A redefinition of Misumenops F.O.Pickard-Cambridge, 1900 (Araneae, Thomisidae) and review of the New World species

2008·1.1 MB·English

by  Lehtinen, P T

Bull.Br.arachnol.Soc.(2008)14(4),173–198 173
A redefinition of Misumenops F. O. Pickard- obesulus(Gertsch&Davis,1940),comb.n.exMisumenops
Cambridge, 1900 (Araneae, Thomisidae) and and Misumenoides vazquezae (Jiménez, 1986), comb. n. ex
Misumenaarepresentedasadditionalnewcombinationsin
review of the New World species
theMisumenini.
Pekka T. Lehtinen*
Introduction
ZoologicalMuseum,CentreforBiodiversity,
FIN-20014UniversityofTurku,Finland According to Platnick’s (2006) catalogue the genus
Misumenops in the subfamily Thomisinae has a world-
and wide distribution and includes 119 species. Most of the
unrevised species still listed there in Misumenops are
Yuri M. Marusik restrictedtotheNewWorld(81)andparticularlytothe
InstituteforBiologicalProblemsoftheNorth, Neotropical Region (60).
PortovayaStr.18,Magadan,685000Russia All genera of Misumenini were still known until now
from the diagnoses provided by Simon (1895) and
Mello-Leitão(1929),whichwerechieflybasedontheeye
Summary
pattern and some other somatic characters. Minor dif-
The type species of Misumenops, Misumena maculis- ferencesbetweenthemaredifficulttoobserveandthere-
sparsaKeyserling,1891fromBrazilisredescribedfromits fore the species of the New World genera Misumenops,
typematerial.Acloselyrelated,butwidelyconfusedspecies,
Misumena, Mecaphesa, Misumessus, Misumenoides,
Misumenops pallidus (Keyserling, 1880) (_\) from Bolivia
andeasternBraziltonorthernArgentina,isredescribedand Runcinioides and Metadiaea have repeatedly been con-
compared with the type species. The revised concept of fused with each other in regard to generic placements,
Misumenopsisbrieflydiscussedanditsdifferencesfromthe and earlier also confused with Diaea Thorell, 1869 and
Eurasian Ebrechtella tricuspidata (Fabricius, 1775) as well Synema Simon, 1864, members of different tribes. The
as the North American Mecaphesa asperata (Hentz, 1847)
number of synonyms of single species is high in groups
and Mecaphesa celer (Hentz, 1847) are summarised.
thathavebeenrevisedaccordingtomodernstandardsof
Misumenopsguianensis(Taczanowski,1872)fromnorthern
South America, Misumenops bellulus (Banks, 1896) from taxonomy (Lehtinen, 2004), based mainly on detailed
Florida and the Caribbean islands, Misumenops temibilis structuresofthecopulatoryorgans,butacceptingslight
(Holmberg, 1876) from southern South America, and infraspecific variation in coloration and leg spination.
Misumenops variegatus (Keyserling, 1880), comb. n. ex
Considering the lack of proper drawings of the type
MisumenafromPeruareredescribed.Lectotypesaredesig-
nated for Diaea pallida Keyserling, 1880 (\), Misumena species and principles applied to the taxonomy of the
pallens Keyserling, 1880 (\) and Misumena maculis-sparsa Misumenini, it is not surprising that the concept of
Keyserling,1891(_).Afemaleneotypeisdesignated(from Misumenopswasquitevagueuntiltherecentrevisionof
recentmaterial)forXysticustemibilisHolmberg,1876and the misumenine genera by Lehtinen (2004). The study
its senior (but homonymous) synonym Thomisus cinereus
of Palaearctic, Oriental, and some African, Pacific and
Nicolet, 1849. The six confirmed Neotropical species of
American species (Lehtinen, 1993, 2004) revealed that
the genus Misumenops are listed, with four species in the
maculissparsus-group: M. maculissparsus, M. pallidus, M. the majority of species assigned to Misumenops are not
guianensisandM.bellulus,thelastbeingtheonlyspeciesof related to the type species and belong elsewhere.
MisumenopswithintheUSA(Florida);theM.temibilis-and The type species of Misumenops has been cited in the
M. variegatus-groups are both monotypic, but are con-
taxonomic literature outside catalogues only twice after
firmed as members of the newly diagnosed Misumenops.
the description of the genus, by Mello-Leitão (1929)
Misumena exanthematica Holmberg, 1876 from Patagonia
andMisumenoidesnicoletiRoewer,1951asanomennovum and Lehtinen (2004). To most European and Asiatic
for Thomisus cinereus Nicolet, 1849 are synonymised with researchers the concept of Misumenops was based on
Misumenopstemibilis(Holmberg,1876).Thejuniorsecond- M. tricuspidatus (Fabricius, 1775) (=Ebrechtella t.), a
ary homonym Misumenops variegatus Mello-Leitão, 1917
specieswidespreadinthePalaearctic.Thisspecieshas67
is regarded as a nomen dubium. The resurrected genus
taxonomicandidentificationentriesinPlatnick’s(2006)
MisumessusBanks,1904isdiagnosedandtheresurrection
of Runcinioides Mello-Leitão, 1929 is confirmed, leading catalogue compared with 3 entries for the type species.
to the revalidated combinations Runcinioides argenteus In the New World two widespread North American
Mello-Leitão, 1929, R. pustulatus Mello-Leitão, 1929, R. species,M.asperatus(Hentz,1847)andM.celer(Hentz,
souzaiSoares,1942,andR.litteratus(Piza,1933),allcomb.
1847), have served as the model for Misumenops
n.exMisumenops.ThesynonymisationofMetadiaeaMello-
(Kaston, 1948; Schick, 1965; Suman, 1970; etc), al-
Leitão,1929withMisumenopsisnotaccepted.Misumenops
pallidus sensu Rinaldi (1983) from Brazil is transferred to though never compared with the type species. The
thestillunrevised‘‘Misumenops’’pallens-group.Themono- conceptsofSoutheastAsian(Barrion&Litsinger,1995)
typicgenus ChorizopsisSimon,1864istreatedasa nomen andparticularlyIndian(e.g.Tikader,1980)Misumenops
dubium,asitstypespeciesisnowalsoregardedasanomen
species were polyphyletic, used without argumentation,
dubium.AllNearcticspecieslistedinMisumenopsbySchick
and based on unclear reasons (cf. Lehtinen, 2004: 177–
(1965, 1970) and Dondale & Redner (1976), 12 Central
AmericanandCaribbeanspeciesofvariousauthors,and21 180).
