Phylogeny of the tribe Colletieae, Rhamnaceae

A cladistic analysis based on 63 morphological characters was carried out on the tribe Colletieae including two presumed closely related genera, Geanothus and Noltea as outgroupS. In addition to a parsimony analysis of the equally weighted characters, analyses investigating the effects of character weighting, removal of a presumed hybrid species as well as the impact of uncertainly scored characters were undertaken. In all analyses Noltea was placed as sister group to a well supported monophyletic Colletieae. Nineteen different in group topologies were found, with the additional analyses mainly supporting two of them. Within the Colletieae a basal dichotomy divides Trevoa and Retanilla from the remainder of the tribe. While the Trevoa-Retanilla clade is fully resolved, the second lacks detailed resolution. Within this clade the Colletia species from a well supported monophyletic group, while monophyly of the disjunct genus Discaria could not be confirmed.     

Reproductive Biology of the Andean Shrub Discaria nana (Rhamnaceae)

Abstract: The widely-held view that the frequency of self-compatibility increases at higher elevations has been questioned for communities of the southern Andes. The study of pollination biology of obligate outcrossers with wide altitudinal range may provide clues on how plants can remain reproductively successful in increasingly hostile environments without resorting to selfing. I studied the phenology, pollinator assemblages, breeding system and maternal success of the perennial entomophile outcrosser, Discaria nana, across a 1200-m elevation span in the Andes of Mendoza and Neuquén, Argentina (34° - 37°S). D. nana behaved as self-incompatible throughout the studied gradient. At the higher site proportionally fewer flowers were pollinated, stigmatic loads were smaller and number of visitor species was lower than at low altitude; however, natural fruit set was 3.6 times higher and seed set 3.3 times higher. As a non-exclusive explanation, it is proposed that increased maternal success reflected better pollination quality at the higher site, to which several factors seemed to contribute. At higher altitude D. nana a) exhibited mutualisms with a high proportion of the available pollen vectors, b) had a higher rate of specialized pollinators (Lepidoptera, obligate nectar feeders) to unspecialized ones (Diptera, pollen and mixed feeders) in the visitor assemblage which would reduce ineffective (i.e. self) pollination because nectar feeders visited both male and female phase flowers and pollen feeders favoured male phase flowers, c) had an extended floral longevity (especially the female phase), and d) showed an increased floral display (as number of flowers per unit plant area). Differential rewarding through more concentrated nectar may explain the change in assemblage composition. The flowers' reverse herkogamy (i.e. the location of the stigma below the anthers) seemed to enhance the beneficial effects of the abundance of more effective pollinators.     

Time course of nodule development in the Discaria trinervis (Rhamnaceae) –Frankia symbiosis

The time course of initiation and development of root nodules was investigated in the South American actinorhizal shrub Discaria trinervis (Rhamnaceae). A local strain of Frankia (BCU110501) which was isolated from D. trinervis nodules, was used as inoculum. Inoculated seedlings were periodically studied under the light microscope after clearing with aqueous NaClO. In parallel, semithin and ultrathin sections were analysed by light and electron microscopy. Infection by Frankia BCU110501 involved intercellular penetration among epidermal and cortical root cells. Nodule primordia were detected from 6 d after inoculation, while bacteria were progressing through intercellular spaces of the outer layers of cortical cells. Invasion of host cells by the symbiont occurred 7–9 d after inoculation, and hypertrophy of the primordium cells was associated with Frankia penetration. Root hairs were not deformed during the early events of nodule formation. From 13 to 16 d after inoculation, the proximal cellular zone of the primordia behaved differently from the other tissues after NaClO treatment and remained darkly pigmented. At the same time, differentiation of Frankia vesicles started to occur inside already infected cells. By 16 d after inoculation, spherical vesicles of BCU110501 were homogeneously distributed in the host cells. These vesicles were septate and surrounded by void space. Frankia spores or sporangia were not observed in the nodule tissue. This study has clarified the mode of Frankia penetration in D. trinervis , one of the Rhamnaceae which also includes Ceanothus . The events involved in infection, nodule induction, host-cell infection and vesicle differentiation have been characterized and identified as time-segregated developmental processes in the ontogeny of D. trinervis root nodules.     

