treatments-xml/data/C0/63/78/C063786AFFA2FFF0FF0D14EE5FFB97AF.xml
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<mods:title id="2EB54BA597E11F4AD6D65EE7EEC5E2A9">New records and description of fifty-four new species of aquatic beetles in the genus Hydraena Kugelann from South America (Coleoptera: Hydraenidae)</mods:title>
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<paragraph id="4875C97CFFA2FFF1FF0D14EE5EFB90B7" blockId="12.[151,474,1501,1527]" box="[151,474,1501,1527]" pageId="12" pageNumber="13">
<heading id="133D7E10FFA2FFF1FF0D14EE5EFB90B7" bold="true" box="[151,474,1501,1527]" fontSize="11" level="1" pageId="12" pageNumber="13" reason="1">
<taxonomicName id="8FCAB2FFFFA2FFF1FF0D14EE5EFB90B7" ID-CoL="8VRR8" authority="Kugelann, 1794" authorityName="Kugelann" authorityYear="1794" box="[151,474,1501,1527]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FF0D14EE5EFB90B7" bold="true" box="[151,474,1501,1527]" pageId="12" pageNumber="13">
<emphasis id="7ABE156EFFA2FFF1FF0D14EE5E3190B7" bold="true" box="[151,272,1501,1527]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
Kugelann, 1794
</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<paragraph id="4875C97CFFA2FFF1FF0D17175B2D9379" blockId="12.[151,1036,1571,1594]" box="[151,1036,1571,1594]" pageId="12" pageNumber="13">
<taxonomicName id="8FCAB2FFFFA2FFF1FF0D17175EFC9379" ID-CoL="8VRR8" authority="Kugelann, 1794: 578" authorityName="Kugelann" authorityPageNumber="578" authorityYear="1794" box="[151,477,1571,1594]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FF0D17175FDD937A" box="[151,252,1572,1594]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
Kugelann, 1794: 578
</taxonomicName>
(
<typeStatus id="977177DEFFA2FFF1FE7017175D369379" box="[490,535,1572,1593]" pageId="12" pageNumber="13">type</typeStatus>
species
<taxonomicName id="8FCAB2FFFFA2FFF1FDF417175B219379" ID-CoL="3N2ZN" authority="Kugelann 1794: 579" authorityName="Kugelann" authorityPageNumber="579" authorityYear="1794" box="[622,1024,1571,1594]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="species" species="riparia">
<emphasis id="7ABE156EFFA2FFF1FDF417175C00937A" box="[622,801,1572,1594]" italics="true" pageId="12" pageNumber="13">Hydraena riparia</emphasis>
Kugelann 1794: 579
</taxonomicName>
).
</paragraph>
<paragraph id="4875C97CFFA2FFF1FF0D17555E669272" blockId="12.[151,1437,1638,2023]" pageId="12" pageNumber="13">
The speciose, cosmopolitan genus
<taxonomicName id="8FCAB2FFFFA2FFF1FDBF17555DB4933E" ID-CoL="8VRR8" box="[549,661,1638,1662]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FDBF17555DB4933E" box="[549,661,1638,1662]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
</taxonomicName>
is well defined by the following autapomorphic characteristics: 1) presence of a labral-mandibular interlocking device (
<bibRefCitation id="2C5BB48DFFA2FFF1FD6F17B95CB193E2" author="Perkins" box="[757,912,1674,1698]" pageId="12" pageNumber="13" refString="Perkins, P. D. (1989). Adelphydraena, new genus, and two new species from Venezuela, and remarks on phylogenetic relationships within the subtribe Hydraenina (Coleoptera: Hydraenidae). Proceedings of the Biological Society of Washington, 102, 447 - 457." type="journal article" year="1989">Perkins 1989</bibRefCitation>
), 2) mentum with an acute median projection anteriorly (
<bibRefCitation id="2C5BB48DFFA2FFF1FE89179D5E8F9386" author="Perkins" box="[275,430,1710,1734]" pageId="12" pageNumber="13" refString="Perkins, P. D. (1989). Adelphydraena, new genus, and two new species from Venezuela, and remarks on phylogenetic relationships within the subtribe Hydraenina (Coleoptera: Hydraenidae). Proceedings of the Biological Society of Washington, 102, 447 - 457." type="journal article" year="1989">Perkins 1989</bibRefCitation>
), 3) various complex sensilla (
