﻿Contributions to Ecuadorian butterworts (Lentibulariaceae, Pinguicula): two new species and a re-evaluation of Pinguiculacalyptrata

﻿Abstract Comparatively few species of the insectivorous genus Pinguicula L. have been recognized in South America so far. In recent years, a number of narrowly endemic taxa from the Andes have been described that simultaneously refined the broad taxonomic concepts of the “historical” species. Here, we describe two striking new species from Southern Ecuador that further condense the circumscription of Pinguiculacalyptrata Kunth. Pinguiculajimburensissp. nov. and P.ombrophilasp. nov. are clearly beyond the taxonomic scope of the known species and consequently described as new to science. The deviating morphological features of the two new taxa are described and illustrated and the remaining morphological spectrum of P.calyptrata in Ecuador is outlined. The two new species add to the exceptional biodiversity in the Amotape-Huancabamba Zone and underline its importance as a biodiversity hotspot in urgent need of protection.


Introduction
Pinguicula L., the second largest genus of the most specious family of carnivorous plants -Lentibulariaceae (Lamiales) -comprises approximately 115 species (Fleischmann 2021). The genus is distributed throughout Eurasia and the Americas, is largely absent from Africa (only Morocco) and not present in Oceania and Antarctica. Roughly half of all taxa occur in Latin America with a clear center of diversity in Mexico, with about 40 endemic species alone. Despite the high diversity in Central America, Mexico and the Caribbean, only a few species are known from South America and restricted to the Andean mountain chain. At the moment, most authors accept seven species here, namely: P. antarctica Vahl, P. calyptrata Kunth, P. elongata Benj., P. involuta Ruiz & Pav., P. jarmilae Halda & Malina, P. nahuelbutensis Gluch and P. rosmarieae Casper, Bussmann & T.Henning. All species belong to the monophyletic sect. Ampullipalatum Casper within the subgenus Isoloba, with the exception of P. elongata (sect. Heterophylliformis (Casper) Fleischmann & Roccia within subgen. Themnoceras).
Significantly, all recently described species are narrow endemics (P. jarmilae, P. nahuelbutensis & P. rosmarieae), whereas the earlier described taxa usually have larger ranges. This already indicates that the relatively low number of taxa known from (Andean) South America might be, at least partially, rather a result of under-collection and a lack of taxonomic studies, than true species poverty.
In Ecuador, the genus is to this date only represented by Pinguicula calyptrata Kunth (Taylor 1975;Jørgensen and León-Yánez 1999). The holotype for this species was collected in the páramo of Saraguro (Loja Province, Ecuador) by Alexander von Humboldt and Aimé Bonpland during their five years expedition to South America (1799)(1800)(1801)(1802)(1803)(1804). It was published in Karl Sigismund Kunth's "Nova Genera et Species Plantarum" (Humboldt et al. 1818) with a brief and superficial description and without an illustration as was customary at the time. The only taxonomic study that subsequently dealt with this group of plants in Ecuador dates back 48 years (Taylor 1975). His treatment of the Lentibulariaceae for the Flora of Ecuador was, however, mostly based on the scarce herbarium material available at the time and could not deliver a critical review of the genus for Ecuador.

Pinguicula in the Amotape-Huancabamba Zone and adjacent Andean regions
The genus Pinguicula is particularly species-rich in Latin America. However, the taxonomic richness south of the genus´ center of diversity: Mexico, Central America and the Caribbean, is limited, and most of the few taxa recognized are reported to have large latitudinal distributions. Lately, the continuous botanical exploration of the Andes and especially of the so-called Amotape-Huancabamba Zone (hereafter AHZ), however, shows a more complex picture for the genus in this area than hitherto believed. This phytogeographical zone has been revealed as an area characterized by an exceptional biodiversity and a noticeable accumulation of narrowly endemic taxa (Weigend 2002(Weigend , 2004. This has been demonstrated for many groups of organisms including numerous angiosperm taxa (e.g. Deanna et al. 2018;Escobar et al. 2018;Acuña-Castillo et al. 2021;Landis et al. 2021;Tejedor and Calatayud 2022;Vieu et al. 2022) and only very recently also for the Lentibulariaceae. With the description of P. rosmarieae, the first narrowly endemic Pinguicula has been documented for the AHZ (Casper et al. 2020). And the recent description of Utricularia amotape-huancabambensis ) revealed the same narrow endemism for the other large genus in the family.
