Carpological Characters Research Articles

Possible phylogenetic relationships of Gymnarrhena (Gymnarrheneae, Gymnarrhenoideae, Asteraceae) based on comparative carpology

The amphicarpic annual Gymnarrhena micrantha (Asteraceae), inhabiting the deserts of North Africa to the Middle East, is interesting not only for its unusual biology, but also for its peculiar morphological structure. The monotypic genus Gymnarrhena, with uncertain position in the Asteraceae for a long time, is currently included in the tribe Gymnarrheneae and a separate subfamily Gymnarrhenoideae. The objective of the study was to examine fruit and seed structure in G. micrantha to consider possible phylogenetic relationships of the genus based on carpological characters. It was found that fruit structural organization in G. micrantha, with simply arranged pericarp and seed coat, parenchymatous and poorly differentiated on the tissue, corresponds to that of the early diverging evolutionary lines of Asteraceae, in which the states of these characters are considered to be plesiomorphic in the family. It has been suggested that taxonomically important structural features of Gymnarrhena cypsela include the structure of the apical region, pericarp and seed coat, the place of pappus attachment and its structure, the type of pappus hygroscopic tissue, and morphophysiological features of receptacular bracts surrounding the cypsela. The study did not confirm the relationship of Gymnarrhena with the tribes Corymbieae and Cichorieae, the seed coat of which differs markedly from the seed coat of Gymnarrhena in the presence of fine porosity of the exotesta cell walls. A set of the anatomical and carpological characters indicates a possible relationship of Gymnarrhena with the basal subtribes Cardopatiinae and Carlininae of the tribe Cardueae. The morphophysiological adaptation of G. micrantha and Cousiniopsis atractyloides cypselae to dissemination in desert conditions, similar in structural details, apparently suggests a close relationship between the subfamilies Gymnarrhenoideae and Carduoideae.

Fruit characters in early divergent Arecaceae-Arecoideae: comparative carpology of Hyophorbe Gaertn. (Chamaedoreeae)

ABSTRACT The phylogenetic relationships of two early divergent tribes of Arecoideae (Chamaedoreeae and Iriarteeae) remain poorly resolved. Nevertheless, in some phylogenetic reconstructions the genus Hyophorbe is recognized as the earliest divergent genus of the subfamily. Hence, the fruit anatomy, histogenesis and organogenesis of Hyophorbe species is studied here, with the aim of revealing the key carpological characters of early divergent taxa of Arecaceae – Arecoideae. The multilayered pericarp consists mostly of the mesocarp (parenchymatous cells with different inclusions) and differentiates into outer and inner zones from its early developmental stages (in H. lagenicaulis an additional 1–3 layered inner subdermal zone develops in the inner mesocarp). The one-layered endocarp is represented either by palisade sclereids, lignification of which begins at late developmental stages, or by almost unspecialized thin-walled epidermal cells. The fruits of Hyophorbe species are referred to two different morphogenetic types: the pyrenariums of the Ilex type with a one-layered lignified endocarp (H. lagenicaulis, H. verschaffeltii, H. indica) and berries of the Nuphar type without a continuous lignified zone in the pericarp (H. vaughanii and H. amaricaulis). It is shown that the lignified palisade endocarp could be interpreted as a plesiomorphic character state of Ceroxyloideae – Arecoideae clade, whereas a non-lignified endocarp could be treated as an apomorphy of this clade or a plesiomorphic character state of the family Arecaceae.

Tetrataenium shrirangii a new species from the Western Ghats, India

Tetrataenium shrirangii C.Rekha, Gosavi, Manudev & Dalavi, a new species of Apiaceae is described here from the lateritic plateau of Chaukul Village in Sindhudurg District in Maharashtra. The new species is closely similar to T. birmanicum (Kurz) Manden., a species found in Assam (northeast India) to Thailand, by its apically inflexed petals, persistent calyx, and style, but is distinguished from the latter by the presence of narrow, oblong‐ovate or ovate‐lanceolate to elliptic or round leaflets, 2–6 ovate involucre bracts with 1–3 lobed or rounded apex, 3–5 elliptic‐oblong involucel, and other carpological characters such as ovoid‐ellipsoid fruits with narrowly winged lateral ribs, 4–6 unequal commissural vittae and with slightly concave commissural face. A detailed description, diagnostic characters, photo plates and illustrations are provided for easy identification of this species.

