Ecological niche differentiation between native and non-native shrimps in the northern Baltic Sea

Invasions of non-native species are modifying global biodiversity but the ecological mechanisms underlying invasion processes are still not well understood. A degree of niche separation of non-native and sympatric native species can possibly explain the success of novel species in their new environment. In this study, we quantified experimentally and in situ the environmental niche space of caridean shrimps (native Crangon crangon and Palaemon adspersus, non-native Palaemon elegans) inhabiting the northern Baltic Sea. Field studies showed that the non-native P. elegans had wider geographical range compared to native species although the level of habitat specialization was similar in both Palaemon species. There were clear differences in shrimp habitat occupancy with P. elegans inhabiting lower salinity areas and more eutrophicated habitats compared to the native species. Consequently, the non-native shrimp has occupied large areas of the northern Baltic Sea that were previously devoid of the native shrimps. Experiments demonstrated that the non-native shrimp had higher affinity to vegetated substrates compared to native species. The study suggests that the abilities of the non-native shrimp to thrive in more stressful habitats (lower salinity, higher eutrophication), that are sub-optimal for native shrimps, plausibly explain the invasion success of P. elegans.     

Dietary habits of the polkadot skate <Emphasis Type="Italic">Dipturus chinensis</Emphasis> in the East China Sea

We investigated the dietary habits of the polkadot skate, Dipturus chinensis (Japanese name: Gangi-ei), in the East China Sea by analyzing the stomach contents of 529 individuals. Of these, 18 (3.40%) had empty stomachs. The mean and maximum fullness indices (FIs) were 0.67% and 6.12%, respectively. The mean FI of female skates > 500 mm in total length (TL) was higher than that of males and smaller females. The most dominant prey category was crustaceans, which included shrimps, crabs, and anomurans, followed by fishes. Cephalopods were additionally observed. Dipturus chinensis fed on a wide variety of crustacean taxa, and among them, Caridea and Galatheidae were particularly common. Fishes that could be identified were mostly demersal fishes, such as Callionymidae, Triglidae, and Rajidae. Some juvenile D. chinensis were also observed, indicating that cannibalism occurs in this species. Even though dental sexual dimorphism was observed in large size classes (mainly > 500 mm TL), the diet of males and females significantly overlapped. Size-related dietary shifts from small crustaceans, such as shrimps and anomurans, to relatively fast-swimming prey, such as fishes, crabs, and cephalopods, were observed in all seasons. The composition of these prey categories varied among seasons. Shannon–Wiener diversity indices calculated for each size class ranged from 2.01 to 2.51, indicating D. chinensis are generalist predators. As size increased, trophic level increased from 3.58 to 3.85, indicating that larger specimens are at a slightly higher trophic level than smaller specimens. These results suggest that D. chinensis are middle-trophic-level generalist predators that utilize diverse prey depending on their size and the season, and thus play an influential role in the demersal ecosystem of the East China Sea.     

Host-use pattern of the shrimp <Emphasis Type="Italic">Periclimenes paivai</Emphasis> on the scyphozoan jellyfish <Emphasis Type="Italic">Lychnorhiza lucerna</Emphasis>: probing for territoriality and inferring its mating system

In symbiotic crustaceans, host-use patterns vary broadly. Some species inhabit host individuals solitarily, other species live in heterosexual pairs, and even other species live in aggregations. This disparity in host-use patterns coupled with considerable differences in host ecology provide opportunities to explore how environmental conditions affect animal behavior. In this study, we explored whether or not symbiotic crustaceans inhabiting relatively large and structurally complex host species live in aggregations. We expected Periclimenes paivai, a small caridean shrimp that lives among the tentacles of the large and morphologically complex scyphozoan jellyfish Lychnorhiza lucerna, to live in groups given that the host traits above constraint host-monopolization behaviors by symbiotic crustaceans. We described the population distribution of P. paivai during a bloom of L. lucerna near the mouth of the Paraíba River estuary in Paraíba, Brazil. The population distribution of P. paivai did not differ statistically from a random Poisson distribution. Male shrimps were most often found dwelling on the surface of L. lucerna individuals as small groups (2–4 individuals), in agreement with expectations. Periclimenes paivai is a sexually dimorphic species with males attaining smaller average body sizes than females and exhibiting no elaborated weaponry (claws). Females, but not males, experience positive allometry in cheliped size and were found living solitarily in small but not large host individuals. The above suggest that females might be territorial or that they might be competing for resources (i.e., food) likely expected to impact their reproductive output. Our results agree, but only partially, with the idea that large and morphologically complex host species should harbor non-territorial gregarious symbiotic crustaceans. Symbiotic crustaceans represent excellent models to improve our understanding about the conditions driving the social behavior of marine organisms.     

