Doronomyrmex pocahontas: not a workerless parasite but still an enigmatic taxon (Hymenoptera,Formicidae)

Summary New data suggest that the North American antDoronomyrmex pocahontas Buschinger 1979, originally described as a workerless inquiline, is not a social parasite. Breeding experiments show a remarkable phenotypic plasticity, which is probably due to hybridization or to intraspecific polymorphism.     

Embryology ofCrocus thomasii (Iridaceae)

The embryology ofCrocus thomasii is described. Male meiosis is of simultaneous type, and gives rise to starchy microspores which develop into lipoid pollen grains; these are two-celled and show a spinulate acolpate, abaculate exine lacking apertures. The tapetum is glandular and its cells become bi- or sometimes multinucleate. The ovule is anatropous and bitegmic; the inner integument forms the micropyle. Megasporogenesis is heteropolar with starch accumulation in the functional chalazal megaspore. Embryo sac development conforms to thePolygonum type. The endosperm development is nuclear. The embryo develops according to the Caryophyllad type. In the ripe seed it is differentiated and enveloped by a starchy cellular endosperm. The embryological characters observed strongly favour a close relation betweenC. thomasii andC. sativus.     

Embryology ofEriocaulon setaceum (Eriocaulaceae)

Eriocaulon setaceum can be characterized by: young microsporangium wall with epidermis, endothecium (with fibrous thickenings), and glandular tapetum (uninucleate cells); pollen grains 3-celled, spiraperturate; embryo sac development according to the Polygonum type and with antipodal cyst; endosperm nuclear; embryo small, with incipient differentiation into cotyledonary and epicotylary loci; seed coat mainly from the inner layers of the integuments; pericarp 2-layered and membranous. Embryologically, theEriocaulaceae are nearer to theXyridaceae than to otherFarinosae. Their elevation to the rank of an order,Eriocaulales, therefore appears justified.     

Embryology of Gomortegaceae (Laurales): characteristics and character evolution

The embryological characteristics of Gomortegaceae, which are poorly understood, were investigated on the basis of Gomortega nitida, the only species of the family, to understand better the evolution of this group within Laurales. Comparisons with other Laurales and Magnoliales (a sister group of Laurales) show that Gomortega has many embryological features in common with the other lauralean families. Notably, Gomortega shares a testa without or with at best only a poorly developed mesotesta as a synapomorphy with all other Laurales. The genus further shares anthers dehisced by valves as a synapomorphy with the other Laurales (except for Calycanthaceae and Monimiaceae), and a non-multiplicative testa and bisporangiate anther as synapomorphies with Atherospermataceae and Siparunaceae (although the non-multiplicative testa occurs as a homoplasy in Monimiaceae, and the bisporangiate anther in Monimiaceae pro parte, Lauraceae pro parte and Hernandiaceae, respectively). Gomortega shows simultaneous cytokinesis to form pollen grains, a one-celled ovule archesporium and non-specialized chalaza, all or part of which may be synapomorphies shared with Atherospermataceae. Gomortega appears to have no embryological autapomorphies, but further comparison with Atherospermataceae is required.Kweon Heo and Yukitoshi Kimoto contributed equally to this work.     

Embryology and relationships ofGyrocarpus andHernandia (Hernandiaceae)

Two representative genera of Hernandiaceae,Gyrocarpus andHernandia, were investigated embryologically to contribute to a better understanding of their respective evolutionary position. Comparisons with other lauralean families using Chloranthaceae or Annonaceae (as a representative of Magnoliales) as an outgroup of Laurales (if present, plus other related taxa) support a lauraceous affinity for the two genera because of the presence of ramified raphal vascular bundles at the chalaza (a synapomorphy), but do not provide evidence for the separation of Hernandiaceae from Lauraceae.Hernandia rather shares with Lauraceae two apomorphies (i.e., the seed pachychalazy and the ruminate seed) which may be homoplasies judged from results of cladistic and molecular studies published elsewhere.Hernandia is greatly divergent from an ancestral line common withGyrocarpus and is even diversified within the genus. Based on evidence from embryology as well as from other sources, it seems best to accept two separate subfamilies in Hernandiaceae as usually have been accepted: one is a derived subfamily Hernandioideae, and the other a less specialized Gyrocarpoideae.     

