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Age matters: substrate-specific colonization patterns of benthic invertebrates on installed large wood

Age matters: substrate-specific colonization patterns of benthic invertebrates on installed large... Aquat Ecol (2020) 54:741–760 https://doi.org/10.1007/s10452-020-09772-y(0123456789().,-volV)(0123456789().,-volV) Age matters: substrate-specific colonization patterns of benthic invertebrates on installed large wood . . Florian Dossi Patrick Leitner Wolfram Graf Received: 24 December 2019 / Accepted: 23 April 2020 / Published online: 6 May 2020 The Author(s) 2020 Abstract Large wood (LW) is an indispensable Our results show that (1) installed LW serves as an element in riverine ecosystems, especially in lower abundantly and heterogeneously colonized habitat, (2) river parts. The presence of LW significantly shapes the state of decay of LW pieces significantly affects local hydraulics, morphology, the nutrient budget; benthic invertebrate colonization in terms of density promotes overall river dynamics; and additionally and diversity and (3) even rare or threatened taxa presents a unique habitat for numerous benthic closely associated to LW were abundantly present on invertebrate species. Therefore, LW is recognized as the installed logs, emphasizing the suitability of the valuable asset for river restoration measures. Experi- chosen approach. ences from previous projects show that ecological responses on LW implementation measures vary Keywords Macroinvertebrates  Xylal  State of greatly. That complicates comparisons and estima- decay  River gradient  LW  Wood condition tions on the success of planned measures. Method- ological inconsistencies and thus reduced transferability of the results is one major issue. Additionally, wood quality aspects are suspected to Introduction be important factors affecting benthic invertebrate colonization patterns. The focus of this study is Large wood (LW) is a key component of natural river therefore to consistently assess the ecological signif- ecosystems. Previous studies have already stressed the icance of installed LW and concrete samples of similar beneficial effects of instream wood structures on local size and shape in terms of benthic invertebrate river hydraulics (e.g., Shields et al. 2001; Mutz 2003; colonization and to further test, if the condition of Manners et al. 2007), hydromorphology (e.g., Gurnell wood affects the benthic invertebrate colonization. et al. 1995, Kail et al. 2007, Blanckaert et al. 2014), nutrient balance (e.g., Bilby and Bisson 1998; Gurnell et al. 2005; Flores et al. 2011) and habitat diversity (e.g., Dudley and Anderson 1982; Hering and Reich Handling Editor: Telesphore Sime-Ngando. 1997). Submerged LW provides essential habitats F. Dossi (&)  P. Leitner  W. Graf which are existential for many xylobiont species IHG - Institute of Hydrobiology and Aquatic Ecosystem (Anderson et al. 1978; Hoffmann and Hering 2000) Management, BOKU - University of Natural Resources and are of increasing importance along the river course and Life Sciences, Gregor-Mendel-Strasse 33, (Dossi et al. 2018). LW further offers vital aquatic– 1180 Vienna, Austria e-mail: florian.dossi@boku.ac.at 123 742 Aquat Ecol (2020) 54:741–760 terrestrial interface areas and oviposition sites, signif- Smith and Smock 1992; Benke 1998). Besides natural icantly promoting the reproductive success of mer- fluctuations of densities due to riverine characteristics olimnic invertebrate species (Dudley and Anderson on local up to regional scales or methodological 1982; Sweeney 1993; Hoffmann and Hering 2000). inconsistencies, especially regarding the quantifica- Due to the wide variety of beneficial aspects, LW is tion of LW pieces and related individual densities, known to generally promote the density and diversity variations due to differing wood quality aspects, such of fish and aquatic invertebrate species (e.g., Dudley as hardness, species and condition, were discussed as and Anderson 1982; Copp 1992; Hering and Reich potentially important factors (Benke and Wallace 1997; Hoffmann and Hering 2000; Pilotto et al. 2003). That suggests a type specific colonization of 2014, 2016) and therefore presents a valuable and wood substrate, especially considering general sub- cost-effective asset for river restoration measures, strate selection processes of invertebrates substantially especially given the large amount of morphologically determining species richness, composition and density degraded river sections and comparably low costs to in freshwater ecosystems (Minshall 1984). Wood conventional measures (Kail and Hering 2005; Kail quality aspects were focused by only a limited number et al. 2007). Rough estimates assume that even in of studies, but most results indicate that the type and densely populated areas such as Central Europe, quality of instream LW affect invertebrate coloniza- approximately one-third of the degraded river sections tion patterns (e.g., Anderson et al. 1978; Kaufman and King 1987; Magoulick 1998; McKie and Cranston could be restored by reintroducing LW structures (Kail and Hering 2005). 1998; O’Connor 1991;Spa¨nhoff et al. 2000). A profound understanding of LW and ecosystem The aim of this study is therefore (1) to consistently interactions is a prerequisite for efficient implemen- assess the general ecological value of LW structures of tations in river management practices. Even though installed wood and concrete samples of comparable LW and its function in river ecosystems have been size and shape in terms of species richness and density, extensively investigated, knowledge gaps and there- (2) to investigate colonization patterns based on the fore implementation shortcomings still persist. Kail condition of the introduced LW pieces and (3) to test if et al. (2007) evaluated 50 restoration projects in the results are consistent within different river Germany and Austria involving LW placement and stretches along the longitudinal gradient of a med- found that only approximately 58% were successful. ium-sized lowland river in Austria. The authors concluded that the key to success lied in the consideration of site-specific characteristics. Hence, profound knowledge on river type specific Materials and methods wood characteristics is one important criterion to promote the success of measures. One challenge, hard Study sites to come by, is the lack of knowledge of the pristine state of LW and related ecological aspects in many Four sites along the Lafnitz River have been investi- European streams due to the long history of active gated. The Lafnitz River is one of the last medium- wood removal (Hering and Reich 1997; Hering et al. sized meandering rivers in the Central Europe with 2000). Additional studies in different areas with near-natural flow-regime and morphodynamics, ripar- remaining intact riparian vegetation and at least ian vegetation and LW accumulations along large near-natural LW dynamics are therefore of utter most parts of its course. It is therefore well suited to study importance to improve the understanding of LW and the importance of LW and its interactions with biota. biota interactions. The Lafnitz lies within the Danube catchment, located Benke and Wallace (2003) called attention to in the southeastern part of Austria (Fig. 1). The river additional difficulties regarding comparability and course has an approximate length of 112 km and transferability of results from different studies. Fun- drains into the Raab River, in Hungary. The Lafnitz damental information such as reported invertebrate River has a catchment size of approximately densities span wide apart from several hundred (e.g., 2000 km at the border of Austria, making it the O’Connor 1992; Rabeni and Hoel 2000) to many 13th largest river in Austria (BMLFUW 2002; Cejka (ten-) thousands of individuals per square meter (e.g., et al. 2005). The spring is located in the federal state of 123 Aquat Ecol (2020) 54:741–760 743 Fig. 1 Overview of the project area and location of the Lafnitz River in Austria (left) and location of the investigation sites along the river course (right); overlay: Ecoregions according to Illies (1978) Styria and originates at an altitude of 940 m above sea size at each site were assessed in March 2014 prior to level (m a.s.l.). Following Illies (1978), the first 36 the installation of the samples. Mean flow velocities river kilometers are situated in the ecoregion 4-Alps, and water depths were derived from averaged transect whereas the following section lies in ecoregion measurements. Grain size distribution were based on 11-Hungarian Plains (Fig. 1). The mean annual the choriotope assessment (Multi-Habitat-Sampling- discharge of the Lafnitz River spans from 2.6 m /s Method (MHS), AQEM Consortium 2002). in the upper section (near Site A) to approximately 6.3 m /s in the lower section near the town Dobersdorf Sample characteristics, sampling design (near Site E). and laboratory work The sampling sites were chosen to cover a variety of different river characteristics along the longitudinal For this colonization experiment three different types gradient to (1) allow general statements on the of substrate, comparable in size, were installed at each potential ecological benefits of installed wood struc- sampling site: (1) concrete bars, (2) fresh logs and (3) tures as well as to (2) assess possible variations in rotten logs (Figs. 2, 3). different river sections. At all four sites naturally The concrete bars were used to mimic lithal deposited large wood accumulations were present to substrates of comparable size and shape to the logs. ensure a comparable colonization potential. The dimensions of each piece were 50 cm 9 5cm 9 Samples have been taken at four sites along the 20 cm (length 9 width 9 height). In each concrete river course (see Table 1), whereas the most upstream bar, a whole was drilled and a rope was attached to be site is located at 648 m a.s.l. and the most downstream fixed at nearby trees (see Figs. 4, 5). site at 244 m a.s.l. The sites were chosen based on All xylal substrate samples (fresh and rotten) significant changes of river characteristics such as originate from the Lafnitz River and were sampled slope, discharge (stream order according to Strahler in February 2014 near the village of Neustift, in the (1957)), substrate composition and morphological middle section of the river. To ensure comparability, characteristics. Figure 1 gives an overview of the site only previously submerged logs with a diameter locations along the river course. Site-specific charac- between 200 mm and 300 mm were used in this teristics are shown in Table 1. study. Each log was cut to a length of 630 mm and Abiotic characteristics at each site including mean analogous to the concrete bars; a hole was drilled in flow velocity, mean water depth and dominant grain each sample log and a rope was attached to fix the log 123 744 Aquat Ecol (2020) 54:741–760 Table 1 Overview of site characteristics including altitude 1957), average water depth (m), average flow velocity (m/s) (meters above sea level), distance from spring (km), mean river and average river width (m) slope (%), dominant grain size (cm), Strahler number (Strahler Site Altitude Distance from Slope Dominant grain Strahler Average water Flow Average River (m asl) Spring (km) (%) size (cm) Number depth (m) Velocity (m/ width (m) s) A 648 8.7 1.7 6.3–20 4 0.2 0.3 8 B 438 26.1 0.9 6.3–40 5 0.3 0.6 15 C 324 52.1 0.3 2–6.3 5 0.5 0.3 10–20 D 244 100.4 \0.1 