Comparative wood anatomy of epacrids (Styphelioideae,Ericaceae s.L.)

The wood anatomy of 16 of the 37 genera within the epacrids (Styphelioideae, Ericaceae s.l.) is investigated by light and scanning electron microscopy. Several features in the secondary xylem occur consistently at the tribal level: arrangement of vessel-ray pits, distribution of axial parenchyma, ray width, and the presence and location of crystals. The primitive nature of Prionoteae and Archerieae is supported by the presence of scalariform perforation plates with many bars and scalariform to opposite vessel pitting. The wood structure of Oligarrheneae is similar to that of Styphelieae, but the very narrow vessel elements, exclusively uniseriate rays and the lack of prismatic crystals in Oligarrheneae distinguish these two tribes. The secondary xylem of Monotoca tamariscina indicates that it does not fit in Styphelieae; a position within Oligarrheneae is possible. Like most Cosmelieae, all Richeeae are characterized by exclusively scalariform perforation plates with many bars, a very high vessel density and paratracheal parenchyma, although they clearly differ in ray width (exclusively uniseriate rays in Cosmelieae vs. uniseriate and wide multiseriate rays in Richeeae). Several wood anatomical features confirm the inclusion of epacrids in Ericaceae s.l. Furthermore, there are significant ecological implications. The small vessel diameter and high vessel frequency in many epacrids are indicative of a high conductive safety to avoid embolism caused by freeze-thaw cycles, while the replacement of scalariform by simple vessel perforation plates and an increase in vessel diameter would suggest an increased conductive efficiency, which is especially found in mesic temperate or tropical Styphelieae.     

Pollen tetrad segregation and pollen ovule ratios in six species of Acrotriche (Styphelioideae: Ericaceae)

Pollen morphology can be useful in the determination of phylogeny and mating systems of plants. Pollen ovule ratios can give an indication of the mating system of plant species, and there are a number of theories as to reductions in response to pollination strategies. Pollen tetrad segregation occurs when a number of pollen grains within a tetrad abort; this process has evolved a number of times in the family Ericaceae. Pollen ovule ratios along with the total number of tetrads, number of viable grains and proportions of each of the tetrad types were investigated in six species of Acrotriche (Styphelioideae: Ericaceae). Pollen ovule ratios were used in comparison with related species to give indications of mating systems and to examine theories regarding reductions in pollen production. The study species can be arranged into distinct groupings based on the proportions of tetrad types that correlate with floral morphology and may be indicative of phylogentic associations.     

Supermatrices, supertrees and serendipitous scaffolding: inferring a well-resolved, genus-level phylogeny of Styphelioideae (Ericaceae) despite missing data

For the predominantly southern hemisphere plant group Styphelioideae (Ericaceae) published sequence datasets of five markers are now available for all except one of the 38 recognised genera. However, several markers are highly incomplete therefore missing data is problematic for producing a genus level phylogeny. We explore the relative utility of supertree and supermatrix approaches for addressing this challenge, and examine the effects of missing data on tree topology and resolution. Although the supertree approach returned a more conservative hypothesis, overall, both supermatrix and supertree analyses concurred in the topologies they returned. Using multiple genes and a dataset of variably complete taxa we found improved support for the monophyly and position of the tribes and genus level relationships. However, there was mixed support for the Richeeae tribe appearing one node basal to the Cosmelieae tribe or vice versa. It is probable that this will only be resolved through further sequencing. Our study supports previous findings that the amount of data is more critical than the completeness of the dataset in estimating well-resolved trees. Our results suggest that a “serendipitous” scaffolding approach that includes a mixture of well and poorly sequenced taxa can lead to robust phylogenetic hypotheses.     

