Phylogeography and systematics of the Malagasy rock‐thrushes (Muscicapidae,Monticola)

Cruaud, A., Raherilalao, M. J., Pasquet, E. & Goodman, S. M. (2011) Phylogeography and systematics of the Malagasy rock‐thrushes (Muscicapidae, Monticola). —Zoologica Scripta, 40, 554–566. The patterns of genetic variation and the systematics of members of the widespread Old World genus Monticola (Family Muscicapidae) occurring on Madagascar remain unresolved. Herein, we address these questions by examining the phylogeography of Malagasy Monticola using two molecular markers (ND2 and ATP6, 1.5 kb) from 60 individuals sampled across their known range. To clarify the relationships within the clade groupings, we use a statistical haplotype network and an analysis of the genetic structure of the different populations sampled. A morphological study was conducted in parallel that used many of the same individuals employed in the molecular study to examine potential differences between the recovered clades. Based on molecular genetics and morphology, M. imerinus is distinct from the M. sharpei complex, which is composed of five phylogroups: Group A (Central Highlands, typical sharpei), Group B (Central West, Bemaraha), Group C (Northern Highlands), Group D (Montagne d’Ambre, erythronotus) and Group E (Southwestern, bensoni). While molecular data show high levels of geographical structure, these differences exhibit low levels of intergroup genetic divergence (0.01–0.07%). We suggest that two species of Monticola occur on Madagascar, imerinus and sharpei, and the forms referable to bensoni and erythronotus, as well as unnamed populations from the Central West (Bemahara), should be considered as part of M. sharpei and are populations that are probably isolated and undergoing incipient speciation.     

A reevaluation of the genus Malassezia by means of genome comparison

Results of a study of the genus Malassezia on the basis of genome characters confirm that two species should be maintained, M. furfur and M. pachydermatis. The two forms associated with skin disease, frequently referred to as Pityrosporum orbiculare and P. ovale, were found to be synonymous, the name M. furfur having priority. Malassezia pachydermatis, hitherto regarded as a strictly zoophilic species, may also be found on humans.     

PHYLOGENY AND SYSTEMATICS OF THE GENUS MONOMORPHINA (EUGLENACEAE) BASED ON MORPHOLOGICAL AND MOLECULAR DATA1

Morphological studies of 16 strains belonging to the genus Monomorphina revealed a single, parietal, orbicular chloroplast in their cells. The chloroplast has a tendency to be perforated and disintegrates in aging populations and thus may appear to be many chloroplasts under the light microscope. A single chloroplast in the cells of Cryptoglena skujae is also parietally located and highly perforated. It never forms a globular and closed structure, but is open from the side of the furrow, resembling the letter C. We have verified the Monomorphina pyrum group (M. pyrum–like) on the basis of phylogenetic analysis of SSU rDNA and morphological data. The strain CCAC 0093 (misidentified as M. reeuwykiana) diverges first on the SSU rDNA phylogenetic tree. The rest of the M. pyrum–like strains form a tight cluster, subdivided into several smaller ones. Because morphological differences between the M. pyrum–like strains (including the strain CCAC 0093) do not conform to the tree topology, we suggest that they all (except the strain CCAC 0093) belong to M. pyrum. We designate a new species, M. pseudopyrum, for the strain CCAC 0093, solely on the basis of molecular characters. We also suggest that M. reeuwykiana and similar species should stay in Phacus and Lepocinclis unless detailed molecular and morphological studies show otherwise. Emended diagnoses of the genera Monomorphina and Cryptoglena and the species M. aenigmatica are also proposed, as well as the delimitation of an epitype for M. pyrum, the type species for the genus Monomorphina.     

New insights into the systematics of Malagasy mongoose‐like carnivorans (Carnivora,Eupleridae, Galidiinae) based on mitochondrial and nuclear DNA sequences

