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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.212.2705" ID-GBIF-Dataset="9729e71b-ef08-4369-bfae-4ea93fc0bc49" ID-PMC="PMC3428703" ID-Pensoft-Pub="1313-2970-212-45" ID-PubMed="22933849" ModsDocAuthor="" ModsDocDate="2012" ModsDocID="1313-2970-212-45" ModsDocOrigin="ZooKeys 212" ModsDocTitle="On immature and adult forms of Trichognathus marginipennis Latreille, 1829 (Coleoptera, Carabidae, Galeritini)" checkinTime="1451248828114" checkinUser="pensoft" docAuthor="Santos, Guilherme Ide Marques dos" docDate="2012" docId="18C2ADE801498E70F89DB8BE56FE6F86" docLanguage="en" docName="ZooKeys 212: 45-62" docOrigin="ZooKeys 212" docSource="http://dx.doi.org/10.3897/zookeys.212.2705" docTitle="Trichognathus marginipennis Latreille 1829" docType="treatment" docVersion="3" lastPageNumber="58" masterDocId="FFF0155DFC18FF8CFF8EFB5B8534FF9C" masterDocTitle="On immature and adult forms of Trichognathus marginipennis Latreille, 1829 (Coleoptera, Carabidae, Galeritini)" masterLastPageNumber="62" masterPageNumber="45" pageNumber="47" updateTime="1668154301997" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>On immature and adult forms of Trichognathus marginipennis Latreille, 1829 (Coleoptera, Carabidae, Galeritini)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Santos, Guilherme Ide Marques dos</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2012</mods:date>
<mods:detail type="volume">
<mods:number>212</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>45</mods:start>
<mods:end>62</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.212.2705</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.212.2705</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-212-45</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="152036498" LSID="urn:lsid:plazi:treatment:18C2ADE801498E70F89DB8BE56FE6F86" httpUri="http://treatment.plazi.org/id/18C2ADE801498E70F89DB8BE56FE6F86" lastPageId="13" lastPageNumber="58" pageId="2" pageNumber="47">
<subSubSection pageId="2" pageNumber="47" type="nomenclature">
<paragraph pageId="2" pageNumber="47">
<taxonomicName LSID="http://species-id.net/wiki/Trichognathus_marginipennis" authority="Latreille, 1829" authorityName="Latreille" authorityYear="1829" class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="2" pageNumber="47" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis Latreille, 1829</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="48" pageId="2" pageNumber="47" type="pupa">
<paragraph pageId="2" pageNumber="47">Pupa.</paragraph>
<paragraph pageId="2" pageNumber="47">
(Figs 1
<normalizedToken originalValue="bd">b-d</normalizedToken>
, 7
<normalizedToken originalValue="eg">e-g</normalizedToken>
) Body length: 11.0-12.0 mm.
</paragraph>
<paragraph lastPageId="3" lastPageNumber="48" pageId="2" pageNumber="47">
Adecticous, exarate. Yellowish white with legs long and thin, antenna with two setae at base of scape. Antenna long, longer than half body length, folded posteriorly and fitted at inferior margin of eye, directed through ventral surface of legs. Eyes prominent, each about
<normalizedToken originalValue="¼">1/4</normalizedToken>
width of head. Labrum with rounded anterior margin. Mandible relatively long and markedly sclerotized at apex. Pronotum subtrapezoidal, anterior margin prominent and rounded at middle; many widely spaced setae, these more concentrated near margins. Mesonotum wider than pronotum with setae at middle, these more concentrated near base. Metanotum longer than mesonotum; higher concentration of setae on median region; metathoracic legs longer than abdomen. Abdomen with eight segments visible in dorsal view. Five pairs of lateral projections, wide at base, extended over a narrow stalk, widened slightly at apex, with a pair of short setae. The first lateral projection at junction of first and second visible abdominal segments,
<pageBreakToken pageId="3" pageNumber="48" start="start">second</pageBreakToken>
from third segment, third from fourth and so on until sixth segment. Terga one to six dorsally with a marked concentration of long setae, these formed in two groups close to the midline. Segments seven and eight with many fewer setae. Spiracles visible in segments one to six, above the base of the lateral projections.