Hawaiian species listed in Misumenops and Synema by Although‘‘Misumenops’’sensuauct.isalmostcosmo-
Suman (1970) are transferred to the newly diagnosed and politan and ‘‘well known’’ (i.e. much cited), its type
delimited Mecaphesa Simon, 1900. The resulting 43 new
species Misumena maculis-sparsa Keyserling, 1891 has
combinations in Mecaphesa are listed. Misumenoides
been illustrated only once (Mello-Leitão, 1929) after its
*Towhomallcorrespondenceshouldbesent. original description and not by the author of the genus
174 RedefinitionofMisumenops
(F.O.P.-Cambridge,1900).ThefiguresofMello-Leitão Queiros’’: information about type preservation from Dr
(1929) are not detailed enough for undoubted specific IsabelaRinaldi;MZSP=MuseudeZoologiadaUniver-
identification, and it is not certain whether he saw the sidade de São Paulo, São Paulo, Brazil (Dr Ricardo da
syntypes in BMNH. The closely related M. pallidus PintaRocha);MZT=ZoologicalMuseum,Universityof
(Keyserling,1880)hasbeendiscussedmoreoften,butits Turku, Turku, Finland (no present curator); MZUM=
identificationhascausedmanyproblems,andsometimes Museo de Zoologia, Universidad de Montevideo,
large samples identified as M. pallidus include up to Uruguay, Dr Roberto Capocasale, Dr Ricardo Perez-
five different species or even no pallidus at all, although Miles; NHRS=Naturhistoriska Riksmuseet, Stockholm,
three other different species are present (E. Reimoser’s Sweden, Dr Torbjörn Kronestedt; NMW=Naturhistor-
labelled samples in NMW collected by himself!). The isches Museum Wien, Vienna, Austria, Dr Jürgen
only revisional study supposed to deal with M. pallidus Gruber,DrVerenaStagl;PGC=CollectionofMsPeggy
(Rinaldi,1983)wasbasedentirelyonmisidentifiedspeci- Gerba, Arizona, USA; PTL=Temporary personal col-
mens with regard to this species, but this is not surpris- lectionofPekkaT.Lehtinenafterretirement;UNAM=
ing because of repeated confusion by other authors in Universidad Nacional Autonoma de México, México
the past. This confusion is a strong argument for the City, Prof. Oscar Francke & Dr Maria L. Jiménez;
necessityofthethreelectotypedesignationsmadeinthis ZMB=Zoologisches Museum der Humboldt Univer-
publication. The syntypes of neither M. pallidus nor M. sität, Berlin, Germany, Dr Jason Dunlop; ZMMU=
pallenswerestudiedbyRinaldi(J.Gruber,pers.comm., Zoological Museum, University of Moscow, Russia, Dr
2006), but only Brazilian samples identified by Mello- Kirill Mikhailov.
Leitão, Piza, Soares, etc. AME=anteriormedianeyes,PME=posteriormedian
Thegoalofthispaperistogiveareviseddefinitionfor eyes, MOT=median ocular trapezium, RTA=
Misumenops, to redescribe the type species and its close retrolateral tibial apophysis, ITA=intermediate tibial
relatives, and preliminarily discuss the concept of New apophysis, VTA=ventral tibial apophysis.
World Misumenops sensu auct. according to modern
principles of taxonomy, and to present a brief com-
Taxonomy
parison of Misumenops with the ‘‘model’’ of Eurasian
Misumenops (=Ebrechtella tricuspidata) and the New
World‘‘model’’M.asperatus(Hentz,1847).Acorrected Misumenops F. O. Pickard-Cambridge, 1900
placementofallNorthAmericanspeciesofMisumenops
Thomisus:Nicolet,1849:396,inpart;Taczanowski,1872:90,inpart.
sensu auct. is also discussed.
Diaea:Keyserling,1880:112,inpart.
Misumena:Holmberg,1876:27,inpart;1881:155;Keyserling,1880:
101,inpart;1891:245;Banks,1896:71,inpart;Simon,1897:9,
Methods
inpart;Petrunkevitch,1911:410,inpart.
Misumenops F. O. Pickard-Cambridge, 1900: 141; Petrunkevitch,
In addition to traditional light microscope analysis
1911:410;Mello-Leitão,1929:77,inpart;Lehtinen,2004:173.
of material (Olympus SZH & Wild M5 with ocular
Misumessus:Banks,1910:50,inpart.
micrometer),scanningelectronmicroscopywithaJEOL Metadiaea:Piza,1933:88,inpart.
5200 was used for micrographs of male palps and other Misumenoides:Roewer,1951:448.
structures,digitisedwithSemAforesoftware.Thedigital References concerning unchecked material of Misumenops pal-
lidusarenotincluded.
photographsweretakenwithanOlympusC5050digital
cameraconnectedwithanOlympusSZX12lightmicro- Type species: Misumena maculis-sparsa Keyserling,
scopeandenhancedusingtheCombineZSsoftware.All 1891 from eastern Brazil.
