Some Morphogenetic Factors in Taxonomy of Dinoflagellates

The formative ability of ectoplasm in situ and in isolation from the living cell is shown to be an autodynamic morphogenetic factor creating sui generis membrane structure in species. Heteromorphic division and formation of specific aberrants, which differ from the parent cell by characters which have been used to separate genera, may be caused by molecular changes in ectoplasm as a primary factor. The subpellicular vacuom reticulum in Gyrodinium and other genera originates possibly from the upset metabolism of protoplasm. It is purely phenotypic. Oxygen deficiency induces in Prorocentrum endogenous cysts and dwarf-cell formation, a plasmolysis-like effect. The morphological norm in dinoflagellates is maintained also by the angle of nuclear fission and ensuing cytoplasm disposition. It is constant in sea and culture in some species, almost constant in sea but multivariable in culture in other species. The static form in some species is related to a single method of division (Exuviaella marina). Polymorphy of Exuviaella, Prorocentrum and other genera is induced by the presence of many types of divisions, which also favours an abundance of aberrants. “Osmomorphoses” originate in critical salinity-temperature conditions owing to flexibility of the pellicle in Gymnodinioideae; it is prevented by the membrane rigidity in the thecate forms. Flexibility of membrane in juvenile cells favours exogenous division. Rigid membranes of adult cells inhibit exogenous, favour endogenous fission. “Cyclomorphoses”, “phagomorphoses” (feeding habit) and different types of aberrants are useful to complete classification in modern and fossil dinoflagellates since morphogenesis rules work regardless of time and space.     

Host Specificity of the Argentine Root-Boring Weevil, Heilipodus ventralis (Coleoptera: Curculionidae), a Potential Biocontrol Agent for Snakeweeds (Gutierrezia: Asteraceae) in Western North American Rangelands—U.S. Quarantine Tests

Native snakeweeds, especially Gutierrezia sarothrae (Pursh) Britton and Rusby and Gutierrezia microcephala (DC.) A. Gray, are among the most widespread and damaging weeds of rangelands in the western United States and northern Mexico. The genus long ago spread to southern South America, where further speciation occurred. We have found several species of insects in Argentina that damage other species of snakeweeds there and are possible candidates for biological control in North America. The first of these, the root-boring weevil, Heilipodus ventralis (Hustache), was tested in Argentina and then sent to the USDA-ARS Insect Quarantine Facility at Temple, Texas, for host specificity testing on North American plants. We tested H. ventralis on 40 species of the family Asteraceae, in 19 tests of five types, using 686 adults and 365 larvae. Host specificity increased from adult feeding, to ovipositional selection, to larval development. At Temple, adults fed mostly on 6 species of the closely related genera Grindelia, Gutierrezia, and Gymnosperma, but with substantial feeding on four other genera of the two preferred subtribes Solidagininae and Machaerantherinae and on Baccharis in the tribe Baccharidinae, with lesser feeding on the subtribe Asterinae, all in the tribe Astereae, and on 1 species in the tribe Anthemideae. Females oviposited primarily on the same 6 species but very little on plants outside the 2 preferred subtribes. Larvae developed only on 9 of the 29 U.S. plant species tested, 6 within the two preferred subtribes and on Brickellia and Aster in other tribes. Only 5 species of three genera appear to be potential true hosts of H. ventralis in North America, on which all stages of the life cycle, adult feeding, oviposition, and larval development, can take place; these are Gymnosperma glutinosum (Spreng.) Less., Gutierrezia grandis Blake, Gut. microcephala, Gut. sarothrae, and Grindelia lanceolata Nutt. None of these genera contain species of economic or notable ecological value; the few rare species appear to be protected by habitat isolation from attack by H. ventralis. H. ventralis, therefore, appears sufficiently host specific for field release in North America. This is the first introduced biocontrol agent to be approved for release in a continental area to control a native weed.     

