扁苞蕗蕨和长柄蕗蕨的细胞分类学研究

本文首次报道了四川产扁苞蕗蕨和长柄蕗蕨的染色体及孢子发生情况。它们的染色体数目分别为 n=26和n=28,都是有性生殖的二倍体。染色体基数26在蕗蕨属的发现进一步证明了它和膜蕨属、厚壁蕨属的密切亲缘关系。这两种蕗蕨的孢子囊通常形成128个孢子及少数256个孢子的情况被描述,它们应该被看成是有性生殖真蕨,尤其是在膜蕨科和薄囊真蕨中的其他较原始的类群中的正常的孢子发生路线。     

A Preliminary Study on Cytology of Chinese Pteris

Cytological and biosystematic studies on the genus Pteris have made a great contribution to the theory of speciation and evolution in ferns. Sino-Japanese area is one of the speciation centers of this genus. But only a few Chinese species have been known cytologically. As a preliminary study, ten species of the genus Pteris and two species of the related genera Pteridium and Histiopteris were observed. All the materials were collected from southwest and south China. Their young sori were fixed in ethanol and glacial acetic acid (3:1), in the field. The preparations were made with acetocarmine squash method. The vouchers are deposited in PE. The results of observation are summarized in Table 1. Of Pteris, four species are agamosporous, eight are polyploid, six are actually the members of species complexes. Only two species are purely sexual diploids. Pteris cretica L. var. nervosa (Thunb.) Ching et S. H. Wu has 58 autobivalents at meiosis of spore mother cell, and usually produces 32 diplospores per sporangium. It is obviously an agamosporous diploid. Apart from 32-spored sporangium, some 4-, 8-, 16-, 64-, 13-, 34-, 36-spored, completely abortive or partly abortive sporangia were also found in the fixed material of a single individual. For explaining these unusual types of sporangia at least nine sporogenesis routes can be inferred. It may not be impossible that besides mainly producing functional diplospores, P. cretica var. nervosa also gives a few functional giant spores with a doubled or multiplied somatic chromosome number, which means that polyploids may be simply derived from the spores produced by their diploid parent. The spores in a sporangium are usually tetrahedral, but a few bilateral ones are also found in some sporangia. Sometimes, the bilateral spores are even more than the tetrahedral ones in a sporangium. Pteris vittata L. from the type locality shows 58 bivalents at diakinesis of meiosis. Without doubt, it is a sexual tetraploid, which was also found in south Guangdong and south Yunnan. However, Its natural ancestral diploid has been widely found in the subtropical regions of China, such as Sichuan, Guizhou, Yunnan and Hubei provinces. A sterile triploid with the chromosome number of n=201Ⅰ+26Ⅱ+5Ⅲ and a sterile tetraploid hybrid with the chromosome number of n=9Ⅰ+45Ⅱ+3Ⅲ+21Ⅴ were found in south Yunnan and south Guangdong respec tively. It is evident that P. vittata L. sensu lato is a species complex which includes several cy totypes. Its hexaploid form was reported from south India by Abraham et al. The distribu tion pattern of different cytotypes in P. vittata complex may indicate that the tropics is more favourable to formation and surviving of polyploid than the subtropics. However, it is most possible that Chinese subtropics is the origin place of the ancestral member in the P. vittata complex. In gross morphology, the tetraploid and triploid forms are only bigger and stronger than the diploid form. But the diploid can also grow rather big in cultivation. They can har dly be distinguished from each other. Therefore, they are not given formal names here. The author fully agree with Prof. T. Reichstein when he said in 1983 that it was hoped that a new nomenclature could be adopted for the cytotypes of species complex in future. The special chromosome number of Pteris deltodon Bak .was counted in this work for the first time. It has 55 bivalents at meiosis of SMC. This number is the only exception in the genus Pteris, and shows that aneuploidy may have taken place in this genus. However, the pos sibility can not be ruled out that P. deltodon is an allotetraploid came from two diploid species respectively with the chromosome number 26 and 29. Pteris gallinopes described by Prof. Ching in i983 is an agamosporous tetraploid with 116 autobivalents at meiosis, different from the related triploid species P. dactylina Hook. and P. henryi Christ. P. ensiformis Burm^。 P. muftifida Poir. and P. semipinnata L. are sexual tetraploids with n=58. P. excelsa Gaud. and P. oshimensis Hieron. var. paraemeiensis Ching are agamosporous triploids with n = 87 autobivalents. P. wallichiana Agardh is a sexual diploid with n=29. The high frequency of polyploids and agamospory among these random sampled Chinese species futher confirms the conclusion that both polyploidy and apomixis have played an important role in speciation and evolution of the genus Pteris. Pteridium aquilimum (L.) Kunh var. latiusculum (Desv.) Underw. ex Heller and Histiopteris incisa (Thunb.) J. Sm. have n=52 and n=96 respectively. They are both sexual tetraploids. Their chromosome numbers show again that the genera Pteridium and Histiopteris are distinquished from the genus Pteris by different basic numbers. This project is supported by the National Natural Science Foundation of China.     

