International Biological Program Technical Reports (1970-1975)

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The Pacific Cooperative Studies Unit & The Hawaii-Pacific Islands Cooperative Ecosystems Studies Unit
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Canopy-associated arthropods in Acacia koa and Metrosideros tree communities along the Mauna loa Transect
(Island Ecosystems IRP, U.S. International Biological Program, 1976-06) Gagne, Wayne C
The spatial distribution and zonation of canopy-associated arthropods of Acacia koa and Metrosideros tree communities along an altitudinal transect on the east flank of Mauna Loa was determined by insecticidal fogging of the canopy with pyrethrum. Eight sites were on the Mauna Loa Transect, which has been intensively sampled by IBP participants in the Island Ecosystems IRP. Two sets of transect zones were determined on the basis of arthropod distribution. The influence of environmental and biotic factors, plant community structure and climate are interpreted according to distribution patterns. The distribution of arthropod groups coincided quite closely with vascular plant communities of the transect as defined by other studies. The composition, spatial distribution, and environmental relationships of arthropod canopy communities along the Mauna Loa Transect are compared with the situation pertaining along other lower elevational transects to sea level in Hawaii Volcanoes National Park as well as with other ecosystems in order to further characterize the arthropod canopy community. Host specificity, vegetation structure, competition between ecological homologs, and climate appeared to have the most important influence on population density and spatial distribution patterns of the arthropod taxa studied.
Spatial Distribution of Bird Species on the East Flank of Mauna Loa
(Island Ecosystems IRP, U.S. International Biological Program, 1975-10) Conant, Sheila
Eleven transects in ten different types of plant communities found along an altitudinal gradient on the east flank of Mauna Loa were sampled monthly using the "count x detectability" method of censusing birds. Eight of these sites were on the Mauna Loa Transect, which has been intensively sampled by IBP participants in the Island Ecosystems IRP. Frequency, density and distribution of the 29 bird species encountered are discussed. Data on species presence and density at each site were used to construct community (sampling site) ordinations based on similarity indices and species ordinations based on two-way tables. The results of these analyses provided the basis for the objective definition of transect zones for those sites located on the Mauna Loa Transect. The transect zones objectively defined by avian community analyses were almost identical to those based on analyses of plant communities on Mauna Loa. The habitats selected by each bird species are discussed. Optimum habitats as a reflection of maximum density are described for each species. At least two types of environmental factors associated with the altitudinal gradient appear to be important influences on patterns of avian spatial distribution: 1) continuously varying environmental factors (e.g., rainfall, temperature, gradual transitions from one plant community to another); 2) habitat discontinuities (narrow ecotones, abrupt changes in vegetation structure). Competition is briefly discussed as a factor that may affect density and distribution of some bird species. In some cases (e. g., 'Apapane, 'I’iwi) it is apparent that very specific factors (e.g., blooming time and amount in certain important food plants such as Metrosideros and Sophora) strongly influence the distribution and local abundance of a few bird species. However, vegetation structure appeared to be the single most important influence on population density and patterns of spatial distribution of the bird species studied.
Ecology of Fungi in Wildland Soils along the Mauna Loa Transect
(Island Ecosystems IRP, U.S. International Biological Program, 1975-11) Stoner, Martin F.; Stoner, Darleen K.; Baker, Gladys E.
The distribution of fungi in soils along the Mauna Loa Transect was determined by an approach employing specific fungal reference genera, selective isolation methods, and a combination of analytical techniques. Two sets of transect zones were determined on the basis of fungal distribution. The influence of environmental factors, particularly those relating to soil, vascular plant communities, and climate, are interpreted according to distribution patterns. The distribution of fungal groups coincided clearly with vascular plant communities of the transect as defined by other studies. Features of the structure, stability, and development of fungal communities, and of the ecological roles of certain fungi are indicated by the results. The composition, spatial distribution, and environmental relationships of fungal communities along the Mauna Loa Transect are compared with situations in other insular and continental ecosystems in order to further characterize and elucidate the ecology of the Hawaiian soil-borne mycoflora. An overall evaluation of the research indicates that the selective methods employed to evaluate fungal distribution represent an effective approach to ecosystem analysis on a broad scale.
