Host-symbiont specificity during onset of symbiosis between the dinoflagellates Symbiodinium spp. and planula larvae of the scleractinian coral Fungia scutaria

Many corals which engage in symbioses with dinoflagellates from the genus Symbiodinium (zooxanthellae) produce offspring which initially lack zooxanthellae. These species must choose their symbionts from numerous genetically distinct strains of zooxanthellae co-occurring in the environment. In most cases, symbiosis onset results in an association between a specific host coral and a specific strain of algal symbiont. This is the first study to examine host-symbiont specificity during symbiosis onset in a larval cnidarian, and the first to examine such events in a scleractinian of any life stage. We infected planula larvae of the solitary Hawaiian scleractinian Fungia scutaria with both homologous zooxanthellae, freshly isolated from F. scutaria adults, and heterologous zooxanthellae, isolated from Montipora verrucosa, Porites compressa, and Pocillopora damicornis, three species of scleractinians which co-occur with F. scutaria. We found that homologous zooxanthellae were better able to establish symbioses with larval hosts than were heterologous isolates, by two separate measures: percent of a larval population infected, and densities of zooxanthellae per larva. We also measured algal densities in larvae over a 4-day period until the onset of settlement and metamorphosis. We found no changes in zooxanthella population densities, regardless of zooxanthella type or the light environment in which they were incubated. Strong infection of host larvae with homologous algae compared to heterologous algae suggests that there is a specificity process which occurs sometime during the early stages of infection between the partners, and which results in the establishment of a specific symbiosis.     

Variation in behavior of lycaenid larvae when attended by different ant species

Partner quality can be crucial for the outcome of a mutualistic interaction. In multi-species associations, the characteristics of potential partners can vary substantially and thus the associated benefits. As a consequence of such variation, one might expect strategies of adjusting investments to the characteristics of a partner. Lycaenid butterfly larvae often interact mutualistically with several ant species of different size and aggressiveness and thereby different ability to protect the larvae. Attending ants are rewarded with nutritious secretions. Both ant behavior and a larva's need for protection are known to influence larval investment in the relationship. This study investigates the effect of six ant species on larval behavior in the lycaenid butterfly, Glaucopsyche lygdamus. The overall level of secretion, as well as the response to varying number of attending ants, were found to be influenced by ant species. This revised version was published online in November 2006 with corrections to the Cover Date.     

Ant protection of a Heteropteran trophobiont against a parasitoid wasp

Trophobiotic associations between Caternaultiella rugosa (Heteroptera; Plataspidae) and two ant species, Camponotus brutus (Formicidae) and Myrmicaria opaciventris (Myrmicinae), were recorded at the base of the trunks of Bridelia micrantha (Euphorbiaceae). The bug colonies were generally sheltered in pavilions built by the ants, but during periods of proliferation, part of the colonies developed outside the pavilions. We examined the protective activity of the two associated ant species against an encyrtid parasitoid wasp that parasitized egg masses of the bug. Egg masses situated outside the pavilions were significantly less parasitized on trees occupied by either ant species than on experimental trees where ants were excluded by a sticky barrier. Egg masses were significantly less parasitized on trees occupied by C. brutus than on those occupied by M. opaciventris, both inside and outside the pavilions. Moreover, they were less parasitized inside than outside the pavilions when protected by C. brutus. Inside the pavilions, parental care of the egg masses and first-instar nymphs was abandoned to the ants. In contrast, outside the pavilions, the females of C. rugosa guarded egg masses and clusters of first-instar nymphs. Furthermore, adults and last-instar nymphs grouped around and above first-instar nymphs, forming a protective shield. These groups formed and disintegrated daily, according to the attending ant's rhythm of activity.     

