Using temporary emigration to inform movement behaviour of cave‐dwelling invertebrates: a case study of a cave harvestman species

1. Understanding of animal movement behaviour is critical for developing appropriate conservation strategies because of its profound implications for predicting species' responses to perturbations. To date there is a substantial knowledge gap of the movement behaviour of subterranean invertebrates. 2. In this study temporary emigration (TE), the probability that an individual is absent from the cave on a given sampling occasion, was used as a method to inform on the movement and behaviour of cave‐dwelling invertebrates. Because these animals are difficult to study with traditional tracking techniques, the capture–recapture (CR) modelling framework was used to assess TE and to account for imperfect detectability and unobservable states. 3. Specifically, the influence of season, sex, and surface weather variables on the TE of the facultative cave‐dwelling harvestman species, Paranemastoma sillii sillii (Herman, 1871), was investigated. Multistate CR models were used while accounting for individual heterogeneity on 999 individually marked adults of two populations inhabiting two caves, located in southwestern Romania. 4. Harvestmen demonstrated clear heterogeneity and seasonality in TE. Findings showed that the TE of harvestmen exhibited variation between caves and there was little support for surface weather variables influencing TE and for sex‐specific TE patterns. 5. These results show that the study of TE patterns is useful to gain valuable insights into movement and intrinsic behavioural processes of cave‐dwelling harvestmen. The method could potentially be used for other invertebrate groups with similar movement characteristics and when traditional tracking techniques are difficult.     

High levels of fluctuating asymmetry in isolated stickleback populations

ABSTRACT: BACKGROUND: Fluctuating asymmetry (FA), defined as small random deviations from the ideal bilateral symmetry, has been hypothesized to increase in response to both genetic and environmental stress experienced by a population. We compared levels of FA in 12 bilateral meristic traits (viz. lateral-line system neuromasts and lateral plates), and heterozygosity in 23 microsatellite loci, among four marine (high piscine predation risk) and four pond (zero piscine predation risk) populations of nine-spined sticklebacks (Pungitius pungitius). RESULTS: Pond sticklebacks had on average three times higher levels of FA than marine fish and this difference was highly significant. Heterozygosity in microsatellite markers was on average two times lower in pond (HE [almost equal to] 0.3) than in marine (HE [almost equal to] 0.6) populations, and levels of FA and heterozygosity were negatively correlated across populations. However, after controlling for habitat effect on heterozygosity, levels of FA and heterozygosity were uncorrelated. CONCLUSIONS: The fact that levels of FA in traits likely to be important in the context of predator evasion were elevated in ponds compared to marine populations suggests that relaxed selection for homeostasis in ponds lacking predatory fish may be responsible for the observed habitat difference in levels of FA. This inference also aligns with the observation that the levels of genetic variability across the populations did not explain population differences in levels of FA after correcting for habitat effect. Hence, while differences in strength of selection, rather than in the degree of genetic stress could be argued to explain habitat differences in levels of FA, the hypothesis that increased FA in ponds is caused by genetic stress cannot be rejected.     

The mid-latitude biodiversity ridge in terrestrial cave fauna

The world's obligate cave-dwelling fauna holds considerable promise for biogeographic analysis because it represents a large number of independent evolutionary experiments in isolation in caves and adaptation to subterranean life. We focus on seven north temperate regions of at least 2000 km², utilizing more than 4300 records of obligate cave-dwelling terrestrial invertebrates. In North America, highest diversity was found in northeast Alabama while in Europe highest diversity was found in Ariège, France, and in southeast Slovenia. Based on these regions as well as more qualitative data from 16 other regions, we hypothesize that a ridge (ca 42°–46° in Europe and 34° in North America) of high biodiversity occurs in temperate areas of high productivity and cave density. This may reflect a strong dependence of cave communities on long term surface productivity (as reflected in actual evapotranspiration), because the subterranean fauna relies almost entirely on resources produced outside caves. This dependence may explain the unique biodiversity pattern of terrestrial cave invertebrates.     

