Entiminae Schoenherr, 1823 503 3.7.5 Entiminae Schoenherr, 1823 latithorax Pierce (Tisoc-Dueñas 1989), seven in Ectemnorhinus spp. (Chown & Scholtz 1989 a) and Adriana E. Marvaldi, Analía A. Lanteri, M. Guadalupe up to 11 in Naupactus leucoloma Boheman (Mat- del Río and Rolf G. Oberprieler thiessen 1991). Females of Entiminae (as well as those of other Distribution. The Entiminae include about 1370 broadnosed weevils classified in Cyclominae) do genera and more than 12,000 species worldwide, not use their rostrum for preparing an oviposi- occurring in all biogeographical regions (Nearc- tion site. Two main oviposition types occur in tic, Palaearctic, Afrotropical, Oriental, Austral- Entiminae: eggs deposited loosely and at random asian, Neotropical and Chile/southern Argentina). (the “Sitona type”) and eggs deposited in batches Among the 55 tribes recognized in the catalogue between adjoining surfaces (the “Brachyderes type”) by Alonso-Zarazaga & Lyal (1999), about 40 are (van Emden 1950, 1952; Marvaldi 1999). Eggs laid represented mainly in a restricted region, about randomly are usually placed on plants, surface lit- ten occur in two biogeographical regions [e.g., ter or soil, and they darken as they develop, whereas Nearctic and Palaearctic, Palaearctic and Orien- eggs laid in batches are covered with an adhesive tal, Afrotropical and Oriental, Afrotropical and glutinous substance secreted during oviposition, Palaearctic, Australasian and Patagonian (Chile/ hidden between adjoining surfaces (leaves, cracks southern Argentina)], and the remaining occur in the soil, crevices, litter or various niches near in more than three regions. As an example, the soil) and remain pale (Marvaldi 1999). Eggs of Agraphi ni, Hormorini and Ophryastini are typi- Ectemnorhinini are laid individually or in small cally Nearctic; Holcorhinini, Otiorhynchini, groups in ground litter below the host plant or Phyllobiini and Sciaphilini are Palaearctic; Any- among plant parts, and they darken as they develop potactini, Entimini, Eudiagogini, Eustylini and (Chown & Scholtz 1989 a). In Pachyrhynchini Lordopini are Neotropical; Embrithini, Oosomini (e.g., Pantorhytes Faust) they are laid singly in the and Tanyrhynchini are Afrotropical; Mesostylini, bark (van Emden 1952; May 1978). When laid in Nastini and Omiini are Oriental; Celeuthetini batches, each batch (Fig. 3.7.5.1 D) consists on an and Ottistirini are Australasian; and Anomoph- agglutinated cluster of 20–80 eggs, a number that thalmini are endemic to Patagonia. Some tribes can vary depending on the food resources available have more restricted ranges, such as Nothog- to the female, time after its eclosion and other envi- nathini, which only occur in India, Ophtalmor- ronmental conditions. The fecundity of entimines rhynchini in Central Africa, Premnotrypini in the is very high compared with that of other weevil high Andes of northern South America, Typhlo- subfamilies. Eggs hatch after 5–20 days, with the rhinini in Madagascar, Ectemnorhinini on small length of the egg stage being affected mainly by islands in the southern Ocean (from Marion and temperature and humidity. Prince Edward Islands to Heard Island), Elytru- Soil-dwelling larvae and polyphagy are char- rini in Polynesia, Laparocerini on the Macarone- acteristic features of the Entiminae, most of them sian islands (Azores, Madeira and Canary Islands), feeding on a variety of plants (spanning different Polycatini in the Philippines and Rhyncogonini angiosperm families) in the adult and larval stages. on various islands of the eastern Pacific (Mar- However, larvae of Ectemnorhinini and Pachy- quesas, Hawaii and Tahiti). One extinct tribe is rhynchini do not feed on roots in the soil. Larval described from the Upper Miocene of Germany. development in the Ectemnorhinini occurs on or [O’Brien & Wibmer 1978; Alonso-Zarazaga & Lyal near the soil surface, usually among plant mats or 1999; Yunakov & Nadein 2006.] detritus (Chown & Scholtz 1989 a), whereas pachy- rhynchine larvae feed arboreally, tunneling in Biology and Ecology (Fig. 3.7.5.1 A–D). Larvae branches, where they also pupate in a chamber lined (Fig. 3.7.5.1 A) of most species of Entiminae live with plant fibers and frass (May 1978). The large freely in the soil, feeding externally on the roots pupal cases of Leptopius duponti (Boisduval) (Lepto- of their host plants. Pupation (Fig. 3.7.5.1 B) also piini) formed in calcareous sands along the South occurs in the soil, in an earthen cell lined with a Australian coast can become calcified after eclosion larval secretion. Adults (Fig. 3.7.5.1 C) feed on the of the weevil and preserved for a long time (Lea aerial green parts of the plants, especially fresh 1925; Tilley et al. 1997). Similar bauxitic pisoliths