Research Article |
Corresponding author: Robert Perger ( robertperger@hotmail.com ) Academic editor: Danilo Harms
© 2020 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.
Citation:
Perger R, Rubio GD (2020) Sympolymnia, a new genus of Neotropical ant-like spider, with description of two new species and indirect evidence for transformational mimicry (Araneae, Salticidae, Simonellini). Zoosystematics and Evolution 96(2): 781-795. https://doi.org/10.3897/zse.96.55210
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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., Sympolymnia 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. lauretta 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. lauretta 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.
Bolivia, jumping spider, mimicry complex, myrmecomorph, polymorphism, South America
Ant-resembling spiders have fascinated many naturalists over centuries and are a promising group to study mimicry and evolution through natural selection (
The genus Synemosyna was established by
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
Sampled locations and ecoregions according to the regionalisation by
Female genitalia were dissected as in
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 follows (curators in parenthesis):
IBSI-Ara Instituto de Biología Subtropical, Misiones, Argentina (G. Rubio).
Distributional records without coordinates were georeferenced 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 regionalisation of Neotropical ecoregions by
In this study, an indirect, correlative method is employed to support mimicry, without studying the impact of receiver 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 (
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 similarity 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).
Tribe Simonellini Peckham, Peckham & Wheeler, 1889
Janus lucasii Taczanowski, 1871 (by original designation)
This genus can be distinguished from the other four genera of Simonellini (Cylistella 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
Comparison of somatic and genitalic characters of Sympolymnia gen. nov., Synemosyna, Fluda and Erica (data for the latter two from
Character | Sympolymnia gen. nov. | Synemosyna | Fluda | Erica |
---|---|---|---|---|
carapace lateral constriction | present | present | absent | absent |
carapace lateral white patches | 2 | 0–1 | 0 | 1 |
abdomen | ovate, about as long as carapace | fusiform, elongated, longer than carapace | ovate to fusiform, about as long as or longer than carapace | ovate, about as long as carapace |
abdomen constriction | absent or dorsoproximal | dorsomedian, distinct, in males also laterally | absent or dorsoproximal, in males of some spp. also laterally | absent |
tibial apophysis of male palp | conspicuously elongated to narrow tooth or spine | at the most developed to broad triangular tooth or subrectangular | broad triangular tooth to conspicuously elongated to narrow tooth or spine | obtuse edge |
epigyne opening(s) | 1 | 1 | 2–3 | 2 |
spermathecae | large, lung-shaped | small, globular, pear- or kidney-shaped | small, globular, pear- or kidney-shaped | not described |
This new genus comprises the species Sympolymnia edwardsi (Cutler, 1985), comb. nov., S. lauretta (Peckham & Peckham, 1892), comb. nov., S. lucasi (Taczanowski, 1871), comb. nov., S. cutleri sp. nov. and S. shinahota sp. nov.
The specific epithet, Sympolymnia, is a combination 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.
Cylistella has a rounded, beetle-like habitus without constrictions and is the morphologically most distinct group within this tribe. Sympolymnia gen. nov. is possibly most related to Synemosyna (Table
1 | Cephalic part as wide as thoracic part | 2 |
– | cephalic part wider than thoracic part | S. lucasi (Taczanowski, 1871), comb. nov. |
2 | Abdomen not constricted (Fig. |
3 |
– | Anterior part of abdomen with a distinct dorsal constriction (Fig. |
S. shinahota sp. nov. |
3 | Males | 4 |
– | Females | 5 |
4 | Tibial apophysis of male palp tooth-like | S. lauretta (Peckham & Peckham, 1892), comb. nov. |
– | tibial apophysis of male palp arcuate spine | S. edwardsi (Cutler, 1985), comb. nov. |
5 | Copulatory ducts with coils | 6 |
– | copulatory ducts without coils | S. edwardsi (Cutler, 1985), comb. nov. |
6 | Epigyne opening small, wider than long (semicircular) (Fig. |
S. cutleri sp. nov. |
– | epigyne opening large, longer than wide (ovate) | S. lauretta (Peckham & Peckham, 1892), comb. nov. |
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.