Fruit and seed structure in the ANA-grade angiosperms: Ancestral traits and specializations.

The representatives of the ANA-grade angiosperms demonstrate a diverse pattern of morphological characters, but their apocarpous gynoecium (except in Nymphaeaceae), composed of at least partly ascidiate carpels, the four-nucleate and four-celled female gametophyte, and the diploid endosperm (except in Amborella) are inferred to be plesiomorphies. Since the structure of fruits in Austrobaileyales is under-investigated, this research aims to fill this gap in these data, describing the carpological characters of ANA-grade taxa, and potentially illuminating the ancestral fruit and seed types of angiosperms. The pericarp and seed coat anatomy was studied with light microscopy. The character optimization was carried out using WinClada software. The fruits of Austrobaileya, Trimenia, Kadsura, and Schisandra are determined to be apocarpous berries of the Schisandra type, with a parenchymatous pericarp and mesotestal (Austrobaileya) or exomesotestal seeds (other genera). Most inferred scenarios of fruit evolution indicate that the apocarpous berry is either the most probable plesiomorphic fruit type of all angiosperms, or that of all angiosperms except Amborellaceae. This inference suggests the early origin of the berry in fruit evolution. The plesiomorphic seed type of angiosperms according to reconstructed scenarios of seed type evolution was either a seed lacking a sclerenchymatous layer or an exotestal seed. The current research indicates that an apocarpous berry, and not a follicle, is a probable plesiomorphic character of the ANA-grade taxa and of angiosperms as a whole.

Taxonomic significance of fruit and seed structure in the subfamily Cassinoideae (Celastraceae)

The paper reports on the macro- and micromorphology and anatomy of fruits and seeds in 19 species representing 13 African and American genera of subfamily Cassinoideae (family Celastraceae). A diversity of carpological characters was identified: fruit type and peculiarities of its dehiscence; presence of trichomes on fruit surface; subtypes of pyrenaria (polymerous, olygomerous, dimerous, pseudomonomerous); number of seeds, morphology of the seed (shape, presence/absent of outgrowths); the ratio of parenchymatous (ground) and sclerenchymatous (supporting) cell elements in the pericarp, cell shape and number of cell layers in the seedcoat. Fruit and seed characters support the differences between genera, also reflect on their possible relationship. The traits revealed are also used to substantiate and support the recent taxonomic revision of many genera, primarily African. The array of revealed fruit and seed traits strongly support previous taxonomic decisions to define most genera of the subfamily Cassinoideae in a narrow sense. It is especially relevant in the taxonomy of the African representatives, characterizing among others the distinct indigenous flora of the Cape Floristic Region.

Carpological Analysis of Two Endemic Italian Species: Pimpinella anisoides and Pimpinella gussonei (Apiaceae).

This study aims to clarify the taxonomic doubts, which have varied over the centuries, on the only two endemic species of the genus Pimpinella growing in Italy: P. anisoides and P. gussonei. For this purpose, the main carpological characters of the two species were examined, analyzing the external morphological traits and their cross-sections. Fourteen morphological traits were identified, obtaining datasets for the two groups using 40 mericarps (20 per species). The obtained measurements were subjected to statistical analysis (MANOVA and PCA). Our results highlight that at least 10 of the 14 morphological traits analyzed support the distinction between P. anisoides and P. gussonei. In particular, the following carpological characters are very significant ways to distinguish between the two species: monocarp width and length (Mw, Ml), monocarp length from base to maximum width (Mm), stylopodium width and length (Sw, Sl), length/width ratio (l/w) and cross-section area (CSa). In particular, the fruit of P. anisoides is larger (Mw 1.61 ± 0.10 mm) than that of P. gussonei (Mw 1.27 ± 0.13 mm), the mericarps of the first species are longer (Ml 3.14 ± 0.32 vs. 2.26 ± 0.18 mm) and the cross-section area (CSa) of P. gussonei is larger (0.92 ± 0.19 mm) than that of P. anisoides (0.69 ± 0.12 mm). The results also highlight the importance of the morphological traits of the carpological structures for the specific discrimination of similar species. The findings of this study contribute to an evaluation of the taxonomic significance of this species within the genus Pimpinella, and also provide valuable information for the conservation of these two endemic species.

