The triple alliance: how a plant-ant, living in an ant-plant, acquires the third partner, a scale insect

Summary. Mutualistic associations between ants and plants often involve a third partner, scale insects (Hemiptera, Coccoidea). In southeast Asia, plant-ants of the genus Cladomyrma live together with coccoids in hollowed twigs of a wide range of ant-plants (myrmecophytes). Established colonies never lack sap-sucking scale insects and the ants appear to be dependent on the honeydew excretions of their trophobionts. Acquisition of scale insects thus seems to be an important step in the successful establishment of a new colony on a host plant. Coccoids may either be transported by a foundress ant on her nuptial flight (vertical transmission via co-dispersal) or ant and coccoid disperse on their own and associate anew during colony initiation (horizontal transmission). Here we test the assumption that founding gynes of Cladomyrma do not transport scale insects during their dispersal flight, and we determine when and how trophobionts are acquired. The results obtained in this study show that although coccoids are not carried by foundresses, their numbers within founding chambers of Cladomyrma increased with the founding stage. In all Cladomyrma species, colony-founding is claustral and the entrance hole into the nest chamber is first plugged with pith debris and later grows over by wound callus. However, access of trophobionts into hollow twigs is facilitated either by additional small openings (secondary slits) produced by the foundress ant after hollowing out a nest chamber or by the maintenance of an orifice at the entrance hole during the founding period. Immature pseudococcids experimentally placed onto twigs close to a Cladomyrma foundation readily entered these slits. Exclusion experiments suggest that foundresses of C. petalae are less successful in rearing their first offspring if entry of coccoids into founding chambers is prevented.Received 28 September 2003; revised 9 May and 11 October 2004; accepted 9 November 2004.     

Host nest usurpation and colony foundation in the European amazon ant,Polyergus rufescens Latr. (Hymenoptera: Formicidae)

Summary The socially parasitic mode of founding new colonies by queens of the European amazon antPolyergus rufescens was analysed in the laboratory. Newly-mated females of this obligatory slave-maker were individually introduced into queenright and queenless artificially established colonies of bothFormica cunicularia (the slave present in the natal dulotic nest) andF. rufibarbis (another potentialServiformica host). Particular attention was devoted to the behavioural patterns displayed by these young queens during the usurpation phases. Our observations, supported also by video-taping, show that the slave-making female, before laying her eggs, must penetrate the host colony, kill the resident queen, become accepted by the adult workers and appropriate the host brood. The parasite was almost always adopted in the colonies ofF. cunicularia, whereas in the presence ofF. rufibarbis it was generally killed in a short time. The failure in the attempt of usurping the colonies ofF. rufibarbis is discussed in relation to the host specificity typical of this slave-maker. Finally, egg-laying byPolyergus successful usurpers, the subsequent eclosion of the brood, and its complete social integration in the newly-established mixed colonies were also recorded.     

A new species of Chamaecrista Moench (Caesalpiniaceae) from interior Bahia,Brazil

Chamaecrista arboae, a species of sect.Absus ser.Absoideae newly discovered on Chapada Diamantina in interior Bahia, Brazil, is described, figured, and discussed.     

Queen size dimorphism in the ant Tetramorium moravicum (Hymenoptera, Formicidae): Morphometric, molecular genetic and experimental evidence

Summary. By combining different methods we evaluate whether the ant Tetramorium rhenanum is specifically separated from T. moravicum or whether it is a conspecific microgyne form. High-precision morphometry shows a clear difference in queen size. Sequence comparison of 1031 bp of COI reveals that T. rhenanum falls into a clade with T. moravicum, which is significantly separated from T. forte and T. chefketi. T. rhenanum shares at least two haplotypes with T. moravicum and is considered as a junior synonym. Sexual behaviour and colony foundation experiments corroborate conspecificity. The queen dimorphism is discussed in the context of social parameters such as queen number and reproductive strategy.Received 10 June 2004; revised 28 October 2004; accepted 12 November 2004.     

