Zoosyst. Evol. 96 (2) 2020, 781-795 | DOI! 10.3897/zse.96.55210 gee BERLIN Sympolymnia, anew genus of Neotropical ant-like spider, with description of two new species and indirect evidence for transformational mimicry (Araneae, Salticidae, Simonellin1) Robert Perger!, Gonzalo D. Rubio? 1 Coleccion Boliviana de Fauna, La Paz, Bolivia 2 National Research Council of Argentina (CONICET), Experimental Station of Agriculture (EEA-INTA), R 14, Km 836, Cerro Azul, Misiones, Argentina http://zoobank.org/CCDFASBA-6E71-4D10-8808-7D39DBB92210 Corresponding author: Robert Perger (robertperger@hotmail.com) Academic editor: Danilo Harms # Received 8 June 2020 # Accepted 25 September 2020 Published 19 November 2020 Abstract Sympolymnia, a new genus of myrmecomorph jumping spider belonging to the tribe Simonellini Peckham, Peckham & Wheeler, 1889, is described. It comprises five species: the type species, Sympolymnia lucasi (Taczanowski, 1871), comb. nov., Sympolymnia lauretta (Peckham & Peckham, 1892), comb. nov., Svmpolymnia edwardsi (Cutler, 1985), comb. nov. and Sympolymnia shinahota sp. nov. and S. cutleri sp. nov. Sympolymnia lauretta (Peckham & Peckham, 1892) is recorded from Bolivia for the first time. Ontogenetic shifts of ant-resemblance are observed: Juveniles of S. cutleri sp. nov. and S. /auretta mimic black ants of the genus Crematogaster Lund, 1831, but those of S. shinahota sp. nov. most closely resemble Pseudomyrmex ethicus (Forel, 1911). Adults of S. cutleri sp. nov., S. /auretta and S. shinahota sp. nov. resemble the ant Camponotus sanctaefidei Dalla Torre, 1892 and orange adults of S. shinahota sp. nov. are putative mimics of Camponotus latangulus Roger, 1863. Key Words Bolivia, jumping spider, mimicry complex, myrmecomorph, polymorphism, South America Introduction Ant-resembling spiders have fascinated many natural- ists over centuries and are a promising group to study mimicry and evolution through natural selection (Nelson and Jackson 2012; Ceccarelli 2013). The morpholog- ical resemblance to ants (or myrmecomorphy) occurs in many spider families (Cushing 1997), but is partic- ularly common in jumping spiders (Maddison 2015; Maddison and Sztits 2019). The ant-like appearance has reached an extreme in Synemosyna Hentz, 1832 (Peckham and Peckham 1892; Oliveira 1988), a genus of the Neotropical tribe Simonellini Peckham, Peckham & Wheeler, 1889. Several species of Synemosyna were cited for their strong resemblance to members of the ant genus Pseudomyrmex Lund, 1831 (see Cushing 1997 for a review). The similarity was attributed to a conspicuous- ly elongated body with short legs and locomotory move- ment with short, rapid lunges (Oliveira 1988). Addition- ally, these forms have a deeply-constricted abdomen, which imitates the separation between the postpetiole and the gaster of Pseudomyrmex ants (Oliveira 1988). The genus Synemosyna was established by Hentz (1846) to accommodate S. formica Hentz, 1846. Three Species were subsequently described in this genus (Peckham and Peckham 1892, 1894). Galiano (1966) re- vised Synemosyna and synonymised Simonella Peckham & Peckham, 1885 with the former, which was accept- ed by subsequent authors (Cutler 1981, 1985, 1993; Cutler and Muller 1991; Makhan 2006) and presently Synemosyna comprises 20 species (WSC 2020). Cutler (1985) emphasised the presence of an ovate abdo- men in Synemosyna lauretta and S. lucasi (besides the large spermathecae and simple copulatory tubes) and Copyright Robert Perger, Gonzalo D. Rubio. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 782 Robert Perger & Gonzalo D. Rubio: Ant-like spiders of Bolivia: Sympolymnia | MM Southwest Amazon forest HM Bolivian Yungas forest (B Bolivian Tucuman forest | BB Chiquitano forest 0 100 200 300km Figure 1. Sampled locations and ecoregions according to the regionalisation by Navarro and Ferreira (2011), map produced with QGIS (version 2.14.3, http:/(www.qgis.org/en/site/). 1. Villa Teresa, La Paz Dept.; 2. Villa Tunari, Cochabamba Dept.; 3. Santa Maria la Antigua, Santa Cruz Dept.; 4. Cafetal, Buena Vista, Santa Cruz Dept.; 5. Cotoca, Santa Cruz Dept.; 6. La Guardia, Santa Cruz Dept.; 7. Bermejo, Santa Cruz Dept.; 8. Santa Rosa de la Mina, Santa Cruz Dept.; 9. Santiago de Chiquitos, Santa Cruz Dept.; 10. Arambulo, Tarija Dept. described Synemosyna edwardsi which also possessed these characters. Those three species differ from the elon- gated and slender species of Synemosyna that resemble Pseudomyrmex ants and are possibly mimics of ants of the genus Crematogaster Lund, 1831 (Cutler 1985). In the present contribution, we revise the taxonomy of these species and report the results of a survey in four forest ecoregions in Bolivia, including the description of two new species, a new country record and indirect evidence of transformational ant mimicry. Methods The sampling of spiders and ants was conducted in 10 locations in primary forest in four forest ecoregions of Bolivia (Amazon, Yungas, Chiquitano and Bolivian Tucuman forest, according to the ecoregion classification by Navarro and Ferreira 2011) (Figs 1, 2). Spiders and ants were collected with a beating tray. Photographs of live spiders and their habitats were taken with a Pana- sonic Lumix GX-80 system camera fitted with macro lenses. Colour