<bibRefCitation id="2C5BB48DFFA2FFF1FC91179D5C859386" author="Perkins" box="[779,932,1710,1734]" pageId="12" pageNumber="13" refString="Perkins, P. D. (1997). Life on the effective bubble: exocrine secretion delivery systems (ESDS) and the evolution and classification of beetles in the family Hydraenidae (Insecta: Coleoptera). Annals of Carnegie Museum, 66 (2), 89 - 207." type="journal article" year="1997">Perkins 1997</bibRefCitation>
), and 4) specialized features of the exocrine secretion delivery system (
<bibRefCitation id="2C5BB48DFFA2FFF1FE5717E15D4893AA" author="Perkins" box="[461,617,1746,1770]" pageId="12" pageNumber="13" refString="Perkins, P. D. (1997). Life on the effective bubble: exocrine secretion delivery systems (ESDS) and the evolution and classification of beetles in the family Hydraenidae (Insecta: Coleoptera). Annals of Carnegie Museum, 66 (2), 89 - 207." type="journal article" year="1997">Perkins 1997</bibRefCitation>
). There is general agreement among current workers that
<taxonomicName id="8FCAB2FFFFA2FFF1FA9517E05A5E93AB" ID-CoL="8VRR8" box="[1295,1407,1747,1771]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FA9517E05A5E93AB" box="[1295,1407,1747,1771]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
</taxonomicName>
is monophyletic, based on these characters (for example,
<bibRefCitation id="2C5BB48DFFA2FFF1FC9017C55CE3924E" author="Jach" box="[778,962,1782,1806]" pageId="12" pageNumber="13" refString="Jach, M. A., Beutel, R., Diaz, J. A. &amp; Kodada, J. (2000). Subgeneric classification, descriptions of head structures, and world check list of Hydraena Kugelann (Insecta: Coleoptera: Hydraenidae). Annalen des Naturhistorischen Museums in Wien, 102 B, 177 - 258." type="journal article" year="2000">Jäch et al. 2000</bibRefCitation>
). The South American species meet these generic criteria.
</paragraph>
<paragraph id="4875C97CFFA2FFF0FF5D160D5AA69424" blockId="12.[151,1437,1638,2023]" lastBlockId="13.[151,1436,151,751]" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13">
The division of
<taxonomicName id="8FCAB2FFFFA2FFF1FEE6160D5ECD9216" ID-CoL="8VRR8" box="[380,492,1854,1878]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FEE6160D5ECD9216" box="[380,492,1854,1878]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
</taxonomicName>
into subgenera is currently a matter of uncertainty. Some groupings of species into large putative monophyletic groups is more certain, but the basal relationships among the major clades, based on recently published DNA analysis, are not well supported (
<bibRefCitation id="2C5BB48DFFA2FFF1FCAA16B55B2592DE" author="Ribera" box="[816,1028,1926,1951]" pageId="12" pageNumber="13" refString="Ribera, I, Castro, A., Diaz, J. A., Garrido, J., Izquierdo, A., Jach, M. A., Valladares, L. F. (2011). The geography of speciation in narrow-range endemics of the ' Haenydra' lineage (Coleoptera, Hydraenidae, Hydraena). Journal of Biogeography 38, 502 - 516." type="journal article" year="2011">
Ribera
<emphasis id="7ABE156EFFA2FFF1FC1C16BB5CE092DE" box="[902,961,1926,1951]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
2011
</bibRefCitation>
: 508). The morphological cladistic analysis of
<bibRefCitation id="2C5BB48DFFA2FFF1FE8716995ECB9282" box="[285,490,1962,1987]" pageId="12" pageNumber="13" refString="Jach, M. A., Beutel, R., Diaz, J. A. &amp; Kodada, J. (2000). Subgeneric classification, descriptions of head structures, and world check list of Hydraena Kugelann (Insecta: Coleoptera: Hydraenidae). Annalen des Naturhistorischen Museums in Wien, 102 B, 177 - 258." type="journal article">
Jäch
<emphasis id="7ABE156EFFA2FFF1FEC3169F5EB49283" box="[345,405,1963,1987]" italics="true" pageId="12" pageNumber="13">et al.</emphasis>
(2000)
</bibRefCitation>
concluded that &quot;the great phylogenetic alternative&quot; to
<taxonomicName id="8FCAB2FFFFA2FFF1FBF216985BF99283" ID-CoL="8VRR8" box="[1128,1240,1963,1987]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FBF216985BF99283" box="[1128,1240,1963,1987]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
</taxonomicName>
(
<emphasis id="7ABE156EFFA2FFF1FB76169F5BD69283" box="[1260,1271,1964,1987]" italics="true" pageId="12" pageNumber="13">s</emphasis>
.
