View metadata, citation and similar papers at core.ac.uk brought to you by CORE

provided by PubMed Central Journal of Science: Vol. 11 | Article 147 Campos and Fernández

Coleopterans associated with that form phytotelmata in subtropical and temperate , Raúl E. Campos1,2a* and Liliana A. Fernández1,2b

1Instituto de Limnología "Dr. Raúl A. Ringuelet," Universidad Nacional de La Plata - CONICET, CC 712 (1900) La Plata, Buenos Aires, Argentina 2Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) Abstract A list of the most common plants that form phytotelmata and their associated coleopterans (aquatic, semi-aquatic and terrestrial) from the northeastern subtropical and temperate area of Argentina, South America with biological and behavioral observations is presented in this study. of (n = 3), (5), Apiaceae (6), Araceae (2), Urticaceae (1), Marantaceae (1), (1), Dipsacaceae (1) and Cyperaceae (1) were identified as phytotelmata. Aquatic species of Scirtidae (2), (2), and Hydrophilidae (4), semi-aquatic Chelonariidae (2), and terrestrial species of Carabidae (3), Staphylinidae (5), Histeridae (1), Elateridae (1), Cantharidae (1), Cleridae (1), Tenebrionidae (1), Meloidae (1), Anthicidae (1), Chrysomelidae (3), Curculionidae (7) and Apionidae (1) were identified from six species of Eryngium L. (Apiales: Apiaceae), two species of Kunth (: Poaceae), distichantha Lemaire (Poales: Bromeliaceae), and from fallen leaves of edulis Martius (: Arecaceae) from the temperate and subtropical area. The highest species richness was recorded in Eryngium phytotelmata. Fifteen species of inhabit Eryngium cabrerae Pontiroli, 11 in E. horridum Malme, 7 in E. stenophyllum Urban, 4 in E. aff. serra Chamisso and Schlechtendal., 3 in E. elegans Chamisso and Schlechtendal, 2 in E. eburneum Decne and E. pandanifolium Chamisso and Schlechtendal. From , 6 species of coleopterans were collected from Guadua trinii (Nees) Nees ex Ruprecht and 4 from G. chacoensis (Rojas) Londoño and Peterson. Three species of aquatic coleopterans were recorded from A. distichantha and only one from E. edulis.

Keywords: Aquatic , Aechmea distichantha, biodiversity, Eryngium species, Guadua species Correspondence: a*[email protected]; [email protected], *Corresponding author Received: 25 August 2010, Accepted: 15 December 2010 Copyright : This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 11, Number 147 Cite this paper as: Campos RE, Fernández LA. 2011. Coleopterans associated with plants that form phytotelmata in subtropical and temperate Argentina, South America. Journal of Insect Science 11:147 available online: insectscience.org/11.147

Journal of Insect Science | www.insectscience.org 1 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández

Introduction Kitching 1983). The other families are mainly terrestrial and at least 20 are known from the Phytotelmata are pools of water impounded Neotropical and Nearctic regions (Mestre et by plants (Vargas 1928). The plants, for the al. 2001; Juncá and Da Silva Borges 2002; most part, are living but the term phytotelmata Frank et al. 2004), although only the species also includes pools of water held ephemerally of Staphylinidae were studied (Frank and on concave surfaces of fallen leaves (Kitching Barrera 2010). 1971; Maguire 1971). A specific community develops in these micro environments, These studies on aquatic beetles of Argentina establishing food chains where not only were mostly taxonomic, but few were carried aquatic species but also terrestrial visiting out on communities. We know of two studies species are involved (Frank and Lounibos involving communities of aquatic beetles of 1983). The phytotelmata are distributed in all temperate and subtropical regions. One of continents except Antarctica, but diversity is them is about the structure and temporal greater in the tropics and subtropics. change in the community of coleopterans Approximately 1500 species where inhabiting permanent, semi-permanent and aquatic may find habitats suitable for temporary puddles (Von Ellenrieder and their development (Fish 1983) are known. Fernández 2000). The other is an inventory of Among the most studied phytotelmata, the species from a National Park in the Bromeliaceae native to the New World, the subtropical region, where new distributions Heliconiaceae from South and Central and species from Argentina are reported America and Indonesia, and the Araceae of (Fernández et al. 2008). A third study, not cosmopolitan distribution are highlighted. exclusively on aquatic coleopterans, refers to Pitcher plants (Sarraceniaceae and the community of organisms that live in Nepenthaceae) are special phytotelmata due to ephemeral pools, referring to beetles as their insectivorous characteristic. However, potential predators of a flood-water mosquito some organisms are able to survive in their (Campos et al. 2004). There are no studies digestive fluid (Beaver 1983). A subset of that consider species lists or communities of phytotelmata is the so-called dendrotelmata, beetles that live in or are associated with cavities in hollow trees and stumps of bamboo phytotelmata in Argentina, except the study of (Poaceae), some of which can hold water for a Montero et al. (2010) which cites coleopterans long time. to family level, associated with Aechmea distichantha Lemaire (Poales: Bromeliaceae). The phytotelmata support a large number of species, mainly insects belonging to the order During previous studies on mosquitoes that Odonata, Plecoptera, Trichoptera, Hemiptera, inhabit phytotelmata, associated beetles with Coleoptera and Diptera. The latter are found each phytotelmata class were collected in all types of phytotelmata, as opposed to without performing an exhaustive search. A other orders (Fish 1983). Few identified preliminary report of phytotelmata and their species of beetles have been reported from aquatic and amphibious beetles collected from phytotelmata, of which only members of the subtropical and temperate Argentina is families Scirtidae, Dytiscidae, and presented here. This study includes occasional Hydrophilidae are aquatic (Frank 1983;

Journal of Insect Science | www.insectscience.org 2 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández terrestrial beetles that use the same plants as these formations (APN 2009) are seen on the refuge. rivers.

