Encyclopedia of Entomology Encyclopedia of Entomology

Edited by John L. Capinera University of Florida

Second Edition

Volume 4 S–Z Professor John L. Capinera Dept. Entomology and Nematology University of Florida Gainesville FL 32611–0620 USA

Library of Congress Control Number: 2008930112

ISBN: 978-1-4020-6242-1 This publication is available also as: Electronic publication under ISBN 978-1-4020-6359-6 and Print and electronic bundle under ISBN 978-1-4020-6360-2

This work is subject to copyright. All rights are reserved, whether the whole or part of the material is con- cerned, speciically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, repro- duction on microilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its cur- rent version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law.

© 2008 Springer Science+Business Media B.V.

The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a spe- ciic statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. springer.com

Editor: Zuzana Bernhart, Dordrecht/ Sandra Fabiani, Heidelberg Development Editor: Sylvia Blago, Heidelberg Production Editor: le-tex publishing services oHG, Leipzig Cover Design: Frido Steinen-Broo, Spanien

Printed on acid-free paper SPIN: 11757993 2109 — 5 4 3 2 1 0 Editorial Board

Cyrus Abivardi Eugene J. Gerberg Swiss Federal Institute of Technnoloy University of Florida Donald R. Barnard Donald W. Hall United States Department of Agriculture University of Florida Jean-Luc Boevé Marjorie A. Hoy Royal Belgian Institute of Natural Sciences University of Florida Drion Boucias John B. Heppner University of Florida Florida State Collection of Arthropods Paul. M. Choate Pauline O. Lawrence University of Florida University of Florida Whitney Cranshaw Heather J. McAuslane Colorado State University University of Florida Thomas C. Emmel James L. Nation University of Florida University of Florida J. Howard Frank Herb Oberlander University of Florida United States Department of Agriculture Severiano F. Gayubo Frank B. Peairs Universidad de Salamanca Colorado State University Acknowledgments

his project is the labor of many people, including some who labored diligently behind the scenes. Among those to whom I am greatly indebted for their ‘behind-the-scenes’ assistance are Pam Howell and Carole Girimont (irst edition) and Pam Howell (second edition) for document processing and editing assistance; Mike Sanford, Pat Hope, and Jane Medley (irst edition) and Hope Johnson (second edition) for assistance with the images, and Marsha Capinera for compiling the list of contributors. Ron Cave, Andrei Sourakov, and Lyle Buss helped greatly by supplying numerous photographs for the second edition. Howard Frank deserves special mention for his editing acumen and assistance. Drion Boucias con- tributed the lengthy unattributed sections on insect pathology. he unattributed biographic sketches with last names beginning with A to J were contributed by Howard Frank. All other unattributed sections were contributed by John Capinera. Preface

Some biologists have called this the ‘Age of Insects.’ Among animals, certainly the diversity of insects is unrivaled. Nearly one million species have been described to date, and some entomologists estimate that as the tropics are fully explored, we will ind that there are actually more than three million insect spe- cies. he large number of insects is oten attributed to the divergence of plants (angiosperms), which provide numerous hosts and places to feed, but if plant feeders are excluded from the tabulation the biodiversity of insects remains unrivaled. Virtually every environment has been exploited by these resil- ient organisms. Even if one dislikes insects, they are impossible to ignore, and a little knowledge about them could be indispensable should one have a ‘close encounter’ of an unpleasant kind. Insects are remarkable biological organisms. hey are small enough to escape the detailed scrutiny of most people, but I have yet to meet anyone whom, once provided the opportunity to examine insects closely (through a microscope) is not completely amazed by the detail and complexity of these exqui- sitely designed (by natural selection) beasties. hey are fascinating in function as well as form. Insects are the only invertebrates to ly, they are disproportionately strong, and their ecological adaptability deies belief. For example, some insects produce their own version of anti-freeze, which allows them to be fro- zen solid yet to regain normal function upon thawing. heir sensory abilities are beyond human com- prehension; a male insect can sometimes locate a female by her ‘perfume’ (pheromone) from several kilometers distance. Although not normally considered intelligent, insects display surprisingly complex behaviors, and altruistic social systems that could well serve as models for human societies. Insects and their close relatives are important for many reasons besides their sheer diversity. heir efect on humans is profound. Insects are our chief competitor for food and iber resources throughout the world. Annual crop losses of 10 to 15% are attributed to insects, with both pre-harvest and post- harvest losses considerably more at times. Insects also are the principal vector of many human, animal, and plant diseases, including viruses, mollicutes, bacteria, fungi, and nematodes. he ability to transmit diseases magniies their efect, and makes it more diicult to manage injury. Over the course of human history, insect-transmitted disease has caused untold human sufering. For example, introduction of lea-transmitted bubonic plague to Europe centuries ago killed millions of people and caused severe disruption to western civilization. hough less dramatic, mosquito-transmitted malaria kills thousands annually throughout the world, and unlike plague, which is now mostly a historical footnote, the toll continues to mount. Advances in technology, particularly the introduction of chemical insecticides, have done much to remove the threat of insect-related damage from the consciousness of most humans. Insecticides are applied preventatively to avoid pre- and post-harvest damage to crops, to our dwellings, and to our land- scape. his is an ot-overlooked but remarkable achievement that has increased stability in the supply and price of resources, and in the lives of resource producers. No longer are people faced with starvation or economic ruin due to the ravages of insects; in almost all parts of the world, the ready availability of insecticides can be used to prevent massive insect population outbreaks. However, we realize increas- ingly that this approach is not without its own set of health, environmental and economic costs, and alleviating dependency on insecticides, or making alternatives to insecticides more readily available, has assumed greater priority. x Preface

We are faced with an interesting dichotomy. here is a wealth of information about insects, but it is known mostly to ‘insect scientists’ (entomologists). he public (non-entomologists or 99.99% of all peo- ple) has little knowledge about insects, and poor access to vital information about these important organisms. So this encyclopedia is presented to bridge the gap – to better enable those with a need to know to ind fundamental information provided by more than 450 experts in the ield of entomology. We provide a broad overview of insects and their close relatives, including taxonomy, behavior, ecology, physiology, history, and management. Importantly, we provide critical links to the entomological litera- ture, much of which presently is unavailable for search electronically. he contributors are distinguished entomologists from around the world. hey hope that the availability of this encyclopedia will help oth- ers to reap the beneits of centuries of discovery, and to discover the wonders that make the study of insects so compelling. It was constructed with college and university students in mind, but others may ind it a handy reference.

