treatments-xml/data/03/A9/87/03A987CF0553A906FF452841FC4B276B.xml

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<mods:title id="568D597B55B363FA32C6B44932E52D62">Little walking leaves from southeast Ecuador: biology and taxonomy of Typophyllum species (Orthoptera, Tettigoniidae, Pterochrozinae)</mods:title>
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<subSubSection id="C31A65520553A90CFF452841FDDD224E" box="[151,633,1055,1080]" pageId="23" pageNumber="24" type="nomenclature">
<paragraph id="8BBF36D90553A90CFF452841FDDD224E" blockId="23.[151,633,1055,1080]" box="[151,633,1055,1080]" pageId="23" pageNumber="24">
<heading id="D0F781B50553A90CFF452841FDDD224E" bold="true" box="[151,633,1055,1080]" fontSize="10" level="2" pageId="23" pageNumber="24" reason="2">
<taxonomicName id="4C004D5A0553A90CFF452841FDDD224E" authority="Enderlein 1917" authorityName="Enderlein" authorityYear="1917" box="[151,633,1055,1080]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="lacinipenne">
<emphasis id="B974EACB0553A90CFF452841FDDD224E" bold="true" box="[151,633,1055,1080]" pageId="23" pageNumber="24">
<emphasis id="B974EACB0553A90CFF452841FE18224E" bold="true" box="[151,444,1055,1080]" italics="true" pageId="23" pageNumber="24">Typophyllum lacinipenne</emphasis>
<bibRefCitation id="EF914B280553A90CFE162841FDDD224E" author="Enderlein" box="[452,633,1055,1080]" pageId="23" pageNumber="24" refString="Enderlein, G. (1917) Neue neotropische Pseudophyllinen. Zoologischer Anzeiger, 49, 17 - 20." type="journal article" year="1917">Enderlein 1917</bibRefCitation>
</emphasis>
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<taxonomicName id="4C004D5A0553A90CFF452837FE0E2208" authority="Caudell 1918" authorityName="Caudell" authorityYear="1918" box="[151,426,1128,1150]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="undulatum">
<emphasis id="B974EACB0553A90CFF452837FEB82208" box="[151,284,1128,1150]" italics="true" pageId="23" pageNumber="24">T. undulatum</emphasis>
<bibRefCitation id="EF914B280553A90CFEF02836FE0E2208" author="Caudell" box="[290,426,1128,1150]" pageId="23" pageNumber="24" refString="Caudell, A. N. (1918) On a collection of Orthoptera (exclusive of the Locustidae) made in central Peru by N. Iconnicoff and C. Schunke. Insecutor Insciiae Menstruus, 6, 1 - 70." type="journal article" year="1918">Caudell 1918</bibRefCitation>
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<taxonomicNameLabel id="A24757B00553A90CFE622836FDA9220B" box="[432,525,1128,1149]" pageId="23" pageNumber="24">
<emphasis id="B974EACB0553A90CFE622836FDA9220B" bold="true" box="[432,525,1128,1149]" pageId="23" pageNumber="24">syn. nov.</emphasis>
</taxonomicNameLabel>
</paragraph>
<paragraph id="8BBF36D90553A90CFF4528D6FE4322EA" blockId="23.[151,1437,1128,1303]" box="[151,487,1158,1181]" pageId="23" pageNumber="24">
<taxonomicName id="4C004D5A0553A90CFF4528D6FE2022EA" authority="Vignon 1925" authorityName="Vignon" authorityYear="1925" box="[151,388,1158,1181]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="acutum">
<emphasis id="B974EACB0553A90CFF4528D6FF5E22EB" box="[151,250,1160,1181]" italics="true" pageId="23" pageNumber="24">T. acutum</emphasis>
Vignon 1925
</taxonomicName>
<emphasis id="B974EACB0553A90CFE5828D9FE4322EA" bold="true" box="[394,487,1159,1180]" pageId="23" pageNumber="24">
<taxonomicNameLabel id="A24757B00553A90CFE5828D9FE4322EA" box="[394,487,1159,1180]" pageId="23" pageNumber="24">syn. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
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http://lsid.speciesfile.org/urn:lsid: 
<taxonomicName id="4C004D5A0553A90CFE3428FBFDFE22CD" ID-CoL="8NKFC" box="[486,602,1189,1211]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="order">Orthoptera</taxonomicName>
.speciesfile.org:TaxonName:5490
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<taxonomicName id="4C004D5A0553A90CFF45289BFE7722AC" authority="Enderlein 1917" authorityName="Enderlein" authorityYear="1917" box="[151,467,1220,1242]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="lacinipenne">
<emphasis id="B974EACB0553A90CFF45289BFE8322AC" box="[151,295,1220,1242]" italics="true" pageId="23" pageNumber="24">T. lacinipenne</emphasis>
: 
<bibRefCitation id="EF914B280553A90CFEE4289AFE7722AC" author="Enderlein" box="[310,467,1220,1242]" pageId="23" pageNumber="24" refString="Enderlein, G. (1917) Neue neotropische Pseudophyllinen. Zoologischer Anzeiger, 49, 17 - 20." type="journal article" year="1917">Enderlein 1917</bibRefCitation>
</taxonomicName>
(
<taxonomicName id="4C004D5A0553A90CFE31289BFDDD22AC" box="[483,633,1220,1242]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="lacinipennis">
<emphasis id="B974EACB0553A90CFE31289BFDDD22AC" box="[483,633,1220,1242]" italics="true" pageId="23" pageNumber="24">T. lacinipennis</emphasis>
</taxonomicName>
): 19, Vignon 125a: 270 (
<taxonomicName id="4C004D5A0553A90CFC52289BFBB322AC" box="[896,1047,1220,1242]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="lacinipennis">
<emphasis id="B974EACB0553A90CFC52289BFBB322AC" box="[896,1047,1220,1242]" italics="true" pageId="23" pageNumber="24">T. lacinipennis</emphasis>
</taxonomicName>
), 
<bibRefCitation id="EF914B280553A90CFBFF289AFB1622AC" author="Vignon" box="[1069,1202,1220,1242]" pageId="23" pageNumber="24" refString="Vignon, P. (1931) Recherches sur les Sauterelles-Feuilles de l'Amerique tropicale. Archives du Museum d'Histoire Naturelle, Paris, 5, 57 - 212." type="journal article" year="1931">Vignon 1931</bibRefCitation>
(
<taxonomicName id="4C004D5A0553A90CFB10289BFAFC22AC" box="[1218,1368,1220,1242]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="lacinipennis">
<emphasis id="B974EACB0553A90CFB10289BFAFC22AC" box="[1218,1368,1220,1242]" italics="true" pageId="23" pageNumber="24">T. lacinipennis</emphasis>
</taxonomicName>
): 135, 
<treatmentCitation id="0AA110C80553A90CFF1528BCFECA228E" author="Beier" box="[199,366,1250,1272]" page="365" pageId="23" pageNumber="24" year="1960">
<bibRefCitation id="EF914B280553A90CFF1528BCFE9E228E" author="Beier" box="[199,314,1250,1272]" pageId="23" pageNumber="24" refString="Beier, M. (1960) Orthoptera, Tettigoniidae, Pseudophyllinae II. Das Tierreich, 74, 1 - 396." type="journal article" year="1960">Beier 1960</bibRefCitation>
: 365
</treatmentCitation>
; 
<taxonomicName id="4C004D5A0553A90CFEAB28BDFD1B228E" authority="Caudell 1918: 53" authorityName="Caudell" authorityPageNumber="53" authorityYear="1918" box="[377,703,1250,1272]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="undulatum">
<emphasis id="B974EACB0553A90CFEAB28BDFE5B228E" box="[377,511,1250,1272]" italics="true" pageId="23" pageNumber="24">T. undulatum</emphasis>
: 
<treatmentCitation id="0AA110C80553A90CFDDE28BCFD1B228E" author="Caudell" box="[524,703,1250,1272]" page="53" pageId="23" pageNumber="24" year="1918">
<bibRefCitation id="EF914B280553A90CFDDE28BCFD3C228E" author="Caudell" box="[524,664,1250,1272]" pageId="23" pageNumber="24" refString="Caudell, A. N. (1918) On a collection of Orthoptera (exclusive of the Locustidae) made in central Peru by N. Iconnicoff and C. Schunke. Insecutor Insciiae Menstruus, 6, 1 - 70." type="journal article" year="1918">Caudell 1918</bibRefCitation>
: 53
</treatmentCitation>
</taxonomicName>
; 
<treatmentCitation id="0AA110C80553A90CFD1828BCFC34228E" author="Vignon" box="[714,912,1250,1272]" page="269" pageId="23" pageNumber="24" year="1925">
<bibRefCitation id="EF914B280553A90CFD1828BCFCF9228E" author="Vignon" box="[714,861,1250,1272]" pageId="23" pageNumber="24" refString="Vignon, P. (1925 a) Essai de classification du genre Typophyllum Serville (Orth. Phasgon.). A propos des quatre especes nouvelles dont les types sont au Musee de Madrid. EOS, Revista Espanola de Entomologia, 1, 249 - 281." type="journal article" year="1925" yearSuffix="a">Vignon 1925a</bibRefCitation>
: 269
</treatmentCitation>
, 
<treatmentCitation id="0AA110C80553A90CFC4828BCFBF1228E" author="Vignon" box="[922,1109,1250,1272]" page="427" pageId="23" pageNumber="24" year="1930">
<bibRefCitation id="EF914B280553A90CFC4828BCFB86228E" author="Vignon" box="[922,1058,1250,1272]" pageId="23" pageNumber="24" refString="Vignon, P. (1930) Chapitre IV: Mimetisme et faits connexes. Introduction a la Biologie Experimentale (Les etres organises: activites, instincts, structures), 310 - 459." type="book chapter" year="1930">Vignon 1930</bibRefCitation>
: 427
</treatmentCitation>
, 
<treatmentCitation id="0AA110C80553A90CFB8D28BCFABD228E" author="Vignon" box="[1119,1305,1250,1272]" page="134" pageId="23" pageNumber="24" year="1931">
<bibRefCitation id="EF914B280553A90CFB8D28BCFB4C228E" author="Vignon" box="[1119,1256,1250,1272]" pageId="23" pageNumber="24" refString="Vignon, P. (1931) Recherches sur les Sauterelles-Feuilles de l'Amerique tropicale. Archives du Museum d'Histoire Naturelle, Paris, 5, 57 - 212." type="journal article" year="1931">Vignon 1931</bibRefCitation>
: 134
</treatmentCitation>
, 
<treatmentCitation id="0AA110C80553A90CFAF628BCFF542361" author="Beier" page="365" pageId="23" pageNumber="24" year="1960">
<bibRefCitation id="EF914B280553A90CFAF628BCFA3C228E" author="Beier" box="[1316,1432,1250,1272]" pageId="23" pageNumber="24" refString="Beier, M. (1960) Orthoptera, Tettigoniidae, Pseudophyllinae II. Das Tierreich, 74, 1 - 396." type="journal article" year="1960">Beier 1960</bibRefCitation>
: 365
</treatmentCitation>
; 
<taxonomicName id="4C004D5A0553A90CFF28295CFD942361" authority="Vignon 1925a: 270" authorityName="Vignon" authorityPageNumber="270" authorityYear="1925" box="[250,560,1281,1303]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="acutum">
<emphasis id="B974EACB0553A90CFF28295CFEFA2361" box="[250,350,1282,1303]" italics="true" pageId="23" pageNumber="24">T. acutum</emphasis>
: 
<treatmentCitation id="0AA110C80553A90CFEB9295FFD942361" author="Vignon" box="[363,560,1281,1303]" page="270" pageId="23" pageNumber="24" year="1925">
<bibRefCitation id="EF914B280553A90CFEB9295FFE592361" author="Vignon" box="[363,509,1281,1303]" pageId="23" pageNumber="24" refString="Vignon, P. (1925 a) Essai de classification du genre Typophyllum Serville (Orth. Phasgon.). A propos des quatre especes nouvelles dont les types sont au Musee de Madrid. EOS, Revista Espanola de Entomologia, 1, 249 - 281." type="journal article" year="1925" yearSuffix="a">Vignon 1925a</bibRefCitation>
: 270
</treatmentCitation>
</taxonomicName>
, 
<treatmentCitation id="0AA110C80553A90CFDEB295FFD562361" author="Vignon" box="[569,754,1281,1303]" page="135" pageId="23" pageNumber="24" year="1931">
<bibRefCitation id="EF914B280553A90CFDEB295FFD642361" author="Vignon" box="[569,704,1281,1303]" pageId="23" pageNumber="24" refString="Vignon, P. (1931) Recherches sur les Sauterelles-Feuilles de l'Amerique tropicale. Archives du Museum d'Histoire Naturelle, Paris, 5, 57 - 212." type="journal article" year="1931">Vignon 1931</bibRefCitation>
: 135
</treatmentCitation>
, 
<treatmentCitation id="0AA110C80553A90CFD2F295FFC072361" author="Beier" box="[765,931,1281,1303]" page="365" pageId="23" pageNumber="24" year="1960">
<bibRefCitation id="EF914B280553A90CFD2F295FFCD42361" author="Beier" box="[765,880,1281,1303]" pageId="23" pageNumber="24" refString="Beier, M. (1960) Orthoptera, Tettigoniidae, Pseudophyllinae II. Das Tierreich, 74, 1 - 396." type="journal article" year="1960">Beier 1960</bibRefCitation>
: 365
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BBF36D90553A90CFF45291AFB1E2079" blockId="23.[151,1437,1348,2021]" pageId="23" pageNumber="24">
Second group acording to 
<bibRefCitation id="EF914B280553A90CFE1B291AFDDB232B" author="Vignon" box="[457,639,1348,1373]" pageId="23" pageNumber="24" refString="Vignon, P. (1925 a) Essai de classification du genre Typophyllum Serville (Orth. Phasgon.). A propos des quatre especes nouvelles dont les types sont au Musee de Madrid. EOS, Revista Espanola de Entomologia, 1, 249 - 281." type="journal article" year="1925" yearSuffix="a">Vignon (1925a)</bibRefCitation>
, with basally broadened hind tibiae. All three described from unique females (photos in OSF). 