Embryology of Siparunaceae (Laurales): characteristics and character evolution

Embryological characters of Siparunaceae, which are poorly understood, were studied on the basis of two constituent genera, an African Glossocalyx and a South American Siparuna, to better understand their evolution within Laurales. These two genera have many embryological characteristics in common with the other lauralean families. Noticeably, they share the multi-celled ovule archesporium (uncertain in Glossocalyx) as a synapomorphy with all the other lauralean families except Lauraceae, the anthers dehisced by valves as a synspomorphy with all the other lauralean families except Calycanthaceae and Monimiaceae, and the bisporangiate anther as a synapomorphy with Gomortegaceae and Atherospermataceae. Siparunaceae are, however, distinct from all other laularean families in having unitegmic ovules that were derived from bitegmic ovules, probably due to an elimination of the outer integument. Likewise, the lack of the testa (i.e., developed outer integument), the "endotegmic" seed coat, and the perichalazal seed at maturity are also characteristics of Siparunaceae. Within the family, Siparuna differs from Glossocalyx in having plural tetrads of megaspores and plural, starchy-rich, one-nucleate, tubular embryo sacs (autapomorphies). On the other hand, Glossocalyx is characterized by having bilaterally flattened seeds (autapomorphy). Although functional aspects of those autapomorphies are uncertain, both Glossocalyx and Siparuna show evolution in different embryological characters.     

Embryology ofMalaxis saprophyta,with comments on the systematic position ofMalaxis (Orchidaceae)

InMalaxis saprophyta, anther wall development corresponds to the Monocotyledonous type. The uninucleate tapetum is of secretory type and the endothecium develops U- and V-shaped thickenings on the inner tangential and radial walls. Cytokinesis is simultaneous; tetrahedral, isobilateral and T-shaped tetrads are formed which are compactly aggregated in pollinia. At anthesis the microspore tetrads are 2-celled. The ovule is anatropous, bitegmic and both integuments are dermal in origin. A single hypodermal cell develops directly into a megaspore mother cell. Embryo sac development is predominantly monosporic and less often bisporic. Irrespective of the type of development, the mature embryo sac is 6-nucleate. Although double fertilization occurs, the primary endosperm nucleus degenerates. Embryogeny is of the Onagrad type. The mature embryo lacks differentiation into cotyledon, plumule and radicle. The reticulate seed coat is formed entirely by the outer layer of outer integument. There are three sterile and three fertile valves in the ovary. Although initially parenchymatous, the entire three sterile valves in the ovary and the upper half of the three fertile valves become sclerified after fertilization. The embryological characters support the disputed systematic position ofMalaxis within subtribeMalaxidinae ofEpidendreae.     

Embryology and relationships ofSuriana maritima L. (Surianaceae)

An intensive study of the embryology ofSuriana maritima L. is made to provide the information for discussing relationships and systematic position of the genus which is uncertain yet. Notable embryological features ofSuriana include: ovular archesporium multi-celled; ovule unitegmic, campylotropous, and crassinucellate with a nucellar cap; obturator not formed; ovule and seed pachychalazal; chalazal haustorium of endosperm formed; mature seed exalbuminous. Comparisons with possibly related families suggest thatSuriana is distinct from any of the families discussed but best resembles Rutaceae (and Simaroubaceae). Until a study onStylobasium and several other genera, which might be closer toSuriana, comes up in future, we tentatively support the placement ofSuriana as a monotypic family Surianaceae in Rutalessens. Takhtajan.     

Embryology and systematic position ofLiparis (Orchidaceae)

The embryology ofLiparis paradoxa andL. rostrata has been studied. The young anther wall consists of an epidermis, endothecium, three middle layers and secretory tapetum with uninucleate cells. In the mature anther, two or three sub-epidermal layers develop fibrous thickenings. The anther wall development corresponds to the massive type. Simultaneous cytokinesis results in decussate, isobilateral, linear, T-shaped and tetrahedral pollen tetrads. The pollinia are compact and at anthesis the pollen grains are 2-celled. Ovules are anatropous, bitegmic and tenuinucellate. Both the integuments are dermal in origin and 2-layered. Development of female gametophyte is of the monosporic, 8-nucleate type. Double fertilization occurs. The primary endosperm nucleus degenerates. The mature embryo is organized from the derivatives of tiers 1, 1, m, and n. Its development conforms to a variation of the Onagrad type. The seed coat is formed entirely by the outer layer of the outer integument. There are three sterile and three fertile valves in the ovary. In the prefertilization stages valves consist of parenchymatous cells. After fertilization, the sterile valves become sclerenchymatous whereas the fertile valves remain parenchymatous. The embryological characters support the disputed systematic position ofLiparis within subtribeLiparidinae ofEpidendreae.     