measurements are given in millimetres. Diagnosis: Misumenops resembles the New World
Abbreviations used in text (present curator in paren- generaMecaphesaSimon,1900andRuncinioidesMello-
theses, if did not participate in loans or information Leitão, 1929 in regard to many somatic characters. The
for this project): BMNH=British Museum, Natural dorsal surface of the carapace of Misumenops has
History, Dr Paul Hillyard, Ms Janet Beccaloni; numerous long rigid setae in and around the ocular
BPBM=BerniceP.BishopMuseum,Honolulu,Hawaii, region and shorter setae in other parts of the carapace
USA, Dr Scott Miller (Dr Ronald Englund); and on the dorsal surface of the abdomen (Figs. 30, 61,
IZPAN=Instytut Zoologiczny, Polska Akademia Nauk 63), in contrast to the presence of only normal hairs in
(Mr Tomasz Huflejt), Ms Dominika Mierzwa; the Old World species of ‘‘Misumenops’’. These rigid
DJC=Collection of Dr D. T. Jennings, Maine, USA; setae are also present, but more conspicuous and also
MACN=Museu Nacional de Ciencias Naturales longer throughout the dorsal surface of the body in
‘‘Bernardino Rivadavia’’, Dr Cristian Grismado; Mecaphesa (Figs. 64–65) and especially in Runcinioides
MNHN=MuséumNationald’HistoireNaturelle,Paris, (Figs.66–67).Thelectotypeandallolectotypeofthetype
France, Dr Michel Hubert, the late Dr Jacqueline species are strongly bleached and also many of the
Heurtault, Dr Christine Rollard; MLP=Museu de La dorsal setae have been worn off; see also p.178. All
Plata,LaPlata,Argentina,DrCristianItuarte,DrLuis speciesofMisumenopsaredifferentiatedfromallspecies
Alberto Pereira, Ms Monica Tassara; MRJ=Museu ofMisumenoidesbythelackofaserratemargin(Fig.36)
Nacional,RiodeJaneiro,Brazil,DrAdrianoKury,Mr on the thoracic part of the male carapace. Dark annu-
Thiago da Silva Moreira; MZLQ=Departamento de lations on male legs I–II are usually present also on the
Zoologia, Escola Superior de Agricultura ‘‘Luiz de tarsi of Misumenops (Figs. 24, 29, 30, 60, 63), but are
P.T.Lehtinen&Y.M.Marusik 175
restrictedtothepatellae,tibiaeandmetatarsiinmalesof provide comparative SEM figures of the male palp
all other genera of Misumenini. The most useful diag- and photographs of both sexes of E. tricuspidata and
nostic features are found in the structure of the genital Mecaphesaasperata.InTable1webrieflysummarisethe
organs.ThetibialapophysesofMisumenopsconsistofa maindifferencesinthepalpandfemalepatternsbetween
separate VTA and a distally narrowed apophysis origi- these three genera.
nating from the complete fusion of RTA and ITA (Fig. SpeciesoftherevisedMisumenopsareseparatedfrom
2).ThetipofRTAhasaserratetolobatemargin(Figs. all other Neotropical species currently listed (Platnick,
37–38, 40, 57b) or subapical concentric ridges in M. 2006) in this genus by different basic structural pat-
variegatus(Fig.49),andacentral,moreorlessmembra- terns of the male and female copulatory organs. Al-
nous part with one conspicuous seta (Figs. 10, 40, 56) though types or authoritatively identified material of
thatislackinginRuncinioidesandMecaphesa.Alltibial many of them have already been checked by us, a
apophyses are lateral processes of a single plate in complete revision of this strongly polyphyletic assem-
Mecaphesa (Figs. 42–43). Males of Misumenops are blage of misumenine species is beyond the scope of this
further differentiated from Runcinioides by having a publication.
relatively short embolus originating mesally (opposite Description:Small(_2.5–3.5mm,\4–6mm)misume-
the tibial apophyses: Figs. 1 & 5) and by asimple nine spiders with distinct sexual dimorphism in size and
tutacular apophysis at the base of the cymbium (Figs. legcoloration.AdultspecimensoftrueMisumenopspale
2–3,55),whileRuncinioideshasatutaculargroovealong yellow to pale brown, with distinct paired pattern in
the tegular margin ending in a fissure in the tegular posterior half of abdomen (Fig. 22). Pattern of leg
margin (Fig. 50). Runcinioides also has a lateral boss on annulations in some male specimens obscure, while
thecymbialmargin,homologoustothewholetutaculum other specimens of same population may have very
in Misumenops. For some diagnostic details of the distinct dark annuli (Figs. 29–30). Dorsum of male
copulatory organs of Misumenops and groups including abdomen often lightly sclerotised, but scutum obscurely
the ‘‘model’’ species of recent authors, see Table 1. limited.
FemalesofMisumenopsarecharacterisedbyamostly Tutacular structures of Misumenops consist of a
small epigynal hood (Figs. 4, 7) (widest in M. bellulus: groove along cymbial margin and a hairy process or
Fig. 58) and the lack of a thin central septum which is knob at base of cymbium (Figs. 2–3). Size of this
characteristic for most species of Mecaphesa (Kaston, tutacular process is a useful specific character. Tibial
1981: figs. 1485, 1497) or wider, posteriorly rounded apophyses of the revised Misumenops consist of fused
septum as in the only species of Misumessus (Kaston, RTA–ITA with serrate, dentate, lobate or ridged apex
1981: fig. 1486). The vulva of Misumenops consists of (Figs. 37–38, 57b) and a small, distinctly separate VTA
tubular U-shaped receptacula with short connecting (Fig. 1); this is essentially different from the single plate
ducts (Figs. 8–9), while the vulva of Runcinioides has withthreedistinctlateralprocesses(RTA,ITA&VTA)
verylongmeanderingductsandaposteriorlyprolonged in Mecaphesa (Figs. 42–43). Mesal face of RTA–ITA
quite narrow epigynal hood (Rinaldi, 1988: figs. 3–4). excavated, more or less membranous, and a long seta
As was pointed out above, the model species of present in centre of this cavity (Figs. 10, 40).
Misumenops were different for European and North Embolic tip usually with subdistal triangular lamina
American arachnologists. The placement of Araneus (Figs. 13, 47, 57a), embolic surface smooth or partly
tricuspidatus Fabricius, 1775 in Ebrechtella was made coarsely striated (Figs. 13, 39, 47–48). Female epigynal
possible by study of the holotype male of the type hood with well sclerotised margin; seminal receptacula
species of Ebrechtella, E. fruhstorferi Dahl, 1908 from long, U-shaped; seminal ducts between receptacula
Java (ZMB); for details, see Lehtinen (2004). Here we short, uncoiled.
