Checklist of the helminth parasites of vertebrates in Costa Rica

Helminth parasites of vertebrates have been studied in Costa Rica for more than 50 years. Survey work on this group of parasites is far from complete. We assembled a database with all the records of helminth parasites of wild and domestic vertebrates in Costa Rica. Information was obtained from different sources such as literature search (all published accounts) and parasite collections. Here we present a checklist with a parasite-host list as well as a host-parasite list. Up to now, 303 species have been recorded, including 81 species of digeneans, 23 monogeneans, 63 cestodes, 12 acanthocephalans, and 124 nematodes. In total, 108 species of vertebrates have been studied for helminths in Costa Rica (31 species of fishes, 7 amphibians, 14 reptiles, 20 birds, and 36 mammals). This represents only 3.8% of the vertebrate fauna of Costa Rica since about 2,855 species of vertebrates occur in the country. Interestingly, 58 species (19.1 %) were recorded as new species from Costa Rica and most of them are endemic to particular regions. Considering the valuable information that parasites provide because it is synergistic with all the information about the natural history of the hosts, helminth parasites of vertebrates in Costa Rica should be considered within any initiatives to accomplish the national inventory of biological resources. Starting with this compilation work, the Colección de Helmintos de Costa Rica (CHCR), hosted at the Facultad de Microbiología, Universidad de Costa Rica, has re-emerged and it is our hope that it will have the standards of quality to assure that it will become the national depository of helminths in the country.     

常州地区蝶类资源及区系组成

在1991～2002年间,笔者对江苏常州地区蠊类资源进行了调查,共采集到蠊类64种(或亚种),隶属于8科46属。其中古北种14种,东洋种10种,古北—东洋共有种37种,广布种3种,分别占总数的21．9％、15．6％、57．8％和4．7％,表明该地区蝴蝶以古北、东洋混生种类为主要成份,并且有由古北区向东洋区过渡的特征。同时还记述了各种蝴蝶的学名和寄主。调查还发现江苏省新记录1属10种．     

Identification of butterfly species with a single neural network system

Growing interest in conservation and biodiversity increased the demand for accurate and consistent identification of biological objects, such as insects, at the level of individual or species. Among the identification issues, butterfly identification at the species level has been strongly addressed because it is directly connected to the crop plants for human food and animal feed products. However, so far, the widely-used reliable methods were not suggested due to the complicated butterfly shape. In the present study, we propose a novel approach based on a back-propagation neural network to identify butterfly species. The neural network system was designed as a multi-class pattern classifier to identify seven different species. We used branch length similarity (BLS) entropies calculated from the boundary pixels of a butterfly shape as the input feature to the neural network. We verified the accuracy and efficiency of our method by comparing its performance to that of another single neural network system in which the binary values (0 or 1) of all pixels on an image shape are used as a feature vector. Experimental results showed that our method outperforms the binary image network in both accuracy and efficiency.     

Responses of butterflies to twentieth century climate warming: implications for future ranges

We analyse distribution records for 51 British butterfly species to investigate altitudinal and latitudinal responses to twentieth century climate warming. Species with northern and/or montane distributions have disappeared from low elevation sites and colonized sites at higher elevations during the twentieth century, consistent with a climate explanation. We found no evidence for a systematic shift northwards across all species, even though 11 out of 46 southerly distributed species have expanded in the northern part of their distributions. For a subset of 35 species, we model the role of climate in limiting current European distributions and predict potential future distributions for the period 2070-2099. Most northerly distributed species will have little opportunity to expand northwards and will disappear from areas in the south, resulting in reduced range sizes. Southerly distributed species will have the potential to shift northwards, resulting in similar or increased range sizes. However, 30 out of 35 study species have failed to track recent climate changes because of lack of suitable habitat, so we revised our estimates accordingly for these species and predicted 65% and 24% declines in range sizes for northern and southern species, respectively. These revised estimates are likely to be more realistic predictions of future butterfly range sizes.     