The cytotaxonomy and origin ofRanunculus yaegatakensis,an endemic taxon of Yakushima Island

Ranunculus yaegatakensis, a close relative of the widespreadR. silerifolius, is endemic to the alpine region (ca. 1,600 m alt.) of Yakushima Island. This species differs morphologically fromRanunculus silerifolius mainly due to its dwarfism and leaf shape, but has an identical karyotype with one (Karatsu-type) of the four cytotype ofR. silerifolius. Ranunculus yaegatakensis is highly cross-compatible with the Karatsu-type plants ofR. silerifolius. Their F₁ hybrids show regular meiotic behaviors and set seeds successfully. In addition, the Karatsu-type ofRanunculus silerifolius is found in individuals that are collected not only from the lowland of Yakushima Island but from its neighboring islands. It is supposed to be more specialized than other cytotypes. This evidence suggests thatRanunculus yaegatakensis has been derived from the Karatsu-type plants ofR. silerifolius. We suggest thatRanunculus yaegatakensis be reduced to a variety ofR. silerifolius.     

Embryology of <Emphasis Type="Italic">Japonolirion</Emphasis> (Petrosaviaceae,Petrosaviales): a comparison with other monocots

Japonolirion osense, the sole species of the genus, endemic to Japan, which is placed together with Petrosavia in the Petrosaviaceae and the order Petrosaviales, is still poorly known with respect to systematic characters. Here I present an embryological study of the anther, ovule, and seed of J. osense. Japonolirion is characterized by a glandular anther tapetum, simultaneous cytokinesis in the microspore mother cell, two-celled mature pollen grains, anatropous and crassinucellate ovules, a two-cell-layered nucellar cap formed early in ovule development, antipodal cells hypertrophied in post-fertilization stages, the ab initio cellular mode of endosperm formation, and exotegmic seeds. Comparisons with the basal monocots Acorus (Acorales) and Araceae (Alismatales), and with the more derived monocots Nartheciaceae (Dioscoreales) and Velloziaceae/Triuridaceae (Pandanales), showed that Japonolirion is clearly distinct from those basal and more derived monocots, supporting a distinct position for Petrosaviaceae or Petrosaviales within the monocots. Extensive comparisons further suggest that the two-cell-layered nucellar cap, whose cells are rich in cytoplasm at the time of fertilization in Japonolirion and thus obviously function as the obturator, is likely to be a common characteristic of the basal monocots and may even be a link with the magnoliids.     

Kingdonia,Embryology and Its Systematic Significance

The embryological studies show that Kingdonia is similar to Circaeaster while different from other members of the Ranunculales (sensu Takhtajan, 1980) in development of endosperm and embryo. We consider that there is close systematic relationship between Kingdonia and Circaeaster.     

Embryology ofCordia sebestena (Boraginaceae)

Sporogeneses and gametogeneses inCordia sebestena are described. Anther wall develops according to the Dicotyledonous type. Pollen grains are three-celled at the time of dehiscence. The occurrence of enlarged, precociously developing, densely cytoplasmic cells toward the proximal end of the anther and between adjacent pollen sacs is a unique feature in this species. The possible functions of these cells are discussed. A placental obturator develops simultaneously with the ovules. The dual origin of parietal tissue (from nucellar epidermis and the archesporial initial) is an unusual feature in this species. Megagametophyte development is monosporic or (occasionally) bisporic. Megaspore tetrads are linear or T-shaped. Double dyads and tetrads are frequently observed.—Maintainance of subfamily status forCordioideae is supported.     

川东獐牙菜(龙胆科)的胚胎学研究

利用石蜡切片技术,对川东獐牙菜(Swertia davidii)的胚胎发育过程进行显微观察,并根据现有资料,对獐牙菜属的几种植物进行了比较胚胎学研究。结果表明:川东獐牙菜花药四室,药壁发育为基本型,绒毡层异型起源,为腺质绒毡层,发育后期药室内观察到的绒毡层核是早期该层细胞有丝分裂凸入药室并原位退化形成的,中层细胞3层,药室内壁退化,花药壁表皮宿存,细胞柱状伸长,纤维状加厚;小孢母细胞减数分裂为同时型,四分体排列方式主要为四面体形和左右对称型,少数为"T"形和十字交叉形,成熟花粉为2-细胞类型;子房2心皮、1室,侧膜胎座;薄珠心,单珠被,倒弯生胚珠,大孢子母细胞减数分裂形成4个大孢子直线形排列,合点端的大孢子具功能,雄配子体发育为蓼型;2个极核在受精前融合为1次生核,合点端3个反足细胞宿存,每个细胞均多核和异常膨大,形成明显的反足吸器,并在胚乳之外形成染色较深的类似"外胚乳"结构;珠孔受精,受精作用属于有丝分裂前类型;胚乳发育为核型,胚胎发育为茄型;果实成熟时,种子发育至心形胚阶段;反足细胞在龙胆科一些短命植物中的宿存与分裂具有重要的生殖适应与进化意义。     

