Invasive <Emphasis Type="Italic">Tamarix</Emphasis> (Tamaricaceae) in South Africa: current research and the potential for biological control

Most species of Tamarix originate in Eurasia and at least five species have become invasive around the world, including South Africa. However, T. usneoides is indigenous to southern Africa, where the potential for biological control of the invasive species is being investigated. Recent research on the invasive species is reviewed here with particular reference to these South African biocontrol efforts. The successful biological control programme against invasive Tamarix in the USA, using several species of “Tamarisk beetle”, is being used as a guide for the South African research. The South African programme is complicated by firstly, the presence of the indigenous T. usneoides which raises the precision of host-specificity required, and secondly, the introduced and indigenous Tamarix have a high intrinsic value for phytoremediation of mine tailings dams in South Africa. The phylogenetic proximity of these Tamarix species to each other has contributed to this challenge, which has nevertheless been successfully addressed by molecular techniques used to separate the species. In addition, classical morphological techniques have been used to separate the Tamarisk beetles, so that now they can generally be matched to Tamarix tree species. Overall, it is concluded that given the broad knowledge now available on the ecology and identity of both the trees and their biocontrol agents, the prospects for successful biological control of Tamarix in South Africa are good.     

Ixodid ticks of African buffalo (<Emphasis Type="Italic">Syncerus caffer</Emphasis>), impala (<Emphasis Type="Italic">Aepyceros melampus</Emphasis>) and elephant (<Emphasis Type="Italic">Loxodonta africana</Emphasis>) in five protected park estates in the Zambezi valley,Zimbabwe

Wildlife hosts many pathogens of economic importance and is considered as a reservoir of important tick-borne diseases of livestock in southern Africa. The species composition of ticks parasitizing buffalo (Syncerus caffer), impala (Aepyceros melampus) and elephant (Loxodonta africana) was investigated in five protected parks in the Zambezi valley, Zimbabwe. A total of 1104 adult ticks was collected from 75 adult animals comprising five buffaloes, five elephants and five impalas drawn from five protected wildlife parks. Five tick species belonging to two genera were recovered, with Rhipicephalus decoloratus being the most prevalent species in all the three animal groups. Amblyomma hebraeum was only recovered from buffaloes whereas Rhipicephalus zambeziensis was recovered from buffalos and elephants. Significant differences in mean tick species distribution and concentration were observed amongst the wildlife parks and these appeared to be influenced by the number of hosts in each park. The study revealed that buffaloes are the major host of R. decoloratus in the Zambezi valley. The presence of these ixodid ticks within the Zambezi valley may have significant ecological and economic impacts on wildlife conservation, domestic animals and human health.     

Reduced ant defenses in <Emphasis Type="Italic">Macaranga</Emphasis> myrmecophytes (Euphorbiaceae) infested with a winged phasmid <Emphasis Type="Italic">Orthomeria cuprinus</Emphasis>

Macaranga is a tree genus that includes many species of myrmecophytes, which are plants that harbor ant colonies within hollow structures known as domatia. The symbiotic ants (plant–ants) protect their host plants against herbivores; this defense mechanism is called ‘ant defense’. A Bornean phasmid species Orthomeria cuprinus feeds on two myrmecophytic Macaranga species, Macaranga beccariana and Macaranga hypoleuca, which are obligately associated with Crematogaster ant species. The phasmids elude the ant defense using specialized behavior. However, the mechanisms used by the phasmid to overcome ant defenses have been insufficiently elucidated. We hypothesized that O. cuprinus only feeds on individual plants with weakened ant defenses. To test the hypothesis, we compared the ant defense intensity in phasmid-infested and non-infested M. beccariana trees. The number of plant–ants on the plant surface, the ratio of plant–ant biomass to tree biomass, and the aggressiveness of plant–ants towards experimentally introduced herbivores were significantly lower on the phasmid-infested trees than on the non-infested trees. The phasmid nymphs experimentally introduced into non-infested trees, compared with those experimentally introduced into phasmid-infested trees, were more active on the plant surface, avoiding the plant–ants. These results support the hypothesis and suggest that ant defenses on non-infested trees effectively prevent the phasmids from remaining on the plants. Thus, we suggest that O. cuprinus feeds only on the individual M. beccariana trees having decreased ant defenses, although the factors that reduce the intensity of the ant defenses remain unclear.     