John L. Capinera, Gainesville (Florida) April, 2008 Coffee Pests and their Management C 961 Coffee Pests and their predatory organisms, which regulate the popula- Management tions of many pests, represented 42% of the total of species collected in a cofee plantation. For this Juan f. Barrera reason, it is important to protect and maintain El Colegio de la Frontera Sur, Tapachula, Chiapas, the natural enemies of pests, avoiding the indis- Mexico criminate use of chemical pesticides and some agronomic practices that are harmful to natural he perennial and evergreen nature of the cofee control. he goal of this section is to describe the plant (Coffea spp.) favors attack by a number of biological and ecological characteristics of the insects and mites (Table 17, Figs. 73 and 74). All main insects and mites of C. arabica L. and portions of the plants are susceptible to attack, C. canephora Pierre ex Froehner, the damage and damage may appear at the seed bed, nursery, caused by these pests, their natural enemies, and plantation, or in the warehouse. Certain pests pest management in cofee growing countries of afect the cofee plant only temporarily, while oth- tropical America. he pests to be described are ers live for several generations on the plant. In listed in Table 17, which also includes the parts of some instances, the attack may cause the death of the plant that are damaged and the development the plant, but in most cases the pests only weaken stage of the cofee plant that they damage. he cri- the plant, reducing yield. When the bean is terion applied to include these organisms in the attacked, quality also may be afected. category of “major pests,” was that they were Insects constitute the most numerous group reported in at least one of the manuals on cofee of cofee pests; of more than 850 species of insects pests that have been published in Brazil, Colom- that feed on cofee in the world, approximately bia, Costa Rica, Cuba, El Salvador, Guatemala, 200 (23.5%) have been reported in the tropical Honduras, Jamaica, Mexico or Venezuela. and sub-tropical areas in America. Out of these, hardly thirty species, mostly indigenous, cause losses considered important. he pests and the Coffee Berry Borer, Hypothenemus seriousness of the problems they cause vary from hampei (Ferrari) (Coleoptera: one country to another, and from one area to Curculionidae: Scolytinae) another. he cofee pest that is considered the most important in tropical America is the cofee Distribution berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Curculionidae: Scolytinae), now cos- his is the most serious insect pest of cofee mopolitan but originating in Africa. he cofee worldwide. It originated in Africa. In the Americas, leaf miner, Leucoptera coffeella Guérin-Méneville it is found in cofee plantations from Mexico to (Lepidoptera: Lyonetiidae), and the root mealy- Brazil, including some countries in the Caribbean bugs (Pseudococcidae) are causing serious prob- region such as Cuba, Jamaica, the Dominican lems in several countries. Bugs of the genus Republic and Puerto Rico. Antestiopsis (Pentatomidae), which are very harm- ful in Africa, have not yet been reported in the American hemisphere. Damage and Economic Importance Most of the insects that are found in cofee plantations are beneicial because they contrib- Cofee berry borer (Fig. 73) is a direct pest ute to plant pollination, degrade organic matter, because it causes direct damage to the product to or feed on phytophagous organisms. A study be harvested, the cofee bean. he attacked green, conducted in Mexico showed that parasitic and ripe and dry fruits or berries usually show a hole 962 C Coffee Pests and their Management Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of cof- fee in tropical America Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Acari: Tarsonemidae Polyphagotarsonemus latus Brazil Nymph, adult Leaves (Banks) Acari: Tenuipalpidae Brevipalpus sp. Brazil, Jamaica, Mexico Nymph, adult Leaves Acari: Tetranychidae Olygonychus coffeae Guatemala, Mexico Nymph, adult Leaves (Nietner) Olygonychus ilicis (McGregor) Brazil, Guatemala Nymph, adult Leaves Olygonychus punicae (Hirst) El Salvador Nymph, adult Leaves Olygonychus yothersi Colombia, Costa Rica, Nymph, adult Leaves (McGregor) Venezuela Coleoptera: Anthribidae Araecerus fasciculatus All coffee growing coun- Larva, adult Bean (DeGeer) tries in America Coleoptera: Cerambycidae Plagiohammus maculosus Costa Rica, El Salvador, Larva Stem, root (Bates) Guatemala, Honduras, Mexico Plagiohammus mexicanus Mexico Larva Stem, root Breuning Plagiohammus spinipennis Mexico Larva Stem, root (Thomson) Coleoptera: Curculionidae Brachyomus quadrinodosus Venezuela Adult Leaves (Lacordaire) Cleistolophus similis Sharp Costa Rica Adult Leaves Compsus sp. Colombia Adult Leaves Epicaerus capetillensis Sharp Guatemala, Honduras, Adult Leaves Mexico Hypothenemus hampei Mexico to Brazil, including Larva, adult Fruit, bean (Ferrari) Cuba, Jamaica, Dominican Republic, and Puerto Rico Lachnopus buchanani Cuba Adult Leaves Marshall Macrostylus boconoensis Colombia, Venezuela Adult Leaves Bordón Pantomorus femoratus Sharp Costa Rica Adult Leaves Coffee Pests and their Management C 963 Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of coffee in tropical America (Continued) Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Pantomorus godmani Brazil Adult Leaves Crotch Steirarrhinus sp. Costa Rica Adult Leaves Xylosandrus morigerus Mexico to Brazil Larva, adult Young stems, (Blandford) branches Coleoptera: Scarabaeidae Anomala sp. El Salvador Larva Root Dyscinetus picipes Burmeister Cuba Larva Root Phyllophaga spp. Widely distributed in coffee Larva Root plantations in America Phyllophaga latipes (Bates) El Salvador Larva Root Phyllophaga menetriesi El Salvador Larva Root (Blanchard) Phyllophaga obsoleta El Salvador Larva Root (Blanchard) Phyllophaga sanjosecola Costa Rica Larva Root Saylor Phyllophaga vicina Moser Costa Rica Larva Root Hemiptera: Aphididae Toxoptera auranti (Boyer de Tropical and sub-tropical Nymph, adult Leaves, buds and Fonscolombe) areas of the Old World. other tender parts of Widely distributed in coffee the plant plantations in America Hemiptera: Coccidae Coccus spp. Mexico Nymph, adult female Aerial part of the plant Coccus hesperidum L. Guatemala, Mexico Nymph, adult female Aerial part of the plant Coccus viridis (Green) Brazil, Colombia, Costa Nymph, adult female Aerial part of the Rica, Cuba, Ecuador, El plant Salvador, Guatemala, Honduras, Jamaica, Mexico, Puerto Rico, Surinam, Venezuela Parasaissetia sp. Colombia Nymph, adult female Aerial part of the plant Parasaissetia nigra (Nietner) El Salvador, Guatemala, Nymph, adult female Aerial part of the Puerto Rico, West Indies plant Saisettia spp. El Salvador, Mexico Nymph, adult female Aerial part of the plant 964 C Coffee Pests and their Management Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of coffee in tropical America (Continued) Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Saisettia coffeae (Walker) Brazil, Costa Rica, Cuba, El Nymph, adult female Aerial part of the Salvador, Guatemala, plant Honduras, Mexico, Dominican Republic, Venezuela Saisettia olae (Oliver) Brazil, Cuba, Guatemala, Nymph, adult female Aerial part of the Mexico plant Toumeyella sp. Venezuela Nymph, adult female Root Toumeyella liriodendri Guatemala Nymph, adult female Root (Gmelin) Hemiptera: Cerococcidae Cerococcus catenarius Brazil Nymph, adult female Aerial part of the Fonseca plant Hemiptera: Diaspididae Chrysomphalus sp. Guatemala Nymph, adult female Aerial part of the plant Chrysomphalus dictyospermi Guatemala Nymph, adult female Aerial part of the (Morgan) plant Ischnaspis longirostris Colombia, Cuba, Nymph, adult female Aerial part of the (Signoret) Guatemala plant Lepidoshaphes beckii Venezuela Nymph, adult female Aerial part of the (Newman) plant Selenaspidus articulatus Colombia, Ecuador, Mexico Nymph, adult female Aerial part of the (Morgan) plant Hemiptera: Margarodidae Icerya purchasi Maskell Venezuela Nymph, adult female Aerial part of the plant Hemiptera: Ortheziidae Insignorthezia insignis Browne Brazil, Colombia Nymph, adult female Aerial part of the plant Praelongorthezia praelonga Brazil Nymph, adult female Aerial part of the (Douglas) plant Hemiptera: Pseudococcidae Brevicoccus sp. Guatemala Nymph, adult female Root Ceroputo sp. Costa Rica Nymph, adult female Root Dysmicoccus sp. Colombia, Ecuador Nymph, adult female Root Dysmicoccus bispinosus Brazil, Guatemala, Hondu- Nymph, adult female Root (Beardsley) ras, Mexico Dysmicoccus brevipes Costa Rica, El Salvador, Nymph, adult female Root (Cockerell) Guatemala, Honduras, Mexico Coffee Pests and their Management C 965 Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of coffee in tropical America (Continued) Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Ferrisia virgata (Cockerell) Brazil, Mexico, West Indies Nymph, adult female Aerial part of the plant Geococcus sp. Mexico, Venezuela Nymph, adult female Root Geococcus coffeae Green El Salvador, Guatemala, Nymph, adult female Root Honduras, Surinam Geococcus radicum Green El Salvador Nymph, adult female Root Neochavesia caldasiae Colombia Nymph, adult female Root (Balachowsky) Rhizoecus sp. Mexico, Venezuela Nymph, adult female Root Rhizoecus andensis Hambleton Colombia Nymph, adult female Root Rhizoecus coffeae Laing Costa Rica Nymph, adult female Root Paraputo sp. Guatemala Nymph, adult female Root Planococcus citri (Risso) Brazil, Colombia, Costa Nymph, adult female Root, aerial part of Rica, Cuba, El Salvador, the plant Guatemala, Honduras, Jamaica, Mexico, Puerto Rico Planococcus halli Ezzat & Guatemala Nymph, adult female Root McLonnell Pseudococcus elisae Guatemala Nymph, adult female Root (Borchsenius) Pseudococcus longispinus Guatemala Nymph, adult female Root (Targioni-Tozzeti) Pseudococcus jongispinus Mexico Nymph, adult female Aerial part of the Targioni-Tozzetti plant Puto sp. Costa Rica Nymph, adult female Root Puto antioquensis (Murillo) Guatemala Nymph, adult female Root Rhizoeccus campestris Guatemala Nymph, adult female Root (Hambleton) Rhizoeccus caticans Guatemala Nymph, adult female Root (Hambleton) Rhizoeccus kondonis Kuwana Guatemala Nymph, adult female Root Rhizoeccus nemoralis El Salvador, Honduras Nymph, adult female Root Hambleton Hymenoptera: Formicidae Acromyrmex spp. Venezuela Adult Leaves Acromyrmex coronatus (F.) Brazil Adult Leaves Acromyrmex octospinosus Trinidad Adult Leaves (Wheeler) 966 C Coffee Pests and their Management Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of coffee in tropical America (Continued) Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Atta spp. Guatemala, Ecuador, Adult Leaves Nicaragua, Venezuela Atta cephalotes (L.) Colombia, Costa Rica, Adult Leaves Mexico, Surinam, Trinidad Atta fervens Say Mexico Adult Leaves Atta insularis Guérin- Cuba Adult Leaves Méneville Atta laevigata Smith Brazil Adult Leaves Atta mexicana (Smith) Guatemala, Mexico Adult Leaves Atta sexdens (L.) Brazil Adult Leaves Atta sexdens rubropilosa Brazil Adult Leaves Forel Lepidoptera: Apateloididae Olceclostera moresca Colombia Larva Leaves (Schaus.) Lepidoptera Arctiidae Estigmene acrea (Drury) Colombia Larva Leaves Lepidoptera: Dalceridae Dalcera abrasa Brazil Larva Leaves Herrich-Schaeffer Zadalcera fumata Schaus Brazil Larva Leaves Lepidoptera: Elachistidae Stenoma cecropia Meyrick Colombia Larva Leaves Lepidoptera: Geometridae Glena sp. Brazil Larva Leaves Oxydia spp. Colombia Larva Leaves Oxydia saturniata Guenée Brazil Larva Leaves Lepidoptera: Limacodidae Phobetron hipparchia Brazil, Colombia Larva Leaves (Cramer) Sibine spp. Colombia Larva Leaves Lepidoptera: Lyonetiidae Leucoptera coffeella (Guérin- Widespread wherever Larva Leaves Méneville) coffee is grown in the Neotropical area Lepidoptera: Megalopygidae Megalopyge lanata (Stoll) Brazil, Colombia Larva Leaves Podalia sp. Brazil Larva Leaves Coffee Pests and their Management C 967 Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of coffee in tropical America (Continued) Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Lepidoptera: Noctuidae Agrotis spp. Colombia, Costa Rica, Larva Stems of small plants Ecuador, El Salvador in germinating seedbeds or recently transplanted plants Agrotis ipsilon (Hufnagel) Brazil Larva Stems of small plants in germinating seedbeds or recently transplanted plants Agrotis repleta Walker Venezuela Larva Stems of small plants in germinating seedbeds or recently transplanted plants Feltia spp. Costa Rica, El Salvador, Larva Stems of small plants Venezuela in germinating seed- beds or recently transplanted plants Pseudoplusia includens Honduras Larva Leaves (Walker) Spodoptera sp. Colombia, Costa Rica, Ecua- Larva Stems of small plants dor, El Salvador in germinating seedbeds or recently transplanted plants Spodoptera eridania (Stoll) Venezuela Larva Stems of small plants in germinating seed- beds or recently transplanted plants Spodoptera frugiperda (Smith) Costa Rica, Brazil Larva Stems of small plants in germinating seedbeds or recently transplanted plants; leaves Trichoplusia ni (Hübner) Colombia Larva Leaves Lepidoptera: Psychidae Oiketicus geyeri (Berg) Brazil Larva Leaves Oiketicus kirbyi Brazil, Cuba Larva Leaves Lucas Lepidoptera: Saturniidae Automeris sp. Brazil, Colombia Larva Leaves Automeris complicata Brazil Larva Leaves Walker 968 C Coffee Pests and their Management Coffee Pests and their Management, Table 17 The most common phytophagous insects and mites of coffee in tropical America (Continued) Taxon (scientific and Country where the insect/ Developmental Plant parts affected common name) mite is reported as coffee stage feeding in/on pest the plant Automeris coresus Boisduval Brazil Larva Leaves Automeris illustris Walker Brazil Larva Leaves Eacles imperialis magnifica Brazil Larva Leaves (Walker) Eacles masoni Schaus Ecuador Larva Leaves Lonomia circunstans (Walker) Brazil Larva Leaves Orthoptera: Gryllidae Paroecanthus guatemalae Guatemala, Honduras Adult female Stem, branch Saussure Paroecanthus niger Saussure El Salvador, Guatemala Adult female Stem, branch Orthoptera: Tettigoniidae Gongrocnemis sp. Guatemala Nymph, adult Leaves, buds, fruit pulp, beans Idiarthron atrispinum (Stål) Costa Rica, Guatemala Nymph, adult Leaves, buds, fruit pulp, beans Idiarthron subquadratum Colombia, Guatemala, El Nymph, adult Leaves, buds, fruit Saussure & Pictet Salvador, Honduras, pulp, beans Mexico

on its apical portion. The hole is located at the which produces ochratoxin A, a potent toxin that center or ring of the berry’s ostiole and the emis- sometimes contaminates green cofee beans, sion of sawdust can be observed through this roasted cofee, and cofee brews, including instant hole. Its attack reduces the yield and affects the cofee. bean quality. Characteristic damage includes the rotting of developing beans as a result of sap- rophytic microorganisms that enter through the Description hole, the drop of young berries due to attack, and the loss of bean weight due to insect feeding. he he egg is elliptical, crystalline and yellowish borer can cause bean yield losses of 30–35% with toward maturity. Its length varies from 0.52 to 100% of perforated berries at harvest time; nev- 0.69 mm. he larva is white-yellowish, without ertheless, damage can be greater if harvest is legs, with a “C”-shaped body and a wide thoracic delayed. All the commercial cofee varieties and region. he head is light brown, with visible and species are attacked by this insect. However, it forward-extending mandibles. Visible hairs shows preference for C. canephora, and its multi- spread over the head and body. Females molt plication is also higher on beans of this cofee twice and males once. he length of the last lar- species. Recently it was suggested that H. hampei val instar is from 1.88 to 2.30 mm. he pre-pupa serves as a vector for Aspergillus ochraceus Wilh., is similar to the larva, but its color is milky-white, Coffee Pests and their Management C 969

Coffee Pests and their Management, Figure 73 Some coffee pests: (a) Coffee berry borer, Hypothenemus hampei (Curculionidae) infesting a coffee berry; (b) Damage of coffee leaf by coffee leaf miner, Leucoptera coffeella (Lyonetiidae); (c) Root mealybugs (Pseudococcidae); (d) Scale insects on coffee leaf (Coccidae). its body is less curved, and it does not feed. he 1.84 to 2.00 mm long. he adult is elongated with pupa is milky-white and yellowish towards a cylindrical body slightly arched towards the maturity. Many of the adult’s characteristics can end of the abdomen. It is about 1.50–1.78 mm be seen in the pupal stage. he pupa varies from long and its body is bright black, although 970 C Coffee Pests and their Management