was described from photos of live spec- imens. Spiders were euthanised with ethyl acetate and stored in 80% ethanol. Preserved specimens were ex- zse.pensoft.net amined under both dissecting microscopes and a com- pound microscope with reflected light and identified us- ing original and re-descriptions and keys (e.g. Peckham and Peckham 1892; Galiano 1966, 1967; Mello-Leitao 1933; Taczanowski 1871; Cutler 1985). Photos of pre- served specimens were taken with the Visionary Digital Passport II Imaging system at the Zoological Museum, University of Hamburg, Germany. Female genitalia were dissected as in Levi (1965), examined after digestion in ~15% sodium hydrox- ide (NaOH) solution and clarified in clove oil to ex- amine the internal structures. Temporary preparations were observed and photographed by GR using a Leica DM500 compound microscope and a Leica M60 ster- eomicroscope. Structures were sketched on incident light photograph models using a computer system for drawing and treatment of the image (Wacom digitiser tablet with GIMP, free software). All measurements, which were obtained with an ocular micrometer, are given in millimetres. Body length (BL) measurement refers to the distance from the anterior margin of the carapace to the posterior margin of the opisthosoma. Morphological terms and description formats follow the main recent studies on similar jumping spiders (Ruiz and Maddison 2015). Zoosyst. Evol. 96 (2) 2020, 781-795 Figure 2. Examples for sampled habitat types; A. Bolivian Yungas forest (forest edge) in Villa Teresa, La Paz Dept.; B. Southwest Amazon forest (forest edge) in Villa Tunari, Cochabamba Dept.; C. Cerrado vegetation in Chiquitano forest, Santiago de Chiquitos, Santa Cruz Dept.; D. Bolivian Tucuman forest, Arambulo, Tarija Dept. (this area is comparably dry in the dry season). Arachnological collections were abbreviated as fol- lows (curators in parenthesis): CBF Coleccion Boliviana de Fauna, La Paz, Boliv- ia (R. Perger). IBSI-Ara Instituto de Biologia Subtropical, Misiones, Argentina (G. Rubio). MNRJ Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil (A. Kury). PAS Museum and Institute of Zoology of the Pol- ish Academy of Sciences, Warsaw, Poland (W. Wawer). SMNK _ Staatliches Museum ftir Naturkunde Karl- sruhe, Germany (H. Hofer). Ecoregion distribution Distributional records without coordinates were geo- referenced via the gazetteers GeoLocator (http://tools. freeside.sk/geolocator/geolocator.html) and GeoNames (http://www.geonames.org/). The ecoregion affinities of the species at a continental level were investigated by visualising the coordinates and shapefiles of the re- gionalisation of Neotropical ecoregions by Olson et al. (2011) by using the geographic information system, QGIS (version 2.14.3, http://www.qgis.org/en/site/). For Bolivian ecoregions, the shapefile from Navarro and Ferreira (2011) was used, as the results were more consistent than those obtained from Olson et al. (2011). Geographic coordinates are shown in decimal degrees with reference datum WGS84 and elevation in metres above sea level (m a.s.1.). Ant mimicry In this study, an indirect, correlative method is employed to support mimicry, without studying the impact of re- ceiver responses on mimic fitness. While correlations do not imply causality, correlative approaches are useful for investigating putative cases of resemblance between taxa and extrapolating the consequences of mimicry beyond a single, well-studied population (de Jager and Anderson 2019). To illustrate adaptive divergence between mimetic and non-mimetic phenotypes, we identified derived traits in the mimic that likely evolved in association with spe- cies-specific models and receivers. For the analysis of ant resemblance, we considered all ants that were collected in the surveyed locations and were about the same body length of the spiders. The sim- ilarity was analysed, based on a qualitative, descriptive assessment of integument colour, shine, development (e.g. appressed, erected, short, long) and colour of hairs and shape of body parts (e.g. abdomen shape: fusiform or ovate; apically pointed or rounded). zse.pensoft.net 784 Robert Perger & Gonzalo D. Rubio: Ant-like spiders of Bolivia: Sympolymnia Table 1. Comparison of somatic and genitalic characters of Sympolymnia gen. nov., Synemosyna, Fluda and Erica (data for the latter two from Galiano (1971) and Peckham and Peckham (1892)). Character | Sympolymnia gen. nov. Erica constriction white patches abdomen ovate, about as long as ovate, about as carapace longer than carapace as or longer than carapace long as carapace abdomen absent constriction tibial apophysis of male palp absent or dorsoproximal | dorsomedian, distinct, in}absent or dorsoproximal, in males males also laterally of some spp. also laterally conspicuously elongated | at the most developed to Subrectangular to narrow tooth or spine | broad triangular tooth or broad triangular tooth to conspicuously elongated to obtuse edge t narrow tooth or spine opening(s) spermathecae large, lung-shaped not described kidney-shaped shaped Results S. lucasi (Yaczanowski, 1871), comb. nov., S. cutleri sp. Tribe Simonellini Peckham, Peckham & Wheeler, 1889 Sympolymnia gen. nov. http://zoobank.org/F30A 2797-7 111-46D4-83A