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Araneae; Hemiptera; Diptera) Inhabiting Webs of the Spider Tengella Radiata (Araneae, Tengellidae)

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Araneae; Hemiptera; Diptera) Inhabiting Webs of the Spider Tengella Radiata (Araneae, Tengellidae) AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3065, 17 pp., 22 figures June 10, 1993 Natural History and Systematics of Arthropod Symbionts (Araneae; Hemiptera; Diptera) Inhabiting Webs of the Spider Tengella radiata (Araneae, Tengellidae) WILLIAM G. EBERHARD,' NORMAN I. PLATNICK,2 AND RANDALL T. SCHUH3 ABSTRACT The webs of the Costa Rican tengellid spider er numbers include the theridiid spider Argyrodes Tengella radiata (Kulczyinski) are regularly inhab- bryantae Exline and Levi, the mysmenid spiders ited by at least four species of arthropod symbi- Mysmenopsis dipluramigo Platnick and Shadab onts: the phorid fly Megaselia scalaris Loew; the and Maymena rica Platnick, new species, and the plokiophilid bug Lipokophila eberhardi Schuh, new plokiophilid bug Lipokophila tengella Schuh, new species; the uloborid spiderPhiloponella vicina (O. species. The behaviors ofthe symbionts and hosts, P.-Cambridge); and the mysmenid spider Mys- and the interactions among them (which include menopsis tengellacompa Platnick, new species. contest competition, parasitism, predation, and Other arthropods found in Tengella webs in small- commensalism) are described. ' Staff Scientist, Smithsonian Tropical Research Institute; Professor, Escuela de Biologia, Universidad de Costa Rica, Ciudad Universitaria, Costa Rica. 2Chairman and Curator, Department of Entomology, American Museum of Natural History; Adjunct Professor, Department of Biology, City College, City University of New York; Adjunct Professor, Department of Entomology, Cornell University. 3Curator, Department of Entomology, American Museum of Natural History; Adjunct Professor, Department of Biology, City College, City University of New York; Adjunct Professor, Department of Entomology, Cornell Uni- versity. Copyright © American Museum of Natural History 1993 ISSN 0003-0082 / Price $2.00 2 AMERICAN MUSEUM NOVITATES NO. 3065 INTRODUCTION The webs oflarge spiders, such as the orb- Other arthropod species that inhabit Ten- weavers Nephila and Cyrtophora, and some gella webs but have been collected in smaller diplurids, psechrids, and social theridiids, are numbers include the theridiid spider Argy- known to harbor a number of species of rel- rodes bryantae Exline and Levi, the mys- atively small kleptoparasites such as ulobor- menid spiders Mysmenopsis dipluramigo id, theridiid, pholcid, and mysmenid spiders, Platnick and Shadab and Maymena rica Plat- chloropid and drosophilid flies, and others nick, new species, and the plokiophilid bug (Exline and Levi, 1962; Thornhill, 1975; Lipokophila tengella Schuh, new species. Vollrath, 1976, 1979a, b, c; Robinson and We thank Drs. F. C. Thompson and R. Robinson, 1977; Robinson and Lubin, 1979; Peterson ofthe Systematic Entomology Lab- Eberhard, 1980; Nyffeler and Benz, 1980; oratory, United States Department of Agri- Rypstra, 1981; Sivinski and Stowe, 1981; Ta- culture, for identifying the phorid flies. In naka, 1984; Griswold, 1985; Nentwig, 1985; addition to material in the American Mu- Sivinski, 1985; Forster and Murphy, 1986; seum of Natural History (AMNH), speci- Whitehouse, 1986; Baert and Murphy, 1987; mens were supplied by Mr. Darrell Ubick of Coyle and Meigs, 1989, 1992; Coyle et al., the California Academy of Sciences (CDU), 1991). In general, these species take advan- Mr. John A. Murphy of Hampton, England tage ofthe fact that the large spiders often do (JAM), and Dr. H. W. Levi of the Museum not respond to the presence of very small of Comparative Zoology, Harvard Univer- animals in their webs (presumably thus sity (MCZ). Help with illustrations was pro- avoiding expenditures of energy that are un- vided by Dr. Mohammad U. Shadab likely to significantly enhance prey intake). (AMNH). We thank Drs. Jonathan A. Cod- Some symbionts capture small prey that fall dington of the National Museum of Natural into the web and are undetected or ignored History and Frederick A. Coyle of Western by the owner, and some feed directly from Carolina University for helpful comments on prey being consumed by the owner. a draft of the manuscript. This paper describes still another situation of this sort, with the unusual complication that four rather than one species ofinsect and NATURAL HISTORY spider regularly inhabit webs of the host, METHODS Tengella radiata (Kulczy'nski). These include a phorid fly, a plokiophilid bug, and repre- Most of the behavioral observations were sentatives of two other spider families (Ulo- made on animals kept on Tengella webs in boridae and Mysmenidae). The interactions 22 x 30 x 10 cm plastic containers with glass among these species are complex, and include tops. This permitted observations of details contest competition, parasitism, predation, of behavior inside the tubular retreats that and commensalism. Two of the species, the would otherwise have been difficult or im- mysmenid spider Mysmenopsis tengellacom- possible. Some additional observations were pa and the plokiophilid bug Lipokophila made in the field near San Antonio de Escazu, eberhardi, are described below as new by the Costa Rica. Counts of symbionts were made second and third authors, respectively. by collecting entire webs, including retreats, Fincke (1981) showed that one of the as- and carefully examining their contents. sociated species, the uloborid spider Philo- ponella vicina (0. P.