Evaluation of genes involved in Norway lobster (<Emphasis Type="Italic">Nephrops norvegicus</Emphasis>) female sexual maturation using transcriptomic analysis

In marine ecosystems, the most significant migration observed in terms of biomass distribution is the one connected with the vertical movements in the water column. In the present study, the vertical profiles of the mesopelagic shrimps Gennadas elegans, Eusergestes arcticus, Sergia robusta, and the epipelagic Parasergestes vigilax in the Balearic Sea (western Mediterranean), during the stratified (summer) and non-stratified (autumn) hydrographic conditions, were investigated through their ontogeny, from the larval to adult stages. The mesopelagic adults were observed to move down to the deeper layers during the night more than during the daylight hours. Most larvae aggregated within the limits of the upper water column. The P. vigilax larvae were collected only during the stratified period. The first two larval stages vertical distribution indicates that the mesopelagic crustacean spawning could occur at greater depths. During the non-stratified period, the larvae of the mesopelagic species tended to remain at about 500 m depth at night, rising towards the upper layers at sunrise. Vertical patterns are discussed, as strategies associated with predator–prey trade-offs. To our knowledge, the present study is the first such attempt to jointly analyze the vertical migrations of the developmental stages of the pelagic shrimps in the Mediterranean Sea.     

The Structure and Density of Aggregations of the Hermit Crab <Emphasis Type="Italic">Pagurus proximus</Emphasis> Komai, 2000 in Epibiosis of the Brown Alga <Emphasis Type="Italic">Costaria costata</Emphasis> (C. Agardh) De A. Saunders, 1895 in the Sublittoral Zone of Vostok Bay,Sea of Japan

A study of epibiosis of the brown alga Costaria costata in the sublittoral zone of Vostok Bay, Sea of Japan during the spring and summer of 2016 showed that hermit crabs of the following four species accounted for approximately 85% of the total biomass: Pagurus proximus, P. minutus, P. brachiomastus, and P. middendorffii. Of these, the most abundant one (96.5% of the total number of hermit crabs) was P. proximus, the mean aggregation density of which on C. costata (429 ± 221 ind./m²) was higher by an order of magnitude than that in the adjacent area of the bouldery bottom. The size ranges of males and females of this species in aggregations on boulders and those on C. costata were similar, with a predominance of larger individuals on algae. In the size–frequency distribution, three groups of individuals corresponding to three size–age cohorts were differentiated. The male-to-female ratio was 1 : 1.7. Hermit crab aggregations on C. costata are ephemeral; their existence is determined by the duration of the period of algae development, that is, from late spring to the middle of the summer.     

Life history traits determine the differential vulnerability of native and invasive apple snails (<Emphasis Type="Italic">Pomacea</Emphasis> spp.) to a shared juvenile-stage predator

The vulnerability of gastropods to their predators varies with life history traits such as morphology, body size, behavior, and growth rates as well as predator size. A recent study suggested that the invasive apple snail, Pomacea maculata, was considerably more vulnerable to crayfish predators than the native Florida apple snail, P. paludosa. The difference was hypothesized to be caused by the relatively small hatchling size of P. maculata. To test this hypothesis, we conducted a series of feeding assays designed to quantify maximum feeding rates and selective foraging of crayfish on apple snails. The rate at which crayfish killed individual P. maculata (i.e., kill rates) decreased with snail size, and kill rates on both species increased with crayfish size. Kill rates on juvenile P. maculata were higher than kill rates on size-matched hatchling P. paludosa, and crayfish fed selectively on P. maculata when offered mixed groups of size-matched snails. Further analyses revealed that hatchling P. paludosa possess shells 1.8× heavier than size-matched P. maculata suggesting differences in vulnerability to crayfish were consistent with interspecific differences in shell defenses. Differences in hatchling size and defensive traits in combination make crayfish kill rates on hatchling P. maculata approximately 15.4× faster than on hatchling P. paludosa, but the relative contribution of hatchling size to differences in apple snail vulnerability was >3× greater than the contribution of defensive traits.     

