a systematic revision of goniosomatinae (arachnida ... · a systematic revision of goniosomatinae...

A systematic revision of Goniosomatinae (Arachnida :Opiliones : Gonyleptidae), with a cladistic analysisand biogeographical notes

Márcio Bernardino DaSilvaA and Pedro GnaspiniA,B

ADepartamento de Zoologia, Instituto de Biociências, Universidade de São Paulo,Rua do Matão, trav. 14, n� 101, 05508-090 São Paulo, SP, Brazil.

BCorresponding author. Email: [email protected]

Abstract. Goniosomatine harvestmen have strongly armed pedipalps, generally large bodies and, commonly, very longlegs (sometimes more than 20 cm), and are distributed in the Brazilian Atlantic forest, from southern Bahia to SantaCatarina. Since they are conspicuous animals and individuals of some species tend to concentrate in caves (and also underrock boulders), they have been (and still are) the target of several studies, especially those focusing on reproductive anddefensive behavior, population ecology, physiology, chromosomes, etc. In spite of their importance for biological studies(some species constitute important and frequently used models for these studies), the taxonomy of Goniosomatinae hasfaced some problems, including misidentification, a large number of undescribed species and the lack of a phylogenetichypothesis for the relationships among its species (which would allow evolutionary studies to be made). The lasttaxonomic changes in the subfamily were made 60 years ago. Considering a taxonomic revision and cladistic analysis ofthe subfamily to be of paramount importance, the main scope of the present paper is to provide a cladistic analysis andtaxonomic revision of the species of Goniosomatinae and a new arrangement of genera (and species). The main taxonomicchanges are given as follows. Six genera are recognised within the subfamily: Goniosoma; the newly described genusPyatan; the reestablished genera Serracutisoma, Heteromitobates and Mitogoniella; and Acutisoma. New genericsynonyms include: Glyptogoniosoma=Goniosomella =Lyogoniosoma =Metalyogoniosoma =Xulapona =Goniosoma,Acutisomelloides =Pygosomoides= Spelaeosoma= Serracutisoma; and Acutisomella =Heteromitobates. Newlydescribed species include: Goniosoma capixaba; G. apoain; Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini;Serracutisoma pseudovarium; S. fritzmuelleri; S. guaricana; Heteromitobates anarchus; H. harlequin; H. alienus;Mitogoniella taquara; M. unicornis; and Acutisoma coriaceum. New combinations include: Goniosoma macracanthum(Mello-Leitão, 1922); G. unicolor (Mello-Leitão, 1932); G. carum (Mello-Leitão, 1936); Serracutisoma proximum(Mello-Leitão, 1922); S. banhadoae (Soares & Soares, 1947); S. molle (Mello-Leitão, 1933); S. thalassinum (Simon,1879); S. catarina (Machado, Pinto-da-Rocha & Ramires, 2002); S. inerme (Mello-Leitão, 1927); S. spelaeum (Mello-Leitão, 1933); Heteromitobates inscriptus (Mello-Leitão, 1922); H. albiscriptus (Mello-Leitão, 1932); Mitogoniellamodesta (Perty, 1833); and M. badia (Koch, 1839). Reestablished combinations include: Mitogoniella indistincta Mello-Leitão, 1936 and Acutisoma longipes Roewer, 1913. New specific synonyms include: Acutisomella cryptoleuca=Acutisomella intermedia =Goniosoma junceum=Goniosoma patruele =Goniosoma xanthophthalmum=Metalyogoniosoma unum=Goniosoma varium, Goniosoma geniculatum =Goniosoma venustum; Goniosomellaperlata =Progoniosoma minense=Goniosoma vatrax, Glyptogoniosoma perditum =Progoniosoma cruciferum =Progoniosoma tijuca =Goniosoma dentipes; Leitaoius iguapensis= Leitaoius viridifrons = Serracutisoma proximum;Acutisoma marumbicola=Acutisoma patens = Serracutisoma thalassinum; Progoniosoma tetrasetae= Serracutisomainerme; and Acutisoma monticola=Leitaoius nitidissimus = Leitaoius xanthomus =Mitogoniella mutila=Acutisomalongipes. The following species are considered species inquirenda: Goniosoma lepidum Gervais, 1844;G. monacanthum Gervais, 1844; G. obscurum Perty, 1833; G. versicolor Perty, 1833; and Mitogoniella badia (Koch,1839). The monotypic genus Goniosomoides Mello-Leitão, 1932 (and its species, G. viridans Mello-Leitão, 1932)is removed from Goniosomatinae and considered incertae sedis.

Introduction

The systematics of Neotropical Opiliones was establishedmainly by the works by Roewer (e.g. 1913, 1923, 1930) andMello-Leitão (e.g. 1923, 1932) during the early 20th century.In the ‘Roewerian system’, some characters were arbitrarily

selected to establish the different families, another arbitrary setof characters was used to establish subfamilies and yet anotherto establish genera and species, creating a closed system. Thissystem yielded a confusing and subjective taxonomy, with alarge number of species and genera, most of them monotypic

� CSIRO 2009 10.1071/IS09022 1445-5226/09/060530

CSIRO PUBLISHING

www.publish.csiro.au/journals/is Invertebrate Systematics, 2009, 23, 530–624

(Kury 1990; Pinto-da-Rocha 1997, 2002). From the 1940s thatscenario began to change with the works by Soares and Soares(e.g. 1948), with an extensive number of synonyms established,and an important effort to reorganise the systematics ofharvestmen (mainly the Neotropical ones). Only recently, in acombined effort to reorganise the knowledge on the taxonomyand evolution of Neotropical harvestmen, a larger number oftaxonomic revisions, mainly based on cladistic analyses, haveappeared at different taxonomic levels (e.g. Pinto-da-Rocha1997, Stygnidae; Pinto-da-Rocha 2002, GonyleptidaeCaelopyginae; Tourinho and Kury 2003, Sclerosomatidae[Eupnoi] genus Jussara; and several unpublished M.Sc. andPh.D. Theses).

The Atlantic rainforest (which includes forested areaslocated in the eastern to south-eastern coast of Brazil) harboursthe highest diversity of harvestmen in the world, with about600 described species (Pinto-da-Rocha et al. 2005). Almost allof this richness is represented by species of the large familyGonyleptidae (with about 830 species – Kury 2003 and somedescriptions published afterwards), which is organised in16 subfamilies, of which nine are exclusive of this biome. Inaddition, most of these species show a high degree of endemism,with distributions mostly related to only one or a few mountainranges or valleys. These characteristics make harvestmen agood model for historical biogeographical studies (e.g. Pinto-da-Rocha et al. 2005). One of these endemic subfamilies isGoniosomatinae (with 46 species recorded before the previousstudy – Kury 2003).

Goniosomatinae include harvestmen with strongly armedpedipalps, generally with large bodies, and many of them withvery long legs (sometimes more than 20 cm – for example, seeGnaspini 1999), distributed in the Brazilian Atlantic forest, fromsouthern Bahia to Santa Catarina. Since they are conspicuousanimals and individuals of some species tend to concentrateinside caves (and also under boulders), they have been thetarget of several studies, especially those focusing onreproductive and defensive behavior, population ecology,physiology, chromosomes, etc. (e.g. Gnaspini 1995, 1996;Gnaspini and Cavalheiro 1998; Machado and Oliveira 1998;Gnaspini et al. 2003; Willemart and Gnaspini 2004a, 2004b;Ferreira et al. 2005; Oliveira et al. 2006). They have alsoprovided important biogeographical information for historicalarea relationships (Pinto-da-Rocha et al. 2005).

In despite of its importance for biological studies, thetaxonomy of Goniosomatinae has faced some problems,including misidentification, a large number of undescribedspecies and the lack of a phylogenetic hypothesis for therelationship among its species (which would allow evolutionarystudies to be made). The last major taxonomic changes madein the subfamily were those by Soares and Soares (1948) and,more recently, the transference of Microgoniosoma toPachylinae (as a junior synonym of Discocyrtus–Ringuelet1954) and the transfer of Heteromitobates from Mitobatinae byKury (2003). A first revision concerning the subfamily wasprepared by Stefanini-Jim (1985, 1995) in her M.Sc. andPh.D. theses, which remain unpublished. Since several speciesof the subfamily are still the focus of, and constitute importantmodels for, several biological studies, a taxonomic revisionand cladistic analysis of the subfamily were considered to be

of paramount importance. Therefore, the main scope of thepresent paper is to provide a cladistic hypothesis ofrelationships and a taxonomic revision of the species ofGoniosomatinae.

Goniosomatinae – a brief historicaland taxonomic review

Mello-Leitão (1935a) established the subfamily Goniosominaewith 13 genera (previously included in Gonyleptinae):Goniosoma Perty, 1832; Progoniosoma Roewer, 1913;Acutisoma Roewer, 1913; Lyogoniosoma Mello-Leitão, 1926;Leitaoius Roewer, 1930; Acutisomella Roewer, 1930;Serracutisoma Roewer, 1930; Microgoniosoma Mello-Leitão,1930; Glyptogoniosoma Mello-Leitão, 1932; AcutisomelloidesMello-Leitão, 1932; Goniosomoides Mello-Leitão, 1932;Spelaeosoma Mello-Leitão, 1933; and Pygosomoides Mello-Leitão, 1933. Four genera were included afterwards: XulaponaMello-Leitão, 1936; Goniosomella Mello-Leitão, 1936;MitogoniellaMello-Leitão, 1936; andMetalyogoniosoma Soares& Soares, 1946. Many of the genera were monotypic. Three mainsynonyms were proposed by B. Soares (1944a) and Soares andSoares (1948), reducing 14 genera to three: Acutisoma(=Leitaoius=Acutisomella=Serracutisoma = Glyptogoniosoma=Acutisomelloides=Pygosomoides=Mitogoniella); Goniosoma(=Progoniosoma=Spelaeosoma); and Lyogoniosoma(=Goniosomella=Xulapona). Therefore, in their largemonographic work about Neotropical harvestmen, Soares andSoares (1948) included only six genera in the subfamily: Acutisoma(20 species), Goniosoma (18 species), Goniosomoides (monotypic),Lyogoniosoma (3 species), Metalyogoniosoma (monotypic) andMicrogoniosoma (monotypic). Since its species Microgoniosomafuscum Mello-Leitão, 1930 was recognised as a junior synonym ofDiscocyrtus testudineus (Holmberg, 1876) by Ringuelet (1954),Microgoniosoma is presently placed in Pachylinae as a juniorsynonym of Discocyrtus Holmberg, 1878.

In addition, the following two genera were once includedin Goniosomatinae and afterwards transferred toanother subfamily. Cadeadoius Mello-Leitão, 1936 wasdescribed in Goniosomatinae and is presently placed inProgonyleptoidellinae (by Soares and Soares 1985).Metagoniosoma Roewer, 1917 was described in Gonyleptinae,transferred to Goniosomatinae by Mello-Leitão (1935b), back toGonyleptinae by B. Soares (1944c) and is presently considered ajunior synonym of Gonyleptes Kirby, 1818 (by Kury 2003).

Stefanini-Jim (1985) made an important historical reviewand reorganisation of species within the subfamily, andcorrected the subfamily name from Goniosominae toGoniosomatinae. She also discussed that the inclusion ofspecies in genera seemed artificial. In her unpublished theses,Stefanini-Jim (1985, 1995) proposed the synonymisation of allgenera in a single genus, Goniosoma Perty, 1833, and severalsynonyms among species, one of the largest cases being thesynonymisation of almost all species from Santa Catarina andParaná with Goniosoma badium Koch, 1839 (which hasbeen already discussed by Gnaspini 1993, 1999). Herproposed synonyms were made by especially considering theresemblance of genitalic characters, and generally disregardingmorphological and colour pattern differences. Although not

Systematic revision of goniosomatine harvestmen Invertebrate Systematics 531

published, her taxonomic changes have been largely used so far(based both on identifications made by that author or byother authors using her 1985 thesis), especially the use ofGoniosoma as the single genus in the subfamily (e.g. Pinto-da-Rocha 1993; Gnaspini and Trajano 1994).

The main historical steps in the taxonomy of Goniosomatinaecan be summarised as follows:

XIX century – 16 species described in the genus Goniosoma – typespecies: G. varium Perty, 1833.

Beginning of XX century –Roewerian era – large number of descriptions(6 genera and 6 species described byRoewer, 8 genera and 25 species byMello-Leitão and 2 species by Piza) – Goniosomatinae Mello-Leitão,1935.

1940 to 1944 – 14 genera and 49 species.1948 – Soares and Soares – several synonyms – 6 genera and 41 species +3 new species.

1985 – Stefanini-Jim (unpublished) – 1 genus and 25 species + 1 newspecies.

Present study – 6 genera and 25 species + 11 new species.

Methods

Material studied

We studied material (including undetermined specimens andtypes) especially from the following Brazilian collections:Museu de Zoologia da Universidade de São Paulo (MZSP);Museu Nacional da Universidade Federal do Rio de Janeiro(MNRJ); collection H. E. M. Soares (HS) (presently in MNRJ)and Instituto Butantan (IBSP); and specimens collected duringthe present study. In addition, we also examined types and othermaterial from the following collections: Institut Royal desSciences Naturelles de Belgique, Brussels, Belgium (ISNB);Naturhistorisches Museum Wien, Austria (NHMW); Museumfür Naturkunde-Universität zu Berlin, Germany (ZMHB);Forschungsinstitut und Naturmuseum Senkenberg, Frankfurt,Germany (SMF); collection C. F. Roewer (ROEWER)(presently in SMF); Escola Superior de Agricultura ‘Luiz deQueiroz’, Universidade de São Paulo, Piracicaba, Brazil(MZLQ); Museu de História Natural Capão da Imbuia,Curitiba, Brazil (MHNC); and Zoologisk Museum, StatensNaturhistoriske Museum, Kobenhavns Universitet, Denmark(ZMUK).

The following types (following the original names) wereexamined.

Acutisoma banhadoae Soares & Soares, 1947 – male paratype – MZSPA. inerme Mello-Leitão, 1927 – male allotype – HSA. inscriptumMello-Leitão, 1922 – syntypes (genotype ofAcutisomella) –MZSP

A. longipes Roewer, 1913 – male holotype – SMFA. marumbicola H. Soares, 1945 – paratypes – MZSPA. monticolum Mello-Leitão, 1922 – female holotype – MZSPA. patens Roewer, 1930 – syntypes – SMFA. proximum Mello-Leitão, 1922 – male lectotype and paralectotypes(genotype of Serracutisoma) – MZSP

Glyptogoniosoma perditumMello-Leitão, 1936 –male holotype –MNRJGoniosoma grossum Koch, 1839 – female syntypes – ZMHBG. modestum Perty, 1832 – syntypes – ZMHBG. xanthophthalmumMello-Leitão, 1931 –male allotype and paratypes –MNRJ

G. varium Perty, 1832 – syntypes – ZMHB

G. vatrax Koch, 1848 – syntypes – ZMHBG. venustum Perty, 1832 – syntypes – ZMHBItatiaya hamata Roewer, 1928 – holotype (genotype) – ISNBMetalyogoniosoma unum Soares & Soares, 1946 – immature maleholotype (genotype) – MZSP

Mitogoniella indistincta Mello-Leitão, 1936 – syntypes (genotype) –

MNRJProgoniosoma calcar Roewer, 1913 – male holotype – SMFProgoniosoma tetrasetae Roewer, 1930 – male holotype – SMFProgoniosoma tijuca Roewer, 1930 – female holotype – SMFSpelaeosoma spelaeumMello-Leitão, 1933 –male holotype (genotype) –MNRJ

Xulapona cara Mello-Leitão, 1936 – male and female syntypes(genotype) – MNRJ

The following type material was not examined.

Goniosoma lepidum Gervais, 1844, G. monacanthum Gervais, 1844and G. obscurum Perty, 1832 – types are lost (after Soares and Soares1948).

Goniosoma albiscriptum Mello-Leitão, 1932, G. badium Koch, 1839,G. dentipes Koch, 1839, G. junceum Perty, 1832, G. patruele Perty,1832,G. roridumPerty, 1832,G. versicolor Perty, 1832,Goniosomellaperlata Mello-Leitão, 1936, Glyptogoniosoma unicolor Mello-Leitão,1932, Leitaoius ornatus Mello-Leitão, 1934 and L. viridifrons Mello-Leitão, 1935 – types are probably lost (we were not able to find them,and some were not deposited in the collection mentioned in theoriginal or subsequent description).

Acutisoma inermeMello-Leitão, 1927, A. marumbicola H. Soares, 1945,Acutisomella cryptoleucaMello-Leitão, 1940,Acutisomella intermediaMello-Leitão, 1937, Goniosoma geniculatum Mello-Leitão, 1931,G. xanthophthalmum Mello-Leitão, 1931, Leitaoius iguapensis Piza,1938, L. guttulatus Mello-Leitão, 1934, L. xanthomus Mello-Leitão,1935, Mitogoniella mutila Piza, 1938, Progoniosoma cruciferumMello-Leitão, 1923, P. macracanthum Mello-Leitão, 1922 andP. minense Mello-Leitão, 1932 – this material was examined byR. Stefanini-Jim (1985) but could not be located; it may have possiblybeen taken with the HS collection to the MNRJ and should be checkedin the future.

Acutisoma acutangulum and A. thalassinum are presently deposited atMuséum National d’histoire Naturelle, Paris, France (MNHN), butwere not lent for study. Dr. A. Muñoz-Cuevas provided photographsof A. acutangulum.

The monotypic genus Goniosomoides Mello-Leitão, 1932 (and itsspecies, Goniosomoides viridans Mello-Leitão, 1932) is removedfrom Goniosomatinae in the present study and considered incertaesedis. Based on the analysis of the original description, we concludedthat it does not belong in Goniosomatinae. However, sincethe type could not be examined and the illustration does not allowa precise identification, we could not precisely place thespecies, although it seems to belong either in Caelopyginae orProgonyleptoidellinae.

Aiming at completeness of information, we provide below a listof species that were described in Goniosoma (or described inAncistrotus and transferred toGoniosoma – in this case they aremarkedwith *) and afterwards transferred to other genera; which mostlyhappened before Goniosomatinae was erected. We indicate under‘comb.’ (or ‘syn.’, where the case is a synonymy) the authorresponsible for the transference to the genera where the species ispresently placed and, if not the same, we indicate under ‘transf.’ theauthor responsible for the first transference from Goniosoma.Since these species belong to genera presently placed in other sub-families or even families, they were not examined during the presentstudy.

Metamitobates squalidus (Perty, 1832) – transf. Koch (1839), comb.Kury (1992) – Gonyleptidae, Mitobatinae.

532 Invertebrate Systematics M. B. DaSilva and P. Gnaspini

Mitobates triangulus Sundevall, 1833 =Mitobates conspersus (Perty,1833) – transf. Koch (1839), syn. Simon (1879) – Gonyleptidae,Mitobatinae.

* Neoancistrotus bifurcatus (Koch, 1839) – transf. Soerensen (1884),comb. Kury (1992) – Gonyleptidae, Mitobatinae.

* Promitobates hexacanthus (Koch, 1839) – comb. Roewer (1913) –Gonyleptidae, Mitobatinae.

Leptocnema sulphurea (Perty, 1832) – comb. Koch (1839) – Gonylepti-dae, Progonyleptoidellinae.

Phareus raptator (Gervais, 1844) – comb. Simon (1879) – Stygnidae.Stygnus ferrugineus (Perty, 1832) – comb. Koch (1839) – Stygnidae.Cranaus chlorogaster (Gervais, 1844)– comb.Simon (1879)–Cranaidae.Cranaus cinnamomeus (Gervais, 1844) – comb. Simon (1879) –

Cranaidae.Phareicranaus calcariferus (Simon, 1879) – comb. Roewer (1913) –Cranaidae.

Santinezia curvipes (Roewer, 1916) =GoniosomapavaniMuñoz-Cuevas,1972 – syn. Pinto-da-Rocha and Kury (2003) – Cranaidae.

Observation and illustration

External characters were studied under a stereomicroscopeand the male genitalia and some other details were studiedunder a scanning electron microscope (SEM). To preparethe genitalia for the SEM analysis, we used the proceduredescribed by Pinto-da-Rocha (2002), but we dried thematerial using a ‘Critical Point Dryer BAL-TEC CPD 030’(Bal-Tec A.G., Balzers, Liechtenstein). Body and appendagemeasurements were taken using a caliper and measurements ofgranules were made under the stereomicroscope.

Terminology

In the present study, we followed traditional names for generalmorphology and description schemes used by previousstudies, especially on laniatoreans (e.g. Pinto-da-Rocha 1997,2002; Shultz and Pinto-da-Rocha 2007). However, since thereare different specifications of terms usually adopted amongharvestman researchers, and since armature and bodyornamentation are important for the establishment of speciesand genera, we present the terms as they are used herein.

granule = short elevation (height = diameter) usually present in largenumbers in the same structure covering it rather homogeneously(see Figs 1–5, 58, 86).

tubercle = blunt elevation somewhat taller than granules when comparedon the same structure (see Figs 27, 58, 59).

spine = pointed elevation generally taller than tubercles (see Figs 31,55, 83).

apophysis = irregularly shaped structure, generally larger than thoseabove, which occurs only at coxa, trochanter and apex of the femurof the appendages (see Figs 12, 58).

Considering these elevations, a structure may be:

armed=with spines.unarmed =without spines, but may present granules or tubercles.smooth =without granules, tubercles or spines.

Please note that setae and apophyses are not considered inthis classification. Therefore, for instance, a coxa may bedescribed as smooth but it will always carry apophyses.

In each leg, six rows of spines/tubercles can be recognised(Fig. 10), and were named as follows (using the femur as anexample): row1= dorsal–retrolateral row, starting near the tooth

of the articulation between trochanter and femur and endingnear the retrolateral apical apophysis; row2 = retrolateralrow, ending near the articulation between femur and patella;row3 and row4 = respectively ventral–retrolateral and ventral–prolateral rows, ending at the ventral membrane of articulationbetween femur and patella; row5 = prolateral row; androw6= dorsal–prolateral row, ending near the prolateral apicalapophysis.

Granules

In some structures, the distribution of granules representsone of the most important characters for taxonomicrecognition and phylogenetic inference. The shape of granulesusually varies independently among different subareas of thesame body region. For instance, the dorsal scute may bear smallgranules whereas its posterior margin may present large ones.

Therefore, granulation was analysed considering the mostpossible number of units of the body, and the combination ofthe description of the granulation in each unit would result inan adequate diagnostic feature for the whole body. The mostimportant areas analysed under a granulation perspectiveand used both in the taxonomic and phylogenetic sectionswere: the dorsal scute areas, central area of the carapace(defined by sulcus I), lateral margin of the dorsal scute,posterior margin of the dorsal scute + free tergites, posteriormargin of the stigmatic area + free sternites and the ventralsurface of coxae IV.

Two independent features were noted: number and size ofgranules. Concerning number, the exact number of granules onthe dorsal scute areas and on the central area of the carapacewere counted, since they are easily observed, and this numbermay be used for species recognition. For the other regions, weestimated the density of granules in three levels: low, mediumand high. For rows of granules (common on margins and onthe free tergites and sternites), we considered density to be lowwhen we detected up to 2 granules per mm, medium for2–4 granules per mm and high for 4 or more granules per mm.For areas (as in coxae IV), we considered density low whenwe detected up to 4 granules per mm2, medium for 4–8 granulesper mm2 and high for 8 or more granules per mm2. Concerningsize, we recognisedfive levels:minute, small, medium, large, andvery large, respectively presented in Figs 1–5.

Pedipalp

For ornamentation of femur, tibia and tarsus of pedipalps,we followed Pinto-da-Rocha (2002): in tibia and tarsus,‘I’ designates high spines and ‘i’ designates short spines(height equal or less than half of the highest spine in thearticle); in femur, ‘I’ designates spines and tubercles and‘i’ designates short tubercles (same as above). However, sincethe armature is more complex in Goniosomatinae than amongCaelopyginae, we included the size of the base of the spines(in tibia and tarsus) in our measurements, contrary to Pinto-da-Rocha (2002), who used only the size of the setae. Since the totalnumber of spines and tubercles is similar among the species, thecharacter that will discriminate among them is the number of ‘I’sin femur (as in character 72). Since some granules, tuberclesand/or spines may be absent in some individuals of the same


species, we describe a ‘general armature pattern’ using thosestructures that appear in at least half of the individuals analysed.

Colour

We recognised three factors that determine the generalcolour pattern (the epidermic pigmentation below the cuticle,the cuticle itself (which varies between green and brown) andthe white dry-mark, but only the last two were used in ourcladistic analysis. Epidermic pigmentation varies largely indistribution and density among individuals (Figs 193, 209,212, 224, 228). It is frequently associated to the areas of thedorsal scute, and, as a consequence, the depression betweenareas and the longitudinal medial line appear completelydepigmented. It is also common to observe a lack of pigmentsforming a characteristic pattern behind the ocularium, with thearea ahead of the ocularium showing a large amount ofpigments, frequently becoming blackish. We here use the ‘dry-mark’, a term defined by A.B. Kury (pers. comm.) for a patch

in the more external serose layer of the cuticle that formswhite drawings in the animal (Fig. 190) when it dries up whenremoved from liquid. On the dorsal scute, it is generallyassociated with the depigmented lines. Since there is a typicalcombination between the epidermic pigmentation andthe dry-mark, generally forming a striped area behind theocularium, this pattern will not be repeatedly described, and itwill be herein named ‘mask’ (Figs 190, 193, 212, 224, 228).

Cladistic analysis

We used, as our ingroup, 36 species recognised as belongingto Goniosomatinae. The names used herein already take intoaccount the resulting cladogram and the new taxonomicscheme resulting from the cladistic analysis and from thesynonyms recognised during the present study. The taxonomicchanges are presented under ‘Taxonomy’, ahead in this paper.As outgroups, we used 11 species from 7 subfamilies ofGonyleptidae: Discocyrtus invalidus Piza, 1938 and Roeweria

1 2 3

4 5 6

7 8 9

Figs 1–9. 1–5, Examples of different sizes of granules on area II of the dorsal scute. 1, Minute (Goniosoma roridum).2, Small (Goniosoma varium). 3, Medium sized (Acutisoma longipes). 4, Large (Serracutisoma pseudovarium, sp. nov.).5, Very large (Serracutisoma inerme). 6–9, Apex of tarsus and detail of claws (cl). 6, 7,Heteromitobates discolor, showingpectinate claws. 8, 9,Goniosoma varium, showing smooth claws. A progression in size of the tarsal process (tp) can be seenfrom Fig. 6 to 8 and to 16 (16: see next plate). Scale = 0.05mm.


virescens (Mello-Leitão, 1940) (Pachylinae); Longipernacancellata (Roewer, 1913) and Promitobates viridigranulatus(Soares & Soares, 1946) (Mitobatinae); Neosadocus variabilis(Mello-Leitão, 1935) and Sphaerobunus pulcher (Mello-Leitão,1922) (Gonyleptinae); Acrogonyleptes unus (Mello-Leitão, 1927) (Hernandariinae); Progonyleptoidellus striatus(Roewer, 1913) (Progonyleptoidellinae); Zortalia leprevosti

Soares & Soares, 1947 (Sodreaninae); and Caelopygus elegans(Perty, 1833) and Metarthrodes pulcherrimus (Mello-Leitão,1931) (Caelopyginae).

We used characters that have been traditionally used incladistic analysis of harvestmen, especially concerningNeotropical groups (e.g. by Pinto-da-Rocha 1997, 2002), andwe added characters that seemed to be useful for the analysis of

10

11

12

15

1813

14

16 17

Figs 10–18. 10, Cross-section of a leg of a generalised goniosomatine showing the six basic rows (numbers) of granules/spines (see text) (unscaled). 11–13, Goniosoma varium. 11, Habitus, dorsal view. 12, Right femur IV (dorsal view);ap = apophysis. 13, Eye mound (posterior view), with a pair of medium-sized spines. 14–16,Goniosoma capixaba, sp. nov.14, Habitus, dorsal view. 15, Right femur IV (dorsal view). 16, Distitarsus IV (lateral view), showing a very large tarsalprocess (tp); cl = tarsal claw. 17, 18,Goniosoma venustum. 17, Habitus, dorsal view. 18, Right tibia IV (ventral view) – row 2and row3 are highlighted. Scale = 4mm, except for 13 = 1mm, 16 = 2mm.


the taxa herein studied. We tried to use as many informativecharacters as possible and we did not do any aprioristicexclusion of characters. For the establishment of charactersused in the analysis, we preferred the use of multistatecharacters (following, for instance, Wilkinson 1995). Thestates were previously ordered, when appropriate, based onsimilarity, and they were polarised simultaneously betweenoutgroups and ingroup (following, e.g. Nixon and Carpenter

1993). Therefore, we added some characters for the ‘resolutionof the outgroup’ in order to find a better relationship betweenoutgroups and ingroup in such a way that it reduces the problemsof character polarisation that may affect the topology of theingroup cladogram (Nixon and Carpenter 1993). Based on thefact that McGhee (1977) and Gnaspini (1999) demonstratedthat morphometric and meristic characters may help on thediagnosis of species among Opiliones (and also within

19

22

23

25

26

242120

27

Figs 19–27. 19–21, Goniosoma roridum. 19, Habitus, dorsal view. 20, Right trochanter and femur IV (dorsal, view).21, Trochanter, femur, patella and base of tibia of right pedipalp (ventral–retrolateral view). 22, 23, Goniosoma vatrax.22, Habitus, dorsal view. 23, Right femur IV (dorsal view). 24–27, Goniosoma macracanthum. 24, Habitus, dorsal view.25, Right femur IV (dorsal view). 26, Joint of right coxa and trochanter IV (prolateral view), showing prolateral apicalapophysis of coxa (paac) and prolateral basal apophysis of trochanter (pbat). 27, Eye mound (posterior view), with a pairof low tubercles (tb). Scale = 4mm, except for 21 = 2mm, 26 = 3mm, 27 = 1mm.


Goniosomatinae – Gnaspini 1999), we decided to incorporatesome of these data in our analysis. For that, continuous andquantitative characters were coded following Thiele (1993).Thus, characters related to counts or ratios (93–97) were codedas 10-state with 1/10 weighting; characters related to sizeand density of granulation were weighted 2/10 (75–77 and 80)or 3/10 (78–79 and 81–83). Most polymorphisms were treated as

ambiguities, and the two states were used for the samepolymorphic species; in character 49, we used the apomorphicstate in order to inform the presence of an apomorphic statein that given species. The characters related to the penis, themorphometric characters, most of those related to ornamentationand armature of legs, and some of those related to colour arevalid only formales; only one character (related to colour) is valid

2830

32

36

3537

33

34

3129

Figs 28–37. 28–31, Goniosoma dentipes. 28, Habitus, dorsal view – notice the reduced prolateral basal apophysis oftrochanter IV (pbat). 29, Right femur IV (dorsal view). 30, Femur of right pedipalp (ventral–retrolateral view). 31,Eye mound (posterior view), with a pair of very large spines (sp). 32–34, Goniosoma ensifer. 32, Apex of coxa andtrochanter IV. 33, Right femur IV (dorsal view). 34, Left tibia IV (retrolateral view); ras = retrolateral apical spine.35–37, Goniosoma apoain, sp. nov. 35, Habitus, dorsal view. 36, Right femur IV (dorsal view). 37, Right tibia and baseof metatarsus IV (ventral–retrolateral view) – notice armature of rows. Scale = 4mm, except for 30 = 2mm, 31 = 1mm.


only for females and the characters related to body granules,pedipalp, and tarsal claws are valid for both sexes. Somecharacters show variation within the species (as can be seen inthe descriptions in the ‘Taxonomy’ section), including somerelated to age (which were not used herein) and to sexualdimorphism (e.g. some related to male armature), and shouldbe viewed with caution to avoid misinterpretations (asdiscussed, e.g. by Goodnight and Goodnight 1953 andGnaspini 1999). Therefore, we tried to recognise ‘generalpatterns’ of each character for each species, and these patternswere used both for the recognition of species limits(in conjunction with other characters, such as colour andgenital morphology, for instance), and for the establishment ofthe states used in the cladistic analysis for those given characters.This is not the case when there is gender polymorphism, since,where appropriate, we used separate characters for males andfemales.

The charactermatrixwas edited usingNDE0.5.0 (Page 2001).The parsimony analysis was carried out using NONA 2.0(Goloboff 1999), with the interface Winclada 0.9.9 (Nixon1999) and PAUP 4.0b10 (Swofford 2002). Since the use ofexact search algorithms was not possible owing to the largenumber of taxa, we opted for a heuristic search algorithm. Weconducted 1000 independent replicates of Wagner additionfollowed by TBR (tree bisection and reconnection). Thecommands ‘hold10 000; mult*1000; hold/1000’ and‘mult*max*’ were used in NONA 2.0, and ‘hsearch addseq =random nreps = 1000 hold = 100’ in PAUP 4.0b10. Bremersupport indices (Bremer 1994) were calculated for each nodeof the proposed phylogenetic hypothesis: we used thecommand ‘constraint’ of PAUP 4.0b10 followed by theparenthetical notation of the clade in question.

List of characters

The characters used in the present cladistic analysis are listedin Appendix 1 and are summarised in the matrix of charactersshown in Table 1. Characters 75 to 83 and 93 to 97 are continuousand/or quantitative; characters 84 to 92 were used only forresolution at the outgroup level and do not vary withinGoniosomatinae. State 0 of each character is the plesiomorphicstate at the node including the whole Goniosomatinaesubfamily as a result of the present phylogenetic study. State 0of continuous characters (75–83 and 93–97) was established asthe smallest value found within the range. For multistatecharacters, we either indicated a hypothesis for thetransformation series between states (whenever we understoodthat we could recognise and hypothesise a linear connectionbetween states – and in this case the character was treated asadditive), or considered non-additive, as noted (but we alsoconducted a cladistic analysis considering all multistatecharacters as non-additive and compared the results). In ourdiscussion, we base the optimisation of characters on thephylogenetic hypothesis obtained herein (as in Fig. 233), andwe used the group+ concept of Amorim (1982) (which may bebriefly explained as follows: a clade may be referred to as thename of the ‘basal’ taxon followed by ‘+’ – example:G. varium+

refers to the clade (G. varium, (G. capixaba, (G. venustum,G. roridum))) as in Fig. 233).

Results of the cladistic analysis

We obtained a single tree (Fig. 233) of 5629 steps, CI= 0.29and RI= 0.68; Bremer support values are given for each clade(these numbers were generally higher at the generic level, and atthe Goniosomatinae node, giving support to the generic schemehere proposed). Table 2 shows the changes in character statefor each clade of the cladistic hypothesis obtained.

We made two additional analyses. When we removed thecontinuous and quantitative characters, we obtained 148 mostparsimonious trees (MPTs) of 4950 steps, CI= 0.30 andRI= 0.69, and the strict consensus tree had 5110 steps,CI= 0.29 and RI = 0.68; in this hypothesis, clade (G. dentipes+G. ensifer)+ (as in Fig. 233) collapses partially [becoming(G. dentipes, G. ensifer, clade G. apoain+)], cladeSerracutisoma collapses partially [becoming (S. proximum,clade S. banhadoae+, S. fritzmuelleri, clade S. catarina+)],clade S. banhadoae+ collapses [becoming (S. banhadoae,S. molle, S. pseudovarium, S. thalassinum)] and clade(H. discolor+H. inscriptus)+ collapses [becoming (H. discolor,H. inscriptus, H. albiscriptus, H. anarchus)]. When weconsidered all multistate characters as non-additive, weobtained 288 MPTs of 5012 steps, CI= 0.32 and RI = 0.66,and the strict consensus tree had 5140 steps, CI= 0.31 andRI= 0.65; in this hypothesis, clade Goniosomatinae isrearranged [becoming (Goniosoma, (Pyatan, Serracutisoma,clade Heteromitobates+))], clade Goniosoma is rearranged[becoming (G. vatrax, G. macracanthum, clade G. dentipes+G. ensifer, (clade G. varium+, (G. unicolor, (G. carum,(G. calcar, G. apoain)))))], clade S. banhadoae+ collapsespartially [becoming (S. banhadoae, S. molle, cladeS. pseudovarium+ S. thalassinum)], and cladeHeteromitobates is rearranged [becoming (H. harlechin,H. alienus, (H. discolor, H. albiscriptus, (H. anarchus,H. inscriptus)))]. We should stress that in both alternativeanalyses, all genera considered herein are preserved.

Characters – main tendencies and evolution

Here we discuss the main tendencies based on thecladistic analysis; the evolution within each character isdiscussed in Appendix 1. From the cladistic analysis, twomain groups may be recognised, each with three subgroups,which, based on shared characters and morphological‘intervals’, were here recognised as genera in our newclassification scheme, namely (Heteromitobates+ (Acutisoma+Mitogoniella)) + (Serracutisoma+ (Pyatan+Goniosoma)).

Considering the male genitalia, the more pronouncedchanges occurred within Goniosoma. There was a tendencyfor ‘simplification’ and reduction of armature, easily observedby the absence and/or reduction of some spines, which arecommon among other Goniosomatinae and also species ofother subfamilies (e.g. characters 5, 6, 7, 10, 11 and 12), thenarrowing of the ventral plate (character 2), and the reductionand further loss of the ventral process (character 13). In this clade,other striking apomorphies appear, such as the convex apicalmargin and the ‘beak’ in the stylus (characters 1 and 14).

In the other groups, the same genitalic patterns observed inother subfamilies of Gonyleptidae are conserved – for instance,the apical margin of the ventral plate is concave, and length and


width are similar. However, there is a tendency for a generalincrease in the number of spines (e.g. characters 7, 10 and 11),mainly in Heteromitobates, which may bear up to 13 spines ineach side of the ventral blade. In the same genus, otherstriking modifications appear: there is a well developeddorsal process and a different ventral process (characters 13and 16). For the other genera, diagnostic apomorphies are thedifferentiation of the stylus (character 15) in Mitogoniella

and Acutisoma, the hexagonal ventral blade in Serracutisoma,and the truncus dorsally invading the ventral blade in Pyatan.

In legs, especially concerning leg IV, there is a tendency toa reduction in size in Goniosoma, aided by a bending offemur IV (characters 18, 96 and 97). At the same time,granulation and armature highly increased, but spinesconcentrate on tibiae and metatarsi (characters 21, 22 and25–33), in addition to an increase in the inner apophysis of

Table 1. Matrix of characters used in the cladistic analysis of Goniosomatinae


coxa IV (which leads to an increased armature of this region)and in the dorsal–retrolateral apical apophysis of femur IV(characters 40 and 46). In turn, in Serracutisoma, spinesconcentrate on femur IV, mainly on the inner face, with a veryconspicuous armature. Other striking apomorphy of this genusis the ‘bent’ femur (characters 20 and 23). In Mitogoniella,length of legs is highly increased, followed by a reduction ingranulation, and absence of armature, which is a tendencyshared with its sister-genus, Acutisoma (characters 21, 22, 24to 34, 96, and 97). In Pyatan, leg IV is very different from the

others, being very strong, with large acuminate tubercles,and strongly armed. Concerning the prolateral apophysisof coxa IV, there is a tendency in the subfamily for itsreduction, and it is kept large only in Serracutisoma. Anotherapomorphy that deserves note is the prolateral apical apophysisof the trochanter in Serracutisoma, Pyatan, and Goniosoma(characters 38 to 45). Moreover, as extremes, we haveMitogoniella, with long legs and small bodies, againstGoniosoma, with short legs and large bodies (characters 96and 97).

Table 1. (Continued )


Table 1. (Continued )


An important character to discuss is the presence of pectinatetarsal claws of legs III and IV (character 50) in all species ofHeteromitobates, previously known only in Caelopyginae whenGonyleptidae is considered (being a diagnostic feature of thissubfamily), and first detected in Heteromitobates discolor byA.B. Kury (1994) (see also discussion under ‘Taxonomy’).

Concerning the general ornamentation of the dorsal scute andthe free tergites, we may observe, from one side, an increase ofgranules followed by an armature of the free tergites and of theposterior margin of the dorsal scute in part of Serracutisoma(especially in S. proximum+) and, from the other side, reduction insize (with almost smooth species) and increase in density ofgranules in Goniosoma, although, at the same time, this genusbears an eye mound armed with very large spines (characters 52to 56, 75 to 83, and 93).

Finally, we should mention the previously recogniseddiagnostic character of Goniosomatinae, which is the stronglyarmed and robust pedipalps.Actually, an increase in this armatureoccurs independently several times within the subfamily(characters 71 to 74).

Taxonomy

Descriptions, diagnoses, identification keys and discussionconcerning the new taxonomic scheme for genera and speciesof Goniosomatinae are shown below. The classification in generapresented as well as the order for the description of generaand species follows the results of the phylogenetic analysispresented above. In order to facilitate comparisons, genericdiagnostic characters and important structures in specific andgeneric descriptions are presented in bold. When the variation inquantitative characters are shown, the type specimen values aregiven in parentheses. In the synonymic list following eachtaxon, the following conventions were used: ‘comb.’ indicatesthat that combination was proposed as new by that author;when followed by ‘type species’, it indicates that that specieswith new combination was used as the type species of that givengenus by that author; ‘comb. reestabl.’ indicates that that authorreestablished a previous combination for that species nameproposed by other author (in this case, this previouslyproposed combination is declared elsewhere in the samesynonymic list); ‘syn.’ indicates to which senior synonym thatspecies/genus name was proposed as a junior synonym by thatauthor. In the lists of ‘specimens examined’, ‘ma’= adult male,‘fe’= adult female, and ‘im’= immature.

GONIOSOMATINAE Mello-Leitão

GoniosominaeMello-Leitão, 1935a: 402. – 1935b: 32, 1936: 32, 1940: 23,1949: 13; Piza, 1938: 139; B. Soares, 1943: 205, 1944a: 311, 1945a:192, 1945b: 231, 1945c: 350, 1946: 495; Soares & Soares, 1945: 252,1946: 233, 1947a: 63, 1947b: 209, 1947c: 249, 1948: 621; H. Soares,1945: 208, 1970: 212, 1974; Soares & Bauab, 1970: 131; Muñoz-Cuevas, 1972: 28; Trajano, 1987: 538–544, 547, 548; Trajano &Gnaspini, 1991a: 383, 388, 393, 399, 402, 1991b: 75, 76; Pinto-da-Rocha, 1993: 235.

Goniosomatinae Stefanini-Jim, 1985: 29 [unpublished thesis].–Pinto-da-Rocha, 1995: 80; Gnaspini, 1996: 418; Pinto-da-Rocha, 1999: 358,379, 382; Sabino &Gnaspini, 1999: 675, 677; Machado & Raimundo,2001: 138, 143;Pinto-da-Rocha et al., 2001: 143; Silva-da-Rocha et al.,2001: 99; Pinto-da-Rocha, 2002: 381, 382; Hara & Gnaspini, 2003:

258, 259, 262, 264, 268; Hara et al., 2003: 441, 443; Willemart, 2002:50, 51; Kury, 2003: 115; Willemart & Gnaspini, 2004a: 16, 21–24,2004b: 222, 229, 231–233; Gnaspini et al., 2004: 32, 34.

Diagnosis

Anterior margin of carapace smooth or with sparse granules;eye-mound wide, with 1 pair of tubercles or spines or a singlespine. Dorsal scute with 3 areas; area II invading area Imedially, dividing it into 2 parts; area I with 1 centraltubercle on both sides; area III with 1 pair of tubercles orspines. All areas and carapace with homogeneously distributedgranules (those of areas generally a little larger than those ofcarapace, while those of margins have different sizes); lateralmargins with 2 longitudinal rows of granules, one close tolateral groove and other to the edge; posterior margin with 1transversal row of granules like those of free tergites. Angles ofposterior margin of dorsal scute and free tergites with orwithout spines or tubercles. Coxae I–III with 1 longitudinal rowof tubercles, decreasing from I to III, and lateral granules,increasing from I to III; coxa IV with larger granules(granules on lateral face larger than those on ventral face);trochanter with 3 tubercles (1 medial and 2 apical) decreasingfrom I to III, and sparse granules, increasing from I to III. Robustpedipalps longer than dorsal scute; femur with a row of largeventral tubercles and 1 or 2 retrolateral setae; tibia with ventralarmature IiIi prolaterally (and sometimes with 1 distinct basaltubercle or spine in addition to standard armature) and IiIIiretrolaterally; tarsus with ventral armature IIi prolaterally andIiIi retrolaterally. Legs I–IV with granulation and armatureorganised in 6 regular longitudinal rows of granules and/orspines; apical end of dorsal rows with 2 pointed apophysesbackward, being retrolateral larger than prolateral; tarsus I–IVof adult with more than 8 segments; tarsus III–IV with 2 smoothor pectinate claws and with tarsal process varying fromvery short to as long as a claw; distitarsus I–II with 3segments. Penis with or without ventral and dorsal processon glans; ventral plate with 2 groups of setae of same size, beingone basal and other apical or subapical (the latter with 1 smallmedial seta).