2–6.3 6 0.6 0.35 20 Fig. 2 Picture of the Lafnitz River at site A (left) and site B (right) Fig. 3 Picture of the Lafnitz River at site C (left) and site D (right) at the chosen sites (see Fig. 4). All sampled logs were based on the summarized decay categories of Robin- classified into the categories ‘‘rotten’’ and ‘‘fresh’’ on son and Beschta (1990) (class 1–3: fresh; class 4–5: site (examples see Fig. 6). Wood classification was rotten) and Grette (1985) (class 1–4 fresh; class 5–6: 123 Aquat Ecol (2020) 54:741–760 745 Fig. 4 Sample schematics and dimensions for fresh and rotten logs (left) and concrete samples (right) Fig. 5 Example of one installed concrete bar at site A rotten). Details on the classification characteristics of The surface area of the sampled LW pieces varied 2 2 wood pieces are shown in Annex see Table 5, 6. between 0.16 m and 0.32 m . The volume of each Length, width and volume of each LW piece was LW piece was measured in an overflow tank and 3 3 measured, and the surface area was calculated using varied between 2.8 dm and 10.2 dm . In total, there the formula of a simplified shape of a truncated cone: were 24 wood samples, corresponding to a total surface area of approximately 5.2 m (12 rotten logs: 2 2 S ¼ r  p þ p  R þ p  h ðÞ r þ R 2 2 2.5 m ; 12 fresh logs: 2.7 m ). All wood sample characteristics are summarized in Table 2. After where S = surface area, r = radius 1, R = radius 2, classification and preparation of the samples, all wood h = sample height. samples were rinsed, cleaned and carefully examined 123 746 Aquat Ecol (2020) 54:741–760 Fig. 6 Example of fresh (left) and rotten logs (right) installed at the Lafnitz River for remaining organisms to ensure a comparable pre- organisms were removed, and each log or concrete bar colonization state of all samples. The logs were then was subsequently reintroduced to the river. Benthic stored in black plastic bags until installation. samples were taken on three different dates in April 2014, June 2014 and March 2015. All samples were The logs and concrete bars were installed in a similar manner at each site. At each of the four sites, taken within 1 day. Due to the long exposition time three concrete bars and six logs (three fresh/three and the high morphodynamics at the Lafnitz River, not rotten) were installed in early March 2014. Three long all samples were consistently available for analysis. guiding ropes (approximately 15–20 m) were tied to Single logs or concrete bars which occasionally riparian trees, and three substrate samples (either logs stranded or were entirely covered with sediments or concrete bars) were subsequently bound to each were excluded from subsequent analysis (Details see guiding rope. The allocation of logs and concrete bars Table 2). on the guiding ropes was randomized, and all samples Sampling was performed with a standardized were installed in comparable orientations, parallel to 500-lm mesh-size kick-net with a frame size of the flow. However, a slight displacement or motion of 25 9 25 cm (surface area per single sample: the samples was possible. The location and orientation 0.0625 m ) and a net length of 1.2 m. Prior to of the guiding ropes was chosen to ensure comparable sampling, all LW pieces were carefully put into the habitat characteristics for each sample within each site kick-net to avoid organisms from drifting. Benthic (e.g., flow velocity, surrounding substrate invertebrates were then carefully brushed and washed composition). from the LW piece into the kick net. All organisms Details on the habitat parameters at each concrete were preserved with formaldehyde (4%) (Table 3). bar or log are given in Table 2. All single logs and The Screening-Taxa list (Ofenbo¨ck et al. 2010) was concrete bars were installed with a minimum distance used as a basis for identification. In many cases of 2 m to each other to allow undisturbed sampling Ephemeroptera, Plecoptera and Trichoptera taxa could conditions at each log or concrete bar. To ensure be identified to a lower taxonomic level (genus/ comparable colonization potentials, all samples had species), whereas Diptera taxa were mainly identified contact with the river bottom. to family level. Oligochaeta were not identified Sampling was performed in spring 2014 and 2015. further. All taxa were counted and weighed (wet To ensure comparable exposition times, all logs and weight). Specimens were then fixed in 70% ethanol concrete bars were cleaned and carefully examined and stored. 6 weeks prior to each sampling run. All attached 123 Aquat Ecol (2020) 54:741–760 747 Table 2 Overview of the sample characteristics including exact sampling point and the availability (only fully submerged 3 2 sample ID, the site, volume (dm ), surface (cm ), the state (f- samples from each sampling date were considered for analysis) fresh, r-rotten, c-concrete), the flow velocity (mean flow of samples at the each sampling date (1-23.04.14, 2-28.06.14, velocity at exact sampling point in m/s), dominant substrate at 3-18.03.15) 3 2 Sample ID Site Volume (dm ) Surface (cm ) State Flow velocity (m/s) Substrate 1 2 3 W01 A 4.8 2161 r 0.3 Psammal ??? W02 A 5.1 2131 r 0.3 Microlithal ??? W03 A 7.2 2416 r 0.1 Psammal ??? W04 A 3.9 1981 f 0.4 Psammal/Akal ??? W05 A 6.1 2462 f 0.2 Psammal ??? W06 A 10.2 3178 f 0.3 Psammal ??? C01 A 5 2700 c 0.7 Akal/Microlithal ??? C02 A 5 2700 c 0.1 Psammal ??? C03 A 5 2700 c 0.7 Akal/Microlithal – ?? W07 B 4.5 2027 r 0.8 Microlithal ??? W08 B 4 1911 r 0.8 Microlithal ??? W09 B 3.9 1880 r 0.8 Microlithal ??? W10 B 5.2 2314 f 0.8 Akal/microlithal ??? W11 B 7 2594 f 0.4 Microlithal ??? W12 B 4.9 2123 f 0.7 Akal/microlithal ??? C04 B 5 2700 c 0.8 Akal/microlithal ??? C05 B 5 2700 c 0.7 Akal/microlithal – ?? C06 B 5 2700 c 0.7 Akal/microlithal ??? W13 C 6 2350 r 0.2 Psammal – ?? W14 C 5.9 2332 r 0.5 Akal/psammal ? – ? W15 C 4 1927 r 0.3 Psammal ??? W16 C 7.1 2600 f 0.2 Psammal – ?? W17 C 2.8 1678 f 0.3 Psammal ??? W18 C 4.2 1952 f 0.2 Psammal ??? C07 C 5 2700 c 0.4 Akal – ?? C08 C 5 2700 c 0.5 Akal/microlithal ? – ? C09 C 5 2700 c 0.5 Akal ? – ? W19 D 3.6 1811 r 0.6 Psammal/akal ??? W20 D 4 1898 r 0.6 Psammal/akal ??? W21 D 3 1629 r 0.6 Psammal/akal ??? W22 D 4.8 2129 f 0.6 Psammal/akal ??? W23 D 6.8 2559 f 0.6 Psammal/akal ??? W24 D 4.1 1944 f 0.6 Psammal/akal ??? C10 D 5 2700 c 0.6 Psammal/akal ?? – C12 D 5 2700 c 0.6 Psammal/akal ??? C13 D 5 2700 c 0.6 Psammal/akal ??? Data analysis test equality of variances between the substrate groups and the Shapiro–Wilk Test to test on normality of the Statistical analyses were performed with the Software data. All abundance and biomass data (from all R-Studio 1.1.456. The Levene’s test was applied to substrates) were converted to densities (Ind/m ). 123 748 Aquat Ecol (2020) 54:741–760 Results Table 3 Overview of the total number of specimen, taxa and families observed at each site and substrate type In total * 43,000 benthic invertebrate specimen and A BCD 108 taxa from 52 families and 12 orders were collected No of specimen (see Annex in Table 7). The most abundant orders were Concrete 3338 2578 796 1918 Diptera (* 32%; mainly Chironomidae), Crustacea Fresh 2221 3270 3887 3562 (* 27%; only Gammarus fossarum) and Trichoptera Rotten 3154 5819 6703 4746 (22%; mainly Limnephilidae). Ephemeroptera com- No of taxa prised a share of * 13% and Plecoptera of * 3%. Concrete 33 29 32 42 Benthic invertebrate species richness gradually Fresh 36 36 34 48 increased along the river course from 49 taxa at site A Rotten 42 49 43 47 to 71 taxa at site D. Taxa richness per sample ranged No of families from 4 to 27 taxa. Details on the number of specimens, Concrete 22 19 22 27 taxa and families at each site and substrate type are Fresh 27 24 23 32 summarized in Table 3. Rotten 30 30 23 31 Density and diversity differences between substrate types Abundances for cluster and NMDS analyses were log Benthic invertebrate colonization showed significant (n ? 1) or presence/absence transformed. For cluster differences between the substrate types (see Fig. 7). analysis ‘‘Bray–Curtis’’ distance measure and Abundance, biomass and species richness were signifi- ‘‘Ward.D2’’ linkage method was applied. NMDS cantly higher (Wilcoxon test:a = 0.01) on wood samples analysis (Kruskal, 1964) was performed with ‘‘Sør- 2 2 (Ind/m : l = 2436.9 ± 241.3; g/m : l = 31.7 ± ensen (Bray–Curtis)’’ distance measure. 5.5; No. of taxa: l =15.5 ± 0.6) compared to concrete 2 2 The affinity of species or taxa to a particular samples (Ind/m : l = 1102.2 ± 139.0; g/m : l = substrate type was performed combining two different 8.1 ± 1.4; No. of taxa: l = 11.8 ± 0.6). Wood sample methods: showed a generally wider variation, as shown by the higher standard errors. Further, significant differences 1. Identification of taxa exclusively present on one 2 2 between fresh (Ind/m : l = 1723.9 ± 219.8; g/m : substrate type and l = 23.7 ± 4.8; No. of taxa: l = 13.9 ± 0.8) and rotten 2. identification of taxa which were significantly 2 2 wood samples (Ind/m : l = 3194.5 ± 401.6; g/m : more abundant on one substrate type based on the l = 40.2 ± 10; No. of taxa: l = 17.3 ± 0.9) were results of an ‘‘Indicator species Analysis (ISA). evident. Only biomass differences between fresh and All taxa with an Indicator value (IV) [ 25 rotten wood did not show significant results. Still, a (Dufrene and Legendre 1997) and a p value consistent pattern with lowest benthic invertebrate of B 0.05 were considered as significantly over- density and diversity on concrete, followed by fresh represented on the corresponding substrate type. wood and peak values on rotten wood, was clearly visible. This two-level approach was chosen in order to The most prominent density differences among the consider that even though some taxa significantly substrate types were found for Gammarus fossarum benefit from the presence of wood are not necessarily 2 2 (concrete: l = 102.2 Ind/m ; fresh: 648 Ind/m ; limited to wood as a habitat. The sole information on rotten: 1086 Ind/m ) (see Fig. 8: left). Diptera (mainly exclusive occurrences would therefore be insufficient. represented by Chironomidae taxa) showed compara- ble densities on concrete bars (487 Ind/m ) and fresh logs (418 Ind/m ) while being distinctly more abun- dant on rotten logs (1206 Ind/m ). Lowest Trichoptera densities were recorded on concrete (167 Ind/m ), followed by fresh (433 Ind/m ) and highest on rotten 123 Aquat Ecol (2020) 54:741–760 749 2 2 Fig. 7 Boxplot based on the number of individuals/m (left), biomass/m (middle) as well as the number of taxa (right) recorded at each single log or concrete bar; f—fresh wood, r—rotten wood; *p B 0.05, **p B 0.01, -p C 0.05 Fig. 8 Mean number of individuals of each order found on concrete bars (c), fresh (f) and rotten logs (r) with additional display of the standard error of the mean (left) and total number of taxa in each order (right) logs (673 Ind/m ). For Ephemeroptera and Plecoptera Trichoptera: Glossosoma sp.). In total, 39 taxa were taxa no particular trend was visible. The number of only found on wood samples (e.g., Plecoptera: taxa for all orders except for Crustacea (G. fossarum) Agnetina elegantula and Coleoptera: Macronychus showed a minor but consistent pattern as shown in quadrituberculatus). A further differentiation between Fig. 8: right. The number of taxa for Ephemeroptera, fresh and rotten wood samples showed nine taxa Plecoptera, Trichoptera and Diptera gradually exclusively present on fresh (e.g., Trichoptera: Rhy- increases from concrete bars to fresh and rotten logs acophila