First Known Neotropical Species Of Brunettia (Diptera: Psychodidae) From Southeastern Brazil, With Description Of A New Subgenus And Systematic Comments On The Genus

10 Pages·1995·4 MB·English

by  Freddy Bravo

FIRST KNOWN NEOTROPICAL SPECIES OF BRUNETTIA (DIPTERA:
PSYCHODIDAE) FROM SOUTHEASTERN BRAZIL, WITH DESCRIPTION
OF A NEW SUBGENUS AND SYSTEMATIC COMMENTS ON THE GENUS
1
Freddy Bravo23
Dalton de Souza Amorim2,4
ABSTRACT
Brunettiacaipira,sp.n.isdescribedfromsoutheasternBrazil.Thisisthefirstdescribedspeciesofthe
genusintheNeotropicalregion. Anewsubgenus,Brunettia(Neobrunettia) isproposedforthisspecies.The
remainingspeciesofthegenusarerestrictedtotheOriental,Afrotropical,Australian,andPalearcticregions,
placedin sevensubgenera. Ananalysisoftherelationships amongthe subgeneraofBrunettia isproceeded,
whichresults in apartial Solution forthephylogenetic relationships among the subgenera. B. (Mirousiella)
appears as the sister-group of the set of the remaining subgenera; B. (Campanulobrunettia) composes a
monophyletic arrangement with B. (Maurobrunettia); B. (Campanulobrunettia), B. (Maurobrunettia), B.
{Brunettia),andB.(Neobrunettia) standasamonophyleticunity.Nosynapomorphywasfoundforthetypical
subgenus, B. (Brunettia), which most probably does not referto a monophyletic arrangement; a polytomy
includingB.(Atrichobrunettia),B.(Plesiobrunettia),B.(Horobrunettia),andthegroupofsubgeneraincluding
B. (Brunettia)remainsunresolved. Also,no synapomorphywasfoundforB. (Mirousiella). The pantropical
distributionforthisgroupsuggeststhatithasatleastaGondwanicorigin.
KEYWORDS.Neotropical,Diptera,Psychodidae,Brunettia,Systematics.
INTRODUCTION
ThegenusBrunettiaAnnandale, 191 asredefinedbyDUCKHOUSE(1966)most
certainlycorrespondtoamonophyletictaxon.Theabsenceofsetaeonthegonocoxiteand
onthetergite9 seemstobe synapomorphicforthegenus. Thepresenceofthese setaein
five species included by DUCKHOUSE (1991) in the subgenus B. {Horobrunettia)
Duckhouse, 1991 is interpreted by him as a secondary acquisition. Other six subgenera
were recognized by DUCKHOUSE (1991): Brunettia {Brunettia) Annandale, 1910, B.
{Atrichobrunettia) Satchell, 1953, B. {Campanulobrunettia) Duckhouse, 1991, B.
{Maurobrunettia) Duckhouse, 1991, B. {Mirousiella) (Vaillant, 1974), and B.
{Plesiobrunettia) Duckhouse, 1991. The monophyletism of these subgenera was not
demonstrated. Thephylogenetic relationships among the subgenera ofBrunettia were
also not determined to date.
Following DUCKHOUSE (1991), the genus Brunettia is absent only from the
1.Contributionn°794,DeptodeZoologia,UFPR.
2.Depto.deBiologia;FaculdadedeFilosofia,CiênciaseLetrasdeRibeirãoPreto/USP;Av.Bandeirantes3900; 14040-901 RibeirãoPretoSP:
Brazil.
3.CAPESfellowship.
4.ResearcherfellowshipofCNPq;partiallymadeunderFAPESPgrant93/0954-9.
Iheringia,Sér.ZooL,PortoAlegre,(79): 149- 158,29 set. 1995
O
,
BRAVO&AMORIM
15
Nearcticregion.TherearesomefewspeciesknownfromsomePacificislands,Japan,and
NewZealand. Mostspeciesdescribedarefoundiníndia,Papua-NewGuinea, Southeast
Ásia,andAustrália.DUCKHOUSE(1991)referredtotwoNeotropicalspecies,butdidnot
describedthem. Anew speciesoíBrunettiafromsoutheasternBrazilisdescribed,based
on males and females, whichis place in anew subgenus,Neobrunettia. Aphylogenetic
analysisisperformedtodeterminedthephylogeneticrelationshipsamongthesubgenera
ofthe genus.