</paragraph>
</subSubSection>
<subSubSection lastPageId="12" lastPageNumber="57" pageId="3" pageNumber="48" type="adults">
<paragraph pageId="3" pageNumber="48">Adults</paragraph>
<paragraph pageId="3" pageNumber="48">
(Figs 2
<normalizedToken originalValue="af">a-f</normalizedToken>
, 3
<normalizedToken originalValue="aj">a-j</normalizedToken>
, 4
<normalizedToken originalValue="ae">a-e</normalizedToken>
, 5
<normalizedToken originalValue="ah">a-h</normalizedToken>
, 6
<normalizedToken originalValue="ad">a-d</normalizedToken>
, 7
<normalizedToken originalValue="bd">b-d</normalizedToken>
, 8a, 8f, 9a). Body length: 16-18.5mm.
</paragraph>
<paragraph pageId="3" pageNumber="48">Body form: dorso-ventrally depressed.</paragraph>
<paragraph pageId="4" pageNumber="49">
<pageBreakToken pageId="4" pageNumber="49" start="start">Color</pageBreakToken>
: head and pronotum from yellowish-brown to dark-brown; elytra dark-greenish with lateral margins and apex yellowish (Fig. 7b), legs from light to dark-brown, femora the same color as pronotum, some specimens with distal third darker;
</paragraph>
<paragraph pageId="5" pageNumber="50">
<pageBreakToken pageId="5" pageNumber="50" start="start">Vestiture</pageBreakToken>
: integument covered dorsally and ventrally with testaceous setae; those of antennae and especially the scape, noticeably longer and more robust than on the other segments.
</paragraph>
<paragraph pageId="5" pageNumber="50">
Head (Figs 2
<normalizedToken originalValue="cd">c-d</normalizedToken>
, 7d): wider behind eyes; two pairs of long orbital setae, short scattered dorsal setae, sparser at middle ventrally, suborbital zone with an elevation near the inner margin bearing 5-10 thick setae, like those of scape and some sparser and shorter. Gula glabrous.
</paragraph>
<paragraph pageId="5" pageNumber="50">Antenna (Fig. 2a): long (approximately 2/3 of the body length) and covered with short setae; antennomere 1 very long, as long as antennomeres 3 and 4 together, robust, club-shaped bearing long ventral thick setae; antennomere 2 short widened at apex, with thick ventral setae, antennomeres 3-11 setiform.</paragraph>
<paragraph pageId="5" pageNumber="50">
Labrum (Figs 2
<normalizedToken originalValue="ef">e-f</normalizedToken>
): subrectangular, lateral margins rounded, lobed anteriorly; six long apical setae: one on each angle and four near middle. Epipharynx with two rounded rows of anterior parapedial setae converged medially, each row bearing approximately 13 setae; median region with thin setae near the thick ones.
</paragraph>
<paragraph pageId="5" pageNumber="50">
Mandibles asymmetrical (Figs 3
<normalizedToken originalValue="ad">a-d</normalizedToken>
): incisor region 3-4 times longer than molar region, terebral margin smooth, scrobe glabrous, anterior retinacular tooth prolonged ventrally; ventral groove (fig. 8f) crosses entire mandible with many ventral microtrichia inserted in each puncture.
</paragraph>
<paragraph pageId="5" pageNumber="50">
Maxilla (Figs 3
<normalizedToken originalValue="ef">e-f</normalizedToken>
): basistipes with lobular projection and about ten spine-shaped setae and a few thin setae; lacinia with tooth-shaped apex with setae concentrated on inner margin and a row of thicker setae, basal pubescence present; galea two segmented, basigalea longer than distigalea with some setae; palpifer with three thick spine-shaped setae; palpomere 1 with one seta and many punctures near the base, palpomeres 2 and 3 with fine pubescence and ventral spine-like setae; palpomere 4 securiform and covered with fine pubescence.
</paragraph>
<paragraph pageId="5" pageNumber="50">
Labium (Figs 3
<normalizedToken originalValue="gi">g-i</normalizedToken>
): submentum-mentum suture present; mentum transverse, emarginate, its lateral lobes with acute apices and rounded tooth between them, palpiger glabrous, longer than palpomere 1, palpomere 2 longer than 3, covered with setae and some long ventral ones; palpomere 3 securiform and covered with fine pubescence; prementum with each paraglossa narrow, separated from glossal sclerite, latter unciform, upwardly directed (Fig. 3g) with two long setae ventrally, on proximal third.