Coffee Pests and their Management, Figure 74 Some additional coffee pests: (a) Coffee branch perforat- ed by Xylosandrus morigerus (Curculionidae); (b) Coffee stem attacked by a stem borer, Plagiohammus maculosus (Cerambycidae); (c) Aphids on coffee leaf; (d) Adults of a katydid, Idiarthron subquadratum; (Tettigoiidae) (e) Oviposition by a bush cricket, Paroecanthus (Gryllidae) on the stem of a coffee bush. Coffee Pests and their Management C 971 yellowish when emerging from the pupa. he adults ind refuge in the black, dry berries. Adult head is ventrally located and is protected by females emerge massively from these old berries the pronotum. he antennae are elbowed and with irst rainfall, initiating the infestation by clubbed at the ends. Mouthparts are the typical attacking berries from the earliest lowerings of chewing type and the elytra are convex and pos- the new harvest. sess longitudinal grooves that alternate with lon- gitudinal series of bristles. Females have well-developed wings that allow them to ly, Natural Enemies while the males’ wings are atrophied. Females are easily diferentiated from males because they Cofee berry borer is attacked by several natural are larger. enemies. Four parasitoid species from Africa are the best known: Prorops nasuta Waterston (from Cameroon, Ivory Coast, Zaire, Kenya, Tanzania, Biology and Ecology Togo, Uganda) and Cephalonomia stephanoderis Betrem (Ivory Coast, Togo) (both Hymenoptera: Adult females initiate the infestation. In general, Bethylidae), and two solitary ectoparasitoids of a berry is infested by a single female. If the cofee the larva, pre-pupa and pupa, Heterospillus coffei- bean is watery or milky, the insect tends to aban- cola Schimideknecht (Hymenoptera: Braconidae) don it and the bean usually rots. But if the bean (Cameroon, Zaire, Kenya, Tanzania, Uganda) (a consistency is hard enough, the founding female free-living wasp that deposits a single egg near a constructs a gallery where she lays the eggs. he borer’s egg cluster in a recently attacked berry) eggs are oviposited one by one, forming small and Phymastichus coffea LaSalle (Hymenoptera: groups within the cofee bean. he female lays Eulophidae) (Togo, Kenya) (a gregarious endop- from 1 to 3 eggs per day during the irst 15–20 arasitoid of H. hampei adults which parasitizes days; aterwards, the egg laying diminishes grad- the borer during the berry perforation). Other ually. Both the founding female and the larvae parasitoids that have been reported attacking build tunnels in the bean, where they also feed. H. hampei include Aphanogmus dictyna (Water- Pupation takes place within the cofee bean ston) (Hymenoptera: Ceraphronidae) (Uganda), where the larva hatched. he duration of the bio- Sclerodermus cadavericus Benoit (Hymenoptera: logical cycle, from egg to adult, varies according Benthylidae) (Uganda, Zaire, Kenya), Cephalono- to the temperature: 21 days at 27°C, 32 days at mia hyalinipennis Ashmead (Mexico) and Cryp- 22°C and 63 days at 19.2°C. As the irst adult of- toxilos sp. (Hymenoptera: Braconidae) (Colombia). spring appear, the population inside an infested In Brazil and Colombia, there are reports of an bean typically consists of 25–30 individuals in undescribed species of Cephalonomia parasitizing all stages of development, of which there are H. hampei. approximately 10 females for each male. Mating Some of the predators that have been recorded is conducted between siblings inside the bean. include Dindymus rubiginosus (F.) (Hemiptera: he mated females leave the bean where they Pyrrhocoridae) (Indonesia), Calliodes, Scoloposcelis developed to look for another where they will (Hemiptera: Anthocoridae) (Colombia), and Lep- oviposit. Several generations occur while berries tophloeus sp. near punctatus Lekovich (Coleoptera: are available. Ater cofee harvest, the borer Laemophloeidae) (Togo, Ivory Coast). However, continues to reproduce in the non-harvested most of the predators of H. hampei reported berries located on the plant and on the ground. from around the world (most of them anecdotal In locations with low rainfall, where there is a records) have been ants (Hymenoptera: Formici- clearly deined period between harvests, the dae), including Azteca instabilis (F. Smith), 972 C Coffee Pests and their Management Crematogaster curvispinosa Mayr, C. torosa Mayr, Management Dolichoderus bituberculatus Mayr, Pheidole radoszkowskii Mayr, and Solenopsis geminata (F.). An integrated pest management strategy is used Unknown species of Azteca, Brachymyrmex, against the cofee berry borer. he principal tactics Paratrechina, Pheidole, Prenolepis and Wasmannia are cultural control, biological control, use of traps have been recorded as well. baited with attractants, and chemical control with Several entomopathogenic fungi attack the cof- synthetic insecticides. Sampling infested berries is fee berry borer, but Beauveria bassiana (Balsamo) used for pest control decision-making. Vuillemin is the most common species infecting H. hampei adults under natural conditions. Other fungi recorded infecting H. hampei are Fusarium oxyspo- Sampling Infested Berries rum Schlechtend, F. avenaceum (Fr.) Sacc., Hirsutella eleutheratorum (Nex ex Gray) Petch., Metarhizium he proportion of infested berries is calculated anisopliae (Metschnikoff) Sorokin, Nomuraea rileyi based on the following sampling protocol: in an (Farlow) Samson, Paecilomyces amoenoroseus (Hen- area of 1–5 ha, 20 uniformly distributed sites are nings) Samson, P. far inosu s (Holm. ex S.F. Gray), P. selected; at each site ive cofee plants in a row are fumosoroseus (Wize) Brown & Smith, P. javanicus selected; 20 berries of each cofee plant are exam- (Friederichs & Bally) Brown & Smith, P. lilacinus ined (without tearing them of), and the number (hom.) Samson, and Verticillium lecanii (Zimmer- of perforated berries is recorded. man). Some of these fungi, such as M. anisopliae and P. lilacinus, have been isolated from H. hampei- infested berries collected from the soil. Cultural Control Metaparasitylenchus hypothenemi Poinar (Tylenchida: Allantonematidae), an entomopatho- here are a number of cultural practices that may genic nematode attacking H. hampei adults, has be used to minimize damage by borers. he ber- been reported in Mexico and appears to have a ries let on the plant before maturity and on the wide distribution in cofee plantation in Mexico ground ater harvest are collected and boiled for and Central America 5 min to eliminate the borers in them. his practice his nematode cause sterility in female bor- is also called “manual control” or “rere.” Weeds are ers. he natural parasitism by an undescribed controlled ater the harvest in order to facilitate species of Panagrolaimus (Rhabditida: Panagro- the collection of berries from the ground and to laimidae) has been reported in H. hampei in India increase the mortality of H. hampei by dehydra- and Mexico. M. hypothenemi and Panagrolaimus tion of the berries. he cofee and shade plants are sp. were found infecting the same H. hampei adults pruned to create less favorable environmental in Mexico. Species from Heterorhabditidae and conditions for multiplication of the borer. Cofee Steinernematidae (Rhabditida) are able to infect plant density is decreased because high sowing H. hampei in the laboratory, but this has not been densities favor infestation. he cofee plants are observed in the ield. fertilized so that they have more uniform lower- In Colombia, infections in the cofee berry ings. Varieties with the same fruiting pattern are borer caused by bacteria such as Bacillus sp. and used because the early lowering varieties are an Serratia sp. were observed. Also, infections of infestation source for late lowering varieties; proteobacterium Wolbachia in H. hampei adults however, cofee varieties or species which lower have been reported from samples around the world. earlier or later than the main variety can be used he microsporidian Mattesia sp. was observed in a as “trap crops,” if managed properly. he harvest is population of laboratory-reared insects. conducted as the fruits ripen. Coffee Pests and their Management C 973 Biological Control to sunlight. Early in the morning is the most efective time to apply it in the ield, when the The natural enemies most often used against borer is starting to penetrate the cofee berry. the borer in tropical America have been the par- asitoids C. stephanoderis, P. na suta and P. coffea, and the entomopathogenic fungus B. bassiana. Insect Traps These three parasitoids were introduced to tropical America from Africa. They are estab- Traps are used for monitoring and control of the lished in most of the countries where they have cofee berry borer. hey are made using 2 L plastic been released. Nevertheless, classical biological bottles into which one or more windows have control with these African parasitoids has not been cut to allow the entry of lying females. Bor- been sufficient to reduce the borer population ers are attracted by a mixture of methanol and below the economic injury level. Yearly inocula- ethanol (1:1 or 3:1) and they are caught and tive and inundative releases of parasitoids have drowned in the water placed at the bottom of the been used with better results. However, inunda- trap. Typically, 16–25 traps are deployed per hect- tive releases are expensive because mass rearing are. Each trap is suspended from a branch of a methods and facilities have not been developed cofee plant at 1.2–1.5 m above the ground. Borers for area-wide releases. Parasitoids are produced captured are removed from the traps and counted for inoculative releases in laboratories where weekly. he best time to use the traps for H. ham- the borer is reared mostly in parchment coffee pei control is ater the harvest, during the massive (35% humidity) for use in rearing the parasi- emergence of females from old berries. Better toids. A rearing system for H. hampei in an arti- results for suppression of insect infestation in the ficial diet has been developed; however, its next harvest can be obtained by combining the application for mass production of parasitoids use of traps with strict sanitation. is not fully employed. An alternative and less intensive rearing system to produce parasitoids for inoculative releases is production of the par- Chemical Control asitoids in rural areas, also known as “parasitoid rural rearing.” In this system, the coffee growers here are several chemical insecticides used for rear the parasitoids at their farms or communi- borer control, among which endosulfan is out- ties. Such rearing is conducted using coffee standing for its ability to cause high mortality of berries infested by the borer in the field. Regard- H. hampei. However, this organochlorine insecti- less of the rearing method used, annual releases cide is being seriously questioned for negative of parasitoids are needed to manage the borer side efects (it is highly toxic to ish and bees, and population. it causes secondary pest outbreaks by eliminating he use of B. bassiana for borer control is the natural enemies); borer resistance (appar- more developed than is the use of parasitoids. Its ently this pest is not resistant to endosulfan in success has resulted from the relatively easy tropical America; nevertheless, there is concern propagation, formulation and application of this about the development of resistance, as in the fungus. Strains of B. bassiana are commonly case of New Caledonia); and sanctions in the collected for mass production from infected international market due to the possible presence H. hampei females in the ield. Rice grains are used of residues in the cofee bean. he insecticide as the propagation substrate for this entomopatho- should only be used if the borer population gen. he fungus requires high relative humidity for reaches the economic threshold. he best time germination of the spores and it is very susceptible for spraying is when the adult borer starts 974 C Coffee Pests and their Management penetrating the fruit, at the so-called semi-con- by leaf miner can cause severe defoliation. In sistency stage of development (about 20% dry Ecuador, defoliation between 70 and 90% has weight in the bean). his period varies, according been reported on C. arabica and from 30 to 40% to the temperature, from 90 to 140 days ater the on C. canephora. he lack of leaves on the plant main lowering. Formerly, treatments were reduces the photosynthetic activity, and conse- throughout the plantation, but now sprays are quently the availability of nutrients for the fruits. directed only at infested areas. In Brazil, when 94–95% of the leaves were mined, a reduction in yield between 68–80% has been observed. Coffee Leaf Miner, Leucoptera coffeella (Guérin-Méneville) (Lepidoptera: Lyonetiidae) Description

