See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/273621871 Morphology and Anatomy of Stem Mines in Cipocereus minensis (Wender.) Ritter (Cactaceae), an Endemic Species to Eastern Brazil Article in Haseltonia · January 2012 DOI: 10.2985/1070-0048-17.1.5 CITATIONS READS 3 278 4 authors: Deusa Abreu Emilia Arruda University of Cambridge Federal University of Pernambuco 5 PUBLICATIONS 83 CITATIONS 37 PUBLICATIONS 218 CITATIONS SEE PROFILE SEE PROFILE Gladys F. A. Melo-de-Pinna J. Hugo Cota-Sánchez University of São Paulo University of Saskatchewan 75 PUBLICATIONS 615 CITATIONS 69 PUBLICATIONS 1,375 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Systematics of Ceropegieae View project FLORAL ANATOMY AND CHEMICAL PROFILE IN EPIPHYTIC CACTI FROM THE BRAZILIAN ATLANTIC RAINFOREST View project All content following this page was uploaded by Gladys F. A. Melo-de-Pinna on 26 May 2015. The user has requested enhancement of the downloaded file. Haseltonia 17: 42–50. 2012 42 MORPHOLOGY AND ANATOMY OF STEM MINES IN CIPOCEREUS MINENSIS (Wender.) RITTER (CACTACeae), AN ENDEMIC SPECIES TO EASTERN BRAZIL Deusa D. Abreu1,2 Emilia Arruda3 G. Flávia A. Melo-de-Pinna1 J. Hugo Cota-Sánchez2,4 1 Universidade de São Paulo, Depto. de Botânica, CEP 05422-970, São Paulo, SP - Brazil 2 University of Saskatchewan, Department of Biology, Saskatoon, SK S7N 5E2 – Canada 3Universidade Federal de Pernambuco, Depto. de Botânica, CEP 50670-901, Recife, PE - Brazil Abstract: Plant mines are structures with the form of a cavity caused by consumption of host plant tissue by the insect’s miner larvae. Plant mines are more common in leaves, but in Cipocereus minensis, a species in which the leaves are modified spines, the miner activity is restricted to the stem. The aim of this paper was to document the morphological and anatomical differences in the infected and uninfected stems of C. minensis due to the feeding habit of the mining agent. Fresh tissue samples of non-mined and mined young stem of C. minensis were collected and examined in transverse sections. We hypothesize that the infection begins follow- ing mating when the females scratch the surface of the stem or while they feed on fruits and lay eggs, which subsequently develop into larvae, invading the cactus stem. The insect’s miner larvae had mostly consumed the parenchyma tissue towards the center of the stem, and periderm formed along the entire path of the in- sect. This meristematic tissue or “wound periderm” is a common response for compartmentalization to isolate the damaged tissue, in this case the incubating chamber, in which the eggs will be placed. There were no signs of consumption of vascular tissue in the infested samples, further suggesting a compartmentalized infestation. The nest chamber was found in the stem pith region, with periderm surrounding an insect’s miner pupa in- side identified as a member of the Cerambycidae. The mining insect depends on a host plant to complete the life cycle; however, the nature of this partnership and the long-term effects of the insect on the plant tissue are unknown. The complex mechanisms by which herbivorous insects control the morphogenesis of the plant host are discussed. We propose that C. minensis has a recognition system to identify insect attack and evaluate the effectiveness of early response triggering compartmentalized defense mechanisms by protecting the injured area with a new layer of periderm. Keywords: Stem anatomy, Cactaceae, Cerambycidae, Cipocereus minensis, controlled morphogenesis, recogni- tion system, stem mines. Introduction defensive mechanisms. However, in spite of the plants’ protective devices, insects are often successful Herbivory occurs in a wide group of insects in in fulfilling dietary and life cycle requirements. The predatory or parasitic forms. Among herbivorous in- relationship between mining insects and plants is ex- sects, mining species are remarkable because they are ceptional in terms of the insects’ specialization and adapted to specific environments and exhibit highly ability to survive in the microclimatic conditions, on specialized feeding habits. The complex nature of a selective and restricted diet provided by the host these plant-insect interactions involves intricate strat- plant (Csóka 2003; Schoonhoven et al. 2005). egies and adaptations that allow plants to overcome Plant mines are structures with the form of a cav- insect attack by deterring or reducing its effects using ity caused by consumption of host plant tissues by 4 Author for correspondence (Email: [email protected]) HASELTONIA VOL. 17. 