Higher mobility of butterflies than moths connected to habitat suitability and body size in a release experiment

Mobility is a key factor determining lepidopteran species responses to environmental change. However, direct multispecies comparisons of mobility are rare and empirical comparisons between butterflies and moths have not been previously conducted. Here, we compared mobility between butterflies and diurnal moths and studied species traits affecting butterfly mobility. We experimentally marked and released 2011 butterfly and 2367 moth individuals belonging to 32 and 28 species, respectively, in a 25 m × 25 m release area within an 11‐ha, 8‐year‐old set‐aside field. Distance moved and emigration rate from the release habitat were recorded by species. The release experiment produced directly comparable mobility data in 18 butterfly and 9 moth species with almost 500 individuals recaptured. Butterflies were found more mobile than geometroid moths in terms of both distance moved (mean 315 m vs. 63 m, respectively) and emigration rate (mean 54% vs. 17%, respectively). Release habitat suitability had a strong effect on emigration rate and distance moved, because butterflies tended to leave the set‐aside, if it was not suitable for breeding. In addition, emigration rate and distance moved increased significantly with increasing body size. When phylogenetic relatedness among species was included in the analyses, the significant effect of body size disappeared, but habitat suitability remained significant for distance moved. The higher mobility of butterflies than geometroid moths can largely be explained by morphological differences, as butterflies are more robust fliers. The important role of release habitat suitability in butterfly mobility was expected, but seems not to have been empirically documented before. The observed positive correlation between butterfly size and mobility is in agreement with our previous findings on butterfly colonization speed in a long‐term set‐aside experiment and recent meta‐analyses on butterfly mobility.     

Insect extinctions on a small denuded Bornean island

We report odonate and butterfly extinctions on Pulau Mengalum (Sabah) between 1928 and 2007. Pulau Mengalum has lost all of its closed-canopy forest; 55.5% of the odonates and 40% of the butterflies present in 1928 have likely been extirpated. Fourteen and five species of odonates and butterflies found by us were new records for the island, respectively. It is unclear if newly recorded species were missed by previous surveyors in 1928 or if they have colonized the island after that time. While our study indicates that deforestation is a serious threat to tropical insects, it remains unclear if deforested areas provide opportunities for new colonists.     

Progressive bias in species status is symptomatic of fine-grained mapping units subject to repeated sampling

Atlas maps of butterflies invariably fail to distinguish the status of records, that is, whether they are observations of breeding populations or vagrant individuals. Yet, for conservation purposes, it is clearly important to know whether records relate to breeding populations in suitable habitats or not. The high mobility of butterfly adults carries two expectations. First, vagrants will frequently be recorded within grid squares, particularly when mapping is fine grained. Second, the frequency of vagrants is likely to be higher in mapping units comprising biotopes with few butterfly habitats, than in those rich in butterfly habitats, if only because there is a greater regional pool of potential vagrants for biotopes depleted in butterfly habitats. It follows that with repeated sampling of squares or biotopes differences in species breeding status will be cumulative between those rich and poor in butterfly habitats. In this paper, these predictions have been tested using data from 30, 1 ha grid squares on Alderley Edge Site of Special Scientific Interest (SSSI) (UK), half of which are in woodland (few butterfly habitats) and half with open habitats (non-woodland; rich in butterfly habitats). Butterflies have been recorded during 38 visits over 4 years. Within squares, vagrants have been distinguished from breeding species on the basis of the presence of larval hostplants and other suitable resources for seasonal maintenance. Owing to the occurrence of vagrants, number of breeding species was found to be significantly fewer than total species in mapping units. Vagrants were found to be significantly more abundant in woodland squares, which contain few species larval hostplants, than in squares dominated by open biotopes with many species larval hostplants. Repeated sampling resulted in a cumulative increase in the frequency of vagrant species. Furthermore, the relative frequency of vagrants increased at a greater rate in woodland squares compared to non-woodland squares. Suggestions are made as to how recording for atlases may be improved.     

Negative relationships between species richness and evenness render common diversity indices inadequate for assessing long-term trends in butterfly diversity

Species richness and evenness, the two principle components of species diversity, are frequently used to describe variation in species assemblages in space and time. Compound indices, including variations of both the Shannon–Wiener index and Simpson’s index, are assumed to intelligibly integrate species richness and evenness into all-encompassing measures. However, the efficacy of compound indices is disputed by the possibility of inverse relationships between species richness and evenness. Past studies have assessed relationships between various diversity measures across survey locations for a variety of taxa, often finding species richness and evenness to be inversely related. Butterflies are one of the most intensively monitored taxa worldwide, but have been largely neglected in such studies. Long-term butterfly monitoring programs provide a unique opportunity for analyzing how trends in species diversity relate to habitat and environmental conditions. However, analyzing trends in butterfly diversity first requires an assessment of the applicability of common diversity measures to butterfly assemblages. To accomplish this, we quantified relationships between butterfly diversity measures estimated from 10 years of butterfly population data collected in the North Saskatchewan River Valley in Edmonton, Alberta, Canada. Species richness and evenness were inversely related within the butterfly assemblage. We conclude that species evenness may be used in conjunction with richness to deepen our understandings of assemblage organization, but combining these two components within compound indices does not produce measures that consistently align with our intuitive sense of species diversity.     