Theterminologyforthethoracicpleural sclerites isreinterpretedbasedonamore
general analysis ofthePsychodomorphamorphology. Terminology formaleterminalia
morphology follows McALPINE (1981) and for female terminalia follows SAETHER
(1977). ThenamesforotherstructuresfollowDUCKHOUSE(1991).Typesaredeposited
attheMuseudeZoologia,UniversidadedeSãoPaulo,SãoPaulo(MZSP).Someparatypes
willbesenttoTheNaturalHistoryMuseum(BMNH).Alimeasurementsareinmillime-
ters.
Brunettia {Neobrunettia), subgen. n.
Typespecies:Brunettia(Neobrunettia)caipira,sp.n.,heredesignated.
This subgenus differs from the remaining subgenera by the following features
(whicharealsodiagnosticforthespecies): subcostalvein(Sc) absent,onlyahairlineof
unsclerotized cuticula (hj) seen under microscope. K diffuse and wide basally, with a
x
ruptureinthebasalfourth.
Brunettia caipira, sp. n.
(Figs. 1-18)
Typelocality:Brazil,StateofSãoPaulo,RibeirãoPreto,CampusdaUniversidadedeSãoPaulo.
Holotype:male,onslide-mounting,withthelabel:Brasil,StateofSãoPaulo,RibeirãoPreto,26.VI.1993,
FreddyBravocol. (MZSP); allotype:female,samedataofholotype(onslide-mounWting)(MZSP).Paratypes:
BRAZIL,RiodeJaneiro:RiodeJaneiro,RepresaRioGrande,22s50'S;43s00' [Malaisetrap], ld, 109,
II.W1968W,M.Alvarengacol.(MZSP);ld,19,IX.1969(MZSP);ld,19,IV.1972(MZSP).SilvaJardim,22WS;
42 [Malaisetrap],19,VII.1974,F.M.Oliveiracol.(8paratypesonslide-mountings)(MZSP).SãoPaulo:
RibeirãoPreto(CampusofUSP),21s15'S;48900'W, ld,29,26.VI.1993,FreddyBravocol.(MZSP); ld,
29.IV.1993(MZSP); 19,03.IH.1993(MZSP);2c?, 19, 10.VII.1993(MZSP); ld,06.VIII.1993(MZSP); ld
27.VIII.1993 (MZSP); ld,39, 19.111.1994(MZSP); ld,22.in.1994(MZSP).
Diagnosis.Subcostalvein(Sc)absent,onlyahairlineofunsclerotizedcuticula(h2)
seenundermicroscope.Rjdiffuseandwidebasally,witharuptureinthebasalfourth.Post-
hypandriumplatedeveloped, covering aedeagus.
Description.Male.Tegumentblack;alarmembranehyaline-black.Body,alarveins
and alarmembrane covered withblack setae. Frons with apatch ofwhite setae. Length
from thorax anteriorlytoposteriorendofabdómen, 2.19.
mm
Head: subcircular,0.38 longfromvertextoclypeusdistally; distancebetween
eyesatmost0.37.Acompleteeye-bridgeaboveantennae(figs. 1,2).Palpus 1.0-3.4-4.0-
4.4. Antenna with 15 articles, 1.19 long; distal flagellomeres shorter that basal ones;
Iheringia,Sér.Zool.,PortoAlegre,(79): 149- 158,29 set. 1995
FirstknownNeotropicalspeciesofBrunettia. 51
flagellomeres eccentric, except for the last two; last flagellomere with apicule; ascoid
paired,C-shaped(fig. 7). Labellafleshy, with6largersetaedistributedondorsalmargin
andsomeshorteronesventrally;probablylabela 1 and2fused(fig.5).Maxillasmall and
round (fig. 4). Cibarium with strong walls (fig. 6).
Thorax: 0.73 long. Episternum I ventrally fused to antero-dorsal extension of
katepisternum.EpimeronIincludingthespiracle,withananteriorprotuberance.Anterior
spiracle withmembrane around. Stigmatic porevisible (fig. 8).
Wing:length, 1.87.Maximumwidth,0.88.Scabsent.Rspectinate.R R andCuA,
wellsclerotized(notsoclearforR inthedrawing). R reaching wing on a1?pex5., Humeral
x 5
and analregions slightly developed (fig. 10).