Fleshy-fruitedness in the New Zealand flora

Aim It has been claimed that the New Zealand flora has an unusually high frequency of fleshy-fruitedness. This paper tests whether fleshy-fruitedness is indeed more common in New Zealand than in other temperate floras, then examines the distribution of fleshy-fruitedness among taxa and floristic elements to determine whether the flora conforms to predictions for a continental island with a relictual floristic element. Lastly, I test the extent to which fleshy-fruitedness has influenced colonization success and subsequent speciation within New Zealand. Methods Information on fruit characteristics for all indigenous seed plants was extracted from the Flora of New Zealand series and analysed with χ² tests. Results Contrary to previous claims fleshy-fruitedness was not unusually common in the New Zealand flora as a whole, when compared with other temperate floras. It is only more common in alpine communities and among trees. I also found no evidence for selective immigration; fleshy-fruited New Zealand genera were not more likely, than dry-fruited genera, to also occur in Australia. Furthermore there is no evidence that the New Zealand environment has favoured fleshy-fruited taxa; there has been no autochthonous evolution of fleshy-fruitedness in New Zealand, fleshy-fruitedness has had no significant effect on speciation within New Zealand, and endemic genera are no more likely to be fleshy-fruited than nonendemic genera. Fleshy-fruitedness in New Zealand is, however, strongly related to floristic elements of the flora. New Zealand is a continental island and therefore, theoretically, those elements of the flora dating from a time when the landmass was less isolated, should show a more balanced representation of dispersal modes. Contrary to this, fleshy-fruitedness is more common among species in Gondwanan taxa or in taxa with pollen records dating to before the Miocene. Main conclusions Fleshy-fruitedness in New Zealand conforms to neither the expectations for an isolated landmasses, namely a disharmonic range of dispersal modes, nor expectations for a continental island. I suggest that this pattern may be a product of selective survival of highly vagile taxa in the low-lying archipelago that was New Zealand during the late Cretaceous to mid-Cenozoic, followed by an invasion by taxa with a broader range of dispersal modes facilitated by the establishment of the circumpolar current.     

The reproductive biology of the New Zealand flora

New Zealand's long isolation from other elements of Gondwanaland, oceanic climate, the unusual combination of ancient Gondwanic, tropical and more recently arrived elements in its flora, and its relatively depauperate pollinator and disperser fauna have set the stage for the evolution of a subtle, complex and distinctive reproductive biology. This contrasts markedly with the neighbouring continent of Australia where the fauna is diverse and the flowers vivid. Recent advances in understanding New Zealand's floral biology include evidence that the ancestor to the anthophytes was cosexual, with insect pollinators receiving stigmatic nectar rewards; the discovery of ground-level bat pollination in an obligate root parasite; the finding that the greater resource sensitivity of fruit set in males than in females may account for sex ratio variation in gynodioecious species; and, evidence for much more pronounced mast seeding at higher altitudes even in the absence of mammalian seed predators.     

Phylogenetic relationships of epacrids and vaccinioids (Ericaceae s. l.) based onmatK sequence data

Cladistic relationships of epacrids and vaccinioids (Ericaceae) are investigated using nucleotide sequence data from the chloroplast encodedmatK gene. Sequences of 56 taxa were aligned and analyzed using parsimony methods. Results show thatVaccinioideae as currently recognized are not monophyletic. The epacrids are sister to a clade that includes theLyonia group, theGaultheria group, and theVaccinieae. Arbutus andPyrola branch early inEricaceae, before the rhododendroid group.Enkianthus is sister to the remainingEricaceae (includingEpacridaceae).Vaccinieae are strongly supported as monophyletic, butVaccinium andAgapetes are polyphyletic.     

Extinction and recolonization of coastal megafauna following human arrival in New Zealand

Extinctions can dramatically reshape biological communities. As a case in point, ancient mass extinction events apparently facilitated dramatic new evolutionary radiations of surviving lineages. However, scientists have yet to fully understand the consequences of more recent biological upheaval, such as the megafaunal extinctions that occurred globally over the past 50 kyr. New Zealand was the world''s last large landmass to be colonized by humans, and its exceptional archaeological record documents a vast number of vertebrate extinctions in the immediate aftermath of Polynesian arrival approximately AD 1280. This recently colonized archipelago thus presents an outstanding opportunity to test for rapid biological responses to extinction. Here, we use ancient DNA (aDNA) analysis to show that extinction of an endemic sea lion lineage (Phocarctos spp.) apparently facilitated a subsequent northward range expansion of a previously subantarctic-limited lineage. This finding parallels a similar extinction–replacement event in penguins (Megadyptes spp.). In both cases, an endemic mainland clade was completely eliminated soon after human arrival, and then replaced by a genetically divergent clade from the remote subantarctic region, all within the space of a few centuries. These data suggest that ecological and demographic processes can play a role in constraining lineage distributions, even for highly dispersive species, and highlight the potential for dynamic biological responses to extinction.     

