treatments-xml/data/03/F3/1C/03F31C15FFE3B30BB5D511F22661FF54.xml

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<mods:title id="A6690546502DEFBE7BA23213E9805F53">Egg morphology update based on new chorionic data of Potamanthus luteus (Linnaeus), Ephemera danica Müller and Oligoneuriella rhenana (Imhoff) (Insecta, Ephemeroptera) obtained by scanning electron microscopy</mods:title>
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<taxonomicName id="4C5AD680FFE3B308B5D511F224C0FB4F" ID-CoL="496DZ" box="[151,436,1191,1217]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriella" kingdom="Animalia" order="Ephemeroptera" pageId="8" pageNumber="23" phylum="Arthropoda" rank="species" species="rhenana">
<emphasis id="B92E7111FFE3B308B5D511F224C0FB4F" bold="true" box="[151,436,1191,1217]" italics="true" pageId="8" pageNumber="23">Oligoneuriella rhenana</emphasis>
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<paragraph id="8BE5AD03FFE3B308B5D511A72773FA82" blockId="8.[151,1436,1266,1972]" box="[151,519,1266,1292]" pageId="8" pageNumber="23">Description of egg morphology</paragraph>
<paragraph id="8BE5AD03FFE3B308B584104F27A6FA22" blockId="8.[151,1436,1266,1972]" pageId="8" pageNumber="23">
<emphasis id="B92E7111FFE3B308B584104F24E9FABA" bold="true" box="[198,413,1306,1332]" italics="true" pageId="8" pageNumber="23">General features:</emphasis>
Oval-like eggs with slightly pointed poles, morphologically characterized by a wavy chorion surface, although apparently smooth at low magnifications, with many attachment structures which are distributed uniformly (
<figureCitation id="1361B186FFE3B308B491103F2769FA0A" box="[467,541,1386,1412]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="8" pageNumber="23">Fig. 4</figureCitation>
A. B). The eggs of this species are big and their dimensions vary between 264–297 µm in length and 161–208 µm in width
</paragraph>
<paragraph id="8BE5AD03FFE3B308B58410EF243FF83A" blockId="8.[151,1436,1266,1972]" pageId="8" pageNumber="23">
<emphasis id="B92E7111FFE3B308B58410EF2656FA5A" bold="true" box="[198,802,1466,1492]" italics="true" pageId="8" pageNumber="23">Attachment structures and chorionic sculpturing:</emphasis>
These two basic characteristics of eggs must be analyzed jointly in this species, because all the chorionic structures observed externally are related to an attachment function, but at the same time constitute the egg sculpturing, since they cover the egg completely. The chorion surface is a fibrous adhesive layer strongly adhering to the rest of the chorion, whose fibers may be distinguished clearly by SEM from 
<date id="FFE48BC3FFE3B308B700130F27F8F9FA" box="[578,652,1626,1652]" pageId="8" pageNumber="23" value="2000-10">2000x</date>
magnifications. The fibers are formed of short and fine filaments (0.11 μm diameter) arranged in parallel forming bundles (
<figureCitation id="1361B186FFE3B308B60313D726FCF912" box="[833,904,1666,1692]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="8" pageNumber="23">Fig. 4</figureCitation>
C, D). In addition, these bundles of filaments may be related with each other following the same direction or intercrossing (
<figureCitation id="1361B186FFE3B308B17713FF2108F94A" box="[1077,1148,1706,1732]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="8" pageNumber="23">Fig. 4</figureCitation>
B, C), causing different ultrastructural levels of organization in the thickness of the fibrous layer. The fiber arrangement also causes the appearance of different 
<typeStatus id="54E113A1FFE3B308B4AA13AF2752F89A" box="[488,550,1786,1812]" pageId="8" pageNumber="23">types</typeStatus>
of external structure, including small orifices (+2 µm of diameter) and fibrous projections (
<figureCitation id="1361B186FFE3B308B4C6127724BFF8B2" box="[388,459,1826,1852]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="8" pageNumber="23">Fig. 4</figureCitation>
C, D). The egg surface presents many such orifices randomly distributed, and often can be observed in them, although with difficulty, that the fibers can be seen to be organized into several layers (
<figureCitation id="1361B186FFE3B308B58B1227247BF802" box="[201,271,1906,1932]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="8" pageNumber="23">Fig. 4</figureCitation>
C). These orifices are probably related into aeropyles, although a detailed study would be necessary to confirm this.