</paragraph>
<paragraph pageId="5" pageNumber="50">
Prothorax (Figs 2
<normalizedToken originalValue="ab">a-b</normalizedToken>
, 7d): subtrapezoidal, height in lateral view similar to width in dorsal view (pronotum slightly convex and prosternum markedly convex), anterior margin slightly larger than posterior one, anterior angles more widely rounded than posterior ones; pronotum covered with fine setae, one or two longer setae on the anterior third of the margins and another one next to posterior angles. Proepisternum with short setae near anterior margin. Prosternum covered with many thick setae and a few thin ones, high concentration of thick setae near the midline. Proepimeron dark with few sparse thin setae.
</paragraph>
<paragraph lastPageId="6" lastPageNumber="51" pageId="5" pageNumber="50">
Mesothorax (Fig. 2b): with mesosternum darker laterally, covered with setae; mesepisternum subrectangular, dark, with margin adjacent to mesocoxae wider than its opposite and covered with thin setae; mesepimerum dark and usually glabrous
<pageBreakToken pageId="6" pageNumber="51" start="start">.</pageBreakToken>
</paragraph>
<paragraph pageId="6" pageNumber="51">Metathorax (Fig. 2b): metasternum wide, darker laterally, pubescence more concentrated laterally and near posterior margin; metepimeron elongate, dark and glabrous with blunt and wide proximal portion that partially covers ventrite I, middle region more tapered and wide inner margin.</paragraph>
<paragraph lastPageId="9" lastPageNumber="54" pageId="7" pageNumber="52">
<pageBreakToken pageId="7" pageNumber="52" start="start">Legs</pageBreakToken>
(Figs 4
<normalizedToken originalValue="ad">a-d</normalizedToken>
; 5
<normalizedToken originalValue="ab">a-b</normalizedToken>
; 9b): fore and median coxae globose covered with thin setae, hind coxa with only few setae at middle and one or two long setae near anterior margin; median coxa with one or two long setae near the posterior margin; trochanter subtriangular, fore trochanter with a few sparse short setae on entire surface and thick and long ones
<pageBreakToken pageId="8" pageNumber="53" start="start">on</pageBreakToken>
ventral margin; median trochanter with one or two long setae ventrally; femur long and covered ventrally with short setae, fore femur broader than others, flattened laterally with outer surface with a band of thicker and longer setae midline, inner surface with more concentration of setae near base, ventral margin with very long and thick spine-like bristles; hind femur longer than others; tibial length subequal to femur, tibia covered with thin setae and two dark spurs at the apex, except the fore tibia with widely separated spurs, only one distally; antenna cleaner on the distal third with a strong dark spur curved
<pageBreakToken pageId="9" pageNumber="54" start="start">at</pageBreakToken>
base and a row of thick setae on the inner surface extended from base to apex; median and hind tibiae with more concentrated pubescence ventrally, especially on posterior half and two dark spurs at apex; tarsomeres 1-4 covered with setae, denser ventrally; fore tarsomeres 1-3 wider in males with rows of adhesive setae with a wide and rough plate (Fig. 9b); tarsomere 5 elongate with only few setae; tarsal claws simple and glabrous.
</paragraph>
<paragraph pageId="10" pageNumber="55">
<pageBreakToken pageId="10" pageNumber="55" start="start">Elytra</pageBreakToken>
(Figs 2a, 4e): stria 1 very short, ranging from level of scutellum to approximately anterior fifth terminated in thick punctuation; fine setae covering entire structure with a few longer bristles in more defined punctations, especially near stria 1; posterior third near external margin and in the apex; striae 4 and 5 very close to one another at the apex, as are striae 6 and 7.
