The Evolution and Distribution of Subfam. Spiraeoideae (Rosaceae) of China,with Special Reference to Distribution of the Subfamily in the World

The subfam. Spiraeoideae, consisting of 22 genera and more than 260 species in the world,is the most primitive subfamily of Rosaceae. It has developed into two groups,i.e. evergreen and deciduous ones, of which eight genera and 100 species in China are totally deciduous. In the present paper, the origin,evolution and distribution of the Chinese genera is discussed mainly, and the distribution of the whole subfamily in the floristic regions of the world is also mentioned. Based on evolutionary trends of morphological characters, Spiraea L. is considered as the most primitive genus in the deciduous group of subfam. Spiraeoideae, from which some genera are been derived, the systematic position and evolutionary relationships between different genera are elucidated in this paper. Through the analysis on the geographical distribution of the genera in China, the areal types may be divided as follows: (1) North Temperate Type: Spiraea, Physocarpus, Aruncus. (2) East Asian and North American Disjunct Type: Sorbaria. (3) Mediterranean, West Asian (or Central Asia) and East Asian Type: Sibiraea. (4) Temperate Asian Type: Exochorda.(5) East Asian Type: (a) Sino Himalayan Distribution: Neillia; (b) Sino Japan Distribution: Stephanandra. After analysis of the distribution of subfam. Spiraeoideae in the world, it is shown that the Eastern Asiatic Region, being the richest in genera, species and endemic species of the world,is not only the center of distribution and differentiation,but also an important region for occurrence and development of some deciduous genera of this subfamily, while in North America, the Madrean Region and Rocky Mountain Region, genera, species and endemic species are abundant, which indicates that the western part of North America is also the distribution center of this subfamily at the present, but it may be the secondary center of distribution. It can be seen that the relatively primitive and evergreen g enera, i.e. Quillaja and Kageneckia, are now confined to South America. The fact implies that the South America may be the region for early differentiation and development of the evergreen genera in Subfam. Spiraeoideae. The analysis of Chinese plants has shown that China has the most members of the subfamily in Eastern Asiatic Region, with eight genera, 82 species and 62 endemic species and that the maximum concentration is in western Sichuan, northwestern Yunnan and their adjacent areas. It is very obvious that the center of distribution and diversity of Subfam. Spiraeoideae in China lies in the Hengduan Mountain Region of Sino Himalayan Forest Subkingdom and the western part of Sino Japan Forest Subkingdom, where may be the birthplace of some genera in China. It may be considered that the deciduous genera of Subfam. Spiraeoideae might have originated in Laurasia.According to the fossil records, the time of origin of Subfam.Spiraeoideae dates back to the Lower Cretaceous.     

中国蔷薇科绣线菊亚科的演化、分布——兼述世界绣线菊亚科植物的分布

绣线菊亚科是蔷薇科最原始的亚科,共有22属260余种, 包括常绿和落叶两大类群,前者是 原始类型。我国有8属100种,全都为落叶性。本文着重讨论中国各属的起源、演化和分布等 ,同时也概述全亚科植物在世界各植物区的分布等问题。绣线菊属Spiraea是该亚科落叶类群中最原始的属,它在早期发生趋异进化,衍生出形态各异而亲缘关系密切 的不同属,本文阐明了中国各属的系统位置和属间的亲缘关系。通过对我国各属地理分布的 分析对比,属的分布区可归纳为5个类型。对全球绣线菊亚科植物在世界各植物区中的属、种数统计表明,东亚区有8属105种,其中有96个特有种,是该亚科植物分布最多而又最集中 地区,具有在系统发育上处于各主要演化阶段的落叶类型,因此,东亚区是全球绣线菊亚科植 物的现代分布和分化中心,也是落叶类群发生和发展的关键地区。在北美洲,从马德雷区至落基山区一带分布着11属46种,均为特有种,显然北美洲西部也是该亚科植物的现代分布中心,但可能是第二分布中心。南美洲至今保存2个较古老的常绿属,即Quillaja和K ageneckia,基于此,南美洲可能是绣线菊亚科某些常绿属早期分化和发展的关键地区 。中国绣线菊亚科植物在东亚区占绝对优势,有8属82种,其中有62个特有种,分别占该区属 、种和 特有种数的100%、82%、和65%, 这些类群分布最密集地区是在中国喜马拉雅森林植物亚区 中的横断山脉地区和中国日本森林植物亚区的西部,这一带是中国绣线菊亚科的现代分布和多样性中心,很可能是某些属的发源地。由此看来,绣线菊亚科的落叶属可能起源于劳亚古陆。据化石记载,该亚科植物的起源时间可以追溯到白垩纪早白垩世。     