The Malagasy carnivorans (Eupleridae) comprise seven genera and up to ten species, depending on the authority, and, within the past decades, two new taxa have been described. The family is divided into two subfamilies, the Galidiinae, mongoose‐like animals, and the Euplerinae, with diverse body forms. To verify the taxonomic status of Galidiinae species, including recently described taxa, as well as some recognized subspecies, we studied intrageneric genetic variation and structure, using both mitochondrial and nuclear markers. Our results suggest the recognition of four species in the Galidiinae, rendering each genus monospecific. We propose to recognize three subspecies in Galidia elegans (G. e. dambrensis, G. e. elegans, and G. e. occidentalis), two subspecies in Mungotictis decemlineata (M. d. decemlineata and M. d. lineata), and two subspecies in Galidictis fasciata (G. f. fasciata and G. f. grandidieri, the latter was recently described as a distinct species). Our results indicate also that Salanoia durrelli should be treated as a junior synonym of Salanoia concolor. Low levels of intraspecific divergence revealed some geographical structure for the Galidiinae taxa, suggesting that environmental barriers have isolated certain populations in recent geological time. All taxa, whether at the species or subspecies level, need urgent conservation attention, particularly those with limited geographical distributions, as all are threatened by forest habitat degradation.     

Molecular phylogeny,biogeography and chromosome evolution of Malagasy dwarf geckos of the genus Lygodactylus (Squamata,Gekkonidae)

We performed a phylogenetic analysis using nuclear (RAG‐1, RAG‐2) and mitochondrial (16S) markers, a statistical Bayesian reconstruction of ancestral distribution areas and a karyological analysis on most Malagasy species of the gekkonid genus Lygodactylus. The phylogenetic analysis largely confirms major basal branching pattern of previous molecular studies, but highlights significant differences concerning both the relationships between different species groups as well as those within groups. The biogeographic analysis supports a Malagasy origin of Lygodactylus, an oversea dispersal to continental Africa and a return to Madagascar. The L. madagascariensis group (also including a new candidate species identified herein) is the most basal clade in Lygodactylus, and the sister group of a clade with all the remaining species. The second most basal clade is the L. verticillatus group, placed as the sister group of a clade comprising African and Malagasy species. The sister lineage of the L. verticillatus group originated the African radiation through an oversea dispersal out of Madagascar. Eventually, the sister lineage of the L. capensis group originated secondary dispersals from Africa to Madagascar. In Madagascar, lineage diversification in different species groups mainly occurred from southern to northern and eastern regions. Dispersal, vicariance and paleoclimatic refugia probably played a relevant role in the evolutionary history of closely related taxa and in speciation mechanisms. The cytogenetic analysis evidenced a high karyotypic variability in Lygodactylus (from 2n = 34 to 2n = 40), which is at least partly consistent with the phylogenetic relationships and the composition of the various species group. Chromosome evolution occurred independently in different lineages, mainly through a reduction in the chromosome number and starting from a putative primitive karyotype of 2n = 40 with all telocentric elements.     

Mastigopterus imperator Smith and Radcliffe, 1913, a senior synonym of M. praetor Smith and Radcliffe, 1913 (Ophidiidae, Ophidiiformes)

 A rare deep-sea ophidiid genus Mastigopterus known to contain two species, M. imperator Smith and Radcliffe, 1913 and M. praetor Smith and Radcliffe, 1913, was reviewed on the basis of six specimens including the holotypes of both species. Dorsal and anal fin ray counts and the size of cephalic sensory pores previously thought to be diagnostic characteristics to discriminate the two species did not suggest the presence of two forms, the large (M. imperator) and the small species (M. praetor), in the genus. Apparently the genus is represented by a single species, M. imperator, known from off Madagascar, the East and South China Seas and Papua New Guinea. Received: September 7, 1999 / Revised: July 9, 2001 / Accepted: November 13, 2001     

Molecular phylogeny of Macrolycus (Coleoptera: Lycidae) with description of new species from China

Macrolycus is a genus of net‐winged beetles with 69 species distributed in the eastern Palearctic and northernmost part of the Oriental region. The first molecular phylogeny of Macrolycus was produced using an rrnL + tRNA‐Leu + nad1 mtDNA fragment. The major lineages and species limits were identified with morphology and molecular data. We propose that Cerceros is a subgenus of Macrolycus to enable identification of all adult specimens in the genus without DNA sequencing. Two species groups are proposed in Macrolycus s. str. and six in Cerceros. Additionally, twelve Macrolycus species are newly described from China: M. aquilinus, M. baihualingensis, M. bicolor, M. guangxiensis, M. jianfenglingensis, M. kuatunensis, M. lizipingensis, M. parvus, M. phoeniceus, M. rhodoneurus, M. rosaceus and M. sichuanensis. Macrolycus holzschuhi is proposed to be a junior subjective synonym of M. jeanvoinei. The highest diversity of Macrolycus is found in southern China. The species from the main islands of Japan are placed in two species groups: M. excellens is a sister to remaining species of the M. murzini group and the M. flabellatus group is a monophylum of closely related species in a sister position to the M. bicolor group.     

