Collembola respond to aphid herbivory but not to honeydew addition

Abstract. 1. Euedaphic collembola alter their soil distribution in response to above‐ground aphid herbivory of Poa annua L. Graminae, a host grass. 2. Two mechanisms potentially underpin this effect. Carbon‐rich aphid honeydew falling onto the soil surface may affect mycophagous collembola; alternatively aphid‐induced changes in root biomass may be necessary to produce changes in collembola abundance. 3. When compared to a plant‐only control, aphid herbivory increased the number of collembola in the top 5 cm of soil, reduced both foliar and root biomass, and increased shoot/root ratio. Honeydew addition had no effect on collembola numbers or any recorded host‐plant parameter. 4. Honeydew deposition is not responsible for the increased numbers of collembola found in the upper soil after aphid herbivory; aphid‐induced reductions in root biomass may be the most important factor explaining knock‐on effects of aphid herbivory on soil fauna.     

Influence of fungi on growth and survival of Onychiurus armatus (Collembola) in a metal polluted soil

Summary The influence of food quantity and quality on growth and survival of Onychiurus armatus (Tullb.) in metal polluted environments has been investigated in laboratory experiments. The Collembola was reared on five species of fungi isolated from a metal polluted soil close to a brass mill in SE Sweden.Survival of O. armatus was improved when fungal biomass was continuously added in a polluted mor (1,300 ppm Zn and 200 ppm Cu), and when specimens were fed metal polluted fungi for 1, 3 and 7 days a week, only those that were starved had increased mortality. Allometric growth, on the other hand, was significantly reduced when Collembola was given surplus of metal polluted fungi, whereas growth losses caused by metals were offset by protein rich food. Hence, sufficient food quantities alone could overcome mortality losses but not growth retardation in a metal polluted environment.Feeding preference of O. armatus was not determined by the protein content of the fungi although this was beneficial for growth. Metals changed the relative palatability of fungal species, but one of the metal tolerant species, Paecilomyces farinosus, which was also protein rich, remained reasonably attractive for O. armatus also when it was metal polluted. The mechanisms by which growth and survival of O. armatus were promoted by a combination of protein and Zn/Cu rich fungi seemed to be crucial in understanding the fate of a population of this species in a metal polluted soil.     

黄淮海平原保护性耕作下玉米季土壤动物多样性

调查了黄淮海平原农田保护性耕作下玉米拔节期(7月)和成熟期(9月)土壤动物的丰富度和多样性.结果表明：玉米成熟期土壤动物丰富度和多样性高于拔节期;拔节期土壤动物类群和数量免耕处理高于翻耕处理.土壤动物主要分布在表层(0～10 cm)土壤中,而翻耕处理下层(10～20 cm)土壤中仍有较大分布,导致翻耕与免耕处理下层土壤动物差异显著.免耕处理下,蜱螨目、小蚓类、双翅目和双尾目（特别是蜱螨目）数量高于翻耕处理.在玉米成熟期,秸秆还田效应显著,秸秆量越大土壤动物丰富度越高,其中弹尾目、蜱螨目、鞘翅目和啮虫目的个体数量均显著增加.弹尾目数量在玉米两个生长期均与秸秆还田量高度正相关,表明弹尾目在秸秆分解及其养分循环中起着重要作用.     

Resistance and resilience of subalpine wetlands with respect to prolonged drought

Vegetation dynamics of subalpine wetlands in the Sierra Nevada, California, were studied from 1988 through 1996. During this period, the region experienced a drought lasting from 1988 to 1994 and reaching its extreme in 1992. Our intention was to analyze the changes in plant species diversity, composition, and biomass, and interpret them in terms of drought resistance and resilience. Four plant communities, dominated respectively byCarex rostrata, Juncus balticus, Scirpus acutus orNuphar polysepalum were clearly discernible in the marsh along the water depth gradient. Species diversity ofCarex rostrata, Scirpus acutus andNuphar polysepalum communities was the highest during the driest year, 1992, while biomass was lowest for all vegetation types in that year. Dominance ofCarex andJuncus has not changed over the years, however,Scirpus almost totally disappeared from the marsh, and theNuphar zone become dominated by the rhizomatous perennial,Hippuris vulgaris, and terrestrial ruderals in dry years. In terms of changes in species composition,Carex andJuncus communities were both resistant and resilient. TheNuphar community seemed to be less resistant and more resilient, while theScirpus community was neither resistant nor resilient. If we consider biomass per plot as a variable of interest, regardless of species composition, thenCarex andJuncus were to some extent resistant and all plots were resilient because they were able to recover quickly to their pre-drought biomass. Life histories of dominant species were a more important determinant of community stability than species diversity.     