leaves or flowers, cutting their edges in a charac- from northern Queensland have been suggested to teristic “notching” pattern. Life cycles usually last represent entimine pupal cases as well (Tilley et al. about a year but in some cases are completed in 1997; Eggleton & Taylor 2008), although without two or three years (Young et al. 1938; May 1994). direct evidence of any weevil constructing them. Larvae overwinter underground (usually as pre- Entiminae are primarily and ancestrally associ- pupae) and pupate in warmer months, and the ated with angiosperm plant taxa (Marvaldi et al. teneral adults emerge from the soil during spring 2002). Associations with gymnosperms or crypto- and summer. The longest part of the cycle is spent gams most likely constitute secondary host shifts in the larval stage, which is the more injurious for and/or, in polyphagous species, expansions of the the plants. The number of instars varies among original host range. Ectemnorhinini mostly feed on species, for intance, four occur in Premnotrypes cryptogams, both as adults and larvae, but some also Authenticated | [email protected] Download Date | 5/8/14 1:22 PM 504 Adriana E. Marvaldi, Analía A. Lanteri, M. Guadalupe del Río and Rolf G. Oberprieler Fig. 3.7.5.1 Life cycle of Entiminae. A, Naupactus leucoloma Boheman, mature larva in soil; B, N. leucoloma, pupa in earth cell; C, N. cervinus Boheman, adult feeding; D, Atrichonotus taeniatulus Berg, posture. feed on bryophytes, lichens, algae and cyanobacteria Rosaceae, Rutaceae, Solanaceae and many others. (Chown 1989, 1994; Chown & Scholtz 1989 a). Legumes are among their main hosts. It is common Members of the genera Canonopsis C. O. Waterhouse, among polyphagous species that adult and larval Christensenia Brink and Ectemnorhinus G. R. Water- stages feed on different plant taxa, i.e., adults on house are the only Ectemnorhinini that feed on leaves of various dicotyledons and larvae on roots of angiosperms, but they also incorporate bryophytes monocotyledons (pastures) (Marvaldi 1998 b). The and other cryptogams in their diet (Chown 1989). frequency of the host shifts depends not only on The only exception is Palirhoeus eatoni (C. O. Water- the physiology of plants and weevils but also on the house) which is oligophagous, restricted to feeding availability (plant apparency) of these plants. For on three species of marine algae (Chown 1994). A dif- instance, adults of Pantomorus ruizi (Brèthes) (Nau- ferent association with cryptogams occurs in some pactini), known to feed on various dicotyledons, species of Gymnopholus Heller (Leptopiini) and Pan- have recently been found consuming pine needles torhytes (Pachyrhynchini) inhabiting humid forests in Patagonia, whereas the larvae probably feed on at high elevations in New Guinea. These weevils live roots of grasses growing around the trees (Gómez in an epizoic symbiosis with cryptogamic plants and & Lanteri 2006), and adults of the Palaearctic Otio- microfauna, the lichens, algae and mosses growing rhynchus kollari Gyllenhal were found feeding on on their pronotum and elytra providing camouflage ferns (Muñiz 1970). and protection for the weevils and food and shel- Even when polyphagous, entimines can show ter for the rotifers, nematodes and mites that live definite host preferences (oligophagy). South Ame- among them (Gressit 1966). rican species of Pandeleteius Schoenherr (Tanyme- Entiminae feed on monocotyledons (e.g., Poa- cini) are predominantly associated with trees and ceae) and a great variety of dicotyledonous fami- shrubs of the families Anacardiaceae (Schinopsis and lies, including Fabaceae, Fagaceae, Malvaceae, Schinus), Asclepiadaceae (Baccharis) and Fabaceae Authenticated | [email protected] Download Date | 5/8/14 1:22 PM Entiminae Schoenherr, 1823 505 (Prosopis), and those of Enoplopactus Heller (Naupac- range and the ability to develop on most gymno- tini) with shrubs of the subfamily Zygophyllaceae sperms and broad-leaved plants have enabled this (Larrea) (Lanteri et al. 2002). Phyllobius oblongus Lin- beetle to establish itself in nurseries, greenhouses naeus (Phyllobiini) is much more common on elms and landscapes around the world (Moorhouse et al. (Ulmus, Ulmaceae) and blackthorn (Prunus spinosa, 1992; van Tol et al. 2004). In Australia it has been Rosaceae) than on other trees (Morris 1976). Spe- introduced over a decade ago and, together with cies of Sitona Germar are much more stenophagous the cribrate weevil – O. cribricollis Gyllenhal, and than most Entiminae, being mainly associated with the Rough Strawberry Weevil – O. rugosostriatus legumes (Fabaceae), the adults feeding on leaves (Goeze), is considered a pest of various berries, veg- and the larvae on roots and nitrogen-fixing root etables and fruit