tristis and Silo pallipes) and 17 exclusively (see Fig. 8: right). on rotten wood samples (e.g., Trichoptera Hydropsy- Differences in species distribution between the che bulbifera and Lype phaeopa) (see Fig. 9: left). substrate are well reflected in the number of substrate- Results from each site and run separately reveal a specific taxa (see Table 4). Considering all samples, comparable pattern to the overall analysis (see Fig. 9: three taxa were exclusively present on concrete bars middle). Lowest number of exclusive taxa were found (e.g., Ephemeroptera: Epeorus alpicola and on concrete (l = 2.7 ± 0.6) followed by fresh- 123 750 Aquat Ecol (2020) 54:741–760 Fig. 9 Bar chart showing the number of exclusive taxa on each run, c—concrete, w—wood, f—fresh wood, r—rotten wood substrate type from all sites (Site A to Site C) (left); number of (middle); number of exclusive taxa on each substrate type at exclusive taxa on each substrate type for each single sampling each site (Site A to Site C) (right) (l = 3.8 ± 0.9) and rotten wood samples allocations patterns based on the substrate type are (l =6±0.6). visible. Wood samples (fresh and rotten) are mainly The results of the indicator species analysis are distributed in the middle of the NMDS plot, with shown in Table 4. The analysis was performed based rotten wood samples being generally allocated within on different taxonomic resolutions (family, genus and closer proximity to each other compared to the other species level), and results of all three taxonomic levels substrate types. Concrete samples are generally showed consistent results. Seven families were iden- oriented to the left side of the plot within each site tified, mainly represented by one dominant taxon group and rotten wood samples to the right. Fresh which was significantly more abundant on fresh and wood samples are generally oriented in between rotten wood, respectively (e.g., Trichoptera: Lepidos- concrete and rotten wood samples. tomatidae: Lepidostoma basale; Limnephilidae: Hale- Comparable results are obtained by the cluster sus sp., Rhyacophilidae: Rhyacophila s.str.sp.) and analysis (see Fig. 12) identifying four distinct groups. two families/genera significantly more abundant One group comprises all substrate types from the two exclusively on rotten wood samples (Coleoptera: upstream investigation sites (A and B), one the Hydraenidae: Hydraena sp.; Crustacea: Gammaridae: concrete samples from both lower investigation sites Gammarus fossarum). On genus/species level one (C and D), one the fresh and rotten wood samples from additional taxon was found to be significantly more site C and the last one both wood samples from site D. abundant on concrete samples (Plecoptera: Perlidae: The slight separation of the rotten wood samples at site B in the second cluster are mainly caused by distinctly Perla sp.) and one on fresh and rotten wood samples (Ephemeroptera: Heptageniidae: Heptagenia higher invertebrate densities on the samples compared longicauda). to those at sites A and B. Following the allocation of samples within the groups from the NMDS analysis, Community composition the cluster analysis shows a closer allocation of the concrete samples from site C and D to the samples of Community analysis shows a clear separation of the first two sites A and B. samples per site (see Fig. 11) indicating that site- specific river characteristics are the predominant factors determining the benthic community. In addi- tion, a distinct longitudinal pattern as well as 123 Aquat Ecol (2020) 54:741–760 751 Table 4 Overview of Order Family Genus Species Preference Type exclusive taxa (Type 1) and taxa significantly more Coleoptera Corixidae Corixidae Gen. sp. C 1 abundant on concrete bars Dytiscidae Dytiscidae Gen. sp. R 1 and fresh/rotten logs (Type Elmidae Esolus sp. F ?R1 2) Elmidae Macronychus quadrituberculatus F ?R1 Gyrinidae Orectochilus villosus F ?R1 Hydraenidae Hydraena* sp.* R 2 Crustacea Gammaridae* Gammarus* fossarum* R2 Diptera Empididae Empididae Gen. sp. R 1 Psychodidae Psychodidae Gen. sp. R 1 Tipulidae Tipulidae Gen. sp. R 1 Ephemeroptera Ephemeridae Ephemera danica R1 Heptageniidae Epeorus alpicola C1 Heptageniidae Heptagenia coerulans R1 Heptageniidae Heptagenia* longicauda* F ?R2 Leptophlebiidae Habroleptoides sp. R 1 Leptophlebiidae Habrophlebia* confusa* R2 Mollusca Tateidae Potamopyrgus antipodarum F1 Odonata Calopterygidae Calopteryx sp. F 1 Gomphidae Ophiogomphus cecilia R1 Platycnemididae Platycnemis pennipes F1 Plecoptera Chloroperlidae Chloroperla sp. F 1 Perlidae Agnetina elegantula F ?R1 Perlidae Perla* sp.* C 2 Perlodidae* Isoperla* sp* F ?R2 Taeniopterygidae Rhabdiopteryx navicula R1 Trichoptera Glossosomatidae Glossosoma sp. C 1 Goeridae Goera pilosa R1 Goeridae Silo pallipes F1 Hydropsychidae Hydropsyche bulbifera F1 Hydropsychidae Hydropsyche dinarica F ?R1 Hydropsychidae Hydropsyche siltalai R1 Hydropsychidae* F ?R2 Hydroptilidae Hydroptila sp. F 1 Lepidostomatidae* Lepidostoma* basale* F ?R2 Leptoceridae Ceraclea dissimilis R1 Leptoceridae Ylodes simulans R1 Limnephilidae* Anabolia furcata F ?R1 Limnephilidae* Chaetopteryx sp. F ?R1 Limnephilidae* Halesus* sp.* F ?R2 Limnephilidae* Melampophylax melampus R1 Polycentropodidae Cyrnus trimaculatus F1 Polycentropodidae Polycentropus flavomaculatus R1 Psychomyiidae Lype phaeopa R1 Psychomyiidae Psychomyia pusilla F ?R1 *Marks the taxonomic level of a significant Rhyacophilidae Rhyacophila s.str.sp.* F ?R2 overrepresentation Rhyacophilidae Rhyacophila tristis F1 (p B 0.05) 123 752 Aquat Ecol (2020) 54:741–760 Fig. 10 Overview of the number of individuals and biomass between the substrate types are shown in the table below: c— per m as well as the number of taxa on each substrate type for concrete, w—wood, f—fresh wood, r—rotten wood; *p B 0.05, each investigation site separately (a–d); significant differences **p B 0.01, -p C 0.05 Fig. 11 Ordination graph of NMDS analysis based on the are shown by name; all other species are only indicated by solid species composition and density from all concrete bars as well as circles to ensure readability; final stress for 2-dimensional fresh and rotten logs (sites A to D) (left) and display of species solution = 0.1912 allocations in the ordination graph (right); species with best fit Site-specific analysis consistently follow the patterns found in the overall dataset (see Fig. 10). Significant differences (Wil- A site-specific approach gives comparable results to coxon test: * = 0.05; ** = 0.01) are shown in Fig. 10. the pooled data. Abundance, biomass and taxa rich- Lowest values were generally found on concrete ness at each site and on each substrate type samples, followed by fresh and rotten wood samples. 123 Aquat Ecol (2020) 54:741–760 753 Fig. 12 Cluster dendrogram (distance measure: Bray–Curtis; group-linkage-method: Ward.D2) based on the species composition at each site (A-D) One sole exception is evident at site A showing higher adjacent lithal habitats (e.g., Benke et al. 1984; Smock abundance on concrete than on wood samples. Highest et al. 1989, 1992; O’Connor 1991; Piegay and Gurnell abundance, biomass and taxa richness, regardless of 1997; Hoffmann and Hering 2000; Spa¨nhoff et al. the substrate types, were evident at the sites B and C. 2000; Grafahrend-Belau and Brunke 2005; Milner and The most considerable colonization differences Gloyne-Phillips 2005; Coe et al. 2009; Pilotto et al. between the substrate types are evident at site C. 2016; Dossi et al. 2018). The number of exclusive taxa found at each site is The high variability of invertebrate density on illustrated in Fig. 9 (right). Lowest values were wood is difficult to compare and interpret. Main consistently observed on concrete samples with only reasons for these differences comprise the lack of a one exclusive taxon at site A, C and D and none at site common experimental and quantification approach as B. The number of exclusive wood taxa varies from site well as general river types or regional differences A to C between seven to ten taxa. At site D, a (Benke and Wallace 2003). The aim of this study considerably higher amount of 24 exclusive wood taxa therefore was to assess the ecological significance of was evident. Fresh wood samples mark an increase installed large wood compared to uniform concrete from three taxa at site A to six at site D, while only one structures for benthic invertebrate communities, with exclusive taxon was found at site B and C. On rotten particular attention on the wood condition. To over- wood samples, an increase of exclusive taxa from four come difficulties in the quantification of heterogenous at site A to eight at site D was found. instream LW pieces, we conducted our study with concrete, fresh and rotten LW of comparable size and shape. Discussion Our results generally agree with the above-men- tioned findings. A discrimination between lithal and Previous studies already stressed the general ecolog- xylal substrates showed distinctly higher benthic ical importance of instream wood structures but most invertebrate density, biomass and diversity on xylal of them focused on naturally deposited logs or wood samples. accumulations of varying sizes. Further, wood char- Besides similar patterns, the majority of the above- acteristics such as the state of decay received little mentioned studies reported higher invertebrate density attention. Detailed results varied greatly but nonethe- on xylal substrates, especially considering those from less consistently showed higher benthic invertebrate North America (see Benke and Wallace 2003). A densities and diversities on wood compared to potential underestimation due to an insufficient 123 754 Aquat Ecol (2020) 54:741–760 exposition time can be ruled out in our study. Samples Information on determining factors of specific were exposed for 6 weeks and previous studies showed benthic invertebrate colonization patterns on logs of that colonization on installed substrates peaks at different decay stages is still sparse, but the results of approximately two to 6 weeks the latest (e.g., Nilsen previous studies suggest that wood surface character- and Larimore 1973; O’Connor 1992; Spanhoff et al. istics present a decisive aspect (e.g., Kaufman and 2000). Comparisons with a previously conducted King 1987; O’Connor 1991; Phillips 1993; Phillips study at the Lafnitz River, which focused on natural and Kilambi 1994; Magoulick 1998; Spanhoff et al. instream wood (see Dossi et al. 2018) further support 2000). LW further develops a productive epixylic our results. Recorded invertebrate density and biomass biofilm which is an essential food source for grazing were on a comparable level but on average higher in taxa (Golladay and Sinsabaugh 1991; Sinsabaugh the present study on the installed logs (1655 Ind/m vs. et al. 1991), which might depend on log surface 2 2 2 2437 Ind/m ; 19.2 g/m vs. 29.6 g/m ). Similar characteristics. Conditioned wood has a softer and differences between installed and natural substrates more diverse, 3-dimensional, structure compared to have already been observed and discussed. While most other habitats. It therefore provides a comparably Spa¨nhoff and Cleven (2010) generally referred to large, smooth and more importantly complex habitat rapid initial colonization processes of newly intro- on a small spatial scale compared to rocks and fresh duced substrates resulting in above-average inverte- LW which is potentially beneficial for benthic inver- brate densities in the first weeks, Spanhoff et al. (2000) tebrate colonization (Kaufman and King 1987; added another aspect that was also observed in the O’Connor 1991; Magoulick 1998). The consistently Lafnitz River. Due to the longer residence time of higher densities and diversities on rotten logs com- naturally deposited, compared to installed logs, larger pared to fresh logs and concrete bars support these parts of their surface tend to be covered by sediments assumptions. Besides overall quantities, consistent which are therefore not (or only partially) suitable for statements on taxa-specific colonization patterns are benthic invertebrate colonization. That affects surface only possible to a limited extend. Even though most density calculations and leads to a slight underesti- taxa were found in higher quantities on rotten wood, mation of benthic invertebrate abundance on naturally only a few taxa showed significant preferences. The deposited logs. most prominent differences were found for G. fos- Site characteristics and the natural longitudinal sarum being significantly more abundant on rotten zonation of aquatic communities along the river logs. G. fossarum being a shredder with preferences course were identified as the dominant factor govern- for low flow velocities might benefit most from the ing the overall composition of the benthic invertebrate more complex surface structure of rotten wood. It community. However, overall species richness and provides flow protection as well as retention areas for density of invertebrates further depended on the wood organic matter on a small spatial scale. The softer condition as already indicated by previous studies surface might further facilitate the fragmentation and (e.g., Magoulick 1998; Spa¨nhoff et al. 2000). An processing of the woody material itself as well as additional distinction of our samples between different epixylic biofilm growth. Most taxa labeled as closely substrates revealed discrete benthic invertebrate den- associated to wood such as L. basale, M. quadritu- sity and diversity differences, with lowest values on berculatus, O. villosus or A. elegantula (Anderson concrete, followed by fresh and peak values on rotten et al. 1978; Hoffmann and Hering 2000; Dossi et al. wood. Only at site A, no clear differences were evident 2018) were found to be significantly more abundant on between the substrate types, which emphasizes the wood but did not show any preference for either fresh results of Dossi et al. (2018). Their study showed that or rotten logs. Assumptions on possible differences the importance of LW as unique habitat structure due to varying biofilm developments on the tested significantly changes along the longitudinal gradient substrate types could not be verified. Grazer taxa were of a river. While invertebrate species showed no equally dominant on the tested substrate types. significant preference for a specific substrate type in Our results further show that artificial LW intro- upper river sections, an increasing diversification of duction, even of comparably small logs as used in our the benthic communities among xylal and lithal study, comprise a valuable element in river restoration substrates was evident along the river course. measures. All samples were consistently colonized by 123 Aquat Ecol (2020) 54:741–760 755 heterogenous invertebrate taxa throughout the sam- also the species of wood becomes of particular pling period and also threatened or rare species, interest. Wood species-specific properties (e.g., firm- closely associated to wood (e.g., M. quadritubercula- ness, structure, texture, stability, chemical composi- tus, H. longicauda; A. elegantula), were frequently tion) and subsequent varying degradation and habitat found on the installed logs (Graf 1997; Bauernfeind characteristics will be investigated in a follow-up and Humpesch 2001; Graf and Kovacs 2002;Ja¨ch research. et al. 2005; Buffagni et al. 2016). Acknowledgements Open access funding provided by Our results emphasize that not only the presence of University of Natural Resources and Life Sciences Vienna LW is of importance. The suitability as a habitat (BOKU). significantly depends on the state of decay. That relates to two important, interrelated aspects, specif- Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which ically the residence time of logs in stream channels permits use, sharing, adaptation, distribution and reproduction and the species of wood. While residence time and the in any medium or format, as long as you give appropriate credit state of decay are clearly connected, species of wood to the original author(s) and the source, provide a link to the may be decisive as well. Besides possible colonization Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are preferences for certain wood species, differing degra- included in the article’s Creative Commons licence, unless dation rates (Spa¨nhoff and Meyer 2004) and thus indicated otherwise in a credit line to the material. If material is variations of the required residence time in rivers add not included in the article’s Creative Commons licence and your another scale to the research topic. These findings are intended use is not permitted by statutory regulation or exceeds of particular importance considering management the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit actions like the large-scale removal of instream LW, http://creativecommons.org/licenses/by/4.0/. which deprived rivers from an essential structural element and habitat and significant changes of riparian vegetation communities (Hering et al. 2000; Annex Hohensinner et al. 2013). Given these initial insights on the effect of wood quality on benthic invertebrate See Tables 5, 6 and 7. colonization patterns further research incorporating Table 5 Overview of decay categories; decay class: decay category according to Robinson and Beschta (1990) Decay class Bark Twigs Surface texture Shape Wood color State 1 Intact Present Intact/firm Round Original n 2 Intact Absent Intact/firm Round Original n 3 Trace Absent Smooth to some surface abrasion Round Original to darkening n 4 Absent Absent Abrasion to some holes and openings Round to oval Dark d 5 Absent Absent Vesicular with many holes and openings Irregular Dark d n not decayed, d decayed Table 6 Overview of Decay class Bark Limbs Surface texture Center State decay categories; decay class: decay category 1 Intact Present Firm Solid n according to Grette (1985) 2 Intact Absent Firm Solid n 3 Loose or absent Absent Firm Solid n 4 Absent Absent Slightly rottend Solid n 5 Absent Absent Extensively rottend Solid d 6 Absent Absent Completely rottend Solid d 7 Absent Absent Completely rottend Rotted d N not decayed, d decayed 123 756 Aquat Ecol (2020) 54:741–760 Table 7 Taxalist of all taxa found at all four sites; ‘‘c’’—concrete bars, ‘‘f’’—fresh logs, ‘‘r’’—rotten logs Order Family Taxon c f r Turbellaria Turbellaria Turbellaria Gen.sp. * * * Gastropoda Hydrobiidae Potamopyrgus antipodarum (J.E. Gray, 1843) – * – Oligochaeta Oligochaeta Oligochaeta Gen.sp. * * * Amphipoda Corophiidae Corophium sp. – * – Gammaridae Gammarus fossarum Koch, 1835 * * * Hydrachnidia Hydrachnidia Hydrachnidia Gen.sp. * * * Ephemeroptera Baetidae Baetis sp. * * * Caenidae Caenis sp. – * – Ephemerellidae Ephemerella ignita (Poda, 1761) * * * Ephemerella mucronata (Bengtsson, 1909) * * * Ephemerella notata Eaton, 1887 * * * Ephemerella sp. * * * Ephemeridae Ephemera danica Muller, 1764 – – * Heptageniidae Ecdyonurus sp. * * * Electrogena sp. – * – Epeorus alpicola (Eaton, 1871) * – – Epeorus assimilis (Eaton, 1871) * * * Heptagenia coerulans Rostock, 1877 – – * Heptagenia flava Rostock, 1877 * * * Heptagenia longicauda (Stephens, 1836) * * * Heptagenia sp. * * * Heptagenia sulphurea (Mu¨ller, 1776) * * * Heptageniidae Gen.sp. * * * Rhithrogena sp. * * * Leptophlebiidae Habroleptoides sp. – – * Habroleptoides confusa Sartori & Jacob, 1986 – – * Habrophlebia sp. – * * Paraleptophlebia sp. – * * Oligoneuriidae Oligoneuriella rhenana (Imhoff, 1852) * * * Potamanthidae Pothamantus luteus (Linnaeus, 1767) * * – Odonata Calopterygidae Calopteryx sp. – * – Gomphidae Gomphus vulgatissimus (Linnaeus, 1758) * * – Onychogomphus sp. * * – Ophiogomphus cecilia (Geoffroy In Fourcroy, 1785) – – * Platycnemididae Platycnemis pennipes (Pallas, 1771) – * – Plecoptera Chloroperlidae Chloroperla sp. – * – Siphonoperla sp. * * * Leuctridae Leuctra sp. * * * Nemouridae Amphinemura sp. * * * Nemoura/Nemurella sp. * * * Protonemura sp. * * * Perlidae Agnetina elegantula (Klapalek, 1905) – * * Dinocras sp. * * * Perla sp. * * * Perlidae Gen.sp. – * * 123 Aquat Ecol (2020) 54:741–760 757 Table 7 continued Order Family Taxon c f r Perlodidae Isoperla sp. * * * Perlodes sp. * * * Taeniopterygidae Brachyptera risi (Morton, 1896) * * * Brachyptera seticornis (Klapalek, 1902) * * * Brachyptera sp. * * * Rhabdiopteryx navicula Theischinger, 1974 – – * Heteroptera Aphelocheiridae Aphelocheirus aestivalis (Fabricius, 1803) * – * Corixidae Corixidae Gen.sp. * – – Coleoptera Dryopidae Pomatinus sp. – – * Dytiscidae Dytiscidae Gen.sp. – – * Elmidae Elmidae Gen.sp. – – * Elmis sp. * * * Esolus sp. – * * Limnius sp. * * * Macronychus quadrituberculatus Mu¨ller, 1806 – * * Gyrinidae Orectochilus villosus (Mu¨ller, 1776) – * * Helophoridae Helophorus sp. – * * Hydraenidae Hydraena sp. * * * Trichoptera Brachycentridae Brachycentrus subnubilus Curtis, 1834 * * * Glossosomatidae Glossosoma conformis Neboiss, 1963 * – * Glossosoma sp. * – – Goeridae Goera pilosa (Fabricius, 1775) – – * Goeridae sp. * * – Silo pallipes (Fabricius, 1781) * * – Hydropsychidae Cheumatopsyche lepida (Pictet, 1834) * – * Hydropsyche bulbifera Mclachlan, 1878 – * – Hydropsyche dinarica Marinkovic, 1979 – * * Hydropsyche instabilis (Curtis, 1834) * * * Hydropsyche pellucidula (Curtis, 1834) * * * Hydropsyche siltalai Do¨hler, 1963 – * * Hydropsyche sp. * * * Hydroptilidae Hydroptila sp. – * – Ithytrichia lamellaris Eaton, 1873 * * * Lepidostomatidae Lepidostoma basale (Kolenati, 1848) * * * Leptoceridae Ceraclea dissimilis (Stephens, 1836) – – * Leptoceridae Gen.sp. – * * Ylodes simulans (Tjeder, 1929) – – * Limnephilidae Allogamus auricollis (Pictet, 1834) * * * Anabolia furcata Brauer, 1857 – * * Chaetopteryx fusca Brauer, 1857 – * * Chaetopteryx sp. – * * Halesus sp. * * * Limnephilidae Gen.sp. * * * Melampophylax melampus (McLachlan, 1867) – – * Potamophylax cingulatus (Stephens, 1837) * * * 123 758 Aquat Ecol (2020) 54:741–760 Table 7 continued Order Family Taxon c f r Potamophylax rotundipennis (Brauer, 1857) * * * Odontoceridae Odontocerum albicorne (Scopoli, 1763) * * * Polycentropodidae Cyrnus trimaculatus (Curtis, 1834) – * – Polycetropodidae Polycentropus flavomaculatus (Pictet, 1834) – – * Psychomyiidae Lype phaeopa (Stephens, 1936) – – * Psychomyia pusilla (Fabricius, 1781) – * * Rhyacophilidae Rhyacophila s.str.sp. * * * Rhyacophila tristis Pictet, 1834 – * – Sericostomatidae Sericostoma sp. * * * Diptera Athericidae Ibisia marginata (Fabricius, 1781) * * * Ceratopogonidae Bezzia-Gruppe sp. – * – Chironomidae Chironomidae Gen.sp. * * * Empididae Empididae Gen.sp. – * * Limoniidae Antocha sp. ** – Hexatoma sp. * * * Limoniidae/ Limoniidae Gen.sp. – * * Pediciidae Pediciidae Dicranota sp. * * * Psychodidae Psychodidae Gen.sp. – – * Simuliidae Prosimulium sp. – * – Simulium sp. * * * Tipulidae Tipulidae Gen.sp. – – * *Present, - absent Blanckaert KJF, Han R, Pilotto F, Pusch MT (2014) Effects of References large wood on morphology, flow and turbulence in a low- land river. 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Age matters: substrate-specific colonization patterns of benthic invertebrates on installed large wood

