Evolutionary size changes in plants of the south‐west Pacific

Aim To investigate evolutionary changes in the size of leaves, stems and seeds of plants inhabiting isolated islands surrounding New Zealand. Location Antipodes, Auckland, Campbell, Chatham, Kermadec, Three Kings and Poor Knights Islands. Methods First, we compared the size of leaves and stems produced by 14 pairs of plant taxa between offshore islands and the New Zealand mainland, which were grown in a common garden to control for environmental effects. Similar comparisons of seed sizes were made between eight additional pairs of taxa. Second, we used herbarium specimens from 13 species pairs to investigate scaling relationships between leaf and stem sizes in an attempt to pinpoint which trait might be under selection. Third, we used herbarium specimens from 20 species to test whether changes in leaf size vary among islands located at different latitudes. Lastly, we compiled published records of plant heights to test whether insular species in the genus Hebe differed in size from their respective subgenera on the mainland. Results Although some evidence of dwarfism was observed, most insular taxa were larger than their mainland relatives. Leaf sizes scaled positively with stem diameters, with island taxa consistently producing larger leaves for any given stem size than mainland species. Leaf sizes also increased similarly among islands located at different latitudes. Size changes in insular Hebe species were unrelated to the average size of the respective subgenera on the mainland. Main conclusions Consistent evidence of gigantism was observed, suggesting that plants do not obey the island rule. Because our analyses were restricted to woody plants, results are also inconsistent with the ‘weeds‐to‐trees’ hypothesis. Disproportionate increases in leaf size relative to other plant traits suggest that selection may favour the evolution of larger leaves on islands, perhaps due to release from predation or increased intra‐specific competition.     

Convergent evolution of gigantism in the flora of an isolated archipelago

Animals often evolve conspicuous differences in body size after colonising isolated islands. However, far less is known about repeated patterns in the evolution of plant size on islands. We tested for convergent evolution of leaf area, seed size and plant stature in the flora of the Chatham Islands (i.e. Rēkohu), which are located 700 km off the east coast of New Zealand. First, we measured leaf area, seed size and plant stature from populations of 22 plant taxa on the Chatham Islands. These data were then compared to analogous measurements from their sister taxa on the mainland to test for evidence of island gigantism, and whether size changes differ between leaves, seeds and stature. Second, we collated data from the literature to test whether size changes in endemic taxa differed among plant growth forms or were correlated with divergence times. Results showed that all three plant traits tended to increase in size on the Chatham Islands. However, field data showed that size increases tended to be more consistent in seeds and leaves than in stature. Data from the literature indicated that size increases also differed among growth forms. Herbaceous species showed the strongest evidence of gigantism, followed by woody plants, while graminoids showed a weak trend towards dwarfism. Insular size increases in seeds and leaves were also positively related to divergence times, indicating that taxa which have resided on the Chatham Islands for longer periods show stronger evidence of gigantism than taxa which arrived more recently. Overall results illustrate that gigantism is a hallmark of the Chatham Island flora, providing a remarkable example of convergent evolution.     

Sexual size dimorphism in island plants: the niche variation hypothesis and insular size changes

The niche variation hypothesis predicts insular populations exhibit increased sexual size dimorphism (SSD), to minimize intraspecific competition. Although many animal taxa conform to this prediction, insular patterns of SSD have yet to be investigated in plants. Here, we tested for differences in SSD of dioecious plants that colonised four island groups (Kermadec, Three Kings, Chatham and Auckland Islands) from New Zealand. Using herbarium collections, we quantified leaf and stem sizes of 263 individuals from 28 dioecious taxa. We developed a novel analytical technique to explore changes in the direction of SSD on islands. Lastly, we tested for evolutionary size changes of male and female plants on islands. The degree of SSD did not vary predictably between insular and mainland taxa, contrary to predictions of the niche variation hypothesis. Furthermore, the direction of SSD was not predictable on islands, while it was consistently female biased on the mainland. Our results suggest that selection favours increased size of both sexes on islands and that SSD is unpredictable for insular plants.     

Determining host plant preferences for the critically endangered Lord Howe Island stick insect (<Emphasis Type="Italic">Dryococelus australis</Emphasis>) to assist reintroduction

The Lord Howe Island stick insect (Dryococelus australis) is one of the world’s rarest insects. However, the opportunity to reintroduce the species to Lord Howe Island, and commence the path to recovery, may occur within the next 5 years. Understanding the insect’s host plant and habitat preferences on Lord Howe Island is critical to maximising the likelihood of reintroduction success. However, very little ecological information was documented before the species became extinct on the island in the 1930s. Here we examine the Lord Howe Island stick insect’s preference for potential host plants, a key aspect of habitat suitability. We conducted preference trials using 15 common plant species found on Lord Howe Island. Both nymphs and adults consumed some but not all of these plant species. Nymphs were able to survive on 7 of these 15 plants for the duration of the 26-day trials although failed to survive on some of the plants most preferred by adults. Overall, these data reveal that there are numerous plants on Lord Howe Island that the stick insect can consume, though their suitability varies with different developmental stages of the insect. These data are encouraging for any future reintroduction attempts and would greatly aid the selection and monitoring of release sites.     

