4 Insects and Spiders — Update 2014 Paul Hanson In the last 15 years there have been a number here, there is still much to learned about the of publications on the insects of cloud forests, arthropods of cloud forests. and it is impossible to include all of them in this update. In particular, it has not been possible to Small insect orders compile all the taxonomic publications that For dragonflies and damselflies (Odonata) include species occurring in Monteverde, a there is a very useful website that provides keys compilation that would be useful since and descriptions of the families, as well as identification is the principal impediment to photos of all the species known from studying insects. Nonetheless, I have attempted Monteverde (Haber 2014). to mention the most important taxonomic For the order Orthoptera there are two publications (of an order, family or subfamily) general publications, one on katydids as well as some examples of specific (Tettigoniidae: subfamily Conocephalinae) investigations. (Naskrecki 2000) and another on grasshoppers As in Hanson (2000), this update is organized (suborder Caelifera) (Rowell 2013), which by taxonomic group. Nonetheless, some provide valuable resources for future research on publications deal with distinct groups and these groups of insects. therefore do not fit this organization, for example the studies of arthropods associated True bugs, spittlebugs, leafhoppers, with epiphytes (Yanoviak et al. 2004, 2006). treehoppers, etc. (order Hemiptera) Before commencing with the individual groups, The vast majority of true bugs (suborder two general comments should be made. First, the Heteroptera) are predatory or phytophagous, but estimations of species numbers in each order are species in the subfamily Triatominae too low, but updated estimations are not (Reduviidae) suck blood from vertebrates. In attempted here. Second, it is important to Costa Rica, one species, Triatoma dimidiata, is emphasize that despite the advances reported the vector of Chagas disease. One of the few studies of this species in the wild was carried out in Monteverde (Salas Peña 2010), where it was Butterflies and moths (order Lepidoptera) found principally on trunks of live trees, although it was more abundant in residential For butterflies (superfamily Papilionoidea) areas. there is a website providing a list of species With respect to the suborder known from Monteverde, with more detailed Auchenorrhyncha, the principal publication in information and photos of the glasswing recent years is one on treehoppers butterflies (Nymphalidae: Ithomiini) (Haber (Membracidae), which includes a guide for 2001). identification, photos of all the genera, and In Hanson (2000) there is a summary of the summaries of their biology (Godoy et al. 2006). biology of Manataria maculata (Nymphalidae- This book should greatly facilitate future studies Satyrinae; now classified as a subspecies of M. of these gaudy insects. A publication on the hercyna), which lays eggs on bamboo in the spittlebugs (Cercopidae) associated with Pacific lowlands (Murillo & Nishida 2003), but pastures (Thompson & León-González 2005) spends most of the year in reproductive diapause will assist in identifying Monteverde species at higher altitudes, including Monteverde. This associated with grasses. is an unusual butterfly in that it is crepuscular Recently, a study was carried out in and uses different communal roosts—protected Monteverde which examined the responses of sites near the ground during the day and in the six species of Auchenorrhyncha to a model of a canopy at night, probably to avoid birds during redstart (Myioborus, Parulidae), a bird species the day and mice during the night (Hedelin & that hunts insects by first frightening and then Rydell 2007). Its Vogel’s organ, situated at the pursuing them. Two Membracidae (with base of the front wing, detects ultrasounds of structural defenses) were the most sensitive, insectivorous bats, allowing this butterfly to whereas two Cixiidae (defended by camouflage) respond with evasive flight (Rydel et al. 2003). were the last to flee; two Cercopidae (with Although Vogel’s organ is found in the majority aposematic coloration) showed an intermediate of Satyrinae, this is the first case where response (Galatowitsch & Mumme 2004). sensitivity to ultrasound has been demonstrated (other Satyrinae probably use this organ to Beetles (order Coleoptera) detect the sounds associated with birds in flight). Some general works that include aids for As mentioned in Hanson (2000), the only identification and summaries of biology are Satyrinae in Monteverde with transparent wings available for Staphylinidae (Navarette-Heredia is Cithaerius pireta (cited as C. menander). et al. 2002), Scarabaeidae-Dynastinae (Ratcliffe Recently, the larval host plant of this species has 2003), Scarabaeidae-Cetoniinae (Solís 2004), been discovered, Philodendron herbaceum Chrysomelidae-Cassidinae (Chaboo 2007) and (Araceae) (Murillo-Hiller 2009). Chrysomelidae-Chrysomelinae (Flowers 2004). Most families of Lepidoptera are One of the largest families of beetles is microlepidopterans, but their biology is Curculionidae, which includes the phytophagous relatively poorly known. The larvae of many weevils. Two groups that are found in species are leaf miners or leaf rollers. The larvae Monteverde and which have been the subjects of of a few species induce galls on plants; for research in recent years are members of the example, an undescribed species of Momphidae subfamily Baridinae associated with Piperaceae produces quite large stem galls on Conostegia (Prena 2010) and those in the tribe Derelomini oerstediana (Melastomataceae). (now known as Acalyptini; subfamily Curculioninae). Various members of the latter Flies (order Diptera) group are pollinators of palms, but some species Of all the principal insect orders, the have changed host plants and can be pollinators inventory of flies is the most advanced, both in of Cyclanthaceae and Anthurium (Araceae) Costa Rica as a whole and in cloud forests in (Franz 2006). particular. The two volumes by Brown et al. (2009, 2010) provide keys for the identification of fly genera and summaries of our knowledge of each genus. This valuable work opens doors attached to its abdomen (Blanco & Barboza for future research on flies. 2005). Many of the specialists who collaborated in the production of the two volumes mentioned Wasps, ants and bees (order above are currently carrying out an inventory of Hymenoptera) the flies found in a cloud forest at Zurquí de There are two books in Spanish on the Moravia (Zurqui All-Diptera Biodiversity Hymenoptera of the Neotropical region, one that Inventory, ZADBI 2014). Although this cloud emphasizes identification (Fernández & Sharkey forest is located in a different mountain range, 2006) and the other with detailed summaries of the results of this project will be very applicable the biology of the order (Hanson & Gauld 2006). to the Monteverde cloud forest. When the results Although these books cover the entire region, of the Zurqui project become available we they provide a good introduction for studies of should have a significantly better understanding Monteverde hymenopterans. of the fly fauna of Costa Rican cloud forests. The vast majority of hymenopterans are The biology of fly larvae is extremely parasitoids of other insects, but in recent years diverse. This order includes the most species- several species have been discovered that have rich family of gall-inducers, Cecidomyiidae made an evolutionary transition to phytophagy. (Hanson & Gómez-Laurito 2005). Although For example, in Monteverde the larvae of there is a large diversity of plant galls in Eurytoma werauhia (Eurytomidae) feed on the Monteverde, there are very few studies of these floral buds of Werauhia gladioliflora insects. In large part this is due to the fact that (Bromeliaceae) (Gates & Cascante-Marín 2004). Cecidomyiidae is probably the largest family of In the Central Valley of Costa Rica the larvae of insects and at the same time probably harbors Allorhogas conostegia (Braconidae) induce galls the greatest proportion of undescribed species in the fruits of Conostegia xalapensis (more than 99%). Other phytophagous fly larvae (Melastomataceae), but it is possible that in include leaf miners, for example, two species of Monteverde another (undescribed) species of the Agromyzidae on Bocconia frutescens same genus produces these galls (Chavarría et (Papaveraceae) (Boucher & Nishida 2014). al. 2009a). This deserves more research, and if The larvae of other flies are predators, one of this hypothesis is correct, it could represent an the best studied families being Syrphidae. The interesting case of speciation. literature gives the impression that nearly all The wasps that are best known to the general predatory syrphid larvae feed on aphids and public are the eusocial wasps which construct other Sternorrhyncha (Hemiptera) on plants. paper (carton) nests (Vespidae, subfamily Nonetheless, the biology of predatory syrphids is Polistinae). The best studied species in probably much more diverse in the Neotropics. Monteverde is Polybia aequatorialis (O'Donnell For example, Ocyptamus luctuosus, a species et al. 2004, Jones et al. 2009), which is the only found in Monteverde, is a predator in the water species of Polistinae that occurs at very high that accumulates in epiphytic bromeliads elevations (more than 3000 m), where they have (Rotheray et al. 2000). Ocyptamus is one of the enormous colonies (Chavarría