Materials and Methods Punta Lara Provincial Park is situated on the Río de La Plata, northeast of Buenos Aires Study area Province. The park area corresponds to the Field study was conducted from 1994 through austral part of the Paranense forest ecoregion 2009 in subtropical and temperate Argentina. (Dinerstein et al. 1995), which in this area is The subtropical areas sampled were Iguazú characterized by a narrow gallery forest on the National Park (25º 39’ S, 54º 18’ W) and river. Under natural circumstances, daily floods Chaco National Park (26º 45’ S and 59º 37’ occur with the tide. On the other side of the W), and the temperate areas were Punta Lara gallery forest, there is an open field. Annual Provincial Park (34º 51’ S and 57º 52’ W), and rainfall is approximately 850 mm, and the Ernesto Tornquist Provincial Park (38º 10’ S temperature is mild. and 62º 8’ W) (Figure 1). “Ernesto Tornquist” Provincial Park is situated Iguazú National Park is located north of southeast of Buenos Aires Province in the Misiones Province, separated from neighboring Austral Pampeana ecoregion (Dinerstein et al. by the Iguazú River. This park is part of 1995). This area is characterized by a plain the Paranense forest ecoregion (Dinerstein et with extensive pastures from which a mountain al. 1995) with a topography and drainage range of Paleozoic origin emerges. The pattern dominated by a basaltic plateau that mountains measure 170 km long with a reaches altitudes of 700 m. The annual rainfall maximum height of 1243 MASL and are varies from 1500 to 2000 mm, the dry season oriented from northwest to southeast occurs in winter and abundant precipitations (Harrington 1947). The climate of the are recorded in summer. The mean temperature mountain is wet-subwet with scarce or null varies between 16 and 22º C. The dominant deficit of water (Burgos and Vidal 1951). vegetation is the subtropical forest (APN Annual mean temperature and precipitations 2009). are 14º C and 896 mm (SMN 1981, 1986) with rains falling predominantly in the austral Chaco National Park is situated in Chaco spring-summer period, and occasional snow Province and is part of the humid “Chaco” falling in winter. ecoregion (Dinerstein et al. 1995), a very gentle slope area dominated by depressed Sampling environments. It has a warm subtropical With the exception of bamboo, all climate with summer rainfalls from 750 to phytotelmata were sampled by extracting the 1300 mm. Within the protected area, forest fluid contents with a pipette attached to a lift environments, with savannahs, marshes and pump. After the first extraction, the plants lagoons are found. The forest consists of trees were flushed twice with clean tap water, and that are 15 m tall, with the bottom layer the aquatic contents were extracted with a covered by bromeliads with strong spines on pipette after each wash. All insects were killed the edge of their leaves, making the forest area in the field and preserved in 80% commercial almost impenetrable. Very dense areas of ethyl alcohol.

Journal of Insect Science | www.insectscience.org 3 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández Table 1. Coleopterns inhabiting phytotelmata (except trehole) at subtropical and temperate Argentina.

A: Adult, L: Larva, Aq: Aquatic, Saq: Semi-aquatic, Te: Terrestrrial, St: Subtropical, T: Temperate Due to the fact that the study was carried out and extracting the sample with a hose in national parks where it was not possible to connected to a lift pump. Stump and broken cut bamboo, samples were taken by making a bamboo were sampled again, and internodes hole with a drill in the middle of the internode were washed twice as with the other

Journal of Insect Science | www.insectscience.org 4 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández phytotelmata. Terrestrial beetles found on arrangement of their leaves delimits axils, as plants in the vicinity of phytotelmata were the peripheral leaves are older than the central collected manually using entomological ones. The internal axils hold free water and forceps. debris, the intermediate a semi-liquid interface composed mainly of debris and slime-fluxes, Results and the oldest, wet slime and debris. Thus, the real cavity where water is contained for a long The identified phytotelmata were: Guadua time is limited to the axils of internal leaves. trinii (Nees) Ness ex Ruprecht, Guadua Water in external cavities is ephemeral and chacoensis (Rojas) Londoño and Peterson, makes favorable habitats for semi-aquatic or Merostachys clausseni Munro (Poaceae), terrestrial macro-invertebrates. Depending on Aechmea distichantha Lemaire, Aechmea the species, the size of the plants and the recurvata (Klotzsch), L.B. Smith, Billbergia water retained by the axils was variable. The nutans Wendland ex Regel, Pseudananas mean diameters were: E. cabrerae 83 cm (± sagenarius (Arruda) Camargo, SD 14.9); E. horridum 77 cm (± SD 0.17); E. friburgensis Mez, (Bromeliaceae), Eryngium stenophyllum 94 cm (± SD 0.23); E. aff. serra sp. (Apiaceae), Alocasia odora (Lindl.) Hoch, 46 cm (± SD 0.15); E. elegans 40 cm (± SD Philodendron bipinnatifidum Schott ex 0.11). The volumes of impound water were: Endlicher, (Araceae), Urera baccifera (L.) 69.4 ml (± SD 43.1); 20.6 ml (± SD 14.2); 7.4 Gaudichaud-Beaupré ex Weddell, ml (± SD 6.6); 19.1 ml (± SD 17.1); 19.1 ml (Urticaceae), Maranta sp. (Marantaceae), (± SD 17.1); 7.7 ml (± SD 12.3), respectively. Euterpe edulis Martius, (Arecaceae), No measures of E. eburneum were recorded. Dipsacus sp. (Dipsacaceae) at Iguazú National Park, Aechmea distichanta and Eryngium sp. Guadua (Figure 4) grow mainly in at Chaco National Park, Eryngium horridum low places near streams, and can reach up to Malme, Eryngium stenophyllum Urb., 12 meters. In the study area we observed two Eryngium aff. serra Cham. and Schltdl., types of bamboo phytotelmata, one in the Eryngium elegans Cham. and Schltdl. at internodes that have one or more holes in the Ernesto Tornquist Provincial Park, wall by the penetration of rain water (Figure Androtrichum giganteum (Kunth) Pfeiff. 5) and the other in the stumps as a result of (Cyperaceae), Guadua trinii, Eryngium cutting bamboo (Figure 6). The longitude of cabrerae Pontiroli, Eryngium eburneum the internodes, inlet holes and the volume of Decne, Eryngium pandanifolium Cham. and water retained into the internodes were Schltdl., and Dipsacus sp. at Punta Lara different in each species of bamboo. Guadua Provincial Park. All except Alocasia odora chacoensis was the biggest, with a mean and Dipsacus sp. are native phytotelmata. longitude, hole diameter and water volume of 23.3 cm (± SD 3.5), 9.1 mm (± SD 4.9) and From the above 21 classes of phytotelmata 366.1 ml (± SD 225), while G. trinii was 23.1 sampled, coleopterans were collected from A. cm (± SD 5.5), 6.1 mm (± SD 3.8) and 45.2 distichantha, E. edulis, six species of ml (± SD 41.1) respectively. Eryngium and two species of Guadua (48%) (Table 1). Eryngium species that are Aechmea distichantha (Figure 7) is epiphytic phytotelmata are morphologically similar to or terricolous and was the dominant bromeliad bromeliads (Figures 2, 3). The imbricate in Chaco and Iguazú subtropical parks. The

Journal of Insect Science | www.insectscience.org 5 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández volume of water impounded in its axils, stages from Eryngium horridum. Larvae of sometimes several liters, was much greater this weevil were observed drilling the stalk of than the volume in the axils of Eryngium. inflorescence and adults living in axils.