Herbivory and the evolution of divaricate plants: Structural defences lost on an offshore island

Many island plants are characterized by unique morphology. For example, the high branching angles and small leaves of divaricate plants are a common feature of the New Zealand flora. The divaricate growth form may be an adaptation to deter browsing by extinct avian herbivores (moa); alternatively aspects of the insular climate may be responsible. However, our understanding of the selective pressures responsible for the high branching angles and small leaves of divaricate plants is incomplete. Here, I tested for differences in traits associated with the divaricate growth form between plants from Chatham Island and the New Zealand mainland. Moa never reached the Chatham Islands and its flora is derived from plants on mainland New Zealand. Therefore, I predicted Chatham Island plants to have lost morphological adaptations that may have deterred moa herbivory. Traits were quantified on 316 individuals in the field, allowing for 12 island‐mainland taxonomic comparisons. Chatham Island plants consistently produced smaller branching angles, larger leaves, shorter internodes and larger stems than related mainland plants. Results are therefore consistent with the hypothesis that selection for small leaves and high angled branching may be relaxed on the Chatham Islands due to an absence of moa. Smaller branching angles and larger leaves may offer a competitive advantage to Chatham Island plants.     

A comparative analysis of island floras challenges taxonomy‐based biogeographical models of speciation

Speciation on islands, and particularly the divergence of species in situ, has long been debated. Here, we present one of the first, complete assessments of the geographic modes of speciation for the flora of a small oceanic island. Cocos Island (Costa Rica) is pristine; it is located 550 km off the Pacific coast of Central America. It harbors 189 native plant species, 33 of which are endemic. Using phylogenetic data from insular and mainland congeneric species, we show that all of the endemic species are derived from independent colonization events rather than in situ speciation. This is in sharp contrast to the results of a study carried out in a comparable system, Lord Howe Island (Australia), where as much as 8.2% of the plant species were the product of sympatric speciation. Differences in physiography and age between the islands may be responsible for the contrasting patterns of speciation observed. Importantly, comparing phylogenetic assessments of the modes of speciation with taxonomy‐based measures shows that widely used island biogeography approaches overestimate rates of in situ speciation.     

Size evolution in island lizards

Aim  The island rule, small animal gigantism and large animal dwarfism on islands, is a topic of much recent debate. While size evolution of insular lizards has been widely studied, whether or not they follow the island rule has never been investigated. I examined whether lizards show patterns consistent with the island rule.
Location  Islands worldwide.
Methods  I used literature data on the sizes of island–mainland population pairs in 59 species of lizards, spanning the entire size range of the group, and tested whether small insular lizards are larger than their mainland conspecifics and large insular lizards are smaller. I examined the influence of island area, island isolation, and dietary preferences on lizard size evolution.
Results  Using mean snout–vent length as an index of body size, I found that small lizards on islands become smaller than their mainland conspecifics, while large ones become larger still, opposite to predictions of the island rule. This was especially strong in carnivorous lizards; omnivorous and herbivorous species showed a pattern consistent with the island rule but this result was not statistically significant. No trends consistent with the island rule were found when maximum snout–vent length was used. Island area had, at best, a weak effect on body size. Using maximum snout–vent length as an index of body size resulted in most lizard populations appearing to be dwarfed on islands, but no such pattern was revealed when mean snout–vent length was used as a size index.
Main conclusions  I suggest that lizard body size is mostly influenced by resource availability, with large size allowing some lizard populations to exploit resources that are unavailable on the mainland. Lizards do not follow the island rule. Maximum snout–vent length may be biased by sampling effort, which should be taken into account when one uses this size index.     

