2005. The Journal of Arachnology 33:456±467 TARSAL SCOPULA SIGNIFICANCE IN ISCHNOCOLINAE PHYLOGENETICS (ARANEAE, MYGALOMORPHAE, THERAPHOSIDAE) Jose Paulo Leite Guadanucci: Museu de Zoologia da Universidade de SaÄo Paulo, Instituto de BiocieÃncias da Universidade de SaÄo Paulo, Av. NazareÂ, 481, Ipiranga, CEP: 04263±000 SaÄo Paulo, SPÐBrazil. E-mail: [email protected] ABSTRACT. Tarsal scopula condition and carapace length were studied for eighteen Ischnocolinae species. For cladistic analysis a matrix of 20 terminals and 30 characters of representatives of Ischnocol- inae, Theraphosinae, Aviculariinae, Harpactirinae and Trichopelmatinae were analyzed using Nona 2.0 computer software. The matrix was analyzed in four different ways: 1. each tarsal scopula (legs I±IV) coded as separate characters; 2. one character with six ordered states; 3. one character with six independent states; 4. without tarsal scopula character. The ®rst two matrices result in one tree with the same indices (L 5 72; CI 5 0.54; RI 5 0.74) and topology: Part of Ischnocolinae is monophyletic (H. rondoni(S. longibulbi(I. algericus1Catumiri))) and the other representatives (Oligoxystre and Genus 1) form a distinct monophyletic group with Theraphosinae, Harpactirinae and Aviculariinae. There are no homoplasies in tarsal scopula evolution in the second cladogram. The other two cladograms show less resolution for the Ischnocolinae than the two ®rst cladorams. The tarsal scopula condition appears to have no relation to spider size (t 520.80433; P 5 0.438247) and should be used in phylogenetic analysis of Ischnocolinae because it provides information on the character variability within the subfamily. Keywords: Phylogeny, South America, cladistics The condition of the tarsal scopula has had subfamily based on a plesiomorphic character an important role in the systematics of the Is- state (divided tarsal scopula), the situation of chnocolinae Simon 1892. The scopula shows the group's systematics is very confusing. Ra- ontogenetic differentiation, being divided in ven (1985) considered Ischnocolinae a para- all juvenile Theraphosidae and becoming en- phyletic group that should have been revised tire in adults of some groups (Pocock 1897; at the generic level and grouped into mono- Gerschman de Pikelin & Schiapelli 1973; phyletic units. However, since the description PeÂrez-Miles, 1994). The condition of the tar- of the type-genus Ischnocolus Ausserer 1871, sal scopula has been considered a good taxo- only a few genera have been revised. Gersch- nomic tool and has already been used to di- man de Pikelin & Schiapelli (1973) revised agnose genera and species groups in the subfamily as a whole but of the 42 genera Theraphosidae. Its use in phylogenetics is included in this study, only 10 are in fact Is- questionable since, within the Theraphosinae, chnocolinae representatives. The remaining 12 the presence of a divided scopula is related to were subsequently synonymyzed or trans- small sized species (PeÂrez-Miles 1994). Char- ferred to other families and subfamilies. Rud- acterized as theraphosids with a divided tarsal loff (1997) revised the genus Holothele scopula (plesiomorphic state), Ischnocolinae Karsch 1879 but did not present a diagnosis is considered a paraphyletic group. Ischnocol- of the genus and its species; the identi®cation inae is the subfamily of Theraphosidae Tho- key does not include all the species and the rell 1869 that shows the broadest geographic structures are poorly illustrated. Smith (1990) distribution, with species occurring in north- published a taxonomic revision of European ern, central and eastern Africa, Seychelles, the and African Ischnocolinae and presented de- Middle-East, the Mediterranean region, cen- scriptions and diagnoses for all genera. tral and south Americas and the Antilles For over a century, the tarsal scopula state (Smith 1990; Rudloff 1997; Vol 2001). Con- ``divided by a longitudinal band of setae'' was sidering that Ischnocolinae was proposed as a used in Theraphosidae taxonomy (PeÂrez-Miles 456 GUADANUCCIÐTARSAL SCOPULA SIGNIFICANCE 457 1994). Gerschman de Pikelin & Schiapelli species used for tarsal scopulae condition and (1973), following Ausserer (1871), considered carapace length: the tarsal scopula condition an important tax- ? onomic character and stated that the divided Holothele rondoni (Lucas & BuÈcherl 1972): 1 , ApiaÂcas, Mato Grosso, Brazil (MZSP 18046); 1 tarsal scopula is present in all juvenile thera- ?, ApiaÂcas, Mato Grosso, Brazil (MZSP 18038); phosids. Juvenile Theraphosinae Thorell 1870 1 ?, ManicoreÂ, Amazonas, Brazil (MZSP 18990). have divided tarsal scopulae that become en- Sickius longibulbi Soares & Camargo 1948: 4 ?, tire in the adult stage, in Ischnocolinae the Itirapina, SaÄo