</paragraph>
<paragraph id="8BE5AD03FFE2B309B58415CD248EFE7C" blockId="9.[151,1436,152,538]" pageId="9" pageNumber="24">
In certain points of the fibrous layer, one or more bundles of filaments rises from the layer surface, turning spirally to produce fibrous projections, that could be classified as “fiber-coils”, of which two 
<typeStatus id="54E113A1FFE2B309B19515952061FF54" box="[1239,1301,192,218]" pageId="9" pageNumber="24">types</typeStatus>
can be distinguished, without a terminal fiber clusters and with a terminal knob. “Fiber-coils without terminal fiber clusters” correspond to a round structure of 6.5 µm diameter and sucker-shape, in which the filaments form a thick edge that delimits a depressed central area (
<figureCitation id="1361B186FFE2B309B7DB146D2795FEDC" box="[665,737,312,338]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="9" pageNumber="24">Fig. 4</figureCitation>
D). On the other hand, “fiber-coils with terminal knob” correspond to a pedunculate structure and mushroom-shape (
<figureCitation id="1361B186FFE2B309B601143526FEFEF4" box="[835,906,352,378]" captionStart="FIGURE 4" captionStartId="9.[151,255,1879,1903]" captionTargetBox="[152,1434,596,1854]" captionTargetId="figure@9.[152,1436,596,1855]" captionTargetPageId="9" captionText="FIGURE 4. Egg morphology of Oligoneuriella rhenana: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm)." httpUri="https://zenodo.org/record/176556/files/figure.png" pageId="9" pageNumber="24">Fig. 4</figureCitation>
E), approximately 7.3 µm height and 5.8 µm diameter in the umbrella-shaped part. This last structure has a more compact appearance than the previous one and its fibrous nature is only observable in the peduncle base, for which reason it has been classified as “fibrous attachment structure”.
</paragraph>
<paragraph id="8BE5AD03FFE2B309B58414AA21BDFD94" blockId="9.[151,1436,152,538]" box="[198,1225,511,538]" pageId="9" pageNumber="24">
<emphasis id="B92E7111FFE2B309B58414AA2421FD97" bold="true" box="[198,341,511,537]" italics="true" pageId="9" pageNumber="24">Micropyles:</emphasis>
These basic chorionic structures were not found in the eggs of this species.
</paragraph>
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<paragraph id="8BE5AD03FFE2B309B5D5120224B4F858" blockId="9.[151,1436,1879,2006]" pageId="9" pageNumber="24">
<emphasis id="B92E7111FFE2B309B5D51202246CF8E1" bold="true" box="[151,280,1879,1903]" pageId="9" pageNumber="24">FIGURE 4.</emphasis>
Egg morphology of 
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<emphasis id="B92E7111FFE2B309B4B61202279BF8E0" box="[500,751,1879,1902]" italics="true" pageId="9" pageNumber="24">Oligoneuriella rhenana</emphasis>
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: A, General shape of egg (scale bar = 50 μm). B, Detail of adhesive layer surface and lateral attachment structures (scale bar = 10 μm). Detailed of orifice in adhesive layer (C) (scale bar = 1 μm). D, Fiber coils without terminal fibers cluster in detail (scale bar = 1 μm). E, Fiber- coils with terminal knob in detail (scale bar = 5 μm).