Photosynthesis,stomatal conductance and terpene emission response to water availability in dry and mesic Mediterranean forests

Key message

Warmer summer conditions result in increased terpene emissions except under severe drought, in which case they strongly decrease.

Abstract

Water stress results in a reduction of the metabolism of plants and in a reorganization of their use of resources geared to survival. In the Mediterranean region, periods of drought accompanied by high temperatures and high irradiance occur in summer. Plants have developed various mechanisms to survive in these conditions by resisting, tolerating or preventing stress. We used three typical Mediterranean tree species in Israel, Pinus halepensis L., Quercus calliprinos and Quercus ithaburensis Webb, as models for studying some of these adaptive mechanisms. We measured their photosynthetic rates (A), stomatal conductance (g _s), and terpene emission rates during spring and summer in a geophysical gradient from extremely dry to mesic from Yatir (south, arid) to Birya (north, moist) with intermediate conditions in Solelim. A and g _s of P. halepensis were threefold higher in Birya than in Yatir where they remained very low both seasons. Quercus species presented 2–3-fold higher A and g _s but with much more variability between seasons, especially for Q. ithaburensis with A and g _s that decreased 10–30-fold from spring to summer. Terpene emission rates for pine were not different regionally in spring but they were 5–8-fold higher in Birya than in Yatir in summer (P < 0.05). Higher emissions were also observed in Solelim for the drought resistant Q. ithaburensis (P < 0.001) but not for Q. calliprinos. α-Pinene followed by limonene and 3-carene were the dominant terpenes. Warmer summer conditions result in increased Terpene emission rates except under severe drought, in which case they strongly decrease.

     

The diet of feral raccoon dog (<Emphasis Type="Italic">Nyctereutes procyonoides</Emphasis>) and native badger (<Emphasis Type="Italic">Meles meles</Emphasis>) and red fox (<Emphasis Type="Italic">Vulpes vulpes</Emphasis>) in Denmark

The raccoon dog (Nyctereutes procyonoides) is an East Asian Canid that has been introduced in Europe. Introduction of alien species is an increasing conservation issue. We examined the diet of a recently established raccoon dog population in Denmark by analysing stomach content in 249 carcasses collected in 2008–2016. Raccoon dog diet was compared to the diet of native badger (Meles meles) and red fox (Vulpes vulpes) in Denmark. The most common food for raccoon dogs were invertebrates (frequency of occurrence, FO 69%), small mammals (FO 68%), birds (FO 41%), fruits (FO 38%), amphibians (FO 36%) and carrions (FO 34%). The occurrence of invertebrates was highest during spring and summer, while fruits, cereals and carrions were eaten most often during autumn and winter. As expected, raccoon dog shared the major food categories with badger and red fox, but generally, it had a wider dietary niche. Overall, dietary overlap between raccoon dog and badger was 0.74 (Pianka index, O_jk). The dietary overlap with red fox was relatively high in all seasons, peaking in summer (O_jk 0.87) and dropping in winter (O_jk 0.79). Despite the dietary overlap between the alien racoon dog and native red fox and badger, the species may coexist due to partitioning of feeding habitats and/or because the red fox is limited by other factors, e.g. diseases and anthropogenic activities. The introduced raccoon dog seems to fit a dietary niche between badger and red foxes in human-dominated landscapes in north-western Europe.     

Differentiation of craniomandibular morphology in two sympatric <Emphasis Type="Italic">Peromyscus</Emphasis> mice (Cricetidae: Rodentia)

In the Santa Cruz Mountains of California, dietary partitioning is believed to allow Peromyscus californicus (California mouse) and Peromyscus truei (pinyon mouse) to occur sympatrically; P. californicus feeds primarily on arthropods, whereas P. truei feeds primarily on acorns. To better understand how these species partition resources, we examine if these dietary differences extend to differences in craniomandibular morphology. We use a geometric morphometric approach to test the hypothesis that P. californicus and P. truei exhibited size and shape differences in craniomandibular morphology, in particular, regions of the skulls that pertain to biting ability and mechanical advantage of the jaw adductor muscles. We found that P. truei exhibited relatively wider zygomatic arches, relatively broader, more robust masseteric fossa and coronoid process, and a higher mechanical advantage of the masseter jaw muscle. These craniomandibular traits suggested that P. truei exhibits a relatively stronger bite force that is more suitable to access hard-shelled acorns despite its smaller body size.     