Neuronal organization of the accessory olfactory bulb of the lizard Podarcis hispanica: Golgi study

The neuronal organization of the accessory olfactory bulb (AOB), which receives sensory information from the vomeronasal organ, was described in a squamate reptile (Podarcis hispanica) by means of light microscopy. Using the Golgi-impregnation method, seven neuronal types could be distinguished: Periglomerular cells constitute a morphologically heterogeneous population of small neurons located between and around the glomeruli. The mitral cells are diffusely distributed in the AOB. Their cell bodies are usually located within the mitral cell layer, but some of them could be also observed in the plexiform layers. Mitral cells were classified into three subgroups on the basis of their sizes and dendritic tree morphologies. Thus, the “outer mitral cells” have the biggest cell bodies, and their distal secondary dendrites are mainly distributed rostrocaudally in the external plexiform layer. The “inner mitral cells” have large cell bodies, and their secondary dendrites are distributed dorsoventrally and are located deeper than those of the other two subgroups. The third type, the “small mitral cells,” is the smallest one among mitral cells in the AOB, and from their cell bodies, only two main dendritic trunks arise. The granule cells are composed of several categories based on their different cell body locations and dendritic tree morphologies. Thus, the “superficial granule cells” are located exclusively in the external plexiform layer and have small dendritic fields. The “middle granule cells” have fusiform cell bodies—situated in the internal plexiform layer—and present a wide dendritic projection area. Finally, the “deep granule cells” are distributed throughout the granule cell layer and include a great variety of dendritic tree morphologies. The distribution and morphological features of all neuronal types constituting the AOB of Podarcis were compared with those reported on other vertebrates. The results suggest that the lamination pattern and neuronal organization of the AOB in lizards are more similar to that of mammals than to that of the remaining vertebrates.     

Beiträge zur Systematischen Anatomie derAnthemideae: Die Spaltöffnungsapparate

The subdivision of theAnthemideae into two subtribes purely on the grounds of the presence or absence of receptacular paleae can no longer be maintained. Anatomical data may serve as a basis for a more adequate division of the tribe. This survey of the stomatal apparatus types within theAnthemideae is based on the investigation of 29 species from 15 genera and the evaluation of literature data: Anomocytic stomatal apparatus occur in all species examined, anisocytic at least in all genera investigated by us. We even found representatives of some rare types, such as polo-, helico- or hemiparacytic apparatus, in nearly all species. Diacytic types were not found inOtanthus, Artemisia, Tripleurospermum, Tanacetum corymbosum, and they also appear to be lacking inSantolina andEriocephalus. Paracytic stomatal apparatus and a new type which links paracytic with actinocytic and cyclocytic was discovered inOtanthus maritimus andArtemisia stellerana.

     

Relationships among Cichorium species and related genera as determined by analysis of mitochondrial RFLPs

Mitochondrial DNA polymorphism was employed to assess cytoplasmic diversity among cytoypes of the genus Cichorium and related genera of the tribe Lactuceae (Asteraceae). Hybridization patterns of total DNA using six restriction enzymes and five heterologous mtDNA probes were examined. From estimates of mtDNA diversity, Cichorium spinosum appeared as an ecotype of C. intybus rather than a separate species. Interspecific mtDNA polymorphism in the genus Cichorium was higher than that observed in Cicerbita Crepis, Lactuca and Tragopogon. Molecular data seemed to indicate that Catananche is very distant from the other genera examined. Intergeneric comparisons allowed the clustering of Cicerbita, Lactuca and Cichorium, genera which belong to different subtribes. However, further molecular investigations on a larger number of genera are needed to clarify the relationships among genera within and between subtribes of the tribe Lactuceae.     

Nuclear DNA Amounts in Palms (Arecaceae)

Nuclear DNA amounts are reported for 83 species and 53 genera of palms, covering all of the six subfamilies. 4C DNA contents range between 3.89 and 55.62 pg in diploids, showing an approximate 14.3-fold variation in genome size. Polyploids have DNA contents of up to 156.40 pg/4c which demonstrates a 40.2-fold variation. Diploids with high DNA contents occur in three subfamilies of palms (Coryphoideae, Calamoideae, Arecoideae), and seem to be further restricted to particular tribes or subtribes (Thrinacinae, Borasseae, Lepidocaryeae, Caryoteae, some subtribes of Areceae). Palms from the subfamilies Nypoideae and Phytelephantoideae have the lowest DNA amounts, followed by the Phoeniceae and the Corypheae: Livistoninae from the subfamily Coryphoideae. Although DNA amounts in some genera and subtribes are usually constant, e.g., in Phoenix, Phytelephas, the Livistoninae, Dypsidinae, diploid Butiinae), considerable variation occurs at the diploid level in some large and apparently actively evolving genera such as Chamaedorea, Pinanga, Cenoma and possibly Bactris. Formaldehyde fixation is recommended for palms, as conventional ethanol-acetic acid fixation has proved to be unsuitable for DNA estimation of Feulgen-stained nuclei by microdensitometry, since it can lead to errors up to 2.5-fold in extent. Chromosome counts are reported for 72 of the species studied, of which 42 are new.     

