Prey preference and gregarious attacks by the invasive flatworm Platydemus manokwari

The flatworm Platydemus manokwari (Tricladida: Rhynchodemidae) preys on various species of land snail, and its introduction to areas outside of its native range is thought to have caused the extinction of native land snails on several Pacific islands. Platydemus manokwari occurs in areas where land snails have been absent since its invasion, suggesting that the flatworm can prey on animals other than land snails. To identify the alternative prey and prey preferences of P. manokwari, I examined the feeding ecology of P. manokwari under field and laboratory conditions. Individuals were observed attacking live earthworms in a forest where land snails are already extinct, on Chichijima, Ogasawara (Bonin) Islands, Japan. I also observed them attacking earthworms and a species of isopod and land snail in the lab. To prey on the worm, similar to other earthworm-eating flatworms, P. manokwari wrapped itself around the prey and fed on it by inserting its pharynx into the earthworm’s body. Large earthworms were able to escape P. manokwari attacks by autotomy, but the autotomized body parts were eaten. Several P. manokwari individuals together attacked earthworms that were larger than themselves. The laboratory experiments showed that such gregarious attacks increased predation success on both large earthworms and land snails. The flatworms also attacked the isopods, although the predation rate was low. These results demonstrate that P. manokwari is a polyphagous predator of slow-moving soil invertebrates (land snails, slugs, earthworms, and isopods) and that invasion by P. manokwari may directly and indirectly impact native soil fauna.     

Experimental Test of Preferences for an Invasive Prey by an Endangered Predator: Implications for Conservation

Identifying impacts of exotic species on native populations is central to ecology and conservation. Although the effects of exotic predators on native prey have received much attention, the role of exotic prey on native predators is poorly understood. Determining if native predators actively prefer invasive prey over native prey has implications for interpreting invasion impacts, identifying the presence of evolutionary traps, and predator persistence. One of the world’s most invasive species, Pomacea maculata, has recently established in portions of the endangered Everglade snail kite’s (Rostrhamus sociabilis plumbeus) geographic range. Although these exotic snails could provide additional prey resources, they are typically much larger than the native snail, which can lead to lower foraging success and the potential for diminished energetic benefits in comparison to native snails. Nonetheless, snail kites frequently forage on exotic snails. We used choice experiments to evaluate snail kite foraging preference in relation to exotic species and snail size. We found that snail kites do not show a preference for native or exotic snails. Rather, snail kites generally showed a preference for medium-sized snails, the sizes reflective of large native snails. These results suggest that while snail kites frequently forage on exotic snails in the wild, this behavior is likely driven simply by the abundance of exotic snails rather than snail kites preferring exotics. This lack of preference offers insights to hypotheses regarding effects of exotic species, guidance regarding habitat and invasive species management, and illustrates how native-exotic relationships can be misleading in the absence of experimental tests of such interactions.     

The Decimation of Endemic Hawai'ian Tree Snails by Alien Predators

Endemic terrestrial tree snails of the Hawai'ian Islands, likethose of other oceanic islands and even some continental areas,are extremely sensitive to disturbance because of their lowpopulation numbers and small geographic ranges. Like many otherplants and animals of oceanic islands, they have evolved nodefenses against introduced predators and competitors. The rangeof Achatinella mustelina, a tree-snail species found only ina short mountain range on the island of O'ahu, typifies thisproblem. Mark-recapture studies at two field locations revealthat the snails exhibit slow growth andlate maturity (3–5years). Fecundity is estimated at about 7 offspring per adultper year. The young are born live at about 4.6 mm. Populationgrowth typically depends on considerable longevity (> 10years). Demographic effects of the depredations by alien predators,rats and a North American predatory snail, Euglandina rosea,were documented in two long-term study sites. The predatorysnail eats all sizes of A. mustelina and can rapidly drive populationsto extinction (less than one year). Rats tend to select largersnails as prey and may leave an area before destroying all ofthe prey snails present; while reproductive output is temporarilydestroyed, populations may survive. Actions necessary to conserveHawai;ian tree snails, or indeed any group of relatively sedentaryinvertebrates with small species ranges, must include predatorabatement, but also preservation and restoration of sufficientlyarge and complex forest habitats that the invertebrates mayfind refuge from alien predators.     

