POLLINATION OF TWO SYMPATRIC SPECIES OF DALECHAMPIA (EUPHORBIACEAE) IN SURINAME BY MALE EUGLOSSINE BEES

Dalechampia brownsbergensis and D. fragrans co-occur in Suriname, and both are pollinated by fragrance-collecting male euglossine bees. Dalechampia brownsbergensis appears to bloom year-round and is pollinated by relatively few species of bees, including Euglossa tridentata and E. gaianii. In contrast, D. fragrans appears to bloom from late October through early December and is visited and pollinated by at least 13 species of euglossines. Field observations of pollination indicated that the two species did not share pollinators. However, when the flowers of D. fragrans were “transplanted” into a population of D. brownsbergensis, the main pollinator of D. brownsbergensis also visited the flowers of D. fragrans. The pollinators of D. fragrans, however, did not visit the flowers of D. brownsbergensis. Partial sharing of pollinators may have only a small negative impact on the two sympatric plant species at this site because they flower simultaneously only part of the year, and they are often spatially separated from one another.     

FLORAL PAEDOMORPHY LEADS TO SECONDARY SPECIALIZATION IN POLLINATION OF MADAGASCAR DALECHAMPIA (EUPHORBIACEAE)

The traditional evolutionary interpretation of Von Baer's “laws” of embryology is that retention of early developmental forms into adulthood (paedomorphosis) leads to the evolution of simpler or more generalized morphology and ecology. Here we show that paedomorphosis can also be involved in an increase in ecological specialization, in this case of plant–pollinator relationships. A paedomorphic transition from generalized pollination (by several functional types of pollinators) to specialized pollination (by one or a few species in one functional type) occurred in a clade of endemic Madagascar vines (Dalechampia spp., Euphorbiaceae). This evolutionary transition involved staminate flowers that fail to develop “normally,” instead holding mature pollen inside virtually unopened, bud‐like flowers. This paedomorphic morphology restricts reward access to “buzz‐pollinating” bees, including Xylocopa species (carpenter bees), which can remove pollen by sonication. This is one of very few reports of paedomorphic specialization, and, as far as we are aware, the first documented case of a rapid reversal to specialized pollination in a lineage of plants that had previously switched from specialized to generalized pollination in conjunction with dispersing to a new region.     

POLLINATION ECOLOGY OF FOUR DALECHAMPIA SPECIES (EUPHORBIACEAE) IN NORTHERN NATAL,SOUTH AFRICA

The pollination ecology of four Dalechampia species was studied in three areas in northern Natal, South Africa. All species were pollinated by resin- and/or pollen-collecting megachilid bees (Hymenoptera: Megachilidae). The most common of these at all study sites was Heriades sp. (Megachilini), which collected both pollen and resin. This bee was the primary pollinator of D. galpinii and D. volubilis, and a secondary pollinator of D. aff. parvifolia and D. capensis. The primary pollinators of these latter two Dalechampia were Pachyanthidium near cucullatum and P. cordatum (Anthididiini), respectively. These are larger bees that collected mainly resin. Other visitors varied among Dalechampia species and among sites. Pairs of Dalechampia species were often found in sympatry (within 30 m of each other) and shared pollinators to varying extents. “Transfer experiments,” in which we placed inflorescences of two Dalechampia species together, reinforced observations of flower discrimination by bees foraging among naturally occurring sympatric pairs. Heriades sp. showed no obvious discrimination between Dalechampia species, but Pachyanthidium spp. “preferred” the Dalechampia species with the greater resin reward. Occasionally, however, Pachyanthidium would visit less rewarding species. Thus, although sympatric African Dalechampia species showed some pollinator partitioning, it was much weaker than found among New World species, and cannot alone explain the general absence of Dalechampia hybrids in northern Natal.     

THE MORPHOLOGY AND RELATIONSHIPS OF DALECHAMPIA SCANDENS (EUPHORBIACEAE)

The circumtropical but preponderantly American genus Dalechampia, comprising nearly 100 species of twining vines (or rarely subshrubs), is strikingly isolated within the Euphorbiaceae because of its distinctive bibracteate inflorescences. There has been considerable taxonomic controversy with regard to the relationships of the genus, and it has been suggested that Dalechampia is allied to the tribe Euphorbieae because of a supposed analogy between its inflorescence and the cyathium in the Euphorbieae. Field and laboratory investigations of the common American species D. scandens, together with a comparative survey of related species, have thrown some light on these problems. The Dalechampia inflorescence seems best interpreted as consisting of a terminal staminate pleiochasium (with part of the lateral branches transformed for nectar production), juxtaposed to a 3-flowered pistillate cyme. The lips of the conspicuous bilabiate involucre are formed by the hypertrophied bracts which subtend the staminate and pistillate cymes. The bisexual inflorescences appear to be distinctly proterogynous, rather than proterandrous, as has been previously suggested. The configuration of the inflorescence—a bilaterally symmetrical pseudanthium—suggests adaptation for crosspollination, but the closing movement of the bracts makes self-pollination probable in the absence of visits by pollinators. The similarity of the Dalechampia inflorescence to the cyathium of the Euphorbieae appears to be entirely superficial, and both reproductive and vegetative data suggest that Dalechampia is related to taxa of tribe Plukenetieae.     

