EFFECT OF NECTAR-FORAGING ANTS AND WASPS ON THE REPRODUCTIVE FITNESS OF TURNERA ULMIFOLIA (TURNERACEAE) IN A COASTAL SAND DUNE IN MEXICO

We evaluated over two years the effect of different ant species on the reproductive fitness (estimated as end-of-season fruit set per treatment) of Turnera ulmifolia. Research was done on a sand dune matorral located on the coast of Veracruz, México. The results show that (i) plants associated with the larger ant species (Camponotus abdominalis) produced more fruits than plants associated with the smaller ant species, and (ii) plants associated with ants were subject to lower levels of herbivory, than plants without ants. Consequently, ant presence is not synonymous of plant protection, and the level of protection by ants will depend on the size of the worker ants in a guild of ant visitors. Moreover, when ants were excluded, wasps and bees increased their visits, exerting a higher level of protection than that offered by the smaller ant species. However, the differential effect of wasps needs further attention.Evaluamos durante dos años el efecto de diferentes especies de hormigas sobre el éxito reproductivo (estimado como número total de frutos por temporada) de Turnera ulmifolia. El estudio se llevó a cabo en el matorral de duna costera en la costa central del estado de Veracruz, México. Los resultados muestran que (i) plantas asociadas con la especie de hormiga más grande (Camponotus abdominalis) produjeron más frutos que plantas asociadas con las especies de hormiga más pequeñas, y (ii) plantas asociadas con las hormigas sufrieron niveles de herbivoría menores, que plantas sin hormigas. Consequentemente, la presencia de hormigas no es sinónimo de protección para una planta, y el nivel de protección por hormigas dependerá del tamaño de las hormigas obreras en un gremio de hormigas visitantes. Más aún, cuando se excluyeron a las hormigas, aumentaron las visitas de avispas y abejas, las que efectuaron un nivel de protección mayor que aquel ofrecido por las especies pequeñas de hormigas. Sin embargo, el efecto diferencial de las avispas requiere atención futura

EFECTO DE UNA FUENTE DE ALIMENTO EXPERIMENTAL SOBRE UNA ASOCIACIÓN HORMIGA-HEMIPTERO

In order to assess if ants attracted to honeydew-producing Hemiptera switched food sources when offered an additional, considerably rich and abundant food source, we studied the interaction between the plant Solanum lycocarpum (Solanaceae), an efn-lacking shrub, the predominantly diurnal ant Camponotus crassus (Hymenoptera: Formicidae), and two species of membracids (Enchenopa spp.; Homoptera: Membracidae). The study was done during September of 1999 at the Reserva Ecológica of IBGE (ca. 35 km S of Brasilia-DF, Brasil). The results show that ants were attracted both to the experimental food source and to the membracids. Moreover, the mean number of ants visiting the experimental nectary was significantly higher than those visiting the membracids. However, the membracids were never left unattended, which suggests that ants, even though presented with an abundant and rich food source, do not abandon other sources. The latter can be explained due to an increment in the honeydew production rate by the membracids, thus the attention levels of ants does not vary Another possibility is the fidelity of worker ants to a food source and its location. To show that ants switch food sources based on quality and/or quantity, and not merely based of their presence and/or absence, requires future research, both for plants with and without extrafloral nectaries.Estudiamos la asociación entre el arbusto sin nectarios extraflorales (NEF) Solanum lycocarpum (Solanaceae), la hormiga Camponotus crassus (Hymenoptera: Formicidae) y dos especies de membrácidos (Enchenopa spp., Hemiptera: Membracidae) productores de ambrosía, para determinar si las hormigas abandonarían a los membrácidos al ofrecerles experimentalmente una fuente de alimento considerablemente rica y abundante. El estudio se realizó en septiembre de 1999 en la Reserva Ecológica del Instituto Brasileiro de Geografía y Estadística (IBGE) (ca. 35 km al sur de Brasilia-DF, Brasil). Los resultados muestran que las hormigas fueron atraídas tanto al nectario experimental como a los membrácidos. Más aún, el número promedio de hormigas visitando a los nectarios experimentales fue significativamente mayor que el de las que visitaron a los membrácidos. Sin embargo, los membrácidos nunca fueron abandonados, lo que sugiere que las hormigas, a pesar de explotar una fuente abundante de alimento, no abandonan otras fuentes. Lo anterior puede explicarse por un incremento en la tasa de producción de ambrosía por los membrácidos, por lo que los niveles de atención de las hormigas hacia ellos no varía aún en la presencia de una fuente alternativa de alimento. Otra posibilidad puede ser la lealtad de las obreras por un recurso y su ubicación; algunos individuos retornan siempre a un sitio con una fuente de alimento. El poder demostrar que las hormigas cambian de recurso alimenticio basadas en cantidad y/o calidad, y no solamente basadas en la presencia o ausencia de los mismos, requiere de investigaciones futuras, tanto para plantas con como sin NEF