D-095BSECCD6AF Type species. Janus lucasii Taczanowski, 1871 (by orig- inal designation) Diagnosis. This genus can be distinguished from the other four genera of Simonellini (Cy/iste/la Simon, 1901, Erica Peckham & Peckham, 1892, Fluda Peckham & Peckham, 1892 and Synemosyna) by the presence of two translucent white patches between the cephalic and thoracic areas (Figs 4 and 10B) and the presence of large, lung-shaped spermathecae (Fig. 6B, G). Additional characters to separate Sympolymnia gen. nov. from Erica, Fluda or Synemosyna are shown in Table 1. Composition. This new genus comprises the spe- cies Sympolymnia edwardsi (Cutler, 1985), comb. nov., S. lauretta (Peckham & Peckham, 1892), comb. nov., Key to adults of Sympolymnia nov. and S. shinahota sp. nov. Etymology. The specific epithet, Sympolymnia, 1s a com- bination of “sym”, meaning “with” in Greek and “Polymnia”, one of the nine Muses of Greek mythology, daughter of Zeus and Mnemosyne and the protector of the divine hymns and mimic arts. The gender of the name is feminine. Remarks. Cy/iste//a has a rounded, beetle-like habitus without constrictions and is the morphologically most dis- tinct group within this tribe. Sympolymnia gen. nov. 1s pos- sibly most related to Synemosyna (Table 1). This relation- ship is indicated by the presence of a constriction between the cephalic and thoracic parts and a single female genital opening. Additionally, in Synemosyna aurantiaca and Syn- emosyna formica, the embolus of the male palp is elongat- ed and winds around the retro-dorsal surface of the dorsum of the cymbium, similar to species of Sympolymnia gen. nov. However, all species of Sympolymnia gen. nov. are distinguished from Synemosyna by consistent differences in genitalic and somatic characters (Table 1), requiring an adequate generic treatment (see also Ruiz and Bustamante 2016; Kanesharatnam and Benjamin 2018; Rubio et al. 2020 for other, recent generic taxonomy in Salticidae). 1 CEBiaNe Pahl as WitleraseMICtaC© MAIS. a6 84,00 cn MlE nek abtecle 2 oe cca Bact Whats t eee ag eazace Ween gett thacc tect rere tek Te oe 2 - ceonalie part widel Aan ThOracl Giana Dec seds: hagas porn teesedele phe ee tba act ecg eee S. lucasi (Taczanowski, 1871), comb. nov. 2 ADCOMEHMOmEONSIMICLE (EIS. LSE) Ae 2c c8 Es Lh Ik SSS es Pe ee ba abil cane De ee Cann cl Mi cant eaee cere acne mgeee amas Laat aene Spore 3 - Anterior part of abdomen with a distinct dorsal constriction (Fig. 3C) ...........ccccceeeeceeeeeeeeeeeeeeeeeaeeeees S. shinahota sp. nov. 3 UCTS ae: need + Ai aca ee ss Con 2 pene E45. se Pion ee eh FO ERS | Ave RL ER | OSE ee Lae: 4 - Pek Na ct maar eS NAC Rae eT ny eA a Pt Te or HT ey a A 5 4 Tibial apophysis of male palp tooth-like ...............cceceeeeeeeeeeeeeeeeee ees S. lauretta (Peckham & Peckham, 1892), comb. nov. _ tibial ‘apophysis.of malejPalP ArCUATES SPINE... ....oc ec ancgendwclsne sontdecgeretqeenenarsbe content S. edwards/ (Cutler, 1985), comb. nov. 5 GO DUlate CCL AAG I COINSLE 540, a5, cen OM aeu Seal sts sere evce teenie ayo R hee dse Oek henge auk oo OR Uae faz ant vagh isha Agente danaba ken derck sea thaader sabecepeaiae oa 6 - CORBMIATORYTOUICTS WIKTIOUTSCOI IS); Fert. oo a3 ewan t eereectee Warm. I8 Pons wee aes 1T Th PUe Powys. ioe S. edwardsi (Cutler, 1985), comb. nov. 6 Epigyne opening small, wider than long (semicircular) (Fig. SF) ..........ccccceeeeeceeeeeeeeceeeee esse eeeeeeseeeeeeeeaees S. cutleri sp. nov. epigyne opening large, longer than wide (ovate) ..............:::::eeeeeeeees S. lauretta (Peckham & Peckham, 1892), comb. nov. zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 781-795 Species accounts Sympolymnia lucasi (Yaczanowski 1871), comb. nov. Fig. 6 Janus lucasii Taczanowski, 1871: 127, pl. 3, fig. 10. Simonella lucasi Peckham, Peckham & Wheeler, 1889: 254, pl. 12, fig. 9. Synemosyna lucasi Galiano, 1966: 366, figs 21, 22, 48 and 60. Synemosyna lucasi Galiano, 1967: 35, figs 29-32. Synemosyna lucasi Cutler, 1985: 90, figs 1, 2. Type deposit. Holotype 92 in PAS; Janus lucasii Taczanowski, 1871 (examined) (Fig. 6). Diagnosis. Thoracic part narrower than cephalic part in dorsal view, distinctly concave posteriorly in later- al view (Fig. 6A); abdomen at most with slight dorsal constriction; bulb of male palp about 65% of cymbium length, tibial apophysis in retro-lateral view with relative- ly broad base (see Galiano 1966 for line drawings), apex with spine-like process; epigyne opening small (width: ~ 0.07 mm), transversely elliptical. Geographical and ecological distribution (Fig. 7). The species was reported from Brazil (Galiano 1967) (the type location “Wassa” reported by Taczanowski [1871] refers to the river or the area of Uac¢a, which was formerly situated in French Guiana, now Brazil), French Guiana (Vedel et al. 2013), Colombia (Cutler and Miller 1991) and Peru (Cutler 1985). Synemosy- na lucasi appears to be associated with semi-deciduous or deciduous forests (Guajira-Barranquilla Xeric shrub, Sint Valley dry forest) and moist Amazon forests (To- cantins Pindaré moist forests, MarajO varzea and Ucay- ali moist forests). Cutler and Edwards (2002) recorded S. cf. lucasi from Trinidad Island (Lesser Antilles). The taxonomic status of these specimens remains to be de- termined. Sympolymnia