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A New Gall-Inducing Shining Leaf Beetle (Coleoptera: Chrysomelidae) from Thailand and Its Relevance to the Evolution of Herbivory in Leaf Beetles

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A New Gall-Inducing Shining Leaf Beetle (Coleoptera: Chrysomelidae) from Thailand and Its Relevance to the Evolution of Herbivory in Leaf Beetles A new gall-inducing shining leaf beetle (Coleoptera: Chrysomelidae) from Thailand and its relevance to the evolution of herbivory in leaf beetles Fredric V. Vencl1 and Kenji Nishida2 Abstract. Extremely few gall-inducing Coleoptera attack Monocotyledonae. We de- scribe and report on a new species of true shining leaf beetle, whose larvae form complex, multilayered galls on the host plant. During a field survey in Western Thai- land, a galling member of the genus Oulema (Criocerinae) was discovered in a pre- montane rainforest attacking Commelina paludosa, a member of the monocotyle- donous dayflowers (Commelinaceae). Our report is the first instance of a galling member of the genus Oulema and is described herein O. reclusa. The galler’s adult habitus exhibits the general traits typical of the genus Oulema: front of the head be- tween the eyes narrow, with the sides forming an angle of more than 90°; emarginate pronotum, moderately constricted below the middle; antennae filiform, longer than half the body length; two small protuberances on the vertex, almost touching; claws connate; 9th elytral stria with a complete row of punctures. The larva is extremely reduced, lacking discernable setae, a dorsal anus, ambulatory warts and ocelli. Our observation of a coleopterous gall-inducer may be the first on a monocot. Despite the fact that the putative sister taxa of the Criocerinae are the Sagrinae and the Bruchidae, gall-inducers and seed feeders respectively, we believe that gall-induction, as well as leaf mining in Oulema, are derived trophic habits within the Criocerinae. We discuss the relevance of this discovery to the evolution of ecto- and endophagy in the Criocerinae, a very basal subfamily within the broad Chrysomelidae radiation. Keywords. gall, Oulema, Commelina, Monocotyledonae, Chrysomelidae, Criocerinae. 1 Ecology and Evolution, Stony Brook University, Stony Brook, ny, 11794-5245 and The Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Ancon, Republic of Panama (from U.S.A.) unit 0948, apo aa 34002. [email protected] 2 Sistema de Estudios de Posgrado en Biología, Escuela de Biología, Universidad de Costa Rica, 2060 San José, Costa Rica. [email protected] A new gall-inducing shining leaf beetle (Coleoptera: Chrysomelidae) from Thailand 247 1. Introduction Extremely few gall-inducing Coleoptera attack the Monocotyledonae. Here we re- port on a true shining leaf beetle, whose larvae form complex, multilayered galls on its monocotyledonous host plant. During a field survey in Thailand, the second author discovered a galling member of the chrysomelid subfamily Criocerinae as- sociated with a monocot in the dayflower genus Commelina (Commelinaceae). The Criocerinae are a cosmopolitan group of nearly 2000 species distinguished by their brilliant, often colorful appearance, narrow, sometimes hour-glass shaped pronotum, and excrement-covered, folivorous larvae. Subsequent inspection sug- gested that this is the first instance of a galling member of the genus Oulema De Gozis, and is described herein. With the exception of the mining Oulema pumilo Vencl and Aiello, and Lema quadrivittata Boheman, the putatively galling Orthole- ma samalkotensis Heinze and O. abnormis Heinze, and several stem bores (Schmitt pers. com.; Vencl and Aiello 1997;Venclet al. 2004), the larvae of the vast majority of shining leaf beetles are external grazers on leaf surfaces of both Monocotyle- donae and Dicotyledonae (Monrós 1960; Schmitt 1988; Jolivet and Hawkswood 1995;Venclet al. 2004). The present case represents a notable exception and as such, an interesting problem for our understanding of the selective factors that influ- enced the evolution of trophic modes in larval Criocerinae in particular, and in exophytic phytophagous insects in general. Here we ask if internal foraging behav- iors, such as galling, boring and mining, were the progenitors of the leaf surface trophic mode, which now predominates within the Criocerinae, or, alternatively, whether internal feeding represents a highly derived habit, perhaps reflecting a re- treat from the high costs associated with the invasion of the leaf-surface adaptive zone (sensu Simpson 1953). We present below our understanding of this remark- able discovery. 2. Study site and habitat Galls were found on a dayflower Commelina paludosa Blume (Commelinaceae), at 962 m at Thong Pha Phum National Park (14°41´29.7N, 098°24´04.5˝E). The Park is located in Kanchanaburi Province, Thailand, which borders Myanmar (Fig. 1). The study site, a large bamboo thicket at bottom of a hill, was shaded, very humid and the ground was moist at the time of the investigation (Plate 29 A). The climate in this area is tropical pre-montane rain forest, with a southeastern monsoon rainy season lasting from May to July, and a northeastern monsoon with a distinct dry season lasting from November to January (Thong Pha Phum National Park 2004). 