Conservation of moths in The Netherlands: population trends, distribution patterns and monitoring techniques of day-flying moths

The loss of butterfly diversity in north-western Europe has been studied extensively but much less is known about the status of other macrolepidoptera. The present paper presents results of the research on mainly day-flying larger moths carried out by Dutch Butterfly Conservation in The Netherlands. Research started in 1992 with a public campaign and a mapping programme, leading to an increase in the number of records of macrolepidoptera. Results from the mapping programme on day-flying moths showed a strong general decrease in The Netherlands, comparable with the loss of butterfly diversity. Transect counts have been undertaken since 2000 for a few species and this method is sufficient to measure trends over time if the moth densities are high enough. Another research component was to focus on marshland habitats in The Netherlands. Fifty-five characteristic marshland moth species were selected and the relative abundance of nearly all these species has declined. These trends were compared with four other countries in the Atlantic biogeographical zone. The relative abundance of the majority of marshland moths in The Netherlands was found to be higher than those in other European countries, so that The Netherlands has a special responsibility for the conservation of the moths of its marshes. Moreover, we emphasise that, next to butterflies, moths can be of great use in nature conservation because of their potential significance as indicator species in endangered habitats.     

The Butterflies of Barro Colorado Island,Panama: Local Extinction since the 1930s

Few data are available about the regional or local extinction of tropical butterfly species. When confirmed, local extinction was often due to the loss of host-plant species. We used published lists and recent monitoring programs to evaluate changes in butterfly composition on Barro Colorado Island (BCI, Panama) between an old (1923–1943) and a recent (1993–2013) period. Although 601 butterfly species have been recorded from BCI during the 1923–2013 period, we estimate that 390 species are currently breeding on the island, including 34 cryptic species, currently only known by their DNA Barcode Index Number. Twenty-three butterfly species that were considered abundant during the old period could not be collected during the recent period, despite a much higher sampling effort in recent times. We consider these species locally extinct from BCI and they conservatively represent 6% of the estimated local pool of resident species. Extinct species represent distant phylogenetic branches and several families. The butterfly traits most likely to influence the probability of extinction were host growth form, wing size and host specificity, independently of the phylogenetic relationships among butterfly species. On BCI, most likely candidates for extinction were small hesperiids feeding on herbs (35% of extinct species). However, contrary to our working hypothesis, extinction of these species on BCI cannot be attributed to loss of host plants. In most cases these host plants remain extant, but they probably subsist at lower or more fragmented densities. Coupled with low dispersal power, this reduced availability of host plants has probably caused the local extinction of some butterfly species. Many more bird than butterfly species have been lost from BCI recently, confirming that small preserves may be far more effective at conserving invertebrates than vertebrates and, therefore, should not necessarily be neglected from a conservation viewpoint.     

Nestedness of butterfly assemblages in the Zhoushan Archipelago,China: area effects,life-history traits and conservation implications

The nested subset pattern (nestedness) of faunal assemblages has been a research focus in the fields of island biogeography and conservation biology in recent decades. However, relatively few studies have described nestedness in butterfly assemblages in oceanic archipelago systems. Moreover, previous studies often quantified nestedness using inappropriate nestedness metrics and random fill algorithms with high Type I errors. The aims of this study are to examine the existence of nestedness and underlying causal mechanisms of butterfly assemblages in the Zhoushan Archipelago, China. We used the line-transect method to determine butterfly occupancy and abundance on 42 study islands from July to August 2014. We obtained butterfly life-history traits (wingspan, body weight and minimum area requirement) by field work and island geographical features (area and isolation) from the literature. We used the recently developed metric WNODF to estimate nestedness. Partial Spearman rank correlation was used to evaluate the associations of nestedness and island geographical features as well as butterfly life-history traits related to species extinction risk and colonization ability. The butterfly assemblages were significantly nested. Island area and minimum area requirement of butterflies were significantly correlated with nestedness after controlling for other independent variables. In contrast, the nestedness of butterflies did not appear to result from passive sampling or selective colonization. However, multi-year studies are needed to confirm that target effects are not muddling these results. Our results indicate that selective extinction may be the main driver of nestedness of butterfly assemblages in our study system. From a conservation viewpoint, we should protect both large islands and species with large area requirement to maximize the number of species preserved.     