lucasi was not observed in the present study area and there is no confirmed record of this species for Bolivia. Sympolymnia lauretta (Peckham & Peckham, 1892), comb. nov. Figs 3B, 4A, 8I-K Synemosyna lauretta Peckham & Peckham, 1892: 79, pl. 7, fig. 1. Synemosyna lauretta Simon, 1901: 509, figs 610, 611. Synemosyna melanura Mello-Leitaéo, 1917: 142. Simonella atrogaster Mello-Leitaéo, 1941: 256. Synemosyna lauretta Galiano, 1966: 362, figs 23, 24, 42, 43, 47, 58. Type deposit. The type of S. /auretta and its synonyms were destroyed in a recent fire (WSC 2020; A. Kury, unpublished). Diagnosis. Abdomen not constricted; bulb about 75% of cymbium length, tibial apophysis in retro-lateral view 785 with broad base, obtusely angulated in the middle, apex triangular, evenly tapering; opening of epigyne very large (width: ~ 0.21 mm), longitudinal-elliptical (see Galiano 1966 for line drawings). Variation. Smaller juveniles (BL < 3.1 mm) of S. lauretta had a shiny black body with pointed abdom- inal apex (Fig. 81). Larger juveniles (BL 3.2—3.9 mm) had a bright and pointed abdomen, but a black, matt carapace. The body of the adults (BL => 3.9 mm) was completely matt, and the abdomen apically more rounded (Figs 3B, 8K). Geographical and ecological distribution (Fig. 7). Sympolymnia lauretta was reported from Brazil (Peckham and Peckham 1892; Galiano 1966; Pod- gaiski et al. 2007; Rodrigues et al. 2016), Argentina (Galiano 1966) and Peru (Cutler 1985). This species is predominately found in seasonal forests south of 15°S. According to the biogeographic regionalisation by Ol- son et al. (2011), the records of this species refer to Araucaria moist forest, Cerrado, Alto Parana Atlantic forest and Serra do Mar coastal forests. In the pres- ent study, S. /auretta was observed in Tucuman (Tarija Department: Arambulo) and Chiquitano forests (Santa Cruz Department: Bermejo, Santa Rosa de la Mina, Santiago de Chiquitos). The records of the present study are the first for Bolivia. In almost all primary forest locations, S. Jauretta was obtained from isolated smaller trees and larger bushes overgrown with climb- ing plants (Fig. 2C, D), which are typically found in early successional forest and along forest edges. Sym- polymnia lauretta was not collected in closed, moist forests in the study area. Cutler and Edwards (2002) recorded S. cf. /auretta from Trinidad Island (Lesser Antilles). The taxonomic status of this population re- mains to be determined. Material examined. Botrvia * 1 4, 1 9; Santa Cruz Department, Bermejo; 18.1361°S, 63.6191°W; 8 Aug 2017; R. Perger leg., CBF + 1 4, 3 9, 1 juv.; Santa Rosa de la Mina; 16.5391°S, 62.4622°W; 9-13 Sep 2016; R. Perger leg.,; CBF * 1 4, 1 9, 1 juv.; Santiago de Chiq- uitos; 18.3225°S, 59.5763°W; 27-28 Dec 2017; R. Perg- er leg., CBF. Brazit * 2 4, 2; Santa Catarina State, Governador Celso Ramos; 27.314°S, 48.570°W; 2 Nov 2016; J.E. Baigorria leg.; IBSI-Ara 0909. ARGENTINA ¢ | 2; Misiones province, General Manuel Belgrano, Reserva Karadya; 25.859°S, 53.960°W; 21 Mar 2016; J.E. Baigorria leg.; IBSI-Ara 0610 * 1 9: Iguazu, Puer- to Iguazu; 25.592°S, 54.569°W; 14 Sep 2016, leg. J.E. Baigorria; IBSI-Ara 0788 « 1 9; Leandro N. Alem, EEA INTA Cerro Azul; 27.654°S, 55.435°W; 20 Oct 2017; G.D. Rubio leg.; IBSI-Ara 0985 * 2 3; same location and collector as for preceding; 22 Jan 2018; IBSI-Ara 0998 + 1 3; same location and collector as for preced- ing; 21 Nov 2017; IBSI-Ara 1013 * 1 3; same data as for preceding; IBSI-Ara 1036 * 1 4; same data as for preceding; IBSI-Ara 1041. zse.pensoft.net 786 Robert Perger & Gonzalo D. Rubio: Ant-like spiders of Bolivia: Sympolymnia rs Bi ie | Figure 3. Dorsal habitus of A. Svnemosyna myrmeciaeformis (Taczanowski, 1871), adult female; B. Sympolymnia lauretta (Peck- ham & Peckham, 1892), adult female; C. S. shinahota sp. nov. (IBSI-Ara 0726), female holotype. Scale bars: 1 cm. Sympolymnia shinahota sp. nov. http://zoobank.org/ABACO0D71-01A5-4E81-A 1 EB-C037E9888747 Figs 3C, 4B, SA-D, 8A-D, 9 Type deposit. Holotype 9°; Bo.ivia: Santa Cruz De- partment, Buena Vista, Cafetal; 17.4658°S, 63.6969°W; 342 ma.s.l.; beating tray sampling; 21 Jan 2016; R. Perg- er leg.; IBSI-Ara 0726. Paratypes 1 <4; Brazit: Amazo- nas State, Manaus, Taruma Mirim; 26 Feb 1988; SMNK- ARA 00364 * 2 2; Bottvia: Cochabamba Department, Villa Tunari; 16.9844°S, 65.4094°W; 335 m a.s.l.; beat- ing tray sampling; 6 Dec 2017; R. Perger leg.; IBSI-Ara 1033 * 1 9; same data as for preceding; CBF. Diagnosis. Sympolymnia shinahota sp. nov. is distin- guished from congeners by a distinct dorsal constriction zse.pensoft.net in the basal half of the abdomen (Figs 3C, 4B, 8A—C); male palp (Fig. 9C—F) with small cymbium, bulb about 80% of cymbium length, tibial apophysis in retro-later- al view narrow, apex evenly tapering (Fig. 9C, D); cop- ulatory opening in female very small and quadrangular (width: ~ 0.05 mm) (Fig. 5C). Comparisons. The male palp of S. shinahota sp. nov. (Fig. 9C—F) resembles that of S. /ucasi. However, the lat- ter is distinguished by a male palp tibial apophysis with a spine-like apex (evenly tapering in S. shinahota sp. nov.), the thoracic part narrower than the cephalic region in dorsal view and distinctly concave posteriorly in lateral view and the epigyne opening