Systematic revision of the genus Everettia Godwin‐Austen, 1891 (Mollusca: Gastropoda: Dyakiidae) in Sabah,northern Borneo

The species of the snail genus Everettia in the Malaysian state of Sabah are superficially similar and difficult to distinguish by their shells. This paper presents new data on the taxonomy and distribution of Everettia in Sabah that have accumulated since the revision by Godwin‐Austen in 1891. By using morphological and molecular phylogenetic approaches, we reveal at least seventeen species of Everettia in Sabah, of which eleven are new to science, namely: Everettia layanglayang sp. nov. , Everettia lapidini sp. nov. , Everettia paulbasintali sp. nov. , Everettia occidentalis sp. nov. , Everettia jasilini sp. nov. , Everettia safriei sp. nov. , Everettia interior sp. nov. , Everettia jucundior sp. nov. , Everettia planispira sp. nov. , Everettia monticola sp. nov. , and Everettia dominiki sp. nov. , and one new subspecies, namely, Everettia corrugata williamsi ssp. nov. Phylogenetic analysis of mitochondrial COI and 16S, and nuclear ITS‐1 sequences demonstrates the monophyly of most of the morphologically well‐defined species. Our results show that certain aspects of classical morphology‐based taxonomy for Everettia species, especially with regard to the unique combination of shell surface sculptures, animal head colour, and mantle pigmentation, are solid. A dichotomous key to the Sabah species and subspecies of Everettia is provided. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 515–550.     

Photosynthesis–nitrogen relationships in species at different altitudes on Mount Kinabalu, Malaysia

In situ photosynthetic nitrogen-use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) was investigated in species that commonly distributed at different altitudes (600–3700m above sea level) on Mount Kinabalu. Photosynthetic nitrogen-use efficiency was lower in species at higher altitudes, with a mean PNUE at 3700m being one-third as large as that at 600m. This difference in PNUE was larger than that explained by the biochemical response to lower air pressures only. Across altitudes a negative correlation between ¹³C abundance (¹³C) and PNUE was found. Species at higher altitudes tended to have higher ¹³C, suggesting that they had a lower conductance for CO₂ diffusion from the air to chloroplasts. The lower conductance might be responsible for the lower PNUE in species at higher altitudes. Although leaf nitrogen content per unit area tended to be higher at higher altitudes, it did not seem to contribute to increasing photosynthetic rates. Thus, the idea that a higher nitrogen content at higher altitudes is a compensation for a lower PNUE was not supported. In contrast to the large difference in PNUE among altitudes, PNUE tended to converge within a narrow range among species growing at the same altitude.     

Genus Colocasiomyia (Drosophilidae: Diptera) in Sabah,Bornean Malaysia: High species diversity and use of host aroid inflorescences

A total of 21 Colocasiomyia species, including 17 undescribed species, are reported from Sabah (Mt. Kinabalu and neighboring areas), Malaysia, based on samples collected from inflorescences of 14 or 15 Araceae species. This number of species is the largest as a local fauna of this genus in the world. The high species diversity is attributed to 12 undescribed species belonging to the Colocasiomyia baechlii species group. A particular breeding habit of Colocasiomyia is sharing of the same inflorescence by a pair of species, with partial niche separation between them: one species uses exclusively the pistillate (lower female‐flower) section of the spadix for oviposition and larval development, whereas the other mostly uses the staminate (upper male‐flower) section. However, the baechlii group species show quite different patterns of host plant use: many (up to eight) species cohabit on the same inflorescence. It is unlikely that they separate their breeding niches micro‐allopatrically within an inflorescence. Instead, species composition and their proportions of individual numbers vary among different localities, seasons and host plants, with partial overlap among them. Such partial separations in local distribution, phenology and host selection would in combination contribute to their coexistence and promote the species diversity of this group.     

Canopy structure of tropical and sub-tropical rain forests in relation to conifer dominance analysed with a portable LIDAR system

Background and Aims

Globally, conifer dominance is restricted to nutient-poor habitats in colder, drier or waterlogged environments, probably due to competition with angiosperms. Analysis of canopy structure is important for understanding the mechanism of plant coexistence in relation to competition for light. Most conifers are shade intolerant, and often have narrow, deep, conical crowns. In this study it is predicted that conifer-admixed forests have less distinct upper canopies and more undulating canopy surfaces than angiosperm-dominated forests.

Methods

By using a ground-based, portable light detection and ranging (LIDAR) system, canopy structure was quantified for old-growth evergreen rainforests with varying dominance of conifers along altitudinal gradients (200–3100 m a.s.l.) on tropical and sub-tropical mountains (Mount Kinabalu, Malaysian Borneo and Yakushima Island, Japan) that have different conifer floras.

Key Results

Conifers dominated at higher elevations on both mountains (Podocarpaceae and Araucariaceae on Kinabalu and Cupressaceae and Pinaceae on Yakushima), but conifer dominance also varied with soil/substrate conditions on Kinabalu. Conifer dominance was associated with the existence of large-diameter conifers. Forests with higher conifer dominance showed a canopy height profile (CHP) more skewed towards the understorey on both Kinabalu and Yakushima. In contrast, angiosperm-dominated forests had a CHP skewed towards upper canopy, except for lowland dipterocarp forests and a sub-alpine scrub dominated by small-leaved Leptospermum recurvum (Myrtaceae) on Kinabalu. Forests with a less dense upper canopy had more undulating outer canopy surfaces. Mixed conifer–angiosperm forests on Yakushima and dipterocarp forests on Kinabalu showed similar canopy structures.

