Host specificity in parasitic mistletoes (Loranthaceae) in New Zealand

1. We quantify the degree of host specificity for the five extant New Zealand loranthaceous mistletoes ( Alepis flavida, Ileostylus micranthus, Peraxilla colensoi, Peraxilla tetrapetala and Tupeia antarctica ).
2. Host specificity is highest for A. flavida, P. colensoi and P. tetrapetala which primarily parasitize species of Nothofagus , and lowest for T. antarctica and especially I. micranthus which parasitize a wide range of host species.
3. These patterns of host specificity support the suggestion that relative host abundance is a key factor determining the degree of host specialization in mistletoes (resource fragmentation hypothesis). While evolutionary history may be important in the specificity of the mistletoe–host relationship in some situations, our data suggest that for New Zealand mistletoes evolutionary history simply reflects the temporal component of relative host abundance.
4. We conclude that it is the stability of host availability through time and space which is the dominant factor determining host specificity patterns.     

Heterogeneity in vertebrate and invertebrate herbivory and its consequences for New Zealand mistletoes

Vertebrate herbivores generally have greater effects than invertebrates on plants. However, few studies have investigated the effects of both invertebrate and vertebrate herbivores on a single plant species. In New Zealand, nationwide declines in mistletoe populations have often been attributed to possum herbivory, but never to insect herbivory. The main goal of the present study was to document levels of vertebrate and invertebrate herbivory on endemic New Zealand mistletoe plants to suggest whether herbivory is leading to mistletoe decline. In the present study, the annual amount of leaf loss from herbivory by the brushtail possum (Trichosurus vulpecula), insect herbivory and leaf abscission were measured in two populations each of three mistletoe species (Alepis flavida, Peraxilla colensoi, and Peraxilla tetrapetala, Loranthaceae). In two populations of each species from February 1997 to February 1998, abscission accounted for the most leaf loss (range 10–84% of total mean leaf area, mean 33%), whereas insects and possums usually removed small and similar amounts (less than 3%). Possum browse caused large amounts of abscission in only one population (A. flavida at Eglinton). Observed possum browse was more heterogeneous than insect browse among branches within a plant (possum coefficient of variation = 2.63, insect CV = 1.98, P < 0.001), among plants in a population (possum CV = 2.15, insect CV = 0.69, P < 0.001), and between populations (possum CV = 1.36, insect CV = 1.09). Moreover, insects damaged 100% of the study plants but never removed more than 16% of leaf area on a single plant, whereas possums only browsed 32% of the study plants but severely defoliated some plants. Thus, while the mean amount of biomass removed across a population may have important consequences for mistletoe survival, the effect of possums on mistletoe populations may also depend on the heterogeneity of browse among individuals in the population.     

Sustainable utilization of woodrose-producing mistletoes (Loranthaceae) in South Africa

Pedistylis galpinii and Erianthemum dregei produce woodrose outgrowths at attachment points with host trees. In some rural areas of southern Africa, woodroses are sold as curios. The socio-economics related to woodrose harvesters were investigated via interview schedules and field survey. Harvesting is not detrimental to hosts as only branches infected with mistletoes are cut. Over half the harvesters interviewed reported harvesting both live and dead mistletoes while 20% used only dead ones. The density of dead mistletoes could satisfy present demand. Such sustainable harvesting practices (only using dead woodroses) allow for woodrose harvesting according to calculated quotas. The median gross income per month generated from woodroses was estimated to be between US $38 and 90. Eleven percent of harvesters relied on woodroses as their only source of income. Marketing was relatively haphazard, involving informal selling at roadsides and unscheduled visits to retailers. The market for woodroses can be expanded and the potential exists to increase supply considerably on a sustainable basis.     