Effects of belowground grazing by collembola on growth,mycorrhizal infection,and P uptake of Geranium robertianum

We hypothesized that the grazing of vesicular-arbuscular mycorrhizal (VAM) hyphae by soil animals could be responsible for the lack of a direct relationship between mycorrhizal infection intensity and nutrient uptake under field conditions. To test this hypothesis, we determined the effect of a range of densities of the collembola, Folsomia candida, on growth, VAM infection, and P uptake in Geranium robertianum, a common forest herb, under greenhouse conditions. Total and aboveground growth were greater at low collembola density than either at higher collembola density or without collembola. These differences were greater when the plants were grown in a high organic content soil mix than when grown in sand. Root mass was not affected by collembola density. In the soil mix, root length decreased with increasing collembola density, but not in the sand. The percent of root length infected with VAM was lower at any collembola density than when collembola were absent. Total infected root length decreased linearly with increasing collembola density. Few significant differences in P uptake or tissue concentration were found. Thus, plant growth (but not P uptake) may be stimulated at low collembola density and inhibited at high. We discuss mechanisms which may be responsible for this non-linear response, and the implications of the pattern of response to studies of plant competition, nutrient turnover, and revegetation.     

Predation of Notiophilus (Coleoptera: Carabidae) on Collembola as a predator-prey teaching model

The carabid beetle Notiophilus preys readily on the collembolan Folsomia Candida Willem in very simple experimental conditions. Some features of this predator-prey system are outlined to emphasize its use as a teaching practical. Experiments with another potential collembolan prey, Onychiurus burmeisteri (Lubbock), are described in the context of developing the method for more advanced studies.     

The microbiological fate of polycyclic aromatic hydrocarbons: carbon and oxygen balances for bacterial degradation of model compounds

A series of pure bacterial strains belonging mainly to theRhodococcus andPseudomonas genera were grown on one of the following polycyclic aromatic hydrocarbons (PAH) supplied as sole carbon and energy source: naphthalene, fluorene, phenanthrene, anthracene, fluoranthene and pyrene. In each case, a quantitative evaluation of the carbon repartition of the PAH degraded into CO₂, biomass and water-soluble metabolites was carried out. In addition, the kinetics of oxygen consumption and of water-soluble metabolite accumulation during PAH biodegradation was followed with respirometric equipment. Satisfactory carbon balances were obtained and the data correlated well with oxygen consumption values. The results show that growth on PAH presents high mineralization yields (from 56% to 77% of carbon) and sizeable production of biomass (from 16% to 35% of carbon) and limited but significant accumulation of metabolites (from 5% to 23% of carbon). The mineralization yields were higher and biomass yields lower in the case of higher PAH. Some differences between strains were also observed.     

吉林中、西部平原区土壤弹尾虫群落结构的比较

2003年7月和9月对吉林省中、西部平原区天然次生林、农田、居民点园地、防护林和天然割草场等典型生境进行了土壤弹尾虫的调查,用Tullgren法提取弹尾虫,应用个体密度、类群数量、群落多样性、丰富度和均匀度等指数,研究土壤弹尾虫群落结构特征,了解土地利用差异对土壤弹尾虫群落结构的影响。共捕获土壤弹尾虫2亚目8科12属5 531头,优势类群为Onychiurus、Folsomia、Acanthocyrtus、Xenylla和Coloburella属, 占总捕获个体数的90.3%。弹尾虫类群空间分布的广狭与个体数量的多寡在吉林省中、西部平原区呈现出明显的一致性;中部平原区与西部平原区土壤弹尾虫群落组成差异主要体现在优势类群个体密度方面;生境类型明显影响弹尾虫个体密度分布,其中农田、居民点园地和防护林土壤弹尾虫个体密度较低,在不同土层间分布比较均匀,但生境差异对弹尾虫类群数的影响不明显;同时农业生产活动也降低了弹尾虫群落多样性。     

