Exoskeletal cuticle of cavernicolous and epigean terrestrial isopods: A review and perspectives

Comparative ultrastructural studies of the integument in terrestrial isopod crustaceans show that specific environmental adaptations of different eco-morphotypes are reflected in cuticle structure. The biphasic molting in isopods is a valuable experimental model for studies of cuticular matrix secretion and degradation in the same animal. The aim of this review is to show structural and functional adaptations of the tergal cuticle in terrestrial isopods inhabiting cave habitats. Exoskeletal cuticle thickness, the number of cuticular layers, epicuticle structure, mineralization, pigmentation and complexity of sensory structures are compared, with greater focus on the well-studied cave trichoniscid Titanethes albus. A large number of thinner cuticular layers in cave isopods compared to fewer thicker cuticular layers in related epigean species of similar body-sizes is explained as a specific adaptation to the cavernicolous life style. The epicuticle structure and composition are compared in relation to their potential waterproofing capacity in different environments. Cuticle mineralization is described from the functional point of view as well as from the aspect of different calcium storage sites and calcium dynamics during the molt cycle. We also discuss the nature and reduction of pigmentation in the cave environment and outline perspectives for future research.     

Spatial distribution of calcite and amorphous calcium carbonate in the cuticle of the terrestrial crustaceans Porcellio scaber and Armadillidium vulgare

The crustacean cuticle is an interesting model to study the properties of mineralized bio-composites. The cuticle consists of an organic matrix composed of chitin–protein fibres associated with various amounts of crystalline and amorphous calcium carbonate. It is thought that in isopods the relative amounts of these mineral polymorphs depend on its function and the habitat of the animal. In addition to the composition, the distribution of the various components should affect the properties of the cuticle. However, the spatial distribution of calcium carbonate polymorphs within the crustacean cuticle is unknown. Therefore, we analyzed the mineralized cuticles of the terrestrial isopods Armadillidium vulgare and Porcellio scaber using scanning electron-microscopy, electron probe microanalysis and confocal μ-Raman spectroscopic imaging. We show for the first time that the mineral phases are arranged in distinct layers. Calcite is restricted to the outer layer of the cuticle that corresponds to the exocuticle. Amorphous calcium carbonate is located within the endocuticle that lies below the exocuticle. Within both layers mineral is arranged in rows of granules with diameters of about 20 nm. The results suggest functional implications of mineral distribution that accord to the moulting and escape behaviour of the animals.     

Cuticle differentiation in the embryo of the amphipod crustacean Parhyale hawaiensis

The arthropod cuticle is a multilayered extracellular matrix produced by the epidermis during embryogenesis and moulting. Molecularly and histologically, cuticle differentiation has been extensively investigated in the embryo of the insect Drosophila melanogaster. To learn about the evolution of cuticle differentiation, we have studied the histology of cuticle differentiation during embryogenesis of the amphipod crustacean Parhyale hawaiensis, which had a common ancestor with Drosophila about 510 million years ago. The establishment of the layers of the Parhyale juvenile cuticle is largely governed by mechanisms observed in Drosophila, e.g. as in Drosophila, the synthesis and arrangement of chitin in the inner procuticle are separate processes. A major difference between the cuticle of Parhyale and Drosophila concerns the restructuring of the Parhyale dorsal epicuticle after deposition. In contrast to the uniform cuticle of the Drosophila larva, the Parhyale cuticle is subdivided into two regions, the ventral and the dorsal cuticles. Remarkably, the boundary between the ventral and dorsal cuticles is sharp suggesting active extracellular regionalisation. The present analysis of Parhyale cuticle differentiation should allow the characterisation of the cuticle-producing and -organising factors of Parhyale (by comparison with the branchiopod crustacean Daphnia pulex) in order to contribute to the elucidation of fundamental questions relevant to extracellular matrix organisation and differentiation. This work was supported by the German Research Foundation (DFG, grant number MO 1714/1-1).     