transversely elliptical. In the other two species with known males, the tibial apo- physis of the male palp is either broader (S. /auretta) or Zoosyst. Evol. 96 (2) 2020, 781-795 787 Figure 4. Lateral habitus of A. Sympolymnia lauretta (Peckham & Peckham, 1892), adult female; B.S. shinahota sp. nov. (IBSI-Ara 0726), female holotype. Scale bars: 1 cm. Arrows indicate the two light patches between the cephalic and thoracic areas (these patches are translucent white in live specimens). narrower (S. edwardsi) than in S. shinahota sp. nov. In all four congeners, the abdomen is not or only indistinctly constricted anteriorly. Description. Female holotype (Figs 3C, 4B, 5A-— D). Total length: 4.25 mm. Carapace length: 2.27 mm; width: 0.90 mm. Integument slightly shiny, orange yel- lowish with dark bands around eyes of last three rows, dorsum with sparse, simple, moderately long, whit- ish-yellow setae, denser and longer on anterior half of cephalic area and posterior half of abdomen; without pubescence. Carapace slender and elongated, cephal- ic portion a little longer than wide (width: 0.85 mm), as wide as widest thoracic part, smooth, marked con- striction (width 0.52 mm) between cephalic and tho- racic part, two translucent white areas at each side of constriction, separated by narrow black area (imitating part of femora I), constriction followed by globular, somewhat wrinkled knob, which is terminated behind by short pedicle which is more slender than the anterior constriction, evenly tapering when seen in lateral view and concave in dorsal view. Eyes arranged in four rows, quadrangle formed by the second and fourth rows of eyes wider than long, anterior eyes large, in contact, oc- cupying entire front of vertical inclination of face; sec- ond pair placed on back behind eyes of first pair, but a little further from each other than distance between ex- ternal borders of first ones, directed sideways; posterior eyes larger, separated by the same distance as those of second row, eyes of the third pair halfway between sec- ond and fourth. Chelicerae light brown, with three teeth on promargin and two on retromargin. Anterior half of sternum yellow and posterior half light brown. Abdo- men length: 1.80 mm; width: 1.10 mm, of same length as carapace, broader, commencing by short pedicle that appears to constitute prolongation of that of thoracic part, dorsally completely covered by scutum, integu- ment smooth, constriction in anterior portion. Slender and comparably long legs, in order 4, 3, 1, 2, third and fourth pair stouter, light brown. Epigyne (Fig. 5A—C): epigynal plate forming part of the epigastric sclerite, copulatory opening in very small and quadrangular (width: ~ 0.05 mm), copulatory ducts long, starting in a chamber, forming a spiral with one loop and entering the spermathecae posteriorly; spermatheca lung-shaped; copulatory ducts anterior to spermathecae. zse.pensoft.net 788 Robert Perger & Gonzalo D. Rubio: Ant-like spiders of Bolivia: Sympolymnia Figure 5. Genitalia and chelicerae of Sympolymnia spp.: S. shinahota sp. nov., female holotype (IBSI-Ara 0726): A. Epigyne in ventral view, cleared (blue line course of copulatory ducts); B. Spermathecae, dorsal view; C. Epigyne, ventral view; D. Chelicera in anterior view. Sympolymnia cutleri sp. nov., female holotype (IBSI-Ara 1072): E. Chelicera in anterior view; F. Epigyne in ventral view, cleared (blue line course of copulatory ducts); G. Spermathecae, dorsal view. LE tite £ tC il Tittle AAsSIA~ Guy ane 14. K: Sold iei B Aetm. HT. Taezanotyben, C Figure 6. Holotype of Sympolymnia lucasi (Taczanowski, 1871): A. Habitus dorsal and lateral views; B. Spermatheca and copula- tory duct; C. Data label. zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 781-795 MS Ss. lucasi ecoregions lS S. cutleri sp. nov. ecoregion GS S. shinahota sp. nov. ecoregions [9 S. lauretta ecoregions Figure 7. Ecoregion distribution: * |§ Sympolymnia lucasi (Taczanowski, 1871); ™@ @ S. /auretta (Peckham & Peckham, 1892). Male paratype (Fig. 9). Total length: 3.81 mm. Cara- pace length: 1.94 mm; width: 0.74 mm. Cephalic portion a little longer than wide (width: 0.71 mm), thoracic con- striction 0.50 mm wide. General form of carapace, chel- icerae and sternum as female, colouration dark brown. Abdomen length: 1.70 mm; width: 0.83 mm, covered dorsally and completely by a scutum, with a conspicu- ous constriction in the anterior portion; colouration dark brown. Palp (Fig. 9C—F): small cymbium, bulb about 80% of cymbium length, irregularly shaped spherical, embolus long, arising at the basal side of the bulb, with- out complete circular revolution and is accommodated on a slight retro-lateral concavity of cymbium, lacking pars pendula, tibial apophysis tooth-like, narrow, moderately tapering, tip directed forward. Variation. Juvenile females had a shiny, dark brown-blackish body surface with dark orange to light brown cephalic part and a pointed abdomen (Fig. 8A). The constriction in the proximal half of the abdomen was marked by a relatively broad, light transverse band. Adult females had a matt, blackish body surface with a dark or- ange or completely black cephalic part, the bright band in the abdominal constriction indistinct or absent (Fig. 8B, C). The holotype (BL 4.25 