Giant Tortoises as Ecological Engineers: A Long-term Quasi-experiment in the Galápagos Islands

Giant tortoises were once a megafaunal element widespread in tropical and subtropical ecosystems. The role of giant tortoises as herbivores and seed dispersers, however, is poorly known. We evaluated tortoise impacts on Opuntia cactus (Cactaceae) in the Galápagos Islands, one of the last areas where giant tortoises remain extant, where the cactus is a keystone resource for many animals. We contrasted cactus populations immediately inside and outside natural habitats where tortoises had been held captive for several decades. Through browsing primarily and trampling secondarily tortoises strongly reduced densities of small (0.5–1.5 m high) cacti, especially near adult cacti, and thereby reduced densities of cacti in larger size classes. Tortoises also caused a shift from vegetative to sexual modes of reproduction in cacti. We conclude that giant tortoises promote a sparse and scattered distribution in Opuntia cactus and its associated biota in the Galápagos Islands.     

Nurse plants vs. nurse objects: effects of woody plants and rocky cavities on the recruitment of the Pilosocereus leucocephalus columnar cactus

BACKGROUND AND AIMS: Most studies on cactus recruitment have focused on the role of woody plants as seedling facilitators. Although the spatial association of cacti with objects had been described, the mechanisms underlying this association remain unknown. The aims of this study were to identify which mechanisms facilitate the establishment of a columnar cactus under the shade and protection of objects and to compare these mechanisms with those involved in plant-plant facilitation. METHODS: Three split-split-plot field experiments were conducted to compare the effects of two microhabitats (inside rocky cavities and beneath plant canopies) on seed removal, germination, seedling survivorship and dry weight. Flat, open spaces were used as the control. For each microhabitat, the effect of seed or seedling protection and substrate limitation were explored; aboveground microclimate and some soil properties were also characterized. KEY RESULTS: The permanence of superficial seeds was greater inside rocky cavities than beneath woody plant canopies or on flat, open areas. Germination was similar in cavities and beneath plant canopies, but significantly higher than on flat, open areas. Seedling survivorship was greater beneath plant canopies than inside cavities or on flat, open spaces. CONCLUSIONS: The mechanisms of plant facilitation are different from those of object facilitation. There are seed-seedling conflicts involved in the recruitment of P. leucocephalus: nurse plants favour mainly seedling survivorship by providing a suitable microenvironment, while nurse objects mainly favour seed permanence, by protecting them from predators.     

Host‐dependent phenotypic plasticity of aedeagus morphology in a pair of cactophilic sibling Drosophila species of the repleta group (Diptera,Drosophilidae)

The rapid evolution of male genital morphology is a characteristic feature of several animal groups. Such rapid divergence makes this trait a useful key for species identification. The aedeagus, the intromittent organ of male genitalia, is considered the main diagnostic trait in the Drosophila repleta group. In this study we analysed phenotypic plasticity and genetic variations associated with aedeagus size and shape in the cactophilic sibling species Drosophila gouveai Tidon‐Sklorz and Sene, 2001 and Drosophila antonietae Tidon‐Sklorz and Sene, 2001. Phenotypic plasticity in aedeagus morphology was evaluated in terms of the response to rearing media prepared with each species’ natural host plant, Pilosocereus machrisii Dawson, 1957 and Cereus hildmannianus Schum, 1890 respectively. Our results show that aedeagal shape differed significantly between species and that both shape and size presented host‐related phenotypic plasticity in both species. Flies reared on P. machrisii had, on average, larger aedeagi than those grown in C. hildmannianus. The general shape of aedeagus also differed significantly between flies that emerged in different host cactus. Patterns of variation in aedeagus morphology are discussed in the light of the current knowledge of evolutionary relationships and host plant use, in the D. buzzatii cluster, an assemblage of species in active cladogenesis.     