<emphasis id="7ABE156EFFA2FFF1FB67169F5A389283" box="[1277,1305,1964,1987]" italics="true" pageId="12" pageNumber="13">str</emphasis>
.) had been found in
<taxonomicName id="8FCAB2FFFFA2FFF1FF6416FD5D0792A6" ID-CoL="8VRR8" authority="(Hydraenopsis)" baseAuthorityName="Hydraenopsis" box="[254,550,1998,2022]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FF6416FD5E4F92A6" box="[254,366,1998,2022]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
(
<emphasis id="7ABE156EFFA2FFF1FEE516FD5D3A92A6" box="[383,539,1998,2022]" italics="true" pageId="12" pageNumber="13">Hydraenopsis</emphasis>
)
</taxonomicName>
. At least some of the South American species of
<taxonomicName id="8FCAB2FFFFA2FFF1FBC216FD5BE992A6" ID-CoL="8VRR8" box="[1112,1224,1998,2022]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA2FFF1FBC216FD5BE992A6" box="[1112,1224,1998,2022]" italics="true" pageId="12" pageNumber="13">Hydraena</emphasis>
</taxonomicName>
are most probably related to species placed by Jäch
<emphasis id="7ABE156EFFA3FFF0FD8111AA5D7495F0" box="[539,597,152,176]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
in
<taxonomicName id="8FCAB2FFFFA3FFF0FDE511AB5C8895EF" ID-CoL="8VRR8" authority="(Hydraenopsis)" baseAuthorityName="Hydraenopsis" box="[639,937,151,176]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FDE511AB5DCE95F0" box="[639,751,152,176]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
(
<emphasis id="7ABE156EFFA3FFF0FC9811AB5CBF95F0" box="[770,926,152,176]" italics="true" pageId="13" pageNumber="14">Hydraenopsis</emphasis>
)
</taxonomicName>
. However, the DNA study by Ribera
<emphasis id="7ABE156EFFA3FFF0FAFB11AA5ABD95F0" box="[1377,1436,152,176]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
included only one unidentified South American species (from the famous
<taxonomicName id="8FCAB2FFFFA3FFF0FC40118F5B1D9594" ID-CoL="3N3JS" box="[986,1084,188,212]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="107" phylum="Arthropoda" rank="species" species="tobogan">Tobogan</taxonomicName>
locality), probably of the
<emphasis id="7ABE156EFFA3FFF0FAF9118E5FDE95B8" italics="true" pageId="13" pageNumber="14">Marginicollis</emphasis>
Group. More DNA studies are needed, especially of the unusual
<taxonomicName id="8FCAB2FFFFA3FFF0FC6C11D25B5395B8" ID-CoL="3N2JH" box="[1014,1138,225,248]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="73" phylum="Arthropoda" rank="species" species="paeminosa">
<emphasis id="7ABE156EFFA3FFF0FC6C11D25B5395B8" box="[1014,1138,225,248]" italics="true" pageId="13" pageNumber="14">Paeminosa</emphasis>
</taxonomicName>
,
<taxonomicName id="8FCAB2FFFFA3FFF0FB1911D35BC395B7" ID-CoL="3N38K" box="[1155,1250,224,247]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="82" phylum="Arthropoda" rank="species" species="curvosa">
<emphasis id="7ABE156EFFA3FFF0FB1911D35BC395B7" box="[1155,1250,224,247]" italics="true" pageId="13" pageNumber="14">Curvosa</emphasis>
</taxonomicName>
, and
<taxonomicName id="8FCAB2FFFFA3FFF0FAB911D35ABD95B8" ID-CoL="3N3YY" box="[1315,1436,224,248]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="86" phylum="Arthropoda" rank="species" species="multispina">
<emphasis id="7ABE156EFFA3FFF0FAB911D35ABD95B8" box="[1315,1436,224,248]" italics="true" pageId="13" pageNumber="14">Multispina</emphasis>