Genome size and ploidy level among wild and cultivated <Emphasis Type="Italic">Prunus</Emphasis> taxa in Slovakia

2C DNA content and ploidy level variation of Prunus spinosa and closely related taxa together with Prunus domestica L. and Prunus insititia L. was studied in Slovakia. The aim of the study was to define genome sizes and find differences between closely related taxa within Prunus spinosa sensu lato mentioned in previous works. According to our results, investigated taxa can be divided into three groups according to ploidy level: Prunus spinosa, Prunus dasyphylla, Prunus ×fruticans, Prunus ×dominii and Prunus ×schurii are tetraploids, Prunus ×fechtneri is pentaploid, and P. domestica and P. insititia are hexaploids. Genome size differences within tetraploid taxa were relatively small (Prunus spinosa: 1.40?±?0.02, P. ×domini: 1.44?±?0.01, P. ×fruticans: 1.48?±?0.02, P. ×schurii: 1.44?±?0.02), but statistically significant. Although further research is needed, it seems that the concept of several taxa as product of hybridization between P. spinosa and cultivated plum species has been supported by our study.     

Hybridization success is largely limited to homoploid <Emphasis Type="Italic">Prunus</Emphasis> hybrids: a multidisciplinary approach

Prunus fruticosa is a rare shrub occurring in Eurasian thermophilous forest-steppe alliances. The species frequently hybridizes with cultivated Prunus species in Europe (allochthonous tetraploid P. cerasus and partly indigenous diploid P. avium). Propidium iodide flow cytometry, distance-based morphometrics, elliptic Fourier analysis and embryology were employed to evaluate the extent of hybridization in six Slovak populations. Flow cytometric analyses revealed three ploidy levels: diploid (P. avium), triploid (P. × mohacsyana) and tetraploid (P. fruticosa, P. × eminens and P. cerasus). In addition, P. fruticosa and P. cerasus, at the tetraploid level, were found to differ in absolute genome size. An embryological evaluation suggested the existence of a triploid block in P. × mohacsyana and significant potential for hybridization among tetraploid taxa (indicated also by a continuous distribution of genome size data and further mirrored by morphometrics). Although hybrids significantly differ in ploidy level and embryological characteristics, they are almost indistinguishable using morphological characters. Hybridization with P. cerasus thus turns out to be a significant threat to wild populations of P. fruticosa compared to the relatively weak influence of P. avium.     

The effects of recent changes in air temperature on trends in airborne <Emphasis Type="Italic">Alternaria</Emphasis>, <Emphasis Type="Italic">Epicoccum</Emphasis> and <Emphasis Type="Italic">Stemphylium</Emphasis> spore seasons in Bratislava (Slovakia)

Over the period 2002–2014, air temperature significantly increased regionally for Bratislava. However, no significant shifts have been observed in other meteorological parameters examined. The aim of this study was to analyse the effect of significant temperature trends on timing, duration and intensity of Alternaria, Epicoccum and Stemphylium spore seasons. Aerobiological monitoring was conducted using a Burkard 7-day volumetric spore trap. Mann–Kendall tau test was used to determine trends in spore seasons characteristics, whereas Spearman’s correlation coefficients were calculated to establish the relationships between temperature and spore season time series. Spore seasons of analysed taxa changed throughout the years of study. Alternaria spore season now starts earlier, ends later and lasts longer. Start, end and peak dates as well as duration of Alternaria spore seasons were significantly correlated with recorded increases in winter temperatures. Despite significant lengthening of Alternaria spore seasons, the lack of rising trend in its spore season index has been registered. This phenomenon could be partly explained by the reduction in the source vegetation due to drop of agricultural land use areas in Bratislava. In contrast, the intensity of Stemphylium spore seasons significantly increased during the study period and was correlated with recorded increases in summer–autumn temperatures. Based on the results of this study, it could be concluded that changes in selected fungal spore season patterns in Bratislava (earlier start date, later end date and longer duration of Alternaria spore seasons and higher Stemphylium spore season indexes) might be caused by the recorded local change in air temperature.     