Competition between the gypsy moth, Lymantria dispar, and the northern tiger swallowtail, Papilio canadensis: interactions mediated by host plant chemistry, pathogens, and parasitoids

The gypsy moth, Lymantria dispar, and the northern tiger swallowtail, Papilio canadensis, overlap geographically as well as in their host ranges. Adult female swallowtails are incapable of distinguishing between damaged and undamaged leaves, and the opportunities for competition between these two species are numerous. We designed field and laboratory experiments to look for evidence of indirect competition between P. canadensis and L. dispar larvae. Swallowtail caterpillars were reared in the laboratory on leaves from gypsy-moth-defoliated and undefoliated trees to explore host-plant effects. We tested for pathogen-mediated interactions by rearing swallowtail larvae on both sterilized and unsterilized leaves from defoliated and undefoliated sources. In addition, we measured the effects of known gypsy moth pathogens, as well as gypsy moth body fluids, on the growth and survival of swallowtail larvae. Field experiments were designed to detect the presence of parasitoid-mediated competition, as well: we recorded parasitism of swallowtail caterpillars placed in the field either where there were no gypsy moth larvae present, or where we had artificially created dense gypsy moth populations. We found evidence that swallowtails were negatively affected by gypsy moths in several ways: defoliation by gypsy moths depressed swallowtail growth rate and survival, whether leaves were sterilized or not; sterilization significantly reduced the effect of defoliation, and gypsy moth body fluids proved lethal; and swallowtail caterpillars suffered significantly increased rates of parasitism when they were placed in the field near gypsy moth infestations.     

Species-specific effects of tending ants on the development of lycaenid butterfly larvae

The transfer of nutrients between organisms is a common feature of mutualism. The production of these food rewards is often assumed to be costly. Estimation of the costs of producing food rewards is important for understanding the overall effects of the interaction on fitness. When food rewards are harvested by several species differing in foraging behavior, costs to the producer may differ. The larvae of many species in the butterfly family Lycaenidae produce secretions consumed by tending ants. Here I report that three North American ant species, Formica perpilosa, Dorymyrmex sp. (smithi complex), and Forelius foetida, had no negative effect on the duration of development and adult size of the lycaenid Hemiargus isola. Moreover, tending by the ant Formica perpilosa significantly enhanced larval growth, resulting in butterflies that were 20% heavier than their untended counterparts. Tending by the ants Dorymyrmex sp. (smithi complex) and Forelius foetida had no effect on butterfly weight. Tended, nonfeeding larvae lost 69% more weight than untended, nonfeeding larvae. Taken together, the results suggest that, although ant tending imposes a physiological cost, H. isola larvae use behavioral or physiological mechanisms to compensate or overcompensate for nutrients lost to ants.     

Ant visitation of extrafloral nectaries of Passiflora: the effects of nectary attributes and ant behavior on patterns in facultative ant-plant mutualisms

Extrafloral nectary (EFN) plants are widespread and can be quite species-rich in some communities. Thus, ants that utilize extrafloral nectar may have the opportunity to discriminate among a wide variety of nectar sources, resulting in variation in the ant attention EFN plants receive. In this study, we compare ant visitation rates of three Passiflora species that coexist in an early successional neotropical forest. These three vine species (Passiflora auriculata, P. biflora, and P. oerstedii) differ in their extrafloral nectary structure and placement, and thus may attract different numbers or species of ants. Through censuses of ants tending extrafloral nectaries, we found that P. auriculata received significantly higher numbers of ant visitors than P. oerstedii, but did not differ significantly from P. biflora in its attractiveness to ants. We also found that termite worker baits (simulating herbivores) placed on P. auriculata and P. biflora were discovered by ants significantly more quickly than baits placed on P. oerstedii. In both ant visitation censuses and in termite bait trials, we found no significant associations between Passiflora species and the species of ant visitors. We also performed experimental manipulations of several characteristics of P. auriculata, which resulted in changes in levels of ant visitation. When petiolar nectaries of P. auriculata were experimentally blocked, visitation by the common ant Ectatomma ruidum declined, even though nectaries on the leaf surfaces were still functional. Connections with other vegetation also had an effect on ant visitation. Though experimental creation of connections between growing P. auriculata shoots and other vegetation did not enhance ant visitation, eliminating connections resulted in a significant decline in the number of ant visitors. The results of this study suggest factors that may contribute to variation in ant visitation of extrafloral nectary plants. In addition, this study demonstrates that extrafloral nectary plants co-occurring in a habitat and available to the same ants may differ in patterns of visitation by ants and perhaps in the quality of protection from herbivores that they receive.     