The Use of Sheep Wool in Nest Construction by Hawaiian Birds
(Island Ecosystems IRP, U.S. International Biological Program, 1975-09) van Riper, Charles III
The utilization of sheep wool as a nesting material was examined from 1969 through 1975 on the island of Hawaii. Of the 10 bird species studied, six incorporated wool into their nests. Both introduced and endemic birds use wool, with a significantly greater usage by endemic birds. Use of wool in nest construction appears correlated with the intricacy of the nest that a species builds, with a significant difference between degree of usage in complex and simple nests. Roughly built nests, like those of the Cardinal (Cardinalis cardinalis), contained no wool whereas the complex nest of the Elepaio (Chasiempis sandwichensis) had a great deal of wool. Wool is apparently used by the birds because it is a readily available material in certain areas, and because of its binding quality. The wool is gathered from tufts that snag on branches as the sheep pass or from dried skins on the ground. The amount of wool utilized in each nest varies both interspecifically and intraspecifically, but in all nests only the body of the nest contained wool, the lining always being of other materials. A separate study was conducted to determine if wool is used only when available nearby or is a sought-after material. Only the Elepaio was found to consistently travel distances to procure wool, whereas the other species studied used it only when available within their territories.
Plant-Pollinator Interactions in Hawaii: Pollination Energetics of Metrosideros collina (Myrtaceae)
(Island Ecosystems IRP, U.S. International Biological Program, 1976-04) Carpenter, F Lynn
The most abundant tree species in much of the undisturbed Hawaiian forests was the subject of a two year study on plant-pollinator interaction and energetics. The purposes of the study were 1) to determine the roles of insects and of some endemic Hawaiian birds in the pollination of the tree, Metrosideros collina, 2) to test the hypothesis that maximal outbreeding and seed set occur at intermediate levels of nectar availability, 3) to understand the adaptive significance of profuse flowering in this species, and 4) to compare the pollination ecology of this species and the degree of specialization in the plant-pollinator community with those of similar mainland systems. Endemic Hawaiian birds (Drepanididae) are essential for high levels of fruit set and outbreeding in M. collina. Fruit set was much higher in redflowered individuals when birds were allowed to use inflorescences than when only insects used them. This is apparently caused by partial self-incompatibility, such that maximal fruit set occurs only with outbreeding, the primary agents of which are the birds. The predominant flower color in the population, the dimensions of floral parts, and copious nectar secretion adapt this species to bird pollination. However, insects effect moderate amounts of pollination and fruit set. The open flower and the color- and scent-variability within the population may be adaptations for insect pollination in the event that bird pollination fails. The population seems to have differentiated along an elevational gradient, with adaptations for bird pollination increasing proportionally with elevation. The generalization of the pollination strategy is on both an individual and populational basis, and enables M. collina to be the good colonizer that successional patterns and its own geographical distribution show it to be. Red-flowered individuals are partially self-compatible, but yellow-flowered individuals are totally self-compatible. The yellow-flowered morph may be evolving autogamy. At this stage its breeding system is intermediate between autogamy and outbreeding, with geitonogamy being encouraged due to 1) increased attractiveness to insects relative to the high energy birds by means of color and scent cues, and 2) increased nectar flow which satiates pollinators. Furthermore, yellow-flowered individuals have a floral structure that facilitates transferal of pollen to stigmas in the same inflorescence by means of small size pollinators, or even without the aid of a pollinator. A model is derived that predicts the selective results of various degrees of pollinator limitation on nectar productivity. Bird numbers are more constant relative to nectar availability than would be expected by random sorting, although temporary deficits and surpluses of these pollinators occur: their inability to respond instantly to changes in the intensity of bloom introduces lags into the system, and these have important consequences for pollination, outbreeding, and gene flow. During some times of the year pollinators are limiting to M. collina, and intraspecific competition occurs. Maximal fruit set and outbreeding do occur at intermediate nectar availabilities. Interspecific competition between species of trees for pollinators is a potential selective force that may explain the character displacement and staggering of flowering seasons of several tree species in the Hawaiian forests. Comparison with a similar but more diverse forest community in New Zealand and with mainland tropical forests suggests that the length of flowering season per tree species is inversely related to the number of tree species competing for pollinators. Profuse flowering in M. collina results in lowered fruit set per inflorescence because of decreased outbreeding, but the total number of fruits set per tree is probably high because of partial self-compatibility in most individuals. Thus, the M. collina system does not help explain profuse flowering in mainland tropical species that are totally self-incompatible. In comparison with mainland communities, the degree of specialization in the plant-pollinator relationship seems to be less in Hawaiian forests, although more information is needed on the Hawaiian lobelias and Sophora chrysophylla before such a statement should be made with any certainty.