Evolution of the Canthonini Longitarsi (Scarabaeidae) in Madagascar

Dung beetle species belonging to the worldwide tribe Canthonini (Scarabaeidae) and occurring in Madagascar are all endemic to that island. The Malagasy Canthonini form three lineages, one of which is the group Longitarsi that includes five genera. The phylogenetic relationships of Malagasy Canthonini are not fully resolved and only few species of Longitarsi have been included in previous studies. Here we infer the phylogenetic relationships within the Longitarsi group using molecular data and together with morphological examination revise the systematics of the group. The five genera of the Longitarsi group form one monophyletic clade and thus we suggest the synonymization of the younger genera Sikorantus, Phacosomoides, Madaphacosoma and Aleiantus; with the oldest genus belonging to this clade Epactoides. We describe two new species: Epactoides jounii sp. n and Epactoides mangabeensis sp. n. Most of the species of Longitarsi inhabit the eastern rainforests, with very low local species diversity and highly restricted geographical ranges. In the group Longitarsi four species are wingless. The loss of wings has evolved at least twice, at high altitude along the mountain range.     

Molecular phylogeny supports the morphologically based taxonomic transfer of the “medicagoid”Trigonella species to the genus Medicago L.

 The taxonomic transfer of the 23 Trigonella species previously known as medicagoids to the genus Medicago L. is reanalyzed on the basis of a molecular phylogenetic approach. The internal and external transcribed spacers of 53 Medicago species s. str. and 10 of the 23 medicagoid species were sequenced. Both maximum parsimony or maximum likelihood criteria joined medicagoid species more confidently with Medicago rather than with Trigonella. The basal position of the medicagoid species within the genus Medicago, together with morphologically atypical Medicago species (such as M. radiata and M. platycarpa), raised the question of the expediency to define a new genus and is discussed, relatively to the support of each clade. Using criteria of monophyly and support, it was concluded that the medicagoids are better joined in Medicago rather than placed in a new genus. Received February 6, 2001 Accepted July 17, 2001     

THE EVOLUTION AND SYSTEMATICS OF BEE-EATERS (MEROPIDAE)

Behavioural and ecological characters are used in addition to structural ones in considering the systematics of the Meropidae. The two species of Nyctyornis are the most primitive extant forms. Meropogon is retained as a monotypic genus for forsteni, and all other bee-eaters are placed in the single genus Merops, being ecologically and morphologically rather uniform except in details of wing and tail structure, which should be considered only of specific importance. M. breweri and M. oreobates are thought to be secondarily rather than primarily forest species. With the submergence of Aerops and Melittophagus, and the monotypic genera Bombylonax and Dicrocercus, which are considered to be closely related to the pusillus species-group, Merops is enlarged to 21 species which are uniform except in tail-shape, wing formula and throat feather structure. These characters are of specific importance only, and have a mosaic distribution within the genus. Their use in the definition of the formerly recognized genera results in an artificial classification. The proposed delimitation of species-groups within Merops differs somewhat from previous arrangements, and affinities argued in the text are summarized in Fig. 15, which shows superspecies and species-groups. The species recognized are formally tabulated below. The Meropidae probably originated in southeast Asian forest and spread through former forest to Africa. Only in Africa was the open-country environment invaded, and speciation in the savanna was in two main directions, producing small sedentary and large migratory species. Representatives of both types returned to Asia in open country. From the present distribution of species it is inferred that speciation has proceeded under the main influences of (1) isolation of a population and its habitat in Pleistocene Africa and (2) isolation of migrants away from their breeding range. The distributions of species with wide or rather limited ranges are discussed in terms of physiological adaptation and ecological competition. Opinion has not been expressed on the validity of subspecies, which have been discussed only in delimiting controversial species. In the following summary, the only subspecies named are those affected by changes from Peters' scheme (cf. Table 1). Superspecies are bracketed. Nyctyornis amicta N. athertoni Meropogon forsteni Merops guloris M. mülleri M. bulocki M. bullockoides M. pusillus M. variegatus (?loringi, oariegatus, lafresnayii, bangweoloensis) M. oreobates M. hirundineus M. breweri M. revoilii M. albicollis M. orientalis M. boehmi M. viridis M. superciliosus (persicus, chrysocercus, superciliosus) M. philippinus (philippinus, salvadorii) M. ornatus M. apiaster M. leschenaulti M. malimbicus M. nubicus (nubicus, nubicoides)     