Effect of directional selection for body size on fluctuating asymmetry in certain morphological traits in <Emphasis Type="Italic">Drosophila ananassae</Emphasis>

Variation in the subtle differences between the right and left sides of bilateral characters or fluctuating asymmetry (FA) has been considered as an indicator of an organism’s ability to cope with genetic and environmental stresses during development. However, due to inconsistency in the results of empirical studies, the relationship between FA and stress has been the subject of intense debate. In this study, we investigated whether stress caused by artificial bidirectional selection for body size has any effect on the levels of FA of different morphological traits in Drosophila ananassae. The realised heritability (h²) was higher in low-line females and high-line males, which suggests an asymmetrical response to selection for body size. Further, the levels of FA were compared across 10 generations of selection in different selection lines in both sexes for sternopleural bristle number, wing length, wing-to-thorax ratio, sex comb-tooth number and ovariole number. The levels of FA differed significantly among generations and selection lines but did not change markedly with directional selection. However, the levels of FA were higher in the G10 generation (at the end of selection) than G0 (at the start of selection) but lower than the G5 generation in different selection lines, suggesting that the levels of FA are not affected by the inbreeding generated during the course of selection. Also, the levels of FA in the hybrids of high and low lines were significantly lower than the parental selection lines, suggesting that FA is influenced by hybridisation. These results are discussed in the framework of the literature available on FA and its relationship with stress.     

Does fluctuating asymmetry constitute a sensitive biomarker of nutritional stress in house sparrows (Passer domesticus)?

Small random deviations from left–right symmetry in bilateral traits, termed fluctuating asymmetry (FA), are theoretically predicted to increase with environmental stress and believed to constitute a potential biomarker in conservation. However, reported relationships between FA and stress are generally weak and variable among organisms, traits and stresses. Here we test if, and to what extent, FA increases with nutritional stress, estimated from independent feather growth measurements, in free-ranging house sparrows (Passer domesticus). Ptilochronological feather marks showed significant heterogeneity among study plots, indicating that house sparrow populations were exposed to variable levels of nutritional stress during development. However, individuals from more stressed populations did not show increased levels of fluctuating asymmetry in tarsus or rectrix length, nor was there evidence for significant between-trait concordance in FA at the individual or the population level. Lack of support for FA in tarsus and rectrix length as estimator of nutritional stress in house sparrows may indicate that developmental instability is insensitive to nutritional stress in this species, poorly reflected in patterns of fluctuating asymmetry due to ecological or statistical reasons, or highly context-specific. Such uncertainty continues to hamper the use of FA as a biomarker tool in conservation planning.     

Is phenotypic canalization involved in the decline of the endemic Aquilegia thalictrifolia? Rethinking relationships between fluctuating asymmetry and species conservation status

Fluctuating asymmetry (FA), the deviation from the normal symmetrical condition of a morphological trait having specific morphological symmetry, increases in response to environmental and genetic stress, is related to phenotypic plasticity and is considered a tool for monitoring a species conservation status. However, FA–stress relations are dependent on measured traits or species‐specific characteristics such as mating system and habitat. This study investigates the relationships between FA, genetic diversity, population size, density and individual fitness traits (plant height, fruit and seed set), in the endemic Aquilegia thalictrifolia, a mixed breeder that is declining, but maintaining high levels of heterozygosity. Leaf and flower FA and other traits were investigated in 10 populations of A. thalictrifolia, the whole species range. As a result, we found similar patterns of FA in leaves and flowers between populations, indicating a homogenous level of stress between populations that differed for other traits. FA and the other traits were not related, including heterozygosity. Heterozygosity was not related to individual fitness traits with the exception of plant height. In accordance with other studies, we found that the role of FA as a tool for assessing the conservation status of a species or population should be reconsidered. However, we conclude that a low level of FA should not automatically be considered an indicator of good conservation status or low level of stress, because in species that evolved in highly stable environments it may indicate a scarce ability to plastically respond to environmental changes, as a consequence of environmental and genetic canalization. Further investigation of this point is needed.     

Evolutionary history shapes the association between developmental instability and population‐level genetic variation in three‐spined sticklebacks

Developmental instability (DI) is the sensitivity of a developing trait to random noise and can be measured by degrees of directionally random asymmetry [fluctuating asymmetry (FA)]. FA has been shown to increase with loss of genetic variation and inbreeding as measures of genetic stress, but associations vary among studies. Directional selection and evolutionary change of traits have been hypothesized to increase the average levels of FA of these traits and to increase the association strength between FA and population‐level genetic variation. We test these two hypotheses in three‐spined stickleback (Gasterosteus aculeatus L.) populations that recently colonized the freshwater habitat. Some traits, like lateral bone plates, length of the pelvic spine, frontal gill rakers and eye size, evolved in response to selection regimes during colonization. Other traits, like distal gill rakers and number of pelvic fin rays, did not show such phenotypic shifts. Contrary to a priori predictions, average FA did not systematically increase in traits that were under presumed directional selection, and the increases observed in a few traits were likely to be attributable to other factors. However, traits under directional selection did show a weak but significantly stronger negative association between FA and selectively neutral genetic variation at the population level compared with the traits that did not show an evolutionary change during colonization. These results support our second prediction, providing evidence that selection history can shape associations between DI and population‐level genetic variation at neutral markers, which potentially reflect genetic stress. We argue that this might explain at least some of the observed heterogeneities in the patterns of asymmetry.     