</paragraph>
</caption>
<paragraph id="8BE5AD03FFE1B30AB5D515CD2411FF3C" blockId="10.[151,1437,152,2018]" box="[151,357,152,178]" pageId="10" pageNumber="25">Partial discussion</paragraph>
<paragraph id="8BE5AD03FFE1B30AB58415952168FDE4" blockId="10.[151,1437,152,2018]" pageId="10" pageNumber="25">
The main morphological aspects of 
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<emphasis id="B92E7111FFE1B30AB73915952677FF57" box="[635,771,192,217]" italics="true" pageId="10" pageNumber="25">
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. rhenana
</emphasis>
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egg were dealt with in depth by 
<bibRefCitation id="EFCBD0F2FFE1B30AB1DE15952013FF54" author="Degrange" box="[1180,1383,192,218]" pageId="10" pageNumber="27" refString="Degrange, C. (1960) Recherches sur la Reproduction des Ephemeropteres. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universite de Grenoble, 51, 71 - 93." type="journal article" year="1960">Degrange (1960)</bibRefCitation>
and later, briefly, by 
<bibRefCitation id="EFCBD0F2FFE1B30AB41E15BD276BFE8C" author="Haybach" box="[348,543,232,258]" pageId="10" pageNumber="27" refString="Haybach, A. (2003) Egg-structure of German mayflies (Insecta, Ephemeroptera). - an overview. Lauterbornia, 47, 41 - 58." type="journal article" year="2003">Haybach (2003)</bibRefCitation>
. Our SEM observations confirm the fibrous nature of the attachment structures (layer surrounding the chorion and lateral projection), that has already been described by 
<bibRefCitation id="EFCBD0F2FFE1B30AB06814452590FEDC" author="Degrange" pageId="10" pageNumber="27" refString="Degrange, C. (1960) Recherches sur la Reproduction des Ephemeropteres. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universite de Grenoble, 51, 71 - 93." type="journal article" year="1960">Degrange (1960)</bibRefCitation>
in this species, which is a very common feature in 
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eggs (
<bibRefCitation id="EFCBD0F2FFE1B30AB103146D202EFEDC" author="Koss" box="[1089,1370,312,338]" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss &amp; Edmunds 1974</bibRefCitation>
). The morphology and arrangement of the fibers of the fibrous layer of 
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. rhenana
</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C5AD680FFE1B30AB1DE1435244CFE2C" authority="Eaton, 1870" authorityName="Eaton" authorityYear="1870" class="Insecta" family="Siphlonuridae" genus="Siphlonurus" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="lacustris">
<emphasis id="B92E7111FFE1B30AB1DE143520E8FEF7" box="[1180,1436,352,377]" italics="true" pageId="10" pageNumber="25">Siphlonurus lacustris</emphasis>
(Eaton, 1870)
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eggs were very similar according to 
<bibRefCitation id="EFCBD0F2FFE1B30AB7AA14DD26C6FE2C" author="Degrange" box="[744,946,392,418]" pageId="10" pageNumber="27" refString="Degrange, C. (1960) Recherches sur la Reproduction des Ephemeropteres. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universite de Grenoble, 51, 71 - 93." type="journal article" year="1960">Degrange (1960)</bibRefCitation>
, but SEM investigations suggest quite the opposite. The fibers of 
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<emphasis id="B92E7111FFE1B30AB4ED14E52742FE47" box="[431,566,432,457]" italics="true" pageId="10" pageNumber="25">
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. rhenana
</emphasis>
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are short and compact bundles, formed by tiny inconspicuous filaments, which may be arranged in parallel or intercrossed, whereas the fibers of 
<taxonomicName id="4C5AD680FFE1B30AB6B6148D210CFE7F" box="[1012,1144,472,497]" class="Insecta" family="Siphlonuridae" genus="Siphlonurus" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="lacustris">
<emphasis id="B92E7111FFE1B30AB6B6148D210CFE7F" box="[1012,1144,472,497]" italics="true" pageId="10" pageNumber="25">S. lacustris</emphasis>
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are very long, relatively thick, and very conspicuous; they are loosely arranged in different directions (
<bibRefCitation id="EFCBD0F2FFE1B30AB10417552023FD94" author="Gaino" box="[1094,1367,512,538]" pageId="10" pageNumber="27" refString="Gaino, E. &amp; Rebora, M. (2001) Synthesis and function of the fibrous layers covering the eggs of Siphlonurus lacustris (Ephemeroptera, Siphlonuridae). Acta Zoologica Stockholm, 82, 41 - 48." type="journal article" year="2001">Gaino &amp; Rebora 2001</bibRefCitation>
). The ultrastructural organization between the fibrous attachment layer and the rest of the chorionic layers is still unknown in 
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. rhenana
</emphasis>
</taxonomicName>
, and a study of the eggshell using TEM is necessary.
