TECTONIC CONTROLS ON FLUVIAL LANDSCAPE DEVELOPMENT IN CENTRAL-EASTERN PORTUGAL: INSIGHTS FROM LONG PROFILE TRIBUTARY STREAM ANALYSES

This study examines the long profiles of tributaries of the Tejo (Tagus) and Zêzere rivers in central eastern Portugal (West Iberia) in order to provide new insights into the patterns, timing and controls on drainage development during the Pleistocene to Holocene incision stage. The long profiles were extracted from lower order tributary streams associated with the trunk drainage of the Tejo River and one main tributary, the Zêzere River (Fig. 1). These streams flow through a landscape strongly influenced by variations in bedrock lithology (mainly granites and metasediments), fault structures delimiting crustal blocks with distinct uplift rates, and a base-level lowering history (tectonic uplift / eustatic). The long profiles of the tributaries of the Tejo and Zêzere rivers record a series of transient and permanent knickpoints. The permanent knickpoints have direct correlation with the bedrock strength, corresponding to the outcropping of very hard quartzites or to the transition from softer (slates/metagreywaques) to harder (granite) basement. The analyzed streams/rivers record also an older transient knickpoint/knickzone separating: a) an upstream relict graded profile, with lower steepness and higher concavity, that reflects a long period of quasi-equilibrium conditions reached after the beginning of the incision stage; and b) a downstream reach displaying a rejuvenated long profile, with steeper gradient and lower concavity, particularly for the final segment, which is often convex (Fig. 2). The rejuvenated reaches testify the upstream propagation of several incision waves that are the response of each stream to continuous or increasing crustal uplift and dominant periods of base-level lowering by the trunk drainages, coeval of low sea level conditions. The long profiles and their morphological configurations enabled spatial and relative temporal patterns of incision to be quantified for each individual tributary stream. The incision values of streams flowing in uplifted blocks of the Portuguese Central Range (PCR) (ca.380-280 m) indicate differential uplift and are higher than the incision values of streams flowing on the adjacent South Portugal planation surface – the Meseta (ca. 200 m). The normalized steepness index, calculated using the method of Wobus et al. (2006), proved to be sensitive to active tectonics, as lower ksn values were found in relict graded profiles of streams located in less uplifted blocks, (e.g. Sertã stream in the PCR), or in those flowing through tectonic depressions. Fig. 1 – Geological map of the study area. 1 – fluvial terraces (Pleistocene); 2 – sedimentary cover (Paleogene and Neogene); 3 – slates and metasandstones (Devonian); 4 – slates and quartzites (Silurian); 5 – quartzites (Ordovician); 6 – slates and metagreywackes (Precambrian to Cambrian); 7 – slates, metagreywackes and limestones (Precambrian); 8 – granites and ortogneisses; 9 – diorites and gabros; 10 - fault. SFf – Sobreira Formosa fault; Sf – Sertã fault; Pf – Ponsul fault; Gf – Grade fault. The differential uplift indicated by the distribution of the ksn values and by the fluvial incision was likely accumulated on a few major faults, as the Sobreira Formosa fault (SFf), thus corroborating the tectonic activity of these faults. Due to the fact that the relict graded profiles can be correlated with other geomorphic references documented in the study area, namely the T1 terrace of the Tagus River (with an age of ca. 1 Myr), the following incision rates can be estimated: a) for the studied streams located in uplifted blocks of the PCR, 0.38 m/kyr to 0.28 m/kyr; b) for the streams flowing on the South Portugal planation surface, 0.20 m/kyr. The differential uplift inferred between crustal blocks in the study area corroborates the neotectonic activity of the bordering faults, which has been proposed in previous studies based upon less robust data. Fig. 2 – Longitudinal profile of the Nisa stream a tributary of the Tejo River. Note the equilibrium relict profile upstream the older transient knickpoint (hatched line) and the downstream rejuvenated profile (continuous line). Legend: tKP – transient knickpoint; rKp – resistant knickpoint; Mt – schist and phyllite; Gr – granite; Hf – hornfels; Og – orthogneisse. In the inset Distance – Slope plots, fill circles correspond to the relict graded profile, crosses correspond to the rejuvenated profile located downstream the older transient knickpoint (tKP).