The fallen floral bracts of the palmetto The greatest diversity of coleopterans (27 Euterpe edulis (Figure 8) were the only type species) was found in Eryngium (Table 1); the of phytotelmata found in this study in non- largest percentage of species was collected living parts of the plant. The bracts, which from E. cabrerae (56%), followed by E. collect rain water in their inner cavity, are horridum (41%), E. stenophyllum (26%), E. woody and more than one meter in length. aff. serra (15%), E. elegans (11%), and E. eburneum and E. pandanifolium (7%). Beetles Sixteen families of coleopterans were were poorly represented in E. edulis, A. recorded, fourteen from Eryngium, five from distichantha and in both Guadua bamboos Guadua, three from A. distichantha, and one (Figure 9). No species inhabiting Eryngium from E. edulis. Hydrophilidae, Histeridae, were observed in Guadua or E. edulis. Only Elateridae, Cantharidae, Cleridae, Liodessus sp. was found both in Eryngium and Tenebrionidae, Meloidae, Anthicidae and in A. distichantha (Table 1). Chrysomelidae were present exclusively in species of Eryngium, while chelonariids were found in Guadua, and Apionidae in A. Discussion distichantha. No exclusive family was found in E. edulis. Dytiscids were present in Among the 21 classes of phytotelmata Eryngium and A. distichantha; scirtids in reported here, 15 correspond to the subtropical Eryngium, Guadua and A. distichantha; northern area, and nine to the temperate area. staphylinids in Eryngium, Guadua and E. Coleopterans were only recorded in edulis, and curculionids in Eryngium and Eryngium, Guadua, A. distichantha and E. Guadua. Among aquatic coleopterans, edulis. The species of Eryngium are dominant Enochrus (Hugoscottia) sp. (Hydrophilidae) in temperate areas (Campos 2010), while the and unidentified Scirtidae were collected in species of Guadua are distributed in the the larval stage, while Copelatus sp., subtropical area, extending in the temperate Liodessus sp. (Dytiscidae), Phaenonotum sp., area along a narrow strip of forest on La Plata Paracymus sp., Tropisternus setiger River. Aechmea distichantha and E. edulis are (Hydrophilidae) were collected as adults subtropical. (Table 1). Semi-aquatic species were represented by Chelonariidae larvae and Five families of aquatic coleopterans have Curculionidae by Tyloderma adults (Table 1). been related to phytotelmata in the Neotropical region. A recent review carried Families of terrestrial beetles were Carabidae, out by Frank and Lounibos (2008) cites Staphylinidae, Histeridae, Tenebrionidae, representatives of the families Scirtidae, Elateridae, Cantharidae, Cleridae, Meloidae, Dytiscidae and Hydrophilidae inhabiting Anthicidae, Chrysomelidae and Curculionidae bromeliads. Other phytotelmata, like bamboo (Table 1). The weevil Heilipodus erythropus and fallen flower bracts of a palm, also (Klug) (Coleoptera, Curculionidae) was the include a few species of the families Elmidae only species collected in both larval and adult (Sanchez and Liria 2009) and Noteridae

Journal of Insect Science | www.insectscience.org 6 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández (Greeney 2004), respectively. On the other Frank and Lounibos 2008) and in fallen hand, the diversity of terrestrial beetles that flower bracts of the stilt-root palm Iriartea visit phytotelmata is higher. A study carried deltoidea Ruiz and Pavon (Arecaceae: out in southern Brazil (Mestre et al. 2001) Iriarteeae), and Laccophillus Leach and shows that the bromeliad Vriesea inflata Thermonectus Dejean in this last phytotelmata (Wawra) Wawra is visited by species of 13 (Greeney 2004). In our study, adults of families of terrestrial coleopterans. However, unidentified species of the Copelatus other authors (Gutierrez Ochoa et al. 1993; and Liodessus Guignot dwelling in the axils of Greeney 2001; Frank et al. 2004) mentioned Eryngium and Liodessus and in the axils of A. other families not cited by Mestre et al. (2001) distichantha were found. As adults were that use bromeliads and other phytotelmata to scarce and no immatures were found, it is hide or find prey, summarizing at least 20 presumed that these could be opportunistic families. In accordance with Frank and species, which use water from the axils when Lounibos (2008), representative species of the pools dry up. three families of aquatic coleopterans were recorded from phytotelmata of Argentina Hydrophilidae – There are two subfamilies other than bromeliads. In contrast with the with species that inhabit phytotelmata, numerous terrestrial coleopterans known, only Sphaeridiinae and Hydrophilinae. Subfamily species of 12 families were recorded, probably Sphaeridiinae is found in a great diversity of due to the lack of an exhaustive search. habitats, mainly terrestrial, but in some cases Several studies show that coleopterans are a some species of this subfamily are secondarily taxon of scarce richness and abundance in the aquatic, while Hydrophilinae are mostly communities living in phytotelmata. The aquatic and are found in most freshwater exceptions were observed in invertebrate environments (Archangelsky 2004). All the communities of the bromeliads Vriesea species specialized in living in Bromeliaceae extensa Sm. (Synon.: Alcantarea extensa are Sphaeridiinae subfamily and belong to the (Sm.) Grant) and Vriesea inflata where a genera Coelostoma Brullé, Lachnodacnum Carabidae (Juncá and Borges 2002) and a Orchymont, Phaenonotum Sharp, Omicrus Scirtidae (Mestre et al. 2001), respectively, Sharp (Frank and Lounibos 2008). Other were dominant although richness was low, members of this subfamily are contrary to the high species richness (in Dactylosternum Wollaston and Pelosoma particular Dytiscidae) observed in fallen floral Mulsant, which were found inhabiting in bracts of a palm (Greeney 2004). In our study, fallen flower bracts of a palm (Greeney 2004). although not quantified, the small number of The genera of Hydrophilinae that were beetles was highly contrasting with dipterans reported from phytotelmata are and other in all sampled Hydrobiomorpha Blackburn, Enochrus phytotelmata. Thomson and Derallus Sharp, all collected as adults from fallen floral bracts of a palm Aquatic Coleopterans (Greeney 2004). The only record of a Dytiscidae – Species of four genera of hydrophilid inhabiting treeholes in the Dytiscidae have been cited in the literature as Neotropic region was quoted by H. F. inhabitants of phytotelmata (sensu lato): Greeney as unpublished data in “The insects Copelatus Erichson and of plant-held water: a review and Babington in bromeliads (Greeney 2001; bibliography” (Greeney 2001), and did not

