BIOSYSTEMATIC INVESTIGATIONS IN THE GENUS AGROPYRON. I. CYTOLOGICAL STUDIES OF SPECIES KARYOTYPES

Schulz -Schaeffer , Jurgen (Montana State Coll., Bozeman), and Peter Jurasits . Biosystematic investigations in the genus Agropyron. I. Cytological studies of species karyotypes. Amer. Jour. Bot. 49(9): 940–953. Illus. 1962.—Twenty-five species of the genus Agropyron are analyzed cytologically in this presentation. Accession numbers, names of collectors, locations where seed was collected, and observed chromosome numbers are listed. Chromosome numbers of A. panormitanum (2n = 28), A. lolioides (2n = 58), A. brachyphyllum (2n = 42), A. ciliatiflorum (2n = 28), A. kosanini (2n = 56), A. pseudorepens (2n = 28), A. squamosum (2n = 42), and A. subulatum (2n = 56) are reported. No previous counts in these species are known to the authors. Chromosome counts of A. caespitosum (2n = 42) and A. elongatiforme (2n = 58), deviate from previous reports. Idiograms of all species and photomicrographs of mitotic metaphase root tip cells of 14 species are presented. The distribution of 11 satellite-chromosome types in 25 Agropyron species is shown in Table 2. The proportions of these 11 satellite-chromosome types are recorded in Table 3. The significance of these satellite chromosomes as indicator chromosomes for genome relationships is discussed together with the pertinent literature.     

蟛蜞菊属和孪花菊属(菊科-向日葵族)的细胞学研究

研究了菊科向日葵族鳢肠亚族蟛蜞菊属(Sphagneticola O. Hoffm.)和孪花菊属(Wollastonia DC. ex Decne.)各2种植物的染色体数目和染色体形态。蟛蜞菊[S. calendulacea (L.) Pruski]的染色体数目为2n=50, 核型公式为2n=18m+30sm+2st,南美蟛蜞菊[S. trilobata (L.) Pruski]的染色体数目为2n=56, 核型公式为2n=24m+28sm+4st, 孪花菊[W. biflora (L.) DC.]的染色体数目为2n=30,核型公式为2n=24m+4sm+2st,山孪花菊[W. montana (Blume) DC.]的染色体数目为2n=74, 核型公式为2n=37m+31sm+6st。根据上述结果并结合以前的有关资料,推测蟛蜞菊属的染色体基数可能为x=14和x=25,而不应是x=15。该属的3个新世界热带种[S. brachycarpa (Baker) Pruski、S. gracilis (Richard) Pruski和南美蟛蜞菊]可能都基于x=14, 其中S. gracilis为二倍体(2n=2x=28), S. brachycarpa和南美蟛蜞菊为四倍体(2n=4x=56); 唯一的亚洲种(蟛蜞菊)可能是基于x=25的二倍体(2n=2x=50)。染色体资料不支持将山孪花菊(x=37)这一植物置于孪花菊属(x=15)中。     

GENOME RELATIONSHIPS OF INTRA- AND INTERSECTIONAL SPECIES HYBRIDS OF ONCIDIUM TRIQUETRUM

Chromosome numbers and meiotic behavior of six interspecific hybrids of Oncidium triquetrum were investigated. The somatic chromosome number of O. triquetrum (section Oncidium) x O. variegatum (section Oncidium) was 2n = 42. O. triquetrum x O. splendidum (section Plurituberculata) and O. triquetrum x O. cebolleta (section Cebolletae) were 2n = 39 and O. triquetrum x O. floridanum (section Planilabra), O. triquetrum x O. flexuosum (section Synsepala) and O. triquetrum x O. onustum, (section Onusta) were 2n = 49. The intersectional species hybrid, O. triquetrum x O. variegatum, showed fairly regular meiosis with a mean of 20.4 bivalents, indicating strong homology of parental genomes. In contrast, highly irregular meiosis with poor chromosome pairing was observed in all intersectional hybrids. The mean number of bivalents ranged from 1.8 to 9.5. On the basis of chromosomal affinity at metaphase I, O. triquetrum showed a close relationship to O. variegatum in the same section, but more distant relationships to O. splendidum, O. cebolleta, O. floridanum, O. flexuosum and O. onustum belonging to different sections.     

