Natural History and Ecology of Soldier Beetles (Coleoptera: Cantharidae) in the Mexican Tropical Dry Forests

Natural History and Ecology of Soldier Beetles (Coleoptera: Cantharidae) in the Mexican Tropical... Abstract Until today, most information about the natural history and ecology of soldier beetles came from temperate zones, mainly from Holarctic areas, while tropical regions have been poorly studied. The aim of this contribution is to compile and synthesize information concerning the natural history and ecology of Cantharidae (Coleoptera) from the Mexican tropical dry forest (TDF), to serve as a starting point for more in-depth study of the group in one of the Mexico’s most endangered ecosystems. All compiled data on the family have been organized into the following topics: distributional patterns and habitat preferences, feeding behavior and host plants, and daily and seasonal activity cycles. For the first time, it was provided a list of host plants for TDF Cantharidae genera and species, and it was also observed a high ecological diversity in the phenology and behavior of TDF Cantharidae assemblages. Further research concerning cantharids and other TDF insects needs to have a more comprehensive and integrated approach toward understanding the patterns of distribution and diversity, and elucidating the role that cantharids play in ecosystems, especially in TDF, which is one of the most endangered ecosystem in the world. plant, insect, interaction, phenology, behavior Cantharids, or soldier beetles (Coleoptera: Cantharidae), are the most diverse group of soft-bodied terrestrial coleopterans (Ramsdale 2002). Cantharidae are usually classified into five subfamilies: Cantharinae, Chauliognathinae, Malthininae, Silinae, and Dysmorphocerinae, following the classification of Brancucci (1980). On the basis of molecular phylogenetic analyses, it has been proposed that, in fact, there should be only four subfamilies (excluding Dysmorphocerinae) (Sagegami-Oba et al. 2007). However, cantharidologists continue using Brancucci’s scheme (Constantin and Chaboo 2016) and an open discussion concerning the phylogeny of Cantharidae still remains. Soldier beetles are strictly terrestrial during their entire life cycle, and are distributed from ecosystems as cold as the Arctic tundra to the warm, dry African grasslands, ranging from the tropical dry forests (TDFs) of the American continent to the tropical rain forest of India. It is possible to find them in forests, meadows, savannahs, and grasslands, almost always among the foliage and inflorescences of their host plants (Ramsdale 2010, Pelletier and Hérbert 2014). Many species are associated with vegetation close to rivers, creeks, and other water bodies (Zaragoza-Caballero 2004, Ramsdale 2010). Members of this family are frequently classified as generalists in their feeding habits and habitats. However, the distribution of most cantharids is much more restricted than it was thought, since even within the same ecosystem each species may occupy a particular habitat, just as Fender described in his review of the North American Malthinini: ‘Each species appears to have a somewhat selective instinct for its preferred habitat’ (Fender 1951). This tendency of habitat specialization is noticeable up to the highest taxonomical hierarchies, although there are no studies concerning this issue. For example, Silini are evidently most diverse in tropical ecosystems, there are 119 Silinae species registered from Nearctic region, whereas in the Neotropical region are 819 (Delkeskamp 1977, Pérez-Hernández 2017); Chauliognathini prefer open areas, savannahs, and meadows within tropical ecosystems (Miskimen 1966), and Ichthyurini are common in tropical deciduous forests (Fender 1960). In contrast, Malthinini and Malthodini are more frequently associated with temperate coniferous forests, particularly on the foliage of coniferous and deciduous trees in shaded and humid environments such as marsh (Fender 1951, Ramsdale 2002). It has been hypothesized that Cantharidae habitat selection in tropical zones, and therefore distributional patterns, strongly depends on the distribution of adult host plants (Miskimen 1966, 1972). Nevertheless, as many species have predatory feeding habits and show an indirect relationship with plant species, another hypothesis is that specificity to particular environments is in fact determined by the abiotic requirements of the larvae (Ramsdale 2002). Unfortunately, there is a lack of information concerning the distributional patterns of Cantharidae and their specificity to certain habitats or their biotic interactions. This novel issue is in need of further study. Until now, there has been a paucity of information as to the natural history and ecology of soldier beetles from tropical regions. The aim of this contribution is to compile and synthesize the most relevant information regarding the natural history and ecology of Cantharidae (Coleoptera) from Mexican TDFs, to serve as a starting point for more in-depth study of the group in one of the most endangered ecosystems in the World (Miles et al. 2006). All compiled data on the family has been organized into the following topics: distributional patterns and habitat preferences, feeding behavior and host plants, and daily and seasonal activity cycles. Most data come from entomological samplings carried out in eight localities on the Mexican Pacific region. Materials and Methods Cantharidae specimens were studied in different years from eight Mexican localities, where TDF is the dominant ecosystem. From north to south, these are San Javier, Sonora; San Buenaventura, Jalisco; Ixtlahuacán, Colima; Sierra de Huautla, Morelos (Tepalcingo and Tlaquiltenango municipalities); Santiago Dominguillo, Oaxaca; Acahuizotla, Guerrero; and Huatulco, Oaxaca (Fig. 1). Light attraction, Malaise traps (except Acahuizotla), aerial nets, and beating of vegetation were all used for entomological samplings 5 d a month in each locality, always coinciding with the new moon. Specific details concerning sampling methods and study areas may be consulted in the studies by González-Ramírez et al. (2017), Noguera et al. (2007, 2012), Pérez-Hernández and Zaragoza-Caballero (2015), Rodríguez-Vélez et al. (2009), and Zaragoza-Caballero et al. (2003, 2009). In pursuance of the observation and documentation of the natural history and ecology of cantharids, Tepalcingo was visited in different months of 2014 and 2015, thus providing most of the information included in this study. Fig. 1. View largeDownload slide Map depicting distribution of TDF ecosystem in Mexico (map modified from Conabio, 1999). Localities and years of Cantharidae (Coleoptera) sampling for this contribution are showed. Fig. 1. View largeDownload slide Map depicting distribution of TDF ecosystem in Mexico (map modified from Conabio, 1999). Localities and years of Cantharidae (Coleoptera) sampling for this contribution are showed. Results Distributional Patterns and Habitat Preferences In Mexico, representatives of all Cantharidae subfamilies have been classified into 19 genera and 223 species (Zaragoza-Caballero and Pérez-Hernández 2014). More than 70 species registered in Mexican TDF belong to the genera Belotus Gorham (Coleoptera: Cantharidae), Chauliognathus Hentz (Coleoptera: Cantharidae), Ichthyurus Westwood (Coleoptera: Cantharidae) (Chauliognathinae), Plectonotum Gorham (Coleoptera: Cantharidae), Caccodes Sharp in Blackburn & Sharp (Coleoptera: Cantharidae) (Malthininae), Discodon Gorham (Coleoptera: Cantharidae), Ditemnus LeConte (Coleoptera: Cantharidae), Polemius LeConte (Coleoptera: Cantharidae), Silis Charpentier (Coleoptera: Cantharidae), and Tytthonyx LeConte (Coleoptera: Cantharidae) (Silinae) (Pérez-Hernández and Zaragoza-Caballero 2015). None of these genera or supraspecific taxa are restricted to TDF, as in other regions of the World (Hanson 2011); in fact, cantharids were observed in a high diversity of habitats from studied localities (Table 1), and with slight differences, congeneric species frequently