SOURCE PLANTS FOR CARBONIFEROUS MICROSPORES: LYCOSPORA FROM PERMINERALIZED LEPIDOSTROBUS

Permineralized Lepidostrobus specimens were examined from the Westphalian A of England, Westphalian B (Breathitt) of the Appalachian Basin, and the Westphalian D (Desmoinesian) of the Illinois Basin and Western Interior Coal Region to correlate species of the dispersed spore genus Lycospora with source cones and cone types (“species”) with parent plants. Based on 51 specimens of Lepidostrobus oldhamius sensu Balbach, three cone types, producing three species of Lycospora, are recognized. Lycospora pellucida occurs in smaller diameter cones with a prominently coronate stele; cones are associated with Lepidophloios harcourtii from the Westphalian A-B. Lycospora pusilla occurs in larger diameter cones with a small “pith” containing elongate, sclerified, tanniferous cells from Westphalian D coal balls. Secretory cells also are present in sporophyll trace sheaths and in subarchesporial parenchyma pads in sporangia. The suggested plant source is Lepidodendron hickii. Lycospora granulata occurs in large cones with a large “pith” composed of thin-walled parenchyma; no secretory cells were observed in the “pith,” sporophyll traces, or subarchesporial pads. These cones are associated with Lepidophloios hallii from the Westphalian D. Association of Lycospora species with source lycopods increases the possibilities of paleoecological interpretation using microspores from Carboniferous coals.     

Classification of the gall midge tribe Aphidoletini (Diptera,Cecidomyiidae: Aphidoletidi) with descriptions of a new genus and a new species from the Kurile Islands

A new classification is proposed for the gall-midge tribe Aphidoletini including 4 subtribes, 10 genera, and 18 species: Aphidoletina—3 genera with 7 species (Aphidoletes, Tribremia, Ligulodiplosis), Monobremiina subtrib. n.—2 genera with 5 species (Monobremia, Shikotanodiplosis gen. n.), Aculeatodiplosina subtrib. n.—2 genera with 2 species (Aculeatodiplosis, Setodiplosis), and Triommatomyiina—3 genera with 4 species (Triommatomyia, Tingidoletes, Chanchudiplosis). New subtribes and a new genus with a new species are described. Shikotanodiplosis sundukovi gen. et sp. n. differs from Monobremia in a wide and curved aedeagus, basally swollen sclerotized hypoproct, and small basal lobes of the gonocoxites. Keys to the subtribes, genera, and species of Aphidoletini are given. The diagnoses of Aculeatodiplosis fasciata Fedotova et Sidorenko, 2005, Setodiplosis unifaria Fedotova, 2006, and Triommatomyia Mamaev, 1961 are supplemented with biometrical characters. Tribremia aphidophaga Marikovskij, 1956 is resurrected from synonyms of Aphidoletes aphidimyza (Rondani, 1847).     

An Internet‐based platform for the estimation of outcrossing potential between cultivated and Chilean vascular plants

