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New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae) Author(s): MICHAEL S. ENGEL and KUMAR KRISHNA Source: American Museum Novitates, Number 3592:1-8. 2007. Published By: American Museum of Natural History DOI: http://dx.doi.org/10.1206/0003-0082(2007)3592[1:NDFEAI]2.0.CO;2 URL: http://www.bioone.org/doi/ full/10.1206/0003-0082%282007%293592%5B1%3ANDFEAI%5D2.0.CO %3B2 BioOne (www.bioone.org ) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non- commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
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Page 1: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors nonprofitpublishers academic institutions research libraries and research funders in the common goal of maximizing access tocritical research

New Dolichorhinotermes from Ecuador and in Mexican Amber(Isoptera Rhinotermitidae)Author(s) MICHAEL S ENGEL and KUMAR KRISHNASource American Museum Novitates Number 35921-8 2007Published By American Museum of Natural HistoryDOI httpdxdoiorg1012060003-0082(2007)3592[1NDFEAI]20CO2URL httpwwwbiooneorgdoifull1012060003-00822820072935925B13ANDFEAI5D20CO3B2

BioOne (wwwbiooneorg) is a nonprofit online aggregation of core research in thebiological ecological and environmental sciences BioOne provides a sustainable onlineplatform for over 170 journals and books published by nonprofit societies associationsmuseums institutions and presses

Your use of this PDF the BioOne Web site and all posted and associated contentindicates your acceptance of BioOnersquos Terms of Use available at wwwbiooneorgpageterms_of_use

Usage of BioOne content is strictly limited to personal educational and non-commercial use Commercial inquiries or rights and permissions requests should bedirected to the individual publisher as copyright holder

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY

CENTRAL PARK WEST A T 79TH STREET NEW YOR K NY 10024

Number 3592 8 pp 4 figures 1 table December 12 2007

New Dolichorhinotermes from Ecuador and inMexican Amber (Isoptera Rhinotermitidae)

MICHAEL S ENGEL1 AND KUMAR KRISHNA2

ABSTRACT

Two new species of the Neotropical termite genus Dolichorhinotermes Snyder and Emerson(Rhinotermitidae Rhinotermitinae) are described and figured Dolichorhinotermes lanciarius Engeland Krishna new species from southeastern Ecuador is the largest species of the genus and isdistinctive in both the major and minor soldier caste Dolichorhinotermes apopnus Engel andKrishna new species preserved in earliest Miocene amber from Chiapas Mexico is similar to Ddominicanus Schlemmermeyer and Cancello in Early Miocene (Burdigalian) amber from theDominican Republic but differs in several significant respects Distinctions between Rhinotermesand Dolichorhinotermes are briefly discussed

RESUMEN

Se describen e ilustran dos especies nuevas pertecientes al genero Dolichorhinotermes Snyder yEmerson (Rhinotermitidae Rhinotermitinae) Dolichorhinotermes lanciarius Engel y Krishnaespecie nueva proveniente del sureste de Ecuador es la especie de mayor tamano del genero y sedistingue tanto en los soldados mayores como en los menores Dolichorhinotermes apopnus Engel yKrishna especie nueva preservada en ambar de la base del Mioceno basal de Chiapas Mexico esmas semejante a D dominicanus Schlemmermeyer y Cancello encontrada en ambar del Miocenobasal (Burdigaliano) de la Republica Dominicana Se presentan algunos comentarios sobre lasdiferencias para distinguir entre Rhinotermes y Dolichorhinotermes

Copyright E American Museum of Natural History 2007 ISSN 0003-0082

1 Division of Invertebrate Zoology American Museum of Natural History and Division of Entomology(Paleoentomology) Natural History Museum and Department of Ecology amp Evolutionary Biology 1501 CrestlineDrive ndash Suite 140 University of Kansas Lawrence Kansas 66049-2811 (msengelkuedu)

2 Division of Invertebrate Zoology American Museum of Natural History (krishnamnhorg)

INTRODUCTION

Termites of the genus DolichorhinotermesSnyder and Emerson are distinctive members ofthe Neotropical rhinotermitid fauna Six modernspecies have hitherto been described from vari-ous localities throughout tropical America rang-ing from Bolivia Peru and Brazil northward toSurinam Guyana French Guiana Trinidadand Panama and Martinique and Haiti in theWest Indies (eg Emerson 1925 Snyder 19241926 1949 Constantino 1991 1998 Ensaf andBetsch 2002 Ensaf et al 2004) In additiona single fossil has been described in Early Mio-cene amberfromtheDominicanRepublic(Schlem-mermeyer and Cancello 2000) The genus hashad a checkered past in that some authors havefelt the distinctions between DolichorhinotermesRhinotermesHagen and Acorhinotermes Emersonwere subtle and intergraded (eg Mathews1977) However Quennedey and Deligne (1975)identified significant morphological charactersof the soldier head capsule particularly featuresof the labrum that justify the recognition ofseparate albeit closely related genera

Herein we provide the description of a re-markable new species of Dolichorhinotermesfrom southeastern Ecuador representing thefirst record of the genus from that country Inaddition we newly record the genus Dolicho-rhinotermes forthesouthernMexicanpaleofaunaThe only other known fossil of the genus is Doli-chorhinotermes dominicanus Schlemmemeyerand Cancello in Early Miocene (Burdigalian)amber from the Dominican Republic TheMexican fossil is remarkably similar to D domi-nicanus but is specifically distinct Previously thetermite fauna of Mexican amber included sixspecies representing four families Mastotermeselectromexicus Krishna and Emerson (Masto-termitidae) Calcaritermes vetus Emerson andIncisitermes krishnai Emerson (Kalotermitidae)Coptotermes sucineusEmerson and Heterotermesprimaevus Snyder (Rhinotermitidae) and Nasu-titermes electrinus Krishna (Termitidae)

SYSTEMATICS

Genus Dolichorhinotermes Snyder and Emerson

Dolichorhinotermes Snyder and Emerson In Snyder1949 374 Type species Rhinotermes longilabiusEmerson 1925 by original designation

DIAGNOSIS Imago with third flagellar ar-ticle shorter than first flagellar article Majorsoldier with labrum distinctly elongate apexof labrum frequently extending to mandibularapex (in Rhinotermes the labrum is subquad-rate and much shorter) Minor soldier withopening of frontal gland at front of head butnot on conspicuous prolongation of headcapsule mandibles vestigial with roundedmargins sides of head in dorsal aspect straightor convex

Dolichorhinotermes lanciarius new species

figures 1ndash2

Dolichorhinotermes hageni Mathur and Thapa1962 29 Maiti 2006 12 Nomen nudum

DIAGNOSIS The new species is distinctlylarger than all previously described species inall comparable metrics particularly the sol-dier which is the largest of all known speciesNoteworthy features include the dentition ofthe major soldier mandibles (fig 1) and theshape of the major and minor soldier labra(figs 1ndash2) Refer also to section on compar-isons (see below)

