Planthopper (Hemiptera Flatidae) Parasitized by Larval Erythraeid Mite

(Trombidiformes Erythraeidae)mdashA Description of Two New Species From

Western Madagascar

Joanna Makol12

Hanna Moniuszko1

Dariusz Swierczewski3

and Adam Stroinski4

1Institute of Biology Department of Invertebrate Systematics and Ecology Wrocław University of Environmental and Life Sciences Kozuchowska 5b 51-631Wrocław Poland2Corresponding author e-mail joannamakolupwrocpl3Department of Zoology and Animal Ecology Jan Długosz University Armii Krajowej 1315 42-201 Czestochowa Poland4Museum and Institute of Zoology Polish Academy of SciencesWilcza 64 00-697 Warsaw Poland

Suject Editor Takumasa Kondo

J Insect Sci 14(194) 2014 DOI 101093jisesaieu056

ABSTRACT Descriptions of Dambullaeus adonis Makol et Moniuszko sp nov (Trombidiformes Erythraeidae Callidosomatinae) andLatois nigrolineata Swierczewski et Stroinski sp nov (Hemiptera Fulgoromorpha Flatidae) from Madagascar are provided The firsthost record for ectoparasitic larvae of Dambullaeus Haitlinger 2001 and the first evidence on hostndashparasite association between flatidadult and erythraeid larvae are given Genus Dambullaeus known exclusively from larvae and now comprising two species ofGondwanan distribution is critically reappraised

Key Words host parasite Fulgoromorpha Parasitengona taxonomy

Madagascar is one of eight important global biodiversity hotspots ow-ing to its unique biota and the high level of threat to its natural habitats(McNeely et al 1990 Myers et al 2000 Ganzhorn et al 2001) It hasevolved as an incredible wealth of biodiversity with thousands of spe-cies that can be found nowhere else on earth On the one hand this isdue to the long isolation of Madagascar from all other landmasses(Storey et al 1995) and on the other several alternative mechanismshave generated local endemism (Vences et al 2009)

The knowledge of erythraeid fauna (Trombidiformes Erythraeidae)of Madagascar is scarce and limited to single records on Abrolophusaitapensis (Southcott 1948) (Abrolophinae) Charletonia adellaeHaitlinger 2007 Charletonia agatae Haitlinger 1987 Charletoniaalarobiaensis Haitlinger 1987 Charletonia arlettae Haitlinger 1987Charletonia dorotae Haitlinger 1987 Charletonia edytae Haitlinger1987 Charletonia iwonae Haitlinger 1987 Charletonia justynaeHaitlinger 1987 Charletonia kibonotensis (Tragardh 1908)Charletonia milloti (Andre 1946) Charletonia tatianae (Tragardh1908) (Callidosomatinae) Leptus aldonae (Tragardh 1908) Leptusmadagascariensis (Andre 1941) and Leptus maranaensis Haitlinger1987 (Leptinae) (Andre 1941 1946 Haitlinger 1987 2007) Of thoseC milloti has been known exclusively from active postlarval formswhich are recognized as predators in parasitengone mites whereas forother taxa the parasitic larva constitutes the only described instarErythraeid larvae parasitize various arthropods Data on hosts are scarceand in majority of cases restricted to unidentified representatives ofhexapod orders

The monotypic genus Dambullaeus Haitlinger 2001 withDambullaeus piae Haitlinger 2001 was described from Sri Lankabased on single specimen collected from plants and since then no fur-ther account of the genus has been published Here we provide a de-scription of a second representative of the genus Dambullaeus adonisMakol et Moniuszko sp nov ectoparasitic the newly described flatidplanthopper of the genus Latois Stal 1866 (Hemiptera Flatidae) fromMadagascar The present distribution pattern of Dambullaeus points tothe history of the genus which may date back to the sundering ofGondwana and separation of India fromMadagascar

Flatidae constitute one of the largest families within planthoppers(Fulgoromorpha Hemiptera) with 1437 species described in 294 gen-era and 12 tribes distributed worldwide (Bourgoin 2013) These phy-tophagous insects are highly diverse in terms of their color and size(from 45 up to 32 mm) and are found on all continents but are espe-cially common and abundant in the tropics (OrsquoBrien 2002) They aredivided into two subfamiliesmdashFlatinae and Flatoidinae which in mostcases can be easily distinguished from each other by the shape of thebody A majority of Flatinae are flattened laterally in contrast toFlatoidinae which hold their wings horizontally (OrsquoBrien and Wilson1985) Madagascan flatid fauna presently consists of 17 genera with 39species of Flatinae and 11 genera with 37 species of Flatoidinae and hasbeen reviewed by Swierczewski and Stroinski (2013)

The flatid planthopper genus Latois Stal 1866 covers six speciesLatois antica (Signoret 1860) Latois suturalis (Signoret 1860) Latoisbicoloripes Karsch 1890 Latois frontalis Melichar 1901 Latoismajor Melichar 1901 and recently discovered Latois nigrofasciataSwierczewski et Stroinski 2012 In this article we describe another spe-cies Latois nigrolineata Swierczewski et Stroinski sp nov from threelocalities in the western part of the island

Materials and Methods

The studied material originating from a few localities in westernMadagascar (see Type Material) comes from the collection of theCalifornia Academy of Sciences in San Francisco (CAS) (Dr N Pennycurator)

Flatid Host Preparation The abdomens were cut off and boiled in10 KOHwith a few drops of chlorazol black for dying the ectodermicgenital ducts based on the method introduced by Carayon (1969) andBourgoin (1993) Dissections and cleaning of genital structures wereperformed in distilled water Final observations and drawings weredone in glycerin using a camera lucida attached to a light microscopeThe photos of the habitus were taken using a stereoscopic microscopeLeica MZ 16 with digital camera IC 3D The photos of female genitalstructures were taken using a light microscope Leica DM5500Bwith digital camera Leica DFC49 Final images were created using

VC The Author 2014 Published by Oxford University Press on behalf of the Entomological Society of America

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (httpcreativecommonsorglicensesby-nc40) which permitsnon-commercial re-use distribution and reproduction in any medium provided the original work is properly cited For commercial re-use please contact journalspermissionsoupcom

Journal of Insect Science

RESEARCH

HELICON 50 and ADOBE PHOTOSHOP software The SEM photo-graphs of uncoated specimens were taken in the Laboratory ofScanning Microscopy MIZ PAS (Warsaw) using a scanning micro-scope HITACHI S-3400N under low vacuum conditions

The following measurements were taken and abbreviations used inthe description of hemipteran host total lengthmdashmeasured (in dorsalview) from the apex of head protrusion to the apex of tegmina ABmdashwidth of vertex measured at the anterior marginlength of vertex meas-ured in mid line CEmdashwidth of frons between eyeslength of frons inmid line DEmdashmaximum width of fronslength of frons in mid lineFBmdashlength of pronotum in mid linelength of vertex in mid lineGFmdashlength of mesonotumlength of pronotum in mid line GBthornFmdashlength of mesonotumcumulative length of vertex and pronotum in midline GHmdashlength of mesonotum in mid linewidth of mesonotumbetween lateral angles IJmdashlength of tegmen measured from the baseto the apical margin in median portionwidth of tegmen measured fromthe apex of clavus to the anterior margin The nomenclature of the malegenitalia follows Bourgoin (1988) and Bourgoin and Huang (1990)and for the female genitalia Bourgoin (1993) Vein nomenclature afterinterpretation proposed by Szwedo and _Zyła (2009)