Structures(*onleftpalp) Misumenopss.str.(maculissparsus) Mecaphesaasperata/celer Ebrechtellatricuspidata
VTA Aswideaslong(Fig.1) Aswideaslong(Figs.42–43) Longerthanwide(Fig.14)
RTA&ITAseparate No(Fig.2) Yes(variableinMecaphesaspp.) Yes(Fig.15)
Embolictipposition Centro-basalpartofcymbium Onprolateralpartofcymbium Upper-centralpartofcymbium
(c.3.30o’clock*)(Figs.1,6) (variableinotherMecaphesaspp.) (c.2o’clock*)(Figs.15,35)
Openingofembolicduct Distal,unclearmargins(Figs.1–2) Distal(Figs.42–43) Subdistalovalpit(Figs.16–17)
Surfaceultrastructureof Coarselystriatedtosmooth(Figs. Variablystriatedtosmooth(Figs. Denselystriated(Figs.16–17)
embolus 13,39,48) 42–43)
Embolicturns 180((halfaturn)(Figs.1,5, >360(intwoplanes(turns)(Fig.42) 360((wholeturn)(Fig.35)
44–45,54)
Embolusorigininmid- 8.30–10o’clock*(Figs.1,5, 6.30–8o’clock(Figs.42–43),variable Fromcentreoftegulum
prolateralpartoftegulum 44–45,54)exceptM.variegatus inotherMecaphesaspp. (c.2o’clock*)(Fig.35)
11.30(Fig.46)
Cymbiumwithtutaculum Yes(Figs.1,3) No(onlytutaculargroove)(Figs. No(Figs.14–15)
42–43)
\carapacewithsublateral Yes(Figs.18,22,29,30,60–61, Variable(Fig.62),otherMecaphesa No(Fig.26)
brownbands 63) spp.(Figs.64–65)
Table1: SomedifferencesbetweenMisumenops,Mecaphesa,andEbrechtellatricuspidata.Mecaphesaasperata(Fig.42),Mecaphesaceler(Fig.
43),E.tricuspidata(Figs.14–17,25–28,34–35,51).
176 RedefinitionofMisumenops
Composition and range: The six species which remain as a separate genus by Lehtinen (2004: table 1), and the
in the revised Misumenops are best characterised by the status of Misumessus Banks, 1904 (type species Mis-
structure of the male and female copulatory organs. umena oblonga Keyserling, 1880), formerly part of the
These six species include a group of four closely related traditional Misumenops, was also discussed by Lehtinen
species (the maculissparsus-group: M. maculissparsus, (2004: 174). The resurrection of these two genera is
M. pallidus, M. guianensis (Taczanowski, 1872) and M. confirmed and they are diagnosed here (pp.190 and
bellulus(Banks,1896),seebelow)aswellastwomoreor 195).
less isolated species, but both sharing the diagnostic The presence of numerous long erect setae on the
characters of Misumenops: M. temibilis (Holmberg, carapace of all North American ‘‘Misumenops’’
1876) and M. variegatus (Keyserling, 1880). Only the (=Mecaphesa)explainstheerroneousdiagnosticcharac-
male of M. variegatus is known to us, but its embolic ters suggested for ‘‘Misumenops’’ by Kaston (1948,
structureanduniquemodificationsofRTA(Fig.49)can 1981),Schick(1965)andSuman(1970).Lehtinen(2004)
be homologised with modifications in the other five emphasisedthedifferencesincolorationbetweentheOld
species. WorldandNewWorldspeciesofMisumenopsauct.This
The known range of the revised Misumenops extends is only partly true, as several Neotropical species also
from Florida and the Caribbean islands (M. bellulus) to have a lot of greenish or greenish grey colour on the
southern Chile and Argentina (M. temibilis). The type carapace, abdomen and legs in living or freshly caught
species of Runcinioides, Mecaphesa and Misumessus specimens. Many of these species must be transferred
have each been found to represent genera outside the from Misumenops to the still unnamed ‘‘Misumenops’’
revised Misumenops. pallens-group(=MetadiaeasensuRinaldi,1988,inpart)
Discussion: The synonymy of Metadiaea Mello- or to other unnamed taxa.
Leitão, 1929 with Misumenops proposed by Rinaldi
(1988) cannot be accepted, as the type species of Meta-
The maculissparsus-group
diaea, M. fidelis Mello-Leitão, 1929 was compared
neither with the type species of Misumenops nor with Thefourspeciesofthisgrouparemoreorlesssimilar
any of its relatives stated here to belong to this genus. in regard to the structure of the male and female
Unfortunately the syntypes, the only known identified copulatory organs, while their somatic characters are
material of M. fidelis, have obviously become lost and also shared by the two species outside the
thusthegroundsforhersynonymisationdidnotinclude maculissparsus-group. The differences between the four
the study of the type species of either genus. speciesconsistofdifferentdistalmodificationsofRTA–
The original descriptions and drawings (Keyserling, ITA, origination of the embolus, as well as ultrastruc-
1880) of two Peruvian species do not exclude the poss- tural details of the embolus. The female epigynes are
iblerelationshipofMisumenapunctataKeyserling,1880 superficiallyverysimilarinthefirstthreespecies,butthe
and Misumena amabilis Keyserling, 1880 with Mis- epigynal hood is distinctly wider in M. bellulus than in
umenops, and the former species was transferred to the the three other species (cf. also Bryant, 1940).