Floral micromorphological, palynological, and carpological characteristics of the rare endemic Korean species Peucedanum miroense (Apiaceae).

Peucedanum miroense K. Kim, H.-J. Suh & J.-H. Song, recently reported as a new species endemic to South Korea, is distributed on mountains Duta-san and Swinŭm-san in Gangwon Province, South Korea. Given its conspicuously limited habitat range and small populations, this species qualifies as an endangered Red List species. In this study, we undertook a detailed investigation of the floral micromorphological, palynological, and carpological characters of P. miroense based on a range of microscopic analyses. Examination of the floral micromorphology of P. miroense revealed that the surface patterns of bractlets have potential utility in taxonomy studies of the genus Peucedanum. In addition, we established that this plant is characterized a papillose conical-tabular rugose pattern of petal epidermal cells and bears sunken nectary slits on the stylopodium. Also identified were simple spherical orbicules bearing psilate surface ornamentation, as well as small tricolporate prolate to perprolate pollen grains characterized by verrucate and gemmate ornamentation. Moreover, we verified pollen wall stratification and presence of pollenkitt. The mericarps had colliculate-striate surface patterns with trichomes and eight or nine vittae, and prismatic crystals were detected in different layers of the pericarp. We believe that the findings of this study will contribute to an evaluation of the taxonomic significance of this species within the genus Peucedanum, and also provide valuable information for the conservation of this rare endemic species. RESEARCH HIGHLIGHTS: We provided the first detailed descriptions of the floral micromorphological, palynological, and carpological characters of the rare endemic Korean species Peucedanum miroense K. Kim, H.-J. Suh & J.-H. Song, based on different microscopic analyses. Our findings provide valuable information regarding the epidermal cell patterns of bractlets, petals, stylopodium, gynoecium, and anthers. We believe our study makes a significant contribution to the literature, as our findings will contribute to enhancing the taxonomical classification of species within the genus Peucedanum, and will provide a basis for the conservation of this rare endemic species. This results will be of value to researchers studying the future biology and taxonomy of species within the family Apiaceae, particularly those using different microscopic techniques, including FE-SEM, digital slide scanning, and TEM.

Morphological Diversity and Taxonomic Importance of Fruits and Seeds of the Genus Spigelia (Loganiaceae) in Mexico

Abstract— The genus Spigelia (Loganiaceae) is a Neotropical genus that includes more than 80 species distributed from the United States to Argentina. Systematic studies of the group have rarely considered the carpological characters, so their taxonomic importance has not been analyzed in detail. The aim of this study is to analyze the morphological characteristics of these organs to provide more information for a better species separation and for a more robust infra-generic classification. We provide a detailed description of Spigelia fruits and seeds of the Mexican species since the country represents one of its main centers of diversification. Fruits and seed samples from 54 individuals across 14 species were measured and photographed to describe each one, maintaining as much detail as possible. With the morphological matrix a DFA, PCA, and a dendrogram using the NJ method was performed. The fruits and seeds of the Mexican Spigelia species proved to be taxonomically useful, allowing the differentiation of four main clusters of species separated by the size of the analyzed structures in the three analyses (DFA, PCA, NJ). The results show the variation in size and shape of the structures analyzed, which shows the taxonomic importance of fruits and seeds of the genus Spigelia. Our work represents an advance in the detailed knowledge of structures, which are rarely considered in the descriptions, and which allows us to have a separation of species. A taxonomic treatment and photographs of the fruits and seeds are also included.