Aquatic macroinvertebrate community structure of aNymphoides peltata-dominated and macrophyte-free site in an oxbow lake

The aquatic macroinvertebrates in two freshwater biotopes,viz. aNymphoides peltata-dominated site and a macrophyte-free site, were studied quantitatively in a shallow alkaline oxbow lake of the river Waal, the main branch of the river Rhine in The Netherlands. The research comprised the analysis of water, sediment and macrophyte samples.In the macrophyte-free site Oligochaeta and Nematocera, particularly of the collector gatherer functional feeding group, dominated the prevailing benthic community. The total macroinvertebrate biomass ranged here from 0.3 to 0.9 g ash-free dry weight per m² of biotope.Species richness, densities, and biomass of macroinvertebrates were considerably higher in the biotope dominated byNymphoides peltata. Many taxa were found associated with the aboveground macrophyte. The sediment compartment, however, contributed most to the total density and biomass of macroinvertebrates. Nematocera and Oligochaeta were the most abundant fauna groups, whereas the largest share in total biomass was provided by clams (Mollusca). The biomass of the total macroinvertebrate community in theNymphoides-dominated site ranged from 6.2 to 7.5 g ash-free dry weight per m² of biotope. The biomass of the aboveground phytophilous fauna ranged from 0.1 to 0.6 g ash-free dry weight per m² of biotope. In September, when theNymphoides peltata vegetation was in its senescent phase, the largest numbers and the highest biomass of phytophilous macroinvertebrates were observed. The contribution of the shredder functional feeding group was high in this period. This, and the overall high abundance of fauna with a detritivorous mode of life, indicates the importance of macrophyte detritus as input to food chains.     

Patchiness and compensatory growth in a fungus-Collembola system

The compensatory growth potential of a grazed fungal biomass was mathematically expressed as a function of patchiness in its distribution and demonstrated in an experiment using the fungivorous collembolan Onychiurus armatus and the soil fungi Verticillium bulbillosum and Penicillium spinulosum. The model addresses the regrowth potential in relation to patch fragmentation, travelling time and consumption rate of the collembolan and the mean relative growth rate of the fungus. It suggests that the mean relative growth rate required for regrowth decreases with patch fragmentation and increases with the mean growth rate of the fungus. The experiments were performed with a system of soil-filled vials provided with fungi and collembolans. The size of the vials and the length of the tubes connecting them were varied to give different patch sizes and travelling times. The respiratory activity of fungi after grazing increased as a unit of mycelium was distributed into smaller connected vials. The slow growing species V. bulbillosum showed a greater but delayed response to grazing in comparison with the fast growing P. spinulosum. An increased travelling time delayed the growth response in both species.     

Barley straw decomposition with varied levels of microbial grazing by Folsomia fimetaria (L.) (Collembola,Isotomidae)

Summary Folsomia fimetaria (L.) were added (0, 5, 10, 20 animals) to 0.100 g barley straw which had been inoculated 10 days (244 h) earlier with a natural soil microflora. Respiration (CO₂ evolution) was monitored continuously. Mass loss, fungal standing crop (total and FDA-active), bacterial and protozoan biomass were estimated 42 days (1,000 h) after microbial inoculation. The degree of surface cover by hyphae was surveyed at regular intervals. No significant differences (P>0.05) were found in respiration, mass loss or microbial biomass, but the density of surface hyphae were reduced by addition of Collembola. Fungal production was low, less than 5% of the estimated microbial production, and could not account for all collembolan growth during incubation. F. fimetaria appeared to consume mainly bacteria and protozoa, and had little impact on carbon mineralization.     

Relationships in theAcanthaceae and related families as suggested by cladistic analysis ofrbcL nucleotide sequences

A parsimony analysis of DNA sequences of the chloroplast-encoded generbcL from twelve members of theAcanthaceae s.l., including members of the sometimes segregateThunbergioideae andNelsonioideae, and other families in theBignoniales sensuThorne (1992) is presented. The results largely agree with the classification of theAcanthaceae presented byBremekamp (1965) andThorne (1992) and supportNelsonioideae as a sister group to the rest of theAcanthaceae. Thunbergioideae are placed as a sister toAcanthaceae s.str.Acanthus andAphelandra, both representatives ofAcanthoideae, form a sister group toRuellioideae. An analysis of branch support found that many branches throughout theBignoniales are weakly upheld. This points to the need for further studies in the group using more sequences ofrbcL as well as other data. None of the families ofBignoniales as presently circumscribed (includingAcanthaceae s.l.) were strongly supported, although the larger clade containing the families of theBignoniales was robust.     