However, despite constantly growing botanical research activities and new collections, taxon delimitation in Pinguicula remains problematic. This is particularly true for Pinguicula calyptrata, which is distributed from Central Colombia, throughout Ecuador and most of the AHZ until the North Peruvian Departamentos Cajamarca, Amazonas and San Martín (all specimens from Bolivia that we are aware of are misidentified P. involuta) and is unsatisfactorily defined in its entirety (Casper et al. 2020). Caspers concept of Pinguicula, in both systematics and alpha-taxonomy, was very flower focused (Fleischmann 2021). Species delimitation and the infrageneric classification of the genus in his works largely founded on morphological character complexes of the corolla, traditionally thought to be evolutionary stable. Instead, they were often subject to rapid diversifications and thereby prone to misinterpretations as recent molecular studies repeatedly revealed (Lustofin et al. 2020;Shimai et al. 2021, see Fleischmann 2021 for a detailed discussion). In the course of studying the Andean taxa, the late Siegfried Jost Casper (1929Casper ( -2021 also based his allocation of new collections/observations on the gross flower morphology and (for the time being) tended to classify any observation from the Northern Andes with more or less spreading rosettes and bluish flowers with a yellow palate as P. calyptrata (pers. correspondence). However, similar to the situation reported for a number of Cuban species (Domínguez et al. 2013), gradual morphological variation is found between most of the populations, varying with latitudinal and longitudinal distribution and elevation. Furthermore, on site-conditions such as e.g. moisture, soil type, associated vegetation and presumably the local pollinator fauna -a commonly ignored aspect in the context of speciation in the mosaic-like microhabitats of the Andes -affect genetic exchange between populations and facilitate diversification (Kay and Sargent 2009;Van der Niet et al. 2014). These dynamic and diverse networks of relationships and interactions can have a catalyzing effect on such character complexes that can be subject to rapid alteration and adaptation. In particular the plasticity of e.g. flower size, shape and color is greater than traditionally assumed and divergent versus convergent developments can be difficult to determine, potentially complicating an unambiguous assignment of taxa (Fleischmann 2021). With the ongoing botanical exploration of the (remaining) remote paramo habitats of Ecuador and Peru (and other Andean countries), the real extent of the morphological spectrum becomes increasingly recognizable and, in the case of P. calyptrata, quickly goes beyond the narrow scope of the protologue.
Besides biogeographical data and habitat preferences, the overall bauplan of Pinguicula provides only a limited number of characters to take into consideration for taxon delimitation. Basically, the plant body in this genus is reduced to leaves and flowers and morphological characterization boils down to a handful of characters delivered by these two organs. In the light of the aforementioned gradual variability between populations and the known uncertainties in species recognition in Andean South America (e.g. P. huilensis -Cuatrecasas, 1945; P. chilensis and P. antarctica -Gluch 2017; Roccia 2018) it appears appropriate that species delimitation must be subject to significant, quantifiable and consistent morphological differences (see stalk length debate in Gluch 2017 andRoccia 2018) and, in the case of the AHZ, must be considered in relation to omnipresent and often sympatric "typical" P. calyptrata. Nevertheless, lumping all taxa of Pinguicula from the Northern Andes together under P. calyptrata due to a superficial similarity in their flower and rosette morphology would not do the actual diversity any justice and could leave a false impression regarding the actual threat status of potential narrow endemic taxa.