Colony structure of a bamboo-dwellingTetraponera sp. (Hymenoptera: Formicidae: Pseudomyrmecinae) from Malaysia

Summary The colony structure of the bamboo-inhabiting SE-Asian pseudomyrmecine antTetraponera sp. PSW-80 nearattenuata F. Smith was investigated at the Ulu Gombak Field Studies Centre, Selangor, West-Malaysia. Based on the dissection of 54 stem internodes from 12 different culms of the large bambooGigantochloa scortechinii Gamble and on the mapping of three colonies, the following demographic characteristics emerge. The colonies are monogynous but highly polydomous (at least up to 36 internodes and up to 9 stems occupied) and very populous for a pseudomyrmecine not involved in an ant-plant mutualism. One completely censused colony had 6953 adult workers and 2079 alates (adults plus pupae). The single queen suppresses gyne development in her own nest and, to a lesser extent, in other nests within the same stem. The overall numerical sex ratio was 0.961 (females:males), the investment sex ratio, 2.931, i.e., almost exactly the 31 ratio expected for a monogynous outbred hymenopteran in which the colony queen also produces all the male offspring. Brood is distributed to all other nest chambers from the queenright chamber. The symbiotic pseudococcids (Kermicus wroughtoni Newstead) are present in all inhabited internodes, with small early instar individuals prevailing numerically by far over the larger stages. The rieht well secluded honeydew supply within the internode and the efficient architectural protection provided by the internode wall (access usually only through a 2 × 3 mm-hole) allowT. sp. PSW-80 to reach an unusually large colony size without being an aggressive and protective plant mutualist like other members of its subfamily with similar demographic features.     

Two new species ofPhyllagathis related toP. tuberculata (Melastomataceae) from Peninsular Malaysia

Phyllagathis tuberculataKing and two closely allied new species,P. magnificaA. Weber andP. stoneiA. Weber, are described and illustrated. Within the genus, these species form a distinct and isolated group which is restricted to Peninsular Malaysia.P. tuberculata occurs in Perak (probably confined to G. Bujang Melaka),P. magnifica andP. stonei are found in the mountains on the Pahang/Selangor border (Genting Highlands, Gombak valley). The distinctive characters of the three species are listed and some general information relating to inflorescence morphology, tubercle anatomy, fruit structure and seed dispersal is provided.     

Two new Artotrogids (Copepoda: Siphonostomatoida) from Madeira Island,Portugal

Two new artotrogids are recorded from Madeira Island. A new species of Cryptopontius, associated with algae, differs from its congeners in the number of antennular segments, setation of the antennal exopod, setation and length of the maxillule lobes, and setation of both rami of the first and fourth legs. The second new artotrogid belongs to Dyspontius.It was associated with Gerardia savagliae Lacaze-Duthiers, 1864 (Cnidaria: Zoanthidea) and differs from its congeners in cephalothorax shape, siphon length and setation of the maxillule lobes.     

Two new species ofInocybe, sectionMarginatae (Agaricales, Cortinariaceae) from Japan

Two new species of the sectionMarginatae are described, namely,Inocybe populea sp. nov. andI. leptoclada sp. nov., from western Japan and eastern Japan, respectively.Inocybe leptoclada is assigned to sectionMarginatae, even though it lacks a marginate bulb.Inocybe populea is placed in the sectionMarginatae because of the presence of a marginate (often indistinct) bulbous base of stipe and the caulocystidia throughout. The discoveries ofI. leptoclada andI. populea provide further evidence that sectionMarginatae and sectionInocybe [=Cortinatae] are poorly defined.     

Two new species and one new variety of Oudemansiella (Agaricales) from Japan

Two new species and one new variety of Oudemansiella (Agaricales) from Japan are described and illustrated: (1) Oudemansiella latilamellata sp. nov. (subgenus Xerula, section Radicatae) has relatively broad lamellae and broadly ellipsoid spores with a subacute apex; (2) Oudemansiella rhodophylla sp. nov. (subgenus Xerula, section Radicatae), growing in deciduous forests of Fagus crenata, Quercus crispula, etc., is characteristic in having lamellae tinted reddish; and (3) Oudemansiella amygdaliformis var. bispora var. nov. (subgenus Xerula, section Albotomentosae) is distinguished from the type variety by having two-spored basidia and grows in evergreen oak forests (Castanopsis sieboldii, Quercus glauca, etc.) or bamboo groves.     

Two new species ofConsolida (Ranunculaceae) from the Flora Iranica area

Two species ofConsolida are described as new:C. lorestanica is distributed in W. Iran (Lorestan), andC. kandaharica is endemic to S. Afghanistan.Dedicated to Hofrat Prof. DrK. H. Rechinger on the occasion of this 80th birthday.     