Journal of Insect Science | www.insectscience.org 7 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández mention the genus. During our sampling, we (Daugherty and Juliano 2003). Scirtid species found Tropisternus setiger (Germar), from South America are little known, and the Paracymus sp. (adults) and Enochrus sp. only genera cited as phytotelmata inhabitants (larva), all Hydrophilinae, and Phaenonotum are Scirtes, and Cyphon collected from sp. (adults) a Sphaeridiinae, living in the axils Bromeliaceae in (Ospina-Bautista of four Eryngium phytotelmata. T. setiger is a et al. 2004) and (Liria 2007), ubiquitous species that lives in ephemeral respectively. Other studies carried out in (Campos et al. 2004), temporary, and Brazil (Mestre et al. 2001), Perú (Louton et al. permanent pools (Von Ellenrieder and 1996) and Argentina (Montero et al. 2010) Fernández 2000) and therefore it cannot be mentioned Scirtidae living in bromeliads and regarded as a specific inhabitant of bamboo without indicating their identity. In phytotelmata. Its habit of oviposition on this study, larvae of three morpho-species aquatic vegetation (Jerez and Moroni 2006) living in Guadua, A. distichantha and limits its possibility of developing in the axils Eryngium phytotelmata in both subtropical of Eryngium or other phytotelmata. Enochrus and temperate areas were found. However, is a widely distributed genus living in they could not be identified because they were temporary and permanent environments, not reared to the adult stage (and because whose females lay egg cases that adhere to the there are no keys to Argentinian scirtid underside of the leaves of the floating Azolla larvae). filiculoides Lamarck (Salviniales: Azollaceae) (Fernández 1992). Eryngium phytotelmata are Hydrophilic or semi-aquatic Coleopterans frequently partially submerged in floodwater Chelonaridae – This family is little known and puddles, where dense vegetation is formed has been found in association with treeholes dominated by A. filiculoides. When the pond (Spangler 1980) and bromeliads, apparently dries up this small plant is retained in the axils feeding on decaying plant matter (Costa et al. (Campos unpublished data), and can contain 1988). However, there is still controversy over larvae or egg cases of Enochrus between its whether or not their larvae are aquatic. leaves. This would explain the presence of Spangler (1980), in contrast with others Enochrus larvae in the leaf axils of E. authors (Boving and Craighead 1931), cabrerae as was observed in this study. concludes that these larvae are not aquatic because they do not have real anal gills and Scirtidae – Is the most abundant family of attributes their presence in aquatic coleopterans that inhabit phytotelmata, mainly environments as a consequence of the moss treeholes (Greeney 2001), bamboo (Louton et being washed off of their habitat. Only a few al. 1996), and bromeliads (Ospina-Bautista et larvae are known and little is known of their al. 2004). There are five genera known from behavior. More research could reveal phytotelmata: Scirtes Illiger (Picado 1913), unknown species that inhabit phytotelmata Cyphon Paykull, Ora Clark (Frank et al 2004), that could be adapted to aquatic life. In this Prionocyphon Redtenbacher (White 1978), study, we reported unidentified Chelonaridae and Flavohelodes Klausnitzer (Stribling and larvae inhabiting bamboo internodes (Guadua Young 1990), whose larval habits are mainly chacoensis and G. trinii). The presence of detritivorous converting the leaf litter into fine these larvae in a habitat of vertical and smooth particles which benefit fine particle feeders walls, such as bamboo internodes where there co-occurring in the same phytotelmata is not an interface between water and

Journal of Insect Science | www.insectscience.org 8 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández decaying matter (as in axils of bromeliads and sensory bristles that work like a trap when treeholes), would suggest that these species capturing prey, was found in the internodes of are better adapted to live in an aquatic Guadua bamboo. This eighth modified environment. Because the behavior of segment (Figure 10) is used to plug the Chelonaridae species is unknown, this opening of the gallery in which they live hypothesis should be tested. (Costa et al. 1988). Our observations revealed that the larvae of Pachyteles sp., reared in Curculionidae - Of the 131 species of semi- bamboo internodes that contained the original aquatic weevils (Curculionidae, Erirhinidae sediment and water, dived to the bottom, from and Dryophthoridae) known from Argentina which they extracted debris with their (Morrone and O'Brien 1999), we collected an mandibles and built a cell compartment on the unidentified specie of the genus Tyloderma. inner wall of the internodes. It was verified Lanteri et al. (2002) reports that 12 species of that they are good predators of Isoptera and Tyloderma in Argentina are associated with other small insects because, when feeding, the aquatic macrophytes. We found abundant body bends towards the last abdominal individuals of Tyloderma living in axils of E. segment and adopts a V form (Costa et al. horridum, but none was immersed in water. 1988). It was also observed that the community of macro-invertebrates living in Terrestrial Coleopterans bamboo internodes is very diverse This group is composed of beetles that use (unpublished data), and the free water space is phytotelmata as refuge or searching for prey, occupied by various terrestrial insects, even and as already mentioned, is represented by by termite and ant nests. For this reason, many families. Some species are closely Pachyteles larvae could be considered one of associated with the plant host, while others are the top predators within this community. On occasional. The families that were found the other hand, the unique individual of the during our study are discussed below. unidentified Carabidae collected from Eryngium cabrerae is a terrestrial species Carabidae – The species which live associated considered a visitor. with phytotelmata were only mentioned as inhabitants of Bromeliaceae and belong to Staphylinidae – This familiy is represented in Agra Fabricius, Calathus Bonelli, Callida communities of bromeliads (e. g. Picado Latreille and Dejean, Colpodes MacLeay, 1913; Lüderwaldt 1915; Zaragoza 1974), Lebia Latreille, Lia Eschscholtz, Onypterigia fallen floral bracts of a palm (Greeney 2004), Dejean, Phloeoxena Chaudoir, Pterostichus and from inflorescences of several species of Bonelli, (Laessle 1961; Zaragoza 1974; Heliconia L. (Zingiberales: Heliconiaceae) Gutiérrez et al. 1993) and Platynus Bonelli (Frank and Barrera 2010). Six species of (Montes de Oca et al. 2007) that were Staphylinidae were collected (Table 1). collected in Jamaica and Mexico. Less Loncovilius sp., Pinophilus sp. and Stenus sp. specialized Carabidae were found were present exclusively in the axils of the occasionally, looking for prey in fallen floral species of Eryngium. Loncovilius sp. was bracts of a palm in Ecuador (Greeney 2004). most abundant mainly in Eryngium cabrerae. During this study, a conspicuous larva of the An unidentified adult and larva were collected genus Pachyteles Perty, characterized by a from species of Guadua and from fallen floral widened eighth abdominal segment and bracts of E. edulis, respectively. All species