Seed dispersal systems in the New Zealand flora

Knowledge of the dispersal mechanisms used by plants is important in phylogenetic, ecological, biogeographical, and conservation studies. Here we attempt to assign dispersal mechanisms to the entire flora-2595 plant species of the New Zealand Botanical Region. Anemochory is the most frequent dispersal mechanism, utilised by 79% of species. The next most frequent mechanisms are endozoochory (33%), hydrochory (28%), epizoochory (26%), and ballistic (8%). Polychory is common, particularly in monocotyledonous and dicotyledonous families and hydrochorous, epizoochorous, and ballistically dispersed species. Epizoochory is more common in New Zealand than in other regions, and species using this dispersal mechanism are over-represented among threatened species. Frugivory is less common than previously reported, and is under-represented among threatened species. Some mechanisms are poorly known, and entanglement and capsulivory are dispersal mechanisms apparently unique to New Zealand. Dispersal mechanisms reflective of New Zealand's distinctive assemblages of large flightless birds and reptiles are not apparent. A pattern of reduction in dispersal-related structures is evident in some genera. The mechanisms utilised by some species are ambiguous. Thus there remains a need for further investigation of the dispersal mechanisms utilised by plants in New Zealand.     

Fossil Ericaceae from New Zealand: Deconstructing the use of fossil evidence in historical biogeography

The Australasian Ericaceae epitomize many problems in understanding the biogeography of the southern hemisphere, especially the relative contributions of Gondwanan vicariance and dispersal. Late Cretaceous fossil pollen of the family suggests extreme antiquity of the group in Australasia, but recent phylogenetic evidence suggests much younger histories for most of the groups in that region. This paper documents two new species of latest Oligocene-Early Miocene macrofossils of Ericaceae from New Zealand. Cyathodophyllum novae-zelandiae G.J.Jord. & Bannister gen. and sp. nov. is the oldest record of the tribe Styphelieae, but is of a clade now extinct in New Zealand, possibly related to the Tasmanian genus Cyathodes. Richeaphyllum waimumuensis G.J.Jord. & Bannister sp. nov. is a member of Richeeae, but it is ambiguous as to whether it is a member of the impressive modern New Zealand radiation in Dracophyllum. These fossils emphasize the fact that at least some of the fossil pollen of Ericaceae may have been derived from extinct lineages and therefore should not be used as evidence for the antiquity of any modern New Zealand clade of Ericaceae. New fossils and/or detailed analysis of fossil and extant pollen may help resolve such uncertainty.     

Honeyeaters and the New Zealand forest flora: The utilisation and profitability of small flowers