mm), collected in Cafetal, Buena Vista (Santa Cruz Department), had a matt, or- ange yellowish body except for some darker patches 789 500 1000 1500 km S. cutleri sp. nov.; S. shinahota sp. nov.; around the second and third eyes and the abdominal apex (Figs 3C, 4B). The abdominal constriction becomes less pronounced with increasing body length. Etymology. The specific epithet, shinahota, refers to a place with many ants or an ant nest in the Yuracaré language (Querejazu 2005), spoken by the Yuracaré people living along the Chapare River in the Amazon Basin of Bolivia. Geographical and ecological distribution (Fig. 7). Sympolymnia shinahota sp. nov. is known from Brazil, Amazonas State, Manaus and from the Bolivian Depart- ments of Cochabamba (Villa Tunari) and Santa Cruz (Buena Vista). The collection locations of this species were situated in moist Amazonas forest regions, includ- ing Uatuma-Trombetas moist forest (Amazonas State, Manaus) and pre-Andean Southwest Amazon rainforest (Villa Tunari, Cochabamba Dept. and Buena Vista, Santa Cruz Dept.). However, in all areas that were surveyed in the present study, S. shinahota sp. nov. was exclusively collected in early successional forests in large tree-fall gaps or secondary forest from isolated, small trees that were densely overgrown with climbing plants, several metres away from the edge of primary forest. Remarks. Photographs of two individuals (Maddison 2018) suggest that this species also occurs in the Amazon zse.pensoft.net 790 Robert Perger & Gonzalo D. Rubio: Ant-like spiders of Bolivia: Sympolymnia Figure 8. Live habitus of Sympolymnia spp. and potential ant models. Please note the ontogenetic shift of shine and abdomen shape in the spiders. Svmpolymnia shinahota sp. nov.: A. Juvenile female, Villa Tunari, Cochabamba Dept. (please note the shiny, pointed abdomen); B, C. Adult females, same location; D. Holotype female, Buena Vista, Santa Cruz Dept.; E. Pseudomyrmex ethicus, Villa Tunari, Cochabamba Dept.; F. Crematogaster sp., Villa Tunari, Cochabamba Dept.; G. Camponotus sanctaefidei, La Guardia, Santa Cruz Dept.; H. C. /atangulus, Buena Vista, Santa Cruz Dept.; Svmpolymnia lauretta: 1. Juvenile, Bermejo, Santa Cruz Dept. (please note the shiny and pointed abdomen); J. Sub-adult male, Santiago de Chiquitos, Santa Cruz Dept.; K. Adult female, Santa Rosa de la Mina, Santa Cruz Dept.; L. Sympolymnia cutleri sp. nov.: adult female, Chairo, La Paz Dept. rainforest in Ecuador (Yasuni National Park). Light variants were also reported in S. /auretta and S. lucasi (Galiano 1967). However, these reports refer to speci- mens that were stored in alcohol or formaldehyde for a longer time and may have faded due to the preservative (Galiano 1967). This study reports for the first time an in situ observation of an orange variant of Sympolymnia. Sympolymnia cutleri sp. nov. http://zoobank.org/8A91D161-DAOF-419C-A 121-9ES50A59BA75F Figs 5F, G, 8L, 10 Type deposit. Holotype 2°; Bo.tvia: La Paz Depart- ment, Nor Yungas Province, Villa Teresa; 16.2019°S, 67.8294°W; 1340 ma.s.l.; beating tray sampling; 17 Jan 2018; R. Perger leg.; IBSI-Ara 1072. Paratypes 2 2; same data as for preceding; IBSI-Ara 1024 * 3 9, | juv.; same data as for preceding; CBF « | 9; La Paz Department, zse.pensoft.net Nor- Yungas province, Coroico; Aug 1993; Metzner leg.; SMNK: 1358. Diagnosis. Sympolymnia_ cutleri sp. nov. and S. /auretta are indistinguishable in their somatic charac- ters. Sympolymnia cutleri sp. nov. can be separated from all congeners by an epigyne with small, semi-circular opening (Figs 5F, G, 10D, E) (opening of epigyne very large and longitudinal-elliptical in S. /auretta). Description. Female holotype. Total length: 4.20 mm. Carapace length: 2.10 mm; width: 0.81 mm. Integument slightly shiny, dark brown, blackish, dorsum with sparse, simple, moderately long, whitish setae, denser and longer on anterior half of cephalic area and posterior half of ab- domen. Carapace slender and elongated, cephalic portion as long as wide (width: 0.81 mm), as wide as widest tho- racic part, smooth, marked constriction (width 0.62 mm) between cephalic and thoracic part, two translucent are- as at each side of constriction, separated by narrow dark Zoosyst. Evol. 96 (2) 2020, 781-795 19.1 0,4 mm 0,3 mm 0.3mm 03mm ~~ # Figure 9. Sympolymnia shinahota sp. nov., paratype male (SMNK-ARA: 00364): A. Dorsal view; B. Lateral view; palp; C. Retrolateral; D. Retroventral; E. Ventral; F. Prolateral (photographs by Hubert Hofer, SMNK, Germany). area (imitating part of femora II), constriction followed by globular, somewhat wrinkled knob, which is terminated behind by short pedicle which is more slender than the anterior constriction, evenly tapering when seen in lateral view and concave in dorsal view. Eyes arranged in four rows, quadrangle formed by the second and fourth rows of eyes wider than long, anterior eyes large, in contact, occupying entire front of vertical inclination of face; sec- ond pair placed on back behind eyes of first pair, but a little further from each other than distance between exter- nal borders of first ones, directed sideways; posterior eyes larger, separated by same distance as those of second row, eyes of the third pair halfway between second and fourth. Chelicerae light