Museum genomics reveals the Xerces blue butterfly (Glaucopsyche xerces) was a distinct species driven to extinction

The last Xerces blue butterfly was seen in the early 1940s, and its extinction is credited to human urban development. This butterfly has become a North American icon for insect conservation, but some have questioned whether it was truly a distinct species, or simply an isolated population of another living species. To address this question, we leveraged next-generation sequencing using a 93-year-old museum specimen. We applied a genome skimming strategy that aimed for the organellar genome and high-copy fractions of the nuclear genome by a shallow sequencing approach. From these data, we were able to recover over 200 million nucleotides, which assembled into several phylogenetically informative markers and the near-complete mitochondrial genome. From our phylogenetic analyses and haplotype network analysis we conclude that the Xerces blue butterfly was a distinct species driven to extinction.     

Vulnerability and body size: tetrapods versus fish

The relationship between body size and risk of extinction has been the focus of much recent attention; however, it has only been stated for a few groups inhabiting restricted areas. Based on the IUCN list and other public records, we suggest that the relationship between vulnerability and body size is a trend among all vertebrates, especially tetrapods. In the case of fish, our results show that body size may be subordinate to another factor that we named concealment. Other plausible explanations about this difference could be the human preference for large terrestrial vertebrates and/or the inappropriateness of the current IUCN listing criteria for fish species.     

Age-related changes in thoracic mass: possible reallocation of resources to reproduction in butterflies

In nectar-feeding butterflies, reproductive potential is usually thought to depend on the size of the reproductive reserves in the abdomen, the adult food quality and, for females, the amount of resources received in the spermatophores at mating. Recent findings show that thorax mass and nitrogen content decrease with age in some butterfly species, and that thorax resources may be used for reproduction in the butterfly Pieris napi , just as in some other insects. In order to determine whether this is a general pattern and ascertain how it relates to the investment of resources in reproduction we studied the dynamics of thorax and abdomen mass changes in 11 Swedish butterfly species. By regressing thorax and abdomen mass on age of field-collected specimens, we show that loss of mass from both the thorax and the abdomen is a common phenomenon among nectar-feeding temperate zone butterflies under natural conditions. We argue that our results indicate that resources from flight muscles can be reallocated to reproduction by these butterflies, thus increasing their reproductive potential. Within species, females use proportionately more resources from the thorax than do males, as expected from the difference in investment of resources in reproduction. Among males we expect species with a higher reproductive investment to have a larger decrease in thorax and abdomen mass, and our data indicate that this is the case. Looking at the change in relative thorax mass, our results suggest that the use of resources from the thorax does not affect flight performance negatively, something that could constrain the use of muscle resources.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 363–380.     

Assessing species’ habitat associations from occurrence records,standardised monitoring data and expert opinion: A test with British butterflies

Accurate knowledge of species’ habitat associations is important for conservation planning and policy. Assessing habitat associations is a vital precursor to selecting appropriate indicator species for prioritising sites for conservation or assessing trends in habitat quality. However, much existing knowledge is based on qualitative expert opinion or local scale studies, and may not remain accurate across different spatial scales or geographic locations. Data from biological recording schemes have the potential to provide objective measures of habitat association, with the ability to account for spatial variation. We used data on 50 British butterfly species as a test case to investigate the correspondence of data-derived measures of habitat association with expert opinion, from two different butterfly recording schemes. One scheme collected large quantities of occurrence data (c. 3 million records) and the other, lower quantities of standardised monitoring data (c. 1400 sites). We used general linear mixed effects models to derive scores of association with broad-leaf woodland for both datasets and compared them with scores canvassed from experts.Scores derived from occurrence and abundance data both showed strongly positive correlations with expert opinion. However, only for occurrence data did these fell within the range of correlations between experts. Data-derived scores showed regional spatial variation in the strength of butterfly associations with broad-leaf woodland, with a significant latitudinal trend in 26% of species. Sub-sampling of the data suggested a mean sample size of 5000 occurrence records per species to gain an accurate estimation of habitat association, although habitat specialists are likely to be readily detected using several hundred records. Occurrence data from recording schemes can thus provide easily obtained, objective, quantitative measures of habitat association.     

Newly arrived or previously overlooked: is there evidence for climate-driven changes in the distribution of molluscs in the Barents Sea?