Pathogenic effect of entomopathogenic nematode-bacterium complexes on terrestrial isopods

In this study, we evaluated the effect of entomopathogenic nematodes (EPNs) Steinernema carpocapsae, Steinernema feltiae and Heterorhabditis bacteriophora, symbiotically associated with bacteria of the genera Xenorhabdus or Photorhabdus, on the survival of eight terrestrial isopod species. The EPN species S. carpocapsae and H. bacteriophora reduced the survival of six isopod species while S. feltiae reduced survival for two species. Two terrestrial isopod species tested (Armadillidium vulgare and Armadillo officinalis) were found not to be affected by treatment with EPNs while the six other isopod species showed survival reduction with at least one EPN species. By using aposymbiotic S. carpocapsae (i.e. without Xenorhabdus symbionts), we showed that nematodes can be isopod pathogens on their own. Nevertheless, symbiotic nematodes were more pathogenic for isopods than aposymbiotic ones showing that bacteria acted synergistically with their nematodes to kill isopods. By direct injection of entomopathogenic bacteria into isopod hemolymph, we showed that bacteria had a pathogenic effect on terrestrial isopods even if they appeared unable to multiply within isopod hemolymphs. A developmental study of EPNs in isopods showed that two of them (S. carpocapsae and H. bacteriophora) were able to develop while S. feltiae could not. No EPN species were able to produce offspring emerging from isopods. We conclude that EPN and their bacteria can be pathogens for terrestrial isopods but that such hosts represent a reproductive dead-end for them. Thus, terrestrial isopods appear not to be alternative hosts for EPN populations maintained in the absence of insects.     

The influence of habitat heterogeneity on host-pathogen population dynamics

The influence of spatial heterogeneity on the population dynamics of a naturally occurring invertebrate host-pathogen system was experimentally investigated. At ten week intervals over a two year period, I quantified the spatial distribution of natural populations of the terrestrial isopod crustacean Porcellio scaber infected with the isopod iridescent virus (IIV). During the seasonally dry periods of summer and early fall in central California, isopod populations were highly aggregated and the degree of patchiness and distance between inhabited patches was greatest. Coincident with increased patchiness and patch spacing was an increase in isopod density within patches. During the wet seasons of winter and spring, isopod population patchiness, inter-patch spacing, and within-patch density was low. Seasonal changes in virus prevalence were negatively correlated with within-patch density, patchiness, and inter-patch spacing. The influence of the spatial distribution of isopods on virus prevalence was also tested in field experiments. The virus was introduced into arrays of artificial habitat patches colonized by isopods in which interpatch distance was varied. The prevalence of resulting infections was monitored at weekly intervals. In addition, dispersal rates between artificial patches and natural patches were quantified and compared. The results showed that isopods in treatments with the smallest inter-patch spacing had the highest virus prevalence, with generally lower prevalence among isopods in more widely spaced patches. The spacing of experimental patches significantly affected virus prevalence, although the experiments did not resolve a clear relationship between patch spacing and virus prevalence. Rates of dispersal between patches decreased with increased patch spacing, and these rates did not differ significantly from dispersal between natural patches. The results suggest that rates of dispersal between isopod subpopulations may be an important component of the infection dynamics in this system. I discuss the consequences of these findings for host-pathogen dynamics in fragmented habitats, and for other ecological interactions in spatially heterogeneous habitats.     

Distribution,density, and habitat use among native and introduced populations of the Australasian burrowing isopod <Emphasis Type="Italic">Sphaeroma quoianum</Emphasis>

The Australasian burrowing isopod (Sphaeroma quoianum) has been introduced to numerous embayments along the Pacific coast of North America. In some bays, populations of S. quoianum can exceed tens of thousands of individuals m⁻³ and bioturbation by the isopods can exacerbate shoreline erosion. Within their native range, however, studies recognize S. quoianum primarily as a woodborer. We measured the distribution, prevalence, habitat use, density, and associated fauna of S. quoianum in two bays within the native range [Tamar estuary (Tamar), Tasmania and Port Phillip Bay (PPB), VIC, Australia] and in one bay where the isopods had been introduced (Coos Bay, OR, USA). Distribution, prevalence, and habitat use were determined from shoreline surveys. Densities and the associated fauna of S. quoianum were measured in three intertidal substrata (marsh bank, wood, and friable rock). In all embayments, S. quoianum occurred primarily between 5 and 30 salinity and 55−68% of sites harbored isopods. Habitat use varied between embayments. Distributional patterns suggest salinity is the primary factor that limits the establishment and spread of S. quoianum. Isopod densities in all substrata were greater in Coos Bay than in the Tamar or PPB, although only densities within marsh banks varied significantly. Similarities in the amount of habitat and food, and the burrow dwelling lifestyle of S. quoianum suggest habitat availability/quality, food levels, predation, and competition are not responsible for the large differences in density. Lack of parasites or disease in populations of S. quoianum introduced to Coos Bay could be responsible for the prolific densities observed.     