General sexual dimorphism in subfamily

Males with granulation of legs stronger than females, with legsIII-IV unarmed (or weakly armed in some species) on females,even if armed on males of the same species. Male femur IV maybe straight, curved or sharply bent, with variable length; femaleswith femur straight or almost straight, generally shorter thanthat of males. Apophyses of coxa and trochanter of leg IV largeron males; females have only the prolateral apical apophysis ofcoxa IV pointed and short, like a robust spine. Female tarsalprocess IV at least half the size of male process. Angles ofposterior margin of dorsal scute and free tergites generallylarger on females, armed with robust spine on each angle;male with angles unarmed or with weak spines.

Male dimorphism

Most species show a large variation in size of males, withdifferent morphs. Different male morphs were observed inmany taxa of the Grassatores infra-order of Laniatores


(Gnaspini et al. 2004), andwerehypothesised as theoccurrenceoftwo adult instars (Gnaspini et al. 2004) but are the result ofdifferent reproductive strategies (Tsurusaki and Fujikawa 2004)caused by territorial disputes between males (Machado andMacías-Ordóñez 2007). Larger males differ from smaller ones

mainly by having more developed and/or more pronouncedsecondary sexual characters. In Goniosomatinae, larger maleshave apophyses and spines of legs IV larger and with a moredefined shape, e.g. more curved, with stronger branches, or morepointed, and legs IV more curved, bent, or longer.

38

44

43

42

39

4047

46

41

45

Figs 38–47. 38–40, Goniosoma unicolor. 38, Habitus, dorsal view. 39, Right femur IV (dorsal view). 40, Rightpedipalp (ventral–prolateral view). 41, 42, Goniosoma calcar. 41, Habitus, dorsal view. 42, Left femur IV (dorsalview) – notice very accentuated curvature. 43–45, Goniosoma carum. 43, Habitus, dorsal view – notice the curvedretrolateral apical apophysis of coxa (raac). 44, Ventral view of coxa IV, showing the bifid ventral apophysis (vac). 45, Rightfemur IV (dorsal view) – notice the bent retrolateral apical apophysis (raaf). 46, 47, Pyatan insperatum, gen. nov., sp. nov.46, Habitus, dorsal view – notice the small ratio between width and length of the dorsal scute. 47, Right femur IV (dorsalview), showing acuminate granules. Scale = 4mm, except for 40, 44 = 2mm.


Distribution

Brazilian Atlantic Forest, from southern Bahia to Santa Catarina.

New proposed classification

GoniosomatinaeGenus Goniosoma Perty, 1833G. varium Perty, 1833 (type)G. capixaba, sp. nov.G. venustum Koch, 1839G. roridum Perty, 1833

G. vatrax Koch, 1848G. macracanthum (Mello-Leitão, 1922), comb. nov.G. dentipes Koch, 1839G. ensifer (Mello-Leitão, 1940)G. apoain, sp. nov.G. unicolor (Mello-Leitão, 1932), comb. nov.G. calcar (Roewer, 1913)G. carum (Mello-Leitão, 1936), comb. nov.G. lepidum Gervais, 1844 species inquirendaG. monacanthum Gervais, 1844 species inquirendaG. obscurum Perty, 1833 species inquirendaG. versicolor Perty, 1833 species inquirenda

48

51

49

50

56

55

52

53

54

Figs 48–56. 48–50, Serracutisoma proximum. 48, Habitus, dorsal view. 49, Right femur IV (dorsal view). 50, Right femurIII (retrolateral view). 51, 52, Serracutisoma banhadoae. 51, Habitus, dorsal view. 52, Right femur IV (dorsal view). 53–56,Serracutisoma molle. 53, Habitus, dorsal view. 54, Right coxa and trochanter IV (retrolateral view); paac = prolateral apicalapophysis of coxa, daat = dorsal apical apophysis of trochanter, raat = retrolateral apical apophysis of trochanter. 55, Rightfemur IV (dorsal view); sp = spine. 56, Apex of right femur III (retrolateral view) – notice armed row3. Scale = 4mm.


Genus Pyatan, gen. nov.P. insperatum DaSilva, Stefanini-Jim & Gnaspini, sp. nov. (type)

Genus Serracutisoma Roewer, 1930 reestabl.S. proximum (Mello-Leitão, 1922), comb. nov. (type)S. banhadoae (Soares & Soares, 1947), comb. nov.S. molle (Mello-Leitão, 1933), comb. nov.S. pseudovarium, sp. nov.S. thalassinum (Simon, 1879), comb. nov.S. fritzmuelleri, sp. nov.S. catarina (Machado, Pinto-da-Rocha & Ramires, 2002), comb. nov.S. inerme (Mello-Leitão, 1927), comb. nov.S. guaricana, sp. nov.S. spelaeum (Mello-Leitão, 1933), comb. nov.

Genus Heteromitobates Roewer, 1913H. discolor (Soerensen, 1884) (type)H. inscriptus (Mello-Leitão, 1922), comb. nov.H. albiscriptus (Mello-Leitão, 1932), comb. nov.H. anarchus, sp. nov.H. harlequin, sp. nov.H. alienus, sp. nov.

Genus Mitogoniella Mello-Leitão, 1936 reestabl.M. indistincta Mello-Leitão, 1936, comb. reestabl. (type)M. taquara, sp. nov.M. unicornis, sp. nov.M. modesta (Perty, 1833), comb. nov.M. badia (Koch, 1839), comb. nov. species inquirenda

Genus Acutisoma Roewer, 1913A. acutangulum (Simon, 1879) (type)A. coriaceum, sp. nov.A. hamatum (Roewer, 1928)A. longipes Roewer, 1913, comb. reestabl.

Key for the genera of Goniosomatinae based on males

1. Femur IV retrolaterally armed; prolateral apophysis of coxa IV falcate(Fig. 54, paac); apical apophysis of trochanter dorsally placed andcurved inward (Fig. 54, raat) ......................... Serracutisoma (p. 563)

Femur IV unarmed or, if armed, spines are larger on ventral rows (row3and row4) (Fig. 89); prolateral apophysis of coxa conic; apicalapophysis of trochanter prolateral or absent .....................................2

2. Tarsal claws III–IV pectinate ...........................Heteromitobates (p. 582)Tarsal claws III–IV simple .....................................................................3

3. Prolateral apophysis of coxa IV pointed, with 1 small subapical process,or blunt (Figs 82, 94, 107); eye-mound with 1 pair of tubercles close toeach other and with the space between them at the same level orhigher than the rest of the eye-mound closer to the eyes (Fig. 109), orwith a single spine (Fig. 100); without prolateral apical apophysison trochanter IV .................................................................................4

Prolateral apophysis of coxa IV pointed, without subapical process butwith a basal process (Fig. 17); small eye-mound with 1 pair of spines(Figs 13, 31) or with very small tubercles far from each other, like smallgranules (Fig. 27); trochanter IV with 2 prolateral apophyses (Fig. 46)(if subapical apophysis is absent, then tibia IV is armedand retrolateral apophysis of coxa IV is large and pointed, as inFigs 32, 34)........................................................................................5

4. Legs very long (more than 10� the body size) and approximately smoothor with minute granules (Fig. 96); trochanter IV without retrolateralbasal apophysis and coxa IV with reduced retrolateral apophysis;prolateral apophysis of coxa IV with blunt apex (Fig. 99)............................................................................................Mitogoniella (p. 588)

Legs relatively short andwith conspicuous granulation; trochanter IVwithretrolateral basal apophysis and coxa with crescentlike or cylindricalmedian-sized retrolateral apophysis (Fig. 107); prolateral apophysis ofcoxa IV with pointed apex (Fig. 104); dorsal–retrolateral apical

apophysis of femur IV straight and inward (Figs 110, 111, raaf) ..................................................................................Acutisoma (p. 592)

5. Femur IV curved outward, robust,with pointed granules and strong ventralarmature (Fig. 47); dorsal apical apophyses of femur IV with samesize .............................................................Pyatan, gen. nov. (p. 560)

Femur IV curved outward and inward (S-shaped) (Fig. 23) or straight(Fig. 20), short and with round granules; femur IV unarmed or withweak tomediumarmature on row3 (Fig. 18); dorsal apical apophyses offemur IV of very different size, retrolateral very large and prolateralminute (Fig. 23) ..................................................Goniosoma (p. 545)

Key for the genera of Goniosomatinae based on females

1. Eye-mound with 1 pair of spines (Fig. 13).............................................2Eye-mound with 1 pair of tubercles or a single spine (Fig. 59) .............3

2. Dry-mark largely covering the dorsal scute: round and filled up betweenspines of area III and on the carapace; and as a frame on rest ofabdominal scute (Fig. 202) ........................Pyatan, gen. nov. (p. 560)

Dorsal scute with other combination of dry-marks ..............................................................................................................Goniosoma (p. 545)

3. Tarsal claws III and IV pectinate (Figs 6, 7) ................................................................................................................Heteromitobates (p. 582)

Tarsal claws III and IV simple (Figs 8, 9)..............................................44. Eye-mound with 1 pair of tubercles close to each other and the space

between them at the same level or higher than the rest of the eye-moundcloser to the eyes (Fig. 109), or with a single spine (Fig. 100) .........5

Eye-mound with 1 pair of tubercles distant from each other and with thespace between them low-leveled, lower than upper margin of eyes(Fig. 59) ......................................................... Serracutisoma (p. 563)

5. Ventral granules on posterior margin of stigmatic area and free sterniteslarger and in larger number than those dorsally on posterior margin ofdorsal scute and free tergites.................................Acutisoma (p. 592)

Ventral granules smaller or of the same size than dorsal ones...........................................................................................Mitogoniella (p. 588)

Genus Goniosoma Perty

(Figs 1, 2, 8, 9, 11–45, 112–140, 190–200, 234, 235)

Goniosoma Perty, 1833: 202, 207, 208. – Koch, 1839: 52, 58, 62, 64,119, 122, 124, 1848: 21; Gervais, 1844: 106–110; Simon, 1879:227–233, 277; Soerensen, 1884: 613; Roewer, 1913: 170, 257–264,1923: 465, 497–499, 1930: 349, 382; Mello-Leitão, 1923: 154, 155,191, 192, 1926: 32, 1931: 124, 125, 1932: 233, 266–268, 479, 1935b:110, 111, 1936: 34; B. Soares, 1944b: 280, 1944c: 178, 1945c: 351,352; Soares & Soares, 1948: 628–632; Soares & Bauab, 1970: 131,133, 134; Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; H. Soares,1974: 483; Trajano, 1987: 538, 541; Trajano & Gnaspini, 1991a: 383,388, 393, 402, 1991b: 75; Gnaspini & Trajano, 1992: 42, 56, 57;Pinto-da-Rocha, 1993: 229, 235, 251, 1995: 80, 81, 1996: 22, 1999:382; Gnaspini, 1995, 1996, 1999; Pellegatti-Franco & Gnaspini,1996: 355, 360; Gnaspini & Cavalheiro, 1998; Machado &Oliveira, 1998, 2002: 1510, 1519, 1521; Gnaspini & Hoenen,1999: 55, 56; Sabino & Gnaspini, 1999; Machado et al., 2000,2001, 2002; Machado & Raimundo, 2001: 138–142; Pinto-da-Rocha et al., 2001: 137, 143, 152, 153; Sessegolo et al., 2001:184; Silva-da-Rocha et al., 2001: 99, 101; Willemart, 2001: 249,251, 2002: 51, 55; Machado, 2002; Santos & Gnaspini, 2002;Gnaspini et al., 2003, 2004: 31–35; Hara & Gnaspini, 2003: 258,259, 262–264, 266, 269, 270, 273; Hara et al., 2003: 441, 442; Kury,2003: 117; Willemart & Gnaspini, 2004a, 2004b.

ProgoniosomaRoewer, 1913: 170, 264–266, 268, 269, 271–275. – 1923:465, 499–504, 627, 1930: 349, 382–387; Mello-Leitão, 1922: 340,1923: 155–158, 192, 1927: 401, 1932: 233, 258–265, 1940: 23;


Bristowe, 1925: 502; B. Soares, 1945a: 192, 1945c: 352; Soares &Soares, 1948: 623, 628–632, 634 (syn. Goniosoma); Moritz, 1971:200, 204. (Type =Goniosoma dentipes Koch, 1839, by originaldesignation.)

LyogoniosomaMello-Leitão, 1926: 33, 55. – 1932: 232, 244, 1935b: 110;Roewer, 1930: 349, 444; B. Soares, 1946: 497; Soares & Soares, 1948:633, 634. (Type =Progoniosoma macracanthum Mello-Leitão, 1922,by original designation.) Syn. nov.

Glyptogoniosoma Mello-Leitão, 1932: 271, 462, 479. – 1936: 36;B. Soares, 1944a: 260 (syn. Acutisoma). (Type =Progoniosomacruciferum Mello-Leitão, 1923, by original designation.) Syn. nov.

GoniosomellaMello-Leitão, 1936: 33. –B. Soares, 1945c: 352; Soares &Soares, 1948: 633 (syn.Lyogoniosoma). (Type =Goniosomella perlataMello-Leitão, 1936, by original designation.) Syn. nov.

Xulapona Mello-Leitão, 1936: 32. – B. Soares, 1945c: 352; Soares &Soares, 1948: 633 (syn. Lyogoniosoma). (Type =Xulapona caraMello-Leitão, 1936, by original designation.) Syn. nov.

Metalyogoniosoma Soares & Soares, 1946: 233. – 1948: 634.(Type =Metalyogoniosoma unum Soares & Soares, 1946, byoriginal designation.) Syn. nov.

Type species:Goniosoma varium Perty, 1833, by subsequent designationof Roewer, 1913.

57

58

64

65

6662

59

6163

60

Figs 57–66. 57–60, Serracutisoma pseudovarium, sp. nov. 57, Habitus, dorsal view. 58, Dorsum, right view;ap = apophysis, gr = granule. 59, Eye mound (posterior view), with a pair of high tubercles (tb). 60, Right femurIV (dorsal view). 61, 62, Serracutisoma thalassinum. 61, Habitus, dorsal view. 62, Right femur IV (dorsal view).63–66, Serracutisoma fritzmuelleri, sp. nov. 63, Habitus, dorsal view. 64, Right trochanter and femur IV(dorsal view). 65, Right femur IV (prolateral view). 66, Trochanter, femur, patella and base of tibia of rightpedipalp (ventral–retrolateral view). Scale = 4mm, except for 58, 66 = 2mm, 59 = 1mm.


Diagnosis

Eye-mound low and generally with one pair of medium-sized or very large spines (exception: tubercle like amedium-sized granule in G. macracanthum). Coxa IV withone pointed prolateral apical apophysis with 1 small sub-basal posterior process, oblique or nearly longitudinal andslightly curved downward; retrolateral apical apophysis large,of same size or larger than trochanter, or, at least, like a spine.Femur short, curved inward (from with a strong curvatureto almost straight) and very granulated; flat granules, mainly onretrolateral row (row2); dorsal rows, on apical region, verygranulated and disorganised and/or with a nearly longitudinalcomb of granules (very close to each other); dorsal–prolateralapical apophysis large and curved backward. Tarsalprocess short, medium-sized or long (from minute to largerthan tarsal claws); if large in male, female has medium-sizedtarsal process. Tarsal claws simple. Penis longer than wider;apical group of setae with four setae on a single longitudinal row(exception: one additional ventral seta, but very small, inG. vatrax); basal group with three setae on longitudinal,L-like or oblique row, and one additional very small, ventralseta; all setae pointed. Glans without or with ventral process;with apical lamina or thin (without lamina); stylus with a dorsalapical ‘beak’.

Distribution

Espírito Santo, southern Minas Gerais, Rio de Janeiro and, inSão Paulo, only Serra da Bocaina and north littoral.

Key for the species of Goniosoma based on males

1. Coxa IV with a bifid and very long ventral apophysis on apex, withthe part beyond dorsal scute longer than trochanter IV...................................................................................................... G. carum (p. 559)

Coxa IV without ventral apophysis ......................................................22. Trochanter IV with 2 prolateral apophyses with subapical larger ........3

Trochanter IV with 1 prolateral apophysis, sub-basal, or 2, but with thesubapical reduced...............................................................................8

3. Eye-moundwith1pair ofmedium-sized spines (<1mm, lower thanpair ofspines of area III); dorsal scute with white dry-mark following groovesand on the carapace, mainly...............................................................4

Eye-mound with 1 pair of large spines (>1mm, higher than pair ofspines of area III); dorsal scute without white dry-mark or with thismore restricted to some regions .........................................................7

4. Retrolateral apophysis of coxa IV large, with nearly same length oftrochanter IV .........................................................G. varium (p. 548)

Retrolateral apophysis small, like a spine.............................................55. Femur IV straight and with a comb of dorsal granules in all apical

half; granulation of posterior margin of dorsal scute and free tergitesnormal ................................................................................................6

Femur IV curved inward, with a small comb; posterior margin of dorsalscute and free tergites with granules much larger than those of rest ofdorsal scute.......................................... G. capixaba, sp. nov. (p. 549)

6. Dark body and dorsal scute all punctate with dry-mark around thegranules ................................................................G. roridum (p. 551)

Light body, with dry-mark following the grooves of areas and on thecarapace, never around the granules .................G. venustum (p. 550)

7. Retrolateral apophysis of coxa IV larger than trochanter IV, pointed;retrolateral apical apophysis of trochanter IV small, like a granule;femur IV with medium curvature .......................G. unicolor (p. 557)

Retrolateral apical apophysis of coxa IV reduced; retrolateral apicalapophysis of trochanter IV large, straight and inward and with roundapex; femur IV with very strong curvature.............G. calcar (p. 558)

8. Prolateral apical apophysis of trochanter IV absent; retrolateralapophysis of coxa IV straight ............................................................9

Prolateral apical apophysis of trochanter IV smaller than basal apophysis;retrolateral apophysis of coxa IV S-shaped ............G. vatrax (p. 552)

9. Area III and eye-mound with spines; retrolateral apophysis of coxa IVsmaller than 2� the size of trochanter IV........................................10

Area III and eye-mound with 1 pair of very small tubercles;retrolateral apophysis of coxa IV very large, 2� to 3� the size oftrochanter IV ............................................G. macracanthum (p. 553)

10. Posterior margin of dorsal scute with medium-sized granules, largerthan granules on rest of dorsal scute; dry-mark shaping as a cross ondorsal scute; tibia IV with a retrolateral apical spine straight, largerthan others........................................................................................11

Dorsal scute only with minute granules; thin dry-mark followingthe grooves of areas; tibia IV without retrolateral spines ....................................................................................G. apoain, sp. nov. (p. 556)

11. Femur IV with strong inward curvature; retrolateral apical apophysisof coxa IV longer than 1.5� the size of trochanter ............................................................................................................G. ensifer (p. 555)

Femur IV with weak inward curvature; retrolateral apophysis of coxaIV with about same size of trochanter IV............G. dentipes (p. 554)

Key for the species of Goniosoma based on females

Goniosoma ensifer, G. macracanthum and G. apoain, sp. nov.are not included in the key because no females are known fromthe collections examined so far.

1. Eye-moundwith 1 pair of medium-sized spines (<1mm, lower than pair ofspines of area III) (Fig. 13) ................................................................2

Eye-mound with 1 pair of large spines (>1mm, higher than pair of spinesof area III) (Fig. 31) ...........................................................................6

2. Dorsal scute completely punctate with dry-mark around the granules(Fig. 195) .............................................................G. roridum (p. 551)

Dorsal scute without punctate dry-mark.................................................33. Posterior margin of dorsal scute and free tergites with few and very large

granules (Fig. 5); posterior angles of these segments stronglyarmed...................................................G. capixaba, sp. nov. (p. 549)

Posterior margin of dorsal scute and free tergites with medium-sizedor smaller granules; posterior angles of these segments unarmed.....4

4. Dry-mark on dorsal scute as a frame; posterior margin of dorsal scuteand free tergites with minute granules ...............................................5

Dry-mark on dorsal scute restricted to other regions; posterior margin ofdorsal scute and free tergites with medium-sized granules and withangles with tubercles...............................................G. vatrax (p. 552)

5. Dry-markon thecarapacebehindeye-moundcup-shaped (Fig.17); generalcolour yellowish brown (Fig. 192) ...................G. venustum (p. 550)

Dry-mark on the carapace behind eye-mound as a frame; general colourdark-brown (Fig. 190)...........................................G. varium (p. 548)

6. Dorsal scute without dry-mark.................................G. unicolor (p. 557)Dorsal scute with dry-mark.....................................................................7

7. Dry-mark cross-shaped, covering the entire area II and from the carapace toarea III; granules of posterior margin of dorsal scute and free tergitesmedium-sized or large..........................................G. dentipes (p. 554)

Dry-mark restricted to other regions; posterior margin of dorsal scuteand free tergites with minute granules or practically smooth ............8

8. General colour black; spines of area III short (about 1/2� the size ofspines of eye-mound)..............................................G. calcar (p. 558)

Colour brown; spines of area III higher than 1/2� the size of spines ofeye-mound..........................................................................................9


9. Dry-mark round between spines of area III; epidermic pigmentation asa frame ................................................................... G. carum (p. 559)

Without round dry-mark between spines of area III; epidermicpigmentation as a frame and reduced to lighter large circles aroundeach granule ...........................................G. apoain, sp. nov. (p. 556)

Goniosoma varium Perty

(Figs 2, 8, 9, 11–13, 112–114, 190)

Goniosoma varium Perty, 1833: 208. – Koch, 1839: 52; Gervais, 1844:107; Simon, 1879: 228; Soerensen, 1884: 613; Roewer, 1913: 258,1923: 497; Mello-Leitão, 1923: 154, 192, 1932: 266; Soares & Soares,1948: 631; Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; Kury, 2003:118.

Goniosoma patruele Perty, 1833: 202. –Koch, 1839: 122; Gervais, 1844:108; Simon, 1879: 233; Soares & Soares, 1948: 630 (comb. reestabl.);Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Syn. nov.

Goniosoma junceum Perty, 1833: 202. – Gervais, 1844: 108 (syn.Goniosoma patruele Perty, 1833). Syn. nov.

Progoniosoma patruele. – Roewer, 1913: 265, 266 (comb.), 1923: 500,501, 1930: 383; Mello-Leitão, 1923: 157, 192, 1932: 259, 263.

Goniosoma xanthophthalmum Mello-Leitão, 1931: 125. – 1935b: 111,1936: 34; B. Soares, 1944c: 178, 1945c: 352; Soares & Soares, 1948:632; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118. Syn. nov.

Goniosoma xatnhophthalmum (typographical error) Mello-Leitão,1931: 125.

Acutisomella intermedia Mello-Leitão, 1937: 294. Syn. nov.Acutisomella cryptoleuca Mello-Leitão, 1940: 26. – B. Soares, 1944c:178 (syn. Goniosoma xanthophthalmum Mello-Leitão, 1931).Syn. nov.

Acutisoma intermedium. – B. Soares, 1945b: 231 (comb.); Soares &Soares, 1948: 625; Kury, 2003: 116.

Metalyogoniosoma unum Soares & Soares, 1946: 234. – 1948: 634;Kury, 2003: 119. Syn. nov.

Goniosoma minense (misidentification). – H. Soares, 1974: 483.

Material examined

Types. ‘Brasilien’, Amazon river (mistaken), ma lectotype and 2 feparalectotypes (here designated) (ZMHB927), Olfers leg. Espírito Santo:Santa Leopoldina (Chaves), im type of Metalyogoniosoma unum(MZSP1801), Vervloet leg., X/1945. Rio de Janeiro: EngenheiroPaulo de Frontin (Morro Azul), fe holotype of Acutisomellaintermedia (IBSP47); Rio de Janeiro (Jacarepaguá), 2 fe and 1 imsyntypes of Goniosoma xanthophthalmum (MNRJ11382), Berla leg.;Rio de Janeiro (Jacarepaguá), ma allotype of Goniosomaxanthophthalmum (MNRJ42224), Berla leg., V/1915; Rio de Janeiro(Jacarepaguá), fe holotype of Acutisomella cryptoleuca (MNRJ58400),R. Arlé.

Other material examined. Espírito Santo: Linhares, 2 ma and 3 fe(HS582), J. Jim leg., XII/1971; Santa Tereza, 1 ma (HS838), O.L. Peixotoet al. leg.,VIII/1981.Riode Janeiro:CasimirodeAbreu (Barra deSão João,Morro de São João), 4 ma and 3 fe (MNRJ17390), Expedicão Arachné leg.,21–24/III/2003; Casimiro de Abreu (São João, Fazenda Reunidas), 4 maand4 fe (MNRJ4302), Exped.UNI-RIO leg., I/1994;Guapimirim (Est. Ecol.Est. Paraíso), 5 ma and 3 fe (MZSP15498), R. Pinto-da-Rocha &R.S. Bérnilsleg., VII/1996; Magé (Inhomirim, Raiz da Serra), 5 ma, 3 fe and 6 im(MZSP15244), R.S. Bérnils leg., X/1996; Nova Iguacu (Tinguá), 3 maand 5 fe (HS302), Peracchi & Izecksohn leg., IX/1965; Rio de Janeiro(Tijuca), 1 ma and 1 fe (HS280), G.B. Milton Valle leg., II/1944;Mangaratiba, 1 ma and 1 im (MNRJ 17391), E. Vasconcelos leg., 10/IV/2003.

Description

Dorsum. Eye-mound with 1 pair of medium-sized, straight,parallel and separate spines. Area I with 1 pair of tubercles ofsize of a medium-sized granule. Area III with 1 pair of largespines slightly backward; these spines are higher than those ofeye-mound. Angles of free tergites and posterior marginunarmed.

Granulation. Carapace: 2–6 small granules. Areas I–III:18–26 small granules. Lateral and posterior margins:medium density of minute granules. Free tergites: low densityofminute granules.Anal operculum practically smooth. Venter:Posterior margin of stigmatic area and free sternitespractically smooth. Coxa I and Coxa IV: medium density ofminute granules.

Chelicerae. Segment I with few granules. Segment II withmany bristles on apex.

Pedipalps. Trochanter with 2–6 dorsal and 3–5 ventralelevations. Femur with 5–7 ventral elevations (standardarmature IiiIiI), 2 retrolateral subapical spines and dorsalgranulation with low density of minute granules. Patellawith 1 weak ventral–retrolateral subapical tubercle. Tibiawith 1 weak ventral–prolateral subapical tubercle, of samesize of that of patella, in addition to the standard armature ofsegment.

Leg I. Trochanter with low density of minute granules.Femur with minute granules. Other segments practicallysmooth.

Leg II. Trochanter with low density of minute granules.Femur with small granules. Other segments practically smooth.

Leg III. Trochanter with low density of minute granules.Femur and patella with small granules. Tibia with minutegranules. Metatarsus practically smooth.

Leg IV. Coxa with medium-sized granules on lateralmargin. Prolateral apical apophysis oblique; retrolateral apicalapophysis pointed, nearly as long as trochanter. Contact zonebetween coxa and the lateral–posterior margin of dorsal scuteextended. Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 apical (latter larger); ventral granulation: low densityof medium-sized granules. Femur with medium inwardcurvature; row1 and row2 with large, flat and longitudinallyelongate granules (mainly on medial third); dorsal rowsdisorganised on apical third with high density of highergranules, and the presence of a longitudinal comb of granuleson apex; higher granules on apex of row3; dorsal apicalapophyses: retrolateral large, curved backward and prolateralminute. Tibia armed with large spines (curved backward) onapical half of row3; row2 with a comb of high granules closeto each other. Metatarsus with few and large basal spinescurved backward. Tarsal process medium-sized.

Tarsal segmentation. 9–11 (10), 20–24 (22), 9–11 (10–11),10–12 (11).

Penis. Ventral plate with lateral margin almost straight(with a light concavity, caused by the wider base) and concaveapical margin; apex apically extended above apical group ofsetae. Apical group with an additional apical seta distant fromthe others, situated on apex of plate and dorsally curved;basal group with an oblique row of setae very close to eachother.Glanswith short ventral processwith a round apical lamina


inserted on a wide stalk. Truncus invading the ventral platemedially on dorsal face.

Colour pattern

Dorsal scute and venter light-brown; carapace, legs IV andspines of area III dark-brown; pedipalps, chelicerae and legs I–III(except trochanter) dark green; light rings adjacent to tibia andmetatarsus of legs. Dry-mark as a frame, on dorsal scute andcarapace, like a wide stripe before the eye-mound; like a roundspot between spines of area III; on apex of coxa, at base oftrochanter and femur.

Female

Angles of free tergites and posterior margin of dorsal scuteas among males.

Measurements (in mm)

Dorsal scute: width: 9.07–10.45 (9.98), length: 8.20–8.94(8.80); leg I: 30.22–33.54 (31.15), II: 56.10–67.55 (60.85), III:44.46–48.41 (46.76), IV: 57.52–64.17 (59.51), femur IV:14.85–16.92 (16.04).

Remarks

Six junior synonyms were recognised for the present species,four of them in the present study. The two main reasons to thelarge number of synonyms are the facts that (1) three of themwere described from females (which are distinct from malesbecause of the dimorphic characters), and (2) the type-localities are in Rio de Janeiro state, where there were manycollections in the past, with many specimens that were describedas new species. However, the unique dry-mark and the medium-sized spine on the eye mound were enough to recognise thespecies and to propose such synonyms, even without part of thetype material. Metalyogoniosoma unum was described from asingle immature specimen, probably a last stage nymph, in whichthe armature of the eye-mound is not completely developed.Soares and Soares (1946) used this character to describe thespecies and its monotypic genus. We examined the dissectedpenis found in the same vial of this specimen and we concludedthat it probably belongs to a different species and that it wasmistakenly included in the same vial, since it is very distinctivefrom the penis of other species of Goniosoma and it is alreadyfullydeveloped tobelong to an immature specimen.Wediscussedelsewhere (see Gnaspini et al. 2004) that the penis is fullydeveloped in adults but it is much less developed in theimmediately previous nymphal stage.

Goniosoma capixaba, sp. nov.

(Figs 14–16, 115–117, 191)

Material examined

Types. Espírito Santo: Apiacá (Fazenda Rio Doce), ma holotypeand allotype (MNRJ6499), R. Baptista & A. Baptista leg., 24–25/VII/1989; 7 ma and 3 fe paratypes (MNRJ6499) (idem); Domingos Martins,1 ma paratype (HS850), E. Izecksohn et al. leg., 9/VIII/1983; 1 ma (HS849)(idem).

Description

Dorsum. Eye-mound with 1 pair of short and separate spines,slightly curved forward. Area I with 1 central pair of smalltubercles. Area III with 1 pair of large spines slightlybackward, of same size of spines of eye-mound. Angles ofposterior margin of dorsal scute and free tergites with 1 or2 high tubercles.

Granulation. Carapace: 2–5 (2) medium-sized granules.Areas I–III: 7–16 (13) large granules. Lateral margin: lowdensity of medium-sized granules; lateral margin of dorsal scutewith small granules. Posterior margin: medium density of verylarge granules. Free tergites: low density of very large granules.Anal operculum: low density of small granules. Venter:Posterior margin of stigmatic area and free sternites: highdensity of medium-sized granules.Coxa I: low density of minutegranules. Coxa IV: low density of small granules; lateral: largegranules.

Chelicerae. Segment I with 1 dorsal–retrolateral apicaltubercle and 2 basal granules. Segment II with sparse granuleson apical half.

Pedipalps. Trochanter with 1–3 (3) dorsal and 2–4 (3)ventral elevations. Femur with standard armature IiiIiIi,7–9 (7) ventral elevations and 2 retrolateral subapical setae.Patella with 1 strong ventral–retrolateral subapical tubercle.Tibia with 1 ventral–prolateral basal spine (half the size ofsetae of segment), in addition to the standard armature.

Leg I. Trochanterwithmediumdensity ofminute granules.Femur with small granules. Other segments practically smooth.

Leg II. Trochanter with high density of small granules.Femur and patella with small granules. Other segmentspractically smooth.

Leg III. Trochanter with high density of small granules.Femur and patella with medium-sized granules. Tibia andmetatarsus with small granules.

Leg IV. Coxa with large and oblique prolateral apicalapophysis; retrolateral apical apophysis pointed, straight,of same size of spine of area III of dorsal scute (smaller thanhalf the size of trochanter). Trochanter with 2 prolateralapophyses, being 1 sub-basal and 1 subapical, latter 2� thesize of former, both slightly curved upward; retrolateral apicalapophysis curved backward and small, with a tubercle nearby;granulation: medium density of small granules. Femur withmedium inward curvature and with medium-sized and roundgranules; row1 and row6 disorganised on apical half withlarge number of high granules and presence of a smalllongitudinal comb on apical region; row3 with large spineson apical region; row4 with small spines on apical half; dorsalretrolateral apical apophysis very large, curved backward, andprolateral apophysis very small. Patella with very largegranules. Tibia with row2 with high granules close to eachother in all extension; row3 with small spines from base toapical fourth, and very large spines on apex; spine stronglycurved backward on apex close to row3. Metatarsus withrow3 and row4 with large granules; the rest with very smallgranules. Tarsal process very long and robust, longer thantarsal claws.

Tarsal segmentation. 9–10 (9), 12–20 (18), 9–12 (10),11–14 (11/12).


Penis. Ventral plate rectangular with an extension abovethe apical group of setae; apical margin straight and wide; regionwhere setae of apical group are inserted is concave.Apical groupwith 4 setae equidistant to each other; basal group with setaeclose to each other, forming an oblique row. Glans with a thinventral process smaller than half the size of stylus. Truncus notinvading ventral plate.

Colour pattern

General dark-brown; carapace black and dorsal scute andventer light-brown; leg IV very dark; joint of tibia andmetatarsus of legs with a lighter ring; granules and tuberclesof posterior margin of dorsal scute and of free tergites yellow.Dry-mark as a frame on dorsal scute and a wide spot dividedat middle between the spines of area III; on carapace like around spot behind eye-mound as wide as in between the eyes(absent in a central triangle and in lateral fragment) and, beforeeye-mound, like a wide spot; on apex of coxa, base of trochanterand base of femur of leg IV.



Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angle of posterior margin of dorsal scutewith a very large granule like others of margin. Angles of freetergites with 1 robust spine each. Anal operculum with lowdensity ofmedium-sized granules.Tarsal process half the size oftarsal claws.

Remarks

This new species is similar to G. venustum and G. roridum. Itcan be distinguished mainly by a curved femur IV and absenceof a comb of granules on dorsum of femur IV (the very largegranules are not aligned), which are plesiomorphic states, andby very large granules on dorsal scute, an autapomorphic state.

Etymology

‘Capixaba’ is a Brazilian word, given in apposition, to indicate anative or inhabitant from Espírito Santo state.

Goniosoma venustum Koch

(Figs 17, 18, 118, 119, 192, 193)

GoniosomavenustumKoch,1839:64.–Simon,1879:233;Roewer, 1913:258, 260, 1923: 497, 498, 1930: 382; Mello-Leitão, 1923: 155, 192,1932: 266, 268; B. Soares, 1944b: 280; Soares & Soares, 1948: 632;Moritz, 1971: 213; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118;Gnaspini et al., 2004: 34, 35.

Goniosoma geniculatum Mello-Leitão, 1931: 125. – 1935b: 110;B. Soares, 1945c: 351; Soares & Soares, 1948: 629; Muñoz-Cuevas,1972: 32;Machado et al., 2001: 22;Machado&Raimundo, 2001: 138;Machado, 2002: 389–391; Kury, 2003: 117. Syn. nov.

Material examined

Types. ’Brasilien’, 1 ma lectotype and 1 fe paralectotype (heredesignated) (ZMHB929). Rio de Janeiro: Rio de Janeiro (Jacarepaguá),ma holotype of Goniosoma geniculatum (MNRJ18207).

Other material examined. Rio de Janeiro: Santa Maria Madalena(Parque Desengano), 2 fe (MNRJ6746), A. & R. Baptista leg., XII/1991;Santa Maria Madalena (Rio do Norte, Desengano), 5 ma, 9 fe and 1 im(MNRJ6741), R. & A. Baptista leg., XII/1991; Petrópolis (Parque Nacionalda Serra dosÓrgãos,Vale do Jacó, base da Pedra doPadre), 7ma, 1 fe and 3 im(MZSP22587), M.B. DaSilva et al. leg., VII/2001.

Description

Dorsum. Eye-moundwith 1 pair ofmedium-sized and separatespines. Area I with 1 pair of tubercles like small granules. AreaIII with spines with nearly same size of spines of eye-mound.Angles of free tergites and posterior margin unarmed.

Granulation. Carapace: 0–5minute granules.Areas I–III:14–35 minute granules. Lateral margin and anal operculum:low density of minute granules. Posterior margin and freetergites: medium density of minute granules. Venter:Posterior margin of stigmatic area and free sternites: highdensity ofminute granules.Coxa I andCoxa IV:mediumdensityof minute granules.

Chelicerae. Segment I with 2 basal granules, being 1 apicaldorsal–retrolateral and 1 prolateral apical. Segment II withmedium density of small granules.

Pedipalps. Trochanter with 2–4 dorsal and 2–5 ventralelevations. Femur with 6–9 ventral elevations (standardarmature IiiIiii), 2 retrolateral subapical setae and lowdensity of minute granules in dorsal granulation. Patella with1 ventral–retrolateral subapical tubercle like a small granule.Tibia with 1 ventral–prolateral subapical tubercle like a smallgranule, in addition to the standard armature of segment.

Leg I. Trochanterwithmediumdensity ofminute granules.Femur, patella and tibia with small granules. Metatarsuspractically smooth.

Leg II. Trochanter with medium density of minutegranules. Femur and patella with medium-sized granules.Tibia with small granules. Metatarsus practically smooth.

Leg III. Trochanter with medium density of minutegranules. Femur and patella with medium-sized granules.Tibia with large granules. Metatarsus with medium-sizedgranules.

Leg IV. Coxa with medium-sized lateral granulation.Prolateral apical apophysis oblique and large; retrolateralapical apophysis pointed and thin, near the size of spines ofarea III. Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 apical (latter larger); 1 retrolateral apical apophysislike a granule, slightly curved backward; ventral granulation: lowdensity of small granules. Femur practically straight with a lightinward medial curvature; granules round; row1 graduallypointing to center, forming a comb on apical half composedby distinct granules, higher and closer to each other; row3 withspines on apical region; dorsal apical apophyses: retrolaterallarge and curved backward and prolateral small (1/3 size offormer). Tibia with row2 with high granules close to eachother in all extension; row3 with small spines from base toapical fourth, where the spines are larger and slightly curvedbackward; spine strongly curved backward on apex close to


row3. Metatarsus with row3 and row4 with large granules; therest with very small granules. Tarsal process medium-sized(type) or very long, longer than tarsal claw.

Tarsal segmentation. 10–14 (12), 24–31, 11–13 (12), 11–14(11–12).

Penis. Ventral plate wide on medial region and base, withconvex apical margin, but with a small medial concavity;extension above apical group of setae with a lateralconstriction on both side. Apical group with the more basalseta situated onmedial region of ventral plate, far from the others;basal group disposed in an inverted-L-shaped row, being 1dorsal, 1 horizontal to former, ventral, and the third more basal.Glans without ventral process and with very small dorsalprocess, like a callus. Truncus not invading the ventral plate.

Colour pattern

Dorsal scute and venter light-brown/yellowish brown,carapace and legs I–III darker; pedipalps and chelicerae darkgreen; leg IV reddish brown; light-coloured rings adjacent tojoint of tibia and metatarsus of legs. Dry-mark as a frame, in around spot divided at middle between the spines of area III,‘cup’-shaped behind of eye-mound and like a wide spot beforeeye-mound; at base of trochanter. Pink articular membranesbetween coxae and trochanters, in live animals.


Dorsal scute: width: 8.06–9.02 (9.02), length: 8.14–8.51(8.50); leg I: 39.41–44.47 (39.60), II: 83.98–95.56, III:59.07–67.74 (59.07), IV: 78.89–86.63 (84.21), femur IV:19.37–22.19 (20.59).

Female

In addition to dimorphic characters already mentioned,the female differs by: Tarsal process always medium-sized(even in same locality where males show it long). Angles offree tergites and posterior margin of dorsal scute unarmed asamong males.

Biology

Machado et al. (2001); Machado and Raimundo (2001);Machado (2002); Gnaspini et al. (2004).

Remarks

This species is very similar toG. roridum. Expressive differencesare just present in colour pattern and granulation. It shares the‘tarsal process longer than claws’ with G. capixaba sp. nov., butthis condition is polymorphic in G. venustum.

Goniosoma roridum Perty

(Figs 1, 19–21, 120, 121, 194, 195)

Goniosoma roridum Perty, 1833: 202. –Koch, 1839: 124; Gervais, 1844:108; Simon, 1879: 233; Soares & Soares, 1948: 631 (comb. reestabl.);Muñoz-Cuevas, 1972: 32; Kury, 2003: 118.

Progoniosoma roridum. – Roewer, 1913: 265, 266, 271 (comb.), 1923:500, 502, 1930: 383, 384; Mello-Leitão, 1923: 158, 192, 1932: 259,264.

Goniosoma inscriptum (misidentification). – Sabino & Gnaspini, 1999:675.

Material examined

Types. Rio de Janeiro: Teresópolis, ma neotype (here designated)(MNRJ4303), R.C.C. Baptista & M.I. Landim leg.

Other material examined. Rio de Janeiro: Teresópolis, 2 ma, 4 fe and2 im(MNRJ4303),R.C.C.Baptista&M.I.Landim leg., IV/1995;Teresópolis(Parque Nacional da Serra dos Órgãos, Rio Paquequer), 5 ma, 3 fe and 1 im(ZMHB35330), M.B. DaSilva et al. leg., VII/2001; Teresópolis (BR116),1ma and2 fe (MZSP27305),M.B.DaSilva et al. leg., VII/2001;Guapimirim(‘Estacão Ecológica Estadual Paraíso’), 2 ma, 9 fe and 5 im (MZSP15497),R. Pinto-da-Rocha & R.S. Bérnils leg., 4–7/VII/1996.

Description

Dorsum. Pair of short, straight and separate spines on eye-mound. Area I with 1 central pair of low tubercles. Area IIIwith 1 pair of large and slightly backward spines. Angles ofposterior margin of dorsal scute and free tergites with a smalltubercle.

Granulation. Carapace: 5–10 (10) small granules. AreasI–III: 51–67 (53) minute granules. Lateral margin: lowdensity of minute granules. Posterior margin and freetergites: medium density of minute granules. Analoperculum: practically smooth. Venter: Genital and stigmaticareas practically smooth. Posterior margin of stigmatic areaand free sternites: high density of minute granules. Coxa I: lowdensity of small granules. Coxa IV: medium density of minutegranules; lateral: medium density of medium-sized granules.

Chelicerae. 2 basal and 1 prolateral granules and 1 apicaldorsal–retrolateral tubercle on segment I. Segment II withsome minute granules on basal region and larger on apicalregion.

Pedipalps. Trochanter with 2–3 (2) dorsal and 4 ventralelevations. Femur with 6–9 (7) ventral elevations (standardarmature IiiIiii), 2 retrolateral subapical setae and dorsalgranulation with low density of small granules. Patella with astrong subapical ventral–retrolateral tubercle. Tibia with 2 sub-basal ventral–prolateral tubercles, like that of patella, in additionto the standard armature.

Leg I. Trochanter with low density of small granules.Femur and patella: minute granules. Other segmentspractically smooth.

Leg II. Trochanter with medium density of smallgranules. Femur with small granules. Other segmentspractically smooth.

Leg III. Trochanter with high density of small granules.Femur, patella, tibia and metatarsus with small granules.

Leg IV. Coxa with large and oblique prolateral apicalapophysis; retrolateral apical apophysis pointed, straight,with same size of spine of area III of dorsal scute (smaller thanhalf the size of trochanter). Trochanter with 2 prolateralapophyses, being 1 sub-basal and 1 subapical, latter 2� thesize of former, both slightly curved upward; retrolateral apicalapophysis curved backward and small, with 1 tubercle close toit; granulation: medium density of small granules. Femurpractically straight with a light inward medial curvature;granules round, row1 gradually pointing to center, forming acomb on apical half composed by distinct granules, higher andvery close to each other; row3with spines on apical region; dorsalapical apophyses: retrolateral large and curved backward andprolateral small (1/3 size of former). Tibia with row2 with high


granules close to each other in all extension; row3 with smallspines from base to apical fourth, where the spines are larger andslightly curved backward; strongly curved backward spine onapex close to row3. Metatarsus with row3 and row4 with largegranules; the rest with granules very small. Tarsal processmedium-sized with about 1/3 size of claws.

