THE ANATOMY OF TUMORS INDUCED ON CITRUS BY CITRUS VEIN-ENATION VIRUS

Citrus vein-enation virus-induced tumors on leaves, thorns, stems, and roots of Citrus auranti-folia (Christm.) Swingle and C. jambhiri Lush. were investigated. Leaf vein tumors are initiated as cytological abnormalities of phloem fiber primordial cells adjacent to protophloem sieve tubes. The tumors then enlarge through hyperplasia of the affected fiber primordia. The mesophyll and epidermal tissues on the abaxial side of affected veins divide less prolifically and contribute to the tumor mass. Leaf vein tumors are determinate, like the protophloem where they originate, and cease enlarging as the leaf matures. Most thorn tumors are initiated in fiber primordia and develop similarly to vein tumors. Stem tumors and some thorn tumors develop from affected cells of the procambial tissue that lies between the metaxylem and metaphloem of vascular bundles. The cambium which differentiates in these areas is composed of affected cells and produces large amounts of abnormal xylem tissue. The resulting woody tumors are indeterminate like the cambium and secondary xylem.     

DIFFERENTIATION OF LATERAL SHOOTS AS THORNS IN ULEX EUROPAEUS

Ulex europaeus is a much-branched shrub with small, narrow, spine-tipped leaves and axillary thorn shoots. The origin and development of axillary shoots was studied as a basis for understanding the changes that occur in the axillary shoot apex as it differentiates into a thorn. Axillary bud primordia are derived from detached portions of the apical meristem of the primary shoot. Bud primordia in the axils of juvenile leaves on seedlings develop as leafy shoots while those in the axils of adult leaves become thorns. A variable degree of vegetative development prior to thorn differentiation is exhibited among these secondary thorn shoots even on the same axis. Commonly the meristems of secondary axillary shoots initiate 3–9 bracteal leaves with tertiary axillary buds before differentiating as thorns. In other cases the meristems develop a greater number of leaves and tertiary buds as thorn differentiation is delayed. The initial stages in the differentiation of secondary shoot meristems as thorns are detected between plastochrons 10–20, depending on vigor of the parent shoot. A study of successive lateral buds on a shoot shows an abrupt conversion from vegetative development to thorn differentiation. The conversion involves the termination of meristematic activity of the apex and cessation of leaf initiation. Within the apex a vertical elongation of cells of the rib meristem initials and their immediate derivatives commences the attenuation of the apex which results in the pointed thorn. All cells of the apex elongate parallel to the axis and proceed to sclerify basipetally. Back of the apex some cortical cells in which cell division has persisted longer differentiate as chlorenchyma. Although no new leaves are initiated during the extension of the apex, provascular strands are present in the thorn tip. Fibrovascular bundles and bundles of cortical fibers not associated with vascular tissue differentiate in the thorn tip and are correlated in position with successive incipient leaves in the expected phyllotactic sequence, the more developed bundles being related to the first incipient leaves. Some secondary shoots displayed variable atypical patterns of meristem differentiation such as abrupt conversion of the apex resulting in sclerification with limited cell elongation and small, inhibited leaves. These observations raise questions concerning the nature of thorn induction and the commitment of meristems to thorns.     

A REVISION OF ARIOCARPUS (CACTACEAE). I. THE STATUS OF THE PROPOSED GENUS ROSEOCACTUS

Anderson, Edward F. (Claremont Graduate School, Claremont, California.) A revision of Ariocarpus (Cactaceae). I. The status of the proposed genus Roseocactus. Amer. Jour. Bot. 47(7) : 582–589. Illus. 1960.—The proposed genus Roseocactus Berger is found to be a subgenus of Ariocarpus. The subgenera Ariocarpus and Roseocactus are basically similar in habitat, seedlings, presence of mucilage systems, tubercle structure, seed structure, fruiting habit, flower origin and structure, alkaloidal properties, trichomes, pollen grains, and chromosome number. They differ in flower color, presence or absence of an areolar groove, tubercle divergence, and trichome texture. A hypothesis is proposed to explain variations in the areoles. The differences in structure of mature tubercles are thought to be due to variability in the location of growth and elongation with respect to the floral and spinous areolar areas.     