-Cambridge), faculta- THE HOST tively builds its orb-webs in the mesh of Tengella radiata is an unusual spider, and threads above Tengella sheet webs, and ap- the family Tengellidae is of uncertain limits. parently prefers such websites over alterna- The association with Tengella ofsuch genera tives. Those Philoponella spiders associated as Calamistrula, Lauricius, Liocranoides, and with Tengella fed more often and disap- Zorocrates proposed by Lehtinen (1967) re- peared less often from their websites than did mains unsupported by synapomorphies, but nonassociated conspecifics. Griswold (cited in Coddington and Levi, 1993 EBERHARD ET AL.: TENGELLA SYMBIONTS 3 Fig. 1. Webs of mature Tengella radiata females. Lateral (a) and end-on (b) views of a newly built web (probably built the preceding evening). Lateral view (c) of an older web, with close-up (d) of outer edge of sheet. The tunnel retreat is at the right in (a) and (c). Arrow in (c) marks long, sagging sticky line. Note also sparse mesh and orbs ofPhiloponella vicina which are present in (c) but nearly completely lacking in (a) and (b). Scale bar = 10 cm in (a), (b), and (c), 2 cm in (d). 1991) has treated tengellids as the sister group (Wolff, 1977; W. Eberhard and M. Santana, of the classical Lycosoidea. Only three spe- unpublished data). Despite its obvious suc- cies of Tengella are known, and two of those cess in varied habitats, however, T. radiata are apparently very rare, being represented is unknown outside Costa Rica. In Panama by a total of four specimens from Mexico, and further south, similar webs in similar Guatemala, and "South America" (Wolff, sites are constructed by mygalomorph spi- 1977). Tengella radiata, however, is ex- ders of the family Dipluridae. What factors tremely common in a wide variety ofhabitats limit the ranges of these apparently ecologi- in Costa Rica, ranging from elevations of cally equivalent groups remain unknown. about 50 m in tropical rain forest up to at The web of T. radiata is generally a large, least 1500 m in secondary growth in and flat sheet turned up at the edges, narrowing around coffee fields in the Valle Central to a tube at the rear, protected side (fig. 1). 4 AMERICAN MUSEUM NOVITATES NO. 3065 During the day, the spider, which is about 2 BEHAVIOR AWAY FROM cm long, is usually found in the tube, whereas THE TENGELLA RETREAT at night it rests at the mouth or on the upper surface of the sheet near the entrance. Con- Both Mysmenopsis tengellacompa and Li- trary to the report of Fincke (1981), spiders pokophila eberhardi were frequently seen on will often attack prey during the day. There the sheet of the host's web. Both walked ei- is a meshwork of threads of variable density ther on top of or under the sheet without and extension above the sheet, and long, sag- apparent difficulty. Most movements were ging lines of cribellum (sticky) silk are often relatively leisurely, but M. tengellacompa in- attached to the meshwork near the edges of dividuals occasionally made brief, quick runs, the sheet (fig. 1). In a newly built web, the especially when fighting among themselves mesh above the sheet was reduced, and there near prey. In both cases, and probably es- were no lines of cribellum silk. When prey pecially in the case of the bug, older individ- falls onto the sheet, the spider usually rushes uals (both males and females) were more like- out, seizes it with its chelicerae, and rushes ly to be found on the sheet away from the back to the tube, where it finishes subduing tube than were younger ones. the prey and feeds. Both ofthese species attacked small insects (e.g., ants about 2 mm long) dropped onto the web. The spiders usually moved to the THE SYMBIONTS prey in a series of short bursts of running or Lipokophila, Mysmenopsis, and Philopo- walking that were interspersed with short nella symbionts were present in appreciable pauses. They often reoriented and moved numbers in the webs of larger Tengella at more directly toward the prey after such both Finca La Selva (elev. 100 m, near Puerto pauses, and in general gave the impression Viejo, in Heredia province) and San Antonio
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