</taxonomicName>
Groups, to better understand the relationships of South American
<taxonomicName id="8FCAB2FFFFA3FFF0FC0910375B22945C" ID-CoL="8VRR8" box="[915,1027,260,284]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FC0910375B22945C" box="[915,1027,260,284]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
</taxonomicName>
. For now, I prefer not to formally assign species to subgenera until more results of DNA analysis are published, and it is shown that all species are monophyletic, and that &quot;
<taxonomicName id="8FCAB2FFFFA3FFF0FE31107F5D669424" box="[427,583,332,356]" class="Insecta" family="Hydraenidae" genus="Hydraenopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FE31107F5D669424" box="[427,583,332,356]" italics="true" pageId="13" pageNumber="14">Hydraenopsis</emphasis>
</taxonomicName>
&quot; is definitely not nested within what is now considered
<taxonomicName id="8FCAB2FFFFA3FFF0FB21107F5A0A9424" ID-CoL="8VRR8" box="[1211,1323,332,356]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FB21107F5A0A9424" box="[1211,1323,332,356]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
</taxonomicName>
(
<emphasis id="7ABE156EFFA3FFF0FAA1107E5A679424" box="[1339,1350,333,356]" italics="true" pageId="13" pageNumber="14">s</emphasis>
.
<emphasis id="7ABE156EFFA3FFF0FACE107E5A519424" box="[1364,1392,333,356]" italics="true" pageId="13" pageNumber="14">str</emphasis>
.).
</paragraph>
<paragraph id="4875C97CFFA3FFF0FF5D10435FFB97AF" blockId="13.[151,1436,151,751]" pageId="13" pageNumber="14">
<taxonomicName id="8FCAB2FFFFA3FFF0FF5D10435E1694C8" ID-CoL="8VRR8" box="[199,311,368,392]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FF5D10435E1694C8" box="[199,311,368,392]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
</taxonomicName>
is perhaps the most speciose water beetle genus (
<bibRefCitation id="2C5BB48DFFA3FFF0FCF1105C5B2594C7" author="Perkins" box="[875,1028,367,391]" pageId="13" pageNumber="14" refString="Perkins, P. D. (1997). Life on the effective bubble: exocrine secretion delivery systems (ESDS) and the evolution and classification of beetles in the family Hydraenidae (Insecta: Coleoptera). Annals of Carnegie Museum, 66 (2), 89 - 207." type="journal article" year="1997">Perkins 1997</bibRefCitation>
,
<bibRefCitation id="2C5BB48DFFA3FFF0FB8A105C5BCD94C7" author="Jach" box="[1040,1260,367,391]" pageId="13" pageNumber="14" refString="Jach, M. A. &amp; Balke, M. (2008). Global diversity of water beetles (Coleoptera) in freshwater. Hydrobiologia 595, 419 - 442." type="journal article" year="2008">Jäch &amp; Balke 2008</bibRefCitation>
,
<bibRefCitation id="2C5BB48DFFA3FFF0FB62105C5AB194C7" author="Perkins" box="[1272,1424,367,391]" pageId="13" pageNumber="14" refString="Perkins, P. D. (2011). New species (130) of the hyperdiverse aquatic beetle genus Hydraena Kugelann from Papua New Guinea, and a preliminary analysis of areas of endemism (Coleoptera: Hydraenidae). Zootaxa 2944, 1 - 417." type="journal article" year="2011">Perkins 2011</bibRefCitation>
). Including the current contribution, the number of described species becomes about 900, but many, many more undescribed species reside in museums (
