Two new species of Salina MacGillivray (Collembola, Paronellidae) with rectangular mucro from South America

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REVISTA

BRASILEIRA

DE

Entomologia

AJournalonInsectDiversityandEvolution

w w w . r b e n t o m o l o g i a . c o m

Systematics,

Morphology

and

Biogeography

Two

new

species

of

Salina

MacGillivray

(Collembola,

Paronellidae)

with

rectangular

mucro

from

South

America

Fábio

Gonc¸

alves

de

Lima

Oliveira

a,∗

_,

_Nikolas

_Gioia

_Cipola

b

a_Laboratory_of_Comparative_Biology_and_Bees,_Department_of_Biology,_Faculdade_de_Filosoﬁa_Ciências_e_Letras_de_Ribeirão_Preto,_Universidade_de_São_Paulo,_Ribeirão_Preto,_SP,_Brazil b_Laboratory_of_Systematics_and_Soil_Invertebrate_Ecology,_Instituto_Nacional_de_Pesquisas_da_Amazônia,_Manaus,_AM,_Brazil

a

r

t

i

c

l

e

i

n

f

o

Articlehistory: Received27July2015 Accepted3January2016 Availableonline23February2016 AssociateEditor:DanielaMaedaTakiya Keywords: Chaetotaxy Cremastocephalini Neotropics Speciesgroup S-chaeta

a

b

s

t

r

a

c

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TwonewspeciesofSalina,S.maculiflorasp.nov.fromBrazilandS.colombianasp.nov.from Colom-biaaredescribedandillustrated.Thecompletedorsalchaetotaxy,includingthespecializedchaetae (S-chaeta),isstudiedinthesenewspecies.Comparisonsbasedonthechaetotaxyofthebasomedian field,abdomenII,andmucroshapearemadebetweenspeciesfromgroupsbeta,celebensis,and borneen-sis.ThisisthefirstrecordofSalinawithrectangularmucro(betagroup)inSouthAmericaandakeyto thesevenNearcticandNeotropicalspeciesisprovided.

Introduction

SalinaMacGillivrayisagenusofparonellidswithcurrently71 nominalspecieswidelydistributedinAmerica,Africa,andsouth Asia(Bellingeretal.,2015).Salinaspeciesresembleothergeneraof Cremastocephalini(sensuMitra,1993),suchasAkabosiaKinoshita, 1919,bytheabsenceofscales,dorsalchaetotaxyanddenswithone spatulatedandenlargeddistalappendix(Szeptycki,1979;Mitra, 1993).However,Salinaisdistinguishedfromothergenerabythe presenceofaninterocellarmacrochaetaand densnotcrenulate (Yoshii,1983;Mitra,1993;Soto-Adames,2010).

Basedontheshapeofthemucro,dentalappendageandtergal macrochaetotaxy,Mitra(1973,1993)proposedtwospeciesgroups withinSalina:celebensisandindica.Posteriorly,Yoshii(1981,1983) studiedthechaetotaxyoftheﬁrsttwoabdominalsegmentsand includedthegroupsbeta,borneensisandsaikehialongside celeben-sis,whileexcludingtheindicagroup.

Soto-Adames(2010)recentlyreviewedthebetagroupand pro-posedanewdiagnosis,mainlycharacterizedbytherectangular mucrowithtwomainteethandthesecondabdominalsegment withoneexternaland onecentralmacrochaeta.Sofar,thebeta groupisexclusivelyfoundin theNew Worldand contains ﬁve species: Salina beta Christiansen and Bellinger, 1980 from the

∗ Correspondingauthor.

E-mail:fabiogloliveira@gmail.com(F.G.deLimaOliveira).

United States, S. ventricolor Gruia, 1983 from Cuba, S. wolcotti Folsom,1927fromPuertoRico,andS.bidentata(Handschin,1927) andS.thibaudiSoto-Adames,2010fromCostaRica.Noneofthemis recordedfromSouthAmerica(Soto-Adames,2010;Bellingeretal., 2015).

Herein,wedescribetwonewspeciesofSalinawith rectangu-larmucrofromSouthAmerica,providingdetailedinformationon dorsalchaetotaxyandanidentiﬁcationkeyforSalinabetaspecies group.

Materialandmethods

Specimenswerepreservedin80%ethanol,clariﬁedwith potas-sium dichromate (K2Cr2O7) and hydrochloric acid (HCl) and

mountedonglassslideswithHoyerliquid.Wefollowedthe pro-ceduresdescribedbyArléandMendonc¸a(1982)andChristiansen andBellinger(1998),respectively.Specimenswerephotographed in ethanol gel using a stereomicroscope (M165C) attached to a DFC420 digital camera. Photographs were digitally corrected usingLeicaApplication SuiteV3.4.1.Typematerialis deposited attheInvertebrateCollection ofInstitutoNacionaldePesquisas daAmazônia (INPA), Manaus, Brazil,and “Instituto deCiencias Naturales”(ICN/UNAL),UniversidadNacionaldeColombia,Bogotá, Colombia.