brown, with five teeth on promargin and three on retromargin. Anterior half of sternum pale yellow and posterior part dark brown, blackish. Abdomen length: 2.00 mm; width: 1.10 mm, of same length as carapace, broader, commencing by a short pedicle that appears to constitute a prolongation of that of thoracic part, covered dorsally and completely by a scutum, without constriction, smooth. Slender and comparably long legs, in the order 4, 3, 1, 2, first pair pale (leg I with dark longitudinal bands on anterior and posterior sides), third and fourth pair stouter, dark brown. Epigyne (Fig. 5F, G): epigynal plate forming part of the epigastric sclerite, with a small semi-circular opening, wider than long (width: ~ 0.17 mm); copulatory ducts starting in a small chamber (hard to see), forming a spiral and entering the spermathecae posteriorly; sper- matheca lung-shaped; copulatory duct anterior to sper- matheca, between them. Male unknown. Variation. One immature (shiny surface, pointed ab- domen) and five female adults (matt surface, rounded abdomen) (Fig. 8L) were collected, showing the same ontogenetic shift in integument shine and abdomen shape as observed in the other congeners. Etymology. The specific epithet, cutleri, is a patronym in honour of Bruce E. Cutler in recognition of his contri- butions to the taxonomy of Simonellini. Geographical and ecological distribution (Fig. 7). Sympolymnia cutleri sp. nov. is exclusively known from the type location in Bolivian Yungas forest. Discussion Species richness and ecoregion distribution This study provides the first records for the tribe Simonellini for the Departments of Cochabamba, Santa Cruz and Tarija and the ecoregions Chiquitano forest and Bolivian Tucuman forest. The presence of previously- unrecorded or unknown species was not surprising, as the Bolivian spider fauna is little known (Cutler 1981; Perger and Perger 2017; Perger and Rubio 2018, 2020), which is consistent with the sampling effort reported for other invertebrate groups (Perger and Santos-Silva 2010, 2018; Perger and Grossi 2013; Perger 2015; Perger and Guerra 2016). With three species, Bolivia and Brazil have the highest species richness of Sympolymnia amongst all of the Neotropical countries. The distribution of Sympolymnia spp. appears to cor- respond to the delineation of humid Andes forest, Ama- zon rain forests and sub-humid, semi-deciduous forests (including Chiquitano forest, Bolivian Tucuman forest, Cerrado and Atlantic forest) (Fig. 7). This pattern sug- zse.pensoft.net 792 Robert Perger & Gonzalo D. Rubio: Ant-like spiders of Bolivia: Sympolymnia E 0,1 mm i Figure 10. Sympolymnia cutleri sp. nov., paratype female (SMNK-ARA: 1358): A, B. Carapace; C. Abdomen lateral; D. Epigyne; E. Spermatheca and copulatory duct (photographs by Hubert Hofer, SMNK, Germany). gests that the significant shift in seasonality and related conditions triggered diversification in Sympolymnia. For example, the Chiquitano forest ecoregion is distinguished by a pronounced seasonality and lower annual precipi- tation than Amazon forests (Ibisch and Mérida 2003). Accordingly, an essential factor contributing to the high species richness in Bolivia might be the meeting of sev- eral biogeographic realms and their corresponding fau- nal elements. High diversity in insect groups (Pearson et al. 1999: Kitching et al. 2001; Wappes et al. 2011) and Castianeirinae spiders (Perger and Perger 2017) in Boliv- ia was explained by high species turnover between a large number of ecoregions. Nevertheless, the comparably high richness of Bolivian species of Sympolymnia is likely partly attributed to low sampling efforts in Brazil, Ecua- dor, Peru, Venezuela and Colombia. Further taxonomic work and sampling campaigns are needed to clarify the species richness patterns of Sympolymnia. Ant mimicry The consideration of all ant species that were collected in the microhabitats of Sympolymnia species allowed a ro- bust preliminary assessment of potential ant models. We identified at least four potential ant model species in three zse.pensoft.net genera and subfamilies (Table 2). These species were amongst the most abundant ants in all microhabitats in which those resembling Sympolymnia spp. were collected (a subjective estimation). Strong indirect support for ant mimicry was provided by: 1) species-specific similarity involving morphological aspects (Table 2, Fig. 8); 2) sympatry: both spiders and ants were found in the same microhabitats; and 3) mimic less abundant than the ant model. Alternative processes, such as convergent evolution, exploitation of perceptual bias, developmental or phylogenetic constraints, spatial autocorrelation, crypsis, or random matching (de Jager and Anderson 2019), were unlikely to be responsible for the resemblance between ants and spiders in the present case. All suggested ant models are known for being well de- fended. The stinger in Crematogaster species is well de- veloped, but the venom is applied topically by wiping on a victim instead of injecting it inside the body (Buren 1959). Camponotus species do not possess stingers, but are well protected by their powerful mandibles and the release of defensive chemical compounds (Fisher and Cover 2007). Mimicry complexes involving adult polymorphic