Distribution he egg is oval, translucent yellow and similar to a lattened volcano in proile. It is 0.28 mm long, 0.18 his species is found in the Neotropics: Mexico, mm wide, and 0.08 mm tall. he larva has a dors- Central America, South America and the Caribbean oventrally lattened body with a more pro- region. It is widespread wherever cofee is grown. nounced flattening of the head and the first thoracic segment. he true legs are found on the 1st, 2nd and 3rd thoracic segments but four pairs Damage and Economic Importance of prolegs occur on the 6th, 7th, 8th and 13th abdominal segments. It has four larval instars. he In some areas of tropical America, the cofee leaf larva attains a length of 4.5 mm. he pupa is white miner is considered to be the principal insect in the initial stage and ochre towards maturity, pest of cofee; certainly this is the case in some except for the dorsal portion, which remains white. cofee-growing areas in Brazil. Leaves are the he pupa is covered by a white cocoon which only plant organs damaged by this insect. he resembles an elongated “H” or “X.” he adult is a damage is caused by the larva. Four larvae per small moth between 2.0 and 3.0 mm long with its leaf may cause leaf drop. he afected leaves show body covered by silvery scales. he antennae are irregular light-brown spots. If the damaged sur- long and thin. he front wings possess a gray oval face of the leaf is rubbed, the leaf separates into point distally, surrounded by a black line and two layers and between them is found a small edged by a yellow stripe that extends along the white worm, from 2 to 5 mm in size. he cofee margin. Males tend to be slightly smaller than leaf miner lesions may be confused with the females. symptoms of Anthracnose (Colletotrichum sp.), but in the latter case the leaf layers do not sepa- rate when rubbed. Four months ater lowering, a Biology and Ecology reduction in the rate of growth of the cofee ber- ries and an increase in leaf production take place; he female usually lays its eggs irregularly on the this allows the plant to compensate for the dam- upper surface of the darkest, most mature leaves, age caused by the miner. But when the fruit particularly on the middle and lower parts of the growth starts again, if there is more than one leaf cofee plant. Eggs are laid individually or in small miner lesion per leaf it will result in economic clusters of up to seven eggs, with a total fecundity damage. he damage increases if simultaneously that varies between 30 and 80 eggs. Upon hatch- the plant is under drought stress. Attack of cofee ing, the larva makes a semi-circular cut at its base Coffee Pests and their Management C 975 and penetrates rapidly into the leaf, where it Management moves about, mining the palisade parenchyma tissue. When ready to pupate, the fully developed here are several useful approaches to manage- larva leaves the gallery very early in the morning, ment of cofee leaf miner. population. Sampling is making a semi-circular cut on the face of the leaf, recommended prior to initiating chemical through which it slips down by a silk thread control. which it secretes from the mouth. Cocoon for- mation and pupation take place on the lower face of the coffee leaf, oten on a curvature of the leaf Sampling Damaged Leaves or close to a protruding vein. he duration of the life cycle, from egg to adult, lasts between 25 and he recommended sampling protocol follows: 75 days, depending on the temperature. Several sampling is initiated when the cofee lowers, and generations occur annually, particularly in cofee is conducted monthly until the berries stop grow- plantations with full sunlight or only lightly ing. he cofee plantation to be sampled is divided shaded. he abundance of L. coffeella is signii- into areas not larger than one hectare. he sam- cantly afected by the onset of rainfall, and by pling is conducted by selecting a zigzag path natural enemies, which are very numerous ater across the cofee plantation and by selecting 12 the end of the dry season. cofee plants at random. From each cofee plant, 25 leaves are selected at random, and the number of leaves with mines is recorded. he irst two Natural Enemies pairs of leaves at the tip of the branches are not sampled. The coffee leaf miner is attacked by a large number of parasitoids; predators and some insect pathogens have also been recorded. More Cultural Control than 20 morphospecies of parasitoids wasps (Hymenoptera) have been reported in tropical he shade canopy of cofee plantation should not America. Eulophidae are the most common par- be trimmed immediately ater harvest; it should asitoids of L. coffeella; this group is largely be thinned only when the onset of the rainy sea- unknown because keys for neotropical species do son is imminent. Adequate soil fertilization is not exist. In Mexico, Neochrysocharis was the important. hick mulch coverage of the soil genus with the greater number of morphospe- should be maintained. High cofee plant densities cies, and also the one that was collected most fre- should be avoided. he cofee plant should be quently. It was followed, in order of abundance, pruned to stimulate vigorous growth. Damaged by Pnigalio, Closterocerus, and Zagrammosoma. leaves should be collected and placed in contain- Of two braconids collected in Mexico, Stiropius ers that allow the escape of parasitoids but not of letifer (Mann) was the most abundant and most the cofee leaf miner. widely distributed. Wasps (Vespidae) are the most important predators of cofee leaf miner in Bra- zil, but in Mexico, the most important predators Biological Control are ants (Formicidae). he bacteria Pseudomonas aeruginosa (Schroeter) Migula and Erwinia her- he introduction of natural enemies into new bicola (Löhnis) Dye, and the fungus Cladospo- areas has not been widely explored. he most rium sp., have been reported infecting L. important action conducted so far has been to coffeella. protect the already existing natural enemies by 976C Coffee Pests and their Management avoiding the use of broad-spectrum, residual Root Mealybugs (Hemiptera: contact insecticides. he natural control exerted Pseodococcidae) by the cofee leaf miner’s natural enemies varies from 2 to 70%; however, in most cases it is unnec- Distribution essary to resort to the use of chemical control. Regrettably, the use of chemical insecticides may Root mealybugs are found in Neotropical coun- eliminate a large portion of the beneicial organ- tries where cofee is grown. he principal root isms, causing pest resurgence and making it dif- mealybugs afecting cofee plants in tropical icult to implement control. In certain countries America are shown in Table 17. like Honduras, high and recurring L. coffeella infestations have diminished signiicantly when the use of chemical control is not applied for sev- Damage and Economic Importance eral years and the beneicial fauna is restored. his supports the idea that cofee leaf miner con- hese insects attack the cofee plant roots and some trol should not be based on use of insecticides in species also afect the foliage. he foliage of attacked order to avoid disrupting the actions of parasi- cofee plants appears withered, the color of the toids and predators. leaves fade, and they have copper, brown or necrotic edges. Additionally, total or partial leaf drop may occur. hese symptoms are more evident during Chemical Control the dry season. In case of serious attacks by Dysmi- coccus bispinosus (Beardly), a thick, cork-like, dark Numerous chemical insecticides can be used for crust covers the main and secondary roots; the suppression of L. coffeella and protection of foli- attacked roots lose their absorbent root hairs. Heav- age. hese products include both organophos- ily attacked plants perish. Infestation appears to be phate and pyrethroid insecticides. hey are associated with ants (Formicidae). he symptoms inexpensive and can be applied at the same time may be confused with the symptoms of fungal with other agrochemicals, but they are highly diseases and with physiological plant problems. toxic and they are more likely to cause ecological In the case of Neorhizoeccus coffeae (Laing) and disturbances. Organophosphorates are oten D. brevipes (Ckll.) infestations, the branches turn applied twice at an interval of 30–45 days, with whitish and the afected root seems to be covered an additional application in cases of severe with lour, the crust separates easily, and consider- attack. In the case of pyrethroids, one or two able deteriorated tissue appears. he attacked plants applications at an interval of 45–60 days are rec- have little anchorage and are easily dislodged. ommended. he application of granular insecti- Root mealybugs have become important cofee cides with systemic action to the soil is also pests in some areas of tropical America during the recommended in cases where it is diicult to last 20 years. In Guatemala, the most harmful species apply foliar sprays. Soil applications interfere is D. bispinosus; in Costa Rica, N. coffeae and D. brevi- much less with the natural enemies of the cofee pes; in El Salvador, D. brevipes, Rhizoeccus nemoralis leaf miner, and this approach can be used to con- Ham. and Geococcus coffeae Green; and in Colombia, trol pests and soil diseases simultaneously. Gran- Chavesia caldasiae (Balachowsky). At some cofee ular insecticides should be shallowly buried at plantations in Colombia, Planococcus citri (Risso) has the drip line of the plant once a year during the also appeared as a pest, causing up to 30% yield loss rainy season. Where this type of product is used, in the attacked trees. Other forms of damage caused it is recommended that harvest occur 90 days by root mealybugs include excessive extraction of ater application. potassium, destruction of the absorbent root hairs, Coffee Pests and their Management C 977 development of small rotting areas which tend to Other species, such as Pseudococcus adonidum (L.) atrophy, and enhanced entry of plant pathogens. his are oviparous. Females die shortly ater the eggs damage creates a general condition of weakness, slow hatch. Upon eclosion, the small nymphs start look- growth and plant death in many cases. ing for an appropriate place to settle on the plant Dysmicoccus brevipes weakens the cofee root; at the selected site, they insert their mouthparts, plants but it rarely kills them. In Costa Rica, and feed by suctioning the sap from the root. Some plants with more than 20 mealybugs per liter of of them settle down permanently on a site until they soil are more susceptible to infection by the fungus reach maturity, and others may change their feeding Cercospora coffeicola Berk & Cooke. Damage is site by moving short distances. Depending on the more apparent on nutrient-deicient soils, and type of soil, the humidity, aeration and age of the where weeds are abundant. Plants in seed beds cofee plant, they usually place themselves between and tree nurseries are also attacked. he varieties 10 and 60 cm under the soil surface, their popula- of C. arabica grown in Central America (e.g., tion diminishing as the soil depth increases. Difer- Caturra, Catuaí, Bourbon) are susceptible to the ent species prefer diferent parts of the root. For mealybug attack, while tolerance has been example, D. brevipes and R. nemoralis prefer the observed on C. canephora, C. dewevrei De Wild. main and the lateral roots, while G. coffeae attacks & Durand, and C. excelsa Chev. the absorbent roots; the smaller species attack the whole root system near the soil surface. As they feed and develop, the nymphs and Description adults excrete their characteristic waxy cover and form compact colonies. Mealybugs excrete sugary Mealybug eggs are small (0.5 mm). he nymphs are substances (honeydew), which supports the growth oval, slightly swollen, usually white, yellow or pink- of fungi (i.e., Bornetina), which contribute to for- colored, and covered by a white waxy-mealy dust mation of the thick, cork-like, dark crust covering with waxy ilaments projecting laterally. he female and sheltering the mealybug colony; a succession of nymphs molt three times, and the males, contrary crusts give a knotty appearance to the root. he sug- to the females, form a waxy cocoon in the third ary substances also attract certain ant species, which instar, where they pupate. he adult females have no live in a symbiotic association (trophobiosis) with wings and they are similar to the nymphs but larger. the mealybugs. In exchange for the sugary foodstuf, Smaller species, such as Geococcus and Rhizoecus, the ants give them protection and transportation are from 1.5 to 2.0 mm long and the larger ones, from one root to another and from one plant to such as Dysmicoccus and Pseudococcus, are from 2.5 another. he ants that associate with mealybugs in to 5.0 mm long. Males are white, fragile-looking, South America and in some of the Caribbean smaller than the females, and they possess a pair of Islands are in the genus Acropyga. In Colombia, the wings and a pair of terminal ilaments. Hope ant (A. robae Donisthorpe) and the Amagá ant (A. fuhrmanni Forel) are associated with N. cof- feae and C. caldasiae, respectively. In Guatemala, D. Biology and Ecology bispinosus seems to be associated with the presence of the ant Solenopsis geminata (F.). P. c itr i does not Mealybugs generally live attached to the cofee root, produce large quantities of sugary excretions when forming numerous colonies. heir reproduction it lives on the plant roots, and is not attractive to may be sexual or parthenogenetic (partial or total). ants. In certain cases, the mealybugs have lived for Eggs are laid in groups and covered by a layer of more than a year in the absence of ants. cotton-like wax or by an egg sac of crystalline wax he life cycle, from egg to adult, requires from ilaments. A single female may deposit 300–600 eggs. 30 to 120 days, according to the species and the 978 C Coffee Pests and their Management temperature. Five generations develop per year in ant nests should be examined critically; from 15 to the case of D. bispinosus. Root mealybugs develop 20 plants/ha should be checked, paying more atten- better during the rainy season, particularly in low tion to those that are close to the ant nests and/or or medium altitude plantations in Central Amer- possess yellow leaves. he surrounding shade trees ica. Other conditions that favor their development and bushes should also be checked. he plants are are sandy, acid pH, and medium moisture soils. In checked by moving the stems in all directions in Colombia, the damage caused by Rhizoecus sp. order to gain visibility of the base of the roots. seems to increase in old, poorly fertilized planta- tions, and in Guatemala D. bispinosus is found most frequently in 1–5 year-old plantations. Mealybugs Cultural Control are polyphagous, also attacking other plants such as shade trees (Inga spp.), cassava (Manihot escu- Mealybugs should not be present in the seed bed lenta Crantz), sugarcane (Saccharum), banana trees and tree nursery. he limits of any infestation (Musa), lemon trees (Citrus) and some herbs that sites should be determined and marked. Adequate grow on the coffee plantation. In Costa Rica, Anred- fertilization should be provided, including era ramosa (Moq.) Eliasson is an alternate host of addition of organic matter to the soil. Physical D. brevipes; in El Salvador, D. bispinosus has been conditions of the soil should be improved in found associated with Lantana camara L. order to avoid loods. Planting cofee trees on land previously supporting plants that are highly susceptible to mealybugs (e.g., cassava, sugarcane) Natural Enemies should be avoided. Alternate host plants should be eliminated from the plantation. Severely dam- In general, the literature on coffee mealybugs in aged plants should be removed and burned. tropical America does not make reference to their natural enemies. In Cuba, Coccidoxenoides peregrinus (Timberlake) (Hymenoptera: Encyrtidae) is cited Biological Control as a solitary, primary endoparasite of the pseudo- coccid complex in coffee. Other natural enemies his is practically unexplored in the cofee grow- of mealybugs reported in Cuba are Diadiplosis ing countries of tropical America. cocci Felton (Diptera: Cecidomyiidae), Leptomastix dactylopii Howard (Hymenoptera: Encyrtidae) and Signiphora sp. (Hymenoptera: Signiphoridae). Plant Resistance to Insects

In Guatemala, some research has been conducted Management which supports the use of plants grated on resis- tant rootstocks of C. canephora (genotypes 3757, here are management options for mealybugs, but 3754, 3751, 3581, 3752 and 3756) and C. dewevrei. insecticides are normally used once pest popula- tions develop. Chemical Control

Sampling Systemic organophosphorate and carbamate insec- ticides produce good results, although they are Sampling should preferably be conducted on young expensive. he presence of mealybugs in seed beds cofee plantations (up to 6 years old). Plants near or on plants younger than 1 year old is suicient Coffee Pests and their Management C 979 justiication for insecticide application. On planta- When Capnodium (sooty mold) and Meliola (black tions older than 3 years, insecticide application is mildew) fungi grow on the honeydew excreted by made if more than 1.6 colonies per plant, on average, the scales, they interfere with photosynthesis. Ants are found. In no case should the damage be allowed are present where scale insects are feeding. to exceed 25% of the absorbent roots. Insecticides In cases of severe attack, a dirty appearance are applied on the drip line of the plant if the dam- on the plant, general weakening, growth delay, yel- age is on the small roots. If the damage is on the lowing and drop of foliage and fruit are observed. main root a funnel-shaped hole should be made With the articulated scale, Selenaspidus articulatus around the tree trunk, the insecticide should be (Morgan), old attacks may be recognized because poured in and the hole should be covered again with the site where the scales were located turns yellow soil, adding also a layer of dead leaves. Application of or discolored, resembling infection by the cofee granular insecticides is made at the beginning of the rust fungus (Hemileia vastatrix Berk. and Br.). rainy season or 3 months before starting the harvest. Some species, such as the green scale, Coccus viridis (Green), are considered to be quite impor- tant to cofee production, though some attack a Scale Insects, Mealybugs and number of diferent cultivated plants. Severe infes- Related Foliage Pests (Hemiptera) tations of C. viridis may kill young tree nursery plants. he incidence of these pests is highest on Distribution cofee plantations lacking adequate shading.