Phylogenetic relationships of theJuglandaceae

A cladistic analysis of molecular data from the chloroplast generbcL was used to examine the taxonomic relationships of the walnut family (Juglandaceae). In addition, chemical and morphological data from a previous study byHufford (1992) were incorporated, expanded, and analyzed independently and in combination with the molecular data. The results of these analyses suggest that theJuglandaceae are more closely related to theFagaceae, Betulaceae, Casuarinaceae, andUrticaceae and their relatives (sensuCronquist 1981) than they are to theAnacardiaceae (sensuThorne 1983). However, sequence data fromrbcL also suggest a relationship between the higherHamamelidae and certain families in theRosidae sensuCronquist 1981 (such asRosaceae andRhamnaceae), an outcome which would add credence to the widely accepted view of the polyphyletic nature of theHamamelidae.     

Biocontrol of the toxigenic plant pathogen <Emphasis Type="Italic">Fusarium culmorum</Emphasis> by soil fauna in an agroecosystem

In 2011 and 2013, a field experiment was conducted in a winter wheat field at Adenstedt (northern Germany) to investigate biocontrol and interaction effects of important members of the soil food web (Lumbricus terrestris, Annelida; Folsomia candida, Collembola and Aphelenchoides saprophilus, Nematoda) on the phytopathogenic fungus Fusarium culmorum in wheat straw. Therefore, soil fauna was introduced in mesocosms in defined numbers and combinations and exposed to either Fusarium-infected or non-infected wheat straw. L. terrestris was introduced in all faunal treatments and combined either with F. candida or A. saprophilus or both. Mesocosms filled with a Luvisol soil, a cover of different types of wheat straw and respective combinations of faunal species were established outdoors in the topsoil of a winter wheat field after harvest of the crop. After a time span of 4 and 8 weeks, the degree of wheat straw coverage of mesocosms was quantified to assess its attractiveness for the soil fauna. The content of Fusarium biomass in residual wheat straw and soil was determined using a double-antibody sandwich (DAS)-ELISA method. In both experimental years, the infected wheat straw was incorporated more efficiently into the soil than the non-infected control straw due to the presence of L. terrestris in all faunal treatments than the non-infected control straw. In addition, Fusarium biomass was reduced significantly in all treatments after 4 weeks (2011: 95–99%; 2013:15–54%), whereupon the decline of fungal biomass was higher in faunal treatments than in non-faunal treatments and differed significantly from them. In 2011, Fusarium biomass of the faunal treatments was below the quantification limit after 8 weeks. In 2013, a decline of Fusarium biomass was observed, but the highest content of Fusarium biomass was still found in the non-faunal treatments after 8 weeks. In the soil of all treatments, Fusarium biomass was below the quantification limit. The earthworm species L. terrestris revealed a considerable potential as an effective biocontrol agent contributing to a sustainable control of a Fusarium plant pathogen in wheat straw, thus reducing the infection risk for specific plant diseases in arable fields.     

Faunistic similarity and endemism of earthworms in east mediterranean region

The analysis presents earthworm distribution in eight designated tectonic units (C—Cyprus, EP—East Pondites, ET—East Taurus block, Sc—Sakarya continent, Kb—Kırşehir block, L—the Levant, TM—Taurus–Menderes block, WP—West Pondites) in the East Mediterranean region (EM). It represents a full list of earthworm species (N = 81) of the region and reveals significant faunal similarities between Sc + WP, and between Kb + TM. The new Sc + WP unit is characterized by the presence of archaic (Spermophorodrilus) as well as modern earthworm fauna. In contrast, the second newly established Kb + TM unit is characterized by poorly presented earthworm autochthonous fauna and by a lack of endemic species. The analysis done on the species list (N = 73) representing the autochthonous family Lumbricidae showed statistically supported separation of L and EP between themselves and from other designated units. The Levant, part of the Arabian tectonic plate, is characterized by a high level of endemism in ancient (Healyella) and modern (Dendrobaena, Perelia) genera, and by the lack of archaic genera. The EP harbors unique species representing archaic (Eophila), ancient (Healyella), and modern (Allolobophora, Aporrectodea, Dendrobaena and Eisenia) earthworm genera in the EM. The analysis also indicates origin of earthworm fauna in TM and faunistic relationships among the designated units. Clearly, the present-day division of the EM earthworm fauna shows recognizable influences of tectonic history.     