Taking these considerations into account, most Ecuadorian populations of Pinguicula fall into the range of P. calyptrata and this species in its (original) unsharp definition is present in many suitable Andean habitats sampled, although a more finegrade analysis of P. calyptrata s.l., applying population genetics, is urgently needed and strongly encouraged. However, in two localities, two distinct taxa could be found growing sympatrically. The collections from the Cerro Plateado Biological Reserve show two taxa, that clearly differ in leaf morphology and rosette shape, albeit having similar flowers. Conversely, collections from Yacuri National Park show two taxa, differing considerably in both character complexes: leaves/rosettes and detailed flower morphology.
Despite the disparate grade of morphological dissimilarity observed for the two taxon pairs, the new taxa are both separated at species level to meet the needs of a conservation assessment under the IUCN criteria (IUCN Standards and Petitions Committee 2022) and facilitate potential protective measures. Whether a different taxonomic concept might ultimately be more appropriate to reflect the close relationship among (some) Andean Pinguicula species remains open to future studies, ideally within the framework of a more comprehensive approach using both phylogenetic and population genetic methods.
The new taxa are morphologically clearly distinguishable and a consistent spatial segregation is evident. Both new taxa share their habitat with rather typical P. calyptrata and are ecologically separated by contrasting habitat requirements, a pattern reported already for P. rosmarieae and P. calyptrata from North Peru (Casper et al. 2020). Similarly, the taxon pairs found in Ecuador also grow in close proximity to each other without any direct spatial overlap. No morphologically intermediate plants were found, indicating that the taxa here described are discrete reproductive units without naturally occurring hybrid individuals (unlike recently reported for Utricularia - Henning et al. 2021).
Consequently, P. jimburensis sp. nov. and P. ombrophila sp. nov. are described as species new to science. All aspects of their biology are presented and discussed. Furthermore, the morphological spectrum of P. calyptrata throughout the known Ecuadorian populations is outlined and illustrated. A key to the Ecuadorian species is provided and an up-to-date assessment of the taxonomic situation of Pinguicula in the Northern Andes is made.

Materials and methods
In the current study, we document two new narrow endemic Pinguicula species from southern Ecuador as a result of recent botanical explorations. We conduct detailed morphological analyses for these new species, provide a taxonomic description, present images for all floral and vegetative plant structures, and provide a distribution map. The conservation status of the newly described species and their relationships with other species are also discussed.
Furthermore, the morphological spectrum of P. calyptrata throughout the known Ecuadorian populations is outlined and illustrated. The populations of this species have been collected from north to south in all the Ecuadorian Andean provinces from 2100 to 4100 m.
We consulted pertinent literature, examined specimens at ECUAMZ, GUAY, LOJA, QCA and QCNE herbaria (Thiers 2023), and high-resolution images of type material for Neotropical taxa (Tropicos database, https://www.tropicos.org/ and the JSTOR Global Plants website http://plants.jstor.org). Diagnosis. Pinguicula jimburensis belongs to Pinguicula sect. Ampullipalatum and is closely allied to the other North Andean species of the section (P. calyptrata, P. ombrophila and P. rosmarieae). With the latter two it shares the lack of involute leaf margins, but clearly differs from them in its terrestrial habit (vs. litho-/epiphytic) and morphologically by the oblong leaves (vs. widely ovate to rounded). The flowers of P. jimburensis lack a distinct yellow palate, a character shared with P. involuta and the Peruvian endemic P. rosmarieae, but the evenly bent, tooth-like spur is unique among related species. First and foremost, P. jimburensis is characterized by erect leaves that are shallow and irregularly lobed. This character is, despite a terminological similarity with the leaves of only distantly related P. elongata, unique among all South American taxa.
Etymology. The specific epithet refers to the type locality, Lagunas Negras de Jimbura, which is part of the Yacuri National Park in the Ecuadorian provinces of Loja and Zamora-Chinchipe.