Diferent scale insects, mealybugs and related foliage pests live on the cofee plant. he geo- Pest Description graphic distribution of some is restricted to a few countries of tropical America, whereas others he following cases are presented as examples: are distributed more widely. Some are reported C. viridis – adult females are motionless, oval, some- attacking the cofee plant only in South America, times asymmetric, very lat and pale yellow. hey others only in Central America or the Caribbean have some black spots centrally, and they tend to (Table 17). be sot and elastic. hey are about 2.2 mm wide and 4.0 mm long. he presence of males is very rare. Saisettia coffeae (Walker) – adult females are Damage and Economic Importance motionless, almost spherically shaped and dark brown. hey are 2.0–3.5 mm in diameter. he males Scale insects, mealybugs and related species attack are winged. P. c itr i – adult females are mobile, oval, the aerial part of the cofee plant and, in some spe- pale yellow or dark orange, with very clear segments cies, also the root (e.g., Planococcus citri [Risso]). on the body, and 4.0 mm in size. hey are covered he leaves, fruit, branches and young tissues of with a dusty white glandular secretion except for a the aerial part of the attacked cofee plant oten longitudinal stripe dorsally. hey have ilaments lat- support colonies or groups of circular, oval or elon- erally. Males are smaller (1.0 mm), violet to yellow gated scales, which may be lattened or swollen, in color, and they have well-developed wings. with a sot or hard consistency. In other cases, colo- nies of insects have a sot body covered with white, cotton-like ilaments. hese insects cause damage Biology and Ecology by removing large quantities of sap, which causes plant malnutrition. Also, sticky honeydew and he biology of these insects varies among spe- blackish molds can be found covering the foliage. cies and can be quite complex. he irst instar 980 C Coffee Pests and their Management has legs and antennae and is very active. To feed, Natural Enemies the insects attach and insert their mouthparts. Ater the irst molt, they generally lose their legs hese insects are susceptible to a large number of and antennae and the insect becomes sessile. By parasites, predators and pathogens as natural then, it begins to secrete a waxy, scale-shaped enemies. layer that covers the body. In the case of scales of the family Diaspididae, this layer of scale is almost always separated from the insect’s body. Management Adult females remain under this cover and they produce their eggs or directly give birth to the Sampling nymphs therein. he location on the plant, and the age of the plant they prefer to attack, depends During the dry season, inspections should be con- on the species of scale: C. viridis is commonly ducted to check for the presence of scales and located along the leaf veins, on the back of the related species in the cofee plantation, as well as leaves, on young buds and on seed bed cofee on other plants cultivated nearby or at the same fruits of nursery plants; S. articulatus is found time. mainly on the leaves and fruits of production plants; the round scale, Parasaissetia sp., mostly Cultural Control attacks the stems and branches of cofee plants younger than 1 year; the black scale, Ischnaspis he nursery shading should be reinforced during longirostris (Signoret), infests the leaves, branches the dry season. Afected plants should not be and fruits of old, poorly attended cofee planta- transplanted. Weeds should be suppressed. he tions; Cerococcus catenarius Fonseca gathers in pests should be kept under control on host plants the form of a line or chain along the trunks and existing in or near the cofee plantations. Sanitary branches; P. c itr i attacks new branches, leaves, pruning should be performed to eliminate (by lower buds, fruit peduncles and fruits; Orthezia burning) old and unproductive branches infested spp. attack branches, leaves and fruits, mostly of by the pests. robusta cofee in Brazil. he males develop very much like the females except that in the last stage, before transforming into adults, they go Biological Control through a pupal stage; the wings develop exter- nally over the pupa. Most of the scales reproduce Natural enemies should be protected and pre- parthenogenetically. Some species are oviparous served, using insecticide only if necessary. (S. coffeae, S. olae [Oliver]) and others are vivipa- rous (Coccus hesperidum L.). he total number of eggs produced per female varies among the Chemical Control species; for example: C. viridis, between 50 and 600 eggs; Orthezia praelonga Douglas, more than Chemical control is directed only at infested plants, 200 eggs; C. catenarius, about 900; S. coffeae can ater checking to determine that the scale colonies lay up to 1,600 eggs. he complete life cycle, from are alive. For better control, mineral oil is added to egg to adult, lasts between 40 and 60 days. he the insecticide solution, with applications made scale insects are more abundant during the dry every 15 days until the problem is corrected. he season and at the onset of the rainy period. Hard oil should not be used during lowering or during rains and natural enemies are important factors sunny periods of the day. During the rainy season, in the mortality of these pests. granulated insecticides may be used. Coffee Pests and their Management C 981 Cutworms and Armyworms Biology and Ecology (Lepidoptera: Noctuidae) Adults are moths that are active at night, laying Distribution their eggs individually (A. ipsilon) or in groups (S. frugiperda). During the irst two larval stages, hese insects are widely distributed in the cofee they feed on leaves that are at soil level, and in plantations of tropical America (Table 17). the last three they act as cutting worms. During the day, they remain hidden in the soil. In some species, such as A. ipsilon, the larvae coil up when Damage and Economic Importance disturbed. Larvae pupate in the soil.

Cutworms and armyworms constitute an economi- cally important pest for many crops. Damage is Natural Enemies caused during the night by the larval stage. he larvae attack the stems of small cofee plants in ger- here are many natural enemies (parasitoids, minating beds, seedbeds or plant nurseries, and predators and pathogens) of these pests. In Ecua- recently transplanted plants. On seed beds and plant dor, the larval parasitoids Bonetia sp. (Diptera: nurseries, plant damage typically takes the form of Tachinidae) and Chelonus sp. (Hymenoptera: plants cut at the soil level or slightly above, or Braconidae) and the predatory ground beetle withered plants. In the case of recently transplanted Calosoma sp. (Coleoptera: Carabidae), are cited. cofee plants, defoliated and sometimes dead cofee plants can be observed. Spodoptera frugiperda (Smith) larvae feed on the stem, causing withering Management and inally death of the small plants during the irst year of their lives. In other cases, the stem breaks at Sampling the site of the ring formed by the larval feeding. When the infestation is severe, many plants are killed Night-time inspection of seedbeds and the young and re-sowings are needed, which increases the cof- plantations should be made to detect initial fee plantation set-up costs. Damage is more frequent infestations. in plantations that are close to ields where corn, beans, vegetables, cassava or pasture are grown. Cultural Control Pest Description Seedbeds or plant nurseries should be kept clean he following species are presented as examples: of weeds and dead leaves, since the larvae seek S. frugiperda – the larvae have a well-contrasted, shelter there. inverted “Y” on the head; neonate larvae are white with a black head, but as they grow they turn dark. Large worms are light brown to dark green in color Mechanical Control and they are about 4.0 cm long. Agrotis ipsilon (Huf- nagel) – small larvae are brown with paler back Larvae should be eliminated by hand during the marks, and large ones, which may be as be as large as night-time inspections. Heavy watering should be 4.0–5.0 cm, are shiny black-gray in color, with a pale applied to get the larvae out of their hiding places, gray line on the back and black tubercles on each of followed by manual elimination. Light traps can the segments. be used to capture the adults. 982 C Coffee Pests and their Management Biological Control mortality of the branches and stems is caused by microorganisms that invade the plant tissues Biological insecticides such as Bacillus thuringiensis through the feeding sites formed by the brown Berliner should be used, particularly at the begin- cofee borer. Ants, termites or mites can be found ning of infestations, when the larvae are small. in the abandoned galleries. Oten the ants cause the death of branches or stems when enlarging the Chemical Control abandoned galleries of X. morigerus to construct their own nests. If death of branches or stems does Insecticides can be incorporated into the soil, before not occur, the yield is reduced as a result of dam- or ater sowing, for cutworm control. Granular age to the lowering and development of the fruit. products are used in a preventive manner. he use he symptoms can be observed more frequently of poisoned baits during the night and dry weather on weak cofee plants, but attacks may also be seen is also recommended. on the young stems of pruned plantations. his pest is particularly important in Ecuador. In Mex- ico, X. morigerus is also an important pest in the Soconusco region in Chiapas. Brown Coffee Borer, Xylosandrus morigerus (Blandford) (Coleoptera: Curculionidae: Scolytinae) Description

Distribution he egg is oval, white and very small. he larva is milky white, with a yellowish head, and lacks legs. his pest comes from the Oriental region, having he pupa is white initially, turning cream to brown its distribution center in the Indomalayan area. It toward maturity. he adult is cylindrical and from was detected in the western hemisphere in 1958– 1.40 to 1.90 mm long. It is diferentiated from 1959, and it is now found from Veracruz, Mexico other species of the same genus by the bright to Brazil. brown-reddish color, by the stouter body and because the declivity commencing only on-third of the elytral length from the base, and by the near Damage and Economic Importance absence of punctures on the sides of the elytra (variable). Females have well-developed wings and Various tree species can be attacked by X. morigerus ly, but males are incapable of light. Females are (e.g., avocado, cacao, cedar, cofee). his insect dis- larger than males. plays a strong preference for attacking robusta cof- fee, C. canephora. Some reports indicate that it may also infest C. arabica; however, this has not Biology and Ecology been conirmed in Mexico. he attacked cofee plant branches and young stems typically display a Mated females take light during the day, leaving few or many holes of about 1.0 mm diameter. the gallery where they developed in search of Blackening of the tissues may be seen around the branches or stems, which they penetrate to perforations. A longitudinal cut of an afected construct the new galleries. The female lays branch reveals a gallery in which the whitish lar- from 20 to 60 eggs in 8–10 days. X. morigerus is an vae can be observed, along with reddish brown ambrosia beetle. he adults and larvae get more adults. he attacked young branches and stems nutrition by feeding on fungi (e.g., Ambrosiaemy- dry up distally and then die. Apparently, the ces zeylanicus Trotter is reported from Ecuador; Coffee Pests and their Management C 983 Raffaelea tritirachium Batra from Mexico) than ethanol have been used for monitoring lying from the coffee plant tissues. hese fungi grow females in robusta cofee plantations. inside the gallery, which is inoculated by the found- ing female. he larvae have three instars. he life cycle, from egg to adult, is 20–40 days. A gallery Cultural Control may contain more than 80 individuals in all stages of development. he sex ratio in galleries is female- Infested vegetative material, particularly in dominated; various studies have found only one young or pruned plantations, should be cut and male for each 7, 11 or 20 females. Mating occurs burned periodically. Adequate fertilization should within, or very close to the gallery. Infestation is be applied. Shade of cofee should be regulated by apparently less evident under drought conditions, pruning. Weeds should be suppressed by shading, because the ambrosia fungi require moisture. Nev- mulching, use of ground cover, and by selective ertheless, reports from Ecuador indicate that the weeding by hand. populations are larger during the dry season of the year. X. morigerus is a pest which frequently attacks healthy plants; however, very strong attacks may be Biological Control observed when the cofee plants have been weak- ened by droughts, malnutrition, nematode attacks his approach has not yet been attempted for this and competition with weeds. he attacks may be species. accompanied by attacks from other Scolytinae.

Chemical Control Natural Enemies his is recommended when the beetle population No native parasitoids of this pest have been has undergone a marked increase and natural and reported in cofee growing countries in tropical cultural control cannot restrain it. Insecticides are America. However, it should be mentioned that in useful only when adults are out of the galleries or Indonesia, a Tetrastichus sp. (Eulophidae) has been are boring on the branches; once they have taken reported, and also probably a bethylid parasitoid. refuge within the galleries, insecticides have little In Ecuador, ants (Formicidae) have been recorded or no efect on X. morigerus. as predators of the brown cofee borer, including species of Crematogaster, Leptothorax, Pheidole, Pseudomyrmex and Solenopsis. he entomopatho- Stem Borers Plagiohammus spp. genic fungus B. bassiana has been reported infecting (Coleoptera: Cerambycidae) this insect pest. Distribution

Management hree species of Plagiohammus have been reported attacking the stem of cofee plants in Mexico and Sampling Central America. P. maculosus (Bates) has the wider geographic distribution (Costa Rica, El Salvador, No sampling methods have been developed in Guatemala, Honduras, Mexico), while P. me x i canu s cofee plantations; however, some studies indicate Breuning and P. spinipennis (homson) have that penetrated branches and stems have an aggre- been recorded attacking cofee only in Mexican gated distribution in the ield. Traps baited with plantations. 984 C Coffee Pests and their Management Damage and Economic Importance ready to pupate, the larva moves close to the excre- tory opening, which has been made close to the A pile of white-yellowish sawdust or powder present ground, and it isolates itself within the stem in a at the base of coffee plants, at the soil level, in a good chamber surrounded by sawdust. he larval period indication of infestation by Plagiohammus. Infested lasts from 2 to 3 years. Adults are more visible at the plants may have a withered, yellow-like and decay- beginning of the rainy season (April through June), ing appearance. Careful observation at the stem base the period when egg laying occurs. he abundance may help identify the hole or holes (ca. 5.0 mm in of these cerambycids is higher in high-altitude cof- diameter), where the sawdust originates. A longitu- fee plantations (>1,000 m) and in places with long dinal cut of the stem and root may uncover a large, summers or with lack of rain. Abandoned cofee white or creamy-colored larva with long gallery plantations are more severely attacked. containing powder; the gallery begins at the stem and may go as low as the tip of the central tap root. hese borers are one of the most destructive cofee Natural Enemies plant pests in certain areas of tropical America. he damage is caused by the larva when it bores into the here is no information on the natural enemies of stem and the root. he borer attack delays the plant the Plagiohammus spp. growth and it may cause death directly by damaging its root, or indirectly, by facilitating stem breakage following wind action or other factors. Management

Sampling Description Cofee plants having sawdust as the base of the he egg is unknown. he larva is creamy-white, with trunk should be searched for. If damage is recent, the thorax wider than the abdomen, and legless. Its the sawdust is white or pale yellow. head is light brown with strong and visible mandibles extended forward. A well-developed larva is about 4.5 cm long. he pupa is brown and similar in size to Cultural Control the adult. he adult has an elongated body, cylindri- cal, from 2.0 to 3.5 cm long by 0.8 cm wide. he body Infested stems should be removed. Adequate is light brown with two white lines on the prothorax fertilization should be applied. Weeds should be and with irregular white spots on the elytra. he managed by shading, mulching, ground cover, antennae are longer than the body (4.0 cm). and mechanical removal.