PATTERNS OF CHARACTER DIVERGENCE AND THE EVOLUTION OF REPRODUCTIVE ECOTYPES OF DALECHAMPIA SCANDENS (EUPHORBIACEAE)

The variation of four floral characters (resin-gland area, gland-stigma distance, gland-anther distance, and anther-stigma distance) was analyzed across 15 populations of Dalechampia scandens occurring sympatrically, in various combinations, with five other congeners. Univariate and multivariate analysis of variance and a posteriori comparison tests indicate that there are significant statistical patterns of character divergence away from sympatric congeners for three of the floral characters. These characters, which on the basis of common garden studies appear to be under genetic control, may not vary independently; i.e., genetic control may be overlapping. The characters appear to be functionally related. Populations of Dalechampia scandens occurring sympatrically with congeners possessing relatively large resin glands, large gland-stigma distances and large anther-stigma distances (e.g., D. dioscoreifolia and D. affinis) have significantly smaller resin glands, gland-stigma distances and anther-stigma distances than do populations occurring sympatrically with congeners with relatively small resin glands, gland-stigma distances, and anther-stigma distances (e.g., D. cissifolia, D. heteromorpha and D. schottii). Populations of D. scandens not sympatric with other Dalechampia species generally have intermediately sized structures. The pattern of bidirectional divergence is consistent with the evolutionary scenario that selection against interspecific pollination has resulted in local ecotypic differentiation and character displacement in populations sympatric with ecologically similar congeners.     

SEED PRODUCTION AND DISPERSAL IN DALECHAMPIA (EUPHORBIACEAE): DIVERGENT PATTERNS AND ECOLOGICAL CONSEQUENCES

Patterns of capsule development and methods of seed dispersal are described and compared for 16 neotropical species of Dalechampia. In 15 species all the capsules in a single inflorescence develop simultaneously. In one species, D. spathulata, the capsules develop sequentially. I suggest that the difference between D. spathulata and the other species is the result of the fact that D. spathulata occurs in a low-light, rainforest environment, whereas the other species grow in high-light environments. Sequential development of capsules appears to reduce the loss of seeds to seed predators in species that occur in light-limited environments. The seeds of all 16 species of Dalechampia are dispersed by explosive dehiscence of the capsules. Within the genus there are several different adaptations that increase the average dispersal distance for the seed crop.     

EUGLOSSINE BEE POLLINATION OF THE ORCHID,COCHLEANTHES LIPSCOMBIAE: A FOOD SOURCE MIMIC

Cochleanthes lipscombiae is pollinated by large male and female euglossine bees. The flowers lack pollinator rewards, but attract bees searching for nectar. The euglossines extend their long tongues and crawl into the gullet-flower. The bees probe the back-swept lateral sepals for nectar. Pollination occurs as a pollinarium laden bee backs out, deposits pollinia on the stigma, and obtains a new pollinarium load by dislodging the anther. Some related orchid species have similar morphological characteristics as those essential to the pollination mechanism of C. lipscombiae. These features may have taxonomic significance at the generic level. Cochleanthes lipscombiae may be a floral mimic of a sympatric legume, but may also receive exploratory visits by bees searching for food resources. The latter may be young, recently emerged naive bees, or individuals seeking new nectar hosts during a period of rapid host species turnover.     