Journal of Insect Science | www.insectscience.org 9 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández could be considered as potential top predators unidentified Cantharidae inhabiting two from the aquatic communities because they species of Eryngium that could be considered can prey on various species of aquatic larvae occasional predators that use phytotelmata to as was observed by Greeney (2004) and Frank hide in or find prey. and Barrera (2010) during studies carried out in communities inhabiting the fallen floral Cleridae and Meloidae – This is the first bracts of a palm, and flower bracts of record of representatives of these families of Heliconia, respectively. coleopterans associated with phytotelmata. We found adults in E. horridum, but Histeridae - The only species of this family unfortunately, their identification was not associated with phytotelmata, Omalodes possible. sobrinus Erichson, was reported from hemisphaerica Lamarck 1783 of Tenebrionidae - There are few reports of Mexico, and was listed as a very scarce associated Tenebrionidae to phytotelmata, all herbivore (Gutiérrez et al. 1993). We reported of them referring to Bromeliaceae. Goniadera an unidentified species associated with E. oculata Champion, Eurymepoton sp. and cabrerae in temperate areas, which was also Eleodes sp. were cited inhabiting A. bracteata, of very low abundance. caputmedusae Morren, and Vriesea chiapensis Matuda, respectively (Zaragoza Elateridae - Few studies have been done on 1974). Stibia sp. was found associated with Elateridae associated with phytotelmata. Bromelia hemisphaerica (Gutiérrez et al. Zaragoza (1974) cited Elateridae inhabiting 1993) and Glyptotus cribratus LeConte to phytotelmata to species level. The species Tillandsia fasciculata Swartz, the latter in the cited were: Platycrepidius boucardi (Salle), Nearctic region (Frank et al. 2004). Megapenthes cincticollis Champion, Ischiodontus anceps Candèze, and Anchastus Another unidentified tenebrionid was found in seminiger Champion, inhabiting the bromeliad terrestrial and epiphytic Vriesea inflata Aechmea bracteata (Swartz) Grisebach in (Mestre et al. 2001). The ecological niche of Mexico. Domingues et al. (1989) reported the phytotelmic species is unknown; however, larvae of unidentified Elateridae inhabiting Zaragoza (1974) classified them as epiphytic bromeliads in Brazil and Mestre et detritivorous because they feed on the debris al. (2001) found unidentified adults of that accumulates in the leaf axils. In the Elateridae in the epiphytic Vriesea inflata present study, we report Rhypasma (Bromeliaceae), but not in terrestrial ones. quadricollis (Fairmaire) associated with While Frank et al. (2004) reported Conoderus Eryngium cabrerae, a phytotelmata different amplicollis (Gyllenhal) inhabiting Tillandsia from bromeliads. recurvata (Linnaeus) L. in USA. We reported an unidentified species associated with the Anthicidae - The only species of this family terrestrial E. horridum (Apiaceae). reported from phytotelmata is Anthicus asphaltinus Champion, collected in Mexico Cantharidae - The only record of Cantharidae from B. hemisphaerica, which was considered is Chauliognathus tripartitus Chevrolat scarcely abundant and its ecological niche inhabiting Vriesea sp. from Mexico (Zaragoza unknown (Gutiérrez et al. 1993). We collected 1974). In our study, we found few larvae of an identified specimen from E. stenophyllum.