Potential for higher invasiveness of the alien Oxalis pes-caprae on islands than on the mainland

The higher vulnerability of islands to invasions compared to mainland areas has been partially attributed to a simplification of island communities, with lower levels of natural enemies and competitors on islands conferring vacant niches for invaders to establish and proliferate. However, differences in invader life-history traits between populations have received less attention. We conducted a broad geographical analysis (i.e. 1050 km wide transect) of plant traits comparing insular and mainland populations to test the hypothesis that alien plants from insular populations have the potential for higher invasiveness than their alien mainland counterparts. For this purpose plants of the annual geophyte Oxalis pes-caprae were grown from bulbs collected in the Balearic islands and the Spanish mainland under common greenhouse conditions. There were no significant differences in bulb emergence and plant survival between descendants from insular and mainland populations. However, Oxalis descendants from insular populations produced 20% more bulbs without reducing allocation to bulb size, above-ground biomass or flowering than descendants from mainland populations. Based on the lack of sexual reproduction in Oxalis and the dependence of invasion on bulb production, our study suggests that the higher occurrence of Oxalis in the Balearic islands than in the Spanish mainland can partially be explained by genetically based higher propagation potential of insular populations compared to mainland populations.     

生境片段化对千岛湖马尾松林内土壤种子库的影响

土壤种子库是森林群落更新的主要来源之一,对森林的演替和恢复等具有重要意义。生境片段化现象正日益严重地影响着森林群落,并可影响森林土壤种子库。研究了千岛湖地区的大陆及岛屿次生马尾松林内土壤种子库的组成及其影响因素(e.g.,岛屿面积,形状指数,隔离度和距岛屿边缘距离等)。根据大陆和岛屿的面积及边缘梯度,采用大数量小样方法,分别在土壤种子库最大化(初冬,2015年12月)和最小化(晚春,2016年4月)时期对马尾松林内土壤进行了机械取样。对土壤样品进行萌发实验,检测了两个时期的土壤种子库上层(0—2 cm)和下层(2—5 cm)种子组成,并通过广义线性混合效应模型等手段分析其影响因素。结果显示:(1)所有316个土壤样本中,萌发出幼苗1422株,隶属于29科、40属、41种。其中,木本植物幼苗占13种1024株,草本占28种398株。(2)Jaccard指数和相关性分析均显示初冬、晚春时期的土壤种子库组成具有很高的相似性;土壤种子库上、下层组成的相似性也很高。(3)广义线性混合效应模型分析显示,在大陆和岛屿上,土壤种子库下层种子含量低于上层;而大陆样地土壤种子库中的木本植物种子数较岛屿样地高。岛屿上,土壤种子库中的种子数随土层的加深而降低;随边缘梯度升高也下降,尤其是草本植物的种子。对于岛屿上的木本植物,不耐阴种的种子数量远大于耐阴种,尤其是土壤下层。表明千岛湖地区马尾松林内土壤种子库组成受到生境片段化的影响,进而可能作用于该类型森林群落的演替。     

Recruitment patterns of scleractinian corals in an isolated sub-tropical reef system

The density of recruits of scleractinian corals on settlement plates at Lord Howe Island, a small isolated sub-tropical island 630 km off the Australian coastline, was within the range of values reported for comparable studies on the Great Barrier Reef. However, there was a difference in the relative abundance of taxonomic groups, with recruitment at Lord Howe Island during the summer of 1990/91 dominated by corals from the Family Pocilloporidae, Family Poritidae, and sub-genus Acropora (Isopora) (in order of abundance). By contrast, on the Great Barrier Reef, recruits are generally predominantly species from the Family Acroporidae (other than the Acropora (Isopora) group). Both the recruits and the established coral communities at Lord Howe Island are dominanted by corals which release brooded planulae, as opposed to the pattern of mass-spawning with external fertilisation more typical of Great Barrier Reef corals. I hypothesise that the release of brooded planulae would be advantageous in an isolated reef community because (a) brooded larvae can travel large distances and survive the journey to the isolated reef and/or (b) brooded larvae have a shorter period before they are competent to settle and are therefore more likely to be retained on the parental reef once a population has been established.     

Insular shifts and trade-offs in life-history traits in pond frogs in the Zhoushan Archipelago, China

Island and mainland populations of animal species often differ strikingly in life-history traits such as clutch size, egg size, total reproductive effort and body size. However, despite widespread recognition of insular shifts in these life-history traits in birds, mammals and reptiles, there have been no reports of such life-history shifts in amphibians. Furthermore, most studies have focused on one specific life-history trait without explicit consideration of coordinated evolution among these intimately linked life-history traits, and thus the relationships among these traits are poorly studied. Here we provide the first evidence of insular shifts and trade-offs in a coordinated suite of life-history traits for an amphibian species, the pond frog Rana nigromaculata . Life-history data were collected from eight islands in the Zhoushan Archipelago and neighboring mainland China. We found consistent, significant shifts in all life-history traits between mainland and island populations. Island populations had smaller clutch sizes, larger egg sizes, larger female body size and invested less in total reproductive effort than mainland populations. Significant negative relationships were found between egg size and clutch size and between egg size and total reproductive effort among frog populations after controlling for the effects of body size. Therefore, decreased reproductive effort and clutch size, larger egg size and body size in pond frogs on islands were selected through trade-offs as an overall life-history strategy. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in life-history traits across a range of terrestrial vertebrate taxa.     