Effect of beetle size on predation of cabbage root fly eggs by ground beetles

Laboratory studies were done to determine the numbers of cabbage root fly (Delia radicum L.) (Diptera: Anthomyiidae) eggs eaten by sixty species of ground beetles (Coleoptera: Carabidae) collected from a range of habitats. For 35 species that were between 2.7 mm and 10 mm long, there was a linear relationship between the numbers of eggs eaten and beetle length. For each 1 mm increase in length above 2.7 mm, an additional 18 eggs were eaten/beetle/day. Eight further species within the above size range did not eat any fly eggs, as these species are known to be either phytophagous or to feed on moving prey. Large numbers of eggs were eaten by beetles in 13 of the 25 genera tested. Based on size, maximum numbers of eggs were eaten by 7 species of Agonum Bonelli, 5 species of Amara Bonelli and seven species of Bembidion Latreille. Only one of the 6 species of Pterostichus Bonelli tested, P. strenuus (Panz.), ate the expected numbers of eggs, the other species being too large to feed from such small prey items as cabbage root fly eggs. The numbers of beetles larger than 10 mm that ate eggs was highly variable and so 10 mm was considered the upper size limit of carabid predators of cabbage root fly eggs.     

A tale of two snails: is the cure worse than the disease?

The giant African snail, Achatina fulica, has been introduced to many parts of Asia as well as to numerous islands in the Indian and Pacific Ocean, and has recently reached the West Indies. It has been widely decried as a disaster to agricultural economies and a threat to human health, leading to a clamor for the introduction of biological control agents. In fact, the lasting impact on agriculture may not be severe, and the human health risk is probably minor. This snail can be an aesthetic atrocity and a nuisance in other ways, however. Wherever A. fulica has achieved high densities, it has subsequently undergone a striking decline. Although this decline has been attributed to introduced predators, there is little evidence for this hypothesis; instead, epizootic disease seems to be at least part of the cause. However, the introduced predators, especially a New World snail, Euglandina rosea, have wrought havoc with the native land snails of many islands. They have already caused many extinctions and will almost certainly cause others. This predator was introduced by government agencies in many areas despite warnings from competent biologists that the effects could be disastrous. Pressures for such actions may become overwhelming in the face of a highly visible invasion, despite policies that should mandate extreme caution.     

The predatory snail Euglandina rosea successfully follows mucous trails of both native and non‐native prey snails

Mucous trail following is a primary means by which many predatory snails locate prey. I compared the ability of individuals of Euglandina rosea to follow mucous trails of two groups of gastropods: those found within its native habitat (southeastern USA), and those found outside its native range (Kansas). Members of E. rosea followed trails for both species found inside and outside its native range equally well. In contrast to previous studies, I found that the predatory snails consistently followed trails in the direction in which they were laid. I quantified the kinematics of trail‐following behavior using inter‐tentacle angle as the primary metric. In both prey groups, there were significant differences in the predator's inter‐tentacle angle when tracking a trail versus not, and when successfully following (in the direction the trail was laid) versus unsuccessfully following (opposite the direction that the trail was laid) trails. In addition, in both prey groups, there were significant differences in the predator's velocity when tracking a trail versus not, and when successfully versus unsuccessfully following trails. This study confirmed that members of E. rosea are robust generalist predators, capable of successfully tracking native and non‐native snails, and should not be introduced as biologic control agents. These results may be useful to managers, as they provide insight into how trail following could be used to trap members of this invasive species.     

Extinction or survival: partulid tree snails in American Samoa

Four partulid tree snail species are known from American Samoa. In 1998, we surveyed the recently established National Park (units on three islands: Tutuila, Tau, Ofu) and neighboring areas for partulids. On Tutuila, Samoana abbreviata, previously considered probably extinct, was extremely rare (15 snails seen); Samoana conica was more common (288 snails) but still rare; Eua zebrina was the most common (1102 snails), at one locality perhaps near its natural abundance. The species have similar distributions within the Park. All three have declined dramatically since the 1920s. Before 1980, when the predatory snail Euglandina rosea was introduced, habitat destruction, and perhaps rat predation and shell collecting, probably caused the decline. The Park provides protection to the Tutuila partulids by protecting habitat, although development is still a potential problem. But predation by E. rosea may yet cause their extinction. On Ofu (only outside the Park), 12 live Samoana thurstoni were found; 31 have now been recorded by western science. Also, an apparently robust population of E. zebrina, previously considered a Tutuila endemic, was found. Euglandina rosea is not on Ofu, so these populations are important remnants of the fauna. No partulids were found on Tau; none has ever been recorded there.     