Chromosomes ofLauxaniidae andChamaemyiidae (Diptera)

Chromosome numbers of 28 species ofLauxaniidae range from 2n=8 to 2n=12 with one having 2n=8, three having 2n=10 and 24 having 2n=12. Total complement length averaged 58.1 μ for 71 complements measured. The species studied in detail are classified according to chromosome number and morphology. The two Chamaemyiid species studied have 2n=6 and 2n=15, the latter including several microchromosomes. The karyotypical reorganizations have clearly involved inversions and translocations and a progressive reduction of chromosome numbers.     

Chromosome counts of some MongolianPotentilla species

Chromosome numbers are reported for 26 collections representing 12 MongolianPotentilla taxa. The first chromosome records are reported forP.[x]agrimonioides Bieb. (2n = 42, 49–50),P. [x]chionea Soják (2n = 64, 74+1B, 84–86, 86),P. [x]stepposa Soják (2n = 62) andP. supina, L. subsp.costata Soják (2n = 42). Tetraploid and pentaploid populations ofP. sericea L. (2n = 28, 34, 35) and hexaploid populations ofP. omithopoda Tausch (2n = 42) were found in Mongolia. Chromosome numbers of the following species were confirmed:P. longifolia Willd. exSchlecht. (2n = 14),P. multifida L. (2n = 28),P. sanguisorba Willd. exSchlecht. (2n = 28),P. sericea L. (2n = 42),P. supina L. subsp.paradoxa (Nutt.) Soják (2n = 28 China) andP. tanacetifolia Willd. exSchlecht. (2n = 28). Taxonomic observations are briefly discussed.     

New Karyotypes and Some Considerations about the Chromosomal Diversification of Ctenomys minutus (Rodentia: Ctenomyidae) on the Coastal Plain of the Brazilian State of Rio Grande do Sul

The dominant mammals occupying the subterranean niche in South America are rodents of the genus Ctenomys, which form a large group of 56 species with chromosome numbers ranging from 2n= 10 to 70. In southern Brazil, Ctenomys minutus is the species with the widest geographic distribution, inhabiting sandy fields and dunes extending from Jaguaruna beach in the state of Santa Catarina to the town of São José do Norte in the state of Rio Grande do Sul. Eleven karyotypes (2n= 42; 2n= 46a; 2n= 46b; 2n= 47a; 2n= 47b; 2n= 48a; 2n= 48b; 2n= 49a; 2n= 49b; 2n= 50a and 2n= 50b) were described for this species and zones of hybridization are also known. A sample of 51 C. minutus specimens was collected from five sampling sites about 20 km apart along the coastal plain of Rio Grande do Sul between the municipalities of Tavares (31°23′S 51°09′W) and São José do Norte (31°52′S 51°54′W). We were able to extend the known geographic distribution of C. minutus by 90 km, from Tavares southwards to São José do Norte. During our study we found five karyotypes (2n= 46b, 47b, 48b, 49b and 50b), four of which (2n= 47b, 48b, 49b and 50b) have not previously been described for this species.     

Meiotic configuration and chromosome number in some Iranian species of Elymus L. and Agropyron Gaertner (Poaceae: Triticeae)

Chromosome numbers and analyses of meiotic metaphase I are reported for the following taxa: Agropyron cristatum subsp. incanum (2n= 42), A. cristatum subsp. pecttnatum (2n=28 – 33), Elymus elongatus subsp. ponticus (2n= 69, 70), E. hispidus var. hispidus (2n= 41 43), var. podperae (2n= 42) and var. villosus (2n= 41, 42), E. libanoticus (2n= 14), E. pertenuis (2n= 28, 28+1B), E. repens (2n= 42), E. transhyrcanus (2n= 40–42), E. hispidus var. villosus x E. cf. repens (2n= 42). Chromosome numbers only are reported for the following taxa: E. gentri (2n= 41, 42), E. nodosus subsp. dorudicus (2n= 28), and E. elongatiformis (2n= 56, 57). The haploid genomic constitution SP is reported for Elymus pertenuis. Variable chromosome numbers (2n= 28–32) were observed in the meiotic metaphase I within single anthers of Agropyron cristatum subsp. pectinatum, and the supernumerary chromosomes in this taxon are assumed to have originated from crosses with hexaploids. Partial elimination of these supernumerary chromosomes probably occurs during archesporial mitotic divisions or at an early stage in the meiotic cycle. A hybrid, morphologically intermediate between E. hispidus and E. repens, was obtained from a seed of E. hispidus collected in the field. The meiotic metaphase I configuration in this E. hispidus hybrid suggests that the pollen parent may itself be a hybrid or hybrid derivative of E. repens x E. hispidus.     

