Cryptic speciation between Juniperus deltoides and Juniperus oxycedrus (Cupressaceae) in the Mediterranean

Analyses of individuals classically treated as Juniperus oxycedrus L. var. oxycedrus from Morocco, Portugal, Spain, France, Italy, Greece and Turkey, using DNA sequencing of nrDNA (ITS 1, 5.8S, ITS 2) plus RAPDs, leaf terpenoids and morphology revealed that two cryptic, genetically distinct but morphologically almost identical species are present. These species, J. oxycedrus L. var. oxycedrus and Juniperus deltoides R.P. Adams, are about as different from each other as Juniperus navicularis and Juniperus macrocarpa are from J. oxycedrus var. oxycedrus. Examination of herbarium specimens revealed that the two species are largely allopatric with J. deltoides occurring from Italy eastward through Turkey into the Caucasus Mts. and Iran. J. oxycedrus var. oxycedrus appears to be largely concentrated west of Italy (France, Spain, Portugal, Morocco). Cryptic speciation is discussed.     

Morphological analysis,chorology and ecology of Juniperus navicularis Gand. in the Iberian Peninsula

Juniperus navicularis is an endemic taxon of Iberian Peninsula that develops on paleodunes of coastal ecosystems and whose distribution range is not well defined. In this paper, we present the results of the morphological analysis in populations attributed to J. navicularis of southern Spain. In order to assess the similarity between Portuguese and Spanish populations, the Mann–Whitney U test was performed, the same test was also applied between this taxon and Juniperus oxycedrus subsp. oxycedrus. In order to distinguish this species from other taxa of the Juniperus oxycedrus group (subsp. oxycedrus, subsp. badia, subsp. macrocarpa) in the southwestern Mediterranean region, a principal component analysis has been carried out. New data about the chorology and ecology of Spanish populations considered as a priority habitat in the Habitat Directive as well as number of individuals and conservation status are provided.     

Large-Scale Patterns of Quercus ilex, Quercus suber, and Quercus pyrenaica Regeneration in Central-Western Spain

In Central-Western Spain, forests and woodlands composed of Quercus sp. support outstanding levels of biodiversity, but there is increasing concern about their long-term persistence due to a lack of regeneration. We hypothesize that this regenerative lack is operating on a large geographic scale; that there are differences in the abundance of regeneration between three oak species; that oak regeneration is governed mainly by forest management and structure; and that shrubs act as important physical protectors of seedlings and saplings. We analyzed whether densities of oak seedlings and saplings in several size classes were related to stand-structure, understory, and physiographic variables potentially affecting regeneration. Data collected at a regional level (1 km × 1 km grid) by the Spanish Forest Inventory were evaluated from 2,816 plots. Results revealed that regeneration failure was common for all size categories, from small seedlings to large saplings, and for the three oak species studied, especially the evergreens. Of the Quercus ilex, Q. suber, and Q. pyrenaica plots studied, 49%, 62%, and 20% were lacking any small seedlings, and 82%, 96%, and 56% did not have any large saplings, respectively. Regeneration was positively correlated with tree cover and density, especially of small and medium-sized trees, and negatively correlated with the presence of large trees, indicating that regeneration failure is mostly associated with more open, uniform, and/or aged woodlands. Regeneration densities of Q. ilex and Q. suber were positively correlated with all understory variables, suggesting that the presence of pioneer shrubs represent a major safe site for early tree recruitment, independent from specific shrub species.     

Variability of the Needle Essential Oils of Juniperus deltoides R.P.Adams from Different Populations in Serbia and Croatia

The essential‐oil compositions of one Croatian and three Serbian populations of Juniperus deltoides R.P.Adams have been determined by GC/MS analysis. In total, 147 compounds were identified, representing 97.3–98.3% of the oil composition. The oils were dominated by monoterpenes, which are characteristic components for the species of the section Juniperus. Two monoterpenes, α‐pinene and limonene, were the dominant constituents, with a summed‐up average content of 49.45%. Statistical methods were used to determine the diversity of the terpene classes and the common terpenes between the newly described J. deltoides populations from Serbia and Croatia. Only reports on several specimens from this species have been reported so far, and there are no studies that treat population diversity. Cluster analysis of the oil contents of 21 terpenes showed high correlation with the geographical distribution of the populations, separating the Croatian from the Serbian populations. The comparison of the essential‐oil compositions obtained in the present study with literature data, showed the separation of J. deltoides and J. oxycedrus ssp. oxycedrus from the western Mediterranean region.     

