Impact of the female and hermaphrodite forms of Opuntia robusta on the plant defence hypothesis.

The optimal defence hypothesis predicts that increased plant defence capabilities, lower levels of damage, and lower investment in vegetative biomass will occur more frequently in sexual forms with higher resource-demanding tissue production and/or younger plant parts. We aimed to examine the effects of sexual form, cladode, and flower age on growth rate, herbivore damage, and 4-hydroxybenzoic acid (4-HBA), chlorogenic acid, and quercetin (QUE) concentrations in Opuntia robusta plants in central Mexico. Our findings demonstrated that hermaphrodite flowers showed faster growth and lesser damage than female flowers. The effect of cladode sexual forms on 4-HBA and QUE occurrence was consistent with the predictions of the optimal defence hypothesis. However, chlorogenic acid occurrences were not significantly affected by sexual forms. Old cladodes exhibited higher QUE and 4-HBA occurrences than young cladodes, and hermaphrodites exhibited higher 4-HBA concentrations than females. Resource allocation for reproduction and secondary metabolite production, and growth was higher and lower, respectively, in females, compared to hermaphrodites, indicating a trade-off between investment in reproduction, growth, and secondary metabolite production. Secondary metabolite concentrations in O. robusta plants were not negatively correlated with herbivore damage, and the two traits were not accurate predictors of plant reproductive output.

First description of extrafloral nectaries in Opuntia robusta (Cactaceae): Anatomy and ultrastructure.

To our knowledge, there are no studies about the structure and ecological function of extrafloral nectaries (EFNs) in Opuntia robusta. This is the first description of EFNs in O. robusta, where young spines have an interesting structure and a secreting function, which are different from EFNs described in other Cactaceae species. We used light, scanning-electron, and transmission-electron microscopy to examine morphology, anatomy, and ultrastructure of the secretory spines in areoles in female and hermaphrodite individuals of O. robusta. Young cladodes develop areoles with modified and secretory spines as EFNs only active during the early growth phase. EFNs are non-vascularized structures, with no stomata, that consist of a basal meristematic tissue, a middle elongation region, and an apical secretory cone formed by large globular epidermal cells, containing nectar and medullar elongated cells. We observed the presence of Golgi apparatus, vesicles and plastids in the medullar and sup-epidermal cells of the spine. We propose that the nectar is stored in the globular cells at the apex of the spine and secreted by breaking through the globular cells or by pores. We recorded a more frequent presence of ants on younger cladode sprouts producing young secreting spines: this result is parallel with the predictions of Optimal Defense Hypothesis, which states that younger plant organs should be better defended than older ones because their loss produces a higher fitness impairment. Although Diaz-Castelazo's hypothesis states that a more complex structure of EFNs correlates with their lower among-organs dispersion, comparing to less complex EFNs, non-vascularized structure of EFNs in O. robusta is not associated with their higher among-organs dispersion likened to O. stricta, which produces vascularized EFNs. We provide evidence that this characteristic is not a good taxonomic feature of Opuntia genus. Moreover, the comparison of EFNs of O. robusta and O. stricta suggests that the hypothesis of Diaz-Castelazo should be revised: it is rather a rule but not a law.

Eimeria atlapetesi nom. nov., a replacement name for Eimeria pileata Soriano-Vargas et al., 2015 (Apicomplexa: Eimeriidae), preoccupied by Eimeria pileata Straneva and Kelley, 1979 (Apicomplexa: Eimeriidae), with observations on histopathology and phylogenetic analysis.

Eimeria pileata Soriano-Vargas, Medina, Salgado-Miranda, García-Conejo, Galindo-Sánchez, Janczur, Berto and Lopes, 2015 is a junior homonym of Eimeria pileata Straneva and Kelley, 1979 and needs to be replaced. This coccidium was described from a rufous-capped brush finch Atlapetes pileatus Wagler in the Nevado de Toluca Natural Protected Area, Mexico. Thus, to maintain the original intent of the specific epithet derived from the scientific name of the type-host, the name Eimeria atlapetesi nom. nov. is proposed as a replacement name. Additionally, the current work reports another rufous-capped brush finch A. pileatus parasitized by E. atlapetesi in co-infection with an Isospora sp., providing observations of histopathology and phylogenetic analysis of 18S ribosomal RNA (rRNA) gene from E. atlapetesi. Endogenous forms of E. atlapetesi and Isospora sp. were observed in intestinal sections. Few oocysts of Isospora sp. were observed; therefore they were not morphologically or molecularly identified. In return, E. atlapetesi was identified and it was phylogenetically close to Eimeria dispersa Tyzzer, 1929 from the domestic turkey Meleagris gallopavo Linnaeus.