Tarsal segmentation. 10–13 (11/12), 23–30 (28), 10–13(12/13), 10–13 (13).

Penis. Ventral plate wide on medial region and base,with apical margin convex and straight on medial region;extension above apical group of spines with a lateralconstriction on both sides. Apical group with the more basalseta far from the others;basal groupwith rowof setae forming aninverted-L, being 1 dorsal, 1 horizontal to former, ventral, and thethird more basal. Glans without ventral process and with verysmall dorsal process, like a callus. Truncus not invading theventral plate.

Colour pattern

General dark-brown; carapace black; coxa, patella andtibia IV very dark; dorsal scute light-brown with blackmargins; ventral face light-brown, as around the joint of tibiaand metatarsus and tarsus. Dry-mark around each granule ofdorsal scute, giving punctate aspect to body, in a round spotdivided at middle between spines of area III and, on carapace,a wide spot before eye-mound and ‘cup’-shaped behind eye-mound.


Dorsal scute: width: 8.15–9.08 (9.08), length: 8.11–8.67(8.67); leg I: 38.11–43.16 (43.16), II: 83.15–95.05 (95.05),III: 59.27–66.82 (66.82), IV: 73.99–86.96 (86.96), femur IV:19.26–22.51 (22.51).

Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of posterior margin of dorsal scutewith 1 large granule each; angles of free tergites with strongtubercles.

Biology

Sabino and Gnaspini (1999).

Remarks

The species is easily recognised by the dry-mark around allgranules of the dorsal scute, giving a white-punctate aspect toits body. Although its type is lost, the original description byPerty (1833), especially concerning colour pattern, and theillustration by Koch (1839) allowed the precise identificationof the species. Therefore, we decided to designate a neotype inthe present study.

Goniosoma vatrax Koch

(Figs 22, 23, 122, 123)

Goniosoma vatrax Koch, 1848: 21. – Simon, 1879: 233; Roewer, 1913:258, 262, 1923: 497, 499; Mello-Leitão, 1923: 154, 192, 1932: 266,267; B. Soares, 1945c: 352; Soares & Soares, 1948: 632;Moritz, 1971:213; Muñoz-Cuevas, 1972: 32; Kury, 2003: 118.

Progoniosoma minense Mello-Leitão, 1932: 259, 265. – B. Soares,1945c: 352. Syn. nov. (Transferred from synonymy withG. xanthophthalmum.)

Goniosomella perlata Mello-Leitão, 1936: 33. – B. Soares, 1945c: 352.Syn. nov.

Goniosoma minense. – Soares & Soares, 1948: 629 (comb.); Muñoz-Cuevas, 1972: 32.

Lyogoniosoma perlatus (misspelling). – Soares & Soares, 1948: 634(comb.).

Lyogoniosoma perlatum. – Kury, 2003: 119.

Material examined

Types. ’Brasilien’, 1 ma lectotype and 1 ma paralectotype (heredesignated) (ZMHB931). Minas Gerais: Santa Bárbara, ma holotype ofGoniosomella perlata (MNRJ42562), Dr. L. Moraes; Ouro Preto, maholotype of Progoniosoma minense (MNRJ1409).

Other material examined. Minas Gerais: Catas Altas (Serra doCaraca), 1 ma (MZSP16376), U. Martins & R. Kloss leg., XI/1961; OuroPreto (Morro de São Sebastião), 1ma and 2 fe (MNRJ), Alceu&F. Peres leg.,VII/1955.

Description

Dorsum. Eye-moundwith 1 pair ofmedium-sized, straight andseparate spines. Area I with 1 pair of tubercles slightly distinctfrom other granules of area.Area IIIwith 1 pair ofmedium-sizedspines slightly backward close to each other (distance smallerthan distance between eyes). Angles of free tergites andposterior margin unarmed.

Granulation. Carapace: 4–10 (5) small granules. AreasI–III: 19–33 (19) small granules. Lateral margin: mediumdensity of small granules. Posterior margin and free tergites:high density of small granules. Anal operculum: mediumdensity of minute granules. Venter: Stigmatic and genitalarea and free sternites: high density of small granules. CoxaI: high density of minute granules. Coxa IV: medium density ofsmall granules.

Chelicerae. Segment Iwith about 8medium-sized basal andlateral granules in a row; II with medium density of minutegranules.

Pedipalps. Trochanter with 4–7 dorsal and 4–7 ventralelevations. Femur with 6–8 ventral elevations (standardarmature IiIiiii), 1 or 2 retrolateral subapical setae and dorsalface with low density of minute granules. Patella with 1ventral–retrolateral subapical tubercle like a small granule.Tibia with 1 short ventral–prolateral subapical spine, inaddition to the standard armature.

Leg I. Trochanter with low density of minute granules.Femurwith small granules. Patellawith minute granules. Tibiaand metatarsus practically smooth.

Leg II. Trochanterwith medium density of small granules.Femur with small granules. Patella and tibia with minutegranules. Metatarsus practically smooth.

Leg III. Trochanter with high density of small granules.Femurwith large pointed granules. Patella and tibia with smallgranules. Metatarsus with minute granules.

Leg IV. Coxa with prolateral apical apophysis almostlongitudinal; retrolateral apical apophysis pointed, a littlelarger than trochanter, oblique and slightly S-shaped.Trochanter with 1 prolateral sub-basal apophysis, 1


prolateral apical apophysis, lower than sub-basal apophysis,and 1 retrolateral apical apophysis with same size of prolateral.Femur with strong inward curvature; row2 and row3 with flatand longitudinally elongate granules (mainly on medial third);dorsal rows very granulated and highly disorganised on apicalthird; retrolateral apical apophysis large and prolateral apicalapophysis minute. Tibia with row3 armed in all extension, butwith large spines only on apical third. Metatarsus with row3and row4 armed with small but robust spines at base. Tarsalprocess medium-sized.

Tarsal segmentation. 9–10 (10), 19–21 (20), 9–10 (9),10–11(11).

Penis. Ventral plate long and narrow with convex apicalmargin and slightly concave lateral margins; apical regionextended above apical group of setae. Apical group with 1additional fifth ventral seta, smaller than others; basal groupwith setae in an oblique row.Glanswith only the stylus.Truncusnot invading the ventral plate.

Colour pattern

Colour of examined material totally lost. Roewer (1913)described it as rust yellowish brown with lateral margins ofdorsal scute black and the granules of this margin yellow.



Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of posterior margin of dorsal scuteand free tergiteswithhighandweak tubercles.Posteriormarginand free tergites with medium-sized granulation. Femur IVcurved inward only at base.

Remarks

H. Soares (1974) synonymised Progoniosoma minense withG. xanthophthalmum. We herein confirmed the observationmade by Stefanini-Jim (1985 – unpublished M.Sc. thesis) thatG.minense is actually a junior synonymofG. vatrax. The speciesis distinguished by the following diagnostic characters: prolateralapical apophysis of trochanter IV smaller than basal apophysisand retrolateral apical apophysis of coxa IV curved, reduced dry-mark, in addition to the endemism in southern Minas Gerais(Serra do Espinhaco).

Goniosoma macracanthum (Mello-Leitão), comb. nov.

(Figs 24–27, 124, 125, 196)

Progoniosoma macracanthum Mello-Leitão, 1922: 340. – 1923: 156.Lyogoniosoma macracanthum. – Mello-Leitão, 1926: 55 (comb., typespecies), 1932: 244; Roewer, 1930: 444; B. Soares, 1946: 497;Soares & Soares, 1948: 634; Kury, 2003: 119.

Material examined

Types. São Paulo: São Paulo (Cantareira), 1 ma holotype (MZSP524).

Other material examined. Minas Gerais: Santa Rita de Ibitipoca,1 ma (MZSP19237), 1997. Rio de Janeiro: Itatiaia (Parque Nacional deItatiaia), 1 ma (MZSP), F.H.S. Santos & M.N. Ferreira leg., XII/2001.

Description

Dorsum. Eye-mound with 1 pair of low, weak, and separatetubercles. Area III higher at the center, each side with 1 pair ofvery small tubercles, like other granules. Angles of free tergitesand posterior margin smooth.

Granulation. Carapace: 16–29 (27) minute granules.Areas I–III: 19–33 (19) minute granules. Lateral andposterior margins: high density of minute granules. Freetergites: medium density of minute granules. Analoperculum: low density of minute granules. Venter:Stigmatic and genital areas and free sternites: high densityof minute granules.Coxa I: low density of small granules.CoxaIV: medium density of minute granules.

Chelicerae. Segment I with 3 basal and 2 retrolateralsmall granules, 1 medial and 1 apical. Segment II with highdensity of minute granules.

Pedipalps. Trochanter with 4–6 (4) dorsal and 4–6 (4)ventral elevations. Femur with 6–8 (6/7) ventral elevations(standard armature IiiIiii), 1 retrolateral subapical seta anddorsal face with low density of minute granules. Patella with1 medium-sized ventral–retrolateral subapical tubercle. Tibiawith 1 ventral–prolateral subapical tubercle larger than that ofpatella, but weaker, in addition to the standard armature.

Leg I. Trochanter with low density of small granules.Femur, patella and tibia with small granules. Metatarsuswith minute granules.

Leg II. Trochanterwith medium density of small granules;posterior apical apophysis larger than other tubercles ofsegment, more robust. Femur, patella and tibia with smallgranules. Metatarsus with minute granules.

Leg III. Trochanter with high density of small granules,large and robust posterior apical apophysis. Femur, patella andtibia with medium-sized granulation (ventral is larger).Metatarsus with minute granules.

Leg IV. Coxa with prolateral apical apophysis insertedalmost dorsally, upward, curved down and outward; retrolateralapical apophysis pointed and very large (2 to 3� the size oftrochanter); contact zone between coxa and the lateral–posteriormargin of dorsal scute extended. Trochanter with 1 smallprolateral sub-basal apophysis and 1 retrolateral apicalapophysis pointed, curved backward, associated with granules.Femur short with strong inward curvature; row2 with flatgranules; dorsal rows disorganised and very granulated onapical third; retrolateral apical apophysis very large andprolateral apical apophysis minute. Tibia with 1 comb of largegranules close to each other on row2; row3 with small spinescurved backward, increasing from base to apex. Metatarsuswith small spines at base of row3.Tarsal processmedium-sized.

Tarsal segmentation. 9, 19–21 (21), 9–10 (10), 10–11 (11).Penis. Ventral plate very narrow, wider on basal region,

narrower on apex; apical region extended, with a lateral subapicalconstriction, convex apical margin; lateral concavity betweenapical and basal groups. Apical group with 4 subapical setae;basal group with setae disposed in a longitudinal row. Glans


with thin ventral process smaller than stylus (half the size).Truncus not invading the ventral plate.

Colour pattern

General brown; legs darker; carapace and apex of coxa IVwith apophysis black; apex of trochanters, apex of retrolateralapical apophysis of coxa IV and eye-mound yellow; groovesof areas and free segments of abdomen slightly red; pedipalpsslightly green. Dry-mark totally absent. Pink articularmembranes between coxae and trochanters, in live animals.


Dorsal scute: width: 10.39–10.97 (10.75), length:9.52–10 (9.52); leg I: 27.84–31.19 (29.1), II: 52.11–60.94(53.24), III: 40.54–47.56 (43.56), IV: 54.70–63.05 (59.14),femur IV: 13.38–16 (13.79).

Remarks

The type locality, São Paulo – Cantareira is probably mistaken,since (1) the species has not been found in recent arachnidexpeditions to Serra da Cantareira, which is still a well-preserved area; (2) the 2 examined specimens were recentlycollected in a distinct area, and such a disjunct distribution(especially considering these areas) is not common among thestudied harvestmen; and (3) this kind of mistake was notuncommon at that time, considering harvestmen and otherarthropods as well.

Although there are a few specimens of this species, itsdiagnostic characters and autapomorphies are veryconspicuous: dorsal scute totally unarmed, very largeretrolateral apical apophysis of coxa IV (almost 3� the size oftrochanter IV) and very curved prolateral apical apophysis ofcoxa IV.

Goniosoma dentipes Koch

(Figs 28–31, 126, 127, 197)

Goniosoma dentipes Koch, 1839: 58. – Simon, 1879: 228; Soares &Soares, 1948: 629 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury,2003: 117.

Goniosoma grossum Koch, 1839: 62; Simon, 1879: 233; Roewer, 1913:265 (syn. Progoniosoma dentipes (Koch, 1839)).

Progoniosoma dentipes. – Roewer, 1913: 265 (comb., type species),1923: 627, 1930: 383; Mello-Leitão, 1923: 156, 192, 1932: 259, 260.

Progoniosoma cruciferum Mello-Leitão, 1923: 155, 192. – Roewer,1930: 383, 384. Syn. nov.

Glyptogoniosoma cruciferum. – Mello-Leitão, 1932: 271 (comb., typespecies).

Acutisoma cruciferum. – B. Soares, 1945c: 350 (comb.); Soares &Soares, 1948: 623; Kury, 2003: 115.

Progoniosoma tijucaRoewer, 1930: 383. –Mello-Leitão, 1932: 259, 263.Syn. nov.

Goniosoma tijuca. – Soares & Soares, 1948: 631 (comb.); Kury, 2003:118.

Glyptogoniosoma perditum Mello-Leitão, 1936: 36. Syn. nov.Acutisoma perditum. – B. Soares, 1945c: 351 (comb.); Soares & Soares,1948: 626; H. Soares, 1974: 481; Kury, 2003: 116.

Progoniosoma grossum. – Moritz, 1971: 200.

Material examined

Types. Brasilien: 2 fe syntypes of Goniosoma grossum (ZMHB928);fe holotype of Progoniosoma tijuca (ROEWER1342). Minas Gerais:Paraopeba (probably wrong record), 1 ma, type of Glyptogoniosomaperditum (MNRJ42705), (no datum of collector). Rio de Janeiro: Rio deJaneiro (Corcovado), fe holotype of Progoniosoma cruciferum(MNRJ1453), E. Moraes Mello leg.

Othermaterial examined. Riode Janeiro:Rio de Janeiro (Alto daBoaVista), 4ma and 7 fe (MZSP1824),Maria Eugênia leg., 1944;Rio de Janeiro(Alto da Boa Vista, Tijuca), 3 ma and 1 fe (HS845), A. Peracchi leg., II/1970;Rio de Janeiro (Floresta da Tijuca), 3 ma and 2 fe (MZSP27299),M.B. DaSilva et al. leg., VII/2001; Rio de Janeiro (São Conrado, base daPedra Branca), 1 ma (MNRJ4301), S. Pott leg., II/1996.

Description

Dorsum. Eye-moundwith 1 pair of very high, divergent spinesslightly curved forward. Area I with 1 pair of round tubercles.Area IIIwith high spines. Angles of free tergites and posteriormargin with robust round tubercles, little larger than othergranules of segment.

Granulation. Carapace: 5–7 medium-sized granules.Areas I–III: 10–17 medium-sized granules. Lateral margin:medium density of small granules. Posterior margin: mediumdensity of medium-sized or large granules. Free tergites: lowdensity of medium-sized or large granules. Anal operculum:low density of small granules. Venter: Stigmatic and genitalareas and free sternites: medium density of minute granules.Coxa I: high density ofminute granules.Coxa IV: low density ofminute granules, small granules on lateral margin.

Chelicerae. Segment I with about 6 medium-sized basalgranules in a row and 1 retrolateral apical granule; II with highdensity of minute granules, denser at apex.

Pedipalps. Trochanter with 2–5 dorsal and 2–4 ventralelevations. Femur with 4–7 ventral elevations (standardarmature IiIii), 1 retrolateral subapical seta and dorsal facewith low density of minute granules. Patellawith 1 high ventral–retrolateral subapical tubercle. Tibia with 1 ventral–prolateralsubapical tubercle like that of patella, in addition to the standardarmature.

Leg I. Trochanter with low density of minute granules.Femur with small granules. Patella and tibia with minutegranules. Metatarsus practically smooth.

Leg II. Trochanterwith medium density of small granules.Femur and patella with small granules. Tibia with minutegranules. Metatarsus practically smooth.

Leg III. Trochanter with medium density of smallgranules. Femur and patella with medium-sized granules.Tibia with small granules. Metatarsus with minute granules.

Leg IV. Coxa with 1 oblique prolateral apical apophysis;retrolateral apical apophysis pointed, with near the size oftrochanter (variation: may be only half-size of trochanter insmaller animals). Trochanter with 1 small prolateral sub-basal apophysis and 1 retrolateral apical apophysis pointed,curved backward and with same size of prolateral apophysis.Femur with medium inward curvature (being almost straight onbasal half); large granules, larger on ventral rows and row5;row2 and apical half of row1 with flat granules; dorsal rowsdisorganised on apical third; retrolateral apical apophysislarge and prolateral apical apophysis minute. Tibia with a


comb of large granules close to each other on row2; large, sparsespines curved backward on row3 and 1 straight and largeretrolateral apical spine. Metatarsus armed at the basal halfof row3 with large (similar size of spines of row3 of tibia) andsparse spines, decreasing from base to apex. Tarsal processmedium-sized.

Tarsal segmentation. 9–11, 18–20, 9–13, 10–12.Penis. Ventral plate very long and narrow; apical region

extended, with 2 lateral subapical constrictions on each side,apical margin convex, but with a small concavity on medialregion. Apical group with subapical setae; basal group withsetae disposed in a longitudinal row. Glans with thin ventralprocess smaller than stylus (half the size).Truncus not invadingthe ventral plate.

Colour pattern

Redish-brown, lighter on venter, on spine of eye-moundand on granules, black on rest; pedipalp slightly green. Dry-mark on dorsal scute forming a large ‘cross’ composed by:a round area behind eye-mound, medial region and lateral–posterior margins of area I, area II, between spines of area IIIand from the lateral margins of the center of area I upto groove III; at base of trochanter IV. Pink articularmembranes between coxae and trochanters, on live animals.


Dorsal scute: width: 8.54–10.48, length: 8.69–9.36; leg I:31.95–36.06, II: 66.75–73.99, III: 49.31–55.12, IV: 63.50–70.32,femur IV: 16.76–19.16.

Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites with smallrobust spines.

Remarks

The type was not found in Senkenberg Museum, Frankfurt,where it was supposed to be deposited. However the species iseasily recognised by its diagnostic characters in descriptionsfrom type-material (e.g. Roewer 1913). It has importantautapomorphies: dry-mark like a cross on dorsal scute and 1distinct retrolateral apical spine on tibia IV, in addition to itsgeneral reddish-brown colour.

Four junior synonyms were recognised for the presentspecies, three of them in the present study. The two mainreasons to the large number of synonyms are the facts that(1) three species were described from females (which aredistinct from males because of the dimorphic characters), and(2) the type-localities are in Rio de Janeiro state, where there wasmany collections in the past, with many specimens that weredescribed as new species.

The type of the synonym Glyptogoniosoma perditum isdistinctly a male (based on the more curved femur IV andlarger retrolateral apophysis of coxa IV, for example).Although its type locality, Paraopeba-MG, is far from thepresent distribution of G. dentipes, we have seen somespecimens of the senior synonym larger and more developedlike the holotype of G. perditum, and we have concluded the

synonymy and the fact that there is a large variation of size andcurvature of femur IV among adult males of this species.We alsobelieve that the type locality of G. perditum (Paraopeba – MG)may be mistaken because the other specimens are distributed onthe littoral of Rio de Janeiro state.

Since we have types of synonyms for comparison, wepreferred not to designate a neotype for this species at this time.

Goniosoma ensifer is very similar to G. dentipes and theyare sympatric. They can be distinguished by the higherdevelopment of some structures in G. ensifer, such as thevery large retrolateral apophysis of coxa IV and verycurved femur IV, for example. Since only a few specimens ofG. ensifer were available for study and since no specimen ofG. dentipes that we examined showed the same pattern asG. ensifer, we have decided to keep the species separate,although a future and more detailed analysis may lead to theconclusion of the synonymy.

Goniosoma ensifer (Mello-Leitão)

(Figs 32–34, 128–130)

Progoniosoma ensifer Mello-Leitão, 1940: 23. – B. Soares, 1945c: 352.Goniosoma ensifer. – H. Soares, 1974: 483 (comb.); Kury, 2003: 117.

Material examined

Rio de Janeiro: Rio de Janeiro (Floresta da Tijuca), 1 ma (MZSP18959),Werner H. S. Lopes leg., I/1953 (very damaged material).

Description

Dorsum. Eye-moundwith 1 pair of very high, divergent spinesslightly forward. Area I with 1 pair of round tubercles. Area IIIwith spines of same size of spines of eye-mound, slightlybackward and close to each other (same distance as betweeneyes). Angles of free tergites and posterior margin withsmall round tubercles.

Granulation. Carapace: 6 medium-sized granules. AreasI–III: 13 small granules. Lateral and posterior margins:low density of small granules. Free tergites: low density ofsmall to medium-sized granules. Anal operculum: lowdensity of minute granules. Venter: Stigmatic and genitalareas and free sternites: low density of small granules.Coxa I: high density of minute granules. Coxa IV: lowdensity of minute granules.

Chelicerae. Segment I with 4 basal granules in a row and1 retrolateral apical granule; II with low density of minutegranules, with a denser apex.

Pedipalps. Trochanter with 4 dorsal and 4 ventralelevations. Femur with 4 or 5 ventral elevations (standardarmature IiIii), 1 retrolateral subapical seta and dorsal facewith low density of minute granules. Patella with 1 shortventral–retrolateral subapical spine. No datum for tibia.

Leg I. Trochanter with low density of minute granules.Femur and patella with small granules. Tibia and metatarsuswith minute granules.

Leg II. Trochanter with low density of minute granules.Femur and patella with small granules. Tibia and metatarsuswith minute granules.


Leg III. Trochanter with medium density of minutegranules. Femur with medium-sized granules. Patella withsmall granules. Tibia and metatarsus with minute granules.

Leg IV. Coxa with 1 oblique prolateral apical apophysis;retrolateral apical apophysis very large (2� the size oftrochanter), pointed and slightly curved inward. Trochanterwith 1 small prolateral sub-basal apophysis and 1retrolateral apical apophysis pointed, curved backward and ofsame size of prolateral apophysis. Femur with strong inwardcurvature (basal half is almost straight); large granules; row1and row2 with flat and longitudinally elongate granules(mainly on medium region); dorsal rows very disorganised andgranulated on apical third; retrolateral apical apophysis large andstrongly curved backward, prolateral apical apophysis minute.Tibia with a comb of high granules close to each other onrow2; large spines on row3 and 1 straight and large retrolateralapical spine. Metatarsus with decreasing and sparse spineson row3. No datum for tarsal process.

Tarsal segmentation. 10, 18, 11, 11.Penis. Ventral plate very long and narrow; apical region

extended, with 2 lateral subapical constrictions on each side,apical margin convex. Apical group with subapical setae; basalgroup with setae disposed in an inverted-L-shaped row. Glanswith thin ventral process smaller than stylus (half the size).Truncus not invading the ventral plate.

Colour pattern

Colours totally lost. Mello-Leitão (1940), in the originaldescription, described it as [translation] ‘general colour brown,washed in black; eye-mound spines yellow; chelicerae andpedipalps yellow, marbled in black; apophysis of legs IVlighter in colour, with red apical third; femora IV lighter incolour’.


Dorsal scute: width: 11.19, length: 10.52 (no datum for legs);measurements of Mello-Leitão (1940): legs: 37, 75, 54, 70;femur IV: 19.37.

Remarks

The type (MNRJ53935) was examined by R. Stefanini-Jimduring her study, but could not be located presently(see ‘Material’). However, the original description andillustration allowed the precise identification of the species.

Soares and Soares (1948) curiously did not cite this speciesin their monographic review.

See also remarks under G. dentipes (previous species)concerning a comparison between the two species.

Goniosoma apoain, sp. nov.

(Figs 35–37, 131, 132)

Material examined

Types. Rio de Janeiro: Nova Friburgo (Rio Grande de Cima), 1 maholotype (MNRJ 6320), A.B. Kury & R. Pinto-da-Rocha leg., X/1988;Nova Friburgo (Mury, Debossan, 950m), 1 ma paratype (MZSP15116),

Bérnils & Labiak leg., VII/1996; Teresópolis (Pedra do Sino), 1 fe paratype(MNRJ6459), A.B. Kury et al. leg., II/1989.

Description

Dorsum. Eye-mound with 1 pair of high and separate spines.Area Iwith 1 pair ofmedium-sized tubercles.Area IIIwith1 pairof slightly curved backward spines, smaller than spines of eye-mound, with base with small diameter (entire spine withpractically same diameter) and close to each other (samedistance as between eyes). Angles of free tergites andposterior margin smooth.

Granulation. Carapace: 11–19 (16) small granules. AreasI–III: 40–61 (40) small granules. Lateral margin: high densityofminute granules.Posteriormargin:mediumdensity ofminutegranules. Free tergites: medium density of small granules.Analoperculum: lowdensity of small granules.Venter:Stigmatic andgenital areas and free sternites practically smooth. Coxa I:medium density of minute granules. Coxa IV: high density ofminute granules.

Chelicerae. Segment I with some very small and sparsegranules; II with very minute granules.

Pedipalps. Trochanter with 3 or 4 (3) dorsal and 2 ventralelevations. Femur with 4–7 (4) ventral elevations (standardarmature IiiIii), 1 or 2 retrolateral subapical setae and dorsalface practically smooth. Patella with 1 ventral–retrolateralsubapical tubercle, like a minute granule. Tibia with smallventral–prolateral subapical tubercle, in addition to thestandard armature.

Leg I. Trochanter with high density of minute granules.Femur, patella, tibia and metatarsus smooth.

Leg II. Trochanter with high density of small granules.Femur and patella with minute granules. Other segmentspractically smooth.

Leg III. Trochanter with high density of small granules.Femur and patella with small granules. Other segmentspractically smooth.

Leg IV. Coxa, lateral granulation: high density of medium-sized granules; prolateral apical apophysis almostlongitudinal; retrolateral apical apophysis straight, pointed,with 2� the size of prolateral apophysis, a little smaller thantrochanter. Trochanter practically smooth with 1 smallprolateral sub-basal apophysis and 1 retrolateral apicalapophysis curved backward and with same size of prolateralapophysis and minute granules associated. Femur short withstrong inward curvature; row1 and row2 with flat andlongitudinally elongate granules (on medial region); row5 withhigher granules mainly on medial region; dorsal rowsdisorganised on apical third; retrolateral apical apophysismedium-sized and curved backward, prolateral apicalapophysis small. Tibia short and slightly curved inward; row3with short spines in all extension, 3 to 4 subapical larger spinescurved backward; granules of row2 little larger than those ofother rows. Metatarsus with row2, row3 and row4 armed withspines decreasing in size toward apex; on row2 present at thebasal third, on row3 at the basal third and on row4 reaching halfof astragalus. Tarsal process minute.

Tarsal segmentation. 8–10 (9), 14–22 (20–22), 10–11,10–12 (12).


Penis. Ventral plate very long and narrow (little wideron subapical region); apical region extended with 2 lateralsubapical constrictions on each side, apical margin triangular.Apical group with subapical setae; basal group with largesetae close to each other in an almost longitudinal row. Glanswith thin ventral process smaller than stylus (half the size).Truncus not invading the ventral plate.

Colour pattern

General brown very homogeneous. Carapace darker brown;apex of coxa, trochanter and base of femur of leg IV and pairof spines of area III of dorsal scute black. Dry-mark on dorsalscute as a very thin frame; absent on carapace; on apex of coxaand at base of trochanter and femur IV.Epidermic pigmentationof dorsal scute as a frame with reduction in large circles aroundeach granule.


Dorsal scute:width: 11.04, length: 9.68; leg I: 30.86, II: 60.10,III: 44.86, IV: 55.67, femur IV: 14.31.

Remarks

Goniosoma apoain, sp. nov. can be distinguished fromG. dentipes mainly by the dry-mark of dorsal scute followingthe grooves of areas, absence of retrolateral apical spine on tibiaIV, strongly curved femur IV and spines of area III close to eachother. The retrolateral apical apophysis of coxa IV has the samesize of trochanter IV.

Etymology

Name given in apposition. From Brazilian Tupi native language,‘apoain’ means ‘short’. Goniosoma apoain, sp. nov. has thelower ‘leg length/body length’ ratio in the subfamily, thereforebeing the species of Goniosomatinae with the shortest legs.

Goniosoma unicolor (Mello-Leitão), comb. nov.

(Figs 38–40, 133–135, 198)

Glyptogoniosoma unicolor Mello-Leitão, 1932: 462, 479.Acutisoma unicolor. – Soares & Soares, 1948: 627 (comb.); Kury, 2003:117.

Material examined

Types. Rio de Janeiro: Itatiaia, ma lectotype and 5 fe paralectotypes(here designated) (MNRJ; damaged, dry and pinned).

Other specimens examined. Rio de Janeiro: Itatiaia (ParqueNacional de Itatiaia, Rio Campo Belo), 3 ma and 1 fe (MZSP22588),F.H.S dos Santos & M.B. DaSilva leg., 9/VIII/2000; Parque Nacionalde Itatiaia, 2 ma and 1 fe (MZSP16375), R. Pinto-da-Rocha leg., IV/1991;Petrópolis (Itaipava), 1 ma and 1 fe (MNRJ6557), A.B. Kury leg.,24/XII/1989; Rio Claro, (MNRJ5530), A.B. Kury et al., I/1997; SãoPaulo: Bananal (SP247), 1 ma and 2 fe (MZSP22589), M.B. DaSilvaet al. leg., VII/2001; Ubatuba (Praia da Fortaleza), 1 ma (MNRJ17372),I. Sazima leg., 9/II/1982.

Description

Dorsum. Eye-moundwith 1 pair of high and divergent spines.Area I with 1 pair of low tubercles. Area III with spines with

same size of spines of eye-mound. Angles of free tergites withlow tubercles; posterior margin unarmed.

Granulation. Carapace: 3–7 (5) small and medium-sizedgranules. Areas I–III: 20–35 (35) small granules. Lateralmargin: medium density of small granules; higher densityof granules on margin of this area. Posterior margin and freetergites: high density of small granules. Anal operculum:high density of minute granules. Venter: Stigmatic andgenital areas and free sternites practically smooth. Coxa I:low density of minute granules. Coxa IV: low densityof minute granules; lateral: low density of medium-sizedgranules.

Chelicerae. Segment I with some very small and sparsegranules; II with many minute granules.

Pedipalps. Trochanter with 2 dorsal and 3 or 4 (3) ventralelevations.Femurwith6 to10 (9/10) ventral elevations (standardarmature IiiIiIi), 2 retrolateral subapical setae and dorsalgranulation with low density of minute granules. Patella with1 ventral–retrolateral subapical tubercle. Tibia with 1 shortventral–prolateral subapical spine, in addition to the standardarmature.

Leg I. Trochanter with low density of minute granules.Femur, patella, tibia and metatarsus smooth.

Leg II. Trochanter with medium density of minutegranules. Femur with small granules. Patella with minutegranules. Other segments practically smooth.

Leg III. Trochanter with high density of small granules.Femur with large granules. Patella with small granules. Tibiawith medium-sized granules. Metatarsus practically smooth.

Leg IV. Coxa with 1 oblique prolateral apicalapophysis; retrolateral apical apophysis pointed and alittle larger than trochanter. Contact zone between coxaand the lateral–posterior margin of dorsal scute extended(as long as distance to apical margin of coxa).Trochanter with 2 prolateral apophyses, being 1 sub-basal and 1 apical (latter larger); retrolateral apicalapophysis curved backward and with same size of atubercle of free tergites; ventral granulation: low density ofmedium-sized granules. Femur with flat granules; curvedinward, except on apical fifth where the femur is slightlycurved outward; dorsal rows disorganised after basal third,with large number of granules; row2 with longitudinallyelongate granules (mainly on medial third); row3 withhigh granules on apical half; row4 with high granules onapical fourth; retrolateral apical apophysis large andstrongly curved backward, prolateral apophysis with sizeof a granule. Row2 of tibia with very large granules; row3with small spines increasing on basal half and large spinescurved backward on apical half. Metatarsus with spines onbasal region of ventral rows. Tarsal process short.

Tarsal segmentation. 9–11 (10), 19–23 (22), 11–12 (11/12),11–14 (12).

Penis. Ventral plate with lateral margin straight; apexextended above apical group of setae with 2 lateralconstrictions on each side; apical margin trapezoid. Apicalgroup with one seta more basally placed and far from theothers; basal group with setae in an oblique row. Glans withthin ventral process smaller than stylus. Truncus not invadingthe ventral plate.


Colour pattern

General yellowish-brown. Pedipalps and femur I slightlygreen. Leg IV and carapace light-red. Dry-mark only at baseof trochanter and femur IV. Epidermic pigmentation ofdorsal scute reticulate and weak, as a frame. Pink articularmembranes between coxae and trochanters.



Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites armed withrobust and short spines; angles of dorsal scute; femur IVslightly curved outward, with basal fourth slightly curvedinward.

Remarks

The type-series, which was supposedly lost, was casually foundamong the material on exhibition at the MNRJ, dry, pinned anddamaged. However, the diagnostic characters, homogeneousyellowish-brown colour, position and size of apophysis of legIV and the large spine of eye-mound, in comparisonwithmaterialrecently collected in the type locality, Itatiaia-RJ, allowed theprecise identification of the species.

Goniosoma calcar (Roewer)

(Figs 41, 42, 136, 137, 199)

Progoniosoma calcar Roewer, 1913: 265, 272. – 1923: 500, 503, 1930:383, 384; Mello-Leitão, 1923: 155, 192, 1932: 259; B. Soares,1945c: 352.

Goniosoma calcar. – Soares & Soares, 1948: 631 (comb.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 117; Gnaspini et al., 2004: 34.

Material examined

Type. Riode Janeiro:Petrópolis, 1ma holotype (SMF884), (no datumof collector).

Other specimens examined. Rio de Janeiro: Teresópolis, 1 ma(SMF1343), (no datum of collector); Teresópolis (Fazenda Revolta), 1 ma(MZSP16367), F.A.G. Mello leg., II/1996; Teresópolis (Parque Nacional daSerra dosÓrgãos), 2 fe (MZSP20647),Rocha et al. leg.,XI/1999;Teresópolis(Parque Nacional da Serra dos Órgãos), 1 ma and 1 fe (MZSP20648), Rochaet al. leg., XI/1999;Teresópolis (ParqueNacional da Serra dosÓrgãos), 1ma(MZSP20649), Rocha et al. leg., XI/1999; Teresópolis (Parque Nacional daSerra dos Órgãos), 1 ma (MZSP20650), Rocha et al. leg., XI/1999.

Description

Dorsum. Eye-moundwith 1 pair of very high, divergent spinesslightly forward. Area I with 1 pair of low and round tubercles,like a medium-sized granule. Area III with 1 pair of low spines(about half the size of spines of eye-mound ), slightly backwardand close to each other (distance smaller than distance betweeneyes). Angles of free tergites and posterior margin practicallysmooth.

Granulation. Carapace: 5–20 (7) small granules. AreasI–III: 18–31 (30) minute granules. Lateral and posteriormargins: high density of minute granules. Free tergites andanal operculum: medium density of minute granules. Venter:Posterior margin of stigmatic area and free sternitespractically smooth. Coxa I: medium density of small granules.Coxa IV: practically smooth; lateral: low density of minutegranules.

Chelicerae. Segment I with about 4 basal granules; II withhigh density of small granules.

Pedipalps. Trochanter with 2–6 (2) dorsal and 4–6 (4/5)ventral elevations. Femur with 5–8 (5/6) ventral elevations(standard armature IiIiIi), 1 or 2 (1) retrolateral subapicalsetae and dorsal granulation with low density of minutegranules. Patella with 1 medium-sized and weak ventral–retrolateral subapical spine. Tibia with 1 ventral–prolateralsubapical spine (like that of patella), in addition to thestandard armature.

Leg I. Trochanter with high density of minute granules.Femur with small granules. Patella and tibia with minutegranules. Metatarsus practically smooth.

Leg II. Trochanter with high density of small granules.Femur and patella with small granules. Tibia with minutegranules. Metatarsus with minute granules.

Leg III. Trochanter with high density of small granules.Femur and patella with medium-sized granules. Tibia withsmall granules. Metatarsus with minute granules.

Leg IV. Coxa long and narrow with pointed prolateralapical apophysis, almost longitudinally placed and slightlyupward; retrolateral apical apophysis small and robust.Trochanter longitudinally placed, with 2 prolateralapophyses, being 1 sub-basal and 1 apical (latter larger);retrolateral apical apophysis very large and robust, straight andinward, with round apex. Femur short and with very stronginward curvature; dorsal rows very granulated and disorganisedon apical third; row2 with flat and longitudinally elongategranules (mainly on medial third); retrolateral apical apophysislarge and strongly curved backward, prolateral apophysis like agranule. Tibia curved inward with a comb of higher granuleson row2; row3 with small sparse spines, being apical larger.Metatarsus with row2, row3 and row4 armed with sparse basalspines, decreasing toward apex; spines of row2 larger andthose of row3 located only at the basal half of metatarsus.Tarsal process medium-sized.

Tarsal segmentation. 8–11 (10), 18–22 (19), 9–10 (10),11–12 (12).

Penis. Ventral plate long and narrow with apex extendedabove apical group of setae with 2 lateral constrictions on eachside; apicalmargin triangular.Apical groupwith subapical setae;basal groupwith setae in an almost longitudinal row.Glanswiththin ventral process smaller than stylus. Truncus not invadingthe ventral plate.

Colour pattern

Animal of very homogeneous black colour; venter, dorsalscute and spines of eye-mound reddish dark-brown;pedipalps lighter. Small, round dry-mark divided at middleon dorsal scute, between spines of area III (variation: dry-mark


absent on dorsal scute); on apex of coxa and base of trochanter ofleg IV.


Dorsal scute: width: 9.72–12.50 (12.50), length: 9.77–11.20(9.80); leg I: 29.50–32.94 (30.90), II: 57.20–63.82 (60.30), III:43.70–49.61, IV: 56.30–63.44 (56.30), femur IV: 14.00–15.78(14.00).

Female

In addition to dimorphic characters already mentioned, thefemale differs by: Granules of dorsal scute and free tergitessmall. Small, round dry-mark divided at middle on dorsal scute,between spines of area III, always present and strong. Angles offree tergites and posterior margin of dorsal scute as amongmales.

Remarks

This species has many autapomorphies and conspicuousdiagnostic characters as the narrow general shape of the bodydue to the longitudinal position of coxa and trochanter IV,trochanter IV with 1 very robust and straight inwardretrolateral apical apophysis, and femur IV strongly curvedinward.

Goniosoma carum (Mello-Leitão), comb. nov.

(Figs 43–45, 138–140, 200)

Xulapona cara Mello-Leitão, 1936: 32. – B. Soares, 1945c: 352. (Typeof Xulapona.)

Lyogoniosoma carum. – Soares & Soares, 1948: 633 (comb.); Kury,2003: 119.

Material examined

Types. Minas Gerais: Vicosa, 1 ma lectotype and 3 ma paralectotypes(here designated) (MNRJ42531), J. Moogen leg.

Other specimens examined. Minas Gerais: Vicosa, 1 ma and 3 fe(MNRJ5146), J. Moogen leg.

Description

Dorsum. General aspect smooth, except for the armature ofcarapace and area III. Eye-mound with very high, divergentforward spines (at least 3� the size of eye-mound).Area Iwith 1pair of tubercles like small granules. Area III with 1 pair ofparallel and slightly backward spines, lower than spines of eye-mound; distance between them same as distance between thespines of eye-mound. Angles of free tergites and posteriormargin smooth.

Granulation. Carapace and Areas I–III: large numberof minute granules. Lateral and posterior margins: highdensity of minute granules. Free tergites and analoperculum: practically smooth. Venter: Posterior marginof stigmatic area, free sternites and coxa IV: high densityof minute granules. Coxa I: medium density of minutegranules.

Chelicerae. Segment I with 4medium-sized basal granules,1 apical dorsal–retrolateral granule and 1 apical granule. SegmentII with medium density of small granules.

Pedipalps. Trochanter with 2–5 (5) dorsal and 2–5 (5)ventral elevations. Femur with 7–10 (10) ventral elevations(standard armature IiiIiiIi), 1 subapical retrolateral spine anddorsal granulation with medium density of small granules.Patella with 1 strong subapical ventral–retrolateral spine(half size of retrolateral spine of femur). Tibia with 1ventral–prolateral subapical spine, similar to that of patella, inaddition to the standard armature of segment.

Leg I. Trochanter with high density of small granules.Femur, patella and tibia with minute granules.

Leg II. Trochanter with high density of small granules.Femur and patella with small granules. Tibia and metatarsuswith minute granules.

Leg III. Trochanter with high density of small granules.Femur with large and pointed granules. Patella and tibia withsmall granules. Metatarsus with minute granules.

Leg IV. Coxa very long, with apex oblique; prolateralapical apophysis oblique and pointed, slightly curveddownward; retrolateral apical apophysis a little larger thanprolateral apophysis with longitudinal pointed apex curvedinward; presence of a retrolateral basal apophysis, close tofree sternites, like a small tubercle; presence of a bifid ventralapical apophysis of similar size of other apical apophysis.Trochanter with 2 prolateral apophyses, being 1 sub-basaland 1 apical (larger); 1 small retrolateral apical apophysissimilar to the basal apophysis of coxa. Femur S-shaped,2/3 basal with medium inward curvature; row1 and row2 withflat and longitudinally elongate granules, except on apicalfourth; dorsal rows very disorganised with medium-sizedgranulation on apical third; other rows with small granules;dorsal apical apophysis: retrolateral large and L-shaped,backward, and prolateral minute. Tibia with row3 armedwith spines in all extension (smaller on basal third); row2with smaller spines on apical third. Metatarsus armed onrow3 with small spines decreasing to apex and on row2with spines at base, the first one large. Tarsal process minute.

Tarsal segmentation. 9–14 (12), 17–31, 10–13 (12), 11–14(11–12).

Penis. Ventral plate square with slightly concave lateralmargins and wide apical margin. Apical group with 2 medialsetae more ventrally; basal group with setae very sparsedisposed in a disorganised oblique row. Ventral face withmany large bristles. Glans with ventral process with a smoothapical lamina inserted by base on a stalk. Truncus invading theventral plate medially on dorsal face.

Colour patterns

Colour totally lost. Mello-Leitão (1936) described it as[translation]: ‘General colour brown, with black spines on areaIII and coxa IV’.Dry-mark as a frame on dorsal scute and roundbetween the spines of area III.




Female

Angles of free tergites and posterior margin of dorsal scuteas among males.

Remarks

This is a very distinct and easily recognisable species inthe subfamily due to its many autapomorphies (very elongatecoxa IV, presence of ventral apophysis of coxa IV, distinct dorsalapical apophysis of femur IV, etc.) in addition to manyplesiomorphic characters of Goniosoma, since it is the basalspecies of the genus.

Since the available specimens are only a few and very old, thedescription is somewhat incomplete.

Goniosoma – species inquirenda

Remarks

The types of G. lepidum Gervais, 1844, G. monacanthumGervais, 1844, G. obscurum Perty, 1833 and G. versicolorPerty, 1833 are lost, and the species can not be recognised byoriginal description and illustration. Thus, Kury (2003) hasconsidered them to be ‘species inquirenda’. Considering thedescriptions, it is not possible to include them into the genusGoniosoma, nor even into Goniosomatinae.

Genus Pyatan, gen. nov.

(Figs 46, 47, 141, 144, 201, 202, 234)

Type species: Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini,sp. nov., here designated, by monotypy.

Diagnosis

Dorsal scute longer than wide. Eye-mound with 1 pair ofmedium-sized spines. Coxa IV with 1 pointed prolateral apicalapophysis with a small posterior sub-basal process; retrolateralapical apophysis reduced. Trochanter with 2 prolateralapophysis, the apical larger. Legs III with large pointedgranules. Femur IV large, robust and curved outward.Tarsal process medium-sized and tarsal claws smooth. Peniswith truncus invading dorsally the ventral plate; latter withthe apical margin almost straight and dorsal process finger-like.

Discussion

This genus shares many characters with its sister genusGoniosoma. It can be distinguished from Goniosoma byhaving a femur IV curved outward and robust, with strong andacuminate granules, a very peculiar penis and distinct shape ofbody.

Distribution

North littoral of São Paulo state and south littoral of Rio deJaneiro state.

Etymology

FromBrazilian Tupi native language, ‘pyatã’means ‘strong’ dueto its robust body, mainly the leg IV.

Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini,sp. nov.