Understanding the Complex Structure of a Plant-Floral Visitor Network from Different Perspectives in Coastal Veracruz, Mexico

Our premise was to understand the basic structure of the flower-flower visitor community at La Mancha in Veracruz, Mexico. We used network analyses to study the structure of this community. In particular, to analyze, (1) if flower color and shape (“as a limited portion of the traditional floral syndromes definition”) were linked to the arrival of certain floral visitors, (2) if visits to flowers were generalist, specific and/or modular; and (3) which plant species, if any, in the core of the network could affect the stability of floral visitors. In order to analyze the organization of the plant-floral visitor community, we prepared network graphics using Pajek, nestedness (as NODF) with Aninhado, and modularity with the SA algorithm. The network obtained was nested suggesting that generalist species (with the most associations) were interacting with specialists (with fewer associations). Furthermore, floral visitors (Hymenoptera, Diptera, Lepidoptera and Trochiilidae) did not exhibit a particular preference for a specific flower color or shape, each pollinator group visited most flowers/colors/shapes considered. The same was similar for all 14 resulting modules. As in other studies, we suggest that pollination leans to generalization rather than to specialization. We suggest that maybe seasonality/food resource could be the factors to analyze as the next step in floral visits which may be the answer to modularity in this seasonal ecosystem

Change in land use, forest management and its implications in the biotic interactions associated with Cedrela odorata Linnaeus, Meliaceae

Este trabajo documenta el efecto del manejo forestal en las interacciones planta-hormiga-herbívoro asociadas al cedro rojo Cedrela odorata L. 1759 en ecosistemas forestales del centro de Veracruz, México. Debido a la elevada demanda como madera preciosa y a la creciente explotación de su ecosistema, las plantaciones de C. odorata han sido promovidas. Asimismo, el establecimiento de plantaciones monoespecíficas y sus prácticas de manejo, ha propiciado el desequilibrio ecológico y el desarrollo de plagas. En las plantaciones de C. odorata la larva del insecto herbívoro conocido como barrenador de los tallos Hypsipyla grandella (Lepidoptera: Pyralidae) se ha convertido en una plaga de gran importancia económica. En este estudio, encontramos que en los sitios donde las prácticas de manejo forestal se efectuaron con mayor intensidad, la complejidad y composición de la vegetación adyacente a los árboles de C. odorata disminuyó, afectando también negativamente a la riqueza de la comunidad de hormigas, pero incrementando su abundancia. En las plantaciones la dominancia se concentra en un bajo número de especies de hormigas características de ambientes perturbados y tolerantes al manejo. La diseminación del daño causada por el barrenador del tallo es mayor en sitios con elevada intensidad de manejo, encontrando un patrón invertido en los sitios mejor conservados.This paper documents the effect of forest management on plant-ant-herbivore interactions associated with red cedar Cedrela odorata L. 1759 in forest ecosystems of central Veracruz, Mexico. Due to the high demand as precious wood and the growing exploitation of its ecosystem, the plantations of C. odorata have been promoted. Likewise, the establishment of monospecific plantations and their management practices has led to ecological imbalance and the development of pests. In the plantations of C. odorata the larvae of the herbivorous insect known as the shoot borer Hypsipyla grandella (Lepidoptera: Pyralidae) have become a pest of great economic importance. In this study, we found that in the places where forest management practices were carried out with greater intensity, the complexity and composition of the vegetation adjacent to the trees of C. odorata decreased, also negatively affecting the richness of the community of ants, but increasing its abundance. In plantations, dominance is concentrated in a low number of ant species characteristic of disturbed and tolerant environments. The dissemination of the damage caused by the shoot borer is greater in sites with high intensity of management, finding an inverted pattern in the best conserved sites