Systematic patterns of foraging for nectar by Amakihi (Loxops virens)
(Island Ecosystems IRP, U.S. International Biological Program, 1975-07) Kamil, Alan C.
The visits of color-banded, territorial Amakihi to marked clusters of mamane bloom were recorded. The results indicated the existence of three systematic patterns in foraging behavior: (1) Amakihi avoid revisiting flower clusters they have recently visited; (2) when revisits do occur, they tend to be temporally spaced so as to allow time for nectar replenishment; (3) at least in some cases, the male and female Amakihi do not feed randomly with respect to each other, but rather a portion of the territory is utilized only by the female. Each of these characteristics of Amakihi foraging for nectar probably increases the foraging efficiency of the birds.
Seasonal abundances of the mamane moth, its nuclear polyhedrosis virus, and its parasites
(Island Ecosystems IRP, U.S. International Biological Program, 1975-05) Conant, Michael
The mamane moth (Uresephita polygonalis, Denis and Schiff.) is a serious pest of the mamane tree (Sophora chrysophylla, Salish.) on the island of Hawaii. The larvae of this moth feed on mamane leaflets sometimes causing serious defoliation. The life cycle and development of U. polygonalis were determined by observation of laboratory reared animals. Results of laboratory tests indicated that Acacia koa was not a host of the larvae. Seasonal abundance of the moth was estimated from monthly counts of eggs and larvae collected from four sampling sites. Apparently there are no positive correlations of population dynamics with rainfall, humidity, temperature and vegetative flushing of mamane. Four parasites were reared from U. polygonalis collected at the sampling site. Only one of these, an ichneumonid (Rorogenes blackburni, Cameron) appeared to be an important parasite, although it did not occur in high enough numbers to seriously affect mamane moth populations. The nuclear polyhedrosis virus, present only at sampling site 4, was a major factor in the regulation of the U. polygonalis population at that site. Laboratory tests indicated that larvae from all sites were highly susceptible to the virus. However, why the virus did not occur at all sites remains to be determined. Possibly the amount of sunlight and ultraviolet radiation reaching the trees and ground beneath them affects the virus which is inactivated by light. Thus, in years when U. polygonalis populations do not reach high levels, the virus is confined to cloud covered areas such as site 4. The virus disease plays a major role in population regulation when it reaches epizootic levels.
Pollination Energetics and Foraging Strategies in a Metrosideros-Honeycreeper Association
(Island Ecosystems IRP, U.S. International Biological Program, 1975-05) Carpenter, F Lynn; MacMillen, Richard E.
Evidence suggests that birds, specifically certain nectar-eating species of Drepanididae, are necessary for proper pollination, outcrossing, and high levels of seed set in Metrosideros collina on the island of Hawaii. Selection pressures acting upon the honeycreeper-M. collina association in the Keauhou Ranch forests on Mauna Loa fluctuate through the year. At the peak of the M. collina blooming season in late spring, nectar is relatively abundant to the honeycreepers, which means that the most important pollinators are potentially limiting to M. collina. Resultant intraspecific competition has selected for high rates of nectar secretion to attract pollinators; a ceiling in the population has been placed on these rates by the fact that seed set declines at very high flow rates, apparently because of decreased outcrossing due to pollinator sedentariness in such trees. At other times of the year when M. collina flowers are less abundant, a surplus of pollinators occurs, which results in high levels of seed set but in competition for food among the nectar-eaters. Loxops virens, and also Himatione sanguinea to a lesser extent, can turn to insectivory under these circumstances and therefore need not possess a highly specialized and efficient nectar-foraging strategy, but the more nectar-dependent Vestiaria sanguinea establishes interspecifically:defended territories in M. collina and visits flowers on its territory in a timed sequence. An hierarchy of interspecific aggression seems to be aligned among the three honeycreeper species in order of degree of nectar dependency.
