An ecological classification of Central European macromoths: habitat associations and conservation status returned from life history attributes

To be used as a predictive conservation tool, classification of animal habitats should rely on actual resource requirements of individual species. Shreeve et al. (J Insect Conserv 5:145–161, 2001) produced a resource-based habitat classification for British butterflies, obtaining habitat association groups, whose constituent species differed in their distribution extent, distribution change and vulnerability in Britain. To test the utility of this approach for a group with a less-known biology, we produced a resource-based classification of habitats of Central European macromoths. We worked with macrolepidopteran moth families, except for the megadiverse Geometridae and Noctuidae. We produced a matrix of 178 life history attributes describing resource use by 164 species, subjected the matrix to ordination analysis, and compared the resulting moths groupings with external ecological information. Five habitat association groups were distinguished: I—close canopy moths, II—open canopy moths, III—grasslands moths, IV—herb-feeding hawk moths, and V—lichen feeders. The classification sustained deleting attributes related to host plants taxonomy. Groups I–III sustained control for taxonomic positions of the moths, whereas IV and V did not. Members of the groups differed in the representation of externally obtained habitat associations, biogeography elements, threat status, and range size. Endangered species were over-represented in groups II and III and underrepresented in group I, in agreement with recent land cover changes across the continent. Species resources can be used to reconstruct their habitat needs, and it is possible to scale up from life histories through habitat use to range structures.     

Validity of <Emphasis Type="Italic">Scolecenchelys aoki</Emphasis>, with a redescription of <Emphasis Type="Italic">Scolecenchelys gymnota</Emphasis> (Anguilliformes: Ophichthidae)

The myrophine ophichthid fishes (worm eels) Muraenichthys aoki Jordan and Snyder 1901 and Muraenichthys gymnotus Bleeker 1857 are redescribed as valid species of Scolecenchelys based on the types and non-type specimens collected from the Indo-Pacific. Because both species are similar to each other in having acute snouts, the posterior margin of the eye before the rictus, and their dorsal-fin origins located slightly posterior to a vertical line through the anus, Scolecenchelys aoki has usually been regarded as a junior synonym of Scolecenchelys gymnota. However, S. aoki is clearly distinguishable from S. gymnota by having a median groove on the ventral side of snout (absent in S. gymnota), uniserial maxillary teeth in smaller specimens (<200 mm TL; vs. biserial), three infraorbital sensory pores at postorbital area (vs. two), and more numerous vertebrae (56–65 in predorsal vs. 51–57; 53–58 in preanal vs. 47–52). Scolecenchelys aoki is restricted to Japanese waters and regarded as a senior synonym of Muraenichthys borealis Machida and Shiogaki 1990. Scolecenchelys gymnota is widely distributed in the Indo-Pacific, from South Africa and the Red Sea to Samoa, north to Okinawa, Japan. Sphagebranchus huysmani Weber 1913 and Muraenichthys fowleri Schultz 1943 are synonymized under S. gymnota.     

Habitat use and dispersal characteristic by <Emphasis Type="Italic">Stethophyma grossum</Emphasis>: the role of habitat isolation and stable habitat conditions towards low dispersal

Species in a highly fragmented environment, such as the intensively used agricultural landscapes of Europe, are expected to be in danger of extinction. We hypothesize according to Kisdi’s theory (Am Nat 159:579–596, 2002) that species in fragmented landscapes with isolated habitats in general tend to possess low dispersal. In order to verify this hypothesis we studied the movement patterns of Stethophyma grossum, a hygrophilous species of wetlands, by mark–release–recapture techniques in a landscape with scattered suitable habitats over 3 years. The study focused on the major population in this landscape (site #1) as dispersal behaviour was assumed to be greatest. Actually, marked individuals of S. grossum were never found in any further suitable habitats in close vicinity to site #1. Despite that the peatland meadow of study site #1 was all over covered with homogenous vegetation only 6% (1.8 ha) of the whole area (30 ha) were occupied by S. grossum. The mean recapture rate over 3 years amounted to 39% with no significant differences between males and females. Both covered little distances within their mean range size of 1.8 ha; the median distances were 36.91 m for males and 26.65 m for females. We confirm the hypothesis that sub-populations of species in longstanding naturally isolated habitats, which habitat conditions have been stable; evolved low dispersal with little movements which are routine movements to find mating partners or food.     