Water availability drives stem growth and stem water deficit of <Emphasis Type="Italic">Pinus canariensis</Emphasis> in a drought-induced treeline in Tenerife

Tree growth of Pinus canariensis at treeline in Tenerife, Canary Islands, is thought to be primarily controlled by wet season precipitation (P) prior to the current year´s growth. Therefore, we investigated the inter-annual variations in stem water deficit (ΔW) and radial growth (RG) during two consecutive years differing in wet season P. ΔW was extracted from stem circumference variations, and the influence of environmental variables was evaluated by Pearson correlation statistics. Wet season P was considerably lower in 2008 than in 2009; despite this difference in P between both years, shallow soil water availability was almost exhausted during both summers. However, the effect of shallow soil drought showing a clear seasonality of ΔW and RG was only detected in 2008. In summer 2009, RG rates were highest during the summer indicating that P. canariensis was able to tap water from deep soil layers originating from P prior to the current year´s growth. The ability to use deep soil water during extended periods of shallow soil water deficit was also reflected in a close positive correlation between RG and whole-tree water use. In our study, the effect of only one hydrological dry year resulted in a severe reduction in annual RG. Thus, when wet season P is low for a number of years, chronic drought may have negative implications for tree growth at treeline in Tenerife.     

Taxonomic significance of pollen morphology for species delimitation in <Emphasis Type="Italic">Psidium</Emphasis> (Myrtaceae)

Previous studies reported Psidium as one of the most difficult genera to delimit within the American Myrtaceae. Even though palynology has improved the taxonomy of Angiosperms, information about the usefulness of pollen morphology for taxonomic purposes in Myrtaceae remains contradictory. Here, we investigate the significance of pollen morphology for Psidium taxonomy with specific focus on its usefulness for determining species groups of taxonomic significance. Pollen traits observed by light and scanning electron microscopy were quantified and examined using cluster and ordination analyses. Average size of pollen grains was visualized by boxplots. Pollen grains of Psidium are isopolar, oblate, peroblate or oblate-spheroidal, 3-syncolporate or 4-syncolporate. The sexine ornamentation is rugulate, granulate or spinulose-granulate and differs between the mesocolpium and apocolpium. Cluster analysis revealed four distinct groups: Psidium cauliflorum (G1) and Psidium oligospermum (G3) as single-species groups; Psidium brownianum, P. oblongatum, P. ovale, P. sartorianum, P. guajava, Psidium sp. 1, Psidium sp. 2 (G2), and Psidium cattleianum, P. longipetiolatum, P. guineense, P. myrtoides (G4). Supported by ordination analysis, three traits better explained these groups: type of exine ornamentation, size of P-EV and pollen shape. The used approach efficiently distinguished related species, as well as explained species groups of taxonomic significance suggesting pollen morphology to be a significant source of information for taxonomic studies in Psidium.     

Few But Relatively Large Prey: Trophic Ecology of <Emphasis Type="Italic">Chauliodus sloani</Emphasis> (Pisces: Stomiidae) in Deep Waters of the Central Mediterranean Sea

The trophic ecology of Chauliodus sloani (Stomiidae) was thoroughly investigated by stomach content analysis for the first time in the Mediterranean. Overall 206 individuals (64.0–260.0 mm SL) were collected from 2013 to 2015. C. sloani can be defined a specialist predator which feeds exclusively on mesopelagic fish belonging to Gonostomatidae, Myctophidae, Paralepididae, Phosichthyidae, Sternoptychidae and Stomiidae. Gonostomatidae (%IRI = 26.02), Myctophidae (%IRI = 24.77) and Sternoptychidae (%IRI = 24.35) were important food items for C. sloani. Maurolicus muelleri (%IRI = 36.43), Cyclothone braueri (%IRI = 28.26) and Vinciguerria attenuata (%IRI = 12.97) were the most important prey. Cases of cannibalism were also observed. The examination of food size spectrum, in relation to predator length, demonstrated that C. sloani developed a feeding strategy aimed to maximizing energy input and based on the capture of few and relatively large prey: more than 50% of prey items exceeded the value of 20% for ratio between prey and predator size (SL) and the 5.4% of prey measured more than 50% of predator size. Differences in food composition across seasons were found; C. braueri and M. muelleri were more abundant in autumn, V. attenuata during spring. Prey items mainly belongs to weakly vertical migrating fauna, usually concentrated at 400 m Deep Scattering Layer.     