Chromosome counts in some Chinese ferns

The spore mother cells of six species of Chinese homosporous ferns have been examined, among them the chromosome numbers of five species, Adiantum capillus-junonis Rupr. (n= 30), Ctenitis rhodolepis (Clarke) Ching (n = 41), Cyclogramma flexilis (Christ) Tagawa (n = 68), Leptogramma scallani (Christ) Ching (n = 36) and Vandenboschia auriculata (B1.) Copel. (n=36), from Emei Shan (Omei) Sichuan province are recorded for the first time. The chromosome number n=68 from the species Cyclogramma flexilis suggests that the base number for this genus is 34. Another species, Athyrium brevifrons Nakai from Wuling Shah Beijing (Peking) has a gameticchromosome number n=40, as already reported by K. Mitui and H. Hirabayashi.     

Ontogeny of strobili, sporangia development and sporogenesis in Equisetum giganteum (Equisetaceae) from the Colombian Andes

Studies on the ontogeny of the strobilus, sporangium and reproductive biology of this group of ferns are scarce. Here we describe the ontogeny of the strobilus and sporangia, and the process of sporogenesis using specimens of E. giganteum from Colombia collected along the Rio Frio, Distrito de Sevilla, Piedecuesta, Santander, at 2200m altitude. The strobili in different stages of development were fixed, dehydrated, embedded in paraffin, sectioned using a rotatory microtome and stained with the safranin O and fast green technique. Observations were made using differential interference contrast microscopy (DIC) or Nomarski microscopy, an optical microscopy illumination technique that enhances the contrast in unstained, transparent. Strobili arise and begin to develop in the apical meristems of the main axis and lateral branches, with no significant differences in the ontogeny of strobili of one or other axis. Successive processes of cell division and differentiation lead to the growth of the strobilus and the formation of sporangiophores. These are formed by the scutellum, the manubrium or pedicel-like, basal part of the sporangiophore, and initial cells of sporangium, which differentiate to form the sporangium wall, the sporocytes and the tapetum. There is not formation of a characteristic arquesporium, as sporocytes quickly undergo meiosis originating tetrads of spores. The tapetum retains its histological integrity, but subsequently the cell walls break down and form a plasmodium that invades the sporangial cavity, partially surrounding the tetrads, and then the spores. Towards the end of the sporogenesis the tapetum disintegrates leaving spores with elaters free within the sporangial cavity. Two layers finally form the sporangium wall: the sporangium wall itself, with thickened, lignified cell walls and an underlying pyknotic layer. The mature spores are chlorofilous, morphologically similar and have exospore, a thin perispore and two elaters. This study of the ontogeny of the spore-producing structures and spores is the first contribution of this type for a tropical species of the genus. Fluorescence microscopy indicates that elaters and the wall of the sporangium are autofluorescent, while other structures induced fluorescence emitted by the fluorescent dye safranin O. The results were also discussed in relation to what is known so far for other species of Equisetum, suggesting that ontogenetic processes and structure of characters sporoderm are relatively constant in Equisetum, which implies important diagnostic value in the taxonomy of the group.     