Dossi, Florian; Leitner, Patrick; Graf, Wolfram
Aquatic Ecology , Volume 54 (3) – Sep 6, 2020
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Abstract

Aquat Ecol (2020) 54:741–760 https://doi.org/10.1007/s10452-020-09772-y(0123456789().,-volV)(0123456789().,-volV) Age matters: substrate-specific colonization patterns of benthic invertebrates on installed large wood . . Florian Dossi Patrick Leitner Wolfram Graf Received: 24 December 2019 / Accepted: 23 April 2020 / Published online: 6 May 2020 The Author(s) 2020 Abstract Large wood (LW) is an indispensable Our results show that (1) installed LW serves as an element in riverine ecosystems, especially in lower abundantly and heterogeneously colonized habitat, (2) river parts. The presence of LW significantly shapes the state of decay of LW pieces significantly affects local hydraulics, morphology, the nutrient budget; benthic invertebrate colonization in terms of density promotes overall river dynamics; and additionally and diversity and (3) even rare or threatened taxa presents a unique habitat for numerous benthic closely associated to LW were abundantly present on invertebrate species. Therefore, LW is recognized as the installed logs, emphasizing the suitability of the valuable asset for river restoration measures. Experi- chosen approach. ences from previous projects show that ecological responses on LW implementation measures vary Keywords Macroinvertebrates  Xylal  State of greatly. That complicates comparisons and estima- decay  River gradient  LW  Wood condition tions on the success of planned measures. Method- ological inconsistencies and thus reduced transferability of the results is one major issue. Additionally, wood quality aspects are suspected to Introduction be important factors affecting benthic invertebrate colonization patterns. The focus of this study is Large wood (LW) is a key component of natural river therefore to consistently assess the ecological signif- ecosystems. Previous studies have already stressed the icance of installed LW and concrete samples of similar beneficial effects of instream wood structures on local size and shape in terms of benthic invertebrate river hydraulics (e.g., Shields et al. 2001; Mutz 2003; colonization and to further test, if the condition of Manners et al. 2007), hydromorphology (e.g., Gurnell wood affects the benthic invertebrate colonization. et al. 1995, Kail et al. 2007, Blanckaert et al. 2014), nutrient balance (e.g., Bilby and Bisson 1998; Gurnell et al. 2005; Flores et al. 2011) and habitat diversity (e.g., Dudley and Anderson 1982; Hering and Reich Handling Editor: Telesphore Sime-Ngando. 1997). Submerged LW provides essential habitats F. Dossi (&)  P. Leitner  W. Graf which are existential for many xylobiont species IHG - Institute of Hydrobiology and Aquatic Ecosystem (Anderson et al. 1978; Hoffmann and Hering 2000) Management, BOKU - University of Natural Resources and are of increasing importance along the river course and Life Sciences, Gregor-Mendel-Strasse 33, (Dossi et al. 2018). LW further offers vital aquatic– 1180 Vienna, Austria e-mail: florian.dossi@boku.ac.at 123 742 Aquat Ecol (2020) 54:741–760 terrestrial interface areas and oviposition sites, signif- Smith and Smock 1992; Benke 1998). Besides natural icantly promoting the reproductive success of mer- fluctuations of densities due to riverine characteristics olimnic invertebrate species (Dudley and Anderson on local up to regional scales or methodological 1982; Sweeney 1993; Hoffmann and Hering 2000). inconsistencies, especially regarding the quantifica- Due to the wide variety of beneficial aspects, LW is tion of LW pieces and related individual densities, known to generally promote the density and diversity variations due to differing wood quality aspects, such of fish and aquatic invertebrate species (e.g., Dudley as hardness, species and condition, were discussed as and Anderson 1982; Copp 1992; Hering and Reich potentially important factors (Benke and Wallace 1997; Hoffmann and Hering 2000; Pilotto et al. 2003). That suggests a type specific colonization of 2014, 2016) and therefore presents a valuable and wood substrate, especially considering general sub- cost-effective asset for river restoration measures, strate selection processes of invertebrates substantially especially given the large amount of morphologically determining species richness, composition and density degraded river sections and comparably low costs to in freshwater ecosystems (Minshall 1984). Wood conventional measures (Kail and Hering 2005; Kail quality aspects were focused by only a limited number et al. 2007). Rough estimates assume that even in of studies, but most results indicate that the type and densely populated areas such as Central Europe, quality of instream LW affect invertebrate coloniza- approximately one-third of the degraded river sections tion patterns (e.g., Anderson et al. 1978; Kaufman and King 1987; Magoulick 1998; McKie and Cranston could be restored by reintroducing LW structures (Kail and Hering 2005). 1998; O’Connor 1991;Spa¨nhoff et al. 2000). A profound understanding of LW and ecosystem The aim of this study is therefore (1) to consistently interactions is a prerequisite for efficient implemen- assess the general ecological value of LW structures of tations in river management practices. Even though installed wood and concrete samples of comparable LW and its function in river ecosystems have been size and shape in terms of species richness and density, extensively investigated, knowledge gaps and there- (2) to investigate colonization patterns based on the fore implementation shortcomings still persist. Kail condition of the introduced LW pieces and (3) to test if et al. (2007) evaluated 50 restoration projects in the results are consistent within different river Germany and Austria involving LW placement and stretches along the longitudinal gradient of a med- found that only approximately 58% were successful. ium-sized lowland river in Austria. The authors concluded that the key to success lied in the consideration of site-specific characteristics. Hence, profound knowledge on river type specific Materials and methods wood characteristics is one important criterion to promote the success of measures. One challenge, hard Study sites to come by, is the lack of knowledge of the pristine state of LW and related ecological aspects in many Four sites along the Lafnitz River have been investi- European streams due to the long history of active gated. The Lafnitz River is one of the last medium- wood removal (Hering and Reich 1997; Hering et al. sized meandering rivers in the Central Europe with 2000). Additional studies in different areas with near-natural flow-regime and morphodynamics, ripar- remaining intact riparian vegetation and at least ian vegetation and LW accumulations along large near-natural LW dynamics are therefore of utter most parts of its course. It is therefore well suited to study importance to improve the understanding of LW and the importance of LW and its interactions with biota. biota interactions. The Lafnitz lies within the Danube catchment, located Benke and Wallace (2003) called attention to in the southeastern part of Austria (Fig. 1). The river additional difficulties regarding comparability and course has an approximate length of 112 km and transferability of results from different studies. Fun- drains into the Raab River, in Hungary. The Lafnitz damental information such as reported invertebrate River has a catchment size of approximately densities span wide apart from several hundred (e.g., 2000 km at the border of Austria, making it the O’Connor 1992; Rabeni and Hoel 2000) to many 13th largest river in Austria (BMLFUW 2002; Cejka (ten-) thousands of individuals per square meter (e.g., et al. 2005). The spring is located in the federal state of 123 Aquat Ecol (2020) 54:741–760 743 Fig. 1 Overview of the project area and location of the Lafnitz River in Austria (left) and location of the investigation sites along the river course (right); overlay: Ecoregions according to Illies (1978) Styria and originates at an altitude of 940 m above sea size at each site were assessed in March 2014 prior to level (m a.s.l.). Following Illies (1978), the first 36 the installation of the samples. Mean flow velocities river kilometers are situated in the ecoregion 4-Alps, and water depths were derived from averaged transect whereas the following section lies in ecoregion measurements. Grain size distribution were based on 11-Hungarian Plains (Fig. 1). The mean annual the choriotope assessment (Multi-Habitat-Sampling- discharge of the Lafnitz River spans from 2.6 m /s Method (MHS), AQEM Consortium 2002). in the upper section (near Site A) to approximately 6.3 m /s in the lower section near the town Dobersdorf Sample characteristics, sampling design (near Site E). and laboratory work The sampling sites were chosen to cover a variety of different river characteristics along the longitudinal For this colonization experiment three different types gradient to (1) allow general statements on the of substrate, comparable in size, were installed at each potential ecological benefits of installed wood struc- sampling site: (1) concrete bars, (2) fresh logs and (3) tures as well as to (2) assess possible variations in rotten logs (Figs. 2, 3). different river sections. At all four sites naturally The concrete bars were used to mimic lithal deposited large wood accumulations were present to substrates of comparable size and shape to the logs. ensure a comparable colonization potential. The dimensions of each piece were 50 cm 9 5cm 9 Samples have been taken at four sites along the 20 cm (length 9 width 9 height). In each concrete river course (see Table 1), whereas the most upstream bar, a whole was drilled and a rope was attached to be site is located at 648 m a.s.l. and the most downstream fixed at nearby trees (see Figs. 4, 5). site at 244 m a.s.l. The sites were chosen based on All xylal substrate samples (fresh and rotten) significant changes of river characteristics such as originate from the Lafnitz River and were sampled slope, discharge (stream order according to Strahler in February 2014 near the village of Neustift, in the (1957)), substrate composition and morphological middle section of the river. To ensure comparability, characteristics. Figure 1 gives an overview of the site only previously submerged logs with a diameter locations along the river course. Site-specific charac- between 200 mm and 300 mm were used in this teristics are shown in Table 1. study. Each log was cut to a length of 630 mm and Abiotic characteristics at each site including mean analogous to the concrete bars; a hole was drilled in flow velocity, mean water depth and dominant grain each sample log and a rope was attached to fix the log 123 744 Aquat Ecol (2020) 54:741–760 Table 1 Overview of site characteristics including altitude 1957), average water depth (m), average flow velocity (m/s) (meters above sea level), distance from spring (km), mean river and average river width (m) slope (%), dominant grain size (cm), Strahler number (Strahler Site Altitude Distance from Slope Dominant grain Strahler Average water Flow Average River (m asl) Spring (km) (%) size (cm) Number depth (m) Velocity (m/ width (m) s) A 648 8.7 1.7 6.3–20 4 0.2 0.3 8 B 438 26.1 0.9 6.3–40 5 0.3 0.6 15 C 324 52.1 0.3 2–6.3 5 0.5 0.3 10–20 D 244 100.4 \0.1 