Land molluscs as food of Malayan rodents and insectivores

Laboratory and field experiments demonstrate that, contrary to previously published stomach analysis data, rats selectively consume land molluscs. Rattus jalorensis and R. argentiventer prefer molluscs as food, while R. exulans and R. r. diardi are less inclined to do so. Macrochlamys resplendens , a land snail, is most favoured, with the slug Micropar-mariort malayanus next in preference. The heavily-shelled Achatina fulica is only occasionally eaten and Quantula striata is ignored. Examination of gastro-intestinal contents of wild rats caught from various habitats in Malaya show that when present in large numbers molluscs form an important part of the natural food of rats, yet laboratory evidence demonstrates that as many as 60% of mollusc-eating rats may fail to show evidence of shells on examination within 12 hours of feeding–an indication that stomach analyses alone may be of limited value.
The Malayan insectivores Echinosorex gymnurus and Suncus murinus often feed on snails and slugs, though they appear not to be suitable hosts for infection with the rat lungworm Angiostrongylus cantonensis.     

The food niches of the invasive Dugesia tigrina (Girard) and indigenous Polycelis tennis Ijima and P. nigra (Müller) (Turbellaria; Tricladida) in a Welsh lake

The invasion of Llyn Coron in North Wales by the American immigrant Dugesia tigrina resulted in the almost entire displacement of the native Polycelis tenuis and P. nigra populations. Because competition for food is the most important factor controlling and regulating populations of British lake-dwelling triclads, the diets of the invasive and native triclads in the lake were examined in an attempt to explain the successful invasion.A serological technique, the precipitin test, identified the gut contents of field-collected triclads. Niche breadth, electivity and niche overlap indices were used in analysis of the data. A broad food niche was recorded for all the triclad species. Oligochaetes predominated in the diet of D. tigrina followed by Asellus and chironomids, in almost equal proportions, snails and caddisflies, with mayflies scarcely eaten. Both Polycelis species fed heavily on oligochaetes followed by Asellus. P. tenuis ate slightly higher proportions of chironomids and caddisflies than snails and mayflies, whilst P. nigra consumed similar proportions of these four prey taxa. Gammarus, scarce in Llyn Coron, and cladocerans were not eaten by the Polycelis species and by only a few D. tigrina.To-date there has been no apparent shift in the diet of the Polycelis species as a consequence of the invasion by D. tigrina. In the absence of any evidence for the partitioning of food on the basis of prey size or condition, the considerable overlap in the diets of D. tigrina and the Polycelis species suggests the potential for severe inter-specific competition for food. The superior exploitation by D. tigrina of the available food resource requires explanation.     

Ecological and evolutionary implications of diet in monitor lizards

A survey of 35 species indicates that monitor lizards (Varanus) typically hunt over large areas, search in particular microhabitats, and feed frequently on a wide variety of prey, many of which are relatively small. There is ontogenetic, seasonal, and geographic variation in diet. With some exceptions, invertebrates are the predominant prey, but rare predation on vertebrates is often energetically significant. A few monitors specialize on prey types that occur as occasional items in the diet of species with more generalized diets; these include crabs, snails, orthopterans, lizards, and large mammals. For most species, prey specialization occurs via habitat selection and a variety of prey types and sizes are eaten, as expected for widely searching predators. Comparisons with other anguimorphans suggest that derived features of Varanus are associated with high body temperature and activity levels; specialized chemoreception; and rapid, skillful capture of hidden and/or potentially hard to catch prey. Occasional ingestion of moderately large prey is primitive for Varanoidca (Helodermatidae +Varanidae), accentuating a trend that is perhaps primitive for anguimorphan lizards. Reduction of very large prey prior to ingestion is a derived attribute within Varanus , seen infrequently in several larger species and commonly in V. komodoensis. This study illustrates the synthesis of comparative natural history in a phylogenetic context, a method that addresses the history of organismal change.     

