifically in eyespots centers on both wings, are still unclear. We used CRISPR-Cas9 to target both genes in Bicyclus anynana butterflies. We show that Antp is essential for eyespot development on the forewings and for the differentiation of white centers and larger eyespots on hindwings, whereas Ubx is essential not only for the development of at least some hindwing eyespots but also for repressing the size of other eyespots. Additionally, Antp is essential for the development of silver scales in male wings. In summary, Antp and Ubx, in addition to their conserved roles in modifying serially homologous segments along the anterior–posterior axis of insects, have acquired a novel role in promoting the development of a new set of serial homologs, the eyespot patterns, in both forewings (Antp) and hindwings (Antp and Ubx) of B. anynana butterflies. We propose that the peculiar pattern of eyespot origins on hindwings first, followed by forewings, could be due to an initial co-option of Ubx into eyespot development followed by a later, partially redundant, co-option of Antp into the same network.     

Frequency discrimination of brief tonal steps as a function of frequency in the lesser bulldog bat

In a two-alternative, forced-choice task lesser bulldog bats were trained to distinguish between a pure tone pulse and a pulse composed of a series of brief tonal steps oscillating between two different frequencies. The tone-step pulse gradually approximates the pure tone pulse as the frequency difference between the steps becomes progressively smaller. Frequency difference limens for the brief tonal frequency steps were determined for a broad range of ultrasonic frequencies. The variation in tone-step difference limens with frequency appears to be correlated to the frequency structure of the bat's short-constant-frequency/frequency-modulated echolocation sound. There was a marked decline in the value of the relative frequency difference limens (Weber ratio) over a fairly narrow range of frequencies above the constant frequency and a sharp increase in threshold above this range. The relative thresholds for frequency discrimination were small and uniform over the frequency range of the frequency-modulated sweep and increased for frequencies below the frequency- modulated sweep. Thus, the most accurate frequency-discrimination abilities occur over a narrow frequency range around the frequency of the constant-frequency component of returning echoes. Frequency discrimination over the range of frequencies of the frequency-modulated component is relatively good. Accepted: 20 March 1999     

宁夏贺兰山自然保护区蝴蝶群落多样性及其环境影响因子

宁夏贺兰山自然保护区蝴蝶群落多样性及其与环境因素的关系,2017年5-9月采用样线法对贺兰山东麓6类生境和不同干扰类型10条样线的蝴蝶群落结构及其多样性季节动态进行调查。共记录蝴蝶5科36属45种,蛱蝶科Nymphalidae的属和物种数最多,为17属19种;凤蝶科Papilionidae最少,仅1属1种。菜粉蝶Pieris rapae、云粉蝶Pontia daplidice、斑缘豆粉蝶Colias erate和小檗绢粉蝶Aporia hippia是该地区的优势种,个体数量分别占总个体数的11.76%、11.63%、11.21%和10.17%。不同生境样线优势类群和常见类群不同。蝴蝶的栖息地偏好与寄主植物有关,蝴蝶的生境分布类型可分为生境广布型、湿润平原型、荒漠半荒漠草原型和山地森林型。蝴蝶群落Shannon-Wiener多样性和丰富度指数以灰榆疏林草地生境最高,优势度最低。各物种在生境内的季节变化趋势与不同生境植被生长季节相关,高峰期为7-8月。不同调查时间蝴蝶的优势种和常见种不同。物种数以7月份调查最多,有33种,占全年调查总物种数的73.33%;5月份调查最少,有20种。蝴蝶群落Shannon-Wiener多样性和丰富度指数以8月份最大,5月份最小。蝴蝶成虫发生类型分为全年发生型、春季型、夏季型和夏秋季型。不同生境和季节发生的优势种可以作为对生境状况进行评估的指示类群。采用CCA分析物种分布与微环境因子的关系,海拔对蝴蝶物种多样性分布格局有显著影响。蝴蝶丰富度与海拔、温度、风速显著正相关。适度干扰有利于蝶类多样性增加,较强的人为干扰会影响蝶类栖息环境,降低蝶类多样性。因此,生境差异性和干扰与蝴蝶群落的物种多样性密切相关,维持贺兰山垂直植被带的生境异质性和保持适度干扰是保护蝴蝶多样性的关键。     

Two different sensory mechanisms for the control of pupal protective coloration in butterflies

The butterflies Graphium sarpedon nipponum Fruhstorfer and Papilio xuthus Linné show pupal protective color polymorphism, but the two species appear to have different sensory mechanisms for determining pupal coloration. When light was of sufficient illumination, the larvae of Graphium sarpedon became bright yellowish green pupae on white pupation boards and reddish brown pupae on black pupation boards. The pupal coloration thus strongly depended on the brightness of the pupation site. In addition, larvae became bright yellowish green pupae in complete darkness. From these results, measurement of the illumination suggested that pupal color is determined by the illuminant difference between incidence light from the dorsal direction and ventral light from a paper board; i.e., the sum of the reflected light of the board plus the penetrated light passing through the board. The illuminant difference required for reddish brown coloration was 40 lux or more. The optical signals received through the stemmata during a critical period before formation of the thorax garter (band string) were important for coloration. By contrast, in Papilio xuthus, successive tactile signals from a rough surfaced pupation site during a critical period before and after formation of the garter were important for determining brown pupal coloration.     