The Island Ecosystems Data Bank
(Island Ecosystems IRP, U.S. International Biological Program, 1975-08) Bridges, Kent W.; Carey, G Virginia
The US/IBP Island Ecosystems IRP has made considerable use of computer processing for analyzing its data. As a result, many sets of machine readable records have been created. These include data, programs, and procedures for running the data with the programs. These items have been archived so that they may be used in the future. This report lists available items, describes the facilities used in processing the data, and discusses some of the aspects of operating a data bank.
User oriented statistical analysis programs: a brief guide
(Island Ecosystems IRP, U.S. International Biological Program, 1975-07) Bridges, Kent W.
A set of seven statistical analysis programs has been made available for use on an interactive mini-computer. These programs are designed for easy use and should allow novice computer users a convenient way to analyze their data. This report presents the information on how to gain access to the computer through a time-sharing terminal and how to use the programs. The programs perform the following types of analyses: simple sample descriptions and histogram plots, t-tests on unpaired samples, one-way analysis of variance, least squares regressions with linear and various equations for curves with plots of the fit equations, polynomial regressions, multiple regressions, and contingency tables.
Parasites of the Hawaii Amakihi (Loxops virens virens)
(Island Ecosystems IRP, U.S. International Biological Program, 1975-04) van Riper, Charles III
This paper deals with the parasites of the Hawaii Amakihi (Loxops virens virens). The study was undertaken in 1969 to determine the types of parasites present and what role they could play in regulating population numbers. Betaparasites and endoparasites were collected from as many birds as possible. A new form for the autopsy of passerine birds was developed and is included as Appendix I. Most of the external parasites I found on the Amakihi still await identification. The most commonly encountered feather louse seems to be Philopterus. Mites of the genus Rhinonyssus were found in the nasal passages of the Amakihi. I found Hippoboscid flies on Amakihi; blowflies often frequent active nests. Few parasites inhabit the nest of the Amakihi. Most of the nest fauna are nonparasitic arthropods associated with nests as saprophages or as predators on other nest arthropods. What appeared to be avian pox was found on five Amakihi. All were inflicted with the dry variety of pox. Four were infected on the head and one was infected on the leg. In two cases the infection was severe enough to cause death to the bird. Coccidia does not appear to be an important disease of wild Amakihi. I found two mild cases of this disease. In four of the 24 birds examined, smears of the heart, liver, spleen, and lungs were made; all were negative. I made 131 peripheral blood smears from 103 different Amakihi. Two slides were positive. Both contained protozoan parasites from the "Plasmodium relictum complex," being either F. giovannolai or P. matutinum. The birds that contracted malaria were caged individuals that had been moved from Puu Laau (7,500 feet) to Kamuela, Hawaii (2,500 feet). These birds died 32 and 45 days, respectively, after capture. Both exhibited ataxia, shivering, and weakness before death. Intestinal worms were the most frequently encountered parasites. Capillaria sp. ova were present in seven of the 24 fecal smears examined and adult worms were found in eight of the birds. Ova of Tetrameres sp. were detected in two individuals; adult worms were found in one bird. Tapeworm eggs were diagnosed in the feces of two birds. Both individuals contained large numbers of adults. Unidentified nematodes were found in three individuals. One bird had three worms in the left atrium and aorta. Puu Laau is a high, arid region. Many of the mosquito borne infections are not present there.
Temporal pattern of gene arrangement frequency in altitudinal populations of Drosophila immigrans on Mauna Loa, Hawaii
(Island Ecosystems IRP, U.S. International Biological Program, 1975-05) Paik, Y.K.; Sung, K.C.
The chromosomal polymorphism of D. immigrans from three different elevations on Mauna Loa, Hawaii was studied from 1971 to 1973. Three inverted gene arrangements in the second chromosome were common at most times of these years. The frequencies of arrangement A in the left arm of this chromosome showed a "spring-summer-winter" pattern of seasonal change in 1973 when collections were made almost monthly, but the other two arrangements in the right arm showed no perceptible changes. The A arrangement also showed significant altitudinal variation. in frequency in 1971. The frequency pattern of gene arrangement A was correlated with dry condition of the environment.