Host-preference and density of woodrose-forming mistletoes (Loranthaceae) on savanna vegetation,South Africa

In the Bushbuckridge region of South Africa host preference and density of two woodrose-forming mistletoes, Erianthemum dregei (Eckl. & Zeyh.) V. Tieghem and Pedistylis galpinii (Schinz ex Sprague) was quantified in relation to land-use (harvested or unharvested), rainfall (high > 660 or low < 660 mm year^–1) and catenal position (top or lower slope). These two mistletoes are generalist hemi-parasites of savanna trees and shrubs occurring on 25 and 17 hosts respectively, seven of which are shared. Thirty-six percent of woody plant species recorded were found to be hosts. Although Sclerocarya birrea (A. Rich.) Hochst. comprised only 4% of woody plant density in the environment, it was the principal host for both mistletoes, accounting for 71% of total E. dregei and 42% of P. galpinii infection. Mistletoe infection relative to density of Ficus stuhlmanii, Trichilia emetica and Cassine transvaalensis indicated that these were preferential hosts to S. birrea. Mistletoe host preference was negatively correlated with host wood density. Mistletoe number per tree had a weak relationship to canopy size. Mistletoes of all size classes were denser at high rainfall relative to low rainfall sites. Interestingly, the overall mistletoe size class distribution was similar between harvested and unharvested sites. The ratio of living to dead mistletoe was 2 to 1 for E. dregei and 1.5 to 1 for P. galpinii. There are sufficient dead mistletoes in unharvested and harvested areas to satisfy present market demand. Living E. dregei predominated in harvested rather than unharvested areas suggesting that current-harvesting levels had little or no negative effect on the population. In contrast, P. galpinii was denser in unharvested areas possibly owing to its higher market value and thus higher harvesting levels.     

Nitrogen concentration and mimicry in some New Zealand mistletoes

Summary The nitrogen concentration in photosynthetic organs of 41 pairs of mistletoes and their respective hosts was compared and found to be higher (in relation to that of their hosts) in cryptic mistletoes and lower in non-cryptic mistletoes. This supports the hypothesis that crypsis may have evolved as a mechanism for avoiding predation by vertebrate herbivores. There were no consistent differences between the nitrogen concentrations of hosts and mistletoes in trees and larger-leaved shrubs (whether native or introduced). The largest differences between mistletoes and their hosts were found on small-leaved divaricate shrubs (Coprosma ssp., Melicope simplex) where the nitrogen concentrations in cryptic mistletoes (Korthalsella spp.) were significantly higher than in their hosts, but significantly lower in prominent mistletoes (Loranthus micranthus, Tupeia antarctica) which apparently advertise their unpalatability. If crypsis in New Zealand mistletoes evolved as a protection against herbivory, then it must have evolved in the absence of mammalian herbivores, and the appropriate selection pressures could have been provided only by moas, extinct ratite birds. Otherwise, alternative explanations, such as differing relationships between water use and nitrogen uptake, must be sought for the observed associations of nitrogen concentration and cryptic mimicry in New Zealand mistletoes.     

Occurrence of the lutein-epoxide cycle in mistletoes of the Loranthaceae and Viscaceae

The lutein-epoxide cycle (Lx cycle) is an auxiliary xanthophyll cycle known to operate only in some higher-plant species. It occurs in parallel with the common violaxanthin cycle (V cycle) and involves the same epoxidation and de-epoxidation reactions as in the V cycle. In this study, the occurrence of the Lx cycle was investigated in the two major families of mistletoe, the Loranthaceae and the Viscaceae. In an attempt to find the limiting factor(s) for the occurrence of the Lx cycle, pigment profiles of mistletoes with and without the Lx cycle were compared. The availability of lutein as a substrate for the zeaxanthin epoxidase appeared not to be critical. This was supported by the absence of the Lx cycle in the transgenic Arabidopsis plant lutOE, in which synthesis of lutein was increased at the expense of V by overexpression of -cyclase, a key enzyme for lutein synthesis. Furthermore, analysis of pigment distribution within the mistletoe thylakoids excluded the possibility of different localizations for the Lx- and V-cycle pigments. From these findings, together with previous reports on the substrate specificity of the two enzymes in the V cycle, we propose that mutation to zeaxanthin epoxidase could have resulted in altered regulation and/or substrate specificity of the enzyme that gave rise to the parallel operation of two xanthophyll cycles in some plants. The distribution pattern of Lx in the mistletoe phylogeny inferred from 18S rRNA gene sequences also suggested that the occurrence of the Lx cycle is determined genetically. Possible molecular evolutionary processes that may have led to the operation of the Lx cycle in some mistletoes are discussed.Abbreviations A antheraxanthin - - and -Car - and -carotene - Chl chlorophyll - -DM dodecyl--d-maltoside - DPS de-epoxidation state of the violaxanthin cycle (= [A+Z]/[V+A+Z]) - Lut lutein - Lx lutein epoxide - Caro total carotenoid concentration - V violaxanthin - VAZ pool size of the violaxanthin cycle (= V+A+Z) - VDE violaxanthin de-epoxidase - Z zeaxanthin - ZE zeaxanthin epoxidase     