(Figs 46, 47, 141–144, 201, 202)

Material examined

Types. Rio de Janeiro: Angra dos Reis (Estrada Angra-Lídice), maholotype and 8 ma, 8 fe and 1 im paratypes (MNRJ5554), A. Kury et al. leg.,II/1997;Mangaratiba (Estrada para Lídice, Sítio DonaMaria), 1 ma and 1 feparatypes (MNRJ6816), R. Sachsse & S. Potsch et al. leg., VIII/1992;Mangaratiba, 5 ma and 1 fe paratypes (MNRJ17386), E. Vasconcelosleg., 10.IV.2003; Itaguaí, 1 ma and 1 fe paratypes (MNRJ17377), Mattos&Maciel leg., 1948; Itaguaí, 1 ma paratype (MNRJ 17374);Angra dos Reis(Cachoeira do RioManbucaba), 1 ma paratype (MZSP22582),M.B. DaSilvaet al. leg., VII/2001;Angra dosReis, 1 ma and 1 fe paratypes (MNRJ42427),no data of collector (‘type’material ofR. Stefanini-Jim).SãoPaulo:Ubatuba(Núcleo Picinguaba, Poco dos Amores), 2 ma and 1 fe paratypes (SMF), M.B. DaSilva & F.H.S. Santos leg., VIII/2000.

Description

Dorsum. Eye-mound low with medium-sized spines far fromeach other. Area I with 1 pair of tubercles like small granules.Area III with 1 pair of slightly backward spines higher thanspines of eye-mound. Angles of free tergites and posteriormargin with tubercles like very large and robust granules.

Granulation. Carapace: 0–7 (2) minute granules. AreasI–III: 10–19 (19) minute granules. Lateral margin and analoperculum: low density of minute granules. Posterior marginand free tergites: high density of minute granules. Venter:Posterior margin of stigmatic area and free sternites:medium density of minute granules. Coxa I: high density ofminute granules. Coxa IV: practically smooth.

Chelicerae. Segment I with 2 basal and 1 dorsal–retrolateralapical small granules. Segment II with medium density ofminute granules.

Pedipalps. Trochanter with 1–3 (3) dorsal and 3 or 4 (3)ventral elevations. Femur with 6–8 (7) ventral elevations(standard armature IiiIiii), 1 retrolateral subapical seta anddorsal granulation with low density of minute granules. Patellawith 1 low ventral–retrolateral subapical tubercle. Tibia with 1high and strong ventral–prolateral subapical tubercle, in additionto the standard armature of segment.Granules of femora, patellae,tibiae and metatarsi of legs pointed and backward.

Leg I. Trochanter with medium density of minutegranules. Femur and patella with small granules. Tibia withminute granules. Metatarsus smooth.

Leg II. Trochanter with medium density of minutegranules. Femur and patella with medium-sized granules.Tibia with small granules. Metatarsus smooth.

Leg III. Trochanter with medium density of minutegranules. Femur and patella with large granules. Tibia withsmall granules.Metatarsus smooth. Femur and tibia with smallapical spines on row3.

Leg IV. Coxa with oblique and pointed prolateral apicalapophysis, with a small basal process, and retrolateral apicalapophysis very small, thin and pointed. Trochanter with 2prolateral apophyses, being 1 sub-basal and 1 apical (2�larger); retrolateral apical apophysis small like a medium-sized granule. Femur robust and curved outward; largegranules, very large on row2 and row4; spines very large,


increasing on apical half of row3; small dorsal apical apophysesvery similar to each other. Tibia with 3 apical spines on row3.Metatarsus with granules, larger on ventral rows. Tarsalprocess medium-sized. Tarsal claws smooth.

Tarsal segmentation. 10–12 (11–12), 18–27 (26/27), 11–12(11), 11–14 (11–12).

Penis. Ventral plate square, with slightly concave lateralmargins and slightly concave apical margin, almost straight.Apical group with 4 setae in a dorsal row (more basal verysmall) and 1 ventral subapical smaller seta; basal groupwith 3 or4 slightly spatulate setae in an oblique row. Ventral face with

many bristles.Glanswith a ventral process with an apical laminawith projections on margin, inserted by base; stylus cylindrical;dorsal process finger-like.Truncus invading dorsally the ventralplate.

Colour pattern

Yellowish-brown; carapace black; leg IV slightly red;pedipalps and chelicerae dark green; venter light-brown.Dry-mark as a frame on abdominal scute; in a round spotbetween spines of area III; round spot behind eye-mound;apex of coxa, base of trochanter and base of femur of leg IV.

68 69

67

73

70

7271

Figs 67–73. 67–70, Serracutisoma inerme. 67, Habitus, dorsal view. 68, 69, Right femur IV (dorsal view) of twodifferent adult males, showing variation in size and development. 70, Femur, patella and base of tibia of left pedipalp(ventral–prolateral view). 71–73, Serracutisoma guaricana, sp. nov. 71, Habitus, dorsal view. 72, Right femur IV(dorsal view). 73, Apex of tibia IV (ventral view), showing armature of rows2, 3 and 4. Scale = 4mm, except for 70 = 2mm.


Pink articular membranes between coxae and trochanters,on live animals.



Remarks

It was recognised as an undescribed species by Stefanini-Jim(1985) in her unpublished M.Sc. thesis, and corroboratedduring the present study.

Etymology

From Latin, ‘unexpected’. Name created by R. Stefanini-Jimin her unpublished M.Sc. thesis and maintained herein in orderto avoid confusions.

74

76

75

77

79

80 81

78

Figs 74–81. 74–77, Serracutisoma spelaeum. 74, Habitus, dorsal view. 75, Right femur IV (dorsal view). 76, Apex ofright femur III (retrolateral view). 77, Right pedipalp (ventral–retrolateral view). 78, Heteromitobates discolor. Habitus,dorsal view. 79, 80, Heteromitobates inscriptus. 79, Habitus, dorsal view. 80, Eye mound (posterior view), with a pairof ‘pawn-shaped’ tubercles. 81, Heteromitobates albiscriptus, apex of coxa, trochanter, femur, patella and base of righttibia IV – this figure may also represent the leg IV of H. inscriptus and H. discolor, since the legs IV of these three speciesare very similar to each other. Scale = 4mm, except for 77 = 2mm, 80 = 1mm.


Genus Serracutisoma Roewer, reestabl.

(Figs 4, 5, 48–77, 145–163, 203–218, 234, 236)

Serracutisoma Roewer, 1930: 349, 447. –Mello-Leitão, 1932: 232, 251;B. Soares, 1944a: 260 (syn. Acutisoma Roewer, 1913).

Acutisomelloides Mello-Leitão, 1932: 233, 272, 479. – 1935b: 110;B. Soares, 1944a: 260 (syn. Acutisoma Roewer, 1913). (Type =Acutisoma inerme Mello-Leitão, 1927, by original designation.)Syn. nov.

SpelaeosomaMello-Leitão, 1933: 139. – 1935b: 110; B. Soares, 1945c:352; Soares & Soares, 1948: 628 (syn. Goniosoma Perty, 1833).(Type = Spelaeosoma spelaeum Mello-Leitão, 1933, by originaldesignation.) Syn. nov.

Pygosomoides Mello-Leitão, 1933: 140. – 1935b: 110; B. Soares,1944a: 263 (syn. Acutisoma Roewer, 1913). (Type =Pygosomoides mollis Mello-Leitão, 1933, by original designation.)Syn.nov.

Type species: Acutisoma proximum Mello-Leitão, 1922, by monotypy.

82

83

area 3sp

85

92

91 93 89

row3

row2

row2

row3

gr

row2

88

90

87

86

84

Figs 82–93. 82–86, Heteromitobates anarchus, sp. nov. 82, Habitus, dorsal view. 83, Dorsum, right view. 84, Rightfemur IV (dorsal view); sp = spine. 85, Right tibia IV (dorsal view). 86, Trochanter, femur, patella and base of tibia ofright pedipalp (ventral–retrolateral view); gr = large granules. 87–90.Heteromitobates harlequin, sp. nov. 87,Habitus, dorsalview. 87, 89, Left femur IV (dorsal and ventral views). 90, Femur, patella and base of tibia of left pedipalp (ventral–prolateralview). 91–93, Heteromitobates alienus, sp. nov. 91, Habitus, dorsal view. 92, Right apex of coxa, trochanter, femur andpatella IV (dorsal view). 93, Right metatarsus IV (dorsal view). Scale = 4mm, except for 83, 86, 90 = 2mm.


Diagnosis

Eye-moundwith 1 pair of tubercles (exception: 1 pair ofmedium-sized spines in S. catarina). Coxa IV with 1 large prolateralapical apophysis with falcate apex; the retrolateral apicalapophysis reduced or like a medium-sized spine. TrochanterIV with 1 dorsal–prolateral apical apophysis curvedinward; the retrolateral apical apophysis small or large, curvedinward or like a large spine, straight. Femur IV straight or sharply

bent inward and upward (exception: slightly curved inward inS. pseudovarium, sp. nov.), armed on row2; dorsal apicalapophyses of near size to each other. Femur III armedventrally on row3. Tarsal process minute and tarsal clawssmooth. Penis with hexagonal ventral plate; apical groupwith dorsal row with 5 setae (the second and the last one, fromapex to base, are smaller) and ventral row with 1 seta; basalgroup transversal (or almost) with 4 setae; all setae spatulateand wide. Glans with ventral process with an apical lamina

94 95

96

97

99

100

102

101

98

raaf paaf

Figs 94–102. 94, Mitogoniella indistincta, habitus, dorsal view. 95–98, Mitogoniella taquara, sp. nov. 95, Habitus,dorsal view. 96, Apex of trochanter and right femur IV (dorsal view). 97, Joint between right femur and patella IV(dorsal view), showing prolateral apical apophysis of femur (paaf) and retrolateral apical apophysis of femur (raaf). 98,Right pedipalp (ventral–retrolateral view). 99–102, Mitogoniella unicornis, sp. nov. 99, Habitus, dorsal view. 100,Eye mound (posterior view), with a single medial spine. 101, Right femur IV (dorsal view). 102, Femur of left pedipalp(ventral–retrolateral view). Scale = 4mm, except for 100 = 1mm, 97, 98, 102 = 2mm.


inserted by base and apical margin with many projections;dorsal process small, like a callus, or absent; styluscylindrical. Truncus invading the ventral plate medially ondorsal face.

Distribution

Santa Catarina, Paraná and southern São Paulo.

Key for the species of Serracutisoma based on males

1. Retrolateral spines of femur IV large and robust reducing from basetoward apex (Fig. 49).........................................................................2

Spines with other distribution, more restricted to specific parts of femurIV, as basal, medial, or subapical regions (Fig. 69) ..........................4

2. Angles of posterior margin of dorsal scute and tergites armed (Fig. 51);dry-mark absent on dorsal scute, region around granules of dorsalscute darker (Fig. 207); femur III with ventral spines very close to eachother (Fig. 56) ....................................................................................3

Angles of posterior margin of dorsal scute and tergites unarmed (Fig. 67);dry-mark as a Y behind eye-mound; femur III with sparse ventralspines.................................................................S. proximum (p. 565)

3. Femur IV bent inward subapically, where the row of retrolateral spinesends; retrolateral apical apophysis of trochanter IV curved backwardand smaller than other apophysis of trochanter ............................................................................................................S. banhadoae (p. 572)

Femur IV straight and with large spines almost reaching the apex;retrolateral apical apophysis of trochanter straight inward and largerthan other apophysis of trochanter IV .........S. thalassinum (p. 575)

4. Ventral spines of femur III sparse with 2 or 3 larger (Fig. 76); tibia IVventrally armed on apex (Fig. 73); prolateral apophysis of coxa IValmost transversal (Fig. 74)................................................................5

Ventral spines of femur III small, with same size and close to each other(Fig. 56); tibia IV unarmed; prolateral apophysis of coxa IV oblique(Fig. 53) .............................................................................................8

5. Retrolateral apical apophysis of trochanter IV like a granule, smaller thanother apophyses of trochanter IV (Fig. 63); eye-mound with 1 pair ofspines; dorsal scute withwhite dry-mark following the grooves of areasand forming a Y behind eye-mound ....................S. catarina (p. 577)

Retrolateral apical apophysis of trochanter IV of similar size or largerthan other apophyses of trochanter IV (Fig. 71); eye-moundwith 1 pairof tubercles; dorsal scute without dry-mark or restricted to the carapace...........................................................................................................6

6. Granules of dorsal scute very large (Fig. 5); femur IV bent inward on apicalthird, with spines increasing toward the apical third; animal withgeneral colour dark green....................................... S. inerme (p. 578)

Granules of dorsal scute small or minute (Figs 1, 2); femur IV mediallybent inward (Fig. 72) .........................................................................7

7. Area III with 1 pair of spines; granules of dorsal scute small; dry-markabsent on dorsal scute ........................S. guaricana, sp. nov. (p. 579)

Area III with 1 pair of very low tubercles; dorsal scute withminute granules, almost not visible; dry-mark behind eye-mound.........................................................................S. spelaeum (p. 580)

8. Dry-mark following the grooves of areas and as a Y behind eye-mound;femur IV armed with 2 or 3 large subapical spines close to each other........................................................S. fritzmuelleri, sp. nov. (p. 576)

Dry-mark on dorsal scute absent, but with darker region around granules(Fig. 207); femur IV with medium-sized or very small spines; dorsalscute with large granules (Fig. 4).......................................................9

9. Femur IV with medium-sized medial spines; area III with 1 pair ofspines.........................................................................S. molle (p. 573)

Femur IV with small subapical spines; area III with 1 pair oftubercles .......................................S. pseudovarium, sp. nov. (p. 574)

Key for the species of Serracutisoma based on females

1. Femur IV armed on basal region of row2...............................................2Femur IV unarmed..................................................................................4

2. Dry-mark as a frame on dorsal scute and as a Y on the carapace; granulesof areas of dorsal scute small (Fig. 204)...........S. proximum (p. 565)

Dry-mark absent on dorsal scute, but with serous accumulation whichallows whitish appearance to the animal; granules of areas of dorsalscute medium-sized or large ..............................................................3

3. Serous accumulation on whole dorsal scute, except around granulesand on lateral region of the carapace; spines on basal region of femurIV near the size of spine of angle of posterior margin of dorsalscute .................................................................S. banhadoae (p. 572)

Serous accumulation on whole dorsal scute, except only around granules;spines on basal region of femur IV much smaller than spine of angleof posterior margin of dorsal scute, with height same as diameter(Fig. 211) .......................................................S. thalassinum (p. 575)

4. Dry-mark as a frame on dorsal scute and as a Y on the carapace ..........5Dry-mark absent or restricted to other regions .......................................6

5. Epidermic pigmentation on abdominal scute, mainly, fragmented, inaddition to frame dry-mark (Fig. 214) .........................................................................................................S. fritzmuelleri, sp. nov. (p. 576)

Epidermic pigmentation on dorsal scute homogeneous, in addition toframe dry-mark ....................................................S. catarina (p. 577)

6. Granules of dorsal scute very large and yellow.......... S. inerme (p. 578)Granules of dorsal scute smaller and, generally, with same colour of rest

of scute ...............................................................................................77. Strong serous layer on whole dorsal scute, except around granules,

which allow grayish aspect to animal; granules of dorsal scutemedium-sized or large........................................................................8

Serous layer normal (homogeneous); small or minute granules .............98. 16 to 41 granules on areas of dorsal scute; spines on area II ...................

..................................................................................S. molle (p. 573)More than 42 granules on areas of dorsal scute; conic tubercles on

area III (Fig. 207).........................S. pseudovarium, sp. nov. (p. 574)9. Dry-mark round on the carapace behind eye-mound................................

........................................................................... S. spelaeum (p. 580)Dry-mark absent on dorsal scute.............S. guaricana, sp. nov. (p. 579)

Serracutisoma proximum (Mello-Leitão), comb. nov.

(Figs 48–50, 145, 146, 203, 204)

Acutisoma proximum Mello-Leitão, 1922: 343. – 1923: 161; B. Soares,1944a: 262 (comb. reestabl.), 1944b: 280, 1945b: 232, 1945c: 351,1946: 496; Soares & Soares, 1948: 627; Ramires & Giaretta, 1994;Machado & Raimundo, 2001: 138; Machado et al., 2001: 17; Kury,2003: 116; Willemart & Gnaspini, 2004a: 16.

Serracutisoma proxima. – Roewer, 1930: 447 (comb.); Mello-Leitão,1932: 251.

Leitaoius ornatus Mello-Leitão, 1934: 412. – 1935b: 111, 1940: 25;B. Soares, 1944a: 262 (syn.AcutisomaproximumMello-Leitão, 1922).

Leitaoius viridifronsMello-Leitão, 1935a: 402. – 1935b: 111, 1940: 25.Syn. nov.

Leitaoius iguapensisPiza, 1938: 139.–Mello-Leitão, 1940: 25.Syn. nov.Acutisoma viridifrons. – Soares & Soares, 1948: 627 (comb.); Kury,2003: 117.

Acutisoma iguapense. – Soares & Soares, 1948: 624 (comb.); Kury,2003: 116.

Goniosoma proximum. – Gnaspini & Trajano, 1994: 566; Pinto-da-Rocha, 1995: 81; Gnaspini, 1995: 138, 1996: 417, 433, 1999;Gnaspini & Cavalheiro, 1998: 82; Willemart, 2001: 251, 2002: 51,55; Machado, 2002: 389–391.


Material examined

Types. São Paulo: ma lectotype and 3 ma and 2 fe paralectotypes(MZSP528) (herein designated, following Stefanini-Jim’s unpublishedM.Sc. Thesis); Iguape (Barra do Rio Ribeira do Iguape), ma lectotypeand ma paralectotype of Leitaoius iguapensis (MZLQ0025), F. Lane leg.,VII/1937.

Other specimens examined. São Paulo:Cubatão (Vale do Quilombo),2 ma and 2 fe (HS1069), A. Giaretta leg., 16/IX/1990; Peruíbe (EstacãoEcológica Juréia-Itatins), 2 ma and 4 fe (MZSP16475), R. Barichelloleg., VI/1998; Miracatu, 1 fe (SMF40501), R.P. de Campos leg.,X/2001; Tapiraí, 4 ma and 5 fe (IBSP308), C. Rheims & V.C. Onofrio

leg., IX/1997; Iporanga (Fazenda Intervales), 2 ma, 7 fe and im’s(MZSP16369), P. Gnaspini leg., VI/1992; Iporanga (Cachoeira doMirante), 2 ma (SMF40502), F.H.S. Santos et al. leg., XI/1999;Iporanga (Parque Estadual Intervales, Cachoeira do Mirante), 1 ma and2 fe (ZMHB35331), F.H.S. Santos & M.B. DaSilva leg., II/2000; Iporanga(Gruta do Tatu), 1 ma and 4 fe (NHMW21179), P. Gnaspini et al. leg.,XI/1999.

Description

Dorsum. Eye-mound lowwith 1 pair of round tubercles a littlesmaller than mound. Area I with the main pair of elevations

103

109

108 107raafraaf

104

110

111

105

106

Figs 103–111. 103, Mitogoniella modesta, habitus, dorsal view. 104–106, Acutisoma coriaceum. 104, Habitus, dorsalview. 105. Right femur IV (dorsal view). 106, Trochanter and femur of right pedipalp (ventral–retrolateral view) – noticemedium-sized granules on the dorsal surface (below). 107, 108, Acutisoma hamatum. 107, Habitus, dorsal view. 108, Leftfemur IV (dorsal view). 109–111, Acutisoma longipes. 109, Eye mound (posterior view), showing that the base betweentubercles is placed above the eye level. 110, 111, Right femur IV (dorsal view), showing polymorphism; raaf = retrolateralapical apophysis of femur. Scale = 4mm, except for 106 = 2mm, 109= 1mm.


indistinct; area III with 1 pair of short, robust and slightlybackward spines. Angles of posterior margin with tubercleslike tubercles of eye-mound; angles of free tergites and analoperculum with spines with 2� the size of tubercles ofposterior margin of dorsal scute.

Granulation. Carapace: 6–13 (11)medium-sized granules.Areas I–III: 17–27 (20) medium-sized granules. Lateralmargin, posterior margin and free tergites: low density ofmedium-sized granules. Anal operculum: low density of small

granules. Venter: Posterior margin of stigmatic area and freesternites: medium density of minute granules. Coxa I: lowdensity of minute granules. Coxa IV: low density of smallgranules; lateral: large granules.

Chelicerae. Segment I with 2 pairs of minute granules,being 1 basal and 1 apical. Segment II with medium density ofsmall granules.

Pedipalps. Trochanter with 2 or 3 (2) dorsal and 2 or 3 (2)ventral elevations. Femur with 6–8 (6/7) ventral elevations

112 113 114

117116115

118 119 120

123122121

Figs 112–123. Male genitalia. 112–114, Goniosoma varium, dorsal and lateral views, and detailed lateral view of glans.115–117, Goniosoma capixaba, sp. nov., dorsal and lateral views, and detailed lateral view of ventral process. 118, 119,Goniosomavenustum, dorsal and lateral views. 120, 121,Goniosomaroridum, dorsal and lateral views. 122, 123,Goniosomavatrax, dorsal and lateral views. Scale = 0.05mm.


(standard armature IiiIiii), 1 retrolateral subapical seta anddorsal granulation with medium density of minute granules.Patella: small ventral–retrolateral subapical tubercle like aminute granule. Tibia with a low prolateral sub-basal tubercle,in addition to the standard armature of segment.

Leg I. Trochanter smooth, except standard tubercles ofsubfamily. Femur and patella with minute granules. Othersegments practically smooth.

Leg II. Trochanter smooth, except standard tubercles ofsubfamily. Femur and patella with minute granules. Othersegments practically smooth. Tibia with sparse minute spineson row3.

Leg III. Femur armed on apical region of row3 with sparsesmall spines. Trochanter with low density of minute granules.Femur, patella and tibia with minute granules. Metatarsuspractically smooth.

124 125

127 128 129

126

132131130

133 134 135

Figs 124–135. Male genitalia. 124, 125, Goniosoma macracanthum, dorsal and lateral views. 126, 127, Goniosomadentipes, dorsal and lateral views. 128–130,Goniosoma ensifer, dorsal and lateral views, and detailed lateral view of glans.131, 132,Goniosoma apoain, sp. nov., dorsal and lateral views. 133–135,Goniosoma unicolor, dorsal and lateral views, anddetailed lateral view of the basal group of spines. Scale = 0.05mm.


Leg IV. Prolateral apical apophysis of coxa oblique;retrolateral apical apophysis like spines of angles of freetergites. Trochanter with dorsal–prolateral apical apophysisupward and retrolateral apical apophysis curved backwardof similar size of retrolateral apophysis of coxa. Femur withsmall and straight granules; row2 armed in all extension(large spines on basal half, decreasing toward apex); row1

armed with large spines on basal third, like those of row2.Patella and tibia with small granules.

Tarsal segmentation. 9–13, 17–23, 10–13, 11–14.Penis. Ventral plate with concave apical margin, like a V;

ventral face with bristles. Apical group of setae with the moreventral seta on medial region. Glans with very small dorsalprocess, like a callus.

136 137 138

141140

143142

145 146 147

144

Figs 136–147. Male genitalia. 136, 137,Goniosoma calcar, dorsal and lateral views. 138–140,Goniosoma carum, dorsaland lateral views, and detailed lateral viewof glans. 141–144,Pyatan insperatum, gen. nov., sp. nov., dorsal and lateral views,detailed lateral view of glans, and ventral–lateral view showing setae (arrows). 145, 146, Serracutisoma proximum, dorsaland lateral views. 147, Serracutisoma banhadoae, dorsal view. Scale = 0.05mm.


Colour pattern

Dorsal scute dark-brown with regions, like margins, slightlygreen; pedipalps and chelicerae dark green; legs I–III andventer reddish-brown; leg IV black; joint of femur and tibia oflegswith a lighter ring; variation: granulesmay be yellow,mainlyon margins of dorsal scute. Dry-mark as a frame on abdominalscute, like a Y behind eye-mound, on apex of coxa, wholetrochanter and base of femur of leg IV. Black articularmembranes between coxae and trochanters.


Dorsal scute: width: 7.89–8.57 (7.94), length: 7.42–8.46(7.72); leg I: 42.88–53.21, II: 120.6–152.4, III: 66.01–77.25,IV: 85.47–99.84, femur IV: 24.14–28.64 (26.94).

Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites with spinesof same size of those of area III, but more robust; angles of

148 149 150

153

156155154

157 158 159

151 152

Figs 148–159. Male genitalia. 148, Serracutisoma banhadoae, lateral view. 149, 150, Serracutisoma molle, dorsal andlateral views. 151, 152, Serracutisoma pseudovarium, sp. nov., dorsal and lateral views. 153, 154, Serracutisomathalassinum, dorsal and lateral views. 155–157, Serracutisoma fritzmuelleri, sp. nov., dorsal and lateral views, anddetailed lateral view of glans. 158, 159, Serracutisoma inerme, dorsal and lateral views. Scale = 0.05mm.


posterior margin of dorsal scute with spines of half the size ofspines of free tergites. Femur IVwith large spines on basal thirdof row1 and row2. Leg IV dark-brown and epidermicpigmentation of dorsal scute fragmented as a frame.

Biology and records

Gnaspini and Trajano (1994) [cav.]; Ramires and Giaretta (1994)[biol.];Gnaspini (1995 [cit.], 1996 [biol.], 1999 [biol.]); Pinto-da-

Rocha (1995) [cav.]; Gnaspini and Cavalheiro (1998) [biol.];Machado and Raimundo (2001) [cit.]; Willemart (2001 [cit.],2002 [cit.]); Machado (2002) [cit.]; Willemart and Gnaspini(2004a) [cit.].

Remarks

Since this species is very abundant in forests and, sometimes,entrances of caves in the Speleological Province of Vale do

160 161 162

165

168

171170

167

169

166

163 164

Figs 160–171. Male genitalia. 160, 161, Serracutisoma guaricana, sp. nov., dorsal and lateral views. 162, 163,Serracutisoma spelaeum, dorsal and lateral views. 164, 165, Heteromitobates inscriptus, dorsal and lateral views. 166,167,Heteromitobates albiscriptus, dorsal and lateral views. 168, 169,Heteromitobates anarchus, sp. nov., dorsal and lateralviews. 170, 171, Heteromitobates harlequin, sp. nov., dorsal and lateral views. Scale = 0.05mm.


Ribeira, itwas included in somenatural history studies about cavegoniosomatines (synonymic list). It was cited as Goniosomaproximum in most above cited publications followingR. Stefanini-Jim’s identification and suggestion (in herunpublished 1985M.Sc. Thesis) that the subfamily wouldhave only the genus Goniosoma. Therefore, this combinationhas never been officialised in a taxonomic review. This speciesis easily recognised by the dry-mark forming a Y behind theeye-mound (shared with S. fritzmuelleri, sp. nov.), very elongatelegs and small body, and femur IV with decreasing spines onrow1 and row2.

Serracutisoma banhadoae (Soares & Soares), comb. nov.

(Figs 51, 52, 147, 148, 205)

Acutisoma banhadoae Soares & Soares, 1947a: 64. – 1947b: 210, 1947c:250, 1948: 625; Kury, 2003: 115.

Material examined

Type. Paraná: Piraquara (Banhado), 1 ma paratype (MZSP1862),Hatschbach & Imaguire leg., IX/1944.

Other specimens examined. Paraná: Piraquara (Banhado), 4 ma(MZSP1026), Imaguire leg., IV/1946; Piraquara (Banhado), 1 ma(MZSP984), Gofferjé leg., XII/1945; Piraquara (Banhado), 1 ma(MZSP1860), Hatschbach leg., XI/1945; São José dos Pinhais (Represade Guaricana), 1 ma (MZSP27300), F.C. Pioker leg., IX/2000; São Josédos Pinhais (Represa de Guaricana, mata), 1 ma (ZMHB35332), R.L. Pintoleg., I/2001; São José dos Pinhais (Represa de Guaricana), 2 ma(NHMW21180), M.R. Hara leg., II/2001; São José dos Pinhais(Represa de Guaricana, mata), 1 ma and 1 fe (MZSP22576),R.H. Willemart et al. leg., VI/2001.

Description

Dorsum. Eye-mound low with 1 pair of separate, smalland round tubercles. Area I with 1 pair of tubercles a littlelarger than granules of area. Area III with 1 pair of short andslightly backward spines. Posterior margin and free tergiteswith nearly 1 pair of medium-sized and elongate laterallygranules, in addition to tubercles of angles. Tubercles ofangles of posterior margin round and high; angles of freetergites with tubercles a little larger than tubercles of angles ofposterior margin.

Granulation. Carapace: 8–14 medium-sized granules.Areas I–III: 21–34 medium-sized granules. Lateral margin:medium density of medium-sized granules. Posterior marginand free tergites: low density of medium-sized granules. Analoperculum: low density of small granules. Venter: Posteriormargin of stigmatic area and free sternites: medium density ofsmall granules. Coxa I: high density of minute granules. CoxaIV: low density of small granules; lateral: medium-sizedgranules.

Chelicerae. Segment II with high density of medium-sizedgranules.

Pedipalps. Trochanter with 2–4 dorsal and 2 or 3 ventralelevations. Femur with 6–8 ventral elevations (standardarmature IiiIiii), 1 retrolateral subapical seta and dorsalgranulation with medium density of medium-sized granules.Patella: tubercle like a small ventral–retrolateral subapical

granule. Tibia with 1 low and round sub-basal prolateraltubercle, in addition to the standard armature of segment.

Leg I. Trochanter with low density of small granules.Femur and patella with small granules. Tibia and metatarsuswith minute granules.

Leg II. Trochanter with low density of small granules.Femur and patella with small granules. Tibia and metatarsuswith minute granules; all extension of row2 of tibia andmetatarsus with sparse minute spines.

Leg III. Femur armed on apical region of row3 with smallspines very close to each other. Trochanter with low density ofsmall granules. Femur and patella with small granules. Tibiaand metatarsus with minute granules.

Leg IV. Coxa: prolateral apical apophysis oblique;retrolateral apical apophysis like a small spine as thetubercles of angles of free tergites. Trochanter withdorsal–prolateral apical apophysis curved inward andretrolateral apical apophysis curved backward, a little largerthan retrolateral apophysis of coxa. Femur slightly bent inwardand upward on apical third; row2 armed with large spines fromsub-basal region to the bend (up to the apical third); row3 withsmall increasing spines at apical direction; dorsal apicalapophyses small, retrolateral apophysis larger. Tibia withsmall spines on apex of row3.

Tarsal segmentation. 9–10, 16–20, 10–11, 10–14.Penis. Ventral plate very wide; apical margin slightly

concave. Apical group with more ventral seta directed to3rd seta of apical group. Glans with dorsal process verysmall, like a callus.

Colour pattern

Animal strongly black; legs I–III dark-brown; dorsal scutevery dark brown; pedipalps and chelicerae green; joint of tibiaand metatarsus of legs with a lighter ring; serous layer on dorsalscute allowing grayish appearance to animal, but, aroundgranules, the layer is weak. Dry-mark on apex of coxa, baseof trochanter and femur of leg IV. Black articular membranesbetween coxae and trochanters. Variation: granules of dorsalscute, mainly those of lateral and posterior margins may beyellow.


Dorsal scute: width: 8.80–9.98, length: 7.77–8.57; leg I:39.02–44.62, II: 89.48–106.33, III: 56.29–63.77, IV:72.81–85.24, femur IV: 21.03–23.41.

Female

In addition to dimorphic characters already mentioned,the female differs by: Granules of lateral and posteriormargins and free tergites are larger (large or very large) andwith higher density. Angles of posterior margin of dorsalscute and free tergites with spines of same size of spines ofarea III, but more robust. Femur IV with spines on row2decreasing in size toward the basal half. Colour pattern:dorsal scute brown with dark green margins, carapace blackand granules yellow; coxa IV, trochanters and base of femurIV brown; rest of legs, pedipalps and chelicerae green. Epidermic


pigmentation of dorsal scute fragmented and as a frame. Dry-mark on whole dorsum of trochanter IV.

Remarks

We could not examine the holotype (deposited in Gofferjé’sprivate collection, in Blumenau, Santa Catarina), but we haveseen the paratype, allowing the precise identification ofthe species. Soares and Soares (1947a) described in details thisspecies and they discussed its affinity with Acutisomamarumbicola H. Soares, 1945. The latter have beensynonymised here with S. thalassinum and it can bedistinguished by the direction of prolateral apical apophysisof coxa IV and distribution of spines on femur IV; inS. banhadoae, these spines end in subapical position.

Serracutisoma molle (Mello-Leitão), comb. nov.

(Figs 53–56, 149, 150, 206, 207)

Pygosomoides mollisMello-Leitão, 1933: 140. – 1935b: 111; B. Soares,1944a: 263.

Acutisoma molle. – B. Soares, 1944a: 263 (comb.), 1945a: 192, 1945b:231, 1945c: 351, 1946: 496; H. Soares, 1945: 208; Soares & Soares,1947a: 64, 1947b: 210, 1947c: 250, 1948: 625;Hara&Gnaspini, 2003:258, 259, 262, 264, 266, 269, 273; Kury, 2003: 116.

Leitaoius guttulatus Mello-Leitão, 1934: 413. – 1935b: 111, 1940: 25;B. Soares, 1943: 205, 1944a: 263 (syn.Acutisomamolle (Mello-Leitão,1933)).

Goniosoma aff. badium. – Pinto-da-Rocha, 1993: 235, 1995: 80, 1996:22; Gnaspini, 1995: 146, 148, 1996: 432, 433; Machado & Raimundo,2001: 138; Machado et al., 2001: 22; Silva-da-Rocha et al., 2001:99, 101; Machado, 2002: 388, 390, 391; Willemart & Gnaspini:2004a: 16.

Goniosoma undesc. sp. aff. badium. – Gnaspini, 1996: 418.‘undescribed species nearG.badium’.–Gnaspini&Cavalheiro, 1998: 82.Goniosoma sp. 1 aff. badium. – Gnaspini, 1999.Goniosoma sp. – Sessegolo et al., 2001: 184.

Material examined

Types. São Paulo: Japira, ma and fe syntypes of Leitaoius guttulatus(IBSP9), Petruski leg.

Other specimens examined. São Paulo: Botucatu (Fazenda São João),2 ma and 1 fe (marked as ‘allotype’ in the label) (HS524), V.C. Jesus leg.,I/1971; Botucatu, 1 ma (HS854) (no data of collector); Itararé (Gruta daBarreira), 2 ma (MZSP22575), F.H.S. Santos leg., II/2002. Paraná: SãoJerônimoda Serra, 2ma and 2 fe (MZSP16368),M. Segalla&G. Skuk leg.,XII/1997; Tunas do Paraná (Parque Estadual de Campinhos), 1 ma(SMF40503), F.H.S. Santos leg., X/2001; Tunas do Paraná (Gruta dosJesuítas), 5ma and2 fe (MZSP18958), P.Gnaspini leg., III/1992;RioBrancodo Sul (Gruta da Lancinha), 1 ma, 2 fe and 2 im (MZSP16833), P. Gnaspinileg., V/1993; Colombo (Gruta de Bacaetava), 1 ma and 1 fe (MZSP18976),R. Pinto-da-Rocha leg., VI/1999; Morretes (Marumbi), 1 ma and 1 fe(MZSP976), Gofferjé leg., XI/1945; Piraquara (Banhado), 1 ma and 1 fe(MZSP1867), Gofferjé leg., XII/1945;Curitiba (Barigui), 1 ma (MZSP309),R. Lange leg., IV/1942; Curitiba, 1 ma (MZSP1866), Gofferjé leg., VIII/1945; Ponta Grossa (Vila Velha), 4 ma, 1 fe and 4 im (MNRJ 17378),Imaguire leg., 17/VIII/1947; Balsa Nova (Serra de São Luiz), 1 ma and 2 fe(MZSP18135), R. Pinto-da-Rocha et al. leg., III/1999; Balsa Nova(Serra de São Luiz), 1 ma and 1 fe (MZSP18140), R. Pinto-of-Rocha et al.leg., III/1999.

Description

Dorsum. Eye-mound low with 1 pair of weak and hightubercles. Area I with 1 pair of tubercles a little stronger thangranules of area. Area III with 1 pair of medium-sized andslightly backward spines. Angles of posterior margin withtubercles a little larger than other granules of margin; angles offree tergiteswith tubercles a little larger than tubercles of anglesof posterior margin.

Granulation. Carapace: 9–21 large granules. Areas I–III:16–41 large granules. Lateral margin, posterior margin andfree tergites: low density of large granules. Anal operculum:medium density of small granules. Venter: Posterior margin ofstigmatic area and free sternites: medium density of smallgranules. Coxa I: low density of minute granules. Coxa IV:low density of small granules.

Chelicerae. Segment Iwith 2 basal and 1 retrolateral apicalsmall granules; II with medium density of small granules onapical half.

Pedipalps. Trochanter with 2–5 dorsal and 2 or 3 ventralelevations. Femur with 7–9 ventral elevations (standardarmature IiiiIiii), 1 or 2 retrolateral subapical setae anddorsal granulation with medium density of minute granules.Patella: tubercle like a medium-sized subapical ventral–retrolateral granule. Tibia with 1 sub-basal prolateral tuberclelike a large granule, in addition to the standard armature ofsegment.

Leg I. Trochanter with low density of minute granules.Femur with small granules. The other segments practicallysmooth.

Leg II. Trochanter with low density of minute granules.Femur and patella with small granules. Tibia with minutegranules. Metatarsus practically smooth. Row3 and row4 oftibia with sparse minute spines.

Leg III. Femur armed on apical region of row3 with smallspines very close to each other. Trochanter with low density ofminute granules. Femur and patella with small granules. Tibiaand metatarsus with minute granules.

Leg IV. Coxa: prolateral apical apophysis oblique andslightly upward; retrolateral apical apophysis straight,pointed and of similar size of spines of area III of dorsal scute.Trochanter with dorsal–prolateral apical apophysis curvedinward with upward apex and 1 retrolateral apical apophysiscurved backward and a little larger than retrolateral apophysisof coxa. Femur slightly bent inward and upward on apicalthird; row2 armed with sparse spines in all extension and 1–3very large spines on half of row; row3 with small spinesincreasing toward apex; dorsal apical apophyses small,retrolateral apophysis larger. Patella and tibia with medium-sized granulation and metatarsus with small granules.

Tarsal segmentation. 8–11, 17–21, 9–12, 10–13.Penis. Ventral plate very wide; apical margin slightly

concave. Apical group with more ventral seta on medialposition. Glans with dorsal process very small, like a callus.

Colour pattern

Dorsal scute dark-brown with margins sometimes darkgreen; legs I–III greenish-brown; pedipalps and cheliceraegreen; coxa, tibia, metatarsus of leg IV and venter brown;


spines of area III and femur IV black; joint of tibia andmetatarsusof legs with a lighter ring; serous layer on dorsal scute allowinggrayish aspect to animal, but this layer is weak around granules.Dry-mark on apex of coxa, base of trochanter and femur of legIV. Black articularmembranes between coxae and trochanters.Variation: white intrusion in cuticle of lateral grooves of areas.


Dorsal scute: width: 7.79–10.29, length: 8.1–9.36; leg I:38.64–46.28, II: 81.13–98.24, III: 55.34–65.26, IV: 72.14–85.53, femur IV: 19.9–24.7.

Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of free tergites with spines of samesize of spines of area III.

Biology and records

Pinto-da-Rocha (1993, 1995, 1996) [cav.]; Gnaspini (1995 [cit.],1996 [cit.], 1999 [cit.]); Gnaspini and Cavalheiro (1998) [biol.];Machado and Raimundo (2001) [cit.]; Machado et al. (2001)[cit.]; Sessegolo et al. (2001) [cav.]; Silva-da-Rocha et al. (2001)[cav.]; Machado (2002) [cit.]; Hara and Gnaspini (2003) [biol.];Willemart and Gnaspini (2004a) [cit.].

Remarks

The type (MNRJ27609) was examined by R. Stefanini-Jimduring her study, but could not be located presently(see ‘Material’). However, the original description andillustration allowed the precise identification of the species.

The cited allotype was not described together with the typeseries, but we found a remark in a label found in the vial.

Since it was identified by Stefanini-Jim in her unpublished1985M.Sc. thesis as Goniosoma badium (in fact, Stefanini-Jim re-described G. badium based on an immature male of thepresent species), and since it is very abundant in some caves ofParaná, Serracutisoma molle has been referred at as G. aff.badium and as G. n. sp. aff. badium in some papers about thenatural history of goniosomatines (see above).

Serracutisoma pseudovarium, sp. nov.

(Figs 4, 57–60, 151, 152, 208, 209)

Goniosoma cf. varium (misidentification). – Gnaspini & Trajano, 1994:561, 562; Pinto-da-Rocha, 1995: 81; Willemart, 2002: 51.

Goniosoma varium (misidentification). – Gnaspini & Cavalheiro, 1998:82; Gnaspini, 1999.

Material examined

Types. São Paulo: Iporanga (Gruta da Barra Bonita), ma holotype(MZSP22577), M.B. DaSilva et al. leg., XI/1999; Iporanga (Gruta Detrás),1maparatype (SMF), F.H.S. Santos&M.B.DaSilva leg., XI/1999; Iporanga(Caverna dos Caboclos I), 2 ma and 1 fe paratypes (MZSP18903), Vanzoliniet al. leg., XI/1955; Iporanga (Parque Estadual Intervales), 2 ma and 3 feparatypes (MZSP), M.B. DaSilva & F.H.S. Santos leg., XI/1999; Iporanga(Gruta da Barra Bonita), 3 ma and 2 fe paratypes (MZSP16891), P. Gnaspinileg., V/1993; Iporanga (PETAR, Parque Estadual Turístico doAlto Ribeira),1 ma and 1 fe paratypes (ZMHB35333), A.C. Lopes leg., VI/2001.

Description

Dorsum. Eye-mound low with 1 pair of low and roundtubercles. Area III with 1 pair of conic and high tubercles.Angles of posterior margin with tubercles a little smaller thantubercle of area III; angles of free tergites with short spines(2� the size of tubercles of posterior margin of dorsal scute).

Granulation. Carapace: 14–21 (21) large granules. AreasI–III: 43–68 (58) large. Lateral margin, posterior margin andfree tergites: low density of large granules. Anal operculum:low density of small granules. Venter: Posterior margin ofstigmatic area and free sternites: medium density of smallgranules. Coxa I: medium density of minute granules. CoxaIV: low density of medium-sized granules; lateral: low density oflarge granules.

Chelicerae. Segment I with 2 basal granules, being 1 smalldorsal apical and 1 prolateral minute; II with medium density ofminute granules.

Pedipalps. Trochanter with 2–4 (3) dorsal and 2 or 3 (3)ventral elevations. Femur with 5–9 (8) ventral elevations(standard armature IiiIii), 1 retrolateral subapical seta anddorsal granulation with low density of medium-sized granules.Patella: ventral–retrolateral subapical tubercle like a minutegranule. Tibia with 1 low and weak prolateral sub-basaltubercle, in addition to the standard armature of segment.

Leg I. Trochanter with medium density of small granules.Femur with small granules. Patella and tibia with minutegranules. Metatarsus practically smooth.

Leg II. Trochanterwith medium density of small granules.Femur and patella with medium-sized granules. Tibia withminute granules. Metatarsus practically smooth.

Leg III. Femur armed on apical region of row3 withsmall spines very close to each other. Trochanter withmedium density of minute granules. Femur and patellawith large granules. Tibia with small granules. Metatarsuswith minute granules. Row4 of tibia and row3 and row4 ofmetatarsus armed with minute spines.

Leg IV. Prolateral apical apophysis oblique and slightlyupward; retrolateral apical apophysis like a spine of about1/3 size of trochanter. Trochanter with dorsal–prolateralapical apophysis curved inward and with its side fused totrochanter; retrolateral apical apophysis a little larger thanretrolateral apophysis of coxa, spine-shaped and straight.Femur slightly curved inward with large normal granules, alittle larger on row3 and row4; row2 armed with about 4 verysmall spines on subapical region (only 1 larger); dorsal rows alittle disorganised on apical fifth; dorsal apical apophysessmall, retrolateral apophysis larger. Patella and tibia withlarge granules and metatarsus with medium-sized granules.


Penis. Ventral plate very wide; apical margin slightlyconcave. Apical group with the most ventral spine on medialposition. Glans with dorsal process very small, like a callus.

Colour pattern

Animal strongly black; legs I–III dark-brown; pedipalps andchelicerae green; serous layer on dorsal scute allowing grayishaspect to animal, but the layer is weak around granules.