<taxonomicName id="4C004D5A0553A90CFE112937FDC123F6" box="[451,613,1383,1408]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="lacinipenne">
<emphasis id="B974EACB0553A90CFE112937FDC123F6" box="[451,613,1383,1408]" italics="true" pageId="23" pageNumber="24">T. lacinipenne</emphasis>
</taxonomicName>
with mostly green tegmina is from central 
<collectingCountry id="F31776490553A90CFBB22939FB1623F6" box="[1120,1202,1383,1408]" name="Bolivia" pageId="23" pageNumber="24">Bolivia</collectingCountry>
(Provincia Sara). 
<taxonomicName id="4C004D5A0553A90CFA552937FEAB23D3" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="undulatum">
<emphasis id="B974EACB0553A90CFA552937FEAB23D3" italics="true" pageId="23" pageNumber="24">T. undulatum</emphasis>
</taxonomicName>
from central 
<collectingCountry id="F31776490553A90CFE7A29D3FE7F23D2" box="[424,475,1421,1444]" name="Peru" pageId="23" pageNumber="24">Peru</collectingCountry>
is brown, slightly smaller, and has the anal margin of the tegmina distally undulated. The specimen of 
<taxonomicName id="4C004D5A0553A90CFE8B29EFFE6323BE" box="[345,455,1457,1480]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="acutum">
<emphasis id="B974EACB0553A90CFE8B29EFFE6323BE" box="[345,455,1457,1480]" italics="true" pageId="23" pageNumber="24">T. acutum</emphasis>
</taxonomicName>
, also from central 
<collectingCountry id="F31776490553A90CFD4929EEFD6A23B1" box="[667,718,1456,1479]" name="Peru" pageId="23" pageNumber="24">Peru</collectingCountry>
(Pozuzo), with distally slightly undulated tegmina, is yellowish brown and was perhaps greenish in life. Since there are only marginal differences, it does not seem very reasonable to maintain three different names. Hopefully the corresponding males will be discovered soon.
</paragraph>
<paragraph id="8BBF36D90553A903FF152A42FD9526A3" blockId="23.[151,1437,1348,2021]" lastBlockId="24.[151,1436,151,213]" lastPageId="24" lastPageNumber="25" pageId="23" pageNumber="24">
<emphasis id="B974EACB0553A90CFF152A42FEE12043" bold="true" box="[199,325,1564,1589]" pageId="23" pageNumber="24">The songs.</emphasis>
With the calling songs of four species added here, there are now documented the songs of eight or nine 
<taxonomicName id="4C004D5A0553A90CFF1D2A61FEFB202E" box="[207,351,1599,1624]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB0553A90CFF1D2A61FEFB202E" box="[207,351,1599,1624]" italics="true" pageId="23" pageNumber="24">Typophyllum</emphasis>
</taxonomicName>
species: 
<taxonomicName id="4C004D5A0553A90CFE1A2A1FFD99202E" box="[456,573,1599,1624]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="bolivari">
<emphasis id="B974EACB0553A90CFE1A2A1FFD99202E" box="[456,573,1599,1624]" italics="true" pageId="23" pageNumber="24">T. bolivari</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C004D5A0553A90CFD9E2A1FFD58202E" box="[588,764,1599,1624]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="mortuifolium">
<emphasis id="B974EACB0553A90CFD9E2A1FFD58202E" box="[588,764,1599,1624]" italics="true" pageId="23" pageNumber="24">T. mortuifolium</emphasis>
</taxonomicName>
, 
<emphasis id="B974EACB0553A90CFCD92A1FFCBC202E" box="[779,792,1601,1624]" italics="true" pageId="23" pageNumber="24">T</emphasis>
. sp. nr. 
<taxonomicName id="4C004D5A0553A90CFCA62A1EFBA7202E" box="[884,1027,1600,1624]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="trapeziforme">
<emphasis id="B974EACB0553A90CFCA62A1EFBA7202E" box="[884,1027,1600,1624]" italics="true" pageId="23" pageNumber="24">trapeziforme</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF914B280553A90CFBC12A61FB42202E" author="Morris" box="[1043,1254,1599,1624]" pageId="23" pageNumber="24" refString="Morris, G. K., Klimas, D. E. &amp; Nickle, D. (1989) Acoustic signals and systematics of false-leaf katydids from Ecuador (Orthoptera, Tettigoniidae, Pseudophyllinae). Transactions of the American Entomological Society, 114, 215 - 264." type="journal article" year="1989">
Morris 
<emphasis id="B974EACB0553A90CFBBB2A1FFB07202E" box="[1129,1187,1599,1624]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
1989
</bibRefCitation>
, the latter could be 
<taxonomicName id="4C004D5A0553A90CFF6A2A3BFE82200A" box="[184,294,1637,1660]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="morrisi" status="sp. nov.">
<emphasis id="B974EACB0553A90CFF6A2A3BFE82200A" box="[184,294,1637,1660]" italics="true" pageId="23" pageNumber="24">T. morrisi</emphasis>
</taxonomicName>
<emphasis id="B974EACB0553A90CFEFF2A3AFE22200B" bold="true" box="[301,390,1636,1661]" pageId="23" pageNumber="24">
<taxonomicNameLabel id="A24757B00553A90CFEFF2A3AFE22200B" box="[301,390,1636,1661]" pageId="23" pageNumber="24" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
), 
<taxonomicName id="4C004D5A0553A90CFE4F2A3BFCC2200A" authority="Montealegre &amp; Morris 1999" authorityName="Montealegre &amp; Morris" authorityYear="1999" box="[413,870,1636,1661]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="zingara">
<emphasis id="B974EACB0553A90CFE4F2A3BFDAA200A" box="[413,526,1637,1660]" italics="true" pageId="23" pageNumber="24">T. zingara</emphasis>
(Montealegre &amp; Morris 1999)
</taxonomicName>
, 
<emphasis id="B974EACB0553A90CFCA02A3BFC21200A" box="[882,901,1637,1660]" italics="true" pageId="23" pageNumber="24">T.</emphasis>
sp. (
<bibRefCitation id="EF914B280553A90CFC6E2A3AFB5A200A" author="Morris" box="[956,1278,1636,1661]" pageId="23" pageNumber="24" refString="Morris, G. K. &amp; Montealegre-Z., F. (2001) Los Tettigoniidae (Orthoptera: Ensifera) del Parque Regional Nacional Ucumari: Aspectos interesantes de comunicacion acustica. Revista Colombiana de Entomologia, 27, 93 - 105." type="journal article" year="2001">Morris &amp; Montealegre 2001</bibRefCitation>
), 
<taxonomicName id="4C004D5A0553A90CFAC32A3BFA31200A" box="[1297,1429,1637,1660]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB0553A90CFAC32A3BFA31200A" box="[1297,1429,1637,1660]" italics="true" pageId="23" pageNumber="24">T. egregium</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C004D5A0553A90CFF452AD7FE9720D6" box="[151,307,1671,1696]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB0553A90CFF452AD7FE9720D6" box="[151,307,1671,1696]" italics="true" pageId="23" pageNumber="24">T. erosifolium</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C004D5A0553A90CFE962AD7FE1020D6" box="[324,436,1673,1696]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="morrisi" status="sp. nov.">
<emphasis id="B974EACB0553A90CFE962AD7FE1020D6" box="[324,436,1673,1696]" italics="true" pageId="23" pageNumber="24">T. morrisi</emphasis>
</taxonomicName>
<emphasis id="B974EACB0553A90CFE6C2AD9FDBE20D6" bold="true" box="[446,538,1671,1696]" pageId="23" pageNumber="24">
<taxonomicNameLabel id="A24757B00553A90CFE6C2AD9FDBE20D6" box="[446,538,1671,1696]" pageId="23" pageNumber="24" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
and 
<taxonomicName id="4C004D5A0553A90CFD842AD7FCA920D6" box="[598,781,1671,1696]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="onkiosternum" status="sp. nov.">
<emphasis id="B974EACB0553A90CFD842AD7FCA920D6" box="[598,781,1671,1696]" italics="true" pageId="23" pageNumber="24">T. onkiosternum</emphasis>
</taxonomicName>
<emphasis id="B974EACB0553A90CFCC42AD9FCD620D6" bold="true" box="[790,882,1671,1696]" pageId="23" pageNumber="24">
<taxonomicNameLabel id="A24757B00553A90CFCC42AD9FCD620D6" box="[790,882,1671,1696]" pageId="23" pageNumber="24" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(this work, 
<figureCitation id="133B2A5C0553A90CFBD32AD9FBF220D6" box="[1025,1110,1671,1696]" captionStart="FIGURE 19" captionStartId="24.[151,250,1520,1542]" captionTargetBox="[176,1415,294,1472]" captionTargetId="figure@24.[151,1435,265,1499]" captionTargetPageId="24" captionText="FIGURE 19. Oscillograms and linear spectrograms of calling songs of four Typophyllum species: A. T. egregium, complete call (field recording, 9.5 ºC, specimen cbt 002 s 21); B. T. onkiosternum sp. nov., one double pulse of longer sequence (cage, 18 ºC, cbt 004 s 01); C. T. morrisi sp. nov., one pulse and below complete call, here consisting of three pulse groups (cage, 23 ºC, cbt 003 s 01); D. T. erosifolium one pulse and below seven double pulses from continuous calling (cage, 20 ºC, cbt 001 s 01); all recordings made at night." httpUri="https://zenodo.org/record/289373/files/figure.png" pageId="23" pageNumber="24">Fig. 19</figureCitation>