Settlement and pigmentation of glass eels (<Emphasis Type="Italic">Anguilla rostrata</Emphasis> Lesueur) in a coastal lagoon

American eel (Anguilla rostrata Lesueur) is born in the Atlantic Ocean, but larvae redistribute throughout diverse habitats of North American estuaries and freshwater streams. We hypothesized that early stage A. rostrata differed in abundance among sites within a single coastal lagoon, Newport Bay (Maryland) for two sampling seasons (March–May in 2007 and 2008). Catch per unit effort (CPUE) of early stage A. rostrata was usually similar between years at a site, except that it was higher at one site in 2008 than in 2007. The CPUE varied among sites within Newport Bay, but not significantly so because of high, intra-annual variance in CPUE at a site. As reported for New Jersey coastal estuaries, variation in the CPUE tended to be higher in brackish water habitats. Intra-annual variation in CPUE from March until May was partially explained by oxygenation and salinity. The CPUE of settling eels was greatest when water was well-oxygenated (dissolved oxygen >8 mg·L⁻¹) and mildly brackish (0.3–3.2 ppt). While larval supply to a coastal estuary may annually influence the magnitude of potential settlers, temporal differences in habitat conditions within Newport Bay also influenced settlement patterns. Differences in habitat conditions can affect the pigmentation rates of settling early stage eels. We measured rates of pigmentation, which corresponded with age of the fish. However, rates did not differ among sites or vary with habitat conditions. Pigmentation levels from March to May increased at a rate of about 0.02% per week. Monitoring programs for early phase American eel should consider the steepness of habitat gradients within estuaries and habitat covariates when assessing population status.     

Mud crabs (Xanthidae) in Chesapeake Bay: claw characteristics and predation on epifaunal bivalves

Abstract. We examined claw characteristics of mud crabs (Eurypanopeus depressus, Rhithropanopeus harrisii) to determine if one crab species was potentially more powerful than the other. We related our findings to the abilities of individuals of each species to open epifaunal mytiliform bivalves (Ischadium recurvum; Mytilopsis leucophaeata) that occur on beds of eastern oysters (Crassostrea virginica) in mesohaline Chesapeake Bay. There were high correlations between claw width or height and claw length, and between claw length and carapace width for both mud crab species. The mechanical advantage or “grip strength’ of the crusher and cutter claws of both species did not change with crab size (carapace width) and did not differ between sexes in each species, nor did the cutter data differ between species. However, individuals of E. depressus had a significantly stronger crusher claw grip than did those of R. harrisii. Data on mechanical advantage for both species were similar to values reported in the literature for members of other xanthid crab species. These values in turn overlapped those reported for calappid, cancrid, majid, and grapsid crabs, and were greater than those of various species of portunid crabs and individual species of fiddler crab, lobster, crayfish, and ghost shrimp. When simultaneously presented with the two species of bivalves, the mud crabs E. depressus chose mussels of M. leucophaeata first and crabs of R. harrisii chose mussels of I. recurvum first about two‐thirds of the time; ultimately, the crabs ate both bivalve species in >50% of the choice experiments. The size range in E. depressus was greater than that in R. harrisii, and crabs of E. depressus opened larger bivalves than did crabs of R. harrisii, although similar‐sized individuals of the two crab species overlapped in their ability to open bivalves of both species. In Mytilopsis leucophaeata, there is probably no size refuge from predation by the mud crabs whereas the larger mussels of I. recurvum do have a refuge in size.     

Release from native root herbivores and biotic resistance by soil pathogens in a new habitat both affect the alien <Emphasis Type="Italic">Ammophila arenaria</Emphasis> in South Africa

Many native communities contain exotic plants that pose a major threat to indigenous vegetation and ecosystem functioning. Therefore the enemy release hypothesis (ERH) and biotic resistance hypothesis (BRH) were examined in relation to the invasiveness of the introduced dune grass Ammophila arenaria in South Africa. To compare plant–soil feedback from the native habitat in Europe and the new habitat in South Africa, plants were grown in their own soil from both Europe and South Africa, as well as in sterilised and non-sterilised soils from a number of indigenous South African foredune plant species. While the soil feedback of most plant species supports the ERH, the feedback from Sporobolus virginicus soil demonstrates that this plant species may contribute to biotic resistance against the introduced A. arenaria, through negative feedback from the soil community. Not only the local plant species diversity, but also the type of plant species present seemed to be important in determining the potential for biotic resistance. As a result, biotic resistance against invasive plant species may depend not only on plant competition, but also on the presence of plant species that are hosts of potential soil pathogens that may negatively affect the invaders. In conclusion, exotic plant species such as A. arenaria in South Africa that do not become highly invasive, may experience the ERH and BRH simultaneously, with the balance between enemy escape versus biotic resistance determining the invasiveness of a species in a new habitat.Plant nomenclature follows Arnold and De Wet (1993)     