Biology and Ecology Biological Control

Not much is known about the bionomics of Plagio- his has not been attempted yet. hammus spp. Adult females lay eggs on the bark of cofee plant stems, at a height below 30.0 cm. Upon eclosion, the larva penetrates the stem and bores Chemical Control longitudinally all the way to the root, while it feeds, grows and develops. he larvae may be found in the In places where the pest appears yearly, a preven- stem, from the base to a height of one meter. When tive insecticide application with a brush or a Coffee Pests and their Management C 985 manual pump is recommended, treating from important, because they may cause death of the plants. the stem base up to 60.0 cm high. Application may he attacks are more severe in plant nurseries, on be repeated once or twice every 20 days. In order recently transplanted cofee bushes and on 1 year-old to kill the larva within the stem, a cotton ball plants, although mature plantations may also sufer soaked in an insecticide can be inserted through the attacks of these pests. In some plantations it is esti- the respiration and excretion opening made by the mated that 2 or 3% of the transplanted cofee plants larva, or insecticide solution can be injected into may be lost to white grub attack. Cofee plantations the opening with a syringe. When treating in this located in the vicinity of pastures are most afected. manner, the oriice is enlarged, the product is applied and the oriice is sealed with mud, clay or any other material that solidiies. his step, Pest Description though efective, can be expensive due to the labor it may require. he egg is white; when recently laid, they are elon- gated, and later on they adopt a round shape. he larva has a milky-white colored body with a “C” White Grubs (Coleoptera: shape, with long thoracic legs covered with hair. he Scarabaeidae) head is dark or light, with strong mandibles. here are three larval stages; the last stage grows up to Distribution 3.5–4.0 cm long. he pupa is brown-golden in color, with a size that varies between 1.8 and 2.0 cm. he Phyllophaga is a well-represented genus of white adult is a strong, heavy bodied scarab. Depending grubs in cofee plantations in tropical America on the species, they may be light or dark brown (Table 17). In El Salvador, P. latipes (Bates), P. men- or reddish-brown in color, measuring from 0.5 to etriesi (Blanch) and P. obs ol eta (Blanch) are found, 2.5 cm in length; the antennae are enlarged distally, whereas in Costa Rica P. sanjosecola Saylor and with the apical expansion consisting of several P. v icina (Moser) are reported. Other white grubs laminated segments. hey are able to ly. recorded in cofee are Anomala sp. (El Salvador) and Dyscinetus picipes Burmeister (Cuba). Biology and Ecology

Adult females, which have twilight habits, come out at the beginning of the rainy season and they Damage and Economic Importance lay their eggs within the irst 10.0 cm of depth in the soil, close to pastures or fodders. he eggs are White grubs attack the cofee plant root. he damage laid one by one or forming small groups. A female is caused by larvae that live in the soil and feed on the may lay up to 200 eggs. Small larvae feed them- root system of the plant. In the seedbed and plant selves with organic matter and small roots, and nursery, the plants wither and die rapidly; in the when they reach the last development stage, they cofee plantation, irregular areas on one or several are voracious root eaters. hey are found at dif- cofee plants, usually young, may be observed, which ferent depths, according to the soil temperature show symptoms of yellowishness, limited growth, and humidity. hey are common in areas that scarce fruits and mummiied fruits. When the afected have been gramineous pastures. he larval stage plants are taken out of the soil, lesions, very few small lasts about 6 months. Pupation takes place in a roots and partial or total bark peeling on the main chamber or cell located in the soil at a depth and secondary roots are observed. In some cofee between 10.0 and 20.0 cm. he duration of the growing areas, these pests can be economically life cycle, from egg to adult, varies from 9 to 10 986 C Coffee Pests and their Management months. Adults are strongly attracted by artiicial manually. Light traps, preferably 40 watt black- light and they feed from the leaves of some plants, light traps, should be used to capture and eliminate such as cassava, African oil palm and Erythrina adults. he use of a trap for every 10–15 ha is rec- trees (Fabaceae). ommended, which should be turned on from 18:00 to 21:00 o’clock. his procedure has the disadvan- tage of attracting a number of other night habit Natural Enemies insect species, which should not be eliminated.

he larval parasitoids Campsomeris, Elis and Tiphia (Hymenoptera: Scoliidae) have been Biological Control reported in cofee plantations. Parasitism of bac- teria Micrococus sp. on larvae and parasitism of Biological control has not been attempted in cof- fungi Spicaria sp. and Metarhizium sp. on pupae fee plantations. has been observed. Nematodes as parasites of lar- vae have been observed as well. A robber ly Diog- mites species (Diptera: Asilidae) has been recorded Chemical Control predating larvae in the soil. Several mammal, reptile and bird species predate on the adults. In the case of plant nurseries and recently trans- planted cofee plants (<1 year old), one larva per plant justiies the use of granular insecticides. Management he application can also be made at sowing time. With three large larvae or seven small larvae per Sampling square meter, insecticides are recommended for young plants. Three year old plants withstand Root and soil samples at a depth of up to 20.0 cm up to eight larvae; for 4 year-old plants, 12–15 should be taken, in order to determine the infes- larvae; well-attended mature plantations with- tation sources. The samples are taken from three stand up to 20 larvae per cofee plant. cofee plants, at 30.0 m intervals. On areas <7 ha, sampling should be taken diagonally and for larger areas sampling should be taken in parallel. Black Citrus Aphid, Toxoptera aurantii (Boyer De Fonscolombe) (Hemiptera: Aphididae) Cultural Control Distribution Weeds should be suppressed principally by shading, by mulching, by ground cover vegeta- his aphid comes from the tropical and sub-tropi- tion, by slashing back and by selective weeding cal areas of the Old World. It is widely distributed by hand. Shade trees should be pruned. in cofee plantations in tropical America.

Physical-Mechanical Control Damage and Economic Importance

During preparation of the seedbed or plant T. aurantii attacks leaves, buds and other tender nursery, the soil to be used for bag illing should parts of the cofee plant. Coiled, deformed and curled be sited, and the larvae found therein killed leaves and tender buds are signs of infestation; Coffee Pests and their Management C 987 also, reduced growth, and leaf and flower drop Natural Enemies occur. Damage may occur in seedbeds, plant nurs- eries, and on adult coffee plants. Yellow, green or More than 70 species of natural enemies have been black insect colonies, more or less round shaped, reported on T. aurantii around the world. In cofee can be found on the lower surface of foliage. hey plantations in tropical America, the following are easily excited, producing a characteristic noise have been reported: the braconid parasitoids which may be audible if the colonies are very large. Diaretus sp. and Lysiphlebus testaceipes (Cresson); he infestation may be accompanied by a fungus, the entomopathogenic fungus Acrostalagmus called sooty mold, on the foliage, and also by the albus Preuss; the coccinellid predators Hippodamia presence of ants. In general, this aphid is not very sp. and Cycloneda sp.; the syrphid predators important as a pest; however, a considerable yield Allograpta sp., Paragus borbonicus Macquart and reduction may appear when severe and prolonged Baccha clavata Fabricius; and the green lacewing attacks occur, particularly if the infestation appears predator Chrysopa sp. (Chrysopidae). during the lowering and fruiting season. he damage is oten more severe in the plant nursery, on growing plants. T. aurantii is reported to be Management responsible for the transmission of pathogens to cofee plants in Costa Rica and Guadeloupe. Sampling

Description Growers are advised to monitor young leaves throughout the dry season for aphids or damage. he nymphs are similar to adults, but smaller and dark-brown in color. he adults have a globoid, dark green or black body, and they may or may not Cultural Control have wings; apterous females are larger (2.0–2.1 mm) than winged ones (1.7–1.8 mm). hey bear a Reinforce shade of cofee plantation during the dry pair of cornicles on the back of the body. season. Afected plants should not be transplanted.

Biology and Ecology Biological Control

Adult females generally reproduce by parthenogen- It is generally acknowledged that natural enemies esis and are viviparous. Males are winged and rarely contribute importantly to prevent T. aurantii from seen. An apterous female may produce 50 female having greater economic impact. Natural enemies nymphs in 7 days. he life cycle, from nymph to should be conserved. adult, is 6 days at 25°C. hese aphids excrete honey- dew on which the sooty mold fungus grows. he fungus gives a blackish appearance to the plant. Chemical Control he honeydew is highly appreciated by ants; hence the association of ants with aphids, providing them with If chemical control becomes necessary, either protection and transport to other plants. T. aurantii insecticidal oil or an insecticide may be used. inds conditions more favorable during the dry sea- Chemical control should only be applied at the son. When conditions are adverse, winged females irst signs of damage during periods of young are produced in order to disperse and colonize new leaves’ growth. Young leaves should be completely plants. Infestations appear in a cyclic manner. moistened ater application of chemicals. 988C Coffee Pests and their Management Leaf-Cutting Ants, Atta and Pest Description Acromyrmex (Hymenoptera: Formicidae) Atta cephalotes (L.) is hereinater described. heir colonies contain three castes: queens, males and Distribution workers. Queens are big (16 mm), with a strong brown-reddish color, and they have wings (although Leaf-cutting ants of the Atta and Acromyrmex they lose them ater the nuptial light); also, they genera are found in Neotropical countries where have a pair of horns on the occipital lobules and coffee is grown. he principal leaf-cutting ant another pair on the lower part of the head, close species afecting cofee plants in tropical America to the mandibles. Males are winged but smaller are shown in Table 17. (13 mm) than the queens, and they do not have the aforementioned horns. Workers are wingless. Sol- dier workers present abundant yellowish hair on the Damage and Economic Importance forehead sides, and they are about 13–15 mm long. Forager workers have less hair and they are about Leaf-cutting ants attack leaves, tender buds and 9–10 mm long; fungus-cultivator workers are lighter lowers of the cofee plant. he leaves of attacked colored and smaller (from 2 to 4 mm). cofee plants have semi-circular cuts or these plants are completely defoliated. Leaf fragments dispersed on the ground are seen around the defoliated plants. In Biology and Ecology recent attacks, the presence of ants carrying leaf and lower pieces may be observed. It is possible to detect Atta and Acromyrmex ants are social insects that earth mounds (nests) nearby or relatively far away. use plant leaves to cultivate symbiotic fungus Situations in which ants are direct plant pests (Leucoagaricus gongylophorus (Möller) Singer; are rare; however, in the tropical and sub-tropical Attamyces spp.), which serves as their foodstuf. areas of America, ants of the Atta and Acromyrmex hey form colonies constituted of three castes: genera can constitute important pests of many cul- queens, males and workers; the latter are sterile tivated and wild plants. In tropical America, cutting and present acute polymorphism and functions ants constitute the dominant group of herbivorous (soldiers, foragers, cultivators). he mating of animals, because they consume much more vegeta- queens with males takes place outside the nest tion than any other animal group. In the case of during the nuptial flight, at the beginning of cofee, these ants are generally considered of minor the first rains. Newly mated females dig their importance. Nevertheless, in some areas like the nests in the soil and begin to cultivate the fungi Turrialba region of Costa Rica, A. cephalotes attacks which will serve as their food and to lay their on cofee plants can be severe in monocultures. he irst eggs. he eggs give birth to the larvae and damage is caused by the worker caste when they cut ater 40–60 days, the irst adult workers emerge. the cofee foliage and lowers with their mandibles. In New colonies have a single tower-like mound of some Atta species, from 5 to 28 colonies/ha have small size of <200 cm2 in area, and with a small been observed, with the possibility of having one entrance hole, whereas older colonies are lat- or more millions of workers in each colony. he tened, with larger entrance holes and a colony nests they construct may have dimensions that vary surface area >200 cm2. he growth of the colony is between 30 and 600 m2. From one day to the other, very slow at the beginning, but during the second one or more cofee plants may be completely defo- and third year it accelerates rapidly and then it liated by these ants. Cofee plantations near woody diminishes as the colony starts production of or weedy areas are attacked more commonly. males and winged queens. Towards the end of the Coffee Pests and their Management C 989 third year, the population is enormous, and it is Cultural Control possible to observe more than 1,000 entrance/exit holes on the nest. he Acromyrmex nests are Queens should be eliminated at the recently formed simpler than those of Atta. In order to reach the nests using a grub hoe. Repellent plants (e.g., sor- plants which serve as their food, the forager work- ghum) should be sown. Because cofee on farms ers move from the nest, which is often in non- with low vegetational diversity is at greater risk of cultivated ields, through narrow paths which can attack by A. cephalotes, it is recommended that go more than 100 m in distance. he workers’ shade trees be planted in order to increase shade activity is more intense during the night. levels and therefore to decrease ant colonization. Also, it may be desirable to plant shade trees that are palatable to leaf-cutting ants, but that should Natural Enemies be either not commercially valuable (e.g., Erythrina poeppigiana [Walp]. Cook, Cordia alliodora (Ruiz Apparently these ants have few natural enemies. & Pav.) Oken, Swietenia macrophylla King, Cedrela Several predators, such as birds, toads, lizards and odorata L.), or that are tolerant of ant attack, in anteaters feed on the queens and males during the order to divert ants from foraging cofee plants. nuptial light. In Colombia, other carnivorous ants have been observed to be predators of leaf-cutting Biological Control ants. he importance of all these natural enemies in regulating leaf-cutting ant populations is unknown. Some entomopathogenic (e.g., Paecilomyces sp. and M. anisopliae) and antagonist (Trichoderma Management viride Persoon ex Gray) fungal strains have proved to be successful against leaf-cutting ant colonies in experimental studies. However, the practicality of he optimal time of the year to control leaf-cutting these fungi has not been assessed in commercial ants has not been determined. However, the nup- cofee plantations. tial light period, a crucial event within the ant life cycle, should be taken into account. Considering the ecological importance of leaf cutting ants as plant population regulators in woody and grazing Chemical Control areas, and taking into account that in certain areas they are eaten by humans, it is recommended Insecticides are applied directly through some of that population regulation, not elimination, be the entrance/exit holes of the nests, taking the the primary goal. precaution of plugging or closing most of them before. he “ant hill beating” procedure may also be used, which consists of digging in the nest Sampling with a shovel in order to uncover the ant brood, and spraying them with insecticide. Also, leaf- No sampling techniques have been developed for cutting ants can be successfully controlled using leaf-cutting ants in cofee plantations. he nests can baits containing insecticides. Treatment results be located by following the narrow paths used by can be improved by basing the amount of insec- the ants. Ant colony density can be estimated by ticide applied on an estimate of the colony sampling four 125-m2 plots at each edge of a farm volume, instead of surface colony area. An (north, west, east and south), and a single 500-m2 experimental study shows that mounds of dump area in the center of the farm (a total area of 1,000 m2). material can be used as a highly effective 990 C Coffee Pests and their Management small-scale deterrent to protect Hibiscus plants fungi may be involved in the damage (e.g., Phoma from defoliation by A. cephalotes, but this method costarricensis Echandi). has not been tested in coffee.