</paragraph>
<paragraph lastPageId="11" lastPageNumber="56" pageId="10" pageNumber="55">
Hind wing (Fig. 3j): fully developed, about 2.5 times longer than wide, rounded anal lobe and tapered apex; anal vein bifurcated near base, proximal branch reaching margin and other branch (AA1+2) fused with cubitus anterior (CuA) at level of poste
<pageBreakToken pageId="11" pageNumber="56" start="start">rior</pageBreakToken>
third, forming a cell; a small branch extended from cell formed to cr branch, but not connected; oblong cell present.
</paragraph>
<paragraph pageId="11" pageNumber="56">Male genital segments (Fig. 5c): tergite IX formed by antecostal region, a tergite and two laterotergites; antecostal region arc shaped, mediotergite IX shorter and less curved; laterotergites IX form a comma-shaped structure.</paragraph>
<paragraph pageId="11" pageNumber="56">
Aedeagus (Figs 5
<normalizedToken originalValue="dh">d-h</normalizedToken>
): median lobe cylindrical, slightly curved downward; phallobase rounded and distal region tapered with very short tip, dorsal surface slightly sclerotized and wrinkled with many punctures except anterior fifth and at the base; left paramere (Fig. 5d) broad, concave, base deflected and left basal projection going over the base; right paramere smaller and markedly attached at base of median lobe.
</paragraph>
<paragraph pageId="11" pageNumber="56">Female genital segments: sternite VIII (Figs 6a, 9a) 2.45 times wider than long, sclerotized, more weakly at medio-basal region; two basal projections (apodemes) almost half size of ventrite length, with rounded apex; fine setae posterior to spiracles and bordering posterior margin; two transverse lateromedian sclerotized stripes from base of projections to distal third of ventrite. Tergum 8 (Fig. 6b) formed by two laterotergites united at middle by a thin membrane, each laterotergite with a wide projection at the anterior margin, slightly bilobed at apex, and a lateral sclerotized band similar to that of sternite VIII, short thin setae at lateroexternal margin and distal margin with many long spine-like setae and a few thinner ones.</paragraph>
<paragraph lastPageId="12" lastPageNumber="57" pageId="11" pageNumber="56">
Female genitalia (Figs 6
<normalizedToken originalValue="cd">c-d</normalizedToken>
, 8a). Ovipositor with gonocoxites 1 subtriangular, falciform; gonocoxites 2 with apex turned out and some setae near inner margin, la
<pageBreakToken pageId="12" pageNumber="57" start="start">terotergite</pageBreakToken>
IX folded upon itself in internal margin, thus half-moon shaped. Internal organs comprised of a wrinkled bursa coiled at base; secondary spermathecal gland globose; spermathecal gland fusiform, attached by long duct, spermatheca 2 of similar size to pygidial glands, but more rounded.
</paragraph>
</subSubSection>
<subSubSection lastPageId="13" lastPageNumber="58" pageId="12" pageNumber="57" type="examined material">
<paragraph pageId="12" pageNumber="57">Examined material.</paragraph>
<paragraph pageId="12" pageNumber="57">
BRAZIL, Amazonas: Rio
<normalizedToken originalValue="Juruá">Jurua</normalizedToken>
, 1902, E. Garbe col., Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♀, 2♂. Mato Grosso: Barra do
<normalizedToken originalValue="Garças">Garcas</normalizedToken>
, IX.1943, Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 2♀; Campo Novo do Parecis, 25.VII - 04. VIII, S. Rosa, S. Casari, G. Ide &amp; L. Prado cols., 4♀, 2♂;
<normalizedToken originalValue="Corumbá">Corumba</normalizedToken>
, XI.1917, E. Garbe col., Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♀, 1♂; Diauarum, 02.XI.-11.XII.1973, G. R. Kloss col., 4♀, 5♂;
<normalizedToken originalValue="Três">Tres</normalizedToken>
Lagoas, Fazenda Floresta, 13-20.IX.1964, Exp. Depto. Zool., 6♀, 1♂; same locality, Fazenda Retiro de Telhas, 16.X.1964, Exp. Depto. Zool. col., 1♀; same locality, left margin Sucuriu river, Fazenda
<normalizedToken originalValue="Canaã">Canaa</normalizedToken>
, XI.1966, F. Lane col., 2♂; same locality, X.1966, F. Lane col., 13♀, 7♂; same locality, IV.1967, F. Lane col., 1♀, 1♂; same locality, I.1967, F. Lane col., 8♀, 5♂; same locality, VI.1967, F. Lane col., 2♀, 3♂. Minas Gerais: Paracatu, VII. 1960, Exp. Formoso col., 1♀.