EVOLUTION OF PLANT POLLINATION SYSTEMS: HYPOTHESES AND TESTS WITH THE NEOTROPICAL VINE DALECHAMPIA

The results of pollination and mating-system studies were integrated with a phylogenetic study of 40 Neotropical species of Dalechampia L. (Euphorbiaceae) to reconstruct the history of evolutionary change in pollination systems. The results of this analysis were treated as a hypothesis and tested for circularity problems and robustness in the face of changes in the data set. The historical hypothesis was used to make specific predictions about details of pollination ecology and reward biochemistry; the predictions were supported by independent observations. I conclude that pollination systems in Dalechampia have been evolutionarily labile, relative to most morphological features, with repeated parallelisms and reversals. Transitions among the three pollination systems evolved by Dalechampia (pollination by resin-collecting bees, fragrance-collecting male euglossine bees, and pollen-collecting bees) have been facilitated by biochemical exaptation (preadaptation). Pollination by male euglossine bees is relatively rare in the genus but has originated independently three to four times. In contrast, pollination by resin-collecting female bees is very common, but has originated only once. Eighty-six to 97% of transitions between pollination systems involved an intermediate phase during which both old and new pollinators were effective, but 3 to 14% of transitions may have been “instantaneous,” lacking the intermediate phase. Clades of species secreting resin rewards are about 10 times as species rich as clades of species secreting fragrance rewards; circumstantial evidence suggests that different extinction rates may be responsible for this difference. Relatively allogamous (cross-pollinating) species have evolved from more autogamous (self-pollinating) species up to 13 times, and autogamous species have evolved from more allogamous ones up to 11 times. Species occurring in disturbed habitats are facultatively autogamous, whereas species of undisturbed habitats are often highly allogamous.     

POLLINATION OF RAFFLESIA (RAFFLESIACEAE)

The genus Rqfflesia includes about 13 species of parasitic flowering plants, among which are the largest known flowers. The flower with subtending scales is the only part of the plant external to the host and is produced solitary on roots (rarely stems) of the genus Tetrastigma (Vitaceae). Field studies were made of the pollination process in R. pricei, a species endemic to the Crocker Range in the Malaysian state of Sabah (northern Borneo). Pollination is mediated by carrion (bluebottle) flies of the genera Lucilla and Chrysomya. Experimental data indicate that both visual and olfactory cues are important in attracting flies to flowers. Flies (mostly female L. papuensis) obtain loads of the viscous liquid pollen matrix by visiting male flowers and entering anther grooves on the central column of the flower, precisely guided by ridges armed with hairs that force the fly into a position in which the pollen is positioned on the dorsal part of the thorax. “Windows” on the inside of the perigone diaphragm apparently help orient their flight inside flowers. Pollen-loaded flies visiting female flowers may enter the infradiscoidal sulcus formed by a broad ring of stigmatic tissue above and the expanded base of the column below. On entering the sulcus the fly is wedged in so tightly that pollen is rubbed off the thorax onto the stigma. Only large flies could be effective in picking up pollen from male flowers and transferring it to female flowers. The pollination syndrome is sapromyophily, in which the flower closely parallels trap flowers of several other plant families, although it is not a trap. The flower provides no reward for pollinators but deceives them by an apparent offering of food and possibly brood place. Rafflesia plants are extremely rare, perhaps in part because of infrequency of pollination, which requires neighboring male and female flowers simultaneously in bloom.     

KLEINODENDRON AND XYLEM ANATOMY OF CLUYTIEAE (EUPHORBIACEAE)

Kleinodendron, a new genus of Euphorbiaceae, was assigned by Smith and Downs to the tribe Cluytieae. A xylem anatomical survey indicates that there are no objections to this placement. Woods of Cluytieae are diverse but may be characterized generally by having pores which average less than 80 μ in diameter and which are well divided between solitary and radial multiple distributions in the same species; simple vessel perforations; alternate intervascular pitting; fiber-tracheids and libriform wood fibers; exclusively uniseriate, or uniseriate and biseriate heterocellular vascular rays in the same species; uniseriate “bridges” linking superposed biseriate ray segments; diffuse, diffuse-in-aggregates, and scanty vasicentric axial parenchyma, sometimes in the same species; and crystal rhomboids. That Microdesmis and Pogonophora diverge sharply from these generalizations in having scalariform vessel perforations and broad vascular rays, is an indication that they may not be closely related to other genera in Cluytieae.     

MOSQUITO POLLINATION OF HABENARIA OBTUSATA (ORCHIDACEAE)

The floral morphology and pollination of Habenaria obtusata (Orchidaceae) are described. Habenaria obtusata occurs in Alaska, Canada, Norway, Siberia, etc., in great numbers and is pollinated by mosquitoes of the genus Aedes. The pollination process is described as observed in glass cages.     