Integrated island ecosystem ecology in Hawaii: spatial distribution of island biota: introduction
(Island Ecosystems IRP, U.S. International Biological Program, 1975-06) Mueller-Dombois, Dieter; Bridges, Kent W.
This portion of the synthesis volume is being reproduced now because it is important that all synthesis volume contributors have these basic materials as they prepare their own contributions. Chapter 6, the subject matter of this report, follows the first five chapters that were contained in Technical Report 54 (Introductory Survey, Part I of the Synthesis Volume). It is expected that there will be some reworking of sections of this report as more data are analyzed and the integration needs of the entire synthesis part become more apparent. Chapter 6 is only the introduction to Part II. Additional material will include the altitudinal distribution of individual organism groups (Chap. 7) and their spatial integration along the Mauna Loa Transect (Chap. 8). This report is a draft manuscript of the introduction to Part II of our proposed synthesis volume. As chapter 6, this introduction forms the continuation to Part I in Technical Report 54, which contains the first five chapters. Part II, which is concerned with an analysis of the spatial distribution of island biota in an ecosystem context, is one of four major synthesis areas of the Island Ecosystems Integrated Research Program of the US/IBP. This report gives the conceptual and physical background to our distribution analysis along a transect on a high-island mountain, Mauna Loa. It further discusses current ecological theory of species distributions along environmental gradients and presents examples together with the analytical techniques that we adopted for all organism groups investigated in spatially integrated community samples.
Climate data for the IBP sites on Mauna Loa, Hawaii
(Island Ecosystems IRP, U.S. International Biological Program, 1975-08) Bridges, Kent W.; Carey, G Virginia
This report contains the fourth set of Tables for the climatic data collected at the US/IBP Island Ecosystems IRP field sites on Mauna Loa, Hawaii. These data were collected during the first six months of 1975 using the same procedures as employed for the data collection reported in Technical Report No. 22, Supplement 1, Technical Report No. 38, and Technical Report No. 59.The figures are not included in this report because the data are for only the first half of the year. No attempt has been made in this report to interpret the Tables that are presented.
Evolution of the endemic Hawaiian cerambycid-beetles
(Island Ecosystems IRP, U.S. International Biological Program, 1975-08) Gressitt, J.L.
The Hawaiian cerambycid fauna presents one of the remarkable examples of great proliferation of species and form with adaptive radiation from a single ancestor. This fauna consists of over 135 species of plagithmysines (Clytini) plus one endemic species each of two somewhat widespread primitive genera (Megopis and Parandra) which breed primarily in rotten wood. There are also 17 species introduced by man. The plagithmysines evolved from a single immigrant ancestor species which probably came from Mexico or southern California a few million years ago. The group feeds in living trees and has remarkably specific host-tree food habits within a species of beetle, but has adapted to a large number of families (27) of trees for the whole group. Body form has diverged greatly, with the result that seven generic names have been proposed to accommodate the species. However, these have recently been reduced to the single genus Plagithmysus, with 5 subgenera, following the discovery of intermediate forms.
Cytogenetics of the Hawaiian Telmatogeton (Diptera)
(Island Ecosystems IRP, U.S. International Biological Program, 1975-03) Newman, Lester J.