Phylogenetic position of Ligusticopsis (Apiaceae, Apioideae): evidence from molecular data and carpological characters.

Ligusticopsis (Apiaceae, Apioideae) is now considered to have an East-Asia and Sino–Himalaya distribution. The genus was not recognized as a natural and separate genus and was treated as a synonym of Ligusticum both in Flora Reipublicae Popularis Sinicae and Flora of China since first established, though Pimenov et al. have made many taxonomic revisions to Ligusticopsis, phylogenetic relationships between Ligusticopsis and Ligusticum have been in dispute. Thirty-four plastomes and 35 nrITS from Apioideae were analysed by RAxML and MrBayes to reconstruct the phylogenetic relationships, along with carpology of 10 species and comparative analyses of 17 plastomes to investigate the evidence supporting the independence of Ligusticopsis. As a result, nine species suggested to be Ligusticopsis formed a highly supported monophyletic branch (Subclade A) inside Selineae both in maximum likelihood and Bayesian inference; the results of the comparative analyses further supported the monophyly of Subclade A, mainly in the location of genes at the IRa/LSC boundary, the sequence diversity exhibited by various genes (e.g. trnH-GUG–psbA and ycf2) and same codon biases in terminator TAA (relative synonymous codon usage = 1.75). Species in Subclade A also had shared characters in mericarps, combined with other characters of the plant, ‘base clothed in fibrous remnant sheaths, pinnate bracts, pinnate bracteoles longer than rays of umbellule, mericarps strongly compressed dorsally, median and lateral ribs filiform or keeled, marginal ribs winged, and numerous vittae in commissure and each furrow’ should be the most important and diagnostic characters of Ligusticopsis. Our phylogenetic trees and other analyses supported the previous taxonomic treatments of Pimenov et al. that Ligusticopsis should be a natural and separate genus rather than a synonym of Ligusticum.

Polyphyletic Trachyspermum (Umbelliferae) revisited: a contribution of molecular and carpological data

Trachyspermum is an economically important genus widely distributed in the Old World. A modern monographic survey is absent, so the number of species and their affinities are treated in the regional Floras extremely contradictorily. In this study, all species names available in the genus Trachyspermum were revised, and 19 validly published names are kept in the genus. To investigate the species relationships in Trachyspermum, which was shown in previous studies to be polyphyletic, nrDNA ITS sequence data were determined for additional taxa, accompanied with study of fruit morphology, micromorphology and anatomy. Our molecular analysis indicates that Trachyspermum is an aggregate of at least seven distant lineages, and that T. ammi (generitype) does not group with other Trachyspermum species. Many carpological characters do not agree with these groups, being rather homoplastic, but we identify that types of hairs in fruit indumentum and microsculpture of hair cuticle are consistent with molecular clades within Trachyspermum. Our investigation provides one more step towards to the taxonomic revision of the genus, and the possibility of splitting of the current Trachyspermum into monophyletic genera is discussed.

Comparative carpological study of Poikilospermum (Urticaceae) in relation to taxonomy

SummaryFruit morphology and the anatomical structure of the pericarp, fruiting perianth, and seed coat were studied in 15 species of Poikilospermum, a genus whose position within the Urticaceae has long been controversial. Possible evolutionary trends of their transformation are suggested for both subgenera; plesiomorphies were found in P. oblongifolium and P. scabrinervium. Structural peculiarities of the fruit connected with its ejection out of the tubular perianth are discussed. The archaism of the fruit in Poikilospermum is revealed, indicated, as in Boehmeria, by the presence of the rudiment of an aborted carpel in the form of a large two-lobed rib. Using carpological anatomical characters, the species studied are classified into informal groups, such characters being able to pull the species within the subgenera into rough groupings where gross morphology has been unable to do so. It is shown that heterobathmy may be strongly associated with the genus Poikilospermum. Each subgenus has its own set of primitive carpological characters: in subgenus Poikilospermum the absence of a fruiting perianth which encases the fruit, and also of capitate inflorescences with swollen receptacles; in subgenus Ligulistigma remnant rudiment of the second carpel and ovary loculus, as well as a primitive, less simplified seed coat. Though the position of Poikilospermum as indicated by molecular data is within Urera, our results suggest that Dendrocnide (the only genus of the Urticeae that has a pyrenarium fruit type) may be closest to Poikilospermum, although the pericarp structure and dissemination behaviour in Poikilospermum are more specialised than those exhibited by Dendrocnide. Seed coat structure is also shown to exhibit traits seen in Moraceae.