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo: Boa
<normalizedToken originalValue="Esperança">Esperanca</normalizedToken>
do Sul, Fazenda
<normalizedToken originalValue="Itaquerê">Itaquere</normalizedToken>
, left margin of
<normalizedToken originalValue="Jacaré-Guaçu">Jacare-Guacu</normalizedToken>
river, under rotten plank, very humid place, 20.VI.1965, K. Lenko col., 2♂; Funil, 28.V.1902, O. Dreher col., Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 2♀, 2♂; Franca, XI.1902, O. Dreher col., Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♀; Indiana, IV.1944, Dirings, 1♀; same locality, 02.III.1935, 1♀, 1♂; same locality, II.1935, Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♂; same locality, II.1934, Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♂; same locality, XI.1935, 2♀; Onda Verde, Fazenda
<normalizedToken originalValue="São">Sao</normalizedToken>
<normalizedToken originalValue="João">Joao</normalizedToken>
, I.1946, F. Lane col., 1♀, 2♂; Pirassununga, Emas, without date, Schubart col., 1♀;
<normalizedToken originalValue="São">Sao</normalizedToken>
Roque, 1961, H. Lane col., Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 2♀; without locality and date, 1♀;
<normalizedToken originalValue="Ribeirão">Ribeirao</normalizedToken>
Preto,
<normalizedToken originalValue="Tamanduá">Tamandua</normalizedToken>
, 15.XI.1954, Barreto col., 1♂; same locality, 1896, Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♀; same locality, X.1954, Barreto col., 1♀.
<normalizedToken originalValue="Paraná">Parana</normalizedToken>
: Curitiba, 1938, Departamento de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♂; Porto
<normalizedToken originalValue="Tibiriçá">Tibirica</normalizedToken>
, Rio
<normalizedToken originalValue="Paraná">Parana</normalizedToken>
, IX.1938, 3♂; same locality, IX.1940, 1♀. Santa Catarina: Anita Garibaldi, VII.1929, Dirings col., 3♀, 1♂; same locality, VI.1929, Dirings, 4♂; Nova
<normalizedToken originalValue="Teutônia">Teutonia</normalizedToken>
(currently Seara), II.1966, F. Plaumann col., 2♀, 1♂; Rio Grande do Sul: Cruz Alta, without date, E. Garbe col., Departamnto de Zoologia
<normalizedToken originalValue="São">Sao</normalizedToken>
Paulo, 1♂; Porto Alegre, II.1929, Dirings, 1♂. PERU - Inca: Chanchamayo, 1200m, XII.1948, Schunke col., 1♀. ARGENTINA, Salta: San Martin, Pocitos, I.1960, A. Martinez leg., 5♀, 1♂; without locality, III. 1958, Martinez Pereira col., 3♀, 4♂. Misiones: Leandro N. Alem, XII.1961, A. Martinez leg., 1♀; Puerto Esperanza, II.1961, A. Martinez leg., 1♀, 1♂; Dos de Mayo, 350m, X.1971, 2♀, 3♂. BOLIVIA, Santa Cruz: Chapare, 400m, Dirings, 1♀, 1♂; Parapeti, X.1960, A. Martinez leg., 1♀; Provincia de Ichilo, Buenavista, II.1950, A. Martinez leg., 1♀, 3♂; Santa Cruz de La Sierra, 08.III.1954, Bruira col., 1♀. PARAGUAY, Presidente Hayes: Puerto Elsa [Puerto Elisa], XI.1936, 1♂; without locality, XII.1950, Dirings, 1♀, 1♂. All material is from MZSP.