Biotic resistance and the spatiotemporal distribution of an invading woodwasp, <Emphasis Type="Italic">Sirex noctilio</Emphasis>

Quantifying the strength of interactions among introduced and native species across space and time is critical in understanding biological invasions as they can attenuate or amplify the magnitude of impact. The European woodwasp, Sirex noctilio F., a global threat to pines, is a recent invader to North America. It attacks and kills primarily Pinus resinosa and Pinus sylvestris, and encounters a diverse assemblage of potential competitors for this resource. We quantified spatial colonization patterns of this woodwasp and resident competitors including scolytine bark beetles, woodboring cerambycid and buprestid beetles, and the fungal root rot diseases Armillaria and Heterobasidion across an 80 year old pine plantation over 4 years. All xylophages were spatially aggregated, but only on a localized scale of 15–20 m. Colonizers occurred non-randomly within trees, with S. noctilio negatively or neutrally associated with all other colonizing agents, whereas all other insect and root rot colonizers were mostly positively co-associated. An autologistic regression with spatially-weighted variables indicated the probability of a dead tree exhibiting symptoms of S. noctilio attack was positively associated with tree density, host species (P. sylvestris), and density of S. noctilio-attacked trees from the current and previous year. Interspecific stand patterns were weaker; probability of attack was negatively associated only with root rot pathogens. Across spatial scales, there were mixed (woodborers) and neutral (bark beetles) associations between S. noctilio and other co-colonizing insects. Our results suggest that competitive interactions with resident species may be contributing to the limited success of S. noctilio in North America, but are unlikely to be driving it by themselves.     

Distribution and abundance of a keystone tree, <Emphasis Type="Italic">Schinziophyton rautanenii</Emphasis>, and factors affecting its structure in Zambia,southern Africa

Schinziophyton rautanenii is a keystone tree species whose fruits are eaten by wildlife and livestock, nuts are used to extract oil for human use and its wood is used for making curios that are sold to tourists. The species occurs in southern African countries of Angola, Namibia, Botswana, Mozambique, Zambia, Zimbabwe and the Democratic Republic of Congo but little is known about its population and conservation status. The objectives of this study were to (i) assess factors determining the distribution and abundance of the species in Zambia, (ii) evaluate its regeneration potential and population status and (iii) assess whether the main determinants of forest and species stand structures also favour S. rautanenii. The study is based on three forest surveys conducted in 2004, 2005–2008 and 2015. In Zambia S. rautanenii abundance and population structure are controlled by soil, climate and human factors. The species is most abundant in southwest Zambia where population recruitment is good but is rare and recruitment is poor elsewhere in the country. Fire appears to be a major cause of tree damage and possibly death. The elephant is a major disperser of S. rautanenii nuts and the low recruitment levels in the country may partly be attributed to the decline in the elephant population from about 250,000 in 1960 to 28,000 in 2008. Different conservation strategies will be required for areas where the species is rare and for those where the species is abundant. Whatever the strategies, it will be important to address issues of fire management, fruit harvesting and the role of wildlife in the conservation of S. rautanenii in Zambia.     