Home-field advantage: native signal crayfish (Pacifastacus leniusculus) out consume newly introduced crayfishes for invasive Chinese mystery snail (Bellamya chinensis)

The introduction of non-indigenous plants, animals and pathogens is one of today’s most pressing environmental challenges. Freshwater ecologists are challenged to predict the potential consequences of species invasions because many ecosystems increasingly support novel assemblages of native and non-native species that are likely to interact in complex ways. In this study we evaluated how native signal crayfish (Pacifastacus leniusculus) and non-native red swamp crayfish (Procambarus clarkii) and northern crayfish (Orconectes virilis) utilize a novel prey resource: the non-native Chinese mystery snail (Bellamya chinensis). All species are widespread in the United States, as well as globally, and recent surveys have discovered them co-occurring in lakes of Washington State. A series of mesocosm experiments revealed that crayfish are able to consume B. chinensis, despite the snail’s large size, thick outer shell and trapdoor defense behaviour. Crayfish exhibited size-selective predation whereby consumption levels decreased with increasing snail size; a common pattern among decapod predators. Comparison of prey profitability curves—defined as the yield of food (weight of snail tissue) per second of feeding time (the time taken to crack the shell and consume the contents)—suggests that small and very large snails may represent the most profitable prey choice. By contrast, previous studies have reported the opposite pattern for crayfish consumption on thin-shelled snails. For all snail size classes, we found that native P. leniusculus and invasive O. virilis consumed greater numbers of snails than invasive P. clarkii. Moreover, P. leniusculus consistently handled and consumed snails at a faster pace compared to both invasive crayfishes across the range of snail sizes examined in our study. These results suggest not only that B. chinensis is a suitable food source for crayfish, but also that native P. leniusculus may ultimately out-consume invasive crayfishes for this new prey resource.     

Predation on endemic Hawaiian land snails by the invasive snail Oxychilus alliarius

The native land snails of the Hawaiian Islands have suffered numerous extinctions and continue to decline rapidly. A leading cause of these extinctions has been the impact of non-native predators. Oxychilus alliarius is a European predatory snail that has been in Hawaii since at least 1937 and has become one of the most abundant invasive snails in the islands. It is a facultative carnivore and will also consume non-animal food. However, this species has been mostly ignored as a threat and the majority of conservation attention has been focused on Euglandina rosea, the Rosy Wolf Snail. Here we report predation by O. alliarius on two rare and endemic Hawaiian snail species: Kaala subrutila (Helicarionidae) and Pleuropoma cf. sandwichiensis (Helicinidae). The wide distribution of O. alliarius in Hawaii, its high abundance, and now compelling evidence that it preys on native (and probably endangered) snails, indicate that it could be a much greater threat than previously thought.     

Integration of an invasive consumer into an estuarine food web: direct and indirect effects of the New Zealand mud snail

Introduced species interact both directly and indirectly with native species. We examine interactions between the introduced New Zealand mud snail (Potamopyrgus antipodarum) and native estuarine invertebrates and predators through experiments and field studies. A widely held management concern is that when P. antipodarum, which has low nutritional value, becomes abundant, it replaces nutritious prey in fish diets. We tested two key components of this view: (1) that fish consume, but get little direct nutritional value from P. antipodarum; and (2) that P. antipodarum has an indirect negative effect on fish by reducing the energy derived from native prey. We also examined predation by the native signal crayfish, Pacifastacus leniusculus. Laboratory feeding trials showed that both crayfish and fish consume P. antipodarum, a direct effect. Crayfish consumed and successfully digested higher numbers of snails than did fish [Pacific staghorn sculpin (Leptocottus armatus), three spine stickleback (Gasterosteus aculeatus), and juvenile starry flounder (Platicthys stellatus)]. P. antipodarum occurred at low frequencies in the stomachs of wild-caught fish. More interesting were the indirect effects of this invader, which ran counter to predictions. P. antipodarum presence was associated with no change or an increase in the amount of energy derived from native prey by predators. The presence of P. antipodarum also led to increased consumption of and preference for the native amphipod Americorophium salmonis over the native isopod Gnorimosphaeroma insulare. This is an example of short-term, asymmetric, apparent competition, in which the presence of one prey species (snails) increases predation on another prey species (the amphipod).     