EVALUATION OF RELATIONSHIPS WITHIN ERIODICTYON (HYDROPHYLLACEAE) USING TRICHOME CHARACTERISTICS

All 13 taxa of Eriodictyon Bentham (Hydrophyllaceae) were examined to determine the range of variation in trichome characters within the genus. Four simple trichome types were found: short and straight, intermediate length and straight, long and straight, long and wavy. Glandular capitate trichomes were also found in some species. Sessile glands were also observed but not included in the study. Most taxa displayed unique combinations of trichome types on stems, leaves, inflorescence axes and flower parts that allowed those taxa to be identified using trichome types alone. Trichome data support most previous taxonomic treatments, but suggest that 1) the two varieties of E. traskiae are extremely similar, 2) the rare E. altissimum is most similar to the widespread E. californicum, 3) the rare E. capitatum is allied with E. crassifolium var. nigrescens or perhaps E. angustifolium, 4) E. crassifolium var. denudatum and E. crassifolium var. nigrescens are essentially indistinguishable using trichome characters and, coupled with variation in other characters, are best combined under the name E. crassifolium var. nigrescens. Trichome characters provide a wealth of taxonomically useful information and may prove useful in the study of related genera of Hydrophyllaceae.     

Thorns as induced defenses: experimental evidence

Summary We report evidence from controlled experiments that long straight thorns deter herbivory by browsers. Cut branches of three woody species that had their thorns removed suffered significantly greater herbivory by a tethered goat than did paired intact branches. Branches on living Acacia seyal plants that had their thorns removed suffered significantly greater herbivory by a wild population of free-ranging giraffes than did intact branches on the same plants. These differences in herbivory resulted in long term losses of branch length in clipped as opposed to control branches. In addition, branches within reach of giraffes produced longer thorns and a greater density of thorns than did higher branches. These results imply that increased thorn length is an induced defense.No significance should be attached to order of authorship     

Increased thorn length in Acacia depranolobium —an induced response to browsing

Summary I report here longer thorns induced by large mammal herbivory on the tree Acacia depranolobium. I compared trees that had been browsed by domestic goats to trees protected from goat browsing. Thorns on browsed branches within the reach of goats (<125 cm above the ground) were significantly longer than thorns from higher branches on the same browsed trees, and significantly longer than branches at similar heights on unbrowsed trees. It appears that increased thorn length was an induced response to large mammal herbivory in Acacia depranolobium, both among and within individual trees.     

Survey of the variation in alar and malar thorns in skates: Phylogenetic implications (Chondrichthyes: Rajoidei)

A total of 62 species, representing 24 of the 28 supraspecific taxa of skates, was surveyed for the presence, distribution, and structure of alar and malar thorns. The survey revealed that alar thorns are almost universally present in mature male skates, and that malar thorns are of two types that may have separate origins. The location and orientation of alar thorns differed in two major clades of skates (groups I and II). They were either exposed on the disc with the crown oriented obliquely to the long axis of the base or imbedded in slits in the integument and oriented parallel to the long axis of the base. Group I possessed the former type of thorns and, with several exceptions, group II possessed the latter type. Group I either lacked malar thorns or possessed malar thorns that were most likely derived from generalized placoid scales. Group II possessed, with several exceptions, malar thorns derived from enlarged placoid scales. Within group I the supraspecific taxa Atlantoraja and Rioraja possessed unique alar thorns that were reclined laterally and associated with lateral grooves in the integument. Within group II the supraspecific taxa Anacanthobatis, Cruriraja, Dipturus, Okamejei, Raja, the North Pacific Assemblage, and the Amphi-American Assemblage had a large barb on the distal section of the crown of the alar thorn that suggests that these taxa form a clade. The crown of the alar thorns of the latter five supraspecific taxa was undulatory, thus supporting their monophyletic status. Another assemblage within group II had ridges that ran most of the length of the crown of the alar thorns corroborating their monophyletic status. The survey also suggested that variation of integumentary grooves associated with the alar thorns in Bathyraja, and presence or absence of malar thorns in Psammobatis, may be phylogenetically informative. © 1996 Wiley-Liss, Inc.     