Population structure and phylogeography of the mistletoes Tristerix corymbosus and T. aphyllus (Loranthaceae) using chloroplast DNA sequence variation

The mistletoe Tristerix corymbosus (Loranthaceae) is present in the temperate forest and Chilean matorral biomes of Chile and northwest Patagonia. The closely related cactus-specific species, T. aphyllus, occurs only in the matorral biome. The population structure of these mistletoes was examined to determine whether the distribution of haplotypes corresponds mostly to geographic zone, biome, or other biotic factors. Samples from 108 individuals in 26 localities of T. corymbosus and 13 individuals in four localities of T. aphyllus were collected. Sequences were obtained from two chloroplast genome regions: the atpB-rbcL spacer and the trnL-F region. Haplotypes were analyzed using parsimony and Bayesian trees as well as parsimony networks. All methods placed the haplotypes in four clades, one of which corresponded to T. aphyllus and the others to T. corymbosus. Within T. corymbosus, the different clades did not correlate with biome, geographical region, host, or any apparent morphological feature of the mistletoe. The morphologically distinct cactus parasite T. aphyllus likely arose in sympatry from an unspecialized tree parasite, T. corymbosus, after a host switch. The present day haplotype distribution is complex and resulted from post-glaciation migrations from multiple Pleistocene refugia.     

Carbon and nitrogen isotope ratios, nitrogen content and heterotrophy in New Zealand mistletoes

The carbon isotope ratio ('¹³C) of New Zealand mistletoes (-29.51ǂ.10‰) and their hosts (-28.89ǂ.12‰) is generally more negative, and shows less difference between mistletoes and their hosts, than found in previous studies. In 37% of the examined pairs, the '¹³C of mistletoes was less negative than that of their hosts. These reversals were not associated with the relative position (proximal or distal) of the host material with regard to the mistletoe. Differences between host and mistletoe tended to be greater on hosts with less negative '¹³C. Both nitrogen content and isotope ratio ('¹⁵N) of the mistletoe leaves were strongly correlated with those of their hosts. Nitrogen contents of mistletoe leaves were similar to those of their hosts at low nitrogen contents but proportionately less on hosts with a high nitrogen content, whereas '¹⁵N of mistletoes was consistently similar to that of their hosts. The '¹³C of mistletoes was related to both host nitrogen content and '¹⁵N, but '¹³C in host tissue was related to neither, suggesting that the mistletoes derived both nitrogen and carbon from their hosts. The '¹³C of both hosts and mistletoes were significantly related to leaf conductance and carbon dioxide concentration but relationships with transpiration and water use efficiency were not significant. In all cases there was no clear separation between the responses of hosts and mistletoes. This may be related to the similarity of stomatal conductance, transpiration and photosynthesis in the studied mistletoes and their hosts and is consistent with the small differences in '¹³C between mistletoes and hosts found in this study. Consequently, the estimation of mistletoe heterotrophy from carbon discrimination is confounded, as the small difference between host and mistletoe carbon discrimination could equally well result from either similarities in photosynthesis and water relations or heterotrophic assimilation of host-derived carbon. The differences between our study and previous studies (which are mostly from seasonally dry or semi-arid to arid environments) may be related to the temperate environment in which these mistletoes grow. Water is freely available so that the mistletoe is able to obtain sufficient water and dissolved nutrients without having to maintain the high transpiration rate and low water potentials that are needed to extract water from a water-stressed host. Similarly, mistletoe photosynthesis is less inhibited by water stress. The physiological similarities between mistletoe and hosts from a temperate environment are reflected in their similar '¹³C values.     