Phylogenetic relationships within the fern genus Hymenophyllum s.l. (Hymenophyllaceae, Filicopsida): contribution of morphology and cytology

The phylogenetic relationships of Hymenophyllum and its segregate genera Cardiomanes, Hymenoglossum, Rosenstockia, Serpyllopsis and Microtrichomanes are addressed, using 31 morphological characters of the sporophyte and one cytological character. As expected, this study reveals considerable morphological heterogeneity within the genus sensu lato, but several apomorphic changes allow support for some clades. Four unresolved taxa, Cardiomanes, Hymenoglossum, Diplophyllum and Mecodium pro parte are probably the most basal elements in Hymenophyllum. The analysis also suggests the polyphyly of Mecodium, and two unexpected associations: Sphaerocionium together with Microtrichomanes; and a broad clade composed of subg. Hymenophyllum, Hemicyatheon and Craspedophyllum, genera Rosenstockia and Serpyllopsis, and subsect. Leptocionium and Amphipterum. These associations appear justified by morphological, cytological or geographical data, and most of them are in agreement with preliminary molecular results.     

IDANOTHEKION GEN. N., A SYNANGIATE POLLEN ORGAN WITH SACCATE POLLEN FROM THE MIDDLE PENNSYLVANIAN OF ILLINOIS

Idanothekion glandulosum gen. et sp. n. is a synangiate pollen organ represented by approximately 30 specimens contained in coal balls from the middle Pennsylvanian of Illinois. Each synangium is composed of seven to nine elongate sporangia that are fused laterally for approximately four-fifths of their length, and are radially arranged about, and fused to, a short central column; the central column is restricted to the proximal one-third of the synangium. Distal to the column the sporangia surround a hollow central area. Dehiscence occurred by means of a longitudinal slit along the mid-line of the inner face of each sporangium. The outer walls of the sporangia have a complex histology involving an external epidermis, a middle presumably glandular layer containing scattered enlarged cells, and an inner layer made up of thin-walled parenchyma. Vascular tissue is present in the central column and outer walls of the sporangia. Each sporangium has a prominent, attenuate, multicellular tip. Large numbers of saccate pollen grains similar to those found in numerous fossil and extant coniferophytes as well as some Mesozoic pter-idosperms were produced in each sporangium. Idanolhekion resembles some synangia assignable to Paleozoic members of the Marattiales; however, the new genus compares most closely with pollen organs believed to have been produced by members of the Pteridospermales. It seems most likely that Idanothekion represents the pollen organ of some member of the Lyginopteridaceae that produced pollen of a type which up to now has not been known from Paleozoic seed ferns.     

Chromosome studies of cheilanthoid ferns (Pteridaceae: Cheilanthoideae) from the western United States and Mexico

Although analyses of chromosome numbers represent a fundamental step in the study of any group of organisms, the xeric-adapted cheilanthoid ferns (Pteridaceae: subfamily Cheilanthoideae) have received little attention from cytogeneticists due to the difficulty in obtaining samples and accurate chromosome counts. In an effort to clarify patterns of chromosomal evolution in this group, we present 131 chromosome counts representing 75 taxa of cheilanthoid ferns from the western United States and Mexico. First reports are provided for 24 taxa, including the first count for the genus Cheiloplecton. Nine other taxa yielded numbers that had not been reported previously. Our data suggest that chromosome base numbers are more stable than previously thought and that much of the reported variation may involve erroneous counts. When coupled with published DNA sequence data, our counts suggest that the plesiomorphic base number of subfamily Cheilanthoideae is x = 30 and that x = 29 has arisen just once or twice among the taxa studied.     

Molecular systematics of the fern genus Hymenophyllum s.l. (Hymenophyllaceae) based on chloroplastic coding and noncoding regions

We investigated the phylogenetic relationships of the filmy fern genus Hymenophyllum s.l. using the rbcL and rps4 genes and the intergenic spacer rps4-trnS. Because of variation in length of the noncoding marker, we tested and compared three methods for integrating indels. They proved to be useful for estimating a phylogeny of the genus. The rps4-trnS marker, with coded indels integrated, produced better resolution than analysis of either rps4 or rbcL, and combining the three data sets allowed us to obtain a well resolved and strongly supported topology. We interpret our data as showing support for the classical bigeneric system for the family, and call into question several classifications proposed in the past century. The segregate genera Cardiomanes, Hymenoglossum, Serpyllopsis, and Rosenstockia are embedded within Hymenophyllum s.l. Although the deepest relationships within the genus remain uncertain, two subgenera described by Morton do have some support: (1) Sphaerocionium, in which the problematic section Microtrichomanes is embedded; and (2) a diverse Hymenophyllum, including species that were placed originally in Serpyllopsis, Rosenstockia, Hemicyatheon, and Craspedophyllum by Copeland. Subgenus Mecodium appears to be polyphyletic; nevertheless, a subgroup within Mecodium is strongly supported. Several unexpected associations gain support from cytological data and certain morphological characters not previously used to distinguish species groups within Hymenophyllum s.l.     