Erythraeid Larvae Preparation Larvae (for the place of origin seeType Material under D adonis Makol et Moniuszko sp nov) attachedto the host specimen were stored in 70 ethanol for 5 yr and after pin-ning the host they were kept dry for a half year in the entomologicalcollection During the host preparation larvae were dyed together withthe host in warm 10 KOH with a few drops of chlorazol black andafter cleaning in distilled water transferred to glycerol Morphologicalexamination of larvae was carried out in LM and in SEM The scanningelectron microscope Zeiss EVO LS15 served for close-up investigationof fine structures in dehydrated and gold-coated specimens Larvae des-tined for light microscopy studies were preserved in 70ndash75 ethanoland fixed on permanent slides in Faurersquos fluid (Walter and Krantz2009) Measurements were taken under a Nikon Eclipse E600 with dif-ferential interference contrast coupled with NIS-Elements Br softwarewhereas drawings (except the SEM photo-based Fig 2) were madeunder a Nikon Eclipse 80i combined with camera lucida All measure-ments are given in micrometers Terminology abbreviations andmodes of measurements follow Makol et al (2012a) and Southcott(1961)

Nomenclature The new taxa names published in this article havebeen registered in ZooBank (wwwzoobankorg) The LSID isurnlsidzoobankorgpub2FF09FDB-700B-418C-818E-7FF0A873E919

Results

Taxonomy Order Hemiptera L 1758Suborder Fulgoromorpha Evans 1946Superfamily Fulgoroidea Latreille 1807Family Flatidae Spinola 1839Subfamily Flatinae Spinola 1839Genus Latois Stal 1866Type speciesNephesa antica Signoret 1860L nigrolineata Swierczewski et Stroinski sp nov (Figs 1ndash33 and 50)

Material Examined

Type Material Holotype (MADAGASCAR MajungaAmbovomamy Belambo 20 km NW of Port Berger 26 Mayndash5 June2007) (15 27070 S 47 36800 E CAS coll R HarinrsquoHala MIrwin F Parker malaise secondary growth on white sand elev 33 mMG-33-23) (CASLOT 044483) (CAS)

Paratypes (MADAGASCAR Majunga Ambovomamy Belambo20 km NW of Port Berger 6ndash12 April 2007 15 27070 S 47 36800

E) (CAS coll R HarinrsquoHala M Irwin F Parker malaise secondarygrowth on white sand elev 33 m MG-33-16) (CASLOT 034286)mdash1$ (CAS) (MADAGASCAR Tulear Province Mikea Forest NW ofManombo elev 30 m 6ndash16 January 2002 22 54220 S 43 28530 E)(coll M Irwin R HarinrsquoHala CAS malaise trapmdashin deciduousdry forest MA-02-18A-09) (CASLOT 044695)mdash1 (CAS)(MADAGASCAR Majunga Prov Besalampy District Analangidrodry forest 17 kmWof Besalampy 14ndash21 December 2007) (16 41490

S 4 31410 E CAS coll M Irwin R HarinrsquoHala malaise dry foreston sand elev 200 ft MG-41-12)ndash1 (CAS)

Etymology The specific epithet refers to black narrow band on dor-sal surface of thorax

Diagnosis The new species differs from other members of the genusin the following characters frons without protrusion in the upper part ofhead median carina absent or as remnant

Description Total length 075ndash077 cmHead truncate with compound eyes (in dorsal view) about the same

width as thorax (Figs 2 5 and 6) Vertex transverse distinctly wider

Figs 1ndash4 L nigrolineata Swierczewski et Stroinski sp nov (1) Frontal view (2) Anterior part of body dorsal view (3) Tegmen (4) Malegenital capsule

2 JOURNAL OF INSECT SCIENCE VOLUME 14

than long at midline proportion AB frac14 264ndash30 posterior part partlycovered by pronotum (Figs 5ndash7) lateral margins slightly arcuate andsubparallel anterior margin ridgeous and convex medially obsoletedisc of vertex in median portion elevated with median carina(Figs 5ndash8)

Frons (Figs 1 and 8ndash10) a little longer than wide the widest belowthe level of compound eyes proportion CE frac14 090ndash100 proportionDE frac14 100ndash108 frons without protrusion in the upper part of headdisc of frons with two well visible arcuate and connected at base lateralcarinae reaching the level of antennae median carina absent or asremnant lateral margins of frons carinate and elevated lateral marginwithout incision (Figs 1 8 and 9) disc of frons irregularly rugosewith sensory and excretory organs (Fig 10) central part (between lat-eral carinae) depressed

Antennal segment II (pedicel) about as long as wide wider at apexsensory organs and sensilla located at the top of pedicel in shallowdepression and partly at upper surface (Figs 7 and 11) Compound eyesoval with very small callus at lower posterior margin lateral ocellipresent

Frontoclypeal suture arcuate clypeus without carinae median por-tion convex (Fig 1) Rostrum reaching hind coxae apical part a bitshorter than the basal one

Thorax Pronotum (Figs 2 and 5ndash8) a bit shorter than vertex (propor-tion FB frac14 080ndash091) anterior margin (in lateral view) at the samelevel as posterior margin anterior margin (in dorsal view) almoststraight partly covering posterior margin of vertex posterior margindeeply incised disc of pronotum with median carina and lateral pitslateral parts of disc with strong and sharp postocular eminences

Mesonotum (Figs 2 and 6) deltoid proportion GF frac14 640ndash813GB thorn F frac14 305ndash361 GH frac14 115ndash118 in lateral view at the samelevel as posterior margin of pronotum lateral angles placed before 1=2of the length of mesonotum at midline disc of mesonotum with threeseparated at base and parallel carinae reaching the posterior margin

Tegmen (Figs 3 and 12ndash16) subrectangular membranous flat sur-face smooth and proportion IJ frac14 181ndash195 costal marginmdashbasalpart to 13 weakly arcuate posterior part almost straight costal anglebluntly rounded apical margin strongly arcuate sutural angle roundedpostclaval sutural margin straight costal area narrower than costal cellat midline widening apicad with dense and numerous transverse vein-lets ending at the level of end of clavus costal cell tapering apicadwith sparse net of veinlets basal cell more twice as long as wide

Longitudinal stem ScthornR arises as extremely short common stemfrom basal cell ScthornRA weakly basally elevated distal part of RPextremely weakly visible M leaving basal cell with a long stalk butshorter than CuA stalk First fork of ScthornRA near the end of costal areafirst fork of RP before end of costal area Location of M1thorn2 fork afterRP fork M3thorn4 fork before M1thorn2 CuA diverging after the level of Mfork Sc ending with 1 terminal RAwith 7ndash9 RP with 6ndash8 M1thorn2 with11ndash16 M3thorn4 with 11ndash15 terminals and CuAwith 5ndash7 terminals endingat postclaval margin Claval veins not elevated connected a little beforeend of clavus transverse veinlets absent

Irregular net of numerous transverse veinlets starting from basalpart of tegmen nodal line absent apical line weakly distinguishabletubercles present mainly on costal area between basal parts of ScthornRAand M veins and on clavus Sensory and secretory structures on thewhole surface of tegmen (Figs 15 and 16)

Figs 5ndash10 L nigrolineata Swierczewski et Stroinski sp nov (5) Anterior part of body dorsal view (6) Head and thorax dorsal view(7) Anterior part of body dorsolateral view (8 9) Anterior part of body frontal view (10) Frons surface

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 3

Femora shorter than tibiae fore and middle tibiae rectangular incross section with carinate margins hind tibia arcuate with two lateralspines after midlength row of seven apical teeth in formula two (lon-ger) thorn five (shorter) basitarsomere as long as cumulative length of sec-ond and hind tarsomeres with arcuate line of seven apical teeth lateralteeth larger than internal ones

Male Anal tube (in lateral view Figs 4 and 17) elongated and dis-tinctly curved at midline basal part about the same width as apical partand anus placed a bit after half of length Anal tube (in dorsal viewFig 18) in a shape of baseball bat basal part distinctly separated androunded apical part tapered shallowly incised medially and anusplaced a bit after half of length