inhabited similar environments. Compiled information describing the habitat preferences of TDF cantharid genera is summarized in the following paragraphs. Table 1. Statewide distribution, habitats, and observed trophic guilds of Cantharidae (Coleoptera) genera from TDF in Mexico Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa aSuggested trophic guild. View Large Table 1. Statewide distribution, habitats, and observed trophic guilds of Cantharidae (Coleoptera) genera from TDF in Mexico Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa aSuggested trophic guild. View Large In Mexican TDF, Malthininae species belong to Caccodes and Plectonotum. Very little is known about Caccodes habitat preferences, except that they can be collected by beating herbaceous and shrub vegetation, as well as through light and Malaise traps; members of the genus are presumably associated to closed habitats near creeks and rivers. Members of Plectonotum have been previously registered in South American rain forests, mountain forests, and paramos, as well as in Ecuador in zones ‘covered with highly diverse vegetation’ (Constantin 2008); consequently, the sole species registered in Mexico from TDF indicates more diverse habitat preferences in the genus, along with a larger geographical distribution in America (Pérez-Hernández and Zaragoza-Caballero 2015). Silinae members distributed in Mexican TDF belong to the Discodon, Ditemnus, Silis, Polemius, and Tytthonyx genera. In North America, Silis mostly inhabit the canopies of hardwood and mixed forests (Pelletier and Hébert 2014), and in Mexico, it is commonly found in the herbaceous stratum of tropical forests. In Mexican TDF, species of this genus are associated to shading or open habitats adjacent to water bodies, which usually present higher humidity than other habitats for most of the year (Fig. 2b). Ditemnus species have similar preferences to those of Silis members, also found more frequently in herbs. Discodon and Polemius species are associated to shady habitats and the canopy of shrubs, and they can usually be found in plants near water bodies or the forest edge (Fig. 2a, c, and d); they also spend most of the d perching on the underside of shrub leaves, such as those of Acacia cochliacantha Humb. & Bonp. ex Willd. (Fabales: Fabacaeae), and other bushes in shaded sites (Fig. 3). Members of Tytthonyx can be remarkably abundant in herbs and short shrubs. They are good flyers and can thus be collected using Malaise traps, which is usually an ineffective method for Cantharidae sampling. Fig. 2. View largeDownload slide Examples of habitats of Discodon (a, c, d), Silis (a), Polemius (c), and Chauliognathus (b) in Mexican TDF. Fig. 2. View largeDownload slide Examples of habitats of Discodon (a, c, d), Silis (a), Polemius (c), and Chauliognathus (b) in Mexican TDF. Fig. 3. View largeDownload slide (a, b) Discodon sp. preying on a conspecific individual; (c) a different species of Discodon, feeding on pollen; (d) Silis female eating a dipteran; (e, h) Chauliognathus forreri feeding on nectar and pollen; (f) C. nigriceps female feeding on nectar (Photo Enrique Ramírez); (g) Discodon perching on the underside of the leaves. Fig. 3. View largeDownload slide (a, b) Discodon sp. preying on a conspecific individual; (c) a different species of Discodon, feeding on pollen; (d) Silis female eating a dipteran; (e, h) Chauliognathus forreri feeding on nectar and pollen; (f) C. nigriceps female feeding on nectar (Photo Enrique Ramírez); (g) Discodon perching on the underside of the leaves. Members of Chauliognathus dwell in a wide range of environmental conditions and habitats, from temperate to tropical regions and from deserts to rainforests (Miskimen 1972). However, they are typically associated with herbs or shrubs in open, sunny areas (Fig. 2b). Belotus species are also associated to herbs or minute flowers, and frequent open fields adjacent to water bodies. Ichthyurus is a genus with a mainly tropical affinity, usually found in the canopy of shrubs. At the species level, one can observe a wide spectrum of habitat preferences, even among congeneric species. For example, members of Chauliognathus emerge with similar seasonality in TDF, and slight differences in habitat preference and seasonality may serve to avoid a more competitive coexistence. In this way, Chauliognathus forreri Gorham (Coleoptera: Cantharidae) has been exclusively observed visiting vines of Serjania triquetra Radlk. (Sapindales: Sapindaceae) at the beginning of the rainy season; halfway through that season, Chauliognathus dispar Champion (Coleoptera: Cantharidae), Chauliognathus nigrocinctus Gorham (Coleoptera: Cantharidae), and Chauliognathus profundus LeConte (Coleoptera: Cantharidae)—as well as C. forreri (Coleoptera: Cantharidae)—can be found on a variety of plants, despite being highly associated to flowers of Asteraceae, Apiaceae, and other herbs (Fig. 3; Table 2). It has also been recorded that Eupatorium L. (Asterales: Asteraceae) plants are known hosts of C. dispar in the subtropical dry forest (Miskimen 1966). Table 2. Association between some plant and Cantharidae species from studied Mexican TDFs localities Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey I, Ixtlahuacán, Colima; T, Tepalcingo, Morelos; Sierra de Huautla, Morelos. View Large Table 2. Association between some plant and Cantharidae species from studied Mexican TDFs localities Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey I, Ixtlahuacán, Colima; T, Tepalcingo, Morelos; Sierra de Huautla, Morelos. View Large Feeding Behavior and Host Plants Studies concerning the feeding behavior of Cantharidae larvae from tropical regions are very scarce (Jiron and Hedström 1985). However, available data depicting this stage indicate that generalist behavior is more frequent than specialization (Traugott 2003). In temperate regions, soldier beetle larvae are commonly registered as predominantly voracious predators whose survival critically depends on the availability of high-quality prey (Gambardella and Vaio 1978, Bilde et al. 2000). They generally feed on eggs, adult insects and other larvae, gastropods and earthworms (Ramsdale 2002, Eitzinger and Traugott 2011), whereas other species are facultatively or predominantly phytophagous, such as those of Silis (Ramsdale 2002). It is highly probable for soldier beetle larvae from TDF to show the same behavior. Most publications concerning the family affirm that most species are omnivorous and predominantly predators (Sunderland et al. 1987, Leksono et al. 2005, Jerinic-Prodanovic et al. 2010). Nevertheless, very few genera seem to present this pattern, such as Rhagonycha Eschscholtz (Coleoptera: Cantharidae), which visits the foliage and inflorescences of a great variety of plants where it feeds equally upon insects and nectar (Ramsdale 2002, Day et al. 2006). Ichthyurus are also mainly predators, and phytophagous to a lesser degree (Miskimen 1961). Members of Podabrini and Cantharidini from temperate regions have frequently been associated to the consumption of aphids (Aphididae) (Ramsdale 2010). In TDF, adults of Belotus, Chauliognathus, and Discodon were commonly observed feeding on nectar and/or pollen, whereas Discodon and Silis were observed feeding on insects (Fig. 3a and d). Chauliognathus adults have developed buccal structures adapted to feeding on nectar and pollen, mainly from plants with small, abundant flowers (Miskimen 1972, Ramsdale 2002), occasionally predate larvae of lepidopterans and adults of other beetles (Fender 1962) (Table 2). In TDF, Chauliognathus nigriceps Gorham (Coleoptera: Cantharidae) commonly feeds on Croton spp. (Malphigiales: Euphorbiaceae), whose flowering time begins a few weeks before the start of the rainy season; C. dispar, C. nigrocinctus, and C. profundus have been observed actively feeding on the nectar and pollen of Bidens odorata var. odorata Cav. (Asterales: Asteraceae), Melampodium divaricatum (Rich.) DC. (Asterales: Asteraceae), Tridax coronopifolia (Kunth) Hemsl. (Asterales: Asteraceae), Sanvitalia