Hierarchical Perception of Stimuli During Case Construction in the Bagworm Moth Eumeta crameri (Lepidoptera: Psychidae)

We studied case construction behavior of bagworm moth, Eumeta crameri (Lepidoptera: Psychidac) in the host plant, Acacia nilotica, at two sites. At the time of the study, Site A had 1-year-old A. nilotica saplings only, whereas Site B had full grown trees. The larva of bagworm moth used either thorns or cut-twigs or both as the materials for building its case. It renovated its case three times, and during each instance it increased the volume of its case by replacing older thorns or cut-twigs by newer and longer ones. However, it exhibited a spectacular predilection for thorns, irrespective of the sites, during the first-instar stage of its development. Thereafter, at Site A it used exclusively cut-twigs, whereas at Site B it preferably used thorns provided they were available in the range of the required length and within the threshold distance. It appears that the bagworm has an ability to process thorn length and distance signals hierarchically to fulfill its priority, that is, case building. Thus it optimizes time and energy expenditures during the period of its larval growth leading to pupation by toggling its preference between thorns and cut-twigs.     

Ecological barriers to early colony establishment in three coexisting acacia-ant species in Kenya

In black cotton uplands in East Africa, four symbiotic acacia-ant species compete for possession of a single swollen thorn tree species, Acacia drepanolobium, and yet coexist at fine spatial scales. Three of the four ant species produce independent foundress queens that establish colonies claustrally within swollen thorns, most often on small saplings. We conducted surveys of such saplings at two sites in 2001 and 2004, and examined foundresses and incipient colonies within their swollen thorns to determine what factors influence their success. Competition among foundresses for nest initiation sites was intense, with an average of over one founding attempt per swollen thorn in all samples, and with living and dead queens significantly hyper-dispersed among available thorns. Combat with other foundresses was the most common cause of death among claustral queens, especially for Tetraponera penzigi. In interspecific battles for nest initiation sites, T. penzigi was dominant over Crematogaster nigriceps and C. mimosae, and C. nigriceps won over 80% of its contests with C. mimosae foundresses. For singleton foundresses, brood parasitism by the braconid wasp Trigastrotheca laikipiensis typically results in the death of the entire ant brood. Host queens defend parasite larvae, pupae and eclosed adults, apparently unable to distinguish the wasps from legitimate offspring. Rates of brood parasitism were as high as 15–20% for incipient colonies of both Crematogaster species, but were extremely low for T. nigriceps in all samples. Although T. penzigi and C. nigriceps foundresses are always solitary, approximately 18% of claustral C. mimosae colonies contain cooperating pleometrotic queens. For unparasitized, claustral C. mimosae colonies, brood production per queen did not differ between solitary and cooperating foundresses. However, the per-capita risks associated with parastitism were reduced for pleometrotic queens. Received 8 March 2005; revised 23 May 2005; accepted 3 June 2005.     

Weapon (thorn) automimicry and mimicry of aposematic colorful thorns in plants

In order to further characterize the function of coloration in plants as defense against herbivory, two types of thorn mimicry are described: (1) A unique type of weapon (thorn) automimicry (within the same individual) that was previously known only in animals, and (2) mimicry of aposematic colorful thorns, by colorful elongated and pointed plant organs (buds, leaves and fruit) that, despite their appearance, are not sharp. Some thorny plants including dozens of species of Agave, one species of Aloe and a palm species have thorn-like imprints or colorations on their leaves, constituting thorn automimicry by giving the impression of more extensive thorns. The mimicry of aposematic colorful thorns is a typical case of Batesian mimicry, but the thorn automimicry is a special intra-organismic Batesian mimicry. I propose that both types of mimicry serve as anti-herbivore mechanisms.     

