Mark A. Schmaedick | Geology Reports

USGS2024

Reproductive ecology and egg parasitism of the Samoan swallowtail butterfly

We investigated the reproductive ecology and effects of egg parasitism on the Samoan swallowtail butterfly ( Papilio godeffroyi ), which survives only on Tutuila Island, American Samoa, after having disappeared from the much larger islands of Upolu and Savai‘i in independent Samoa. During monthly surveys of its only known host plant, Micromelum minutum , across eight sites in 2013 and 2014, we collected eggs, eggshells, larvae, pupae, and pupal exuviae. Live specimens were reared under laboratory conditions to determine reproductive outcomes, developmental rates, and sex ratios, as well as parasitoid attack frequencies, brood sizes, and sex ratios. Sixty-six of 448 (14.7%) eggs produced larvae, 47 of which became adults. The sex ratio was approximately even overall and within each developmental stage. Eggs were slightly larger on individual host trees and in host tree stands that yielded more eggs per unit of foliage, indicating that ovipositing females responded to some features of host trees and stands. Eggs hatching female or male larvae were similar in size, and the sexes developed at similar rates. A newly described species of parasitoid wasp, Ooencyrtus pitosina (Encyrtidae), emerged from 73.6% of 382 butterfly eggs that failed to hatch in the laboratory (62.7% of 448 eggs overall). Forty-one other eggs contained dead parasitoid larvae. An additional, unidentified Ooencyrtus wasp species emerged from a single P. godeffroyi egg. No parasitoids were reared from P. godeffroyi larvae or pupae. Of 656 P. godeffroyi eggshells collected in the field and examined in the laboratory, 62.2% showed signs of having been parasitized by O. pitosina. There was no evidence that parasitism rates were density-dependent. O. pitosina brood sizes ranged from 1 to 5, with the sex ratio skewed toward females (2.40 F:1.00 M). Larger parasitoid broods were associated with slightly larger host eggs, indicating that female wasps may adjust brood size according to host egg size or that fewer wasp larvae are able to complete development in smaller eggs. Techniques used to rear both P. godeffroyi and O. pitosina in the laboratory could be applied to a captive-rear, wild-release program, which may facilitate reestablishment of the species in Samoa.

Tutuila Island↗

USGS2023

Reproductive response of the Samoan swallowtail butterfly to variability in host plant and habitat characteristics

The Samoan swallowtail butterfly ( Papilio godeffroyi ) has become restricted to Tutuila Island, American Samoa. Factors driving its extirpation on other islands may be partly due to the availability and suitability of habitat, given the singular association we observed of P. godeffroyi with its host plant, Micromelum minutum. We expected that as a host plant specialist, P. godeffroyi might respond to variation in the physical traits and habitat context of M. minutum , which could help identify conservation opportunities throughout its present and former range. Because the ecology of P. godeffroyi is largely unknown, we conducted systematic surveys during 15 months in 2013 and 2014 on Tutuila to investigate its reproductive response to the physical and phenological variability of individual host trees and to the structural and geographical variability of the stands in which they occurred. We searched foliage for eggs, larvae, and pupae and monitored seasonal trends in herbivory and production of leaves, flowers, and fruits on 117 host trees in 8 rainforest stands, mostly within the National Park of American Samoa (NPSA), starting in March 2013 and ending in August 2014. We observed high variability in butterfly reproductive output between individual trees and stands, due largely to a strong negative relationship between leaf biomass (at both tree and stand levels) and oviposition density (number of eggs per foliar biomass) and frequency (number of eggs per survey). Distribution patterns of larvae and pupae were less distinct but generally followed trends in oviposition. Our findings indicate that the butterfly is reproducing widely across Tutuila and is likely to respond positively to management to increase the availability of small host trees in low-density stands. Thus, the species is a good candidate for conservation management focusing on habitat.