A high incidence of parasitism ofHeliothis SPP. [Lep.: Noctuidae] larvae in cotton in Southeastern Arkansas

During 1981 and 1982, bollworm,Heliothis zea (Boddie), and tobacco budworm,H. virescens (F.), larvae (n=3,666) were collected from 41 cotton fields near Portland, Arkansas (USA) to assess the occurrence of parasitism. Three strategies were employed to controlHeliothis spp. in these fields: (1) release ofTrichogramma pretiosum Riley; (2) insecticidal control; or (3) inaction (check). Insecticide use in nonchemical control fields was reduced, but not eliminated.Heliothis spp. larvae collected in cotton had higher parasitism rates in 1981 (30.9%) and 1982 (50.1%) than had been reported for cotton since the advent of organochlorine insecticide usage. Four species of larval parasites and 1 species of larval-pupal parasite were recorded. The larval parasiteMicroplitis croceipes (Cresson) comprised 90.6% and 94.5% of all parasitic insects reared from field collectedHeliothis spp. in 1981 and 1982, respectively. No difference (P>0.05) in level of parasitism existed betweenH. zea andH. virescens. Differences between treatments occurred only in 1982 whenH. zea larvae were parasitized at a greater (P<0.05) rate in check fields (68.3%) than in insecticidal control fields (44.3%). Higher levels of larval parasitism in cotton fields may be a consequence of reduced insecticide usage and changes in materials applied, particularly the pyrethroids. Mention of a trademark or proprietary product does not constitute a guarantee or warranty of the product by the U.S. Dept. of Agriculture and does not imply its approval to the exclusion of other products that may also be suitable.     

Field evaluation of insect resistance in a wild tomato and its effects on insect parasitoids

Populations ofHelicoverpa (=Heliothis) zea (Boddie),Heliothis virescens (F.),Manduca sexta (L.) andM. quinquemaculata (Haw.) and their egg and larval parasitoids were sampled in field plots of the: insect-resistant wild tomato,Lycopersicon hirsutum f. glabratum C. H. Mull, accession PI 134417; susceptible commercial tomato cultivar ‘Better Boy’; F₁ hybrid; and selected, moderately resistant backcross genotype. Densities ofH. zea andH. virescens eggs and small larvae were higher on resistant genotypes than on susceptible genotypes, but densities of large larvae were similar on all genotypes. Densities ofManduca spp. larvae were too low to permit similar analyses of the effects of plant genotype. Rates of egg parasitism byTrichogramma spp. andTelenomus sphingis (Ashmead) were reduced on insect-resistant genotypes. Rates of parasitism by the larval parasitoidsCampoletis sonorensis (Cameron) andCotesia congregata (Say) were reduced on resistant genotypes. No consistent effects on parasitism rates byCotesia marginiventris (Cresson) were observed and parasitism rates byCardiochiles nigriceps Viereck were unaffected.     

Larval niche selection and evening exposure enhance adoption of a predacious social parasite, Maculinea arion (large blue butterfly), by Myrmica ants

Larvae of the butterfly genus Maculinea feed briefly on a foodplant before being adopted as social parasites into Myrmica ant nests. Each Maculinea species typically survives only with a single Myrmica species, yet the eggs are laid across the overlapping territories of 3-5 Myrmica species and several other ants. The ability of Maculinea arion - a 'predatory' species of Maculinea - to influence its adoption into host Myrmica colonies was studied for the first time in the field. Some earlier reports, involving captive non-host ants, suggested that larvae of the predatory Maculinea follow ant trails into host nests or wander some distance from their foodplant before being discovered and (after a long interaction) carried away by Myrmica foragers. No dispersal from foodplants occurred in wild Maculinea arion larvae. Instead, they increased by >100-fold their probability of being discovered and adopted by Myrmica spp., rather than by ants of other genera, by exposing themselves in the micro-niche occupied by Myrmica foragers at their time of day of peak foraging. Despite a complex, hour-long interaction with Myrmica workers before being carried to the nest, Maculinea arion did not enhance its adoption by host species of Myrmica. Eggs were laid without bias in Myrmica sabuleti (host) and Myrmica scabrinodis (non-host) territories; larval survival on Thymus was the same in both ants' territories; larvae waited to be found beneath their foodplant rather than seek their host; Myrmica sabuleti and Myrmica scabrinodis foraged in the same vertical and temporal niches, and had the same probability of discovering larvae; both ants behaved identically after finding larvae and took the same time to adopt them; and the ratio of wild larvae taken into Myrmica sabuleti or Myrmica scabrinodis nests was the same as the distribution between these ants of Thymus, eggs and pre-adoption larvae.     