Adaptive radiation of photosynthetic physiology in the Hawaiian lobeliads: dynamic photosynthetic responses

Hawaiian lobeliads have radiated into habitats from open alpine bogs to densely shaded rainforest interiors, and show corresponding adaptations in steady-state photosynthetic light responses and associated leaf traits. Shaded environments are not uniformly dark, however, but punctuated by sunflecks that carry most of the photosynthetically active light that strikes plants. We asked whether lobeliads have diversified in their dynamic photosynthetic light responses and how dynamic responses influence daily leaf carbon gain. We quantified gas exchange and dynamic light regimes under field conditions for ten species representing each major Hawaiian sublineage. Species in shadier habitats experienced shorter and less numerous sunflecks: average sunfleck length varied from 1.4 ± 1.7 min for Cyanea floribunda in shaded forest understories to 31.2 ± 2.1 min for Trematolobelia kauaiensis on open ridges. As expected, the rate of photosynthetic induction increased significantly toward shadier sites, with assimilation after 60 s rising from ca. 30% of fully induced rates in species from open environments to 60% in those from densely shaded habitats. Uninduced light use efficiency—actual photosynthesis versus that expected under steady-state conditions—increased from 10 to 70% across the same gradient. In silico transplants—modeling daily carbon gain using one species’ photosynthetic light response in its own and other species’ dynamic light regimes—demonstrated the potential adaptive nature of species differences: understory Cyanea pilosa in its light regimes outperformed gap-dwelling Clermontia parviflora, while Clermontia in its light regimes outperformed Cyanea. The apparent crossover in daily photosynthesis occurred at about the same photon flux density where dominance shifts from Cyanea to Clermontia in the field. Our results further support our hypothesis that the lobeliads have diversified physiologically across light environments in Hawaiian ecosystems and that those shifts appear to maximize the carbon gain of each species in its own environment. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

GLANDULAR STRUCTURES OF HOLOCARPHA AND THEIR ONTOGENY

Carlquist , Sherwin . (Rancho Santa Ana Botanic Garden, Claremont, California.) Glandular structures of Holocarpha and their ontogeny. Amer. Jour. Bot. 46(4): 300–308. Illus. 1959.—Two types of advanced glandular structures occur in the 4 species of the genus Holocarpha. Sessile disk-shaped glands occur at the tips of upper leaves and of involucral and receptacular bracts. Unlike all other glandular structures of Madinae which have been investigated, these originate from several protodermal initials rather than a single one. These glands, however, represent modifications of a glandular trichome. The other type of glandular structure, termed hollow-stalked trichome here, occurs on the outer surface of involucral and receptacular bracts. These trichomes originate from a single cell but differ from others in the formation of a hollow stalk, the wall of which is one cell in thickness. Mesophyll of the bract, often with an included vascular bundle, is present as an intrusion into the base of the hollow stalk. Corresponding to the advanced nature of the glandular structures, the leaves show specializations in the “inrolling” of margins. Upper leaves have a cylindrical organization of vascular tissue, whereas basal leaves are “normal” and leaves of the main stem are intermediate. The species of Holocarpha differ in certain details of leaf anatomy and structure of hollow-stalked trichomes. The systematic distribution of these is given. The essential unity of the various glandular structures of Madinae is discussed both in terms of mature structure and ontogeny, and the steps in the evolution of these are suggested.     

The Potential Anti-Herbivory Role of Microorganisms on Plant Thorns

Thorns, spines and prickles are some of the anti-herbivore defenses that plants have evolved. They were recently found to be commonly aposematic (warning coloration). However, the physical anti-herbivore defense executed by these sharp structures seems to be only the tip of the iceberg. We show that thorns of various plant species commonly harbor an array of aerobic and anaerobic pathogenic bacteria including Clostridium perfringens the causative agent of the life-threatening gas gangrene, Bacillus anthracis, and Pantoea agglomerans. Septic inflammation caused by plant thorn injury can result not only from bacteria. Medical literature indicates that thorns, spines or prickles also introduce pathogenic fungi into animals or humans. Dermatophytes that cause subcutaneous mycoses are unable to penetrate the skin and must be introduced into the subcutaneous tissue by a puncture wound. The common microorganism-thorn combinations seem to have been an important contributor to the fact that so many plant thorns are aposematically colored, as a case of convergent evolution of aposematism in these organisms.Key Words: aposematism, herbivory, pathogen, spine, thorn, bacillus anthracis, clostridium perfringens, sporotrichosis, Mycetoma, subcutaneous mycotic disease     