Reproductive biology of Colletia spinosissima (Rhamnaceae) in Argentina

 We studied the reproductive behavior of Colletia spinosissima Gmel. at six sites in two regions of Argentina during the 1998–2000 flowering seasons. Flowering takes place in winter. The flowers are homogamous, fragrant, and entomophilous. There was evidence of self-incompatibility and cross-incompatibility between nearby partners. Fruit set of open-pollinated flowers (13.3%) seemed pollen-limited, since low fecundity was consistently associated with small stigmatic pollen loads and with low numbers of successful pollen tubes, and fecundity increased with abundance of efficient pollinators. Reproductive success (percent ovules becoming viable seeds) was 3.6. It is hypothesized that homogamy was established in Colletia because in an impoverished pollination environment – such as the one likely prevailing when this lineage evolved – achieving pollen export and receipt in a single pollinator visit has selective advantages. A large stigma and an extragynoecial compitum would compensate for the homogamy-driven, increased pollen-stigma interference. Received March 8, 2001 Accepted July 19, 2001     

The inflorescences structure of Cyperus L. section Luzuloidei Kunth (Cyperaceae)

In the present work the inflorescences of 12 species of Cyperus sect. Luzuloidei Kunth sensu Kükenthal were analyzed using the methodology and terminology of Troll's school. All the inflorescences studied are polytelic (indeterminate). The inflorescences are anthela-like or capitate, and can be terminal or pseudolateral. Below the main florescence a paracladial zone is present. In this zone three types of branching were observed, i.e. normal, accessory-axillar and prophyllar branching, whose position vary among the different species. In the paracladial zone a subzone of long paracladia and a subzone of short paracladia were observed. In the short paracladial subzone the spikelets are clustered in fascicles, which can be serial, prophyllar or mixed. Variation in length and position of pherophylls, length of prophyll, number and degree of branching were noticed. The systematic value of the inflorescences in the section Luzuloidei is discussed.     

Phylogenetic analysis of the tribe Bovini (Mammalia: Artiodactyla)

A matrix of 57 (mainly cranial) characters and 32 taxa of fossil and Recent Bovini (buffaloes, bison and cattle) has been analysed using the parsimony program HENNIG86. Among the best established results are the exclusion of Parabos , polyphyly of Leptobos , monophyly of the Bos sensu lato + buffaloes group (a clade including all Recent Bovini), probable monophyly of the Bubalina and Syncerina, and the close relationship between bison and yak. Some other interesting questions raised include the apparent absence of a close link between Pliocene African bovines (except "Leptobos" syrticus ) and later African buffaloes, and the possible monophyly of Pelorovis oldowayensis+Bos sensu lato .     

Quantitative analysis of the phenotypic variability of shoot architecture in two grapevine (Vitis vinifera) cultivars

BACKGROUND AND AIMS: Plant architecture and its interaction with agronomic practices and environmental constraints are determinants of the structure of the canopy, which is involved in carbon acquisition and fruit quality development. A framework for the quantitative analysis of grapevine (Vitis vinifera) shoot architecture, based on a set of topological and geometrical parameters, was developed for the identification of differences between cultivars and the origins of phenotypic variability. METHODS: Two commercial cultivars ('Grenache N', 'Syrah') with different shoot architectures were grown in pots, in well-irrigated conditions. Shoot topology was analysed, using a hidden semi-Markov chain and variable-order Markov chains to identify deviations from the normal pattern of succession of phytomer types (P0-P1-P2), together with kinematic analysis of shoot axis development. Shoot geometry was characterized by final internode and individual leaf area measurements. KEY RESULTS: Shoot architecture differed significantly between cultivars. Secondary leaf area and axis length were greater for 'Syrah'. Secondary leaf area distribution along the main axis also differed between cultivars, with secondary leaves preferentially located towards the basal part of the shoot in 'Syrah'. The main factors leading to differences in leaf area between the cultivars were: (a) slight differences in main shoot structure, with the supplementary P0 phytomer on the lower part of the shoot in 'Grenache N', which bears a short branch; and (b) an higher rate and duration of development of branches bearing by P1-P2 phytomers related to P0 ones at the bottom of the shoot in 'Syrah'. Differences in axis length were accounted for principally by differences in individual internode morphology, with 'Syrah' having significantly longer internodes. This trait, together with a smaller shoot diameter, may account for the characteristic 'droopy' habit of 'Syrah' shoots. CONCLUSIONS: This study highlights the architectural parameters involved in the phenotypic variability of shoot architecture in two grapevine cultivars. Differences in primary shoot structure and in branch development potential accounted for the main differences in leaf area distribution between the two cultivars. By contrast, shoot shape seemed to be controlled by differences in axis length due principally to differences in internode length.     

Differences in proleptic and epicormic shoot structures in relation to water deficit and growth rate in almond trees (Prunus dulcis)

Background and Aims

Shoot characteristics differ depending on the meristem tissue that they originate from and environmental conditions during their development. This study focused on the effects of plant water status on axillary meristem fate and flowering patterns along proleptic and epicormic shoots, as well as on shoot growth rates on ‘Nonpareil’ almond trees (Prunus dulcis). The aims were (1) to characterize the structural differences between proleptic and epicormic shoots, (2) to determine whether water deficits modify shoot structures differently depending on shoot type, and (3) to determine whether shoot structures are related to shoot growth rates.