Reaching the canopy on the ground: incidence of infection and host-use by mistletoes (Loranthaceae and Viscaceae) on trees felled for timber in Amazonian rainforests

There is a profound absence of knowledge of infestation prevalence and host-use by mistletoes of mature South American tropical rainforests. In this study, we fill this gap using information gathered from felled trees at a logging concession area in Amazonian Brazil. We sampled individuals of 18 tree species, which occurred in two forest physiognomies; open forest with canopy interrupted by palm trees and closed, denser forest, with emergent trees. We hypothesized that infection incidence would be higher in open than in closed forest, irrespective of the mistletoe species involved. In addition, we expected that mistletoe parasitism would be higher on host species that were more abundant, taller, deciduous, and had less dense wood. We sampled 870 individual trees in both sites combined. All but one host species was infected by at least one species of mistletoe. We found 13 mistletoe species/morphospecies, Loranthaceae (7) and Viscaceae (6), parasitizing very different hosts. Mistletoe infection incidence was higher in the closed forest (10.3%) than in the open forest (5.4%). In the closed forest, host height influenced incidence positively, while deciduousness had a negative influence. Our results show that mistletoes are common in the canopy of pristine tropical forests and, contrary to expectations, that infection incidence was higher in the closed forest. The positive relation between infection incidence and host height in this forest type suggests that emergent trees have higher chances of being infected than individuals of correspondent species in the lower forest layers.     

A rejoinder onOryctina (Loranthaceae)

Both of the two main generic features ofOryctina (Loranthaceae), i.e., its dioeciousness and its absence of floral bracteoles, are based on misinterpretations. Instead, both species appear to be monoecious, and both are bracteolate. Nevertheless,Oryctina should be maintained as a distinct genus, probably most closely related toMaracanthus.     

Validation and redescription ofOryctina (Loranthaceae)

The formerly obscure genusOryctina v.Tiegh. is redescribed and validated. It is shown to include two species, an old one, previously known asOryctanthus scabridus Eichl., nowOryctina scabrida (Eichl.) v.Tiegh., and a new one,Oryctina subaphylla Rizz., both from Brazil.     

Effectiveness of short‐tongued bees as pollinators of apparently ornithophilous New Zealand mistletoes

Abstract The flowers of two species of threatened New Zealand mistletoes (Peraxilla tetrapetala and Peraxilla colensoi, Loranthaceae) have explosive buds that do not open unless force is applied by birds or two species of native short‐tongued bees. Opened flowers are visited by a variety of birds and insects. Although both species of Peraxilla conform to a pollination syndrome of ornithophily, bees may be effective alternative pollinators. We investigated the effectiveness of bees and birds as pollinators of P. colensoi at one site and P. tetrapetala at two sites in the South Island. Bees and other insects outnumbered birds as flower visitors at all three sites. By excluding birds with wire cages, we showed that two bee species regularly open flowers of P. tetrapetala, but only rarely open flowers of P. colensoi. Few pollen grains were deposited when either birds or bees opened buds, so opening buds was not by itself sufficient for adequate pollination. Instead, pollen continued to accumulate over the next 6 or 7 days, even inside cages that excluded birds. Both populations of P. tetrapetala were regularly pollen‐limited, but in different ways. At Ohau, opened flowers gained enough pollen to produce seeds, but many buds were not opened and hence failed to set seed. In contrast, at Craigieburn, nearly all buds were opened, but many of these did not receive enough pollen. These results demonstrate that native bees can partially replace birds as pollinators of mistletoes, despite their apparent ornithophilous syndrome. Ongoing reductions in New Zealand forest bird numbers means that the service bees provide may be important for the long‐term future of these plants.     

New species of Acizzia Heslop-Harrison (Hemiptera: Psyllidae) from Australian mistletoe (Loranthaceae)

Three new species of Acizzia are described from Australian mistletoe (Loranthaceae: Amyema spp.). These are: A. loranthacae sp. n., A. amyemae sp. n. and A. pendulae sp. n. They are characterised by: male proctiger with prominent posterior lobe; antenna 1.9–2.9 times width of head; forewing with costal break and pterostigma, with pattern in A. amyemae and A. pendulae but not in A. loranthacae ; and female proctiger simple in A. amyemae and A. pendulae but with prominent dorso-apical process in A. loranthacae . The Loranthaceae is a new host family record for Acizzia . Given the probable radiation of Acizzia on mimosaceous hosts, its occurrence on eucalypt-inhabiting mistletoe, yet its apparent absence from mistletoes on Acacia, is discussed.     

