Ant exclusion and reproduction of Croton sarcopetalus (Euphorbiaceae)

A total of 20 insect species were observed on the extrafloral nectaries (EFNs) of Croton sarcopetalus. The most frequent ant species were Zacryptocerus sp., Crematogaster brevispinosa, C. scelerata, and Linepithema humile. Ants continuously patrolled the plants taking extrafloral nectar. Experimental data showed that there were no significant differences in either the degree of herbivory or in the reproductive output between control stems (with ants) and treated ones (without ants). We found no significant evidence of protection by ants mediated by EFNs in C. sarcopetalus. It is possible that the plant has other mechanisms than ant protection to prevent herbivore damage (e.g., hairs, latex, chemical defense). As this species occupies the southernmost distribution for the genus, the presence of EFNs in this species may be the remnant of a mutualistic interaction with the ants in tropical regions, where the abundance and diversity of ants and herbivores are comparatively higher. To draw accurate patterns in the genus Croton, further investigations in other species from tropical regions of South America are needed.

Interference competition and scavenging by Crematogaster ants (Hymenoptera: Formicidae) associated with the webs of the social spider Anelosimus eximius (Araneae: Theridiidae) in the central Amazon

Crematogaster cf. victima is a common inhabitant in the sheet web nests of the social spider Anelosimus eximius in the central Amazon basin near Manaus. A number of other ant species were found foraging on the non-sticky webs of A. eximius, but none of these reached the web occupation frequency found in C. cf. victima, nor, with the exception of an unidentified species of Pheidole, did they form satellite nests in the web, as did this species. Many prey which escaped the knock-down threads of the sheet web of A. eximius colonies were captured by ants in the lower web portions which they dominated. Furthermore, prey which were rejected by A. eximius, especially large, heavily sclerotized beetles, were also consumed by this ant. Repeated observations and experiments suggest that C. cf. victima is able to deter A. eximius activity through aerial venom release. Resources lost by A. eximius colonies to ants, especially C. cf. victima, in colonial web area and prey, may pose significant costs and may reduce colony growth.

Revisão taxonômica do grupo limata de Crematogaster Lund, 1831 (Formicidae: Myrmicinae: Crematogastrini)

O gênero Crematogaster é um grupo diverso, com características peculiares e taxonomia complexa. Cerca de 780 espécies e subespécies foram descritas para este gênero, e constantes mudanças são feitas ao longo do tempo devido a sua ampla distribuição geográfica e variedade morfológica. Poucas revisões foram feitas para este gênero, sendo, as existentes, feitas com fauna regional. Neste trabalho, foi feita a revisão taxonômica do grupo de espécies Crematogaster limata, onde foram redescritas sete espécies: Crematogaster brasiliensis Mayr, 1878, C. carinata Mayr, 1862, C. levior Longino, 2003, C. limata Smith, 1858, C. longispina Emery, 1890, C. tenuicula Forel, 1904 e quatro novas espécies foram descritas: Cremastogaster sp. nov. 1, Cremastogaster sp. nov. 2, Crematogaster sp. nov. 3, Crematogaster sp. nov. 4. A variedade Crematogaster longispina var. boliviana foi elevada à categoria de espécie e redescrita. Mapas de distribuição geográfica para todas as espécies e uma chave de identificação específica baseada em operárias são apresentados.

Sperm Bundles in the Seminal Vesicle of the Crematogaster victima (Smith) Adult Males (Hymenoptera: Formicidae)

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Composition of Extrafloral Nectar Influences Interactions between the Myrmecophyte Humboldtia brunonis and its Ant Associates

Ant-plant interactions often are mediated by extrafloral nectar (EFN) composition that may influence plant visitation by ants. Over a 300 km range in the Indian Western Ghats, we investigated the correlation between the EFN composition of the myrmecophytic ant-plant Humboldtia brunonis (Fabaceae) and the number and species of ants visiting EFN. EFN composition varied among H. brunonis populations and between plant organs (floral bud vs. young leaf EFN). In general, EFN was rich in sugars with small quantities of amino acids, especially essential amino acids, and had moderate invertase activity. In experiments at the study sites with sugar and amino acid solutions and with leaf or floral bud EFN mimics, dominant EFN-feeding ants differentiated between solutions as well as between mimics. The castration parasite Crematogaster dohrni (northern study site) was the least selective and did not exhibit any clear feeding preferences, while the largely trophobiont-tending non-protective Myrmicaria brunnea (middle study site) preferred higher sucrose concentrations and certain essential/non-essential amino acid mixtures. The mutualistic Technomyrmex albipes (southern study site) preferred sucrose over glucose or fructose solutions and consumed the leaf EFN mimic to a greater extent than the floral bud EFN mimic. This young leaf EFN mimic had low sugar concentrations, the lowest viscosity and sugar: amino acid ratio, was rich in essential amino acids, and appeared ideally suited to the digestive physiology of T. albipes. This preference for young leaf EFN may explain the greater protection afforded to young leaves than to floral buds by T. albipes, and may also help to resolve ant-pollinator conflicts. The differential response of dominant ants to sugar, amino acids, or solution viscosity suggests that plants can fine-tune their interactions with local ants via EFN composition. Thus, EFN can mediate local partner-choice mechanisms in ant-plant interactions.