Distribution, habitat and associated vegetation. Specimens of P. jimburensis have so far only been collected around the Lagunas Negras de Jimbura in the Yacuri National Park in the province of Loja. As a result, P. jimburensis is endemic and thus far only known growing between the grass and shrubby paramo vegetation around the lagoon complex, especially in swampy areas (Figs 2, 3). According to the Ministerio del Ambiente del Ecuador (2013), this locality lies within a much larger zone dominated by the ecosystem named arbustal siempreverde montano alto del Páramo del sur. Inhabitants of this area call this type of vegetation "paramillo" and the environment is characterized by a constant cloudiness and drizzle with strong winds. The vegetation is dominated by the herbs Oritrophium sp. Paepalanthus lodiculoides Moldenke, Chusquea spp. and Phlegmariurus spp., between the shrubby vegetation dominated by Chuquiraga jussieui J.F. Gmel., Monticalia peruviana (Pers.) C. Jeffrey and Miconia spp. Additionally, Pinguicula calyptrata has been collected in this locality (Pérez et al. 8690,8755,QCA;Figs 2,3). The border between Ecuador and Peru is only about 3 km in a straight line from the type locality, hence P. jimburensis could also occur in Peru.
Conservation status. Only one population of approximately 50 mature individuals of this species was discovered at the type locality at the Yacuri National Park. The habitat is very close to the shore of the Laguna Negra and exposed to human activities related to spiritual rituals. Additionally, the trail that connects the lagoon complex closely passes by the population of this new species. According to the IUCN Red List criteria (IUCN Standards and Petitions Committee 2022) this species is assessed as Vulnerable (VU, Criterion D2).
Affinities. P. jimburensis apparently is a close ally of P. calyptrata as are all Ecuadorian and Andean taxa. A similar atypical leaf orientation, with erect, elongated leaves can only be found in P. elongata Benj. from Colombia and Venezuela. This strange species is, contrary to traditional placements (Ernst 1961, Casper 1966, not closely related to the other Southern American taxa, but represents an isolated lineage with affinities to Mexican and European clades, as a recent molecular phylogenetic analysis has shown (Shimai et al. 2021). Moreover, the leaves of P. jimburensis are only slightly elongated and otherwise flat with a distinct abaxial and adaxial leaf lamina and not involute as the almost filiform leaves of P. elongata, with the abaxial leaf surface sometimes completely hidden. Furthermore, the leaf margins of P. jimburensis are irregularly, shallowly lobed, a unique character among Pinguicula from South America so far. With the lack of a yellow palate and the evenly narrowed spur that resembles the ripper tooth of a predatory mammal in shape, the flowers appear more similar to those of P. involuta than to typical flowers of P. calyptrata. The latter is characterized by the presence of a yellow palate and the spur, although very variable, usually has a stubby apex preceded by a narrower or at least straight section that is rather abruptly angled downwards after the throat. Both P. jimburensis and P. involuta lack a yellow palate and the thorn-shaped, regular curved spurs in both taxa are evenly narrowed towards the apex. However, P. involuta has a very different leaf morphology and is overall a much smaller taxon distributed much further south. P. jimburensis cannot be confused with any other American Pinguicula species based on the exceptional leaf morphology and orientation.
Additional specimens examined (paratypes). Ecuador. Loja. Cantón Espíndola, Parroquia Jimbura, Parque Nacional Yacuri, Lagunas Negras de Jimbura, 04°42'S, Diagnosis. Pinguicula ombrophila belongs to Pinguicula sect. Ampullipalatum and is closely allied to the other North Andean species of the section (P. calyptrata, P. jimburensis and P. rosmarieae). It differs from P. calyptrata and P. jimburensis in its lithophytic (vs. terrestrial) habit and the combination of broad rosettes with flat leaves (unlike P. jimburensis) without curled margins (unlike P. calyptrata). The flowers of P. ombrophila are similar to those of P. calyptrata and a distinct yellow palate -absent in P. rosmarieae and P. jimburensis -is present. It differs from all three other taxa by having very short flower scapes that barely reach leaf-length.
Etymology. The specific epithet was chosen to point at the particular habitat requirements of the plants. They prefer very wet conditions where they not only receive constant moisture from the surrounding waterlogged paramo-soil, but are also fully exposed to the high amounts of precipitation and fog typical for this area. The name ombrophila signifies "rainloving" from the two latin words "ombros" (rain) and "philos" (that loves/is fond of ).