Conclusions

The results generally supported the prediction, suggesting that lower growth of angiosperm trees (except L. recurvum on Kinabalu) in cold and nutrient-poor environments results in a sparser upper canopy, which allows shade-intolerant conifers to co-occur with angiosperm trees either as emergents or as codominants in the open canopy.     

Three new species of Mitrephora (Annonaceae) from Sabah, Malaysia

Three new species of Mitrephora (Blume) Hook. f. & Thomson, M. clemensiorum, M. vittata and M. woodii , are described from Sabah, Malaysia. Mitrephora clemensiorum is related to M. korthalsiana Miq., but is distinguished by having long inflorescences, long flowering and fruiting pedicels, and fruits with sessile monocarps. Mitrephora vittata is related to M. reflexa Merr., but has lanceolate, subcoriaceous leaves, and densely pubescent inflorescence rachides, flowering pedicels and bracts. Mitrephora woodii is similar to M. heyneana (Hook. f. & Thomson) Thwaites, but has smaller flowers and fruits with sessile monocarps     

Depreissia decipiens,an enigmatic canopy spider from Borneo revisited (Araneae,Salticidae), with remarks on the distribution and diversity of canopy spiders in Sabah,Borneo

Depreissia is a little known genus comprising two hymenopteran-mimicking species, one found in Central Africa and one in the north of Borneo. The male of Depreissia decipiens is redescribed, the female is described for the first time. The carapace is elongated, dorsally flattened and rhombus-shaped, the rear of the thorax laterally depressed and transformed, with a pair of deep pits; the pedicel is almost as long as the abdomen. The male palp is unusual, characterized by the transverse deeply split membranous tegulum separating a ventral part which bears a sclerotized tegular apophysis and a large dagger-like retrodirected median apophysis. The female epigyne consists of one pair of large adjacent spermathecae and very long copulatory ducts arising posteriorly and rising laterally alongside the spermathecae continuing in several vertical and horizontal coils over the anterior surface. Relationships within the Salticidae are discussed and an affinity with the Cocalodinae is suggested. Arguments are provided for a hypothesis that Depreissia decipiens is not ant-mimicking as was previously believed, but is a mimic of polistinine wasps. The species was found in the canopy in the Kinabalu area only, in primary and old secondary rainforest at 200–700 m.a.s.l. Overlap of canopy-dwelling spider species with those in the understorey are discussed and examples of species richness and endemism in the canopy are highlighted. Canopy fogging is a very efficient method of collecting for most arthropods. The canopy fauna adds an extra dimension to the known biodiversity of the tropical rainforest. In southeast Asia, canopy research has been neglected, inhibiting evaluation of comparative results of this canopy project with that from other regions. More use of fogging as a collecting method would greatly improve insight into the actual species richness and species distribution in general.     

Variations in the reproductive cycle of Bornean montane tree species along elevational gradients on ultrabasic and non-ultrabasic soils

Although lowland tree species in the ever-wet regions of Southeast Asia are characterized by the supra-annual cycle of reproduction, the reproductive phenology of montane tree species remains poorly understood. In this study, we investigated the reproductive phenology of montane tree species using litter samples that were collected every 2 weeks from six rainforest sites, consisting of three elevations (1700, 2700, and 3100 m), on Mount Kinabalu, Borneo. At each elevation, one site was on infertile ultrabasic soil and one was on relatively fertile non-ultrabasic soil. We used a composite sample from 10 or 20 litter traps per site and sorted it by species. Therefore, the obtained data captured reproductive phenology in the population of each species rather than in an individual tree. Ten-year time series of flower and fruit litterfall were obtained for 30 and 39 tree species, respectively. Fourier analysis was used to identify the dominant cycle of each time series. The most abundant cycle across species was supra-annual, followed by sub-annual, and annual cycles. Many species at higher elevations showed supra-annual cycles of flower litterfall, whereas species in the 1700 m sites often showed annual or sub-annual cycles regardless of soil types. No systematic differences were found among sites for fruit litterfall. Mechanisms underlying these elevational patterns in reproductive cycle remain unclear but may include more severe El Niño droughts, lower primary productivity, lower soil fertility, and the absence of some sub-annually or annually reproducing families at higher elevations.     

Pollination mutualism between Alocasia macrorrhizos (Araceae) and two taxonomically undescribed Colocasiomyia species (Diptera: Drosophilidae) in Sabah, Borneo

Two taxonomically undescribed Colocasiomyia species were discovered from inflorescences of Alocasia macrorrhizos in Kota Kinabalu City, Sabah, Borneo, Malaysia. The aims of this study were to investigate the reproductive ecology of the flies and the plant, ascertain the importance of the flies as pollinators and examine the intimate association between flowering events and life history of the flies. We conducted sampling, observations and field pollination experiments. The flies were attracted by the odour of female‐phase inflorescences in the early morning on the first day of anthesis. They fed, mated and oviposited in the inflorescences for 1 day. On the second day, the flies, covered with pollen grains, left the male‐phase inflorescences for the next female‐phase inflorescences. The immature forms of both fly species hatched, developed and pupated within the infructescences without damaging the fruits, and developed adults emerged when the mature infructescences dehisced. The flowering events and fly behaviours were well synchronized. In field pollination experiments, inflorescences bagged with a fine mesh (insect exclusion) produced almost no fruits, whereas those bagged with a coarse mesh (bee exclusion) produced as many fruits as the open‐pollinated controls. These results indicate that these flies are the most efficient and specialised pollinators for their host, A. macrorrhizos. These flies, in return, depend on A. macrorrhizos for food and habitat through most of their life cycle. This study provides a deeper insight into the less recognised, highly intimate pollination mutualism between Araceae plants and Colocasiomyia flies.