Taxonomy and pathogenicity of <Emphasis Type="Italic">Leptographium</Emphasis> species associated with <Emphasis Type="Italic">Ips subelongatus</Emphasis> infestations of <Emphasis Type="Italic">Larix</Emphasis> spp. in northern China,including two new species

The larch bark beetle (Ips subelongatus), which occurs in larch plantations over a vast area of eastern Asia, infects both dying and fallen trees. When its population reaches a high density, the beetle may also infect healthy trees, resulting in tree decline and, eventually, death. Leptographium spp., in both their sexual and asexual states, are mainly associated with conifer-infesting bark beetles; some species are important tree pathogens. The aims of this study were to identify the Leptographium spp. associated with I. subelongatus infestations of Larix spp. in northern China and to examine their pathogenicity towards the tree. Morphological studies and phylogenetic approaches based on multilocus DNA sequence data (ITS2- partial r28S, partial β-tubulin, and EF-1α gene regions) showed that three Leptographium species occur in association with I. subelongatus in the areas investigated: Leptographium taigense, which is recorded in China for the first time, and two new species, namely L. innermongolicum sp. nov. and L. zhangii sp. nov. Leptographium innermongolicum is closely related to L. taigense, whereas L. zhangii belongs to the Grosmannia piceaperda species complex. The pathogenicity of these Leptographium species towards mature Larix spp. was tested by stem inoculation in forests. All inoculations only resulted in small lesions on the inner bark; therefore, the three Leptographium species were not considered to be pathogenic.     

Contribution to the study of the genus <Emphasis Type="Italic">Lemniscomys</Emphasis> (Rodentia: Muridae). Morphometric and molecular approaches

A total of 204 specimens belonging to eight species of the genus Lemniscomys were examined with standard morphometric measurements. Our results show that the seven Sub-Saharan species seem to follow a latitudinal gradient from the Center to the South of the African continent. The only North African species L. barbarus looks close to L. griselda and L. rosalia. We also applied a molecular analysis through PCR (Polymerase Chain Reaction) method for the amplification of the 16S rRNA gene. For the purpose of constructing a phylogenetic tree with Maximum Likelihood method, we extracted eight sequences from the GenBank library; seven belonging to the genus Lemniscomys and one to the genus Arvicanthis used as outgroup. We managed to identify a region comprised of 458 nucleotides of which 388 were common for all species and 70 were variable. The phylogenetic tree shows us that the sister group L. bellieri and L. macculus, is the most basal, while L. striatus and L. rosalia appears to be close to the sister group L. barbarus and L. zebra. We also noticed a difference between morphometric and molecular results; the latter are more in agreement with pelage patterns subdivision between Lemniscomys species. These differences can be explained by a high rate of phenotypic evolution that can surpass the molecular counterpart as in the case of the genus Gerbillus.     

Mycorrhizal detection of native and non-native truffles in a historic arboretum and the discovery of a new North American species, <Emphasis Type="Italic">Tuber arnoldianum</Emphasis> sp. nov.

During a study comparing the ectomycorrhizal root communities in a native forest with those at the Arnold Arboretum in Massachusetts (USA), the European species Tuber borchii was detected on the roots of a native red oak in the arboretum over two successive years. Since T. borchii is an economically important edible truffle native to Europe, we conducted a search of other roots in the arboretum to determine the extent of colonization. We also wanted to determine whether other non-native Tuber species had been inadvertently introduced into this 140-year-old Arboretum because many trees were imported into the site with intact soil and roots prior to the 1921 USDA ban on these horticultural practices in the USA. While T. borchii was not found on other trees, seven other native and exotic Tuber species were detected. Among the North American Tuber species detected from ectomycorrhizae, we also collected ascomata of a previously unknown species described here as Tuber arnoldianum. This new species was found colonizing both native and non-native tree roots. Other ectomycorrhizal taxa that were detected included basidiomycetes in the genera Amanita, Russula, Tomentella, and ascomycetes belonging to Pachyphlodes, Helvella, Genea, and Trichophaea. We clarify the phylogenetic relationships of each of the Tuber species detected in this study, and we discuss their distribution on both native and non-native host trees.     