Pygofer (in lateral view Figs 4 and 17) higher than wide dorsalpart narrower than ventral posterior margin arcuate Posterodorsalangle widely rounded without process

Genital styles (in lateral view Figs 4 and 17) L-shaped dorsal mar-gin basally almost straight before capitulum distinctly rising and form-ing fold capitulum short and sharp posterior margin arcuate ventralmargin weakly arcuate apically with sharp and short process

Phallic complex periandrium (in lateral view Figs 19 and 20)excluding median ventral keel about the same width and distinctlycurved lateral split surpassing half of length of periandrium dorsal partlonger than ventral apically (in lateral view) with narrow incisionlower part of dorsal periandrium with spiniferous microsculpture andtwo well sclerotized single processes oriented ventro-basad lowerprocess shorter than upper upper part of periandrium apically well scle-rotized with membranous pleated layers (Fig 20) ventral part of peri-andrium shorter than dorsal bluntly rounded with wide keel (Fig 19)

Aedeagus (in lateral view Fig 21) shaft as long as dorsal part ofperiandrium weakly arcuate and apically bluntly rounded posteroven-tral part near apex with well sclerotized bulb-like appendage ventralfold (Fig 22) with deeply incised apex and spiniferous margins

Female Pregenital sternite (Fig 25) with wide and well-sclerotizedmedian portion lateral parts narrower and weakly separated frommedian part anterior margin almost straight posterior margin weaklyconvex and covered by additional narrow and convex lobe

Anal tube (in lateral view Fig 26) oval ventral margin strongly arc-uate posterior part bluntly rounded anus placed before midlength analtube extending a bit beyond the posterior margin of the gonoplac

Anal tube (in dorsal view Fig 27) oval anterior margin weaklyconcave posterior margin in median portion with distinct incision lat-eral margins arcuate anus placed before midlength

Gonoplac unilobate elongately oval (Fig 29) posterior margin in34 with row of 14 well developed elongated dense teeth posteroven-tral angle with membranous smooth narrow and elongate lobe dorsalpart of ventral margin with long setae

Gonapophysis VIII (in lateral view of the external side Figs 30and 31) triangular and laterally flattened tapering apicad with jaggedventral margin apical part with four horizontal and one vertical den-ticles endogonocoxal process huge as long as gonapophysis taperingapicad membranous with spiniferous microsculpture and marginalhairs

Gonospiculum as in Figs 32 and 33Bursa copulatrix (Fig 23) with single pouch oval cells without

sclerites Spermatheca (Fig 24) well developed ductus receptaculi lon-ger than diverticulum ductus basal part smooth and narrow remaining

Figs 11ndash16 L nigrolineata Swierczewski et Stroinski sp nov (11) Antenna (12ndash14) Tegmen (15) Tegmen surface (16) Tegmen sensorystructures

4 JOURNAL OF INSECT SCIENCE VOLUME 14

part widened and ribbed diverticulum ductus smooth basal part nar-rower than apical part

Coloration (Figs 1ndash4) General color milky white with pinkishtinge black or dark brown band with yellow boundaries extendingfrom the anterior margin of vertex through the dorsal part of head andthorax terminating at the end of scutellum with big black spot lateralmargins of frons clypeus and legs yellowish tegmina marginated withbrownish-yellow band

Distribution Madagascar Mahajanga and Toliara provinces (Fig 50)Order Trombidiformes Reuter 1909Suborder Prostigmata Kramer 1877Cohort Parasitengona Oudemans 1909Superfamily Erythraeoidea Robineau-Desvoidy 1828Family Erythraeidae Robineau-Desvoidy 1828Subfamily Callidosomatinae Southcott 1957GenusDambullaeusHaitlinger 2001Type speciesDambullaeus piaeHaitlinger 2001DambullaeusHaitlinger 2001

Diagnosis Larva Scutum oval in outline with straight anterior mar-gin Two pairs of trichobothria and three pairs of nonsensillary setaeAnterior pair of sensilla (ASens) located at c half length of scutumposterior sensilla (PSens)mdashclose to the posterior margin of scutumThe second pair of nonsensillary setae (ML) distinctly elongated morethan three times longer than nonsensillary setae of first (AL) and third(PL) pair One pair of circular eyes placed laterally to scutum Odontusbifid subterminally Leg segmentation formula 7-7-7 Coxal setation

formula 1-2-2 All tarsi terminated with double claws and claw-likeempodium Anterior claw fimbriated

Deutonymph and adult Not knownDistribution Sri Lanka andMadagascarD adonisMakol et Moniuszko sp nov (Figs 34ndash50)

Material Examined

Type Material Holotype larva (MADAGASCAR MajungaAmbovomamy Belambo 20 km NWof Port Berger 6ndash12 April 200715 27070 S 47 36800 E) (CAS coll R HarinrsquoHala M Irwin FParker malaise secondary growth on white sand elev 33 m MG-33-16) (CASLOT 034286) ectoparasitic on single specimen (female) ofL nigrolineata Swierczewski et Stroinski (CAS) paratypes larvae thesame data as for holotypemdasheight specimens (CAS) four specimens(collection of the Department of Invertebrate Systematics and EcologyInstitute of Biology Wrocław University of Environmental and LifeSciences Wrocław Poland)

Etymology The specific epithet adonis refers to the name of thedivinity in Greek mythology

Diagnosis Larva Scutum rounded posteriorly Setae AL ML andPL located in anterior half of scutum ML more than three times longerthan AL Odontus with stout subapical splinter on lateral face

The new species differs from Dambullaeus piae Haitlinger 2001 inthe shape of dorsal scutum (posterior half of scutum in the shape ofsemicircle in D adonis in D piae scutum narrowing posteriorly withtruncated posterior termination which encompasses the bases ofPSens) LW ratio (c 1 inD adonis lt 1mdashinD piae)

Figs 17ndash24 L nigrolineata Swierczewski et Stroinski sp nov (17) Genital capsule lateral view (18) Anal tube dorsal view (19) Periandriumlateral view (20) Periandrium ventral view (21) Aedeagus lateral view (22) Aedeagus ventral view (23) Female bursa copulatrix lateralview (24) Spermatheca

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 5

Description For morphometric data see Table 1Gnathosoma Cheliceral bases stout movable claw distinctly curved

(Fig 34) Mouth externally surrounded by flaps which facilitate theattachment to the host internallymdashwith lamellar fimbriae (Figs 35 38and 39) A pair of setulated thread-like at termination adoral setae (cs)(c 28) covered with minute barbs placed dorsally in anteromedianposition (Fig 38) Club-shaped supracoxal setae (elcp) (c 5) locatedposterolaterally at gnathosoma base (Fig 38) Ventral side of gnatho-soma with a pair of barbed setae (as) shorter (c 18) than cs and a pairof subcapitular setae (bs) (c 55) covered with setules in proximal halfof the stem (Fig 39) Pedipalp formula (Figs 36 and 37) 0-B-B-BBB-BBBBoz Odontus with stout subapical splinter on lateral face of theclaw Palp tarsus with four normal setae covered with setules one sole-nidion (o) located in proximal position and one distal eupathidium (z)

Dorsal side of idiosoma Scutum (Figs 38 and 41) with fine linearwisps of cuticle (character visible in SEM) Setae AL ML and PLplaced in anterior half of scutum PL at the widest part of the scleriteASens located just behind the level of PL PSens markedly longer thanASens shifted to the posterior margin of scutum Both ASens andPSens covered with setules arising along distal half of the stem Singleeye lenses (17 mm in diameter) placed on indistinct sclerites close tothe level of posterior margin of scutum The remaining part of idiosomacovered with folded in line cuticle Dorsal setae (Fig 40) covered withsetules distributed along the stem stem inserted in setal base in theshape of truncated cone fD frac14 30ndash35 (N frac14 13)