Holotype ♀ in
Thoracic part narrower than cephalic part in dorsal view, distinctly concave posteriorly in lateral view (Fig.
The species was reported from Brazil (
Synemosyna lauretta Peckham & Peckham, 1892: 79, pl. 7, fig. 1.
Synemosyna lauretta Simon, 1901: 509, figs 610, 611.
Synemosyna melanura Mello-Leitão, 1917: 142.
Simonella atrogaster Mello-Leitão, 1941: 256.
Synemosyna lauretta Galiano, 1966: 362, figs 23, 24, 42, 43, 47, 58.
The type of S. lauretta and its synonyms were destroyed in a recent fire (
Abdomen not constricted; bulb about 75% of cymbium length, tibial apophysis in retro-lateral view with broad base, obtusely angulated in the middle, apex triangular, evenly tapering; opening of epigyne very large (width: ~ 0.21 mm), longitudinal-elliptical (see
Smaller juveniles (BL ≤ 3.1 mm) of S. lauretta had a shiny black body with pointed abdominal apex (Fig.
Sympolymnia lauretta was reported from Brazil (
Bolivia • 1 ♂, 1 ♀; Santa Cruz Department, Bermejo; 18.1361°S, 63.6191°W; 8 Aug 2017; R. Perger leg.;
Holotype
♀; Bolivia: Santa Cruz Department, Buena Vista, Cafetal; 17.4658°S, 63.6969°W; 342 m a.s.l.; beating tray sampling; 21 Jan 2016; R. Perger leg.; IBSI-Ara 0726. Paratypes 1 ♂; Brazil: Amazonas State, Manaus, Taruma Mirím; 26 Feb 1988;
Sympolymnia shinahota sp. nov. is distinguished from congeners by a distinct dorsal constriction in the basal half of the abdomen (Figs
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).
The male palp of S. shinahota sp. nov. (Fig.
Female holotype (Figs
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.
Male paratype (Fig.
Juvenile females had a shiny, dark brown-blackish body surface with dark orange to light brown cephalic part and a pointed abdomen (Fig.
The specific epithet, shinahota, refers to a place with many ants or an ant nest in the Yuracaré language (
Sympolymnia shinahota sp. nov. is known from Brazil, Amazonas State, Manaus and from the Bolivian Departments of Cochabamba (Villa Tunari) and Santa Cruz (Buena Vista). The collection locations of this species were situated in moist Amazonas forest regions, including 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.
Photographs of two individuals (
Holotype
♀; Bolivia: La Paz Department, Nor Yungas Province, Villa Teresa; 16.2019°S, 67.8294°W; 1340 m a.s.l.; beating tray sampling; 17 Jan 2018; R. Perger leg.; IBSI-Ara 1072. Paratypes 2 ♀; same data as for preceding; IBSI-Ara 1024 • 3 ♀, 1 juv.; same data as for preceding;
Sympolymnia cutleri sp. nov. and S. lauretta are indistinguishable in their somatic characters. Sympolymnia cutleri sp. nov. can be separated from all congeners by an epigyne with small, semi-circular opening (Figs
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 abdomen. Carapace slender and elongated, cephalic portion as long as wide (width: 0.81 mm), as wide as widest thoracic part, smooth, marked constriction (width 0.62 mm) between cephalic and thoracic part, two translucent areas at each side of constriction, separated by narrow dark 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; second pair placed on back behind eyes of first pair, but a little further from each other than distance between external 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.
One immature (shiny surface, pointed abdomen) and five female adults (matt surface, rounded abdomen) (Fig.
Live habitus of Sympolymnia spp. and potential ant models. Please note the ontogenetic shift of shine and abdomen shape in the spiders. Sympolymnia 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. latangulus, Buena Vista, Santa Cruz Dept.; Sympolymnia lauretta: I. 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.