Plant Responses to Extreme Climatic Events: A Field Test of Resilience Capacity at the Southern Range Edge

The expected and already observed increment in frequency of extreme climatic events may result in severe vegetation shifts. However, stabilizing mechanisms promoting community resilience can buffer the lasting impact of extreme events. The present work analyzes the resilience of a Mediterranean mountain ecosystem after an extreme drought in 2005, examining shoot-growth and needle-length resistance and resilience of dominant tree and shrub species (Pinus sylvestris vs Juniperus communis, and P. nigra vs J. oxycedrus) in two contrasting altitudinal ranges. Recorded high vegetative-resilience values indicate great tolerance to extreme droughts for the dominant species of pine-juniper woodlands. Observed tolerance could act as a stabilizing mechanism in rear range edges, such as the Mediterranean basin, where extreme events are predicted to be more detrimental and recurrent. However, resistance and resilience components vary across species, sites, and ontogenetic states: adult Pinus showed higher growth resistance than did adult Juniperus; saplings displayed higher recovery rates than did conspecific adults; and P. nigra saplings displayed higher resilience than did P. sylvestris saplings where the two species coexist. P. nigra and J. oxycedrus saplings at high and low elevations, respectively, were the most resilient at all the locations studied. Under recurrent extreme droughts, these species-specific differences in resistance and resilience could promote changes in vegetation structure and composition, even in areas with high tolerance to dry conditions.     

Habitat requirements for the conservation of wood cricket (Nemobius sylvestris) (Orthoptera: Gryllidae) on the Isle of Wight, UK

In the UK, wood cricket (Nemobius sylvestris) is a ‘Species of Conservation Concern’, being restricted to only three areas in southern England. Little information is available on the specific habitat requirements of this species. In 2006, a field investigation within three woodlands on the Isle of Wight was undertaken to identify its habitat preferences. Factors positively influencing wood cricket presence within woodlands included the presence of a well-developed leaf litter layer, relatively low ground vegetation cover and height, low canopy cover and relatively short distances between individual populations. Regression models identified the degree of isolation and variables describing vegetation structure as the main predictors for wood cricket presence within woodland fragments. The results of this study indicate the preference of wood cricket for open wooded edges. Conservation efforts for this species should focus on continuation of regular management activities aimed at providing permanent open edge habitat within woodlands, to maintain viable populations.     

Genome size and ploidy levels in highly fragmented habitats: the case of western Mediterranean Juniperus (Cupressaceae) with special emphasis on J. thurifera L.

Mediterranean junipers are of special ecological importance as key components of resource islands in semi-arid mountain ecosystems of the Mediterranean basin. The fragmentation of their habitat, which was primarily natural and driven by climatic drought conditions, is currently being aggravated by anthropogenic pressure. In the framework of this concern, the present work aims to contribute establishing a genomic profile of Juniperus in its western Mediterranean range, with a special emphasis placed on J. thurifera. DNA contents were assessed by flow cytometry in 43 populations of nine taxa within their Mediterranean range (first reports for J. navicularis, J. thurifera subsp. africana and J. thurifera subsp. thurifera). Chromosome numbers were determined by orcein staining in eight taxa (first counts for J. oxycedrus subsp. badia, J. phoenicea subsp. phoenicea, J. phoenicea subsp. turbinata, of 2n?=?2x?=?22, and for J. thurifera subsp. thurifera, of 2n?=?4x?=?44). Tetraploid cytotypes have been the only ones found in the 19 populations of J. thurifera studied, this being the first report of a Juniperus species exclusively polyploid. In J. thurifera, C-value does not respond to habitat fragmentation, in the same way that genetic diversity within populations was previously shown to be unaltered, suggesting that this factor has not had, at least to date, a significant impact on populations at genomic and genetic levels. Habitat fragmentation leads to deeply age-biased populations with a male-biased imbalanced sex ratio (lack of females), indicating an urgent need to improve regeneration within the populations of this species.     