Karyological data of 47 accessions of 28 Artemisia (Asteraceae, Anthemideae) species from Iran, with first new reports for Iranian populations and first absolute counts in three species

Somatic chromosome numbers of 47 accessions representing 28 Artemisia species are provided from Iran. Two basic chromosome numbers, x = 8, 9, each with diploid, tetraploid and hexaploid levels, were found. Different chromosome numbers, 2n = 16, 16 + 1B, 16 + 5B, 32, 48, and 2n = 18, 18 + 1B, 19, 36, 36 + 1B, 36 + 2B, 37, 49 + 2B, 49 + 3B, 51 + 6B, 54, 54 + 1B, 54 + 3B, 54 + 5B, in studied accessions were identified. Chromosome numbers are reported for the first time in three species, counts in four species are new for Iran, and other counts have been thoroughly compared to previous data. Forty percent of the studied accessions are polyploid and B-chromosome(s) are reported in 17 % of accessions. Aneuploidy and aneusomy are other relevant cytological phenomena. Some karyological parameters, total karyotype length, karyotype formula, mean value of centromeric indices, mean arm ratio, A ₁ and A ₂ indices, were estimated to characterize the karyotypes numerically. A ₁ and A ₂ indices showed that karyotypes tend to be asymmetric in polyploid and dysploid taxa. PCA analysis of all karyological parameters has shown some systematic and evolutionary implications. The consideration of all these chromosome numbers and cytogenetic mechanisms has led us to infer the different patterns of chromosomal evolution in the genus.     

中国桤木属植物的细胞学研究(Ⅰ)

对桦木科桤木属自然分布于中国西南地区的4个种进行了细胞学研究,结果分别为:川滇桤木K(2n)=56=30m+24sm+2st;桤木K(2n)=56=38m+16sm+2M;蒙自桤木K(2n)=56=28m+26sm+2st;毛桤木K(2n)=56=42m+14sm;其中川滇桤木的核型属于2A型,其余均为2B型;上述种类染色体数目均为2n=56。4种材料的核型均为首次报道。     

Chromosome number variability in central european members of the<Emphasis Type="Italic">Festuca ovina</Emphasis> and<Emphasis Type="Italic">F. pallens</Emphasis> groups (sect.<Emphasis Type="Italic">Festuca</Emphasis>)

Chromosome numbers for 98 plants ofF. pallens, 19 ofF. psammophila, F. belensis andF. vaginata, and 44 ofF. ovina (originating from Austria, the Czech Republic, Germany, Slovakia and Latvia) are given. In addition to theF. ovina andF. pallens groups, chromosome counts for the following taxa are also reported:F. alpestris (2n=14) reported for the first time in this work,F. amethystina subsp.amethystina (2n=28),F. brevipila (2n=42),F. cinerea (2n=28),F. rupicola subsp.rupicola (2n=42) andF. versicolor subsp.versicolor (2n=14).InF. pallens, two ploidy levels (2n=2x=14+0-1B, 2n=4x=28+0-1B) as well as two natural triploid plants (2n=21+0-1B), were found. In addition to the fourF. pallens types that have been distinguished in Austria, one new tetraploid type (F. pallens “scabrifolia”) from the Czech Republic and Germany is reported and its taxonomy is discussed. The distributions of the Oberösterreich-Niederösterreich and Pannonisches-HügellandF. pallens types outside of Austria are documented.Only the diploid chromosome number (2n=14) was found inF. psammophila andF. vaginata. Chromosome numbers forF. psammophila subsp.muellerstollii andF. belensis (both 2n=14) were determined here for the first time. Two ploidy levels, 2n=14+0-5B corresponding toF. ovina subsp.ovina and 2n=28 corresponding toF. ovina subsp.guestphalica andF. cf.duernsteinensis were confirmed inF. ovina. Differences in chromosome structure (simple and multiple secondary constrictions) betweenF. pallens as opposed toF. psammophila andF. vaginata are discussed. A complete survey of published chromosome counts for Central European species from theF. ovina andF. pallens groups is included.     