New Zealand flowers are frequently considered unspecialised allowing easy access to pollen and nectar by a wide range of visitors. Most conform with a syndrome of insect pollination (entomophily). Pollination of forest flowers by birds has been described for a range of species whose flowers are morphologically ornithophilous. On Kapiti Island and Little Barrier Island, all three species of New Zealand honeyeaters have been described feeding on flowers currently assumed to be entomophilous or where the pollination system is unknown. The persistence and regularity of visits suggests that the birds are obtaining suitable rewards in the form of nectar and could be serving as pollinators. We measured the nectar energetic value from flowers of three ornithophilous and five entomophilous species. Nectar production over 24 hours was highest in ornithophilous species, but the standing crop of nectar overlapped—ornithophilous species: Metrosideros fulgens (standing crop 6.6 J), Metrosideros excelsa (22 J), and Fuchsia excorticata (1.8 J); and entomophilous: Pittosporum crassifolium (23 J), Pseudopanax arboreus (1.5 J), Dysoxylum spectabile (3.7 early flowers -6.7 J late flowers), Pittosporum eugenioides (2.7 J) and Geniostoma rupestre (1.8 J). The entomophilous species present the flowers in aggregation and as result birds can visit a large number flowers per minute. We found that the average estimated nectar consumption rate for all the entomophilous species except G. rupestre was enough to sustain the two smaller New Zealand honeyeaters (hihi energy requirements= 0.12 kJ min(-1), median energy obtained: 0.16 kJ min(-1) D. spectabile—0.57 kJ min(-1) P. crassifolium); bellbird energy requirements = 0.10 kJ min(-1), median energy obtained: 0.14 kJ min(-1) D. spectabile—0.68 kJ min(-1) P. crassifolium). However, we estimate that if the birds are able to selectively forage on the flowers with most nectar, the energetic returns of all species may be sufficient for hihi and bellbird (hihi: 0.18 kJ min(-1) G. rupestre—0.93 kJ min(-1); P. crassifolium; bellbird: 0.12 kJ min(-1) G. rupestre 1.11 kJ min(-1) P. crassifolium). If tui (energy requirements: 0.25 kJ min(-1), forages randomly, only P. crassifolium (0.80 kJ min(- 1)) and D. spectabile late in the season (0.30 kJ min(-1)) provide sufficient returns, but if selective, P. arboreus (0.45 kJ min(-1)) may also suffice. We suggest that because (a) the nectar produced by entomophilous flowers provides sufficient energy to sustain the energetic requirements of birds, and (b) these plants flower in the cooler months when insect activity is reduced, birds might have played a wider role in pollination than previously considered. This finding is of particular importance because the abundance of New Zealand honeyeaters on the mainland has decreased considerably since human colonisation and this could be affecting forest regeneration.     

Periodicity of extinction and recolonization of the West Indian topshell Cittarium pica in the Quaternary of Bermuda (Gastropoda: Trochoidea)

Hermit‐crab transported shells of the West Indian top shell Cittarium pica occur in numerous terrestrial fossil deposits on Bermuda, which is the most remote outpost of this Caribbean species. Cittarium is so far known only from deposits of interglacial ages corresponding to marine isotope stages (MIS) 11, 9, 5e, and 1 (Holocene). In at least the cases of MIS 11 and 5e, Cittarium appears at the very beginning of the interglacial. The species is definitely absent from well‐stratified cave deposits of the last glacial (MIS 4–2). It is hypothesized that colder sea‐surface temperatures (SSTs) during glacial periods caused Cittarium to be extirpated on Bermuda, and that it has made at least four independent colonizations of the island in the past 400 000 years. Because of the limited larval life of Cittarium, the window for colonization may be a very narrow period at the onset of interglacials when the SST has risen sufficiently to make Bermuda habitable but the velocity of the Gulf Stream is still sufficiently rapid to transport viable larvae from the Bahamas. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 235–243.     

中国贵州杜鹃属两新种

报道了中国贵州杜鹃属植物2新种--枇杷叶杜鹃(Rhododendron eriobotryoides Xiang Chen & Jia Y.Huang)和金波杜鹃(R.jinboense Xiang Chen & X.Chen).枇杷叶杜鹃与皱叶杜鹃(R.denudatum)相似,但本种叶背毛被较薄而稀疏,花丝基部密被微柔毛,花柱通体被腺体.金波杜鹃与短脉杜鹃(R.brevinerve)相似,但前者的叶长圆状椭圆形,叶片上卷呈U形,花序具较多花,花冠较长,花丝基部密被微柔毛,子房密被绒毛.目前,仅发现2新种各有1个含数株的种群,资源量十分稀少,依据IUCN红色名录标准均属于极危物种(CR),需要加以重点保护.     

Potential plant genetic resources of the New Zealand flora

New Zealand harbours a highly endemic flora of some 2000 species of vascular plants, but except for numerous ornamentals, only four indigenous plants are cultivated for the production of fibre (Phormium tenax, pharmaceutical properties (Solanum laciniatum andS. aviculare), and as a vegetable (Tetragonia tetragonioides). Another 15 native species are relatives of common commercial plants and are considered potential genetic resources. Of the six early Polynesian plant introductions to New Zealand, only the kumara (sweet potato) is still of some economic importance today.