The system used in the descriptions of the species follows Soto-Adames (2010), except Mitraand Dallai (1980)for mucro description. The labial chaetotaxy follows Fjellberg (1999). http://dx.doi.org/10.1016/j.rbe.2016.01.002

0085-5626/© 2016 Sociedade Brasileira de Entomologia. Published by Elsevier Editora Ltda. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

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F.G.L.Oliveira,N.G.Cipola/RevistaBrasileiradeEntomologia60(2016)128–136 129

Nomenclature of the head dorsal chaetotaxy follows the AMS system of genera Entomobrya, Rondani 1861, Seira Lubbock, 1870 and Trogolaphysa Mills, 1938 (Jordana and Baquero, 2005; Soto-Adames, 2008; Soto-Adames and Taylor, 2013), body follows Szeptycki (1979) based on the genus Akabosia, and tergal specialized chaetae (S-chaeta) of Entomobryidae accordingtoZhangandDeharveng(2015).Chaetaeofuncertain homology are followed by a question mark (?). Abbreviations used in the text: Abd.=abdominal segment, Ant.=antennal segment, Th.=thoracic segment, mac.=macrochaeta(e), mes.=mesochaeta(e),mic.=microchaeta(e),sens.=sensilla(e).

Results

Salinamaculiﬂorasp.nov.

SeeFigs.1–27.

Diagnosis.Distinguishedbyﬂower-shapedbluespotsonTh.III toAbd.IV;dorsalheadwithchaetaA5;Ant.Idorsallywith4mac.; basomedianﬁeldwithtwociliatedchaetae(M2andE);Th.IItoAbd. IIrespectivelywith1(p2),4(p1-3,p5),2(m3-4)and3(m3,m3e, m5)centralmac.perside;Abd.IVwithoutA4mac.;collophore anteriorsidewith8+8chaetaeandposteriorsidewith3+3chaetae (Figs.1,3,5,12,14–17,19,24,25).

Description.Totallengthoftheholotype1.7mm.Habitus typ-icalofparonellidsofCremastocephalinitribe(sensuMitra,1993) (Fig.1).Specimenspredominantlypaletolightyellowaspect,with bluepigmentcoveringthedistalregionofAnt.II-III,Ant.Ibase, anteriorandlateralsidesofhead;lateralofTh.IItoAbd.V;Th.III with3+3rosette-likebluespots,Abd.I,IIandIVwith2+2,4+4 and6+6ﬂower-shapedspots,respectively;Abd.IIIand.IVwith bluespotsposteriorly.FemurI-IIIwithoneandtibiotarsiI-IIIwith twolightbluespotsdistally;eyepatchareablack(Fig.1).

Head.Eyepatchesoval,withlargestocelliAandBandsmallest GandH,withthreeinterocularciliatedchaetae(Fig.3).Head dor-salchaetotaxyasinFig.3;antennalseries‘An’with6+6chaetae,

An1a,An1,An2,An3aandAn3asmac.,An2aasspine-likemic.; anteriorseries‘A’withA5mes.;medio-ocellarseries‘M’with3+3 chaetae,M4asmac.,M0andM2asmes.;suturalseries‘S’with3+3 chaetae,S2andS3asmac.,S7asmes.;interocularserieswith3+3 chaetae,rasmac.,pandtasmic.;postoccipitalanteriorseries‘Pa’ with3+3chaetae,Pa5andonetypicalchaeta(P5a)asmac.,Pa6

asbothriotricum;postoccipitalmedianseries‘Pm’with2+2mac. (Pm1andPm3);postoccipitalposteriorseries‘Pp’with1+1mac. (Pp1);postoccipitalexternalseries‘Pe’withmac.Pe3present.Ant. IVwithasimpleapicalbulbandsmoothandciliatedchaetae.Ant.

Fig.1.Salinamaculiﬂorasp.nov.:habitusofaspecimeninethanol(Ant.IIIandIV missing).