</paragraph>
<paragraph id="8BE5AD03FFE1B30AB584172D21D3FB4C" blockId="10.[151,1437,152,2018]" pageId="10" pageNumber="25">
The lateral attachment structures in 
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. rhenana
</emphasis>
</taxonomicName>
eggs have been classified as fiber-coils with terminal fiber clusters, although two forms have been distinguished, one apparently evolved from the other (
<bibRefCitation id="EFCBD0F2FFE1B30AB19D17F520E3FD34" author="Degrange" box="[1247,1431,672,698]" pageId="10" pageNumber="27" refString="Degrange, C. (1960) Recherches sur la Reproduction des Ephemeropteres. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universite de Grenoble, 51, 71 - 93." type="journal article" year="1960">Degrange 1960</bibRefCitation>
; 
<bibRefCitation id="EFCBD0F2FFE1B30AB5D5179D24DEFD6C" author="Koss" box="[151,426,712,738]" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss &amp; Edmunds 1974</bibRefCitation>
). In our SEM study, two forms of fiber-coils were also differentiated; however, in relation with the forms already described, morphological differences are evident but we found no clear indication concerning which one evolves from the other. In fact, if we simply look at the organization of the distal end, we can differentiate two 
<typeStatus id="54E113A1FFE1B30AB48616152776FCD4" box="[452,514,832,858]" pageId="10" pageNumber="25">types</typeStatus>
of fiber-coils: those with a terminal knob and those without a terminal fiber cluster. Our fiber-coils with a terminal knob only have a lateral attachment structure which arises from the eggshell surface as a projection, and which is only a part of the attachment structures included in the 
<typeStatus id="54E113A1FFE1B30AB02B16C520E8FC24" box="[1385,1436,912,938]" pageId="10" pageNumber="25">type</typeStatus>
described by 
<bibRefCitation id="EFCBD0F2FFE1B30AB47616ED2704FC5C" author="Koss" box="[308,624,952,978]" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss and Edmunds (1974)</bibRefCitation>
. The fiber-coils without terminal fiber clusters involve the remaining lateral attachment structures, which appear to terminate within themselves, forming a sucker-shaped structure. A similar structure to this last one has already been described by 
<bibRefCitation id="EFCBD0F2FFE1B30AB6E1115D21ABFBAC" author="Koss" box="[931,1247,1032,1058]" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss and Edmunds (1974)</bibRefCitation>
for 
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eggs, but not in 
<taxonomicName id="4C5AD680FFE1B30AB4D911652757FBC7" box="[411,547,1072,1097]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriella" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="rhenana">
<emphasis id="B92E7111FFE1B30AB4D911652757FBC7" box="[411,547,1072,1097]" italics="true" pageId="10" pageNumber="25">
<collectingCountry id="F34DED93FFE1B30AB4D9116524C7FBC7" box="[411,435,1072,1097]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. rhenana
</emphasis>
</taxonomicName>
eggs. In addition, having obtained SEM evidences of the morphology of fiber-coils in eggs of 
<taxonomicName id="4C5AD680FFE1B30AB4D3110D2647FBFC" authority="Gaino &amp; Rebora 2001" authorityName="Gaino &amp; Rebora" authorityYear="2001" box="[401,819,1112,1138]" class="Insecta" family="Siphlonuridae" genus="Siphlonurus" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="lacustris">
<emphasis id="B92E7111FFE1B30AB4D3110D2760FBFF" box="[401,532,1112,1137]" italics="true" pageId="10" pageNumber="25">S. lacustris</emphasis>
(
<bibRefCitation id="EFCBD0F2FFE1B30AB761110D265EFBFC" author="Gaino" box="[547,810,1112,1138]" pageId="10" pageNumber="27" refString="Gaino, E. &amp; Rebora, M. (2001) Synthesis and function of the fibrous layers covering the eggs of Siphlonurus lacustris (Ephemeroptera, Siphlonuridae). Acta Zoologica Stockholm, 82, 41 - 48." type="journal article" year="2001">Gaino &amp; Rebora 2001</bibRefCitation>