Communal Roost Site Selection in a Neotropical Harvestman: Habitat Limitation vs. Tradition

Many species have been reported to form roosting (resting, sleeping) aggregations at ‘traditional’ sites, but the alternative hypothesis that specific sites are used repeatedly because of habitat limitation is rarely tested. We studied the roosting behavior of a species of harvestman (Opiliones, Prionostemma sp.) at a lowland rainforest site in Nicaragua. Both sexes roosted by day in spiny palm trees, dispersed at dusk to forage, and rejoined aggregations just before dawn. The distribution of harvestmen among spiny palms was significantly clumped, and harvestman density did not correlate with spiny palm density. Aggregations formed repeatedly in a small subset of the available spiny palms and the same sites were used in two different years (2001, 2003). Nevertheless, the membership of aggregations was fluid; individual harvestmen were found at multiple roosts and moved up to 0.2 km per night. Translocated animals often returned to the roost where they had been released or nearby roosts but were never found at previously unused sites. The high consistency of site use but low site fidelity of individuals suggests that roost sites differed conspicuously (to the harvestmen) from sites that were not used. We found no univariate or multivariate differences between used and unused sites, however, in the characteristics of the trees or microclimate. These results conflict with the habitat limitation hypothesis but are consistent with the traditional site use hypothesis. The tradition may be mediated by a site‐labeling chemical, a mechanism that does not require individual site fidelity. We discuss these results in relation to the proposed functions of roosting aggregations.     

The effect of diet quality on physiological and life-history traits in the harvestman Pachylus paessleri

The balance between energy acquisition and expenditure is critical to the survival and reproductive success of animals. Here we investigate the long-term effects of diet quality on physiological and life-history flexibility in the harvestman, Pachylus paessleri. We used cow meal as a protein-rich diet and potatoes as a carbohydrate-rich diet in order to reproduce two extreme conditions regarding food quality in harvestmen natural habitat. As proxy variables of the energy expenditure process, we quantified standard metabolic rate (maintenance), changes in body mass (somatic condition), and fecundity (reproduction). We found that animals consuming the protein-rich diet were able to increase both their body condition and fecundity. However, the increment in these two life-history traits was correlated with higher maintenance costs. In contrast, the carbohydrate-rich diet did not provide enough specific nutrients for reproductive events, although it may have allowed animals to survive for a long time. Thus, according to the quality of the diet available in the environment, harvestman females can adopt different life-history strategies correlated with phenotypic adjustments at anatomical and physiological levels. In the Mediterranean region, spatial and temporal changes in food quality are typical, so greater phenotypic flexibility is expected to cope with this kind of environmental variation.     

Hedgerows support rich communities of harvestmen (Opiliones) in upland agricultural landscape

Hedgerows play important roles in agricultural landscapes and they increase biodiversity by providing habitat refugia for species sensitive to agricultural disturbance. We have studied the characteristics of the communities of harvestmen (Opiliones) inhabiting hedgerows. Harvestmen are terrestrial arachnids associated with soil surface and subsurface and sensitive to land-use and disturbance. We were specifically interested in quantifying how hedgerow characteristics (e.g., elevation, length, width, connectivity, and plant diversity) affect harvestman diversity and community composition. We expected harvestman diversity and community composition to be positively related to both hedgerow size (area, length, width) and biological attributes of hedgerows (e.g., connectivity, plant community diversity). We surveyed hedgerow characteristics and harvestman communities of 20 hedgerows in an upland agricultural landscape in the Western Carpathians. Hedgerow characteristics were measured in the field or derived from GIS layers and we used correlation and ordination methods to relate them to harvestman community metrics. We found surprisingly high taxonomic richness of harvestmen within the studied hedgerows (15 species). Importantly, the Shannon index of harvestman communities was positively related to hedgerow length and hedgerow tree layer species richness and diversity. Harvestman community composition varied with hedgerow area and width. Despite their small total area, hedgerows represented an important habitat for diverse harvestman communities and hedgerow attributes such as size and tree diversity significantly affected the composition and the Shannon index of harvestman communities. Thus, greater lengths and widths of hedgerows can provide habitats for a higher Shannon index of harvestman communities, within surrounding agricultural landscapes.     