The specific epithet, cutleri, is a patronym in honour of Bruce E. Cutler in recognition of his contributions to the taxonomy of Simonellini.
Sympolymnia cutleri sp. nov. is exclusively known from the type location in Bolivian Yungas forest.
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 (
The distribution of Sympolymnia spp. appears to correspond to the delineation of humid Andes forest, Amazon rain forests and sub-humid, semi-deciduous forests (including Chiquitano forest, Bolivian Tucuman forest, Cerrado and Atlantic forest) (Fig.
The consideration of all ant species that were collected in the microhabitats of Sympolymnia species allowed a robust preliminary assessment of potential ant models. We identified at least four potential ant model species in three genera and subfamilies (Table
Possible co-occurring ant models for Bolivian species of Sympolymnia and shared characters: S. lauretta (I) BL ≤ 3.1 mm; (II) BL 3.65–4.05 mm; S. cutleri sp. nov. (I) BL ≤ 3.55 mm; (II) BL 4.02–4.32 mm; S. shinahota sp. nov. (I) 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 | S. lauretta | S. cutleri sp. nov. | S. shinahota sp. nov. | Shared characters | ||||
I | II | I | II | I | II | III | ||
Crematogaster sp. (Myrmicinae) (BL 2.7–3.6 mm) | + | * | + | * | + | * | * | integument black, shiny, without pubescence, abdominal apex pointed |
Camponotus sanctaefidei (Formicinae) (BL 3.4–3.8 mm) | * | + | * | + | * | + | * | integument relatively dull, blackish, single setae, without pubescence, abdominal apex rounded |
Pseudomyrmex ethicus (Pseudomyrmecinae) (BL 3.1–3.6 mm) | + | * | * | cephalic part orangish, remaining integument shiny black; without pubescence, abdomen dorsally constricted and apically pointed | ||||
Camponotus latangulus (Formicinae) (BL 3.65–5 mm) | * | * | + | integument relatively dull, orange, without pubescence, single setae, abdominal apex rounded |
Strong indirect support for ant mimicry was provided by: 1) species-specific similarity involving morphological aspects (Table
All suggested ant models are known for being well defended. The stinger in Crematogaster species is well developed, but the venom is applied topically by wiping on a victim instead of injecting it inside the body (
Mimicry complexes involving adult polymorphic jumping spiders were described for Synemosyna aurantiaca (Mello-Leitão, 1917) (reviewed by
The present study is the first describing putative mimicry complexes involving ontogenetic shifts of ant resemblance (transformational mimicry) in Simonellini. It may be hypothesised that transformational mimicry commonly 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-frequency distribution. However, the genus Sympolymnia appears to include species with and without transformational mimicry, possibly depending on the maximum attainable body size or the presence of suitable ant models.
The reported individuals of S. lucasi (female holotype 3.66 mm; male allotype 3.3 mm, juveniles unknown) have about the same body size as Crematogaster-resembling juveniles of S. lauretta and S. cutleri sp. nov. Considering the small size, it is likely that S. lucasi lacks transformational 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 (
The presence of Sympolymnia throughout South America and the sympatry with Synemosyna species in many locations (e.g.
In addition to similarity, sympatry of models and mimics is considered a critical factor in the selection for mimicry (
We extend many thanks to Wioletta Wawer (Museum and Institute of Zoology of the Polish Academy of Sciences, Warsaw, Poland) and Hubert Höfer (Staatliches Museum für Naturkunde Karlsruhe, Germany) for providing photographs 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. Cutler (University of Kansas, Kansas, U.S.A.), Joseph Schubert (Monash University, Clayton, Australia), Tamás Szűts (University of Veterinary Medicine Budapest, Budapest, Hungary) and Danilo Harms (University of Hamburg, Hamburg, Germany) are thanked for commenting on the manuscript and Mike Skinner for language editing.