Abdomen: length, 1.46. Tergites and sternites mostly black, with a lighterregion
anteriorly.
Terminalia (figs. 15-18): length, 0.20; width, 0.25. Tergite 9 and gonocoxites
without setae, gonostyles with only two distai setae. Cerei with apical retinacula, distai
oneslongerwithfringedtips,andtheothers,morebasally,shortandstraigth(figs. 16-17).
Tergite9withtwosmallaperturebasally(fig. 18). Sternite9withapost-hypandrialplate
coveringaedeagus(fig. 15). Sternite 10withmicropilosity(fig. 18).Aedeagussymmetri-
cal, apicallybifid. Parameres divergent.
Female. Similartomaleexceptasfollows. Length,fromthoraxanteriorlytoendof
abdómen2.27 (fig. 12). EpimeronIwithoutananteriorprotuberance (fig. 9). Eyebridge
incomplete(fig.3).Winglength1.82,width0.80;humeralandanallobesseeninthemales
absent(fig. 11).Gonocoxite8coveringsternite9(fig. 14).Cercuswithamicrosetosebasal
protuberance (fig. 13).
Distribution: Brazil, States ofRio deJaneiro and São Paulo.
Philogenetic analysis of Brunettia
Thefollowing features,withconditionfortheterminaltaxa synthesized intable I,
wereobtainedinananalysisoftheintragenericrelationshipsofBrunettia.Thegenerality
ofeachcharacterandtheconclusionsconcerningtherelationshipsamongthe subgeneric
taxa of Brunettia are in figure19. The analysis was made aecording to the Hennigian
argumentation(HENNIG, 1966; AMORIM, 1994a). The mathrix was also submittedto
anumerical analysis using the Hennig86program, ie* Option.
TableI.Charactermathrixofthephylogeneticanalysisofthephylogeneticrelationshipsamongthesubgeneric
taxaofBrunettia.Thesequenceofcharactersfollowthatinthelistandinthecladogram.
outgroup 00000 00000 00000 0000
Mirousiella 11000 10000 00011 1000
Atrichobrunettia 11000 11000 00011 1100
uni
Neobrunettia 11000 10110 1101
"Brunettia" 11000 10000 11011 1101
Campanulobrunettia 11100 10001 11011 1101
Maurobrunettia 11011 10001 11011 1101
Plesiobrunettia 11100 10000 00011 1100
Horobrunettia 11000 10000 00010 1110
Iheringia,Ser.Zool.,PortoAlegre.(79): 149- 158,29 set. 1995
BRAVO&AMORIM
152
Figs. 1-11.Brunettiacaipirasp.n.Cephalicstructures: 1-2,5-7holotyped_;3allotype. 9.Head: 1. anterior,
2, posterior, 3, anterior; 4,maxila(diagramatic); 5,labrum, lateral; 6,hypopharynx; 7, antennae, lastthree
Segments.Thoracicstructures: 8.lateral(dparatype,RibeirãoPreto);9.frontal(9paratype,RibeirãoPreto).
Wing (rigth): 10, d paratype, Ribeirão Preto; 11, 9 paratype, Ribeirão Preto. (Am, anepimeron; AnP,
antepronotum;As,anepisternum;Cs,cardo-stipes;CvS,cervicalsclerite;Ep,epimeronI;Es,episternumI;Kt,
katepisternum;La,lacinia;Lb,labella1+2;Me,meronII;Mn,metanotum;Mp,metapleuron;Pa,maxillarpalpus;
PnP,postnotalphragma;PsP,postpronotum). Scales:A,0.20mm(1-3);B,0.20mm(5); C,0.50mm(8-9);D,
0.20mm(6-7);E, l.OOmrn(10-11).
Iheringia,Sér.Zool.,PortoAlegre,(79): 149- 158,29 set. 1995
FirstknownNeotropicalspeciesofBrunettia... 53
Figs. 12-18. Brunettiacaipirasp.n. 12, 9paratype,RibeirãoPreto,lateral. 13.cercus.2 paratype.Ribeirão
Preto, lateral. 14, 9 allotype, terminalia, dorsal. Terminalia of c holotype: 15, S9, aedeagus, parameres,
gonocoxitesandgonostiles; 16,cercustip; 17,terminalia,lateral; 18,T9,S10andcercus.(Ae,aedeagus:Ce.
cercus;Gc,gonocoxite;Gs,gonostylus;Pa,paramere;S8,sternite8;S9,sternite9;S10,sternite 10;T9,tergite
9).Scales:A,0.10mm(13);B,0.10mm(14);C,0.10mm(15-18); D, 1.00mm(12).