A national‐scale study of outcrossing potential within Chilean vascular flora was conducted using an upgraded algorithm, which adds parameters such as pollinator agents, climate, and geographic conditions. Datasets were organized and linked in a Web platform ( www.flujogenico.cl ), in which the development of a total outcrossing potential (TOP) predictor was formulated. The TOP predictor is the engine in the Web platform, which models the effect of a type of agricultural practice on others (coexistence calculation mode) and on the environment (biodiversity calculation mode). The scale for TOP results uses quintiles in order to define outcrossing potential between species as “very low,” “low,” “medium,” “high,” or “very high.” In a coexistence analysis considering 256 species (207 genera), the 10 highest TOP values were for genera Citrus, Prunus, Trifolium, Brassica, Allium, Eucalyptus, Cucurbita, Solanum, Lollium, and Lotus. The highest TOP for species in this analysis fell at “high” potential, 4.9% of the determined values. In biodiversity mode, seven out of 256 cultivated species (2.7%) were native, and 249 (97.3%) corresponded to introduced species. The highest TOP was obtained in the genera Senecio, Calceolaria, Viola, Solanum, Poa, Alstroemeria, Valeriana, Vicia, Atriplex, and Campanula, showing “high” potential in 4.9% of the values. On the other hand, 137 genetically modified species, including the commercial and pre‐commercial developments, were included and represented 100 genera. Among these, 22 genera had relatives (i.e., members of the same genus) in the native/introduced group. The genera with the highest number of native/introduced relatives ranged from one (Ipomea, Limonium, Carica, Potentilla, Lotus, Castanea, and Daucus) to 66 species (Solanum). The highest TOP was obtained when the same species were coincident in both groups, such as for Carica chilensis, Prosopis tamarugo, and Solanum tuberosum. Results are discussed from the perspective of assessing the possible impact of cultivated species on Chilean flora biodiversity. The TOP predictor ( http://epc.agroinformatica.cl/ ) is useful in the context of environmental risk assessment.     

ESSENTIAL LEAF OILS IN REPRESENTATIVES OF THE AURANTIOIDEAE (RUTACEAE)

The distribution of essential leaf oils in representatives of 12 genera in the Aurantioideae has been studied by gas chromatography. Of the individual oil components, 40 were identified by retention times on two columns, augmentation, and some by IR spectrophotometry. Twelve groupings of components were established and these compared between the tribes Clauseneae and Citreae , among the subtribes clauseninae , balsamocitbinae , and citrinae , and between the “primitive” and the “near citrus group.” The essential oils conform to the botanical groupings, except for Pleiospermium in the citrinae and Murraya in the clauseneae. Fortunello crassifolia (Citbeae ), a possible garden hybrid, seems to be more closely related to the Clauseneae , the remote citroids, than to the near Citrus group. Atalantia also might be an intertribal hybrid with remote or other citroids. A specimen of Aeglopsis chevalieri (balsamocitbinae ) from Florida appears to be of hybrid origin with the parental source close to Citrus.     

Wiederaufstellung der Gattung Sericopelma Ausserer, 1875 (Araneida,Theraphosidae, Theraphosinae)

After the examination of the types of Mygalarachne brevipes Ausserer, 1871, and Sericopelma rubronitens Ausserer, 1875, we declare M. brevipes, a juvenile female, as a species “incertae sedis” since the specimen doesn't present sufficient characteristics which allow a secure identification of the genera and species.     

POLLEN MORPHOLOGY AND SYSTEMATIC RELATIONSHIPS AMONG NEW WORLD SPECIES IN TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)

The morphology of mature pollen grains among 12 New World genera in tribe Antirrhineae has been examined with light and scanning electron microscopy. Pollen from 29 American species have mean polar diam that range from 17 to 26 μm, have a tectate structure with perforate, microreticulate, or reticulate surface pattern, and are subspheroidal to prolate, and trizonocolporate with fusiform or narrowly oblong colpi that are free or occasionally fused at the poles. Among the genera, pollen size is positively associated with style length but not haploid chromosome number. Three major morphological categories are recognized based on grain shape, exine structural pattern, and colpus shape. Two of these morphotypes each characterize individual genera (Mohavea, Linaria). The other morphological category includes ten New World genera and has been divided into three subcategories based on variation in lumina diam and intermittent occurrence of fused colpi. Only Antirrhinum and Pseudorontium have species or populations with pollen in more than one subcategory. Concordance of morphological, anatomical, chromosomal, and palynological data suggests that Linaria and Mohavea are best placed in separate subtribes from other native American genera. Compared to Old World taxa in tribe Antirrhineae, the most common pollen type (subspheroidal, microreticulate, apocolpate, fusiform colpi) may be the most primitive. Pollen grains with fused colpi (Antirrhinum p.p., Pseudorontium) and perforate (Mohavea) or reticulate (Holmgrenanthe) structural patterns probably represent derived types.     