DESCRIPTION Imago UnknownMajor soldier Head thorax and abdomen

yellow legs and antennae pale yellow mandi-ble yellow except apical half reddish brownand teeth dark reddish brown Head withsparsely-scattered relatively short erect tosuberect pale yellow setae Labrum with a fewerect pale yellow setae at apex dorsal surfacelargely devoid of setae except for a fewsparsely scattered short erect pale yellowsetae Pronotum with scattered short erect tosuberect pale yellow setae remaining thoracicsclerites sparsely setose Legs with scatteredpale yellow setae setae minute and intermixedwith a few short to moderate-length setaeAbdomen sparsely setose Head broaderposteriorly with lateral margins gently round-ed and converging anteriorly posterior bordergently rounded Mandibles large left mandi-ble with two stout teeth first just apical ofmidpoint second just basal of midpoint firsttooth distinctly longer than second tooth firsttooth slightly curved along posterior surfacesecond tooth straight right mandible with twoteeth near midpoint first tooth shorter than

2 AMERICAN MUSEUM NOVITATES NO 3592

second tooth and both shorter than those ofleft mandible second tooth extending straightout from inner mandibular surface first toothrunning closer along mandibular surface(fig 1) Labrum elongate slightly wider atapex apex rounded without emarginationwith slight dorsal longitundinal depressionextending posteriorly from apical marginabout one-half of labral length Palpi andpostmentum as depicted in figure 1 Antennawith at least 14 articles (total number un-certain) third flagellar article distinctly short-er than first flagellar article pedicel andsecond through fourth flagellar articles rough-ly equal in length Pronotum saddle-shapedanterior margin broadly rounded mediallylateral margins converging posteriorly poste-

rior margin relatively straight Legs relativelyslender and long Measurements provided intable 1 for holotype and paratype

Minor soldier Head thorax and abdomenpale yellow sparsely setose setae erect tosuberect and pale yellow Head with lateralborders weakly and gently convex sidesconverging anteriorly and slightly convergingfrom midpoint posteriorly to a gently roundedposterior border Labrum with several erectpale yellow setae at apex otherwise dorsalsurface with scattered short pale yellow setaesetae slightly more numerous in apical halfthan basal half Antenna with 16 articlesthird flagellar article distinctly shorter thanthe first Mandibles vestigial representingby broad fleshy lobes bearing a minute

Fig 1 Dolichorhinotermes lanciarius Engel and Krishna new species major soldier A Dorsal aspect ofhead and pronotum B Ventral aspect of head Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 3

sclerotized point (fig 2) Labrum narrowelongate forked at apex lateral processesof fork relatively widely separated Rangeof measurements for paratypes provided intable 1

HOLOTYPE Major soldier Ecuador Gua-laquiza 6xi1934 [6 November 1934] coll Wv Hagen [Gualaquiza at 3u249S 78u339Wis a village in Morona-Santiago Provinceon the eastern slopes of the Andes about97 km northeast of Loja] The holotype ispreserved in the Division of InvertebrateZoology American Museum of NaturalHistory

PARATYPE One major soldier several mi-nor soldiers several workers same data asholotype Deposited in the Division ofInvertebrate Zoology American Museum ofNatural History

ETYMOLOGY The specific epithet is takenfrom the Latin for lsquolsquolancerrsquorsquo in reference to thelance-like labrum of soldiers in the genus

COMPARISONS Dolichorhinotermes lanciar-ius can be readily separated from all otherspecies in the genus by its larger bodyproportions (refer to table 1) The species isgenerally quite similar to D japuraensisConstantinto from Brazil but aside from size

Fig 2 Dolichorhinotermes lanciarius Engel and Krishna new species minor soldier A Dorsal aspect ofhead and pronotum B Lateral aspect of head Scale bar 5 05 mm

4 AMERICAN MUSEUM NOVITATES NO 3592

differs by the rounded labral apex in majorsoldiers (very weakly medioapically concave inD japuraensis) sparse setation (setae of headmore numerous and elongate in D japuraen-sis) more distinctly convex anterior margin ofthe pronotum (very weakly convex in Djapuraensis) right mandible with two well-developed teeth (one stout tooth medially andone minute tooth at basal third in Djapuraensis) and by the more pronouncedapical labral forking in the minor soldier(weakly concave at the apex of the labrum inD japuraensis)

Dolichorhinotermes apopnus new species

figures 3ndash4

DIAGNOSIS The Mexican amber species isremarkably similar to D dominicanus exceptthat D apopnus lacks the integumental wrin-kles lateral to the postclypeal groove (imbri-cate in the Mexican fossil) the general bodyproportions are distinctly smaller (head widthwith compound eyes in D dominicanus160 mm while it is 130 mm in the new fossil

the compound eye is 032 mm in D dominica-nus only 026 mm in D apopnus length ofmetatibia 16 mm in D dominicanus 125 mmin D apopnus) and the integument of the headis darker

DESCRIPTION Imago Length of head toapex of clypeus 102 mm width of head withcompound eyes 130 mm maximum diameterof compound eye with ocular sclerite026 mm separation of compound eye fromlower margin 003 mm length of ocellus013 mm width of ocellus 006 mm ocellocu-lar distance 003 mm median length ofpronotum 087 mm width of pronotum102 mm length of metatibia 125 mm lengthof forewing scale 077 mm length of hindwing scale 049 mm Head large broadlyrounded distinctly wider than long withupwardly lifted frontoclypeal region wherebypostclypeus forms wide projecting proboscis(figs 3ndash4) Postclypeus projected forward toform broad proboscis width of extensiongreater than compound eye diameter withwide dorsal mediolongitudinal grooveFontanelle small located at base of upliftedproboscis and at base of dorsal groove

TABLE 1Measurements of soldiers of Dolichorhinotermes lanciarius new species

(all measurements in millimeters)

Metric

Major soldier

Holotype

Major soldier

Paratype

Minor soldier

Paratypes

Number of specimens 1 1 3

Length of head to side base of mandibles 158 158 069ndash071

Length of head with mandibles 260 255 na

Length of head to apex of labrum 250 255 155ndash158

Maximum width of head 168 168 069

Maximum height of head (without

postmentum) 095 097 049ndash051

Length of left mandible 127 132 vestigial

Length of right mandible 122 127 vestigial

Length of labrum along midline 077 079 079ndash081

Width of labrum at apex 023 026 013ndash015

Width of labrum at base 020 023 008

Maximum width of postmentum 043 046 030

Minimum width of postmentum 033 038 026

Length of postmentum 077 077 036

No of antennal articles ndash 14+a 15andash16

Median length of pronotum 043 048 036

Maximum width of pronotum 082 084 051

Length of metatibia 135 137 094ndash096

aThe antenna is broken after the 14th article in the major soldier paratype and after the 15th in one of the minor soldier

paratypes and so the total number is unknown for these individuals

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 5

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 2: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY

CENTRAL PARK WEST A T 79TH STREET NEW YOR K NY 10024

Number 3592 8 pp 4 figures 1 table December 12 2007

New Dolichorhinotermes from Ecuador and inMexican Amber (Isoptera Rhinotermitidae)

MICHAEL S ENGEL1 AND KUMAR KRISHNA2

ABSTRACT

Two new species of the Neotropical termite genus Dolichorhinotermes Snyder and Emerson(Rhinotermitidae Rhinotermitinae) are described and figured Dolichorhinotermes lanciarius Engeland Krishna new species from southeastern Ecuador is the largest species of the genus and isdistinctive in both the major and minor soldier caste Dolichorhinotermes apopnus Engel andKrishna new species preserved in earliest Miocene amber from Chiapas Mexico is similar to Ddominicanus Schlemmermeyer and Cancello in Early Miocene (Burdigalian) amber from theDominican Republic but differs in several significant respects Distinctions between Rhinotermesand Dolichorhinotermes are briefly discussed

RESUMEN

Se describen e ilustran dos especies nuevas pertecientes al genero Dolichorhinotermes Snyder yEmerson (Rhinotermitidae Rhinotermitinae) Dolichorhinotermes lanciarius Engel y Krishnaespecie nueva proveniente del sureste de Ecuador es la especie de mayor tamano del genero y sedistingue tanto en los soldados mayores como en los menores Dolichorhinotermes apopnus Engel yKrishna especie nueva preservada en ambar de la base del Mioceno basal de Chiapas Mexico esmas semejante a D dominicanus Schlemmermeyer y Cancello encontrada en ambar del Miocenobasal (Burdigaliano) de la Republica Dominicana Se presentan algunos comentarios sobre lasdiferencias para distinguir entre Rhinotermes y Dolichorhinotermes

Copyright E American Museum of Natural History 2007 ISSN 0003-0082

1 Division of Invertebrate Zoology American Museum of Natural History and Division of Entomology(Paleoentomology) Natural History Museum and Department of Ecology amp Evolutionary Biology 1501 CrestlineDrive ndash Suite 140 University of Kansas Lawrence Kansas 66049-2811 (msengelkuedu)

2 Division of Invertebrate Zoology American Museum of Natural History (krishnamnhorg)

INTRODUCTION

Termites of the genus DolichorhinotermesSnyder and Emerson are distinctive members ofthe Neotropical rhinotermitid fauna Six modernspecies have hitherto been described from vari-ous localities throughout tropical America rang-ing from Bolivia Peru and Brazil northward toSurinam Guyana French Guiana Trinidadand Panama and Martinique and Haiti in theWest Indies (eg Emerson 1925 Snyder 19241926 1949 Constantino 1991 1998 Ensaf andBetsch 2002 Ensaf et al 2004) In additiona single fossil has been described in Early Mio-cene amberfromtheDominicanRepublic(Schlem-mermeyer and Cancello 2000) The genus hashad a checkered past in that some authors havefelt the distinctions between DolichorhinotermesRhinotermesHagen and Acorhinotermes Emersonwere subtle and intergraded (eg Mathews1977) However Quennedey and Deligne (1975)identified significant morphological charactersof the soldier head capsule particularly featuresof the labrum that justify the recognition ofseparate albeit closely related genera

Herein we provide the description of a re-markable new species of Dolichorhinotermesfrom southeastern Ecuador representing thefirst record of the genus from that country Inaddition we newly record the genus Dolicho-rhinotermes forthesouthernMexicanpaleofaunaThe only other known fossil of the genus is Doli-chorhinotermes dominicanus Schlemmemeyerand Cancello in Early Miocene (Burdigalian)amber from the Dominican Republic TheMexican fossil is remarkably similar to D domi-nicanus but is specifically distinct Previously thetermite fauna of Mexican amber included sixspecies representing four families Mastotermeselectromexicus Krishna and Emerson (Masto-termitidae) Calcaritermes vetus Emerson andIncisitermes krishnai Emerson (Kalotermitidae)Coptotermes sucineusEmerson and Heterotermesprimaevus Snyder (Rhinotermitidae) and Nasu-titermes electrinus Krishna (Termitidae)

SYSTEMATICS

Genus Dolichorhinotermes Snyder and Emerson

Dolichorhinotermes Snyder and Emerson In Snyder1949 374 Type species Rhinotermes longilabiusEmerson 1925 by original designation

DIAGNOSIS Imago with third flagellar ar-ticle shorter than first flagellar article Majorsoldier with labrum distinctly elongate apexof labrum frequently extending to mandibularapex (in Rhinotermes the labrum is subquad-rate and much shorter) Minor soldier withopening of frontal gland at front of head butnot on conspicuous prolongation of headcapsule mandibles vestigial with roundedmargins sides of head in dorsal aspect straightor convex

Dolichorhinotermes lanciarius new species

figures 1ndash2

Dolichorhinotermes hageni Mathur and Thapa1962 29 Maiti 2006 12 Nomen nudum

DIAGNOSIS The new species is distinctlylarger than all previously described species inall comparable metrics particularly the sol-dier which is the largest of all known speciesNoteworthy features include the dentition ofthe major soldier mandibles (fig 1) and theshape of the major and minor soldier labra(figs 1ndash2) Refer also to section on compar-isons (see below)

DESCRIPTION Imago UnknownMajor soldier Head thorax and abdomen

yellow legs and antennae pale yellow mandi-ble yellow except apical half reddish brownand teeth dark reddish brown Head withsparsely-scattered relatively short erect tosuberect pale yellow setae Labrum with a fewerect pale yellow setae at apex dorsal surfacelargely devoid of setae except for a fewsparsely scattered short erect pale yellowsetae Pronotum with scattered short erect tosuberect pale yellow setae remaining thoracicsclerites sparsely setose Legs with scatteredpale yellow setae setae minute and intermixedwith a few short to moderate-length setaeAbdomen sparsely setose Head broaderposteriorly with lateral margins gently round-ed and converging anteriorly posterior bordergently rounded Mandibles large left mandi-ble with two stout teeth first just apical ofmidpoint second just basal of midpoint firsttooth distinctly longer than second tooth firsttooth slightly curved along posterior surfacesecond tooth straight right mandible with twoteeth near midpoint first tooth shorter than

2 AMERICAN MUSEUM NOVITATES NO 3592

second tooth and both shorter than those ofleft mandible second tooth extending straightout from inner mandibular surface first toothrunning closer along mandibular surface(fig 1) Labrum elongate slightly wider atapex apex rounded without emarginationwith slight dorsal longitundinal depressionextending posteriorly from apical marginabout one-half of labral length Palpi andpostmentum as depicted in figure 1 Antennawith at least 14 articles (total number un-certain) third flagellar article distinctly short-er than first flagellar article pedicel andsecond through fourth flagellar articles rough-ly equal in length Pronotum saddle-shapedanterior margin broadly rounded mediallylateral margins converging posteriorly poste-

rior margin relatively straight Legs relativelyslender and long Measurements provided intable 1 for holotype and paratype