High diversity and regional endemism in land snails of eastern Tanzania

In February/March 1995 we collected land snails (including slugs) at 12 stations in eastern Tanzania. A total of 571 person-hours yielded 9174 snails assigned to 159 morpho-species. The richest two sites each (<4ha of uniform forest) had 50 species (Amboni Cave) and 48 species (near Amani, Usambaras), nearly as great as the most species-rich sites known in the world; sieving of litter and soil would probably yield more species. In lowland (coastal) forests, both diversity and endemism seemed to decrease from north to south. Most snail species were found within only one of four coastal or one montane geographic regions, indicating substantial regional endemism. Only one species (Achatina fulica) appeared in all five regions, and 84% of all other species were found in only one (61%) or two regions (23%). The predatory streptaxids comprised about half the species and a third of the individuals at the Usambara site, an extremely high ratio of carnivores. Small snails (< 5mm greatest adult shell dimension) – many of which are probably undescribed species – comprise a substantial proportion of Tanzanian molluscan diversity; more surveys are needed, especially because of human pressures on the few forest patches remaining.     

INVESTIGATING THE BIASES IN THE USE OF HARD PREY REMAINS TO IDENTIFY DIET COMPOSITION USING ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) IN CAPTIVE FEEDING TRIALS

The analysis of pinniped scats has been used to quantify their diet, using prey remains to identify species and to estimate the numbers and sizes of prey consumed. There are, however, potential biases involved with scat analysis and, for this method to be used successfully, these biases need to be quantified. Thirty-six Antarctic fur seals ( Arctocephalus gazella ) were fed meals of exclusively either fish, squid, or krill and their scats were collected and analyzed. The major sources of error in the analysis of prey remains from scats were the differential erosion and passage rate of items in relation to their size. However, using simple correction functions, such as those which model otolith erosion, it is possible to reduce these errors. Using plastic beads as dietary markers showed recovery rates were negatively related to their size. Larger squid beaks had lower recovery rates than smaller beaks, but there was no size-related bias in the recovery rates of krill carapaces. Only 33% of the squid beaks and 27% of the otoliths originally fed were recovered from the scats. Recovery rates were greater for squid (77%) and fish (50%) eye lenses and these structures gave a better estimate of numbers eaten. Differences found between experimentally derived and published regression equations (used to calculate prey sizes eaten from prey remains) highlights the need for regression equations based on local prey characteristics, if these are to be used with any success to describe the prey eaten.     

Trait-based plant mortality and preference for native versus non-native seedlings by invasive slug and snail herbivores in Hawaii

Non-native herbivores may alter plant communities through their preferential consumption of seedlings of different species. We assessed seedling herbivory by two invasive gastropod species in Hawaii, the giant African snail (Achatina fulica) and the Cuban brown slug (Veronicella cubensis). We hypothesized that six native species would suffer greater gastropod herbivory than four non-native species, and that species with short stature, thin leaves, and lacking physical defenses would suffer the greatest mortality from gastropods. Herbivory was measured during 13-day preference trials using enclosures that each contained four different woody species (two native, two non-native) and were assigned to one of three treatments: giant African snail, Cuban brown slug, or control (no gastropod). Discriminant function analysis was used to predict gastropod-induced seedling mortality from a suite of seedling characteristics. Native species did not always experience greater herbivory than non-natives species, and seedling mortality was 0–100 %. Native Pipturus albidus and Clermontia parviflora suffered 100 % mortality from V. cubensis herbivory, and P. albidus, Psychotria hawaiiensis, and Myrsine lessertiana suffered ≥80 % mortality from A. fulica. Two non-natives (Fraxinus uhdei, Clidemia hirta), and two natives (Metrosideros polymorpha, Diospyros sandwicensis), suffered little damage and no mortality. Non-native Ardisia elliptica suffered 10–30 % gastropod mortality, and non-native Psidium cattleianum mortality was 0–50 %. Leaf thickness best predicted species mortality caused by slugs and snails; some thicker-leaved species suffered most. Invasive snails and slugs threaten some native and non-native seedlings by directly consuming them. Current and future plant community structure in Hawaii may in part reflect the feeding preferences of invasive gastropods.     