Seasonal variations in moisture use in a piñon–juniper woodland

In water-limited environments of the intermountain region of North America, summer precipitation may play a role in the structure and function of aridland communities and ecosystems. This study examined the potential reliance on summer precipitation of two widespread, coexisting woody species in the southwestern United States, Pinus edulis Englmn. (Colorado piñon) and Juniperus osteosperma (Torr) Little (Utah juniper). The current distributions of P. edulis and J. osteosperma are highly suggestive of different dependencies on summer rainfall. We hypothesized that P. edulis was dependent on summer precipitation, utilizing summer precipitation even during extremely dry summers, whereas J. osteosperma was not dependent, using summer precipitation only when amounts were above some minimum threshold. Using sap flux and stable isotopic methods to assess seasonal water sources and water use efficiency, we examined the response of these two species to seasonal variations in moisture at a site located near the northern limits of the North American monsoon. Both sap flux and isotopic results indicated that P. edulis was responsive to summer rain, while J. osteosperma was not. Following summer rain events, sap flux density increased in P. edulis for several days, but not in J. osteosperma. Isotopic evidence indicated that P. edulis took up summer-derived moisture to a greater extent than J. osteosperma. Values of the natural abundance stable isotope ratio of carbon of leaf soluble carbohydrates increased over the summer for P. edulis, indicative of assimilation at higher water use efficiency, but were invariant for J. osteosperma. Our results supported the hypothesis that P. edulis and J. osteosperma are differentially sensitive to summer precipitation and are discussed in the light of potential changes in the seasonality of precipitation associated with climate change.     

Dietary Flexibility and Feeding Strategies of <Emphasis Type="Italic">Eulemur</Emphasis>: A Comparison with <Emphasis Type="Italic">Propithecus</Emphasis>

Despite the great variety of habitats in Madagascar, Eulemur has successfully populated most forested habitats on the island. Although the high dietary flexibility of Eulemur is often credited as one of the drivers of its evolutionary success, other behavioral evidence suggests a limited capacity for dietary switching. To shed light on the feeding strategies of Eulemur, we compared the dietary flexibility between populations of this genus with that of another widespread lemur taxon, Propithecus. We hypothesized that Eulemur would show greater dietary flexibility than Propithecus, which has a digestive system specialized for folivory, and that Eulemur living in dry forests would switch its diet from fruit to other food seasonally. To examine these hypotheses, we performed a phylogenetic least-squares analysis on 10 populations of Eulemur and 7 of Propithecus to assess the contribution of environmental variables and body mass on their dietary flexibility while controlling for phylogenetic relatedness. Eulemur relied heavily on fruit and did not show large variations in primary food over the year. Propithecus consumed leaves and fruits equally and exhibited considerable flexibility across seasons. Therefore, in contrast to our predictions, the anatomical specialization for fiber digestion heightens dietary flexibility in Propithecus. At the intrageneric level, we found similar ecogeographic variation; populations of both genera with heavier body mass consumed more fruit. As we predicted, Eulemur in drier habitats switched the diet from fruit to alternative food more frequently. To compensate for low dietary flexibility, Eulemur mostly adopts a power-feeding strategy by which it increases energy expenditure to acquire patchily distributed fruit resources.     

Size structures and comparative phenology of syntopic populations of <Emphasis Type="Italic">Hirudo verbana</Emphasis> and <Emphasis Type="Italic">Hirudo medicinalis</Emphasis> in eastern Ukraine

Ranges of different species of medicinal leeches (Hirudo spp.) are nearly mutually exclusive with very few areas of overlapping. This biogeographical pattern suggests different ecological requirements for the two species. However, two habitats in Hungary and Ukraine where populations of H. verbana and H. medicinalis coexist were found. The aim of this research was to reveal differences in size structures and phenologies between the two species in a steppe pond with two populations. The study was conducted from March to September in 2009 and 2010 in Horila Dolyna, Kharkiv Region. Leeches were collected several times during the season, weighed and released. The abundance of H. verbana was always higher. The Mann-Whitney U-test and the Wilcoxon test were applied. Significant differences between sequential samples within the species were found. The large weight classes of H. verbana emerged by the end of summer and caused a higher variance in contrast to H. medicinalis. This suggests the higher growth rate and the more r-strategic biology of H. verbana in comparison with the largely K-strategic H. medicinalis.