Guide Advances in Ephemeroptera Biology

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Journal of the Washington Academy of Sciences. Filshie, B. Design of an insect cuticle associated with osmoregulation; the porous plates of chloride cells in an mayfly nymph. Tissue Cell. Handlirsch, A. Ordung Ephemerida Leach. Handbuch der Entomologie. Band III Kapitel 9. Harker, J. Australian Ephemeroptera. Part I. Taxonomy of New South Wales species and evaluation of taxonomic characters. The Ephemeroptera of eastern Australia. Transactions of the Royal Entomological Society of London.

Horridge, G. The dorsal eye of the mayfly Atalophlebia Ephemeroptera. Humpesch, U. Egg development of non-diapausing exopterygote aquatic insects occuring in Europe. Hynes, H. The Ecology of Running Waters. Liverpool : Liverpool University Press. Symbiocladius aurifodinae sp.

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Diptera, Chironomidae , a parasite of nymphs of Australian Leptophlebiidae Ephemeroptera. Landa, V. Comparative anatomy of mayfly larvae Ephemeroptera. Acta Entomologica Bohemoslovaca.

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Phylogeny and higher classificatin of the order Ephemeroptera: a discussion from the comparative anatomical point of view. Studie czhsl. Lestage, J. Marchant, R. Life spans of two species of tropical mayfly nymph Ephemeroptera from Magela Creek, Northern Territory. Life histories of some benthic insects from the La Trobe River, Victoria. McCafferty, W. The higher classification of the Ephemeroptera and its evolutionary basis.

Annals of the Entomological Society of America. Nagell, B. Overwintering strategy of Cloeon dipterum L. Needham, J. Pearson, R. First records of Prosopistoma sedlaceki in Australia Ephemeroptera: Prosopistomatidae. Peters, W. This partial regression upheld the conclusion that body size was an important factor in species description rates Table 2.

Most of the species were described from holotypes collected in the southeastern 78 species and northern regions 74 of Brazil, with 43 being recorded in the South, 16 in the Midwest, and 11 in the Northeast. The Northeast was represented by 13 fewer descriptions than the South, on average, and 18 fewer than the Southeast Fig. Values with different letters are significantly different. The extreme model Fig.

As the number of known species is lower than the total number predicted by the asymptote, this model indicates that many new species of Brazilian ephemeropteran have yet to be described. According to the model, in fact, a total of 11, species are predicted to occur in Brazil, of which, 10, have yet to be described. While ecological and taxonomic studies of mayflies have advanced considerably in Brazil over the past few decades, major gaps still exist in the scientific understanding of the diversity of this order Shimano et al.

One of the principal gaps found in the present analysis was the reduced number of descriptions of nymphs in comparison with those available for adults. This predominance of descriptions of adults may be related to the relative simplicity of identifying and describing the adults in comparison with the nymphs, in most ephemeropteran species, given the number of morphological characters in both cases.

This is demonstrated clearly by the matrix of morphological traits used by Nieto for the analysis of South American baetid species, which was based on characters for the nymphs, but only 14 for the adults. The nymphs are often very similar to one another in many morphological parameters, and their descriptions and identification keys often depend on highly specific characteristics, which permit the differentiation of species, but also hamper the whole process of the identification and description of taxa e.

The presence of atrophied mouthparts and vestigial legs in adults, related to reproductive adaptations Da-Silva and Salles, , also contributes to the reduced number of traits available for the description of this stage. Despite this, three families - Baetidae, Leptohyphidae, and Oligoneuriidae - are characterized by a predominance of species descriptions based on the analysis of the nymphs. We believe that this apparent discrepancy can be accounted for by the fact that most of these descriptions were derived from ecological studies, which tend to focus on the nymphs Shimano et al.

Ultimately, the studies of these families may be more refined simply because there are more specialists in Brazil. The similarity in the number of male and female specimens used in the species descriptions probably reflects the efforts of taxonomists to balance the sampling, given that the early studies were based primarily on female specimens. Given this, we believe that the specialists, recognizing this deviation, have spent more time describing males, with the aim of balancing the available data, and subsequently focused more on the males once they realized that the male genitalia provides more diagnostic features for the differentiation of species than that of the females.

Advances in Ephemeroptera Biology

This would have led to the overall equilibrium between the sexes in species descriptions Fig. It is important to note here that the effective classification of taxa depends on the diagnosis of all life stages, given that many structures are modified or become more or less visible during these different stages. In the specific case of the Ephemeroptera, we believe that the lack of data on the characteristics of one life stage or one of the sexes, in the case of the adults, may often result in errors of identification, especially under field conditions, when ecologists, rather than taxonomists identify the specimens.