procumbens Lam. (Asterales: Asteraceae) and other herbs (Table 2). C. forreri has been observed visiting a wide spectrum of plant species (Table 2) and also robbing nectar by chewing a hole in the base of the Hamelia patens Jacq. (Gentianales: Rubiaceae) flowers to avoid conspecific competitors. Belotus present feeding behavior similar to that of Chauliognathus species, although consuming different plant species. For example, they have been collected feeding on Croton fragilis Kunth (Malphigiales: Euphorbiaceae) in Ixtlahuacán, Colima and on unidentified Apiaceae in the TDF from Sierra Gorda, Querétaro. One Discodon species was observed feeding on the nectar of Croton ciliatoglandulifer Ortega (Malphigiales: Euphorbiaceae) and C. fragilis in Tepalcingo, Morelos and Ixtlahuacán, Colima, respectively, as well as that of Acacia cochliacantha and other species of this genus, and also other unidentified shrubs either before or during the rainy season, depending on the species. It is of interest to mention that one Discodon was observed preying on a conspecific cantharid, while Silis sp. was found feeding on dipterans which were visiting the flowers of herbs. Compiled data on Polemius, Ditemnus, and Tytthonyx feeding preferences are scarce and cannot be treated with certainty. Daily and Seasonal Activity Cycles Most records describing adult habits point toward diurnal activity (e.g., Chauliognathus), with few records of nocturnal or crepuscular behavior (e.g., Discodon). However, we do not yet know which percentage of species is nocturnal, mainly because most studies have been more focused on those which are diurnal. Nocturnal species are usually collected using light traps, which makes it impossible to observe their natural behavior. In Mexican TDF, greater activity of adult cantharids was observed between 1100 and 1600 hours, their usual activities in that period are search for food resources or mates (Fig. 4). At ~1700 hours, cantharids cease activity and either return to the position they had occupied that morning or remain on inflorescence peduncles; the same pattern has been documented in South American TDF (Cerana 2004). Although Belotus has been observed in copula at 1400 hours, the mating of diurnal species such as Chauliognathus usually occurs at the end of the day between 1600 and 1800 hours, just as in the Northern Hemisphere (McLain 1984). Fig. 4. View largeDownload slide (a) Daily activity of Cantharidae species in the TDF of Tepalcingo, Morelos; (b) asynchronous emergence of Chauliognathus species at San Buenaventura TDF, Jalisco, during 1997; (c) asynchronous emergence of Cantharidae genera at Santiago Dominguillo TDF, Oaxaca, during 1998. Fig. 4. View largeDownload slide (a) Daily activity of Cantharidae species in the TDF of Tepalcingo, Morelos; (b) asynchronous emergence of Chauliognathus species at San Buenaventura TDF, Jalisco, during 1997; (c) asynchronous emergence of Cantharidae genera at Santiago Dominguillo TDF, Oaxaca, during 1998. Data from studied TDF localities show diverse patterns of daily adult activity among genera and species (Table 3, Fig. 4), mainly associated with a 24-h variation in temperature. For example, Belotus, Chauliognathus, and Ichthyurus species usually start their activities after 1000 hours at temperature of ~20°C, possibly pausing at midday if temperature reaches 30°C, perching in the receptacle of their host flowers or under the leaves of herbs. Some Discodon become active after 1400 hours, but most of them are crepuscular or nocturnal and initiate their activities at 1700 hours, just as Polemius do. Silis are active at midday, at temperature of up to 24°C. As previously mentioned, mating in diurnal Cantharidae from TDF commonly occurs at the end of the day. Also, copula has usually been observed during the last part of the emergence period. Table 3. Daily activity and seasonal periods of Cantharidae (Coleoptera) genera from Mexican TDF Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. View Large Table 3. Daily activity and seasonal periods of Cantharidae (Coleoptera) genera from Mexican TDF Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. View Large In Mexican TDF, seasonal cantharid activity patterns are more diverse than daily cycles (Fig. 4). This great diversity of seasonal patterns has even been observed within the same assemblage, producing high temporal turnover in species composition (Pérez-Hernández and Zaragoza-Caballero 2016; Fig. 4b). For example, adults of several genera and species can be found active during the entire year, especially those associated to predaceous habits (e.g., Silis). Nevertheless, seasonal patterns in soldier beetles are predominant and adult emergence is generally associated with seasonal changes in spring or summer (Miskimen 1972, Gambardella and Vaio 1978, Pérez-Hernández and Zaragoza-Caballero 2016) (Figs 4 and 5). Different phenomena seem to be involved in those temporal patterns: variation in precipitation, asynchronous emergence, and trophic guild. Fig. 5. View largeDownload slide Total abundance of Cantharidae genera collected in different years at seven localities from Mexican TDF. Months of dry season are usually: November to April; and those of rainy season: May to October. Fig. 5. View largeDownload slide Total abundance of Cantharidae genera collected in different years at seven localities from Mexican TDF. Months of dry season are usually: November to April; and those of rainy season: May to October. The emergence of particular genera is clearly associated with precipitation: Silis, Ditemnus, and Polemius adults (Silinae) emerge during the first week of summer, from the beginning to the first half of the rainy season, while Chauliognathus, Discodon, and Ichthyurus adults are predominant during the entire rainy season (usually from May to October), being more abundant halfway through the season (Pérez-Hernández and Zaragoza-Caballero 2016) (Fig. 5). The asynchronous emergence of genera and congeneric species in the same place is common (Fig. 4b). For example, in Tepalcingo, Morelos, Chauliognathus nigriceps emerges before other congeneric species in synchrony with Croton ciliatoglandulifer Ortega flowering, while C. forreri takes advantage of an anticipated emergence associated to the flowering of Serjania triquetra Radlk. (Sapindaceae) growing on Acacia cochliacantha, which are among the first flowering plants of the season. It is suggested that this phenomenon could be a phenological strategy to achieve greater protection for the less abundant species (Machado and Araújo 2001, Pérez-Hernández and Zaragoza-Caballero 2016). It has also been observed that the emergence period for the adults of several Cantharidae species and the maturity of their host plants are practically synchronized. In Colombian TDF, Chauliognathus proteus Gorham (Coleoptera: Cantharidae) and congeneric species were observed to have emerged almost in coincidence with the blossoming of their host plants, and were also seen to be progressively synchronized from higher to lower altitudes (Miskimen 1972). In Mexican TDF, the main portion of cantharid emergence usually coincides with the start of the rainy season and with the flowering of herbaceous plants and vines. Synchronization between adult emergence and the entire flowering period is not always very strict because several cantharids simply move to other flowering plants when their host plants show signs of desiccation (Machado and Araújo 2001). In other cases, cantharid populations conclude their emergence period even though their host plants are still flowering (Miskimen 1972). It is relevant to mention that, even though soldier beetles are not considered ideal for pollen dispersion, they undoubtedly play that role for specialized plants. For example, many collected individuals (e.g., Belotus, Chauliognathus, and Discodon) carried little to abundant quantities of pollen from their host plants, likely feeding on nutritious floral tissues as a reward. This warrants further investigation. Discussion In general, adults of