NUCLEAR DNA CONTENT VARIATION WITHIN THE ROSACEAE

Nuclear DNA content has been estimated using flow cytometry for 17 species and eight cultivars of Malus and for 44 species of 29 other genera within the Rosaceae. Compared to other angiosperms, diploid genome sizes vary little within the family Rosaceae and within the genus Malus. C-values of genera within the subfamilies Spiraeoideae and Rosoideae are among the smallest of flowering plants thus far reported. In general, the Maloideae have the largest diploid genomes of the family, consistent with their higher chromosome numbers and presumed polyploid origin.     

Parasite relationships and disposition of Filipendula

Filipendula spp. are attacked by rust fungi on the genusTriphragmium. Several related genera (tribe Phragmidieae) occur wholly or partly on Rosaceae, and all are restricted to genera in Rosoideae. It appears, by inference, thatFilipendula also belongs to this subfamily. The only recorded rusts of Spiraeoideae are distantly related, and appear, from morphology and total host relationship, to be of much earlier origin.     

VARIATION IN STRUCTURE AMONG FRUITS OF MALOIDEAE (ROSACEAE)

Fruits of Rosaceae subfamily Maloideae are known as pomes, a fruit type unique to the Maloideae. Using light microscopy we examined the pomes of 173 species, broadly representing the taxonomic diversity of the subfamily. Significant variation occurs among pomes in retention, orientation, and texture of calyx lobes on the distal end of the fruit; density and distribution of sclereids in the flesh; heterogeneity among flesh parenchyma cells; carpel number and number of ovules per carpel; degree of connation among the carpels and adnation between ovary and hypanthium; and texture of the core. Phenetic clustering closely links congeneric fruits. Sorbus is a notable exception; fruits of the compound-leaved species form a cluster unrelated to fruits of the simple-leaved species. Fruits of Malus are also heteromorphic. Division of the Maloideae into two tribes, Crataegeae and Sorbeae, based largely on texture of the core is not substantiated. We hypothesize that the ancestral pome had five carpels, minimal connation of carpels, minimal adnation of ovaries to the hypanthium, two ovules per carpel, and a leathery core. Various adaptations for frugivore dispersal of the seeds partially explain trends in pome evolution.     

Phylogeny and classification of Rosaceae

Phylogenetic relationships among 88 genera of Rosaceae were investigated using nucleotide sequence data from six nuclear (18S, gbssi1, gbssi2, ITS, pgip, and ppo) and four chloroplast (matK, ndhF, rbcL, and trnL-trnF) regions, separately and in various combinations, with parsimony and likelihood-based Bayesian approaches. The results were used to examine evolution of non-molecular characters and to develop a new phylogenetically based infrafamilial classification. As in previous molecular phylogenetic analyses of the family, we found strong support for monophyly of groups corresponding closely to many previously recognized tribes and subfamilies, but no previous classification was entirely supported, and relationships among the strongly supported clades were weakly resolved and/or conflicted between some data sets. We recognize three subfamilies in Rosaceae: Rosoideae, including Filipendula, Rubus, Rosa, and three tribes; Dryadoideae, comprising the four actinorhizal genera; and Spiraeoideae, comprising Lyonothamnus and seven tribes. All genera previously assigned to Amygdaloideae and Maloideae are included in Spiraeoideae. Three supertribes, one in Rosoideae and two in Spiraeoideae, are recognized.     