Bursaphelenchus pinophilus Brzeski & Baujard, 1997 (Nematoda: Parasitaphelenchinae) associated with nematangia on Pityogenes bidentatus (Herbst, 1783) (Coleoptera: Scolytinae), from the Czech Republic

The occurrence of Bursaphelenchus species in the Czech Republic is poorly known, the first report of the genus being made by Kubátová et al. (2000) who reported the association of B. eremus with the hyphomycetous microfungus, Esteya vermicola, and the bark beetle, Scolytus intricatus, collected from Quercus robur, in central Bohemia. To date, four other species have been reported from the country, namely B. fungivorus (Braasch et al., 2002), B. hofmanni (see Braasch, 2001), B. mucronatus (see Braasch, 2001) and B. vallesianus (Gaar et al., 2006). More recently, a survey for Bursaphelenchus species associated with bark- and wood-boring insects in the Czech Republic identified B. pinophilus Brzeski & Baujard, 1997 from the Moravia region. Although this represents a new country record, it was also associated with nematangia on the hind wings of a new insect vector. A total of 404 bark- and wood-boring insects were collected from declining or symptomatic trees and screened for the presence of Bursaphelenchus. Bark and longhorn beetles were captured manually after debarking parts of the trunk displaying symptoms of insect attacks. Longhorn beetle larvae were also collected together with logs cut from the trunk. Logs were kept at room temperature in the laboratory until insect emergence. Each adult insect was individually dissected in water and examined for nematodes. All nematodes resembling dauer juveniles of Bursaphelenchus were collected and identified by molecular characterisation using a region of ribosomal DNA (rDNA) containing the internal transcribed spacer regions ITS1 and ITS2. ITS-RFLP analyses using five restriction enzymes (AluI, HaeIII, HinfI, MspI, RsaI) were performed to generate the species-specific profile according to Burgermeister et al. (2009). Species identification was also confirmed by morphological data after culture of the dauers on Botrytis cinerea Pers. ex Ft., growing in 5% malt extract agar. During this survey, only species belonging to the Curculionidae, subfamily Scolytinae, revealed the presence of nematodes belonging to Bursaphelenchus. Dauers of this genus were found aggregated under the elytra in nematangia formed at the root of the hind wings (Fig. 1). The dauers were identified from 12 individuals of Pityogenes bidentatus (Herbst, 1783) (Coleoptera: Scolytinae) collected under the bark of Pinus sylvestris trunks. Each insect carried ca 10-100 dauers. The ITS-RFLP patterns of the dauers so obtained confirmed the identification of B. pinophilus associated with this insect species. Bursaphelenchus pinophilus has been found mainly in Europe and has been reported from various countries such as Poland (Brzeski & Baujard, 1997), Germany (Braasch, 2001), and Portugal (Penas et al., 2007). The recent detection of this species associated with dead P. koraiensis in Korea (Han et al., 2009) expands its geographical distribution and potential importance. It has been found associated only with Pinus species, but very little is known about the insect vector. The bark beetle, Hylurgus ligniperda, was initially suggested as the insect vector by Pe-nas et al. (2006), although the nematode associated with this insect was later reclassified as B. sexdentati by morphological and molecular analysis (Penas et al., 2007). According to the literature, P. bidentatus has been cited as a vector of Ektaphelenchus sp. (Kakuliya, 1966) in Georgia, and an unidentified nematode species in Spain (Roberston et al., 2008). Interestingly, B. pinophilus was found in the nematangia formed at the root of the hind wings of P. bidentatus. Although this phenomenon is not so common in other Bursaphelenchus species, B. rufipennis has been found recently in such a structure on the hind wings of the insect Dendroctonus rufipennis (Kanzaki et al., 2008). Although other nematode species (e.g., Ektaphelenchus spp.) are frequently found associated within the same nematangia (see Kanzaki et al., 2008), in this particular case, only dauers of B. pinophilus were identified. The association between B. pinophilus and P. bidentatus represents the first report of this biological association and the association with the Scolytinae strengthens the tight and specific links between this group of Bursaphelenchus species and members of the Scolytinae (Ryss et al., 2005).