jump- ing spiders were described for Synemosyna aurantiaca (Mello-Leit&éo, 1917) (reviewed by Cushing 1997) and Myrmarachne in Australia (Pekar et al. 2017). Gilbert (2005) proposed that polymorphism in a mimetic species Zoosyst. Evol. 96 (2) 2020, 781-795 F983 Table 2. Possible co-occurring ant models for Bolivian species of Sympolymnia and shared characters: S. /auretta (1) BL < 3.1 mm; (II) BL 3.65-4.05 mm; S. cutleri sp. nov. (1) BL < 3.55 mm; (II) BL 4.02-4.32 mm; S. shinahota sp. nov. (1) BL < 3.5 mm; (II) BL 3.81—4.05 mm, black form; (III) BL 4.25 mm, orange form. +, co-occurring, resembling; *, co-occurring, not resembling (there were no differences in co-occurrence patterns between locations). Potential ant model nov. Shared characters I II I I Crematogaster sp. + x + x + (Myrmicinae) (BL 2.7-3.6 mm) % + x + S. lauretta| S. cutleri |S. shinahota sp. Sp. nov. I Il x integument black, shiny, without pubescence, abdominal apex pointed x integument relatively dull, blackish, single setae, without pubescence, abdominal apex rounded x cephalic part orangish, remaining integument shiny black; without pubescence, abdomen dorsally constricted and apically pointed Camponotus sanctaefidei aay (Formicinae) (BL 3.4-3.8 mm) Pseudomyrmex ethicus + (Pseudomyrmecinae) Camponotus latangulus x (Formicinae) (BL 3.65-5 mm) increases the protection against predation because it re- duces the number of mimics per model. When a morph increases too much in frequency within the habitat, it may lose its mimetic protection and be exposed to greater predation (Gilbert 2005). The present study is the first describing putative mim- icry complexes involving ontogenetic shifts of ant resem- blance (transformational mimicry) in Simonellini. It may be hypothesised that transformational mimicry common- ly occurs amongst ant-mimicking spiders, assuming that juvenile spiders face at least the same level of predation pressure as adult spiders and considering that ant models have castes occupying a discrete mode in the size-fre- quency distribution. However, the genus Sympolymnia appears to include species with and without transforma- tional mimicry, possibly depending on the maximum at- tainable body size or the presence of suitable ant models. The reported individuals of S. /ucasi (female holotype 3.66 mm; male allotype 3.3 mm, juveniles unknown) have about the same body size as Crematogaster-resembling juveniles of S. /auretta and S. cutleri sp. nov. Considering the small size, it is likely that S. /ucasi lacks transforma- tional mimicry and mimics exclusively Crematogaster ants. In the Bolivian species of Sympolymnia, larger body size and the lack of large Crematogaster species likely favoured the resemblance to different ant models. Transformational mimicry involving Crematogaster and Camponotus ants was proposed for myrmecomorphic sac spiders of the genus Myrmecium Latreille, 1824 in the Amazon forest (Oliveira 1988). Amongst ant-mimick- ing jumping spiders, transformational mimicry was ob- served in several African (Edmunds 1978) and Australi- an (Ceccarelli 2010) species of Myrmarachne MacLeay, 1839 and may occur in the majority of Myrmarachne species (Wanless 1978). Amongst Neotropical jumping spiders, transformational mimicry is only known for Zuniga magna Peckham & Peckham, 1892. The juveniles of Z. magna resemble Camponotus ants, while the adult males imitate Pseudomyrmex gracilis (Fabricius, 1804) and the females Neoponera villosa (Fabricius, 1804) (Oliveira 1988). The limited knowledge of transforma- tional mimicry in myrmecomorphic spiders, in general, is + integument relatively dull, orange, without pubescence, single setae, abdominal apex rounded likely explained by the fact that many species are known only from adult individuals. Conclusion and outlook The presence of Sympolymnia throughout South America and the sympatry with Synemosyna species in many loca- tions (e.g. Galiano 1966; Podgaiski et al. 2007; Rodrigues et al. 2016) suggests an early split and the subsequent ra- diation of the two lineages, accompanied by the selection for Pseudomyrmecinae-resemblance in adult Synemosyna and Myrmicinae-, Formicinae- and Dolichoderinae-re- semblance in adult Sympolymnia. In addition to similarity, sympatry of models and mimics is considered a critical factor in the selection for mimicry (de Jager and Anderson 2019). A more detailed analysis of co- occurrence patterns is needed to investigate the importance of polychromatism, transformational mimicry and mimicry complexes for ant mimicry in Sympolymnia. Additionally, the influence of macro-environmental conditions should be considered, as the distributional patterns of Sympolymnia Species suggest that such factors may have contributed to generating divergent selection pressures in this genus. Acknowledgements We extend many thanks to Wioletta Wawer (Museum and Institute of Zoology of the Polish Academy of Sciences, Warsaw, Poland) and Hubert Hofer (Staatliches Museum fur Naturkunde Karlsruhe, Germany) for providing pho- tographs of essential specimens. We are very grateful to Adriano B. Kury (Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil) for providing information on the status of type material. Bruce E. 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