title: Nectar secretion on fern fronds associated with lower levels of herbivore damage: field experiments with a widespread epiphyte of Mexican cloud forest remnants.
abstract: OBJECTIVE
  The oldest group of plants in which nectar secretions have been observed are the Polypodiopsida (ferns sensu lato). Nectaries have been reported in a dozen extant genera. The function of these nectaries has been investigated in several fern species, and in some circumstances has been demonstrated to have an antiherbivore role, attracting and maintaining biotic defence (ants and/or other predatory arthropods). This study documents foliar nectaries in Pleopeltis crassinervata, a widespread Central American epiphyte growing on a variety of trees in cloud forest areas of Veracruz, Mexico. This is a new record for this genus and species.
  
  
  METHODS
  As previous experimental work on epiphytic species of Polypodium has demonstrated a protective role of ants for developing fronds, we conducted similar experiments (using nylon nail polish to cover nectaries rather than excluding ants with bands of sticky resin as in earlier work). The fronds of Pl. crassinervata developed over 6 weeks, at which time damage was assessed. The experiment was simultaneously conducted on a sympatric species lacking nectaries, Polypodium furfuraceum. Herbivore placement experiments were conducted with large and small caterpillars on both of these ferns.
  
  
  RESULTS
  Fronds with nectaries covered suffered greater damage from herbivores over the course of their development, compared with fronds that had uncovered nectaries functioning normally. The parallel experiment on Po. furfuraceum showed no difference between manipulated and control fronds. Six species of ants (Brachymyrmex minutus, Crematogaster formosa, Paratrechina longicornis, Solenopsis geminata, S. picea and Wasmannia auropunctata) were observed visiting nectaries of Pl. crassinervata; most were effective in removing herbivore larvae placed on the fronds.
  
  
  CONCLUSIONS
  The long evolutionary history of ferns may explain why some previous studies of fern nectaries have shown little or no benefit to ferns from nectary visitors, as any coevolved herbivores are those resistant to ant defence. The results suggest that ants protect Pl. crassinervata fronds against herbivory. The presence of nectaries, and the relationship with ants, may contribute to this fern's widespread occurrence and persistence in the face of disturbance, though many other factors also play a role. Ant defence may be more likely to benefit a widespread species of disturbed habitats that encounters a wide range of non-adapted herbivores.
authors.affiliation: Department of Biological Sciences, Florida International University, Miami, FL 33199, USA. kopturs@fiu.edu
authors.forename: Suzanne
authors.initials: S
authors.lastname: Koptur
authors.forename: Mónica
authors.initials: M
authors.lastname: Palacios-Rios
authors.forename: Cecilia
authors.initials: C
authors.lastname: Díaz-Castelazo
authors.forename: William P
authors.initials: WP
authors.lastname: Mackay
authors.forename: Víctor
authors.initials: V
authors.lastname: Rico-Gray
mesh_terms: D000818:Animals; D060434:Herbivory; D007313:Insecta; D007814:Larva; D008800:Mexico; D057048:Plant Nectar / Q000378:metabolism*; D032494:Polypodium / Q000502:physiology*; D013559:Symbiosis
chemical_list: D057048:Plant Nectar
keywords: Extrafloral nectar; Formicidae; Mexico; Pleopeltis crassinervata; Polypodium furfuraceum; ant protection; antiherbivore defence; cloud forest; epiphyte; extrasoral nectar; ferns; field experiment; herbivory; nectaries; pteridophytes