Methods

A hidden semi-Markov model of the axillary meristem fate and number of flower buds per node was built for two shoot types growing on trees exposed to three plant water status treatments. The models segmented observed shoots into successive homogeneous zones, which were compared between treatments. Shoot growth rates were calculated from shoot extension measurements made during the growing season.

Key Results

Proleptic shoots had seven successive homogeneous zones while epicormic shoots had five zones. Shoot structures were associated with changes in growth rate over the season. Water deficit (1) affected the occurrence and lengths of the first zones of proleptic shoots, but only the occurrence of the third zone was reduced in epicormic shoots; (2) had a minor effect on zone flowering patterns and did not modify shoot or zone composition of axillary meristem fates; and (3) reduced growth rates, although patterns over the season were similar among treatments.

Conclusions

Two meristem types, with different latency durations, produced shoots with different growth rates and distinct structures. Differences between shoot type structure responses to water deficit appeared to reflect their ontogenetic characteristics and/or resource availability for their development. Tree water deficit appeared to stimulate a more rapid progression through ontogenetic states.     

Phylogenetic analysis of the pearlfish tribe Carapini (Pisces: Carapidae)

Fishes of the tribe Carapini (Encheliophis and Carapus) share a noteworthy peculiarity: they shelter in holothurian echinoderms or bivalve hosts. Some species are considered parasitic, others commensal. This study focuses on the phylogeny of the tribe, using two other Carapidae species as an outgroup (Snyderidia canina and Onuxodon fowleri). Insofar as possible, the selected anatomical and behavioural characters where chosen in an ecomorphological perspective, as features that could be responses to various lifestyle‐related constraints. Our character selection also took into account the fact that some features are (presumably) linked. Such features were grouped together as a single trait to avoid their overvaluation. This methodology enabled commensals to be separated from parasites, the former belonging to Carapus and the latter to Encheliophis. Carapus species reflect in their morphology the constraints imposed by a diet of hard, mobile, elusive prey, showing predator‐type features: a strong dentition, a wide mouth opening, a robust food intake apparatus. On the other hand, the endoparasitic Encheliophis species show a generally weaker buccal apparatus and narrow mouth opening, in relation to the different constraints of their lifestyle where the diet constraints are less pronounced: they eat body parts of their host. Changes in both generic diagnoses are proposed and three species are transferred from Encheliophis to Carapus.     

The architecture of Mourera fluviatilis (Podostemaceae): developmental morphology of inflorescences,flowers, and seedlings

Mourera fluviatilis from northern South America is a spectacular member of the Podostemaceae (river-weeds). Its raceme-like inflorescences are up to 64 cm long and have 40–90 flowers arranged in two opposite rows. Inflorescence development starts with the initiation of a double-sheathed (dithecous) bract in a terminal position. All lateral bracts (again dithecous) are initiated in basipetal order along the two flanks of the inflorescence. Each gap between two neighboring bracts contains a single flower. The flowers are bisexual, each with a whorl of 16–20 ligulate tepals and 14–40 stamens, which are arranged in one or two whorls. Floral development starts with the formation of a girdling primordium rim around a two-lobed primordial gynoecium. Stamen and tepal initiation is centrifugal on the girdling primordium. The anthers are introrse or extrorse, depending on stamen position. Seedlings develop two entire, threadlike cotyledons, followed by forked filamentous leaves, which arise from the plumular pole. The radicular pole of the hypocotyl develops into a claw-shaped holdfast that fixes the young plant to the rock. The developmental morphologies of Mourera fluviatilis and other members of the Mourera group (including Lonchostephus and Tulasneantha) fit well with the Podostemoideae bauplan known from other New World genera, such as Apinagia and Marathrum.     

Morphological association between spermatophores and male genitalia in carabid beetles of the tribe Pterostichini (Coleoptera: Carabidae)

A morphological association between genitalia and ejaculates could provide insight into the function and evolution of genitalia. In this study, the morphologies of the ejaculates and male genitalia of 15 species of Pterostichini and two species of Platynini (Coleoptera: Carabidae) are described. All the species examined formed a spermatophore, the morphology of which could be classified into three types based on its relative volume in the female vaginal cavity and the presence or absence of a pluglike conformation. Male genital morphology could be divided into two types by the direction of the endophallus and gonopore. Species with a strongly bent endophallus invariably formed a pluglike spermatophore. The results suggest that the peculiar shape of endophallus found in some species of Pterostichini may function in forming the pluglike structure of the spermatophore.     