oldMisumenopss.lat.byPetrunkevitch(1911).Accord- Discussion: Although M. maculissparsus, M. pallidus
ing to the original description M. amabilis has dark andM.guianensisarerathereasytoidentify,whentheir
annuli also on the tarsi, although not extending to the type material was studied, the type species by light
tip, as in all species placed here in Misumenops. Al- microscopy, the others by SEM (Figs. 1–2, 10–12,
though no dark annuli were described for tarsi I–II in 38, 45), the variation of several palpal and epigynal
M. punctata, the palpal tibia was described as having a characters was found to be high in large samples from
very thick anteriorly directed apophysis with a small ParaguayandnorthernArgentina,possiblyrepresenting
tubercle at its rounded end. The original drawing does single populations. Therefore the final taxonomic revi-
not give such detailed information about the palpal sion of this group must still await additional material
apophyses, but the German text provides no possibility fromdifferentpartsoftherangeofthisspeciesgroupto
of a different interpretation. Both species are also de- exclude the possibility of strong clinal variation in the
scribedashavingscatteredlongsetaeonthecarapacein copulatoryorgans.Mello-Leitão(1929:224–225)placed
apatternmoreorlesssimilartothatinourMisumenops M.maculissparsusandM.guianensisfarfromeachother
spp. in his key, based on presence or absence of an abdomi-
A proper revision of several Neotropical species, nal pattern, most probably without personal study of
including M. punctata and M. amabilis, has been im- anymaterialofM.maculissparsus,butpresentedawide
possible because of the loss of the type material. Ms distribution for M. guianensis within Brazil.
Mierzwa (IZPAN) has informed us that the type collec- A fifth, possibly separate taxon of this group, repre-
tions of Taczanowski (1872) and Keyserling (1880) in sented by several males in the large sample from
theWarsawMuseumdonotincludeanythomisidtypes. Paraguay,maybeworthyofspecificstatus.Theembolic
The spider collection of NMW has undergone several base of these specimens is surrounded by a raised,
phases of possible confusion in the past, owing to marginally crenated outgrowth of the tegulum, and the
careless relabelling and possible changes in the contents tip of the RTA is serrated only on one side. More
ofsinglevials(DrJ.Gruber,pers.comm.,March2006). informationandmaterialisnecessarybeforedescription
Runcinioides Mello-Leitão, 1929 (type species R. of such a ‘‘new’’ taxon, as all its specimens were found
argenteus Mello-Leitão, 1929) has already been treated within a large sample of M. guianensis.
P.T.Lehtinen&Y.M.Marusik 177
Misumenops maculissparsus (Keyserling, 1891) (Figs. (Buenos Aires) kindly informed us (pers. comm., 2006)
1–4, 18–20, 29, 32) that Taquara do Mundo Novo is a well known locality
in Brazil, in Estado do Rio de Janeiro—a fact that
Misumenamaculis-sparsaKeyserling,1891:245,pl.10fig.186.
Misumenops maculissparsus: F. O. Pickard-Cambridge, 1900: 141; seems to have been unknown to Mello-Leitão (1929),
Mello-Leitão, 1929: 232; Lehtinen, 2004: 173 (all figures who obviously included both Taquaras in the range of
referredtoasM.aff.maculissparsusareM.pallidus). this species, as no other material was mentioned.
Types: Lectotype _ designated here from Brazil, The orthography maculissparsus conforms to the cur-
TaquaradoMundoNovo,leg.Drv.Ihering,paralecto- rent Code (ICZN, 1999: Art. 32.5.2.3).
type \ with same data, both in BMNH. Other material examined: , Jujuy, 1 _, leg.
Notes: There are places called Taquara both in Prov- E.Reimoser,1907(NMW,identifiedbyReimoserasM.
inces Rio Grande de Sul and Estado do Rio de Janeiro, pallidus). Some specimens of the large sample from
bothclosetotheeasterncoastofBrazil.TherangeofM. Paraguay in NMW and discussed here under M. guian-
maculissparsuswasstatedtobe‘‘RioGrandedoSulaté ensispossiblybelongtoM.maculissparsusasthissample
Rio de Janeiro’’, which means from Rio Grande do Sul probablyconsistsofsamplesfromdifferenthabitats,but
‘‘to’’ Rio de Janeiro, as interpreted by Mello-Leitão it is difficult to identify all specimens individually with-
(1929: 232), but not ‘‘or’’. However, Dr C. Grismado out detaching a palpus from each male.
Figs.1–9: Leftmalepalpandfemaleepigyne.1–4Misumenopsmaculissparsus;5–9M.pallidus.1,5Palp,ventral;2,6Ditto,retrolateral;3Part
of palp, retrolateral; 4, 7, 8 Epigyne, ventral; 9 Ditto, dorsal. Scale lines=0.1mm. Eh=epigynal hood; Em=embolus; Eo=epigyne
opening; Ep=embolic base pocket; Re=receptaculum; RT=retrolateral tibial apophysis; Sd=seminal duct; Tr=tegular rim;
Tu=tutaculum;VT=ventraltibialapophysis.
178 RedefinitionofMisumenops
Diagnosis: This species can be distinguished from the tarsi typical of all its relatives more or less totally
sibling species M. pallidus by the thicker (wider than faded in lectotype, but present in specimen from Jujuy,
long) lateral tibial apophysis (Fig. 2 cf. Fig. 6), closely Argentina. Femur I with 5 dorsal and 4 prolateral
spaced tibial apophyses, lower position of the embolic spines. Tibial and metatarsal spines weak, indistinct.
base (9.30o’clock: Fig. 1 cf. Fig. 5), and by the colora- Palp as in Figs. 1–3, light coloured; tibia with two
tion of both sexes (Figs. 18–21 & 29 cf. Figs. 22–25 & apophyses, tegulum without apophyses, embolus mak-
30). The general appearance of the epigynes is practi- ing half a circle. Ventral tibial apophysis short, wider
callyindistinguishable(cf.Figs.4&7and32–33).Males thanlong,inmidpartoftibia.FusedRTA–ITAstrongly
of M. maculissparsus, M. pallidus and M. guianensis all swollen (wider than long), pointed and with some distal
have coarse ridges close to the narrowed apex of the modifications which could not be figured in detail,
embolus (Fig. 13), while M. temibilis has a smooth becausenoSEMmountscouldbedonefromsinglemale
embolus, except for the widely triangular lateral lamina in type material. Cymbium with small slightly ridged
close to the apex (Fig. 39). basal tutaculum. Tegulum round, with heavily sclero-
Description: Female: Total length 6.0. Carapace 2.25 tisedmarginalrim,followingcourseofembolus;baseof
long, 2.08 wide (Figs. 18–20), light coloured with two seminal duct thinner than embolic base. Embolus starts
wide brownish bands, ocular area white, clearly separ- from tegular pocket at 9.30o’clock position (in left
atedfromrestofcarapace.Abdomenpentagonal,with3 palp), its base clearly separated from tegulum. Embolic
pairs of brownish spots in basal half, sides without base relatively wide, width of embolus continuously
pattern, venter with dark spot resembling ‘‘W’’ behind tapering along its course, its tip flat and not pointed.