Iheringia.Sér.Zool.,PortoAlegre.(79): 149- 158.29 set. 1995
BRAVO&AMORIM
154
$
^3
ss; <ay > I
K
*$ s£t I I
O
a *Ca) 8 |
-SS
'? •Si S
*>
ti ***>
11,12,19
17
,2,6,14,15,16
Fig. 19.CladogramoíBrunettia,depictingthephylogeneticrelationshipsamongthesubgeneraofthegenus.
Characternumbersfollowtext.
Iheringia,Sér.Zool.,PortoAlegre,(79): 149- 158,29 set. 1995
FirstknownNeotropicalspeciesofBrunettia... 155
1. Flagellum with 14 flagellomeres / 13 flagellomeres.
Thischaracterhasnoproblemofpolarization.Theground-planconditioninDiptera
forthenumberofflagellomeresis 14.Theapomorphicconditionofthischaracterisshared
by ali members of Brunettia. Although there are other groups in Psychodidae also
presenting 13flagellomeres(aswellasmanyothergroupsinDiptera),theyarenotclosely
relatedtoBrunettia. Hence,this shallbeunderstoodas ahomoplastic similaritybetween
groups ofspecies in Psychodidae.
2.Frontalpatchofsetaepartiallydividedbyaventralnudemesalarea/completelydivided.
The mesal patch of frontal setae is undivided in the genera of Maruinini sensu
DUCKHOUSE(1991). In MormiaEnderlein,1937, thispatchispartiallydividedbya
ventral, short, mesal denudedline. In ali species ofBrunettia the division ofthis patch
is complete or almost complete, with amesal denuded line.
3. Ascoid elongate, simple/ascoidpalmate.
DUCKHOUSE (1987) indicates thatthe ascoidis elongatedin Setomima + Mor-
miiniandisshortforBrunettiaandMormia.Theshortconditionoftheascoidsisproposed
byDUCKHOUSE (1987) as apomorphic. However, Gerobrunettiaisplesiomorphicfor
this character. This, however, conflicts with his proposition for the monophyly of
Gerobrunettia+Brunettia.Theshort,unbranchedconditionoftheascoidisacceptedasthe
ground-plan condition ofthe ascoid in Brunettia. The palmate, obviously apomorphic
shape of the ascoid must have been achieved twice inside Brunettia s.L, once in B.
(Plesiobrunettia) andonceinB. {Campanuiobrunettia).Theapomorphicfeatures shared
by the group of subgenera (Neobrunettia, "Brunettia," Campanulobrunettia,
Maur-obrunettia)areindicativethatthissimilarityintheshapeoftheascoidwouldbebetter
interpretedas ahomoplasy.
4. Scapecylindrical/scapeelongated, exceeding theborderofthe eyes.
5. Pedicel subcircular/pedicel elongated.
ThesetwofeaturesareautapomorphicforB.(Maurobrunettia)anddemonstratethe
monophyly of the subgenus. B. (M.) consobrina andB. (M.) salaxdo notpresent these
modificationsonthefirsttwoarticlesoftheantenna,butthereareotherclearlyapomorphic
features which are shared by these two species and by other species ofMaurobrunettia
withelongatedscapeandpedicel. This indicates thatthe shortscape andpedicel inthese
two species can be understood as a case of secondary loss of the characters. B. (A.)
albrifrons and B. (C.) uncinata also present elongated basal antennal articles, but they
certainly do not belong to B. (Maurobrunettia), so the features must have appeared
homoplastically in the genus.
6. Maxilla with the same length ofthe labrum/maxillareduced.
Inthe ground-planofPsychodidae, the maxillahas about the same length than the
labrum. The reduction in the length ofthe maxilla seems a synapomorphy ofBrunettia.
However, a more detailed study of other genera may still show that the feature is not
restrictedtoBrunettia.
Iheringia,Sér.Zool.,PortoAlegre,(79): 149- 158,29 set. 1995
.
BRAVO&AMORIM
156
7. Presence of hairs and scales onthewing/hairs and scales absentonthewing.
This feature is an autapomorphy ofB. (Atrichobrunettia).