A cytogenetic analysis of the marine and freshwater species of the Hawaiian Telmatogeton was conducted to determine their evolutionary relationships. Telmatogeton torrenticola, which occupies the islands of Molokai, Maui, and Hawaii, is divided into separate species based on differences in chromosome number, karyotype, and fixed inversions. T. torrenticola-Molokai (n = 4) has three pairs of metacentric chromosomes and a pair of dot chromosomes. T. torrenticola-Hawaii (n = 3/4) has a differentiated sex chromosome system. Males have two pairs of metacentric autosomes, two acrocentric X-chromosomes, and a metacentric Y-chromosome. Females have two pairs of metacentric autosomes and two pairs of acrocentric X-chromosomes. T. torrenticola from East Maui and from West Maui have the same karyotype of six pairs of acrocentric and a pair of dot chromosomes, but they differ from each other by at least six fixed inversions. In a model of the evolution of the Hawaiian Telmatogeton it is proposed that a marine form was ancestral to the freshwater species. The model also proposes that one of these species may have returned to the sea. Chromosome evolution has involved reduction in chromosome number by centric fusion and the fixation of inversions. T. fluviatilis (n = 7) on Oahu is proposed to be the ancestral freshwater species. It gave rise to T. abnormis (n = 4) and T. hirtus (n = 3/4) on Kauai. T. fluviatilis also gave rise to T. torrenticola-Maui, the ancestor of T. torrenticola-Molokai (n = 4), T. torrenticola-E. Maui (n = 7), and T. torrenticola-Hawaii (n = 3/4). It is suggested that the marine species T. pacificus (n = 4) may have returned to the sea with T. torrenticola-Molokai as its ancestor since the two have identical karyotypes.
Threshold model of feeding territoriality: a test with an Hawaiian honeycreeper
(Island Ecosystems IRP, U.S. International Biological Program, 1975-04) Carpenter, F.L.; MacMillen, R.E.
A cost-benefit model predicting threshold requirements for establishment of feeding territories in nectarivorous birds was tested by comparing 24-hour energy production of nectar with energy budgets in territorial and nonterritorial Hawaiian honeycreepers. The test was positive and demonstrated that nectar productivity was more important in determining establishment of territories than intrusion pressure and floral dispersion.
Integrated island ecosystem ecology in Hawaii: introductory survey
(Island Ecosystems IRP, U.S. International Biological Program, 1975-02) Mueller-Dombois, Dieter
This report is a draft manuscript of Part I (Introductory Survey) of the synthesis volume of the US/IBP Island Ecosystems Stability and Evolution Subprogram. Further parts to follow are: Part II Spatial Distribution of Island Biota; Part III Temporal Relationships of Island Biota; Part IV Community Structure and Niche Differentiation; Part V Genetic variation within Island Species; Part VI General Conclusions.
Soil algal relationships to Onychiurus folsomi, a minute arthropod
(Island Ecosystems IRP, U.S. International Biological Program, 1975-03) McGurk, Linda-Lee
Elucidation of the roles of the soil algae is a natural goal of the algal component study of the Island Ecosystems IRP. Thus, the present study investigated the possibility that the algae could serve as food for the 5 to 6 mm long insect, Onychiurus folsomii, the most ubiquitous of the soil arthropods in the Hawaii Volcanoes National Park. This goal was sought through a series of laboratory experiments. In some algae grown on Bold's basal medium solidified with agar it was found that 15 species of algae in unialgal culture were ingested. Some of the algal species grew after transiting the insects and perhaps their progeny were recycled. While no algae were seen in the gut of any wild insect, the algae and insects occur together. It was found, however, that in the laboratory the insects grew and reproduced more when algae were present than when on agar medium alone. Using carbon-14 marked algal cells it was demonstrated that labeled material from the algae passed out into the insect tissues increasingly with time as the cells passed through insect gut. It is concluded such soil insects as Onychiurus can grow and reproduce on algae found in their environment and may do so in nature.
Acari on murine rodents along an altitudinal transect on Mauna Loa, Hawaii
(Island Ecosystems IRP, U.S. International Biological Program, 1975-04) Radovsky, Frank J.; Tenorio, JoAnn M.; Tomich, P. Quentin; Jacobi, James D.
Rodents were trapped seasonally during a two-year period at 14 primary sites from 840 to 2440 m on a transect (with additional collecting to 2895 m), and intensively collected in the Kilauea Forest near the transect. The sampled habitats were not in the proximity of human habitations. Three of the four murine species present in the Hawaiian Archipelago were taken: Mus musculus, Rattus rattus, and R. exulans. Ectoparasites were recovered from rodents by a standardized washing technique. Mammalogical and parasitological data were analyzed by computer. The occurrence, host associations, and spatial distribution of some Acari are treated here. Significant correlations, including those for some permanently parasitic mites, are partially independent of host factors and are associated with local differences in climate.

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