Embryology of Chaoborus-induced spines in Daphnia pulex

Daphnia pulex (Crustacea: Cladocera) embryos were found to be sensitive to a chemical cue (kairomone) in an extract of the predator Chaoborus americanus (Insecta:Diptera). Sensitivity of embryos to the kairomone remains throughout embryonic development. Apparently declining sensitivity as development proceeds may be due to the amount of time the embryos are exposed to the kairomone. Male embryos were also found to be sensitive to the kairomone. The smallest eggs within a brood produced small offspring, which showed the antipredator morphology to a significantly lower degree than largest eggs. The production of the neckteeth is described, at the developmental stage in the maturation of the Daphnia coinciding approximately with the escape of the embryos from the brood chamber.     

Systematic placement of an enigmatic Southeast Asian taxon Eupetes macrocerus and implications for the biogeography of a main songbird radiation, the Passerida

Biogeographic connections between Australia and other continents are still poorly understood although the plate tectonics of the Indo-Pacific region is now well described. Eupetes macrocerus is an enigmatic taxon distributed in a small area on the Malay Peninsula and on Sumatra and Borneo. It has generally been associated with Ptilorrhoa in New Guinea on the other side of Wallace's Line, but a relationship with the West African Picathartes has also been suggested. Using three nuclear markers, we demonstrate that Eupetes is the sister taxon of the South African genus Chaetops, and their sister taxon in turn being Picathartes, with a divergence in the Eocene. Thus, this clade is distributed in remote corners of Africa and Asia, which makes the biogeographic history of these birds very intriguing. The most parsimonious explanation would be that they represent a relictual basal group in the Passerida clade established after a long-distance dispersal from the Australo-Papuan region to Africa. Many earlier taxonomic arrangements may have been based on assumptions about relationships with similar-looking forms in the same, or adjacent, biogeographic regions, and revisions with molecular data may uncover such cases of neglect of ancient relictual patterns reflecting past connections between the continents.     

Phylogenetic relationships of the enigmatic ornate shiner,Cyprinella ornata,a species endemic to Mexico (Teleostei: Cyprinidae)

The systematics and taxonomy of North Americancyprinid fishes has historically been said tobe in a chaotic state of affairs. Much of theconfusion as to relationships of species restsin the lack of explicit phylogenetic hypothesesof species and reliance upon degree ofdistinctiveness of taxa or their overallsimilarity for generic placement. Some specieshave had more turbulent or variable taxonomichistories than others. The ornate shiner, Cyprinella ornata, is one of those species,having been placed in the genera Notropis,Codoma, and Cyprinella within the last20 years and found in current texts in any ofthese three genera. Most of the confusionregarding placement of this species has beenrelated to lack of explicit phylogenetichypotheses to formulate its classification, butto a certain degree some researchers haveignored phylogenetic studies and preferred torely upon its morphological distinctiveness orreproductive behavioral traits of questionablehomology to place it in a monotypic genus andalign it with the genera Pimephales andOpsopoeodus.I present a phylogenetic analysis based oncomplete Cytochrome b sequences fromornate shiners, other species of Cyprinella, purported relatives, and severalother species of the Shiner Clade to determinethe phylogenetic affinities of this enigmaticspecies. Molecular analysis reveals the ornateshiner to be more closely related to species ofCyprinella than to a Pimephalesplus Opsopoeodus clade as previouslydiscussed in one morphological analysis, or asargued by Page and Johnston (1990), Johnstonand Page (1992) and Page and Ceas (1989) basedon observations of spawning behaviors andhypotheses of homology between crevice-spawningand egg-clumping behaviors. This molecularanalysis is more consistent with earliermorphological phylogenetic hypotheses of Mayden(1989) wherein these two clades are notconsidered to be closely related and the ornateshiner is a member of the genus Cyprinella.