Minor soldier Head thorax and abdomenpale yellow sparsely setose setae erect tosuberect and pale yellow Head with lateralborders weakly and gently convex sidesconverging anteriorly and slightly convergingfrom midpoint posteriorly to a gently roundedposterior border Labrum with several erectpale yellow setae at apex otherwise dorsalsurface with scattered short pale yellow setaesetae slightly more numerous in apical halfthan basal half Antenna with 16 articlesthird flagellar article distinctly shorter thanthe first Mandibles vestigial representingby broad fleshy lobes bearing a minute

Fig 1 Dolichorhinotermes lanciarius Engel and Krishna new species major soldier A Dorsal aspect ofhead and pronotum B Ventral aspect of head Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 3

sclerotized point (fig 2) Labrum narrowelongate forked at apex lateral processesof fork relatively widely separated Rangeof measurements for paratypes provided intable 1

HOLOTYPE Major soldier Ecuador Gua-laquiza 6xi1934 [6 November 1934] coll Wv Hagen [Gualaquiza at 3u249S 78u339Wis a village in Morona-Santiago Provinceon the eastern slopes of the Andes about97 km northeast of Loja] The holotype ispreserved in the Division of InvertebrateZoology American Museum of NaturalHistory

PARATYPE One major soldier several mi-nor soldiers several workers same data asholotype Deposited in the Division ofInvertebrate Zoology American Museum ofNatural History

ETYMOLOGY The specific epithet is takenfrom the Latin for lsquolsquolancerrsquorsquo in reference to thelance-like labrum of soldiers in the genus

COMPARISONS Dolichorhinotermes lanciar-ius can be readily separated from all otherspecies in the genus by its larger bodyproportions (refer to table 1) The species isgenerally quite similar to D japuraensisConstantinto from Brazil but aside from size

Fig 2 Dolichorhinotermes lanciarius Engel and Krishna new species minor soldier A Dorsal aspect ofhead and pronotum B Lateral aspect of head Scale bar 5 05 mm

4 AMERICAN MUSEUM NOVITATES NO 3592

differs by the rounded labral apex in majorsoldiers (very weakly medioapically concave inD japuraensis) sparse setation (setae of headmore numerous and elongate in D japuraen-sis) more distinctly convex anterior margin ofthe pronotum (very weakly convex in Djapuraensis) right mandible with two well-developed teeth (one stout tooth medially andone minute tooth at basal third in Djapuraensis) and by the more pronouncedapical labral forking in the minor soldier(weakly concave at the apex of the labrum inD japuraensis)

Dolichorhinotermes apopnus new species

figures 3ndash4

DIAGNOSIS The Mexican amber species isremarkably similar to D dominicanus exceptthat D apopnus lacks the integumental wrin-kles lateral to the postclypeal groove (imbri-cate in the Mexican fossil) the general bodyproportions are distinctly smaller (head widthwith compound eyes in D dominicanus160 mm while it is 130 mm in the new fossil

the compound eye is 032 mm in D dominica-nus only 026 mm in D apopnus length ofmetatibia 16 mm in D dominicanus 125 mmin D apopnus) and the integument of the headis darker

DESCRIPTION Imago Length of head toapex of clypeus 102 mm width of head withcompound eyes 130 mm maximum diameterof compound eye with ocular sclerite026 mm separation of compound eye fromlower margin 003 mm length of ocellus013 mm width of ocellus 006 mm ocellocu-lar distance 003 mm median length ofpronotum 087 mm width of pronotum102 mm length of metatibia 125 mm lengthof forewing scale 077 mm length of hindwing scale 049 mm Head large broadlyrounded distinctly wider than long withupwardly lifted frontoclypeal region wherebypostclypeus forms wide projecting proboscis(figs 3ndash4) Postclypeus projected forward toform broad proboscis width of extensiongreater than compound eye diameter withwide dorsal mediolongitudinal grooveFontanelle small located at base of upliftedproboscis and at base of dorsal groove

TABLE 1Measurements of soldiers of Dolichorhinotermes lanciarius new species

(all measurements in millimeters)

Metric

Major soldier

Holotype

Major soldier

Paratype

Minor soldier

Paratypes

Number of specimens 1 1 3

Length of head to side base of mandibles 158 158 069ndash071

Length of head with mandibles 260 255 na

Length of head to apex of labrum 250 255 155ndash158

Maximum width of head 168 168 069

Maximum height of head (without

postmentum) 095 097 049ndash051

Length of left mandible 127 132 vestigial

Length of right mandible 122 127 vestigial

Length of labrum along midline 077 079 079ndash081

Width of labrum at apex 023 026 013ndash015

Width of labrum at base 020 023 008

Maximum width of postmentum 043 046 030

Minimum width of postmentum 033 038 026

Length of postmentum 077 077 036

No of antennal articles ndash 14+a 15andash16

Median length of pronotum 043 048 036

Maximum width of pronotum 082 084 051

Length of metatibia 135 137 094ndash096

aThe antenna is broken after the 14th article in the major soldier paratype and after the 15th in one of the minor soldier

paratypes and so the total number is unknown for these individuals

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 5

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 3: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

INTRODUCTION

Termites of the genus DolichorhinotermesSnyder and Emerson are distinctive members ofthe Neotropical rhinotermitid fauna Six modernspecies have hitherto been described from vari-ous localities throughout tropical America rang-ing from Bolivia Peru and Brazil northward toSurinam Guyana French Guiana Trinidadand Panama and Martinique and Haiti in theWest Indies (eg Emerson 1925 Snyder 19241926 1949 Constantino 1991 1998 Ensaf andBetsch 2002 Ensaf et al 2004) In additiona single fossil has been described in Early Mio-cene amberfromtheDominicanRepublic(Schlem-mermeyer and Cancello 2000) The genus hashad a checkered past in that some authors havefelt the distinctions between DolichorhinotermesRhinotermesHagen and Acorhinotermes Emersonwere subtle and intergraded (eg Mathews1977) However Quennedey and Deligne (1975)identified significant morphological charactersof the soldier head capsule particularly featuresof the labrum that justify the recognition ofseparate albeit closely related genera

Herein we provide the description of a re-markable new species of Dolichorhinotermesfrom southeastern Ecuador representing thefirst record of the genus from that country Inaddition we newly record the genus Dolicho-rhinotermes forthesouthernMexicanpaleofaunaThe only other known fossil of the genus is Doli-chorhinotermes dominicanus Schlemmemeyerand Cancello in Early Miocene (Burdigalian)amber from the Dominican Republic TheMexican fossil is remarkably similar to D domi-nicanus but is specifically distinct Previously thetermite fauna of Mexican amber included sixspecies representing four families Mastotermeselectromexicus Krishna and Emerson (Masto-termitidae) Calcaritermes vetus Emerson andIncisitermes krishnai Emerson (Kalotermitidae)Coptotermes sucineusEmerson and Heterotermesprimaevus Snyder (Rhinotermitidae) and Nasu-titermes electrinus Krishna (Termitidae)