Rosaceae chemotaxonomy and the origins of the Pomoideae

The results of a chemotaxonomic survey of the Rosaceae for the occurrence of flavone C-g!ycosides, together with a general literature review of Rosaceae chemotaxonomy, are compared with the hypotheses of classical botanical taxonomy relating to the origins and evolution of the subfamilies of the Rosaceae. A tentative phyiogenetic scheme is presented, based upon all available taxonomic evidence. The chemotaxonomic data is consistent with the hypothesis that the Pomoideae are of allopolyploid origin, produced by ancient hybridization between primitive forms of the subfamilies Prunoideae and Spiraeoideae, although the possibility cannot completely be excluded that primitive Spiraeoideae only were involved. Flavone C-glycosides, hitherto thought to be restricted in the Rosaceae to Crataegus , have also been detected in the following genera: Pyracantha, Osteomeles, Aronia, Hesperomeles, Malacomeles, Chamaemetes, Dichotomanthes (Pomoideae); Quillafa (Spiraeoideae); Agrimonia, Adenostoma , doubtful traces in Sanguisorba, Potentilla, Alchemilla (Rosoideae). None could be detected in the Prunoideae. The chemotaxonomic evidence supports the hypothesis that Quillaja is a relict of an apocarpous ancestor of the Pomoideae and that Dichotomanthes is a relict of the primitive Pomoideae.     

Photinia taishunensis sp. nov. (Rosaceae) from Zhejiang,eastern China

Photinia taishunensis G. H. Xia, L. H. Lou & S. H. Jin (Rosaceae, Spiraeoideae) from Zhejiang, China, is described and illustrated. It has previously been confused with P. lochengensis Yü, but differs in details of habit, leaf size, leaf shape, inflorescence, flower size and pubescence. The pollen grains of P. taishunensis are monads, elliptical with trisulcate apertures, and the exine is rugulose, dotted with sparse punctures.     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

FLORAL DEVELOPMENT AND VASCULATURE IN HYDROCLEIS NYMPHOIDES (BUTOMACEAE)

The flower of Hydrocleis nymphoides consists of three sepals which arise in spiral succession, three simultaneously arising petals, numerous stamens and staminodia which arise in centrifugal order, and six carpels. A residual apex remains at maturity. The first-formed members of the androecium are stamens and the later-formed members are staminodia which develop below the stamens and which become outwardly displaced during expansion of the receptacle. The androecium is supplied by branching vascular trunk bundles. The carpels are completely open but the ventral margins are slightly conduplicately appressed basally. A single dorsal bundle provides the stigmatic area with vascular tissue, and a network of small placental bundles supplies the numerous laminar ovules. There are no clearly defined ventral bundles. It is suggested that Hydrocleis nymphoides is neither the most primitive nor the most advanced member of the family. A pattern of phylogenetic reduction in the androecium and receptacle is suggested for the entire family.     

ONTOGENY AND ANATOMY OF THE FLOWER OF LIMNOCHARIS FLAVA (BUTOMACEAE)

The flowers of Limnocharis flava (L.) Buch. are borne in an indeterminate umbel and each consists of three sepals, three yellow petals, and about 18 carpels surrounded by numerous stamens and staminodia. The androecium is centrifugally developed, and the last-formed members are staminodial; it is supplied by branching vascular systems. Carpels arise almost simultaneously, and a prominent residual floral apex remains. The carpels are partially conduplicately closed and are also primitive in possessing laminar placentation and in lacking differentiation of a style. The gynoecium is essentially apocarpous, but there are slight fusions of adjacent carpels near their ventral margins where they are attached to the receptacle. It is suggested that the Limnocharis flower is the most primitive in the family.     