Prey selection by thaidid gastropods: some observational and experimental field tests of foraging models

Summary Field observations and experiments revealed that predatory intertidal gastropods of the genus Thais (or Nucella) were able both to recognize the expected food value of encountered prey (expected energy or growth potential gained per unit handling time) and to monitor their average yield over time (average energy or growth potential gained per unit foraging time). They appeared to discriminate not only among prey species, but also among different sized individuals of the same prey species. The evidence supporting these interpretations included: 1) field observations of snails feeding preferentially on prey types of higher expected food value even though lower value prey types were available and abundant, 2) a very limited number of direct underwater observations of foraging snails rejecting encountered items that were either of lower expected value than the item finally eaten or not measurably different from it, and 3) field (=arena) experiments in which both average yield, and the distribution and abundance of potential prey were controlled: snails conditioned at a high average yield fed preferentially on high value items, while those animals conditioned at a low yield consumed prey in the proportions that they were encountered. These behaviors are all consistent with a prey-selection decision motivated by energetic considerations. Further, the field experiments indicated that these predatory gastropods could select items from a diverse array of prey so as to maximize growth in their natural environment. The behaviors were not consistent with three alternative foraging hypotheses: non-selective foraging, frequency-dependent foraging on prey types (here, sizes of particular prey species), and frequency-dependent foraging on prey species. Deviations from some of the quantitative predictions of optimal foraging theory appeared related to learning and risk.     

Feral Cats Are Better Killers in Open Habitats,Revealed by Animal-Borne Video

One of the key gaps in understanding the impacts of predation by small mammalian predators on prey is how habitat structure affects the hunting success of small predators, such as feral cats. These effects are poorly understood due to the difficulty of observing actual hunting behaviours. We attached collar-mounted video cameras to feral cats living in a tropical savanna environment in northern Australia, and measured variation in hunting success among different microhabitats (open areas, dense grass and complex rocks). From 89 hours of footage, we recorded 101 hunting events, of which 32 were successful. Of these kills, 28% were not eaten. Hunting success was highly dependent on microhabitat structure surrounding prey, increasing from 17% in habitats with dense grass or complex rocks to 70% in open areas. This research shows that habitat structure has a profound influence on the impacts of small predators on their prey. This has broad implications for management of vegetation and disturbance processes (like fire and grazing) in areas where feral cats threaten native fauna. Maintaining complex vegetation cover can reduce predation rates of small prey species from feral cat predation.     

Feeding selectivity of four species of sympatric stickleback in brackish-water habitats in eastern Canada

Although sympatric populations of sticklebacks occur commonly along the north-eastern coast of North America, very few studies have considered the interspecific relationships within such populations. This paper investigates prey selectivity, feeding behaviour and potential prey availability within sympatric populations from five brackish-water sites in New Brunswick. Differences between the diets of species from different sites were related to differences in the range of prey available. Fish collected from sites with abundant aquatic vegetation consumed greater numbers of prey than those from vegetation-poor sites. The proportions of prey types eaten by both adults and juveniles reflected prey species abundance at the different sites; thus shifts in diet over time were found to be related to changes in the abundance of potential prey and not to shifts in prey preference on the part of the fish. The size of prey consumed was related to mouth size; adult and juvenile Gasterosteus wheatlandi , having the smallest mouths, consumed the smallest prey. Differences in prey size detected in the four species studied were due to differences in the types of prey consumed. Laboratory experiments on feeding behaviour showed that adult Apeltes quadracus and Pungitius pungitius were more efficient at capturing benthic prey than were Gasterosteus aculeatus and G. wheatlandi. Competition for food is thought not to occur in these populations, because of the abundance of potential prey and the morphological constraints on feeding behaviour.     

Patterns of introduction of non-indigenous non-marine snails and slugs in the Hawaiian Islands

The native snails of the Hawaiian Islands are disappearing. One cause is predation by introduced carnivorous snails. Habitat destruction/modification is also important, facilitating the spread of other non-indigenous snails and slugs. Eighty-one species of snails and slugs are recorded as having been introduced. Thirty-three are established: 12 freshwater, 21 terrestrial. Two or three species arrived before western discovery of the islands (1778). During the nineteenth century about one species per decade, on average, was introduced. The rate rose to about four per decade during the twentieth century, with the exception of an especially large number introduced in the 1950s as putative biocontrol agents against the giant African snail, Achatina fulica. The geographical origins of these introductions reflect changing patterns of commerce and travel. Early arrivals were generally Pacific or Pacific rim species. Increasing trade and tourism with the USA, following its annexation of Hawaii, led to an increasing proportion of American species. More general facilitation of travel and commerce later in the twentieth century led to a significant number of European species being introduced. African species dominated the 1950s biological control introductions. The process continues and is just part of the homogenization of the unique faunas of tropical Pacific islands.