Tutuila Island↗

USGS2016

Effects of Climate and land use on diversity, prevalence, and seasonal transmission of avian hematozoa in American Samoa

The indigenous forest birds of American Samoa are increasingly threatened by changing patterns of rainfall and temperature that are associated with climate change as well as environmental stressors associated with agricultural and urban development, invasive species, and new introductions of avian diseases and disease vectors. Long term changes in their distribution, diversity, and population sizes could have significant impacts on the ecological integrity of the islands because of their critical role as pollinators and seed dispersers. We documented diversity of vector borne parasites on Tutuila and Ta‘u Islands over a 10-year period to expand earlier observations of Plasmodium, Trypanosoma , and filarial parasites, to provide better parasite identifications, and to create a better baseline for detecting new parasite introductions. We also identified potential mosquito vectors of avian Plasmodium and Trypanosoma , determined whether land clearing and habitat alterations associated with subsistence farming within the National Park of American Samoa can influence parasite prevalence, and determined whether parasite prevalence is correlated with seasonal changes in rainfall, temperature and wind speed. Three taxonomically distinct lineages of Plasmodium were identified from mosquito vectors and forest birds based on partial sequence data from parasite mitochondrial genes. All three have been described from passerine and galliform birds in Australasia. Two lineages, SCEDEN01 and ORW1, had elongate gametocytes and large schizonts that were consistent with species of Plasmodium in the subgenus Giavannolaia, but were taxonomically distinct from known morphological species of Plasmodium based on a Bayesian phylogenetic analysis of a 478 bp region of the parasite cytochrome b gene. Both are candidates for description as new species. The third lineage (GALLUS02) was detected only in mosquito vectors on Tutuila and was similar in cytochrome b sequence to P. juxtanucleare , a pathogenic species of Plasmodium from chickens and other galliform birds from Australasia, Africa, and South America. Plasmodium relictum , the malarial parasite that has had such a devastating impact on Hawaiian forest birds, was not detected. We observed large, striated trypanosomes in avian hosts from both Tutuila and Ta‘u Islands that fell within the same taxonomic clade as T. corvi and T. culicavium based on 18S ribosomal DNA sequence. We also observed sheathed microfilariae with pointed tails that had some morphological similarities to microfilaria from species of Pelecitus, Struthiofilaria and Eulimdana , but identification will require recovery and examination of adult filarial worms from the connective tissue or body cavities of infected birds. We also observed one or more species of haemococcidians (Isospora, synonym = Atoxoplasma ) within circulating lymphocytes from multiple avian host species. Overall prevalence of Plasmodium was higher on Ta‘u (22%, 75/341) than Tutuila (9.2%, 27/294), with most infections occurring in Polynesian starlings, Samoan starlings, Wattled honeyeaters, and Cardinal honeyeaters. Prevalence was relatively constant from year to year and between seasons at individual study sites, but varied among study sites, with highest rates of infection in areas with agricultural activity at Faleasao (37.4%, 73/195, Ta‘u Island) and Amalau Valley (9.7%, 21/216, Tutuila Island). Prevalence in more remote areas of the National Park of American Samoa was lower, ranging from 1.4% (2/146) at Laufuti and Luatele on Ta‘u to 7.7% (6/78) at Olo Ridge on Tutuila. Similar trends were evident for infections with Trypanosoma and filarial worms. Overall prevalence was not influenced significantly by warmer, wet (summer) or cooler, dry (winter) season. We detected Plasmodium infections in Culex sitiens and C. quinquefasciatus through either salivary gland and midgut dissections or PCR amplification of parasite cytochrome b genes in pooled or individual samples of mosquitoes that were collected on Tutuila. Pooled or individual Aedes oceanicus, A. polynesiensis, A. tutuilae, A. upolensis, A. nocturnus, Aedes (Finlaya) (mixed pools of A. samoanus, A. oceanicus, A. tutuilae), Aedes (Stegomyia) (mixed pools of A. aegypti, A. upolensis, A. polynesiensis ), and C. annulirostris were negative for Plasmodium , but we detected infections with Trypanosoma through midgut and salivary gland dissections in a single C. sitiens from Amalau Valley, Tutuila and three A. oceanicus from Faleasao, Ta‘u. Two of the A. oceanicus from Faleasao amplified successfully with Trypanosoma primers, but sequences were distinctly different from those obtained from avian hosts. We found a strong association between land use and prevalence of mosquito-transmitted parasites on Ta‘u Island with odds of being infected more than 20 times greater in agricultural plots than more remote native forest. This relationship was evident on Tutuila Island but not statistically significant because of the close proximity of study sites and observed movement of birds between native forest and agricultural land. Our data support previous studies that have suggested that Plasmodium and other vector-borne parasites are part of the indigenous parasite fauna in American Samoa. Transmission dynamics appear to be affected by environmental changes associated with land use practices.

HI↗