8. R uniformely wide/R wideinthebase
L t
9. Rj without arupture/witharupture inthebasal fourth.
These are two apomorphic features found in B. {Neobrunettia). No other known
species of Brunettiahas thesefeatures.
10. Rj andR morestrongly sclerotizedthanR /R andR sclerotizedasremainingR
5 2+3 { 5 s
veins.
The apomorphic condition of this feature is actually a reversion of an earlier
acquisitionofthecharacter. The strong sclerotizationinveinsRI?R5, andCuA1 isfound
inothergeneraofMaruinini. NeitherB. {Campanulobrunettiá) noxB. (Maurobrunettia)
presentR andR strongerthantheremainingradialveins. Hence, weinterpretherethat
t 5
atthisleveitheconditionsharedbyB.{Brunettia)and#.(Neobrunettia)isplesiomorphic.
The fact that both, B. (Campanulobrunettiá) and B. (Maurobrunettia), present CuA
1
strongerthantheremainingveinsseemstobeanevidencethatthisinterpretationiscorrect.
11 R forkingnearmiddleofthewing/R forkingbasallytothemiddleofthewing.
2D+3UCKHOUSE(1991)includesthisch2a+r3acterasakeyfeaturetoBrunettia(Brune-
ttia), but compares the position of the radial fork to the position of the medial fork.
M
However, the fork of also presents a basal displacement in some groups, so many
specieswithlongforkof1+R2 donotfitinthekeybecausetheforkofM isalsolong.It
is out ofdoubt that the sho2r+t3 fork ofR is plesiomorphic, as can be v1+e2rified with the
2+3
examinationofthewingsofotherPsychodidaeandofotherbasalDiptera.Theapomorphic
condition of this feature is shared by B. (Neobrunettia), B. (Brunettia), B.
(Campanulobrunettiá), andß. (Maurobrunettia).
12. Males withcostal and anal cells normal/costal and analcells developed.
ItiswellknownthatmalesofsomespeciesofBrunettiapresentenlargedcostaland
anal cells, giving to the wing arather different shape. This is obviously an apomorphic
condition thatis shared at leastby the species included by DUCKHOUSE (1991) inB.
(Brunettia),B. (Campanulobrunettiá),B. (Maurobrunettia), as wellbythenewNeotro-
pical species herein described. B. (M.) consobrina - which is also plesiomorphic for
characterthe shapeofthescapeandpedicel-doesnotpresenttheapomorphiccondition
ofthis character,possiblydueto anotherreversion.
13. Subcostal vein incomplete, but clearly present / Sc absent, only a hair line of
unsclerotizedcuticula(h ) seenundermicroscope.
This is a very obxviously autapomorphic condition of the new subgenus B.
(Neobrunettia).
14. Setaenormallydistributedonthegonocoxites/gonocoxites completenudeofsetae.
15. Tergite9 with setae/setae ontergite 9 completelylost.
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FirstknownNeotropicalspeciesofBrunettia... 157
16. Gonostyle with setae on entire length/no setae basally on gonostyles.
17. 4-5 largersetae apically onthe gonostyle/2 setae apically on the gonostyle in outer
side.
The lost setae on the gonocoxites, tergite 9, and on most ofthe extension ofthe
gonostyles are convincing apomorphies. Hence, the whole set of taxa included by
DUCKHOUSE (1991) inBrunettia mostcertainly compose amonophyletic group. The
only exception is B. (Horobrunettia), which present some setae on the tergite 9.
DUCKHOUSE
(1991) interprets this as a reversion to the plesiomorphic state of the
character, position also adopted here. It is noteworthy thatB. (Mirousiella) present 4-5
terminallargersetaeonthegonostyle,whilealiremainingsubgenerapresenttypicallyonly
2apical setae. Itseems clearthatthere is atransformation seriesreducingthenumberof
setae on the gonostyle. Hence, the condition found inB. (Mirousiella) is plesiomorphic
whencomparedtothatwith2setae.Thisindicatesasister-grouprelationshipbetweenthis
subgenus andtheremaining subgenera.
18. Numberofretinacula above 15 /retinaculanotmore than 5.
19. Retinaculastraight/retinaculamesally folded.
The number ofretinacula in the cerei in most species ofBrunettia is rather high,
about 15,withtheexceptionofthespeciesofB. (Horobrunettia),whichpresentareduced
number. The species of B. (Neobrunettia), "B. (Brunettia)", B. (Campanulobrunettia),
andB. (Maurobrunettia) presentverytypical, elongatedretinaculawith amesal fold, an
additional apomorphy onthe strueture.