Different genome-specific chromosome stabilities in synthetic Brassica allohexaploids revealed by wide crosses with Orychophragmus

Background and Aims

In sexual hybrids between cultivated Brassica species and another crucifer, Orychophragmus violaceus (2n = 24), parental genome separation during mitosis and meiosis is under genetic control but this phenomenon varies depending upon the Brassica species. To further investigate the mechanisms involved in parental genome separation, complex hybrids between synthetic Brassica allohexaploids (2n = 54, AABBCC) from three sources and O. violaceus were obtained and characterized.

Methods

Genomic in situ hybridization, amplified fragment length polymorphism (AFLP) and single-strand conformation polymorphism (SSCP) were used to explore chromosomal/genomic components and rRNA gene expression of the complex hybrids and their progenies.

Key Results

Complex hybrids with variable fertility exhibited phenotypes that were different from the female allohexaploids and expressed some traits from O. violaceus. These hybrids were mixoploids (2n = 34–46) and retained partial complements of allohexaploids, including whole chromosomes of the A and B genomes and some of the C genome but no intact O. violaceus chromosomes; AFLP bands specific for O. violaceus, novel for two parents and absent in hexaploids were detected. The complex hybrids produced progenies with chromosomes/genomic complements biased to B. juncea (2n = 36, AABB) and novel B. juncea lines with two genomes of different origins. The expression of rRNA genes from B. nigra was revealed in all allohexaploids and complex hybrids, showing that the hierarchy of nucleolar dominance (B. nigra, BB > B. rapa, AA > B. oleracea, CC) in Brassica allotetraploids was still valid in these plants.

Conclusions

The chromosomes of three genomes in these synthetic Brassica allohexaploids showed different genome-specific stabilities (B > A > C) under induction of alien chromosome elimination in crosses with O. violaceus, which was possibly affected by nucleolar dominance.Key words: Synthetic Brassica allohexaploids, Orychophragmus violaceus, intergeneric hybrids, genomic in situ hybridization, amplified fragment length polymorphism, single-strand conformation polymorphism, chromosome elimination, chromosome stability, nucleolar dominance     

Chromosome numbers of dominican compositae

Chromosome numbers of 34 species of Dominican Compositae in 26 genera and nine tribes are reported. First counts are given forCoreopsis buchii (2n = 64),Lagascea mollis (2n = 34),Spilanthes urens (n = 16),Liabum subacaule (n = 18),Eupatorium sciatraphes (2n = 40),Hieracium gronovii (2n = 18),Vernonia buxifolia (2n = 34),V. sprengeliana (2n = 34),V. racemosa (2n = 28), andChaptalia leiocarpa (2n = 48,ca. 58). Our report forNarvalina domingensis (2n = 120) is the first for any species of this genus.     

RAPD-Based Analysis of Introgression of Barley Genetic Material into the Genome of Alloplasmic Wheat Lines (Hordeum geniculatumAll./Triticum aestivum L.)

Genomes of three alloplasmic wheat lines obtained on the basis of barley–wheat hybrid Hordeum geniculatumAll. (2n = 28) ×Triticum aestivumL. (2n = 42)(Pyrotrix 28) were examined using random amplified polymorphic DNA (RAPD) analysis. Line L-29 was obtained after first backcross of the initial hybrid with the wheat variety Pyrotrix 28 and ten subsequent self-pollinating generations. This line was represented by euploid plants with typical to the common wheat chromosome number (2n = 42), as well as by aneuploids, which contained an additional telocentric chromosome in the main karyotype (2n = 42 + t). Lines L-26 and L-27 were obtained by two backcrosses of one BC₁ plant with the wheat variety Novosibirskaya 67 and one subsequent self-polination of one BC₃ plant. Chromosome number in all these plants corresponded to 2n = 40 + 4t. RAPD analysis was carried out using seven primers, which were previously proved to be effective for identification of the barley genome fragments within hybrid genomes of alloplasmic lines. The presence of barley genome fragments in line L-29 was revealed by use of five primers, while in lines L-26 and L-27 these fragments were detected by use of one primer. The significant difference in the number of barley RAPD fragments in the genomes of alloplasmic lines obtained at different backcrossing stages suggests more intense displacement of barley genome during backcrossing compared to self-pollination in BC₁ plants.     