SYSTEMATICS

Genus Dolichorhinotermes Snyder and Emerson

Dolichorhinotermes Snyder and Emerson In Snyder1949 374 Type species Rhinotermes longilabiusEmerson 1925 by original designation

DIAGNOSIS Imago with third flagellar ar-ticle shorter than first flagellar article Majorsoldier with labrum distinctly elongate apexof labrum frequently extending to mandibularapex (in Rhinotermes the labrum is subquad-rate and much shorter) Minor soldier withopening of frontal gland at front of head butnot on conspicuous prolongation of headcapsule mandibles vestigial with roundedmargins sides of head in dorsal aspect straightor convex

Dolichorhinotermes lanciarius new species

figures 1ndash2

Dolichorhinotermes hageni Mathur and Thapa1962 29 Maiti 2006 12 Nomen nudum

DIAGNOSIS The new species is distinctlylarger than all previously described species inall comparable metrics particularly the sol-dier which is the largest of all known speciesNoteworthy features include the dentition ofthe major soldier mandibles (fig 1) and theshape of the major and minor soldier labra(figs 1ndash2) Refer also to section on compar-isons (see below)

DESCRIPTION Imago UnknownMajor soldier Head thorax and abdomen

yellow legs and antennae pale yellow mandi-ble yellow except apical half reddish brownand teeth dark reddish brown Head withsparsely-scattered relatively short erect tosuberect pale yellow setae Labrum with a fewerect pale yellow setae at apex dorsal surfacelargely devoid of setae except for a fewsparsely scattered short erect pale yellowsetae Pronotum with scattered short erect tosuberect pale yellow setae remaining thoracicsclerites sparsely setose Legs with scatteredpale yellow setae setae minute and intermixedwith a few short to moderate-length setaeAbdomen sparsely setose Head broaderposteriorly with lateral margins gently round-ed and converging anteriorly posterior bordergently rounded Mandibles large left mandi-ble with two stout teeth first just apical ofmidpoint second just basal of midpoint firsttooth distinctly longer than second tooth firsttooth slightly curved along posterior surfacesecond tooth straight right mandible with twoteeth near midpoint first tooth shorter than

2 AMERICAN MUSEUM NOVITATES NO 3592

second tooth and both shorter than those ofleft mandible second tooth extending straightout from inner mandibular surface first toothrunning closer along mandibular surface(fig 1) Labrum elongate slightly wider atapex apex rounded without emarginationwith slight dorsal longitundinal depressionextending posteriorly from apical marginabout one-half of labral length Palpi andpostmentum as depicted in figure 1 Antennawith at least 14 articles (total number un-certain) third flagellar article distinctly short-er than first flagellar article pedicel andsecond through fourth flagellar articles rough-ly equal in length Pronotum saddle-shapedanterior margin broadly rounded mediallylateral margins converging posteriorly poste-

rior margin relatively straight Legs relativelyslender and long Measurements provided intable 1 for holotype and paratype

Minor soldier Head thorax and abdomenpale yellow sparsely setose setae erect tosuberect and pale yellow Head with lateralborders weakly and gently convex sidesconverging anteriorly and slightly convergingfrom midpoint posteriorly to a gently roundedposterior border Labrum with several erectpale yellow setae at apex otherwise dorsalsurface with scattered short pale yellow setaesetae slightly more numerous in apical halfthan basal half Antenna with 16 articlesthird flagellar article distinctly shorter thanthe first Mandibles vestigial representingby broad fleshy lobes bearing a minute

Fig 1 Dolichorhinotermes lanciarius Engel and Krishna new species major soldier A Dorsal aspect ofhead and pronotum B Ventral aspect of head Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 3

sclerotized point (fig 2) Labrum narrowelongate forked at apex lateral processesof fork relatively widely separated Rangeof measurements for paratypes provided intable 1

HOLOTYPE Major soldier Ecuador Gua-laquiza 6xi1934 [6 November 1934] coll Wv Hagen [Gualaquiza at 3u249S 78u339Wis a village in Morona-Santiago Provinceon the eastern slopes of the Andes about97 km northeast of Loja] The holotype ispreserved in the Division of InvertebrateZoology American Museum of NaturalHistory

PARATYPE One major soldier several mi-nor soldiers several workers same data asholotype Deposited in the Division ofInvertebrate Zoology American Museum ofNatural History

ETYMOLOGY The specific epithet is takenfrom the Latin for lsquolsquolancerrsquorsquo in reference to thelance-like labrum of soldiers in the genus

COMPARISONS Dolichorhinotermes lanciar-ius can be readily separated from all otherspecies in the genus by its larger bodyproportions (refer to table 1) The species isgenerally quite similar to D japuraensisConstantinto from Brazil but aside from size

Fig 2 Dolichorhinotermes lanciarius Engel and Krishna new species minor soldier A Dorsal aspect ofhead and pronotum B Lateral aspect of head Scale bar 5 05 mm

4 AMERICAN MUSEUM NOVITATES NO 3592

differs by the rounded labral apex in majorsoldiers (very weakly medioapically concave inD japuraensis) sparse setation (setae of headmore numerous and elongate in D japuraen-sis) more distinctly convex anterior margin ofthe pronotum (very weakly convex in Djapuraensis) right mandible with two well-developed teeth (one stout tooth medially andone minute tooth at basal third in Djapuraensis) and by the more pronouncedapical labral forking in the minor soldier(weakly concave at the apex of the labrum inD japuraensis)

Dolichorhinotermes apopnus new species

figures 3ndash4

DIAGNOSIS The Mexican amber species isremarkably similar to D dominicanus exceptthat D apopnus lacks the integumental wrin-kles lateral to the postclypeal groove (imbri-cate in the Mexican fossil) the general bodyproportions are distinctly smaller (head widthwith compound eyes in D dominicanus160 mm while it is 130 mm in the new fossil

the compound eye is 032 mm in D dominica-nus only 026 mm in D apopnus length ofmetatibia 16 mm in D dominicanus 125 mmin D apopnus) and the integument of the headis darker

DESCRIPTION Imago Length of head toapex of clypeus 102 mm width of head withcompound eyes 130 mm maximum diameterof compound eye with ocular sclerite026 mm separation of compound eye fromlower margin 003 mm length of ocellus013 mm width of ocellus 006 mm ocellocu-lar distance 003 mm median length ofpronotum 087 mm width of pronotum102 mm length of metatibia 125 mm lengthof forewing scale 077 mm length of hindwing scale 049 mm Head large broadlyrounded distinctly wider than long withupwardly lifted frontoclypeal region wherebypostclypeus forms wide projecting proboscis(figs 3ndash4) Postclypeus projected forward toform broad proboscis width of extensiongreater than compound eye diameter withwide dorsal mediolongitudinal grooveFontanelle small located at base of upliftedproboscis and at base of dorsal groove

