Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae

Nepenthaceae, the pitcher plants of the Old World tropics show a remarkable diversity in SE Asia, especially on the islands of Borneo and Sumatra. This region is considered as a secondary center of diversity. Sequence analysis of the cpDNA TRNK intron supports this hypothesis showing the species of the Malay Archipelago as neighbour group to the isolated species from Sri Lanka, the Seychelles, and Madagascar. Based on phylogenetic reconstructions an origin of recent Nepenthaceae in the Indian subcontinent is assumed. A recent investigation focused on a non-plastid, translocated copy of the TRNK intron has revealed an incongruence to tree topology based on the cpDNA TRNK intron. Although the translocated copy emerged as insufficient for phylogenetic reconstruction of Nepenthaceae some taxa showed, contrary to the cpDNA dataset, relatively high distances to the rest of the taxa. These results indicated that the phylogeny of the TRNK intron could not reflect true phylogenetic relationships. We investigated the peptide transferase 1 (PTR1), to develop a phylogenetic marker that is based on a nuclear low copy gene in Nepenthes. All sequences obtained were probably functional, indicated by the ratio of point mutations of the single codon positions in exon and intron regions. Comparative analysis showed that this locus is of similar variability as the cpDNA TRNK intron and, contrary to the translocated copy of TRNK, potential useful for phylogenetic reconstruction. While in parts congruent to the plastid TRNK intron phylogeny, a higher divergence of some sequences in PRT1 and in the previously reported, non cpDNA dataset indicates that remnants of an older species stock persisted east of Wallace's line and on the Sunda Shelf. This suggests that plastid haplotypes existing today in the main distribution center of the Nepenthaceae could be descendants of more recently dispersed lineages that had been transmitted to an old species stock.     

First record of Coleura (Chiroptera: Emballonuridae) on Madagascar and identification and diagnosis of members of the genus

Abstract

The genus Coleura contains two species: C. seychellensis restricted to the Seychelles Archipelago and C. afra broadly distributed on the African continent and the Arabian Peninsula. We recently captured and collected specimens of Coleura from Madagascar, an island from which it was previously unknown. Using museum specimens and the new material from Madagascar, we review morphological variation in members of this genus and present characters for species identification. On the basis of these comparisons, specimens from Madagascar are referable to C. afra. We present information on the natural history of the recently discovered Malagasy population.     

A phylogenetic analysis of the palm subtribe Oncospermatinae (Arecaceae) based on morphological characters

Subtribe Oncospermatinae (Arecaceae: Arecoideae: Areceae) is a diverse group of spiny Old World palms. The subtribe includesOncosperma, a widespread Asian genus of five species, along with seven monotypic genera, all endemic to the Seychelles and Mascarene Islands of the western Indian Ocean. A phylogenetic analysis was conducted in order to test the monophyly of subtribe Oncospermatinae with respect to other Old World genera of tribe Areceae. A matrix of 38 morphological characters was scored for 29 taxa, including 11 species of the Oncospermatinae. A single most parsimonious tree was found, resolving the subtribe as a polyphyletic group of two distinct clades. One clade containingAcanthophoenix, Deckenia, Oncosperma, andTectiphiala was placed as sister to a large group that includes members of subtribes Archontophoenicinae, Arecinae, Iguanurinae, and Ptychospermatinae. The other clade of Oncospermatinae, including the Seychelles endemic generaNephrosperma, Phoenicophorium, Roscheria, andVerschaffeltia, was resolved as sister to the Madagascar endemic subtribe Masoalinae, and may have arisen in the western Indian Ocean region.     

Diversity of Siphonaria Sowerby I, 1823 (Gastropoda,Siphonariidae) in the Seychelles Bank and beyond