the family have been characterized as common inhabitants of either the herbaceous stratum (Miskimen 1972, Borror et al. 1989, Pelletier and Hébert 2014) or the canopy stratum (Leksono et al. 2005, Hawkeswood and Turner 2008, Pelletier and Hébert 2014). In Mexican cloud forest, cantharids are mainly linked to the shrub layer, and to a lower degree, to the canopy or the ground layer (Deloya and Ordóñez-Reséndiz 2008). In this study, a slightly different pattern has been observed in Mexican TDF, where most genera and species are mainly associated to shrub and herbaceous layers, as well as vines (Tables 1 and 2). Patterns of the vertical distribution of Cantharidae seem to depend on the vegetation type; for example, cantharid species from eastern Canada and the United States are mostly associated with the canopy of temperate forest, while in deciduous hardwood forest, a strong association of soldier beetles with either the canopy or the ground layer has been identified (Hardersen et al. 2014, Pelletier and Hébert 2014). Therefore, it is possible that cantharid species have stratum specialization. If this is correct, the restriction of cantharids to one specific stratum may mark them as a high conservation priority (Hammond et al. 1997). Up to this work, phenological patterns of Cantharidae had not been very deeply explored. Most information concerning seasonal rhythms has been obtained from the labels of specimens from collections (Diéguez et al. 2006, Pelletier and Hébert 2014), and to a lesser degree through inventory or systematic collection (Zaragoza-Caballero et al. 2010, Pérez Hernández and Zaragoza-Caballero 2016). The study of temporal patterns in the diversity of Cantharidae, as well as factors which determine them, is an issue with high potential, especially if the synchronization of the patterns of nectarivorous and pollinivorous cantharids with the phenology of host plants is included. Cantharids are little-specialized insects, as mentioned above, but several species associated to flower plants are able to perform relevant functions such as pollination of native plants or biological control. For example, although they are considered secondary pollinators (Willmer 2011), it would be relevant to evaluate the importance of groups such as this in the current scenario of the global pollination crisis of primary pollinators. This particular issue is currently being analyzed by the author in an ongoing manuscript. Because of their relationship with plants and the frequency with which they inhabit croplands, cantharid populations also suffer because of the effects of the use of pesticides and insecticides (Alexander 2003, Bruck et al. 2006); this is another scarcely explored but evidently relevant issue. In this study, it was provided a list of host plants for TDF Cantharidae genera and species for the first time, and it was also observed a high ecological diversity in the phenology and behavior of TDF Cantharidae assemblages. Nevertheless, further research concerning cantharids and other TDF insects needs to have a more comprehensive and integrated approach, in such a way that it can be applied toward understanding patterns, both current and historical, spatial and temporal, of distribution and diversity, and at species and supraspecific levels, also aimed at identifying the factors which determined them and elucidating the role that cantharids play in ecosystems. TDFs are the second largest type of tropical forests, and encompass nearly 42% of tropical ecosystems around the globe (Miles et al. 2006). Unfortunately, it is also one of the most threatened in the world due mainly to the loss of more than 60% of its original distribution area, as well as to the fragmentation and the land use change that dramatically affects the TDF remaining areas and therefore species diversity and their ecological interactions, among other processes (Miles et al. 2006, Dirzo et al. 2011, Quesada et al. 2011).However, the great biodiversity harbored by TDF is usually studied through inventories and, to a lesser degree, through the analysis of its determining patterns and processes. Hence, study of the natural history and ecology of species inhabiting TDF is one of the main steps toward the comprehension of how different phenomena and ecosystems work, and consequently toward the design of better conservation strategies in order to preserve the ecosystem functioning in the TDF. Acknowledgments The author express their special thanks to S. Zaragoza-Caballero for his comments about the behavior of soldier beetles and interactions; to José Luis Villaseñor and Gabriel Flores Franco for the identification of host plants from Tepalcingo, Morelos; and to Sonia Helen Ponce Wainer for the revision of the English version of the manuscript. She also expresses her special thanks to the Programa de Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México (UNAM) and the Consejo Nacional de Ciencia y Tecnología (CONACyT) for supporting the doctoral project to which this manuscript pertains. References Cited Alexander , K. N. A . 2003 . Changing distributions of Cantharidae and Buprestidae within Great Britain (Coleoptera) , pp. 87 – 91 . In Proceedings, 13th International Colloquium European Invertebrate Survey , 2–5 September 2001 , Leiden, The Netherlands . Bilde , T. , J. A. Axelsen , and S. Toft . 2000 . The value of Collembola from agricultural soils as food for a generalist predator . J. Appl. 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A. Noguera , E. González-Soriano , E. Ramírez-García , and A. Rodríguez-Palafox . 2010 . Insectos del bosque tropical caducifolio del oeste de México , pp. 195 – 214 . In G. Ceballos , J. Bezaury and E. Espinoza-Medinilla (eds.), Diversidad, amenazas y regiones prioritarias para la conservación de las selvas secas del Pacífico de México . CONABIO, WWF, Universidad Nacional Autónoma de México , México . © The Author(s) 2018. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com. This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/about_us/legal/notices) http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Environmental Entomology Oxford University Press

Natural History and Ecology of Soldier Beetles (Coleoptera: Cantharidae) in the Mexican Tropical Dry Forests

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Abstract

Abstract Until today, most information about the natural history and ecology of soldier beetles came from temperate zones, mainly from Holarctic areas, while tropical regions have been poorly studied. The aim of this contribution is to compile and synthesize information concerning the natural history and ecology of Cantharidae (Coleoptera) from the Mexican tropical dry forest (TDF), to serve as a starting point for more in-depth study of the group in one of the Mexico’s most endangered ecosystems. All compiled data on the family have been organized into the following topics: distributional patterns and habitat preferences, feeding behavior and host plants, and daily and seasonal activity cycles. For the first time, it was provided a list of host plants for TDF Cantharidae genera and species, and it was also observed a high ecological diversity in the phenology and behavior of TDF Cantharidae assemblages. Further research concerning cantharids and other TDF insects needs to have a more comprehensive and integrated approach toward understanding the patterns of distribution and diversity, and elucidating the role that cantharids play in ecosystems, especially in TDF, which is one of the most endangered ecosystem in the world. plant, insect, interaction, phenology, behavior Cantharids, or soldier beetles (Coleoptera: Cantharidae), are the most diverse group of soft-bodied terrestrial coleopterans (Ramsdale 2002). Cantharidae are usually classified into five subfamilies: Cantharinae, Chauliognathinae, Malthininae, Silinae, and Dysmorphocerinae, following the classification of Brancucci (1980). On the basis of molecular phylogenetic analyses, it has been proposed that, in fact, there should be only four subfamilies (excluding Dysmorphocerinae) (Sagegami-Oba et al. 2007). However, cantharidologists continue using Brancucci’s scheme (Constantin and Chaboo 2016) and an open discussion concerning the phylogeny of Cantharidae still remains. Soldier beetles are strictly terrestrial during their entire life cycle, and are distributed from ecosystems as cold as the Arctic tundra to the warm, dry African grasslands, ranging from the tropical dry forests (TDFs) of the American continent to the tropical rain forest of India. It is possible to find them in forests, meadows, savannahs, and grasslands, almost always among the foliage and inflorescences of their host plants (Ramsdale 2010, Pelletier and Hérbert 2014). Many species are associated with vegetation close to rivers, creeks, and other water bodies (Zaragoza-Caballero 2004, Ramsdale 2010). Members of this family are frequently classified as generalists in their feeding habits and habitats. However, the distribution of most cantharids is much more restricted than it was thought, since even within the same ecosystem each species may occupy a particular habitat, just as Fender described in his review of the North American Malthinini: ‘Each species appears to have a somewhat selective instinct for its preferred habitat’ (Fender 1951). This tendency of habitat specialization is noticeable up to the highest taxonomical hierarchies, although there are no studies concerning this issue. For example, Silini are evidently most diverse in tropical ecosystems, there are 119 Silinae species registered from Nearctic region, whereas in the Neotropical region are 819 (Delkeskamp 1977, Pérez-Hernández 2017); Chauliognathini prefer open areas, savannahs, and meadows within tropical ecosystems (Miskimen 1966), and Ichthyurini are common in tropical deciduous forests (Fender 1960). In contrast, Malthinini and Malthodini are more frequently associated with temperate coniferous forests, particularly on the foliage of coniferous and deciduous trees in shaded and humid environments such as marsh (Fender 1951, Ramsdale 2002). It has been hypothesized that Cantharidae habitat selection in tropical zones, and therefore distributional patterns, strongly depends on the distribution of adult host plants (Miskimen 1966, 1972). Nevertheless, as many species have predatory feeding habits and show an indirect relationship with plant species, another hypothesis is that specificity to particular environments is in fact determined by the abiotic requirements of the larvae (Ramsdale 2002). Unfortunately, there is a lack of information concerning the distributional patterns of Cantharidae and their specificity to certain habitats or their biotic interactions. This novel issue is in need of further study. Until now, there has been a paucity of information as to the natural history and ecology of soldier beetles from tropical regions. The aim of this contribution is to compile and synthesize the most relevant information regarding the natural history and ecology of Cantharidae (Coleoptera) from Mexican TDFs, to serve as a starting point for more in-depth study of the group in one of the most endangered ecosystems in the World (Miles et al. 2006). All compiled data on the family has been organized into the following topics: distributional patterns and habitat preferences, feeding behavior and host plants, and daily and seasonal activity cycles. Most data come from entomological samplings carried out in eight localities on the Mexican Pacific region. Materials and Methods Cantharidae specimens were studied in different years from eight Mexican localities, where TDF is the dominant ecosystem. From north to south, these are San Javier, Sonora; San Buenaventura, Jalisco; Ixtlahuacán, Colima; Sierra de Huautla, Morelos (Tepalcingo and Tlaquiltenango municipalities); Santiago Dominguillo, Oaxaca; Acahuizotla, Guerrero; and Huatulco, Oaxaca (Fig. 1). Light attraction, Malaise traps (except Acahuizotla), aerial nets, and beating of vegetation were all used for entomological samplings 5 d a month in each locality, always coinciding with the new moon. Specific details concerning sampling methods and study areas may be consulted in the studies by González-Ramírez et al. (2017), Noguera et al. (2007, 2012), Pérez-Hernández and Zaragoza-Caballero (2015), Rodríguez-Vélez et al. (2009), and Zaragoza-Caballero et al. (2003, 2009). In pursuance of the observation and documentation of the natural history and ecology of cantharids, Tepalcingo was visited in different months of 2014 and 2015, thus providing most of the information included in this study. Fig. 1. View largeDownload slide Map depicting distribution of TDF ecosystem in Mexico (map modified from Conabio, 1999). Localities and years of Cantharidae (Coleoptera) sampling for this contribution are showed. Fig. 1. View largeDownload slide Map depicting distribution of TDF ecosystem in Mexico (map modified from Conabio, 1999). Localities and years of Cantharidae (Coleoptera) sampling for this contribution are showed. Results Distributional Patterns and Habitat Preferences In Mexico, representatives of all Cantharidae subfamilies have been classified into 19 genera and 223 species (Zaragoza-Caballero and Pérez-Hernández 2014). More than 70 species registered in Mexican TDF belong to the genera Belotus Gorham (Coleoptera: Cantharidae), Chauliognathus Hentz (Coleoptera: Cantharidae), Ichthyurus Westwood (Coleoptera: Cantharidae) (Chauliognathinae), Plectonotum Gorham (Coleoptera: Cantharidae), Caccodes Sharp in Blackburn & Sharp (Coleoptera: Cantharidae) (Malthininae), Discodon Gorham (Coleoptera: Cantharidae), Ditemnus LeConte (Coleoptera: Cantharidae), Polemius LeConte (Coleoptera: Cantharidae), Silis Charpentier (Coleoptera: Cantharidae), and Tytthonyx LeConte (Coleoptera: Cantharidae) (Silinae) (Pérez-Hernández and Zaragoza-Caballero 2015). None of these genera or supraspecific taxa are restricted to TDF, as in other regions of the World (Hanson 2011); in fact, cantharids were observed in a high diversity of habitats from studied localities (Table 1), and with slight differences, congeneric species frequently inhabited similar environments. Compiled information describing the habitat preferences of TDF cantharid genera is summarized in the following paragraphs. Table 1. Statewide distribution, habitats, and observed trophic guilds of Cantharidae (Coleoptera) genera from TDF in Mexico Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa aSuggested trophic guild. View Large Table 1. Statewide distribution, habitats, and observed trophic guilds of Cantharidae (Coleoptera) genera from TDF in Mexico Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa Genus Statewise distribution Main habitat and stratum Trophic guild Chauliognathinae  Belotus Campeche, Chiapas, Chihuahua, Colima, Distrito Federal, Durango, Hidalgo, Jalisco, Morelos, Michoacán, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Tabasco, Tamaulipas, Veracruz Riparian vegetation from tropical forests; herbaceous stratum Nectarivorous  Chauliognathus Almost entire country, except Baja California, Baja California Sur, Yucatán, or Zacatecas Open fields from tropical forests. Vines, herbaceous and shrub layers Nectarivorous, pollinivorous  Ichthyurus Chiapas, Colima, Oaxaca, Veracruz Canopy of shrubs at tropical forests Omnivorous Malthininae  Caccodes Chiapas, Chihuahua, Colima, Durango, Hidalgo, Oaxaca, Sinaloa, Sonora, Veracruz Foliage of shrubs and canopy from tropical forest Nectarivorousa  Plectonotum Guerrero Unknown Nectarivorousa Silinae  Discodon Almost entire country, except Chihuahua, Sinaloa, Zacatecas, Tamaulipas, Tlaxcala, or Baja California and Yucatán peninsulas. Foliage of shrubs and canopy from tropical forest Pollinivorous, nectarivorous, predator  Ditemnus Colima, Jalisco, Morelos, Oaxaca, Sonora Herb foliage at open fields, near water bodies Predator  Polemius Neotropical region and Durango Foliage of shrubs from tropical forest Nectarivorousa  Silis Entire country Herbaceous near water bodies from tropical forests Predator, nectarivorous  