Molekulare Studien zur Evolution der Myrmekophytie in der Gattung Macaranga (Euphorbiaceae)

Im Rahmen der vorliegenden Arbeit wurde eine detaillierte phylogenetische Analyse der Ameisenpflanzen aus der Gattung Macaranga (Euphorbiaceae) und ihres verwandtschaftlichen Umfelds mit Hilfe von AFLP-Fingerprinting („amplified fragment length polymorphisms“) sowie vergleichender Analyse von mehreren nichtkodierenden Chloroplasten-DNA-Loci vorgenommen. Anhand dieser Untersuchungen sollten im Wesentlichen die folgenden Fragen geklärt werden: (1) Wie stellen sich die Verwandtschaftsverhältnisse zwischen den myrmekophytischen Macaranga-Sektionen Pachystemon, Winklerianae und Pruinosae dar? (2) Wie sind die einzelnen Arten dieser Sektionen miteinander verwandt? (3) Wie oft ist die Lebensweise ”Myrmekophytie” unabhängig voneinander entstanden? Gibt es Hinweise auf Reversionen? (4) Wo liegt genealogisch und auch geographisch der Ursprung der Symbiose zwischen den myrmekophytischen Macaranga-Arten und ihren Partnerameisen? (5) Welche Bedeutung spielen koevolutive Entwicklungen für das Macaranga-Crematogaster-Symbiosesystem? Ist Myrmekophytie im Sinne einer Schlüsselinnovation (Givnish, 1997) als Stimulus für eine adaptive Radiation zu betrachten? (1) Für die AFLP-Analyse wurden 108 Proben aus 43 Macaranga-Arten und 5 unbeschriebenen Morphospezies in die phylogenetische Untersuchung einbezogen. Auf der Basis von 426 Merkmalen wurden Phänogramme sowie Kladogramme rekonstruiert. Zur statistischen Absicherung wurden Bootstrap-Analysen durchgeführt und im Falle der Kladogramme darüber hinaus der „consistency“-Index bestimmt. Die AFLP-Datensätze wurden zusätzlich einer Hauptkomponentenanalyse unterzogen. Mit Hilfe der verschiedenen Untersuchungsmethoden konnten weitgehend übereinstimmende Gruppierungen bzw. evolutive Linien identifiziert werden. Die Sektionen Pachystemon und Pruinosae bilden eine jeweils gut gestützte monophyletische Gruppe. Beide sind vermutlich Schwestergruppen und damit gleich alt. Für die Monophylie der nur aus zwei Arten bestehenden Sektion Winklerianae ergab sich keine Unterstützung. Die Arten der Sektion Pruinosae sind im AFLP-Baum gut aufgelöst. Die nicht myrmekophytische M. gigantea sitzt dabei an der Basis und ist Schwestergruppe zu den myrmekophytischen Arten. Innerhalb der Sektion Pachystemon wurden mit Hilfe der AFLP-Analyse vier gut gestützte Gruppen identifiziert. Für die puncticulata-Gruppe konnte hier erstmals auf molekularer Ebene eine Zugehörigkeit zur Sekt. Pachystemon nachgewiesen werden. Der von Davies (2001) vorgenommene Ausschluss von M. recurvata aus der Sekt. Pachystemon konnte bestätigt werden. Die Verwandtschaftsbeziehungen einzelner Arten zueinander sind in den AFLP-Bäumen nicht aufgelöst. (2) Für die vergleichende Chloroplasten-Sequenzierung wurden nach Maßgabe der Sequenzvariabilität in Testsequenzierungen die Bereiche atpB-rbcL und psbI-trnS für die phylogenetische Untersuchung ausgewählt. Für die