Phylogenetic relationships in Rosaceae inferred from chloroplast matK and trnL-trnF nucleotide sequence data

 Phylogenetic relationships in Rosaceae were studied using parsimony analysis of nucleotide sequence data from two regions of the chloroplast genome, the matK gene and the trnL-trnF region. As in a previously published phylogeny of Rosaceae based upon rbcL sequences, monophyletic groups were resolved that correspond, with some modifications, to subfamilies Maloideae and Rosoideae, but Spiraeoideae were polyphyletic. Three main lineages appear to have diverged early in the evolution of the family: 1) Rosoideae sensu stricto, including taxa with a base chromosome number of 7 (occasionally 8); 2) actinorhizal Rosaceae, a group of taxa that engage in symbiotic nitrogen fixation; and 3) the rest of the family. The spiraeoid genus Gillenia, not included in the rbcL study, was strongly supported as the sister taxon to Maloideae sensu lato. A New World origin of Maloideae is suggested. The position of the economically important genus Prunus and the status of subfamily Amygdaloideae remain unresolved. Received February 27, 2001 Accepted October 11, 2001     

Fruit fossils of Rosoideae (Rosaceae) from the late Pliocene of northwestern Yunnan,Southwest China

The subfamily Rosoideae Focke (Rosaceae) has a good fossil record in the Northern Hemisphere, but these fossil records are confined mainly to a few genera, whereas the majority, in particular those with herbaceous members, are still under‐represented. In this study, we describe new fruit fossils of Rosoideae, including Fragaria achenes and Rubus endocarps, from the late Pliocene of northwestern Yunnan, Southwest China. These fossils add new accounts to the fossil archive of Rosoideae and provide the first fossil record of Fragaria in East Asia. The new fossil findings provide a historical backdrop for the modern diversity and distribution of the subfamily in northwestern Yunnan, a topographically complex area accommodating a high diversity for many plant groups. Our Rubus fossils, in combination with other nearby coeval occurrences of the genus, suggest that Rubus was already establishing its modern diversity in northwestern Yunnan during the late Pliocene. This finding enriches our knowledge of the post‐Neogene diversification of flowering plants in northwestern Yunnan, which is thought to be largely driven by dramatic mountain uplifts and environmental complications associated with the southeastern extension of the Tibetan Plateau.     

Evolution of growth habit, inflorescence architecture, flower size, and fruit type in Rubiaceae: its ecological and evolutionary implications

During angiosperm evolution, innovations in vegetative and reproductive organs have resulted in tremendous morphological diversity, which has played a crucial role in the ecological success of flowering plants. Morindeae (Rubiaceae) display considerable diversity in growth form, inflorescence architecture, flower size, and fruit type. Lianescent habit, head inflorescence, small flower, and multiple fruit are the predominant states, but arborescent habit, non-headed inflorescence, large flower, and simple fruit states occur in various genera. This makes Morindeae an ideal model for exploring the evolutionary appearances and transitions between the states of these characters. We reconstructed ancestral states for these four traits using a bayesian approach and combined nuclear/chloroplast data for 61 Morindeae species. The aim was to test three hypotheses: 1) self-supporting habit is generally ancestral in clades comprising both lianescent and arborescent species; 2) changes from lianescent to arborescent habit are uncommon due to "a high degree of specialization and developmental burden"; 3) head inflorescences and multiple fruits in Morindeae evolved from non-headed inflorescences and simple fruits, respectively. Lianescent habit, head inflorescence, large flower, and multiple fruit are inferred for Morindeae, making arborescent habit, non-headed inflorescence, small flower, and simple fruit derived within the tribe. The rate of change from lianescent to arborescent habit is much higher than the reverse change. Therefore, evolutionary changes between lianescent and arborescent forms can be reversible, and their frequency and trends vary between groups. Moreover, these changes are partly attributed to a scarcity of host trees for climbing plants in more open habitats. Changes from large to small flowers might have been driven by shifts to pollinators with progressively shorter proboscis, which are associated with shifts in breeding systems towards dioecy. A single origin of dioecy from hermaphroditism is supported. Finally, we report evolutionary changes from headed to non-headed inflorescences and multiple to simple fruits.     