CONCLUSIONS
Themathrixanalysisshows fourequallyparsimoniouscladograms.Thedifferences
amongthemconcerns onlythepositionofB. (Mirousiella) ânáB. (Horobrunettia). That
is,B. (Neobrunettia), "B. (Brunettia)", B. (Campanulobrunettia) eB. (Maurobrunettia)
always compose a monophyletic group separate from B. (Plesiobrunettia) and B.
(Atrichobrunettia). This group is named here provisionallyB. (Neobrunettia)* using by
AMORIM
(1994b) artifactforphylogenetic sequenced classifications with polytomies.
Thealternativetopologiescorrespondtodistinctinterpretationsoftheevolutionof
characters 15 and 17. Ifthe numberofsetae in the gonostyle is taken as having a single
origin, ashereinadopted, B. (Mirousiella)comesoutasthesistergroupoftheremainder
ofthe genus, B. (Horobrunettia) being kept in a polytomy with B. (Plesiobrunettia), B.
(Atrichobrunettia)andthegroupB.(Neobrunettia)*.Ifthereduetionofthesetaeontergite
9isacceptedashavingasingleorigin, B. (Horobrunettia)wouldbethesistergroupofthe
setoftheothersubgenera,withß. (Mirousiella) inapolytomywithB. (Plesiobrunettia),
B. (Atrichobrunettia) andthegroupB.(Neobrunettia)*. Ifweadmitahomoplasticarrisal
ofthenumberofsetae inthegonostyle, B. (Horobrunettia) wouldbe the sistergroup of
therestofthegenus,withß. (Mirousiella) asthesistergroupofthis subsetof Brunettia.
Finally, ifan homoplastic arrisal ofthe reduetion ofsetae on tergite 9 is considered, B.
(Mirousiella)wouldbethesistergroupoftheremainderofthegenus,andB (Horobrunettia)
.
Iheringia,Sér.Zool.,PortoAlegre,(79): 149- 158.29 set. 1995
BRAVO&AMORIM
158
wouldbe the sistergroup oftherestofsubgenerainsidethis small group.
The analysis made here corroborates thatBrunettia sensu DUCKHOUSE (1991)
corresponds to a monophyletic unity. The character analysis presented above also
demonstratethatalisubgeneraintheliteraturebutBrunettia{Brunettia)aremonophyletic.
Moreover, they allow a partial Solution for the phylogenetic relationships among the
subgenerainthe genus (fig. 19). Evidently, amoredetailed study ofB. (Horobrunettia)
andofB. (Mirousiella) mayhelptobetterdefinethebasalrelationships inthe genus.
The subgenus B. (Brunettia) presently contains a group of species which most
certainlydonotcomposeamonophyleticunity. Noapomorphywasfoundsharedbythe
Australian-Papuan species placed in the subgenus and neither any apomorphic feature
gathering these species andthose in the Afrotropical regions couldbe detected. Maybe
someofthespeciesinB. (Brunettia)wouldfitbetterinB. (Campanulobrunettia)orinB.
(Maurobrunettia).Actually,thesystematicsoftheOrientalandAfrotropicalspeciesofthe
genus shouldberevisedbeforeadditionalconclusionsabouttheinternairelationshipsin
theBrunettia canbe inferred.
The new species herein presented under a new subgenus has a number of
synapomorphicfeaturesthatmayshowtobemoregeneralwhenotherNeotropicalspecies
come to be described. These features, absent in the Australian/Oriental/Afrotropical
speciesofthegroup,seemtojustifythepropositionofanewsubgenusforthisregion.The
discovery ofNeotropical species ofBrunettia (see also DUCKHOUSE, 1991) certainly
changesthebiogeographicalinterpretationofthegroup.Ifthedistributionofthegenuswas
restrictedtotheOldWorld,onecouldacceptaratherrecentoriginforthegroup.However,
the presence ofNeotropical and Australian species together in the same monophyletic
group (Neobrunettia, Brunettia, Campanulobrunettia, Maurobrunettia) would indicate
thatitispossiblyGondwanicinorigin; stemsoriginatedinhigherleveisinthephylogeny
ofthe genus maybe Pangaeic.
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