Ventral side of idiosoma (Fig 39) fCx frac14 1-2-2 Supracoxal setaeelc I present on dorsal side in terminal part of coxae I Setae 1b 2b andc 3b and c located on coxae I II III respectively Setae 1a 2a 3aplaced between coxal plates I II and III fV frac14 7ndash12 (N frac14 13) the total

number of setae in fD and fV formula (NDV) frac14 40ndash46 (N frac14 13)Ventral setae more sharpened apically than the dorsal setae

Legs (Figs 42ndash49) Leg segmentation formula 7-7-7 For leg chae-totaxy see Table 2 Normal setae on legs setulated Proximal solenidion(j) on tibia I adjoined by companala (z) Famulus (e) on tarsus I locateddistally to solenidion (o) Dorsal eupathidium (z) on tarsus I and subter-minal eupathidia (z) on tarsi IndashIII covered with barbs Microsetae (ves-tigialae k) on genu I tibia I and genu II club shaped All tarsiterminated with double claws and claw-like empodium which is longerand slender than claws Two claws similar in length without terminalhooks onemdashfimbriated along the ventral edge the other one smooth

Distribution Madagascar Mahajanga province (Fig 50)

Discussion

All species of Latois Stal 1866 described so far are confined tomoist habitats (littoral and humid forests) in the eastern part ofMadagascar and the Comoros Islands In contrast the newly describedspecies L nigrolineata Swierczewski et Stroinski is widely distributedin the western part of the island and associated with the vegetation ofsandy areas near streams and rivers

Interestingly here we describe for the first time the phenomenon oferythraeid mite parasitizing the representative of Flatidae familyHostndashparasite associations between Hemiptera and Erythraeidae arenot well documented and the overview of data referring to this topic iscontained in the article by Stroinski et al (2013) Of 826 species ofErythraeidae known for science (Makol and Wohltmann 2012 2013and present data) the larva has been described for 554 species and for181 species the host remains unknown Up to the present 66 erythraeidspecies have been reported to parasitize members of Hemiptera

Figs 25ndash33 L nigrolineata Swierczewski et Stroinski sp nov female (25) Pregenital sternite flattened (26) Anal tube lateral view (27) Analtube dorsal view (28) Anal tube ventral surface (29) Gonoplac external view (30) Gonapophysis VIII lateral view (31) Same apical part(32) Gonapophyses IX and gonospiculum bridge lateral view (33) Same dorsal view

6 JOURNAL OF INSECT SCIENCE VOLUME 14

Table 1 Morphometric data on larvae of D adonis Makol et Moniuszko sp nov

Character D adonis Makol et Moniuszko sp nov D piae Haitlinger 2001

Samplesize

Mean(mm)

Range(mm)

Data for holotype (originaldescriptionpresent study)

GnathosomaGL 13 112 99ndash137 c 120120GW 9 88 81ndash101 ndash71Pa Tr (L) 13 20 16ndash24 ndash24Pa Fe (L) 13 33 27ndash40 ndash26Pa Ge (L) 13 18 14ndash21 ndash33Pa Ti (L) 13 16 11ndash20 ndash21Odontus (L) 13 28 23ndash32 ndash25Pa Ta (L) 13 13 10ndash17 ndash15

IdiosomaIL 13 245 209ndash303 11301127IW 13 185 165ndash206 c 762778ILIW 13 13 12ndash15 c 1514L 12 74 69ndash79 8285W 12 74 65ndash80 104105LW 12 10 09ndash11 0808ASens 10 30 27ndash34 c 4034SBa 12 18 15ndash21 1414PSens 9 49 43ndash56 5259SBp 13 13 11ndash14 1416ASBa 12 40 32ndash46 5849PSBp 12 4 2ndash5 45AL 13 30 23ndash32 3442AW 13 46 43ndash50 ndash50ML 13 106 96ndash112 100110MW 12 39 34ndash44 5240PL 13 30 27ndash35 5256PW 13 53 49ndash58 7069ISD 13 30 26ndash33 2028AWAL 13 16 13ndash20 ndash12AWISD 13 16 13ndash17 ndash18ALPL 13 10 09ndash11 ndash08PWAW 13 11 11ndash12 ndash14DS max 13 35 29ndash40 36ndash60 (DS)52VS max 13 28 26ndash29 ndash332a 12 34 26ndash39 ndash323a 12 26 22ndash32 ndash403a2a 12 08 06ndash10 ndash13

LEGSCx I (L) 13 55 47ndash64 7476Tr I (L) 13 33 26ndash37 4436bFe I (L) 13 65 57ndash73 5664tFe I (L) 13 64 58ndash72 6052Ge I (L) 13 101 93ndash112 9897Ti I (L) 13 134 123ndash145 124127Ta I (L) 13 118 102ndash129 120120Ta I (W) 13 21 18ndash25 ndash19LEG I (

P) 13 571 538ndash601 576572

Cx II (L) 13 58 44ndash67 6865Tr II (L) 13 38 30ndash43 4443bFe II (L) 13 62 56ndash67 6260tFe II (L) 13 64 61ndash72 6052Ge II (L) 13 98 93ndash109 10298Ti II (L) 13 124 111ndash129 130128Ta II (L) 13 108 93ndash124 120123Ta II (W) 13 19 16ndash26 ndash16LEG II (

P) 13 553 524ndash578 586569

Cx III (L) 13 54 47ndash67 6881Tr III (L) 13 36 27ndash48 4434bFe III (L) 13 76 70ndash85 8474tFe III (L) 13 79 70ndash91 9085Ge III (L) 13 114 102ndash127 102102Ti III (L) 12 178 162ndash188 190187Ta III (L) 12 123 108ndash132 128117Ta III (W) 12 18 11ndash23 ndash21LEG III (

P) 12 659 630ndash683 706680

IP 12 1779 1692ndash1844 18681821Ti IAW 13 29 26ndash32 ndash25Ti IGe I 13 13 12ndash14 1313Ti IIPW 13 24 21ndash26 1919

(continued)

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 7

Table 1 Continued

Character D adonis Makol et Moniuszko sp nov D piae Haitlinger 2001

Samplesize

Mean(mm)

Range(mm)

Data for holotype (originaldescriptionpresent study)

Ti IIGe II 13 13 11ndash14 1313Ti IIIAW 12 39 35ndash42 ndash37Ti IIIGe III 12 16 15ndash18 1918Ti IIITi I 12 13 12ndash14 1515

AL length of anterior nonsensillary seta on scutum ASBa distance between the anterior margin of scutum and the level of anterior sensilla ASens length ofanterior sensillum on scutum AW distance between the bases of anterior nonsensillary setae on scutum bFehellip(L) length of basifemur Cxhellip(L) length of coxaDS length of dorsal idiosomal seta Gehellip(L) length of genu GL length of gnathosoma GW width of gnathosoma IL length of idiosoma ISD distance betweenthe level of anterior and posterior sensilla on scutum IW width of idiosoma L length of scutum ML length of nonsensillary seta of second pair on scutumMW distance between the bases of nonsensillary setae of second pair on scutum Odontus (L) length of palp tibial claw (odontus) Pa Fe (L) length of palpfemur Pa Ge (L) length of palp genu Pa Ta (L) length of palp tarsus Pa Ti (L) length of palp tibia Pa Tr (L) length of palp trochanter PL length of posteriornonsensillary seta on scutum PSBp distance between the posterior margin of scutum and the level of posterior sensilla PSens length of posterior sensillum onscutum PW distance between the bases of posterior nonsensillary setae of second pair on scutum SBa distance between the bases of anterior sensillaSBp distance between the bases of posterior sensilla Tahellip(L) length of tarsus Tahellip(W) width of tarsus tFehellip(L) length of telofemur Tihellip(L) length of tibiaTrhellip(L) length of trochanter VS length of ventral idiosomal seta W width of scutum 2a medial seta at the level of coxa II 3a medial seta at the level ofcoxa III