Iwith4+4dorsalmac.;dorsalbasewith5+5to6+6 sensillae-likesmoothmic.andventralbasewith12+12(Figs.5and6).Four prelabralciliatedchaetaeand14labralsmoothchaetae(4/5/5),four anterior(a1-2),a1spine-like;fivemedian(m0-2),andfive poste-rior(p0-2)(Figs.7and8).Labialpalpwithfivesmoothproximal chaetaeandsixpapillae(A-EandH),AandCsimple,Hwithtwo smoothguardappendages(h1-2),Bwithfive(a1,b1-4),Dwith three(d1-2andd4),andEwithlateralprocess(l.p.)smallerthan thepapillaandtwosmoothguardappendages(e1-2)(Fig.9). Maxil-laryouterlobewithapicalappendage(a.a.)andbasalchaeta(b.c.) ofsamelength,bothsmooth,sublobal platewiththreeinternal smoothappendages(Fig.10).Rightmandible(ventralview)with fourincisiveteeth,leftmandiblewithfiveteeth;bothmandibles withsixstrongermolarteeth(Fig.11).Basolateraland basome-dianfieldwithchaetaeA1-5smooth(A5thickest),M2,E,L1and

L2ciliated,randM1absent(Fig.12).Cephalicgroove(CG)with 8+8 chaetae, 5+5 ciliated(CG1-3, CG5 and CG7),1+1 smooth (CG8),and2+2asspine-likemic.(CG4andCG6);medialpostlabial (PLM)with1+1ciliatedchaetae(PLM1);externalpostlabial(PLE) with4+4ciliatedchaetae(PLE1-4).Headventralchaetotaxyasin Fig.13.

Thoraxdorsalchaetotaxy.Th.IIasinFig.14;anteriorseries‘a’ with10+10unnamedmac.ofanteriorcollar,oneS-microchaeta (ms),andoneanterolateralsens.(al);medialseries‘m’with3+3 chaetae,m7asmac.,m1andm2asmes.presentorabsent,plus 1+1unnamedmac.laterally;posteriorseries‘p’with2+2chaetae,

p2asmac.,p1asmes.presentorabsent.Th.IIIdorsalchaetotaxy asinFig.15;series‘p’with6+6mac.(p1-3,p5and2+2unnamed chaetae),andoneanterolateralsens.(al).

Abdomen dorsal chaetotaxy. Abd. I as in Fig. 16; series ‘m’ with 3+3 chaetae, m3 and m4 as mac., m5 as mes., and one S-microchaeta(ms).Abd.IIasFig.17;series‘a’witha5as bothri-otricum;series‘m’with4+4chaetae,m3,m3eandm5asmac.,m2

asbothriotricum;series‘p’with2+2chaetae,p6asmac.presentor absent,p7asmes.pluselmac.Abd.IIIasinFig.18;series‘a’with 3+3chaetae,a5asbothriotricum,a7andam6asmes.;series‘m’ with2+2bothriotricha(m2andm5);series‘p’with3+3chaetae,

p6andpm6asmac.,p7iasmes.Oneanterosubmedialsens.(as) present.Abd.IVasinFig.19;series‘A’with3+3chaetae,A1and

A6asmac.,A3asmes.;series‘Ae’withAe1asmac.;series‘B’with 4+4chaetae,B3-5asmac.,B6asbothriotricum;series‘Be’with

Be1asmac.;series‘C’withC1asmac.;series‘T’with2+2 both-riotricha(T2andT4),andoneS-microchaeta(ms);series‘E’with 2+2mac.(E1-2);series‘F’with4+4mac.(F1-3p);series‘Fe’with 5+5chaetae,Fe1-3asmac.,Fe4-5asbothriotricha;posterior mar-ginwith5+5unnamedciliatedmes.,andthreesens.presents(ps andtwounnamed).Abd.VasinFig.20;series‘a’with3+3chaetae,

a5asmac.,a3anda6asmes.;series‘m’with4+4or5+5mac.

(m2-5e),m5apresentorabsent;series‘p’with5+5or6+6chaetae,p1, p3-5andap6(presentorabsent)asmac.,pp6asmes.;one antero-submedialsens.(as)andtwoaccessorysens.(acc.p4andacc.p5)

present.

TergalS-chaetaeformula1,0|1,0,0,1,0(ms)and1,1|0,0,1, 3,3(sens)ofTh.IItoAbd.V(Fig.14–20).

Legs.Trochanteralorganwith14spine-likechaetae(Fig.21). UnguisI-II(samemorphology)withthreeunpairedinnerteeth,a basal,amedianandoneattheapex(Fig.22).UnguisIIIwithone unpairedinnertoothneartheapex(Fig.23).Unguiculi trilame-late, lamellaeinnertruncate, lamellaeexternalacuminate, both smoothedges.Tenenthairscapitate,weaklyciliated,presenton pretarsus.Alltibiotarsiwithonesocketmonocondyleattheapex (Figs.22and23).TibiotarsusIIIwithasmoothdistalchaeta,near unguiculusbase(Fig.23).

Collophore.Anteriorsidewith3+3longciliatedmac.and5+5 smoothmic.;posteriorsidewith2+2smoothmic.and1+1 cili-ated;lateralsidewith14+14smoothmic.(Figs.24and25).