The biogeography and genetic relationships of Juniperus oxycedrus and related taxa from the Mediterranean and Macaronesian regions

Despite Juniperus spp. being an important component of Mediterranean arid and semi‐arid ecosystems, there is a lack of complex studies on their biogeographical patterns. Using 16 morphological cone and seed traits and three nuclear microsatellite markers, we investigated the morphological and genetic variability of seven Mediterranean and Macaronesian Juniperus taxa (J. oxycedrus ssp. oxycedrus, J. oxycedrus ssp. badia, J. brevifolia, J. cedrus, J. deltoides, J. macrocarpa and J. navicularis) to identify biogeographical trends and interspecific genetic relationships. The highest gene diversity was measured in J. oxycedrus ssp. oxycedrus (H_E = 0.716) and the lowest in J. brevifolia (H_E = 0.441). The west Mediterranean was characterized by a higher level of genetic diversity than the east Mediterranean. A lack of significant genetic differences between European and African populations of J. oxycedrus suggests that the Strait of Gibraltar was not a significant barrier to gene flow, but has promoted some morphological differentiation. The genetic and morphological results strongly support the recognition of J. macrocarpa, J. navicularis and J. deltoides at the species rank, whereas J. oxycedrus ssp. badia should be included in J. oxycedrus. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 637–653.     

Streptanthus bracteatus (Brassicaceae), a rare annual woodland forb, thrives in less cover: evidence of a vanished habitat?

One of the challenges of understanding habitat requirements of endangered species is that the remaining populations may not be in optimal habitat, requiring experimentation to determine optimal habitat and to guide management. A better knowledge of its habitat requirements is important for the conservation of Streptanthus bracteatus, a rare annual of central Texas woodlands. The habitat requirements of a rare, declining species like S. bracteatus can also provide insights into anthropogenic habitat degradation and into previous disturbance regimes. We conducted a garden experiment and a transplant experiment to determine the effect of different light environments on the growth and reproduction of S. bracteatus. Higher levels of light improved S. bracteatus performance, especially fecundity. The optimum level of combined canopy and understory cover at the height of a S. bracteatus plant (??0.5?m above ground) was less than 50?%. The remaining populations of S. bracteatus are in sub-optimal habitat because it is not open enough. The results are consistent with the hypothesis that this species was a ??fire-follower.?? The results also support the hypotheses that central Texas woodlands were once more open and that fire played an ecological role in these woodlands, an example of the ecological requirements of a rare species revealing past community structure and dynamics.     

Essential oil yield,composition, bioactivity and leaf morphology of Juniperus oxycedrus L. from Bulgaria and Serbia

Juniperus oxycedrus L. (Cupressaceae Bartlett) is widely distributed in countries with a Mediterranean climate. All plant parts contain highly aromatic essential oil (EO) and recently there have been efforts to introduce it as a cultivated crop. The species is known for its large morphological and chemical variation and its debatable taxonomic status. This study aimed to (1) compare content, composition, and antimicrobial activity of J. oxycedrus EO samples from plants growing in Bulgaria and Serbia, and (2) quantify morphological variations of leaves. Тhe EO content (yield) in dried juniper leaves varied from 0.06% (Кopaonik, Serbia) to 0.24% (Markovo, Bulgaria). We identified 51 EO constituents, belonging to monoterpenes, sesquiterpenes, and diterpenes. The class monoterpenes (monoterpene hydrocarbons and oxygenated monoterpenes) were the predominant compounds, representing 38.6–65.4% of the total EO, consisting primarily of α-pinene, limonene, sabinene, β-pinene, and β-myrcene. In addition, α-pinene was the major oil constituent in plants from all locations. Sesquiterpenes (sesquiterpene hydrocarbons and oxygenated sesquiterpenes) were the second largest class of constituents, which represented 19.3% tо 33.6% of the total EO. γ-Elemene was found only in the EO of J. oxycedrus from Bulgaria, while a high concentration of α-curcumene was found only in samples from Serbia (7.5–7.8%). Significant differences in antimicrobial activity of the EO were found in bacterial strains Bacillus cereus and Streptococcus pneumoniae. There was no significant difference among the mean leaf width of the six combinations location x sex, and the overall leaf mean width was 1.24 mm. However, there was a significant difference between the mean leaf lengths. In this study, none of the studied populations had a higher concentration of limonene than of α-pinene, indicating that the flora of the two countries include J. oxycedrus and not the previously reported J. deltoides. The results revealed significant variation in EO profile that may contribute to the development of new cultivars of J. oxycedrus.     