Dry-mark on apex of coxa, base of trochanter and femur of legIV. Black articularmembranes between coxae and trochanters.Variation: granules of dorsal scutemaybeyellow,mainly thoseoflateral and posterior margins.



Female

In addition to dimorphic characters already mentioned,the female differs by: Granules near to lateral groove of areas,on lateral margin, and those of posterior margin and freetergites yellow. Angles of free tergites armed with robustspines, 2� the size of tubercles of angles of posterior marginof dorsal scute.

Biology and records

Gnaspini and Trajano (1994) [cav.]; Pinto-da-Rocha (1995)[cav.]; Gnaspini and Cavalheiro (1998) [biol.]; Gnaspini(1999) [biol.]; Willemart (2002) [biol.].

Remarks

This species often lives near the entrances inside caves of Valedo Ribeira region. It is similar to S. thalassinum. It can bedistinguished from the latter and the remaining species of thegenus by femur IV short and curved inward, with subapicalsmall spines on row2.

Etymology

The species is so named due to its misidentification asGoniosoma varium in lists of cave fauna and natural historypublications.

Serracutisoma thalassinum (Simon), comb. nov.

(Figs 61, 62, 153, 154, 210–212)

Goniosoma thalassinum Simon, 1879: 229.Acutisoma thalassinum. – Roewer, 1913: 277, 278 (comb.), 1923: 505,506, 1930: 387; Mello-Leitão, 1932: 275; Soares & Soares, 1948: 627;Kury, 2003: 117.

Acutisoma patens Roewer, 1930: 388, 389. – Mello-Leitão, 1932: 275,276; Soares & Soares, 1948: 626; Kury, 2003: 116. Syn. nov.

Acutisoma marumbicola H. Soares, 1945: 208, 212. – Soares & Soares,1947a: 64, 68, 1947b: 210, 1948: 625; Kury, 2003: 116. Syn. nov.

Acustisoma marumbicola (typographical error). – H. Soares, 1945: 213.

Material examined

Type. ma and fe syntypes of Acutisoma patens (ROEWER1344)(no data of collection). Paraná: Morretes (Marumbi), holotype ofAcutisoma marumbicola (MHNCI7), R. Lange leg., IV/1944.

Other specimens examined. Paraná: Morretes (Marumbi), 1 ma‘paratype’ (included only in the label, but not published as so) ofAcutisoma marumbicola (MZSP1863), R. Lange leg., IV/1944; Morretes(Marumbi), 1 fe ‘paratype of allotype’ (in the label) of Acutisomamarumbicola (MZSP1800), Imaguire leg., VII/1945; Morretes (ParqueEstadual de Marumbi), 2 ma and 3 fe (MZSP18742), R. Pinto-da-Rocha &

A. Chagas leg., IV/1999; Morretes (Parque Estadual de Marumbi), 1 ma(MZSP18760), R. Pinto-da-Rocha & A. Chagas leg., IV/1999. SantaCatarina: Joinville, 1 ma, 2 fe and 1 im (MNRJ17375), Gofferjé leg.,VIII/1946; Blumenau (Parque Ecológico Spitzkopf), 2 ma and 1 fe(MZSP16581), Bonaldo et al. leg., II/1996; Blumenau (Parque EcológicoSpitzkopf), 2 ma (MZSP18344), R. Pinto-da-Rocha et al. leg., III/1999.

Description

Dorsum. Eye-mound low with 1 pair of very low andround tubercles, close to eyes. Main pair of elevations of areaI indistinct from other large present granules. Area III with1 pair of slightly backward spines. Angles of free tergiteswith spines of similar size of spines of area III, but weaker;angles of posterior margin with a little smaller spines slightlyupward (about 45�).

Granulation. Carapace: 6–20 large granules. Areas I–III:20–36 large granules. Lateral and posterior margins: lowdensity of large granules (variation: granules may be verylarge). Free tergites: low density of large granules. Analoperculum: low density of small granules. Venter: Posteriormargin of stigmatic area, free sternites and coxa I: mediumdensity of small granules.Coxa IV: low density ofmedium-sizedgranules; lateral: low density of large granules.

Chelicerae. Segment I with 2–3 basal and 1 dorsal–retrolateral apical minute granules; II with high density ofminute granules on apical half.

Pedipalps. Trochanter with 2–4 dorsal and 2 ventralelevations. Femur with 4–8 ventral elevations (standardarmature IiiIiii), 1 retrolateral subapical seta and dorsalgranulation with low density of medium-sized granules.Patella: tubercle like a small ventral–retrolateral subapicalgranule. Tibia with 1 prolateral sub-basal tubercle like a largegranule, in addition to the standard armature of segment.

Leg I. Trochanter with medium density of small granules.Femur with small granules. Patella, tibia and metatarsuspractically smooth.

Leg II. Trochanterwith medium density of small granules.Femurwithmedium-sized granules.Patellawith small granules.Tibia and metatarsus practically smooth, except by minutespines on ventral rows (row3 and row4).

Leg III. Femur armed on apical region of row3 withsmall spines very close to each other. Trochanter with lowdensity of medium-sized granules. Femur and patella withmedium-sized granules. Tibia with small granules.Metatarsus with minute granules.

Leg IV. Prolateral apical apophysis almost transversaland slightly upward; retrolateral apical apophysis robust andpointed, 2� the size of spines of angles of free tergites.Trochanter with dorsal–prolateral apical apophysiscurved inward and with its side fused with trochanter;retrolateral apical apophysis 2� the size of retrolateralapophysis of coxa, spine-shaped and straight. Femur straight,armed with large spines in all extension of row2, being apicaland basal smaller; row3 with large granules; dorsal apicalapophyses small, retrolateral apophysis larger. Patella withlarge granules, tibia with medium-sized granules (larger onrow2) and metatarsus with small granules.

Tarsal segmentation. 9–11, 17–20, 9–11, 10–12.


Penis. Ventral plate very wide; apical margin slightlyconcave. Apical group with more ventral seta on medialposition. Glans with dorsal process very small, like a callus.

Colour pattern

Animal strongly black; legs I–III dark-brown; dorsal scutevery dark brown; pedipalps and chelicerae green; joint of tibiaand metatarsus of legs with a lighter ring; serous layer ondorsal scute allowing grayish appearance to animal, but thelayer is weak around granules. Dry-mark on apex of coxa,base of trochanter and femur of legs IV. Black articularmembranes between coxae and trochanters. Variation:granules of dorsal scute may be yellow, mainly those of lateraland posterior margins.



Female

In addition to dimorphic characters already mentioned, thefemale differs by: Femur IVwith short and weak spines on basalthird. Dorsal scute brownwith dark green posterior margins; leg Idark green too. Epidermic pigmentation on dorsal scute as afragmented frame.

Remarks

The type in MNHN was not lent for this study. The originaldescription and later illustrations based on this type(e.g. Roewer 1913) allowed its identification. It can bedistinguished by armature of femur IV and direction ofprolateral apophysis of coxa IV.

B. Soares (1945a) discussed the affinity of two juniorsynonyms of this species, A. patens with A. marumbicola.They are distinguished by the direction of the prolateral apicalapophysis of coxa IV, oblique in A. marumbicola and transversalin A. patens. However, this condition in A. patens was probablycreated by misinterpretation of B. Soares (1945a) by theperspective of the illustration by Roewer (1930) in the originaldescription.

The type locality of A. patens is Caldas-MG, that ofA. marumbicola is Marumbi-PR, and that of A. thalassinumwas not indicated. The localities of recently collectedspecimens are in Paraná or Santa Catarina too. Therefore, weconsider that the type locality ofA. patens is mistaken, a commonfact in old descriptions.

Serracutisoma fritzmuelleri, sp. nov.

(Figs 63–66, 155–157, 213, 214)

Material examined

Types. Santa Catarina: Blumenau (Parque Ecológico Spitzkopf), maholotype and 4 ma and 1 fe paratypes (MZSP14785), Bonaldo et al. leg., 3/II/1996; Blumenau (Parque Ecológico Spitzkopf), 4 ma and 4 fe paratypes(MZSP18321), R. Pinto-da-Rocha et al. leg., 31/III/1999; Blumenau(Parque Ecológico Spitzkopf), 1 ma and 1 fe paratypes (MZSP18329),

R. Pinto-da-Rocha et al. leg., 29/III/1999; Blumenau (Parque EcológicoSpitzkopf), 2 ma paratypes (MZSP18382), R. Pinto-da-Rocha et al. leg.,31/III/1999.

Description

Dorsum. Eye-mound low with 1 pair of straight tubercles.Area I with 1 pair of low central tubercles; III with 1 pairof low spines. Posterior margin, free tergites and analoperculum practically smooth, with sparse minute granules.

Granulation. Carapace: 9–14 (9) small granules. AreasI–III: 27–43 (30) small granules. Lateral margin: high densityof small granules. Venter: Posterior margin of stigmaticarea and free sternites: medium density of minute granules.Coxa I: sparse minute granules, in addition to medial row oftubercles.Coxa IV: low density ofminute granules; lateral: smallgranules.

Chelicerae. Dorsal–retrolateral subapical and largersubapical prolateral granules on segment I. Segment II withaccumulation of dorsal granules in all extension.

Pedipalps. Trochanter with 1–2 (2) dorsal and 1 or 2 (1)ventral elevations. Femur with 5 or 6 (6) ventral elevations(standard armature IiiIi), 1 retrolateral subapical seta anddorsal granulation with low density of small granules. Patella:small subapical ventral–retrolateral granule. Tibia with 1 lowsub-basal prolateral tubercle, in addition to the standardarmature of segment.

Leg I. Trochanter practically smooth. Femur with minutegranules. Patella, tibia and metatarsus practically smooth.

Leg II. Trochanter with low density of minute granules.Femur and patella with minute granules. Other segmentspractically smooth.

Leg III. Femur armed on apical region of row3 withsmall spines very close to each other. Trochanter with lowdensity of small granules. Femur, patella and tibia with smallgranules. Metatarsus practically smooth.

Leg IV. Coxa with some minute granules; prolateralapical apophysis oblique; retrolateral apical apophysis likea granule. Trochanter with high density of small ventralgranules; large apical dorsal–prolateral apophysis curvedinward; retrolateral apical apophysis curved backward, with2� the size of basal apophysis. Femur slightly bent upwardon apical third; row2 armed with 1–4 (3) spines close to eachother, near bend, 1 or 2 (2) very large more apical spines; row3armed with smaller spines which grow from basal fourth to apex;row4with large granules; dorsal rows verydefined;dorsal apicalapophyses very small, very similar to each other. Patella, tibiaand metatarsus with small granules.


Penis. Ventral plate very wide; apical margin slightlyconcave. Apical group with more dorsal seta on subapicalposition. Glans with very small dorsal process, like a callus.

Colour pattern

Entirely black with lighter trochanter and venter of femur ofpedipalps; margins of trochanters of legs reddish-brown. Dry-mark as a frame on dorsal scute with thin line; V-shaped behindeye-mound; on apex of coxa IV and base of femur.




Female

In addition to dimorphic characters already mentioned,the female differs by: Posterior margin of dorsal scute andfree tergites smooth, except by large and robust spine on eachangle, those of free tergites larger. General colour black.Brown on abdominal scute, darker on margins. Epidermicpigmentation of dorsal scute and coxae reticulate and as aframe.

Remarks

This new species can be distinguished by a very typicalretrolateral armature of femur IV with 2 or 3 medial–apicalspines close to each other. It shares the dry-mark as a Ybehind eye-mound with S. proximum.

Etymology

This species is named in honor of Fritz Müller, aGerman–Brazilian naturalist from the 19th century. He lived inBlumenau, type locality of the species, where he helped toerect the city and to develop the evolutionary and Darwinisttheories as well as the general zoology.

Serracutisoma catarina (Machado, Pinto-da-Rocha &Giaretta, 2001), comb. nov.

Goniosoma catarina Machado, Pinto-da-Rocha & Giaretta, 2001. –Machado, 2002: 388–391; Kury, 2003: 117.

G. aff. inermes. – Machado & Raimundo, 2001: 138.

Material examined

Types. Santa Catarina: Santo Amaro da Imperatriz, 1 ma holotypeand 1 ma paratype (MZSP16699), A. A. Giaretta. leg., XII/1997;

Description

Dorsum. Eye-mound with 1 pair of medium-sized anddivergent spines. Area I with 1 pair of medium-sizedtubercles. Area III with 1 pair of weak spines of same heightof spines of eye-mound. Angles of posterior margin unarmedand of free tergites with small and weak tubercles.

Granulation. Carapace: 6–14 (9) small granules. AreasI–III: 22–39 (27) small granules. Lateral margin and freetergites: low density of small granules. Posterior margin:2 small granules. Anal operculum: low density of minutegranules. Venter: Posterior margin of stigmatic area, freesternites, coxa I and coxa IV: low density of minute granules.

Chelicerae. Segment I with 2 apical and 1 basal granules.Segment II with high density of minute granules, from half toapex.

Pedipalps. Trochanter with 2 (2) dorsal and 2 (2) ventralelevations. Femur with 6–8 (8) ventral elevations (standardarmature IiiiIiii), 1 (1) retrolateral subapical seta and dorsalgranulation with low density of minute granules. Patella:

ventral–retrolateral subapical tubercle like a small granule.Tibia with 1 high, but weak, prolateral sub-basal tubercle, inaddition to the standard armature of segment.

Leg I. Trochanter with low density of minute granules.Femurwithminute granules.Other segments practically smooth.

Leg II. Trochanter with low density of minute granules.Femur, patella, tibia and metatarsus practically smooth.Minute sparse spines on row4 and row 5 of tibia and row4 ofmetatarsus.

Leg III. Femur armed on apical region of row3 with sparsespines, with 2 or 3 larger.Trochanterwith low density ofminutegranules. Femur and patella with minute granules. Tibia andmetatarsus practically smooth.

Leg IV. Coxa: prolateral apical apophysis long, almosttransversal, with basal region slightly curved downward;retrolateral apical apophysis like a granule. Trochanterwith dorsal–prolateral apical apophysis upward;retrolateral apical apophysis round and similar to a smallgranule. Femur bent upward after its half; row2 armed withsmall and sparse spines, larger near angle; row3 with spinesincreasing toward the region near angle; dorsal apicalapophyses small, of same size. Tibia with few spines on apexof row2, row3 and row4 (larger on this row).

Tarsal segmentation. 10–11 (11), 16–21 (18/20), 11–12(11/12), 11–14 (13).

Penis. Ventral plate with slightly concave apical margin.Apical group with more ventral seta directed toward the secondseta. Glans without dorsal process.

Colour pattern

General light-brown, with leg IV red; carapace, apophysesof coxa and trochanter IV black. Dry-mark as a frame onabdominal scute and forming a Y behind eye-mound; on apexof coxa, base of trochanter and femur of leg IV.



Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of free tergites with robust spines.

Biology and records

Machado et al. (2001) [biol.]; Machado and Raimundo (2001)[biol.]; Machado (2002) [cit.].

Remarks

It is similar to S. inerme, but it can be distinguished by the lessarmed femur IV, pair of spines on eye-mound, not tubercles as inthe remaining species of Serracutisoma, dry-mark as a frame ondorsal scute, absent in S. inerme, and granules of dorsal scutesmall, not very large.


Serracutisoma inerme (Mello-Leitão), comb. nov.

(Figs 5, 67–70, 158, 159, 215)

Progoniosoma badium (part/misidentification) Roewer, 1913: 265,269.–1923: 500, 502, 1930: 383; Mello-Leitão, 1923: 155, 192,1927: 401, 1932: 259, 261; B. Soares, 1945c: 352. Syn. nov.

Acutisoma inerme Mello-Leitão, 1927: 416. – B. Soares, 1945c: 351;Soares & Soares, 1948: 624 (comb. reestabl.); Soares & Bauab, 1970:131, 133; Machado et al., 2001: 17, 18; Kury, 2003: 116.

Progoniosoma tetrasetaeRoewer, 1930: 383, 385. –Mello-Leitão, 1932:259, 262; B. Soares, 1945a: 192. Syn. nov.

Acutisomella inermis. – Roewer, 1930: 445, 447 (comb.).Acutisomelloides inermis. – Mello-Leitão, 1932: 272 (comb., typespecies).

Goniosoma tetrasetae. – Soares & Soares, 1948: 631 (comb.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 118.

Material examined

Types. Santa Catarina: Blumenau, fe holotype (MNRJ1454).Paraná: Curitiba, type ma of Progoniosoma tetrasetae Roewer, 1930(SMF1341), (no data of collector).

Other specimens examined. No detailed locality (only ‘Brasilien’): ma‘allotype’ of G. badium (Roewer, 1913), (SMF/ROEWER1806) (designatedby Roewer, 1913). Santa Catarina: Rio dos Cedros (Alto Palmeiras), ma‘allotype’ (HS388) (designated by Soares and Bauab, 1970), J. Jim leg., VIII/1969; São Bento do Sul (Ano Bom), 4 ma, 2 fe and 2 im (MZSP18652),R. Pinto-da-Rocha et al. leg., IV/1999; São Bento do Sul (Ano Bom), 2 maand 1 fe (MZSP18651), R. Pinto-da-Rocha et al. leg., IV/1999.

Description

Dorsum. Eye-mound with 1 pair of high and separatetubercles. Area I with 1 pair of tubercles like those of eye-mound.Area IIIwith 1 pair of weak spines. Angles of posteriormargin and free tergites unarmed like the rest of segment(only with granules).

Granulation. Carapace: 2–14 (7) large and very largegranules. Areas I–III: 16–36 (22) large and very largegranules. Lateral margin: medium density of small granules.Posterior margin and free tergites: low density of smallgranules. Anal operculum: low density of minute granules.Venter: Posterior margin of stigmatic area and freesternites and coxa IV: medium density of minute granules.Coxa I: low density of minute granules.

Chelicerae. Segment II with high density of minutegranules on the second half.

Pedipalps. Trochanterwith 2 or 3 (2/3) dorsal and2or 3 (2)ventral elevations. Femur with 5–8 (8) ventral elevations(standard armature IiiIiii), 1/2 (1/2) retrolateral subapicalseta and dorsal granulation with low density of small granules.Patella: ventral–retrolateral subapical tubercle like a smallgranule. Tibia with 1 high, but weak, prolateral sub-basaltubercle, in addition to the standard armature of segment.

Leg I. Trochanter with low density of minute granules.Femur with minute granules. Other segments practicallysmooth.

Leg II. Trochanter with low density of minute granules.Femur, patella, tibia and metatarsus with minute granules.Ventral rows of tibia andmetatarsus armed with minute sparsespines on row4.

Leg III. Femur armed on apical region of row3 withsparse spines, with 2 or 3 larger. Trochanter with low densityof minute granules. Femur and patella with small granules.Tibia with minute granules. Metatarsus smooth.

Leg IV. Coxa: prolateral apical apophysis long, almosttransversal, with basal region slightly curved downward;retrolateral apical apophysis like a granule. Trochanterwith dorsal–prolateral apical apophysis upward;retrolateral apical apophysis curved backward, pointed andhalf the size of dorsal–prolateral apophysis. Femur bent inwardand upward in the apical half; row2 armed with large andsparse spines, increasing from base to angle; row3 withsmaller spines closer to each other, increasing toward theregion near the angle; dorsal apical apophyses small, of samesize. Tibia with few spines on row2, row3 and row4 (largeron this row).

Tarsal segmentation. 9–11 (9/10), 17–19 (17/18), 10–13(10), 9–13 (11/12).

Penis. Ventral plate with slightly concave apical margin.Apical group with more ventral seta placed on medial regionof apical group. Glans without dorsal process.

Colour pattern

Dorsal scute, pedipalps and legs I–III grayish-greenish-brown, with epidermic pigmentation as a frame (yellow colouraround grooves); trochanters of legs I–III, apex of coxa IV andfemur IV dark-red; venter light-brown. Dry-mark only at apexof coxa, base of trochanter and femur of leg IV.


Dorsal scute: width: 8.21–11.02 (8.24), length: 7.69–9.81(7.69); leg I: 42.75–55.23, II: 92.02–120.23, III: 63.68–86.01,IV: 83.74–107.49, femur IV: 24.51–32.92.

Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites with spines likespines of area III, but more robust.

Remarks

Despite the name (= unarmed), it is very armed on legs IIIand IV. However it was described by Mello-Leitão (1927)from a female, which is naturally unarmed. Studying the typefemale, Soares and Bauab (1970) later described a male allotype.

Roewer (1913) designated a male allotype of Goniosomabadium Koch, 1839 since he considered its holotype to be afemale. We have examined this specimen (from SMF) and wehave concluded that it is actually a male Serracutisoma inerme.In addition, we have also examined the original description andillustration of G. badium and we have concluded that it isactually a male from a species of the genus Mitogoniella (seeremarks under Mitogoniella badium). Since the species ofMitogoniella have almost smooth legs, as it is the general caseof females of the other genera of Goniosomatinae, Roewer(1913) probably confounded it with a female specimen.

Due to Roewer’s misinterpretation, all subsequent authorshave misidentified many specimens of Serracutisoma asG. badium. In addition, in her unpublished M.Sc. Thesis,


Stefanini-Jim (1985) proposed to synonymise many differentspecies of Serracutisoma under the name G. badium. Therefore,different species of Goniosomatinae were identified by her asG. badium, as can be seen in cave fauna surveys and somesynonymic lists in the present paper.

Serracutisoma guaricana, sp. nov.

(Figs 71–73, 160, 161, 216)

‘undescribed species nearG.badium’.–Gnaspini&Cavalheiro, 1998: 82.Goniosoma sp. 2 aff. badium. – Gnaspini, 1999.undescribed goniosomatine. – Gnaspini et al., 2004: 34.

Material examined

Types. Paraná: São José dos Pinhais (Usina Hidroelétrica deGuaricana, mata), ma holotype and allotype (MZSP22576), R.L. Pinto leg.,I/2001; São José dos Pinhais (Represa de Guaricana), 2 ma, 5 fe and 1 imparatypes (MZSP18165), A. Kury et al. leg., III/1999; São José dos Pinhais(Usina Hidroelétrica de Guaricana), 1 ma and 1 fe paratypes (MZSP18091),A. Kury et al. leg., III/1999; São José dos Pinhais (Represa de Guaricana),4 ma, 5 fe and 8 im paratypes (MZSP16460), P. Gnaspini leg., III/1993;Piraquara (Banhado), 2 ma and 1 im paratypes (MZSP18674), R. Pinto-da-Rocha &A. Chagas leg., IV/1999;Morretes (Parque Estadual deMarumbi),1 ma and 1 fe paratypes (MZSP18763), R. Pinto-da-Rocha &A. Chagas leg.,IV/1999; Morretes (Pinheirinho, Engenheiro Lange), 2 ma, 4 fe and 3 imparatypes (MNRJ 17388), Imaguire leg., XII/1945; Morretes (EngenheiroLange), 2 fe and 3 im paratypes (MNRJ17389), Gengnagel leg., XII/1947;Morretes (Engenheiro Lange), 2 ma and 2 fe paratypes (MNRJ17376),Gengnagel leg., XII/1947.

Description

Dorsum. Eye-mound with 1 pair of weak tubercles withnear the size of eye-mound. Area I with 1 pair of tubercleslike a medium-sized granule. Area III with 1 pair of weak andstraight upward medium-sized spines. Angles of posteriormargin and of free tergites with small tubercles.

Granulation. Carapace: 0–13 (9) small granules. AreasI–III: 16–34 (34) small granules. Lateral margin, posteriormargin, free tergites and anal operculum: low density ofminute granules. Venter: Posterior margin of stigmatic areaand free sternites: medium density of minute granules. Coxa Iand coxa IV: practically smooth.

Chelicerae. Segment II with medium density of smallgranules.

Pedipalps. Trochanter with 1–3 dorsal and 2 or 3 ventralelevations. Femur with 6 to 9 ventral elevations (standardarmature IiiiIiii), 1 retrolateral subapical seta and dorsalgranulation with low density of minute granules. Patella:ventral–retrolateral subapical tubercle like a small granule.Tibia with 1 low prolateral sub-basal tubercle, in addition tothe standard armature of segment.

Leg I. Trochanter practically smooth, except by tuberclesof standard ornamentation of segment. Other segmentspractically smooth.

Leg II. Trochanter with low density of minute granules.Femur, patella and tibia with minute granules. Ventral rows oftibia armed with sparse minute spines (those of row4 larger).Metatarsus smooth.

Leg III. Femur armed on apical region of row3 withsparse spines, with 2 or 3 larger. Trochanter with low densityof minute granules. Femur, patella and tibia with minutegranules. Metatarsus smooth.

Leg IV. Coxa: prolateral apical apophysis long andalmost transversal; retrolateral apical apophysis like a strongtubercle. Trochanter with dorsal–prolateral apical apophysisupward; 1 retrolateral apical apophysis curved backward,pointed and with half the size of dorsal–prolateral apophysis.Femur bent inward and upward on medial region; row2armed with large and sparse spines on basal half, increasingfrom base to the angle; row3 with smaller spines closer toeach other, increasing toward the apex, in all extension; dorsalapical apophyses small, with similar size. Tibia with a fewspines on row2, row3 and row4 (larger on this row).

Tarsal segmentation. 10–11 (10/11), 17–23 (19), 10–13(11/12), 12–15 (13/14).

Penis. Ventral plate with slightly concave apical margin.Apical group with more ventral seta placed on medial regionof apical group (variation: the more apical seta of group may beabsent). Glans without dorsal process.

Colour pattern

Dorsal scute grayish-brown, with epidermic pigmentationas a frame (yellow around grooves); legs I–III and tibiaand metatarsus IV with similar colour; carapace, spines ofarea III, apophysis of coxa and trochanter IV andpedipalps black; leg IV and trochanters of other legsreddish–brown; venter of body and pedipalp light–brown.Dry-mark only on apex of coxa, base of trochanter and femurof leg IV. Strong pink articular membranes between coxaeand trochanters.


Dorsal scute: width: 7.39–10.70 (9.74), length: 7.34–10.11(9.41); leg I: 43.85–56.06 (51.35), II: 92.61–126.21 (115.45),III: 63.12–84.62 (77.28), IV: 80.88–102.87 (96.18), femurIV: 25.10–27.54 (25.10).

Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of free tergites with spines a littlesmaller than spines of area III.

Biology

Gnaspini and Cavalheiro (1998) [biol.]; Gnaspini (1999) [biol.];Gnaspini et al. (2004) [biol.].

Remarks

This species is very similar toS. inerme, but it canbedistinguishedby the general reddish-brown colour, femur IV with bend placedbefore medial region where the retrolateral spines and smallgranules on dorsal scute end.


Etymology

Name given in apposition as a reference to the Guaricana dam,the type locality, where a large population can be found, andhas been studied from a natural history point of view.

Serracutisoma spelaeum (Mello-Leitão), comb. nov.

(Figs 74–77, 162, 163, 217, 218)

SpelaeosomaspelaeumMello-Leitão, 1933:139.–1935b: 111;B.Soares,1945c: 352.

Goniosoma spelaeum. – Soares & Soares, 1948: 631 (comb.); Muñoz-Cuevas, 1972: 32; Trajano, 1987: 538, 541; Trajano & Gnaspini,1991a: 388, 1991b: 76; Gnaspini & Trajano, 1992: 42, 56, 57;Gnaspini & Trajano, 1994: 562, 564, 566–570; Gnaspini, 1995,1996, 1999; Pinto-da-Rocha, 1995: 81; Pellegatti-Franco &Gnaspini, 1996: 360; Gnaspini & Cavalheiro, 1998; Machado &Oliveira, 1998: 364, 365; Gnaspini & Hoenen, 1999: 155; Sabino& Gnaspini, 1999: 677; Machado et al., 2000: 592, 593, 2001: 17, 18,22;Machado&Raimundo, 2001: 138, 140, 142;Willemart, 2001: 251,2002: 51; Machado, 2002: 388–391; Santos & Gnaspini, 2002;Gnaspini et al., 2003; Hara & Gnaspini, 2003: 263; Hara et al.,2003: 441; Kury, 2003: 118; Willemart & Gnaspini, 2004a: 16,22–24, 2004b: 230, 232; Gnaspini et al., 2004: 31–33, 35.

Acutisoma inerme (misidentification). – Trajano, 1987: 539, 543.Goniosoma sp. – Trajano & Gnaspini, 1991a: 393.

Material examined

Type. São Paulo: Iporanga, ma holotype (MZSP27173) (no data ofcollector).

Other specimens examined. São Paulo: Iporanga (Gruta daCassununga, Braco da Pescaria), 2 ma, 2 fe and 6 im (MZSP19693),Vanzolini et al. leg., XI/1955; Iporanga (Parque Estadual Intervales),3 ma (MZSP22578), F.H.S. Santos leg., XI/1999; Iporanga (Gruta daTemimina), 5 ma and 6 fe (MZSP19694), G. Collet leg., VI/1971;Iporanga (Gruta do Moquem), im’s (MZSP14603), P. Gnaspini leg.,VII/1992; Iporanga (Fazenda Intervales), 1 ma and 5 fe (MZSP14590),P. Gnaspini leg., III/1993; Iporanga (Gruta do Moquem), 1 ma(MZSP14597), P. Gnaspini leg., III/1992.

Description

Dorsum. Dorsal scute with smooth and unarmed generalaspect, but with high density of minute granules, mainly onareas and behind eye-mound. Eye-mound with 1 pair of lowand round tubercles. Area III with 1 pair of very low and weaktubercles, slightly conic. Angles of posterior margin and freetergites smooth.

Granulation. Carapace, areas I–III and lateral margin:high density of minute granules. Posteriormargin, free tergitesand anal operculum: medium density of minute granules.Venter: Posterior margin of stigmatic area and freesternites: high density of minute granules. Coxa I: practicallysmooth. Coxa IV: practically smooth; lateral: minute granules.

Chelicerae. Segment I with minute granules. Segment IIwith high density of minute granules.

Pedipalps. Trochanter with 2–4 dorsal and 2 ventralelevations. Femur with 7–9 ventral elevations (standardarmature IiiiIiii), 1 retrolateral subapical seta and dorsalgranulation with low density of minute granules or practicallysmooth. Patella: tubercle like a small ventral–retrolateral

subapical granule. Tibia with 1 high, but weak, prolateralsub-basal tubercle, in addition to the standard armature ofsegment.

Leg I. Trochanter practically smooth, except by tuberclesof standard armature of segment. Other segments practicallysmooth.

Leg II. Trochanter with low density of minute granules.Femur, patella and tibia practically smooth. Row6 of tibiaarmed with minute sparse spines. Metatarsus smooth, exceptby minute sparse spines on row4.

Leg III. Femur armed on apical region of row3 with sparsespines, with 2 or 3 larger.Trochanterwith low density ofminutegranules. Femur, patella, tibia and metatarsus practicallysmooth.

Leg IV. Coxa: prolateral apical apophysis long,almost transversal; retrolateral apical apophysis like a smallgranule. Trochanter with dorsal–prolateral apical apophysisupward; retrolateral apical apophysis curved backward,pointed and of same size of dorsal–prolateral apophysis.Femur bent inward and upward on medial region; row2armed with small spines on basal half, reaching the angle,where there are 2 or 3 very large spines; row3 with smallerspines, increasing toward the apex, from basal third; dorsalapical apophyses small, of similar size. Tibia with few spineson row2 and on row4 (latter row with spines from basalregion); on apical region with about 3 spines larger.

Tarsal segmentation. 11–12, 20–25, 12–14, 13–16.Penis. Ventral plate with concave apical margin. Apical

group with more ventral seta placed on medial region of apicalgroup. Glans without dorsal process.

Colour pattern

Light-brown, almost yellow with dorsal scute andventer yellow; femur and lateral of coxa IV red; prolateralapophysis of this coxa and carapace (except the eye-mound)black. Dry-mark round only behind eye-mound, on dorsalscute; on apex of coxa, base of trochanter and base of femur ofleg IV. Pink articular membranes between coxae andtrochanters.



Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of posterior margin of dorsal scutewith robust and round tubercles; angles of free tergites withstrong and robust spines.

Biology and records

Trajano (1987) [cav.]; Trajano and Gnaspini (1991a [cav.],1991b [biol.]); Gnaspini and Trajano (1992) [biol.]; Gnaspiniand Trajano (1994) [cav.]; Gnaspini (1995 [biol.], 1996 [biol.],1999 [biol.]); Gnaspini and Hoenen (1999) [cit.]; Pinto-da-Rocha (1995) [cav.]; Pellegatti-Franco and Gnaspini (1996)[biol.]; Gnaspini and Cavalheiro (1998) [biol.]; Machado


and Oliveira (1998) [cit.]; Sabino and Gnaspini (1999) [cit.];Machado et al. (2000) [cit.]; Machado et al. (2001) [cit.];Machado and Raimundo (2001) [cit.]; Willemart (2001 [cit.],2002 [cit.]); Machado (2002) [cit.]; Machado and Oliveira(2002) [cit.]; Santos and Gnaspini (2002) [biol.]; Hara andGnaspini (2003) [cit.]; Hara et al. (2003) [cit.]; Willemart andGnaspini (2004a, 2004b) [cit.]; Gnaspini et al. (2004) [cit.].

Remarks

It is similar to S. guaricana, sp. nov. but it can be distinguishedby the lower armature of femur IV, practically smooth dorsalscute with unarmed area III and yellow colour. It is probably thebest studied Neotropical species of harvestman consideringnatural history.

172 173 174

177

180

183182181

178 179

175 176

Figs 172–183. Male genitalia. 172–174, Heteromitobates alienus, sp. nov., dorsal and lateral views, and detailed apicalview of glans. 175, 176,Mitogoniella indistincta, dorsal and lateral views. 177, 178,Mitogoniella taquara, sp. nov., dorsaland lateral views. 179, 180, Mitogoniella unicornis, sp. nov., dorsal and lateral views. 181–183, Mitogoniella modesta,dorsal and lateral views, and detailed lateral view of glans. Scale = 0.05mm.


Genus Heteromitobates Roewer, reestabl.

(Figs 6, 7, 78–93, 164–174, 219–228, 234, 237)

Heteromitobates Roewer, 1913: 348. – 1923: 535, 1931: 122; Mello-Leitão, 1926: 362, 1932: 384, 1935b: 107; Soares&Soares, 1948: 575;Willemart, 2002: 50, 51, 55; Kury, 2003: 115 (syn. Acutisoma);Willemart & Gnaspini, 2004a: 16, 24.

Acutisomella Roewer, 1930: 349, 445, 447. – Mello-Leitão, 1932: 234,278, 279, 1935b: 110, 1937: 294, 1940: 26; B. Soares, 1944a: 260(syn. Acutisoma). (Type =Acutisoma inscriptum Mello-Leitão, 1922,by original designation.) Syn. nov.

Type species: Mitobates discolor Soerensen, 1884, by monotypy.

Diagnosis

Eye-mound with 1 pair of tubercles either simple or withconical base and knob-like-apex (herein called ‘pawn-shapedtubercle’ as a reference to the shape of pawns in Chess game).Coxa IV with 1 pointed or blunt prolateral apical apophysis,with a small posterior subapical process; retrolateral apicalapophysis small or like a medium-sized spine. Femur IVstraight or curved outward, with pointed or normal granules;dorsal apical apophysis of similar size. Tarsal process minute andtarsal claws pectinate. Penis with ventral plate with generalwidth and length nearly the same and with lateral and apicalmargins concave; apical group with 5 spines in a dorsal row(2nd, from apex to base, displaced to venter) and 3 spines in aventral row; basal group with 4 spines in an oblique row.Glans with ventral process mushroom-shaped withprojections on margins and 1 dorsal sac-shaped process;stylus cylindrical.

Distribution

North littoral of São Paulo and south of Rio de Janeiro, northeastof Serra do Mar at São Paulo and Serra da Bocaina.

Key for the species of Heteromitobates based on males

1. Prolateral apophysis of coxa IV very robust, curved downward and withlength of nearly size of width of dorsal scute; metatarsus IV thickerthan rest of leg; area III with 1 pair of tubercles ................................................................................................H. alienus, sp. nov. (p. 587)

Prolateral apophysis of coxa IV normal; metatarsus IV with diametersmaller than that of tibia IV; area III with 1 pair of spines (Fig. 83)...........................................................................................................2

2. Prolateral apical apophysis of trochanter IV absent; dry-mark behind eye-mound like a triangle; dry-mark present on lateral margin of thecarapace..............................................................................................3

Prolateral apical apophysis of trochanter IV present, smaller than basalapophysis; dry-mark behind eye-mound more complex; dry-markabsent on lateral margins ...................H. harlequin, sp. nov. (p. 586)

3. Retrolateral apophysis of coxa IV like a medium-sized spine; angles ofposterior margin of dorsal scute and free tergites armed.................................................................................H. anarchus, sp. nov. (p. 585)

Retrolateral apophysis of coxa IV reduced; angles of posterior margin ofdorsal scute and free tergites unarmed (Fig. 79)................................4

4. Colour light and green; dry-mark like a full triangle behind eye-mound;more than 16 granules on areas I–III .................................................5

Colour black; dry-mark like an empty triangle behind eye-mound; 8 to 15granules on areas I–III ...................................H. albiscriptus (p. 584)

5. Dorsal scute with 16 to 28 small granules; carapace dark; without lightrings adjacent to joint of legs...............................H. discolor (p. 582)

Dorsal scutewithmoreminutegranules; carapace light; light rings adjacentto joint of tibiae and metatarsi ..........................H. inscriptus (p. 583)

Key for the species of Heteromitobates based on females

Heteromitobates alienus, sp. nov. and H. anarchus, sp. nov. arenot included in the key because no females are known fromthe collections examined.

1. Dry-mark behind eye-mound like a triangle; dry-mark present on lateralmargin of the carapace .......................................................................2

Dry-mark behind eye-mound more complex; dry-mark absent on lateralmargins...............................................H. harlequin, sp. nov. (p. 586)

2. Dry-mark behind eye-mound like a full triangle; more than 16 granuleson areas I–III ......................................................................................3

Dry-mark behind eye-mound like an empty triangle; 8 to 15 granules onareas I–III .......................................................H. albiscriptus (p. 584)

3. Dorsal scute with 16 to 28 small granules; carapace dark; without lightrings adjacent to joint of legs...............................H. discolor (p. 582)

Dorsal scute with more minute granules; carapace light; light ringsadjacent to joint of tibiae and metatarsi ............H. inscriptus (p. 583)

Heteromitobates discolor (Soerensen), comb. reestabl.

(Figs 6, 7, 78, 219, 220)

Mitobates discolor Soerensen, 1884: 611.Heteromitobates discolor. – Roewer, 1913: 348 (comb., type species),1923: 535, Mello-Leitão, 1932: 385; Soares & Soares, 1948: 575;Willemart, 2002: 50, 51, 55; Willemart & Gnaspini, 2004a: 16, 24.

Goniosoma discolor. – Machado, 2002: 389–391.Acutisoma discolor. – Kury, 2003: 115 (comb.).

Material examined

Type. Brasil, fe holotype (ZMUC; damaged, dry and pinned, with label‘Bematistes discolor W.S.’).

Other specimens examined. Rio de Janeiro: Angra dos Reis(Cachoeira do Rio Mambucaba) 2 ma and 1 im (MZSP22583),M.B. DaSilva et al. leg., VII/2001; Parati (trilha do Corisco), 4 ma and1 fe (MZSP22584),M.B. DaSilva et al. leg., VII/2001. São Paulo:Ubatuba,2 ma and 7 fe (MZSP 18955), R.L.G. Raimundo leg., IV/2000; Ubatuba(Picinguaba, trilha do Corisco), 1 ma (MZSP16371), A.C. Marques leg.,XI/1991; Ubatuba (Rio-Santos Km 3), 2 ma (SMF40503), F.H.S. Santos &M.B. DaSilva leg., VIII/2000; Ubatuba (trilha do Instituto Oceanográficoda USP), 1 ma (MZSP35334), F.H.S. Santos leg., I/2001; Ubatuba (trilhado Instituto Oceanográfico da USP), 3 ma and 1 fe (NHMW21181),M.B. DaSilva & F.H.S. Santos leg., VIII/2000; Ubatuba, 3 fe(MZSP27301), P. Fiaschi & J.P.V. Atuí leg., III/2000.

Description

Dorsum. Eye-mound low with 1 pair of ‘pawn-shaped’tubercles. Area I with 1 pair of medium-sized tubercles. AreaIII with 1 pair of very high spines, slightly curved backwardand with base with large diameter. Angles of posterior marginof dorsal scute with small granules and of free tergites with 1medium-sized granule in each one.

Granulation. Carapace: 4–10 small granules.Areas I–III:16–32 small granules. Lateral margin: medium density ofsmall granules. Posterior margin: high density of minutegranules. Free tergites and anal operculum: practically


smooth. Venter: Posterior margin of stigmatic area, freesternites and coxa I: medium density of minute granules.Coxa IV: medium density of small granules; lateral: medium-sized granules.

Chelicerae. Segment I with 2 apical, 2 basal and 1 medialretrolateral granules, and additional 1 or 2 sparse granules.Segment II with large number of small granules.

Pedipalps. Trochanter with 2–4 dorsal and 2 ventralelevations. Femur with standard armature IiIii and 5–8 ventralelevations (these elevations are generally low tubercles) and1 or 2 retrolateral subapical setae. Patella with 1 smallventral–retrolateral subapical tubercle. Tibia with 1 long andthin tubercle and 1 medium-sized granule, both prolateral sub-basal, in addition to the standard armature.

Leg I. Trochanter with low density of minute granules.Femur and patella with minute granules. The other segmentspractically smooth.

Leg II. Trochanterwith medium density of small granules.Femur and patella with medium-sized granules. The othersegments with minute granules.

Leg III. Trochanterwithmediumdensity of small granules.Femur andpatellawith large granules.Tibiawithmedium-sizedgranules.Metatarsus with small granules. Granules from femurto metatarsus are pointed (character more conspicuous on largegranules). Those of apex of row3 and row4 of femur and tibiaare larger.

Leg IV. Prolateral apical apophysis of coxa almosttransversal and robust with a small posterior subapical process;retrolateral apical apophysis small like a tubercle andpointed. Trochanter with 1 robust prolateral sub-basalapophysis and 1 small retrolateral apical apophysis (withsame size of retrolateral apical apophysis of coxa) withassociated granules. Femur slightly curved outward, withlarge, strong and pointed granules, larger on region of ventralrows. Tibia with medium-sized granules, generally large onventral rows and small spines on apical third of row3.Metatarsus with small granules.

Tarsal segmentation. 10–12, 21–25, 11–13, 13–15.Penis. Ventral plate with the region where the basal

group of setae is inserted wider. Apical group with 5thseta much smaller and far from the apical group, placedmedially on ventral plate; all setae, except the medial, slightlyspatulate. Truncus invading the ventral plate medially ondorsal face.

Colour pattern

Dorsal scute brown; carapace, area I and apex of coxaIV black; legs I–III, posterior margin of dorsal scute andcenter of area III dark-greenish-brown; femur IV and venterreddish-brown and the rest of leg IV dark-brown; pedipalpsand chelicerae green. Epidermic pigmentation on dorsalscute as a fragmented frame. Dry-mark as a frame onabdominal scute; like a full triangle behind eye-mound; ina longitudinal narrow stripe anteriorly to eye-mound;punctate on lateral margin of carapace; on apex of coxa,from base to apex of trochanter and base of femur of legIV. Strong pink articular membranes between coxae andtrochanters.



Female

In addition to dimorphic characters already mentioned,the female differs by: Angle of posterior margin of dorsalscute with 1 short spine; angle of free tergites with large andstrong spines (on 2nd tergite the pair of spines is larger, like thatof area III). Colour pattern: area I, area III and posterior marginof abdominal scute washed in green.

Biology and records

Machado (2002) [biol.]; Willemart (2002) [biol.]; Willemartand Gnaspini (2004a) [cit.].

Remarks

Since its original description, this species has been allocated in4 subfamilies. It was described from a female by Soerensen(1884) as Mitobates discolor, the type-genus of Mitobatinae.Roewer (1913) transferred it to Caelopyginae (at that time,Coelopyginae) as the type of the monotypic genusHeteromitobates due to the ‘pectinate tarsal claws of legs IIIand IV’. Cunha-Filho (1955), in an unpublished thesis, allocatedthe species in Heterocranainae. Finally, in his unpublishedPh.D. Thesis, Kury (1991) transferred the species toGoniosomatinae, creating the first record of pectinate tarsalclaws in the subfamily, formalising it in Kury (2003).However, Kury’s transfer was previously published byMachado (2002).

This species is very abundant in the north littoral of São Paulostate and there are many specimens in Brazilian collections(mainly MZSP and MNRJ), in which it was misidentified asAcutisoma inscriptum Mello-Leitão, 1922 since they are verysimilar species.