A new species of Mimosa section Mimosa (Leguminosae) from the Paraguayan Cerrado

Mimosa schininii (Leguminosae), a new species from the Amambay region, in the Cerrado ecoregion of northeastern Paraguay, is described and illustrated. This new species belongs to Mimosa L. section Mimosa series Mimosa subseries Brevipedes Barneby, and is similar to M. brevipes Benth. and M. sceptrum Barneby, but differs from them by shorter inflorescence, secondary leaflets venation pattern and fruit dehiscence. This new entity resembles, in some characters, other subseries of series Mimosa, such as Mimosa section Mimosa series Mimosa subseries Pedunculosae (Benth.) Barneby and Hirsutae (Benth.) Barneby, but can be distinguished adequately from them by vegetative, floral, and carpological characters. Illustration, distribution map, and a key to distinguish M. schininii from close relatives are provided.

Phylogeography of a west-Beringian endemic plant: An ancient seed of Stellaria jacutica Schischk. detected in permafrost deposits of the last interglacial

Beringia, the landmass connecting Eurasia and North America during Pleistocene sea level low stands, played a crucial part in the phylogeographic history of northern plants, animals and humans. Beringia was not only route for migration between both continents but, as the only unglaciated sector of the Arctic, it was also an important refugial area for northern biota during the Pleistocene. The finding of a fossil seed of Stellaria jacutica Schischk., endemic in Yakutia and the Russian Far Eastern Magadan Oblast, in permafrost deposits near Batagay demonstrates the existence of that species in the Yakutian Yana Highlands already during the last Pleistocene interglacial, about 125 ka ago. The biogeographical, ecological and seed morphological characters of S. jacutica allow for conclusions on its phylogenetic relationships. Stellaria jacutica is the only large seeded Stellaria species occurring in northeast Siberia today. According to carpological characters, S. jacutica is closely related to south Siberian large-seeded Stellaria species, which have a centre of diversity in the Altai-Sayan mountain range. This kinship is also suggested by corresponding ecological preferences and occurrence in mountain steppes and rock steppe communities mainly in the alpine belt of mountains. The spread of S. jacutica's ancestor north-eastwards assumedly happened during a Pleistocene cold stage, when the climate in Siberia was cooler, drier and more continental than during warm stages. Spread and following allopatric speciation hence occurred prior to the last interglacial, i.e. during or prior to the middle Pleistocene.

Cytogeography and morphological characterisation of a taxonomic, polyploid complex of Mimosa (Leguminosae) from subtropical South America

Mimosa subseries Dolentes Barneby and Brevipedes Barneby are ecologically and morphologically high-diversified infrageneric taxa of this genus in southern South America. We performed a cytogenetical and morphological analysis of both subseries. Chromosome numbers from accessions throughout the area of distribution were studied. The chromosome numbers 2n=8x=104 for M. dolens subsp. callosa (Benth.) Barneby, M. dolens subsp. acerba varieties acerba (Benth.) Barneby, latifolia (Benth.) Barneby and rudis (Benth.) Barneby, M. dolens subsp. rigida var. rigescens (Benth.) Barneby, var. anisitsii (Lindm.) Barneby and var. foliolosa (Benth.) Barneby; and 2n=4x=52 for M. sceptrum Barneby, M. aff. custodis Barneby and M. dolens var. pangloea Barneby are presented for the first time. Their karyotypes were symmetric, with small chromosomes. There were several areas with taxa growing in sympatry, occasionally with intermediate forms. No diploids were found, which suggests the presence of either a declining polyploid complex or ancient polyploidy in the clade of southernmost representatives of M. series Mimosa, the most derived of the genus. Some vegetative, inflorescence and carpological characters seem to be associated with chromosome duplication. The distribution pattern of the cytotypes suggests events of chromosome duplication in centres of diversity and expansion of octoploids to the southernmost areas of distribution. Our findings support the importance of polyploidy in the morphological diversity, distribution and speciation of this complex.