)
</taxonomicName>
and in 
<taxonomicName id="4C5AD680FFE1B30AB6C8110D2164FBFF" box="[906,1040,1112,1137]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriella" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="rhenana">
<emphasis id="B92E7111FFE1B30AB6C8110D2164FBFF" box="[906,1040,1112,1137]" italics="true" pageId="10" pageNumber="25">
<collectingCountry id="F34DED93FFE1B30AB6C8110D26D6FBFF" box="[906,930,1112,1137]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. rhenana
</emphasis>
</taxonomicName>
, we were able to verify that these are clearly different, both in morphology and in arrangement unlike that indicated by 
<bibRefCitation id="EFCBD0F2FFE1B30AB1F011D5259CFB4C" author="Koss" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss and Edmunds (1974)</bibRefCitation>
, in which 
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<emphasis id="B92E7111FFE1B30AB42311FD2497FB4F" box="[353,483,1192,1217]" italics="true" pageId="10" pageNumber="25">S. lacustris</emphasis>
</taxonomicName>
fiber-coils are really of the 
<typeStatus id="54E113A1FFE1B30AB66B11FD2628FB4C" box="[809,860,1192,1218]" pageId="10" pageNumber="25">type</typeStatus>
with terminal fiber clusters.
</paragraph>
<paragraph id="8BE5AD03FFE1B30AB58411852758F9DC" blockId="10.[151,1437,152,2018]" pageId="10" pageNumber="25">
The extrachorion could not be determined in the eggs of 
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<emphasis id="B92E7111FFE1B30AB61C11852697FB67" box="[862,995,1232,1257]" italics="true" pageId="10" pageNumber="25">
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. rhenana
</emphasis>
</taxonomicName>
studied, although this cover is probably present and could be the thin layer that 
<bibRefCitation id="EFCBD0F2FFE1B30AB7DD11AD2613FA9C" author="Degrange" box="[671,871,1272,1298]" pageId="10" pageNumber="27" refString="Degrange, C. (1960) Recherches sur la Reproduction des Ephemeropteres. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universite de Grenoble, 51, 71 - 93." type="journal article" year="1960">Degrange (1960)</bibRefCitation>
described enveloping the attachment structures externally which coagulated in eggs preserved with ethanol. We cannot explain why we did not observe this layer in 
<emphasis id="B92E7111FFE1B30AB5BE101D24FEFAEF" box="[252,394,1352,1377]" italics="true" pageId="10" pageNumber="25">
<taxonomicName id="4C5AD680FFE1B30AB5BE101D24F2FAEF" box="[252,390,1352,1377]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriella" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="rhenana">
<collectingCountry id="F34DED93FFE1B30AB5BE101D2460FAEF" box="[252,276,1352,1377]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. rhenana
</taxonomicName>
,
</emphasis>
having studied eggs preserved in ethanol and obtained from nymphs, even though the extrachorion was observed in the eggs of other species preserved in ethanol (
<bibRefCitation id="EFCBD0F2FFE1B30AB115102520FAFA04" author="Ubero-Pascal" box="[1111,1422,1392,1418]" pageId="10" pageNumber="29" refString="Ubero-Pascal, N., Fortuno, J. M. &amp; Puig, M. A. (2005) New application of air-drying techniques for studying Ephemeroptera and Plecoptera eggs by Scanning Electron Microscopy. Microscopy Research and Technique, 68, 264 - 271." type="journal article" year="2005">
Ubero-Pascal 
<emphasis id="B92E7111FFE1B30AB04110252048FA07" box="[1283,1340,1392,1417]" italics="true" pageId="10" pageNumber="25">et al</emphasis>
. 2005
</bibRefCitation>
); according to 
<bibRefCitation id="EFCBD0F2FFE1B30AB47710CD270CFA3C" author="Koss" box="[309,632,1432,1458]" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss and Edmunds (1974)</bibRefCitation>