四十五种叶蝉的染色体研究（同翅目：叶蝉总科）

研究观察了45种中国雄性叶蝉的减数分裂,其中44种的核型为首次报道,染色体数目变化在2n=12～26之间,性别决定均为XO型。从叶蝉总科的组型图来看,该科染色体数目变化在2n=8～28之间,峰值为2n=18(16+XO),另外几种类型2n=16,20,22也有较高的出现频率。科内染色体数目的进化不具有明显的方向性,2n=22(20+XO)是该科的原始核型,易位导致的不均等互换可能是染色体数目进化的主要机制。从精子发生来看叶蝉总科与角蝉总科的关系较为密切,两者的共同特点是：①精母细胞体积较大,显著不同于沫蝉和蝉科;②减数分裂行为及精子变态过程相似;③染色体数目较少,染色体体积较大;④减数分裂前期具有典型的花束期,没有弥散期,因而不同于蜡蝉。但是由于叶蝉总科的染色体变异范围明显大于角蝉总科,而角蝉总科的核型相对较为保守,从核型上来说角蝉总科是比叶蝉总科较为原始的类群。     

SPORE MORPHOLOGY IN THE CYATHEACEAE. I. THE PERINE AND SPORANGIAL CAPACITY: GENERAL CONSIDERATIONS

The literature on cyatheaceous spore morphology relative to the presence of a perine layer is reviewed, and evidence based on a sodium-hydroxide assay is presented indicating that the outer scultpine layer in certain cyatheaceous spores is perine. Perine so defined characterizes Metaxya, paleotropical and certain neotropical species of Sphaeropteris, nearly all species of Alsophila, all species of Nephelea, and certain species of Trichipteris and Cyathea. It is lacking in Lophosoria, many species of Trichipteris and Cyathea, and all species of Cnemidaria. Two major patterns of spore number per sporangium in the family are reported. Lophosoria, Sphaeropteris, Trichipteris, Cyathea, Cnemidaria, and probably Metaxya are characterized by 64-spored sporangia, whereas most species of Alsophila and all species of Nephelea are characterized by 16-spored sporangia. The congruence of this generic distribution of sporangial-capacity types with Tryon's phyletic arrangement of cyatheaceous genera supports the naturalness of his system. The intrasporangial germination of spores retained in dehisced and dispersed sporangia supports the suggestion that decreased spore number per sporangium in Alsophila and Nephelea may relate to the role of the sporangia as dispersal units. The decreased number of spores per sporangium is associated with a trend toward increase in the number of sporangia per sores, with the highest known count approaching 1000 sporangia per sorus. The Alsophila-Nephelea evolutionary line has probably not been ancestral in the phylogeny of the more advanced groups of ferns.     

鳞毛蕨科四个属的染色体数目

报道了鳞毛蕨科的4个属(毛枝蕨属Leptorumohra、石盖蕨属Lithostegia、黔蕨属Phanerophlebiopsis和柳叶蕨属Cyrtogonellum)6种植物的染色体数目及生殖方式。其中柳叶蕨Cyrtogonellum fraxinellum 'n'=123, 2n=123; 离脉柳叶蕨C. caducum 'n'=123; 斜基柳叶蕨C. inaequalis 2n=123; 四回毛枝蕨Leptorumohra quadripinnata n=41; 石盖蕨Lithostegia foeniculacea 2n=164; 长叶黔蕨Phanerophlebiopsis neopodophylla 2n=82。石盖蕨属、黔蕨属和柳叶蕨属的染色体数目为首次报道。结果表明这3个属的染色体基数和鳞毛蕨科中其他属一样均为x=41。细胞学证据支持将石盖蕨属、黔蕨属置于鳞毛蕨科的处理。本文还发现柳叶蕨属与贯众属Cyrtomium的一些种一样, 具有无融合生殖方式, 而其他3个属仅具有在蕨类植物中较为常见的有性生殖方式。     

Chromosome numbers, meiotic behavior, and pollen viability of species of Vriesea and Aechmea genera (Bromeliaceae) native to Rio Grande do Sul, Brazil