Tytthonyx Baja California Sur, Chiapas, Durango, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Oaxaca, Querétaro, Sinaloa, Veracruz Canopy and herbaceous layers from tropical forests Nectarivorousa aSuggested trophic guild. View Large In Mexican TDF, Malthininae species belong to Caccodes and Plectonotum. Very little is known about Caccodes habitat preferences, except that they can be collected by beating herbaceous and shrub vegetation, as well as through light and Malaise traps; members of the genus are presumably associated to closed habitats near creeks and rivers. Members of Plectonotum have been previously registered in South American rain forests, mountain forests, and paramos, as well as in Ecuador in zones ‘covered with highly diverse vegetation’ (Constantin 2008); consequently, the sole species registered in Mexico from TDF indicates more diverse habitat preferences in the genus, along with a larger geographical distribution in America (Pérez-Hernández and Zaragoza-Caballero 2015). Silinae members distributed in Mexican TDF belong to the Discodon, Ditemnus, Silis, Polemius, and Tytthonyx genera. In North America, Silis mostly inhabit the canopies of hardwood and mixed forests (Pelletier and Hébert 2014), and in Mexico, it is commonly found in the herbaceous stratum of tropical forests. In Mexican TDF, species of this genus are associated to shading or open habitats adjacent to water bodies, which usually present higher humidity than other habitats for most of the year (Fig. 2b). Ditemnus species have similar preferences to those of Silis members, also found more frequently in herbs. Discodon and Polemius species are associated to shady habitats and the canopy of shrubs, and they can usually be found in plants near water bodies or the forest edge (Fig. 2a, c, and d); they also spend most of the d perching on the underside of shrub leaves, such as those of Acacia cochliacantha Humb. & Bonp. ex Willd. (Fabales: Fabacaeae), and other bushes in shaded sites (Fig. 3). Members of Tytthonyx can be remarkably abundant in herbs and short shrubs. They are good flyers and can thus be collected using Malaise traps, which is usually an ineffective method for Cantharidae sampling. Fig. 2. View largeDownload slide Examples of habitats of Discodon (a, c, d), Silis (a), Polemius (c), and Chauliognathus (b) in Mexican TDF. Fig. 2. View largeDownload slide Examples of habitats of Discodon (a, c, d), Silis (a), Polemius (c), and Chauliognathus (b) in Mexican TDF. Fig. 3. View largeDownload slide (a, b) Discodon sp. preying on a conspecific individual; (c) a different species of Discodon, feeding on pollen; (d) Silis female eating a dipteran; (e, h) Chauliognathus forreri feeding on nectar and pollen; (f) C. nigriceps female feeding on nectar (Photo Enrique Ramírez); (g) Discodon perching on the underside of the leaves. Fig. 3. View largeDownload slide (a, b) Discodon sp. preying on a conspecific individual; (c) a different species of Discodon, feeding on pollen; (d) Silis female eating a dipteran; (e, h) Chauliognathus forreri feeding on nectar and pollen; (f) C. nigriceps female feeding on nectar (Photo Enrique Ramírez); (g) Discodon perching on the underside of the leaves. Members of Chauliognathus dwell in a wide range of environmental conditions and habitats, from temperate to tropical regions and from deserts to rainforests (Miskimen 1972). However, they are typically associated with herbs or shrubs in open, sunny areas (Fig. 2b). Belotus species are also associated to herbs or minute flowers, and frequent open fields adjacent to water bodies. Ichthyurus is a genus with a mainly tropical affinity, usually found in the canopy of shrubs. At the species level, one can observe a wide spectrum of habitat preferences, even among congeneric species. For example, members of Chauliognathus emerge with similar seasonality in TDF, and slight differences in habitat preference and seasonality may serve to avoid a more competitive coexistence. In this way, Chauliognathus forreri Gorham (Coleoptera: Cantharidae) has been exclusively observed visiting vines of Serjania triquetra Radlk. (Sapindales: Sapindaceae) at the beginning of the rainy season; halfway through that season, Chauliognathus dispar Champion (Coleoptera: Cantharidae), Chauliognathus nigrocinctus Gorham (Coleoptera: Cantharidae), and Chauliognathus profundus LeConte (Coleoptera: Cantharidae)—as well as C. forreri (Coleoptera: Cantharidae)—can be found on a variety of plants, despite being highly associated to flowers of Asteraceae, Apiaceae, and other herbs (Fig. 3; Table 2). It has also been recorded that Eupatorium L. (Asterales: Asteraceae) plants are known hosts of C. dispar in the subtropical dry forest (Miskimen 1966). Table 2. Association between some plant and Cantharidae species from studied Mexican TDFs localities Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey I, Ixtlahuacán, Colima; T, Tepalcingo, Morelos; Sierra de Huautla, Morelos. View Large Table 2. Association between some plant and Cantharidae species from studied Mexican TDFs localities Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey Cantharidae species Host plant species Interaction Chauliognathinae  Belotus bicolor Croton fragilis Kunth (Euphorbiaceae)I Eating nectar Unknown ApicaeaeT Eating nectar  Chauliognathus dispar Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar Crusea sp. (Rubiaceae)T Eating nectar Gomphrena serrata L. (Amaranthaceae)T Eating nectar Lagascea rigida (Cav.) Stuessy (Asteraceae)T Eating nectar Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Salvia aff. rhyacophila (Fernald) (Lamiaceae)T Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus forreri Bidens odorata Cav. var. odorata (Asteraceae)T Feeding on petals and nectar Croton sp. (Euphorbiaceae)H Eating nectar and pollen (?) Cynanchum foetidum (Cav.) Kunth (Apocynaceae)T Eating nectar and pollen Dalea foliolosa Barneby (Fabaceae)T Eating nectar Dicliptera peduncularis Knees (Acanthaceae)T Eating nectar Euphorbia graminea Jacq. (Euphorbiacaeae)T Eating nectar Hamelia patens Jacq. (Rubiaceae)T Eating nectar Nissolia fruticosa Jacq. (Fabaceae)T Eating nectar and pollen Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Serjania triquetra Radlk. (Sapindaceae)T Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Trichilia hirta L. (Meliaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar and petals  Chauliognathus nigriceps Croton ciliatoglandulifer Ortega (Euphorbiaceae)T, H Eating nectar  Chauliognathus nigrocinctus Bidens odorata var. odorata Cav. (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar  Chauliognathus profundus Bidens odorata Cav. var. odorata (Asteraceae)T Eating nectar and pollen Melampodium divaricatum (Rich.) DC. (Asteraceae)T Eating nectar and pollen Sanvitalia procumbens Lam., 1972 (Asteraceae)T Eating nectar and pollen Spermacoce ocymoides Burm. f. (Rubiaceae)T Eating nectar Tridax coronopifolia (Kunth) Hemsl. (Asteraceae)T Eating nectar Silinae  Discodon Acacia cochliacantha Humb. & Bonpl. ex Willd. (Fabaceae)T Eating pollen (?) Bunchosia canescens (Aiton) DC. (Malpighiaceae)T Eating pollen Croton ciliatoglandulifer Ortega (Euphorbiaceae)T Nectarivorous Croton fragilis Kunth (Euphorbiaceae)I, T Nectarivorous Hamelia patens Jacq. (Rubiaceae)T Perching on leaves Trichilia hirta L. (Meliaceae)T Eating pollen (?)  