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Figs.2–8.Salinamaculiﬂorasp.nov.:(2)symbolsusedindetailedchaetotaxyschemes;(3)headdorsalchaetotaxy;(4)apicalbulbofAnt.IV(dorsalview);(5)Ant.Idorsal chaetotaxy(right);(6)Ant.Iventralchaetotaxy;(7)prelabralchaetae;(8)labralchaetotaxy.

Furcula.Denswithrowsofciliatedchaetaeandmucrowithtwo mainteeth(d1andv1)andonereducedaccessoryteethpresentor absent(ap)(1outof14specimens)(Figs.26and27).

Typematerial.Holotypefemale,onslideN◦ COLLE040/INPA: Brazil, Amazonas, Rio Preto da Eva municipality, “Aprisco Pásargada” farm (02◦4226.6S; 59◦4253.2W), Amazon forest, 13.ix.2013,45m,Malaisetrap,BGOliveiracoll.Paratypesonslide N◦COLLE040A-G/INPA:3males,7femalesinslides,and4 spec-imensinethanol,samedataasholotype.3paratypesinethanol, samedataasholotype,exceptdate:13.xii.2013.

Otherexaminedmaterial.1immatureonslide,Brazil,Amazonas, PresidenteFigueiredomunicipality,AM-270,Km18(02◦0248S; 59◦5201W), 30.viii.2014,146m, dish trap,NGCipola and FGL Oliveira coll. 2 females on same slide, Manaus municipality, “Raifran” farm, entrance to “Brasileirinho” Km 7 (03◦0208S; 59◦5216W),29.viii.2013,38m,BG-Malariatrap,LBLealcoll. 1 femaleonslide,NovoAirãomunicipality,“MatoGrosso”stream (02◦4900S;60◦5508W),28-31.iii.2013,dishtrap,JTCâmaraand ACMaldanercoll.AlldepositedinINPA.

Etymology.Referstoitsdistinctcolorpatternresemblingflowers (fromLatin:macula=spotandflos=flower)(Fig.1).

Distributionandhabitat.Thisspecieswasfoundonlyinthe Ama-zonforest,AmazonasState,BrazilandinGood’sbiogeographiczone 26ofNeotropicalregion,HighlandsofEasternBrazil:North Brazil-ian(Good,1974).Theclimateoftheareaisequatorialmonsoonal (Am)(Kotteketal.,2006).ThisistheﬁrstspeciesofSalinawith rect-angularmucrodescribedfromSouthAmerica,andtheﬁrstspecies ofSalinadescribedfromBrazil.

Salinacolombianasp.nov.

SeeFigs.28–41.

Diagnosis.Distinguishedbystrongyellowcoloronbody;dorsal headwithS6mes.andwithoutchaetaeAn1aandM0;Ant.Idorsally with7mac.;basomedianﬁeldwithtwociliatedchaetae(M2and

E);Th.IItoAbd.IIrespectivelywith7(a5,m1-2,m4,p1-3),6

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Figs.9–20.Salinamaculiﬂorasp.nov.:(9)labialpalp;(10)maxillaryouterlobe;(11)rightandleftmandibles(rightmolarplateomitted);(12)basomedianandbasolateral ﬁeldoflabium;(13)headventralchaetotaxy;trunkdorsalchaetotaxy:(14)Th.II,(15)Th.III,(16)Abd.I,(17)Abd.II,(18)Abd.III,(19)Abd.IV,(20)Abd.V.

withA3andA4mac.;collophoreanteriorsidewith10+10chaetae andposteriorsidewith5+5chaetae(Figs.28–31,33–37,40,41).

Description.Totallengthoftheholotype2.25mm.Habitus typi-callyofparonellidsofCremastocephalinitribe(sensuMitra,1993) (Fig.28).Specimenswithstrongyellowcolorcoveringthewhole body;eyepatchareablack(Fig.28).

Head.Eyepatchoval,withlargestocelliAandBandsmallestG andH,withthreeinterocularciliatedchaetae(Fig.29).Headdorsal chaetotaxyasinFig.29;antennalseries‘An’with5+5chaetae,An1, An2,An3aandAn3asmac.,An2aasspine-likemic.;anteriorseries ‘A’withA5mes.;medio-ocellarseries‘M’with2+2chaetae,M2as

mes.andM4asmac.;suturalseries‘S’with4+4chaetae,S2andS3

asmac.,S6andS7asmes.;interocularserieswith3+3chaetae,r

asmac.,pandtasmes.;postoccipitalanteriorseries‘Pa’with3+3 or4+4chaetae,Pa2?,Pa5andonetypicalchaeta(P5a)asmac.,