). From two other species of 
<taxonomicName id="4C004D5A0553A90CFF452AF2FE9B20B3" ID-CoL="8PSS6" box="[151,319,1708,1733]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="subFamily" subFamily="Pterochrozinae">Pterochrozinae</taxonomicName>
the songs are known as well: 
<taxonomicName id="4C004D5A0553A90CFD7D2AF3FBDA20B2" authority="Morris &amp; Beier 1982" authorityName="Morris &amp; Beier" authorityYear="1982" box="[687,1150,1708,1733]" class="Insecta" family="Phaneropteridae" genus="Mimetica" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="incisa">
<emphasis id="B974EACB0553A90CFD7D2AF3FCC320B2" box="[687,871,1709,1732]" italics="true" pageId="23" pageNumber="24">Mimetica incisa</emphasis>
(
<bibRefCitation id="EF914B280553A90CFCA82AF2FBD220B2" author="Morris" box="[890,1142,1708,1733]" pageId="23" pageNumber="24" refString="Morris, G. K. &amp; Beier, M. (1982) Song structure and description of some Costa Rican katydids (Orthoptera: Tettigoniidae). Transactions of the American Entomological Society, 108, 287 - 314." type="journal article" year="1982">Morris &amp; Beier 1982</bibRefCitation>
)
</taxonomicName>
and 
<taxonomicName id="4C004D5A0553A90CFB6C2AF2FA3820B2" box="[1214,1436,1708,1733]" class="Insecta" family="Phaneropteridae" genus="Cycloptera" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="arcuata">
<emphasis id="B974EACB0553A90CFB6C2AF2FA3820B2" box="[1214,1436,1708,1733]" italics="true" pageId="23" pageNumber="24">Cycloptera arcuata</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF914B280553A90CFF4C2A8EFDA7209E" author="Braun" box="[158,515,1743,1768]" pageId="23" pageNumber="24" refString="Braun, H. (2002) Die Laubheuschrecken (Orthoptera, Tettigoniidae) eines Bergregenwaldes in Sud-Ecuador - faunistische, bioakustische und okologische Untersuchungen. Doctoral thesis, University of Erlangen-Nurnberg, 142 pp." type="book" year="2002">Braun 2002, recordings in OSF</bibRefCitation>
). In all species the spectrum of the carrier frequencies is extraordinarily narrow, in some species composed of almost pure sine waves. In the large 
<taxonomicName id="4C004D5A0553A90CFCBE2AAAFC4D217B" box="[876,1001,1780,1805]" class="Insecta" family="Phaneropteridae" genus="Cycloptera" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB0553A90CFCBE2AAAFC4D217B" box="[876,1001,1780,1805]" italics="true" pageId="23" pageNumber="24">Cycloptera</emphasis>
</taxonomicName>
male the peak is at 10 kHz, and in 
<taxonomicName id="4C004D5A0553A90CFAAF2AABFF7E2146" class="Insecta" family="Phaneropteridae" genus="Mimetica" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="species" species="incisa">
<emphasis id="B974EACB0553A90CFAAF2AABFF7E2146" italics="true" pageId="23" pageNumber="24">M. incisa</emphasis>
</taxonomicName>
, with the male about the size of the females of small 
<taxonomicName id="4C004D5A0553A90CFCB52B49FC532146" box="[871,1015,1815,1840]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="23" pageNumber="24" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB0553A90CFCB52B49FC532146" box="[871,1015,1815,1840]" italics="true" pageId="23" pageNumber="24">Typophyllum</emphasis>
</taxonomicName>
species (
<quantity id="4CF89B3C0553A90CFBB52B46FB422159" box="[1127,1254,1816,1840]" metricMagnitude="-2" metricUnit="m" metricValue="2.8" metricValueMax="2.9" metricValueMin="2.7" pageId="23" pageNumber="24" unit="mm" value="28.0" valueMax="29.0" valueMin="27.0">27–29 mm</quantity>
tegmen length, 
<bibRefCitation id="EF914B280553A90CFF452B62FE8F2122" author="Vignon" box="[151,299,1852,1877]" pageId="23" pageNumber="24" refString="Vignon, P. (1931) Recherches sur les Sauterelles-Feuilles de l'Amerique tropicale. Archives du Museum d'Histoire Naturelle, Paris, 5, 57 - 212." type="journal article" year="1931">Vignon 1931</bibRefCitation>
), 14 kHz. In the so far studied 
<emphasis id="B974EACB0553A90CFD402B62FCB02123" box="[658,788,1852,1877]" italics="true" pageId="23" pageNumber="24">Typophyllm</emphasis>
males the fequency ranges from 9.5 to 23 kHz, with the smallest species calling at around 20 kHz, which is slightly above the boundary to ultrasound. In these species only very close to a calling male a faint sound is audible to the unaided ear, possibly rather a suppressed fundamental frequency than low-energy components close to the real carrier frequency. With an ultrasound detector, calling males can be heard over fairly long distances. Provided that the females can hear equally well, these maximum broadcast distances probably cover more than the range a phonotactically approaching female can move in one night through understory vegetation.
</paragraph>
<caption id="DF7F6651055CA903FF4529AEFE3D20F7" httpUri="https://zenodo.org/record/289373/files/figure.png" pageId="24" pageNumber="25" targetBox="[176,1415,294,1472]" targetPageId="24">
<paragraph id="8BBF36D9055CA903FF4529AEFE3D20F7" blockId="24.[151,1436,1520,1665]" pageId="24" pageNumber="25">
<emphasis id="B974EACB055CA903FF4529AEFE852073" bold="true" box="[151,289,1520,1542]" pageId="24" pageNumber="25">FIGURE 19.</emphasis>
Oscillograms and linear spectrograms of calling songs of four 
<taxonomicName id="4C004D5A055CA903FC6129AEFB912070" box="[947,1077,1520,1542]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB055CA903FC6129AEFB912070" box="[947,1077,1520,1542]" italics="true" pageId="24" pageNumber="25">Typophyllum</emphasis>
</taxonomicName>
species: 
<emphasis id="B974EACB055CA903FB4429AEFB0B2073" bold="true" box="[1174,1199,1520,1541]" pageId="24" pageNumber="25">A.</emphasis>
<taxonomicName id="4C004D5A055CA903FB6529AFFA942070" box="[1207,1328,1521,1542]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055CA903FB6529AFFA942070" box="[1207,1328,1521,1542]" italics="true" pageId="24" pageNumber="25">T. egregium</emphasis>
</taxonomicName>
, complete call (field recording, 9.5ºC, specimen cbt002s21); 
<emphasis id="B974EACB055CA903FD772A51FD192052" bold="true" box="[677,701,1551,1572]" pageId="24" pageNumber="25">B.</emphasis>
<taxonomicName id="4C004D5A055CA903FD142A4EFCCF2053" box="[710,875,1551,1573]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="onkiosternum" status="sp. nov.">
<emphasis id="B974EACB055CA903FD142A4EFCCF2053" box="[710,875,1551,1573]" italics="true" pageId="24" pageNumber="25">T. onkiosternum</emphasis>
</taxonomicName>
<emphasis id="B974EACB055CA903FCA62A51FC632052" bold="true" box="[884,967,1551,1572]" pageId="24" pageNumber="25">
<taxonomicNameLabel id="A24757B0055CA903FCA62A51FC632052" box="[884,967,1551,1572]" pageId="24" pageNumber="25" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, one double pulse of longer sequence (cage, 18ºC, cbt004s01); 
<emphasis id="B974EACB055CA903FE862A73FEC82035" bold="true" box="[340,364,1581,1603]" pageId="24" pageNumber="25">C.</emphasis>
<taxonomicName id="4C004D5A055CA903FEA12A71FE722032" box="[371,470,1583,1604]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="morrisi" status="sp. nov.">
<emphasis id="B974EACB055CA903FEA12A71FE722032" box="[371,470,1583,1604]" italics="true" pageId="24" pageNumber="25">T. morrisi</emphasis>
</taxonomicName>
<emphasis id="B974EACB055CA903FE0E2A70FD892035" bold="true" box="[476,557,1582,1603]" pageId="24" pageNumber="25">
<taxonomicNameLabel id="A24757B0055CA903FE0E2A70FD892035" box="[476,557,1582,1603]" pageId="24" pageNumber="25" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, one pulse and below complete call, here consisting of three pulse groups (cage, 23ºC, cbt003s01); 
<emphasis id="B974EACB055CA903FEC42A12FE8B2017" bold="true" box="[278,303,1612,1633]" pageId="24" pageNumber="25">D.</emphasis>
<taxonomicName id="4C004D5A055CA903FEE52A13FE602014" box="[311,452,1612,1634]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055CA903FEE52A13FE602014" box="[311,452,1612,1634]" italics="true" pageId="24" pageNumber="25">T. erosifolium</emphasis>
</taxonomicName>
one pulse and below seven double pulses from continuous calling (cage, 20ºC, cbt001s01); all recordings made at night.