Vegetation Thresholds for the Occurrence and Dispersal of <Emphasis Type="Italic">Microcebus griseorufus</Emphasis> in Southwestern Madagascar

Primate populations are declining the world over due to anthropogenic threats, including habitat loss and degradation. This raises the important question of how much habitat degradation a species can cope with. Habitat degradation is pronounced in Madagascar, where most of the human population depends on the direct exploitation of natural resources. We aimed to identify the response of Microcebus griseorufus (the gray-brown mouse lemur) to forest degradation and to define the structural traits of the vegetation that might be crucial for the species’ occurrence in anthropogenic landscapes. We documented the occurrence of Microcebus griseorufus in relation to vegetation structures along a gradient of forest degradation, at the edge of and west of Tsimanampetsotsa National Park in April and May 2007 and from October to December 2015. We confirmed the occurrence of Microcebus griseorufus using trapping and visual surveys, and measured vegetation structure. Logistic regression models showed that Microcebus griseorufus has a threshold response to tree density and the diameter of thick trees. The thresholds of occurrence were at 10–15% of the tree density recorded in intact forest and a mean diameter of trees with a diameter at breast height of >10 cm of 14.3 cm. The definition of such thresholds might help to maintain suitable habitat for this species and other primates living in anthropogenic landscapes, providing connectivity between isolated protected areas and allowing dispersal between populations.     

Biologically feasible gene trees,reconciliation maps and informative triples

Background

The history of gene families—which are equivalent to event-labeled gene trees—can be reconstructed from empirically estimated evolutionary event-relations containing pairs of orthologous, paralogous or xenologous genes. The question then arises as whether inferred event-labeled gene trees are biologically feasible, that is, if there is a possible true history that would explain a given gene tree. In practice, this problem is boiled down to finding a reconciliation map—also known as DTL-scenario—between the event-labeled gene trees and a (possibly unknown) species tree.

Results

In this contribution, we first characterize whether there is a valid reconciliation map for binary event-labeled gene trees T that contain speciation, duplication and horizontal gene transfer events and some unknown species tree S in terms of “informative” triples that are displayed in T and provide information of the topology of S. These informative triples are used to infer the unknown species tree S for T. We obtain a similar result for non-binary gene trees. To this end, however, the reconciliation map needs to be further restricted. We provide a polynomial-time algorithm to decide whether there is a species tree for a given event-labeled gene tree, and in the positive case, to construct the species tree and the respective (restricted) reconciliation map. However, informative triples as well as DTL-scenarios have their limitations when they are used to explain the biological feasibility of gene trees. While reconciliation maps imply biological feasibility, we show that the converse is not true in general. Moreover, we show that informative triples neither provide enough information to characterize “relaxed” DTL-scenarios nor non-restricted reconciliation maps for non-binary biologically feasible gene trees.

     

Forest decline after a 15-year “perfect storm” of invasion by hemlock woolly adelgid,drought, and hurricanes

Invasions by introduced pests can interact with other disturbances to alter forests and their functions, particularly when a dominant tree species declines. To identify changes after invasion by the insect hemlock woolly adelgid (Adelges tsugae; HWA), coinciding with severe droughts and hurricanes, this study compared tree species composition of eastern hemlock (Tsuga canadensis) forests on 11 plots before (2001) and 15 years after (2016) invasion in the southern Appalachian Mountains, USA. Losses of hemlock trees after HWA invasion were among the highest reported, with a 90% decline in density, 86% decline in basal area, and 100% mortality for individuals ≥ 60 cm in diameter. In contrast to predictions of theoretical models, deciduous tree density declined after HWA invasion, while basal area changed little, at least during the initial 15 years after invasion. Overall, forest density declined by 58%, basal area by 25%, and tree species richness by 8%. Factors additional to HWA likely exacerbating forest decline included: droughts before (1999–2001) and after HWA invasion (2006–2008); tree uprooting from hurricane-stimulated winds in 2004; pest-related declines of deciduous tree species otherwise likely benefitting from hemlock’s demise; death of deciduous trees when large hemlocks fell; and competition from aggressive understory plants including doghobble (Leucothoe fontanesiana), rosebay rhododendron (Rhododendron maximum), and Rubus spp. Models of forest change and ecosystem function should not assume that deciduous trees always increase during the first decades after HWA invasion.     