Chloroplasten-Phylogenie wurden für jeden Locus mehr als 100 Sequenzen analysiert. Neben 29 Pachystemon-Arten inkl. vier unbekannter Morphospezies, acht Pruinosae-Arten inkl. eines möglichen Hybriden und den beiden Arten der Sekt. Winklerianae wurden 22 weitere Macaranga- und 10 Mallotus-Arten in die Untersuchung einbezogen. Zwischen den südostasiatischen Arten bestanden nur geringe Sequenzunterschiede. Maximum-Parsimonie-Kladogramme wurden rekonstruiert und die Sequenzen der beiden Loci wurden sowohl einzeln, als auch kombiniert ausgewertet. Indels wurden kodiert und als separate Merkmalsmatrix an die Sequenzdaten angehangen. Innerhalb von Macaranga konnten nur wenige abgesicherte Gruppen identifiziert werden. Deutlich war die Zusammengehörigkeit der afrikanischen Arten und ihr Entstehung aus den südostasiatischen Arten. Die von Davies (2001) der Sektion Pruinosae zugeordnete M. siamensis steht deutlich außerhalb dieser Sektion. Die Arten der Sektionen Pruinosae, Pachystemon und Winklerianae bilden keine statistisch gesicherten monophyletischen Gruppen. Während der Pilotstudien stellte sich heraus, dass die Chloroplastensequenzen nahe verwandter Arten der Sektion Pachystemon weniger nach den Artgrenzen, sondern vielmehr nach geographischen Kriterien gruppierten. (3) Es wurde daher zusätzlich eine phylogeographische Analyse der Chloroplasten-Sequenzen auf der Basis eines Parsimonie-Netzwerks durchgeführt. Neben dem atpB-rbcL-Spacer und einer Teilsequenze des psbI-trnS-Locus (ccmp2) wurde dafür zusätzlich der ccmp6-Locus (ein Abschnitt des ycf3-Introns) sequenziert. Die phylogeographische Untersuchung wurde mit 144 Proben aus 41 Macaranga-Arten durchgeführt. Darin enthalten waren 29 Arten (inkl. vier Morphospezies) mit 112 Proben der Sektion Pachystemon, sieben 7 Arten (inkl. eines potentiellen Hybriden) mit 22 Proben der Sekt. Pruinosae und zwei Arten mit 5 Proben der Sekt. Winklerianae. Das voll aufgelöste statistische Parsimonie-Netzwerk umfasste 88 Haplotypen. Die Sektionen Pachystemon und Pruinosae bilden jeweils eine monophyletische Gruppe. Das geographische Arrangement der Haplotypen unabhängig von der Artzugehörigkeit könnte durch Introgression und/oder „lineage sorting“ bedingt sein. Mit Hilfe der im Rahmen dieser Arbeit gewonnenen Ergebnisse kann man davon ausgehen, dass eine enge Ameisen-Pflanzen-Symbiose innerhalb der Gattung Macaranga mindestens drei-, möglicherweise viermal unabhängig voneinander entstanden ist Eine Reversion hat mindestens einmal, möglicherweise häufiger in der bancana-Gruppe stattgefunden. Ob sich die Symbiose dabei in Westmalaysia oder in Borneo entwickelt hat, kann man nicht sicher sagen; Ob und inwieweit die große Artenzahl in der bancana-Gruppe als eine Folge der Myrmekophytie anzusehen ist, bleibt zunächst offen. Wesentliche Teile der vorliegenden Arbeit liegen bereits in publizierter Form vor (AFLP-Analyse: Bänfer et al. 2004; Chloroplasten-Analyse: Vogel et al. 2003; Bänfer et al. 2006).