Pa6asbothriotricum;postoccipitalmedianseries‘Pm’with2+2 mac.(Pm1andPm3);postoccipitalposteriorseries‘Pp’with1+1 mac.(Pp1);postoccipitalexternalseries‘Pe’withmac.Pe3present. Ant.IVwithasimpleapicalbulbandsmoothandciliatedchaetae (similartoS.maculiﬂorasp.nov.,Fig.4).Ant.Iwith7+7dorsalmac. anddorsalbasewith6+6sensillae-likesmoothmic.(Fig.30);Four prelabralciliatedchaetaeand14labralsmoothchaetae(4/5/5),four

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Figs.21–27.Salinamaculiﬂorasp.nov.:(21)trochanteralorgan;(22)distaltibiotarsusandfootIcomplex;(23)distaltibiotarsusandfootIIIcomplex;(24)collophore anteriorside;(25)posteriorandlateralsideofcollophore;(26)distaldensandmucrowithtwoteeth;(27)distaldensandmucrowiththreeteeth.22-23,arrowsindicate themonocondyleofthetibiotarsus.

Fig.28.Salinacolombianasp.nov.:habitusofaspecimeninethanol.

anterior(a1-2),a1spine-like;fivemedian(m0-2),andfive poste-rior(p0-2)(similartoS.maculiflorasp.nov.,Figs.7and8).Labial palpwithfivesmoothproximalchaetaeandsixpapillae(A-Eand H),AandCsimple,Hwithtwosmoothguardappendages(h1-2), Bwithfive(a1,b1-4),Dwiththree(d1-2andd4),andEwith lat-eralprocess(l.p.)smallerthanthepapillaandtwosmoothguard appendages(e1-2)(similartoS.maculiflorasp.nov.,Fig.9). Maxil-laryouterlobewithapicalappendage(a.a.)andbasalchaeta(b.c.) ofsamelength,bothsmooth,sublobalplatewiththreeinternal smoothappendages(similartoS.maculiflorasp.nov.,Fig.10).Right mandible(ventralview)withfourincisiveteeth,leftmandiblewith fiveteeth;bothmandiblewithsixmolarteeth(Fig.11).Basolateral

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Figs.29–41.Salinacolombianasp.nov.:(29)headdorsalchaetotaxy;(30)Ant.Idorsalchaetotaxy(left);(31)basomedianandbasolateralﬁeldoflabium;(32)headventral chaetotaxy;trunkdorsalchaetotaxy:(33)Th.II,(34)Th.III,(35)Abd.I,(36)Abd.II,(37)Abd.IV,(38)Abd.V;(39)trochanteralorgan;(40)collophoreanteriorside;(41) posteriorandlateralsideofcollophore.

andbasomedianﬁeldwithchaetaeA1-5smooth(A5thickest),M2, E,L1andL2ciliated,randM1absent(Fig.31).Cephalicgroove (CG)with9+9or10+10chaetae,7+7ciliated(CG1-3,CG5, CG8-10),CG10asciliatedorspine-likemic.,CG4,CG6andCG7(present orabsent)asspine-likemic.;medialpostlabial(PLM)with2+2 cil-iatedchaetae(PLM1-2);externalpostlabial(PLE)with2+2ciliated chaetae(PLE1-2).HeadventralchaetotaxyasinFig.32.

Thoraxdorsal chaetotaxy.Th. IIas inFig. 33; series‘a’ with 11+11unnamed chaetae of anterior collar, a5 mac.present; S-microchaeta(ms),andoneanterolateralsens.(al);medialseries‘m’ with4+4mac.(m1,m2,m4andm7)and3+3unnamedchaetae laterally;posteriorseries‘p’ with3+3mac.(p1-3). Th.IIIasin Fig.34; series‘p’with7+7 mac.(p1-5and oneunnamed);one anterolateralsens.(al),andthreeunnamedmes.

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Abdomendorsalchaetotaxy.Abd.IasinFig.35;series‘m’with 4+4or5+5mac.(m2-5andoneunnamed),m2presentorabsent, andoneS-microchaeta(ms).Abd.IIasFig.36;series‘a’witha5

asbothriotricum;series‘m’with4+4chaetae,m3,m3eandm5

asmac.,m2asbothriotricum;series‘p’with2+2chaetae,p6as mac.,p7asmes.;chaetaelabsent.Abd.IIIasinFig.18(thesame toS.maculiﬂorasp.nov.);series‘a’with3+3chaetae,a5as both-riotricum,a7andam6asmes.;series‘m’with2+2bothriotricha (m2andm5);series‘p’with3+3chaetae,p6andpm6asmac.,

p7iasmes.Oneanterosubmedialsens.(as)present.Abd.IVasin Fig.37;series‘A’with4+4mac.(A1,A3,A4andA6);series‘Ae’ withAe1asmac.;series‘B’with4+4chaetae,B3-5asmac.,B6