Description Long-Horned Grasshoppers or Katydids (Orthoptera: he eggs of I. subquadratum are brown in color, Tettigoniidae) elongated, with a hard chorion; they are oviposited in compact clusters. here are six nymphal instars. Distribution Newly emerged nymphs are fragile and gray in color. Nymphs resemble adults, but are smaller, Two Idiarthron species, I. subquadratum Saussure lighter colored, and lack wings. Nymphs and adults & Pictet and I. atrispinum (Stål), and one unknown have strong and large mandibles and the antennae Gongrocnemis species have been reported attacking are very thin and longer than the body. Adults have coffee in Mexico and Central America; apparently, a heavy set, more or less cylindrical body, greenish, I. subquadratum is present in Colombia too. Of brown-gray or light gray in color, with females these, I. subquadratum is the most important katy- from 5.0 to 6.0 cm long. Males are smaller. With did pest in coffee because very high infestations their thorny, strong and long back legs, they can have been reported in some coffee plantations in jump. heir ability to ly is limited and in general El Salvador and Mexico. Most of the information their movements are clumsy. Females have an ovi- available on katydids comes from this species. positor, from 1.0 to 2.3 cm long, at the tip of the abdomen, which looks like a spur or a knife point.

Damage and Economic Importance Biology and Ecology Attacked coffee leaves show irregular holes mar- ginally and centrally; feeding can also be observed Idiarthron subquadratum is arboreal, polyphagous, on the tender buds, shoots and branch tips. A char- and nocturnally active. Both nymphs and adults acteristic symptom of katydid damage is the leave their daytime shelters at night and disperse appearance of green and ripe fruits with damaged by jumping between tree and bush canopies. hey pulp, so that the coffee beans are exposed. he hide in shady places, such as dead leaves, rotten damage is caused by both the nymphs and adults. trunks and weeds; in particular, they take refuge in In cofee plantations where heavy attacks of I. sub- plants of wind-breaks, izote (Yucca guatemalensis quadratum occur, complete destruction of leaves, Baker) barriers, banana plants (Musa spp.) and buds and small branches, the fall of tender fruits Sanseviera sp. his species feeds on leaves and and the destruction of fruits can be observed. In fruits of several plants, including cofee (Coffea general, I. subquadratum is not an economically spp.), banana (Musa spp.), orange (Citrus spp.), important problem in cofee, although it some- chayote [Sechium edule (Jacq.) Swartz], and pacaya times may create some concern in certain areas of (Chamaedorea sp.). Mating occurs in plant cano- Central America and Mexico. In southeastern pies at night or in daytime shelters. Adult females Mexico (Siltepec, Chiapas), the most critical attack place their eggs in the soil and, in some cases, period is from June through November. Damage is under the bark. he eggs are placed in a mass more important in very shaded and abandoned (from 5 to 50), and one female may lay several cofee plantations. In addition to direct damage hundred. In Siltepec, Chiapas, Mexico, mating caused by I. subquadratum, plant pathogenic usually occurs in October, and oviposition occurs Coffee Pests and their Management C 991 in November and December. Adults are killed by the insects are killed manually and/or used as food low temperatures in January and February, and for domestic animals (e.g., chickens and dogs). eggs undergo diapause. At the beginning of the rainy season, between May and June, nymphs emerge and start to feed on coffee plants. Genera- Biological Control tions are overlapping in warmer regions. he life cycle from egg to adult is about 80 days at 28°C. In Some strains of B. bassiana kill nymphs in the lab- El Salvador, this pest is especially common in high oratory. However, the use of this biocontrol agent altitude cofee plantations. has not been attempted in ield.

Natural Enemies Chemical Control

Birds, spiders, parasitic nematodes and an unknown When infestations are heavy, the application of tachinid ly species (Diptera: Tachinidae) have chemical insecticides at the places of refuge is recom- been reported in Mexico. mended. Toxic baits placed inside the bamboo traps are also recommended. he most convenient period for chemical control is 1 month ater the beginning of Management rainfall and before oviposition takes place. Because high infestations of the pest have been related to low Sampling populations of natural enemies, insecticide use should be avoided in order to conserve natural control. “Shelter traps” made with a 10-cm-diameter by 30-cm-long bamboo (Bambusa vulgaris Schrad.) internode closed at one end, can be used for sam- Bush Crickets, Paroecanthus spp. pling I. subquadratum. he bamboo traps are (Orthoptera: Gryllidae) placed on cofee bushes upon the irst rainfall events, and during daytime are checked every Distribution week for captured insects. Bush crickets appear sporadically, afecting cofee Cultural Control plants and shade trees in cofee plantations in some areas of Central America and Mexico. he reported Weed control should be applied. he shade should species are Paroecanthus guatemalae Saussure be regulated. Trash and rotten trunks in the cofee (Guatemala, Honduras) and P. niger Saussure (El plantation should be prevented. Dry banana and Salvador, Guatemala). he Paroecanthus species plantain leaves should be eliminated. in Mexico remains unknown. Recently, high infes- tations of bush crickets have been reported in Honduras. Mechanical Control

Bamboo traps as described earlier for sampling Damage and Economic Importance can be used for elimination of I. subquadratum. he traps are placed in dark spots of the plantation Paroecanthus spp. attack ligniied stems and and in the vicinity of the plants that are normally branches of cofee bushes and shade trees. he used as refuge. he traps are checked weekly and afected plants show small marks or holes, 3.0 mm 992 C Coffee Pests and their Management in diameter by 1.0 mm in depth, distributed in line cofee plant. hree or four generations appear throughout the affected stems and branches. his per year. he attacks are more severe in unshaded mark along the stem gives it the appearance of a cofee plantations. In Honduras, acute infestations lute; hence this damage is known as “lute disease.” have been reported in the dry season in plantations If the stem or branch bark is lited right below each located between 900 and 1,250 m above sea level. hole, an “X” shaped scar on the wood may be observed. he damage is caused when the adult female of the cricket lays its eggs. Heavy attacks of Natural Enemies the bush cricket (when there are many holes), may cause physiological disorders in the cofee plant, An egg parasitoid wasp, Acmopolynema sp. which afects its development. he cricket can be a (Hymenoptera: Mymaridae), has been reported in pathogen vector, or perhaps the lesions may favor Honduras and Mexico. the penetration of diseases. A severe infestation can kill the cofee plant. In Honduras, where high infes- Management tations have been reported, the afected plants develop a yellowish color and they lose leaves and Sampling fruits. Scouting should be conducted to determine the Description limits of the infestation during the dry season. Upon detection of damage, the trunk bark should be he egg is white with elongated shape (1.0 by scratched in search of the insect’s eggs. If the damage 5.0 mm). Nymphs are similar to adults, but their is recent, the perforations are white and unhatched wings are not well developed and they are smaller eggs shall be observed; if the damage is old, the than adults. he adult has a cylindrically shaped perforations are dark and the eggs have hatched. body and is 2.0–2.5 cm long. he legs are yellowish in color and the abdomen is dark brown. he Cultural Control antennae are iliform and their length is almost twice the size of the body. In the female, the wings Weeds should be controlled within and on the do not cover all of the abdomen, which at its tip edges of the plantation. Severely damaged plants shows the cerci and a long pin-shaped ovipositor. should be re-planted, or pruned of the afected stems, and burned thereater to eliminate the eggs. Biology and Ecology

he bush cricket is active at night, while during Biological Control daytime it takes refuge in dark places in the weeds, dead leaves and some plants such as bananas (Musa his has not been attempted. spp.) and izotes (Y. guatemalensis). Only on cloudy days and when it is very abundant can it be seen during the day. he female lays about eight eggs in Chemical Control each oviposition hole, distributing two on each end of the scar it makes on the wood, in an “X” shape. In plantations that are close to the afected planta- he nymphs emerge in about 3 weeks, and they go tions, a preventive insecticide application with a brush through several molts for 3 months before becom- or a manual pump is recommended, treating from ing adults. Nymphs and adults can feed from the the stem base up to 60.0 cm high. Application may be Coffee Pests and their Management C 993 repeated once or twice every 20 days. Also, an insecti- species, such as Phobetron hipparchia (Cramer), Sib- cidal dust can be directed to the main stem, to the soil, ine spp., Olceclostera moresca (Schaus), Megalopyge and to the plantation edges during the dry season. lanata (Stoll & Cramer) and Automeris sp., among others, have urticating hairs which cause painful lesions to anyone touching them. Measuringworms, Leaf-Eating Caterpillars which are active nocturnally, possess camoulage (Lepidoptera) which allows them to go unnoticed during the day. In general, the pupation takes place in the soil. In Distribution Ecuador, Automeris sp. and Eacles masoni Schaus appear cyclically during the rainy season. he adults here is a large and diverse group of leaf-eating or moths have nocturnal habits. Insecticide abuse caterpillar species in tropical American countries and climatological changes can afect the natural afecting cofee. he principal leaf-eating caterpil- enemies of leaf-eating caterpillars, so their popula- lars are shown in Table 17. tions may increase and become damaging.

Damage and Economic Importance Natural Enemies

Cofee bushes afected by leaf-eating caterpillars here are many natural enemies of leaf-eating cat- show totally or partially consumed leaves. Some- erpillars. Among them, birds, parasitic Hymenoptera times the fruits are also afected. Eventually, and Diptera, and fungal, bacterial and viral dis- voracious worms or caterpillar larvae, as well as, eases are notable. their feces, can be observed. Some of these are urticating caterpillars. hese insects are frequently mentioned in the cofee pest manuals of South Management American countries, such as Brazil and Colombia. Some species even defoliate entire sections of Sampling the cofee plantation. Regular inspection of the cofee plantation should be made to detect initial infestation sources. Description

As example of leaf-eating caterpillars, Oxidia sp. Mechanical Control (Geometridae), is described. When small, they are black, and when large, they are light gray. hese he larvae of urticating worms should be elimi- caterpillars attain a length of 5.0–6.0 cm. he lar- nated manually using gloves. In the case of mea- vae are called inchworms or measuringworms. suringworms, the same can be done, but at night. Adults should be eliminated with light traps.

Biology and Ecology Biological Control Adults lay their eggs individually or in groups on the foliage of various plants. he larvae or worm feeds Bacillus thuringiensis Berliner can be used, partic- on the foliage. he caterpillar goes through several ularly at the beginning of infestations when the molts, and as it grows, feeds itself voraciously. Some caterpillars are small. 994C Coffee Pests and their Management Chemical Control Biology and Ecology

Some organophosphates and pyrethroids are Adults feed from the foliage of cofee and other recommended. In general, the use of chemical plants. A distinctive characteristic of these weevils insecticides is not necessary, because the natural is that when they feel threatened they contract enemies provide regulation of the populations of their legs and snout and let themselves fall to the these leaf-eating caterpillars. hus, it is important to ground where they seemingly disappear. heir preserve the natural enemies, and use insecticides eggs are laid in the soil and the larvae lead a sub- only in extreme cases. terranean life (between 10.0 and 20.0 cm deep), feeding from weed roots, including the cofee plant root. he weevil populations are higher from June Leaf Weevils (Coleoptera: through August in Honduras. In Brazil, Pantomo- Curculionidae) rus leucoloma (Boheman) is more frequent in the summer and it attacks both C. arabica and Distribution C. canephora. In Honduras, the most frequent attacks appear in the highest altitude areas. Very Various leaf weevils are present in cofee planta- weedy areas favor infestation. tions in tropical America (Table 17). Natural Enemies

Damage and Economic Importance Predation by assassin bugs (Hemiptera: Reduvii- dae) is reported in Costa Rica. Leaf weevils attack the cofee bush leaves. he leaves show irregular holes, tearing and notches on their edges, oten beginning at the tip and from Management the edge towards the vein. he most afected parts are new leaves and shoots. he damage is caused Sampling by the adults, which feed on the cofee foliage. he attack of these weevils can become important Tender buds and new leaves of the cofee plants when they afect the buds of recently pruned plants should be checked. When the damage only appears and of trees <1 year old. he lesions caused by on old leaves and not new ones, no control mea- this pest on the leaves may favor the infection of sure should be initiated. P. costarricensis.