TABLE 1Measurements of soldiers of Dolichorhinotermes lanciarius new species

(all measurements in millimeters)

Metric

Major soldier

Holotype

Major soldier

Paratype

Minor soldier

Paratypes

Number of specimens 1 1 3

Length of head to side base of mandibles 158 158 069ndash071

Length of head with mandibles 260 255 na

Length of head to apex of labrum 250 255 155ndash158

Maximum width of head 168 168 069

Maximum height of head (without

postmentum) 095 097 049ndash051

Length of left mandible 127 132 vestigial

Length of right mandible 122 127 vestigial

Length of labrum along midline 077 079 079ndash081

Width of labrum at apex 023 026 013ndash015

Width of labrum at base 020 023 008

Maximum width of postmentum 043 046 030

Minimum width of postmentum 033 038 026

Length of postmentum 077 077 036

No of antennal articles ndash 14+a 15andash16

Median length of pronotum 043 048 036

Maximum width of pronotum 082 084 051

Length of metatibia 135 137 094ndash096

aThe antenna is broken after the 14th article in the major soldier paratype and after the 15th in one of the minor soldier

paratypes and so the total number is unknown for these individuals

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 5

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 4: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

second tooth and both shorter than those ofleft mandible second tooth extending straightout from inner mandibular surface first toothrunning closer along mandibular surface(fig 1) Labrum elongate slightly wider atapex apex rounded without emarginationwith slight dorsal longitundinal depressionextending posteriorly from apical marginabout one-half of labral length Palpi andpostmentum as depicted in figure 1 Antennawith at least 14 articles (total number un-certain) third flagellar article distinctly short-er than first flagellar article pedicel andsecond through fourth flagellar articles rough-ly equal in length Pronotum saddle-shapedanterior margin broadly rounded mediallylateral margins converging posteriorly poste-

rior margin relatively straight Legs relativelyslender and long Measurements provided intable 1 for holotype and paratype

Minor soldier Head thorax and abdomenpale yellow sparsely setose setae erect tosuberect and pale yellow Head with lateralborders weakly and gently convex sidesconverging anteriorly and slightly convergingfrom midpoint posteriorly to a gently roundedposterior border Labrum with several erectpale yellow setae at apex otherwise dorsalsurface with scattered short pale yellow setaesetae slightly more numerous in apical halfthan basal half Antenna with 16 articlesthird flagellar article distinctly shorter thanthe first Mandibles vestigial representingby broad fleshy lobes bearing a minute

Fig 1 Dolichorhinotermes lanciarius Engel and Krishna new species major soldier A Dorsal aspect ofhead and pronotum B Ventral aspect of head Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 3

sclerotized point (fig 2) Labrum narrowelongate forked at apex lateral processesof fork relatively widely separated Rangeof measurements for paratypes provided intable 1

HOLOTYPE Major soldier Ecuador Gua-laquiza 6xi1934 [6 November 1934] coll Wv Hagen [Gualaquiza at 3u249S 78u339Wis a village in Morona-Santiago Provinceon the eastern slopes of the Andes about97 km northeast of Loja] The holotype ispreserved in the Division of InvertebrateZoology American Museum of NaturalHistory

PARATYPE One major soldier several mi-nor soldiers several workers same data asholotype Deposited in the Division ofInvertebrate Zoology American Museum ofNatural History

ETYMOLOGY The specific epithet is takenfrom the Latin for lsquolsquolancerrsquorsquo in reference to thelance-like labrum of soldiers in the genus

COMPARISONS Dolichorhinotermes lanciar-ius can be readily separated from all otherspecies in the genus by its larger bodyproportions (refer to table 1) The species isgenerally quite similar to D japuraensisConstantinto from Brazil but aside from size

Fig 2 Dolichorhinotermes lanciarius Engel and Krishna new species minor soldier A Dorsal aspect ofhead and pronotum B Lateral aspect of head Scale bar 5 05 mm

4 AMERICAN MUSEUM NOVITATES NO 3592

differs by the rounded labral apex in majorsoldiers (very weakly medioapically concave inD japuraensis) sparse setation (setae of headmore numerous and elongate in D japuraen-sis) more distinctly convex anterior margin ofthe pronotum (very weakly convex in Djapuraensis) right mandible with two well-developed teeth (one stout tooth medially andone minute tooth at basal third in Djapuraensis) and by the more pronouncedapical labral forking in the minor soldier(weakly concave at the apex of the labrum inD japuraensis)

Dolichorhinotermes apopnus new species

figures 3ndash4

DIAGNOSIS The Mexican amber species isremarkably similar to D dominicanus exceptthat D apopnus lacks the integumental wrin-kles lateral to the postclypeal groove (imbri-cate in the Mexican fossil) the general bodyproportions are distinctly smaller (head widthwith compound eyes in D dominicanus160 mm while it is 130 mm in the new fossil

the compound eye is 032 mm in D dominica-nus only 026 mm in D apopnus length ofmetatibia 16 mm in D dominicanus 125 mmin D apopnus) and the integument of the headis darker

DESCRIPTION Imago Length of head toapex of clypeus 102 mm width of head withcompound eyes 130 mm maximum diameterof compound eye with ocular sclerite026 mm separation of compound eye fromlower margin 003 mm length of ocellus013 mm width of ocellus 006 mm ocellocu-lar distance 003 mm median length ofpronotum 087 mm width of pronotum102 mm length of metatibia 125 mm lengthof forewing scale 077 mm length of hindwing scale 049 mm Head large broadlyrounded distinctly wider than long withupwardly lifted frontoclypeal region wherebypostclypeus forms wide projecting proboscis(figs 3ndash4) Postclypeus projected forward toform broad proboscis width of extensiongreater than compound eye diameter withwide dorsal mediolongitudinal grooveFontanelle small located at base of upliftedproboscis and at base of dorsal groove

TABLE 1Measurements of soldiers of Dolichorhinotermes lanciarius new species

(all measurements in millimeters)