<Emphasis Type="Italic">Neoskrjabinolepis nuda</Emphasis> n. sp. from shrews on Sakhalin Island,Russia, with a taxonomic review of <Emphasis Type="Italic">Neoskrjabinolepis</Emphasis> Spasskii, 1947 (Cestoda: Cyclophyllidea: Hymenolepididae)

Neoskrjabinolepis (Neoskrjabinolepidoides) nuda n. sp. is described from the shrews Sorex unguiculatus (type-host), S. gracillimus, S. isodon and S. caecutiens on Sakhalin Island, Russia. The new species is characterised by: rostellar hooks 40–44 μm long and provided with small epiphyseal thickening of the handle; a long (95–100 μm) cirrus consisting of basal region with claw-shaped spines, a parabasal region with thin needle-shaped spines and an unarmed distal region; a cirrus-sac extending well into the median field; and 15–22 eggs per gravid uterus. A review of the species of Neoskrjabinolepis Spasskii, 1947 is presented. Currently, this genus includes nine species and is divided in two subgenera on the basis of strobilar development, which is gradual in the subgenus Neoskrjabinolepis (four species) and serial in the subgenus Neoskrjabinolepidoides Kornienko, Gulyaev & Mel’nikova, 2006 (five species). An amended generic diagnosis and an identification key to Neoskrjabinolepis spp. are presented.     

Redescription and designation of a neotype for <Emphasis Type="Italic">Aphthalmichthys kuro</Emphasis> Kuroda 1947, and its placement in <Emphasis Type="Italic">Callechelys</Emphasis> (Anguilliformes: Ophichthidae)

Aphthalmichthys kuro, described by Kuroda (1947) based on a specimen from Suruga Bay, Japan, is referred to Callechelys (family Ophichthidae, subfamily Ophichthinae). It differs from its congeners in its vertebral number (142–146), tail length (2.0–2.1 in TL), and coloration (brownish black). The holotype is lost; we herein designate a neotype and redescribe and illustrate the species based on 5 specimens from Japan and 2 from Taiwan.     

Are fish early growth and condition patterns related to life-history strategies?

Life-history studies provide a global framework for comparison of fish species responses and trade-offs facing ecological and environmental constrains. A broad comparison among fishes’ early growth and condition traits is performed in order to determine ecological patterns of early development regarding latitudinal distribution, habitat use and life-history strategies. Based on Winemiller and Rose (1992) classification of life-history strategies, data on early growth and condition indices of 46 fish species worldwide was analysed. Available information on fishes’ early features, namely first year length percentage (relative to species maximum theoretical length), age at maturation and Fulton’s condition index (K), provided a good segregation of species by latitudinal distribution and habitat use, and evidenced the categories of the three-endpoint model. Higher larvae and juvenile growth rates and condition indices (K, mean RNA–DNA ratios and protein contents) were associated with tropical and temperate fish species that occur in complex or variable habitats (respectively coral reefs and estuaries). These species selected for the opportunistic and periodic strategies, investing highly in rapid growth in order to increase survival probability to counter high mortality rates during early stages or unstable habitat conditions. Later age at maturation, slower larvae and juvenile growth as well as lower mean condition indices were consistent with fish species from more stable or predictable environments, as polar regions and freshwater habitats, which selected for the equilibrium strategy. Nonetheless, differences in energy allocation strategies during early stages were not observed, evidencing the scarcity of available data regarding condition indices and/or the importance of integrating life-history intermediate strategies. Future research into condition indices and other physiological processes, for a broader set of species and for a wider latitudinal and habitat range including seasonal variability (particularly for species from tropical and polar regions), is essential to better understand or test current theories of species ecological patterns. The use of direct quantitative measures of young fishes’ metabolic investment and fitness constitutes a new approach for life-history studies, and should be fundamental for predicting species’ responses to acute environmental or human constrains, especially in a global climate change scenario that is expected to affect distribution and abundance of fish species worldwide.     