asbothriotricum;series‘Be’withBe1asmac.;series‘C’withC1

asmac.;series‘T’with2+2bothriotricha(T2andT4),andone S-microchaeta(ms);series‘E’with3+3mac.(E1-2andE4);series ‘F’with4+4chaeta,F1-3asmac.,F3pasmes.;series‘Fe’with4+4 chaetae,Fe1andFe3asmac.,Fe4andFe5asbothriotricha;median regionwithoneunnamedmes.;posteriormarginwith3+3to5+5 unnamedmes.,andfoursens.present(psandthreeunnamed).Abd. VasinFig.38;series‘a’with2+2or3+3chaetae,a5-6asmac.,a3

asmac.ormic.presentorabsent;series‘m’with3+3mac.(m2-3 andm5);series‘p’with5+5mac.(p1,p3-5andap6);one antero-submedialsens.(as)andtwoaccessorysens.(acc.p4andacc.p5)

present.

TergalS-chaetaeformula1,0|1,0,0,1,0(ms)and1,1|0,0,1, 4,3(sens)ofTh.IItoAbd.V(Figs.33–38).

Legs.Trochanteralorganwith18spine-likechaetae(Fig.39). UnguisI-II(samemorphology)withthreeunpairedinnerteeth,a basal,amedianandoneattheapex(similartoS.maculiﬂorasp.nov., Fig.22).UnguisIIIwithoneunpairedinnerteethneartheapex (sim-ilartoS.maculiﬂorasp.nov.,Fig.23).Unguiculitrilamelate,lamellae innertruncate,lamellaeexternalacuminate,bothsmoothedges. Tenenthairscapitate,weaklyciliated,presentonpseudotarsus.All tibiotarsiwithonesocketmonocondyleattheapexoftibiotarsus (Figs.22and23).TibiotarsusIIIwithasmoothdistalchaeta,near unguiculusbase(Fig.23).

Collophore.Anterior side with3+3 long ciliated mac., 1+1 ciliated mes. and 6+6 smooth mic.; posterior side with 4+4 smoothmic.and1+1ciliated;lateralsidewith13+13smoothmic. (Figs.40and41).

Furcula.Denswithrowsofciliatedchaetaeandmucrotwoteeth,

d1andv1(4outof5specimens)andsometimeswithonereduced accessoryteethpresent(ap)(1outof5specimens)(Figs.26and27). Typematerial.Holotypefemale,onslidedepositedinICN/UNAL: Colombia,Nari ˜no,SanAndresdeTumacomunicipality,University campus(01◦48N;78◦452W),AndeanRegion,4-14.iii.2015,3m, Malaisetrap,AnimalTaxonomystudentscoll.Paratypesonslides depositedinICN/UNAL:1male,2femalesandinalcohol31 speci-mensinethanol,samedataasholotype.ParatypesonslideN◦COLLE 041A-C/INPA:3femalesandin10specimensinethanol,samedata asholotype.

Otherexamined material.1 female and 1immature onslide, Colombia,Guaviare Department, SanJosé delGuaviare munici-pality(02◦3406N;72◦4136W),18.iv.2013,192m,Malaisetrap, AnimalTaxonomystudentscoll.1maleand1femaleonslideand 7specimensin ethanol,Playa Guio,14.iv.2013.Alldeposited in ICN/UNAL.

Etymology.Referstoitstypelocality,inColombia.

Distributionandhabitat.Thisspecieswasfoundinareasof Ama-zonforest,GuaviareDepartmentandlowAndesRegion,Tumaco municipality,bothinequatorialareaofColombia.Theselocations aresituatedrespectivelyinGood’sbiogeographiczone26and28 ofNeotropicalregion(Good,1974).Theclimateoftheareais“Af” tropicalrainﬂorestclimate,characterizedbyhighhumidity(Kottek etal.,2006).ThenewspeciesrepresentstheﬁrstrecordofSalina withrectangularmucrofromColombia.

Discussion

Recently,Zhangetal.(2015)proposedagreatchangein Ento-mobryoideataxonomy bysupporting tergalspecialized chaetae (S-chaeta)asthestrongestmorphologicalcharacterforseparating genera,whatledtomanyclassificationchangeswithinitsfamilies andsubfamilies.Salinaisalsoincludedinthiscontextandits S-chaetaepatternofTh.IItoAbd.V=1,1|0,0,1,3(sens)(excluding Abd.IV,seeZhangetal.2015),wascorroboratedinS.maculiflorasp. nov.andS.colombianasp.nov.Thischaracterandotherfeatures, suchastheS-microchaetaepatternofTh.IItoAbd.V=1,0|1,0,0, 1,0(ms),revealedhereforthefirsttimeinSalina,haveoftenbeen omittedinmanyspeciesdescriptions(Mitra,1973,1993;Yoshii, 1981,1983;Soto-Adames,2010).Therefore,wearguetheyshould befurtherinvestigatedinthegenus,astheyhavebeenusefulto separateotherEntomobryoidea(Jantaritetal.,2013;Zhangand Deharveng,2015).