Metric

Major soldier

Holotype

Major soldier

Paratype

Minor soldier

Paratypes

Number of specimens 1 1 3

Length of head to side base of mandibles 158 158 069ndash071

Length of head with mandibles 260 255 na

Length of head to apex of labrum 250 255 155ndash158

Maximum width of head 168 168 069

Maximum height of head (without

postmentum) 095 097 049ndash051

Length of left mandible 127 132 vestigial

Length of right mandible 122 127 vestigial

Length of labrum along midline 077 079 079ndash081

Width of labrum at apex 023 026 013ndash015

Width of labrum at base 020 023 008

Maximum width of postmentum 043 046 030

Minimum width of postmentum 033 038 026

Length of postmentum 077 077 036

No of antennal articles ndash 14+a 15andash16

Median length of pronotum 043 048 036

Maximum width of pronotum 082 084 051

Length of metatibia 135 137 094ndash096

aThe antenna is broken after the 14th article in the major soldier paratype and after the 15th in one of the minor soldier

paratypes and so the total number is unknown for these individuals

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 5

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 5: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

sclerotized point (fig 2) Labrum narrowelongate forked at apex lateral processesof fork relatively widely separated Rangeof measurements for paratypes provided intable 1

HOLOTYPE Major soldier Ecuador Gua-laquiza 6xi1934 [6 November 1934] coll Wv Hagen [Gualaquiza at 3u249S 78u339Wis a village in Morona-Santiago Provinceon the eastern slopes of the Andes about97 km northeast of Loja] The holotype ispreserved in the Division of InvertebrateZoology American Museum of NaturalHistory

PARATYPE One major soldier several mi-nor soldiers several workers same data asholotype Deposited in the Division ofInvertebrate Zoology American Museum ofNatural History

ETYMOLOGY The specific epithet is takenfrom the Latin for lsquolsquolancerrsquorsquo in reference to thelance-like labrum of soldiers in the genus

COMPARISONS Dolichorhinotermes lanciar-ius can be readily separated from all otherspecies in the genus by its larger bodyproportions (refer to table 1) The species isgenerally quite similar to D japuraensisConstantinto from Brazil but aside from size

Fig 2 Dolichorhinotermes lanciarius Engel and Krishna new species minor soldier A Dorsal aspect ofhead and pronotum B Lateral aspect of head Scale bar 5 05 mm

4 AMERICAN MUSEUM NOVITATES NO 3592

differs by the rounded labral apex in majorsoldiers (very weakly medioapically concave inD japuraensis) sparse setation (setae of headmore numerous and elongate in D japuraen-sis) more distinctly convex anterior margin ofthe pronotum (very weakly convex in Djapuraensis) right mandible with two well-developed teeth (one stout tooth medially andone minute tooth at basal third in Djapuraensis) and by the more pronouncedapical labral forking in the minor soldier(weakly concave at the apex of the labrum inD japuraensis)

Dolichorhinotermes apopnus new species

figures 3ndash4

DIAGNOSIS The Mexican amber species isremarkably similar to D dominicanus exceptthat D apopnus lacks the integumental wrin-kles lateral to the postclypeal groove (imbri-cate in the Mexican fossil) the general bodyproportions are distinctly smaller (head widthwith compound eyes in D dominicanus160 mm while it is 130 mm in the new fossil

the compound eye is 032 mm in D dominica-nus only 026 mm in D apopnus length ofmetatibia 16 mm in D dominicanus 125 mmin D apopnus) and the integument of the headis darker

DESCRIPTION Imago Length of head toapex of clypeus 102 mm width of head withcompound eyes 130 mm maximum diameterof compound eye with ocular sclerite026 mm separation of compound eye fromlower margin 003 mm length of ocellus013 mm width of ocellus 006 mm ocellocu-lar distance 003 mm median length ofpronotum 087 mm width of pronotum102 mm length of metatibia 125 mm lengthof forewing scale 077 mm length of hindwing scale 049 mm Head large broadlyrounded distinctly wider than long withupwardly lifted frontoclypeal region wherebypostclypeus forms wide projecting proboscis(figs 3ndash4) Postclypeus projected forward toform broad proboscis width of extensiongreater than compound eye diameter withwide dorsal mediolongitudinal grooveFontanelle small located at base of upliftedproboscis and at base of dorsal groove

TABLE 1Measurements of soldiers of Dolichorhinotermes lanciarius new species

(all measurements in millimeters)

Metric

Major soldier

Holotype

Major soldier

Paratype

Minor soldier

Paratypes

Number of specimens 1 1 3

Length of head to side base of mandibles 158 158 069ndash071

Length of head with mandibles 260 255 na

Length of head to apex of labrum 250 255 155ndash158

Maximum width of head 168 168 069

Maximum height of head (without

postmentum) 095 097 049ndash051

Length of left mandible 127 132 vestigial

Length of right mandible 122 127 vestigial

Length of labrum along midline 077 079 079ndash081

Width of labrum at apex 023 026 013ndash015

Width of labrum at base 020 023 008

Maximum width of postmentum 043 046 030

Minimum width of postmentum 033 038 026

Length of postmentum 077 077 036

No of antennal articles ndash 14+a 15andash16

Median length of pronotum 043 048 036

Maximum width of pronotum 082 084 051

Length of metatibia 135 137 094ndash096

aThe antenna is broken after the 14th article in the major soldier paratype and after the 15th in one of the minor soldier

paratypes and so the total number is unknown for these individuals

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 5

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 6: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

differs by the rounded labral apex in majorsoldiers (very weakly medioapically concave inD japuraensis) sparse setation (setae of headmore numerous and elongate in D japuraen-sis) more distinctly convex anterior margin ofthe pronotum (very weakly convex in Djapuraensis) right mandible with two well-developed teeth (one stout tooth medially andone minute tooth at basal third in Djapuraensis) and by the more pronouncedapical labral forking in the minor soldier(weakly concave at the apex of the labrum inD japuraensis)

Dolichorhinotermes apopnus new species

figures 3ndash4

DIAGNOSIS The Mexican amber species isremarkably similar to D dominicanus exceptthat D apopnus lacks the integumental wrin-kles lateral to the postclypeal groove (imbri-cate in the Mexican fossil) the general bodyproportions are distinctly smaller (head widthwith compound eyes in D dominicanus160 mm while it is 130 mm in the new fossil

the compound eye is 032 mm in D dominica-nus only 026 mm in D apopnus length ofmetatibia 16 mm in D dominicanus 125 mmin D apopnus) and the integument of the headis darker

DESCRIPTION Imago Length of head toapex of clypeus 102 mm width of head withcompound eyes 130 mm maximum diameterof compound eye with ocular sclerite026 mm separation of compound eye fromlower margin 003 mm length of ocellus013 mm width of ocellus 006 mm ocellocu-lar distance 003 mm median length ofpronotum 087 mm width of pronotum102 mm length of metatibia 125 mm lengthof forewing scale 077 mm length of hindwing scale 049 mm Head large broadlyrounded distinctly wider than long withupwardly lifted frontoclypeal region wherebypostclypeus forms wide projecting proboscis(figs 3ndash4) Postclypeus projected forward toform broad proboscis width of extensiongreater than compound eye diameter withwide dorsal mediolongitudinal grooveFontanelle small located at base of upliftedproboscis and at base of dorsal groove

TABLE 1Measurements of soldiers of Dolichorhinotermes lanciarius new species

(all measurements in millimeters)