Amorphous and crystalline calcium carbonate distribution in the tergite cuticle of moulting Porcellio scaber (Isopoda, Crustacea)

The main mineral components of the isopod cuticle consists of crystalline magnesium calcite and amorphous calcium carbonate. During moulting isopods moult first the posterior and then the anterior half of the body. In terrestrial species calcium carbonate is subject to resorption, storage and recycling in order to retain significant fractions of the mineral during the moulting cycle. We used synchrotron X-ray powder diffraction, elemental analysis and Raman spectroscopy to quantify the ACC/calcite ratio, the mineral phase distribution and the composition within the anterior and posterior tergite cuticle during eight different stages of the moulting cycle of Porcellio scaber. The results show that most of the amorphous calcium carbonate (ACC) is resorbed from the cuticle, whereas calcite remains in the old cuticle and is shed during moulting. During premoult resorption of ACC from the posterior cuticle is accompanied by an increase within the anterior tergites, and mineralization of the new posterior cuticle by resorption of mineral from the anterior cuticle. This suggests that one reason for using ACC in cuticle mineralization is to facilitate resorption and recycling of cuticular calcium carbonate. Furthermore we show that ACC precedes the formation of calcite in distal layers of the tergite cuticle.     

Biochemical composition of the isopod Mesidotea entomon (Linnaeus) from the western Arctic

Summary Changes in biochemical composition were determined for the isopod Mesidotea entomon (Linnaeus) from two differing ecological habitats in the western Arctic. Isopods from Pauline Cove, Yukon Territory, showed quantitative differences in total lipid and carbohydrate compared with those of similar body size from Dolomite Lake, Northwest Territories. Total chitin, ash, and protein were similar in isopods from both sites. A seasonal accumulation of protein and lipid was observed in isopods from Dolomite Lake prior to the onset of winter. The roles of ecological habitat, season, parasitism, nutritional and reproductive cycles in the organic composition of M. entomon are discussed.     

Role of cuticular lipids and water-soluble compounds in tick susceptibility to Metarhizium infection

The arthropod cuticle acts as a physiochemical barrier protecting the organism from pathogens' entry. Entomopathogenic fungi actively penetrate the cuticles of arthropod hosts and are therefore directly affected by cuticle composition. Previously we have observed that Metarhizium spp. developing on resistant ticks ultimately die without penetrating tick's cuticle, suggesting that the cuticles of resistant ticks have antifungal compounds. In the present study, lipids and water-soluble cuticular components were extracted from engorged female tick cuticles, of one susceptible and one resistant tick species to Metarhizium spp. While conidia exposed to lipids from the susceptible tick, Rhipicephalus annulatus, germinated and differentiated into appressorium, conidia exposed to lipids from the resistant tick, Hyalomma excavatum, were inhibited. Soluble cuticular component extracts from both susceptible and resistant ticks stimulated conidial germination but not appressorium differentiation. A comparative analysis of the fatty acid profile in lipid extract of each tick exhibited similar compositions, but the relative abundance of C16:0, C18:0, C18:1ω9C and C20:0 was 2–5 times higher in the extracts from resistant ticks. All of these fatty acids inhibited conidial germination in vitro at 1% and 0.1% w/v concentration, but C20:0 stimulated appressorium differentiation at low concentration. This is the first report demonstrating a possible link between the presence of antifungal compounds in a specific concentration in tick cuticle and tick resistance to infection.     

The cuticle of the enigmatic arthropod Phytophilaspis and biomineralization in Cambrian arthropods

Lin, J.‐P., Ivantsov, A.Y. & Briggs, D.E.G. 2011: The cuticle of the enigmatic arthropod Phytophilaspis and biomineralization in Cambrian arthropods. Lethaia, Vol. 44, pp. 344–349. Many non‐trilobite arthropods occur in Cambrian Burgess Shale‐type (BST) biotas, but most of these are preserved in fine‐grained siliciclastics. Only one important occurrence of Cambrian non‐trilobite arthropods, the Sinsk biota (lower Sinsk Formation, Botomian) from the Siberian Platform, has been discovered in carbonates. The chemical compositions of samples of the enigmatic arthropod Phytophilaspis pergamena Ivantsov, 1999 and the co‐occurring trilobite Jakutus primigenius Ivantsov in Ponomarenko, 2005 from this deposit were analysed. The cuticle of P. pergamena is composed of mainly calcium phosphate and differs from the cuticle of J. primigenius, which contains only calcium carbonate. Phosphatized cuticles are rare among large Cambrian arthropods, except for aglaspidids and a few trilobites. Based on recent phylogenetic studies, phosphatization of arthropod cuticle is likely to have evolved several times. □arthropod cuticle, Burgess Shale‐type preservation, fossil‐diagenesis, phosphatization.     