Paulo, Brazil (MZSP 22756). scopulae remain divided into adulthood (Po- Genus 1: 3 ?, Colinas do Sul, Serra da Mesa, cock 1897; Gerschman de Pikelin & Schia- GoiaÂs, Brazil (MZSP 18992). pelli 1973; PeÂrez-Miles 1994). Although this Catumiri petropolium Guadanucci 2004: 1 ?, Pe- ontogenetic differentiation was detected, the troÂpolis, Rio de Janeiro, Brazil (IBSP 8596). ? divided condition continued to be used caus- Catumiri chicaoi Guadanucci 2004: 1 , Una, Ba- hia, Brazil (IBSP 9514). ing the inclusion of juvenile Theraphosinae Catumiri uruguayense Guadanucci 2004: 1 ?, Lav- within Ischnocolinae. This problem remained alleja, A guas Blancas, Uruguay (IBSP 9491). unresolved until Raven (1985) considered Is- Catumiri argentinenese (Mello-LeitaÄo 1941): 1 ?, chnocolinae a paraphyletic group that presents Jujuy, Yuto, El Pantanoso, Argentina (MACN poorly developed tarsal scopulae. The tarsal 6424). scopula as a phylogenetic character was used Genus 2, sp. 1: 3 ?, JaraguaÂ, GoiaÂs, Brazil (MPEG for the ®rst time by PeÂrez-Miles (1992) in a 1677) ? preliminary cladistic analysis of the subfamily Genus 2, sp. 2: 1 , Serra Norte, ParaÂ, Brazil Theraphosinae. In this paper he shows that the (MPEG 1678). Genus 2, sp. 3: 1 ?, Ilha Marajo Breves, ParaÂ, Bra- entire tarsal scopula is synapomorphic for zil (MPEG 1679). some genera of this subfamily. Later, PeÂrez- Genus 2, sp. 4: 1 ? (IBSP 11083), 1 ? (IBSP Miles (1994) discussed the value of the tarsal 11086), 1 ? (IBSP 11087), Pimenta Bueno, Ron- scopula in Theraphosinae systematics and doÃnia, Brazil; 1 ?, Mineiros, GoiaÂs, Brazil (IBSP concluded that the scopula condition is related 8070). to spider size, although some exceptions exist. Genus 2, sp. 5: 1 ?, Linhares, EspõÂrito Santo, Brazil Considering that the role of the tarsal scopula (IBSP 8654); 1 ?, Linhares, EspõÂrito Santo, Bra- ? in Theraphosidae systematic remains obscure, zil (IBSP 7987); 1 , Porto Seguro, Bahia, Brazil (IBSP 11084); 1 ?, IlheÂus, Bahia, Brazil (IBSP the goal of this study is to vary the use of the 11085). character ``tarsal scopula'' in a phylogenetic Oligoxystre new species 1: 1 ? (IBSP 9488), 1 ? analysis for Ischnocolinae and discuss the re- (IBSP 9489), 1 ? (IBSP 9486), 1 ? (IBSP 9484), sults. Central, Bahia, Brazil. METHODS Cladistic analysis.ÐThe cladistic analysis was carried out using Nona version 2.0 (Go- The material examined belongs to the fol- loboff 1993). Search strategy was mult* with lowing institutions: Instituto Butantan, SaÄo 100 replications. The data matrix included 20 Paulo (IBSP); American Museum of Natural terminal taxa and 30 characters and was con- History, New York (AMNH); Museo Argen- structed with NDE (Nexus Data Editor) ver- tino de Ciencias Naturales Bernardino Riva- sion 0.5.0 (Page 2001). The out-group was davia, Buenos Aires (MACN); Museo de la chosen based on the phylogenetic relation- Plata, La Plata (MLP); Museu de Zoologia, ships of Mygalomorphae presented by Golo- Universidade de SaÄo Paulo, SaÄo Paulo boff (1993). In order to avoid an excess of (MZSP); Zoological Museum University of missing entries, we preferred used a Bary- Copenhagen (ZMUC); Museu Paraense Emi- chelidae Simon 1889 rather than a Paratropi- lio Goeldi, BeleÂm (MPEG). didae Simon 1889 as the out group, since the Tarsal scopula condition was observed un- last family presents some incomparable char- der a stereomicroscope. Following PeÂrez- acters with Theraphosidae. Character polarity Miles (1994), a few isolated long and thin was read straight from the preferred clado- hairs in the tarsal scopula were not considered gram following Nixon & Carpenter (1993). divided. Carapace length was used to estimate Below is a list of species used in the cla- spider size. Below is a list of Ischnocolinae distic analysis: 458 THE JOURNAL OF ARACHNOLOGY Figures 1±2.ÐRelationship hypothesis between Ischnocolinae and other Theraphosidae groups. 1. (L 5 72; CI 5 0.54; RI 5 0.74). Tarsal scopula coded in four characters (22±25), one for each leg. 2. (L 5 72; CI 5 0.54; RI 5 0.74). Tarsal scopula coded as one character with six ordered states. GUADANUCCIÐTARSAL SCOPULA SIGNIFICANCE Table 1.ÐMatrix composed of 20 terminals and 31 characters. Characters 0 1 2 3 4 56789101112131415161718192021222324252627282930 Barychelinae 0 0 0 0 0 000000 0 0 0000Ð0000000000000 Reichlingia annae 0 1 0 1 0 000000 0 0 0210Ð1000000000010 Holothele rondoni 0 1 0 0 0 000010 0 0 0212 0 0010000000110 Sickius longibulbi 0 1 0 1 0 000010ÐÐ0210Ð0010000010110 Ischnothele algericus 1ÐÐ0Ð100011 1 0 0101 0 0010000010110 Catumiri petropo- lium 0 1 1 0 0 000010 0 0 1103 0 0000000010110 Catumiri chicaoi 1ÐÐ0Ð000010 ? ? 1103 1 0100120010113 Catumiri uruguay- ense 0 1 0 0 0 000010 0 0 1003 1 0100000010110 Catumiri argenti- nense 0 0 1 0