Uncorrelated mistletoe infection patterns and mating success with local host specialization in <Emphasis Type="Italic">Psittacanthus calyculatus</Emphasis> (Loranthaceae)

Mistletoe infection between conspecific and interspecific hosts can be restricted by seed dispersal, host-mistletoe compatibility and abiotic factors, yet no studies have linked mistletoe infection patterns and pollination together for understanding mistletoe distribution at a local scale. Psittacanthus calyculatus (Loranthaceae) is a hemiparasitic plant with a broad host range across its geographic distribution. The potential for local host adaptation has been shown using cross-inoculation experiments, in which plants of mistletoe seeds collected from a given host are more likely to survive when they are inoculated on conspecific host trees compared with those inoculated on other host provenances. Here we evaluate host adaptation by describing the local patterns of infection (prevalence and intensity) of P. calyculatus mistletoes on three native host tree species (Alnus acuminata, Quercus crassipes, Salix bonplandiana) and one introduced species (Populus alba) and carried out cross-pollination experiments to examine how pollination affects infection patterns of different host species. Mistletoe infection prevalence (proportion of infection) and infection intensity (mean number of mistletoes per tree) were in general disproportional with respect to the availability of native host tree species but higher to that of non-native host tree species. Cross-pollination experiments showed higher mating success on the native host tree species, suggesting higher local adaptation to specially Q. crassipes. The observed spatial distribution of host tree species and mistletoe infection along with the non-random mating could contribute to local genetic structuring of mistletoe populations.     

Species tree phylogeny,character evolution,and biogeography of the Patagonian genus <Emphasis Type="Italic">Anarthrophyllum</Emphasis> Benth. (Fabaceae)

Geologic events promoting the aridization of southern South America contributed to lineage divergences and species differentiation through geographic (allopatric divergence) and biotic and abiotic factors (ecological divergence). For the genus Anarthrophyllum, which is distributed in arid and semi-arid regions of Patagonia, we assessed how these factors affected species diversification and reconstructed its possible biogeographic history in South American arid environments. Sequences were obtained from two molecular markers: the ITS nuclear region and the trnS-trnG plastid region. Using Parsimony, Maximum likelihood and Bayesian inference individual gene trees were reconstructed, and a species tree was obtained using multi-species coalescent analysis. Divergence times among species were estimated using secondary calibrations. Flexible Bayesian models and stochastic character mapping were used to elucidate ancestral geographic distributions and the evolution of the floral and vegetative phenotypes in the genus. Gene trees and species tree analyses strongly support Anarthrophyllum as monophyletic; all analyses consistently retrieved three well-supported main clades: High Andean Clade, Patagonian Clade 1, and Patagonian Clade 2. Main diversification events occurred concomitant with the Andean uplift and steppe aridization; the Andean mountain range possibly acted as a species barrier for the High Andean Clade. Vegetative traits showed adaptations to harsh climates in some clades, while pollinator-related floral features were associated with independent diversification in bee- and bird-pollinated clades within both Patagonian Clades. In conclusion, evolutionary and biogeographic history of Anarthrophyllum resulted from the action of ecological, historical, and geographic factors that acted either alternatively or simultaneously.     