Island biogeography and the reproductive ecology of great tits Parus major

Island biogeography theory has contributed greatly to both theoretical and applied studies of conservation biology (e.g., design of nature reserves, minimum viable population sizes, extinction risk) and community composition. However, little theoretical and empirical work has addressed how island isolation and size affect reproductive ecology. We investigated the reproductive ecology of great tits (Parus major) on one offshore and one nearshore island, as well as on the Danish mainland. Tits breeding on the offshore island bred later, laid smaller clutches, and laid larger eggs than those on the nearshore island and mainland. In addition, the level of ectoparasite infestation in nests was highest on the offshore island, intermediate on the nearshore island, and lowest on the mainland. These insular effects may occur due to lower food abundance on islands, to density-dependent effects, or to effects related to low genetic diversity within island populations. Whatever the cause, the results emphasize that future studies of forest fragmentation/population isolation should consider not only gross measures of reproductive success, but also fine-scale measures such as clutch size, timing of breeding, and parasite prevalence. Received: 10 November 1997 / Accepted: 9 March 1998     

Resource utilization by two insular endemic mammalian carnivores,the island fox and island spotted skunk

We compared resource utilization of two insular endemic mammalian carnivores, the island spotted skunk and island fox, along niche dimensions of space, food, and time on Santa Cruz Island. We predicted that resource use by foxes and skunks would differ along one or more niche dimensions, and that both species would have broader niches or higher densities compared with mainland relatives. Island foxes and island spotted skunks differed to some extent in habitat use, diets, and circadian activity, which may account for their long-term coexistence. Nonetheless, substantial overlap between skunks and foxes in spatial, dietary, and temporal dimensions suggests that competition between the two species does occur. Moreover, competition may be asymmetric, affecting skunks more than foxes. Compared with mainland foxes, island foxes have smaller body size, smaller home range, increased population density, increased diurnal activity, and behavior that is more highly inquisitive and less flightprone all common features of insular faunas. Island skunks, however, apparently have not developed these changes, perhaps due to asymmetric competition with foxes in conjunction with severe ecosystem disturbances caused by feral sheep.     

Mainland size variation informs predictive models of exceptional insular body size change in rodents

The tendency for island populations of mammalian taxa to diverge in body size from their mainland counterparts consistently in particular directions is both impressive for its regularity and, especially among rodents, troublesome for its exceptions. However, previous studies have largely ignored mainland body size variation, treating size differences of any magnitude as equally noteworthy. Here, we use distributions of mainland population body sizes to identify island populations as ‘extremely’ big or small, and we compare traits of extreme populations and their islands with those of island populations more typical in body size. We find that although insular rodents vary in the directions of body size change, ‘extreme’ populations tend towards gigantism. With classification tree methods, we develop a predictive model, which points to resource limitations as major drivers in the few cases of insular dwarfism. Highly successful in classifying our dataset, our model also successfully predicts change in untested cases.     

Intra- and interspecific variation in body size ofProtohermes (Megaloptera: Corydalidae)

The life history of three populations ofProtohermes grandis and two populations ofProtohermes immaculatus (Megaloptera: Corydalidae) was compared. In general, the larvae lived in stream riffles for 2 years and the adults appeared in summer. Adult body size differed between these closely related species and also between the populations ofP. grandis. Dwarfism occurred inP. immaculatus, a species that is endemic to the small, isolated island, Amami Island. The population ofP. grandis on Yaku Island, located between Amami Island and the mainland Kyushu, had an intermediate body size between that ofP. immaculatus and the mainland population ofP. grandis. Despite being an insular population,P. grandis on Tsushima Island had a similar body size to mainlandP. grandis. In these populations with large adults, some larvae lived in the streams for 3 years. The size distribution of benthic animals, which are the prey available toProtohermes larvae, differed between the streams studied. The density of large prey was lowest on Amami Island, intermediate on Yaku Island, and highest on the mainland and Tsushima Island. Different size distributions of available prey may be caused by the differences of benthic fauna; most of Ecdyonuridae and Ephemerellidae (large mayflies) and Perlidae (large stoneflies) were not found on Amami and Yaku Islands. Thus, there is a tendency to dwarfism in the populations ofProtobermes inhabiting streams where the density of large prey is low.     