</paragraph>
</caption>
<paragraph id="8BBF36D9055CA902FF152AF7FC2E27DB" blockId="24.[151,1437,1704,2017]" lastBlockId="25.[151,1437,151,429]" lastPageId="25" lastPageNumber="26" pageId="24" pageNumber="25">
A somewhat broader carrier frequency spectrum show the partly audio-range 
<taxonomicName id="4C004D5A055CA903FBFF2AF7FB1520B6" box="[1069,1201,1705,1728]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055CA903FBFF2AF7FB1520B6" box="[1069,1201,1705,1728]" italics="true" pageId="24" pageNumber="25">T. egregium</emphasis>
</taxonomicName>
and the ultrasonic 
<taxonomicName id="4C004D5A055CA903FA552AF7FE8B2093" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="onkiosternum" status="sp. nov.">
<emphasis id="B974EACB055CA903FA552AF7FE8B2093" italics="true" pageId="24" pageNumber="25">T. onkiosternum</emphasis>
</taxonomicName>
<emphasis id="B974EACB055CA903FEE82A92FE332093" bold="true" box="[314,407,1740,1765]" pageId="24" pageNumber="25">
<taxonomicNameLabel id="A24757B0055CA903FEE82A92FE332093" box="[314,407,1740,1765]" pageId="24" pageNumber="25" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(
<figureCitation id="133B2A5C055CA903FE782A92FE5B2093" box="[426,511,1740,1765]" captionStart="FIGURE 19" captionStartId="24.[151,250,1520,1542]" captionTargetBox="[176,1415,294,1472]" captionTargetId="figure@24.[151,1435,265,1499]" captionTargetPageId="24" captionText="FIGURE 19. Oscillograms and linear spectrograms of calling songs of four Typophyllum species: A. T. egregium, complete call (field recording, 9.5 ºC, specimen cbt 002 s 21); B. T. onkiosternum sp. nov., one double pulse of longer sequence (cage, 18 ºC, cbt 004 s 01); C. T. morrisi sp. nov., one pulse and below complete call, here consisting of three pulse groups (cage, 23 ºC, cbt 003 s 01); D. T. erosifolium one pulse and below seven double pulses from continuous calling (cage, 20 ºC, cbt 001 s 01); all recordings made at night." httpUri="https://zenodo.org/record/289373/files/figure.png" pageId="24" pageNumber="25">Fig. 19</figureCitation>
A,B). The distribution of 
<taxonomicName id="4C004D5A055CA903FCF92A93FC162092" box="[811,946,1741,1764]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055CA903FCF92A93FC162092" box="[811,946,1741,1764]" italics="true" pageId="24" pageNumber="25">T. egregium</emphasis>
</taxonomicName>
reaches up to almost 
<quantity id="4CF89B3C055CA903FB6A2A93FAB42092" box="[1208,1296,1741,1765]" metricMagnitude="3" metricUnit="m" metricValue="3.0" pageId="24" pageNumber="25" unit="m" value="3000.0">3000 m</quantity>
(
<figureCitation id="133B2A5C055CA903FAF12A92FADD2093" box="[1315,1401,1740,1765]" captionStart="FIGURE 20" captionStartId="25.[151,250,1392,1414]" captionTargetBox="[166,1423,495,1344]" captionTargetId="figure@25.[150,1436,481,1370]" captionTargetPageId="25" captionText="FIGURE 20. Altitudinal distribution of A. Typophyllum egregium and B. T. erosifolium; investigated gradient from 1000 – 3400 m divided in 100 - m intervals, acoustic records based on one survey per locality, the two individuals of T. erosifolium between 1200 m and 1300 m were found at sites where no calling could be detected (Bombuscaro and Alto Nangaritza)." httpUri="https://zenodo.org/record/289374/files/figure.png" pageId="24" pageNumber="25">Fig. 20</figureCitation>
A), where it lives in patches of low elfin woodland (
<figureCitation id="133B2A5C055CA903FD062AAEFC8D217E" box="[724,809,1776,1801]" captionStart="FIGURE 21" captionStartId="26.[151,250,1719,1741]" captionTargetBox="[169,1423,211,1686]" captionTargetId="figure@26.[151,1435,193,1698]" captionTargetPageId="26" captionText="FIGURE 21. Habitat photos: A. at upper distribution limit of Typophyllum egregium in Cajanuma; B, C. understory vegetation and small creek in Alto Nangaritza, in this area were found T. morrisi sp. n., T. erosifolium and T. mortuifolium." httpUri="https://zenodo.org/record/289375/files/figure.png" pageId="24" pageNumber="25">Fig. 21</figureCitation>
A). 
<taxonomicName id="4C004D5A055CA903FC852AAFFBAA217F" box="[855,1038,1776,1801]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="onkiosternum" status="sp. nov.">
<emphasis id="B974EACB055CA903FC852AAFFBAA217F" box="[855,1038,1776,1801]" italics="true" pageId="24" pageNumber="25">T. onkiosternum</emphasis>
</taxonomicName>
<emphasis id="B974EACB055CA903FBCB2AAEFBD2217F" bold="true" box="[1049,1142,1776,1801]" pageId="24" pageNumber="25">
<taxonomicNameLabel id="A24757B0055CA903FBCB2AAEFBD2217F" box="[1049,1142,1776,1801]" pageId="24" pageNumber="25" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is known from very few indivudals that were found around 
<quantity id="4CF89B3C055CA903FDC92B4AFDD6215A" box="[539,626,1812,1837]" metricMagnitude="3" metricUnit="m" metricValue="2.15" pageId="24" pageNumber="25" unit="m" value="2150.0">2150 m</quantity>
. The species that produce pure tone songs, like 
<taxonomicName id="4C004D5A055CA903FB502B4BFABE215B" box="[1154,1306,1812,1837]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055CA903FB502B4BFABE215B" box="[1154,1306,1812,1837]" italics="true" pageId="24" pageNumber="25">T. erosifolium</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C004D5A055CA903FAFA2B4BFA38215B" box="[1320,1436,1812,1837]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="bolivari">
<emphasis id="B974EACB055CA903FAFA2B4BFA38215B" box="[1320,1436,1812,1837]" italics="true" pageId="24" pageNumber="25">T. bolivari</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C004D5A055CA903FF1E2B67FED32126" box="[204,375,1848,1872]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="mortuifoium">
<emphasis id="B974EACB055CA903FF1E2B67FED32126" box="[204,375,1848,1872]" italics="true" pageId="24" pageNumber="25">T. mortuifoium</emphasis>
</taxonomicName>
, all occur in lower elevations, where the trees are high and the understory vegetation is relatively sparse. With increasing elevation, the density of the understory vegetation increases as well, whereas the size of the trees decreases. A similar correlation with habitat structure, that open-habitat species produce songs with a broader spectrum, is reported for birds (e.g. 
<bibRefCitation id="EF914B28055CA903FD022BFAFC7F21CB" author="Bertelli" box="[720,987,1956,1981]" pageId="24" pageNumber="25" refString="Bertelli, S. &amp; Tubaro, P. L. (2002) Body mass and habitat correlates of song structure in a primitive group of birds. Biological Journal of the Linnean Society, 77, 423 - 430. http: // dx. doi. org / 10.1046 / j. 1095 - 8312.2002.00112. x" type="journal article" year="2002">
Bertelli &amp; 
<collectingCountry id="F3177649055CA903FC9A2BFAFC3C21CB" box="[840,920,1956,1981]" name="Tuvalu" pageId="24" pageNumber="25">Tubaru</collectingCountry>
2002
</bibRefCitation>
). Forest bird songs tend to be more pure tone, probably due to a comparatively stable atmosphere, which permits selection to favour mechanisms to increase the sound intensity (
<bibRefCitation id="EF914B28055DA902FE5C2CC6FD8926C6" author="Morton" box="[398,557,151,176]" pageId="25" pageNumber="26" refString="Morton, E. S. (1975) Ecological sources of selection on avian sounds. The American Naturalist, 109, 17 - 34. http: // dx. doi. org / 10.1086 / 282971" type="journal article" year="1975">Morton 1975</bibRefCitation>
). By concentrating all the energy on a very narrow frequency band, the transmission range can be maximized (
<bibRefCitation id="EF914B28055DA902FDB32CE2FCDA26A2" author="Wiley" box="[609,894,188,213]" pageId="25" pageNumber="26" refString="Wiley, R. H. &amp; Richards, D. G. (1978) Physical constraints on acoustic communication in the atmosphere: implications for the evolution of animal vocalizations. Behavioral Ecology and Scociobiology, 3, 69 - 94. http: // dx. doi. org / 10.1007 / BF 00300047" type="journal article" year="1978">Wiley &amp; Richards 1978</bibRefCitation>
). This may be also important for the sparsely distributed forest-dwelling katydids. Broad-band songs might evolve as an adaptation for distance ranging by differential degradation of higher versus lower frequencies in dense vegetation (Montealegre &amp; Morris 2004). This could be particularly important for females of 
<taxonomicName id="4C004D5A055DA902FD652D77FC9B2736" box="[695,831,297,320]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055DA902FD652D77FC9B2736" box="[695,831,297,320]" italics="true" pageId="25" pageNumber="26">T. egregium</emphasis>
</taxonomicName>
in the upper range of the species, where its habitat comprises patches of elfin forest. These are fairly fragmented and separated by areas with low vegetation of a small bamboo species and terrestrial bromeliads, which provide little protection against the harsh weather conditions. So it could be hazardous to approach a male calling from too far away.
</paragraph>
<caption id="DF7F6651055DA902FF45292EFAFD23B5" httpUri="https://zenodo.org/record/289374/files/figure.png" pageId="25" pageNumber="26" targetBox="[166,1423,495,1344]" targetPageId="25">
<paragraph id="8BBF36D9055DA902FF45292EFAFD23B5" blockId="25.[151,1436,1392,1475]" pageId="25" pageNumber="26">
<emphasis id="B974EACB055DA902FF45292EFE8623F3" bold="true" box="[151,290,1392,1414]" pageId="25" pageNumber="26">FIGURE 20.</emphasis>
Altitudinal distribution of 
<emphasis id="B974EACB055DA902FDED292EFDFC23F3" bold="true" box="[575,600,1392,1413]" pageId="25" pageNumber="26">A.</emphasis>
<taxonomicName id="4C004D5A055DA902FDB3292EFCEF23F0" box="[609,843,1392,1414]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055DA902FDB3292EFCEF23F0" box="[609,843,1392,1414]" italics="true" pageId="25" pageNumber="26">Typophyllum egregium</emphasis>
</taxonomicName>
and 
<emphasis id="B974EACB055DA902FC51292EFC3E23F3" bold="true" box="[899,922,1392,1413]" pageId="25" pageNumber="26">B.</emphasis>
<taxonomicName id="4C004D5A055DA902FC71292FFB9523F0" box="[931,1073,1392,1414]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055DA902FC71292FFB9523F0" box="[931,1073,1392,1414]" italics="true" pageId="25" pageNumber="26">T. erosifolium</emphasis>
</taxonomicName>
; investigated gradient from 1000– 3400 m divided in 100-m intervals, acoustic records based on one survey per locality, the two individuals of 
<taxonomicName id="4C004D5A055DA902FADC29CEFA3823D2" box="[1294,1436,1422,1445]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055DA902FADC29CEFA3823D2" box="[1294,1436,1422,1445]" italics="true" pageId="25" pageNumber="26">T. erosifolium</emphasis>
</taxonomicName>
between 1200 m and 1300 m were found at sites where no calling could be detected (Bombuscaro and Alto Nangaritza).