Molecular phylogenetic studies have shown that the characters of the reduced shell of the false limpets of the genus Siphonaria Sowerby I, 1823 are highly variable and often insufficient for species delimitation. The taxonomy and distribution of Siphonaria in the Indian Ocean are poorly known. We sampled Siphonaria in the Seychelles Bank to check the occurrence of recorded species using DNA sequences and to study the paths through which Siphonaria species have colonised the Seychelles Bank by reconstructing their phylogenetic relationships. Analyses of a dataset comprising 16 S rRNA gene sequences of 33 specimens from the Seychelles Bank and 300 additional Siphonaria sequences from other regions from GenBank with various methods for species delimitation resulted in 19–102 primary species hypotheses. Assemble Species by Automatic Partitioning provided a conservative estimate of the species number (42) in which several indisputable species were lumped. The results of Automatic Barcode Gap Discovery depended strongly on the assumed prior maximum intraspecific divergence, whereas the tree-based methods Generalised Mixed Yule Coalescent and Poisson Tree Processes resulted in high overestimates. The specimens from the Seychelles Bank represent three clades, belonging to the Siphonaria ‘atra’ group, the Siphonaria ‘normalis’ group and a possibly undescribed species recorded previously only from Hainan. At least two of the three species recorded from the Seychelles Bank came from the east, i.e., from the Coral Triangle in the Indo-Australian Archipelago, the region with the highest marine biodiversity worldwide. A major transport mechanism across the Indian Ocean was probably the South Equatorial Current.     

Tracking island colonization history and phenotypic shifts in Indian Ocean bulbuls (Hypsipetes: Pycnonotidae)

Molecular phylogenies of island organisms provide useful systems for testing hypotheses of convergent or parallel evolution, since selectively neutral molecular characters are likely to be independent of phenotype, and the existence of similar environments on multiple isolated islands provides numerous opportunities for populations to evolve independently under the same constraints. Here we construct a phylogenetic hypothesis for Hypsipetes bulbuls of the western Indian Ocean, and use this to test hypotheses of colonization pattern and phenotypic change among islands of the region. Mitochondrial sequence data were collected from all extant taxa of the region, combined with sequence data from relevant lineages in Asia. Data are consistent with a single Hypsipetes colonization of the western Indian Ocean from Asia within the last 2.6 Myr. The expansion of Hypsipetes appears to have occurred rapidly, with descendants found across the breadth of its western Indian Ocean range. The data suggest that a more recent expansion of Hypsipetes madagascariensis from Madagascar led to the colonization of Aldabra and a secondary colonization of the Comoros. Groupings of western Indian Ocean Hypsipetes according to phenotypic similarities do not correspond to mtDNA lineages, suggesting that these similarities have evolved by convergence or parallelism. The direction of phenotypic change cannot be inferred with confidence, since the primary expansion occurred rapidly relative to the rate of mtDNA substitution, and the colonization sequence remains uncertain. However, evidence from biogeography and comparison of independent colonization events are consistent with the persistence of a small grey continental bulbul in India and Madagascar, and multiple independent origins of large size and green plumage in insular island populations of the Comoros, Mascarenes and Seychelles. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 271–287.     

Phylogenetics and biogeography of Mascarene angraecoid orchids (Vandeae, Orchidaceae)

The large angraecoid orchid clade (subtribe Angraecinae sensu lato) has undergone extensive radiation in the western Indian Ocean, which includes Africa, Madagascar, and a number of Indian Ocean islands, such as the Mascarene Archipelago. To investigate systematics and biogeography of these Mascarene orchids, phylogenetic relationships were inferred from four plastid DNA regions, trnL intron, trnL-F intergenic spacer, matK gene, and rps16 intron. Parsimony and Bayesian analyses provided identical sets of relationships within the subtribe; the large genus Angraecum as currently circumscribed does not form an exclusive clade. Bonniera, an endemic genus to Reunion, is shown to be embedded in part of Angraecum. Evidence from our research supports the main origin of Mascarene Angraecinae from Madagascar, and although there were many independent colonizations, only a few of the lineages radiated within the Mascarene Archipelago.     

Eastward from Africa: palaeocurrent-mediated chameleon dispersal to the Seychelles islands

Madagascar and the Seychelles are Gondwanan remnants currently isolated in the Indian Ocean. In the Late Cretaceous, these islands were joined with India to form the Indigascar landmass, which itself then split into its three component parts around the start of the Tertiary. This history is reflected in the biota of the Seychelles, which appears to contain examples of both vicariance- and dispersal-mediated divergence from Malagasy or Indian sister taxa. One lineage for which this has been assumed but never thoroughly tested is the Seychellean tiger chameleon, a species assigned to the otherwise Madagascar-endemic genus Calumma. We present a multi-locus phylogenetic study of chameleons, and find that the Seychellean species is actually the sister taxon of a southern African clade and requires accomodation in its own genus as Archaius tigris. Divergence dating and biogeographic analyses indicate an origin by transoceanic dispersal from Africa to the Seychelles in the Eocene–Oligocene, providing, to our knowledge, the first such well-documented example and supporting novel palaeocurrent reconstructions.     