epigastric furrow and 5 pairs of dark spots (=muscle Distribution:KnownfromeasternBrazilandnorthern
apodemes). Legs pale, spines and strong macrosetae of Argentina, possibly also from Paraguay.
femora I–II surrounded by brown spots, apical parts of
patellae, basal and apical parts of tibiae with wide
brownish rings (Fig. 20). Femur I with 3 or 4 prolateral Misumenops pallidus (Keyserling, 1880) (Figs. 5–9, 10–
and 1–3 dorsal weak spines (one dorsal spine almost 13, 22–25, 30, 33)
prolateral). Tibia I with 6 pairs of ventral spines, meta-
DiaeapallidaKeyserling,1880:117,pl.2fig.65.
tarsus I with 8 pairs of ventral spines. For length of leg
Misumenapallida:Keyserling,1891:245.
segments, see Table 2. Misumenopspallidus:F.O.Pickard-Cambridge,1900:141(transferto
Epigyne as in Figs. 4 & 32, with triangular anterior Misumenops);Mello-Leitão,1929:229,figs.27&27a–b;notM.
hood(apicalpocket)andtransverseelongatereceptacula pallidussensuRinaldi,1983norpallidus:Lehtinen(2004:156&
163,figs.27–28&76).
visible through integument. Hood with posterior more
?MetadiaeavulgarisPiza,1933:88,fig.1,syn.Mello-Leitão,1941.
sclerotisedrim.Partofreceptaculumthattouchescuticle
Misumenops exanthematicus: Mello-Leitão, 1941: 164, misidentifica-
appears darker. Receptacula long, horizontal U-shaped tion(materialinMACNchecked!).
(Fig. 4). Receptaculum makes one 180( turn. Misumenopsaff.maculissparsus:Lehtinen,2004:figs.5,24–25,75.
Male:Totallength3.0.Carapace1.38long,1.48wide
Types:FoursyntypesfromBrazil(NMW)andseveral
(Figs. 21, 29), light brown, with two wide brownish
females from Colombia [New Granada] originally in
submedian bands; bands without distinct margins; eyes
Keyserling’s personal collection, later preservation un-
surrounded by white pigment rings, ocular area not
known.
separated from rest of cephalic part by white pigmenta-
OnefemalefromBrazil,leg.Helmreich(NMWacqui-
tion as in female. Abdomen ovoid, light coloured, with
sition no. 1847 II 20) is designated here as lectotype for
series of 5 pairs of brownish spots decreasing in size to
Diaea pallida Keyserling, 1880; the vulva of another
spinnerets;someofspotsfusedinline;sideswithbrown
syntype from the same vial has been used for a vulval
stripe;venterwithrectangulargrey-brownishspot;dark
slide mount for comparison with other samples of this
spot with 4 pairs of dots (=muscle apodemes) (Fig. 19).
species group.
Legs light yellow, femora I–II with numerous brown
Othermaterialexamined:BOLIVIA:SantaCruzProv.:Amboro
spots (Fig. 29). Legs I–II with darkened terminal half N.P.,_\,juv.;Montero,_\,juv.;Guarne´,RioSelva,many_\,juv.,
of metatarsus and tibia (Fig. 29); dark distal half of allinlowvegetation,11–19August2007,leg.P.T.Lehtinen(PTL).
PARAGUAY:Foncière:numerous_\,leg.Reimoser,1908(NMW).
URUGUAY:Montevideo,1_2\,PuntaEspinillo,bushesandtrees,
Female Fe Pa Ti Mt Ta Total 10October1996(leg.R.Perez-Miles&M.Perez)(ZMTandZMUM),
6\ 2 juv., 10 November 1996, leg. R. Perez-Miles & M. Perez
I 2.75 1.0 2.25 2.08 1.1 9.18 (ZMUM). ARGENTINA: Salta, Juramento, 1_ 1 subad. _, leg.
II 2.6 1.0 2.13 2.08 1.1 8.91 MaximilianoBiraben,March1939(MLP14781);Salta,29\&numer-
III 1.4 0.75 1.13 1.0 0.63 4.91 ous juv., leg. E. Reimoser, 1907 (NMW); Cordoba, 5\ 2 juv.,
IV 1.75 0.8 1.25 1.2 0.67 5.67 Calamuchita,December1941,leg.J.M.Viana,A.Lisedet.(MACN
10889),1_1\,1985,leg.Viana(MACN);BuenosAiresProv.:Glew,
Male Fe Pa Ti Mt Ta Total 3\ [large juv. not conspecific], leg. Carpintero, 1974, det. A. Lise
(MACN10892);LomasdeZamora,V.Fiorito,1_,March1991,leg.
I 2.25 0.83 1.9 1.9 0.93 7.81
C.Grismado(MACN10893),1_3\,February1992,leg.C.Grismado
II 2.13 0.8 1.73 1.75 0.85 7.26
(MACN 1089?2), February 1992, leg. C. Grismado (MACN); Santa
III 0.95 0.43 0.75 0.7 0.4 3.23
Fé,5October1962(MACN10891).