</paragraph>
</caption>
<paragraph id="8BBF36D9055DA902FF1529B5FBE521C5" blockId="25.[151,1436,1514,1971]" pageId="25" pageNumber="26">
In males of the species that produce pure tones, the speculum on the base of the right tegmen, that serves to amplify and spread the sound, seems to be rather stiff and is merely translucent (
<figureCitation id="133B2A5C055DA902FBFE2A50FBD42050" box="[1068,1136,1550,1575]" captionStart="FIGURE 7" captionStartId="9.[151,250,1698,1720]" captionTargetBox="[176,1394,205,1651]" captionTargetId="figure@9.[170,1409,195,1662]" captionTargetPageId="9" captionText="FIGURE 7. Typophyllum morrisi sp. nov.: A, B. male holotype, lateral and dorsal view (cbt 003 s 04); C. male (cbt 003 s 14); D. female (cbt 003 s 10); E. atypical female with uniform anal margin of tegmina (cbt 003 s 12) (see Fig. 5 C for size difference between tiny male and large female)." httpUri="https://zenodo.org/record/289361/files/figure.png" pageId="25" pageNumber="26">Fig. 7</figureCitation>
B, photos of 
<taxonomicName id="4C004D5A055DA902FAD12A51FAD62050" box="[1283,1394,1551,1574]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="morrisi" status="sp. nov.">
<emphasis id="B974EACB055DA902FAD12A51FAD62050" box="[1283,1394,1551,1574]" italics="true" pageId="25" pageNumber="26">T. morrisi</emphasis>
</taxonomicName>
<emphasis id="B974EACB055DA902FAA82A50FF6C203D" bold="true" pageId="25" pageNumber="26">
<taxonomicNameLabel id="A24757B0055DA902FAA82A50FF6C203D" pageId="25" pageNumber="26" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
and 
<taxonomicName id="4C004D5A055DA902FED22A6AFE38203D" box="[256,412,1586,1611]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055DA902FED22A6AFE38203D" box="[256,412,1586,1611]" italics="true" pageId="25" pageNumber="26">T. erosifolium</emphasis>
</taxonomicName>
in OSF). Males of 
<taxonomicName id="4C004D5A055DA902FDAB2A6AFD59203D" box="[633,765,1588,1611]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055DA902FDAB2A6AFD59203D" box="[633,765,1588,1611]" italics="true" pageId="25" pageNumber="26">T. egregium</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C004D5A055DA902FCE52A6AFC48203D" box="[823,1004,1586,1611]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="onkiosternum" status="sp. nov.">
<emphasis id="B974EACB055DA902FCE52A6AFC48203D" box="[823,1004,1586,1611]" italics="true" pageId="25" pageNumber="26">T. onkiosternum</emphasis>
</taxonomicName>
<emphasis id="B974EACB055DA902FC262A6CFBEA203D" bold="true" box="[1012,1102,1586,1611]" pageId="25" pageNumber="26">
<taxonomicNameLabel id="A24757B0055DA902FC262A6CFBEA203D" box="[1012,1102,1586,1611]" pageId="25" pageNumber="26" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, whose songs have a broader spectrum, have a fully transparent speculum (
<figureCitation id="133B2A5C055DA902FD602A08FD5E2019" box="[690,762,1622,1647]" captionStart="FIGURE 4" captionStartId="6.[151,250,1445,1467]" captionTargetBox="[160,1424,203,1410]" captionTargetId="figure@6.[150,1436,193,1423]" captionTargetPageId="6" captionText="FIGURE 4. Typophyllum egregium: A. green female (specimen cbt 002 s 09), B, C. brown female (- s 10), D, E. brown male (- s 03); in live females ovipositor usually concealed by tegmina; note transparent mirror in right tegmen of male (all photos to same scale)." httpUri="https://zenodo.org/record/289358/files/figure.png" pageId="25" pageNumber="26">Fig. 4</figureCitation>
E). This was also found in the neotropical katydid genus 
<taxonomicName id="4C004D5A055DA902FF452A24FE8D20E5" ID-CoL="8NWMZ" box="[151,297,1658,1683]" class="Insecta" family="Tettigoniidae" genus="Panacanthus" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB055DA902FF452A24FE8D20E5" box="[151,297,1658,1683]" italics="true" pageId="25" pageNumber="26">Panacanthus</emphasis>
</taxonomicName>
, with pure tone resonant songs as ancestral state, where in species with non-resonant songs the speculum membranes are glassy and thin (Montealegre &amp; Morris 2004). Such a thin speculum could be more capable of vibrating in intrinsic modes as a result of successive transient tooth-scraper impacts, while the thicker and less transparent speculum of resonant singers is more suitable for a mechanism where the membrane is driven by tooth-contact rate (op. cit.). In males of 
<taxonomicName id="4C004D5A055DA902FD582B52FD5E2155" box="[650,762,1804,1827]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="morrisi" status="sp. nov.">
<emphasis id="B974EACB055DA902FD582B52FD5E2155" box="[650,762,1804,1827]" italics="true" pageId="25" pageNumber="26">T. morrisi</emphasis>
</taxonomicName>
<emphasis id="B974EACB055DA902FCD62B54FCFB2155" bold="true" box="[772,863,1802,1827]" pageId="25" pageNumber="26">
<taxonomicNameLabel id="A24757B0055DA902FCD62B54FCFB2155" box="[772,863,1802,1827]" pageId="25" pageNumber="26" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
and 
<taxonomicName id="4C004D5A055DA902FC482B52FB922155" box="[922,1078,1802,1827]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055DA902FC482B52FB922155" box="[922,1078,1802,1827]" italics="true" pageId="25" pageNumber="26">T. erosifolium</emphasis>
</taxonomicName>
the speculum membrane looks very homogenous and must oscillate uniformly and smoothly between the solid surround. The teeth on the stridulatory file of these two species are microscopically small and very densely packed, compared to the nonresonant 
<taxonomicName id="4C004D5A055DA902FF2F2B29FE2521F8" box="[253,385,1911,1934]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055DA902FF2F2B29FE2521F8" box="[253,385,1911,1934]" italics="true" pageId="25" pageNumber="26">T. egregium</emphasis>
</taxonomicName>
, where they are not so small and tooth density and number are lower. Whereas one male of 
<taxonomicName id="4C004D5A055DA902FA552B29FEB121C5" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055DA902FA552B29FEB121C5" italics="true" pageId="25" pageNumber="26">T. erosifolium</emphasis>
</taxonomicName>
has almost 300 teeth on its file, a male of 
<taxonomicName id="4C004D5A055DA902FD222BC2FCD021C5" box="[752,884,1948,1971]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055DA902FD222BC2FCD021C5" box="[752,884,1948,1971]" italics="true" pageId="25" pageNumber="26">T. egregium</emphasis>
</taxonomicName>
has less than 150.
</paragraph>
<caption id="DF7F6651055EA901FF452AE9FB5B209A" httpUri="https://zenodo.org/record/289375/files/figure.png" pageId="26" pageNumber="27" targetBox="[169,1423,211,1686]" targetPageId="26">
<paragraph id="8BBF36D9055EA901FF452AE9FB5B209A" blockId="26.[151,1436,1719,1772]" pageId="26" pageNumber="27">
<emphasis id="B974EACB055EA901FF452AE9FEBB20BA" bold="true" box="[151,287,1719,1741]" pageId="26" pageNumber="27">FIGURE 21.</emphasis>
Habitat photos: 
<emphasis id="B974EACB055EA901FE1A2AE9FE4520BA" bold="true" box="[456,481,1719,1740]" pageId="26" pageNumber="27">A.</emphasis>
at upper distribution limit of 
<taxonomicName id="4C004D5A055EA901FCDF2AE9FC5120BB" box="[781,1013,1719,1741]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055EA901FCDF2AE9FC5120BB" box="[781,1013,1719,1741]" italics="true" pageId="26" pageNumber="27">Typophyllum egregium</emphasis>
</taxonomicName>
in Cajanuma; 
<emphasis id="B974EACB055EA901FB5B2AE9FB3E20BA" bold="true" box="[1161,1178,1719,1740]" pageId="26" pageNumber="27">B</emphasis>
, 
<emphasis id="B974EACB055EA901FB732AE9FB1D20BB" bold="true" box="[1185,1209,1719,1741]" pageId="26" pageNumber="27">C.</emphasis>
understory vegetation and small creek in Alto Nangaritza, in this area were found 
<taxonomicName id="4C004D5A055EA901FD212A89FCF3209A" box="[755,855,1751,1772]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="morrisi" status="sp. nov.">
<emphasis id="B974EACB055EA901FD212A89FCF3209A" box="[755,855,1751,1772]" italics="true" pageId="26" pageNumber="27">T. morrisi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A24757B0055EA901FC8F2A89FC31209A" box="[861,917,1751,1772]" pageId="26" pageNumber="27" rank="species">sp. n.</taxonomicNameLabel>
, 
<taxonomicName id="4C004D5A055EA901FC722A89FB8F209A" box="[928,1067,1750,1772]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055EA901FC722A89FB8F209A" box="[928,1067,1750,1772]" italics="true" pageId="26" pageNumber="27">T. erosifolium</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C004D5A055EA901FB8E2A89FB5D209A" box="[1116,1273,1750,1772]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="mortuifolium">
<emphasis id="B974EACB055EA901FB8E2A89FB5D209A" box="[1116,1273,1750,1772]" italics="true" pageId="26" pageNumber="27">T. mortuifolium</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BBF36D9055EA900FF152B46FB7C2713" blockId="26.[151,1437,1816,2021]" lastBlockId="27.[151,1437,151,2013]" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
<emphasis id="B974EACB055EA901FF152B46FD8A2147" bold="true" box="[199,558,1816,1841]" pageId="26" pageNumber="27">The curious mating behaviour</emphasis>
. Pre-copulatory riding of the tiny male on one side of the female’s tegmina, as observed in 
<taxonomicName id="4C004D5A055EA901FEF72B62FD8C2122" box="[293,552,1852,1877]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055EA901FEF72B62FD8C2122" box="[293,552,1852,1877]" italics="true" pageId="26" pageNumber="27">Typophyllum egregium</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C004D5A055EA901FDB62B60FCA42123" box="[612,768,1852,1877]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB055EA901FDB62B60FCA42123" box="[612,768,1852,1877]" italics="true" pageId="26" pageNumber="27">T. erosifolium</emphasis>
</taxonomicName>
, is also recorded for 
<taxonomicName id="4C004D5A055EA901FC292B60FB0C2123" box="[1019,1192,1853,1877]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="trapeziforme">
<emphasis id="B974EACB055EA901FC292B60FB0C2123" box="[1019,1192,1853,1877]" italics="true" pageId="26" pageNumber="27">T. trapeziforme</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF914B28055EA901FB6B2B62FF45210F" author="Xiberras" pageId="26" pageNumber="27" refString="Xiberras, S. &amp; Ducaud, P. (2014 b) Nouvelles synonymies dans le genre Typophyllum (Orthoptera: Tettigoniidae: Pseudophyllinae: Pterochrozini). Lambillionea, 114 (3), 227 - 244." type="journal article" year="2014" yearSuffix="b">Xiberras &amp; Ducaud 2014b</bibRefCitation>
), 
<taxonomicName id="4C004D5A055EA901FF262B3CFD19210F" authority="Montealegre &amp; Morris 1999" authorityName="Montealegre &amp; Morris" authorityYear="1999" box="[244,701,1888,1913]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="zingara">
<emphasis id="B974EACB055EA901FF262B3CFEC1210F" box="[244,357,1890,1913]" italics="true" pageId="26" pageNumber="27">T. zingara</emphasis>
(Montealegre &amp; Morris 1999)
</taxonomicName>
, 
<taxonomicName id="4C004D5A055EA901FD1B2B3CFC9F210F" box="[713,827,1888,1913]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="bolivari">
<emphasis id="B974EACB055EA901FD1B2B3CFC9F210F" box="[713,827,1888,1913]" italics="true" pageId="26" pageNumber="27">T. bolivari</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C004D5A055EA901FC992B3CFC5C210F" box="[843,1016,1888,1913]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="mortuifolium">
<emphasis id="B974EACB055EA901FC992B3CFC5C210F" box="[843,1016,1888,1913]" italics="true" pageId="26" pageNumber="27">T. mortuifolium</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C004D5A055EA901FBD52B3CFB2C210F" box="[1031,1160,1890,1913]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="trigonum">
<emphasis id="B974EACB055EA901FBD52B3CFB2C210F" box="[1031,1160,1890,1913]" italics="true" pageId="26" pageNumber="27">T. trigonum</emphasis>
</taxonomicName>
, as well as for 
<taxonomicName id="4C004D5A055EA901FAE02B3EFE8421EA" class="Insecta" family="Phaneropteridae" genus="Roxelana" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="crassicornis">
<emphasis id="B974EACB055EA901FAE02B3EFE8421EA" italics="true" pageId="26" pageNumber="27">Roxelana crassicornis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C004D5A055EA901FE8F2BDAFDEE21EB" box="[349,586,1924,1949]" class="Insecta" family="Phaneropteridae" genus="Pterochroza" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="ocellata">
<emphasis id="B974EACB055EA901FE8F2BDAFDEE21EB" box="[349,586,1924,1949]" italics="true" pageId="26" pageNumber="27">Pterochroza ocellata</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF914B28055EA901FD892BDBFCDE21EB" author="Castner" box="[603,890,1924,1949]" pageId="26" pageNumber="27" refString="Castner, J. L. &amp; Nickle, D. A. (1995 a) Observations on the behavior and biology of leaf-mimicking katydids (Orthoptera: Tettigoniidae: Pseudophyllinae: Pterochrozini). Journal of Orthoptera Research, 4, 93 - 98. http: // dx. doi. org / 10.2307 / 3503463" type="journal article" year="1995" yearSuffix="a">Castner &amp; Nickle 1995a</bibRefCitation>
; whereas this is doubtful for the latter species, where the male reaches almost the same size as the female, and which apparently shows a different behaviour— P. Ducaud pers. comm.). In 
<taxonomicName id="4C004D5A055EA901FE672B90FD9D2192" box="[437,569,1997,2021]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055EA901FE672B90FD9D2192" box="[437,569,1997,2021]" italics="true" pageId="26" pageNumber="27">T. egregium</emphasis>
</taxonomicName>
the riding of the male on the female’s tegmina preceded mating, and the two always separated after having accomplished the protracted transfer of the spermatophore. Likewise proceeds 
<taxonomicName id="4C004D5A055FA900FA552CC7FE8226A2" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="trapeziforme">
<emphasis id="B974EACB055FA900FA552CC7FE8226A2" italics="true" pageId="27" pageNumber="28">T. trapeziforme</emphasis>
</taxonomicName>
, where the male also rides several days and always leaves the female after mating (P. Ducaud pers. comm.). However, Castner &amp; Nickle (who unfortunately neither assign the behaviour to a certain species nor provide details of their nine field and more than 100 cage observations) report the male remounting the female after mating and resume riding for several hours to several days. This seems to conflict most of the speculations about the adaptiveness of this peculiar behaviour already discussed by the authors themselves (op. cit.).