Phylogeography of Ptychadena mascareniensis suggests transoceanic dispersal in a widespread African-Malagasy frog lineage

Aim The Mascarene ridged frog, Ptychadena mascareniensis, is the only African amphibian species thought to occur on Madagascar and on the Seychelles and also Mascarene islands. We explored its phylogenetic relationships and intraspecific genetic differentiation to contribute to the understanding of transoceanic dispersal in amphibians. Methods Fragments of the mitochondrial 16S rRNA gene were sequenced from specimens collected over most of the distribution area of P. mascareniensis, including populations from Madagascar, Mascarenes and Seychelles. Results We identified five deeply divergent clades having pairwise divergences >5%, which probably all represent cryptic species in a P. mascareniensis complex. One of these seems to be restricted to Madagascar, the Mascarenes and the Seychelles. Sequences obtained from topotypic material (Réunion) were identical to the most widespread haplotype from Madagascar. The single Mauritian/Seychellean haplotype differed by only one mutation from a Malagasy haplotype. Main conclusions It is likely that the Mascarene and Seychellean populations were introduced from Madagascar by humans. In contrast, the absence of the Malagasy haplotypes from Africa and the distinct divergences among Malagasy populations (16 mutations in one divergent hapolotype from northern Madagascar) suggest that Madagascar was populated by Ptychadena before the arrival of humans c. 2000 years ago. Because Madagascar has been separated from Africa since the Jurassic, this colonization must have taken place by overseas rafting, which may be a more widespread dispersal mode in amphibians than commonly thought.     

Are the native giant tortoises from the Seychelles really extinct? A genetic perspective based on mtDNA and microsatellite data

The extinction of the giant tortoises of the Seychelles Archipelago has long been suspected but is not beyond doubt. A recent morphological study of the giant tortoises of the western Indian Ocean concluded that specimens of two native Seychelles species survive in captivity today alongside giant tortoises of Aldabra, which are numerous in zoos as well as in the wild. This claim has been controversial because some of the morphological characters used to identify these species, several measures of carapace morphology, are reputed to be quite sensitive to captive conditions. Nonetheless, the potential survival of giant tortoise species previously thought extinct presents an exciting scenario for conservation. We used mitochondrial DNA sequences and nuclear microsatellites to examine the validity of the rediscovered species of Seychelles giant tortoises. Our results indicate that the morphotypes suspected to represent Seychelles species do not show levels of variation and genetic structuring consistent with long periods of reproductive isolation. We found no variation in the mitochondrial control region among 55 individuals examined and no genetic structuring in eight microsatellite loci, pointing to the survival of just a single lineage of Indian Ocean tortoises.     

The biogeographic and tectonic history of India

Aim To present an up to date account of the Mesozoic history of India and its relationship to the other Gondwana continents and to Eurasia. Location Continents surrounding the Western Indian Ocean. Methods Utilization of recent evidence of continental relationships based upon research in stratigraphy, palaeomagnetism, palaeontology, and contemporary biotas. Results The physical data revealed a sequence of events as India moved northward: (1) India–Madagascar rifted from east Africa 158–160 Ma (million years ago), (2) India–Madagascar from Antarctica c. 130 Ma, (3) India–Seychelles from Madagascar 84–96 Ma, (4) India from Seychelles 65 Ma, (5) India began collision with Eurasia 55–65 Ma and (6) final suturing took place c. 42–55 Ma. However, data from fossil and contemporary faunas indicate that, throughout the late Cretaceous, India maintained exchanges with adjacent lands. There is an absence in the fossil record of peculiar animals and plants that should have evolved, had India undergone an extended period of isolation just before its contact with Eurasia. Main conclusions The depiction of India in late Cretaceous as an isolated continent is in error. Most global palaeomaps, including the most recent one, show India, as it moves northward, following a track far out in the Indian Ocean. But the evidence now indicates that India's journey into northern latitudes cannot have taken place under such isolated circumstances. Although real breaks among the lands were indicated by the physical data, faunal links were maintained by vagile animals that were able to surmount minor marine barriers. India, during its northward journey, remained close to Africa and Madagascar even as it began to contact Eurasia.     