Effects of an aerial 1080 possum poison operation using carrot baits on invertebrates in artificial refuges at Whirinaki Forest Park, 1999– 2002

The effects of an aerial 1080 possum poison operation using carrot baits on invertebrates in Whirinaki Forest Park are described from an un-replicated study of artificial refuges attached to tree trunks. Auckland tree weta (Hemideina thoracica), cave weta (Pharmacus sp. and Isoplectron sp.), cockroaches, spiders and harvestmen, and leaf-veined slugs (Athoracophorus bitentaculatus) were the most frequent occupants, but snails, millipedes, centipedes, flatworms, lepidopteran larvae, glowworm larvae (Arachnocampa luminosa), peripatus, slaters and beetles were also present occasionally. Invertebrate numbers were monitored every second or third month for a year before the poison operation, and for two years afterwards. Numbers of tree weta, cave weta, cockroaches, spiders and harvestmen, and leaf-veined slugs did not decline substantially in refuges in the treatment area relative to those in the non-treatment area immediately after the poison operation. Our results, and those from two other similar studies, suggest that aerial 1080 poison operations are unlikely to have a detrimental effect on invertebrates that occupy cavities above ground.     

Harvestman (Opiliones) fauna associated with Maine lowbush blueberry fields in the major production areas of Washington and Hancock counties

Over a period of 19 yr, the harvestman (Opiliones) community associated with the lowbush blueberry agro-ecosystem in Maine was studied. Eight species representing five genera, four subfamilies, and two families of harvestmen belonging to the suborder Eupnoi were collected. The harvestman community was dominated by two introduced, synanthropic species: Phalangium opilio in all but 1 yr (that year dominated by Rilaena triangularis). Rilaena was recorded for the first time from eastern North America. Relative abundance of harvestman adults increases throughout the season and the temporal pattern of trap capture does not refute speculated life cycles of the harvestmen being univoltine with overwintering eggs. Some blueberry management practices were found to affect trap capture. We did find that on average (with opposite results 1 yr) trap captures are greater in pruned fields than in fruit-bearing fields. Organic fields were found to have higher relative abundance of harvestmen than conventionally managed fields. Conventionally managed fields with reduced-risk insecticides showed no difference in harvestmen relative abundance compared with those conventionally managed fields using the older more persistent organophosphate insecticides. Insecticide trials with common insecticides used in blueberry insect pest management showed that the organophosphate insecticide, phosmet, and the pyrethroid insecticide, esfenvalerate, were detrimental to P. opilio adults when exposed to leaf residues, whereas the reduced-risk insecticide, spinosad, showed no negative effects compared with nonsprayed foliage.     

Land use influences harvestman diversity in northern Spain: a case study involving secondary grasslands and forest plantations (Arachnida: Opiliones)

Harvestmen have a general distribution pattern with more species and higher abundance in forests than in open habitats, as previously verified in mountain Cantabrian areas of northern Spain, of the Orocantabrian Province. The study of harvestmen of the low Cantabrian areas of the adjacent biogeographic Cantabro-Atlantic Province is highly appropriate to determine the influence of biogeographic position on the main diversity parameters and the effect of different land uses on harvestman assemblages. The main types of managed habitats (forest plantations and secondary grasslands), together with natural forests, non-planted young forests, shrublands and habitat boundaries were continuously sampled with seven pitfall traps during 1 year at 28 sites. The harvestman assemblages were classified using six different analyses, and indicator species were identified. The spatial patterns of harvestman diversity in low areas differed from those of mountain areas, although they have 15 species in common. Remarkably, higher average harvestman species richness was measured in low Cantabrian areas than in mountain areas. Shrublands and boundaries were the most species-rich habitats. Forested areas were the poorest in abundance, and were not the habitats richest in harvestman species, though they had higher average richness than mountain Cantabrian forests. Grasslands had a unique harvestman composition with significant extraordinary abundances, in particular of Homalenotus quadridentatus (indicator species of this habitat) and H. laranderas. Interestingly, H. laranderas, Paroligolophus agrestis and Ischyropsalis hispanica, indicator species of some open habitats in low Cantabrian areas, have recently been shown to be indicators of shady forests in mountain Cantabrian territories.     