</paragraph>
<subSection pageId="12" pageNumber="57" type="key for the immatures of galeritini with known pupa">
<paragraph pageId="12" pageNumber="57">
Key for the immatures of
<taxonomicName lsidName="" pageId="12" pageNumber="57" rank="tribe" tribe="Galeritini">Galeritini</taxonomicName>
with known pupa
</paragraph>
<paragraph pageId="12" pageNumber="57">
<table pageId="12" pageNumber="57">
<tr pageId="12" pageNumber="57">
<td colspan="1" pageId="12" pageNumber="57" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Galerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galerita carbonaria" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="carbonaria">Galerita carbonaria</taxonomicName>
</td>
</tr>
<tr pageId="12" pageNumber="57">
<td colspan="1" pageId="12" pageNumber="57" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
</td>
</tr>
</table>
</paragraph>
</subSection>
<subSection pageId="12" pageNumber="57" type="relationships">
<paragraph pageId="12" pageNumber="57">Relationships</paragraph>
<paragraph pageId="12" pageNumber="57">
In his monograph, Reichardt (1967) included a pre-cladistic reconstructed phylogeny in which
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Trichognathus</taxonomicName>
was proposed as the sister group of the Old World Tropical
<taxonomicName class="Insecta" family="Carabidae" genus="Eunostus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eunostus" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Eunostus</taxonomicName>
Castelnau, 1835. According to Ball (1985) and Hunting (2008), however,
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Trichognathus</taxonomicName>
is postulated to be sister group of
<taxonomicName class="Insecta" family="Carabidae" genus="Galerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galerita sensu subsp. lato" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="subSpecies" species="sensu" subSpecies="lato">Galerita sensu lato</taxonomicName>
(
<taxonomicName class="Insecta" family="Carabidae" genus="Galerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galerita" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Galerita</taxonomicName>
sensu stricto +
<taxonomicName class="Insecta" family="Carabidae" genus="Progaleritina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Progaleritina" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Progaleritina</taxonomicName>
), together forming the sister group of
<taxonomicName class="Insecta" family="Carabidae" genus="Ancystroglossus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ancystroglossus" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Ancystroglossus</taxonomicName>
Chaudoir, 1862. Detailed studies on immature characters may contribute to the resolution of phylogenetic relationships within the group.
</paragraph>
</subSection>
<subSection pageId="12" pageNumber="57" type="natural history notes">
<paragraph pageId="12" pageNumber="57">Natural history notes</paragraph>
<paragraph pageId="12" pageNumber="57">
Adults and larvae of
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Trichognathus</taxonomicName>
are fast running predators and go in search of their prey at night usually on sandy beaches along rivers and small streams in grassy areas, as well as in gallery forests (Arndt and Drechsel 1998). Adults and larvae are seen hunting together. No one knows for sure what kind of prey they eat, but based on detailed study of mouthparts, Ball (1985) postulated that the very specialized mouthparts play an important role in the capture and manipulation of food.
</paragraph>
<paragraph pageId="12" pageNumber="57">
Larvae and adults were found together on sandy substrate along a river margin, where they were active from sunset until about two hours later; after this period I found no more carabid larvae and adults. Sharing the same environment, some carabid adults of the genus
<taxonomicName class="Insecta" family="Carabidae" genus="Tetracha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tetracha" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Tetracha</taxonomicName>
Westwood (Fig. 7a) were observed hunting. Possibly the same fact was observed by Arndt and Drechsel (1998) that indicated a frequent association with individuals of the carabid
<taxonomicName class="Insecta" family="Carabidae" genus="Megacephala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Megacephala" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Megacephala</taxonomicName>
Latreille, but now the genus
<taxonomicName class="Insecta" family="Carabidae" genus="Megacephala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Megacephala" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Megacephala</taxonomicName>
is divided into seven genera and
<taxonomicName class="Insecta" family="Carabidae" genus="Tetracha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tetracha" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Tetracha</taxonomicName>
includes only Neotropical species. The group currently accepted as
<taxonomicName class="Insecta" family="Carabidae" genus="Megacephala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Megacephala" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="genus">Megacephala</taxonomicName>
occurs only in sub-Saharan region (Naviaux 2007).
</paragraph>
<paragraph pageId="12" pageNumber="57">
Gregarious behavior was observed (Fig. 7b), as well as cannibalism among adults of
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
in the laboratory. During the pre-pupal stage (about two days before the pupal stage), the larva, usually agile, stayed motionless, writhing only when prodded; the pupal period lasted about one week. After death, the darkening of the head and pronotum of adults was also noticed.