</paragraph>
<paragraph id="8BBF36D9055FA900FF152D2EFB3E244A" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="28">
One possible explanation for the pre-copulatory riding might be associated with the low probability for males and females to meet, due to the low population density, a widely acknowledged feature of arthropods in tropical forests (e.g. 
<bibRefCitation id="EF914B28055FA900FEFA2DE9FE0E27A6" author="Elton" box="[296,426,439,464]" pageId="27" pageNumber="28" refString="Elton, C. S. (1973) The structure of invertebrate populations inside neotropical rainforest. Journal of Animal Ecology, 42, 55 - 104. http: // dx. doi. org / 10.2307 / 3406" type="journal article" year="1973">Elton 1973</bibRefCitation>
, 
<bibRefCitation id="EF914B28055FA900FE6A2DE6FD7127A6" author="Novotny" box="[440,725,440,464]" pageId="27" pageNumber="28" refString="Novotny, V. &amp; Basset, Y. (2000) Rare species in communities of tropical insect herbivores: Pondering the mystery of singletons. Oikos, 89, 564 - 572. http: // dx. doi. org / 10.1034 / j. 1600 - 0706.2000.890316. x" type="journal article" year="2000">Novotny &amp; Basset 2000</bibRefCitation>
). Therefore, one or both partners might for some economical reason not always be disposed to mate immediately. But since the male does not feed while atop the female, he could only reallocate internal resources to produce a spermatophore. The female should not have any problems with receptivity, since adult female tettigoniids have a spermatheca to store sperm until use.
</paragraph>
<paragraph id="8BBF36D9055FA900FF152E19FA8A25F6" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="28">
Another potential explanation discussed by 
<bibRefCitation id="EF914B28055FA900FD062E16FBA82416" author="Castner" box="[724,1036,583,608]" pageId="27" pageNumber="28" refString="Castner, J. L. &amp; Nickle, D. A. (1995 a) Observations on the behavior and biology of leaf-mimicking katydids (Orthoptera: Tettigoniidae: Pseudophyllinae: Pterochrozini). Journal of Orthoptera Research, 4, 93 - 98. http: // dx. doi. org / 10.2307 / 3503463" type="journal article" year="1995" yearSuffix="a">Castner &amp; Nickle (1995a)</bibRefCitation>
, mate-guarding by the male, can probably also be excluded, as the peaceful contiguous riding and subsequent mating of three males with the same female in caged 
<taxonomicName id="4C004D5A055FA900FE9D2ECFFE7724DE" box="[335,467,657,680]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055FA900FE9D2ECFFE7724DE" box="[335,467,657,680]" italics="true" pageId="27" pageNumber="28">T. egregium</emphasis>
</taxonomicName>
showed. Furthermore, it does not seem to be necessary because of rarity in the natural habitat—‘chance encounters’ of male and female (op.cit.) are highly unlikely and the males have to rely on their calling song to bring about a rendezvous. However, if the male should actually resume the piggyback position after mating (op.cit.), it could be adaptive for the pair to stay together. It would save the female future toilsome and possibly dangerous phonotactic approaches, and the male the calling, which might attract predators or parasitoids. But if the probability for the male to attract another female before his previous mate uses up all his sperm is higher than to attract the interest of an eavesdropping bat, he should desert her (or perhaps resume calling atop).
</paragraph>
<paragraph id="8BBF36D9055FA900FF152FD2FB592243" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="28">
According to another assumption (op.cit.) the female might evaluate the male by assessing his weight. But why should she venture a phonotactic approach at all (calling already can convey information on male quality, e.g. 
<bibRefCitation id="EF914B28055FA900FF452F8AFE9A259A" author="Gwynne" box="[151,318,980,1004]" pageId="27" pageNumber="28" refString="Gwynne, D. T. (2001) Katydids and bush-crickets: reproductive behavior and evolution of the Tettigoniidae. Cornell University Press, 317 pp." type="book" year="2001">Gwynne 2001</bibRefCitation>
), and then carry him around for several days instead of retrieving on the spot a more or less nutritious spermathophore and get rid of him? An instance of a male 
<taxonomicName id="4C004D5A055FA900FC482FA7FBB92266" box="[922,1053,1017,1040]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055FA900FC482FA7FBB92266" box="[922,1053,1017,1040]" italics="true" pageId="27" pageNumber="28">T. egregium</emphasis>
</taxonomicName>
, who descended after five days of riding on the female without mating, rather suggests the possibility of the male assessing the female.
</paragraph>
<paragraph id="8BBF36D9055FA900FF152861FBBE2346" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="28">
Possibly the piggyback behaviour of the little walking leaves is one facet of their sophisticated crypticity (see next section). 
<bibRefCitation id="EF914B28055FA900FEE5283AFDF9220B" author="Castner" box="[311,605,1124,1149]" pageId="27" pageNumber="28" refString="Castner, J. L. &amp; Nickle, D. A. (1995 a) Observations on the behavior and biology of leaf-mimicking katydids (Orthoptera: Tettigoniidae: Pseudophyllinae: Pterochrozini). Journal of Orthoptera Research, 4, 93 - 98. http: // dx. doi. org / 10.2307 / 3503463" type="journal article" year="1995" yearSuffix="a">Castner &amp; Nickle (1995a)</bibRefCitation>
already presumed that a male sitting on the wings of a female might foil the search image of predators. But perhaps it is the female alone who obliges the male to ride on her. If the curious ensemble attracts the attention of a predator despite its camouflage, the small male ‘sitting on a leaf’ is spotted first, while the female can escape. For instance in stick insects survival of females can improve considerably with dorsally coupled males (
<bibRefCitation id="EF914B28055FA900FE6A28AAFDFA237A" author="Sivinski" box="[440,606,1268,1293]" pageId="27" pageNumber="28" refString="Sivinski, J. (1983) Predation and sperm competition in the evolution of coupling durations, particularly in the stick insect Diapheromera veliei. In: Gwynne, D. T. &amp; Morris, G. K. (Eds.), Orthopteran Mating Systems: Sexual Competition in a Diverse Group of Insects. Westview Press, Boulder, Colorado, 147 - 162." type="book chapter" year="1983">Sivinski 1983</bibRefCitation>
). Probably the time the male can spend riding (up to five days in 
<taxonomicName id="4C004D5A055FA900FA5528ABFEA42346" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB055FA900FA5528ABFEA42346" italics="true" pageId="27" pageNumber="28">T. egregium</emphasis>
</taxonomicName>
) is limited by starvation, unless he descends from time to time to feed.
</paragraph>
<paragraph id="8BBF36D9055FA900FF152962FB1923B6" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="28">
Likewise mysterious as the reason for this behaviour remains presently the question whether the sexual size dimorphism in 
<taxonomicName id="4C004D5A055FA900FE962901FE70230E" box="[324,468,1375,1400]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB055FA900FE962901FE70230E" box="[324,468,1375,1400]" italics="true" pageId="27" pageNumber="28">Typophyllum</emphasis>
</taxonomicName>
, which is extraordinarily pronounced for a katydid, might be an adaptation to it. This seems to be the case in certain stick insects with prolonged mating, were the female carries around the smaller male. In those species the sexual size difference is especially distinctive (
<bibRefCitation id="EF914B28055FA900FC1829F9FBCD23B6" author="Sivinski" box="[970,1129,1447,1472]" pageId="27" pageNumber="28" refString="Sivinski, J. (1978) Intrasexual aggression in the stick insects Diapheromera veliei and D. covillea and sexual dimorphism in the Phasmatodea. Psyche, 85, 395 - 405. http: // dx. doi. org / 10.1155 / 1978 / 35784" type="journal article" year="1978">Sivinski 1978</bibRefCitation>
, 
<bibRefCitation id="EF914B28055FA900FBA629F6FB1423B6" author="Sivinski" box="[1140,1200,1448,1472]" pageId="27" pageNumber="28" refString="Sivinski, J. (1983) Predation and sperm competition in the evolution of coupling durations, particularly in the stick insect Diapheromera veliei. In: Gwynne, D. T. &amp; Morris, G. K. (Eds.), Orthopteran Mating Systems: Sexual Competition in a Diverse Group of Insects. Westview Press, Boulder, Colorado, 147 - 162." type="book chapter" year="1983">1983</bibRefCitation>
).