SEQUENTIAL RADIATIONS AND PATTERNS OF SPECIATION IN THE HAWAIIAN CRICKET GENUS LAUPALA INFERRED FROM DNA SEQUENCES

The tremendous diversity of endemic Hawaiian crickets is thought to have originated primarily through intraisland radiations, in contrast to an interisland mode of diversification in the native Hawaiian Drosophila. The Hawaiian cricket genus Laupala (family Gryllidae) is one of several native genera of flightless crickets found in rain-forest habitat across the Hawaiian archipelago. I examined the phylogenetic relationships among mitochondrial DNA (mtDNA) sequences sampled from 17 species of Laupala, including the 12S ribosomal RNA (rRNA), transfer RNA (RNA)^val and 16S rRNA regions. The distribution of mtDNA variants suggests that species within Laupala are endemic to single islands. The phylogenetic estimate produced from both maximum likelihood and maximum parsimony supports the hypothesis that speciation in Laupala occurred mainly within islands. The inferred biogeographical history suggests that diversification in Laupala began on Kauai, the oldest rain-forested Hawaiian island. Subsequently, colonization to younger islands in the archipelago resulted in a radiation of considerable phylogenetic diversity. Phylogenetic patterns in mtDNA are not congruent with prior systematic or taxonomic hypotheses. Hypotheses that may explain the conflict between the phylogenetic patterns of mtDNA variation and the species taxonomy are discussed.     

POLLEN MORPHOLOGY,TRICHOME TYPES,AND RELATIONSHIPS OF THE GRONOVIOIDEAE (LOASACEAE)

The pollen of all four genera of Gronovioideae—Cevallia, Fuertesia, Gronovia, and Petalonyx—was examined in light microscopy, and scanning and transmission electron microscopy. The pollen of Cevallia, of Fuertesia, and of Gronovia can be easily distinguished from each other and from all remaining Loasaceae. Only Petalonyx, with a striate tectum, shows a clear relationship to the Mentzelioideae and Loasoideae, the vast majority of which have striate-reticulate or striate tecta. The trichome data are mostly congruent with the pollen data: Cevallia, Fuertesia, and Gronovia each have a distinctive trichome not known to occur elsewhere in the family, while Petalonyx has only the common types. A cladistic analysis of Gronovioideae utilizing Mentzelia as the outgroup proposes that Cevallia, Gronovia, and Fuertesia are a sister group to Petalonyx within the subfamily. The relationships of the four genera to each other and of Gronovioideae to the Loasaceae are discussed.     

Ants,Stem Borers,and Pubescence in Endospermum in Papua New Guinea1

The stems of the ant-plant, Endospermum labios Schodde, serve as colonization sites for the ant, Camponotus quadriceps F. Smith. They are also subject to damage by insect borers. We sampled young E. labios trees in distutbed forest to compare evidence of stem boring insect and stem miner damage in plants with and without colonies of C. quadriceps. Dissections of a subsample of plants showed that dipteran stem borers and stem miner damage were significantly more common in plants lacking C, quadriceps colonies than in plants with established colonies. Evidence from these dissections and from field counts of meristem damage caused by emerging borers suggested that coleopteran stem borers were also more abundant when ants were not present. In addition to the incidence of stem boring insects and ant colonies, we examined relative levels of leaf pubescence by measuring trichome density and leaf size for E. labios trees. We found that trichome density was significantly greater in trees with evidence of prior shoot damage (presumably from stem borer emergence at the meristem) but was not significantly related to the presence or absence of an ant colony. This prompted us to compare trichome density on leaves of nearby small trees and of different branches of the same tree, pairing a stem/branch that appeared damaged with one that appeared healthy. Trichome densities on leaves from damaged stems and branches were significantly greater than were trichome densities on healthy branches and stems. Based on these empirical data, we present several possible explanations for the patterns of association between ants, stem borers, and pubescence. Although feeding preference tests with a common folivore showed no effect of trichome densities on leaves, we suggest further study on how trichomes may affect ovipositing stem borers.     