Field predation of twospotted spider mite in a New Zealand strawberry crop

Predation by naturally occurring predatory arthropods was investigated to explain variations in population numbers of twospotted spider mite (Tetranychus urticae Koch) between first and second season strawberry crops. Araneomorph spiders. European harvestman [Phalangium opilio (L.)], Tasmanian lacewing [Micromus tasmaniae (Walker)] and Pacific damsel bug [Nabis kinbergii Reuter] were the only predators found in high numbers. However, spiders and harvestment were more prevalent than lacewings and nabids. Laboratory feedings trials indicated spiders build horizontal webs in the plants and prey predominantly on small flying insects that shelter in the crops. Similar feeding trials cofirmed the palatability of TSSM to spiders and harvestmen. Immunological testing for proteins of TSSM, aphids, Collembola and leafrollers in the intestines of field collected European harvestman, spiders, Tasmanian lacewing and Pacific damsel bug confirmed European harvestman includes TSSM in its diet, but not in large enough quantities to exert a significant regulating pressure on TSSM populations. Lacewings and nabids include TSSM in their diets but only in very small quantities (2% and 1% respectively). Spiders do not take TSSM unless they drop or spin down onto the spider webbing. The immunological testing also showed European harvestman to be a polyphagous and opportunistic feeder. Prey residues were detected more frequently in harvestmen intestines at times of prey abundance which indicated a seasonality to harvestmen diet.      

Proximate factors and potential benefits influencing selection of Psychotria suterella for shelter by the harvestman Jussara spec.

Harvestmen (Arachnida, Opiliones) are arachnids that rely on chemicals for communication and are particularly dependent on high humidity. The harvestman Jussara spec. (Sclerosomatidae) clearly prefers to rest on the stem and leaves of Psychotria suterella Muell. Arg. (Rubiaceae), a plant having a complex architecture of overlapping branches. So far, few studies have focused on understanding how harvestmen find their host plant and the benefits associated with the selected plant. Here, we investigated cues harvestmen may exploit to find P. suterella and the potential benefits for harvestmen of this interaction. To address how harvestmen find the plant, we tested two non‐exclusive hypotheses: harvestmen use chemicals from conspecifics, and/or harvestmen use chemicals from plants. For the first hypothesis, we assessed the number of harvestmen choosing P. suterella with or without chemicals of Jussara spec. on the main stem. We did not find evidence that Jussara spec. uses chemicals from conspecifics to select P. suterella, at least in isolation, without any mechanical stimuli such as stem texture or size. For the second hypothesis, we tested harvestman behavior exposed to volatiles from P. suterella, a non‐preferred host plant – Impatiens walleriana Hook.f. (Balsaminaceae) – and a blank treatment, both in a triangular arena and in a Y‐tube olfactometer. We also found no evidence that the harvestmen use plant volatiles to find it. We also tested two non‐exclusive hypotheses regarding the benefits of selecting P. suterella for harvestmen. The first hypothesis is that P. suterella offers a higher humidity than other plants in the micro‐environment surrounding the leaves because of a high leaf transpiration rate. Harvestmen could benefit from this because they rest with the body in contact with the leaf or at less than 5 mm from it. We did not find evidence that leaf transpiration rate is important for Jussara spec., as P. suterella did not present high rates compared to other local species. The second hypothesis is that the leaves of P. suterella provide a better shadow than other plants, acting as a sun shade due to its complex architecture with overlapping leaves. We measured light passage through the leaves of P. suterella and three other local species and found that less light passes through the leaves of P. suterella, which therefore provides darker shadow. This potentially provides a favorable micro‐climate for harvestmen, which might help to explain the micro‐habitat preference of Jussara spec.     

Population ecology ofGoniosoma spelaeum, a cavernicolous harvestman from south-eastern Brazil (Arachnida: Opiliones: Gonyleptidae)

Goniosoma spelaeum (Mello-Leitão) is a widespread trogloxene harvestman in caves of the Ribeira Valley in São Paulo State. It inhabits walls and ceilings near cave entrances. Populations inhabiting six selected caves of the region were analysed during this study. Mark-recapture techniques were used to estimate population sizes. Populations vary from tens to hundreds of individuals in each cave, and vary throughout the year. Barra Bonita cave had the largest population and was used to identify other ecological features of the species. These harvestmen show a high degree of philopartry, and recapture rate was high. They remain motionless inside the caves during the day, leaving them after dark to forage, and return just after dawn. They always feed outside the caves and do not carry food into the caves. Soft-bodied insects were the main food observed in the field. In the laboratory, they accepted plant and animal items and industrial foods. Their main predators are the spider Ctenus fasciatus , the heteropteran Zelurus travassosi , and the marsupial Philander opossum , and their main parasites are dipteran larvae (probably Phoridae).     