The Invasion of the Chinese Mitten Crab (<Emphasis Type="Italic">Eriocheir sinensis</Emphasis>) in the United Kingdom and Its Comparison to Continental Europe

Owing to its catadromous lifestyle, the Chinese mitten crab, Eriocheir sinensis, allows comparison between a coastal and an inland biological invasion of the same species. Information about the distribution of this species in the United Kingdom has been collected from sightings made by governmental agencies, The Natural History Museum (London) collection, literature, and from the general public. This information indicated that the range of the species has expanded since the species’ arrival in 1973. The spread has been most marked along the east coast northwards to the river Tyne, on the south coast westwards to the river Teign. The expansion range was quantified and compared using geographic information software, and then compared to recorded spread in Europe. Mitten crabs dispersed along the coast at an average rate of 78 km per year (1976–1999), with a recent sharp increase to 448 km per year (1997–1999). These values are comparable with the historic outbreak in continental Europe where the average rate of dispersion along the Baltic Sea coast (1928–1935) was 416 km per year. Comparable figures for the North Sea coast (1923–1954) were 75 km per year with a peak of 168 km per year in 1927–1937. The upstream spread along rivers in the United Kingdom was 16 km per year in 1973–1998 with a marked increase since 1995 to 49 km per year (1995–1998). These data, in combination with population data published for the river Thames, indicate that the population has been increasing since the early 1990s, causing further range expansion into previously uninvaded river systems. The comparison of the spreading behaviour of the ongoing invasion in the United Kingdom with the historic invasion in northern Europe suggests that E. sinensis in future has the potential to establish itself in all major UK estuaries.     

A new species of <Emphasis Type="Italic">Penopus</Emphasis> (Teleostei: Ophidiidae) from the Ryukyu Trench,the West Pacific,with remarks on <Emphasis Type="Italic">Penopus microphthalmus</Emphasis>

The bathyal genus Penopus (Ophidiidae) is revised based on 23 specimens. One specimen from the Ryukyu Trench represents a new species, Penopus japonicus. The remaining 22 specimens were found on both sides of the Atlantic Ocean, and neither 28 meristic and morphometric characters nor several morphological characters show any differences between the two populations. This supports Séret (1988), who considered Penopus macdonaldi Goode and Bean 1896 a junior synonym of Penopus microphthalmus (Vaillant 1888). The new Japanese species can be separated from the Atlantic species by having fewer rays in the dorsal (117 vs. 135–158) and anal (89 vs. 106–122) fins, the squamation of the head restricted to the middle part of the preopercle versus the squamation covering the dorsum, preopercle and the opercle in part, a distinct spine behind the posterior nostril versus the spine hardly visible and 9 spines on the hind margin of the preopercle versus 4–7 spines.     

Growth patterns of Mytilopsis leucophaeata,an invasive biofouling bivalve in Europe

Abstract

For the first time, growth of Mytilopsis leucophaeata, an important European fouling species, was investigated. By means of growth cages, individual shell growth of three cohorts, with, respectively, initial shell lengths of ≤5 mm, 10 mm and 15 mm, was monitored in the harbour of Antwerp, Belgium, during 2003 – 2004. M. leucophaeata followed an oscillatory growth pattern with a single summer growing period per year (May to August). Growth decreased during wintertime, but never ceased completely. M. leucophaeata has an average growth rate of <3 – 6 mm year^?1. Temperature was found to be the main environmental factor affecting growth. The von Bertalanffy growth function was used to model growth of individuals ≤5 mm, resulting in L_∞ = 16.7 mm and K = 0.56. Based on a combination of growth of all three cohorts, the hypothetical growth of an average individual mussel could be modelled over a 5-year period, resulting in a maximum length >19 mm with a growth rate of 0.41. Its longevity (more than 5 years) and the positive effect of higher water temperatures on growth, combined with its high resistance to chlorination, provides M. leucophaeata with a high potential for severe and long-lasting biofouling     

Comparative toxicity of chlorine and peracetic acid in the biofouling control of Mytilopsis leucophaeata and Dreissena polymorpha embryos (Mollusca,Bivalvia)