</paragraph>
<paragraph id="8BBF36D9055FA900FF152992FC8321E2" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="28">
<emphasis id="B974EACB055FA900FF152992FD4D2393" bold="true" box="[199,745,1484,1509]" pageId="27" pageNumber="28">Why are the little walking leaves so leaf-like?</emphasis>
Already early orthopterists were intrigued by the perfection of mimetism in which the 
<taxonomicName id="4C004D5A055FA900FE0229B1FDDC207E" ID-CoL="8PSS6" box="[464,632,1519,1544]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="subFamily" subFamily="Pterochrozinae">Pterochrozinae</taxonomicName>
excel. While the tegmina with their venation are predestined for the imitation of a leaf, as many other katydid species apparently use as simple camouflage, the little walking leaves additionally exhibit various details that mimic a real separate leaf in various stages of decay. These include tiny transparent windows, brownish patches and margins in otherwise green individuals, undulated or excised margins, light greyish spots, and minute whitish or dark tubercles—all even through a stereomicroscope looking like real holes caused by small herbivorous insects, necrotic spots, lichens, or fungi. In the larger females the dorsal edge of the tegmina is sometimes slightly twisted, so that in dorsal view the midline of the tegmina is s-shaped, just like a withered leaf. Brown body parts sometimes are partly green and “mossy”. Nymphs, that cannot mimic a leaf due to the undeveloped tegmina, frequently have light green and ramified processes on the abdominal tergites, that look like real moss (
<figureCitation id="133B2A5C055FA900FE932B6AFE2B213A" box="[321,399,1844,1869]" captionStart="FIGURE 9" captionStartId="11.[151,250,1970,1992]" captionTargetBox="[173,1412,1003,1888]" captionTargetId="figure@11.[151,1436,981,1924]" captionTargetPageId="11" captionText="FIGURE 9. Typophyllum onkiosternum sp. nov.: A. female (paratype, cbt 005 s 01); B. female nymph (cbt 002 s 16)." httpUri="https://zenodo.org/record/289363/files/figure.png" pageId="27" pageNumber="28">Figs. 9</figureCitation>
B, 11C). The antenna tips are often curled (
<figureCitation id="133B2A5C055FA900FCA22B6AFC6E213A" box="[880,970,1844,1869]" captionStart="FIGURE 11" captionStartId="13.[151,250,1460,1482]" captionTargetBox="[162,1425,203,1426]" captionTargetId="figure@13.[151,1435,193,1438]" captionTargetPageId="13" captionText="FIGURE 11. Typophyllum onkiosternum sp. nov. (A – C): A. female (paratype, tegmen length 24 mm), B. male (holotype, tegmen length 14.5 mm), C. female nymph (cbt 002 s 12); Typophyllum sp., unknown male of species described from females only or new species (D – F, cbt 003 s 08): D. nymph, E, F. adult; G. Typophyllum bolivari, nymph; photos of live individuals staged during day and approximately to scale." httpUri="https://zenodo.org/record/289365/files/figure.png" pageId="27" pageNumber="28">Figs. 11</figureCitation>
D, 15G, 16), reminiscent of a tiny vine, as in other camouflaged katydids, like the little lichen dragon 
<taxonomicName id="4C004D5A055FA900FC962B09FBE72106" box="[836,1091,1879,1904]" class="Insecta" family="Phaneropteridae" genus="Lichenodraculus" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="matti">
<emphasis id="B974EACB055FA900FC962B09FBE72106" box="[836,1091,1879,1904]" italics="true" pageId="27" pageNumber="28">Lichenodraculus matti</emphasis>
</taxonomicName>
(
<taxonomicName id="4C004D5A055FA900FB872B09FAB42106" ID-CoL="8NKZD" box="[1109,1296,1879,1904]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="subFamily" subFamily="Phaneropterinae">Phaneropterinae</taxonomicName>
, 
<taxonomicName id="4C004D5A055FA900FACF2B09FA342106" ID-CoL="8PT46" box="[1309,1424,1879,1904]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Dysoniini">Dysoniini</taxonomicName>
), that was also found in the investigation area (
<bibRefCitation id="EF914B28055FA900FD402B22FCBF21E2" author="Braun" box="[658,795,1916,1940]" pageId="27" pageNumber="28" refString="Braun, H. (2011) The Litte Lichen Dragon - an extraordinary katydid from the Ecuadorian Andes (Orthoptera, Tettigoniidae, Phaneropterinae, Dysoniini). Zootaxa, 3032, 33 - 39." type="journal article" year="2011">Braun 2011</bibRefCitation>
).
</paragraph>
<paragraph id="8BBF36D9055FA907FF152BFEFF722783" blockId="27.[151,1437,151,2013]" lastBlockId="28.[151,1437,151,2013]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
Back in 1894, the eminent orthopterist Brunner von Wattenwyl considered these numerous details as exuberance and an effort far in excess of the necessary, which could not be attributed exclusively to natural selection. 
<bibRefCitation id="EF914B280558A907FED82CC9FE1F26C6" author="Vignon" box="[266,443,151,176]" pageId="28" pageNumber="29" refString="Vignon, P. (1925 a) Essai de classification du genre Typophyllum Serville (Orth. Phasgon.). A propos des quatre especes nouvelles dont les types sont au Musee de Madrid. EOS, Revista Espanola de Entomologia, 1, 249 - 281." type="journal article" year="1925" yearSuffix="a">Vignon (1925a)</bibRefCitation>
concurred with this view and thought the luxury of accessories to be rather contrary to real mimetism. Still in his fine and beautifully illustrated monograph (1931) he came to the conclusion that the detailed resemblance to decayed leaves, or leaves apparently mined or eaten by caterpillars, was useless, his reason being that other species with the much simpler likeness to uninjured leaves are able to hold their own in the struggle with greater success, as shown by their comparative abundance. He considered the details as a decoration unnecessary in the life of the insects, a multiform comedy, mimetism and aesthetics being two aspects of a sole mystery. In contrast 
<bibRefCitation id="EF914B280558A907FE532D2EFDBF27FE" author="Karny" box="[385,539,368,392]" pageId="28" pageNumber="29" refString="Karny, H. (1914). Schutzanpassung der Heuschrecken. Jahresbericht des K. K. Maximilian-Gymnasiums in Wien, 3 - 14." type="book chapter" year="1914">Karny (1914)</bibRefCitation>
already assumed that to deceive sharp-eyed insect-feeding birds it requires an imitation of the leaf in colour and shape up to the smallest detail. He still entertained some doubt that selection alone was sufficient for such crypticity and contemplated Lamarck’s (1809) theory of active acquisition of useful traits.
</paragraph>
<paragraph id="8BBF36D90558A907FF152DA1FE21259A" blockId="28.[151,1437,151,2013]" pageId="28" pageNumber="29">
The different birds of mixed species flocks in the investigation area actually searched very meticulously bark, foliage, bromeliads and other epiphytes, including all sorts of crevices and possible hiding places of arthropod prey (pers. obs.). Since the little walking leaves furthermore come in multifarious outfits (as shown particularly in 
<taxonomicName id="4C004D5A0558A907FA552E17FEA424F2" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="egregium">
<emphasis id="B974EACB0558A907FA552E17FEA424F2" italics="true" pageId="28" pageNumber="29">T. egregium</emphasis>
</taxonomicName>
), it becomes even more difficult for birds as the main visual predators to develop a search image, a term introduced by 
<bibRefCitation id="EF914B280558A907FEE82ED1FDA524DE" author="Tinbergen" box="[314,513,655,680]" pageId="28" pageNumber="29" refString="Tinbergen, L. (1960) The natural control of insects in pine woods: Vol. I. Factors influencing the intensity of predation by songbirds. Archives Neerlandaises de Zoologie, 13, 265 - 343. http: // dx. doi. org / 10.1163 / 036551660 X 00053" type="journal article" year="1960">Tinbergen (1960)</bibRefCitation>
. Such polymorphism in colouration has been recorded in a variety of cryptic prey species (
<bibRefCitation id="EF914B280558A907FF292EEAFD5B24BB" author="Bond" box="[251,767,692,717]" pageId="28" pageNumber="29" refString="Bond, A. B. &amp; Kamil, A. C. (2002) Visual predators select for crypticity and polymorphism in virtual prey. Nature, 415, 609 - 613. http: // dx. doi. org / 10.1038 / 415609 a" type="journal article" year="2002">Bond &amp; Kamil 2002 and references therein</bibRefCitation>
) and is shown to be the potential outcome of frequency dependent selection (op. cit.), since predators have to allocate their attention economically (
<bibRefCitation id="EF914B280558A907FB0B2E89FEC32563" author="Dukas" pageId="28" pageNumber="29" refString="Dukas, R. (2004) Causes and consequences of limited attention. Brain, Behavior and Evolution, 63, 107 - 210. http: // dx. doi. org / 10.1159 / 000076781" type="journal article" year="2004">Dukas 2004 and references therein</bibRefCitation>
). The underlying mechanisms are complicated and not fully understood (
<bibRefCitation id="EF914B280558A907FB6C2EA2FF70254E" author="Gray" pageId="28" pageNumber="29" refString="Gray, S. M. &amp; McKinnon, J. S. (2007) Linking color polymorphism maintenance and speciation. Trends in Ecology and Evolution 22, 71 - 79. http: // dx. doi. org / 10.1016 / j. tree. 2006.10.005" type="journal article" year="2007">Gray &amp; McKinnon 2007</bibRefCitation>
). Through the piggyback behaviour males and females stay together during day and can perceive colour, so even some sort of “assortative” mating is thinkable. The little walking leaves held in cages did not choose their daily roosting places according to their own colour, although differently coloured substrates were available, not confirming observations for 
<taxonomicName id="4C004D5A0558A907FE3D2FD3FC1A25D2" authority="Bellwood 1988" authorityName="Bellwood" authorityYear="1988" box="[495,958,908,933]" class="Insecta" family="Phaneropteridae" genus="Mimetica" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="mortuifolia">
<emphasis id="B974EACB0558A907FE3D2FD3FD4225D3" box="[495,742,908,933]" italics="true" pageId="28" pageNumber="29">Mimetica mortuifolia</emphasis>
(Bellwood 1988)
</taxonomicName>
, but consistent with other observations (
<bibRefCitation id="EF914B280558A907FF4C2FEEFE6525BE" author="Castner" box="[158,449,943,968]" pageId="28" pageNumber="29" refString="Castner, J. L. &amp; Nickle, D. A. (1995 a) Observations on the behavior and biology of leaf-mimicking katydids (Orthoptera: Tettigoniidae: Pseudophyllinae: Pterochrozini). Journal of Orthoptera Research, 4, 93 - 98. http: // dx. doi. org / 10.2307 / 3503463" type="journal article" year="1995" yearSuffix="a">Castner &amp; Nickle 1995a</bibRefCitation>
). And why should a little leaf as an object of its own have the same colour as the background anyway?
</paragraph>
<paragraph id="8BBF36D90558A907FF152FA6FC5723EB" blockId="28.[151,1437,151,2013]" pageId="28" pageNumber="29">
In the investigation area the little walking leaves were quite rare. This fits into an additional line of reasoning (
<bibRefCitation id="EF914B280558A907FF4C2842FEFE2242" author="Hochkirch" box="[158,346,1052,1077]" pageId="28" pageNumber="29" refString="Hochkirch, A. (2001) Der Zwang zu sparen - oder: Was hat Biodiversitat mit Pradation zu tun? 14. Jahrestagung der Gesellschaft fur Tropenokologie. Tagungsband." type="book chapter" year="2001">Hochkirch 2001</bibRefCitation>
): a high predation pressure combined with habitat stability might favour individuals that invest in high life expectancy (a 
<taxonomicName id="4C004D5A0558A907FE4D281FFD9D222E" box="[415,569,1087,1112]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="erosifolium">
<emphasis id="B974EACB0558A907FE4D281FFD9D222E" box="[415,569,1087,1112]" italics="true" pageId="28" pageNumber="29">T. erosifolium</emphasis>
</taxonomicName>
male caught as adult lived for nine more months, calling almost every night) and produce small clutches (one egg per site in unspecified 
<taxonomicName id="4C004D5A0558A907FCE8283AFC41220B" ID-CoL="8PSS6" box="[826,997,1124,1149]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Pterochrozinae">Pterochrozinae</taxonomicName>
, 
<bibRefCitation id="EF914B280558A907FC23283AFB5C220B" author="Caster" box="[1009,1272,1124,1149]" pageId="28" pageNumber="29" refString="Castner, J. L. &amp; Nickle, D. A. (1995 a) Observations on the behavior and biology of leaf-mimicking katydids (Orthoptera: Tettigoniidae: Pseudophyllinae: Pterochrozini). Journal of Orthoptera Research, 4, 93 - 98. http: // dx. doi. org / 10.2307 / 3503463" type="journal article" year="1995" yearSuffix="a">Caster &amp; Nickle 1995a</bibRefCitation>
), therefore, do not have to feed and move much. The resulting low reproduction rate would explain the low population density, which in turn complicates prey specialization by predators. Low population densities also affect evolution. Speciation might be accelerated by rarity, small distribution areas and low dispersal abilities (
<bibRefCitation id="EF914B280558A907FB162891FAF8229E" author="Chown" box="[1220,1372,1231,1256]" pageId="28" pageNumber="29" refString="Chown, S. L. (1997) Speciation and rarity: separating cause from consequence. In: Kunin, W. E. &amp; Gaston, K. J. (Eds.), The Biology of Rarity. Chapman &amp; Hall, London, 91 - 109." type="book chapter" year="1997">Chown 1997</bibRefCitation>
). The underlying (and controversial, op.cit.) logic concerns gene flow, which tends to prevent speciation in abundant species with broad distribution areas (
<bibRefCitation id="EF914B280558A907FD9A2946FD6E2346" author="Mayr" box="[584,714,1303,1328]" pageId="28" pageNumber="29" refString="Mayr, E. 1963. Animal species and evolution. Harvard University Press, Cambridge. http: // dx. doi. org / 10.4159 / harvard. 9780674865327" type="book" year="1963">Mayr 1963</bibRefCitation>
, 
<bibRefCitation id="EF914B280558A907FD052949FCCB2346" author="Stanley" box="[727,879,1303,1328]" pageId="28" pageNumber="29" refString="Stanley, S. M. (1979) Macroevolution. Pattern and Process. W. H. Freeman, San Francisco." type="book" year="1979">Stanley 1979</bibRefCitation>
). Similar predictions make the theory of shifting balance (
<bibRefCitation id="EF914B280558A907FF2D2962FE302322" author="Wright" box="[255,404,1340,1365]" pageId="28" pageNumber="29" refString="Wright, S. (1931) Evolution in Mendelian populations. Genetics, 16, 97 - 159." type="journal article" year="1931">Wright 1931</bibRefCitation>
, 
<bibRefCitation id="EF914B280558A907FE762962FE7B2322" author="Wright" box="[420,479,1340,1364]" pageId="28" pageNumber="29" refString="Wright, S. (1977) Evolution and the Genetics of Populations. Vol. 3 Experimental Results and Evolutionary Deductions. University of Chicago Press, Chicago." type="journal volume" year="1977">1977</bibRefCitation>
, 
<bibRefCitation id="EF914B280558A907FE3C2962FD3D2323" author="Eldredge" box="[494,665,1340,1365]" pageId="28" pageNumber="29" refString="Eldredge, N. (1985) Unfinished synthesis: biological hierarchies and modern evolutionary thought. Oxford University Press, Oxford." type="book" year="1985">Eldredge 1985</bibRefCitation>
, 
<bibRefCitation id="EF914B280558A907FD7A2962FC862322" author="Holt" box="[680,802,1340,1365]" pageId="28" pageNumber="29" refString="Holt, R. D. (1997) Rarity and evolution: some theortical considerations. In: Kunin, W. E. &amp; Gaston, K. J. (Eds.), The Biology of Rarity. Chapman &amp; Hall, London, 209 - 234. http: // dx. doi. org / 10.1007 / 978 - 94 - 011 - 5874 - 9 _ 12" type="book chapter" year="1997">Holt 1997</bibRefCitation>
). The potentially creative role of random genetic drift permits small populations to move between adaptive peaks, while in large populations adaptive evolution is less likely and they are in some measure evolutionary inert because of gene flow.