Cynanchum and the Cynanchinae (Apocynaceae – Asclepiadoideae): a molecular, anatomical and latex triterpenoid study

The phylogeny of the genus Cynanchum s. str. is studied using cpDNA spacers and ITS. Morphological, anatomical and latex triterpenoid data are interpreted in light of the molecular results, and discrepancies are discussed. Vegetative characters are better indicators of relationship than floral characters, especially corona characters. The monophyly of all Malagasy species and, nested within the latter, of all stem-succulent taxa is ascertained and the genera Folotsia, Karimbolea, Platykeleba and Sarcostemma are subsumed under Cynanchum. One African species, C. galgalense, is excluded from Cynanchum.     

Biosystematics of the Muellerianella complex (Homopters,Delphacidae), hybridization studies

Crosses were made between three closely related species of the planthopper genus Muellerianella, M. brevipennis, M. extrusa and M. fairmairei. Most hybrids are produced in crosses between M. extrusa and M. fairmairei. Most of the hybrid females in these crosses and some of the hybrid males are fertile and can be successfully backcrossed with the parental species. Crosses of M. brevipennis with either M. fairmairei or M. extrusa yield less progeny, of which all males are sterile. Backcrosses of the hybrid females with the parental species are partly successful. The possible evolutionary consequences of mating readiness and interfertility between the three species under field conditions are discussed. Efforts to resynthesize triploid gynogenetic forms by backcrossing hybrid females of M. fairmairei and M. brevipennis with males of M. fairmairei, as was reported by Drosopoulos (Evolution 32: 916–920, 1978), failed. To produce offspring, the existing gynogenetic forms have to mate with males of one of the three bisexual species.     

Reproduction in Coquerel's dwarf lemur (Mirza coquereli)

Most nocturnal Malagasy primates, as well as many diurnal species, are highly endangered in their natural habitat. Captive breeding programs have been established for many species, but detailed information on reproduction is only available for three nocturnal taxa: the mouse lemur (Microcebus murinus), the dwarf lemur (Cheirogaleus medius), and the greater dwarf lemur (Cheirogaleus major). In this study, we present data for another nocturnal Malagasy primate, Coquerel's dwarf lemur (Mirza coquereli), which has been propagated since 1982 at the Duke University Primate Center. Unlike all other Malagasy primates bred in captivity, M. coquereli cycles throughout the year, and is clearly less seasonal in its birth distribution than is C. medius or M. murinus. Estrous intervals ranged between 19 and 30.5 days. Estrus lasted no longer than 1 day. After an average gestation length of 89.2 days, litters of one or two were born. Females cycled for the first time between 8 and 15 months of age, and gave birth for the first time between 12.8 and 33.5 months of age. The earliest mating of a male leading to conception was observed at the age of 17 months. For males and females, an increase in daylength appears to trigger pubertal development. Males had seasonal changes in testicular volume which were not explained by covariation with body weight. Maximum testis size occurred in spring, when breeding activity was highest. The occurrence of year-round reproduction in M. coquereli, and the absence of seasonal fattening and/or hibernation, along with their specialized winter diet, may be correlated. © 1995 Wiley-Liss, Inc.     

Phylogeny and taxonomy of rhizobia

Rhizobia are bacteria that form nitrogen-fixing nodules on the roots, or occasionally the shoots, of legumes. There are currently more than a dozen validly named species, but the true number of species is probably orders of magnitude higher. The named species are listed and briefly discussed. Sequences of the small subunit ribosomal RNA (SSU or 16S rRNA) support the well-established subdivision of rhizobia into three genera: Rhizobium, Bradyrhizobium, and Azorhizobium. These all lie within the alpha subdivision of the Proteobacteria, but on quite distinct branches, each of which also includes many bacterial species that are not rhizobia. It has been clear for several years that Rhizobium, on this definition, is still too broad and is polyphyletic: there are many non-rhizobia within this radiation. Recently, therefore, it has been suggested that this genus should be split into four genera, namely Rhizobium (R. leguminosarum, R. tropici, R. etli), Sinorhizobium (S. fredii, S. meliloti, S. teranga, S. saheli), Mesorhizobium (M. loti, M. huakuii, M ciceri, M. tianshanense, M. mediterraneum), and a fourth, unnamed, genus for the current R. galegae. The evidence and pros and cons are reviewed.     