2–6.3 6 0.6 0.35 20 Fig. 2 Picture of the Lafnitz River at site A (left) and site B (right) Fig. 3 Picture of the Lafnitz River at site C (left) and site D (right) at the chosen sites (see Fig. 4). All sampled logs were based on the summarized decay categories of Robin- classified into the categories ‘‘rotten’’ and ‘‘fresh’’ on son and Beschta (1990) (class 1–3: fresh; class 4–5: site (examples see Fig. 6). Wood classification was rotten) and Grette (1985) (class 1–4 fresh; class 5–6: 123 Aquat Ecol (2020) 54:741–760 745 Fig. 4 Sample schematics and dimensions for fresh and rotten logs (left) and concrete samples (right) Fig. 5 Example of one installed concrete bar at site A rotten). Details on the classification characteristics of The surface area of the sampled LW pieces varied 2 2 wood pieces are shown in Annex see Table 5, 6. between 0.16 m and 0.32 m . The volume of each Length, width and volume of each LW piece was LW piece was measured in an overflow tank and 3 3 measured, and the surface area was calculated using varied between 2.8 dm and 10.2 dm . In total, there the formula of a simplified shape of a truncated cone: were 24 wood samples, corresponding to a total surface area of approximately 5.2 m (12 rotten logs: 2 2 S ¼ r  p þ p  R þ p  h ðÞ r þ R 2 2 2.5 m ; 12 fresh logs: 2.7 m ). All wood sample characteristics are summarized in Table 2. After where S = surface area, r = radius 1, R = radius 2, classification and preparation of the samples, all wood h = sample height. samples were rinsed, cleaned and carefully examined 123 746 Aquat Ecol (2020) 54:741–760 Fig. 6 Example of fresh (left) and rotten logs (right) installed at the Lafnitz River for remaining organisms to ensure a comparable pre- organisms were removed, and each log or concrete bar colonization state of all samples. The logs were then was subsequently reintroduced to the river. Benthic stored in black plastic bags until installation. samples were taken on three different dates in April 2014, June 2014 and March 2015. All samples were The logs and concrete bars were installed in a similar manner at each site. At each of the four sites, taken within 1 day. Due to the long exposition time three concrete bars and six logs (three fresh/three and the high morphodynamics at the Lafnitz River, not rotten) were installed in early March 2014. Three long all samples were consistently available for analysis. guiding ropes (approximately 15–20 m) were tied to Single logs or concrete bars which occasionally riparian trees, and three substrate samples (either logs stranded or were entirely covered with sediments or concrete bars) were subsequently bound to each were excluded from subsequent analysis (Details see guiding rope. The allocation of logs and concrete bars Table 2). on the guiding ropes was randomized, and all samples Sampling was performed with a standardized were installed in comparable orientations, parallel to 500-lm mesh-size kick-net with a frame size of the flow. However, a slight displacement or motion of 25 9 25 cm (surface area per single sample: the samples was possible. The location and orientation 0.0625 m ) and a net length of 1.2 m. Prior to of the guiding ropes was chosen to ensure comparable sampling, all LW pieces were carefully put into the habitat characteristics for each sample within each site kick-net to avoid organisms from drifting. Benthic (e.g., flow velocity, surrounding substrate invertebrates were then carefully brushed and washed composition). from the LW piece into the kick net. All organisms Details on the habitat parameters at each concrete were preserved with formaldehyde (4%) (Table 3). bar or log are given in Table 2. All single logs and The Screening-Taxa list (Ofenbo¨ck et al. 2010) was concrete bars were installed with a minimum distance used as a basis for identification. In many cases of 2 m to each other to allow undisturbed sampling Ephemeroptera, Plecoptera and Trichoptera taxa could conditions at each log or concrete bar. To ensure be identified to a lower taxonomic level (genus/ comparable colonization potentials, all samples had species), whereas Diptera taxa were mainly identified contact with the river bottom. to family level. Oligochaeta were not identified Sampling was performed in spring 2014 and 2015. further. All taxa were counted and weighed (wet To ensure comparable exposition times, all logs and weight). Specimens were then fixed in 70% ethanol concrete bars were cleaned and carefully examined and stored. 6 weeks prior to each sampling run. All attached 123 Aquat Ecol (2020) 54:741–760 747 Table 2 Overview of the sample characteristics including exact sampling point and the availability (only fully submerged 3 2 sample ID, the site, volume (dm ), surface (cm ), the state (f- samples from each sampling date were considered for analysis) fresh, r-rotten, c-concrete), the flow velocity (mean flow of samples at the each sampling date (1-23.04.14, 2-28.06.14, velocity at exact sampling point in m/s), dominant substrate at 3-18.03.15) 3 2 Sample ID Site Volume (dm ) Surface (cm ) State Flow velocity (m/s) Substrate 1 2 3 W01 A 4.8 2161 r 0.3 Psammal ??? W02 A 5.1 2131 r 0.3 Microlithal ??? W03 A 7.2 2416 r 0.1 Psammal ??? W04 A 3.9 1981 f 0.4 Psammal/Akal ??? W05 A 6.1 2462 f 0.2 Psammal ??? W06 A 10.2 3178 f 0.3 Psammal ??? C01 A 5 2700 c 0.7 Akal/Microlithal ??? C02 A 5 2700 c 0.1 Psammal ??? C03 A 5 2700 c 0.7 Akal/Microlithal – ?? W07 B 4.5 2027 r 0.8 Microlithal ??? W08 B 4 1911 r 0.8 Microlithal ??? W09 B 3.9 1880 r 0.8 Microlithal ??? W10 B 5.2 2314 f 0.8 Akal/microlithal ??? W11 B 7 2594 f 0.4 Microlithal ??? W12 B 4.9 2123 f 0.7 Akal/microlithal ??? C04 B 5 2700 c 0.8 Akal/microlithal ??? C05 B 5 2700 c 0.7 Akal/microlithal – ?? C06 B 5 2700 c 0.7 Akal/microlithal ??? W13 C 6 2350 r 0.2 Psammal – ?? W14 C 5.9 2332 r 0.5 Akal/psammal ? – ? W15 C 4 1927 r 0.3 Psammal ??? W16 C 7.1 2600 f 0.2 Psammal – ?? W17 C 2.8 1678 f 0.3 Psammal ??? W18 C 4.2 1952 f 0.2 Psammal ??? C07 C 5 2700 c 0.4 Akal – ?? C08 C 5 2700 c 0.5 Akal/microlithal ? – ? C09 C 5 2700 c 0.5 Akal ? – ? W19 D 3.6 1811 r 0.6 Psammal/akal ??? W20 D 4 1898 r 0.6 Psammal/akal ??? W21 D 3 1629 r 0.6 Psammal/akal ??? W22 D 4.8 2129 f 0.6 Psammal/akal ??? W23 D 6.8 2559 f 0.6 Psammal/akal ??? W24 D 4.1 1944 f 0.6 Psammal/akal ??? C10 D 5 2700 c 0.6 Psammal/akal ?? – C12 D 5 2700 c 0.6 Psammal/akal ??? C13 D 5 2700 c 0.6 Psammal/akal ??? Data analysis test equality of variances between the substrate groups and the Shapiro–Wilk Test to test on normality of the Statistical analyses were performed with the Software data. All abundance and biomass data (from all R-Studio 1.1.456. The Levene’s test was applied to substrates) were converted to densities (Ind/m ). 123 748 Aquat Ecol (2020) 54:741–760 Results Table 3 Overview of the total number of specimen, taxa and families observed at each site and substrate type In total * 43,000 benthic invertebrate specimen and A BCD 108 taxa from 52 families and 12 orders were collected No of specimen (see Annex in Table 7). The most abundant orders were Concrete 3338 2578 796 1918 Diptera (* 32%; mainly Chironomidae), Crustacea Fresh 2221 3270 3887 3562 (* 27%; only Gammarus fossarum) and Trichoptera Rotten 3154 5819 6703 4746 (22%; mainly Limnephilidae). Ephemeroptera com- No of taxa prised a share of * 13% and Plecoptera of * 3%. Concrete 33 29 32 42 Benthic invertebrate species richness gradually Fresh 36 36 34 48 increased along the river course from 49 taxa at site A Rotten 42 49 43 47 to 71 taxa at site D. Taxa richness per sample ranged No of families from 4 to 27 taxa. Details on the number of specimens, Concrete 22 19 22 27 taxa and families at each site and substrate type are Fresh 27 24 23 32 summarized in Table 3. Rotten 30 30 23 31 Density and diversity differences between substrate types Abundances for cluster and NMDS analyses were log Benthic invertebrate colonization showed significant (n ? 1) or presence/absence transformed. For cluster differences between the substrate types (see Fig. 7). analysis ‘‘Bray–Curtis’’ distance measure and Abundance, biomass and species richness were signifi- ‘‘Ward.D2’’ linkage method was applied. NMDS cantly higher (Wilcoxon test:a = 0.01) on wood samples analysis (Kruskal, 1964) was performed with ‘‘Sør- 2 2 (Ind/m : l = 2436.9 ± 241.3; g/m : l = 31.7 ± ensen (Bray–Curtis)’’ distance measure. 5.5; No. of taxa: l =15.5 ± 0.6) compared to concrete 2 2 The affinity of species or taxa to a particular samples (Ind/m : l = 1102.2 ± 139.0; g/m : l = substrate type was performed combining two different 8.1 ± 1.4; No. of taxa: l = 11.8 ± 0.6). Wood sample methods: showed a generally wider variation, as shown by the higher standard errors. Further, significant differences 1. Identification of taxa exclusively present on one 2 2 between fresh (Ind/m : l = 1723.9 ± 219.8; g/m : substrate type and l = 23.7 ± 4.8; No. of taxa: l = 13.9 ± 0.8) and rotten 2. identification of taxa which were significantly 2 2 wood samples (Ind/m : l = 3194.5 ± 401.6; g/m : more abundant on one substrate type based on the l = 40.2 ± 10; No. of taxa: l = 17.3 ± 0.9) were results of an ‘‘Indicator species Analysis (ISA). evident. Only biomass differences between fresh and All taxa with an Indicator value (IV) [ 25 rotten wood did not show significant results. Still, a (Dufrene and Legendre 1997) and a p value consistent pattern with lowest benthic invertebrate of B 0.05 were considered as significantly over- density and diversity on concrete, followed by fresh represented on the corresponding substrate type. wood and peak values on rotten wood, was clearly visible. This two-level approach was chosen in order to The most prominent density differences among the consider that even though some taxa significantly substrate types were found for Gammarus fossarum benefit from the presence of wood are not necessarily 2 2 (concrete: l = 102.2 Ind/m ; fresh: 648 Ind/m ; limited to wood as a habitat. The sole information on rotten: 1086 Ind/m ) (see Fig. 8: left). Diptera (mainly exclusive occurrences would therefore be insufficient. represented by Chironomidae taxa) showed compara- ble densities on concrete bars (487 Ind/m ) and fresh logs (418 Ind/m ) while being distinctly more abun- dant on rotten logs (1206 Ind/m ). Lowest Trichoptera densities were recorded on concrete (167 Ind/m ), followed by fresh (433 Ind/m ) and highest on rotten 123 Aquat Ecol (2020) 54:741–760 749 2 2 Fig. 7 Boxplot based on the number of individuals/m (left), biomass/m (middle) as well as the number of taxa (right) recorded at each single log or concrete bar; f—fresh wood, r—rotten wood; *p B 0.05, **p B 0.01, -p C 0.05 Fig. 8 Mean number of individuals of each order found on concrete bars (c), fresh (f) and rotten logs (r) with additional display of the standard error of the mean (left) and total number of taxa in each order (right) logs (673 Ind/m ). For Ephemeroptera and Plecoptera Trichoptera: Glossosoma sp.). In total, 39 taxa were taxa no particular trend was visible. The number of only found on wood samples (e.g., Plecoptera: taxa for all orders except for Crustacea (G. fossarum) Agnetina elegantula and Coleoptera: Macronychus showed a minor but consistent pattern as shown in quadrituberculatus). A further differentiation between Fig. 8: right. The number of taxa for Ephemeroptera, fresh and rotten wood samples showed nine taxa Plecoptera, Trichoptera and Diptera gradually exclusively present on fresh (e.g., Trichoptera: Rhy- increases from concrete bars to fresh and rotten logs acophila