Metric

Major soldier

Holotype

Major soldier

Paratype

Minor soldier

Paratypes

Number of specimens 1 1 3

Length of head to side base of mandibles 158 158 069ndash071

Length of head with mandibles 260 255 na

Length of head to apex of labrum 250 255 155ndash158

Maximum width of head 168 168 069

Maximum height of head (without

postmentum) 095 097 049ndash051

Length of left mandible 127 132 vestigial

Length of right mandible 122 127 vestigial

Length of labrum along midline 077 079 079ndash081

Width of labrum at apex 023 026 013ndash015

Width of labrum at base 020 023 008

Maximum width of postmentum 043 046 030

Minimum width of postmentum 033 038 026

Length of postmentum 077 077 036

No of antennal articles ndash 14+a 15andash16

Median length of pronotum 043 048 036

Maximum width of pronotum 082 084 051

Length of metatibia 135 137 094ndash096

aThe antenna is broken after the 14th article in the major soldier paratype and after the 15th in one of the minor soldier

paratypes and so the total number is unknown for these individuals

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 5

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 7: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

Fig 3 Photomicrograph of holotype imago of Dolichorhinotermes apopnus Engel and Krishna newspecies (AMNH Ch-50) Scale bar 5 10 mm

6 AMERICAN MUSEUM NOVITATES NO 3592

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 8: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

positioned at level of ocelli Compound eyeslarge bulging exophthalmic (figs 3ndash4) Ocelliraised large elongate (longer than wide) nottouching compound eyes Antennae incom-plete left antenna broken at pedicel rightantenna broken at fourth flagellar articlethird flagellar article slightly shorter than firstflagellar article flagellar articles with severalerect setae Integument of head smooth exceptslightly imbricate (but not wrinkled) lateral topostclypeal groove Head and antennae darkbrown nearly black Head with scatterederect setae such setae not located anterior tolevel of ocelli a few setae distinctly clusterednear posterior border of compound eyePronotum wider than long anterior marginslightly convex medially integument brownwith sparse erect to suberect setae some setaedirected posteriorly Remainder of thorax andabdomen brown Legs brown with scatteredshorter setae pretarsal ungues simple aroliumabsent Wing membrane hyaline strongly

reticulate veins very faintly brown except Cand Sc brown forewing scale longer than hindwing scale forewing scale apex overlappingbasal half of hind wing scale when in reposescale with several erect to suberect setaesuberect setae directed toward wing apex

HOLOTYPE Imago AMNH Ch-50 amberMexico Chiapas Simojovel The holotype isdeposited in the Amber Fossil CollectionDivision of Invertebrate Zoology AmericanMuseum of Natural History

ETYMOLOGY The specific epithet is theGreek word apopnus meaning lsquolsquodeceasedrsquorsquo

ACKNOWLEDGEMENTS

We are grateful to Ismael A Hinojosa-Dıazfor translating our abstract into Spanish toSteve Thurston for assistance in the prepara-tion of the figures and to Torsten WapplerDavid Kistner and Valerie Krishna forcarefully reading the manuscript Support for

Fig 4 Photomicrographic detail of head pronotum and basal portions of wings of holotype imago ofDolichorhinotermes apopnus Engel and Krishna new species (AMNH Ch-50) Scale bar 5 10 mm

2007 ENGEL AND KRISHNA NEW DOLICHORHINOTERMES 7

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592

Page 9: New Dolichorhinotermes from Ecuador and in Mexican Amber (Isoptera: Rhinotermitidae)

this work was provided by a GuggenheimFellowship from the John Simon GuggenheimMemorial Foundation (to MSE) This iscontribution No 3469 of the Division ofEntomology University of Kansas NaturalHistory Museum

REFERENCES

Constantino R 1990 [1991] Two new species oftermites (Insecta Isoptera) from westernBrazilian Amazonia Boletim do MuseuParaense Emılio Goeldi serie Zoolgia 6(1) 3ndash9

Constantino R 1998 Catalog of the living termitesof the New World (Insecta Isoptera) Arquivosde Zoologia (Sao Paulo) 35(2) 135ndash231

Emerson AE 1925 The termites of KartaboBartica District British Guiana Zoologica6(4) 291ndash459

Ensaf A and J-M Betsch 2002 Dolicho-rhinotermes neli n sp nouvelle espece pourla science et Cornicapritermes mucronatusEmerson 1950 nouvelle espece de termite pourla Guyane francaise (Isopt Rhinotermitidaeet Termitidae) Bulletin de la Societe Ento-mologique de France 107(4) 346ndash348

Ensaf A RE Garrouste J-M Betsch and A Nel2004 Dolichorhinotermes longilabius (Emerson1925) new species for French Guiana witha preliminary list of the termites in FrenchGuiana Mitteilungen der Schweizerischen

Entomologischen Gesellschaft [Bulletin de laSociete Entomologique Suisse] 77(3ndash4) 277ndash288

Maiti PK 2006 A taxonomic monograph onthe world species of termites of the familyRhinotermitidae (Isoptera Insecta) Memoirsof the Zoological Survey of India 20(4) 1ndash272

Mathews AGA 1977 Studies on termites fromthe Mato Grosso State Brazil Rio de JaneiroAcademia Brasileira de Ciencias 267 pp

Mathur RN and RS Thapa 1962 A revisedcatalogue of Isoptera (white ants) of theEntomological Reference Collection at theForest Research Institute Dehra Dun IndianForest Leaflet (Entomology) 167 1ndash122

Quennedey A and J Deligne 1975 Lrsquoarmefrontale des soldats de termites I Rhinoter-mitidae Insectes Sociaux 22(3) 243ndash267

Schlemmermeyer T and EM Cancello 2000 Newfossil termite species Dolichorhinotermes domin-icanus from Dominican amber (Isoptera Rhi-notermitidae Rhinotermitinae) Papeis Avulsosde Zoologia (Sao Paulo) 41(20) 303ndash311

Snyder TE 1924 An extraordinary newRhinotermes from Panama Proceedings of theBiological Society of Washington 37(1) 83ndash86

Snyder TE 1926 Termites collected on theMulford Biological Exploration to theAmazon Basin 1921ndash1922 Proceedings of theUnited States National Museum 68(14) 1ndash76

Snyder TE 1949 Catalog of the termites(Isoptera) of the world Smithsonian Miscel-laneous Collections 112 1ndash490

This paper meets the requirements of ANSINISO Z3948-1992 (Permanence of Paper)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide

Web site httplibraryamnhorgpubs Inquire about ordering printed copies via e-mail from

scipubsamnhorg or via standard mail from American Museum of Natural History

LibrarymdashScientific Publications Central Park West at 79th St New York NY 10024 TEL

(212) 769-5545 FAX (212) 769-5009

8 AMERICAN MUSEUM NOVITATES NO 3592


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