A study of the endogenous activity rhythms of the marine isopod Exosphaeroma truncatitelson

The endogenous activity rhythms of the South African endemic isopod Exosphaeroma truncatitelson were explored. Isopods were collected on days coinciding with either a neap or spring tide. High- and low-energy beaches, which vary in the intensity of wave action, were selected as study sites. Isopods, in groups of 10–30, were placed in glass tanks with different experimental conditions: darkness + no sand, darkness with sand and natural ambient light (no sand). Activity of the isopods was measured for 50 h by recording the number of isopods swimming actively during a 30-s period. These isopods exhibited an endogenous activity rhythm which coincided with the tidal cycle even though removed from their natural habitat. Cosine curves were fitted to the observed activity data and statistical differences across variables were investigated. Peak activity was shown to occur shortly after high tide. Similar activity cycles were observed for both neap and spring tides and for different times of the year. Results also indicated that the activity cycles at both low- and high-energy beaches were similar. Two survival advantages of this rhythm are proposed: prevention of stranding on high shores and competitive avoidance with other local isopod species.     

Comparative response of Metarhizium brunneum to the cuticles of susceptible and resistant hosts

Earlier studies demonstrated that Metarhizium brunneum, usually a broad-host pathogen of arthropods, is unable to complete its pathogenic life cycle when inoculated on the fungus-resistant tick, Hyalomma excavatum engorged females. While the fungus penetrates the cuticle of fungus-susceptible tick, Rhipicephalus annulatus females, it is unable to penetrate the cuticle of fungus-resistant tick, and even perishes on its surface. This is probably due to high concentration of antifungal fatty acids and probably also due to a hypersensitive-like response of the tick. To understand the metabolic pathways occurring in the fungal hyphae upon encountering the cuticles, we compared the response of the fungus to cuticle from susceptible and resistant tick cuticles by 2D-gels. The intracellular proteomes of M. brunneum Mb7 exposed to cuticle of the fungus-susceptible tick, R. annulatus, and to the fungus-resistant tick, H. excavatum engorged females were compared after exposure to either cuticles. By means of liquid chromatography-mass spectrometry/mass spectrometry we identified in both proteomes common proteins involved in biological processes as well as unique proteins identified only in the proteome of fungus exposed to fungus-resistant tick cuticle. These proteins were identified in high probability as heat shock proteins, four key enzymes of the glyoxylate cycle, and proteins associated with hypoxia, and exposure to antifungal drugs. These findings are discussed within the M. brunneum-tick pathosystem in relation to tick resistance and host resistance in general.     

Response of nematode-trapping fungi to organic substrates in a coastal grassland soil

To understand why Arthrobotrys oligospora and other nematode-trapping fungi are common and sometimes abundant in the coastal grassland soils of the Bodega Marine Reserve (BMR, Sonoma County, CA), we examined how resident trapping fungi responded to the addition of eight organic substrates (lupine leaves, grass leaves, dead isopods, dead moth larvae, isopod faeces, deer faeces, shrimp shells, and powdered chitin). We were especially interested in the effects of dead isopods because isopods are abundant at BMR and because previous studies had documented strong responses of A. oligospora to other arthropods (dead moth larvae). Soil from BMR was packed into vials (40 g dry mass equivalent per vial with water potential at −230 kPa and bulk density at 0.9 g cm⁻³), and one substrate or no substrate was added to the soil surface. After 30 d at 20 °C, trapping fungi were quantified by dilution plating and most probable number procedures. The response of A. oligospora was inversely related to substrate carbon:nitrogen (C:N) ratio: substrates with low C:N ratios (dead isopods, lupine leaves, dead moth larvae) usually caused large increases in A. oligospora whereas those with higher C:N ratios (isopod faeces, deer faeces, grass leaves) did not. An exception was chitin powder, which had a low C:N ratio, but which did not cause A. oligospora to proliferate. Responses of A. oligospora were directly related to the quantity of nitrogen added with each substrate, and those substrates that caused large increases in resident nematodes usually caused large increases in A. oligospora. Other trapping fungi did not respond as strongly as A. oligospora.     