Journal of Insect Science | www.insectscience.org 10 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández with Eryngium species in Argentina. We also Apionidae: The genus Apion, classified as report, Napactus dives Klug associated with E. Curculionidae and Brentidae by other authors, horridum, and Aramigus tessellates (Say) was reported from A. bracteata, Vriesea associated with E. horridum, E. stenophyllum chiapensis (Bromeliaceae) (Zaragoza 1974), and E. cabrerae. We also report an and T. fasciculata (Frank et al. 2004). We unidentified species of Curculionidae from report a species of the same genus inhabiting Guadua bamboo. Heilipodus erythropus is a A. distichantha, in the subtropical area. species extensively distributed in Argentina and neighboring countries (Lanteri et al. Chrysomelidae – Although most of the 2002). It was the most abundant in our study Chrysomelidae are terrestrial, some and was collected from five species of (Donaciinae, Chrysomelinae and Galerucinae) Eryngium. Its larvae drill the inside of the exhibit some degree of adaptation to aquatic stem of the inflorescence of Eryngium life (Konstantinov 2003). The few species of paniculatum Cavanilles and Dombey, where Chrysomelidae reported as inhabitants of they develop their life cycle and later pupate phytotelmata are of terrestrial habits and feed in the root (Ríos de Saluso 1997). De Saluso on the leaves of the host plant (Frank and mentioned that this weevil was found Lounibos 2008). Acentroptera pulchella exclusively on Eryngium, but an old report Guérin-Méneville (Mantovani et al. 2005) of cited this Curculionidae causing damage to Brazil, Calliaspis rubra (Olivier) of Peru Bromeliacaea (Silva et al. 1968). Although H. (Burgess et al. 2003), and Pachybrachys sp. of erythropus is not cited by Morrone and Mexico (Gutierrez et al. 1993) were reported O'Brien (1999) as a semi-aquatic species, it from Bromeliaceae, while some species of was observed in the field that when the adult Systena Chevrolat were found associated with is disturbed, it dives into the water where it Eryngium (Apiaceae) (Jolivet and remains for several minutes until the danger Hawkeswood 1995). No aquatic disappears (Campos unpublished data). This Chrysomelidae were reported from any behavior was observed several times, even phytotelmata. The three species collected without touching the plant. The mere shadow during the present study were associated with of our approach made them take refuge in the Eryngium spp. One of these was an water. Tonesia argentinica was reported unidentified cassidine; the others, Chrysodina inhabiting E. paniculatum in subtropical cuprescens Boheman and Systena sp., are Chaco Province. We found an identified polyphagous, but only the latter was cited species of this genus associated with E. feeding on the leaves of Eryngium (Jolivet and cabrerae in temperate Buenos Aires Province. Hawkeswood 1995). Species of Elateridae, Meloidea, Curculionidae – This family is the most Chrysomelidae, and Curculionidae were extensive of the terrestrial coleopterans observed not only in the axils of Eryngium associated with phytotelmata, and its species phytotelmata, but also on inflorescence during were reported mainly from Bromeliaceae, a the hottest hours, taking refuge in the axils in major pest of these plants (Frank 1999). In the morning or during drier periods. These their 2002 review, Lanteri et al. reported that species might not be interacting in the aquatic H. erythropus and Tonesia argentinica community, except for their input of organic Hustache were the only curculionid associated matter as a consequence of their stools.

Journal of Insect Science | www.insectscience.org 11 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández the exploration of the subtropical forest of the The terrestrial beetle community associated Yungas, located in the northwest. We believe with phytotelmata in the subtropical and that the same will happen with the diversity of temperate region of Argentina was richer in plants that function as phytotelmata. However, species than the aquatic community. Most the development of comprehensive lists could species of terrestrial beetles found in this be limited by the scarce taxonomic knowledge study are ubiquitous, being associated with of the Neotropical coleopterans. other plants that are not phytotelmata. However, some staphylinids species seem to Acknowledgements be exclusive inquilines of Eryngium, interacting with the aquatic community as We thank the Administración de Parques predators. Another specific beetle is the Nacionales, Delegación Regional Noreste weevil H. erythropus whose life cycle takes (APN), Centro de Investigaciones Ecológicas place in different parts of the plant. Among Subtropicales (CIES), and Administración de the aquatic beetles that inhabit Eryngium, Áreas Protegidas y Conservación de la none of them was specific. All the species Biodiversidad, Ministerio de Asuntos Agrarios found in this study use aquatic environments de la Provincia de Buenos Aires for logistic other than phytotelmata. A possible help and licence conceded. We also thank hypothesis is that the presence of ubiquitous Motoyoshi Mogi (Saga University, Japan), species in phytotelmata could be a strategy of Victoria Sy (ILPLA, UNLP-CONICET), and displacement to other aquatic environments, Eduardo Lestani (Universidad Nacional de using these plants occasionally and for a short Córdoba) for their help in field work, and the time. Richness of beetles in Guadua bamboo following scientists for taxonomic was low, however the most common and determination of specimens: Susana Martínez abundant inhabitant was the carabid (Eryngium species) (Facultad de Ciencias Pachyteles. For their behavior and their habit Exactas y Naturales, Universidad de Buenos of constructing cells within internodes, one Aires), Analía Lanteri (Curculionidae), Nora could hypothesize that it is a species closely Cabrera (Chrysomelidae) (Facultad de Ciencias associated with this type of micro-habitat Naturales y Museo de La Plata, UNLP), interacting with phytotelmata through the Mariana Chani Posse (Staphylinidae), Gustavo removal of sediment and providing nutrients Flores (Tenebrionidae) (Instituto Argentino de as a product of excrements. On the other hand, Investigaciones en Zonas Áridas, IADIZA - the scarcity of water beetles could be due to CONICET). Finally, we are grateful to Mónica the relative smallness of the inlets which, Caviglia for help with the English translation. being located in the side wall of the This research was supported by Grant 5924, internodes, could not be detected by most and International Cooperatión Project D 1369 ubiquitous species. The only water beetles from Consejo Nacional de Investigaciones found in the stumps were scirtids, which are Científicas y Técnicas (CONICET), Argentina. also common inhabitants of treeholes. Their This is a contribution of the Instituto de presence could be due to the similarity of Limnología Journal Series No 885. bamboo stumps with treeholes. References The number of coleopterans associated with phytotelmata in Argentina may increase with

Journal of Insect Science | www.insectscience.org 12 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández APN 2009. Administración de Parques Cave RD. 2005. Trigonopeltastes delta Nacionales. Available online, (Coleoptera: Scarabaeidae: Cetoniinae) larva http://www.parquesnacionales.gov.ar developing in a bromeliad terrarium. Coleopterist Bulletin 59: 527-528. Archangelsky M. 2004. Higher-level phylogeny of Hydrophylinae (Coleoptera: Costa C, Vanin SA, Casari Chen SA. 1988. Hydrophilidae) based on larval, pupal and Larvas de Coleoptera do Brasil. Museum de adult characters. Systematic Entomology 29: Zoología, Universidade de São Paulo, Brazil. 188-214. Daugherty MP, Juliano SA. 2003. Leaf Beaver RA. 1983. The communities living in scraping beetle feces are a food resource for Nepenthes pitcher plants: fauna and food tree hole mosquito larvae. The American webs. In: Frank JH and Lounibos LP, Editors. Midland Naturalist 150: 181-184. Phytotelmata: terrestrial plants as hosts of aquatic insects communities, pp. 29-54. Plexus Dinerstein E, Olson DM, Graham DJ, Webster Inc. AL, Primm SA, Bookbinder MP, Ledec G. 1995. Una evaluación del estado de Boving AG, Craighead FC. 1931. Illustrated conservación de las ecoregiones terrestres de synopsis of the principal larval forms of the América Latina y el Caribe. WWF and World order Coleoptera. Entomologica Americana 11: Bank, Washington D.C. 1-352. Domingues RAP, Pugialli HRL, Dietz JM. Burgess J, Burgess E, Lowman M. 2003. 1989. Densidade e diversidade de fauna Observations of a beetle herbivore on a fitotelmata em bromélias de quatro tipos de bromeliad in Peru. Journal of the Bromeliad florestas degradadas. Revista Brasileira de Society 53:221-224. Biologia 49: 125-129.