Distribution, habitat and associated vegetation. The Cerro Plateado Biological Reserve is one of the governmental protected areas along the Cordillera del Cóndor range in Ecuador, protecting around 26 000 ha of mature forest from 850 to 3100 m in the province of Zamora-Chinchipe. The Cordillera del Cóndor runs 150 km northsouth along the border of Ecuador and Peru. This mountain range is isolated from the main Andean chain and is geologically distinct, formed with an intermixture of limestone, quartzitic sandstone, and igneous rock of the Hollin Formation (Gregory-Wodzicki 2000; Neill 2005). The geology of these mountains is similar to the tepuis of the Guyana shield in northwest South America. In fact, a number of angiosperm genera once thought to be endemic to the tepuis of the Guyana shield have also been found along the Cordillera del Cóndor (Berry et al. 1995;Schulenberg and Awbrey 1997).
Pinguicula ombrophila, together with P. calyptrata (Neill et al. 17467, ECUAMZ;Pérez et al. 10170, 11711, QCA;Figs 2, 3), were collected during the first botanical expedition to the Cerro Plateado led by Dr. David A. Neill in 2012 and later in 2016 and 2021 by the first author of this paper. Both species were collected at the summit of this Andean tepui, very close to the Peruvian border, that correspond to the highest sandstone plateau of the Cordillera del Cóndor, growing as a lithophyte on exposed quartzitic sandstone rocks with acidic and nutrient poor soils and in an extremely wet environment (Figs 2, 3). The vegetation is paramo-like, dominated by the grass Chusquea cf. nana and the bromeliad Vriesea sp. along with scattered shrubs of Clethra concordia D.A.Neill, H.Beltrán & Quizhpe, Diplostephium sp., Gaultheria lanigera Hook. and Valeriana plateadensis sp. nov. Á.J. Pérez, C. Persson & J.N. Zapata (Persson et al. 2023).
Conservation status. Only one population with ca. 15 mature individuals of this species was discovered at the type locality at the summit of the Cerro Plateado. It is an isolated area and difficult to access; nevertheless, climate change could affect the environmental requirements of this species. According to the IUCN Red List criteria (IUCN Standards and Petitions Committee 2022) this species is assessed as Vulnerable (VU, Criterion D2).
Affinities. P. ombrophila is superficially very similar to P. rosmarieae with which it shares the wide leaves without involute margins and the resulting large and relatively flat rosettes. Interestingly, both P. ombrophila and P. rosmarieae are lithophytic (the latter rarely epiphytic) plants that prefer the wettest of all non-submerse locations available in the generally very moist paramo habitats. The rosette morphology likely is an adaptation to that and might be subject to environmental constraints. P. rosmarieae shows, however, a different flower morphology with a peculiar box-like corolla-spur complex and the lack of a yellow palate. The flowers of P. ombrophila are more similar to P. calyptrata and the three species are obviously closely related to each other. P. ombrophila is furthermore characterized by a very short flower stalk compared to all other north-Andean taxa. The flowers barely exceed the tips of the freshly developing leaves, a character so far only known and typical for, for example, the south-Andean P. nahuelbutensis (Gluch 2017).