</paragraph>
<paragraph id="8BBF36D90558A907FF1529F9FE8C2003" blockId="28.[151,1437,151,2013]" pageId="28" pageNumber="29">
These merely indicated mechanisms might have contributed to the extraordinarily perfect imitation of tiny leaves, including all the microscopic accessories like factitious fungi cultures on the tegmina. If the tropical diversity is not already very old compared to temperate regions, then the speciation rate must be especially high (e.g. 
<bibRefCitation id="EF914B280558A907FF012A4AFE47205A" author="Chown" box="[211,483,1556,1581]" pageId="28" pageNumber="29" refString="Chown, S. L. &amp; Gaston, K. J. (2000) Areas, cradles and museums: the latitudinal gradient in species richness. Trends in Ecology and Evolution, 15, 311 - 315. http: // dx. doi. org / 10.1016 / S 0169 - 5347 (00) 01910 - 8" type="journal article" year="2000">Chown &amp; Gaston 2000</bibRefCitation>
). Additionally the predation pressure in the tropics is thought to be higher than in temperate ecosystems (
<bibRefCitation id="EF914B280558A907FE492A69FDDA2026" author="Hawkins" box="[411,638,1591,1616]" pageId="28" pageNumber="29" refString="Hawkins, B. A., Cornell, H. V. &amp; Hochberg, M. E. (1997) Predators, parasitoids and pathogens as mortality agents in phytophagous insect populations. Ecology, 78, 2145 - 2152. http: // dx. doi. org / 10.1890 / 0012 - 9658 (1997) 078 [2145: PPAPAM] 2.0. CO; 2" type="journal article" year="1997">
Hawkins 
<emphasis id="B974EACB0558A907FDD52A67FD9B2026" box="[519,575,1591,1616]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
1997
</bibRefCitation>
and references therein), probably again as a result of rarity of potential prey species.
</paragraph>
<paragraph id="8BBF36D90558A906FF152A21FC4B276B" blockId="28.[151,1437,151,2013]" lastBlockId="29.[151,1436,151,285]" lastPageId="29" lastPageNumber="30" pageId="28" pageNumber="29">
<emphasis id="B974EACB0558A907FF152A21FCDB20EE" bold="true" box="[199,895,1663,1688]" pageId="28" pageNumber="29">Where did the walking leaves originate in the tree of life?</emphasis>
A recent molecular phylogeny of 
<taxonomicName id="4C004D5A0558A907FADB2A21FA3820EE" ID-CoL="8NKR2" box="[1289,1436,1663,1688]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Tettigoniidae</taxonomicName>
includes a remarkable result: a clade consisting of 
<taxonomicName id="4C004D5A0558A907FD0A2AFBFC0D20CB" box="[728,937,1700,1725]" class="Insecta" family="Phaneropteridae" genus="Mimetica" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="tuberata">
<emphasis id="B974EACB0558A907FD0A2AFBFC0D20CB" box="[728,937,1700,1725]" italics="true" pageId="28" pageNumber="29">Mimetica tuberata</emphasis>
</taxonomicName>
from 
<collectingCountry id="F31776490558A907FC222AFAFBCF20CB" box="[1008,1131,1700,1725]" name="Costa Rica" pageId="28" pageNumber="29">Costa Rica</collectingCountry>
and 
<taxonomicName id="4C004D5A0558A907FB762AFAFA9020CB" box="[1188,1332,1700,1725]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB0558A907FB762AFAFA9020CB" box="[1188,1332,1700,1725]" italics="true" pageId="28" pageNumber="29">Typophyllum</emphasis>
</taxonomicName>
sp. from 
<collectingCountry id="F31776490558A907FF452A97FF6F2096" box="[151,203,1737,1760]" name="Peru" pageId="28" pageNumber="29">Peru</collectingCountry>
is found to be a sister group to all other katydids (
<bibRefCitation id="EF914B280558A907FCD02A99FC5B2096" author="Mugleston" box="[770,1023,1735,1760]" pageId="28" pageNumber="29" refString="Mugleston, J. D., Song, H. &amp; Whiting, M. F. (2013) A century of paraphyly: A molecular phylogeny of katydids (Orthoptera: Tettigoniidae) supports multiple origins of leaf-like wings. Molecular Phylogenetics and Evolution, 69, 1120 - 1134. http: // dx. doi. org / 10.1016 / j. ympev. 2013.07.014" type="journal article" year="2013">
Mugleston 
<emphasis id="B974EACB0558A907FC502A97FC1F2096" box="[898,955,1735,1760]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2013
</bibRefCitation>
). Traditionally the group was mostly treated as tribe 
<taxonomicName id="4C004D5A0558A907FE992AB2FE462173" ID-CoL="8PTJ3" box="[331,482,1772,1797]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Pterochrozini">Pterochrozini</taxonomicName>
within 
<taxonomicName id="4C004D5A0558A907FDEF2AB2FD522173" ID-CoL="8NL27" box="[573,758,1772,1797]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Pseudophyllinae">Pseudophyllinae</taxonomicName>
(e.g. 
<bibRefCitation id="EF914B280558A907FCEE2AB2FC1A2173" author="Beier" box="[828,958,1772,1797]" pageId="28" pageNumber="29" refString="Beier, M. (1960) Orthoptera, Tettigoniidae, Pseudophyllinae II. Das Tierreich, 74, 1 - 396." type="journal article" year="1960">Beier 1960</bibRefCitation>
), sometimes as subfamily (e.g. 
<bibRefCitation id="EF914B280558A907FAFE2AB2FF76215E" author="Gorochov" pageId="28" pageNumber="29" refString="Gorochov, A. V. (2012) Systematics of the American katydids (Orthoptera: Tettigoniidae). Communication 1. Proceedings of the Zoological Institute RAS, 316, 3 - 21." type="journal article" year="2012">Gorochov 2012</bibRefCitation>
, there under subfamily group 
<taxonomicName id="4C004D5A0558A907FDF82B51FD4C215E" box="[554,744,1807,1832]" class="Insecta" family="Pseudophyllidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Pseudophyllidae</taxonomicName>
), and it was proposed to place it directly under 
<taxonomicName id="4C004D5A0558A907FADB2B51FE94213B" ID-CoL="8NKR2" authority="Braun 2015" authorityName="Braun" authorityYear="2015" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Tettigoniidae (Braun 2015)</taxonomicName>
. One indication that the 
<taxonomicName id="4C004D5A0558A907FD912B6AFD4F213B" ID-CoL="8PSS6" box="[579,747,1844,1869]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Pterochrozinae">Pterochrozinae</taxonomicName>
might be unrelated to 
<taxonomicName id="4C004D5A0558A907FC3B2B6AFB06213B" ID-CoL="8NL27" box="[1001,1186,1844,1869]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Pseudophyllinae">Pseudophyllinae</taxonomicName>
is the development of the non-functional stridulatory file on the right tegmen. It appears as an unsclerotized and inconspicuous vestigial copy of the functional one on the left tegmen, like in all other subfamilies investigated regarding this character, except for the true pseudophyllines, that typically have it completely reduced (
<bibRefCitation id="EF914B280558A907FBFE2BFEFA3521CE" author="Chamorro-Rengifo" box="[1068,1425,1952,1977]" pageId="28" pageNumber="29" refString="Chamorro-Rengifo, J., Braun, H. &amp; Lopes-Andrade, C. (2014) The secret stridulatory file under the right tegmen in katydids (Orthoptera, Ensifera, Tettigonioidea). Zootaxa, 3821 (5), 590 - 596. http: // dx. doi. org / 10.11646 / zootaxa. 3821.5.7" type="journal article" year="2014">
Chamorro-Rengifo 
<emphasis id="B974EACB0558A907FADF2BFFFAED21CF" box="[1293,1353,1952,1977]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2014
</bibRefCitation>
). However, another characteristic share 
<taxonomicName id="4C004D5A0558A907FD9A2B9AFD7C21AB" box="[584,728,1988,2013]" class="Insecta" family="Phaneropteridae" genus="Typophyllum" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B974EACB0558A907FD9A2B9AFD7C21AB" box="[584,728,1988,2013]" italics="true" pageId="28" pageNumber="29">Typophyllum</emphasis>
</taxonomicName>
species and 
<taxonomicName id="4C004D5A0558A907FCBB2B9AFB8321AB" ID-CoL="8NL27" box="[873,1063,1988,2013]" class="Insecta" family="Tettigoniidae" kingdom="Animalia" order="Orthoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Pseudophyllinae">Pseudophyllinae</taxonomicName>
: The functional file is located all along a fairly massive cubital vein that projects basally beyond the lower tegmen surface. Another indication for an early phylogenetic origin is the pure-tone resonant stridulation, which is thought to represent an ancestral character state in katydids (Montealegre-Z. &amp; Morris 2004, Montealegre-Z. 2009). Future studies on katydid phylogeny will hopefully substantiate how long ago these fascinating insects did branch off.
</paragraph>
</subSubSection>
</treatment>
</document>