Heteromitobates inscriptus (Mello-Leitão), comb. nov.

(Figs 79, 80, 164, 165, 221)

Acutisoma inscriptum Mello-Leitão, 1922: 341. – 1923: 159, 191;B. Soares, 1946: 495; Soares & Soares, 1948: 624 (comb. reestabl.);Machado et al., 2001: 17; Kury, 2003: 116.

Acutisomella inscripta. – Roewer, 1930: 445 (comb., type species);Mello-Leitão, 1932: 279.

Material examined

Types. São Paulo: Ilha Bela, fe lectotype and 4 fe paralectotypes(here designated) (MZSP1503), Bicego leg., 1896.

Other specimens examined. São Paulo: Ilha Bela (Riacho da Lage),2 ma (MZSP16858), R.J. Sawaia leg., V/1997; Ilha Bela (Riacho da Lage),1 ma and 1 fe (MZSP16853), R.J. Sawaia leg., IV/1997; São Sebastião(Baraquecaba), 2 ma and 2 fe (MZSP16488), R. Pinto-da-Rocha leg., I/1992;São Sebastião, 2 ma, 1 fe and 2 im (MZSP16374), F.A.G. Mello &N. Carneiro leg., X/1994.


Description

Dorsum. Eye-mound low with 1 pair of ‘pawn-shaped’tubercles. Area I with 1 pair of very small tubercles. Area IIIwith 1 pair of medium-sized and thin spines, slightly backward.Angles of posterior margin of dorsal scute and of free tergiteswith 1 pair of very small tubercles like tubercles of area I.

Granulation. Dorsal scute, free tergites and analoperculum: high density of minute granules. Venter:Posterior margin of stigmatic area and free sternites: highdensity of minute granules. Coxa I: medium density of minutegranules. Coxa IV: high density of small granules.

Chelicerae. Segment I with few sparse minute granules.Segment II with low density of small granules, except on apexwhere it has high density.

Pedipalps. Trochanter with 2–5 (2) dorsal and 2 or 3 (3)ventral elevations. Femur with 5–9 ventral elevations (standardarmature IiiIiii), 2 retrolateral subapical setae and dorsalgranulation with low density of minute granules. Patella with1 high and weak ventral–retrolateral subapical tubercle. Tibiawith 1 prolateral sub-basal tubercle like that of tibia, in additionto the standard armature.

Leg I. Trochanter with low density of minute granules.Femur, patella and tibia with minute granules. Metatarsussmooth.

Leg II. Trochanter with low density of minute granules.Femur with medium-sized granulation. Patella with smallgranules. Tibia with minute granules. Metatarsus smooth.

Leg III. Trochanter with medium density of minutegranules. Spines small increasing on apex of row3 on femurand on tibia. Femurwith large granules.Tibiawith small dorsaland large ventral granules. Metatarsus with minute dorsal andmedium-sized ventral granules.

Leg IV. Prolateral apical apophysis of coxa transversal,robust, with pointed apex, but with a small subapical processand slightly curved downward; retrolateral apical apophysissmall like a very small and thin spine. Trochanter with 1prolateral sub-basal apophysis and 1 retrolateral apicalapophysis like a medium-sized granule. Femur slightlycurved outward; granules very large and pointed; row3armed on apical fifth. Tibia with small spines increasing onapical half of row3, and row4with smaller spines on apical fourth.

Tarsal segmentation. 9–11 (11), 22–28, 10–13 (10/11),12–15 (12).

Penis. Ventral plate with the region where the basal groupof setae is inserted wider. Apical group with 5th seta muchsmaller and far from the apical group, placed near basal group;basal group with all setae, except medial, slightly spatulate.Truncus invading the ventral plate medially on dorsal face.

Colour pattern

Dorsal scute and venter brown; carapace and apex of coxa IVblack; femur IV reddish-brown and the rest of leg IV dark-brown; pedipalps, chelicerae and posterior margin of dorsalscute green; legs I–III greenish-brown; light rings adjacent tojoint of tibia and metatarsus. Epidermic pigmentation ondorsal scute as a fragmented frame. Dry-mark as a frame onabdominal scute; like a full triangle behind eye-mound; in alongitudinal narrow stripe before eye-mound; punctate on lateral

margin of carapace; on apex of coxa, at base and apex oftrochanter (variation: the two trochanter marks may beunited to each other) and base of femur on leg IV; aroundstigmatic area.


Dorsal scute: width: 8.14–9.23 (6.92), length: 7.85–9 (6.80);leg I: 38.29–44.91 (33.77), II: 84.22–102.66, III: 59.19–69.24,IV: 79.65–94.13, femur IV: 21.81–25.21 (20.34).

Remarks

This species is very similar toH. discolor. It can be distinguishedby the lighter colour, higher armature on leg IV and smallergranules in higher quantity on dorsal scute.

Heteromitobates albiscriptus (Mello-Leitão), comb. nov.

(Figs 81, 166, 167, 222–224)

Goniosoma albiscriptumMello-Leitão, 1932: 266, 268, 479. – Soares &Soares, 1948: 628; Soares & Bauab, 1970: 131, 133, 134; Muñoz-Cuevas, 1972: 32; Hara et al., 2003: 441; Hara &Gnaspini, 2003: 258,259, 262, 264, 266, 269;Kury, 2003: 117;Willemart&Gnaspini, 2003:354, 356, 357, 359, 362, 2004a, 2004b.

Goniosoma cf. proximum (misidentification). – Gnaspini & Trajano,1994: 574; Pinto-da-Rocha, 1995: 81; Willemart & Gnaspini,2004a, 2004b.

Material examined

Type. São Paulo: Salesópolis (Estacão Biológica de Boracéia), 1 maneotype (here designated) (MZSP), M.B. DaSilva leg., IV/2001.

Other specimens examined. SãoPaulo:Salesópolis (EstacãoBiológicadeBoracéia), 1ma and1 fe, (MZSP),M.B.DaSilva leg., IV/2001;Salesópolis(Estacão Biológica de Boracéia, Rio dos Pilões), 1 ma (NHMW21182),M.B. DaSilva leg., IV/2001; Salesópolis (Estacão Biológica de Boracéia),1ma and 2 fe (MZSP18073), C.I. Yamamoto leg., 10/II/1997;RibeirãoPires(Gruta da Quarta Divisão), 2 ma and 3 fe (MZSP16370), E. Trajano &P. Gnaspini leg., 30/VIII/1992.

Description

Dorsum. Eye-mound low with 1 pair of ‘pawn-shaped’tubercles. Area I with 1 pair of low tubercles. Area III with 1pair of very high spines, slightly curved backward and withbase with very large diameter. Angles of posterior margin ofdorsal scute smooth and of free tergites with 1 medium-sizedgranule each one.

Granulation. Carapace: 2–12 (5) small granules. AreasI–III: 8–15 (11) small granules. Lateral margin: mediumdensity of small granules. Posterior margin, free tergites andanal operculum practically smooth. Venter: Posterior marginof stigmatic area: high density of minute granules. Freesternites: medium density of minute granules. Coxa I: lowdensity of minute granules. Coxa IV: high density of medium-sized granules; lateral less dense.

Chelicerae. Segment I with 1 larger dorsal–retrolateralapical granule and 1 or 2 sparse granules. Segment II with1 larger medial granule and large number of smaller sparsegranules.

Pedipalps. Trochanter with 1–3 (2) dorsal and 2 or 3 (2)ventral elevations. Femur with standard armature IiiIii and 5–8


(7/8) ventral elevations (these elevations are generally lowtubercles) and 1 or 2 (2) retrolateral subapical setae. Patellawith 1 large ventral–retrolateral subapical granule. Tibia with2 medium-sized prolateral sub-basal granules, in addition to thestandard armature.

Leg I. Trochanter with low density of minute granules.Femur with minute granules. The other segments practicallysmooth.

Leg II. Trochanter with high density of small granules.Femur and patella with small granules. The other segmentspractically smooth.

Leg III. Trochanter with high density of small granules.Femur, patella and tibia with medium-sized granules.Metatarsus with small granules. Granules from femur tometatarsus are pointed (character more conspicuous on largegranules). Row3 and row4 with large granules on femur.

Leg IV. Prolateral apical apophysis of coxa almosttransversal and robust with a small posterior subapical process;retrolateral apical apophysis small like a tubercle andpointed. Trochanter with 1 robust prolateral sub-basalapophysis and 1 small retrolateral apical apophysis (withsame size of retrolateral apical apophysis of coxa) withassociated granules. Femur slightly curved outward, withlarge, strong and pointed granules, larger on region of ventralrows. Tibia with medium-sized granules, generally large onventral rows and small spines on apical third of row3.Metatarsus with medium-sized granules.

Tarsal segmentation. 10–13 (11/12), 24–30 (30), 11–15(15), 13–17 (16/17).

Penis. Ventral plate with the region where the basalgroup of setae is inserted wider. Apical group with5th seta smaller and far from the apical group, placed mediallyon ventral plate; basal group may have 1 additional seta nearthe most dorsal seta; all setae, except the medial, slightlyspatulate. Truncus invading the ventral plate medially ondorsal face.

Colour patterns

General colour black. Dorsal scute and metatarsi of legsvery dark reddish-brown; pedipalps dark green, except coxaand basal half of trochanter, which are yellow; spines of eye-mound yellow too. Epidermic pigmentation in simple frame.Dry-mark as a frame, punctate on lateral margin of carapace,on a longitudinal stripe before eye-mound and an empty trianglebehind eye-mound, on dorsal scute; around stigmatic andgenital areas; on apex of coxa IV, on dorsal face of trochanterIV and base of femur IV.



Female

In addition to dimorphic characters already mentioned,the female differs by: Angle of posterior margin of dorsalscute with 1 short spine; angle of free tergites with large and

strong spines (on 2nd tergite the pair of spines is curved inward).Posterior margin with low density of minute granules andfree tergites with low density of minute granules. Colourpattern: brown with black carapace and area III; posteriormargin of dorsal scute, apical half of femur, patella, tibia andmetatarsus of leg IV, patella, tibia and tarsus of pedipalps,chelicerae and legs I–III dark green; coxa, trochanter andfemur of pedipalps and the pair of main elevations of eye-mound yellow. Epidermic pigmentation of dorsal scute as afragmented frame.

Biology and records

Gnaspini and Trajano (1994) [cav.]; Pinto-da-Rocha (1995)[cav.]; Hara et al. (2003) [cit.]; Hara and Gnaspini (2003)[biol.]; Willemart and Gnaspini (2003) [biol.]; Willemart andGnaspini (2004a, 2004b) [biol.].

Remarks

There is no indication where the type may be deposited and itswhereabouts are unknown. Soares and Bauab (1970) have madean [translation]: ‘addition of other peculiar characters of thisbeautiful species’ and a very detailed description of the colourpattern of the species. That study and the photograph in theoriginal description (Mello-Leitão 1932) allowed theidentification of the species. The type locality is ‘Itatiaya’.However, other specimens have been found in Serra do Mar inSão Paulo state. Since the ranges of distribution of harvestmanspecies are often restricted (see discussion in ‘Biogeography’),we consider the type locality to be wrong.

Heteromitobates anarchus, sp. nov.

(Figs 82–86, 168, 169, 225)

Material examined

Type. São Paulo: Bananal (Parque Nacional da Serra da Bocaina),ma holotype (MZSP25326), P. Fiaschi leg., IV/2000.

Description

Dorsum. Carapace and areas I and III of dorsal scute high.Eye-moundwith 1 pair of ‘pawn-shaped’ tubercles.Area Iwith 1 pairof tubercles like a medium-sized granule in height.Area IIIwith1 pair of very high spines, with large diameter at base and slightlybackward. Angles of posterior margin of dorsal scute withtubercles like a medium-sized granule and of free tergitesarmed with short spines.

Granulation. Carapace: 6 minute granules. Areas I–III:8 minute granules. Lateral margin, posterior margin, freetergites and anal operculum: low density of minute granules.Venter: Posterior margin of stigmatic area and free sternites:high density of minute granules. Coxa I: low density of minutegranules. Coxa IV: medium density of small granules; lateral:low density of medium-sized granules.

Chelicerae. Segment I with 1 medium-sized basal granuleand few smaller granules. Segment II with medium density ofminute granules.

Pedipalps. Trochanter with 3 dorsal and 2 ventralelevations. Femur with 5–6 ventral elevations (standard


armature IiiiI), 2 retrolateral subapical setae and dorsalgranulation with low density of large granules. Patella with 1ventral–retrolateral subapical tubercle like a medium-sizedgranule. Tibia with 1 weak sub-basal tubercle, in addition tothe standard armature.

Leg I. Practically smooth.Leg II. Trochanter with high density of minute granules.

Femur and patella with minute granules. Other segmentspractically smooth.

Leg III. Trochanter with high density of minute granules.Granules of femur and patella increasing from base to apex,mainly on ventral rows. Femur, patella and tibia with smallgranules. Metatarsus with minute granules.

Leg IV. Prolateral apical apophysis of coxa almosttransversal, with apex pointed, but with 1 small subapicalprocess; retrolateral apical apophysis of same size and like aspine. Trochanter with 1 prolateral sub-basal apophysis and1 retrolateral apical apophysis of half the size of prolateralapophysis. Granules of femur, patella, tibia and metatarsuspointed. Femur armed with small and sparse spines on entirerow2; spines on row3 on apical third and on row4 on apical fifth;dorsal apical apophyses small (retrolateral larger and almoststraight). Tibia with small increasing spines on row3 from baseand on row4 on apical third. Metatarsus with medium-sizedgranules, larger on ventral rows.

Tarsal segmentation. 10–11, 25–26, 14–15, 16–17.Penis. Ventral plate with the region where the basal

group of setae is inserted wider. Apical group with 5th setafar from the apical group, placed medially on ventral plate; basalgroup with setae of basal group slightly spatulate. Truncus notinvading the ventral plate.

Colour patterns

General colour black; coxa, trochanter and half of femurof pedipalps and trochanter I yellow. Dry-mark as a frameon abdominal scute; like an empty triangle behind eye-mound;in a wide longitudinal stripe before eye-mound; punctateon lateral margin of carapace; on apex of coxa, from base toapex of trochanter and base of femur on leg IV; around stigmaticarea.


Dorsal scute: width: 7.16, length: 6.57; leg I: 26.55, II: 65.51,III: 44.30, IV: 59.07, femur IV: 16.42.

Remarks

This species can be easily distinguished by the heavilyarmed body and legs, mainly on row2 and row3 of femur IV,on area III of dorsal scute and angles of posterior margin ofdorsal scute and free tergites and by the presence of a retrolateralapical apophysis on coxa IV of same size of prolateral apophysisand a straight and inward retrolateral apical apophysis offemur IV.

Etymology

Latin name derived from the fact that its dry-mark on dorsalscute forms an ‘A’ as the anarchism symbol.

Heteromitobates harlequin, sp. nov.

(Figs 87–90, 170, 171, 226)

Material examined

Types. SãoPaulo:SantoAndré (Paranapiacaba),maholotypeand1maparatype (MZSP22585), M.B. DaSilva & R.H. Willemart leg., IV/1999;Santo André (Paranapiacaba, Gruta do Quarto Patamar), 1 ma and 1 feparatypes (MZSP), F.H.S. Santos & M.B. DaSilva leg., I/2000; Salesópolis(Estacão Biológica de Boracéia), 1 ma paratype (SMF), M.B. DaSilva leg.,III/2001.

Description

Dorsum. Eye-mound low with 1 pair of tubercles like amedium-sized granule; whole carapace very granulated,including the anterior margin. Area I with 1 pair of lowtubercles of same size of a medium-sized granule. Area IIIwith 1 pair of very high spines, slightly backward and withbase with large diameter. Angles of posterior margin ofdorsal scute smooth and of free tergites unarmed, granulessimilar to those of the rest of the segment.

Granulation. Carapace: 11–19 (11) small granules. AreasI–III: 40–61 (47)minute granules.Lateralmargin: high densityof small granules. Posterior margin, free tergites and analoperculum: high density of minute granules. Venter:Posterior margin of stigmatic area and coxa I: high densityofminutegranules.CoxaIV:mediumdensityofminutegranules;lateral: small granules.

Chelicerae. Segment I with many small sparse granules.Segment II with many medium-sized and large granules.

Pedipalps. Trochanter with 3 or 4 (3) dorsal and 2 ventralelevations. Femur with standard armature IiIiIi and 4–7 (4/7)ventral elevations and 1 retrolateral subapical seta. Patellawith 1 ventral–retrolateral subapical tubercle like a smallgranule. Tibia with 1 weak prolateral sub-basal tubercle, inaddition to the standard armature.

Leg I. Trochanter with high density of small granules.Femur and patella with small granules. Tibia with minutegranules. Metatarsus smooth.

Leg II. Trochanter with high density of minute granules.Femurwithmedium-sized granules.Patellawith small granules.Tibia with minute granules. Metatarsus smooth.

Leg III. Trochanter with medium density of minutegranules. Femur with medium-sized granules. Patella andtibia with small granules. Metatarsus with minute granules.

Leg IV. Prolateral apical apophysis of coxa almosttransversal, slightly curved backward, with blunt apex and asmall subapical process; retrolateral apical apophysis smalllike a small spine, smaller than 1/3 of trochanter (highly variablesize; type: small like a granule). Trochanter with 2 prolateralapophyses of almost similar size, being sub-basal a little larger;1 retrolateral apical apophysis like a medium-sized granule.Femur slightly curved outward; row2 armed with very smallspines, alternating with granules in all extension; row3 withsmall spines at the apical half. Tibia with small spines onrow3, except on basal third; row2 with granules higher thanthose of other rows. Metatarsus practically smooth.

Tarsal segmentation. 9, 18–21 (21), 10–11 (10), 10–12 (11).


Penis. Apical group with 5th seta much smaller andfar from the apical group, placed medially on ventral plate;basal group forming an almost transversal row; all setae,except the medial, slightly spatulate. Ventral face of platewith bristles. Truncus invading the ventral plate medially ondorsal face.

Colour pattern

Body medium-brown; trochanter and femur IV slightly red;legs, from half of femur to apex, black; pedipalps and cheliceraegreen; epidermic pigmentation on dorsal scute as a fragmentedframe. Dry-mark as a frame on dorsal scute behind eye-mound;in a wide longitudinal stripe before eye-mound; on apex ofcoxa, at base and apex of trochanter and base of femur of legIV. Dark pink articular membranes between coxae andtrochanters, on live animals.



Remarks

This is the only species of the genus that has a prolateral apicalapophysis on trochanter IV. It has a very conspicuous colourwith a well-developed dry-mark. It is common in granitic caves,in Serra do Mar in São Paulo state.

Etymology

This is a multicoloured species as the character (Harlequin[=Arlecchino, Arlequin]) from traditional European comedyand pantomime. Name given in apposition.

Heteromitobates alienus, sp. nov.

(Figs 91–93, 172–174, 227, 228)

Material examined

Types. São Paulo: São José do Barreiro (Rio Mambucaba),ma holotype (MZSP17714), Pinto-da-Rocha et al. leg., III/1997.

Description

Dorsum. Dorsal scute much narrower than long. Carapacevery granulated, mainly on lateral margins; row with smallgranules on anterior margin. Eye-mound like 2 high domeswith 1 pair of very small tubercles above; entire carapacevery granulated, including the anterior margin. Area I with thepair of main elevations indistinct from other granules. Area IIIwith 1 pair of high tubercles close to each other (smaller thandistance between eyes). Angles of posterior margin and freetergites unarmed, granules similar to those of the rest of thesegment.

Granulation. Carapace: 14 small granules. Areas I–III:52 small granules. Lateral margin: high density of smallgranules. Posterior margin, free tergites and analoperculum: medium density of minute granules. Venter:Posterior margin of stigmatic area and free sternites: high

density of minute granules or practically smooth. Coxa I: lowdensity of minute granules.Coxa IV: medium density of minutegranules; lateral: high density of small granules.

Chelicerae. Segment I with 1 small basal granule and someother smaller sparse granules. Segment II with high density ofminute granules and small from basal third.

Pedipalps. Trochanter with 5 or 7 dorsal and 2 ventralelevations. Femur with 8 ventral elevations (standard armatureIiiIiii), 1 retrolateral subapical seta and dorsal granulationwith low density of minute granules. Patella smooth ventrally.Tibia with 1 weak prolateral sub-basal tubercle, in addition tothe standard armature.

Leg I. Trochanter with high density of minute granules.Femur and patella with minute granules. Other segmentspractically smooth.

Leg II. Trochanter with high density of minute granules.Femur and patella with minute granules. Other segmentspractically smooth.

Leg III. Trochanter with medium density of minutegranules. Femur and patella with small granules. Tibia withminute granules. Metatarsus practically smooth.

Leg IV. Prolateral apical apophysis of coxa very large,same size of length of trochanter, blunt, with a small subapicalprocess, almost transversal, slightly curved backward anddownward; retrolateral apical apophysis small like a thinspine with almost same size of trochanter. Trochanter with1 large prolateral sub-basal apophysis (ending on position ofprolateral apophysis of coxa); 1 retrolateral apical apophysiswith same size of tubercle of area III. Femur short S-shaped,largest outward curvature in basal third; higher granules onrow2 and on apical third of row4; row3 with increasingspines at the apical half. Tibia armed with large spines,increasing from base to apex on row3; dorsal rowsdisorganised; row2 with higher granules than those of otherrows; 1 spine curved backward on apex, close to row3.Metatarsus with astragalus with diameter higher than that oftibia and calcaneus; medium-sized granules and with rowsreasonably disorganised; row3 and row4 with large granules atbase.

Tarsal segmentation. 10, 20, 10–11, 12.Penis. Apical group with 5th seta much smaller and

far from the apical group, placed medially on ventral plate.Ventral face of plate with bristles. Truncus not invading theventral plate.

Colour pattern

Dorsal scute and venter light-brown; legs dark-brown;lateral of coxa and half of femur of leg IV reddish-brown;apophysis of coxa IV black; pedipalps and legs I–III darkgreen. Epidermic pigmentation on dorsal scute as afragmented frame. Dry-mark as a frame on dorsal scute andbehind eye-mound; in a narrow longitudinal stripe before eye-mound; on apex of coxa, at base and apex of trochanter andbase of femur of leg IV.


Dorsal scute: width: 7.50, length: 8.45; leg I: 32.27, II: 60.73,III: 44.40, IV: 58.23, femur IV: 13.88.


Remarks

This species has many autapomorphies so that it is very distinctfrom other species of the genus and of the subfamily in generalaspects. For example, it has a very large prolateral apicalapophysis of coxa IV curved backward, narrower dorsal scute,more robust astragalus of metatarsus IV and femur IV stronglycurved outward (but the basal region is curved inward).Nevertheless, it is placed in the genus Heteromitobatesbecause of the pectinate tarsal claws, reticulate epidermicpigmentation on dorsal scute and only one prolateralapophysis on trochanter IV, among others. Moreover, weshould stress that, first, the genus has a considerably highBremmer support value (see Fig. 233), and, second, thespecies shares several apomorphies with H. harlequin, forminga clade with Bremer support value higher than that of the otherclade within Heteromitobates (see Fig. 233 and Table 2).

Etymology

From Latin ‘alienus’, that means strange or adverse, because ofits very distinct aspect and many autapomorphies.

Genus Mitogoniella Mello-Leitão, reestabl.

(Figs 94–103, 175–183, 229, 230, 234, 238)

MitogoniellaMello-Leitão, 1936: 35.–Piza, 1938: 140;B. Soares, 1944a:260 (syn. Acutisoma).

Type species: Mitogoniella indistincta Mello-Leitão, 1936, by originaldesignation.

Diagnosis

Eye-mound with 1 pair of tubercles very close to each other,with the space between them high nearly the height ofeyes, above or below the eyes level (exception: 1 singlemedial spine on the eye-mound in M. unicornis, sp. nov.).Legs practically smooth, very long and thin. Coxa IV with1 blunt prolateral apical apophysis, transversal and slightlycurved backward, with a small posterior subapical process;retrolateral apical apophysis reduced. Trochanter IV withoutprolateral apical apophysis, with 1 prolateral sub-basalapophysis. Dorsal apical apophysis of femur IV similar toeach other and minute. Tarsal process minute and tarsal clawssmooth (variation: some specimens of M. modesta havepectinate tarsal claws). Penis with ventral plate with widthalmost same as length, with very concave apical margin,lateral margins straight; apical group with 4 or 5 setae in adorsal row and 2 or 3 setae in a ventral row; basal group with4 setae in an oblique row. Ventral face with bristles. Glanswith a large ventral process with an apical lamina with marginwith projections; stylus with a 90� angle on apical third,presence of scales on this angle and cylindrical apex.Dry-mark in dots on the entire lateral margin of dorsalscute. Truncus invading the ventral plate medially on dorsalface.

Distribution

Minas Gerais, Rio de Janeiro (Itatiaia) and southern Bahia.

Remarks

When describing this genus, Mello-Leitão (1936) commentedthat [translation] ‘this genus makes a transition into Mitobatinae’because of the extremely elongate and smooth legs of males,a very conspicuous character.

Key for the species of Mitogoniella based on malesor females

Mitogoniella badia is not included in the key because the typeis lost, and, therefore, we could not characterise it adequately(see below).

1. Eye-mound with 1 single medial spine...M. unicornis, sp. nov. (p. 590)Eye-mound with 1 pair of tubercles ........................................................2

2. Dorsal scute entirely punctate with dry-mark around granules ........................................................................................M. indistincta (p. 588)

Dry-mark on dorsal scute following the grooves of areas (punctate onlyon lateral margins) .............................................................................3

3. Less than 20 generally small granules on areas I–III; space betweentubercles of eye-mound much larger than height of eye..................................................................................................M. modesta (p. 591)

20 or more generally medium-sized granules on areas I–III; spacebetween tubercles of eye-mound same or smaller than height of eyes..............................................................M. taquara, sp. nov. (p. 589)

Mitogoniella indistincta Mello-Leitão, comb. reestabl.

(Figs 94, 175, 176, 229)

Progoniosoma roridum (misidentification). – Bristowe, 1925: 502.Mitogoniella indistincta Mello-Leitão, 1936: 35.Acutisoma indistinctum. – B. Soares, 1945c: 350 (comb.); Soares &Soares, 1948: 624; Kury, 2003: 116.

Goniosoma roridum (misidentification). –Gnaspini & Cavalheiro, 1998:81 (after Bristowe 1925).

Goniosoma indistinctum. – Machado, 2002: 389–391.

Material examined

Types. Minas Gerais: Santa Bárbara, 1 ma lectotype and 1 feparalectotype (here designated) (MNRJ42563), L. Moraes leg.

Other specimens examined. Minas Gerais: Santa Bárbara, 18 ma, 11fe and 4 im (MNRJ), Dr. L. Moraes leg.; Santa Bárbara, 6 ma and 8 fe(MNRJ17545), Nunes, 1960; Brumadinho (Fazenda dos Carvalhos), 4 maand 2 fe (MNRJ17371), D. Pamplona & E.M. Vasconcelos leg., XII/1998;Catas Altas (Gruta do Centenário), 1 ma and 1 fe (MZSP22586), L.S. Hortaleg., IV/1996;CatasAltas (Caraca), 1 fe (MZSP),M.N.Ferreira leg.,X/2001;Alto Caparaó (Parque Nacional do Caparaó), 2 ma and 2 fe (MZSP18956),G. Machado & S. Koehler leg., I/2000; Jaboticatubas (Serra do Cipó, Km126), 1maand1 fe (MNRJ17392), I. Sazimaet al. leg., 1–7/IX/1973;Santanado Riacho, 1 ma (MNRJ17380), I. Sazima leg., VIII/1972.

Description

Dorsum. Eye-mound with 1 pair of tubercles close toeach other; space between tubercles about the height of eyes.Area I with 1 pair of tubercles like a large granule. Area IIIwith 1 pair of medium-sized and weak spines, slightly backward.Angles of posterior margin of dorsal scute and free tergiteswith small tubercles a little larger than other granules ofsegment.

Granulation. Carapace: 7–21 (13)medium-sized granules.Areas I–III: 29–68 (31) medium-sized granules. Lateral


margin, posterior margin, free tergites and anal operculum:lowdensity ofmedium-sizedgranules.Venter:Posteriormarginof stigmatic area and free sternites: high density of smallgranules.Coxa I and coxa IV: mediumdensity of small granules.

Chelicerae. Segment I with 1 pair of basal granules and1 retrolateral subapical granule and 2 prolateral apical granules.Segment II with medium density of small granules on apicalhalf.

Pedipalps. Trochanter with 4 or 5 (4) dorsal and 2 or 3 (2)ventral elevations. Femur with 4 to 9 (8/9) ventral elevations(standard armature IiiiIiii) and 1 retrolateral subapical seta.Patella: ventral–retrolateral subapical tubercle like a granule.Tibia with 1 weak and high prolateral sub-basal tubercle, inaddition to the standard armature of segment.

Leg I. Trochanter with low density of minute granules.Femur, patella and tibia with minute granules. Metatarsuspractically smooth.

Leg II. Trochanter with low density of minute granules.Femur, patella, tibia and metatarsus with minute granules.Row3 of tibia and metatarsus with minute sparse spines.

Leg III. Trochanter with low density of minute granules.Femur, patella, tibia and metatarsus with minute granules.

Leg IV. Coxa with retrolateral apical apophysis likea small granule. Trochanter with retrolateral apicalapophysis like a small granule. Femur almost straight, withminute granules (giving a smooth aspect). Patella and tibiawith minute granules and metatarsus practically smooth.


Penis. Apical group with 5 setae in a dorsal row (morebasal smaller) and 2 in a ventral row; basal group with 4slightly spatulate setae.

Colour pattern

Animal entirely brown; dorsal scute and venter yellowish-brown and carapace and legs I–III darker; leg IV reddish-brown.Dry-mark as a frame and covering all granules on abdominalscute; as a frame behind eye-mound; on apical margin of coxa, atbase and apex of trochanter and base of femur on leg IV.


Dorsal scute:width: 6.55–8.11 (7.08), length: 6.85–8.38 (7.7);leg I: 42.55–61.45, II: 98.7–143.59, III: 62.69–96.23, IV:80.56–116.91, femur IV: 23.94–35.19.

Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites with very robustspines slightly curved inward, of nearly size of spines of area III.

Biology

Bristowe (1925) [biol.]; Gnaspini and Cavalheiro (1998) [cit.];Machado (2002) [biol.].

Remarks

It can be easily distinguished from other species ofMitogoniellaby an interesting autapomorphy: a dry-mark around all granulesof dorsal scute, forming many conspicuous white dots on

the animal. Because of this, it was misidentified as Goniosomaroridum by Bristowe (1925) since they share a similar punctatedry-mark. Our interpretation of Bristowe’s identification as amisidentification is based on the fact that he described theanimals as having very long legs (what is not the case ofG. roridum) and because his record (Ouro Preto, MinasGerais) lies within the range of distribution of Mitogoniellaindistincta.

Mitogoniella taquara, sp. nov.

(Figs 95–98, 177, 178, 230)

Goniosoma sp. – Machado et al., 2003.

Material examined

Types. Minas Gerais: Pains (Gruta São Lourenco), 1 ma paratype(MZSP18953), S.M. Fortes leg., 26/V/2000; Pains (Ressurgência da LocaD’Água), 1ma paratype (SMF), P. Gnaspini et al. leg., IX/2001;Pains (Grutado Isaías), 1 ma and 1 fe paratypes (MZSP27302), P. Gnaspini et al. leg.,IX/2001; Pains (Gruta do Zé da Fazenda), 1 ma paratype (ZMHB35335),P. Gnaspini et al. leg., IX/2001;Doresópolis (caverna da Fazenda Zé GarciaPereira), 1 ma and 3 fe paratypes (MZSP18951), L. Horta leg., 23/IX/1997;Itamonte, 1 ma holotype and 6 ma and 9 fe paratypes (MNRJ5579) (idem);Doresópolis (Gruta Poplotá), 1 ma paratype (MZSP18950), L. Horta leg.,14/IX/1997. Rio de Janeiro: Itatiaia (Parque Nacional de Itatiaia, trilha dosTrês Picos), 1 ma and 1 fe paratypes (NHMW21183), M.B. DaSilva &F.H.S. Santos leg., VIII/2000; Itatiaia (Parque Nacional de Itatiaia), 1 maparatype (MZSP27303), F.H.S. Santos & M.N. Ferreira leg., XII/2001.

Description

Dorsum. Eye-mound with 1 pair of low tubercles close toeach other, being the space between them high, a little lowerthan height of eyes. Area I with 1 pair of low tubercles (2� thesize of granules of areas). Area III with 1 pair of high and thinspines. Angles of posterior margin of dorsal scute and freetergites with 1 small granule each.

Granulation. Carapace: 4–10 (4) small granules. AreasI–III: 21–35 (21) medium-sized granules. Lateral margin:medium density of small granules; higher density of minutegranules on margin of this area. Posterior margin and freetergites: low density of minute granules. Anal operculum:practically smooth. Venter: Posterior margin of stigmaticarea and free sternites: practically smooth. Coxa I: highdensity of small granules. Coxa IV: medium density of minutegranules.

Chelicerae. Segment I practically smooth. Segment II withminute granules on apical half.

Pedipalps. Trochanter with 1 or 2 (2) dorsal and 2 ventralelevations. Femur with 6–9 (7/9) ventral elevations (standardarmature IiiIiIii; these elevations are short and with roundapex), 1 or 2 (1/2) retrolateral subapical setae and dorsalgranulation with low density of minute granules. Patella:small ventral–retrolateral subapical granule. Tibia with 2 shortprolateral sub-basal spines, in addition to the standard armatureof segment.

Leg I. Trochanter with low density of minute granules.Femur, patella and tibia with minute granules. Metatarsuspractically smooth.


Leg II. Trochanter with medium density of minutegranules. Femur and patella with minute granules. Othersegments practically smooth.

Leg III. Trochanter with high density of small granules.Femur, patella and tibia with small granules.Metatarsus withminute granules.

Leg IV. Coxa with retrolateral apical apophysis roundand like a granule (may be from minute to large; being minutein type). Trochanter with retrolateral apical apophysis roundand like a large granule. Femur very long, almost straight(slightly curved outward), with small and pointed granules.Tibia with same granulation as that of femur. Metatarsuswith minute granules.


Penis. Apical groupwith 5 setae in a dorsal row (more basalsmaller) and 2 in a ventral row; basal group with 4 slightlyspatulate spines organised in an oblique row.

Colour pattern

Dorsal scute light-brown, pedipalps and legs I slightly green,with legs III and IV darker; ventral face very light brown,almost yellow. Dry-mark as a frame on dorsal scute and in anarrow medial longitudinal stripe behind eye-mound ondorsal scute; on apical margin of coxa, at base and apex oftrochanter and base of femur, dorsal on leg IV; in addition,from other specimens: in a narrow medial longitudinal stripebefore eye-mound, in dots on whole lateral margin and joiningthe dry-mark of base and apex of trochanter IV. Light pinkarticular membranes between coxae and trochanters of legIV. Legs II–IV red on live animal.



Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of posterior margin of dorsal scutewith small granules; angles of tergite I with 1 very large androbust spine, II and III with thinner, high spines curved inward.Dorsal scute with epidermic pigmentation stronger, allowingdarker aspect.

Biology

Machado et al. (2003) [biol.].

Remarks

It can be distinguished from other species of the genus bymedium-sized granules in smaller quantity on dorsal scute andheight of fusion of base of spines of eye-mound lower than theheight of eyes. This species also has a higher ‘legs length/body length’ ratio. It is very common in its range ofdistribution, mainly in caves, and, although so far undescribed,this species is also frequently found in the museum collectionsexamined.

Etymology

From Brazilian Tupi native language, ‘taquara’ is the name ofa thin bamboo, in reference to the very long and thin legs. Namegiven in apposition.

Mitogoniella unicornis, sp. nov.

(Figs 99–102, 179, 180)

Goniosoma n. sp. – Machado, 2002: 389–391.

Material examined

Types. Bahia: Itororó (Serra do Oricana), ma holotype and 2 ma and5 fe paratypes (MZSP18957), G. Machado leg., III/2000.

Description

Dorsum. Eye-mound with 1 single spine on center, large andcurved forward.Area Iwith 1 pair of tubercles likemedium-sizedgranules. Area III with 1 pair of large and slightly divergentspines (same size of spine of eye-mound). Angles of posteriormargin of dorsal scute and free tergites unarmed, but the rest ofthe segments have granules.

Granulation. Carapace: 0–4 (4) small granules. AreasI–III: 28–39 (30) small granules. Lateral margin: mediumdensity of small granules; higher density of minute granules atthe margins. Posterior margin, free tergites and analoperculum: high density of small granules. Venter: Posteriormargin of stigmatic area and free sternites: high density ofminute granules. Coxa I: high density of small granules. CoxaIV: high density of minute granules; lateral: small granules.

Chelicerae. Segment I with 2 minute basal, 2 minute apicaland 1 larger prolateral apical granules. Segment II with highdensity of small granules.

Pedipalps. Trochanter with 1 or 2 (2) dorsal and 2 ventralelevations. Femur with 7–8 (7/8) ventral elevations (standardarmature IiIiIiii), 1 or 2 (1/2) retrolateral subapical setae anddorsal granulation with low density of minute granules. Patella:ventral–retrolateral subapical tubercle like a granule. Tibia with1 weak prolateral sub-basal tubercle, in addition to the standardarmature of segment.

Leg I. Trochanter with high density of small granules.Femur, patella and tibia with minute granules. Metatarsuspractically smooth.

Leg II. Trochanter with high density of minute granules.Femurwith minute granules. Patellawith small granules. Tibiawith minute granules and minute sparse spines on row3.Metatarsus smooth except by minute sparse spines on row3.

Leg III. Trochanter with high density of minute granules.Femur, patella and tibia with minute granules. Metatarsuspractically smooth.

Leg IV. Coxa with retrolateral apical apophysis like asmall granule. Trochanter with retrolateral apical apophysislike a large granule. Femur very long, almost straight, withminute granules. Tibia with same granulation as that of femur.Metatarsus with minute granules.

Tarsal segmentation. 9–11 (11), 18–22 (19/20), 11–13(13), 12–14 (14).

Penis. Apical group with 4 setae in a dorsal row (morebasal smaller and far from the others) and 3 in a ventral row;


basal group with 4 slightly spatulate setae organised in anoblique row.

Colour pattern

General colour dark-brown; venter and spine of eye-moundyellow; pedipalps slightly green. Dry–mark as a frame onabdominal scute, in a narrow longitudinal medial stripe behindand before eye-mound on dorsal scute; on apical margin of coxa,at base and apex of trochanter and base of femur of leg IV.


Dorsal scute: width: 7.35–8.37 (8.01), length: 7.51–8.55(8.25); leg I: 53.70–66.23 (61.64), II: 141.6–181.4 (181.4),III: 80.39–93.74 (91.34), IV: 101.12–114.86 (114.10), femurIV: 29.61–34.89 (34.74).

Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites armed with largeand robust spines; angles of posterior margin of dorsal scutewith 1 small spine.

Biology

Machado (2002) [biol.].

Remarks

This species has an interesting autapomorphy: the spines ofthe eye-mound are entirely fused, what makes it the onlyspecies of the subfamily with a single spine on the eye-mound.The distributions of this new species and Mitogoniella modestaare separated from those of other species of the subfamily, whatmay indicate the lack of collections in the remaining area.

Etymology

From Latin, ‘unicornis’ means ‘one horn’. This is the onlyspecies in the whole subfamily with a single spine on the eye-mound.

Mitogoniella modesta (Perty), comb. nov.

(Figs 103, 182, 183)

Goniosoma modestum Perty, 1833: 202. – Koch, 1839: 119; Gervais,1844: 108; Simon, 1879: 233; Soares &Soares, 1948: 630 (comb.reestabl.); Muñoz-Cuevas, 1972: 32; Kury, 2003: 117.

Progoniosoma modestum. –Roewer, 1913: 275 (comb.), 1923: 500, 503,1930:383;Mello-Leitão, 1923:157,1932:259,264;Moritz, 1971:204.

Material examined

Types. Bahia: ma lectotype and 1 ma paralectotype (ZMHB930) (heredesignated), Freir leg.

Other specimens examined. Bahia: Santa Luzia (Pedra do Sino), 3 maand 1 fe (MZSP18902), B.S. Santos leg., X/1997.

Description

Dorsum. Eye-mound with the space between the pair oftubercles higher than height of eyes (variation: this height varylargely among specimens); the tubercles are weak and close toeach other. Area I with 1 pair of tubercles like medium-

sized granules. Area III with 1 pair of very large, divergent,and slightly pointed backward spines. Angles of posteriormargin of dorsal scute with tubercles like large granules;angles of free tergites with tubercles 2� larger than those ofangles of posterior margin.

Granulation. Carapace: smooth, in addition to thetubercles of eye-mound. Areas I–III: 10–17 (12) smallgranules. Lateral margin: high density of small granules.Posterior margin and free tergites: medium density of smallgranules (variation: these granules may be medium-sized). Analoperculum: medium density of minute granules. Venter:Posterior margin of stigmatic area and free sternites: highdensity of small granules. Coxa I and coxa IV: medium densityof small granules.

Chelicerae. Segment I with basal granules and 1 prolateralapical granule. Segment II withmediumdensity ofmedium-sizedgranules on apical 2/3.

Pedipalps. Trochanter with 1 or 2 (2) dorsal and 2 or 3 (3)ventral elevations. Femur with 7–9 (7/8) ventral elevations(standard armature Iiiiiiii) and 1 or 2 retrolateral subapicalsetae. Patella: low and weak ventral–retrolateral subapicaltubercle. Tibia with 1 prolateral sub-basal tubercle like that ofpatella, in addition to the standard armature of segment.

Leg I. Trochanter with medium density of small granules.Femurwithminute granules.Other segments practically smooth.

Leg II. Trochanter with medium density of minutegranules. Other segments practically smooth.

Leg III. Trochanter with low density of minute granules.Femur with minute granules. Other segments practicallysmooth.

Leg IV. Coxawith retrolateral apical apophysis like a smallgranule. Trochanter with retrolateral apical apophysis like asmall granule. Femur almost straight, with minute granules(aspect smooth). Patella and tibia with minute granules andmetatarsus practically smooth. Tarsal claws smooth orpectinate.


Penis. Apical group with 4 setae in a dorsal row (morebasal smaller and far from the others) and 2 in a ventral row;basal group with 5 slightly spatulate setae organised in anoblique row.

Colour pattern

Colouringverydamaged.Dry-mark as a frameonabdominalscute, in a narrow medial longitudinal stripe behind and beforeeye-mound on dorsal scute; on apical margin of coxa, at base andapex of trochanter and base of femur on leg IV.

Female

In addition to dimorphic characters already mentioned,the female differs by: Posterior margin and free tergiteswith medium-sized granules. Angles of posterior marginof dorsal scute with short spines; angles of free tergiteswith larger and more robust spines (half the size of spines ofarea III).



The measurements of the type specimen are given betweenparentheses but they were not included in species variationbecause it is much smaller than the other specimens. Dorsalscute: width: 6.1–6.61 (5.47), length: 6.61–6.64 (5.6); leg I:56.15–57.19 (36.55), II: 135.7–157.3 (78.85), III: 83.8–84.7(59), IV: 105.9 (74.1), femur IV: 32.52–32.54 (21.8).

Remarks

The species ofMitogoniella are very similar to each other. Theycan be distinguished only by the granulation and ornamentationof dorsal scute. For example, M. modesta can be distinguishedfrom other species by the space between the pair of tuberclesmuch higher than the height of eyes.

Mitogoniella badia (Koch), comb. nov.,species inquirenda

Goniosoma badium Koch, 1839: 65. – Simon, 1879: 233; Soares &Soares, 1948: 628 (comb. reestabl.); Muñoz-Cuevas, 1972: 32; Kury,2003: 117.

Progoniosoma badium. – Roewer, 1913: 269 (comb.), 1923: 502, 1930:383;Mello-Leitão, 1923: 155, 1927: 401, 1932: 261; B. Soares, 1945c:352.

Remarks

Although the type of Goniosoma badium Koch, 1839 is lost, wehave identified it as a species of the genus Mitogoniella but wecan not recognise it as a synonym of some species or as a fifthspecies in the genus. Therefore, it is here considered a ‘speciesinquirenda’.

Roewer (1913) designated a male allotype of Goniosomabadium Koch, 1839 since he considered its holotype to be afemale. We have examined this specimen (from SMF) and wehave concluded that it is actually a male Serracutisoma inerme(see remarks under Serracutisoma inerme). In addition, wehave also examined the original description and illustrationof G. badium and we have concluded that it is actually amale from a species of the genus Mitogoniella. Since thespecies of Mitogoniella have almost smooth legs, as it is thegeneral case with females of the other genera of Goniosomatinae,Roewer (1913) probably confounded it with a female specimen.