India–Madagascar vicariance explains cascade beetle biogeography

India and Madagascar drifted apart more than 80 Mya, yet few taxonomic groups currently found in these regions bear any signature of this split. When drifting in isolation, extensive volcanic activity covered almost half of India in lava flows, likely triggering widespread extinction on the island. Consequently, most of India's rich extant flora and fauna are considered to be the result of recent Cenozoic dispersal, and no lineages are conclusively a result of ancient vicariance. Many of Madagascar's lineages also stem from either Cenozoic diversification or dispersal events, with the latter being a result of the close proximity of the island with mainland Africa. In the present study, we focus on two remarkable lineages of cascade beetles in the genera Scoliopsis and Tritonus (Coleoptera, Hydrophilidae), respectively, dwelling in the mountains of south India and Sri Lanka, as well as in Madagascar. Based on a molecular phylogeny of the family Hydrophilidae dated with eight fossils, we show that these two lineages are sister taxa, and diverged when Madagascar and Greater India (India, Sri Lanka, Seychelles) separated, suggesting a pattern of Gondwanean vicariance. The results of the present study show that, despite geological upheaval, the present‐day fauna of India still retains traces of its Gondwanan past.     

Phylogeography of the Indo‐West Pacific maskrays (Dasyatidae,Neotrygon): a complex example of chondrichthyan radiation in the Cenozoic

Maskrays of the genus Neotrygon (Dasyatidae) have dispersed widely in the Indo‐West Pacific being represented largely by an assemblage of narrow‐ranging coastal endemics. Phylogenetic reconstruction methods reproduced nearly identical and statistically robust topologies supporting the monophyly of the genus Neotrygon within the family Dasyatidae, the genus Taeniura being consistently basal to Neotrygon, and Dasyatis being polyphyletic to the genera Taeniurops and Pteroplatytrygon. The Neotrygon kuhlii complex, once considered to be an assemblage of color variants of the same biological species, is the most derived and widely dispersed subgroup of the genus. Mitochondrial (COI, 16S) and nuclear (RAG1) phylogenies used in synergy with molecular dating identified paleoclimatic fluctuations responsible for periods of vicariance and dispersal promoting population fragmentation and speciation in Neotrygon. Signatures of population differentiation exist in N. ningalooensis and N. annotata, yet a large‐scale geological event, such as the collision between the Australian and Eurasian Plates, coupled with subsequent sea‐level falls, appears to have separated a once homogeneous population of the ancestral form of N. kuhlii into southern Indian Ocean and northern Pacific taxa some 4–16 million years ago. Repeated climatic oscillations, and the subsequent establishment of land and shallow sea connections within and between Australia and parts of the Indo‐Malay Archipelago, have both promoted speciation and established zones of secondary contact within the Indian and Pacific Ocean basins.     

Diversity of the slippery zone microstructure in pitchers of nine carnivorous Nepenthes taxa

Using a cryo-scanning electron microscope, we studied microstructure of the slippery zone in nine Nepenthes taxa. For N. fusca, N. macrophylla, N. mirabilis, N. ventricosa, N. dicksoniana, and N. veitchii, it was examined here for the first time. Three types of the slippery zone were distinguished among the studied taxa: (1) with well-developed crystalline wax coverage, (2) with greatly reduced wax coverage, and (3) without wax crystals. These data were combined with morphometrical measurements of the two pitcher zones primarily relevant to prey catching and retaining: the slippery zone and the peristome. In species with fully developed wax coverage, the slippery zone was longer and the peristome was narrower compared to those with reduced or lacking crystalline wax. We found statistically significant negative correlation between the relative length of the slippery zone and the relative width of the peristome. Based on the analysis of the relationship between the microstructure of the slippery zone and pitcher macromorphology, two main types of pitchers in Nepenthes are proposed: (a) traps based predominantly on the waxy slippery zone and (b) peristome-based traps.     