Chromosome number, meiotic behavior, and pollen viability were analyzed in 15 species of two genera, Vriesea and Aechmea, native to Rio Grande do Sul, Brazil. This study is the first cytogenetic analysis of these taxa. The chromosome numbers are all n = 25, consistent with the proposed base number of x = 25 for Bromeliaceae. All examined taxa displayed regular bivalent pairing and chromosome segregation at meiosis. Observed meiotic abnormalities include univalents in metaphase I; missing or extra chromosomes and precocious division of centromeres in metaphase II; laggards in telophase I and anaphase II/telophase II. The high pollen viability (>88%) reflects a regular meiosis.     

Spores in situ and problems of the classification of Mesozoic tree ferns

Although tree ferns dominated the Mesozoic flora, their taxonomic relationships are poorly understood at the generic level, and next to nothing can be said of evolutionary trends within the group. At least five genera are recognized based on the remains of spore-bearing structures. However, the dispersed spores belong to the same generalized morphotype, and cannot be assigned to genera based on macroscopic remains of fertile leaves. Electron microscopy of spores in situ may partly resolve these problems providing additional criteria for classification of spore-bearing structures and disperse spores. We studied in situ spores of the Early Cretaceous Alsophilites nipponensis (Oishi) Krassilov, which are comparable to dispersed spores Cyathidites minor Coup. Spore wall micromorphology and ultrastructure indicate their affinities with the modern genus Alsophila R. Brown. Only occasional poorly preserved striate sculptures survive the standard treatment of maceration of the perispore. Our data confirm the primitive status of the species with a great number of spores per sporangium, thick unsculptured exospore consisting of two ultrastructural layers, and the possibility that whole sporangia with unshed spores can function as dispersal units.     

Meiosis in gray voles of the subgenus microtus (rodentia, arvicolinae) and in their hybrids

The results of light and electron microscopic (EM) studies of meiosis in Microtus arvalis males of the karyoform "arvalis" (2n = 46, NFa = 80), in hybrids between the chromosomal forms arvalis and obscurus (2n = 46, NFa = 68), in M. rossiaemeridionalis voles (2n = 54, NFa = 54), and in a hybrid between the species M. rossiaemeridionalis and M. kermanensis (2n = 54, NFa = 54) are presented. SC (synaptonemal complex) karyotypes of the parental forms and the hybrids were constructed on the basis of measurements of the length ofautosomal SCs revealed by the EM analysis in spermatocytes at the stage of middle pachytene. The SC karyotypes of M. arvalis and the hybrids female obscurus x male arvalis consist of 22 synaptonemal complexes of autosomal bivalents and the axial elements of the synaptonemal complexes of the sex chromosomes X and Y. The SC karyotypes of M. rossiaemeridionalis and the hybrid M. rossiaemeridionalis x M. kermanensis consist of 26 synaptonemal complexes of autosomal bivalents and a sex bivalent; they differ only in the length of the Y chromosome axis (Y chromosome in the hybrid was inherited from M. kermanensis). Asynaptic configurations of the autosomal SCs were not observed in the hybrids. The SC axial elements of the X and Y chromosomes in the parental forms and in the hybrids were located close to each other throughout pachytene, but they did not form a synaptic region. The normal synapsis in sterile hybrids (M. rossiaemeridionalis x M. kermanensis) and the behavior of the sex chromosomes in meiosis in fertile and sterile hybrids are discussed in the context of specific features of meiosis and reproductive isolation.     

M chromosome of the wild silkworm, Bombyx mandarina (n = 27), corresponds to two chromosomes in the domesticated silkworm, Bombyx mori (n = 28).

Chromosomes of Bombyx mori (n = 28) and of Bombyx mandarina (n = 27) were studied cytogenetically to resolve the origin of the large M chromosome in the Japaneses type of B. mandarina. In the F1 progeny from the reciprocal cross between B. mandarina and B. mori, the mitotic chromosome number was 2n = 55, and a chromosome configuration of 26 bivalents plus 1 trivalent was observed at metaphase I of germ cells. The trivalent chromosome consisted of the M chromosome from B. mandarina and two chromosomes from B. mori. When males of B. mori were mated to the F1 females, nuclei with two types of chromosome number (2n = 55 and 2n = 56) and two sets of chromosome pairs (26 bivalents plus 1 trivalent versus 28 bivalents) were observed in the metaphase I stage. Linkage analysis showed that the 14th chromosome of B. mori was involved in these two types of chromosome segregation. This result indicates that the M chromosome in B. mandarina arose from a fusion between a chromosome corresponding to the 14th linkage group and another, yet unidentified linkage group.     