The Cerebral Neurosecretory System,Secretory End-Foot System and Infracerebral Gland—a Probable Neuroendocrine Complex in Nephtys (Annelida; Polychaeta)

Abstract The brain of Nephtys contains four neurosecretory cell types with distinctive cytoplasmic inclusions, a cells are located uniquely in a single pair of ganglionic nuclei and b cells are represented by a single pair of cells, whereas c cells and d cells have a scattered distribution. Their axons form two types of secretory release structure. First, possible axon collaterals synapse upon slender “dentritic twigs” in the core of the brain. Secondly, two tracts descend to the brain floor to form a “neurosecretory neuropile” (or storage and release complex) in contact with the inner surface of the brain capsule. Other neurosecretory fibres penetrate through the capsule, branch extensively, and terminate in contact with its ventral surface in close association with the “infracerebral gland”. The gland is derived from the pericapsular epithelium and exhibits signs of specialization for glandular function. In contrast to certain other polychaetes, it does not contain secretory neuron perikarya. The secretory end-foot system is poorly developed. Its terminals are located adjacent to the neurosecretory neuropile, which they encircle. The cell bodies are probably represented by four e cells which, like the terminals, contain many mitochondria.     

A worldwide phylogenetic classification of the Poaceae (Gramineae) III: An update

We present an updated worldwide phylogenetic classification of Poaceae with 11 783 species in 12 subfamilies, 7 supertribes, 54 tribes, 5 super subtribes, 109 subtribes, and 789 accepted genera. The subfamilies (in descending order based on the number of species) are Pooideae with 4126 species in 219 genera, 15 tribes, and 34 subtribes; Panicoideae with 3325 species in 242 genera, 14 tribes, and 24 subtribes; Bambusoideae with 1698 species in 136 genera, 3 tribes, and 19 subtribes; Chloridoideae with 1603 species in 121 genera, 5 tribes, and 30 subtribes; Aristidoideae with 367 species in three generaand one tribe; Danthonioideae with 292 species in 19 generaand 1 tribe; Micrairoideae with 192 species in nine generaand three tribes; Oryzoideae with 117 species in 19 genera, 4 tribes, and 2 subtribes; Arundinoideae with 36 species in 14 genera and 3 tribes; Pharoideae with 12 species in three generaand one tribe; Puelioideae with 11 species in two generaand two tribes; and the Anomochlooideae with four species in two generaand two tribes. Two new tribes and 22 new or resurrected subtribes are recognized. Forty-five new (28) and resurrected (17) genera are accepted, and 24 previously accepted genera are placed in synonymy. We also provide an updated list of all accepted genera including common synonyms, genus authors, number of species in each accepted genus, and subfamily affiliation. We propose Locajonoa, a new name and rank with a new combination, L. coerulescens. The following seven new combinations are made in Lorenzochloa: L. bomanii, L. henrardiana, L. mucronata, L. obtusa, L. orurensis, L. rigidiseta, and L. venusta.     

Sympatric speciation and radiative evolution of socially parasitic ants - Heretic hypotheses and their factual background

According to current hypotheses the main types of social parasitism among ants, namely slavery, temporary parasitism, and inquilinism, arose from such features as predation on other ants, or territorial behavior, both presumed precursors of slavemaking, and polygyny, a presumed precursor of temporary parasitism and inquilinism. The latter is believed also to represent a final instar in several evolutionary pathways leading from slavery, temporary parasitism, and xenobiosis to this permanently parasitic, workerless condition. Speciation, the origin of parasitic species from their usually closely related host species, is suggested to occur due to temporary geographic isolation and subsequent transition of one of the newly formed daughter species to parasitism in the nests of the other. Evidence is presented suggesting that the main types of social parasitism originated independently of each other. 15 ant genera are parasitized exclusively by inquilines, Eve other genera exclusively by temporary parasites. Only four groups of non-parasitic ant species (Formica, Tet-ramorium, Leptothorax subgenera Leptothorax and Myrafant) have parasites of several types each. Within these roups, however, there is little evidence of evolutionary transitions from one type to another. The few exceptions, mainly workerless species of the genera Epimyrma and Chalepoxenus, represent parasites which clearly derive from slave-making congeners, but differ from ordinary inquilines in that they eliminate the host colony queens like their actively dulotic ancestors. The new hypothesis suggests that all forms of interspecific true social parasitism (excluding xenobiosis) orginated from a common “preparasitic” stage, a subpopulation of reproductives in polygynous colonies and species, with diverging sexual behavior (near-nest mating vs. swarming) and caste ratios (production of more sexuals vs. workers). Arguments for sympatric speciation are compiled. Various features of the ancestral, and then host species (colony sizes, population density and structure, transition from polygyny to monoyny, etc.), and of the “preparasite” (production of few, or no workers, etc.) may shape the developing parasite to become a slave-maker, inquiline, or temporary parasite. These features usually leave open only one, or in a few genera, several options. The different types of parasitism within one host species group thus may have developed in a radiative manner from the common, preparasitic stage, which explains that independent colony foundation is a common feature of all true social parasites among ants.     