IV 0.95 0.4 0.83 0.7 0.38 3.26
Notes: Only references to material that could be
Table2: Misumenopsmaculissparsus,legmeasurements. personally checked or was reliably represented by
P.T.Lehtinen&Y.M.Marusik 179
unambiguousdrawingshavebeenincluded.Widespread Mello-Leitão(1929)remainsdoubtful.Thetypematerial
confusionabouttheconceptofM.pallidushasprevailed of Diaea pallida was not available to Rinaldi (1983)
and, e.g., Mello-Leitão (1929) presented more or less and all her figures presented for M. pallidus (figs. 3–4,
identical synonymic lists for M. pallidus and Misumena 7–8, 11–12, 14) seem to refer to some species of the
pallens Keyserling, 1880, the latter obviously not con- M. pallens-group. Although Mello-Leitão (1929) gave
generic with M. pallidus. The identity of fig. 27 in the same lists of references under the names of the
Figs.10–17: Micrographsofleftmalepalp.10–13Misumenopspallidus(Uruguay,PuntaEspinillo);14–17Ebrechtellatricuspidata(Ukraine).10,
14 Ventral; 11–12, 15 Retrolateral; 13, 16–17 Tip of embolus. Em=embolus; Ep=embolic base pocket; RT=retrolateral tibial
apophysis;Tr=tegularrim;Tu=tutaculum;VT=ventraltibialapophysis.Scalelines=0.1mm(10–11,14–15),0.01mm(12),0.005mm
(13,16–17).
180 RedefinitionofMisumenops
speciesforbothM.pallensandM.pallidus,hisdrawings (1983), but no references to localities for specimens
refer correctly to these two species. He mentions the depicted by her were presented and it is possible that
wide range and common occurrence of M. pallidus she simply repeated the synonymy established by
in Brazil, although most probably he also confused Mello-Leitão (1941). The uncertainty of the identifica-
M. maculissparsus, M. pallidus and M. guianensis with tions of M. pallidus by Mello-Leitão culminated in
each other. Later he (Mello-Leitão, 1941) synonymised the latter publication, where material identified by him
Metadiaea vulgaris Piza, 1933 with his Misumenops as M. exanthematicus and checked by us is actually
pallidus and this synonymy was repeated by Rinaldi M. pallidus.
Figs. 18–28: 18–21 Misumenops maculissparsus, lectotype _ and paralectotype \; 22–25 M. pallidus (Uruguay, Punta Espinillo); 26–28
Ebrechtellatricuspidata(Russia,SakhalinArea,MoneronIs.).18,22,26Female,dorsal;19,23Ditto,ventral;20,24Femalelegs
Iⅈ25,27Femalecarapace,frontal;21,28Male,ventral.
P.T.Lehtinen&Y.M.Marusik 181
Figs.29–31: Dorsalviewofmale.29Misumenopsmaculissparsus,lectotype;30M.pallidus(Uruguay,PuntaEspinillo);31Ebrechtellatricuspidata
(Russia,SakhalinArea,MoneronIs).
Two females and one juvenile specimen from 0.68 + 1.05 + 0.9 + 0.68, total 4.81; IV: 1.68 + 0.68 +
Colombia,leg.Nolken(NMWacquisitionno.1873I15) 1.1 + 1.18 + 0.65, total 5.29.
mostprobablyformedpartoftheoriginalsyntypes,but CarapacepatternasinFigs.22&25,yellowwithtwo
as all the original labels were changed 60–70 years ago, wide submarginal brown bands; median yellow band
the possible original handwriting of Keyserling could withbrownmedianstripe,eyefieldwhite.Longsetaeon
notbechecked.However,thesespecimensbelongeither carapace concentrated on clypeus, U-shaped row along
to ‘‘Misumena’’ pallens Keyserling, 1880 or to a closely lightcentralbandofcarapaceandtransverserowbehind
related species. this. Very short hair-like setae present on margin of
Misumenops pallidus and ‘‘Misumena’’ pallens appear carapace and around ocular tubercles of lateral eyes.
more different when seen through a dissecting micro- Basal part of chelicerae with low raised tubercle, setae
scopethanbyjustlookingatstronglypressedspecimens onanteriorfaceshort.Sternumandmouthpartsyellow.
on slides. ‘‘Misumena’’ pallens does not share the most Abdomenpentagonal,withpatternformedbydarkand
importantgenericcharactersofMisumenops:darktarsal whitish transverse stripes and bands; lateral band dark;
annulionmalelegsI–II,alongsetaontheinnerfaceof venter light coloured with wide blackish median band
RTA, etc. It has a modified tip of RTA (as many other between epigastric furrow and spinnerets (Fig. 23),
Misumenini, including Runcinia and Runcinioides), but femalewithillustratedepigynehaspostepigastricareaas
the ultrastructure is essentially different, rather a distal in M. maculissparsus. Legs light brown, ventral side of
rod with ridges than a serrate margin of the tip (cf. also femora I and II with numerous dark spots, ventral side
Rinaldi, 1983: figs. 13–14). oftibiaealsowithsomespots,segmentsfrompatellaeto
The coloration is variable in all species of Mis- metatarsi with dark distal parts (Fig. 24), but not on
umenini, but the weak dorsal patterns of the generally tarsi as in male. Subcutaneous guanine pattern extends
pale‘‘M.’’pallensandverydistinctbrownpatternofM. to posterior part of carapace as a small spot and
pallidushardlyoverlap,whenthedetailsarestudied.The irregularly around and inside dark distal areas of patel-
presence of repeated confusion makes the lectotype lae and tibiae I–II. Leg spination: femur I with 3
selection for both Misumena pallens (see p.196) and prolateral and 1 or 2 dorsal spines, femur II with 1 or 2
Diaea pallida necessary. dorsalspines;tibiaIwith5or6pairs,metatarsusIwith
Diagnosis: This species can easily be distinguished 6 or 7 pairs of ventral spines. Epigyne as in Figs. 7–9 &
from the type species by the thinner (longer than wide) 33, with triangular apical hood and transverse elongate
tibialapophysis(RTA–ITA)closertoVTA(Figs.5–6cf. receptacula.