Morphological variability of Tragopogon pterodes Pančić ex Petrović achenes: taxonomic evaluation of heterocarpy

This study presents the carpological heterogeneity of Tragopogon pterodes, an endemic plant species from Balkan Peninsula. The goal of this paper was to determine the variability of morphological characters of achenes and assess the degree of their morphological differentiation relative to the position they occupy in the receptacle. Analysis included four different populations of T. pterodes and examination of 18 (8 qualitative and 10 quantitative) characters of achenes in total. Data analysis employed basic statistic, univariate (ANOVA, Tukey HSD) and multivariate (CDA, CA) analyses to check the level of heterocarpy in this species. The results indicate possible morphological differentiation of three groups of achenes (outer, median and inner achenes), depending of their position on the receptacle. The members of all three types were described, illustrated and compared. The variability of investigated carpological characters in this species is providing important information prior to their further taxonomic treatment.

Chorological and taxonomic notes on African plants, 2

The taxonomy of complicated native African taxa, Biscutella maritima (Brassicaceae) and annual species of Sesuvium (Aizoaceae) is discussed. The distribution of B. maritima is widened to the most of the coastal areas of NE Algeria and N Tunisia. Morphological differences with regard to other North African members of Biscutella ser. Biscutella are reported to facilitate the identification of B. maritima. We propose to accept four annual Sesuvium species (instead of the one previously accepted species, S. sesuvioides sensu amplissimo) with different distributions in Africa: S. digynum, S. hydaspicum, S. sesuvioides s.str. and S. nyasicum. A delimitation key based on morphological and carpological characters is provided. Lectotypes of S. digynum, S. digynum var. angustifolium, S. hydaspicum, S. nyasicum, Diplochonium sesuvioides (Sesuvium sesuvioides) and Trianthema polysperma (synonym of Sesuvium hydaspicum) are selected. Other discussed taxa belong to the alien elements of the flora. Atriplex semibaccata (Amaranthaceae-Chenopodiaceae) is reported as a new species for Cape Verde. Gaillardia × grandiflora (Asteraceae) is discovered as a new species for Africa found in Morocco. Prosopis velutina (Fabaceae) is collected for the first time in Northern Africa (Morocco). Mollugo verticillata (Molluginaceae) is recorded as a new species for DR Congo. Its morphology, including seed ornamentation, is discussed in reference to other similar-looking Molluginaceae. Incidence of American species Heterotheca subaxillaris (Asteraceae) in North Africa is discussed. Vernonanthura polyanthes (Asteraceae) is recorded as a naturalized invasive species in eastern Zimbabwe. It seems to be the first documented discovery of this ergasiophyte in Africa.

Taxonomic revision and distribution of herbaceous Paramollugo (Molluginaceae) in the Eastern Hemisphere.

The genus Paramollugo with the type species Paramollugo nudicaulis (≡Mollugo nudicaulis) has recently been described after molecular investigations. Here we report two new endemic Malagasy species: Paramollugo simulans and Paramollugo elliotii, and transfer a forgotten New Caledonian endemic Mollugo digyna to Paramollugo (Paramollugo digyna). Consequently, the number of Paramollugo species in the Eastern Hemisphere is increased from three to six. Almost all genus representatives (except Paramollugo nudicaulis, which has a wide distribution in Southern Asia, Arabia and tropical Africa) are endemic to Madagascar, Somalia, or New Caledonia. Since the type of seed coat ornamentation is crucial for species delimitation, a diagnostic key with new taxonomically significant carpological characters and other new traits is provided for all the herbaceous Paramollugo. The distribution patterns of Paramollugo nudicaulis s.str., Paramollugo simulans and Paramollugo elliotii are presented.