Topological structure of plant-bee networks in four Mexican environments

We analyzed the topological structure (e.g., links per species, connectance, core-periphery analyses, specialization, nestedness, modularity) of plant-bee interactions of four areas in Mexico. We used qualitative data (binary networks). Mexico exhibits a complex geography and community variation that can affect bee networks. Network architecture is variable within large spatial scales, thus our results should vary according to site characteristics (La Mancha and Totula in Veracruz, Carrillo Puerto in Quintana Roo, and the Tehuacan-Cuicatlan valley, in Puebla), type of vegetation, endemism, altitude, size of area sampled. Network topology varied among sites, and the presence of nested or modular patterns was analyzed for robustness to simulated species extinctions. The lowest species richness was recorded for the Quintana Roo site (15 plant, 25 bee species), and the highest for the Tehuacan-Cuicatlan valley site (88 plant, 231 bee species). There was a tendency to have more connected networks when species richness was low and networks with greater species richness had a higher number of interactions. The distribution of interactions differed between environments but not due to network size and all were significantly nested. The robustness to cumulative extinctions showed a different pattern at each site; the most robust network was at Carrillo Puerto, which also was the site with less species. Sites with more endemic species (e.g. Tehuacan) had more specialized interactions, and were more susceptible to extinction

Inherited biotic protection in a Neotropical pioneer plant

Chelonanthus alatus is a bat-pollinated, pioneer Gentianaceae that clusters in patches where still-standing, dried-out stems are interspersed among live individuals. Flowers bear circum-floral nectaries (CFNs) that are attractive to ants, and seed dispersal is both barochorous and anemochorous. Although, in this study, live individuals never sheltered ant colonies, dried-out hollow stems - that can remain standing for 2 years - did. Workers from species nesting in dried-out stems as well as from ground-nesting species exploited the CFNs of live C. alatus individuals in the same patches during the daytime, but were absent at night (when bat pollination occurs) on 60.5% of the plants. By visiting the CFNs, the ants indirectly protect the flowers - but not the plant foliage - from herbivorous insects. We show that this protection is provided mostly by species nesting in dried-out stems, predominantly Pseudomyrmex gracilis. That dried-out stems remain standing for years and are regularly replaced results in an opportunistic, but stable association where colonies are sheltered by one generation of dead C. alatus while the live individuals nearby, belonging to the next generation, provide them with nectar; in turn, the ants protect their flowers from herbivores. We suggest that the investment in wood by C. alatus individuals permitting stillstanding, dried-out stems to shelter ant colonies constitutes an extended phenotype because foraging workers protect the flowers of live individuals in the same patch. Also, through this process these dried-out stems indirectly favor the reproduction (and so the fitness) of the next generation including both their own offspring and that of their siblings, alladding up to a potential case of inclusive fitness in plants