BEETLE POLLINATION OF DIEFFENBACHIA LONGISPATHA (ARACEAE)

Dieffenbachia is an monoecious understory herb of tropical rain forests that exhibits a complex and specialized relationship with its beetle pollinators. The erect protogynous inflorescence has the spadix divided, with the female flowers in the basal half and male flowers in the upper half. Dieffenbachia longispatha Engler & Krause is pollinated by scarab beetles in the genera Cyclocephala and Erioscelis. The enveloping spathe of the inflorescence opens in the evening, but no flowers are sexually functional until the stigmas become receptive about 24 hr later. Beetles fly to the inflorescence in darkness, suggesting that floral odors play a role as an attractant. Beetles remain in the inflorescence for 24 hr, eating protein-rich staminodia that surround the stigmas. On the evening of the third day the anthers dehisce and beetles become covered with pollen as they crawl up the spadix in the process of leaving. Beetles fly an average of 80 m between inflorescences, usually to the nearest female inflorescence, although distances of 400–1,000 m have been observed. Minimal estimate of genetic neighborhood sizes are large for D. longispatha (750 to 8,900 plants) and neighborhood areas encompass 41,000 to 67,000 m². Experiments demonstrate that the species is self-compatible and that fruit production is pollinator limited.     

POLLINATION BIOLOGY OF LISTERA CORDATA (ORCHIDACEAE)

Pollination biology of northern California populations of Listera cordata (L.) R. Br. is presented. The flowers are allogamous, self-compatible, and protandrous. At anthesis, access to the stigma is blocked by the flap-like rostellum. When an insect contacts the touch-sensitive trigger hairs of the rostellum, the pollinia are suddenly released and become cemented to the insect's body. One to two days after the pollinia are removed, the rostellum rises and exposes the receptive stigma. Visitors are attracted to the flowers by their foul odor and minute quantities of nectar presented on the surface of the labellum. Pollination efficiency is high with 61–78% of flowers from several localities setting fruit. The most important and abundant pollinators of L. cordata at our study sites are fungus gnats (Sciaridae and Mycetophylidae). Other less frequently encountered pollinators are members of these families: Braconidae, Ichneumonidae, and Tipulidae. At present, there is no compelling evidence that the floral characteristics of L. cordata have evolved as a result of an evolutionary interaction with any specific group of insects. Nor is there any indication that the flowers are sapromyophilous, despite their putrid odor.     

中国大戟科珠子木属植物的整理研究

珠子不属(Phyllanthodendron Hemsl.)在我国主要分布于贵州、云南、广西和广东。经研究,作者于本文将其划分为2组10种,其中包括一个新组,4个新种及一个新组合。     

ONTOGENY OF THE COTYLEDONARY REGION OF CHAMAESYCE MACULATA (EUPHORBIACEAE)

Development of the cotyledonary region in Chamaesyce maculata is described from germination of the seed through formation of the dense mat of branches which characterize this common weed. The cotyledonary node is trilacunar with split-lateral traces. Epicotyl development is limited to a pair of leaves (“V-leaves”) inserted directly above and decussate to the cotyledons. The two V-leaves are also vascularized by three traces and insertion of these traces relative to the vasculature at the immediately subjacent cotyledonary node is asymmetrical; four of the six V-leaf traces arise on one side of the intercotyledonary plane and two arise on the opposite side. Further shoot development is limited to lateral branches that develop sequentially from cotyledonary axillary buds, and then from de novo buds which arise at bases of previously developed lateral branches. The first axillary bud to develop is located on that half of the seedling which supplies the V-leaves with four traces. Development or insertion of the third and fourth branches (first and second de novo branches) relative to the first and second (cotyledonary) branches occurs in two patterns, termed cis and trans. Subsequent de novo branches generally develop between the two most recently developed branches on that half of the seedling, gradually forming a large branch plexus at the cotyledonary region. In mature robust specimens, however, the sequence of lateral branch development may become irregular. Structure of the cotyledonary region of C. maculata does not readily support widely held concepts of homology between the pleiochasium of Euphorbia subgenus Agaloma and the lateral branch system of Chamaesyce.     

BEETLE POLLINATION OF CYCLANTHUS BIPARTITUS (CYCLANTHACEAE)

The morphology, inflorescence phenology, and insect visitors of Cyclanthus were observed during two reproductive seasons at Finca La Selva in the Atlantic lowlands of Costa Rica. The inflorescence of Cyclanthus is an elongate spadix that is subtended and enclosed by four large, cymbiform bracts. Staminate and pistillate flowers are arranged in separate cycles along the length of the spadix and show marked dichogamy. Pollinators of Cyclanthus are beetles of the genus Cyclocephala (Scarabaeidae). The beetles arrive at the inflorescence while it is in the pistillate phase, during the first evening of the 2-day flowering period. They remain in the inflorescence for 24 hrs, until the end of staminate anthesis. The bracts of the inflorescence produce specialized tissue that the beetles consume along with pollen. Chemical analysis of the food tissue indicates that it is almost 50% lipid by dry weight. The scarabs use the inflorescence as an aggregation site for mating. Experiments with bagged inflorescences showed that no seeds are produced in the absence of pollinators, which suggests that Cyclanthus is obligately allogamous.