Nutritional function of replete workers in the pharaoh's ant, Monomorium pharaonis (L.)

Summary: Queens of the pharaoh's ant Monomorium pharaonis (L.), like several other ant species, feed on larval secretions as their main nourishment and their fecundity is positively correlated with the number of large larvae present in the nest. The surplus of secretions produced by larvae is stored in a temporary caste of replete workers, which comprises young workers who remain in the nest and store liquid nourishment. Repletes are characterised by a conspicuously large gaster, caused by large amounts of liquid food stored in the crop, from which it may be regurgitated and distributed among colony members. In this study, repletes of pharaoh's ants were demonstrated to be functioning as buffers, smoothing fluctuations in availability of high quality food to the reproductive queens when larvae are scarce or missing, thus temporarily keeping up the egg production of queens.¶In undisturbed two-queen colonies with 20 large worker larvae and 30 workers (15 young and 15 old workers), approximately 10 repletes developed (one replete per two larvae). Development of older workers into repletes, when some or all repletes had been removed from the colonies, demonstrated that their temporal polyethism exhibits great plasticity in this trait.¶This study confirmed that, in pharaoh's ants, the regulation of fecundity depends not only on the food flow to the queen from larvae or from repletes but also on an unknown larval stimulus.¶The term crop repletes is suggested for replete workers which use their crop to store nourishment, as opposed to fat-body repletes, which store nourishment in their fat body.¶The presence of brood tending crop repletes in nests in several European ant species of Leptothorax, Myrmica, and Lasius, show that repletism is a common trait in ants, and that it may play an important role in regulation of nutrition in ant colonies, as demonstrated in Monomorium pharaonis.     

Prevalence of the parasite Strepsiptera in <Emphasis Type="Italic">Polistes</Emphasis> as detected by dissection of immatures

Summary. hough the paper wasp genus, Polistes, is well studied, we know little of the incidence of parasitism in this group. Here we present details of 45 nest dissections for 4 species: P. dominulus (Christ), P. gallicus (L.), P. stabilinus Richards and P. carnifex (F.) to detail levels of parasitism of colony members by the obligate parasitic group of insects, the Strepsiptera. All 4 species showed evidence of parasitism among immature members. For 3 species, more than 50% of inspected nests were parasitized and the levels of parasitism among brood (larvae and pupae) was very high and did not differ significantly between parasitized nests. One species, P. stabilinus, suffered very low levels of parasitism, which may be related to its habitat choice. The number of parasites per host was positively related to the proportion of infected brood (parasite prevalence) and in some cases reached phenomenally high levels, which casts doubt on previ ously assumed mechanisms of infection for nest-making Hymenoptera, i.e. phoresy. We also document cases of egg parasitism and encapsulation in Polistes nests. Our data show that parasitism levels greatly varied among areas. Finally, the recent debate on the competitive advantage of P. dominulus in its introduced range, USA, has credited an absence of strepsipteran parasites of this species in facilitating its spread. For the first time, we document levels of parasitism for this species in its nature P range and this would appear to corroborate previous claims. We place our work in the context of other studies of parasitism of social insects and posit that the genus Polistes may have much to offer to this field.     