</paragraph>
</subSection>
<subSection pageId="13" pageNumber="58" type="comparisons">
<paragraph pageId="13" pageNumber="58">
<pageBreakToken pageId="13" pageNumber="58" start="start">Comparisons</pageBreakToken>
</paragraph>
<paragraph pageId="13" pageNumber="58">
The description of the third instar larva of
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
made by Arndt and Drechsel (1998) is quite complete and a full redescription is not necessary. However, unlike the original description of the larva, I observed that the abdominal sternites are distinct (Fig. 8b) and not fused as previously described. Furthermore, four bristles were found on the right side of the ventral surface of the head (Figure 8e) in one specimen; usually, three are present on each side.
</paragraph>
<paragraph pageId="13" pageNumber="58">
Comparing the pupae of
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
with those of
<taxonomicName class="Insecta" family="Carabidae" genus="Galerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galerita carbonaria" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="carbonaria">Galerita carbonaria</taxonomicName>
, showed that they are quite similar, with the following differences: each lateral abdominal extension has a pair of setae at the apex and all thoracic segments with some short setae in
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
, whereas in
<taxonomicName class="Insecta" family="Carabidae" genus="Galerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galerita carbonaria" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="carbonaria">Galerita carbonaria</taxonomicName>
there are no such setae. Moreover, the eyes of the pupa of
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
are apparently a bit larger.
</paragraph>
<paragraph pageId="13" pageNumber="58">As noted by Reichardt (1964), there are no consistent geographical patterns among the described morphs noted above. The genitalia of some females were dissected and the fairly long spermathecal gland, considered a remarkable character by Hunting (2008), was not observed, appearing only slightly elongated (Figure 6c). The sexual dimorphism of dilated tarsi was observed more pronounced in some males than in others.</paragraph>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 1.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
: a habitus of third instar larva in dorsal view; habitus of pupa b dorsal view c ventral view d lateral view.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 2.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
(adult male): a habitus dorsal b thorax and abdomen in ventral view c head in dorsal view d head in front view; e. labrum, dorsal view f epipharynx, ventral view.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 3.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
, male: a left mandible in dorsal view b right mandible in dorsal view c right mandible in ventral view d left mandible in ventral view; maxilla e dorsal view; f. ventral view g glossa in lateral view h labium in ventral view i labial palp in dorsal view j hind wing - AP: anal posterior AA anal anterior CuA cubitus anterior MP media posterior.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 4.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
, male. Fore leg a external view b internal view; median leg c internal view d external view e elytra apex.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 5.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
, male. Hind leg: a external view b internal view c male genital segments (median lobe in dash line) d left paramere in lateral view (d1 and d2 lateral views d3 ventral view d4 dorsal view) e median lobe in side view; aedeagus f lateral view g ventral view h ventro-lateral view.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 6.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
, female a sternite VIII b tergites VIII (laterotergites IX, gonocoxites 1 and 2 in dash line) c female genitalia and pygidial glands d gonocoxites 2.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 7. a
<taxonomicName class="Insecta" family="Carabidae" genus="Tetracha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tetracha" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="genus">Tetracha</taxonomicName>
sp. b,c,d,e
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
a
<taxonomicName class="Insecta" family="Carabidae" genus="Tetracha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tetracha" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="genus">Tetracha</taxonomicName>
sp. collected with
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
b dorsal view of adults c living larva in dorsal view d head and prothorax of the adult in lateral view; pupa e dorsal view f lateral view g ventral view.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 8.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
: a gonocoxites from the outside b abdominal segments of the larva in ventral view c head of the larva in dorsal view d larva in dorsal view e head of the larva in ventral view f detail of the mandible filtering apparatus.
</paragraph>
</caption>
<caption pageId="13" pageNumber="58">
<paragraph pageId="13" pageNumber="58">
Figure 9.
<taxonomicName class="Insecta" family="Carabidae" genus="Trichognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trichognathus marginipennis" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="marginipennis">Trichognathus marginipennis</taxonomicName>
(adult): a sternite VIII with pygidial glands b male fore tarsomeres 1-4 in ventral view c Rio Verde view from Fazenda Soledad.
</paragraph>
</caption>
</subSection>
</subSubSection>
</treatment>
</document>