Description of <Emphasis Type="Italic">Gosztonyia antarctica</Emphasis>, a new genus and species of Zoarcidae (Teleostei: Perciformes) from the Antarctic Ocean

A new genus and species of zoarcid fish, Gosztonyia antarctica, is described on the basis of four specimens collected from the Bellingshausen Sea, Antarctic Ocean, at a depth of 615 m. Gosztonyia is placed in the subfamily Lycodinae and can be distinguished from all other zoarcid genera by the following combination of characters: seven branchiostegal rays, interdigitating ceratohyal–epihyal articulation, palatal arch reduced, posterior hyomandibular ramus longer than anterior, cranium narrowed, supratemporal commisure and occipital pores absent. A new species, Gosztonyia antarctica, is described and the relationships of the new genus are discussed.     

Effects of forest fragmentation on genetic variation in endemic understory forest birds in central Madagascar

Summary This study focuses on some genetic consequences of habitat fragmentation in populations of four endemic bird species (Monticola sharpei, Terpsiphone mutata, Foudia omissa, andFoudia madagascariensis) living in the understory of forests in the Réserve Spéciale d'Ambohitantely on the Central High Plateau of Madagascar. The four species differ in their dependency on forest habitats, which may be related to their migration abilities between isolated forest fragments. Genetic variation was analysed on the basis of multi-locus fingerprints in order to study the influence of migration and habitat size on the genetic variability of local populations. There was no evidence that forest fragmentation affects any of the four species with respect to genetic variability.

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Effekte von Waldfragmentation auf die genetische Variabilität endemischer Vogelarten in Zentral-Madagaskar

Zusammenfassung Rund 53 % der Vogelarten Madagaskars sind endemisch. Viele dieser Arten sind an Waldhabitate gebunden und durch die Verinselung der madagassischen Wälder hochgradig bedroht. In dem vorliegenden Projekt wird die Auswirkung der Verinselung auf die genetische Variabilität und die genetische Differenzierung von vier endemischen Vogelarten am Beispiel des Reservats von Ambohitantely im zentralen Hochland Madagaskars untersucht. Ambohitantely beinhaltet mehr als 500 Waldfragmente von 0,64 ha bis 1250 ha. Untersucht wurden: 1. die WaldartFoudia omissa, 2.Monticola (früherPseudocossyphus)sharpei, eine Art mit starker Bevorzugung für primäre Waldlebensräume, die aber gelegentlich auch in Sekundärvegetation auftritt, sowie 3.Terpsiphone mutata und 4.Foudia madagascariensis, beides Arten, die sowohl in Wäldern als auch in offenen Landschaften vorkommen. Die unterschiedliche Abhängigkeit dieser Arten von Waldhabitaten und der damit verbundenen Möglichkeit, offene Landschaften zwischen den Waldfragmenten zu überbrücken, führte zur Frage, ob die Verinselung die genetische Vielfalt der untersuchten Populationen beeinflußt. Für die genetischen Analysen wurden Blutproben von Tieren aus drei Gebieten mit 1250 ha, 136 ha und 28 ha gesammelt. Die Proben wurden mit Multilocus-Fingerprints genetisch charakterisiert. Zum Untersuchungszeitpunkt konnte kein Einfluß der Fragmentierung auf die genetische Populationsstruktur der vier untersuchten Arten nachgewiesen werden.