Coexistence and host use by a large community of Pierid butterflies: habitat is the templet

Summary We report studies on the butterfly-hostplant communities in the species-rich area of west central Morocco. Pieridae feeding on Capparales form two distinct ecological guilds: inflorescence feeders and folivores. Several members of each guild may synchronously occur in sympatry. Substantial levels of cannibalism and inter-specific predation occur amongst the inflorescence feeders. No evidence was obtained for segregation of butterfy species on different hosts. Host plants included in the diet of specialists were also used by generalists. Despite substantial differences observed in laboratory trials of larval survivorship on different hostplants, results were congruent for all Pieridae, suggesting that little one-to-one insect-host coevolution has occurred. Host specialization was instead related to the year-to-year stability of host numbers in an area. High levels of pierid infestation occurred on host species with numerically stable populations. Host numerical stability was correlated with habitat type. There is little evidence for segregation of competing inflorescence feeders by hostplant species, but some evidence for segregation by habitat type (particularly by shading levels). We interpret our results as indicating that the hostplant affiliations of Moroccan Capparales-feeding Pieridae are subject to (at best) diffuse coevolutionary effects from hosts and competitors, and are strongly influenced by habitat characteristics.     

Host plant defences and voltinism in European butterflies

Abstract.  1. With respect to seasonal availability for herbivores, plants defended by synthesising qualitative compounds differ from those protected by accumulation of quantitative macromolecules, leaf toughness, and low water and/or nutrient content. While the palatability of the former plants remains relatively constant during the season, the palatability of the latter group decreases with leaf age.
2. It was hypothesised that in seasonal temperate environments, quantitative plant defences should restrict the annual numbers of insect generations. To test this hypothesis, European butterflies were used as a model, both non-corrected regressions and tests controlling for phylogeny were carried out, and potentially confounding factors such as body size or occurrence in short-season environments were treated as covariables.
3. Non-phylogenetically controlled regressions corroborated that butterflies feeding on quantitatively protected hosts (woody plants + grasses) form fewer generations than species feeding on qualitatively protected forbs. Plant defences fitted voltinism better than butterfly size, and remained significant even after controlling for short seasons. Using independent contrasts, feeding on woody plants plus grasses, and feeding on woody plants only, predicted fewer generations. These patterns, however, applied exclusively for foliage-feeding species.
4. The association between plant defences and voltinism represents a hitherto overlooked pattern in the ecology of temperate herbivores. It may explain why large insects tend to form fewer generations and feed on structurally complex hosts, and why some species remain monovoltine although they are not restricted by short season.     

Occurrence of Hylonycteris underwoodi (Chiroptera,Phyllostomidae) and Thyroptera tricolor (Chiroptera,Thyropteridae) in Honduras

The geographic distributions of Hylonycteris underwoodi and Thyroptera tricolor in Honduras have been established with specimens collected more than 50 years ago. Here, we describe the occurrence of these bat species captured in the historic city of Ciudad Blanca, Gracias a Dios in northeastern Honduras.     

Modulation rate transfer functions in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>) with normal hearing and high-frequency hearing loss

Envelope following responses were measured in two bottlenose dolphins in response to sinusoidal amplitude modulated tones with carrier frequencies from 20 to 60 kHz and modulation rates from 100 to 5,000 Hz. One subject had elevated hearing thresholds at higher frequencies, with threshold differences between subjects varying from ±4 dB at 20 and 30 kHz to +40 dB at 50 and 60 kHz. At each carrier frequency, evoked response amplitudes and phase angles were plotted with respect to modulation frequency to construct modulation rate transfer functions. Results showed that both subjects could follow the stimulus envelope components up to at least 2,000 Hz, regardless of carrier frequency. There were no substantial differences in modulation rate transfer functions for the two subjects suggesting that reductions in hearing sensitivity did not result in reduced temporal processing ability. In contrast to earlier studies, phase data showed group delays of approximately 3.5 ms across the tested frequency range, implying generation site(s) within the brainstem rather than the periphery at modulation rates from 100 to 1,600 Hz. This discrepancy is believed to be the result of undersampling of the modulation rate during previous phase measurements.