Polemius Commelina sp. (Commelinaceae) Perching on leaves  Silis Senna uniflora (Mill.) H.S. Irwin & Barneby (Fabaceae)T Searching for prey Senna hirsuta (L.) H.S. Irwin & Barneby (Fabaceae)T Perching on leaves; searching for prey I, Ixtlahuacán, Colima; T, Tepalcingo, Morelos; Sierra de Huautla, Morelos. View Large Feeding Behavior and Host Plants Studies concerning the feeding behavior of Cantharidae larvae from tropical regions are very scarce (Jiron and Hedström 1985). However, available data depicting this stage indicate that generalist behavior is more frequent than specialization (Traugott 2003). In temperate regions, soldier beetle larvae are commonly registered as predominantly voracious predators whose survival critically depends on the availability of high-quality prey (Gambardella and Vaio 1978, Bilde et al. 2000). They generally feed on eggs, adult insects and other larvae, gastropods and earthworms (Ramsdale 2002, Eitzinger and Traugott 2011), whereas other species are facultatively or predominantly phytophagous, such as those of Silis (Ramsdale 2002). It is highly probable for soldier beetle larvae from TDF to show the same behavior. Most publications concerning the family affirm that most species are omnivorous and predominantly predators (Sunderland et al. 1987, Leksono et al. 2005, Jerinic-Prodanovic et al. 2010). Nevertheless, very few genera seem to present this pattern, such as Rhagonycha Eschscholtz (Coleoptera: Cantharidae), which visits the foliage and inflorescences of a great variety of plants where it feeds equally upon insects and nectar (Ramsdale 2002, Day et al. 2006). Ichthyurus are also mainly predators, and phytophagous to a lesser degree (Miskimen 1961). Members of Podabrini and Cantharidini from temperate regions have frequently been associated to the consumption of aphids (Aphididae) (Ramsdale 2010). In TDF, adults of Belotus, Chauliognathus, and Discodon were commonly observed feeding on nectar and/or pollen, whereas Discodon and Silis were observed feeding on insects (Fig. 3a and d). Chauliognathus adults have developed buccal structures adapted to feeding on nectar and pollen, mainly from plants with small, abundant flowers (Miskimen 1972, Ramsdale 2002), occasionally predate larvae of lepidopterans and adults of other beetles (Fender 1962) (Table 2). In TDF, Chauliognathus nigriceps Gorham (Coleoptera: Cantharidae) commonly feeds on Croton spp. (Malphigiales: Euphorbiaceae), whose flowering time begins a few weeks before the start of the rainy season; C. dispar, C. nigrocinctus, and C. profundus have been observed actively feeding on the nectar and pollen of Bidens odorata var. odorata Cav. (Asterales: Asteraceae), Melampodium divaricatum (Rich.) DC. (Asterales: Asteraceae), Tridax coronopifolia (Kunth) Hemsl. (Asterales: Asteraceae), Sanvitalia procumbens Lam. (Asterales: Asteraceae) and other herbs (Table 2). C. forreri has been observed visiting a wide spectrum of plant species (Table 2) and also robbing nectar by chewing a hole in the base of the Hamelia patens Jacq. (Gentianales: Rubiaceae) flowers to avoid conspecific competitors. Belotus present feeding behavior similar to that of Chauliognathus species, although consuming different plant species. For example, they have been collected feeding on Croton fragilis Kunth (Malphigiales: Euphorbiaceae) in Ixtlahuacán, Colima and on unidentified Apiaceae in the TDF from Sierra Gorda, Querétaro. One Discodon species was observed feeding on the nectar of Croton ciliatoglandulifer Ortega (Malphigiales: Euphorbiaceae) and C. fragilis in Tepalcingo, Morelos and Ixtlahuacán, Colima, respectively, as well as that of Acacia cochliacantha and other species of this genus, and also other unidentified shrubs either before or during the rainy season, depending on the species. It is of interest to mention that one Discodon was observed preying on a conspecific cantharid, while Silis sp. was found feeding on dipterans which were visiting the flowers of herbs. Compiled data on Polemius, Ditemnus, and Tytthonyx feeding preferences are scarce and cannot be treated with certainty. Daily and Seasonal Activity Cycles Most records describing adult habits point toward diurnal activity (e.g., Chauliognathus), with few records of nocturnal or crepuscular behavior (e.g., Discodon). However, we do not yet know which percentage of species is nocturnal, mainly because most studies have been more focused on those which are diurnal. Nocturnal species are usually collected using light traps, which makes it impossible to observe their natural behavior. In Mexican TDF, greater activity of adult cantharids was observed between 1100 and 1600 hours, their usual activities in that period are search for food resources or mates (Fig. 4). At ~1700 hours, cantharids cease activity and either return to the position they had occupied that morning or remain on inflorescence peduncles; the same pattern has been documented in South American TDF (Cerana 2004). Although Belotus has been observed in copula at 1400 hours, the mating of diurnal species such as Chauliognathus usually occurs at the end of the day between 1600 and 1800 hours, just as in the Northern Hemisphere (McLain 1984). Fig. 4. View largeDownload slide (a) Daily activity of Cantharidae species in the TDF of Tepalcingo, Morelos; (b) asynchronous emergence of Chauliognathus species at San Buenaventura TDF, Jalisco, during 1997; (c) asynchronous emergence of Cantharidae genera at Santiago Dominguillo TDF, Oaxaca, during 1998. Fig. 4. View largeDownload slide (a) Daily activity of Cantharidae species in the TDF of Tepalcingo, Morelos; (b) asynchronous emergence of Chauliognathus species at San Buenaventura TDF, Jalisco, during 1997; (c) asynchronous emergence of Cantharidae genera at Santiago Dominguillo TDF, Oaxaca, during 1998. Data from studied TDF localities show diverse patterns of daily adult activity among genera and species (Table 3, Fig. 4), mainly associated with a 24-h variation in temperature. For example, Belotus, Chauliognathus, and Ichthyurus species usually start their activities after 1000 hours at temperature of ~20°C, possibly pausing at midday if temperature reaches 30°C, perching in the receptacle of their host flowers or under the leaves of herbs. Some Discodon become active after 1400 hours, but most of them are crepuscular or nocturnal and initiate their activities at 1700 hours, just as Polemius do. Silis are active at midday, at temperature of up to 24°C. As previously mentioned, mating in diurnal Cantharidae from TDF commonly occurs at the end of the day. Also, copula has usually been observed during the last part of the emergence period. Table 3. Daily activity and seasonal periods of Cantharidae (Coleoptera) genera from Mexican TDF Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. View Large Table 3. Daily activity and seasonal periods of Cantharidae (Coleoptera) genera from Mexican TDF Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. Genera/species Daily activity Seasonality Sampling method Chauliognathinae  Belotus Diurnal (10:00–15:00 h) Almost all the year at certain sites; more abundant in rainy season Light attraction, but mainly aerial net and beating vegetation  Chauliognathus Diurnal (10:00–17:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps  Ichthyurus Diurnal (10:00–16:00 h) Rainy season Mainly through aerial net, beating vegetation, to a lesser degree using light attraction and poorly collected by Malaise traps Malthininae  Caccodes Nocturnal Rainy season Aerial net, beating vegetation, and light attraction  Plectonotum Rainy season Aerial net, beating vegetation Silinae  Discodon Crepuscular, nocturnal (up to 17:00 h) Rainy season Aerial net, beating vegetation, to a lesser degree in Malaise traps; mainly attracted to light traps Ditemnus All year at certain sites; more abundant before rainy season Mainly beating vegetation; light attraction Polemius Crepuscular, nocturnal Rainy season Mainly beating vegetation; light attraction Silis Diurnal to crepuscular (10:00–18:00 h) All year at certain sites; more abundant in first months of rainy season Beating vegetation and light attraction Tytthonyx Diurnal All year in Tlaquiltenango; very rare at other three sites Mainly Malaise traps; sometimes, beating vegetation. Not attracted to lights. View Large In Mexican TDF, seasonal cantharid activity patterns are more diverse than daily cycles (Fig. 4). This great diversity of seasonal patterns has even been observed within the same assemblage, producing high temporal turnover in species composition (Pérez-Hernández and Zaragoza-Caballero 2016; Fig. 4b). For example, adults of several genera and species can be found active during the entire year, especially those associated to predaceous habits (e.g., Silis). Nevertheless, seasonal patterns in soldier beetles are predominant and adult emergence is generally associated with seasonal changes in spring or summer (Miskimen 1972, Gambardella and Vaio 1978, Pérez-Hernández and Zaragoza-Caballero 2016) (Figs 4 and 5). Different phenomena seem to be involved in those temporal patterns: variation in precipitation, asynchronous emergence, and trophic guild. Fig. 5. View largeDownload slide Total abundance of Cantharidae genera collected in different years at seven localities from Mexican TDF. Months of dry season are usually: November to April; and those of rainy season: May to October. Fig. 5. View largeDownload slide Total abundance of Cantharidae genera collected in different years at seven localities from Mexican TDF. Months of dry season are usually: November to April; and those of rainy season: May to October. The emergence of particular genera is clearly associated with precipitation: Silis, Ditemnus, and Polemius adults (Silinae) emerge during the first week of summer, from the beginning to the first half of the rainy season, while Chauliognathus, Discodon, and Ichthyurus adults are predominant during the entire rainy season (usually from May to October), being more abundant halfway through the season (Pérez-Hernández and Zaragoza-Caballero 2016) (Fig. 5). The asynchronous emergence of genera and congeneric species in the same place is common (Fig. 4b). For example, in Tepalcingo, Morelos, Chauliognathus nigriceps emerges before other congeneric species in synchrony with Croton ciliatoglandulifer Ortega flowering, while C. forreri takes advantage of an anticipated emergence associated to the flowering of Serjania triquetra Radlk. (Sapindaceae) growing on Acacia cochliacantha, which are among the first flowering plants of the season. It is suggested that this phenomenon could be a phenological strategy to achieve greater protection for the less abundant species (Machado and Araújo 2001, Pérez-Hernández and Zaragoza-Caballero 2016). It has also been observed that the emergence period for the adults of several Cantharidae species and the maturity of their host plants are practically synchronized. In Colombian TDF, Chauliognathus proteus Gorham (Coleoptera: Cantharidae) and congeneric species were observed to have emerged almost in coincidence with the blossoming of their host plants, and were also seen to be progressively synchronized from higher to lower altitudes (Miskimen 1972). In Mexican TDF, the main portion of cantharid emergence usually coincides with the start of the rainy season and with the flowering of herbaceous plants and vines. Synchronization between adult emergence and the entire flowering period is not always very strict because several cantharids simply move to other flowering plants when their host plants show signs of desiccation (Machado and Araújo 2001). In other cases, cantharid populations conclude their emergence period even though their host plants are still flowering (Miskimen 1972). It is relevant to mention that, even though soldier beetles are not considered ideal for pollen dispersion, they undoubtedly play that role for specialized plants. For example, many collected individuals (e.g., Belotus, Chauliognathus, and Discodon) carried little to abundant quantities of pollen from their host plants, likely feeding on nutritious floral tissues as a reward. This warrants further investigation. Discussion In general, adults of the family have been characterized as common inhabitants of either the herbaceous stratum (Miskimen 1972, Borror et al. 1989, Pelletier and Hébert 2014) or the canopy stratum (Leksono et al. 2005, Hawkeswood and Turner 2008, Pelletier and Hébert 2014). In Mexican cloud forest, cantharids are mainly linked to the shrub layer, and to a lower degree, to the canopy or the ground layer (Deloya and Ordóñez-Reséndiz 2008). In this study, a slightly different pattern has been observed in Mexican TDF, where most genera and species are mainly associated to shrub and herbaceous layers, as well as vines (Tables 1 and 2). Patterns of the vertical distribution of Cantharidae seem to depend on the vegetation type; for example, cantharid species from eastern Canada and the United States are mostly associated with the canopy of temperate forest, while in deciduous hardwood forest, a strong association of soldier beetles with either the canopy or the ground layer has been identified (Hardersen et al. 2014, Pelletier and Hébert 2014). Therefore, it is possible that cantharid species have stratum specialization. If this is correct, the restriction of cantharids to one specific stratum may mark them as a high conservation priority (Hammond et al. 1997). Up to this work, phenological patterns of Cantharidae had not been very deeply explored. Most information concerning seasonal rhythms has been obtained from the labels of specimens from collections (Diéguez et al. 2006, Pelletier and Hébert 2014), and to a lesser degree through inventory or systematic collection (Zaragoza-Caballero et al. 2010, Pérez Hernández and Zaragoza-Caballero 2016). The study of temporal patterns in the diversity of Cantharidae, as well as factors which determine them, is an issue with high potential, especially if the synchronization of the patterns of nectarivorous and pollinivorous cantharids with the phenology of host plants is included. Cantharids are little-specialized insects, as mentioned above, but several species associated to flower plants are able to perform relevant functions such as pollination of native plants or biological control. For example, although they are considered secondary pollinators (Willmer 2011), it would be relevant to evaluate the importance of groups such as this in the current scenario of the global pollination crisis of primary pollinators. This particular issue is currently being analyzed by the author in an ongoing manuscript. Because of their relationship with plants and the frequency with which they inhabit croplands, cantharid populations also suffer because of the effects of the use of pesticides and insecticides (Alexander 2003, Bruck et al. 2006); this is another scarcely explored but evidently relevant issue. In this study, it was provided a list of host plants for TDF Cantharidae genera and species for the first time, and it was also observed a high ecological diversity in the phenology and behavior of TDF Cantharidae assemblages. Nevertheless, further research concerning cantharids and other TDF insects needs to have a more comprehensive and integrated approach, in such a way that it can be applied toward understanding patterns, both current and historical, spatial and temporal, of distribution and diversity, and at species and supraspecific levels, also aimed at identifying the factors which determined them and elucidating the role that cantharids play in ecosystems. TDFs are the second largest type of tropical forests, and encompass nearly 42% of tropical ecosystems around the globe (Miles et al. 2006). Unfortunately, it is also one of the most threatened in the world due mainly to the loss of more than 60% of its original distribution area, as well as to the fragmentation and the land use change that dramatically affects the TDF remaining areas and therefore species diversity and their ecological interactions, among other processes (Miles et al. 2006, Dirzo et al. 2011, Quesada et al. 2011).However, the great biodiversity harbored by TDF is usually studied through inventories and, to a lesser degree, through the analysis of its determining patterns and processes. Hence, study of the natural history and ecology of species inhabiting TDF is one of the main steps toward the comprehension of how different phenomena and ecosystems work, and consequently toward the design of better conservation strategies in order to preserve the ecosystem functioning in the TDF. Acknowledgments The author express their special thanks to S. 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Published: Feb 27, 2018

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