Thorn-dwelling ants provide antiherbivore defence for camelthorn trees, Vachellia erioloba, in Namibia

Ants are widely employed by plants as an antiherbivore defence. A single host plant can associate with multiple, symbiotic ant species, although usually only a single ant species at a time. Different plant-ant species may vary in the degree to which they defend their host plant. In Kenya, ant–acacia interactions are well studied, but less is known about systems elsewhere in Africa. A southern African species, Vachellia erioloba, is occupied by thorn-dwelling ants from three different genera. Unusually, multiple colonies of all these ants simultaneously and stably inhabit trees. We investigated if the ants on V. erioloba (i) deter insect herbivores; (ii) differ in their effectiveness depending on the identity of the herbivore; and (iii) protect the tree against an important herbivore, the larvae of the lepidopteran Gonometa postica. We show that experimental exclusion of ants leads to greater levels of herbivory on trees. The ants inhabiting V. erioloba are an effective deterrent against hemipteran and coleopteran, but not lepidopteran herbivores. Defensive services do not vary among ant species, but only Crematogaster ants exhibit aggression towards G. postica. This highlights the potential of the V. erioloba–ant mutualism for studying ant–plant interactions that involve multiple, simultaneously resident thorn-dwelling ant species.

Arboreal thorn-dwelling ants use domatia morphology to partition nest sites on the savanna ant-plant, Vachellia erioloba

Nest site selection in arboreal, domatia-dwelling ants, particularly those coexisting on a single host plant, is little understood. To examine this phenomenon we studied the African savannah tree Vachellia erioloba, which hosts ants in swollen-thorn domatia. We found four ant species from different genera (Cataulacus intrudens, Tapinoma subtile, Tetraponera ambigua and an unidentified Crematogaster species). In contrast to other African ant plants, many V. erioloba trees (41 % in our survey) were simultaneously co-occupied by more than one ant species. Our study provides quantitative field data describing: (1) aspects of tree and domatia morphology relevant to supporting a community of mutualist ants, (2) how ant species occupancy varies with domatia morphology and (3) how ant colony size varies with domatia size and species. We found that Crematogaster sp. occupy the largest thorns, followed by C. intrudens, with T. subtile in the smallest thorns. Thorn age, as well as nest entrance hole size correlated closely with ant species occupant. These differing occupancy patterns may help to explain the unusual coexistence of three ant species on individual myrmecophytic trees. In all three common ant species, colony size, as measured by total number of ants, increased with domatia size. Additionally, domatia volume and species identity interact to predict ant numbers, suggesting differing responses between species to increased availability of nesting space. The proportion of total ants in nests that were immatures varied with thorn volume and species, highlighting the importance of domatia morphology in influencing colony structure.

Ant diversity (Hymenoptera: Formicidae) and predation by ants on the different stages of the sugarcane borer life cycle Diatraea saccharalis (Lepidoptera: Crambidae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Contrasting responses to induction cues by ants inhabiting Maieta guianensis (Melastomataceae)

As duas formigas que mais comumente se associam com a mirmecofita de sub-bosque Maieta guianensis (Melastomataceac) respondem de maneiras diferentes a simulac (o) over tilde es experimentais de herbivoria. Enquanto Pheidole minutula & induzida tanto por danos fisicos e extratos de tecido foliar, Crematogaster laevis 6 induzida apenas pelos danos fisicos. Sugerimos que tal variac (a) over tildeo interespecifica em respostas induzidas e comum e pode influenciar a qualidade das defesas contra herbivoros que as formigas oferecem as plantas que elas ocupam.

Interspecific variation in the defensive responses of obligate plant-ants: experimental tests and consequences for herbivory

The aggressive behavior of ants that protect plants from herbivores in exchange for rewards such as shelter or food is thought to be an important form of biotic defense against herbivory, particularly in tropical systems. To date, however, no one has compared the defensive responses of different ant taxa associated with the same plant species, and attempted to relate these differences to longer-term efficacy of ant defense. We used experimental cues associated with herbivory-physical damage and extracts of chemical volatiles from leaf tissue-to compare the aggressive responses of two ant species obligately associated with the Amazonian myrmecophyte Tococa bullifera (Melastomataceae). We also conducted a colony removal experiment to quantify the level of resistance from herbivores provided to plants by each ant species. Our experiments demonstrate that some cues eliciting a strong response from one ant species elicited no response by the other. For cues that do elicit responses, the magnitude of these responses can vary interspecifically. These patterns were consistent with the level of resistance provided from herbivores to plants. The colony removal experiment showed that both ant species defend plants from herbivores: however, herbivory was higher on plants colonized by the less aggressive ant species. Our results add to the growing body of literature indicating defensive ant responses are stimulated by cues associated with herbivory. However, they also suggest the local and regional variation in the composition of potential partner taxa could influence the ecology and evolution of defensive mutualisms in ways that have previously remained unexplored.

Association of the stilt bug Jalysus ossesae Henry (Hemiptera : Heteroptera : Berytidae) with myrmecophytic plants of the genus Maieta (Melastomataceae) in an upland forest area in central Amazon, Brazil

Maieta guianensis Aubl. and M. poeppigii Mart. ex. Triana (Melastomataceae) are among the most common myrmecophytic plants in the Amazonian forest understory. These myrmecophytes are colonized exclusively by the ants Pheidole minutula Mayr or Crematogaster sp. and usually host two other arthropods, the spider Faiditus subflavus Exline and Levi and the recently described stilt bug Jalysus ossesae Henry. In this study, the association between J ossesae and the myrmecophytic plants M. guianensis and M. poeppigii in an upland forest area in central Amazon, Brazil, is described. The presence of the stilt bugs on M. guianensis and M. poeppigii and on plants around these myrmecophytes was recorded in five transects. The number and position of the stilt bugs on the leaf surface (upper or lower) and leaf type (with or without domatia) of these myrmecophytes, as well as their behavioral acts, were recorded. Jalysus ossesae was found only on the myrmecophytic plants M. guianensis and M. poeppigii. The stilt bug occurred at similar frequencies on M. guianensis and M. poeppigii, and the number of leaves significantly influenced the presence and number of stilt bugs on these myrmecophytes. Feeding, agonistic interaction between males, and mating were observed. Our data indicate that J. ossesae uses the myrmecophytes M. guinanensis and M. poeppigii as reproductive and foraging sites.

Predaceous ant fauna in new sugarcane fields in the state of São Paulo, Brazil

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)