Inthissense,thespeciesgroupsofSalinarecentlyproposedby Soto-Adames(2010),mustalsobefurtherphylogenetically ana-lyzed,especiallyinaphylogeneticcontext,becausemorphological characteristicsusedintheirdiagnosesareunstable(Table1).For example,inthebetagroup(specieswithrectangularmucro),the basomedianﬁeldhasonlysmoothchaetae,butthetwonewspecies, whichalsopresentrectangularmucro,S.maculiﬂorasp.nov.andS. colombianasp.nov.havetwociliatedchaetae(M2andE)(typical celebensisgroupsensuYoshii,1981,1983).

AnotherexampleisseenonAbd.IIdorsalchaetotaxy,whichwas ﬁrstproposedbyMitra(1973),modiﬁedandYoshii(1981,1983) andpartlyfollowedbySoto-Adames(2010)fortheseparationof Salinagroups,wasbasedonthefollowingcentralmac.formulas: 1,1(m3andm5);2,1(m3,m3eandm5)and2,2(m3,m3e,m5,a6?)

forbeta,celebensisandborneensisgroup,respectively(thelasttwo haveasquaremucro).InS.maculiﬂorasp.nov.andS.colombiana sp.nov.theAbd.IIhas2,1centralmac.(typicalofcelebensisgroup), whileinS.fasciata(Handschin,1928),fromIndonesia,andwitha rectangularmucro(afterYoshii,1983:23)theAbd.IIhas2,2central mac.(typicalofborneensisgroup)(Table1).

Thus,itisclearthattheseparationofspeciesgroupsbasedon thepresenceorabsenceoftwochaetae(a6?,m3e)onAbd.IIis insufﬁcient,sincetheremainingcharacteristics,suchas basome-dianﬁeldchaetaetype,arevariablewithingroups.Therefore,for instance,theonlycharacterabletoseparateSalinaspeciesgroups maybethemucro:rectangularmucrowithtwomainteeth(d1and

v1)andonereducedorabsentaccessorytooth(ap)(betagroup),or squaremucrowiththreemainteeth(d1,apandv1)andareduced orabsentbasaltooth(v2)(celebensisgroup).

WithinthebetagroupandapartfromAbd.IImac.formula,S. maculiflorasp.nov.resemblesS.wolcottiFolsom,1927(see Soto-Adames, 2010) in the color pattern with flower-shaped spots, collophoreanteriorsidewith3+3longciliatedmac.,dorsal chaeto-taxyofTh.III,andAbd.Irespectivelywith4+4and2+2centralmac. (Table1).However,S.maculiflorasp.nov.differsfromthisspecies byAnt.Idorsallywith4mac.(3inS.wolcotti)and6sensillae-like smoothmic.(4inS.wolcotti),fourprelabralchaetae(twoinS. wol-cotti),andcollophoreposteriorsidewith3+3smoothchaetae(1+1 inS.wolcotti)(Table1).Inaddition,S.maculiflorasp.nov.also dif-fersinTh.IIdorsalchaetotaxywith1+1(p2)centralmac.(5+5inS. wolcotti),andTh.IIIwith1+1extranumerarymac.anterolaterally (noneinS.wolcotti).

Salinacolombianasp.nov.resemblesS.thibaudiSoto-Adames, 2010bythechaetotaxypatternofTh.II-IIIwith7and6central mac.,respectively,Abd.Iwith3-4centralmac.,andcephalicgroove (CG)andpostmediallabial(PLM)chaetotaxy(Table1).Conversely, S.colombianalacksdark-coloredpatternsonthebody(presentin S.thibaudi),andAnt.Iwith7mac.(4inS.thibaudi),Abd.Iwith 2+2or3+3centralmac.(m2-4)anteriortoS-microchaeta(ms)

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Table1

ComparisonofmorphologicalcharactersamongselectedspeciesofSalinaandtheirspeciesgroups.

Species Species

group

Ant.IDM Th.IICM Th.IIICM Abd.ICM Abd.IICM Collophore posterior chaetae Unguiculus shape Mucro shape/No ofteeth S.maculiﬂorasp.nov. BR

beta 4 1 4 2 2,1[m3,m3e,m5] 3+3 Truncate Rectangular

2–3teeth S.colombianasp.nov.