Bone growth marks reveal protracted growth in New Zealand kiwi (Aves,Apterygidae)

The presence of bone growth marks reflecting annual rhythms in the cortical bone of non-avian tetrapods is now established as a general phenomenon. In contrast, ornithurines (the theropod group including modern birds and their closest relatives) usually grow rapidly in less than a year, such that no annual rhythms are expressed in bone cortices, except scarce growth marks restricted to the outer cortical layer. So far, cyclical growth in modern birds has been restricted to the Eocene Diatryma, the extant parrot Amazona amazonica and the extinct New Zealand (NZ) moa (Dinornithidae). Here we show the presence of lines of arrested growth in the long bones of the living NZ kiwi (Apteryx spp., Apterygidae). Kiwis take 5–6 years to reach full adult body size, which indicates a delayed maturity and a slow reproductive cycle. Protracted growth probably evolved convergently in moa and kiwi sometime since the Middle Miocene, owing to the severe climatic cooling in the southwest Pacific and the absence of mammalian predators.     

Effects of forest edges on herbivory in a New Zealand mistletoe,

This study examined how forest edges influenced leaf and floral herbivory, as well as seed predation, in a native New Zealand mistletoe species, Alepis flavida. Plants growing on forest edges and in forest interior were compared, and effects of plant size and the neighbouring conspecific plant community were also examined. Leaf herbivory by possums was significantly greater on forest edges than in forest interior in a year of high possum damage, but not in a year with low damage levels. Insect leaf herbivory did not differ between forest edges and interior. Although equal numbers of plants on edges v. interior experienced some floral damage by a specialist caterpillar, there were significantly higher levels of damage on plants growing in the forest interior than on forest edges. Plants with floral damage were larger than plants without damage, and distance to neighbouring mistletoe plants was positively correlated with amount of floral damage, but only for plants in the interior. Significantly greater numbers of plants on edges than in the interior exhibited seed predation by the same specialist caterpillar that caused floral damage, suggesting greater fruit abortion rates in the interior. Amounts of seed damage were inversely correlated with plant size. Forest edges had much stronger effects on leaf herbivory by possums, as well as floral herbivory and seed predation, than did plant size or the neighbouring plant community.     

Developing a forest biodiversity monitoring approach for New Zealand

There is a lack of comprehensive and consistent information to inform policy makers about the status of New Zealand’s forest biodiversity. Three reasons for collecting such information are: assessing the effectiveness of management, reporting on the status of biodiversity under national and international requirements, and improving our knowledge of ecosystem dynamics for designing effective management systems. The challenge is to design monitoring systems that address these multiple needs simultaneously, and at a range of spatial and temporal scales. This article first considers principles for designing enduring monitoring systems based on past experiences, assessing how effectively these principles were implemented in designing New Zealand’s Carbon Monitoring System (CMS), and finally, suggesting future directions for forest biodiversity monitoring in New Zealand. At a national scale we support an unbiased, systematic sample of forests as implemented in several countries (e.g. Austria and the U.S.A.). We consider it best practice to monitor shifts in the fundamental compositional, structural and functional characteristics of ecosystems and use these to derive indicators. We suggest forest biodiversity indicators should include forest area and spatial arrangement, tree mortality and recruitment, exotic weeds, introduced herbivore impacts, and woody debris. Principles discussed in this paper are relevant to biodiversity monitoring in a wider range of ecosystems than forests. Without spatially extensive, robustly designed, biodiversity monitoring systems, New Zealand will remain in a relatively weak position nationally, and internationally, to report on the effectiveness of biodiversity conservation.     

A second species of Ligaria (Loranthaceae)

The combinationLigaria teretiflora (Rizzini) Kuijt is proposed, based onPsittacanthus teretiflorus Rizzini, adding a second species to the genus.Ligaria teretiflora differs fromL. cuneifolia (Ruiz & Pavón) Van Tieghem in having articulated pedicels and obovate leaves which lack a necrotic apex.     

A re-evaluation of Phrygilanthus nudus (Loranthaceae)

New information on the fruit, inflorescence, flower, and pollen ofPhrygilanthus nudus Molina indicates that it belongs toPsittacanthus, leading to the new combinationPsittacanthus nudus (Molina) Kuijt & Feuer.