<bibRefCitation id="2C5BB48DFFA3FFF0FDFF10845C66948F" author="Jach" box="[613,839,439,463]" pageId="13" pageNumber="14" refString="Jach, M. A. &amp; Balke, M. (2008). Global diversity of water beetles (Coleoptera) in freshwater. Hydrobiologia 595, 419 - 442." type="journal article" year="2008">Jäch &amp; Balke 2008</bibRefCitation>
). The tremendous &quot;success&quot; of
<taxonomicName id="8FCAB2FFFFA3FFF0FB35108B5A3E9490" ID-CoL="8VRR8" box="[1199,1311,440,464]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FB35108B5A3E9490" box="[1199,1311,440,464]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
</taxonomicName>
appears to be, in large part, a result of the evolution of an exocrine secretion delivery system (ESDS), which is a combination of specialized cuticular structures, exocrine glands, and behavior that together function as a microbial defense system (
<bibRefCitation id="2C5BB48DFFA3FFF0FF4A13175E48977C" author="Perkins" box="[208,361,548,572]" pageId="13" pageNumber="14" refString="Perkins, P. D. (1997). Life on the effective bubble: exocrine secretion delivery systems (ESDS) and the evolution and classification of beetles in the family Hydraenidae (Insecta: Coleoptera). Annals of Carnegie Museum, 66 (2), 89 - 207." type="journal article" year="1997">Perkins 1997</bibRefCitation>
). About 50 genera are currently recognized for the family. The number of species is quite modest in all but three of these genera, and only these three have an ESDS:
<taxonomicName id="8FCAB2FFFFA3FFF0FC45137B5B6E9720" ID-CoL="8VRR8" box="[991,1103,584,608]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FC45137B5B6E9720" box="[991,1103,584,608]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
</taxonomicName>
,
<taxonomicName id="8FCAB2FFFFA3FFF0FBC5137B5BFF9720" ID-CoL="8VRWR" box="[1119,1246,584,608]" class="Insecta" family="Hydraenidae" genus="Ochthebius" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FBC5137B5BFF9720" box="[1119,1246,584,608]" italics="true" pageId="13" pageNumber="14">Ochthebius</emphasis>
</taxonomicName>
, and
<taxonomicName id="8FCAB2FFFFA3FFF0FABA137B5AB59720" ID-CoL="8VRSL" box="[1312,1428,584,608]" class="Insecta" family="Hydraenidae" genus="Limnebius" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FABA137B5AB59720" box="[1312,1428,584,608]" italics="true" pageId="13" pageNumber="14">Limnebius</emphasis>
</taxonomicName>
.
<taxonomicName id="8FCAB2FFFFA3FFF0FF0D135F5E2697C4" ID-CoL="8VRR8" box="[151,263,620,644]" class="Insecta" family="Hydraenidae" genus="Hydraena" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="7ABE156EFFA3FFF0FF0D135F5E2697C4" box="[151,263,620,644]" italics="true" pageId="13" pageNumber="14">Hydraena</emphasis>
</taxonomicName>
, with perhaps 1000+ species, is the largest and qualifies to be regarded as hyperdiverse (sensu
<bibRefCitation id="2C5BB48DFFA3FFF0FAD6135F5FF597E7" author="Wilson" pageId="13" pageNumber="14" refString="Wilson, E. O. (2003). Pheidole in the new world: a dominant, hyperdiverse ant genus. Harvard University Press, Cambridge, Massachusetts. iii + 1 - 794 pp." type="book chapter" year="2003">Wilson 2003</bibRefCitation>
). The small size of these water beetles certainly contributes to the ESDS diversity equation, through effectively limited dispersal capabilities. Many other groups of insects are tiny, and disperse poorly, but are not hyperdiverse.
</paragraph>
</subSubSection>
</treatment>
</document>