This may result in shortcomings and misinterpretations in many areas of research Mariano and Polegatto, , given the fundamental need for an adequate classification of organisms Marques and Lamas, The observed tendency for larger-bodied species to be described first was expected, given the limitations of the equipment available historically for the examination of small insects. In fact, the lack of information on the biology of the group and collection procedures may have imposed certain limitations on the early studies.

The first species recorded in Brazil was C. While body size has influenced description rates, other factors may also be important, including the number of researchers involved in studies of ephemeropteran in Brazil, the effectiveness of the materials used historically for collection of specimens and other data, and the lack of information on the distribution of these insects, which is still poorly known in much of the country.

In the older studies, in fact, a little or no information was provided on the habitat in which the specimens were collected, and this alone may have hampered the collection of additional studies for the development of more detailed studies on the taxonomy of a given species or genus. In recent years, taxonomic reviews and re-evaluations have led to changes in the listing of ephemeropteran species, the diagnostic characteristics of the different taxa, and known distributions.

The use of advanced microscopy techniques has permitted the discovery of new species, which are often of relatively small body size. Cooperation between research institutions, both in Brazil and abroad, has also helped to reduce Linnaean and Wallacean deficits, contributing to improvements in the reliability and efficiency of mayfly studies. Body size also has an effect on other fields of research - Orlofske and Baird , for example, found that, in bio-monitoring studies, whereas larger specimens tend to be identified to genus or even species, smaller specimens are typically identified to no more than the family level.

Another aspect of the results is the geographic distribution of the holotypes, in particular, the lack of studies from the Brazilian Northeast, a region that includes nine states and makes up almost a fifth In fact, no holotypes have been described from five of these states, and the total number of species known from this region is highly disproportionate to that of the South and Southeast, which are smaller in area. The paucity of data from the Northeast almost certainly reflects the lack of ephemeropteran researchers based in this region, although in recent years, specialists have become established in the states of Bahia and Pernambuco.

Research in this region began only recently, with the first record of a Baetidae being reported by Lugo-Ortiz et al. This emphasizes the need for further studies in the region, given that most of the area has been surveyed only superficially, when at all. A scarcity of research in the Brazilian Northeast is also typical of other macroinvertebrates, such as the Odonata De Marco and Vianna, By contrast, the largest number of ephemeropteran researchers are based in the Brazilian Southeast, and this is reflected in the numbers of species descriptions.

While fewer researchers are based in the northern region, the vast Amazon basin Ab'Saber, attracts scientists from other regions, and around the world. This does not mean, however, that these other regions of the country have been surveyed adequately, given the vast size of the country and the complexity of its hydrographic network. Unfortunately, the number of researchers and the resources available for the study of ephemeropteran diversity are still far from adequate, given the apparent complexity of this group of insects.

For this reasons, efforts should be focused on the description of species as a basic parameter for the understanding of conservation priorities. The large number of as yet undescribed Brazilian species predicted by the results of the present study was unexpected, however, especially considering that Da-Silva and Salles estimated a total of only species for the whole planet, and only for Brazil, in contrast with 10, species predicted for this country here. It is important to remember, however, that a certain amount of variation was found among the estimates, among both methods non-linear and groups, probably related to the reduced number of known species and the steepness of the curve.

The paucity of data on Brazilian mayflies is a fundamental problem here, and this is reflected in the broad confidence interval recorded in the present study Table 2. Whatever the reliability of the estimate presented here - it will almost certainly be altered as data accumulate - what is certain is that many mayfly species have yet to be described in Brazil.

The factors that contribute to the considerable disparity between this estimate and the number of known species include i the recent development of ephemeropteran research in Brazil, ii the paucity of specialists, iii the lack of financial resources for fieldwork, iv the shortage of government incentives for research, and v the long period of time required to train specialists. In addition, the paucity of scientific collections and taxonomic studies means that most tropical species are still unknown Da-Silva and Salles, Focusing on the zoogeography of the mayflies may contribute to the management of areas that have suffered high rates of loss of biodiversity, as well as elucidating the relationship between these organisms and the environments they inhabit.

In addition to this research, some known species require revision in order to refine the original descriptions. Raising nymphs until the adult stage would also be a way of identifying taxa more precisely. This type of approach would also provide more reliable and informative data for studies in other areas, such as ecology and systematics. We would like to thank two anonymous referees for their constructive comments.

Supplementary data associated with this article can be found, in the online version, at doi Ab'Saber, A.

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