Slaying dragons: limited evidence for unusual body size evolution on islands

Aim Island taxa often attain forms outside the range achieved by mainland relatives. Body size evolution of vertebrates on islands has therefore received much attention, with two seemingly conflicting patterns thought to prevail: (1) islands harbour animals of extreme size, and (2) islands promote evolution towards medium body size (‘the island rule’). We test both hypotheses using body size distributions of mammal, lizard and bird species. Location World‐wide. Methods We assembled body size and insularity datasets for the world’s lizards, birds and mammals. We compared the frequencies with which the largest or smallest member of a group is insular with the frequencies expected if insularity is randomly assigned within groups. We tested whether size extremes on islands considered across mammalian phylogeny depart from a null expectation under a Brownian motion model. We tested the island rule by comparing insular and mainland members of (1) a taxonomic level and (2) mammalian sister species, to determine if large insular animals tend to evolve smaller body sizes while small ones evolve larger sizes. Results The smallest species in a taxon (order, family or genus) are insular no more often than would be expected by chance in all groups. The largest species within lizard families and bird genera (but no other taxonomic levels) are insular more often than expected. The incidence of extreme sizes in insular mammals never departs from the null, except among extant genera, where gigantism is marginally less common than expected under a Brownian motion null. Mammals follow the island rule at the genus level and when comparing sister species and clades. This appears to be driven mainly by insular dwarfing in large‐bodied lineages. A similar pattern in birds is apparent for species within orders. However, lizards follow the converse pattern. Main conclusions The popular misconception that islands have more than their fair share of size extremes may stem from a greater tendency to notice gigantism and dwarfism when they occur on islands. There is compelling evidence for insular dwarfing in large mammals, but not in other taxa, and little evidence for the second component of the island rule – gigantism in small‐bodied taxa.     

An example of phenotypic adherence to the island rule? – Anticosti gray jays are heavier but not structurally larger than mainland conspecifics

The island rule refers to the tendency of small vertebrates to become larger when isolated on islands and the frequent dwarfing of large forms. It implies genetic control, and a necessary linkage, of size and body‐mass differences between insular and mainland populations. To examine the island rule, we compared body size and mass of gray jays (Perisoreus canadensis) on Anticosti Island, Québec, located in the Gulf of St. Lawrence, with three mainland populations (2 in Québec and 1 in Ontario). Although gray jays on Anticosti Island were ca 10% heavier, they were not structurally larger, than the three mainland populations. This suggests that Anticosti jays are not necessarily genetically distinct from mainland gray jays and that they may have achieved their greater body masses solely through packing more mass onto mainland‐sized body frames. As such, they may be the first‐known example of a proposed, purely phenotypic initial step in the adherence to the island rule by an insular population. Greater jay body mass is probably advantageous in Anticosti's high‐density, intensely competitive social environment that may have resulted from the island's lack of mammalian nest predators.     

Population genetic and behavioural variation of the two remaining colonies of Providence petrel (<Emphasis Type="Italic">Pterodroma solandri</Emphasis>)

Knowledge of the dispersal capacity of species is crucial to assess their extinction risk, and to establish appropriate monitoring and management strategies. The Providence petrel (Pterodroma solandri) presently breeds only at Lord Howe Island (~32,000 breeding pairs) and Phillip Island-7 km south of Norfolk Island (~20 breeding pairs). A much larger colony previously existed on Norfolk Island (~1,000,000 breeding pairs) but was hunted to extinction in the 18th Century. Differences in time of return to nesting sites are presently observed between the two extant colonies. Information on whether the Phillip Island colony is a relict population from Norfolk Island, or a recent colonization from Lord Howe Island, is essential to assess long-term sustainability and conservation significance of this small colony. Here, we sequenced the mitochondrial cytochrome b gene and 14 nuclear introns, in addition to genotyping 10 microsatellite loci, to investigate connectivity of the two extant P. solandri populations. High gene flow between populations and recent colonization of Phillip Island (95 % HPD 56–200 ya) are inferred, which may delay or prevent the genetic differentiation of these insular populations. These results suggest high plasticity in behaviour in this species and imply limited genetic risks surrounding both the sustainability of the small Phillip Island colony, and a proposal for translocation of Lord Howe Island individuals to re-establish a colony on Norfolk Island.     

The Micro-caddisflies of Lord Howe Island (Hydroptilidae: Trichoptera: Insecta)

Hydroptilid species are described from Lord Howe Island for the first time. All four species are in the endemic Australian genus Orphninotrichia, and all are endemic to Lord Howe Island. Modifications exhibited by these new species to the normally conservative Orphninotrichia features are suggestive of sympatric speciation and, for several, of a shift in larval niche.