2012 43 insect miner larvae living inside the plant. Insect the subfamily Opuntioideae, which has numerous mines are more common in leaves, but their occur- invasive species, in particular outside of their native rence has also been reported in root, stem, petiole, geographic range of distribution. In this regard, the flower, and fruit (Hering 1951; Weis and Berenbaum larvae of Cactoblastis cactorum Bergroth (Lepidop- 1989). The larva of the leaf miner is endophytic, lives tera) have been used for the biological control of inside the epidermal tissues, and depends on a host Opuntia monacantha (Willd.) Haw. in places where plant for shelter and food. The diet of the developing this species has colonized large areas (Lenzi et al. larva includes a selection of nutrients from the sur- 2006). However, the occurrence of cactus-feeding rounding subepidermal tissues of the host plant, in insects associated with members of the other cac- which the epidermis remains intact, and as it feeds, tus subfamilies, e.g., Cactoideae, Pereskioideae, and the larva forms small channels or even a small hollow Maihuenoideae, remains a mystery. Zimmerman and space, where it lives for part of its life cycle (Csóka Granata (2002) indicate that in the Cactaceae inter- 2003). nal cactus-feeding insects are more common than Mining insects have been documented in ap- external feeders, nearly 75% versus only 3%, respec- proximately 10,000 species and ca. 50 families en- tively. The remainder of insects associated with cacti compassing the orders Coleoptera (beetles), Dip- includes sapsuckers (15%) and fruit feeders (6%). In tera (flies), Hymenoptera (wasps), and Lepidoptera turn, mining insects represent a very small propor- (moths) (Connor and Taverner 1997; Csóka 2003). tion (only 1%) of the insect diversity associated with The species in these taxonomic orders undergo sev- cacti (reviewed in Zimmerman and Granata 2002). eral stages (instars) of metamorphosis during their The literature discussing the occurrence of mines life cycle. The largest order of insects, Coleoptera in tropical plants is restricted, and the anatomical (beetles), exhibits a great deal of ecological and mor- changes and effects on mined versus non-mined leaf/ phological diversity; however, few species have the stem are yet to be investigated. Much less is known mining habit. In fact, the most species-rich order about the anatomy and characteristics of the nest- of leaf-mining species is the Lepidoptera, but leaf- ing chambers of mining insects, in plants in general mining beetles have been reported in three families, and in cacti in particular. With the exception of a including jewel beetles (Buprestidae), leaf beetles few examples, the lack of factual information limits (Chrysomelidae), and weevils (Curculionidae) our understanding of the mechanism of action of (Csóka 2003). the mining agent and the degree of morphological Mines in plants have been documented early in transformation in the plant parts and tissues. Here the literature in several gymnosperm and angiosperm we present the case of a mining insect and convey families, such as the Rosaceae (Nawa 1916), Faga- information comparing the anatomy of mined and ceae (Suenaga 1938), Boraginaceae, Chenopodiaceae, non-mined tissues in the stem of Cipocereus minen- Dipsacaceae, Solanaceae (Hering 1951), Asteraceae sis (Wender.) Ritter, a columnar cactus endemic (Hering 1951; Melo-de-Pinna et al. 2002), Pinaceae, to eastern Brazil and easily identified by the bluish Betulaceae (Kumata 1964), Clusiaceae (Almeida- coloration of the fruits. Our goals were 1) to report Cortez and Melo-de-Pinna 2006), Malpighiaceae the occurrence of mines in the stem of C. minensis (Flinte et al. 2006), and the Apiaceae, Balsamina- caused by endophytic insect larvae, and 2) to high- ceae, Caryophyllaceae, Crassulaceae, Euphorbiaceae, light the morphological differences in the infected Fabaceae, Gentianaceae, Liliaceae, Poaceae, Rubia- and uninfected plants and to describe the nesting ceae, Salicaceae, Violaceae, among others (see Csóka area and the stem’s anatomical transformation due 2003). In spite of the vast number of mining insects to the feeding activity of the mining agent. A brief in plants, most studies have focused on the econom- discussion of the ecological and evolutionary advan- ic importance because several plant parts, e.g., leaves tages of this plant-insect association and the complex and fruits, of agricultural and ornamental species are mechanisms by which endophytic herbivorous insect damaged. On the other hand, mining insects have larvae control the morphogenesis of their host, is been used in the biological control of certain plants. also provided. In New Zealand, for example, Oemona hirta F. (Ce- rambycidae) causes the death of the stem of Car- Materials and Methods michaelia R. Br. (Fabaceae),