CO

beta 7 7 6 3(4) 2,1[m3,m3e,m5] 5+5 Truncate Rectangular

2–3teeth S.fasciatab

ID

beta? ? 5 8 6 2,2[m3,m3e,m5,a6?] ? Truncate Rectangular 3teeth S.betae

USA

beta 2 4 5 4 1,1[m3,m5] 1 Truncate Rectangular

3teeth S.thibaudie

CR

beta 4 7(8) 6(5–9) 4(3–6) 1,1[m3,m5] 3+3(3–6) Truncate Rectangular 2–3teeth S.wolcottie

PR

beta 2–3 3(3–5) 4(2–4) 2(2–3) 1,1[m3,m5] 1+1 Truncate Rectangular 2–3teeth S.celebensisa,e

CR,PR

celebensis ? 1 2 2 2,1[m3,m3e,m5] 8+8 Truncate Square3 teeth S.dedorisc,e

CO

celebensis 4 6 8 3 2,1[m3,m3e,m5] 2+2(0–2) Truncate Square3–4 teeth S.mulcahyaed,e

USA

borneensis ? ? ? 13 2,2[m3,m3e,m5,a6?] ? Lanceolate Square3 teeth S.saikehib

ID

borneensis ? 0 2 4 2,2[m3,m3e,m5,a6?] 9+9 Truncate Square3 teeth Abbreviationsandsymbolsusedtorepresentthemorphologicalcharacteristics:DM,dorsalmac.;CM,centralmac.;BR,Brazil;CO,Colombia;CR,CostaRica;ID,Indonesia; PAN,Panama;PR,PuertoRico;USA,UnitedStates;?,unknown;(),chaetalvariation;[],chaetalnomenclature.

a_Yoshii₍₁₉₈₁₎_. b_Yoshii₍₁₉₈₃₎_. c _Mari-Mutt₍₁₉₈₇₎_.

d _Christiansen_and_Bellinger₍₁₉₉₈₎_. e_Soto-Adames₍₂₀₁₀₎_.

(4+4inS.thibaudi),andAbd.IVwith8+8lateralmac.(10+10in S.thibaudi).

Afterthenew species described here,Salina now hasseven speciesofbetagroupfromtheNewWorld,whichcanbeidentiﬁed inthefollowingkey.

KeytoNearcticandNeotropicalspeciesofSalinawith rect-angularmucro(inpart,afterSoto-Adames,2010)

1.Abd.IIwith2+2centralmac.(m3andm3e)(Figs.7and37)...2 1_._Abd._II_with₁₊₁_central_mac._(m3)_._._.₃

2.Th.IIwith1+1centralmac.(p2);Th.IIIwith4+4centralmac.(p1,p2,p3 andp5)(Figs.4and5);Brazil...S.maculiﬂorasp.nov.

2_._Th._II_with₇₊₇_central_mac._(a5,_m1,_m2,_m4,_p1,_p2_and_p3);_Th._III_with

6+6centralmac.(p1,p2,p2a,p3,p3eandp5)(Figs.34and35);Colombia ...S.colombianasp.nov.

3.Abd.IIandIVwithdarkandlightbluetransversalbands,respectively;Th.II with4+4centralmac.;Th.IIIwith5+5centralmac.;USA...S.beta

ChristiansenandBellinger,1980

3.Abd.IIandIVwithouttransversalbands;Th.IIwith3+3to8+8central mac.,butif4+4thenTh.IIIwith4+4orlessmac....4

4.Th.IItoAbd.Iwithmac.formula5,5,3;Cuba...S.ventricolorGruia,1983 4_._Th._II_to_Abd._I_with_mac._formula_never_5,_5,₃_._._.₅

5.Th.II-IIIwith3+3to5+5and2+2to4+4centralmac.,respectively, generalformula3,4;collophorewith1+1posteriorchaetae;PuertoRico... S.wolcottiFolsom,1927

5_._Th._II-III_with₅₊₅_or_more_central_mac.;_collophore_with₂₊₂_to₆₊₆

posteriorchaetae...6

6.Prelabralchaetaetwo;Th.IIIwith5+5to6+6centralmac.;collophorewith 2+2posteriorchaetae;CostaRicaandUSA...S.bidentata(Handschin,1927) 6_._Prelabral_chaetae_four;_Th._III_with₅₊₅_to₉₊₉_central_mac.;_collophore_with

3+3to6+6posteriorchaetae;CostaRica...S.thibaudiSoto-Adames,2010

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgements

WethankBrunoGarciaOliveiraandothercollectorsfromINPA forprovidingspecimensforthisstudy.FernandoFernández(UNAL)

fortheloanofmaterialfromColombia.MarlonB.C.S.daGrac¸afor thehelpwithEnglish.Accesstoequipmentandadditionalfunding fromCAPESPro-Equipamentos.NeusaHamada/CNPq,Laboratório deSistemáticaeEcologiadeInvertebradosdoSolo (ElizabethF. Chilson), CBio, INPA, and Laboratório deBiologia Comparada e Abelhas (Eduardo A. B. Almeida) from USP. The ﬁrst author is grantedbyFAPESP,SãoPauloResearchFoundation, #2013/26335-9/#2011/09477-9.Thesecondauthoris grantedafellowshipby CNPq.

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