Genus Acutisoma Roewer

(Figs 3, 104–111, 184–189, 231, 232, 234, 237)

Acutisoma Roewer, 1913: 170, 276–278, 280. – 1923: 465, 505, 506,1930: 348, 387, 388; Mello-Leitão, 1922: 341–343, 1923: 158–161,191, 192, 1926: 32, 1927: 389, 416, 1932: 234, 274–278, 1935b: 110,1937: 294; Bristowe, 1925: 502; Piza, 1942: 404; B. Soares, 1944a:260, 262, 263, 1944b: 280, 1944c: 311, 1944d: 221, 1945a: 192, 1945b:231, 232, 1945c: 350, 351, 1946: 495, 496; Soares & Soares, 1945:252–254, 1947a: 63, 64, 67, 68, 1947b: 209, 210, 214, 1947c: 249, 250,1948: 621–627; H. Soares, 1945: 208; Trajano, 1987: 539, 542–544,547, 548;Ramires&Giaretta, 1994;Machado et al., 2001: 17–18;Hara&Gnaspini, 2003: 258, 259, 262, 264, 266, 269, 273; Kury, 2003: 115.

Itatiaya Roewer, 1928: 125. (Non Itatiaya Mello-Leitão, 1915; pre-occupied name.)

Leitaoius Roewer, 1930: 443, 445 (replacement for ItatiayaRoewer, 1928). – Mello-Leitão, 1932: 232, 255, 1934: 412, 414,1935a: 402, 404, 1935b: 111, 1940: 25; Piza, 1938: 139; B. Soares,1944a: 260, 262 (syn. Acutisoma). (Type = Itatiaya hamata Roewer,1928, by monotypy.)

Type species: Goniosoma acutangulum Simon, 1879, by originaldesignation.

184 185 186

187 188 189

Figs 184–189. Male genitalia. 184, 185, Acutisoma coriaceum, sp. nov., dorsal and lateral views. 186, 187, Acutisomahamatum, dorsal and lateral views. 188, 189, Acutisoma longipes, dorsal and lateral views. Scale = 0.05mm.


DiagnosisEye-mound with 1 pair of tubercles very close to each otherwith the space between them high, nearly the height of eyes,above or below the eyes level. Coxa IV with a pointed or bluntprolateral apical apophysis with a small posterior subapicalprocess; retrolateral apical apophysis medium-sized,crescentlike (prolateral curvature) or cylindrical withpointed apex. Trochanter IV with 1 retrolateral basal

apophysis similar to prolateral apophysis, but theretrolateral is slightly curved forward. Femur IV withdorsal–retrolateral apical apophysis straight and inwardor curved backward. Tarsal process minute and tarsal clawssmooth. Penis with ventral plate with concave apical marginand convex lateral margin, with wide base and medial regionsand narrower apical region. Setae slightly spatulate, except themedial one; apical group with 4 setae on dorsal row (the more

190 191 192

193

196

195194

Figs 190–196. Photographs. 190, Goniosoma varium, female (observe dry-mark in mask – in abdominal scute andcarapace). 191, Goniosoma capixaba, sp. nov., male. 192, 193, Goniosoma venustum, female, and example of epidermicpigmentationof the type ‘homogeneousmask’. 194, 195,Goniosomaroridum,male and female (194: photo byG.Machado).196, Goniosoma macracanthum, male.


basal very far from the others and minute) and 2 setae on ventralrow; basal group with 4 setae placed in oblique and disorganisedrow. Glans with stylus with a 90� angle pointed to venter onapical region and with cylindrical apex; ventral process with anapical lamina inserted by base and apical margin with manyprojections; without dorsal process. Dry-mark as dots on theentire lateral margin of dorsal scute. Truncus invading theventral plate medially on dorsal face.

Distribution

São Paulo and Serra da Mantiqueira (in São Paulo, Minas Geraisand Rio de Janeiro states).

Key for the species of Acutisoma based on malesor females

No females of Acutisoma coriaceum, sp. nov. are knownfrom the collections examined. Acutisoma acutangulum wasnot included in the key because we were not able to study thetype and, therefore, we can not assure that it is not a synonym.

1. Retrolateral apophysis of coxa IV cylindrical and with conical apex;dorsal–retrolateral apical apophysis of femur IV curved backward;minute granules on dorsal scute........A. coriaceum, sp. nov. (p. 594)

Retrolateral apophysis of coxa IV with retrolateral size curved (crescent-like) (Fig. 107); dorsal–retrolateral apical apophysis of femur IV largeand straight and inward (Fig. 111); small or medium-sized granuleson dorsal scute ...................................................................................2

2. Colour reddish-black, very homogeneous; dorsal scute with smallgranules (Fig. 2) .................................................A. hamatum (p. 597)

Colour on dorsal scute yellowish-brown or may be much darker;pedipalps, chelicerae and legs I–III dark green; dorsal scute, generally,with medium-sized granules (Fig. 3) ...................A. longipes (p. 601)

Acutisoma acutangulum (Simon)

Goniosoma acutangulum Simon, 1879: 230.Acutisoma acutangulum. – Roewer, 1913: 277 (comb., type species),1923: 505, 1930: 387; Mello-Leitão, 1923: 191, 1932: 275; B. Soares,1945c: 350; Soares & Soares, 1948: 627; Kury, 2003: 115.

Remarks

The type of this species is a female from the Paris Museum(MNHN). We did not have access to it. However, Profs.A. Muñoz-Cuevas and W. Lourenco kindly sent usphotographs of the animal, which allowed us to identify it as aspecies belonging to the same clade as A. hamatum, A. longipesand A. coriaceum, sp. nov. Based on its general aspects, weconsider it to be a distinct species, but we just had photographsof a dry and pinned female to base our analysis. Several malecharacters, important for a precise species identification,were missing, which also did not allow us to include thisspecies in the cladistic analysis.

Acutisoma coriaceum, sp. nov.

(Figs 104–106, 184, 185)

Material examined

Types. São Paulo: Salesópolis (Estacão Biológica de Boracéia),ma holotype (MZSP16485), L. Fontes & P. Terra leg., 24–30/I/1979;

Salesópolis (Estacão Biológica de Boracéia), 1 ma paratype (MZSP),P. Gnaspini leg., V/2001.

Description

Dorsum. General aspect of dorsal scute smooth. Eye-moundwith slightly divergent tubercles close to each other, withround apex. Area I with 1 pair of small central granules. AreaIII with 1 pair of medium-sized spines (same height ofelevations of eye-mound), slightly backward and with its basewith large diameter. Angles of posterior margin of dorsal scuteand free tergites with a low and round tubercle.

Granulation. Carapace: 2–8 (2) small granules. AreasI–III: 18–25 (18) minute granules. Lateral margin: mediumdensity of small granules (larger on margin adjacent to area IIand higher density adjacent to carapace). Posterior margin andfree tergites: medium density of minute granules. Analoperculum: practically smooth. Venter: Stigmatic and genitalareas with some sparse minute granules. Posterior margin ofstigmatic area: high density of medium-sized granules. Freesternites: medium density of medium-sized granules. Coxa I:low density of small granules. Coxa IV: medium density ofsmall granules.

Chelicerae. Segment I with 2 dorsal–retrolateral, 1 apical,1 basal, and 1 dorsal–prolateral basal granules. Segment II withstronger granules from apical half.

Pedipalps. Trochanter with 2–3 (3) dorsal and 3 ventralelevations. Femur with 8–9 (9) ventral elevations (standardarmature Iiiiiii), 1 or 2 (1/2) retrolateral subapical setae anddorsal granulation with medium density of medium-sizedgranules. Patella: small ventral–retrolateral subapical granule.Tibia with 2 ventral–prolateral sub-basal tubercles, in addition tothe standard armature.

Leg I. Trochanter with low density of small granules.Femur and patella: minute granules. Other segmentspractically smooth.

Leg II. Trochanter with low density of small granules.Femur and patella with small granules. Other segmentspractically smooth.

Leg III. Trochanterwithmediumdensity of small granules.Femur, patella and tibia with large granules. Metatarsuswith small granules.

Leg IV. Retrolateral apical apophysis of coxa cylindrical,with slightly conical apical region, slightly curved outwardand with 2/3 size of prolateral apophysis. Latter almosttransversal, with blunt apex, slightly curved downward, andwith a small posterior subapical process. Trochanter with 1prolateral basal apophysis, 1 retrolateral basal apophysisa little larger than prolateral apophysis and 1 retrolateral apicalapophysis very small; high density of minute dorsal and smallventral granules. Femur with large and strong granules withround apex, larger on row1 and row2, gradually decreasingtoward apex; presence of 1 additional row of minute granulesbetween these rows; dorsal apical apophyses small, very similarto each other. Tibiawith more pointed granules, mainly on row1and row2. Metatarsus with pointed granules, larger on row2separated by smaller, decreasing toward apex.

Tarsal segmentation. 9–11 (9/10), 19–22 (19/20), 11–12(11/12), 13–15 (13/14).


Penis. Same description as the diagnosis for the genusAcutisoma.

Colour pattern

Entirely black with reticulate pedipalps and chelicerae withdark-reddish-brown background, like tibia and metatarsi oflegs and margin of carapace close to opening of the defensivegland. Punctations of dry-mark of lateral margin of dorsalscute more numerous on carapace; on apex of coxa, at base

and apex of trochanter and at base of femur. Black articularmembranes between coxae and trochanters.



197

198

199 200

203

204

202

201

Figs 197–204. Photographs. 197, Goniosoma dentipes, male. 198, Goniosoma unicolor, male (photo by F.H.S. Santos).199,Goniosoma calcar, male. 200,Goniosoma carum, male. 201, 202,Pyatan insperatum, sp. nov., male and female (photoby R. Pinto-da-Rocha). 203, 204, Serracutisoma proximum, male and female.


Remarks

This species from Serra do Mar of São Paulo state is similar toA. longipes. It can be distinguished by shorter and straight femurIV, strong black colour without dry-mark on grooves of areasand central region of carapace, cylindrical and slightly curvedoutward retrolateral apical apophysis of coxa IV, and minute

granules on the dorsal scute, what gives its leathery aspect to theintegument.

Etymology

From Latin, this name is related to the leathery aspect of theintegument of the dorsal scute.

206 207 210

211

208

205

209 212

Figs 205–212. Photographs. 205, Serracutisoma banhadoae, male. 206, 207, Serracutisoma molle, male (photo byF.H.S. Santos) and example of serous layer reduced around granules of dorsal scute. 208, 209, Serracutisoma pseudovarium,sp. nov., male (photo by G. Machado) and example of epidermic pigmentation of the type homogeneous pigmentation.210–212, Serracutisoma thalassinum, male and female (photos by R. Pinto-da-Rocha), and example of epidermicpigmentation of the type ‘fragmented mask’.


Acutisoma hamatum (Roewer)

(Figs 107, 108, 186, 187, 231)

Itatiaya hamata Roewer, 1928: 125. (Itatiaya Roewer, 1928 = pre-occupied name by Itatiaya Mello-Leitão, 1915.)

Leitaoius hamatus. –Roewer, 1930: 443;Mello-Leitão, 1932: 255, 1934:414, 1940: 25.

Acutisoma hamatum. – B. Soares, 1945c: 350 (comb.); Soares & Soares,1948: 624; Kury, 2003: 115.

Material examined

Types. Rio de Janeiro: Itatiaia (Itatiaya, 1170m), 1 ma holotype(ISNB), P. Brien leg., X/1922.

Other specimens examined. Minas Gerais: Rio Preto (Serra SãoLourenco), 1 ma and 2 fe (MZSP15641), R.S. Bérnils leg., VI/1996;Rio Preto (Ribeirão do Funil), 1 ma and 1 fe (MZSP15605), R.S. Bérnilsleg., V/1996. Rio de Janeiro: Itatiaia (Parque Nacional de Itatiaia,Rio Campo Belo, Piscina da Maromba), 3 ma and 2 fe (MZSP),

213 214 215

217

218

216

Figs 213–218. Photographs. 213, 214, Serracutisoma fritzmuelleri, sp. nov., male and female. 215, Serracutisomainerme, male. 216, Serracutisoma guaricana, sp. nov., male (photo by F.H.S. Santos). 217, 218, Serracutisomaspelaeum, male.


F.H.S. Santos & M.B. DaSilva leg., VIII/2000; Itatiaia (ParqueNacional de Itatiaia), 2 ma and 3 fe (MZSP16375), R. Pinto-da-Rochaleg., IV/1991.

Description

Dorsum. Eye-mound round; tubercles weak and high.Dorsal scute with velvet like aspect. Area I with 1 pair oftubercles, a little distinct from other granules of area. Area IIIwith 1 pair of spines with large diameter at base and slightly

backward. Angles of posterior margin of dorsalscute and free tergites with tubercles like medium-sizedgranules.

Granulation. Carapace: 7–21 (21) small granules. AreasI–III: 17–48 (48) small granules. Lateral margin: lowdensity of small granules; marginal rim: high density.Posterior margin: medium density of small granules. Freetergites: low density of small granules. Anal operculum:low density of minute granules. Venter: Very granulated.Posterior margin of stigmatic area and free sternites: high

219 220 221

225223

222

224

Figs 219–225. Photographs. 219, 220, Heteromitobates discolor, male and female (photos by G. Machado).221, Heteromitobates inscriptus, male. 222–224, Heteromitobates albiscriptus, male, female, and detail showing anexample of epidermic pigmentation of the type ‘homogeneous mask’. 225, Heteromitobates anarchus, sp. nov., male.


density of medium-sized granules. Coxa I: medium density ofminute granules. Coxa IV: medium density of medium-sizedgranules.

Chelicerae. Segment I with 2 basal and 1 smalldorsal–retrolateral apical granules. Segment II with highdensity of small granules.

Pedipalps. Trochanter with 2–3 (3) dorsal and 3 ventralelevations. Femur with 5–8 (6/7) ventral elevations (standard

armature Iiiiiii), 1 retrolateral subapical seta and dorsalgranulation with low density of medium-sized granules.Patella: ventral–retrolateral subapical tubercle like a smallgranule. Tibia with 2 short ventral–prolateral sub-basal spines,in addition to the standard armature.


226 227 230

232

228

229

231

Figs 226–232. Photographs. 226, Heteromitobates harlequin, sp. nov., male. 227, 228, Heteromitobates alienus,sp. nov., male (photo by R. Pinto-da-Rocha), and example of epidermic pigmentation of the type ‘fragmented mask’.229,Mitogoniella indistincta, male. 230,Mitogoniella taquara, sp. nov., male. 231, Acutisoma hamatum, female (photo byG. Machado). 232, Acutisoma longipes, male.


Leg II. Trochanter with low density of minute granules.Femur and patella with minute granules. Other segmentspractically smooth.

Leg III. Trochanter with low density of minute granules.Femur, patella and tibia with small granules.Metatarsus withminute granules.

Leg IV. Apical apophysisofcoxaof similar size:prolateralmore robust, pointed, oblique, and with a small posteriorsubapical process; retrolateral almost longitudinal and

crescentlike (prolateral curvature). Trochanter with basalapophyses with same size; retrolateral apical apophysisvery small. Femur medium-sized and slightly curved inward,with medium-sized granules (larger on row2 and apical fifth ofrow3); retrolateral apical apophysis medium-sized, straight,and transversal; prolateral apical apophysis small. Tibia andmetatarsus with small granules, but row2 with higher granules.


Fig. 233. Hypothesis of phylogenetic relationship among species of Goniosomatinae. Only one most parsimonious tree –97 characters, L = 5629, CI= 0.29, RI= 0.68. Changes in character state for each clade are given in Table 2. Bremersupport numbers are given for internodes (bold numbers refer to genera level). Bars to the right identify species groupedinto each genus. The box at the right shows the abbreviations of the geographic areas were each species was collected(see abbreviations and discussion in ‘Biogeography’).



Colour pattern

Animal strongly black with reddish-black dorsal scute. Dry-mark on apex of coxa, at base and apex (variation: only at base) oftrochanter and at base of femur, in addition to on dorsal scute(diagnosis of the genus). Black articular membranes betweencoxae and trochanters.


Dorsal scute: width: 8.14–9.99 (9.91), length: 7.82–9.54(9.15), leg I: 30.74–37.03, II: 67.87–75.54, III: 50.12–58.23(57.49), IV: 67.16–78.82 (75.27), femur IV: 19.43–23.76(23.76).

Female

In addition to dimorphic characters already mentioned, thefemale differs by: Angles of posterior margin of dorsal scutewith short spines; angles of free tergites with robust spines.Dry-mark as a frame on abdominal scute and in a narrow stripebehind eye-mound; on the entire dorsum of trochanter.

Remarks

This species is very similar to A. longipes but it can bedistinguished by the colour pattern and smaller granules on thedorsal scute.

Acutisoma longipes Roewer, comb. reestabl.

(Figs 3, 109–111, 188, 189, 232)

Acutisoma longipes Roewer, 1913: 277, 280. – 1923: 505, 506, 1930:388; Mello-Leitão, 1923: 191, 1932: 275, 277; B. Soares, 1946: 495;Soares & Soares, 1948: 625; Machado et al., 2001: 17; Kury, 2003:116.

Acutisoma monticola Mello-Leitão, 1922: 342. – 1923: 160, 191, 1932:275, 278, 1937: 294; Roewer, 1930: 388; Piza, 1942: 404; Kury, 2003:116. Syn. nov.

Leitaoius xanthomusMello-Leitão, 1935a: 404. – 1935b: 111, 1940: 25;B. Soares, 1944a: 262 (syn. Acutisoma monticolum Mello-Leitão,1922). Syn. nov.

Leitaoius nitidissimus Mello-Leitão, 1940: 25. – B. Soares, 1944a: 262(syn. Acutisoma monticolum Mello-Leitão, 1922). Syn. nov.

Mitogoniella mutila Piza, 1938: 140. – B. Soares, 1944d: 221(syn. Acutisoma monticolum Mello-Leitão, 1922). Syn. nov.

Acutisoma monticolum. – B. Soares, 1944a: 262, 1944d: 221, 1945c:351, 1946: 496; Soares & Soares, 1945: 252–254, 1948: 626.

Acutisoma sp. – Trajano, 1987: 547, 548.Goniosoma longipes. –Gnaspini & Trajano, 1994: 573; Pinto-da-Rocha,1995: 81; Gnaspini, 1996: 433; Machado & Oliveira, 1998, 2002:1510, 1519, 1521; Gnaspini, 1999; Sabino & Gnaspini, 1999: 677;Machado et al., 2000; Machado et al., 2001: 22; Machado andRaimundo, 2001: 138–142; Willemart, 2001: 249, 251, 2002: 51;Machado, 2002; Santos & Gnaspini, 2002: 179; Gnaspini et al.,2003: 33; Hara & Gnaspini, 2003: 258, 262, 266, 269, 273; Haraet al., 2003: 441, 442; Willemart & Gnaspini, 2004a: 16, 21, 22, 24,2004b: 230, 232.

Goniosoma cf. longipes. – Sabino & Gnaspini, 1999.

Material examined

Types. São Paulo: Franca, ma lectotype and 1 fe paralectotype(here designated) (ROEWER889); Campos do Jordão, type fe ofAcutisoma monticolum (MZSP1505), III/1906; São Paulo (Serra daCantareira), type fe of Mitogoniella mutila (MZLQ0026), Worontzow leg.,1935.

Other specimens examined. Minas Gerais: São Sebastião do Paraíso,1 ma (MZSP1797), Araújo leg., II/1945; Pocos de Caldas (Fonte dosAmores), 4 ma and 5 fe (MNRJ), J. Becker & O.A. Roppa leg., X/1963;Pocos de Caldas (Córrego do Meio), 1 ma (MZSP), J. Becker et al. leg.,VII/1967. São Paulo: Analândia (Fazenda da Toca, gruta de arenito), 3 maand 3 fe (MZSP22581), A. Mesa leg., IX/1982; Ipeúna (Gruta Fazendão),1 ma (MZSP22579), E. Trajano leg., XI/1999; Monte Alegre do Sul, 1 ma(HS253), J.L. Lima leg., 1947; Campos do Jordão, 1 ma (IBSP769),E.R. Pedroso leg., VI/1992; Campos do Jordão (Engenheiro Lefevre),3 ma (MZSP16372), III/1963; Pindamonhangaba (Fazenda SãoSebastião do Ribeirão Grande), 2 fe (IBSP766), R. Martins leg., XII/1994;Atibaia (Parque Florestal de Itapetinga), 1 ma and 2 fe (MZSP14703),G. Machado leg., III/1994; Atibaia (Pedra Grande, mata), 1 ma(MZSP27304), R.H. Willemart leg., VI/2001; Atibaia (Gruta do Fundo),1 fe (MZSP22580), F.H.S. Santos leg., IX/2000; Atibaia (Gruta do Fundo),1 ma (MZSP), F.H.S. Santos leg., X/2000; Itu, 1 ma (IBSP765),R. D’Ávila leg., VIII/1997; São Paulo (Serra da Cantareira), 1 ma(MZSP52), XII/1938.

Description

Dorsum. Eye-mound round with the space between thetubercles very high (about 2� the size of eye-mound);tubercles weak and high. Area I with 1 pair of tubercles alittle distinct from the other granules of area. Area III with1 pair of weak spines close to each other (same distance asbetween eyes) and slightly backward. Angles of posteriormargin of dorsal scute and free tergites with weak and hightubercles.

Granulation. Carapace: 3–20 (6) medium-sized granules.Areas I–III: 17–42 (22) medium-sized granules. Lateralmargin and free tergites: medium/low density of medium-sized granules. Posterior margin: low/medium density ofmedium-sized granules. Anal operculum: mediumdensity of small granules. Venter: Very granulated. Posteriormargin of stigmatic area and free sternites: high/mediumdensity of medium-sized granules. Coxa I: low density ofsmall granules. Coxa IV: medium density of medium-sizedgranules.

Chelicerae. Segment I with 2 basal, 1 dorsal–retrolateralapical and 1 prolateral apical granules. Segment II with mediumdensity of small granules.

Pedipalps. Trochanter with 1–3 (3) dorsal and 2 ventralelevations. Femur with 6–11 (8/11) ventral elevations(standard armature Iiiiiii), 1 or 2 (2) retrolateral subapicalsetae and dorsal granulation with low density of medium-sizedgranules. Patella: ventral–retrolateral subapical tubercle like asmall granule. Tibia with 1 low ventral–prolateral subapicaltubercle, in addition to the standard armature.


Leg II. Trochanterwith medium density of small granules.Femur with medium-sized granules. Patella with smallgranules. Tibia with minute granules. Metatarsus smooth.


Leg III. Trochanterwithmediumdensity of small granules.Femur, patella and tibia with medium-sized granules.Metatarsus with small granules.

Leg IV. Apical apophyses of coxa of similar size, beingprolateral more robust, pointed, almost transversal (variation:oblique); retrolateral almost longitudinal crescentlike(prolateral curvature). Trochanter with basal apophyses withsame size; retrolateral apical apophysis very small. Femurwith slightly inward curvature (variation: medium curvature),with medium-sized granules (larger on row2 and apical fifth ofrow3); retrolateral apical apophysis medium-sized, straightand transversal; prolateral apical apophysis small. Tibiaand metatarsus with small granules, but row2 with highergranules.

Tarsal segmentation. 9–11 (10/11), 16–22 (22), 11–14(11/12), 12–16 (13/14).


Colour pattern

Dorsal scute yellowish-brown; pedipalps, chelicerae andlegs I–III greenish-black; the rest light-reddish-brown. Dry-mark in traces of frame at the abdominal scute; on apex ofcoxa, at base of trochanter and at base of femur, in addition toon lateral margin of dorsal scute (diagnosis of the genus). Purplearticular membranes between coxae and trochanters.


Dorsal scute: width: 8.62–10.8 (9.83), length: 7.91–10.12(9.06); leg I: 28.78–44.33 (37.56), II: 60.53–89.20 (74.98),

Table 2. Changes in character state for each clade of the cladistic hypothesis obtained (as in Fig. 233)For each branch, non-homoplastic changes are given in bold, reversions are noted with an ‘r’, ‘+’ and ‘–’ denote increase or decrease of the values of the

continuous or quantitative characters, and italics indicates that ambiguous changes are shown using ACCTRAN

Clade Changes in character states

Goniosomatinae 49(0) 51(1) 55(1) 60(1) 64(1) 70(1) 72(1) 73(1) 74(1) 76(–) 89(0) 95(+) 96(+) 97(+)Serracutisoma+ 8(1) 34(1) 39(1) 43(2) 72(2) 81(–) 94(+)Goniosoma+ 1(1) 26(2) 38(3) 52(1) 68(1) 75(–) 77(–) 78(+) 95(+)

Goniosoma 5(1) 6(1) 10(3) 11(1) 12(1) 13(1) 14(1) 18(2) 21(1) 22(2) 24(1) 32(1) 33(1) 34(0) 40(2) 46(2)48(1) 72(3) 82(+) 96(–) 97(–)

(G. varium+ +G. vatrax+)+ 3(1) 4(1) 13(2) 17(1) 28(1) 49(1) 67(0) 71(0) 75(+) 96(–)G. varium+ 2(2) 21(2) 32(0r) 59(1) 66(2) 67(4) 68(2) 71(0) 78(–) 82(–)G. capixaba+ 22(1) 24(0) 33(0r) 40(1) 79(–) 94(–) 95(+) 96(+) 97(+)G. venustum+G. roridum 1(2) 3(2) 8(0r) 9(3) 13(3) 16(1) 18(0r) 21(3) 26(1) 30(1) 48(0r) 72(2) 75(–) 82(+) 94(–)

95(+) 96(+)G. vatrax+ 1(2) 18(3) 43(2) 72(2) 95(–) 96(–) 97(–)G. macracanthum+ 3(2) 9(2) 26(3) 33(0r) 43(0r) 68(0r) 71(1) 75(–) 79(+)(G. dentipes+G. ensifer)+ 2(1) 3(3) 42(2) 52(2) 73(2)

G. dentipes+G. ensifer 29(1) 64(2) 67(5) 68(1) 75(+) 76(+) 77(+) 78(–) 79(–) 82(–) 83(–) 93(–) 94(–) 96(+) 97(+)G. apoain+ 1(3) 31(1) 39(2) 49(0r) 72(3) 74(2) 94(+)G. unicolor+G. calcar 33(1) 42(0) 43(1r) 64(0r) 71(0) 80(–)

Pyatan 16(2) 17(2) 19(1) 22(3) 30(1) 79(–) 93(–) 94(–) 95(+)Serracutisoma 2(3) 4(1) 7(1) 9(1) 20(1) 23(3) 35(1) 42(1) 43(3) 67(1) 78(–) 96(+)

S. fritzmuelleri+ 16(1) 58(2) 83(–) 93(+) 96(+)S. proximum+ 20(0) 23(1) 40(1) 59(1) 75(+) 76(+) 77(+) 79(–) 81(+)S. banhadoae+ 20(2) 36(1) 37(1) 57(1) 62(1) 63(0) 64(0r) 67(0) 80(+) 96(–)S. molle+ 23(0) 45(1) 58(1r) 75(+) 76(+) 77(+) 96(–)S. pseudovarium+ S. thalassinum 20(0r) 24(1) 45(2) 56(1) 81(+)

S. catarina+ 23(4) 27(1) 34(3) 37(1) 39(0) 97(+)S. spelaeum+ 25(1) 45(1) 64(0r) 67(0) 77(–) 83(+)S. guaricana+ S. inerme 26(1) 63(0)

Heteromitobates+ 10(1) 19(1) 38(1) 61(2) 67(3) 69(1) 71(0) 82(+) 95(+) 96(+)Heteromitobates 7(2) 10(2) 13(4) 16(3) 22(3) 26(1) 50(1) 57(2) 58(2) 66(1) 72(2)

(H. discolor+H. inscriptus)+ 53(1) 61(3) 95(+)H. discolor+H. inscriptus 78(+)H. albiscriptus+H. anarchus 57(0r) 67(2) 93(–) 95(+)

H. harlequin +H. alienus 17(1) 38(2) 40(1) 67(4) 69(0r) 79(+) 81(–) 93(+) 94(–)Mitogoniella+Acutisoma 2(1) 15(1) 24(1) 42(1) 54(1) 69(2) 76(+) 80(+)

Mitogoniella 15(2) 21(4) 22(4) 38(2) 67(4) 79(+) 83(+) 94(–) 95(–) 96(+) 97(+)M. unicornis+ 36(1) 37(1) 66(1) 72(2) 80(–) 81(–) 93(+)M. indistincta+M. taquara 7(1) 8(1) 54(0r) 75(+) 78(–) 79(–) 83(–) 95(+)

Acutisoma 2(2) 4(1) 40(1) 44(1) 64(0r) 79(–) 80(+) 96(–)A. coriaceum+ 18(1) 19(0r) 39(1) 41(1) 46(2) 47(1) 81(+) 93(+)


III: 43.77–68.85 (56.97), IV: 60.37–90.52 (73.03), femur IV:17.22–27.75 (21.28).

Female

In addition to dimorphic characters already mentioned,the female differs by: Angles of free tergites with robustspines and of similar size of spines of area III. Dry-mark asan entire frame on abdominal scute.

Biology and records

Trajano (1987) [cav.]; Gnaspini and Trajano (1994) [cav.]; Pinto-da-Rocha (1995) [cav.]; Gnaspini (1996) [cit.]; Machado andOliveira (1998 [biol.], 2002 [cit.]); Gnaspini (1999) [biol.];Sabino and Gnaspini (1999) [biol.]; Machado et al. (2000

[biol.], 2001 [cit.]); Machado and Raimundo (2001) [cit.];Willemart (2001 [cit.], 2002 [cit.]); Machado (2002)[cit.]; Santos and Gnaspini (2002) [cit.]; Gnaspini et al. (2003)[cit.]; Hara and Gnaspini (2003) [biol.]; Hara et al. (2003) [cit.];Willemart and Gnaspini (2004a, 2004b) [cit.].

Remarks

The combination Goniosoma longipes has never beenformalised in a taxonomic publication. However, it has beenused in papers about natural history and faunistic surveys,following the identification and suggestion from Stefanini-Jim’s unpublished M.Sc. Thesis. Therefore, we considered thatthis was a valid combination and proposed that the combination

G GoniosomaP PyatanS SerracutisomaH HeteromitobatesM MitogoniellaA Acutisoma

171 Km

800 m1200 m

25°

40°45°50°

20°

Fig 234. Map of distribution of the genera of Goniosomatinae.


Acutisoma longipes should be considered a reestablishedcombination.

The distribution of Acutisoma longipes is much wider thanthat of the other species in the subfamily, since it includes theinterior of São Paulo state and the eastern region of Serra daMantiqueira of São Paulo and Minas Gerais states. In addition,there is a high morphological variation related to size andcurvature of femur IV, width of coxa IV and colour pattern.Therefore, it is possible that there is a complexof species related toA. longipes. However, this variation overlaps and forms a

continuum when many populations are analysed. Thus weconsider it to be a single species for the time being.

Acutisoma hamatum is very similar to A. longipes butwithout morphological or geographic overlapping. Thus, wekeep them as distinct species, although we understand that it ispossible that a future analysis may conclude that theyare synonyms, what would enlarge even more themorphological variation and geographic distribution ofA. longipes. It is also a well-studied species from a naturalhistory point-of-view.

800 m

1200 m

800 m

1200 m

800 m

1200 m

800 m

1200 m

171 Km

171 Km

171 Km

171 Km

45° 40°

20°

50°

25°

20° 40° 23°

25°

20°

45° 24°

20°

47°30′

22°30′

47°30′

235 237

236

238

Figs 235–238. Map of distribution of the species of the genera. 235, Goniosoma (species: 1 =G. varium, 2 =G. capixaba,3 =G. venustum, 4 =G. roridum, 5 =G. vatrax, 6 =G. macracanthum, 7 =G. dentipes, 8 =G. ensifer, 9 =G. apoain, 0 =G. unicolor,A=G. calcar, B=G. carum). 236, Serracutisoma (species: 1 = S. proximum, 2 = S. banhadoae, 3 = S. molle, 4 = S. pseudovarium,5 = S. thalassinum, 6 = S. fritzmuelleri, 7 = S. catarina, 8 = S. inerme, 9 = S. guaricana, 0 = S. spelaeum). 237, Heteromitobates andAcutisoma (species: 1 =H. discolor, 2 =H. inscriptus, 3 =H. albiscriptus, 4 =H. anarchus, 5 =H. harlequin, 6 =H. alienus,7 =A. coriaceum, 8 =A. hamatum, 9 =A. longipes). 238, Mitogoniella (species: 1 =M. indistincta, 2 =M. taquara,3 =M. unicornis, 4 =M. modesta).


BiogeographyGoniosomatinae harvestmen live in the Atlantic rainforest in itsmost humid areas, mainly in ombrophylous dense forests, in anarrow area on the coastal lowlands and related mountains,from Bahia state (in the northeastern region of Brazil) to SantaCatarina state (in the southern region of Brazil) (Fig. 234). A fewspecies occupy all (Goniosoma carum, G. vatrax andMitogoniella indistincta) or part of its range (Mitogoniellataquara, sp. nov., Acutisoma longipes and Serracutisomamolle) in transitions to semi-deciduous seasonal forest. Allspecies have a restricted range, much smaller than the wholerange of the subfamily or of each genus (Figs 235–238).Figures 233 (see box on the right) and 239 respectively show theareas of distribution plotted over the cladogram of the subfamily,and an area cladogram plotted against a map of the region.

Pinto-da-Rocha et al. (2005) used phylogenetic hypotheses ofGoniosomatinae (here proposed),Caelopyginae (Pinto-da-Rocha2002), Sodreaninae and Progonyleptoidellinae (Pinto-da-Rocha,unpubl. data) to present a historical biogeographic hypothesis of

biota diversification in the Atlantic rainforest. They showed thehigh degree of endemism in harvestmen by a similarity analysisamong several well-sampled localities and pointed out theconsiderable influence of geomorphologic history in isolationand speciation of harvestmen. The historical biogeographicanalysis with the four subfamilies cited above resulted ineleven areas of endemism from Bahia to Santa Catarina. Theanalysis showed possible past vicariance events following anorth-to-south sequence that separated the fauna of these areas.

Among the subfamilies studied, Goniosomatinae is the mostinformative for historical biogeography, both by its greatnumber of species and its highest degree of endemism. Thirtytwo of its 36 species are restricted to only one area of endemismproposed by Pinto-da-Rocha et al. (2005) (half of the totalendemic species) and all areas harbor at least one endemicgoniosomatine (Fig. 239). Two areas, southern São Paulo(SSP) and Serra do Espinhaco (SEsp), were only delimited byendemism of 2 and 3 species of Goniosomatinae respectively(Fig. 239).

45° 40°

25°

20°

800 m

1600 m

Fig 239. Area cladogram (modified from Pinto-da-Rocha et al. 2005, and showing the relationship amongthe areas of endemism within the general area of distribution of Goniosomatinae), in which we plotted thenumber of species in each area (between parentheses). See text for abbreviations.


In addition to their importance for defining areas ofendemism, Goniosomatinae seems to contain great historicalinformation for the phylogenetic relationships of its species toinfer relationship among areas. The distribution maps of genera(Fig. 234) and the area cladogram of Goniosomatinae (Figs 233,239) show a structured pattern of early disjunction ranges in abasal level in its phylogeny. Mitogoniella is distributed in thenorth-eastern region and interior areas of Minas Gerais state(‘Bahia’ (BA), ‘Serra do Espinhaco’ (SEsp), and ‘Serra daMantiqueira’ (Mnt)) – the latter harbors Acutisoma, its sister-genus.Heteromitobates ranges in a narrow strip on Serra do Marridge and adjacent coast (‘Serra do Mar de São Paulo’ (SMSP),‘north coast of SãoPaulo and southofRio de Janeiro’ (LSRJ), and‘Serra da Bocaina’ (Boc)). In the other clade, Goniosoma livescloser to the coast in central areas of endemism (‘Espírito Santo’(ES), ‘Serra dos Órgãos’ (Org), and Boc+LSRJ, SEsp and Mnt).Serracutisoma has a range separated from Goniosoma+Pyatan,with its species having the southernmost distribution within thesubfamily (SMSP to ‘Santa Catarina’ (SC)). Even with somesympatry occurring among Acutisoma, Mitogoniella andHeteromitobates, mainly in the SMSP area, these patterns ofdisjunctions may indicate that important early vicariance eventsaffected Goniosomatinae, agreeing with the area cladogramproposed by Pinto-da-Rocha et al. (2005) (see Fig. 239).These patterns lead to the following possible explanationsabout the geographic evolution of goniosomatines.

First, Goniosomatinae probably differentiated in at leasttwo main lineages (Goniosoma+Pyatan+ Serracutisoma andHeteromitobates +Mitogoniella +Acutisoma) before the mainvicariance events proposed by Pinto-da-Rocha et al. (2005)occurred in the Atlantic forest, and these lineages werewidespread over the area (although one should note that thefirst group does not reach the southernmost distribution ofthe subfamily). Afterwards, an important vicariance eventoccurred around the area that includes Boc, LSRJ, and SMSP,in a different way (i.e. with probably different vicariance barriers)for each one of the two main goniosomatine lineages. In the firstmain clade, there was a separation between a northern(Goniosoma+Pyatan) from a southern (Serracutisoma) biota,whereas in the second clade there was a separation between thecoastal (Heteromitobates) and the interior (Mitogoniella +Acutisoma) regions. Yet, it may be related to the appearanceof great rivers or their valleys (e.g. Rio Doce and Rio Paraíba doSul), and posterior marine transgressions, and uplifting ofSerra do Mar and Serra da Mantiqueira mountain ranges(Pinto-da-Rocha et al. 2005).

Second, the high degree of replication of areas of endemismand their relationships, or the paralogous pattern showed in thephylogeny in a specific level (Fig. 233), indicates many dispersalevents among closer areas of endemism alternated with allopatricand/or some sympatric speciation events. The more recentspeciation events were processes that kept these populationsrestricted to the areas of endemism.

Third, almost all recent species seem to have a smalldistribution range, staying endemic to single areas (Figs 233,235–238). Most species maintain this restriction even in areaswhere there is no barrier present for the expansion of their speciesrange (e.g. Org/LSRJ, LSRJ/SMSP, SMSP/SSP). Some

biological characteristics may explain this low vagility forharvestmen in general, and for Goniosomatinae in particular.Firstly, as a general rule, harvestmen live in humid places, andthey are cryptical and nocturnal animals (Pinto-da-Rocha et al.2007). Secondly, and more importantly, some studieshave shown that goniosomatines have high philopatry(e.g. Gnaspini 1996; Santos and Gnaspini 2002; Willemart andGnaspini 2004b; has shown that individuals of populations ofSerracutisoma spelaeum associated to different entrances of thesame cave did not move from one population to the other, evenknowing that these populations are only a few meters apartfrom each other when they rest inside the cave, and that eachspecimen may take a long route outside the cave duringnocturnal foraging activities), which is related togregariousness and sub-social behavior (the latter in turnprobably related to the presence of parental care in specieswith short ovipositors, as is the case with laniatoreans) (e.g.Machado 2002; Willemart and Gnaspini 2004a; Ferreira et al.2005; Buzatto et al. 2007). Moreover, Oliveira et al. (2006)showed a large variation in the number of chromosomes ofS. spelaeum (2n = 92–109; from the same population), whichis not common and may be related to this high philopatry and‘low’ population numbers. Concerning phylogeographicstudies focusing on other groups of harvestmen, a probablepoor and low dispersing ability is reported to result in the veryhigh variation in genetic patterns detected among differentpopulations of the same species of a cyphophthalmid (Boyeret al. 2007) and also of a triaenonychid laniatorean (Thomasand Hedin 2008), which, in combination with the highlyconservative morphology observed among the differentpopulations, led those authors to discuss whether they weredealing with cryptic species. Therefore, goniosomatines mayalso be good targets for phylogeographic studies.

Three species (Acutisoma longipes, Mitogoniella taquara,sp. nov. and Serracutisoma molle) are here hypothesised tohave had recent range expansions. These species ranges spreadover the boundaries of the coastal Atlantic forest areas ofendemism into semi-deciduous seasonal forest areas. It seemsthat the distribution patterns of these species need otherexplanations in addition to the historical scenario proposed byPinto-da-Rocha et al. (2005), whichwas based on the congruencebetween geographic distribution and phylogenetic relationships.We suggest that these three species have recently dispersedtoward the interior region from their original Atlantic forestareas of endemism, because they probably have lower climaterestrictions and could have dispersed to drier ecosystems (Santos2003 has studied the ecophysiology of cave and epigeangoniosomatines and other gonyleptids, for comparative reasonsin his unpublished Ph.D. thesis).

Although this subfamily had a major importance forunderstanding the historical biogeography of the Atlanticrainforest (as previously discussed), most samplingconcentrated close to main cities in the south-eastern region,mainly São Paulo, Rio de Janeiro and Curitiba. Considering thegeographical extension of the Atlantic forest (with a large areastill unsampled) and the high level of endemism presented bythe known species of the subfamily (see discussion above), thereare still large gaps with no knowledge about the harvestman


fauna, especially in the north-eastern part of Brazil and effortsshould be made to cover these gaps of information, which areimportant for a better understanding of the history ofdiversification of the Atlantic forest and its fauna.

Acknowledgements

This study was partially supported by a FAPESP (Fundacão de Amparo àPesquisa do Estado de São Paulo) research grant # 00/04686-4 and a researchfellowship from CNPq (Conselho Nacional de Desenvolvimento Científico eTecnológico) # 300326/94-7, Brazil, to PG, and by aCNPqn� 112091/1998-9andaFAPESPn�00/00270-8 research fellowships toMBS.We thankRicardoPinto-da-Rocha, Adriano Brilhante Kury, and the anonymous reviewers forcriticisms and suggestions to the manuscript.

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Manuscript received 2 April 2009, accepted 21 October 2009


Appendix 1. List of characters and character states

(Continued next page)

1–Penis, ventral plate– shape of apicalmargin (fromadorsal view) [0-1-2-3]:0 – concave1 – straight2 – convex3 – triangular

0

1

2

3

The apical margin becomes straight at nodeGoniosoma +Pyatan; it becomes

convex twice independently withinGoniosoma; and it becomes triangular(from a convex lineage) in a group of 3 species (G. apoain, G. calcar, andG.unicolor).A reversion to the concave state occurs inG.varium. Theotherfour genera maintained the ancestral state (concave apical margin),although that concavity is less pronounced than that of the outgroup.

2 – Penis, ventral plate – shape of lateral margins [0-1-2;0-3]:0 – concave1 – straight2 – convex3 – enlarged/angular

0 213 The ‘enlarged/angular’ margin was considered derived from ‘concave’

because that type of plate shows a slight concavity in some species. Theconcave margin changes to a straight margin three times: at nodeMitogoniella+Acutisoma, and in two groups of species of Goniosoma.In this genus the change to a straight plate is followed by a narrowing of theventral plate, which does not occur in Mitogoniella+Acutisoma. InAcutisoma it becomes convex; and the concave margin becomesenlarged at the node of Serracutisoma.

3 – Penis, ventral plate – apex [0-1-2-3]:0 – without extension1 – with extension, without constriction2 – with extension, with one constriction3 – with extension, with two constrictions

0

12 3

The apical region of the ventral plate can be extended above the apical groupof setae. In this case, it can present one or two lateral constrictions. Thischaracter only changes within Goniosoma. State 1 appears in Goniosoma(exceptG. carum); state 2 appears twice, inG. roridum+G. venustum andin G. macracanthum+, in which, later, state 3 appears in(G. dentipes+G. ensifer)+.

4 – Penis, ventral plate – bristles on ventral face:0 – present1 – reducedThe bristles are absent in Acutisoma, Serracutisoma (with a reversion inS. proximum), and inGoniosoma (exceptG. carum). In addition, they werenot observed in Mitogoniella taquara. There is a high variation indensity, distribution, and size of the bristles, although the cases ofreduction can be considered to be similar.

5 – Penis, ventral plate – setae – shape (mainly the basal group):0 – spatulate1 – pointed

0 1

Pointed setae are a synapomorphy of Goniosoma. However, there is a highvariation in the width of spatulate setae, and some of them are almostpointed in other genera.

6 – Penis, ventral plate – setae – pattern of the apical group:0 – 2 rows1 – 1 row

0 1

The setae of ventral plate can be divided in a basal and an apical group. In itsturn, the apical group can be divided in a dorsal and a ventral longitudinalrow. The former is present in all taxa examined. The presence of a dorsalrow alone (i.e. the absence of a ventral row) is a synapomorphy ofGoniosoma, with a reversion in G. vatrax. The absence of a ventral rowfollows a general path towards a reduction in the number of setae.