Morphological and architectural features of young Nothofagus pumilio (Poepp. & Endl.) Krasser (Fagaceae)

The morphology and architecture of young Nothofagus pumilio plants growing as understorey and in a cleared area were assessed. Length, number of internodes, presence of branches and apex condition were registered for each growth unit (GU) in the sampled plant. GUs were grouped according to branching order and age of the bearing axis. Understorey plants were seven years old and had a short trunk bearing 2–3 orders of branches; all of their most recent GUs were horizontal and of comparable size. Plants growing in the cleared area were 10–16 years old and had a vigorous vertical and dominant trunk bearing 4–5 orders of less vigorous, horizontal branches. Plants from the cleared area showed signs of having had comparable-size GUs in all of their axes at a younger stage of growth. N. pumilio plants appear to start their growth with the production of short GUs in all of their axes irrespective of the light conditions. In contrast, the development of a vertical, vigorous trunk, seems to be restricted to plants growing in open environments.     

Branching patterns of root systems: quantitative analysis of the diversity among dicotyledonous species

Background and Aims

Root branching, and in particular acropetal branching, is a common and important developmental process for increasing the number of growing tips and defining the distribution of their meristem size. This study presents a new method for characterizing the results of this process in natura from scanned images of young, branched parts of excavated roots. The method involves the direct measurement or calculation of seven different traits.

Methods

Young plants of 45 species of dicots were sampled from fields and gardens with uniform soils. Roots were separated, scanned and then measured using ImageJ software to determine seven traits related to root diameter and interbranch distance.

Results

The traits exhibited large interspecific variations, and covariations reflecting trade-offs. For example, at the interspecies level, the spacing of lateral roots (interbranch distance along the parent root) was strongly correlated to the diameter of the finest roots found in the species, and showed a continuum between two opposite strategies: making dense and fine lateral roots, or thick and well-spaced laterals.

Conclusions

A simple method is presented for classification of branching patterns in roots that allows relatively quick sampling and measurements to be undertaken. The feasibilty of the method is demonstrated for dicotyledonous species and it has the potential to be developed more broadly for other species and a wider range of enivironmental conditions.     

Morphological phylogeny of gall‐forming aphids of the tribe Eriosomatini (Aphididae: Eriosomatinae)

Aphids of the tribe Eriosomatini are typically associated with the tree genera Ulmus and Zelkova as the primary host, on which they induce several types of leaf gall. To elucidate evolutionary changes in the aphid–host associations and gall morphology, phylogenetic relationships were inferred using 36 species (28 in the ingroup) and based on 52 morphological characters. Phylogenetic analysis with equal character weighting led to hundreds of most‐parsimonious trees, and the strict consensus of these was poorly resolved. However, the successive weighting of characters yielded three most‐parsimonious trees. The strict consensus of these supported the monophyly of the Eriosomatini and the monophyly of most genera. Reconstruction of the aphid–host associations on the consensus tree indicated that the ancestral Eriosomatini were associated with Zelkova and that the association with Ulmus evolved twice independently. Ancestral reconstruction suggests that galls of the leaf‐roll type are ancestral to those of the completely and incompletely closed pouch type, and that each type of pouch galls evolved independently. It is suggested that despite early diversification of the Eriosomatini on Zelkova species, Zelkova‐associated eriosomatines had become extinct or survived as relict parthenogens on the secondary host due to the elimination of Zelkova in most regions since the late Tertiary. In contrast, two large genera in the Eriosomatini, Eriosoma and Tetraneura, are associated with the largest elm section Ulmus whose elements are distributed widely in Eurasia, including boreal regions. Therefore, the available evidence suggests that the present species diversity and distribution patterns of the eriosomatines have been largely affected by the diversification and extinction of their host plants during the late Tertiary and Quaternary.     

Phylogeny of the Neotropical moth tribe Josiini (Notodontidae: Dioptinae): a hidden case of Miillerian mimicry

The Neotropical moth tribe Josiini (Notodontidae: Dioptinae) contains over 100 described species in 11 genera. All are diurnal, with brightly-coloured, presumably aposematic wing patterns. Larval hostplants are exclusively in the genus Passiflora (Passifloraceae) except for two new records, reported here, from Tumera (Turneraceae). A comparative morphological study of 26 representative josiine species yielded 86 characters from adults, larvae and pupae, all of which are figured and discussed. Phylogenetic analysis of these data produced a single most-parsimonious cladogram. According to the phylogenetic results: (1) monophyly of the Josiini is strongly supported; (2) the currently accepted generic classification is in disarray; (3) morphological character variation is extensive, and adult traits reflect phylogeny more effectively than do those of immature stages; (4) wing pattern types have undergone convergent evolution. A rare phenotype, longitudinal wing stripes, appears in two widely divergent clades, suggesting the evolution of Miillerian mimicry within the Josiini.