Two new species of the genusAmblyeleotris (Pisces: Gobiidae) from the Ryukyu Islands,Japan

Two new shrimp-associated gobies,Amblyeleotris yanoi sp. nov. andA. masuii sp. nov. are described on the basis of specimens from Iriomote-jima Island and Okinawa-jima Islands, Okinawa Prefecture, Japan.A. yanoi is distinguished from other members of the genus by the combination of the following characters: 14 anal fin soft rays, 19 pectoral fin rays, 97–103 longitudinal scales, a candle flame-shaped marking on the caudal fin, a very low membrane connecting the pelvic fins and absence of a ventral frenum.A. masuii differs from all other congeners by having 92–97 longitudinal scales, the length of the interpelvic connecting membrane relative to the longest pelvic fin ray (0.43–0.66), black blotches on the sides of the chin, and blue spots on the opercle and preopercle.     

Integration raids in the Amazon ant Polyergus rufescens (Hymenoptera, Formicidae)

Summary. Groups of enslaved Formica fusca workers from mixed colonies of Polyergus rufescens with numerous slave workforce tend to split off and found small and almost homospecific nests around the main nest, with at least some of them connected with the latter with underground passages. Their inhabitants are able, at least temporarily, to adopt young F. fusca gynes. P. rufescens invades these satellite nests in a manner similar to the normal slave raids, and carries the slaves back to the main nest. The supposed evolutionary cause of this behaviour is to keep integrity of mixed colonies and prevent possible emancipation of slaves.Received 18 August 2004; revised 27 September 2004; accepted 11 October 2004.     

Two new species of Serjania (Sapindaceae) from southeastern Brazil

One new species ofSerjania (S. erythrocaulis) belonging to theS. paradoxa group ofSerjania sect.Eurycoccus is described and illustrated bringing the total number of species in this group to eight. A second new species,Serjania divaricocca, belonging to sect.Serjania is also described and illustrated.

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Resumen  Una especie neuva deSerjania (S. erythrocaulis) perteneciente al grupoS. paradoxa de la secciónEurycoccus es descrita e ilustrada. Con esta especie, el numero total de especies conocidas para este grupo aumenta a ocho. Una segunda nueva especie,Serjania divaricocca, perteneciente a la secciónSerjania es también descrita e ilustrada.

     

Two new species of the orientalis group of Rheocricotopus (Psilocricotopus) from China (Diptera: Chironomidae)

Rheocricotopus (Psilocricotopus) calviculus sp. n. and R. (P.) villiculus sp. n. from China are described as male imagines. The species are closely related to the previously described R. (P.) orientalis Wang forming a new group, the orientalis group, which eventually may deserve the rank of subgenus or even genus. The group is distinguished by lacking a humeral pit, numerous sensilla clavata on third palpomere, cuneiform wings, no or few setae on squama, long legs, banded abdomen, very long inferior volsella, gonocoxite with apical extensions, and gonostylus with a median triangular projection.     

Two new species of Mendoncia (Acanthaceae) from Bahia, Brazil

Two new species, Mendoncia bahiensis and Mendoncia blanchetiana, of the subgenus Bremekampia are described and illustrated. A key to four species represented in Bahia and commentary on their taxonomic affinities are provided.     

Nest structure and colony cycle of the Allegheny mound ant,Formica exsectoides Forel (Hymenoptera: Formicidae)

Summary The nest structure and colony cycle of a population of Allegheny mound ants,Formica exsectoides, were examined in central Michigan. The dispersion pattern of mounds was random. Nest structure and presence of brood were primarily determined by excavation of twenty-three nests over three intervals from June through September. Additional excavations of five nests in 1990 and ten nests in 1991 provided further details on nest structure and colony cycle. Most galleries occurred within the mound and upper 30 cm of soil, but some activity reached depths of 100 to 270 cm. Depth of nests showed little correlation with external measurements of height and diameter. Immature stages were recovered from two strata: the upper 20 cm of nest and mound and the lowest nest depths. Alate sexual forms were found in or near the mound in July, and numerous dealate queens were collected in September from peripheral galleries near the soil surface.     

Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.