tristis and Silo pallipes) and 17 exclusively (see Fig. 8: right). on rotten wood samples (e.g., Trichoptera Hydropsy- Differences in species distribution between the che bulbifera and Lype phaeopa) (see Fig. 9: left). substrate are well reflected in the number of substrate- Results from each site and run separately reveal a specific taxa (see Table 4). Considering all samples, comparable pattern to the overall analysis (see Fig. 9: three taxa were exclusively present on concrete bars middle). Lowest number of exclusive taxa were found (e.g., Ephemeroptera: Epeorus alpicola and on concrete (l = 2.7 ± 0.6) followed by fresh- 123 750 Aquat Ecol (2020) 54:741–760 Fig. 9 Bar chart showing the number of exclusive taxa on each run, c—concrete, w—wood, f—fresh wood, r—rotten wood substrate type from all sites (Site A to Site C) (left); number of (middle); number of exclusive taxa on each substrate type at exclusive taxa on each substrate type for each single sampling each site (Site A to Site C) (right) (l = 3.8 ± 0.9) and rotten wood samples allocations patterns based on the substrate type are (l =6±0.6). visible. Wood samples (fresh and rotten) are mainly The results of the indicator species analysis are distributed in the middle of the NMDS plot, with shown in Table 4. The analysis was performed based rotten wood samples being generally allocated within on different taxonomic resolutions (family, genus and closer proximity to each other compared to the other species level), and results of all three taxonomic levels substrate types. Concrete samples are generally showed consistent results. Seven families were iden- oriented to the left side of the plot within each site tified, mainly represented by one dominant taxon group and rotten wood samples to the right. Fresh which was significantly more abundant on fresh and wood samples are generally oriented in between rotten wood, respectively (e.g., Trichoptera: Lepidos- concrete and rotten wood samples. tomatidae: Lepidostoma basale; Limnephilidae: Hale- Comparable results are obtained by the cluster sus sp., Rhyacophilidae: Rhyacophila s.str.sp.) and analysis (see Fig. 12) identifying four distinct groups. two families/genera significantly more abundant One group comprises all substrate types from the two exclusively on rotten wood samples (Coleoptera: upstream investigation sites (A and B), one the Hydraenidae: Hydraena sp.; Crustacea: Gammaridae: concrete samples from both lower investigation sites Gammarus fossarum). On genus/species level one (C and D), one the fresh and rotten wood samples from additional taxon was found to be significantly more site C and the last one both wood samples from site D. abundant on concrete samples (Plecoptera: Perlidae: The slight separation of the rotten wood samples at site B in the second cluster are mainly caused by distinctly Perla sp.) and one on fresh and rotten wood samples (Ephemeroptera: Heptageniidae: Heptagenia higher invertebrate densities on the samples compared longicauda). to those at sites A and B. Following the allocation of samples within the groups from the NMDS analysis, Community composition the cluster analysis shows a closer allocation of the concrete samples from site C and D to the samples of Community analysis shows a clear separation of the first two sites A and B. samples per site (see Fig. 11) indicating that site- specific river characteristics are the predominant factors determining the benthic community. In addi- tion, a distinct longitudinal pattern as well as 123 Aquat Ecol (2020) 54:741–760 751 Table 4 Overview of Order Family Genus Species Preference Type exclusive taxa (Type 1) and taxa significantly more Coleoptera Corixidae Corixidae Gen. sp. C 1 abundant on concrete bars Dytiscidae Dytiscidae Gen. sp. R 1 and fresh/rotten logs (Type Elmidae Esolus sp. F ?R1 2) Elmidae Macronychus quadrituberculatus F ?R1 Gyrinidae Orectochilus villosus F ?R1 Hydraenidae Hydraena* sp.* R 2 Crustacea Gammaridae* Gammarus* fossarum* R2 Diptera Empididae Empididae Gen. sp. R 1 Psychodidae Psychodidae Gen. sp. R 1 Tipulidae Tipulidae Gen. sp. R 1 Ephemeroptera Ephemeridae Ephemera danica R1 Heptageniidae Epeorus alpicola C1 Heptageniidae Heptagenia coerulans R1 Heptageniidae Heptagenia* longicauda* F ?R2 Leptophlebiidae Habroleptoides sp. R 1 Leptophlebiidae Habrophlebia* confusa* R2 Mollusca Tateidae Potamopyrgus antipodarum F1 Odonata Calopterygidae Calopteryx sp. F 1 Gomphidae Ophiogomphus cecilia R1 Platycnemididae Platycnemis pennipes F1 Plecoptera Chloroperlidae Chloroperla sp. F 1 Perlidae Agnetina elegantula F ?R1 Perlidae Perla* sp.* C 2 Perlodidae* Isoperla* sp* F ?R2 Taeniopterygidae Rhabdiopteryx navicula R1 Trichoptera Glossosomatidae Glossosoma sp. C 1 Goeridae Goera pilosa R1 Goeridae Silo pallipes F1 Hydropsychidae Hydropsyche bulbifera F1 Hydropsychidae Hydropsyche dinarica F ?R1 Hydropsychidae Hydropsyche siltalai R1 Hydropsychidae* F ?R2 Hydroptilidae Hydroptila sp. F 1 Lepidostomatidae* Lepidostoma* basale* F ?R2 Leptoceridae Ceraclea dissimilis R1 Leptoceridae Ylodes simulans R1 Limnephilidae* Anabolia furcata F ?R1 Limnephilidae* Chaetopteryx sp. F ?R1 Limnephilidae* Halesus* sp.* F ?R2 Limnephilidae* Melampophylax melampus R1 Polycentropodidae Cyrnus trimaculatus F1 Polycentropodidae Polycentropus flavomaculatus R1 Psychomyiidae Lype phaeopa R1 Psychomyiidae Psychomyia pusilla F ?R1 *Marks the taxonomic level of a significant Rhyacophilidae Rhyacophila s.str.sp.* F ?R2 overrepresentation Rhyacophilidae Rhyacophila tristis F1 (p B 0.05) 123 752 Aquat Ecol (2020) 54:741–760 Fig. 10 Overview of the number of individuals and biomass between the substrate types are shown in the table below: c— per m as well as the number of taxa on each substrate type for concrete, w—wood, f—fresh wood, r—rotten wood; *p B 0.05, each investigation site separately (a–d); significant differences **p B 0.01, -p C 0.05 Fig. 11 Ordination graph of NMDS analysis based on the are shown by name; all other species are only indicated by solid species composition and density from all concrete bars as well as circles to ensure readability; final stress for 2-dimensional fresh and rotten logs (sites A to D) (left) and display of species solution = 0.1912 allocations in the ordination graph (right); species with best fit Site-specific analysis consistently follow the patterns found in the overall dataset (see Fig. 10). Significant differences (Wil- A site-specific approach gives comparable results to coxon test: * = 0.05; ** = 0.01) are shown in Fig. 10. the pooled data. Abundance, biomass and taxa rich- Lowest values were generally found on concrete ness at each site and on each substrate type samples, followed by fresh and rotten wood samples. 123 Aquat Ecol (2020) 54:741–760 753 Fig. 12 Cluster dendrogram (distance measure: Bray–Curtis; group-linkage-method: Ward.D2) based on the species composition at each site (A-D) One sole exception is evident at site A showing higher adjacent lithal habitats (e.g., Benke et al. 1984; Smock abundance on concrete than on wood samples. Highest et al. 1989, 1992; O’Connor 1991; Piegay and Gurnell abundance, biomass and taxa richness, regardless of 1997; Hoffmann and Hering 2000; Spa¨nhoff et al. the substrate types, were evident at the sites B and C. 2000; Grafahrend-Belau and Brunke 2005; Milner and The most considerable colonization differences Gloyne-Phillips 2005; Coe et al. 2009; Pilotto et al. between the substrate types are evident at site C. 2016; Dossi et al. 2018). The number of exclusive taxa found at each site is The high variability of invertebrate density on illustrated in Fig. 9 (right). Lowest values were wood is difficult to compare and interpret. Main consistently observed on concrete samples with only reasons for these differences comprise the lack of a one exclusive taxon at site A, C and D and none at site common experimental and quantification approach as B. The number of exclusive wood taxa varies from site well as general river types or regional differences A to C between seven to ten taxa. At site D, a (Benke and Wallace 2003). The aim of this study considerably higher amount of 24 exclusive wood taxa therefore was to assess the ecological significance of was evident. Fresh wood samples mark an increase installed large wood compared to uniform concrete from three taxa at site A to six at site D, while only one structures for benthic invertebrate communities, with exclusive taxon was found at site B and C. On rotten particular attention on the wood condition. To over- wood samples, an increase of exclusive taxa from four come difficulties in the quantification of heterogenous at site A to eight at site D was found. instream LW pieces, we conducted our study with concrete, fresh and rotten LW of comparable size and shape. Discussion Our results generally agree with the above-men- tioned findings. A discrimination between lithal and Previous studies already stressed the general ecolog- xylal substrates showed distinctly higher benthic ical importance of instream wood structures but most invertebrate density, biomass and diversity on xylal of them focused on naturally deposited logs or wood samples. accumulations of varying sizes. Further, wood char- Besides similar patterns, the majority of the above- acteristics such as the state of decay received little mentioned studies reported higher invertebrate density attention. Detailed results varied greatly but nonethe- on xylal substrates, especially considering those from less consistently showed higher benthic invertebrate North America (see Benke and Wallace 2003). A densities and diversities on wood compared to potential underestimation due to an insufficient 123 754 Aquat Ecol (2020) 54:741–760 exposition time can be ruled out in our study. Samples Information on determining factors of specific were exposed for 6 weeks and previous studies showed benthic invertebrate colonization patterns on logs of that colonization on installed substrates peaks at different decay stages is still sparse, but the results of approximately two to 6 weeks the latest (e.g., Nilsen previous studies suggest that wood surface character- and Larimore 1973; O’Connor 1992; Spanhoff et al. istics present a decisive aspect (e.g., Kaufman and 2000). Comparisons with a previously conducted King 1987; O’Connor 1991; Phillips 1993; Phillips study at the Lafnitz River, which focused on natural and Kilambi 1994; Magoulick 1998; Spanhoff et al. instream wood (see Dossi et al. 2018) further support 2000). LW further develops a productive epixylic our results. Recorded invertebrate density and biomass biofilm which is an essential food source for grazing were on a comparable level but on average higher in taxa (Golladay and Sinsabaugh 1991; Sinsabaugh the present study on the installed logs (1655 Ind/m vs. et al. 1991), which might depend on log surface 2 2 2 2437 Ind/m ; 19.2 g/m vs. 29.6 g/m ). Similar characteristics. Conditioned wood has a softer and differences between installed and natural substrates more diverse, 3-dimensional, structure compared to have already been observed and discussed. While most other habitats. It therefore provides a comparably Spa¨nhoff and Cleven (2010) generally referred to large, smooth and more importantly complex habitat rapid initial colonization processes of newly intro- on a small spatial scale compared to rocks and fresh duced substrates resulting in above-average inverte- LW which is potentially beneficial for benthic inver- brate densities in the first weeks, Spanhoff et al. (2000) tebrate colonization (Kaufman and King 1987; added another aspect that was also observed in the O’Connor 1991; Magoulick 1998). The consistently Lafnitz River. Due to the longer residence time of higher densities and diversities on rotten logs com- naturally deposited, compared to installed logs, larger pared to fresh logs and concrete bars support these parts of their surface tend to be covered by sediments assumptions. Besides overall quantities, consistent which are therefore not (or only partially) suitable for statements on taxa-specific colonization patterns are benthic invertebrate colonization. That affects surface only possible to a limited extend. Even though most density calculations and leads to a slight underesti- taxa were found in higher quantities on rotten wood, mation of benthic invertebrate abundance on naturally only a few taxa showed significant preferences. The deposited logs. most prominent differences were found for G. fos- Site characteristics and the natural longitudinal sarum being significantly more abundant on rotten zonation of aquatic communities along the river logs. G. fossarum being a shredder with preferences course were identified as the dominant factor govern- for low flow velocities might benefit most from the ing the overall composition of the benthic invertebrate more complex surface structure of rotten wood. It community. However, overall species richness and provides flow protection as well as retention areas for density of invertebrates further depended on the wood organic matter on a small spatial scale. The softer condition as already indicated by previous studies surface might further facilitate the fragmentation and (e.g., Magoulick 1998; Spa¨nhoff et al. 2000). An processing of the woody material itself as well as additional distinction of our samples between different epixylic biofilm growth. Most taxa labeled as closely substrates revealed discrete benthic invertebrate den- associated to wood such as L. basale, M. quadritu- sity and diversity differences, with lowest values on berculatus, O. villosus or A. elegantula (Anderson concrete, followed by fresh and peak values on rotten et al. 1978; Hoffmann and Hering 2000; Dossi et al. wood. Only at site A, no clear differences were evident 2018) were found to be significantly more abundant on between the substrate types, which emphasizes the wood but did not show any preference for either fresh results of Dossi et al. (2018). Their study showed that or rotten logs. Assumptions on possible differences the importance of LW as unique habitat structure due to varying biofilm developments on the tested significantly changes along the longitudinal gradient substrate types could not be verified. Grazer taxa were of a river. While invertebrate species showed no equally dominant on the tested substrate types. significant preference for a specific substrate type in Our results further show that artificial LW intro- upper river sections, an increasing diversification of duction, even of comparably small logs as used in our the benthic communities among xylal and lithal study, comprise a valuable element in river restoration substrates was evident along the river course. measures. All samples were consistently colonized by 123 Aquat Ecol (2020) 54:741–760 755 heterogenous invertebrate taxa throughout the sam- also the species of wood becomes of particular pling period and also threatened or rare species, interest. Wood species-specific properties (e.g., firm- closely associated to wood (e.g., M. quadritubercula- ness, structure, texture, stability, chemical composi- tus, H. longicauda; A. elegantula), were frequently tion) and subsequent varying degradation and habitat found on the installed logs (Graf 1997; Bauernfeind characteristics will be investigated in a follow-up and Humpesch 2001; Graf and Kovacs 2002;Ja¨ch research. et al. 2005; Buffagni et al. 2016). Acknowledgements Open access funding provided by Our results emphasize that not only the presence of University of Natural Resources and Life Sciences Vienna LW is of importance. The suitability as a habitat (BOKU). significantly depends on the state of decay. That relates to two important, interrelated aspects, specif- Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which ically the residence time of logs in stream channels permits use, sharing, adaptation, distribution and reproduction and the species of wood. While residence time and the in any medium or format, as long as you give appropriate credit state of decay are clearly connected, species of wood to the original author(s) and the source, provide a link to the may be decisive as well. Besides possible colonization Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are preferences for certain wood species, differing degra- included in the article’s Creative Commons licence, unless dation rates (Spa¨nhoff and Meyer 2004) and thus indicated otherwise in a credit line to the material. If material is variations of the required residence time in rivers add not included in the article’s Creative Commons licence and your another scale to the research topic. These findings are intended use is not permitted by statutory regulation or exceeds of particular importance considering management the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit actions like the large-scale removal of instream LW, http://creativecommons.org/licenses/by/4.0/. which deprived rivers from an essential structural element and habitat and significant changes of riparian vegetation communities (Hering et al. 2000; Annex Hohensinner et al. 2013). Given these initial insights on the effect of wood quality on benthic invertebrate See Tables 5, 6 and 7. colonization patterns further research incorporating Table 5 Overview of decay categories; decay class: decay category according to Robinson and Beschta (1990) Decay class Bark Twigs Surface texture Shape Wood color State 1 Intact Present Intact/firm Round Original n 2 Intact Absent Intact/firm Round Original n 3 Trace Absent Smooth to some surface abrasion Round Original to darkening n 4 Absent Absent Abrasion to some holes and openings Round to oval Dark d 5 Absent Absent Vesicular with many holes and openings Irregular Dark d n not decayed, d decayed Table 6 Overview of Decay class Bark Limbs Surface texture Center State decay categories; decay class: decay category 1 Intact Present Firm Solid n according to Grette (1985) 2 Intact Absent Firm Solid n 3 Loose or absent Absent Firm Solid n 4 Absent Absent Slightly rottend Solid n 5 Absent Absent Extensively rottend Solid d 6 Absent Absent Completely rottend Solid d 7 Absent Absent Completely rottend Rotted d N not decayed, d decayed 123 756 Aquat Ecol (2020) 54:741–760 Table 7 Taxalist of all taxa found at all four sites; ‘‘c’’—concrete bars, ‘‘f’’—fresh logs, ‘‘r’’—rotten logs Order Family Taxon c f r Turbellaria Turbellaria Turbellaria Gen.sp. * * * Gastropoda Hydrobiidae Potamopyrgus antipodarum (J.E. Gray, 1843) – * – Oligochaeta Oligochaeta Oligochaeta Gen.sp. * * * Amphipoda Corophiidae Corophium sp. – * – Gammaridae Gammarus fossarum Koch, 1835 * * * Hydrachnidia Hydrachnidia Hydrachnidia Gen.sp. * * * Ephemeroptera Baetidae Baetis sp. * * * Caenidae Caenis sp. – * – Ephemerellidae Ephemerella ignita (Poda, 1761) * * * Ephemerella mucronata (Bengtsson, 1909) * * * Ephemerella notata Eaton, 1887 * * * Ephemerella sp. * * * Ephemeridae Ephemera danica Muller, 1764 – – * Heptageniidae Ecdyonurus sp. * * * Electrogena sp. – * – Epeorus alpicola (Eaton, 1871) * – – Epeorus assimilis (Eaton, 1871) * * * Heptagenia coerulans Rostock, 1877 – – * Heptagenia flava Rostock, 1877 * * * Heptagenia longicauda (Stephens, 1836) * * * Heptagenia sp. * * * Heptagenia sulphurea (Mu¨ller, 1776) * * * Heptageniidae Gen.sp. * * * Rhithrogena sp. * * * Leptophlebiidae Habroleptoides sp. – – * Habroleptoides confusa Sartori & Jacob, 1986 – – * Habrophlebia sp. – * * Paraleptophlebia sp. – * * Oligoneuriidae Oligoneuriella rhenana (Imhoff, 1852) * * * Potamanthidae Pothamantus luteus (Linnaeus, 1767) * * – Odonata Calopterygidae Calopteryx sp. – * – Gomphidae Gomphus vulgatissimus (Linnaeus, 1758) * * – Onychogomphus sp. * * – Ophiogomphus cecilia (Geoffroy In Fourcroy, 1785) – – * Platycnemididae Platycnemis pennipes (Pallas, 1771) – * – Plecoptera Chloroperlidae Chloroperla sp. – * – Siphonoperla sp. * * * Leuctridae Leuctra sp. * * * Nemouridae Amphinemura sp. * * * Nemoura/Nemurella sp. * * * Protonemura sp. * * * Perlidae Agnetina elegantula (Klapalek, 1905) – * * Dinocras sp. * * * Perla sp. * * * Perlidae Gen.sp. – * * 123 Aquat Ecol (2020) 54:741–760 757 Table 7 continued Order Family Taxon c f r Perlodidae Isoperla sp. * * * Perlodes sp. * * * Taeniopterygidae Brachyptera risi (Morton, 1896) * * * Brachyptera seticornis (Klapalek, 1902) * * * Brachyptera sp. * * * Rhabdiopteryx navicula Theischinger, 1974 – – * Heteroptera Aphelocheiridae Aphelocheirus aestivalis (Fabricius, 1803) * – * Corixidae Corixidae Gen.sp. * – – Coleoptera Dryopidae Pomatinus sp. – – * Dytiscidae Dytiscidae Gen.sp. – – * Elmidae Elmidae Gen.sp. – – * Elmis sp. * * * Esolus sp. – * * Limnius sp. * * * Macronychus quadrituberculatus Mu¨ller, 1806 – * * Gyrinidae Orectochilus villosus (Mu¨ller, 1776) – * * Helophoridae Helophorus sp. – * * Hydraenidae Hydraena sp. * * * Trichoptera Brachycentridae Brachycentrus subnubilus Curtis, 1834 * * * Glossosomatidae Glossosoma conformis Neboiss, 1963 * – * Glossosoma sp. * – – Goeridae Goera pilosa (Fabricius, 1775) – – * Goeridae sp. * * – Silo pallipes (Fabricius, 1781) * * – Hydropsychidae Cheumatopsyche lepida (Pictet, 1834) * – * Hydropsyche bulbifera Mclachlan, 1878 – * – Hydropsyche dinarica Marinkovic, 1979 – * * Hydropsyche instabilis (Curtis, 1834) * * * Hydropsyche pellucidula (Curtis, 1834) * * * Hydropsyche siltalai Do¨hler, 1963 – * * Hydropsyche sp. * * * Hydroptilidae Hydroptila sp. – * – Ithytrichia lamellaris Eaton, 1873 * * * Lepidostomatidae Lepidostoma basale (Kolenati, 1848) * * * Leptoceridae Ceraclea dissimilis (Stephens, 1836) – – * Leptoceridae Gen.sp. – * * Ylodes simulans (Tjeder, 1929) – – * Limnephilidae Allogamus auricollis (Pictet, 1834) * * * Anabolia furcata Brauer, 1857 – * * Chaetopteryx fusca Brauer, 1857 – * * Chaetopteryx sp. – * * Halesus sp. * * * Limnephilidae Gen.sp. * * * Melampophylax melampus (McLachlan, 1867) – – * Potamophylax cingulatus (Stephens, 1837) * * * 123 758 Aquat Ecol (2020) 54:741–760 Table 7 continued Order Family Taxon c f r Potamophylax rotundipennis (Brauer, 1857) * * * Odontoceridae Odontocerum albicorne (Scopoli, 1763) * * * Polycentropodidae Cyrnus trimaculatus (Curtis, 1834) – * – Polycetropodidae Polycentropus flavomaculatus (Pictet, 1834) – – * Psychomyiidae Lype phaeopa (Stephens, 1936) – – * Psychomyia pusilla (Fabricius, 1781) – * * Rhyacophilidae Rhyacophila s.str.sp. * * * Rhyacophila tristis Pictet, 1834 – * – Sericostomatidae Sericostoma sp. * * * Diptera Athericidae Ibisia marginata (Fabricius, 1781) * * * Ceratopogonidae Bezzia-Gruppe sp. – * – Chironomidae Chironomidae Gen.sp. * * * Empididae Empididae Gen.sp. – * * Limoniidae Antocha sp. ** – Hexatoma sp. * * * Limoniidae/ Limoniidae Gen.sp. – * * Pediciidae Pediciidae Dicranota sp. * * * Psychodidae Psychodidae Gen.sp. – – * Simuliidae Prosimulium sp. – * – Simulium sp. * * * Tipulidae Tipulidae Gen.sp. – – * *Present, - absent Blanckaert KJF, Han R, Pilotto F, Pusch MT (2014) Effects of References large wood on morphology, flow and turbulence in a low- land river. 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