Cultural Control Description Weeding should not be complete, so that adult and Larvae are whitish and legless. he color of adults larvae weevils have a feeding source and abstain varies according to the species, being of-white from attacking the cofee plants. (Compsus sp.), light brown with yellow spots (Macrostylus sp.), grayish, light brown or black (Epicaerus capetilensis Sharp.) or green. heir size Biological Control varies from 9.0 to 13.0 mm. he snout is fairly well developed in these insects. It has not been attempted. Coffee Pests and their Management C 995 Chemical Control 5.0 to 7.0 mm. hey have about ive molts. Pupal size varies from 3.0 to 4.0 mm. he adult is oval, When the populations are large, applications of with an arched body, covered by hairs and with a insecticides to the foliage and then to the soil are length of 2.5–4.5 mm. he head has round promi- recommended. nent eyes, with a short, wide, curved-downwards “snout” and the mouthparts distally. Coffee Bean Weevil, Araecerus fasciculatus (De Geer) (Coleoptera: Biology and Ecology Anthribidae) Adult females lay eggs on the parchment cofee Distribution grooves, placing one per grain and approximately three per day. he average number of eggs laid is Present in all coffee growing countries in America. 52. Larvae create galleries in the seed, and they pupate there also. he life cycle, from egg to adult, is 35–40 days. Between 55 and 74% of the descen- Damage and Economic Importance dants are composed of females. Infestation is more acute on sotened cofee beans. Up to ten genera- Araecerus fasciculatus attacks stored coffee beans. tions are reported per year. Coffee beans stored in warehouses, coffee mills and other places used to gather the harvest will show per- forations and irregular and relatively large galleries Natural enemies caused by this weevil. Accumulation of a fine yellowish powder is also observed. Highly infested In Colombia, the following natural enemies of warehouses will have a large number of little beetles, A. fasciculatus have been reported: Anisep- +the walls, roofs and windows. his weevil, which toromalus calanadrae (Howard) (Hymenoptera: attacks a wide variety of grain in storage, is considered Pteromalidae), Cephalonomia gallicola (Ash- as one of the few economically important pests of mead) (Hymenoptera: Bethylidae), Cheyletus sp. stored cofee in the American countries, particularly (Acari: Cheyletidae) and Monieziella sp. (Acari: in South America. It creates problems in warehouses Tyroglyphidae). that store poorly processed cofee containing more than 12% humidity. he damage is caused by the weevil larvae, which live in and feed on the grains. Management he attack is also favored when the warehouse tem- perature is higher than 27°C and relative humidity is Sampling above 60%. In 6 months of infestation, losses of 30% have been estimated. C. arabica apparently is more Fortnight visits should be made to the storehouses susceptible than C. canephora. he fruits that remain to check the presence of weevils, particularly in the on the plant ater harvest may also be attacked. wet season and in places with very humid weather.

Description Cultural Control he larva is without legs, white, with a “C” shaped body and a relatively wide thorax. he head is Adequate fertilizing, harvesting and pulp extrac- small, light brown in color. hey measure from tion should be conducted. Cofee should be stored 996C Coffee Pests and their Management with adequate humidity. Warehouses and storage a magnifying glass, little animals moving on the leave places should be kept clean. Infested lots should be can be observed, and in general, silky threads which set aside and placed in the sunlight. retain dust and other residues. he damage, which consists of the destruction of supericial cells of the leaf, is caused by immature and adult mites when they Bilogical Control feed. Spider mites may be especially important in some areas of tropical American countries during It is not conducted. abnormally dry weather. In severe attacks the leaf functions are interrupted and they may drop. Leaf defoliation and yield decreases may occur when more Chemical Control than 30 mites per leaf are present, particularly under dry weather conditions. he economic importance of In the case of preventive treatments and the treat- some species of mites on cofee, for example, Polyphag- ment of infested lots, fumigation is recommended. otarsonemus latus (Banks) (Acari: Tarsonemidae) in Ater fumigation, spraying of a 3-month residual Brazil, is unknown. efect pyrethroid with motorized equipment is recommended. he preventive treatment should be conducted when there are 1–2 weevils/m2 of Description sacks. Treatment of the walls, loor and roof of the warehouse where the cofee is going to be stored is he egg is elliptic or spherical, bright orange, red- also recommended. dish or red in color, depending on the species. Its length varies from 0.100 to 0.127 mm. he larva has three pairs of legs, an almost circular body, Spider Mites (Acari) and according to the species, orange or yellow col- ored when hatching, turning green-yellowish as Distribution they feed. hey are from 0.15 to 0.16 mm long. Nymphs (protonymph and deutonymph) have At least, six spider mite species have been four pairs of legs, and they are ovoid and about recorded in cofee in tropical America (Table 17). 0.20 mm long. In the deutonymph, which is larger, Olygonychus (Acari: Tetranychidae) is the most females (0.20–0.26 mm) can be diferentiated from representative genus. males (0.18–0.23 mm). Adult females are larger (0.28–0.50 mm) and more oval than the males (0.25–0.35 mm). Color varies according to the Damage and Economic Importance species and the sex; however, colors such as red and orange are blended, and in some cases the Spider mites attack cofee foliage in all their stages of mites have spots. he broad mite, P. l atu s , has a development. Attacked plants present yellowish, white-milky color and it is smaller than the other brown or copper colored leaves, with more undulated species (0.15–0.20 mm). edges. Sometimes the attacked leaves may dry up and fall. Also, the leaves lose their shine and present a dirty appearance. he symptoms take place in large patches Biology and Ecology in the cofee plantation, and more frequently in old, poorly attended cofee plantations, and near the roads. Adult females reproduce sexually and parthenoge- hese symptoms are easily recognized at a distance. netically. he eggs are laid one by one, preferentially Upon examination of the upper face of the leave with on the upper face of leaves, close to the veins, Coffee Pests and their Management C 997 although P. l atu s , unlike the others, prefers the determined by making parallel inspection routes lower side of the leaves. he eggs may be ixed to 25 m apart from each other, and examining 24 the leaf with the silk threads (cobweb) produced by leaves at random from four cofee plants every 25 m. the mites and which serves for protection and for moving from one leaf to another. Unlike the Tet- Cultural Control ranychidae, Tenuipalpidae (Brivipalpus sp.) do not produce silky threads. Egg laying, in the case of Shade trees should be planted in very sunlit areas. Olygonychus coffeae (Nietner), occurs at a rate of Weed control should be conducted. Adequate fer- 4–6 eggs/day/female for 2 or 3 weeks. Upon eclo- tilizing should be applied. sion, the larvae feed from cells that they puncture with their chelicerae, and in time they become prot- onymphs and the latter become deutonymphs. At Biological Control the end of their development, both protonymphs and deutonymphs go through an inactive stage Not applied. called “quiescence.” An accumulation of various residual materials such as dust and the old exuviae of spider mites can be observed in the cobweb pro- Chemical Control ducing species. he egg to adult life cycle varies from 8 to 28 days, according to the temperature. Some pesticides have a selective action, affect- Females mate with one or more males, and a male ing only mites, and others (non-selective) kill may fertilize several females. Females, which are mites and insects. In case of a simultaneous more abundant than males, disperse from one leaf attack by mites and leaf miners, non-selective to another and from one cofee plant to another, by products are recommended. However, the over- the use of silk threads. However, the factors that use of this practice can negatively affect the contribute the most to dispersion are the wind, beneficial parasitoids and predators. Applica- humans and other animals. Spider mites prefer to tions should be made only to infested areas. colonize the sunlit cofee plants and the older leaves, Various pesticides are recommended, making although in severe infestations they also attack the one application and sometimes a second one. A young leaves. population of 30–40 spider mites per leaf in the dry season cause defoliation, so this density must be avoided. Natural Enemies

Predators such as ladybirds (Coleoptera: Coccinel- References lidae) and rove beetles (Coleoptera: Staphylinidae) are reported. However, the literature on cofee pests Barrera JF (ed) (2002) Tres plagas del café en Chiapas. El is not clear about the predator species present. Colegio de la Frontera Sur. México, 198 pp Cárdenas-M R, Posada-F FJ (2001) Los insectos y otros habi- tantes de cafetales y platanales. Comité Departamental Management de Cafeteros del Quindío. Armenia, Colombia, 250 pp Castillo-Ponce G, Contreras-J A, Zamarripa-C A, Méndez-L I, Vázquez-M M, HolguínM F, Fernández-R A (1996) Tec- Sampling nología para la producción de café en México. Instituto Nacional de Investigaciones Forestales y Agropecuarias. he plantation should be checked during the México. 88 pp. Primera reimpresión. Folleto Técnico Núm 8 summer or dry periods, preferably on roadsides. Cofee Industry Development Company Ltd (1986) Growing The infestation of a coffee plantation plot is cofee in Jamaica. Jamaica, 103 pp 998 C Cold Tolerance in Insects García-G A, Campos-A O, Barrera-S CA, Meoño-R JE (1998) Classification of Cold Tolerance Manual de caicultura. Tercera edición. Asociación Nacional del Café, Guatemala, 218 pp Le Pelley RH (1973) Las plagas del café. Editorial Labor, SA, Insect cold tolerance classiications have tradition- Barcelona, 693 pp ally been divided into freezing tolerance and freeze Matiello JB (1991) O café. Do cultivo ao consumo. Publica- intolerant strategies. his division has been criti- ções Globo Rural. Coleção do Agricultor. Grãos. Editora Globo, SA, Brasil, 320 pp cized in recent years by a number of investigators. Muñoz-H R (2001) Plagas insectiles del cafeto. In: Manual de he arguments for the classiication scheme have caicultura. Instituto Hondureño del Café, Honduras, pp depended on the deinitions applied to the two 115–142 Páliz-S V, Mendoza-M J (1993) Plagas del cafeto. In: Manual terms, and it is how freezing tolerant and freeze de caicultura. Estación Experimental Pichilingue. GTZ, intolerant species have been deined that evokes the FUNDAGRO, Quevedo, Ecuador, pp 144–166 controversy. For example, in freeze tolerance, these insects are said to be capable of withstanding ice formation in some or nearly all parts of the body Cold Tolerance in Insects and associated luids. Most insects in this grouping usually freeze at temperatures between −5 and david rivers −10°C, though others require lower temperatures. Loyola College in Maryland, Baltimore, MD, USA Once frozen, these species can tolerate cooling to much lower temperatures, and upon thawing, the Exposure to low temperatures is among the most insects recover and apparently resume normal important abiotic factors limiting the range of development and behaviors. Some experts, how- insects in temperate climates. he relationship ever, have contended that this example of freeze tol- between insects and cold is dynamic, particularly erance is at the extreme end of cold tolerance and when considering the actual temperature at the only represents insects which are most suited to surface of the integument versus internal and/or survive low temperatures. An examination of some ambient conditions, the length of exposure to low 60–70 species of insects classiied as freezing toler- temperature, and the degree of temperature luc- ant has led to the suggestion that there are distinct tuation over a deined period of time (e.g., day, freeze tolerance strategies that allow insects to be week or winter season). hese issues make it chal- grouped based on supercooling points (SCPs) and lenging to categorize insect tolerance to a speciic lower lethal temperature (LLT): (i) partially freeze set of temperatures, particularly in terms of sur- tolerant species that survive a small portion of their vival. As poikilotherms, although some are hetero- body water converted to ice; (ii) moderately freeze thermic under speciic conditions, insects have tolerant species, if the exposure is suiciently long, adapted to cold environments resulting in exten- die at temperatures <10° below their SCP; (iii) sion of locomotor and/or reproductive activity strongly freezing tolerant insect species display LLT during low temperature exposure, enhancement of twenty degrees or more below their SCP; and (iv) metabolic rate, and maintenance of a positive freezing tolerant species possess very low SCPs and energy balance. he implications to many of these freeze at extremely low temperatures. Insects in this insects are a lengthening of the life cycle and a latter group are capable of surviving at tempera- requirement for individuals to overwinter one or tures a few degrees below their SCP. more times. he actual mechanisms associated Insects that are not tolerant of any ice forma- with these adaptations have received extensive tion in their bodies are generally termed freeze study in recent years, including attempts to deci- intolerant species. he natural tendency has been pher the underlying genetic basis of individual to assume that these insects will die if tissues or and population responses to low temperatures and body luids freeze, and presumably if they avoid seasonal change. the frozen state, these insects will survive. Such