Effects of European Earthworm Invasion on Soil Characteristics in Northern Hardwood Forests of Minnesota, USA

European earthworms are colonizing worm-free hardwood forests across North America. Leading edges of earthworm invasion in forests of northern Minnesota provide a rare opportunity to document changes in soil characteristics as earthworm invasions are occurring. Across leading edges of earthworm invasion in four northern hardwood stands, increasing total earthworm biomass was associated with rapid disappearance of the O horizon. Concurrently, the thickness, bulk density and total soil organic matter content of the A horizon increased, and it’s percent organic matter and fine root density decreased. Different earthworm species assemblages influenced the magnitude and type of change in these soil parameters. Soil N and P availability were lower in plots with high earthworm biomass compared to plots with low worm biomass. Decreases in soil nitrogen availability associated with high earthworm biomass were reflected in decreased foliar nitrogen content for Carex pensylvanica, Acer saccharum and Asarum canadense but increased foliar N for Athyrium felix-femina. Overall, high earthworm biomass resulted in increased foliar carbon to nitrogen ratios. The effects of earthworm species assemblages on forest soil properties are related to their feeding and burrowing habits in addition to effects related to total biomass. The potential for large ecosystem consequences following exotic earthworm invasion has only recently been recognized by forest ecologists. In the face of rapid change and multiple pressures on native forest ecosystems, the impacts of earthworm invasion on forest soil structure and function must be considered.     

Recent increase of filamentous algae in shallow Swedish bays: Effects on the community structure of epibenthic fauna and fish

A summary is presented of estimates of distribution and growth of filamentous algae and its effect on the structure and functioning of epibenthic fauna and fish communities in shallow bays on the Swedish west coast. As a consequence of coastal eutrophication vegetation cover has gradually increased during the last decade, and during 1990's most bays in the Skagerrak-Kattegat area were variously covered with filamentous algae during spring and summer (May–July). In some areas filamentous algae (mainlyCladophora andEnteromorpha) completely covered the bottom. In field studies it was demonstrated that increased cover and dominance of filamentous algae result in structural changes of the epibenthic fauna community. Field studies showed that species richness and biomass of epibenthic fauna increased in a sandy bay with a moderate increase (30 to 50%) of filamentous algae cover. At higher cover (90%), biomass of epibenthic fauna was reduced, however, to the same level as for the sandy habitat, although the dominant epibenthic species were different. Heavy growth of epiphytic filamentous algae on eelgrass resulted in reduced biomass and a shift in the species composition of the epibenthic fauna community. Fish assemblage structure was also related to changes in vegetation. In eelgrass beds, fish species numbers were reduced with increasing cover of epiphytic filamentous algae, and at rocky bottoms with kelp algae (dominated byFucus), fish biomass decreased with increasing cover of attached filamentous algae. Further, foraging efficiency of juvenile cod and settling success of plaice were reduced as a response to increasing dominance of filamentous algae.     

Funsulinid assemblages and facies of the Bombaso Fm. and basal meledis Fm. (Moscovian-Kasimovian) in the central Carnic Alps (Austria/Italy)