A first attempt at determining larval growth in three Antarctic fish from otoliths and RNA/DNA ratios

Otolith growth and RNA/DNA ratios of larval stages of Notolepis coatsi, Gobionotothen gibberifrons and Trematomus scotti, three Antarctic fish, were studied during early 1998. RNA/DNA ratios were significantly different between the notothenioids and paralepidids (P<0.01), but similar among notothenioids (P>0.05). The ratios were independent of larval total length and water temperature. Recent otolith growth (based on the last five increments) and biochemical indices correlated slightly, but not significantly. N. coatsi inhabited deeper and colder waters of the Weddell Sea and showed less growth than the other species, which are associated with the Antarctic Peninsula. In all three species, some larvae had very good growth rates though most were rather poor. Recent growth indices might allow the detection and back-dating of growth changes in Antarctic larvae, and provide insight into a crucial yet poorly understood life-phase of these fish.     

Competition, predation, and the distributions of four desert anurans

Several studies have shown that larval competition and susceptibility to predation affect distributions of amphibian assemblages across ephemeral and perennial habitats. However, few studies have examined mechanisms affecting distribution patterns and site use of anurans adapted to highly ephemeral habitats. This study examines hypotheses about competition and predation as mechanisms creating non-overlapping patterns of site use in four anurans that breed in highly ephemeral habitats: Scaphiopus couchii (Couch's spade-foot toad), Gastrophryne olivacea (narrow-mouthed toad), Bufo speciosus (Texas toad), and Bufo punctatus (red-spotted toad). These four anurans showed a significantly nonrandom pattern of co-occurrence. Only 12% of 95 ephemeral breeding sites surveyed were occupied by more than one species. We tested the hypotheses that non-overlapping use of breeding sites was due to activity rates of their tadpoles that in turn reflect their competitive ability and susceptibility to predation. Tadpoles of S. couchii were significantly more active and more susceptible to predation than were tadpoles of G. olivacea, B. speciosus, and B. punctatus. The masses of G. olivacea, B. speciosus, and B. punctatus were less when they were reared with S. couchii, demonstrating the possible competitive dominance of S. couchii. These results suggest that the competitive ability of S. couchii may play a role in excluding G. olivacea, B. speciosus, and B. punctatus from very ephemeral breeding sites, and that susceptibility to predation could play a role in excluding S. couchii from breeding sites of longer duration that are more likely to be colonized by aquatic predators.     

Patterns of the <Emphasis Type="Italic">Crematogaster-Macaranga</Emphasis> association: The ant partner makes the difference

Summary. One of the most species-rich ant-plant mutualisms worldwide is the palaeotropical Crematogaster-Macaranga system. Although the biogeography and ecology of both partners have been extensively studied, little is known about the temporal structuring and the dynamics of the association. In this study we compared life-history traits of the specific Crematogaster (Decacrema) partner-ants and followed the development of ant colonies on eight different Macaranga host plant species, from colony founding on saplings to adult trees in a snapshot fashion. We found differences in the onset of alate production, queen number and mode of colony founding in the ant species and examined the consequences of these differences for the mutualism with the host plant. The lifespan of some host plants and their specific ant partners seemed to be well matched whereas on others we found an ontogenetic succession of specific partner ants. The partner ants of saplings or young plants often differed from specific partner ants found on larger trees of the same species. Not all specific Crematogaster species can re-colonize the crown region of adult trees, thus facilitating a change of ant species. Therefore lifespan of the ant colony as well as colony founding behaviour of the different partner ant species are important for these ontogenetic changes. The lifespan of a colony of two species can be prolonged via secondary polygyny. For the first time, also primary polygyny (pleometrosis) is reported from this myrmecophytic system.     