Chlorination is the most common antifouling procedure, but the search for alternatives is ongoing. Although concentrations that kill adults will also be effective against larvae, it is advisable to evaluate the toxicity of any candidate toxicant against the combatable life stage. For mussels, the earliest life stages are the most vulnerable ones and thus may require the lowest doses biocides. Since the period of larval presence is restricted to a couple of months, a pointed dosage of biocides during this period will be as effective as a continuous dosage throughout the year. This study reports on the lethal acute toxicity of sodium hypochlorite and peracetic acid to 4 h old embryos of Mytilopsis leucophaeata and Dreissena polymorpha. Chlorination was found effective against M. leucophaeata from a concentration of 0.6 mg/l onwards, even at short exposure times. Commercial peracetic acid showed to be a very good alternative in both species although the most appropriate level still has to be determined.     

Attack responses of the southern Australian whelk, Lepsiella vinosa (Lamarck, 1822) (Gastropoda: Muricidae), to novel bivalve prey: an experimental approach

In their native ranges, muricid gastropods feed on similar prey, often bivalves and barnacles, which they usually drill. Throughout its wide southern Australian distribution, the intertidal Lepsiella vinosa feeds on a range of prey from barnacles and littorinid gastropods in the southeast to mussels in the southwest. A number of muricids have been introduced throughout the world, either with oysters or in ship ballast water. In their new environments, they switch to native prey but their feeding responses to them have never been studied in the laboratory. The object of this study was to study the feeding of L. vinosa on a suite of non-native species. Australian Lepsiella vinosa was taken to Hong Kong, offered five different possible prey species and allowed to feed to satiation for many weeks. Replacement of consumed prey items by similar-sized conspecifics was undertaken until trends emerged. Lepsiella vinosa readily attacked thick- and thin-shelled bivalves from Hong Kong’s sandy beaches, Anomalocardia squamosa and Caecella chinensis, respectively, and from rocky shores, Septifer virgatus and Hormomya mutabilis, again respectively. It attacked them all, as it does its major prey item, Xenostrobus inconstans, in its native southwestern Australia, by either drilling or marginal probing with its proboscis. It also preferred intermediate sized prey (10–15 mm shell length), as with its natural prey. It quickly attacked the sandy shore species, and Hormomya mutabilis and Mytilopsis sallei, the latter two being closely similar in shell form and size to its natural prey X. inconstans. Hormomya mutabilis was the most favored prey, and was most similar in shell form and thickness to X. inconstans. This study therefore suggests that if introduced elsewhere, L. vinosa could radically affect intertidal community structure.     

Phylogenetic Position of <Emphasis Type="Italic">Corchoropsis</Emphasis> Siebold &; Zucc. (Malvaceae s.l.) Inferred from Plastid DNA Sequences

Recent phylogenetic research suggests that Malvaceae s.l. comprises formerly Tiliaceae, Byttneriaceae, Bombacaceae, and Sterculiaceae. Corchoropsis is traditionally included in Tiliaceae or Sterculiaceae and is distributed in China, Korea, and Japan. One to three species have been recognized for this genus. Phylogenetic relationships among the Malvacean taxa have been intensively studied with molecular data, and the evolution of their morphological characteristics has been re-interpreted accordingly. However, no Corchoropsis species have been included for their phylogenetic position. Here, three chloroplast coding regions—rbcL, atpB, and ndhF, from Corchoropsis psilocarpa and Corchoropsis crenata—were amplified and sequenced, then compared with other Malvacean taxa. This analysis of the three plastid gene sequences now places Corchoropsis species in Dombeyoideae, as previously proposed by Takeda (Bull Misc Inform Kew 365, 1912), Tang (Cathaya 4:131–150, 1992), and Bayer and Kubitzki (2003). Within Dombeyoideae, Corchoropsis forms a strongly supported sister relationship with the Dombeya–Ruizia clade.     