Ultrastructure and cuticle formation of the carapace in the myodocopan ostracod exemplified by Euphilomedes japonica (Crustacea: Ostracoda)

The ultrastructure and formation of the cuticle of a myodocopan ostracod, Euphilomedes japonica, are investigated utilizing scanning and transmission electron microscopy. The outer lamella cuticle consists of four layers; epicuticle, exocuticle, endocuticle, and membranous layer like in the cuticle of other arthropods. The exocuticle and endocuticle are well-calcified and the organic matrix develops within the both cuticles. The outermost layer of new cuticle (epicuticle) is secreted first and the inner layers (exocuticle, endocuticle and membranous layer) are added proximally in the pre-, and postmoult stages. The calcification takes place in the whole area of carapace at the same time together with the synthesis of organic matrix within the endocuticle. This study demonstrates that the ultrastructure and formation of the cuticle in myodocopans are different from those in podocopans, and that the myodocopan carapaces have achieved a structural diversity for adaptation to different lifestyles.     

Five in situ observations of live oarfish Regalecus glesne (Regalecidae) by remotely operated vehicles in the oceanic waters of the northern Gulf of Mexico

As part of the SERPENT Project, five observations of apparently healthy oarfish Regalecus glesne by remotely operated vehicles are reported from the northern Gulf of Mexico. Regalecus glesne were observed between 2008 and 2011 at depths from within the epipelagic and mesopelagic zones. These observations include the deepest verified record of R. glesne (463–492 m) and the first record of an arthropod ectoparasite (isopod).     

Gas permeability of plant cuticles

Cuticles from the adaxial surface of Citrus aurantium L. leaves and from the pericarp of Lycopersicon esculentum L. and Capsicum annuum L. were isolated enzymatically and their oxygen permeability was determined. Isolated cuticles were mounted between a gaseous and an aqueous compartment with the physiological outer side of the membrane facing the gaseous compartment. Permeability for oxygen was characterized by permeability (P) and diffusion (D) coefficients. P and D were independent of the driving force (gradient of oxygen concentration) across the cuticle, thus, Henry's law was obeyed. P values for the diffusion of oxygen varied between 3·10^-7 (Citrus), 1.4·10^-6 (Capsicum), and 1.1·10^-6 (Lycopersicon) m·s^-1. Extraction of soluble lipids from the cuticles increased the permeability. By treating the cutin matrix and the soluble lipids as resistances in series, it could be demonstrated that the soluble lipids were the main resistance for oxygen permeability in Citrus cuticles. However, in Lycopersicon and Capsicum, both the cutin matrix and the soluble lipids determined the total resistance. P values were not affected by either the proton concentration (pH 3–9) or the cations (Na⁺, Ca²⁺) present at the morphological inner side of the cuticles. It is concluded that the water content of cuticles does not affect the permeability properties for oxygen. Partition coefficients indicated a high solubility of oxygen in the cuticle of Citrus. The data suggest a solubility process in the cuticle of Citrus with respect to oxygen permeation.Abbreviations CM cuticular membrane - MX cutin polymer matrix - SCL soluble cuticular lipids     

Density dependent grazing effects of the isopod Idotea baltica Pallas on Fucus vesiculosus L in the Baltic Sea

In the Baltic Sea, abiotic factors are often supposed to explain the distribution of the key species Fucus vesiculosus. Still, in many areas, decline of F. vesiculosus has coincided with mass occurrence of the herbivorous isopod Idotea baltica. The aim of this work was to examine whether, how and at what densities I. baltica can affect the distribution of F. vesiculosus in the central Baltic proper. Both large-scale field surveys and a two-week grazing experiment have been performed.In the field survey there was a correlation between density of I. baltica and reduction in depth penetration of F. vesiculosus. At 80 animals per 100 g F. vesiculosus wet weight, the depth penetration of the F. vesiculosus belt was reduced by 2.5 m within a year. In the grazing experiment there was a correlation between density of I. baltica and loss of F. vesiculosus biomass and meristems. In the controls biomass and number of meristems increased by 50%, while at 20 animals per 100 g of F. vesiculosus there was no net growth of F. vesiculosus. Intensity of grazing did not differ between isopod densities of 20, 40 and 60/100 g. At isopod densities of 80 and 100/100 g though, biomass and meristems decreased by 50%, indicating a threshold for the survival of F. vesiculosus in the experiment. At all densities the isopods preferred younger tissue to older.Our results indicate that grazing by Idotea baltica is an important structuring factor in the Baltic Fucus vesiculosus populations.     