Figs. 1–2), higher position of the embolic base Male:Totallength2.35.Carapace1.1long,1.15wide
(10o’clock), relatively thicker seminal duct (as wide as (Fig.30),patternsimilartofemale.LegII:1.9 + 0.65 +
embolic base) (Fig. 5 cf. Fig. 1), longer tutaculum (Fig. 1.6 + 1.43 + 0.75, total 6.33; III: 0.75 + 0.33+0.68 +
11 cf. Fig. 3), and by the coloration of both sexes (Figs. 0.55 + 0.43, total 2.74; IV: 0.88 + 0.3 + 0.65 + 0.6 +
22–25 & 30 cf. Figs. 18–21 & 29). The general appear- 0.43, total 2.86. Abdomen ovoid, with pattern of trans-
ance of the epigynes is practically indistinguishable (cf. verse wide dark bands, sides with dark band; venter
Figs. 7 & 4). Misumenops pallidus differs from the other yellow with pair of small dark spots behind epigastric
closely related species M. guianensis by the less and furrow and larger spot just before spinnerets. Terminal
differently modified tip of RTA and the presence of parts of legs I and II segments darkened. Femur II with
brown spots on femora I–II (Fig. 24), and from M. 5 dorsal spines. Palp as in Figs. 5–7 & 10–13, light
temibilis also by the shorter and partly striated embolus coloured; tibia with two apophyses, tegulum without
(Fig. 13 cf. Fig. 39). apophyses, embolus making half a circle. Ventral tibial
Description: Female: Total length 5–5.75. Carapace apophysis short, wider than long, close to lateral apo-
2.1–2.25 long, 2.0–2.25 wide. Leg I: femur 2.8, patella physisatbaseoftibia(Fig.5).Lateralapophysisslightly
1.25,tibia2.2,metatarsus1.98,tarsus1.3,total9.36;II: swollen near tip, longer than wide, mesally with large
2.55 + 1.13 + 2.05 + 1.88 + 1.07, total 8.68; III: 1.5 + oval cavity, tip gently serrate (Fig. 12). Cymbium with
182 RedefinitionofMisumenops
Figs.32–35: 32Misumenopsmaculissparsus,paralectotype\;33M.pallidus(Uruguay,PuntaEspinillo);34–35Ebrechtellatricuspidata(Moneron
Is.).32–34Epigynalarea;35Leftmalepalp,ventral.
basal tutaculum longer than in type species. Tegulum 2006bythestaffandmostprobablylost,asmanyother
round, with heavily sclerotised apical rim, starting near types of Taczanowski. However, a neotype is not desig-
baseofembolusandfollowingwholecourseofembolus. nated, as at least the male of this species seems to be
Seminal duct thick, as broad as embolic base. Embolus identifiable.
starts from pocket at 10o’clock position (in left palp) Material examined: VENEZUELA: Guarico, Miranda: 1_ 1
(Figs. 5 & 10), its base clearly separated from tegulum. subad. \, Hato Masaguaral, dry meadow, 29 November 1977; 1_,
Can˜oCarascol,galleryforestofRioGuarico,29November1977;1\
Embolicbaserelativelywide,widthofemboluscontinu-
2juv.,NofCorozoPando,savanna,29–30November1977,allleg.&
ouslynarrowingalongitscourse,tipflatandnotpointed
det.P.T.Lehtinen(PTL).PARAGUAY:Fonciére,leg.E.Reimoser,
(Fig. 13). 1908,44_10\+numerousjuvs;1_withdeformedrightpalp,1_with
Distribution: Bolivia, Paraguay, Uruguay, Brazil and outstandingprocessonRTA(NMW,MLP,MNHN,MZT&PTL).
northern Argentina (Colombia not checked).
Diagnosis: The colour pattern of both sexes of M.
Biology: Some samples were collected from flowering
guianensis is usually simpler than that of both M.
garden bushes, as many other Misumenini.
maculissparsus and M. pallidus. Legs I–II of male with
Discussion:Differencesbetweensomespecimensofthe
darkannulations,butnospottedareas.Thelongitudinal
twosiblingspeciesincolourpatternmaypartlybedueto
dark bands on the carapace have more or less parallel
infraspecific variation, and partly to fading of the old
margins, not irregularly serrate as in M. maculissparsus
type material. Great colour variation is exceptionally
and especially M. pallidus. The abdominal pattern is
foundbetweenfreshspecimensofasinglepopulation.A
quite weak or lacking in females, simple in males. The
largesamplefromRioSelva,Bolivia,collectedinAugust
tip of RTA is surrounded by a continuous lamina on
2007 included, e.g. several annulation patterns of male
both sides (Fig. 40), consisting of tightly placed very
legsandevenanadultmalewithuniformlypalelegs,as
distinct lobules with narrow clefts between them (Fig.
well as some females without dorsal pattern. The only
38),incontrasttotheserrationsononlytheoutersidein
reliable diagnostic characters are found in the structure
M. temibilis (Fig. 37) and covering a much wider area
of the male palp. It is not clear if there are any distinct
than the corresponding modifications in M. pallidus
differences in the vulvae, because we did not dissect the
(Fig. 12) and probably M. maculissparsus (but no SEM
epigyneoftheparalectotypeof M.maculissparsus.
micrographs possible for type). The male embolus
is mostly smooth, distally tapering and has only
a triangular subapical lamina and some indistinct
Misumenops guianensis (Taczanowski, 1872) (Figs. 38, ridges in the distal third (Fig. 47); it originates from a
40, 45, 47) tegular pocket at 9o’clock (in left palp) (Fig. 45). The
tutacular apophysis is reduced to an indistinct knob
ThomisusguianensisTaczanowski,1872:90(_\).
Diaeaguyanensis:Keyserling,1880:112,pl.2,fig.62(_\). with a row of transversely placed setae. The epigynal
Misumenopsguianensis:F.O.P.-Cambridge,1900:141. hood is small, as in the related species, and thus not
Misumenopsguyannensis:Mello-Leitão,1929:233:commoninBrazil! diagnosticincomparisonwithM.maculissparsusandM.
(misspelling).
pallidus.
Types: Male and female syntypes from French Description (_/\): Total length 3.1–3.3/5.7. Carapace
Guiana, originally in IZPAN, not found in February 1.57/2.48long,1.38/2.05wide.Abdomen1.81/3.04long,