, eggs from nymphs display a thicker extrachorion. Neither did we observe a micropyle in eggs of 
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<emphasis id="B92E7111FFE1B30AB748109527E4FA57" box="[522,656,1472,1497]" italics="true" pageId="10" pageNumber="25">
<collectingCountry id="F34DED93FFE1B30AB74810952756FA57" box="[522,546,1472,1497]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. rhenana
</emphasis>
</taxonomicName>
, although this chorionic structure is “tagenoform-type” and one or two units appear near one of the polar areas (
<bibRefCitation id="EFCBD0F2FFE1B30AB7EB10BD2617F98C" author="Degrange" box="[681,867,1512,1538]" pageId="10" pageNumber="27" refString="Degrange, C. (1960) Recherches sur la Reproduction des Ephemeropteres. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universite de Grenoble, 51, 71 - 93." type="journal article" year="1960">Degrange 1960</bibRefCitation>
). Probably, as already described in other species displaying a fibrous attachment layer (
<bibRefCitation id="EFCBD0F2FFE1B30AB72313452604F9A4" author="Gaino" box="[609,880,1552,1578]" pageId="10" pageNumber="27" refString="Gaino, E. &amp; Rebora, M. (2001) Synthesis and function of the fibrous layers covering the eggs of Siphlonurus lacustris (Ephemeroptera, Siphlonuridae). Acta Zoologica Stockholm, 82, 41 - 48." type="journal article" year="2001">Gaino &amp; Rebora 2001</bibRefCitation>
), the micropyle can only be observed by SEM when this layer has been removed.
</paragraph>
<paragraph id="8BE5AD03FFE1B30BB58413352661FF54" blockId="10.[151,1437,152,2018]" lastBlockId="11.[151,1436,150,218]" lastPageId="11" lastPageNumber="26" pageId="10" pageNumber="25">
The fibrous adhesive layer and “fiber-coils” as lateral attachment structures seem to be common characteristics of the morphological patterns of 
<taxonomicName id="4C5AD680FFE1B30AB72213DD2664F92C" box="[608,784,1672,1698]" class="Insecta" family="Oligoneuriidae" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="family">Oligoneuriidae</taxonomicName>
eggs. From a taxonomic point of view, the density and distribution of the lateral attachment structures around the egg could help differentiate species, although their use in genera taxonomy is not clear from light microscopy (
<bibRefCitation id="EFCBD0F2FFE1B30AB638138D21E9F97C" author="Koss" box="[890,1181,1752,1778]" pageId="10" pageNumber="28" refString="Koss, R. W. &amp; Edmunds, G. F. Jr. (1974) Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society, 55, 267 - 349." type="journal article" year="1974">Koss &amp; Edmunds 1974</bibRefCitation>
). Descriptions of the chorion morphology in 
<taxonomicName id="4C5AD680FFE1B30AB4EF12552729F894" box="[429,605,1792,1818]" class="Insecta" family="Oligoneuriidae" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="family">Oligoneuriidae</taxonomicName>
using SEM are scarce, very concise or merely represented by a photograph (
<bibRefCitation id="EFCBD0F2FFE1B30AB5AB127D2745F8CC" author="Pescador" box="[233,561,1832,1858]" pageId="10" pageNumber="28" refString="Pescador, M. L. &amp; Edmunds, G. F. Jr. (1994) New genus of Oligoneuriidae (Ephemeroptera) from South America. Annals of the Entomological Society of America, 87, 263 - 269." type="journal article" year="1994">Pescador &amp; Edmunds 1994</bibRefCitation>
; 
<bibRefCitation id="EFCBD0F2FFE1B30AB77C127D262FF8CC" author="Soldan" box="[574,859,1832,1858]" pageId="10" pageNumber="28" refString="Soldan, T. &amp; Thomas, A. G. B. (1983) New and little-known species of mayflies (Ephemeroptera) from Algeria. Acta entomologica bohemoslovaca, 80, 356 - 376." type="journal article" year="1983">Soldan &amp; Thomas 1983</bibRefCitation>
), while our description of 