Temperature and genotypic effects on life history and fluctuating asymmetry in a field strain of Culex pipiens

Fluctuating asymmetry (FA) has been proposed as a tool to measure levels of stress experienced by populations of organisms during development. To be of value as a bio-marker to highlight conditions at particular sites, it is important that variation in FA is due to environmental (eg pollution) variation and not genetic variation among populations and families, in other words heritability for FA should be very close to zero. A full-sib design was set up in which families of Culex pipiens mosquitoes collected from the field were reared at three different developmental temperatures. The effects of temperature and family on developmental rate, egg to adult survival and four wing morphological measures were assessed. There was both a temperature and a family effect on development rate and survival. Temperature affected all four wing traits, but an influence of family was only evident in two of the wing traits. Two separate measures of FA for each of the wing traits were obtained. The mean estimates of FA were mainly around 1% of the value of the character measured. There was evidence of an increase in FA with increase in temperature stress. Heritability was estimated for the wing traits and wing trait FA's using restricted estimation maximum likelihood. The estimates of heritability for the wing traits were small and, individually, did not differ significantly from zero. There was also no evidence of heritable genetic variation for any of the wing trait FA's. The results are discussed in relation to other studies where FA heritabilities have been estimated and in relation to the use of FA as an indicator of environmental stress.     

Marsupials from space: fluctuating asymmetry, geographical information systems and animal conservation

We report the development of a new quantitative method of assessing the effects of anthropogenic impacts on living beings; this method allows us to assess actual impacts and to travel backwards in time to assess impacts. In this method, we have crossed data on fluctuating asymmetry (FA, a measure of environmental or genetic stress), using Didelphis albiventris as a model, with geographical information systems data relating to environmental composition. Our results show that more impacted environments resulted in statistically higher levels of FA. Our method appears to be a useful and flexible conservation tool for assessing anthropogenic impacts.     

Relationship between fluctuating asymmetry and fitness within and between stressed and unstressed populations of the wolf spider Pirata piraticus

Although developmental instability, measured as fluctuating asymmetry (FA), is expected to be positively related to stress and negatively to fitness, empirical evidence is often lacking or contradictory when patterns are compared at the population level. We demonstrate that two important properties of stressed populations may mask such relationships: (i) a stronger relationship between FA and fitness, resulting in stronger selection against low quality (i.e. developmental unstable) individuals and (ii) the evolution of adaptive responses to environmental stress. In an earlier study, we found female wolf spiders Pirata piraticus from metal exposed populations to be characterized by both reduced clutch masses and increased egg sizes, the latter indicating an adaptive response to stress. By studying the relationship between these two fitness related traits and levels of FA at individual level, we here show a significant negative correlation between FA and clutch mass in metal stressed populations but not in unstressed reference populations. As a result, levels of population FA may be biased downward under stressful conditions because of the selective removal of developmentally unstable (low quality) individuals. We further show that females that produced larger eggs in stressed populations exhibited lower individual FA levels. Such interaction between individual FA and fitness with stress may confound the effect of metal stress on FA, resulting in an absence of relationships between FA, fitness and stress at the population level.     

THE DUAL ROLE OF SELECTION AND EVOLUTIONARY HISTORY AS REFLECTED IN GENETIC CORRELATIONS

The patterns of genetic correlations between a series of eye and antenna characters were compared among two sets of spring-dwelling and cave-dwelling populations of Gammarus minus. The two sets of populations originate from different drainages and represent two separate invasions of cave habitats from surface-dwelling populations. Matrix correlations, using permutation tests, indicated significant correlations both between populations in the same basin and from the same habitat. The technique of biplot, which allows for the simultaneous consideration of relationships between different genetic correlations and different populations, was used to further analyze the correlation structure. A rank-3 biplot indicated that spring and cave populations were largely differentiated by eye-antennal correlations, whereas basins were differentiated by both eye-antennal and antennal-antennal correlations. Eye-antennal correlations, which are likely to be subject to selection, were most similar within habitats, which are likely to have similar selective regimes.