Intra‐individual variation of flowers in Gunnera subgenus Panke (Gunneraceae) and proposed apomorphies for Gunnerales

A study of inflorescence and flower development in 12 species from four of the six subgenera of Gunnera (Gunneraceae) was carried out. In the species of subgenus Panke, initiation of floral apices along the partial inflorescences is acropetal but ends up in the late formation of a terminal flower, forming a cyme at maturity. The terminal flower is the largest and the most complete in terms of merosity and number of whorls and thus it is the most diagnostic in terms of species‐level taxonomy. The lateral flowers undergo a basipetal gradient of organ reduction along the inflorescence, ranging from bisexual flowers (towards the distal region) to functionally (i.e. with staminodia) and structurally female flowers (towards the proximal region). Our results show that the terminal structure in Gunnera is a flower rather than a pseudanthium. The terminal flower is disymmetric, dimerous and bisexual, representing the common bauplan for Gunnera flowers. It has a differentiated perianth with two sepals and two alternate petals, the latter opposite the stamens and carpels. Comparisons with other members of the core eudicots with labile floral construction are addressed. We propose vegetative and floral putative synapomorphies for the sister‐group relationship between Gunneraceae and Myrothamnaceae. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 262–283.     

Pseudoracemes in papilionoid legumes: their nature, development, and variation

Pseudoracemes in papilionoid legumes: their nature, development, and variation. Cymelike partial inflorescences called fascicles have been reported in the inflorescences of several papilionoid tribes. The total inflorescence is termed a ‘pseudoraceme’ because of the multiple flowers in each bract axil. Pseudoraceme development has been studied in 22 taxa in five papilionoid tribes (Abreae, Desmodieae, Millcttieae, Phaseoleae and Psoraleeae). Two to twelve flowers occur per bract axil among various taxa, with three the most common number.Pongamia pinnata and Clitoris fairchildiana have only two flowers per axil; Vigna radiata, Phaseolus vulgaris, and Apios americana have four to five commonly, and Dioclea aff.ucayalina and Abrus precalorius have up to 12. The ‘fascicle’ usually consists of a triad of three flowers; each triad resembles a dichasial cyme in that the middle flower appears terminal. The middle flower however is subtended by a bract on the abaxial side, so that the middle flower is technically lateral. When the first-order axis elongates, each triad may either remain intact or be separated by axis intervalS. Many variations on the basic triad pattern occur in the species studied: 1.one or two flowers may develop while others that are initiated remain suppressed; 2. Additional flowers may be produced that replicate the first triad; 3. Additional flowers may form medianly only, on the abaxial side. The second-order inflorescence axis which has produced the three flowers persists to produce more flowers in replication of the triad pattern in several taxa (Apios americana, Vigna radiata, Phaseolus vulgaris, and Dioclea aff.ucayalina). In Butea monosperma the second-order inflorescence apex produces subsequent flowers (after the triad) in a helix. In Erylhrina perrieri, there is no indication of a persistent second-order inflorescence apex after the central flower; such a condition could be interpreted as a cyme, except for the abaxial subtending bract. The triad in Psoralea pinnata is a true cyme; the middle flower lacks a subtending bract other than that subtending the entire fascicle. Developmentally, the difference between a cyme and an early-determinate raceme (as in the triad type of pseudoraceme) is rather slight. Comparison of the types of inflorescences described here may indicate how the transition may have occurred between racemes and cymes in the evolution of legumes.     

Selective fruit abortion in Prunus mahaleb (Rosaceae)

In Prunus mahaleb (Rosaceae), only a small proportion of those flowers that initially develop give rise to fruits, and the probability of producing fruit differs between flowers. The results of this study indicate that the probability of producing fruit is about 60% for the first (i.e., most proximal) flower to develop in each inflorescence, but very low for the last (i.e., most distal). This pattern is maintained regardless of between-inflorescence differences in flowering phenology, suggesting that each inflorescence can be viewed as an independent subunit with respect to allocation of resources for reproduction. These results are compatible with the ovary reserve and selective abortion hypotheses for the adaptive value of low fruit set in P. mahaleb.