7 –Penis, ventral plate – setae – number and organisation of setae of the dorsalrow in the apical group [non-additive]:

0 – 4 setae1 – 5 setae, 1 displaced2 – 5 setae on a row

1

2

0


Appendix 1. (Continued)

The dorsal row of the apical group may have 4 or 5 setae. In the last case, theadditional seta (which is always the smallest one) can be aligned with ordisplaced from the other setae. The character was coded as non-additivebecause there is no evidence for the position of this additional seta when itfirst appeared. Thus, the displaced seta is considered a synapomorphy ofHeteromitobates and the aligned setae appear twice independently, inSerracutisoma and in Mitogoniella indistincta+M. taquara.

8 – Penis, ventral plate – setae – position of more basal seta of apical group:0 – far from the others1 – equidistant to the others

1

0

This more basal seta is the smallest of the apical group. It becomes

‘equidistant’ in the large clade Serracutisoma+ and in Mitogoniellaindistincta+M. taquara, occurring 3 reversions within Goniosoma: inG. unicolor, in G. vatrax, and in G. roridum+G. venustum.

9 – Penis, ventral plate – setae – organisation of basal group [0-1;0-2-3]:0 – oblique row1 – transversal row2 – longitudinal row3 – inverted-L-shaped row

0

1

2 3

The basal group of setae is organised in a single row, which direction vary inrelation to the longitudinal axis of theventral plate. It becomes transversal inSerracutisoma, longitudinal inG.macracanthum+ andchanges to inverted-L-shaped row in G. ensifer; there is a reversion to oblique row inG. unicolor. Yet, there is a direct change from state 0 to state 3 inG. roridum+G. venustum.

10 – Penis, ventral plate – setae – number at the apical ventral group [0-1-2;0-3]:

0 – 1 seta1 – 2 setae2 – 3 setae3 – 0 seta

0

1

2

3

These ventral setae are lost in Goniosoma (synapomorphy; reversion inG. vatrax). Two setae appear in Heteromitobates+, and a third setaappears independently in Heteromitobates and Mitogoniella unicornis.

11 – Penis, ventral plate – setae – number in the basal group [0-1;0-2]:0 – 4 setae1 – 3 setae2 – 5 setae

1 2

0

From the plesiomorphic number (4), a seta is lost inGoniosoma (state 1), andan additional seta appears only in Mitogoniella modesta (state 2).

12 – Penis, ventral plate – setae – small ventral seta, close to basal group:0 – absent1 – present

Although this seta is considered apart from the basal group, it could beconsidered the fourth seta of character 11; in this case, state 0of character 11should have been divided into 2 states. The equivalence between thisreduced seta and that fourth seta was corroborated by the parsimonyanalysis: the fourth seta of the basal group was lost (character 11, state1) and the reduced seta appeared (character 12, state 1) at the node ofGoniosoma. This codification was preferred for matters of simplification.

13 – Penis, glans – ventral process [0-1-2-3;0-4]:0 – with apical lamina inserted by base, with projections1 – with apical lamina inserted by base, without projections2 – thin, without apical lamina3 – absent4 – with apical lamina inserted by center, with projections

041

2 3

The ventral process can be absent or present, with variation in the shapeof the apical lamina and in its insertion on the stem. State 1 appears asa synapomorphyofGoniosoma, with a reduction to thin (state 2) at the nodeGoniosoma varium+ +G. vatrax+ (ACCTRAN), and the process is lost(state 3), twice: in G. roridum+G. venustum and in G. vatrax. InHeteromitobates the process changes from state 0 to 4.

14 – Penis, glans – stylus – shape of apex:0 – round1 – with a dorsal–apical beak




0 1 Synapomorphy of Goniosoma.

15 – Penis, glans – stylus – shape [0-1-2]:0 – straight1 – 90� angle2 – 90� angle with scales

1

2

0

The sharply bent stylus is a synapomorphy of Acutisoma+Mitogoniella, andthe scales appear in Mitogoniella.

16 – Penis, glans – dorsal process [non-additive]:0 – absent1 – small, callus-like2 – finger-like3 – large, sac-like

123 0

The large dorsal process appears only as a synapomorphy ofHeteromitobates.Independently, the callus-like appears in S. fritzmuelleri+ (with a reversionin S. pseudovarium) and in G. roridum+G. venustum. The finger-likeprocess appears autapomorphically in Pyatan insperatum.

17 – Penis, glans – Truncus [0-1;0-2]:0 – medially invading the base of the ventral plate on dorsal face1 – not invading the ventral plate2 – entirely invading the base of the ventral plate on dorsal face

01 2

State 1 appears twice: in Goniosoma except G. carum (with a reversion inG. varium) and in Heteromitobates alienus+H. harlequin. State 2appears autapomorphically in Pyatan insperatum.

18 – Leg IV, femur – inward curvature [0-1-2-3-4]:0 – absent1 – light

2 – medium3 – strong4 – very strong

01

23 4

In dorsal view, femur IV may be straight or from slightly curved to sharplybent. The curved femurhas a slight ‘S’ shape,with the outward curvedbasalregion smaller than the inward curved apical one. The variation of thischaracter concerns only the apical curvature. The curvature of the femurappears three times independently: inAcutisomahamatum+A. longipes, inS. pseudovarium, and as a synapomorphy of Goniosoma. In Goniosoma,state 2 appears directly from state 0, and within this genus it changes tostrong inG. vatrax+ and to very strong only inG. calcar; there is a reversionto state 0 in G. roridum+G. venustum and two reversions to state 2 inG. dentipes and in G. unicolor.

19 – Leg IV, femur – outward curvature [0-1-2]:0 – absent1 – light2 – strong

0 12

The light curvature appears twice, in Heteromitobates+ and as anautapomorphy of Pyatan. The strong curvature appearsautapomorphically in H. alienus. A reversion occurs in Acutisomahamatum+A. longipes, which femora have an inward curvature(character 18).

20 – Leg IV, femur – position of bend [non-additive]:0 – absent1 – medial2 – medial to apical



0

1

2

A bent femur has a straight basal region and an oblique apical region,displaced upward and inward. This condition was considered ‘straight’ incharacters 18 and 19. That bend can be medial or more apical, and wewere not able to identify a transformation series between them. The bendis a synapomorphy of Serracutisoma (using ACCTRAN); it changes threetimes to state 2: in S. catarina, in S. inerme and in S. banhadoae+, with 2reversions (in S. proximum and in S. thalassinum+ S. pseudovarium).

21 – Leg IV, femur – irregularity on dorsal rows [0-1-2-3;0-4]:0 – very low1 – medium (more than 1/3 apical)2 – high with a little developed comb3 – absent, with only a well developed comb4 – absent

0 12 3

4

As cited in ‘Methods’, there is a regular pattern of rows of granules and spineson the legs, but thedorsal rows (row1and row6)may showavariation in thatpattern. Row1 may change its direction, confounding with row6 on theapical region, and these irregularities may reach the basal region. In thesecases, the rowsbecomea combof highgranules very close to eachother.Anirregularity of the dorsal rows (state 1) appears as a synapomorphy ofGoniosoma, changing to state 2 twice (inG.varium+ and inG.unicolor) andto state 3 in G. roridum+G. venustum. The irregularity disappears inMitogoniella (from state 0 to state 4), following a general tendency ofreduction in the ornamentation of legs.

22 – Leg IV, femur – granules of row2 [0-1-2;0-3;0-4]:0 – normally shaped1 – flat2 – flat and longitudinally elongate3 – pointed4 – minute to absent

01

3

4

There is a high variation in the ornamentation of legs, mainly on femur IV ofmales. That variation is more conspicuous on row2, which may representthe general ornamentation of the other rows and legs. The possiblehypothesis that normal granules change to flat granules before theychange to flat and longitudinally elongate ones was rejected in ourcladistic analysis. State 2 appears in Goniosoma, changing to state 1 inG. capixaba+, in G. macracanthum, and in G. dentipes. State 3 appearstwice independently: in Pyatan and as a synapomorphy ofHeteromitobates (using ACCTRAN). Granules reduce a single timeas a synapomorphy of Mitogoniella.

23 – Leg IV, femur – armature of row2 [0-1-2;0-3-4;0-5]:0 – absent1 – large spines in all extension, except on the apical region2 – large spines in all extension3 – large spines only on the medial–apical region4 – spines increasing from base to angle5 – spines only on subapical region

53 41 2

Granules may be replaced by spines with different sizes and distributionpatterns. We can suggest the armature of row2 as a synapomorphy ofSerracutisoma since all species of this genus possess it. However, there is avariation in the distribution of spines among specieswhich renders difficultto establish a series of transformation between the observed states, what ledus to propose independent acquisition directly from an absent state. Underthis hypothesis, state 1 appears in S. proximum+ changing to state 2 inS. proximum; independently, state 2 appears in S. thalassinum; state 3appears only in S. fritzmuelleri; state 4 appears independently inS. catarina+ and in S. molle; and state 5 appears only in S. pseudovarium.

24 – Leg IV, femur – armature of row3:0 – with spines1 – absent (only granules)

0

row3

Elevations of row3 increase towards the apical direction in all species. On thethird apical region these elevations may be granules or spines, sometimesforming a grade. This armature is reduced in Acutisoma+Mitogoniella,in Goniosoma (with a reversion in G. capixaba+), in Heteromitobatesalbiscriptus, in Serracutisoma proximum, and in S. pseudovarium+S. thalassinum.




25 – Leg IV, tibia – armature of row2:0 – absent1 – present

1fila2row2

It appears independently as a synapomorphy of Serracutisoma spelaeum+

and as an autapomorphy of Goniosoma carum.

26 – Leg IV, tibia – armature of row3 [0-1-2-3]:0 – absent1 – straight spines2 – curve spines3 – curve spines, in all extension

1

2

3

fila3

fila3row3

row3

The present series of transformation was proposed because curve spines arelarger than straight ones. In our analysis, the plesiomorphic state inGoniosomatinae may be either ‘absent’ or ‘straight spines’. If weconsider ‘absent’ as the plesiomorphic state, straight spines appear as asynapomorphy of Heteromitobates, and independently in Goniosomaroridum+G. venustum, in G. vatrax, in Serracutisomainerme+ S. guaricana, and in S. banhadoae. Curve spines appear inPyatan+Goniosoma. State 3 appears in (G. dentipes+G. ensifer)+ (with areversion to state 2 in G. apoain) and as an autapomorphy of G. carum.

27 – Leg IV, tibia – armature of row4:0 – absent1 – present

1

row4

It appears independently in Serracutisoma catarina+, in Heteromitobatesinscriptus, and in H. anarchus.

28 – Leg IV, tibia – comb of high granules on row2:0 – absent1 – present

1

row2

It appears in Goniosoma except in the more basal species, G. carum. Thischaracter follows the general trend of increasing of armature tibia IV in thisgenus. In this case, highergranules appears insteadof spines. It is interestingto notice that G. carum, the only species of the genus without this comb,has spines on row2 (character 25).

29 – Leg IV, tibia – retrolateral apical spine:0 – absent1 – present

1

This is a distinct, large, straight, and unique retrolateral apical spine close torow2. It appears only in Goniosoma dentipes+G. ensifer.

30 – Leg IV, tibia – ventral apical spine:0 – absent1 – present

1

This is a distinct, large and backward curved spine, close to row3. Itappears four times: in Heteromitobates alienus, in Pyatan, in G. vatrax,and in G. roridum+G. venustum.

The next three characters concern metatarsus IV, in which the strongestarmature occurs on its basal region, reducing towards the apex until itdisappears.

31 – Leg IV, metatarsus – armature of row2:0 – absent1 – present



row2

row3 row4

It appears as an autapomorphy ofGoniosoma carum and inG. apoain+ (witha reversion in G. unicolor).

32 – Leg IV, metatarsus – armature of row3:0 – absent1 – presentIt appears as a synapomorphy ofGoniosoma, with a reversion inG. varium+.

33 – Leg IV, metatarsus – armature of row4:0 – absent1 – presentIt appears as a synapomorphy of Goniosoma (using ACCTRAN), with a

reversion in G. capixaba+ and in G. macracanthum+.

34 – Legs II and III – armature [0-1-2-3]:0 – unarmed1 – spines close to each other2 – sparse spines3 – sparse spines with 2 or 3 larger ones

1

2

3

These spines are distinct from other granules that increase in size apicallyon ventral rows. The proposed series of transformation was rejected inthe analysis sincewe observed an acquisition of state 3 directly from state 1.Considering Serracutisoma+, state 1 appears either independently inPyatan and as a synapomorphy of Serracutisoma (using DELTRAN),or in Serracutisoma+ with a subsequent reversion in Goniosoma (usingACCTRAN). Inside Serracutisoma, it changes independently to state 2in S. proximum, to state 3 in S. catarina+, and with a reversion inS. pseudovarium. State 1 still appears independently in Heteromitobatesinscriptus.

35 – Leg II – armature of tibia – row 4:0 – absent1 – presentSynapomorphy of Serracutisoma (using ACCTRAN), with independent

reversions in S. spelaeum, S. fritzmuelleri, and S. banhadoae.

36 – Leg II – armature of tibia – row 3:0 – absent1 – presentIt appears four times: inM. unicornis+, in S. banhadoae+ (with a reversion in

S. pseudovarium), in S. catarina, and in S. guaricana.

37 – Leg II – armature of metatarsus – ventral rows:0 – absent1 – presentIt appears three times, with an evolution pattern similar to character 36: in

S. banhadoae+ (with a reversion in S. molle, using ACCTRAN), inM. unicornis+, and in S. catarina+ (with a reversion in S. guaricana).

The last three characters suggest two independent appearance of armature ofleg II, in the genera Mitogoniella and Serracutisoma.

38 –Apophyses of leg IV – shape of prolateral apical apophysis of coxa [non-additive]:

0 – falcate apex1 – with a small subapical process2 – blunt with a small subapical process3 – pointed with a small sub-basal process

01 2 3

processprocess

process

This apophysis has a high variation in shape and size. The differences aregenerally related to the presence and position of a sub-basal process. Forexample, the falcate apophysis is divided by a straight basal region untilthis process, after which the apex is curved, falcate. Due to the largemorphological difference between the falcate apophysis and the others,we prefer to consider this character as non-additive. State 1 appears atthe node of Heteromitobates, Acutisoma andMitogoniella, changing tostate 2 twice, in Mitogoniella and in H. alienus+H. harlequin. Pointedapophysis appears a single time in Pyatan+Goniosoma. The falcateapophysis was maintained only in Serracutisoma.

39 – Apophyses of leg IV – direction of the prolateral apical apophysis ofcoxa [0-1-2]:

0 – almost transversal1 – oblique, between 30� and 60�

2 – almost longitudinal

0 1 2

The direction of this apophysis is related to the longitudinal axis of theanimal. One hypothesis is that it becomes oblique in Serracutisomafritzmuelleri+ and in Pyatan+Goniosoma. An alternative hypothesisindicates a single appearance in Serracutisoma+ with a reversion inS. catarina+. In Goniosoma, it changes to state 2 in G. apoain+ (with areversion inG. unicolor). State 1 also appears independently in Acutisomahamatum+A. longipes (it is polymorphic in the latter).

40 – Apophyses of leg IV – retrolateral apical apophysis of coxa [0-1-2-3]:0 – absent or like a granule or tubercle1 – like a medium spine, pointed2 – near the size of that of trochanter IV3 – much larger than that of trochanter IV (more than 1.5x)

0 1 2 3




In most species this apophysis only varies in size, and in some species itvaries in shape too (variation in shape was addressed in another character41). State 1 appears either four times, in Heteromitobates alienus+H. harlequin, in H. anarchus, in Acutisoma, andin S. proximum+

(using DELTRAN), or twice, in Heteromitobates+ (with threereversions, in Mitogoniella, in H. discolor+H. inscriptus, and inH. albiscriptus) and in S. proximum+ (using ACCTRAN). State 2appears directly from state 0 as a synapomorphy of Goniosoma, and itchanges to state 1 in G. capixaba+ and in G. calcar rejecting our originalhypothesis of transformation. State 3 appears independently inG. macracanthum and in G. ensifer.

41 –Apophyses of leg IV – retrolateral apical apophysis of coxa crescent-like(prolateral curvature):

0 – absent1 – present

This character is a modification of state 1 of character 40. It appears a single

time in Acutisoma hamatum+A. longipes.

42 – Apophyses of leg IV – prolateral apophysis at the basal region oftrochanter [0-1;0-2-3]:

0 – sub-basal and large1 – basal2 – sub-basal and small3 – absent

2 01

This apophysis varies in size and position. It becomes basal twice, inSerracutisoma and in Acutisoma+Mitogoniella, and it becomes smallerin Heteromitobates harlequin, in Goniosoma dentipes+G. ensifer, and inG. apoain. The absent state is present only in the outgroup.

43–Apophyses of leg IV–prolateral apical apophysis of trochanter [0-2-1-3]:0 – absent1 – larger than basal apophysis, prolateral2 – smaller than basal apophysis, prolateral3 – larger than basal apophysis, dorsal–prolateral

0 2 1 3

Our original hypothesis of transformation that a larger apophysis wouldappear from a smaller one was rejected by our analysis. State 1 appearsin Serracutisoma+. The apophysis reduces (state 2) in G. vatrax+,disappearing as a reversion soon after, in G. macracanthum+. Areversion to state 1 also occurs in G. calcar+G. unicolor. State 3appears from state 1 in Serracutisoma. A smaller apophysis (state 2) ofthe same size of the sub-basal apophysis appears independently inHeteromitobates harlequin.

44 – Apophyses of leg IV – retrolateral basal apophysis of trochanter:0 – absent1 – present

This apophysis is curved upward and forward, cylindrical, appearing a singletime as a synapomorphy of Acutisoma.

45 –Apophyses of leg IV – retrolateral apical apophysis of trochanter [0-1-2]:0 – small, of same size of granules1 – much larger than granules, curved backward2 – much larger than granules, straight

State 1 appears twice in Serracutisoma: in S. spelaeum+ and in S. molle+,changing to state 2 in S. pseudovarium+ S. thalassinum.

46 – Apophyses of leg IV – apical dorsal apophyses of femur IV – size ofretrolateral apophysis:

0 – absent1 – small2 – large

Both apophyses of femur IV, retrolateral and prolateral, are present in allspecies of Goniosomatinae and are inserted on the apex of row1 and row6,respectively. Both apophyses vary in size and only the retrolateral varies indirection (character 47). The retrolateral apophysis is large in Acutisomahamatum+A. longipes and as a synapomorphy of Goniosoma. Moreover,it presents different morphology in those groups: it is straight inward inAcutisoma, and it maintains the same shape of the smallest one, beingcurved backward (character 47) in Goniosoma.

47 –Apophyses of leg IV – apical dorsal apophyses of femur IV – curvature ofretrolateral apophysis:

0 – backward1 – straight inward

It is a synapomorphy ofAcutisoma hamatum andA. longipes. It seems that theincrease of the apophysis (character 46) and its straightening forwardprobably occurred in a single event.

48 – Apophyses of leg IV – apical dorsal apophyses of femur IV – size ofprolateral apophysis:

0 – small1 – reduced



The reductionof this apophysis is congruentwith an increaseof the retrolateralapophysis (character 46) as a synapomorphy ofGoniosoma. However thereis a reversion in G. roridum+G. venustum, in addition to an independentappearance in Heteromitobates harlequin.

49 – Tarsi – tarsal process of leg IV – size [0-1-2]:0 – minute (Fig. 6)1 –medium, more than 1=4 and less than

1=2 the size of the tarsal claws (Fig. 8)2 – large, with the same size of the tarsal claws (Fig. 16)It increases (state 1) in Goniosoma (except G. carum), with an additional

increase in G. capixaba and polymorphically in G. venustum, withreversions in G. unicolor and in G. apoain.

50 – Tarsi – tarsal claws of legs III and IV:0 – smooth (Fig. 9)1 – pectinate (Fig. 7)Within Gonyleptidae, pectinate claws were considered to occur only in

Caelopyginae, when A.B. Kury (1994, unpublished note) first observedit in the Goniosomatinae Heteromitobates discolor (see discussion in‘Characters – main tendencies and evolution’). In the present study werecognised it in other species as a synapomorphy of the genusHeteromitobates; and it appears independently as a polymorphism inMitogoniella modesta.

51 – Dorsal scute – area II invading area I medially0 – absent1 – present (e.g. Fig. 224)Synapomorphy of Goniosomatinae.

52 – Dorsal scute – pair of main elevations of eye mound [0-1-2]:0 – small granules or tubercle1 – medium spine (<1mm)2 – large spine (>1mm)

Spines only appear at node of Pyatan+Goniosoma, increasing to ‘largespines’ twice: inG.carumand in (G.dentipes+G.ensifer)+with a reversionto tubercles inG. macracanthum. In addition, spines independently appearalso in Serracutisoma catarina.

53 – Dorsal scute – main elevations of eye mound – knobbed tubercle:0 – absent1 – present

This tubercle has a very different shape, with the normal conical base butwith a knob-like, cylindrical apex. It appears in (Heteromitobatesdiscolor+ H. inscriptus) + (H. albiscriptus+H. anarchus).

54 –Dorsal scute –main elevations of eye mound – elevated region betweentubercles:

0 – absent1 – present

This region is considered to be elevatedwhen it nearly reaches or it is above theeyelevel. In this case tubercles are very close to each other or are totallyfused (the latter is an autapomorphy of Mitogoniella unicornis). Thiscondition is present in Acutisoma+Mitogoniella with a reversion inM. indistincta+M. taquara, although the tubercles of these species arealso close to each other.

55 – Dorsal scute – pair of main elevations of area III:0 – tubercles1 – spines

Synapomorphy of Goniosomatinae, with 5 independent and autapomorphicreversions (Heteromitobates alienus, Serracutisoma spelaeum,S. fritzmuelleri, S. pseudovarium, and Goniosoma macracanthum).

56 –Dorsal scute – angles of posteriormargins of dorsal scute and free tergites(males):

0 – with granules or small tubercles1 – with strong tubercles or spines

In the past, the presence of spines was used as the only diagnostic character toseparate the two ‘main’ genera of the subfamily, Acutisoma andGoniosoma, from each other (see discussion in ‘Taxonomy’). Femaleshave anglesmore armed than those ofmales, but amongmales there ismorevariation. These spines appear in males in Serracutisomapseudovarium+ S. thalassinum and autapomorphically in S. proximumand in Heteromitobates anarchus.

57 – Epidermic pigmentation – dorsal scute of male [0-1;0-2]:0 – with a frame, with homogeneous pigmentation (Figs 193, 224)1 – totally homogeneous (without a frame) (Fig. 209)2 – with a frame, with fragmented pigmentation (Figs 212, 228)Epidermic pigmentation can be distributed homogeneously or as a frame (see‘Terminology’). In the this case, it can be fragmented or homogeneous inthe frame. The frame disappears, allowing an homogeneous pigmentationin Serracutisoma banhadoae+ and in Acutisoma coriaceum. Thefragmented pigmentation appears in Heteromitobates with a reversion inH. albiscriptus+H. anarchus. It is interesting to note that this reversionis followed by a general increase in pigmentation, since these speciesare black.




58 – Epidermic pigmentation – dorsal scute of female [0-1;0-2]:0 – with a frame, with homogeneous pigmentation (Figs 193, 224)1 – totally homogeneous (without a frame) (Fig. 209)2 – with a frame, with fragmented pigmentation (Figs 212, 228)There is a difference in coloration between males and females in some

species. In females, the fragmented pigmentation appears inHeteromitobates and in Serracutisoma fritzmuelleri+, with reversions inS.molle and inS. pseudovarium. The reversionobserved in themales of twospecies of Heteromitobates was not detected among females. InSerracutisoma, while males of S. banhadoae+ show ‘homogeneouspigmentation’, females of S. fritzmuelleri+ have ‘fragmented framepigmentation’ (with a reversions in S. molle and S. pseudovarium).Thus, this species-group of Serracutisoma has the greatest variation incolour between sexes. State 1 was not observed among females ofGoniosomatinae, being restricted to the outgroup.

59 – Epidermic pigmentation – light rings adjacent to the joint of tibia andmetatarsus of legs:

0 – absent1 – presentThese rings are formed by a lack of pigmentation on the apex of the tibia and

the base of the metatarsus. It appears in Serracutisoma proximum+, inGoniosoma varium+, and in Heteromitobates inscriptus.

60 – Coloration, dry-mark – on apex of coxa and base of femur IV:0 – absent1 – presentSynapomorphy of Goniosomatinae.

61 – Coloration, dry-mark – on trochanter IV [0-1-2-3]:0 – absent1 – on base2 – on base and apex, separated3 – from base to apexState 1 appears as a synapomorphyofGoniosomatinaewith an autapomorphic

reversion (to state 0, which is also observed in the outgroups) inSerracutisoma fritzmuelleri. Afterwards, state 2 appears inHeteromitobates+, changing to state 3 in (H. discolor+H. inscriptus)+(H. albiscriptus+H. anarchus) and inM. taquara. A reversion to state 1occurs in A. hamatum+A. longipes, although this condition ispolymorphic. In addition, state 3 independently appears in S. proximum.

62 – Coloration, dry-mark – serous layer reduced around granules of dorsalscute:

0 – absent1 – present (Fig. 207)This layer allows a grayish aspect to the animal. In S. banhadoae+, this layer

reduces around granules, forming dark circles. This character changestogether with character 57 (homogeneous pigmentation) and character 63(absence of dry-mark), giving a very homogeneous coloration to theseanimals.

63 – Coloration, dry-mark – on dorsal scute:0 – present1 – absentPresence of dry-mark will be detailed in the next 6 characters. This character

was used to code only once for the absence of a dry-mark instead of sixtimes, if we considered the next six characters. Anyway, no difference intopology was observed using one method or the other.

64 – Coloration, dry-mark – following grooves of areas [0-1-2]:0 – absent1 – as a frame (e.g. Fig. 190)2 – as a large cross (Fig. 197)

Dry-mark following grooves of areas of abdominal scute, i.e. as a frame, is asynapomorphy of Goniosomatinae. Reversions occur in Serracutisomaspelaeum+, in S. banhadoae+, in Acutisoma coriaceum, in Goniosomamacracanthum, and inG. calcar+G. unicolor. Dry-mark in cross appearsin G. dentipes.

65 – Coloration, dry-mark – around granules:0 – absent1 – present (Fig. 195)This type of dry-mark gives a punctate aspect to the body, and it appearsautapomorphically twice: in Mitogoniella indistincta and in Goniosomaroridum.

66 – Coloration, dry-mark – before the eye mound [0-1-2]:0 – absent1 – narrow stripe (Fig. 224)2 – as wide as the eye mound (Fig. 226)This region presents a longitudinal stripe of dry-mark in some species. State 1appears twice: in Heteromitobates and in Mitogoniella unicornis+ (with areversion in M. indistincta). State 2 appears, from state 1, in H. harlequinand H. anarchus, and, independently, directly from state 0, in Goniosomavarium+.

67 – Coloration, dry-mark – on the carapace, behind eye mound [non-additive]:

0 – absent1 – Y (Fig. 204)2 – empty triangle (Fig. 224)3 – full triangle (Fig. 230)4 – as a frame (Fig. 190)5 – round spot (Fig. 202)State 3 appears in Heteromitobates+, changing twice to state 4: in

H. alienus+H. harlequin and in M. modesta+; a single time to state 2,in H. albiscriptus+ H. anarchus; and a single time to state 5, inA. coriaceum. State 1 appears independently at the node ofSerracutisoma (using ACCTRAN), with 2 changes to ‘absent’ soonafter in S. spelaeum+ and in S. banhadoae+, with state 5 appearing inS. spelaeum. State 5 appears additionally twice, in Pyatan and inGoniosoma carum (using DELTRAN). State 4 appears in G. varium+,and state 5 appears independently in G. dentipes+G. ensifer.

68 – Coloration, dry-mark – round spot in between main elevations ofarea III [non-additive]:

0 – absent1 – present, not divided (Fig. 202)2 – present, divided longitudinally (Fig. 190)State 1 occurs in the group Pyatan+Goniosoma. It changes to state 2 inG. varium+ and it is lost in G. macracanthum and in G. apoain+, with areappearance of state 2 in G. calcar.

69 – Coloration, dry-mark – dots on the lateral margins of the dorsal scute[0-1-2]:

0 – absent1 – in the carapace (Fig. 202)2 – in the entire margin (Fig. 232)State 1 occurs inHeteromitobates+. A reversion occurs in the two more basalspecies of Heteromitobates (H. alienus+H. harlequin). State 2 appear inthe genera Mitogoniella+Acutisoma, and, independently in Goniosomaroridum. Actually, in this species, the whole dorsal scute is covered withdots (see character 65).

70 – Pedipalps – long pedipalp (longer than the dorsal scute) and robust:0 – absent1 – present (e.g. Figs 196 and 231)



Synapomorphy of Goniosomatinae. Although the subfamilies Caelopyginae,Progonyleptoidellinae and Sodreaninae have long pedipalps (see character86), those of Goniosomatinae were coded in a different manner since thereare shorter but more robust, and the structure seems to be different.

71 – Pedipalps – number of lateral setae of femur [0-1-2;0-3]:0 – 11 – 1 or 22 – 23 – absent

These setae are placed at the retrolateral subapical area of the femur and arevery conspicuous. The second seta (states 1 and 2) ismuch shorter andmoreapically placed. We decided to code a different state for polymorphismbecause: (1) polymorphic species present similar rates of femorawith 1 or 2setae; (2) the polymorphism may occur in the same animal, when bothpedipalps are observed; and (3) it appears in a great number of species. Thepolymorphic condition appears in Heteromitobates+ with autapomorphicreversions in H. alienus, in A. hamatum, and in M. indistincta. State 2appears in H. inscriptus and H. anarchus (although this species is knownonly from one specimen, the male holotype), and, independently, inGoniosoma varium+. State 1 also occurs in S. molle, in G. vatrax, and inG. calcar+G. unicolor, changing to state 2 in G. unicolor. State 3 appearsonly in the outgroup.

72 – Pedipalps – pattern of ornamentation of the ventral region of the femur[0-1-2-3]:

0 – 1 I1 – 1 I, but with i’s like tubercles2 – 2 I’s3 – 3 I’s

There is a great variation in the general ornamentation of ventral femur, andthis variation could only be addressed with the number of high elevations.State 1 is a synapomorphy of Goniosomatinae, in which it only persistsin Acutisoma and in Mitogoniella modesta. State 2 appears inHeteromitobates, in the rest of Mitogoniella, and in the large cladeSerracutisoma+. State 3 appears independently in H. discolor, inM. unicornis, in M. taquara, in G. carum, in G. varium, in G. capixaba,and in G. apoain+.

73 – Pedipalps – ventral–retrolateral region of patella [0-1-2]:0 – smooth1 – with a tubercle2 – with a spineThis elevation is the largest in patella and can be easily distinguish from other

granules of this segment. State 1 is a synapomorphy of Goniosomatinae,following the tendency in the subfamily to bear a strongly armed pedipalp.State 2only appears inGoniosomacarum and in (G.dentipes+G.ensifer)+,with reversions in G. unicolor and G. dentipes.

74 – Pedipalps – sub-basal ventral–prolateral region of tibia [0-1-2]:0 – smooth1 – with a tubercle2 – with a spineThis is the only elevation of tibia which varies within the subfamily. It can beused for species diagnosis and phylogenetic purposes, while the rest of thearmature is similar in all species. State 1 is a synapomorphy ofGoniosomatinae. State 2 appears synapomorphically in G. apoain+, andautapomorphically in five more species (three in Goniosoma).

Characters75–83are related togranulation (andare illustrated inFigs1–5) andshould be considered continuous. As already stated, state 0 and the laststate of continuous characters represent respectively the minimum andthe maximum values observed in the range in these characters. Theplesiomorphic state at the node of Goniosomatinae is discussed aftereach character.

75 – Granules – size of granules of areas and carapace:0 – minute1 – small2 – medium-sized3 – large4 – very largeState 1 is the plesiomorphic state in the subfamily. In general, there is atendency for a reduction of these granules in Pyatan+Goniosoma.Granules increase in Serracutisoma proximum+ and in Mitogoniellaindistincta+M. taquara (state 2), and in S. molle+ (state 3).

76 – Granules – size of granules at the posterior margin and free tergites:0 – absent (almost smooth segments)1 – minute2 – small3 – medium-sized4 – large5 – very largeIt shows an evolution pattern similar to that of the last character. However,a reduction to minute (state 1) occurs in Heteromitobates and inPyatan+Goniosoma.

77 – Granules – size of granules of lateral margins:0 – minute1 – small2 – medium-sized3 – largeIt also shows a evolution pattern similar to that of last characters, but with anadditional reduction in Serracutisoma guaricana+ S. spelaeum.

78 – Granules – density of granules at the posterior margin and free tergites:0 – low1 – medium2 – highState 1 is the plesiomorphic state in the subfamily. State 0 occurs inSerracutisoma and in Mitogoniella indistincta+M. taquara, and state 2occur in Pyatan+Goniosoma (with a reversion in G. varium+) and inHeteromitobates discolor+H. inscriptus.

79 – Granules – density of granules of lateral margin:0 – low1 – medium2 – highState 0 is the plesiomorphic state in the subfamily (using DELTRAN).State 1 occurs in Goniosoma (with a reversion in G. capixaba), and,with increasing density, in Heteromitobates and in Mitogoniella.




80 –Granules, venter – size of granules at the posteriormargin of the stigmaticarea and free sternites:

0 – absent (almost smooth segments)1 – minute2 – small3 – medium-sizedState 1 is the plesiomorphic state in the subfamily. States 2 and 3 occur

respectively in Serracutisoma banhadoae+ and in Acutisoma. A reductionof granules occur in Goniosoma calcar+G. unicolor.

81 – Granules, venter – size of granules of coxa IV:0 – minute or almost smooth1 – small2 – medium-sizedState 1 is the plesiomorphic state in the subfamily. Reductions occur in the

large group Serracutisoma+ (with a reversion in S. proximum+), inHeteromitobates alienus+H. harlequin and in Mitogoniella unicornis+

(with a reversion in M. indistincta). State 2 occur in Acutisomahamatum+A. longipes and in Serracutisomapseudovarium+ S. thalassinum.

82 – Granules, venter – density of granules at the posterior margin of thestigmatic area and free sternites:

0 – low (�2)1 – medium (>2, <4)2 – high (�4)State 1 is the plesiomorphic state in the subfamily. Increase in density of

granulesoccurs inHeteromitobates+, inGoniosoma (with 3 reversions) andagain inG. roridum+G. venustum. Reduction in density of granules occurstwice, autapomorphically, in G. dentipes and in Serracutisoma inerme.

83 – Granules, venter – density of granules of coxa IV:0 – low (�2)1 – medium (>2, <4)2 – high (�4)State 1 is the plesiomorphic state in the subfamily.Reduction in density occurs

in Serracutisoma fritzmuelleri+ and in Goniosoma dentipes+G. ensifer.Increase in density occurs in Goniosoma carum, in two species ofSerracutisoma, three of Heteromitobates, and two of Mitogoniella.

Characters 84–92 were used to address the outgroup relationship.

84 – Tarsus of pedipalp biconvex:0 – absent1 – presentState 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus

(Caelopyginae), Progonyleptoidellus striatus (Progonyleptoidellinae),Zortalia leprevosti (Sodreaninae), and Acrogonyleptes unus(Hernandariinae).

85 – Short ventral spines in two medial rows on tarsus of pedipalp:0 – absent1 – presentSame distribution as in the previous character.

86 – Elongate, cylindrical, and thin pedipalps:0 – absent1 – present2 – extremely elongatedState 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus

(Caelopyginae) and Progonyleptoidellus striatus(Progonyleptoidellinae).Zortalia leprevosti (Sodreaninae) presents state 2.

87 – Articles of tarsus IV semispherical:0 – absent1 – present

State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus(Caelopyginae) and Progonyleptoidellus striatus (Progonyleptoidellinae).

88 – Anal operculum with subcuticular white mark:0 – absent1 – presentState 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus(Caelopyginae).

89 – Setae of apical group of penis long and helicoid:0 – absent1 – presentState 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus

(Caelopyginae), Progonyleptoidellus striatus (Progonyleptoidellinae),Zortalia leprevosti (Sodreaninae), Acrogonyleptes unus(Hernandariinae), and Neosadocus variabilis and Sphaerobunus pulcher(Gonyleptinae).

90 – number of areas in the dorsal scute:0 – 41 – 3State 1 appears in Caelopygus elegans and Metarthrodes pulcherrimus(Caelopyginae), Progonyleptoidellus striatus (Progonyleptoidellinae),Zortalia leprevosti (Sodreaninae), Acrogonyleptes unus(Hernandariinae), Neosadocus variabilis and Sphaerobunus pulcher(Gonyleptinae), Roeweria virescens (Pachylinae), and Goniosomatinae.

91 – Dry-mark on free tergites:0 – absent1 – presentState 1 appears in Longiperna cancellata and Promitobates viridigranulatus(Mitobatinae).

92 – Leg IV of male much longer than leg II:0 – absent1 – presentState 1 appears in Longiperna cancellata and Promitobates viridigranulatus(Mitobatinae).

Characters 93–97 are related tomorphometric andmeristic characteristics andshould also be considered continuous. We followed the coding methodpresented by Thiele (1993). States represented by a single value indicate anautapomorphy of a species. Again, state 0 and the last state of continuouscharacters represent the minimum and the maximum values of the rangeobserved in these characters. For each species, we used 10 specimens(whenever possible) in order to obtain an averagenumber that couldbe usedas representative of the species.

93 – number of granules on areas:0 – 8 to 12.621 – 13.5 to 13.82 – 21.5 to 21.543 – 22.33 to 26.44 – 29 to 31.255 – 33.16 – 42.838 – 52.2 to 53.99 – 59 to 61

94 – distance between coxae of leg IV/length of dorsal scute rate:0 – 1.14 to 1.171 – 1.222 – 1.25 to 1.323 – 1.33 to 1.384 – 1.40 to 1.48



5 – 1.49 to 1.536 – 1.627 – 1.728 – 1.919 – 2.01

95 – number of articles of tarsus II:0 – 91 – 11.5 to 122 – 143 – 154 – 16.2 to 185 – 18.3 to 206 – 20.2 to 227 – 22.5 to 22.68 – 24.8 to 26.19 – 27 to 28.9

96 – length of leg II/length of dorsal scute rate:0 – 5.34 to 5.711 – 5.84 to 6.21

2 – 7.19 to 7.313 – 7.66 to 8.54 – 8.74 to 9.555 – 9.97 to 11.26 – 11.7 to 12.57 – 13.1 to 13.78 – 17.4 to 18.79 – 19.4 to 20.6

97 – length of femur IV/length of dorsal scute rate:0 – 1.251 – 1.37 to 1.512 – 1.62 to 1.823 – 1.86 to 2.124 – 2.19 to 2.505 – 2.52 to 2.896 – 3.09 to 3.417 – 3.8 to 3.958 – 4.08 to 4.579 – 5.64 to 5.81


Appendix 2. Index of taxonomic names

Index of generic names included in GoniosomatinaeAcutisoma page 592Acutisomella =Heteromitobates, syn. nov. – page 582Acutisomelloides =Serracutisoma syn. nov. – page 563Glyptogoniosoma =Goniosoma, syn. nov. – page 546Goniosomella =Lyogoniosoma=Goniosoma, syn. nov. – page 546Goniosoma page 545Heteromitobates page 582Itatiaya Roewer, 1928 (pre-occupied) replaced by Leitaoius =Acutisoma – page 592Leitaoius =Acutisoma – page 592Lyogoniosoma =Goniosoma, syn. nov. – page 546Metalyogoniosoma =Goniosoma, syn. nov. – page 546Mitogoniella page 588Progoniosoma =Goniosoma – page 545Pyatan gen. nov. – page 560Pygosomoides =Serracutisoma, syn. nov. – page 563Serracutisoma page 563Spelaeosoma =Serracutisoma, syn. nov. – page 563Xulapona =Lyogoniosoma=Goniosoma, syn. nov. – page 546

Index of specific names included in Goniosomatinaeacutangulum Acutisoma acutangulum – page 594albiscriptus Heteromitobates albiscriptus, comb. nov. – page 584alienus Heteromitobates alienus, sp. nov. – page 587anarchus Heteromitobates anarchus, sp. nov. – page 585apoain Goniosoma apoain, sp. nov. – page 556badium Mitogoniella badia – species inquirenda – page 592banhadoae Serracutisoma banhadoae, comb. nov. – page 572calcar Goniosoma calcar – page 558capixaba Goniosoma capixaba, sp. nov. – page 549carum Goniosoma carum, comb. nov. – page 559catarina Serracutisoma catarina, comb. nov. – page 577coriaceum Acutisoma coriaceum, sp. nov. – page 594cruciferum (Progoniosoma) =Goniosoma dentipes, syn. nov. – page 554cryptoleuca (Acutisomella) =Goniosoma xanthophthalmum=Goniosoma varium,

syn. nov. – page 548dentipes Goniosoma dentipes – page 554discolor Heteromitobates discolor – page 582ensifer Goniosoma ensifer – page 555fritzmuelleri Serracutisoma fritzmuelleri, sp. nov. – page 576geniculatum (Goniosoma) =Goniosoma venustum, syn. nov. – page 550grossum (Goniosoma) =Goniosoma dentipes – page 554guaricana Serracutisoma guaricana, sp. nov. – page 579guttulatus (Leitaoius) =Serracutisoma molle – page 573hamatum Acutisoma hamatum – page 597harlequin Heteromitobates harlequin, sp. nov. – page 586iguapensis (Leitaoius) =Serracutisoma proximum, syn. nov. – page 565indistincta Mitogoniella indistincta – page 588inerme Serracutisoma inerme, comb. nov. – page 578inscriptus Heteromitobates inscriptus, comb. nov. – page 583insperatum Pyatan insperatum, sp. nov. – page 560intermedia (Acutisomella) =Goniosoma varium, syn. nov. – page 548junceum (Goniosoma) =Goniosoma patruele=Goniosoma varium,

syn. nov. – page 548lepidum Goniosoma lepidum – species inquirenda – page 560longipes Acutisoma longipes, comb. nov. – page 601macracanthum Goniosoma macracanthum, comb. nov. – page 553marumbicola (Acutisoma) =Serracutisoma thalassinum, syn. nov. – page 575minense (Progoniosoma) =Goniosoma vatrax, syn. nov. – page 552modesta Mitogoniella modesta, comb. nov. – page 591molle Serracutisoma molle, comb. nov. – page 573monacanthum Goniosoma monacanthum – species inquirenda – page 560


monticola (Acutisoma) =Acutisoma longipes, syn. nov. – page 601mutila (Mitogoniella) =Acutisoma monticolum=Acutisoma longipes,

syn. nov. – page 601nitidissimus (Leitaoius) =Acutisoma monticolum=Acutisoma longipes,

syn. nov. – page 601obscurum Goniosoma obscurum – species inquirenda – page 560ornatus (Leitaoius) =Serracutisoma proximum – page 565patens (Acutisoma) =Serracutisoma thalassinum, syn. nov. – page 575patruele (Goniosoma) =Goniosoma varium, syn. nov. – page 548perditum (Glyptogoniosoma) =Goniosoma dentipes, syn. nov. – page 554perlata (Goniosomella) =Goniosoma vatrax, syn. nov. – page 552proximum Serracutisoma proximum – page 565pseudovarium Serracutisoma pseudovarium, sp. nov. – page 574roridum Goniosoma roridum – page 551spelaeum Serracutisoma spelaeum, comb. nov. – page 580taquara Mitogoniella taquara, sp. nov. – page 589tetrasetae (Progoniosoma) =Serracutisoma inerme, syn. nov. – page 578thalassinum Serracutisoma thalassinum, comb. nov. – page 575tijuca (Progoniosoma) =Goniosoma dentipes, syn. nov. – page 554unicolor Goniosoma unicolor, comb. nov. – page 557unicornis Mitogoniella unicornis, sp. nov. – page 590unum (Metalyogoniosoma) =Goniosoma varium, syn. nov. – page 548varium Goniosoma varium – page 548vatrax Goniosoma vatrax – page 552venustum Goniosoma venustum – page 550versicolor Goniosoma versicolor – species inquirenda – page 560viridifrons (Leitaoius) =Serracutisoma proximum, syn. nov. – page 565xanthomus (Leitaoius) =Acutisoma monticolum=Acutisoma longipes,

syn. nov. – page 601xanthophthalmum (Goniosoma) =Goniosoma varium, syn. nov. – page 548



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