Trichoderma species form endophytic associations within Theobroma cacao trichomes

Trichoderma species are usually considered soil organisms that colonize plant roots, sometimes forming a symbiotic relationship. Recent studies demonstrate that Trichoderma species are also capable of colonizing the above ground tissues of Theobroma cacao (cacao) in what has been characterized as an endophytic relationship. Trichoderma species can be re-isolated from surface sterilized cacao stem tissue, including the bark and xylem, the apical meristem, and to a lesser degree from leaves. SEM analysis of cacao stems colonized by strains of four Trichoderma species (Trichoderma ovalisporum-DIS 70a, Trichoderma hamatum-DIS 219b, Trichoderma koningiopsis-DIS 172ai, or Trichoderma harzianum-DIS 219f) showed a preference for surface colonization of glandular trichomes versus non-glandular trichomes. The Trichoderma strains colonized the glandular trichome tips and formed swellings resembling appresoria. Hyphae were observed emerging from the glandular trichomes on surface sterilized stems from cacao seedlings that had been inoculated with each of the four Trichoderma strains. Fungal hyphae were observed under the microscope emerging from the trichomes as soon as 6 h after their isolation from surface sterilized cacao seedling stems. Hyphae were also observed, in some cases, emerging from stalk cells opposite the trichome head. Repeated single trichome/hyphae isolations verified that the emerging hyphae were the Trichoderma strains with which the cacao seedlings had been inoculated. Strains of four Trichoderma species were able to enter glandular trichomes during the colonization of cacao stems where they survived surface sterilization and could be re-isolated. The penetration of cacao trichomes may provide the entry point for Trichoderma species into the cacao stem allowing systemic colonization of this tissue.     

Microornamentation of leaf chameleons (Chamaeleonidae: Brookesia,Rhampholeon, and Rieppeleon)—with comments on the evolution of microstructures in the chamaeleonidae

Chameleons (Chamaeleonidae) feature many adaptations to their arboreal lifestyle, including zygodactylous feet, a prehensile tail, and epidermal microstructures. In arboreal tree chameleons, the substrate‐contacting site of the feet and tail is covered by microscopic hair‐like structures (setae) of 6–20 µm length. Their friction enhancing function has been shown in recent studies. Leaf chameleons and one representative of the tree chameleons (Chamaeleo namaquensis) secondarily have become ground‐dwelling. Because leaf chameleons are paraphyletic, one could expect that in the three leaf chameleon genera Brookesia, Rhampholeon, and Rieppeleon and the tree chameleon Ch. namaquensis, epidermis has adapted independently to terrestrial locomotion. Using scanning electron microscopy, we investigated the substrate‐contacting surfaces of the feet (subdigital) of 17 leaf chameleon species and five tree chameleon species that have not yet been examined. Additionally, surfaces not involved in locomotion, the flanks (dorsolateral), and scale interstices, were examined. Although the subdigital microstructures in leaf chameleons are more diverse than in tree chameleons, we found some features across the genera. The subdigital microornamentation of Rhampholeon spinosus consists of long thin setae and spines, comparable to those of tree chameleons. All other Rhampholeon species have spines or short but broad setae. Rh. spectrum had tooth‐like structures instead of setae. Subdigital scales of Brookesia have either thorns or conical scale‐tops in the center and feature honeycomb microstructures. In Rieppeleon, subdigital scales have a thorn. Scale surfaces are covered by honeycombs and short hair‐like structures (spines). As subdigital scales with a thorn in the center and honeycomb microstructures were also found in the terrestrial tree chameleon Ch. namaquensis, one can assume that this geometry is a convergent adaptation to terrestrial locomotion. Despite the great number of genus‐specific traits, the convergent evolution of honey‐comb structures in Brookesia, Rieppeleon, and Ch. namaquensis and the high variability of spines and setae in Rhampholeon suggests a rapid adaptation of subdigital microornamentation in Chamaeleonidae. J. Morphol. 276:167–184, 2015. © 2014 Wiley Periodicals, Inc.