Isospora cardellinae n. sp. (Apicomplexa: Eimeriidae) from the red warbler Cardellina rubra (Swainson) (Passeriformes: Parulidae) in Mexico.

A new coccidian species (Protozoa: Apicomplexa: Eimeriidae) collected from the red warbler Cardellina rubra (Swainson) is reported from the Nevado de Toluca National Park, Mexico. Isospora cardellinae n. sp. has subspherical oöcysts, measuring on average 26.6 × 25.4 µm, with smooth, bi-layered wall, c.1.3 µm thick. Micropyle, oöcyst residuum, and polar granule are absent. Sporocysts are ovoidal, measuring on average 19.0 × 12.0 µm, with a knob-like Stieda body, a trapezoidal sub-Stieda body and sporocyst residuum composed of scattered spherules of different sizes. Sporozoites are vermiform with one refractile body and a nucleus. This is the fourth description of an isosporoid coccidian infecting a New World warbler.

Eimeria pileata n. sp. (Apicomplexa: Eimeriidae) from the rufous-capped brush finch Atlapetes pileatus Wagler (Passeriformes: Emberizidae) in Mexico.

A new coccidian species (Protista: Apicomplexa: Eimeriidae) collected from the rufous-capped brush finch Atlapetes pileatus Wagler in the Nevado de Toluca Natural Protected Area, Mexico. Oöcysts of Eimeria pileata n. sp. are ellipsoidal, measuring on average 16.5 × 14.1 µm, with a smooth, bi-layered wall. Micropyle and oöcyst residuum are absent, but a polar granule is present. Sporocysts are ellipsoidal, measuring on average 9.0 × 5.4 µm. Stieda and sub-Stieda bodies are both present. A sporocyst residuum is present as a compact mass of granules. This is the third description of an eimeriid coccidian infecting passerines.

Isospora celata n. sp. (Apicomplexa: Eimeriidae) from the orange-crowned warbler Oreothlypis celata (Say) (Passeriformes: Parulidae) in Mexico.

A new coccidian species (Protista: Apicomplexa: Eimeriidae) is described from the orange-crowned warbler Oreothlypis celata (Say) collected in the Nevado de Toluca National Park, Mexico at 3,000 metres above sea level. Isospora celata n. sp. has subspheroidal oöcysts, measuring 28.4 × 26.4 µm, with smooth, bi-layered wall c.1.2 µm thick. Micropyle and polar granule are absent, but oöcyst residuum is present as a compact mass. Sporocysts are ovoidal, 18.2 × 12.8 µm. Stieda body knob-like and sub-Stieda body irregular and barely discernible. Sporocyst residuum is composed of granules of different sizes. Sporozoites are vermiform with one refractile body and a nucleus. This is the third description of an isosporoid coccidian infecting a New World warbler.

Chemical and physical defense traits in two sexual forms of Opuntia robusta in Central Eastern Mexico.

Sexually dimorphic plants provide an excellent opportunity for examining the differences in the extent of their defense against herbivores because they exhibit sex-related differences in reproductive investment. Such differences enable comparison of the sex with high reproduction expenses with the sex that expends less. The more costly sex is usually also better defended against herbivores. Generally, females are considered more valuable than hermaphrodites in terms of fitness; however, hermaphrodites are more valuable if they can produce seed by autonomous selfing, provided that the inbreeding depression is low and pollen is limited. We studied a gynodioecious population of Opuntia robusta from Central-Eastern Mexico, which has been reported to be trioecious, dioecious, or hermaphrodite, and addressed the following questions: 1) Is the hermaphrodite's reproductive output higher than the female's, and are hermaphrodites thus better defended? 2) Are plant tissues differentially defended? 3) Do trade-offs exist among different physical defense traits? and 4) among physical and chemical defense traits? We found that 1) hermaphrodites had a higher seed output and more spines per areola than females and that their spines contained less moisture. Non-reproductive hermaphrodite cladodes contained more total phenolic compounds (TPCs) than female ones. In addition, 2) hermaphrodite reproductive cladodes bore more spines than female cladodes, and 3) and 4) we found a negative relationship between spine number per areola and areola number per cladode and a positive relationship between spine number per areola per plant and TPC concentration per plant. Non-reproductive hermaphrodite cladodes contained a higher concentration of TPCs than female cladodes, and parental cladodes contained fewer TPCs than both reproductive and empty cladodes.