Morphological variation of Juniperus oxycedrus subsp. oxycedrus (Cupressaceae) in the Mediterranean region

The intra- and inter-population variation of Juniperus oxycedrus subsp. oxycedrus were examined biometrically on the basis of morphological characters of needles, seeds and cones. Cluster analysis of Euclidean distances and discriminant analysis were performed to verify the hypothesis of geographic differentiation and origin from the West- and East-Mediterranean Pleistocene refugial regions. The West-Mediterranean populations of J. oxycedrus subsp. oxycedrus differed from the eastern ones. The intra-populational differentiation of three of the West-Mediterranean populations was significantly higher than within the other West- and all the East-Mediterranean ones.Results confirm the hypothesis of probable different origins of the East- and West-Mediterranean populations of the taxon.     

Post‐fire development of faunal habitat depends on plant regeneration traits

The concept that vegetation structure (and faunal habitat) develops predictably with time since fire has been central to understanding the relationship between fire and fauna. However, because plants regenerate after fire in different ways (e.g. resprouting from above‐ground stems vs. underground lignotubers), use of simple categories based on time since fire might not adequately represent post‐fire habitat development in all ecosystems. We tested the hypothesis that the post‐fire development of faunal habitat structure differs between ecosystems, depending on fire regeneration traits of the dominant canopy trees. We measured 12 habitat components at sites in foothill forests (n = 38), heathy woodlands (n = 38) and mallee woodlands (n = 98) in Victoria, Australia, and used generalised additive models to predict changes in each variable with time since fire. A greater percentage of faunal habitat variables responded significantly to time since fire in mallee woodlands, where fires typically are stand‐replacing, than in foothill forests and heathy woodlands, where canopy tree stems generally persist through fire. In the ecosystem with the highest proportion of epicormic resprouters (foothill forests), only ground cover and understorey vegetation responded significantly to time since fire, compared with all but one variable in the ecosystem dominated by basal resprouters (mallee woodlands). These differences between ecosystems in the post‐fire development of key habitat components suggest there may also be fundamental differences in the role of fire in shaping the distribution of fauna. If so, this challenges the way in which many fire‐prone ecosystems currently are categorised and managed, especially the level of dependence on time since fire and other temporal surrogates such as age‐classes and successional states. Where time since fire is a poor surrogate for habitat structural development, additional complexity (e.g. fire severity, topography and prior land‐use history) could better capture processes that determine faunal occurrence in fire‐prone ecosystems.     

Effects of landscape features on gene flow of valley oaks (<Emphasis Type="Italic">Quercus lobata</Emphasis>)

Landscape features affect habitat connectivity and patterns of gene flow and hence influence genetic structure among populations. We studied valley oak (Quercus lobata), a threatened species of California (USA) savannas and oak woodlands, with a distribution forming a ring around the Central Valley grasslands. Our main goal was to determine the role of topography and land cover on patterns of gene flow and to test whether elevation or land cover forms stronger barriers to gene flow among valley oak populations. We sampled valley oaks in 12 populations across the range of this species, genotyped each tree at eight nuclear microsatellite loci, and created a series of resistance surfaces by assigning different resistance values to land cover type and elevation. We also estimated recent migration rates and evaluated them with regard to landscape features. There was a significant but weak relationship between Euclidian distance and genetic distance. There was no relationship between genetic distances and land cover, but a significant relationship between genetic distances and elevation resistance. We conclude that gene flow is restricted by high elevations in the northern part of the valley oak range and by high elevations and the Central Valley further south. Migration rate analysis indicated some gene flow occurring east–west but we suggest that the high connectivity in the northern Central Valley is facilitating the formation of these links. We predict that southern populations may become more differentiated in the future through genetic isolation and local adaptation taking place in the face of climate change.     

Comparative Field Summer Stress of Three Tree Species Co-occurring in Mediterranean Coastal Dunes