     

A revision of the genus Monstrillopsis Sars (Crustacea: Copepoda: Monstrilloida) with description of a new species from Chile

In the course of a comparative analysis of the species included in the copepod genus Monstrillopsis Sars, a new species of the genus was discovered. The new species, Monstrillopsis chilensis n. sp., was found in a plankton sample collected in coastal waters off central Chile in the Southeastern Pacific. The adult female is similar to females of the type species M. dubia Scott and related forms but differs from its congeners by a combination of characters including: (1) a pair of small cephalic processes present between bases of antennules; (2) deep corrugations present on proximo-lateral surface of fifth pedigerous somite; (3) exopodal lobe of fifth leg not distally elongated; and (4) endopodal lobe of fifth legs short. This is the first record of the genus in the Southeastern Pacific and the second record of any monstrilloid in Chile. A comparative analysis of the species assigned to Monstrillopsis allowed a clarification of the generic characters and limits. The most important apomorphic characters in the genus are the presence of four caudal setae and modifications of the male antennule; other potentially useful characters are the pattern of antennule segmentation and the uniform presence of a reduced inner lobe of the fifth leg. The female of Monstrilla reticulata Davis should be retained in Monstrilla, but the purportedly conspecific male is probably a Monstrillopsis. Monstrillopsis ciqroi Suárez-Morales and M. angustipes Isaac should be transferred to Monstrilla. Haemocera filogranarum Malaquin is probably a species of Monstrillopsis. Monstrillopsis zernowi is tentatively retained in this genus as an aberrant form. A key for the identification of the species of this genus is included.     

Molecular discrimination of six species of Bagrid catfishes from Indus river system using randomly amplified polymorphic DNA markers

Bagrid catfishes constitute a very important group of fishes having immense commercial importance in south-east countries. The phylogenetic relationships and genome specificity among six species of Bagrid catfishes (Mystus bleekeri, M. cavasius, M. vittatus, M. tengara, M. aor and M. seenghala) were investigated using RAPD markers as discriminating characters for the first time. 511 RAPD fragments were generated using ten decamer primers of arbitrary nucleotide sequences. Amplification reactions resulted in fragments ranging in length between 92 and 2,863 bp, which were assigned to 155 RAPD loci. Clearly resolved and repeatable bands were scored for their presence or absence in a binary matrix. Different RAPD profiles were observed for all the six Mystus species. In the present study three group diagnostic, eleven group exclusive and 18 species-specific markers were generated. Thus six Mystus species can be successfully differentiated on the basis of these 18 species-specific RAPD markers. UPGMA dendrogram constructed on the basis of genetic distance formed two distinct clusters, M. seenghala and M. aor form one separate cluster from other four species i.e., M. tengara, M. cavasius, M. bleekeri and M. vittatus. The inferences drawn from the above study clearly showed their genetic distinctness from the other four Mystus species and supported their inclusion into a separate genus, Sperata.     

Molecular evidence for the polyphyly of Olearia (Astereae: Asteraceae)

 Analyses of ITS sequences for 49 species of Olearia, including representatives from all currently recognised intergeneric sections, and 43 species from 23 other genera of Astereae, rooted on eight sequences from Anthemideae, provide no support for the monophyly of this large and morphologically diverse Australasian genus. Eighteen separate lineages of Olearia are recognised, including seven robust groups. Three of these groups and another eight species are placed within a primary clade incorporating representatives of Achnophora, Aster, Brachyscome, Calotis, Camptacra, Erigeron, Felicia, Grangea, Kippistia, Lagenifera, Minuria, Oritrophium, Peripleura, Podocoma, Remya, Solidago, Tetramolopium and Vittadinia. The remaining four groups and three individual species lie within a sister clade that also includes Celmisia, Chiliotrichum, Damnamenia, Pleurophyllum and Pachystegia. Relationships within each primary clade are poorly resolved. There is some congruence between this molecular estimate of the phylogeny and the distribution of types of abaxial leaf-hair, which is the basis of the present sectional classification of Olearia, but all states appear to have arisen more than once within the tribe. It is concluded that those species placed within the second primary clade should be removed from the genus, but the extent to which species placed within the first primary clade constitute a monophyletic group can only be resolved with further sequence data. Received November 12, 2001; accepted April 29, 2002 Published online: November 22, 2002 Addresses of authors: Edward W. Cross, Centre for Plant Biodiversity Research, CSIRO, GPO Box 1600, Canberra, ACT 2601, Australia (E-mail: ed.cross@csiro.au); Christopher J . Quinn, Royal Botanic Gardens, Mrs Macquaries Rd., Sydney, NSW 2000, Australia; Steven J. Wagstaff, Landcare Research, PO Box 69, Lincoln 8152, New Zealand.