Figs 34ndash37 D adonis Makol et Moniuszko sp nov (34) Chelicerae (35) Anterior part of gnathosoma cheliceral sheath (36) Palp medialface (37) Palp lateral face

8 JOURNAL OF INSECT SCIENCE VOLUME 14

Figs 38ndash40 D adonis Makol et Moniuszko sp nov (38) Dorsal side of the body (legs omitted beyond trochanters) (39) Ventral side of thebody (legs omitted beyond trochanters) (40) Dorsal idiosomal seta

Figs 41 D adonis Makol et Moniuszko sp nov (41) Details of scutum

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 9

Figs 42ndash49 D adonis Makol et Moniuszko sp nov (42) Leg I (tibiamdashtarsus) (43) Leg I (trochantermdashgenu) (44) Leg II (tibiamdashtarsus) (45) LegII (trochantermdashgenu) (46) Leg III (tarsus) (47) Leg III (tibia) (48) Leg III (genu) (49) Leg III (trochantermdashtelofemur)

Table 2 Leg chaetotaxy of D adonis Makol et Moniuszko sp nov larvae

Leg segment Chaetotaxy

D adonis Makol et Moniuszko sp nov (N frac14 13) D piae Haitlinger 2001 data for holotype(original descriptionpresent study)

Cx I 1 n 1 supracoxal seta 1 n1 nTr I 1 n 1 n1 nbFe I 4 n 4 n4 ntFe I 5 n 5 n5 nGe I 12 n 1 s 1 k 12 n 1 s12 n 1 s 1 kTi I 17ndash19 n 2 j 1 z 1 k 15 n 2 j 1 k15 n 2 j 1 z 1 kTa I 22ndash26 n 2 z 1 o 1 e 24 n 2 z 1 o24 n 2 z 1 o 1 eCx II 2 n 2 n2 nTr II 1 n 1 n1 nbFe II 4 n 4 n4 ntFe II 5 n 5 n5 nGe II 12 n 1 k 12 n12 n 1 kTi II 15ndash19 n 2 j 14 n 2 j16 n 2 jTa II 23ndash28 n 1 z 1 o 21 n 1 z 1 o21 n 1 z 1 oCx III 2 n 2 n2 nTr III 1 n 1 n1 nbFe III 2 n 2 n2 ntFe III 5 n 5 n5 nGe III 12 n 12 n12 nTi III 18ndash20 n 1 j 15 n 1 j15 n 1 jTa III 24ndash28 n 1 z 23 n23 n 1 z

10 JOURNAL OF INSECT SCIENCE VOLUME 14

In many cases however the host is defined at the level of order whichhinders further conclusions on host specificity Only one species hith-erto recorded from Madagascar ie L (L) madagascariensis isknown to parasitize heteropteran bug Euchelichir longipes Jeannel1942 (Andre 1941) whereas for the remaining taxa the representativesof unidentified Odonata Orthoptera Neuroptera and Lepidoptera areexploited as host or the host remains unknown The limited knowledgeof host spectrum but also the various level of host specificity anticipatedfor erythraeid larvae (Wohltmann 2000 Łaydanowicz andMakol 2010Makol et al 2012b) make each record with precised specific affiliationof host and parasite seemmuch more important

Representatives of Dambullaeus are reported from Madagascar forthe first time All larvae were found attached to abdomen of the femalehost between tergites and sternites The only hitherto record concern-ing the distribution of the genus originated from Sri Lanka The distri-bution encompassing Sri Lanka and Madagascar set apart from eachother for c 4000 km may support the hypothesis on the Gondwananvicariance The time of splitting the IndoMadagascar subcontinent andseparation of Indiarsquos from Madagascar has been variably dated from

100 to 803 Ma (Yoder and Nowak 2006 and references therein) Thetransoceanic dispersal seems unlikely even when larvae constitutingthe instar with the highest dispersal potential facilitated by host areconcerned

Other erythraeid genera known from Madagascar ie AbrolophusBerlese 1891 Charletonia Oudemans 1910 Leptus Latreille 1796are represented in continental Africa and in India However these gen-era of world distribution comprise 114 115 and 275 nominal speciesrespectively (Makol and Wohltmann 2012 2013) and for all these gen-era the taxonomic revisions should precede the conclusions concerningbiogeographic patterns In this light the data concerning Dambullaeuscan constitute a contribution to the discussion on the origin ofMalagasy fauna and also on the age of main parasitengone taxa

Acknowledgments

We are grateful to Dr Ryszard Haitlinger for the loan of holotypeof Dambullaeus piae Haitlinger 2001 JM was partly supported bythe National Science Centre (grant in aid of research N N303-301737)

Fig 50 L nigrolineata Swierczewski et Stroinski sp nov and D adonis Makol et Moniuszko sp nov distribution map

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 11

Table 1 Continued

Character D adonis Makol et Moniuszko sp nov D piae Haitlinger 2001

Samplesize

Mean(mm)

Range(mm)

Data for holotype (originaldescriptionpresent study)

Ti IIGe II 13 13 11ndash14 1313Ti IIIAW 12 39 35ndash42 ndash37Ti IIIGe III 12 16 15ndash18 1918Ti IIITi I 12 13 12ndash14 1515

AL length of anterior nonsensillary seta on scutum ASBa distance between the anterior margin of scutum and the level of anterior sensilla ASens length ofanterior sensillum on scutum AW distance between the bases of anterior nonsensillary setae on scutum bFehellip(L) length of basifemur Cxhellip(L) length of coxaDS length of dorsal idiosomal seta Gehellip(L) length of genu GL length of gnathosoma GW width of gnathosoma IL length of idiosoma ISD distance betweenthe level of anterior and posterior sensilla on scutum IW width of idiosoma L length of scutum ML length of nonsensillary seta of second pair on scutumMW distance between the bases of nonsensillary setae of second pair on scutum Odontus (L) length of palp tibial claw (odontus) Pa Fe (L) length of palpfemur Pa Ge (L) length of palp genu Pa Ta (L) length of palp tarsus Pa Ti (L) length of palp tibia Pa Tr (L) length of palp trochanter PL length of posteriornonsensillary seta on scutum PSBp distance between the posterior margin of scutum and the level of posterior sensilla PSens length of posterior sensillum onscutum PW distance between the bases of posterior nonsensillary setae of second pair on scutum SBa distance between the bases of anterior sensillaSBp distance between the bases of posterior sensilla Tahellip(L) length of tarsus Tahellip(W) width of tarsus tFehellip(L) length of telofemur Tihellip(L) length of tibiaTrhellip(L) length of trochanter VS length of ventral idiosomal seta W width of scutum 2a medial seta at the level of coxa II 3a medial seta at the level ofcoxa III

Figs 34ndash37 D adonis Makol et Moniuszko sp nov (34) Chelicerae (35) Anterior part of gnathosoma cheliceral sheath (36) Palp medialface (37) Palp lateral face

8 JOURNAL OF INSECT SCIENCE VOLUME 14

Figs 38ndash40 D adonis Makol et Moniuszko sp nov (38) Dorsal side of the body (legs omitted beyond trochanters) (39) Ventral side of thebody (legs omitted beyond trochanters) (40) Dorsal idiosomal seta