An alkaloid fraction extracted from the cactus Trichocereus terscheckii affects fitness in the cactophilic fly Drosophila buzzatii (Diptera: Drosophilidae)

The host‐plant environment of phytophagous insects directly affects various aspects of an insect's life cycle. Interestingly, relatively few insect groups have specialized in the exploitation of plants in the Cactaceae family, potentially because of the chemical and ecological challenges imposed by these plants. The cactophilic Drosophila buzzatii Patterson & Wheeler, 1942 is a well‐studied model in evolutionary ecology, partially because of its ability to exploit toxic cactus hosts. Previous studies have shown a negative effect on performance when flies are reared in an alternative columnar cactus host of the genus Trichocereus, relative to its primary cactus host, Opuntia. These observations were attributed to the presence of alkaloids in Trichocereus tissues, a chemical deterrent to herbivores that indirectly affects Drosophila larvae; however, the putative toxic effect of alkaloids has never been tested directly in D. buzzatii. The present study is the first attempt to relate chemical extracts in Trichocereus terscheckii Britton & Rose, 1920 with detrimental effects on D. buzzatii. We assessed the effects of a crude alkaloid extract, rich in phenylethylamines, and a ‘non‐alkaloid fraction’ on viability and adult wing morphology. Our results indicate that rearing larvae on an artificial diet containing different concentrations of the crude alkaloid extract decreased pupal viability and adult size in a concentration‐dependent manner. We discuss the role of cactus alkaloids in the evolution of host‐plant use in cactophilic flies. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 342–353.     

Stem tilting,pseudocephalium orientation,and stem allometry in <Emphasis Type="Italic">Cephalocereus columna-trajani</Emphasis> along a short latitudinal gradient

Cephalocereus columna-trajani is a giant columnar cactus endemic of the Tehuacán-Cuicatlán Valley in Central Mexico. Stem tilting and northward pseudocephalium azimuth in C.␣columna-trajani have functional advantages in terms of interception of direct solar radiation at the northernmost portions of its range. Since the success of both characters strongly depends on the apparent position of the sun during the growing season, in this paper we test the hypothesis that the occurrence of such columnar morphology is restricted geographically and imposes mechanical restrictions that limit column height. Following a latitudinal gradient along the Tehuacán-Cuicatlán Valley, we selected five populations, recorded tilting angle and pseudocephalium azimuth, and carried out allometric and biomechanical analyses of height–diameter relationships. Northern populations showed higher tilting angles. Pseudocephalium azimuth significantly differed among populations, and pseudocephalium orientation was consistently North-Northwestern. Stem allometry showed that the stems of the southern populations increased in height at a far greater rate with respect to diameter than the northern populations. The southernmost population showed the lowest safety factor. These results support the hypothesis that stem tilting in C.␣columna-trajani is functionally advantageous in a restricted geographical range, and imposes mechanical restrictions to column height.     

Pereskiophaga brasiliensis,a natural enemy of the invasive alien cactus Pereskia aculeata,is not suitably host specific for biological control in South Africa

The stem-mining weevil, Pereskiophaga brasiliensis, was a candidate biological control agent for the invasive cactus Pereskia aculeata in South Africa. In host specificity trials, it developed on two indigenous test plant species under choice and no-choice conditions. Pereskiophaga brasiliensis is therefore not suitably host specific for release in South Africa.     

Effect of rearing Cactoblastis cactorum on an artificial diet on the behaviour of Apanteles opuntiarum