In recent years, there have been frequent reports of invertebrate species newly recorded from particular areas of the Northeastern Atlantic, and it has often been suggested that these are the result of changes in species ranges due to recent warming. These suggestions make three assumptions: (1) that we have a good knowledge of the fauna of these areas; (2) that new records of “southern” species are more frequent than new records of “northern” species; (3) that climate change is the only factor affecting species range. I tested these assumptions on published records of 30 benthic molluscan species which have been found alive for the first time in the Russian part of the Barents Sea since 2006. Some of the discussed species are warm-water species and may have extended their ranges northward in response to climate change. However, our baseline knowledge of the molluscan fauna of this area before 2006 is limited by the frequent lack of molluscan specialists to study the available material, by the frequent lack of detailed publication and by changes in sampling and processing methods. New records of “southern” species are in fact not significantly commoner than new records of “northern” species. Also reasons other than climate change for observed changes in species distribution should be considered.     

Addendum to the checklist of the helminth parasites of vertebrates in Costa Rica

In this work, we present an addendum to the "Checklist of the helminth parasites of vertebrates in Costa Rica" with a parasite-host list as well as a host-parasite list. This addendum updates the available information on this group of parasites in Costa Rica, since very recently a new input has been made to describe the helminth fauna of vertebrates, particularly at the Area de Conservación de Guanacaste. In this paper, we add 33 records, representing 23 species. This raises the number of helminth species described in vertebrates from Costa Rica to 325, represented by 89 species of digeneans, 23 of monogeneans. 63 of cestodes, 13 of acanthocephalans, and 137 of nematodes. In total, 133 species of vertebrates have been studied for helminths in Costa Rica (31 species of fishes. 7 amphibians, 18 reptiles, 40 birds, and 37 mammals). Currently, 67 species (20.6 %) have been recorded as new species from Costa Rica and most of them are endemic to particular regions. The Colecci6n de Helmintos de Costa Rica, housed at the Universidad de Costa Rica (UCR), San José, is the depositary of only 23% of the species recorded so far; however this situation is changing and people recognizes.     

Frugivorous butterfly species in tropical forest fragments: correlates of vulnerability to extinction

Insects exhibit a variety of population-level responses to forest fragmentation, ranging from population increase to extinction. However, the biological attributes that underlie differences in extinction vulnerability among insects have been little-studied. Using the frugivorous butterfly community of tropical dry forest in Venezuela, we studied body size, population density and colonization ability as attributes that might underlie the range of responses of insects to forest fragmentation. The study was carried out in a set of forest fragments in the reservoir Lago Guri, formed by the damming of the Caroni River in eastern Venezuela. Results show that larger butterfly species were more vulnerable to extinction from habitat fragments than smaller ones. Rarer species were not more vulnerable to extinction, showing that rarity may not be an important correlate of vulnerability to extinction amongst insects. Contrary to expectation, faster-flying species were more and not less vulnerable to extinction from small habitat fragments. We speculate on the possible reasons for the observed patterns in extinction vulnerability using additional observations on behavioural patterns and larval host plant distributions of some of the butterfly species.     

Ithomiini Butterflies (Lepidoptera: Nymphalidae) of Antioquia, Colombia

Colombia is one of the most biodiverse countries on the planet. However, economic and scientific investment in completing inventories of its biodiversity has been relatively poor in comparison with other Neotropical countries. Butterflies are the best studied group of invertebrates, with the highest proportion of known to expected species. More than 3,200 species of butterflies have been recorded in Colombia, although the study of the still many unexplored areas will presumably increase this number. This work provides a list of Ithomiini butterflies collected in the department of Antioquia and estimates the total number of species present, based on revision of entomological collections, records in the literature and field work performed between 2003 and 2011. The list includes 99 species and 32 genera, representing 27% of all Ithomiini species. We report 50 species of Ithomiini not formerly listed from Antioquia, and found the highest diversity of ithomiine species to be at middle elevations (900–1,800 m). The mean value of the Chao2 estimator for number of species in Antioquia is 115 species, which is close to a predicted total of 109 based on known distributions of other Ithomiini not yet recorded from the department. Nine species are potentially of particular conservation importance because of their restricted distributions, and we present range maps for each species. We also highlight areas in Antioquia with a lack of biodiversity knowledge to be targeted in future studies. This paper contributes to mapping the distribution of the Lepidoptera of Antioquia department in particular and of Colombia in general.