Figs 41 D adonis Makol et Moniuszko sp nov (41) Details of scutum

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 9

Figs 42ndash49 D adonis Makol et Moniuszko sp nov (42) Leg I (tibiamdashtarsus) (43) Leg I (trochantermdashgenu) (44) Leg II (tibiamdashtarsus) (45) LegII (trochantermdashgenu) (46) Leg III (tarsus) (47) Leg III (tibia) (48) Leg III (genu) (49) Leg III (trochantermdashtelofemur)

Table 2 Leg chaetotaxy of D adonis Makol et Moniuszko sp nov larvae

Leg segment Chaetotaxy

D adonis Makol et Moniuszko sp nov (N frac14 13) D piae Haitlinger 2001 data for holotype(original descriptionpresent study)

Cx I 1 n 1 supracoxal seta 1 n1 nTr I 1 n 1 n1 nbFe I 4 n 4 n4 ntFe I 5 n 5 n5 nGe I 12 n 1 s 1 k 12 n 1 s12 n 1 s 1 kTi I 17ndash19 n 2 j 1 z 1 k 15 n 2 j 1 k15 n 2 j 1 z 1 kTa I 22ndash26 n 2 z 1 o 1 e 24 n 2 z 1 o24 n 2 z 1 o 1 eCx II 2 n 2 n2 nTr II 1 n 1 n1 nbFe II 4 n 4 n4 ntFe II 5 n 5 n5 nGe II 12 n 1 k 12 n12 n 1 kTi II 15ndash19 n 2 j 14 n 2 j16 n 2 jTa II 23ndash28 n 1 z 1 o 21 n 1 z 1 o21 n 1 z 1 oCx III 2 n 2 n2 nTr III 1 n 1 n1 nbFe III 2 n 2 n2 ntFe III 5 n 5 n5 nGe III 12 n 12 n12 nTi III 18ndash20 n 1 j 15 n 1 j15 n 1 jTa III 24ndash28 n 1 z 23 n23 n 1 z

10 JOURNAL OF INSECT SCIENCE VOLUME 14

In many cases however the host is defined at the level of order whichhinders further conclusions on host specificity Only one species hith-erto recorded from Madagascar ie L (L) madagascariensis isknown to parasitize heteropteran bug Euchelichir longipes Jeannel1942 (Andre 1941) whereas for the remaining taxa the representativesof unidentified Odonata Orthoptera Neuroptera and Lepidoptera areexploited as host or the host remains unknown The limited knowledgeof host spectrum but also the various level of host specificity anticipatedfor erythraeid larvae (Wohltmann 2000 Łaydanowicz andMakol 2010Makol et al 2012b) make each record with precised specific affiliationof host and parasite seemmuch more important

Representatives of Dambullaeus are reported from Madagascar forthe first time All larvae were found attached to abdomen of the femalehost between tergites and sternites The only hitherto record concern-ing the distribution of the genus originated from Sri Lanka The distri-bution encompassing Sri Lanka and Madagascar set apart from eachother for c 4000 km may support the hypothesis on the Gondwananvicariance The time of splitting the IndoMadagascar subcontinent andseparation of Indiarsquos from Madagascar has been variably dated from

100 to 803 Ma (Yoder and Nowak 2006 and references therein) Thetransoceanic dispersal seems unlikely even when larvae constitutingthe instar with the highest dispersal potential facilitated by host areconcerned

Other erythraeid genera known from Madagascar ie AbrolophusBerlese 1891 Charletonia Oudemans 1910 Leptus Latreille 1796are represented in continental Africa and in India However these gen-era of world distribution comprise 114 115 and 275 nominal speciesrespectively (Makol and Wohltmann 2012 2013) and for all these gen-era the taxonomic revisions should precede the conclusions concerningbiogeographic patterns In this light the data concerning Dambullaeuscan constitute a contribution to the discussion on the origin ofMalagasy fauna and also on the age of main parasitengone taxa

Acknowledgments

We are grateful to Dr Ryszard Haitlinger for the loan of holotypeof Dambullaeus piae Haitlinger 2001 JM was partly supported bythe National Science Centre (grant in aid of research N N303-301737)

Fig 50 L nigrolineata Swierczewski et Stroinski sp nov and D adonis Makol et Moniuszko sp nov distribution map

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 11

Figs 38ndash40 D adonis Makol et Moniuszko sp nov (38) Dorsal side of the body (legs omitted beyond trochanters) (39) Ventral side of thebody (legs omitted beyond trochanters) (40) Dorsal idiosomal seta

Figs 41 D adonis Makol et Moniuszko sp nov (41) Details of scutum

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 9

Figs 42ndash49 D adonis Makol et Moniuszko sp nov (42) Leg I (tibiamdashtarsus) (43) Leg I (trochantermdashgenu) (44) Leg II (tibiamdashtarsus) (45) LegII (trochantermdashgenu) (46) Leg III (tarsus) (47) Leg III (tibia) (48) Leg III (genu) (49) Leg III (trochantermdashtelofemur)

Table 2 Leg chaetotaxy of D adonis Makol et Moniuszko sp nov larvae

Leg segment Chaetotaxy

D adonis Makol et Moniuszko sp nov (N frac14 13) D piae Haitlinger 2001 data for holotype(original descriptionpresent study)

Cx I 1 n 1 supracoxal seta 1 n1 nTr I 1 n 1 n1 nbFe I 4 n 4 n4 ntFe I 5 n 5 n5 nGe I 12 n 1 s 1 k 12 n 1 s12 n 1 s 1 kTi I 17ndash19 n 2 j 1 z 1 k 15 n 2 j 1 k15 n 2 j 1 z 1 kTa I 22ndash26 n 2 z 1 o 1 e 24 n 2 z 1 o24 n 2 z 1 o 1 eCx II 2 n 2 n2 nTr II 1 n 1 n1 nbFe II 4 n 4 n4 ntFe II 5 n 5 n5 nGe II 12 n 1 k 12 n12 n 1 kTi II 15ndash19 n 2 j 14 n 2 j16 n 2 jTa II 23ndash28 n 1 z 1 o 21 n 1 z 1 o21 n 1 z 1 oCx III 2 n 2 n2 nTr III 1 n 1 n1 nbFe III 2 n 2 n2 ntFe III 5 n 5 n5 nGe III 12 n 12 n12 nTi III 18ndash20 n 1 j 15 n 1 j15 n 1 jTa III 24ndash28 n 1 z 23 n23 n 1 z

10 JOURNAL OF INSECT SCIENCE VOLUME 14

In many cases however the host is defined at the level of order whichhinders further conclusions on host specificity Only one species hith-erto recorded from Madagascar ie L (L) madagascariensis isknown to parasitize heteropteran bug Euchelichir longipes Jeannel1942 (Andre 1941) whereas for the remaining taxa the representativesof unidentified Odonata Orthoptera Neuroptera and Lepidoptera areexploited as host or the host remains unknown The limited knowledgeof host spectrum but also the various level of host specificity anticipatedfor erythraeid larvae (Wohltmann 2000 Łaydanowicz andMakol 2010Makol et al 2012b) make each record with precised specific affiliationof host and parasite seemmuch more important

Representatives of Dambullaeus are reported from Madagascar forthe first time All larvae were found attached to abdomen of the femalehost between tergites and sternites The only hitherto record concern-ing the distribution of the genus originated from Sri Lanka The distri-bution encompassing Sri Lanka and Madagascar set apart from eachother for c 4000 km may support the hypothesis on the Gondwananvicariance The time of splitting the IndoMadagascar subcontinent andseparation of Indiarsquos from Madagascar has been variably dated from

100 to 803 Ma (Yoder and Nowak 2006 and references therein) Thetransoceanic dispersal seems unlikely even when larvae constitutingthe instar with the highest dispersal potential facilitated by host areconcerned