3. The host plant A perennial, C. paludosa Blume has erect, sometimes smooth, distally branched stems, which grow to 1 m. The leaves are lanceolate to ovate-lanceolate, sessile, 248 Fredric V. Vencl and Kenji Nishida Figure 1. Collection site of Oulema reclusa, sp. nov., Thong Pha Phum National Park, located in Kanchanaburi Province, Western Thailand (14°41´29.7˝N, 98°24´4.5˝E). generally glabrous, with sheathing petioles, reaching a length of 6 cm. The chief characteristics of C. paludosa are a three stamen-three staminoid configuration, two blue enlarged upper petals, and connate, spath-like involucral bracts (Plate 29 B). Galls may induce branching as they were frequently found at stem junctions. Growing in well-drained areas at mid-elevations, C. paludosa is distributed broadly from South Central China, Nepal to Malaysia (Shu 2000). 4. The gall Galls were collected at the beginning of the dry season in late October and early November 2005. Galls occurred on the host’s stems and induced stem branch points at leaf bases of the sheathing petioles and axillary shoots, possibly causing forma- tion of a new shoot. Although the host plants were common along the bamboo thicket, galls were uncommon and no plants had more than one gall. Of the seven at our disposal, mature galls were globose to spindle shaped, with an average (± se) length × width of 22.19 ± 2.44 × 15.79 ± 0.083 mm (Plate 29 C, D). The un-galled part of the stem (normal stems) averaged 2.5 to 3.0 mm in diameter. The outer sur- face was reddish in color and cracked in some galls (Plate 29 C). Each gall contained a single, narrow chamber with an average (± se) chamber length of 8.06 ± 0.71 mm, containing a single larva, pupa or adult. A relatively irregular margin of large, A new gall-inducing shining leaf beetle (Coleoptera: Chrysomelidae) from Thailand 249 succulent white to translucent ‘nutritive’ tissue surrounded the chamber, which appears to be modified parenchyma (Plate 29 E, F). The mature gall which con- tained a pupa or an adult had a brown (dried) circular, thin layer of the plant epidermis at the exit orifice. It is most likely that the mature larva constructs this circular exit orifice prior to pupation. 5. The adult Oulema reclusa Vencl and Nishida, new species Plate 29 G Diagnosis The adult habitus is 5.26 mm in length and of a uniform, pale yellow- ish-tan. The head is divided and distinctly bituberculate at the vertex. The head and pronotum are narrower than elytra; front of the head between the eyes nar- row, with the sides forming an angle of more than 90°; the claws are connate; the elytra lack any color patterning; and the 9th stria of elytral punctation is complete. Description: Form sub-cylindrical. Head: yellowish tan, shiny, moderately con- stricted behind the eyes and head much narrower than the width of the pronotum; clypeus, labrum and mouth parts with pale tannish-yellow ground color and finely setose; frons separated from the vertex by deep grooves forming an ‘X’ with the angle between them less than 90°; front of head between the eyes narrow, with sides forming an angle of more than 90°; the antennal tubercles almost touching and much closer together than the vertex-frons intersection; vertex weakly divided by a broad, longitudinal depression; antennae are of a uniform tan color, filiform, with 11 segments and longer than 1/2 the body length; segments 5 and 6 equal in length; eyes black, deeply notched to 1/3 their diameter; mandibles black at apices. Pronotum: Pale yellowish-tan, shiny, glabrous and emarginate, with a shallow, sub- medial constriction, 96% of apical width, near base; disc with a long, curved lat- eral seta projecting from each corner; a fine median pit in sulcus. Elytra: Shiny, uniformly yellowish-tan, with a pronounced basal depression; punctation coarse and uniformly spaced, becoming somewhat irregular to obsolete toward the apex; punctation of the 9th stria complete; scutellum tan. Venter: Uniformly yellowish- tan, evenly setose; minutely and evenly punctose; legs uniformly pale yellowish- tan; metafemora slightly swollen; two apical metatibial spurs; tibiae and tarsi very finely setose; claws connate and black. Holotype Female: Reared from a stem gall on Commelina paludosa, Thong Pha Phum National Park (14°41´29.7˝N, 098°24´04.5˝E), 962 m, Thailand, 2005. The pinned holotype is deposited in the U.S. National Museum with its gall voucher (dried). Emergence phenology The single gall to produce an adult was collected on 21 October 2004 and eclosed 25 days later. Host plant Commelina paludosa Blume (Commelinaceae). Distribution The only location known so far is the holotype locality, a mid- elevation, pre-montane rainforest above 500 m. 250 Fredric V. Vencl and Kenji Nishida Etymology Derived from Middle English, from Old French reclus, meaning hid- den or secretive from the world; originally from the Latin reclasus, past participle of recludare, to shut up or to close.
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