Annotated chromosome counts of czechoslovak plants (31–60) (Materials for “Flóra ČSSR”—3)

Chromosome counts of the following 30 taxa (106 populations) are given:Betonica officinalis (2n=16);Bidens frondosus (2n=48);Calamagrostis arundinacea (2n=28+0–2B);Dianthus carthusianorum subsp.latifolius (2n=30);Festuca gigantea (2n=42, 42+2B);Hypericum perforatum (2n=32);Koeleria macrantha (2n=28);Kohlrauschia prolifera (2n=30);Lilium martagon (2n=24+0–2B);Melica ciliata (2n=18);Poa remota (2n=14);Ranunculus polyanthemos (2n=16);R. sardous subsp.sardous (2n=16);Roegneria canina (2n=28+0–1B);Rudbeckia laciniata (2n=76);Scabiosa canescens (2n=16);Serratula tinctoria (2n=22);Seseli elatum subsp.heterophyllum var.beckii (2n=18);S. hippomarathrum (2n=20);Thlaspicaerulescens caerulescens subsp.tatrense (2n=14);Trifolium alpestre (2n=16);T. avense (2n=14);T. medium (2n=79, 80+0–2B, 82);T. rubens (2n=16);Veronica officinalis subsp. alpestris (2n=36);Vincetoxicum hirundinaria (2n=22);Vulpia bromoides (2n=14);Zerna benekenii (2n=28)Z. monoclada (2n=28+0–8B);Z. ramosa (2n=42). Remarks on taxonomy, nomenclature and chorology for some of these taxa are given.     

Chromosome numbers and taxonomic notes for Mexican coreopsis,sections electra and pseudoagarista (compositae: heliantheae)

Notes on morphology and chromosome numbers are given for several species of MexicanCoreopsis, most of which were poorly known prior to recent collections.Coreopsis parvifolia (sectionElectra) is a large, shrubby, octoploid (2n=112) species apparently restricted to a small area in Puebla. It andC. cuneifolia (a diploid with 2n=28) appear to be closely related but differ by a number of morphological features in addition to ploidy level. A first chromosome report forC. pringlei (sectionPseudoagarista) shows this rare species to be diploid (2n=26), a fact which is in agreement with the base number of the section, i.e.,x=13. Additional collections of the very rareC. rudis andC. mcvaughii show them to be similar yet distinct species. Chromosome determinations forC. petrophila from Nayarit and Durango agree with previous counts for Jalisco populations of the species, i.e., 2n=26. Considerable morphological variation exists within this species but no subspecific entities are recognized.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.     

Meiotic configuration and chromosome number in some Iranian species of Ely mus L. and Agropyron Gaertner (Poaceae: Triticeae)

Chromosome numbers and analyses of meiotic metaphase I are reported for the following taxa: Agropyron cristatum subsp. incanum (2 n = 42), A. cristatum subsp. pecttnatum (2 n =28 – 33), Elymus elongatus subsp. ponticus (2 n = 69, 70), E. hispidus var. hispidus (2 n = 41 43), var. podperae (2 n = 42) and var. villosus (2 n = 41, 42), E. libanoticus (2 n = 14), E. pertenuis (2 n = 28, 28+1B), E. repens (2 n = 42), E. transhyrcanus (2 n = 40–42), E. hispidus var. villosus x E. cf. repens (2 n = 42). Chromosome numbers only are reported for the following taxa: E. gentri (2 n = 41, 42), E. nodosus subsp. dorudicus (2 n = 28), and E. elongatiformis (2 n = 56, 57). The haploid genomic constitution SP is reported for Elymus pertenuis. Variable chromosome numbers (2 n = 28–32) were observed in the meiotic metaphase I within single anthers of Agropyron cristatum subsp. pectinatum , and the supernumerary chromosomes in this taxon are assumed to have originated from crosses with hexaploids. Partial elimination of these supernumerary chromosomes probably occurs during archesporial mitotic divisions or at an early stage in the meiotic cycle. A hybrid, morphologically intermediate between E. hispidus and E. repens , was obtained from a seed of E. hispidus collected in the field. The meiotic metaphase I configuration in this E. hispidus hybrid suggests that the pollen parent may itself be a hybrid or hybrid derivative of E. repens x E. hispidus.     