Other erythraeid genera known from Madagascar ie AbrolophusBerlese 1891 Charletonia Oudemans 1910 Leptus Latreille 1796are represented in continental Africa and in India However these gen-era of world distribution comprise 114 115 and 275 nominal speciesrespectively (Makol and Wohltmann 2012 2013) and for all these gen-era the taxonomic revisions should precede the conclusions concerningbiogeographic patterns In this light the data concerning Dambullaeuscan constitute a contribution to the discussion on the origin ofMalagasy fauna and also on the age of main parasitengone taxa

Acknowledgments

We are grateful to Dr Ryszard Haitlinger for the loan of holotypeof Dambullaeus piae Haitlinger 2001 JM was partly supported bythe National Science Centre (grant in aid of research N N303-301737)

Fig 50 L nigrolineata Swierczewski et Stroinski sp nov and D adonis Makol et Moniuszko sp nov distribution map

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 11

Figs 42ndash49 D adonis Makol et Moniuszko sp nov (42) Leg I (tibiamdashtarsus) (43) Leg I (trochantermdashgenu) (44) Leg II (tibiamdashtarsus) (45) LegII (trochantermdashgenu) (46) Leg III (tarsus) (47) Leg III (tibia) (48) Leg III (genu) (49) Leg III (trochantermdashtelofemur)

Table 2 Leg chaetotaxy of D adonis Makol et Moniuszko sp nov larvae

Leg segment Chaetotaxy

D adonis Makol et Moniuszko sp nov (N frac14 13) D piae Haitlinger 2001 data for holotype(original descriptionpresent study)

Cx I 1 n 1 supracoxal seta 1 n1 nTr I 1 n 1 n1 nbFe I 4 n 4 n4 ntFe I 5 n 5 n5 nGe I 12 n 1 s 1 k 12 n 1 s12 n 1 s 1 kTi I 17ndash19 n 2 j 1 z 1 k 15 n 2 j 1 k15 n 2 j 1 z 1 kTa I 22ndash26 n 2 z 1 o 1 e 24 n 2 z 1 o24 n 2 z 1 o 1 eCx II 2 n 2 n2 nTr II 1 n 1 n1 nbFe II 4 n 4 n4 ntFe II 5 n 5 n5 nGe II 12 n 1 k 12 n12 n 1 kTi II 15ndash19 n 2 j 14 n 2 j16 n 2 jTa II 23ndash28 n 1 z 1 o 21 n 1 z 1 o21 n 1 z 1 oCx III 2 n 2 n2 nTr III 1 n 1 n1 nbFe III 2 n 2 n2 ntFe III 5 n 5 n5 nGe III 12 n 12 n12 nTi III 18ndash20 n 1 j 15 n 1 j15 n 1 jTa III 24ndash28 n 1 z 23 n23 n 1 z

10 JOURNAL OF INSECT SCIENCE VOLUME 14

In many cases however the host is defined at the level of order whichhinders further conclusions on host specificity Only one species hith-erto recorded from Madagascar ie L (L) madagascariensis isknown to parasitize heteropteran bug Euchelichir longipes Jeannel1942 (Andre 1941) whereas for the remaining taxa the representativesof unidentified Odonata Orthoptera Neuroptera and Lepidoptera areexploited as host or the host remains unknown The limited knowledgeof host spectrum but also the various level of host specificity anticipatedfor erythraeid larvae (Wohltmann 2000 Łaydanowicz andMakol 2010Makol et al 2012b) make each record with precised specific affiliationof host and parasite seemmuch more important

Representatives of Dambullaeus are reported from Madagascar forthe first time All larvae were found attached to abdomen of the femalehost between tergites and sternites The only hitherto record concern-ing the distribution of the genus originated from Sri Lanka The distri-bution encompassing Sri Lanka and Madagascar set apart from eachother for c 4000 km may support the hypothesis on the Gondwananvicariance The time of splitting the IndoMadagascar subcontinent andseparation of Indiarsquos from Madagascar has been variably dated from

100 to 803 Ma (Yoder and Nowak 2006 and references therein) Thetransoceanic dispersal seems unlikely even when larvae constitutingthe instar with the highest dispersal potential facilitated by host areconcerned

Other erythraeid genera known from Madagascar ie AbrolophusBerlese 1891 Charletonia Oudemans 1910 Leptus Latreille 1796are represented in continental Africa and in India However these gen-era of world distribution comprise 114 115 and 275 nominal speciesrespectively (Makol and Wohltmann 2012 2013) and for all these gen-era the taxonomic revisions should precede the conclusions concerningbiogeographic patterns In this light the data concerning Dambullaeuscan constitute a contribution to the discussion on the origin ofMalagasy fauna and also on the age of main parasitengone taxa

Acknowledgments

We are grateful to Dr Ryszard Haitlinger for the loan of holotypeof Dambullaeus piae Haitlinger 2001 JM was partly supported bythe National Science Centre (grant in aid of research N N303-301737)

Fig 50 L nigrolineata Swierczewski et Stroinski sp nov and D adonis Makol et Moniuszko sp nov distribution map

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 11

In many cases however the host is defined at the level of order whichhinders further conclusions on host specificity Only one species hith-erto recorded from Madagascar ie L (L) madagascariensis isknown to parasitize heteropteran bug Euchelichir longipes Jeannel1942 (Andre 1941) whereas for the remaining taxa the representativesof unidentified Odonata Orthoptera Neuroptera and Lepidoptera areexploited as host or the host remains unknown The limited knowledgeof host spectrum but also the various level of host specificity anticipatedfor erythraeid larvae (Wohltmann 2000 Łaydanowicz andMakol 2010Makol et al 2012b) make each record with precised specific affiliationof host and parasite seemmuch more important

Representatives of Dambullaeus are reported from Madagascar forthe first time All larvae were found attached to abdomen of the femalehost between tergites and sternites The only hitherto record concern-ing the distribution of the genus originated from Sri Lanka The distri-bution encompassing Sri Lanka and Madagascar set apart from eachother for c 4000 km may support the hypothesis on the Gondwananvicariance The time of splitting the IndoMadagascar subcontinent andseparation of Indiarsquos from Madagascar has been variably dated from

100 to 803 Ma (Yoder and Nowak 2006 and references therein) Thetransoceanic dispersal seems unlikely even when larvae constitutingthe instar with the highest dispersal potential facilitated by host areconcerned

Other erythraeid genera known from Madagascar ie AbrolophusBerlese 1891 Charletonia Oudemans 1910 Leptus Latreille 1796are represented in continental Africa and in India However these gen-era of world distribution comprise 114 115 and 275 nominal speciesrespectively (Makol and Wohltmann 2012 2013) and for all these gen-era the taxonomic revisions should precede the conclusions concerningbiogeographic patterns In this light the data concerning Dambullaeuscan constitute a contribution to the discussion on the origin ofMalagasy fauna and also on the age of main parasitengone taxa

Acknowledgments

We are grateful to Dr Ryszard Haitlinger for the loan of holotypeof Dambullaeus piae Haitlinger 2001 JM was partly supported bythe National Science Centre (grant in aid of research N N303-301737)

Fig 50 L nigrolineata Swierczewski et Stroinski sp nov and D adonis Makol et Moniuszko sp nov distribution map

2014 MAKOL ET AL DESCRIPTION OF NEW SPECIES FROM WESTERN MADAGASCAR 11

References CitedAndre M 1941 Sur une nouvelle forme larvaire drsquoAcarien parasite drsquoun

Hemiptere de Madagascar Revue Francaise drsquoentomologie 8 188ndash195Andre M 1946 Un Erythraeus (Acarien) nouveau recueilli a Madagascar