Updates on East Asian <Emphasis Type="Italic">Asterostroma</Emphasis> (Russulales,Basidiomycota): new species and new records from Thailand and China

Two new species from northern Thailand, Asterostroma bambusicola and A. vararioides, are described and illustrated. Asterostroma bambusicola is characterized by globose, echinulate, and amyloid basidiospores and growing on rotten bamboo. Asterostroma vararioides is distinguished by the presence of Vararia-like dichohyphae, subglobose, smooth, and amyloid basidiospores and growing on bark of living angiosperm trees. In the phylogenetic tree inferred from a combined dataset of ITS and nLSU sequence data of Peniophoraceae, A. bambusicola forms a distinct lineage in the sect. Asterostroma clade, whereas A. vararioides and A. laxum form the sect. Laevispora clade. Asterostroma andinum, reported from China for the first time, forms a distinct lineage sister to Gloiothele spp. and Scytinostroma portentosum group. Asterostroma muscicola is reported from Thailand and China for the first time. A key to the species of Asterostroma from Thailand and China is provided.     

Identifying the ecological and societal consequences of a decline in <Emphasis Type="Italic">Buxus</Emphasis> forests in Europe and the Caucasus

The potential impact of new invasive tree pests and diseases is usually quantified in economic terms. The ecological and social impacts are less often assessed. Using a comprehensive literature review we assess the potential ecological and social impact of two non-native invasive species (the box tree moth, Cydalima perspectalis and the fungus Calonectria pseudonaviculata) that threaten the survival of box tree, Buxus spp. in forests in Europe and the Caucasus. A total of 132 fungi, 12 chromista (algae), 98 invertebrate and 44 lichens were found to use Buxus spp. Of these, 43 fungi, 3 chromista and 18 invertebrate species have only been recorded on Buxus spp., suggesting that these species are obligate on Buxus spp. and are most at risk from in the loss of Buxus spp. due to these invasive pest and disease species. Buxus spp. was shown to be important for soil stability and water quality but there was no information on other ecosystem functions provided by Buxus spp. Buxus was found to be of considerable historical cultural importance but there was very limited information on current social values and uses. Buxus trees, wood and leaves are associated with different folklore and sacred rites which are still particularly important in the Caucasus. While we could not find any assessment of the economic value of Buxus forests the biodiversity, cultural and social values of Buxus identified here indicate that its loss could have major indirect and non-market economic effects. This work highlights the importance of studying the ecological and societal implications of biological invasions.     

Stomatal conductance in mature deciduous forest trees exposed to elevated CO<Subscript>2</Subscript>

Stomatal conductance (g _s) of mature trees exposed to elevated CO₂ concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g _s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO₂ compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO₂ treatment. During the 2003 drought in the third treatment year, the CO₂ effect became reversed in Carpinus, resulting in higher g _s in trees exposed to elevated CO₂ compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO₂ enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO₂-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO₂.     

Savanna chimpanzee (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) nesting ecology at Bagnomba (Kedougou,Senegal)

We studied the nesting behavior of the critically endangered West African chimpanzee (Pan troglodytes verus). We assumed that the nesting data stemmed from a single, unhabituated community at the Bagnomba hill site in the savanna-woodlands of southeastern Senegal. The aim of this study was to examine chimpanzees’ nesting habits in terms of the tree species utilized and sleeping nest heights. We recorded a total of 550 chimpanzee nests at Bagnomba between January 2015 and December 2015. The chimpanzees here made nests in particular tree species more often than others. The majority of nests (63%) were in two tree species: Diospyros mespiliformis and Pterocarpus erinaceus. The average height of nesting trees was 10.54 m (SD 3.91, range, 0.0–29.0 m) and average nest height was 7.90 m (SD 3.62, range, 0.0–25.0 m). The result of a linear regression analysis (r = 0.7874; n = 550; p < 0.05) is consistent with a preference for nesting at a particular height. Bagnomba chimpanzees rarely made ground nests (0.36% of nests), but the presence of any ground nesting was unexpected, given that at least one leopard (Panthera pardus) also occupied the hill. This knowledge will enable stakeholders involved in the protection of chimpanzees specifically and of biodiversity in general to better understand chimpanzee ecology and inform a conservation action plan in Senegal where the survival of this species is threatened.