Species of <Emphasis Type="Italic">Teratorhabditis</Emphasis> (Osche, 1952) Dougherty, 1953 (Nematoda:Rhabditida) from sewage and manure samples in India including a discussion on relationships within the genus

Three species of the genus Teratorhabditis (Osche, 1952) Dougherty, 1953 collected from sewage and manure samples of North India are described. The species T. andrassyi Tahseen & Jairajpuri (1988) synonymised by Sudhaus (1991) has been reinstated and is characterized by oviparous females having bluntly rounded lips with labial sensilla not raised beyond labial contour; lip margins and axils heavily cuticularised; three lines in lateral field; considerably wide and tubular stoma having swollen metastegostomal walls with prismatic inner lining bearing five teeth on each plate; basal bulb with usually single haustrulum; tail conoid with a terminal spike; eggs with ridged shells and males with 40–45% distally-fused spicules and a smooth, non-indented bursa having slender papillae in 2/4 + p + 3 configuration. Other populations identified as T. synpapillata and T. palmarum Gerber and Giblin-Davis (1990) are reported for the first time from India. An identification key to species is provided. Relationships within the genus are discussed. Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users Handling editor: K. Martens     

Hypericum sp.: essential oil composition and biological activities

Phytochemical composition of Hypericum genus has been investigated for many years. In the recent past, studies on the essential oils (EO) of this genus have been progressing and many of them have reported interesting biological activities. Variations in the EO composition of Hypericum species influenced by seasonal variation, geographic distribution, phenological cycle and type of the organ in which EO are produced and/or accumulated have also been reported. Although many reviews attributed to the characterization as well as biological activities of H. perforatum crude extracts have been published, no review has been published on the EO composition and biological activities of Hypericum species until recently (Crockett in Nat Prod Commun 5(9):1493–1506, 2010; Bertoli et al. in Global Sci Books 5:29–47, 2011). In this article, we summarize and update information regarding the composition and biological activities of Hypericum species EO. Based on experimental work carried out in our laboratory we also mention possible biotechnology approaches envisaging EO improvement of some species of the genus.     

Habitat selection and segregation by two sympatric lagomorphs: the case of European hares (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) and Eastern cottontails (<Emphasis Type="Italic">Sylvilagus floridanus</Emphasis>) in northern Italy

In northern Italy, the native European hare (Lepus europaeus) and the introduced Eastern cottontail (Sylvilagus floridanus) can occur together at a local scale, as a result of cottontail introduction and expansion into the European hare range. Hare populations are limited in Italy by habitat loss, diseases, and most important by overhunting, and many areas within hare range in northern Italy are undergoing increasing anthropogenic impact. Therefore, quantitative studies on resource selection and exploitation by both species will be of great interest to evaluate the degree of habitat overlap and to search for exploitation competition evidences. We studied habitat selection during resting time by both species in two areas where they occur alone and in one area where they occur together. Habitat selection by the two species was examined at micro- and macro-habitat scales during autumn–winter and spring–summer. Both species selected ecotonal zones between arboriculture stands and crops and between arboriculture stands and spontaneous vegetation (i.e., herbaceous, bush, and woody permanent species), which were the less available in the area of sympatry. No habitat shifts were evident at macro-habitat level because the two species showed a differential micro-habitat use within patches. On the whole, it seems that habitat heterogeneity promoted daytime segregation between the two species. In particular, edges between crops and canopy habitats should be improved, thus reducing chance of intra- and inter-specific encounters.     

Changes in the distribution area of vascular plants in Flanders (northern Belgium): eutrophication as a major driving force

In large parts of Western Europe agricultural intensification after World War II has led to an increased use of fertilisers. The resulting nutrient enrichment (=eutrophication) has a huge impact on the occurrence and distribution of plant species and is one of the main pressures on native plant communities. We used the distribution maps (grid size: 16 km2) of individual plant species, obtained through two consecutive survey projects (1939–1971 and 1972–2004) in Flanders (northern Belgium), to estimate the relative change in their distribution area. The comparison of changes in range size among groups of taxa classified according to habitat preference and Ellenberg indicator values, demonstrated a marked decline in distribution area in species that are characteristic for nutrient-poor habitats. To assess geographic patterns in the change of species assemblages, we calculated the mean Ellenberg N- and R-values for every grid cell during each of both survey periods. Differences between these values were analysed in relation to soil type and estimates of atmospheric nitrogen deposition. The largest shifts in Ellenberg N-values, reflecting a decline of species from nutrient-poor conditions and/or an increase of nitrophilous plants, were observed in areas with nutrient-poor, acid sandy soils and high nitrogen deposition rates. Hence, shifts in species composition were modulated by geographic variation in soil type and levels of nitrogen deposition. As the levels of atmospheric nitrogen deposition are still very high in Flanders, it is likely that species from nutrient-poor habitats such as heathlands, will further decline in the near future. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Nomenclature—Lambinon et al. (2004) for vascular plants.