Summary The Bombaso Formation and basal Meledis Formation in the central Carnic Alps near Straniger Alm and Zollnersee (Austria/Italy) unconformably overlie the folded Variscan basement and consist of shallow marine clastic and carbonate sediments which are arranged to form two fining and deepening upward sequences. Particularly limestones and even breccias of the Bombaso Formation yielded a rich fusulinid fauna composed of 34 species which are attributed to the following zones:Quasifusulinoides quasifusulinoides-Protriticites ovatus; Protriticites pseudomontiparus, andMontiparus montiparus. Breccias of the Bombaso Formation west of Straniger Alm contain the oldest fusulinid fauna of the Carnic Alps, belonging to theQuasifusulinoides quasifusulinoides —Protriticites ovatus zone. The fauna is composed ofQuasifusulinoides quasifusulinoides, Q. fallax, Q. intermedius, Protriticites cf.ovoides, andPr. ovatus. This assemblage is most similar to that of the Peskovskaya Formation of the Myachkovian Horizon in the Moscow Basin indicating uppermost Moscovian age. Limestones from depositional sequence 1 at Zollnersee also contain fusulinids of the uppermost Moscovian which are characterized by a more diverse assemblage:Schubertella donetzica, Fusiella lancetiformis, Beedeina ulitinensis, B. consobrina, B. nytvica, B. siviniensis, Quasifusulinoides pakhrensis, Q. fallax, Q. kljasmicus, Q. quasifusulinoides, Fusulinella rara, andProtriticites ovatus. Limestones and calcareous sandstones-siltstones of the basal Meledis Formation of depositional sequence 2 near Zollnersee and at Cima Val di Puartis are characterized by fusulinids of theProtriticites pseudomontiparus zone (Protriticites globulus, Pr. pseudomontiparus, Pr. sphaericus, Pr. rotundatus, Pr. ovoides, Pr. lamellosus, andPraeobsoletes burkemensis) and byMontiparus paramontiparus zone (Praeobsoletes pauper, P. burkemensis, Obsoletes timanicus, O. obsoletes, Montiparus paramontiparus, M. umbonoplicatus, M. montiparus, M. likharevi, M. rhombiformis andM. priscus) indicating lower to middle Kasimovian age (Krevyakinskian and Khamovnicheskian Horizons of the Russian Platform). In memoriam FranzKahler (1900–1995)     

Copper ions and the regulation ofSaccharomyces cerevisiae metallothionein genes under aerobic and anaerobic conditions

We have previously reported that theSaccharomyces cerevisiae CRS5 metallothionein gene is negatively regulated by oxygen. The mechanism of this repression was the focus of the current study. We observed that the aerobic repression ofCRS5 is rapid and occurs within minutes of exposing anaerobic cultures to air. Furthermore, theCUP1 metallothionein gene ofS. cerevisiae was found to be subject to a similar down-regulation of gene expression. We provide evidence that the aerobic repression of yeast metallothioneins involves copper ions and Ace1, the coppertrans-activator ofCUP1 andCRS5 gene expression. A functional Ace1 binding site was found to be necessary for the aerobic repression ofCRS5. Moreover, the aerobic down-regulation of the metallothioneins was abolished when cells were treated with elevated levels of copper. Our studies show that anaerobic cultures accumulate higher levels of copper than do aerobic cells and that this copper is rapidly lost when cells are exposed to air. In fact, the kinetics of this copper loss closely parallels the kinetics ofCUP1 andCRS5 gene repression. The yeast metallothionein genes, therefore, serve as excellent markers for variations in copper accumulation and homeostasis that occur in response to changes in the oxidative status of the cell.     

Earthworm community structure and diversity in experimental agricultural watersheds in Northeastern Ohio

Earthworms are known to have an important impact on soil fertility but much remains to be known about the factors that influence earthworm abundance and species diversity in agricultural soils and the impact of earthworm diversity on soil processes in those soils. We have studied factors that influence earthworm community structure and biodiversity in experimental agricultural watersheds at the North Appalachian Experimental Watershed near Coshocton, Ohio. We sampled earthworm communities in seven such watersheds from 1990 to 1992. Six earthworm species were present:Aporrectodea caliginosa, A. trapezoides, A. tuberculata, Lumbricus rubellus, Lumbricus terrestris and Octolasion tyrtaeum. The total earthworm biomass ranged from 2 to 32 g m^-2 and population levels ranged from 10 to 350 worms m^-2. Earthworm community structure and diversity differed among watersheds and was influenced by cropping patterns, geographic location and tillage. The greatest earthworm diversity and highest earthworm population levels occurred in a no-tillage watershed and a watershed that had previously been in ryegrass and long-term no-till. High earthworm populations were also observed in a chisel-plowed watershed. Watersheds that had been managed identically for 50 years had much different earthworm communities, indicating that factors other than management were important in determining species composition. The three watersheds with the lowest populations and diversity were adjacent to one another at one end of the station, indicating that at this site geographic location had the predominant influence on earthworm communities. A severe drought in 1991 greatly reduced earthworm populations and biomass. However, earthworm species differed in their response to drought withLumbricus rubellus appearing to be the most drought-sensitive andAporrectodea spp. the most drought-tolerant.