Extreme life history plasticity and the evolution of invasive characteristics in a native ant

Disturbance resulting from urbanization is a leading cause of biotic homogenization worldwide. Native species are replaced with widespread non-native species and ants are among the world’s most notorious invaders. To date, all documented cases of ant invasions involve exotic introduced species that are spread around the world by human-mediated dispersal. I investigated the effect of urbanization on the evolution of invasive characteristics in a native ant species, the odorous house ant, Tapinoma sessile (Say). Colony social structure, life history traits, and the spatial pattern of nest distribution were compared by sampling T. sessile across a gradient of three distinct habitats: natural, semi-natural, and urban. Results demonstrate a remarkable transition in colony social and spatial structure and life history traits between natural and urban environments. In natural habitats, T. sessile colonies are comprised of small, monogyne (single queen), and monodomous (single nest) colonies. In urban areas, T. sessile often exhibit extreme polygyny and polydomy, form large supercolonies, and become a dominant pest. Results also suggest that urban T. sessile colonies may have a negative impact on native ant abundance and diversity. In the natural environment T. sessile coexisted with a wide array of other ant species, while very few ant species were present in the urban environment invaded by T. sessile. Habitat degradation and urbanization can lead to extreme changes in social and spatial colony structure and life history traits in a native ant species and can promote the evolution of invasive characteristics such as polygyny, polydomy, and supercolonial colony structure.     

Odorous House Ants (Tapinoma sessile) as Back-Seat Drivers of Localized Ant Decline in Urban Habitats

Invasive species and habitat disturbance threaten biodiversity worldwide by modifying ecosystem performance and displacing native organisms. Similar homogenization impacts manifest locally when urbanization forces native species to relocate or reinvade perpetually altered habitat. This study investigated correlations between ant richness and abundance in response to urbanization and the nearby presence of invasive ant species, odorous house ants (Tapinoma sessile), within its native region. Surveying localized ant composition within natural, semi-natural, and urban habitat supported efforts to determine whether T. sessile appear to be primary (drivers) threats as instigators or secondary (passengers) threats as inheritors of indigenous ant decline. Sampling 180 sites, evenly split between all habitats with and without T. sessile present, yielded 45 total species. Although urbanization and T. sessile presence factors were significantly linked to ant decline, their interaction correlated to the greatest reduction of total ant richness (74%) and abundance (81%). Total richness appeared to decrease from 27 species to 18 when natural habitat is urbanized and from 18 species to 7 with T. sessile present in urban plots. Odorous house ant presence minimally influenced ant communities within natural and semi-natural habitat, highlighting the importance of habitat alteration and T. sessile presence interactions. Results suggest urbanization releases T. sessile from unknown constraints by decreasing ant richness and competition. Within urban environment, T. sessile are pre-adapted to quickly exploit new resources and grow to supercolony strength wherein T. sessile drive adjacent biodiversity loss. Odorous house ants act as passengers and drivers of ecological change throughout different phases of urban ‘invasion’. This progression through surviving habitat alteration, exploiting new resources, thriving, and further reducing interspecific competition supports a “back-seat driver” role and affects pest management strategies. As demonstrated by T. sessile, this article concludes native species can become back-seat drivers of biodiversity loss and potentially thrive as “metro-invasive” species.     

Interspecific hybridisations in natural populations of ants by example of a regional fauna (Hymenoptera, Formicidae)

Summary: The identification of hybrids by high-precision stereomicroscopy and chorological data is described in detail by example of the ant species Formica bruni and pressilabris. In a synopsis of heterogenous data, the overall hybrid frequency in the ant fauna of Central Europe is estimated. 17 of the 164 ant species of this region are demonstrated and further 2 species strongly suspected to hybridise. The low-frequency hybridisers, showing overall hybrid ratios < 3%, were native elements of the Central European fauna before the onset of human cultivation and experienced direct interspecific contact for longer periods of their natural history. They developed more effective mechanisms of reproductive isolation beginning at the prezygotic level. Extremely high (12-31%) local hybridisation ratios occurred in species that invaded the area after anthropogenic changes in landscape structure. The segregated distribution of invaders and autochthonous species in the precultural period apparently did not impose the need to evolve more effective mechanisms for reproductive isolation. Prezygotic mechanisms in particular are deficient. In local situations, 19% of Lasius jensi matings and 44% of Leptothorax albipennis matings leading to successful nest foundations were matings with heterospecific partners. Signs for a dissolution of interspecific phenotypic differences are not detectable in most of the species. The dispersal of hybrid genotypes is apparently inhibited. Factors that probably stabilise the genomic integrity of hybridizing parent species are: (a) inability to produce hybrid queens (in L. jensi 2 umbratus), (b) aneuploidy of F₁ females with inability to perform a balanced meiosis (in hybrids of L. albipennis with 3 other species), and (c) selection against hybrids in the epigenetic environment of alleles (Leptothorax nylanderi 2 slavonicus). Extreme ratios of heterospecific matings would mean a dangerous drain of genetic material. A mechanism to reduce these losses is postulated, consisting of a 'cleptogamy' (theft of heterospecific sperm) by queens that missed a conspecific male, an establishment of a functioning colony of F₁ hybrid workers, a depression of the rearing or fertility of hybrid queens, and a maximum production of own sons by the queen. This mechanism could explain the persistence of rare species (Lasius jensi or Leptothorax albipennis) in an environment of more populous heterospecific hybridisation partners.     