Burgess J, Vidal AL. 1951. Los climas de la Fernández LA. 1992. Descripción de la ooteca república Argentina según la nueva y estadios larvales de Enochrus (Hugoscottia) clasificación de Thornthwaite. Meteoros 1:3- scutellaris (Bruch) y Enochrus (Methydrus) 32. vulgaris (Steinheil) (Coleoptera, Hydrophilidae). Revista de la Sociedad Campos RE. 2010. Eryngium (Apiaceae) Entomológica Argentina 50: 343-349. phytotelmata and their macro-invertebrates communities, including a review and Fernández LA, Archangelsky M, Manzo V. bibliography. Hydrobiologia 652: 311-328. 2008. Coleópteros acuáticos del Parque Provincial Salto Encantado y valle del Cuña Campos RE, Fernández LA, Sy VE. 2004. Pirú (Misiones, Argentina). Revista de la Study of insects associated with the floodwater Sociedad Entomológica Argentina 67: 87-98. mosquito Ochlerotatus albifasciatus (Diptera: Culicidae) and their posible predators in Fish D 1983. Phytotelmata: flora and fauna. In: Buenos Aires Province, Argentina. Phytotelmata: terrestrial plants as hosts of Hydrobiologia 524: 91-102. aquatic insects communities. In: Frank JH and Lounibos LP, Editors. Phytotelmata: terrestrial

Journal of Insect Science | www.insectscience.org 13 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández plants as hosts of aquatic insects communities, Public Museum Contributions in Biology and pp. 1-27. Plexus Inc. Geology 101: 1-20.

Frank JH. 1983. Bromeliad phytotelmata and Gutiérrez Ochoa M, Camino Lavin M, their biota, especially mosquitoes. In: Frank JH Castrejon Ayala F, Jiménez Pérez A. 1993. and Lounibos LP, Editors. Phytotelmata: Arthropods associated with Bromelia terrestrial plants as hosts of aquatic insects hemisphaerica (Bromeliales: Bromeliaceae) in communities, pp. 101-128. Plexus Inc. Morelos, Mexico. Entomologist 76: 616-621. Frank JH. 1999. Bromeliad-eating weevils. Selbyana 20: 40-48. Harrington H. 1947. Explicación de las hojas 33m y 34m: Sierra de Curamalal y Ventana, Frank JH, Barrera R. 2010. Natural history of Provincia de Buenos Aires. Dirección de Belonuchus Normann spp. and allies Minas y Geología Buenos Aires Boletín 51: 33- (Coleoptera: Staphylinidae) in Heliconia L. 34. (Zingiberales: Heliconiaceae) flower bracts. Insecta Mundi 110: 1-11. Jerez V, Moroni J. 2006. Diversity of freshwater beetle of Chile. Gayana 70: 72-81. Frank JH, Lounibos LP. 1983. Phytotelmata: terrestrial plants as hosts of aquatic insects Jolivet P, Hawkeswood TJ. 1995. Host-plants communities. Plexus Inc. of Chrysomelidae of the world. Backhuys Publishers. Frank JH, Lounibos LP. 2008. Insects and allies associated with bromeliads: a review. Juncá FA. Da Silva Borges C. 2002. Fauna Terrestrial Rewiews 1: 125-153. associada a bromélias terrícolas da Serra da Jibóia, Bahia. Sitientibus série Ciências Frank JH, Sreenivasan S, Benshoff PJ, Deyrup Biológicas 2: 73-81. MA, Edwards GB, Halbert SE, Hamon AB, Lowman MD, Mockford EL, Scheffran RH, Kitching RL. 1971. An ecological study of Steck GJ, Thomas MC, Walker TJ, Welbourn water-filled tree-holes and their position in the WC. 2004. Invertebrate extracted woodland ecosystem. Journal of from native Tillandsia (Bromeliales: Ecology 40: 281-302. Bromeliaceae) in Sarasota County, Florida. Florida Entomologist 87: 176-185. Konstantinov AS. 2003. Chrysomelidae: aquatic leaf beetles of China (Coleoptera). In: Greeney HF. 2001.The insects of plant-held Jäch MA, Ji L Editors. Water beetles of China, waters: a review and bibliography. Journal of Volume 3. pp. 563-572. Zoologisch- Tropical Ecology 17: 241-260. Botanische Gesellschaft in Österreich and Wiener Coleopterologenverein Press. Greeney HF. 2004. Fallen flower bracts of the ! stilt-root palm Iriartea deltoidea (Palmae: Laessle AM. 1961. A micro-limnological Iriarteeae) as phytotelmata habitats in a study of Jamaican bromeliads. Ecology 42: lowland Ecuadorian rainforest. Milwaukee 499-517.