Diet and Feeding Behavior of the Horned Guan (Oreophasis derbianus) in Mexico

The Horned Guan (Oreophasis derbianus) is endemic to humid montane forests of southern Mexico and Guatemala. This species is considered endangered because of their small populations, the loss and fragmentation of habitat, illegal trade, and overexploitation by subsistence hunters. We update information about the species' diet and foraging behavior by integrating the results generated during two and a half decades of research on the Horned Guan's ecology at the El Triunfo Biosphere Reserve, in Chiapas, Mexico, with additional published information compiled from other areas. Based on nearly 450 hrs of direct observations of free-ranging guans and 530 discrete feeding events, we found that during the breeding season Horned Guans feed primarily on fruits from six species of plants and leaves from one species. Horned Guans were not observed eating animal matter, corroborating its specialized frugivore-folivore habits. Our study increases the known plant taxa found in the Horned Guan's diet in El Triunfo from 40 to 63 (Supplemental Material), and globally to 101 species (Supplemental Material). For 48 taxa in El Triunfo, only fruits were consumed, while for eleven taxa consumption was restricted to leaves, and to flowers for one species; for four taxa both fruits and leaves were consumed. We found significant differences between males and females in the location of foraging on trees and diet composition. Young birds are fed fruits of Citharexylum mocinnii and leaves of Solanum appendiculatum by their mothers, both of which are rare in the diet of adult males. The conservation of the Horned Guan requires the long-term protection of suitable habitat that maintains the plant species important in their diet

The Ecology of a Keystone Seed Disperser, the Ant Rhytidoponera violacea

Rhytidoponera violacea (Forel) (Hymenoptera: Formicidae) is a keystone seed disperser in Kwongan heathl and habitats of southwestern Australia. Like many myrmecochorous ants, little is known about the basic biology of this species. In this study various aspects of the biology of R. violacea were examined and the researchers evaluated how these characteristics may influence seed dispersal. R. violacea nesting habits (relatively shallow nests), foraging behavior (scramble competitor and lax food selection criteria), and other life history characteristics complement their role as a mutualist that interacts with the seeds of many plant species

Visitantes florales y potencial polinizador de trichocentrum stramineum orquídea amenazada y endémica de Veracruz

Trichocentrum stramineum es una orquídea endémica del centro de Veracruz y en categoría de riesgo como amenazada según la NOM-059-SEMARNAT-2010 por la reducción poblacional donde habita. Poca información se ha generado de esta orquídea en la literatura especializada y hasta antes del presente trabajo se desconocían sus visitantes florales y su potencial polinizador. De 2014 a 2017, se realizaron observaciones directas e indirectas a las inflorescencias de T. stramineum para registrar los visitantes florales y determinar al potencial polinizador en dos localidades de su distribución geográfica. Se registraron dos especies de visitantes florales en la localidad 1 y tres en la localidad 2. La abeja Centris nitida (Apidae) fue observada efectuando la remoción y deposición de los polinios de T. stramineum en ambas localidades. La abeja Centris tarsata (Apidae) fue observada un par de ocasiones en cada localidad sin que realizara remoción ni deposición de los polinios. La avispa Polybia sp., fue observada en dos ocasiones sólo en la localidad 2 sin llevar a cabo la remoción de polinios. La orquídea T. stramineum presenta un sistema especializado de polinización al depender solo de una especie para la fecundación de las flores. Los registros del polinizador de T. stramineum son los primeros en obtenerse y documentarse para esta orquídea

Interaction Intimacy Affects Structure and Coevolutionary Dynamics in Mutualistic Networks

The structure of mutualistic networks provides clues to processes shaping biodiversity [1-10]. Among them, interaction intimacy, the degree of biological association between partners, leads to differences in specialization patterns [4, 11] and might affect network organization [12]. Here, we investigated potential consequences of interaction intimacy for the structure and coevolution of mutualistic networks. From observed processes of selection on mutualistic interactions, it is expected that symbiotic interactions (high-interaction intimacy) will form species-poor networks characterized by compartmentalization [12, 13], whereas nonsymbiotic interactions (low intimacy) will lead to species-rich, nested networks in which there is a core of generalists and specialists often interact with generalists [3, 5, 7, 12, 14]. We demonstrated an association between interaction intimacy and structure in 19 ant-plant mutualistic networks. Through numerical simulations, we found that network structure of different forms of mutualism affects evolutionary change in distinct ways. Change in one species affects primarily one mutualistic partner in symbiotic interactions but might affect multiple partners in nonsymbiotic interactions. We hypothesize that coevolution in symbiotic interactions is characterized by frequent reciprocal changes between few partners, but coevolution in nonsymbiotic networks might show rare bursts of changes in which many species respond to evolutionary changes in a single species. © 2007 Elsevier Ltd. All rights reserved