Chlorophyll a fluorescence, water potential (_s), and root system of Juniperus oxycedrus ssp. macrocarpa, Juniperus phoenicea ssp. turbinata, and Pinus pinea were studied in Mediterranean coastal dunes of SW Spain during summer drought and after fall rains in 1999, the driest year in the 90's. A strong and reversible depression in the photochemical efficiency of photosystem 2 of the three species was recorded, which happened concomitantly with the diurnal increase and decrease in radiation. J. phoenicea, with superficial root system, was the most affected species by summer drought. It showed high rates of down-regulation of photosynthesis by photoinhibition and positive correlation between _s and F_v/F_p, with _s lower than -7 MPa. However, it tolerated this high stress, showing a fast recovery of its physiological state after fall rains. On the other hand, J. oxycedrus and P. pinea, both with deep root systems, kept their _s values up to -3 MPa, showing lower stress during summer drought. On the other hand, J. oxycedrus and J. phoenicea were more sensible to changes in edaphic water content than P. pinea. These specific responses to summer drought would be determined by their root distributions and stomatal control of transpiration, conditioning the efficiency in getting and using the available water resources. Ecophysiological responses indicate that these species are well-adapted to long periods of drought in Mediterranean climate areas, developing different strategies: J. phoenicea tolerates high stress with a fast recovery after fall rains, while J. oxycedrus and P. pinea are less affected by summer drought since their deep root systems would allow them to reach deep water resources.     

Where do nivicolous myxomycetes occur? – Modeling the potential worldwide distribution of Physarum albescens

To identify potentially suitable areas for the mostly alpine ecological guild of nivicolous (snowbank) myxomycetes, the worldwide distribution of a distinct morphospecies, Physarum albescens, was modelled with a correlative spatial approach using the software MaxEnt from 537 unique occurrence points. Three models were developed, first with only the 19 bioclimatic variables plus elevation from the WorldClim database, second with regularization to correct for pseudo-absence, and third with additional categorical environmental layer on snow cover. All three models showed high mean AUC (area under the curve) values (>0.970). Output maps were comparable, with the third model perhaps the most realistic. For this model, snow cover, precipitation of the coldest quarter (of the year), and elevation predicted best the distribution of Ph. albescens. Elevation alone is a good predictor only in some regions, since (i) elevation of the occurrence points decreases with increasing latitude, and (ii) elevation wrongly predicts the species’ occurrence in arid mountain ranges. The model showed mountains in humid climates with highest incidence, which confirmed field studies: a long-lasting snow cover fluctuating with comparatively mild summers is the decisive factor. As such, the model can serve as a predictive map where fructifications of nivicolous myxomycetes can be expected. Limitations of the model are discussed: cryptic speciation within a morphospecies, including the evolution of reproductively isolated units which may lead to local adaptation and niche differentiation, and wider ranges for myxamoebal populations.     

Natural regeneration of trees in urban woodlands

Abstract. We studied tree regeneration, a key process for the existence of urban woodlands. We hypothesized that, besides the usual biological factors, anthropogenic ones (fragmentation, wear, pollution etc.) determine the regeneration success of tree species in urban woodlands. To test this hypothesis, within an observational setting, we collected data from 30 urban woodlands in the cities of Helsinki and Vantaa, Finland. We defined the number of living saplings (30–200 cm in height) as an indicator of regeneration success and used regression analysis to test different factors as independent variables. The results showed that different tree species responded differently to urban pressure. The regeneration of Picea abies decreased with increasing fragmentation of the forest landscape, whereas for the other most common (deciduous) species, regeneration increased. Wear, measured as total path area per study site, had a negative effect on regeneration success. An a posteriori examination of the data suggested that coarse woody debris might promote regeneration. We conclude that, although tree regeneration in general is not threatened in urban woodlands in the area we studied, the species composition may gradually change. We discuss some management implications for counteracting the urban pressures on tree regeneration.     

Morphologic variation in northern marginal Juniperus oxycedrus L. subsp. oxycedrus populations in Istria

Marginal tree populations are believed to be more differentiated and host less variation than central tree populations. The aim of this study was to perform a detailed morphometric study of J. oxycedrus L. subsp. oxycedrus in northern marginal populations in order to establish its phenotypic variation and geographical differentiation and to examine the eventual presence of putative species J. deltoides in Istria. Morphologic variation was studied in 16 northern marginal populations in Istria. Nine morphological traits from a minimum of 50 leaves from each of 206 individuals and two morphological traits from 30 to 50 seed cones in each of 103 females were measured. Phenotypic variation in Istrian populations was high and of a similar magnitude to that found in the centre of the distribution area. Gender dimorphism in leaf morphology was detected, but the study failed to confirm its uniform distribution pattern. Significant and surprisingly high among-population differentiation (8–30%) was revealed. Average leaf shape in the study area is clearly “delta-shaped” characterised by a relatively wide leaf base. However, no significant differences in any of the studied traits were found between the marginal Istrian populations and the more central populations analysed in our earlier studies.     