The role of controlled propagation on an endangered species: demographic effects of habitat heterogeneity among captive and native populations of the socorro isopod (Crustacea: Flabellifera)

The Socorro isopod, Thermosphaeroma thermophilum, became the focus of a novel conservation strategy when apparent extirpation of the species from the wild prompted construction of the Socorro Isopod Propagation Facility (SIPF) near the native spring in Socorro, New Mexico, USA. We subjected captive populations to 4 habitat treatments over 2 consecutive 50-month experimental trials. Native populations of T. thermophilum remained significantly more abundant than captive subpopulations during the 100-month study. Population trends among artificial habitats differed slightly between the trials, but the magnitude of this difference was independent of linear flow within either pool series, suggesting that captive isopods responded to habitat quality. Isopod age structure was most heterogeneous in pools with plants suggesting that vertical structural diversity is necessary to maintain long-term viability of T. thermophilum in captivity. Observations of `breeding huddles' in the SIPF pools implied that controlled propagation may affect the social basis of the species' mating system. This could be a behavioral response to selection on body size that may reduce predation risks from larger male cannibals, or to increase fitness of both sexes under spatially altered sex ratios of artificial environments. Our results accentuate the need for continued research on species-specific management practices for geographically restricted invertebrate taxa such as the endangered Socorro isopod.     

Structure and Evolution of the Atypical Mitochondrial Genome of <Emphasis Type="Italic">Armadillidium vulgare</Emphasis> (Isopoda,Crustacea)

The crustacean isopod Armadillidium vulgare is characterized by an unusual ∼42-kb-long mitochondrial genome consisting of two molecules co-occurring in mitochondria: a circular ∼28-kb dimer formed by two ∼14-kb monomers fused in opposite polarities and a linear ∼14-kb monomer. Here we determined the nucleotide sequence of the fundamental monomeric unit of A. vulgare mitochondrial genome, to gain new insight into its structure and evolution. Our results suggest that the junction zone between monomers of the dimer structure is located in or near the control region. Direct sequencing indicated that the nucleotide sequences of the different monomer units are virtually identical. This suggests that gene conversion and/or replication processes play an important role in shaping nucleotide sequence variation in this mitochondrial genome. The only heteroplasmic site we identified predicts an alloacceptor tRNA change from tRNA^Ala to tRNA^Val. Therefore, in A. vulgare, tRNA^Ala and tRNA^Val are found at the same locus in different monomers, ensuring that both tRNAs are present in mitochondria. The presence of this heteroplasmic site in all sequenced individuals suggests that the polymorphism is selectively maintained, probably because of the necessity of both tRNAs for maintaining proper mitochondrial functions. Thus, our results provide empirical evidence for the tRNA gene recruitment model of tRNA evolution. Moreover, interspecific comparisons showed that the A. vulgare mitochondrial gene order is highly derived compared to the putative ancestral arthropod type. By contrast, an overall high conservation of mitochondrial gene order is observed within crustacean isopods.     

Drift and upstream movement of invertebrates in a springbrook community ecosystem

Structure, drift, and upstream movement of populations of benthic macroinvertebrates, in particular Synurella dentata Hubricht and Lirceus fontinalis Raf., were examined within a temperate spring ecosystem. Chemical and physical aspects of the springbrook were also investigated and life histories of the gammarids and asellids noted.Chemically and physically the spring proved both constant and predictable, much more so than other lotic systems.Species diversity was low from November through February and increased in March, April, and May. Equitability followed the same trends as species diversity. Both indices were most affected by large fluctuations in the populations of aquatic insects.Significant changes in the numbers of amphipods, isopods, and total macroinvertebrates was evident over a seven month period. Males were present in the isopod population year-round, but only from November to January in the amphipod population. Breeding by the isopods occurred throughout the year and peaked during winter. Amphipods copulated only in the late fall and early winter.Significant diel peaks in the amphipod, isopod, and total invertebrate drift negatively correlated with light intensity levels. Amphipods and isopods did not exhibit any preferential upstream movement during either the day or night; however, total macroinvertebrate upstream movement was greater at night. The total number of invertebrates moving upstream were lower than values reported from other lotic environments.