(E Milloti n sp) Bulletin du Museum drsquohistoire naturelle Paris 2e ser 18268ndash269

Berlese A 1891 Acari Myriapoda et Scorpiones hucusque in Italia repertaPadova Fasc 59

Bourgoin T 1988 A new interpretation of the homologies of the Hemipteramale genitalia illustrated by the Tettigometridae (HemipteraFulgoromorpha) pp 113ndash120 In C Vidano and A Arzone (eds) 6thAuchenorrhyncha Meeting 7ndash11 September 1987 Turin Italy CN R-IPRATurin Italy

Bourgoin T 1993 Female genitalia in Hemiptera Fulgoromorphamorphological and phylogenetic data Ann Soc Entomol Fr (NS) 29225ndash244

Bourgoin T 2013 FLOW (fulgoromorpha lists on the web) a knowledge anda taxonomy database dedicated to planthoppers (Insecta HemipteraFulgoromorpha Fulgoromorpha) Version 8 updated (3 November 2013)(httphemiptera-databasesorgflow)

Bourgoin T and J Huang 1990 Morphologie compare des genitaliamales des Trypetimorphini et remarques phylogenetiques (HemipteraFulgoromorpha Tropiduchidae Ann Soc Entomol Fr (NS) 26 555ndash564

Carayon J 1969 Emploi du noir chlorazol en anatomie microscopique desinsectes Ann Soc Entomol Fr (NS) 5 179ndash193

Ganzhorn J U P P Lowry II G E Schatz and S Sommer 2001 Thebiodiversity of Madagascar one the worldrsquos hottest hotspots on its way outOryx 35 346ndash348

Haitlinger R 1987 Larval Erythraeidae (Acari Prostigmata) fromMadagascar Polskie Pismo Entomol 57 701ndash723

Haitlinger R 2001 Dambullaeus piae genn spn (Acari ProstigmataErythraeidae) from Sri Lanka Zeszyty Naukowe Akademii Rolniczej weWrocławiu Seria Zootechnika XLVIII 429 51ndash54

Haitlinger R 2007 New records of mites (Acari Prostigmata Erythraeidae)from Africa with descriptions of four new species Zeszyty NaukoweUniwersytetu Przyrodniczego we Wrocławiu Seria Biologia i HodowlaZwierzat LV 559 55ndash69

Karsch F 1890 Afrikanische Fulgoriden Berliner Entomol Zeitschrift 3557ndash70

Latreille P A 1796 Precis de caracteres generiques des Insectes disposes dansun ordre naturel Prevot Paris xiiithorn201 pp

Łaydanowicz J and J Makol 2010 Correlation of heteromorphic life instarsin terrestrial Parasitengona mites and its impact on taxonomy ndash the case ofLeptus molochinus (C L Koch 1837) and Leptus ignotus (Oudemans 1903)(Acari Trombidiformes Prostigmata Erythraeidae) J Nat Hist 44669ndash697

Makol J and A Wohltmann 2012 An annotated checklist of terrestrialParasitengona (Actinotrichida Prostigmata) of the World excludingTrombiculidae and Walchiidae Ann Zool 62 359ndash562

Makol J and AWohltmann 2013 Corrections and additions to the checklistof terrestrial Parasitengona (Actinotrichida Prostigmata) of the World ex-cluding Trombiculidae and Walchiidae Ann Zool 63 15ndash27

Makol J M Felska H Moniuszko and G Zalesny 2012a Redescriptionof Leptus kattikus Haitlinger 2009 (Actinotrichida ParasitengonaErythraeidae) and molecular identification of its host from DNA barcodingZootaxa 3569 67ndash78

Makol J A Kłosinska and J Łaydanowicz 2012b Hostndashparasite interac-tions within terrestrial Parasitengona (Acari Trombidiformes Prostigmata)Int J Acarol 38 18ndash22

McNeely J A K R Miller W V Reid R A Mittermeier and T BWerner 1990 Conserving the worldrsquos biological diversity InternationalUnion for the Conservation of Nature and Natural Resources WorldResources Institute Conservation International WWF mdash US and WorldBanks Gland 193 pp

Melichar L 1901 Monographie der Acanaloniiden und Flatiden (Homoptera)(Fortsetzung) Ann des kk Naturhistorischen Hofmuseums Wien 16178ndash258

Myers N R A Mittermeier C G Mittermeier GAB da Fonseca and JKent 2000 Biodiversity hotspots for conservation priorities Nature 403853ndash858

OrsquoBrien L 2002 The wild wonderful world of Fulgoromorpha Denisia (NF)4 83ndash102

OrsquoBrien L and S W Wilson 1985 Planthopper systematics and externalmorphology pp 61ndash102 In L R Nault and J G Rodriguez (eds) Theleafhoppers and planthoppers JohnWiley amp Sons New York NY

Oudemans A C 1910 Acarologische Aanteekeningen XXXI EntomolBerichten 3 43ndash51

Signoret V 1860 Faune des hemipteres de Madagascar 1ere partieHomopteres Ann Soc Entomol Fr (Ser 3) 8 177ndash206

Southcott R V 1948 Larval Smarididae (Acarina) from Australia and NewGuinea Proc Linn Soc N S W 72 252ndash264

Southcott R V 1961 Studies on the systematics and biology of theErythraeoidea (Acarina) with a critical revision of the genera and subfami-lies Aust J Zool 9 367ndash610

Stal C 1866 Hemiptera Homoptera Latr Hemiptera Africana 4 1ndash276Storey M J J Mahoney A D Sounders R A Duncan S P Kelley and

M F Coffin 1995 Timing of hot spot-related volcanism and the breakup ofMadagascar and India Science 267 852ndash855

Stroinski A M Felska and J Makol 2013 Host-parasite associationsbetween Hemiptera (Insecta) and Erythraeidae (Trombidiformes) Ann Zool63 195ndash221

Swierczewski D and A Stroinski 2012 A new species of the genus LatoisStal 1866 from Madagascar (Hemiptera Fulgoromorpha Flatidae) ActaZool Cracoviensia 55 65ndash77

Swierczewski D and A Stroinski 2013 Madagascar Flatidae (HemipteraFulgoromorpha) state-of-the-art and research challenges pp 293ndash301 In APopov S Grozeva N Simov and E Tasheva (eds) Advances in hemi-pterology Pensoft Publishers Sofia vol 319

Szwedo J and D _Zyła 2009 New Fulgoridiidae genus from the UpperJurassic Karabastau deposits Kazakhstan (Hemiptera FulgoromorphaFulgoroidea) Zootaxa 2281 40ndash52

Tragardh I 1908 Arachnoidea Acari Wissenschaftliche Ergebnisse derSchwedischen Zoologischen Expedition nach dem Kilimandjaro dem Meruund den Umgebenden Massaisteppen Deutsch-Ostafrikas 1905ndash1906 unterLeitung von Prof Dr Yngve Sjostedt 20 31ndash57

Vences M K C Wollenberg D R Vieites and D C Lees 2009Madagascar as a model region of species diversification Trends Ecol Evol24 456ndash465

Walter D E and G W Krantz 2009 Collection rearing and preparingspecimens pp 83ndash96 In G W Krantz and D E Walter (eds) A manualof acarology 3rd ed Texas Tech University Press Lubbock

Wohltmann A 2000 The evolution of life histories in Parasitengona (AcariProstigmata) Acarologia 41 145ndash204 [2001]

Yoder A D and N D Nowak 2006Has vicariance or dispersal been the pre-dominant biogeographic force in Madagascar Only time will tell AnnuRev Ecol Evol Syst 37 405ndash431

Received 21 April 2013 accepted 30 November 2013

12 JOURNAL OF INSECT SCIENCE VOLUME 14