Cactoblastis cactorum's unintended arrival to Florida and its expansion in North America represent a threat to Opuntia-based agriculture and natural ecosystems in the United States and Mexico. Apanteles opuntiarum attacks C. cactorum and is a potential biocontrol agent due to its specificity, wide distribution and occurrence. Laboratory rearing methods using excised cladodes for C. cactorum as host larvae of A. opuntiarum were developed, but require a continuous supply of cactus with a risk of microorganisms compromising the rearing. Host cues —including odour of host metabolic subproducts like faeces and chemicals emitted by the attacked plant— are the most important signals that help a parasitoid locate a host. Little attention has been paid to behavioural differences of parasitoids in the presence of hosts reared on artificial diet. Thus, the aims of the present work were to determine the effect of meridic diet for C. cactorum on parasitoid behaviour and to determine whether prior experience (previous exposure to stimuli) influences the response of A. opuntiarum towards hosts. Parasitism rates were assessed using cladodes or meridic diet as larvae food source. Behavioural experiments also evaluated the effect of prior experience (larvae or frass from larvae fed on cactus or diet) on host searching, encounter and attack of different stimulus (larvae or frass from larvae fed on cactus or diet). Parasitism behaviour of A. opuntiarum was negatively affected by the use of meridic diet to feed host larvae. Presenting parasitoids with larvae rather than their frass influenced excitatory behavioural responses in terms of contact, probing and stinging in A. opuntiarum females, whereas the effect of prior experience on the behaviours was not quite consistent. For laboratory mass-rearing procedures of A. opuntiarum, we recommend previous contact of females with frass from cactus-fed larvae and a piece of cactus, which showed enhanced parasitism rates.     

Interaction between the cactus Neobuxbaumia tetetzo and the nurse shrub Mimosa luisana

Abstract. Pattern analysis and association analysis showed that recruitment of the giant columnar cactus Neobuxbaumia tetetzo in the semi-arid valley of Zapotitlán is largely limited to areas beneath the canopies of perennial shrubs, acting as nurse plants. Chi-square and Haberman tests revealed that young cacti were more frequently found beneath canopies of leguminous shrubs, especially Mimosa luisana, than were older cacti. Segregation analysis indicated a positive association (= negative segregation) of young cacti and M. luisana canopies, but older cacti were randomly distributed with respect to M. luisana. This, and the height class distributions of cacti associated withM luisana or not, suggested the replacement of M. luisana by N. tetetzo. Mimosa luisana plants in association withiV. tetetzo had greater amounts of dead basal area thanM luisana lacking associated N tetetzo. This suggested competition from JV. tetetzo promotes the replacement process. The roots of N tetetzo are ≤ 30 cm deep. Interception of soil water before penetration to deeper roots of M luisana may be the mode of competition between the two species.     

Pollination biology of Harrisia portoricensis (Cactaceae), an endangered Caribbean species

Traits associated with self-pollination are common features of island plant communities. In this work, we studied the pollination biology and the breeding system of Harrisia portoricensis, an island columnar cactus, to test for the presence of inbreeding and inbreeding depression. For H. portoricensis, which bears flowers with typical outcrossing morphology, the results from 322 h of direct observations and videotaping showed that visits to flowers by animals were uncommon. Controlled pollinations demonstrated that H. portoricensis has a partially self-compatible breeding system that it is not autogamous and thus requires an external mechanism for the movement of pollen to set fruit. We detected differences in seed size, seed mass, germination success, and multiplicative fitness estimates between self- and cross-pollination treatments. We found that progeny resulting from natural and self-pollination treatments showed signs of inbreeding depression compared with progeny resulting from cross-pollination; however, the magnitude of the inbreeding depression was less than 50%. Our combined results suggest that for this species an endogamous breeding system should be favored by natural selection.     

<Emphasis Type="Italic">In vitro</Emphasis> organogenesis of <Emphasis Type="Italic">Pelecyphora aselliformis</Emphasis> erhenberg (cactaceae)

Summary In vitro propagation of Pelecyphora aselliformis, a Mexican cactus which is considered rare and is highly valued in the commercial market, was initiated using seeds as explants. The longitudinal explants from seedlings germinated in vitro were cultivated on Murashige and Skoog medium containing 8.8 μM benzyladenine (BA) or 4.6 μM kinetin at pH 7.0. After 120 d, each explant gave rise to five shoots and this number of shoots increased 20–25% after subculture. The hyperhydricity was similar in both media, but callus formation was lower on the medium with BA. The shoot development, in terms of epicotyl length, and fresh and dry weight after 6 wk, was also recorded. The epicotyl length was similar on shoot-forming media but the quality of shoots was better on media containing BA. In about 1 yr, 500–600 well-defined shoots were obtained. The rooting of shoots was very slow and a vigorous radical system was observed after 1 yr of culture.