Successional trends in standing dead biomass in Mediterranean basin species

Question: Landscape models of fire occurrence in ecosystems assume that the time since the last fire determines vegetation flammability by enabling the accumulation of dead biomass. In this study we ask if Mediterranean basin shrublands respond to these models or, on the contrary, if initial successional stages in these ecosystems could be more flammable than later stages. Location: Mediterranean shrubland in the Valencia region, eastern Spain. Methods: Using different stages of vegetation development (5, 9, 14 and 26 years since the last fire), we first study the structural comiosition of the above‐ground biomass in 375 individuals of nine woody species. Then, we measure how the standing dead biomass varies during succession, taking into account the surface cover of each species and the quantity of total dead biomass accumulated in different successional stages (3, 9, 14 and 26 years since the last fire). Results: The largest amount of standing dead biomass at the plant community level is observed in the middle stages of the succession. Early successional species, such as Cistus spp., Ulex parviflorus and Pinus halepensis, have a higher percentage of standing dead biomass at earlier stages in the succession than species typical of later successional stages, e.g. Juniperus oxycedrus, Quercus coccifera and Quercus ilex. Conclusions: The results suggest that monotonic increase in fire hazard with increasing stand age is not necessarily the rule in Mediterranean basin shrublands, since early successional species may accumulate large amounts of standing dead biomass and thus promote fire at early successional stages.     

Spatial and temporal woodland patterns along the lower Turkwel River, Kenya

Spatial and temporal patterns of riverine woodlands in arid regions of Africa are poorly documented despite their considerable conservation value. We studied 1540 ha of riverine woodland in the lower Turkwel River floodplain, Kenya, between 1990 and 1998. Forty‐one woodland patches were mapped and their soil physical and chemical characteristics, tree species diversity, woody cover, tree density, wood volume and woodland regeneration were determined. The riverine woodland comprised nine vegetation types and a total of 14 woody species. Woodland patch mosaics were associated with microtopographical features and selected soil attributes. The most important woody species were Hyphaene compressa H. Wendl., Acacia tortilis (Forssk.) Hayne and Cadaba rotundifolia Forssk. The exotic Prosopis chilensis (Mol.) St. was invading parts of the riverine woodland. Overall, woody species diversity was low compared to similar riverine woodlands in East Africa. Tree density, wood volume and woody plant regeneration declined over the 8‐year study period, while woody cover was unchanged. Reduced tree density, wood volume and regeneration of woody species might be linked to changes in river flood patterns following the impoundment of the Turkwel Gorge Dam. It is suggested that spatially heterogeneous and temporally stochastic regeneration events, together with occasional tree mortality caused by channel abandonment, create the complex pattern of woodland patches in the lower Turkwel River floodplain. The mapped woodland patches may serve as monitoring units, which in future could reveal the interplay between changes in flooding patterns as a result of dam impoundment, anthropogenic disturbance and the well‐being of the riverine woodlands.     

“Ghosts of the past”: flightless saproxylic weevils (Coleoptera: Curculionidae) are relict species in ancient woodlands

Historic maps show that the Central European landscape was influenced by exploitive human land-use during the middle ages and in the following centuries. A mixture of ancient woodlands, which survived the period of woodland destruction, and recent woodlands, which were established after 1800, cover about 10% of the study area in NW Germany today. Weevils (Coleoptera: Curculionidae) of the subfamily Cryptorhynchinae with the genera Acalles, Kyklioacalles, Ruteria and of the subfamily Molytinae, tribe Acicnemidini with the genus Trachodes are all flightless and possibly influenced by landscape history. The aims of this investigation are (1) to examine the spatial distribution of flightless saproxylic weevils in ancient and recent woodlands in NW Germany and (2) to test the frequency of possible relict species in relation to historical and current woodland size. Based on a field study in 29 deciduous woodlands and species records in collections and literature, six flightless saproxylic weevils were found to be associated with ancient woodlands in NW Germany. None of these were recorded in any of the 14 recent woodlands studied. The frequency of these relict species is correlated with historical, but not with current, woodland size. Distribution maps for Lower Saxony and data on the phenology of the relict species are presented. These weevils are relict species of ancient woodland, because they were unable to colonise isolated woods that were established after 1800. All of them are dependent on dead or dying wood for larval development. The results show that ancient broadleaved woodlands with long-lasting habitat continuity are of high conservation value for invertebrate species such as saproxylic weevils.