<taxonomicName id="4C5AD680FFE1B30AB1D6127D206EF8CF" box="[1172,1306,1832,1857]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriella" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="rhenana">
<emphasis id="B92E7111FFE1B30AB1D6127D206EF8CF" box="[1172,1306,1832,1857]" italics="true" pageId="10" pageNumber="25">
<collectingCountry id="F34DED93FFE1B30AB1D6127D21D8F8CF" box="[1172,1196,1832,1857]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. rhenana
</emphasis>
</taxonomicName>
eggs demonstrates that SEM can show new details of the chorionic structure which could be taxonomically useful, for example for differentiating from 
<taxonomicName id="4C5AD680FFE1B30AB768122D21D5F81C" authority="Dakki &amp; Giudicelli, 1980" authorityName="Dakki &amp; Giudicelli" authorityYear="1980" box="[554,1185,1912,1938]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriopsis" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="skounate">
<emphasis id="B92E7111FFE1B30AB768122D262EF81F" box="[554,858,1912,1937]" italics="true" pageId="10" pageNumber="25">Oligoneuriopsis skounate</emphasis>
Dakki &amp; Giudicelli, 1980
</taxonomicName>
. This species occurs with 
<taxonomicName id="4C5AD680FFE1B30AB59012F52423F837" box="[210,343,1952,1977]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriella" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="rhenana">
<emphasis id="B92E7111FFE1B30AB59012F52423F837" box="[210,343,1952,1977]" italics="true" pageId="10" pageNumber="25">
<collectingCountry id="F34DED93FFE1B30AB59012F5259EF837" box="[210,234,1952,1977]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. rhenana
</emphasis>
</taxonomicName>
in the Segura river basin and, although we only have a general photograph of its egg (
<bibRefCitation id="EFCBD0F2FFE1B30AB00A12F5242EF86C" author="Soldan" pageId="10" pageNumber="28" refString="Soldan, T. &amp; Thomas, A. G. B. (1983) New and little-known species of mayflies (Ephemeroptera) from Algeria. Acta entomologica bohemoslovaca, 80, 356 - 376." type="journal article" year="1983">Soldan &amp; Thomas 1983</bibRefCitation>
), both species could be differentiated by the “fiber-coils”. 
<taxonomicName id="4C5AD680FFE1B30AB144129D21D5F86F" box="[1030,1185,1992,2017]" class="Insecta" family="Oligoneuriidae" genus="Oligoneuriopsis" kingdom="Animalia" order="Ephemeroptera" pageId="10" pageNumber="25" phylum="Arthropoda" rank="species" species="skhounate">
<emphasis id="B92E7111FFE1B30AB144129D21D5F86F" box="[1030,1185,1992,2017]" italics="true" pageId="10" pageNumber="25">
<collectingCountry id="F34DED93FFE1B30AB144129D216AF86F" box="[1030,1054,1992,2017]" name="Iceland" pageId="10" pageNumber="25">O</collectingCountry>
. skhounate
</emphasis>
</taxonomicName>
has eggs with “fibercoils a with terminal fiber clusters”, very similar in morphology to the egg of 
<taxonomicName id="4C5AD680FFE0B30BB16715CD21DCFF3F" box="[1061,1192,152,177]" class="Insecta" family="Siphlonuridae" genus="Siphlonurus" kingdom="Animalia" order="Ephemeroptera" pageId="11" pageNumber="26" phylum="Arthropoda" rank="species" species="lacustris">
<emphasis id="B92E7111FFE0B30BB16715CD21DCFF3F" box="[1061,1192,152,177]" italics="true" pageId="11" pageNumber="26">S. lacustris</emphasis>
</taxonomicName>
, while the two 
<typeStatus id="54E113A1FFE0B30BB01C15CD20E8FF3C" box="[1374,1436,152,178]" pageId="11" pageNumber="26">types</typeStatus>
of “fiber-coils” of 
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<emphasis id="B92E7111FFE0B30BB42B1595249AFF57" box="[361,494,192,217]" italics="true" pageId="11" pageNumber="26">
<collectingCountry id="F34DED93FFE0B30BB42B159524F5FF57" box="[361,385,192,217]" name="Iceland" pageId="11" pageNumber="26">O</collectingCountry>
. rhenana
</emphasis>
</taxonomicName>
egg are clearly different.
</paragraph>
</subSubSection>
</treatment>
</document>