Genome size and chromosome number in the New Zealand species of Schoenus (Cyperaceae)

Schoenus (Cyperaceae) has holocentric chromosomes. Chromosome numbers were counted and nuclear DNA amounts were measured for all the New Zealand species of the genus. Chromosome numbers ranged from 2n = 8 to c. 2n = 90. Two chromosome races, with 2n = 28 and 2n = 56, were found in S. pauciflorus. Flow cytometry using propidium iodide‐stained nuclei was used to measure genome size. A 14.8‐fold variation in 2C DNA content was found, with values ranging from 1.33 to 19.71 pg/2C nucleus. Phylogenetic trees based on sequence variation in the internal transcribed spacer (ITS) region of the 45S ribosomal DNA locus were constructed using several phylogenetic models to reveal possible evolutionary relationships among the New Zealand Schoenus spp. and a sample of Australian Schoenus spp. Analysis revealed heterogeneity of chromosome number, size and DNA C value within clades. Meiosis in four species showed only bivalent formation at metaphase I. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 555–564.     

Features of the formation of self-fertile euploid lines (2n = 42) by self-pollination of the 46-chromosome barley-wheat BC1 hybrid Hordeum marinum subsp. gussoneanum Hudson (= H. geniculatum All.) (2n = 28) × Triticum aestivum L. (2n = 42)

We studied some features of the development of self-fertile 42-chromosome lines on the base of self-pollination progeny of 46-chromosome plants obtained by backcrossing of barley-wheat hybrids Hordeum marinum subsp. gussoneanum Hudson (= H. geniculatum All.) (2n = 28) × Triticum aestivum L. (2n = 42). The stabilization of karyotypes, resulting in 42-chromosome plants of the wheat type was generally completed by generation BC₁F₁₀. The plants of all self-pollination progenies, including BC₁F₁₀, showed some phenotypic traits characteristic of wild barley. Plants of BC₁F₁₀ with the chromosome sets 2n = 42 and 2n = 42 + t were analyzed by RAPD with a set of 115 primers. Fragments of the wild barley genome were detected in RAPD patterns with 19 primers. Cross-hybridization confirmed that these fragments belonged to the wild barley genome. We raised four phenotypically different 42-chromosome lines from grains obtained from plants of generation BC₁F₁₀, and these lines proved to be cytogenetically stable and self-fertile when grown in the field.     

Chromosome numbers of Orthocarpus and related monotypic genera (Scrophulariaceae: Subtribe castillejinae)

A chromosome number ofn=12 is reported for the three monotypic genera of subtribe Castillejinae:Clevelandia beldingii, Gentrya racemosa, andOpicopephalus angustifolius. Chromosome numbers ofOrthocarpus correspond mostly with current infrageneric classification. SubgenusTriphysaria hasn=11.Orthocarpus sectionsCastillejoides andCordylanthoides, which are closely related toCastilleja (x=12) and the three monotypic genera above, haven=12 with aneuploid reductions ton=10 inO. linearilobus andn=11 inO. lacerus (a species also withn=12). Tetraploids are found in two species.O. brevistylus (n=24) andO. hispidus (n=12, 24). The polyploid.O. laciniatus (n=36, 48) of Peru is postulated to be of hybrid origin between a species ofCastilleja andOrthocarpus attenuatus. SubgenusOrthocarpus sectionOrthocarpus, which hasn=14 in all species except.O. bracteosus (n=15), stands apart both morphologically and in chromosome number from the remainder of the genus.