Integrative taxonomy and phylogeny of leafless Vanilla orchids from the South-West Indian Ocean region reveal two new Malagasy species

The leafless Vanilla species complex from the South-West Indian Ocean (SWIO) region has long been a taxonomic challenge, due to limited patterns of morphological differentiation and an absence of variation within chloroplast sequences. This complex includes seven known morphospecies: V. madagascariensis, V. bosseri, V. decaryana, and V. perrieri endemic to Madagascar, V. humblotii presumed as endemic to the Comoros Archipelago, but also present in Madagascar, V. roscheri from the East African coast, and V. phalaenopsis endemic to Seychelles. A previous population genetic study using microsatellite markers allowed us to distinguish, in addition to the five recognized Malagasy taxa, two other genetic clusters present in the East of the island. An integrative taxonomy approach was therefore conducted by combining microsatellite and morphological data used in the previous study with new data sets, and by adding ITS sequencing data, to validate the taxonomic level of these Malagasy genetic clusters and unravel phylogenetic relationships between SWIO species. As a result, based on phylogenetic, genotypic and morphological evidence, nine species were discriminated in the SWIO region, including seven in Madagascar, with two new eastern species. The leafless Vanilla group originated and diversified in Madagascar, from an ancestor of African descent, with three subsequent independent colonization events from Madagascar to the other territories of SWIO within the two main lineages (white versus yellow flower species). The new Malagasy species, V. allorgeae Andriamihaja & Pailler sp. nov., and V. atsinananensis Andriamihaja & Pailler sp. nov., are described and a new identification key is proposed.     

Pollination by fungus gnats (Diptera: Mycetophilidae) and self-recognition sites in Tolmiea menziesii (Saxifragaceae)

Coleeae (Bignoniaceae) are a tribe almost entirely restricted to Madagascar. Coleeae have previously been placed in neotropical Crescentieae due to species with indehiscent fruits, a character otherwise unusual in Bignoniaceae. A phylogeny based on three chloroplast regions (ndhF, trnT-L spacer, trnL-F spacer) identifies a monophyletic Coleeae that is endemic to Madagascar and surrounding islands of the Indian Ocean (Seychelles, Comores and Mascarenes). African Kigelia is not a member of Coleeae, rather it is more closely related to a subset of African and Southeast Asian species of Tecomeae. The molecular phylogeny indicates that indehiscent fruit have arisen repeatedly in Bignoniaceae: in Coleeae, Kigelia and Crescentieae. The characteristic fleshy fruits of species of Coleeae likely arose autochthonously in Madagascar. Within Coleeae Colea and Ophiocolea are sisters, Phyllarthron is sister to Colea + Ophiocolea, and Rhodocolea is sister to the rest of the tribe.     

A spectacular new species of Nepenthes L. (Nepenthaceae) pitcher plant from central Palawan,Philippines

A new species of Nepenthes L., N. attenboroughii (Nepenthaceae), from Palawan Island in the Philippines, is described and illustrated. It is restricted to rocky, ultramafic soils that comprise the summit region of Mount Victoria, Municipality of Narra, where it occurs in isolation from other members of the genus. On the basis of the morphological features, this new taxon appears to be related to both N. mira Jebb & Cheek of Palawan and N. rajah Hook.f. of Borneo. Its substantial size places it among the largest of known pitcher plants. The diagnostic morphological characters are discussed and an updated key is provided for a revised complex of Nepenthes species from the Palawan and North Borneo phytogeographical region. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 195–202.     

Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean

Rowson, B., Tattersfield, P. & Symondson, W. O. C. (2010). Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean. —Zoologica Scripta, 40, 85–98. A phylogeny is presented for the speciose, near pan‐tropical, carnivorous achatinoid land‐snail superfamily Streptaxoidea inferred from DNA sequences (two nuclear and two mitochondrial regions) from 114 taxa from Africa, the Indian Ocean, Asia, South America and Europe. In all analyses, Streptaxidae are monophyletic, while the (two to six) previously recognised subfamilies are polyphyletic, as are several genus‐level taxa including the most speciose genus Gulella, necessitating major taxonomic review. The Asian Diapheridae are sister to Streptaxidae, which forms several well‐supported clades originating in a persistent basal polytomy. Divergence dating estimates, historical biogeography, and the fossil context suggest a Cretaceous origin of these families, but suggest Gondwanan vicariance predated most radiation. The basal polytomy dates to the Paleogene and may correspond to a rapid radiation in Africa. There is evidence for multiple Cenozoic dispersals followed by radiation, including at least two from Africa to South America, at least two from Africa to Asia and at least two from Africa to Madagascar, indicating Cenozoic turnover in tropical snail faunas. The endemic Seychelles and Mascarene streptaxid faunas each are composites of early Cenozoic lineages and more recent dispersals from Africa, with no direct evidence for an Asian origin as currently proposed. Peak streptaxid diversity in East Africa is explained by Neogene speciation among a large number of coexisting ancient lineages, a phenomenon most pronounced in the Eastern Arc‐Coastal Forests centre of endemism. This includes Miocene diversification in Gulella, a primarily East and South‐East African group which remains strikingly diverse even after unrelated ‘Gulella’ species are reclassified.     

Phylogeography of the introduced species Rattus rattus in the western Indian Ocean,with special emphasis on the colonization history of Madagascar

Aim To describe the phylogeographic patterns of the black rat, Rattus rattus, from islands in the western Indian Ocean where the species has been introduced (Madagascar and the neighbouring islands of Réunion, Mayotte and Grande Comore), in comparison with the postulated source area (India). Location Western Indian Ocean: India, Arabian Peninsula, East Africa and the islands of Madagascar, Réunion, Grande Comore and Mayotte. Methods Mitochondrial DNA (cytochrome b, tRNA and D‐loop, 1762 bp) was sequenced for 71 individuals from 11 countries in the western Indian Ocean. A partial D‐loop (419 bp) was also sequenced for eight populations from Madagascar (97 individuals), which were analysed in addition to six previously published populations from southern Madagascar. Results Haplotypes from India and the Arabian Peninsula occupied a basal position in the phylogenetic tree, whereas those from islands were distributed in different monophyletic clusters: Madagascar grouped with Mayotte, while Réunion and Grand Comore were present in two other separate groups. The only exception was one individual from Madagascar (out of 190) carrying a haplotype that clustered with those from Réunion and South Africa. ‘Isolation with migration’ simulations favoured a model with no recurrent migration between Oman and Madagascar. Mismatch distribution analyses dated the expansion of Malagasy populations on a time‐scale compatible with human colonization history. Higher haplotype diversity and older expansion times were found on the east coast of Madagascar compared with the central highlands. Main conclusions Phylogeographic patterns supported the hypothesis of human‐mediated colonization of R. rattus from source populations in either the native area (India) or anciently colonized regions (the Arabian Peninsula) to islands of the western Indian Ocean. Despite their proximity, each island has a distinct colonization history. Independent colonization events may have occurred simultaneously in Madagascar and Grande Comore, whereas Mayotte would have been colonized from Madagascar. Réunion was colonized independently, presumably from Europe. Malagasy populations may have originated from a single successful colonization event, followed by rapid expansion, first in coastal zones and then in the central highlands. The congruence of the observed phylogeographic pattern with human colonization events and pathways supports the potential relevance of the black rat in tracing human history.     

First record of the fledging period of Temminck's Courser,Cursorius temminckii

The Seychelles Fody, Foudia sechellarum, is a ploceid weaver occurring naturally on three islands in the Seychelles group in the Indian Ocean. The population on Cousine Island was studied between 30 June and 25 August 1997. The size of the population on the island was estimated at 458–614 individuals and densities varied in different habitat types. As Seychelles Fodies in non-breeding plumage are difficult to sex, we provide sexing criteria based on wing length. Breeding pairs form small, probably temporary, territories that are defended by both partners against other fodies, including the introduced Madagascar Fody, Foudia madagascariensis. The Seychelles Fody often breeds semi-colonially and we observed up to five nests close together. The birds are socially monogamous and both sexes share in nest building, nest defense and provisioning of the young. Only females incubate. Many nests were deserted before eggs were laid, including some that had been accepted by the female. Additional males and females were sometimes seen helping provision the young, but this cooperative breeding behaviour appeared to be uncommon. Non-breeding individuals congregate in large flocks, sometimes joined by breeding birds.