TYPES OF LATICIFERS AND CRYSTALS IN JATROPHA AND THEIR TAXONOMIC IMPLICATIONS

Anatomical examination of 37 species of Jatropha (Euphorbiaceae) has revealed the occurrence of two distinct types of laticifers—articulated, and nonarticulated—in addition to idioblasts that are irregularly shaped individual cells. With the notable exception of J. augustii Pax & K. Hoffm. (section Peltatae (Pax) Dehgan & Webster), these idioblasts occur in the two most advanced sections of the subgenus Curcas (Adans.) Pax, namely Loureira (Cav.) Muell. Arg. ex Pax and Mozinna (Ortega) Pax. The presence of two laticifer types and “idioblastic laticifers” in the genus, in association with the morphological reduction series found with evolutionary advancement are, therefore, significant in the delimitation of sections and subsections. Further evidence is presented by the occurrence of “chambered crystalliferous cells” in subgenus Curcas, but not in subgenus Jatropha (=Adenorhopium Griseb.). Reexamination of laticifers in other genera of the Euphorbiaceae is suggested as a possible means of alleviating the long-standing taxonomic dilemma of this large and morphologically diverse family.     

Involvement of Delta and Nodal signals in the specification process of five types of secondary mesenchyme cells in embryo of the sea urchin, Hemicentrotus pulcherrimus

Secondary mesenchyme cells (SMCs) of the sea urchin embryo are composed of pigment cells, blastocoelar cells, spicule tip cells, coelomic pouch cells and muscle cells. To learn how and when these five types of SMCs are specified in the veg₂ descendants, Notch or Nodal signaling was blocked with γ‐secretase inhibitor or Nodal receptor inhibitor, respectively. All types of SMCs were decreased with DAPT, while sensitivity to this inhibitor varied among them. Pulse‐treatment revealed that five types of SMCs are divided into “early” (pigment cells and blastocoelar cells) and “late” (spicule tip cells, coelomic pouch cells and muscle cells) groups; the “early” group was sensitive to DAPT up to the hatching, and the “late” group was sensitive until the mesenchyme blastula stage. Judging from timing of the shift of Delta‐expressing regions, it was suggested that the “early” group and “late” groups are derived from the lower and the middle tier of veg₂ descendants, respectively. Interestingly, numbers of SMCs were also altered with SB431542; blastocoelar cells, coelomic pouch cells and circum‐esophageal muscles decreased, whereas pigment cells and spicule tip cells increased in number. Pulse‐treatment showed that the “early” group was sensitive up to the mesenchyme blastula stage, while the “late” group up to the onset of gastrulation. Thus, it became clear that precursor cells of the “early” and “late” groups, which are located in different regions in the vegetal plate, receive Delta and Nodal signals at different timings, resulting in the diversification of SMCs. Based on the obtained results, the specification processes of five types of SMCs are diagrammatically presented.     

A systematic histological study of palm fruits. VI. Subtribe Linospadicinae (Arecaceae)

The pericarp structure of representative species of the four genera of the subtribe Linospadicinae is described and compared. Tissues found in the pericarp of this group are similar to those found in other subtribes of the Areceae, with no characters unique to the subtribe and nothing to suggest its closest affinities. The four genera, as well as each species examined, do show distinctive pericarp features, however.Laccospadix andLinospadix are similar, both with a single series of fibrous, bundles and an outer series of prominent raphide-bearing cells dominating the pericarp. In bothHowea andCalyptrocalyx, a complex exocarp forms from a series of fibrous bundles and brachysclereids, but each genus has other distinctive characters.Howea has vascular bundles in the exocarp zone, an outer series of raphide-bearing cells, and a conspicuously thickened locular epidermis. Based on a limited sample of four species, raphide-bearing cells appear to be always interior to the exocarp inCalyptrocalyx, and the locular epidermis is thin.