Taxonomy and chorology of Corbichonia (Lophiocarpaceae s.l.) with further description of a new species from Southern Africa

A re-examination of the herbarium material has allowed to describe a new species, Corbichonia exellii, occurring in Southern Africa (Angola, Namibia and South Africa). C. exellii represents the third species in the genus. Morphological differences between all three Corbichonia species (C. decumbens, C. rubriviolacea, and C. exellii) are provided. The description of the genus is defined using the newly discovered characters (reproductive features). The taxonomy of all Corbichonia species (synonyms included) is provided, as well as maps for all species. The lectotypes of Orygia decumbens (basionym of Corbichonia decumbens) and O. mucronata (synonym of C. decumbens) are designated on specimens preserved, respectively, at BM and K. The genus Corbichonia, recently placed in Lophiocarpaceae, is markedly different from the core genus Lophiocarpus on the basis of embryological, morphological and carpological characters, and deserves further investigation concerning its taxonomic status.

One-seeded fruits in the core Caryophyllales: their origin and structural diversity.

The core Caryophyllales consist of approximately 30 families (12 000 species) distributed worldwide. Many members evolved one-seeded or conjoined fruits, but their origin and structural diversity have not been investigated. A comparative anatomical investigation of the one-seeded fruits within the core Caryophyllales was conducted. The origin of the one-seeded fruits and the evolutionary reconstructions of some carpological characters were traced using a tree based on rbcl and matK data in order to understand the ancestral characters and their changes. The one-seeded fruit type is inferred to be an ancestral character state in core Caryophyllales, with a subsequent increase in the seed number seen in all major clades. Most representatives of the ‘Earlier Diverging’ clade are distinguished in various carpological traits. The organization of the pericarp is diverse in many groups, although fruits with a dry, many-layered pericarp, consisting of sclerenchyma as outer layers and a thin-walled parenchyma below, with seeds occupying a vertical embryo position, are likely ancestral character states in the core Caryophyllales clade. Several carpological peculiarities in fruit and seed structure were discovered in obligate one-seeded Achatocarpaceae, Chenopodiaceae, Nyctaginaceae, Seguieriaceae and Sarcobataceae. The horizontal embryo evolved in only certain groups of Chenopodiaceae. The bar-thickening of endotegmen cells appears to be an additional character typical of core Caryophyllales. The syncarpy-to-lysicarpy paradigm in Caryophyllaceae needs to be reinterpreted.

Distinguishing Nonpareil Marketing Group Almond Cultivars through Multivariate Analyses

More than 80% of the world's almonds are grown in California with several dozen almond cultivars available commercially. To facilitate promotion and sale, almond cultivars are categorized into marketing groups based on kernel shape and appearance. Several marketing groups are recognized, with the Nonpareil Marketing Group (NMG) demanding the highest prices. Placement of cultivars into the NMG is historical and no objective standards exist for deciding whether newly developed cultivars belong in the NMG. Principal component analyses (PCA) were used to identify nut and kernel characteristics best separating the 4 NMG cultivars (Nonpareil, Jeffries, Kapareil, and Milow) from a representative of the California Marketing Group (cultivar Carmel) and the Mission Marketing Group (cultivar Padre). In addition, discriminant analyses were used to determine cultivar misclassification rates between and within the marketing groups. All 19 evaluated carpological characters differed significantly among the 6 cultivars and during 2 harvest seasons. A clear distinction of NMG cultivars from representatives of the California and Mission Marketing Groups was evident from a PCA involving the 6 cultivars. Further, NMG kernels were successfully discriminated from kernels representing the California and Mission Marketing Groups with overall kernel misclassification of only 2% using 16 of the 19 evaluated characters. Pellicle luminosity was the most discriminating character, regardless of the character set used in analyses. Results provide an objective classification of NMG almond kernels, clearly distinguishing them from kernels of cultivars representing the California and Mission Marketing Groups.