白菜型油菜(B. Chinensis)与海甘蓝(Crambe abyssinica)属间杂种的细胞学观察

白菜型油菜与海甘蓝属间杂种F₁代染色体数目早期为2n=55。经多代组培无性繁殖后,杂种染色体大量减少,数目变化于25～28条间,平均为26条。杂种PMC减数分裂中,平均配对构型为0.06Ⅲ+11.26Ⅱ+3.80Ⅰ,二价体数目变化于8～13之间,以10Ⅱ、11Ⅱ、12Ⅱ的细胞出现频率较高,分别为26.60%,23.91%和30.98%,单价体数目变化于0～8个之间。杂种染色体数目减少,二价体个数较高,这说明海甘蓝染色体在大量丢失的同时,白菜型油菜的染色体组发生了自然加倍。在后期Ⅱ观察到三分体、染色体落后、染色体桥等异常现象。      

Sporangium Exposure and Spore Release in the Peruvian Maidenhair Fern (Adiantum peruvianum,Pteridaceae)

We investigated the different processes involved in spore liberation in the polypod fern Adiantum peruvianum (Pteridaceae). Sporangia are being produced on the undersides of so-called false indusia, which are situated at the abaxial surface of the pinnule margins, and become exposed by a desiccation-induced movement of these pinnule flaps. The complex folding kinematics and functional morphology of false indusia are being described, and we discuss scenarios of movement initiation and passive hydraulic actuation of these structures. High-speed cinematography allowed for analyses of fast sporangium motion and for tracking ejected spores. Separation and liberation of spores from the sporangia are induced by relaxation of the annulus (the ‘throwing arm’ of the sporangium catapult) and conservation of momentum generated during this process, which leads to sporangium bouncing. The ultra-lightweight spores travel through air with a maximum velocity of ~5 m s^-1, and a launch acceleration of ~6300g is measured. In some cases, the whole sporangium, or parts of it, together with contained spores break away from the false indusium and are shed as a whole. Also, spores can stick together and form spore clumps. Both findings are discussed in the context of wind dispersal.     

十二种沫蝉的染色体研究 (同翅目： 沫蝉总科)

观察了同翅目头喙亚目12种沫蝉雄性的染色体数目和减数分裂行为。通过对沫蝉总科现有核型资料的分析,认为沫蝉总科的核型特点是：①染色体较小,数目较多,总科内染色体数目变化范围较大,众数为2n=26（24+XO）;②染色体的易位现象极为普遍,因此可以推测,通过染色体的易位导致染色体数目增加是核型进化的主要机制;③减数分裂前期Ⅰ具有典型的花束期,但没有弥散期。因此从精子发生来看沫蝉总科与叶蝉总科、角蝉总科和蝉总科的关系更为密切,而与蜡蝉总科的关系较远。头喙亚目的亲缘关系可能是：蜡蝉总科+{蝉总科+［沫蝉总科+（叶蝉总科+角蝉总科）］}。     

Karyotype evolution by centromeric fission inZamia (Cycadales)

The chromosome numbers of several species ofZamia from Mexico are reported.Z. paucijuga, distributed from central Oaxaca to Nayarit, has been found to have 2n = 23, 25, 26, 27 and 28. 2n = 28 is the highest chromosome number yet found in the cycads. Karyotypes of this species differ principally in the number of telocentric and metacentric chromosomes present in each; 2n = 23, 25, 26, 27 and 28 were found to have 5, 3, 2, 1 and 0 metacentric and 8, 12, 14, 16 and 18 telocentric chromosomes, respectively.Z. fischeri has been found to be 2n = 16,Z. furfuracea andZ. loddigesii 2n = 18.Zamia paucijuga on the basis of morphological and ecological characteristics, is considered to be an advanced member of this genus. Chromosome and karyotype evolution inZ. paucijuga may have occurred by centromeric fission of metacentric chromosomes; the karyotypes ofZ. paucijuga are strongly asymmetrical, suggesting that they evolved recently.