Journal of Insect Science | www.insectscience.org 14 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández Lanteri A, Marvaldi A, Suárez S. 2002. Brazilian Archives of Biology and Technology Gorgojos de la Argentina y sus plantas 44: 89-94. huéspedes. Tomo I: Apionidae y Curculionidae. Publicación Especial de la Montero G, Feruglio C, Barberis IM. 2010. Sociedad Entomológica Argentina 1: 1-98. The phytotelmata and foliage macrofauna assemblages of a bromeliad species in Liria J. 2007. Fitotelmata fauna on the different habitats and seasons. Insect bromeleans Aechmea fendleri André y Conservation and Diversity 3: 92–102. Hohenbergia strellata Schult of the San Esteban Nacional Park, Venezuela. Revista Montes de Oca E, Ball GE, Spence JR. 2007. Peruana de Biología 14: 33-38. Diversity of Carabidae (Insecta, Coleoptera) in epiphytic Bromeliaceae in central Veracruz, Louton J, Gelhaus J, Bouchard R. 1996. The Mexico. Environmental Entomology 36: 560- aquatic macrofauna of water-filled bamboo 568. (Poaceae: Bambusoideae: Guadua) internodes in a Peruvian lowland tropical forest. Morrone JJ, O'Brien CW. 1999. The aquatic Biotropica 28: 228-242. and semiaquatic weevils (Coleoptera: Curculionoidea: Curculionidae, Dryopthoridae Lüderwaldt H. 1915. Insekten und sinstiges and Erirhinidae) of Argentina, with indication Tierleben an brasilianischen Bromeliaceen. of their host plants. Physis 57: 25-37. Zeitshrift für wissenschaftliche. Insektenbiologie 11: 78-84. Ospina-Bautista F, Estévez-Varón JV, Betencur J, Realpe-Rebolledo E. 2004. Maguire B. 1971. Phytotelmata: Biota and Estructura y composición de la comunidad de community structure determination in plant- macro invertebrados acuáticos asociados a held waters. Annual Review of Ecology and Tillandsia turneri Baker (Bromeliaceae) en un Systematics 2: 439-464. bosque alto andino Colombiano. Acta Zoológica Mexicana (Nueva Serie) 20: 153- Mantovani A, Magalhães N, Teixeira ML, 166. Leitão G, Staines CL, Resende B. 2005. First report on host plants and feeding habits of the Picado C. 1913. Les broméliacees éphiphytes leaf beetle Acentroptera pulchella Guérin- considérées come milieu biologique. Bulletin Méneville (Chrysomelidae, Hispinae). In: Scientifique de la France et de la Belgique Contributions to systematics and biology of 47 : 215-360. beetles. Papers celebrating the 80th birthday of Igor Konstantinovich Lopatin. Pensoft Ríos de Saluso MLA. 1997. Artrópodos Series Faunistica No. 43, pp. 153-157. asociados al “Caraguatá,” Eryngium Pensoft. paniculatum (Apiaceae). Revista Científica Agropecuaria 1: 35-37. Mestre LAM, Aranha JMR, Esper MdeLP. 2001. Macroinvertebrate fauna associated to Sanchez E, Liria J. 2009. Relative abundance the bromeliad Vriesea inflata of the Atlantic and temporal variation of macroinvertebrates forest (Paraná State, Southern Brazil). in a Venezuelan could forest habitat.

Journal of Insect Science | www.insectscience.org 15 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández International Journal of Tropical Insect Science 29: 3-10. White DS. 1978. Aquatic Coleoptera. In: Merritt RW, Cummins KW, Editors. An Silva AGDA, Gonçalves CR, Monteiro introduction to the aquatic insects of North Galvão D, Gonçalves AJL, Gomes J, Do America, 20: 399-473. Kendall and Hunt Press. Nacimento Silva M, De Simoni L. 1968. Quarto catálogo dos insectos que vivem nas Zaragoza CS. 1974. Coleópteros de algunas plantas do Brasil, seus parasitos e predatores. epífitas y doce nuevos registros de Ministerio da Agricultura, Departamento de especies para la fauna Mexicana. Anales del Defensa a Inspecão Agropecuaria, Serviço de Instituto de Biología, Universidad Nacional Defensa Sanitaria Vegetal, Laboratorio Autónoma de México (Serie Zoología) 45: Central de Patología Vegetal. Río de Janeiro, 111-118. GB, Brazil.

SMN. 1981. Estadísticas climatológicas. 1961- 1970. Servicio Meteorológico Nacional, Fuerza Aérea Argentina. Available online, www.smn.gov.ar

SMN. 1986. Estadísticas climatológicas. 1971- 1980 Servicio Meteorológico Nacional, Fuerza Aérea Argentina. Available online, www.smn.gov.ar

Spangler PJ. 1980. Chelonariid larvae, aquatic or not? (Coleoptera: Chelonariidae). Coleopterists Bulletin 34: 105-114. Figure 1. Location of sampling sites in Argentina, South America. Subtropical area, INP: Iguazú National Park (Misiones Province); Stribling JB, Young DK. 1990. Descriptions of CNP: Chaco National Park (Chaco Province); Temperate area, PLP: Punta Lara Provincial Park; ETP: “Ernesto Tornquist” Provincial Park the larva and pupa of Flavohelodes thoracica (Buenos Aires Province).High quality figures are available online. (Guérin-Méneville) with notes on a phytotelma association (Coleoptera: Scirtidae). Proceedings of Entomological Society of Washington 92: 765-770.

Vargas, L. 1928. Ein interessanter biotope der bioconöse von wasser organismen. Biologisches Zentralblatt 48: 143-162.

Von Ellenrieder N, Fernández LA. 2000. Aquatic coleoptera in the Subtropical- Pampasic ecotone (Argentina, Buenos Aires): species composition and temporal changes. Coleopterists Bulletin 54: 23-35. Figure 2. Eryngium horridum Malme (Apiaceae) at the field site in Sierra de la Ventana, Buenos Aires Province. High quality figures are available online.

Journal of Insect Science | www.insectscience.org 16 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández

Figure 3. Eryngium aff. serra Cham. and Schltdl. (Apiaceae) at the Figure 4. Guadua chacoensis (Rojas) Londoño and Peterson field site in Sierra de la Ventana, Buenos Aires Province. High quality (Poaceae) at the forest site in Iguazú National Park, Misiones figures are available online. Province. High quality figures are available online.

Figure 5. Internodes of Guadua chacoensis (Rojas) Londoño and Peterson (Poaceae) with a hole in the wall. High quality figures are Figure 6. Stumps of Guadua chacoensis (Rojas) Londoño and available online. Peterson (Poaceae). High quality figures are available online. Journal of Insect Science | www.insectscience.org 17 Journal of Insect Science: Vol. 11 | Article 147 Campos and Fernández

Figure 8. Fallen floral bracts of the palmetto Euterpe edulis Martius (Arecaceae) at the field site in Iguazú National Park, Misiones Province. High quality figures are available online.

Figure 7. Aechmea distichantha Lemaire (Bromeliaceae) at the field site in Iguazú National Park, Misiones Province. High quality figures are available online.

Figure 9. Number of species of coleopterans by class of phytotelmata from subtropical and temperate Argentina. High quality Figure 10. Habitus of larva of Pachyteles sp. (Carabidae), A: dorsal ; figures are available online. B: lateral. High quality figures are available online. Journal of Insect Science | www.insectscience.org 18