Spatial and temporal niche partitioning in grassland ants

We examined whether the species composition of a community influences the persistence of larval Ambystoma maculatum in assemblages composed of two larger intraguild predators (A. opacum and A. jeffersonianum) and an alternative prey species (tadpoles of Rana sylvatica). We predicted a priori that A. maculatum would have higher survival in more diverse communities containing alternative species of prey and top predators (A. opacum), the latter of which may lower the abundance of intermediate predators (A. jeffersonianum) via intraguild predation. In a factorial experiment, we manipulated the presence of larval A. opacum, A. jeffersonianum, and R. sylvatica in replicated artificial ponds containing larval A. maculatum. The presence of all three species significantly depressed biomass production in A. maculatum: biomass was highest in ponds lacking the other species and was zero in ponds initially containing all four species. Tadpoles severely reduced the growth of filamentous algae in the ponds. This, in turn, may have affected the abundance of some herbivorous prey of larval salamanders, although this possibility was not tested. The presence of congeneric predators severely restricted the presence in the water column of larval A. maculatum, which otherwise exhibited significant diel patterns of activity in the absence of predators. Together, the presence of tadpoles and a predator-mediated reduction in activity patterns may have limited foraging opportunities for A. maculatum, thus exacerbating the direct impact of predation on survival in this species. These results suggest that diverse assemblages consisting of these particular species may actually inhibit, rather than promote, inclusion of A. maculatum in some communities of pond-breeding amphibians.     

Experimental tests of the mechanism for ant-enhanced growth in an ant-tended lycaenid butterfly

In a previous laboratory study, larvae of the ant-tended lycaenid butterfly Hemiargus isola developed into larger adults when reared with the ant Formica perpilosa than when reared without ants. Ants neither fed butterfly larvae nor significantly delayed developmental duration. We investigated two non-exclusive hypotheses for the mechanism of this effect: larvae tended by F. perpilosa (1) consume more food, and (2) digest the food they consume more efficiently, than those reared without ants. Larvae reared in the laboratory with F. perpilosa ants became significantly heavier adults but produced a significantly lighter fecal mass than their untended counterparts, suggesting that greater food consumption was not the primary mechanism for the higher growth rates of ant-tended larvae. Tended and untended larvae were equally proficient at digesting the contents of pollen (a major natural food source) throughout the tended portion of the life cycle. Taken together, the results suggest that neither greater consumption nor higher assimilation accounts for the larger size of F. perpilosa-tended larvae. We propose that tended larvae may expend less energy than their untended counterparts. Received: 3 January 1997 / Accepted: 18 June 1997     

Three species groups in American trilliums as revealed by compatibility with an Asian species

American trilliums are all diploid and are classified into three groups with regard to compatibility withTrillium kamtschaticum, a single diploid Asian species: (1) Compatible pedicellate-flowered species, i.e.T. erectum, T. gleasoni, andT. vaseyi. (2) Incompatible pedicellate-flowered species, i.e.T. grandiflorum. (3) Incompatible sessile-flowered species, i.e.T. decunbens, T. luteum, T. sessile andT. stamineum. HybridsT. kamtschaticum×T. erectum flowered for the first time 9–10 years after sowing F₁ seeds. The F₁ were completely sterile in pollen and in seed-setting.