Discussion
The results presented in this study show that the assessment of the neotropical biodiversity is far from complete. Even in well-known groups such as the carnivorous plants, new taxa are continuously discovered and described, in particular from remote areas that become accessible in the course of the unlimited urban sprawl. This is both encouraging and worrying at the same time. While there are evidently still pristine habitats left that inhabit an unknown biodiversity, the fact that these ecosystems are now at an accessible distance to human infrastructure puts them under immediate threat of exploitation and destruction. In particular, the eastern slopes and Andean foothills facing Amazonia in the northern Andes border such remaining microhabitats in the form of isolated paramo sites and cloud forest fragments. Their seclusion in relation to similar neighboring habitats by the rugged terrain led to a mosaic of small scale organismal communities that are often self-contained to a certain extent. In the case of flowering plants, the degree of isolation often is determined by the associated pollen vector, usually the pollinating fauna. While spatial isolation of plant-pollinator communities is considered the main driver of diversification in angiosperms, intrapopulation competition for resources can also play an important role. In the case of insectivorous Pinguicula two potentially limited resources appear to be essential: pollinators and prey. Striving for these resources between neighboring populations can lead to rapid phenotypic diversifications in morphologically plastic parts of the plant body such as leaves and flowers. Furthermore, in co-occurring taxa, these changes can be strongly accentuated towards a differentiation by a phenomenon known as character displacement (Beans 2014). An adaptation for different prey by e.g. the different shape and orientation of the leaves in the sympatric taxon pairs found can only be assumed. However, the differences observed in the flower morphology between populations, both sympatric and throughout the entire distribution range of P. calyptrata s.l., point at effective reproductive barriers. The fact that, as far as is known, all species of the section (i.e. Ampullipalatum) can be artificially hybridized, but no hybrids have been reported from the wild (Fleischmann 2021) indicates that there are no genetic barriers, but other factors in effect. Plant-pollinator interactions can be very specific and are often controlled by multiple factors, especially in heterogeneous mosaic-like landscapes such as the high Andes. In addition to the morphological adaptations of flowers (e.g., color, size, spur length), traits such as nectar quantity and sugar concentration are often precisely matched to a particular pollinator taxon (Ackermann and Weigend 2006), and even pollen presentation can be the subject of complex and specific plantpollinator interactions (Henning et al. 2018). Although speculative, such processes likely play a key role in the ongoing diversification of Pinguicula and evidently in the phenotypic divergence observed in the Ecuadorian taxa. Studies on the pollination biology of Pinguicula are scarce, the specificity of their pollination ecology is largely unknown, and appropriate research is strongly encouraged. However, published pollinator observations as e.g. in the recently described P. warijia from Mexico (Zamudio et al. 2023), provide evidence for a single principal pollinator (butterflies) and a respective close interaction.

Morphological variation of Pinguicula calyptrata Kunth
Pinguicula calyptrata shows a great morphological variability in all phenological aspects throughout its distributional range and the different populations considerably vary in size, shape and color of all parts of the plant body (Figs 2, 3). Since the protologues of the "historical" taxa are usually based on a single collection and neither provide measurements, nor by any means outline a morphological spectrum, the real taxonomic extent of Pinguicula remains hidden in the broad taxonomic concept pursued ever since (Casper and Hellwig 2019;Casper et al. 2020). This is particularly true for Pinguicula calyptrata and likely holds true for other South American taxa not yet studied in the necessary depth. The present description of the two new species should be understood as the continuation of an ongoing taxonomic extension of the genus as anticipated once a thorough molecular study including material from as many populations as possible can be conducted. Such a study has recently shed light on the taxonomy and nomenclature of the P. vulgaris aggregate in Europe, which, based on conflicting taxonomic concepts and different methodological approaches, were subject to scientific debates for decades (Majeský et al. 2022). The earlier separations of P. jarmilae, P. nahuelbutensis and P. rosmarieae from their putative sister taxa (P. involuta, P. antarctica & P. calyptrata respectively) were already founded on the evaluation of new collections and their morphological divergence from the broad circumscriptions of the historical taxa. In the case of P. jarmilae, which was later and hence invalidly also described as P. chuquisacensis by Beck et al. (2008), the latter authors already conducted a molecular study to underscore species delimitation and reveal the relationships among the South American taxa. Their key findings were that, contrary to their initial assumption, not morphologically more similar P. calyptrata but P. involuta is the closest relative and that the new species showed an unexpectedly high amount of autapomorphies in its sequence data (plastid markers). Furthermore, their preliminary dataset revealed a high genetic divergence among the South American taxa relative to those found in other clades of Pinguicula. These findings contrast with the comparatively low number of taxa described from Andean South America to this day. We therefore expect that the number of taxa will grow with the availability of molecular data and a concomitant better understanding of the taxon boundaries and speciation processes in this group of plants. For the time being, we must confine ourselves to describe those taxa as new to science, whose delimitation based on morphology and biogeography alone appears objectively evident.
The newly described species and the still unsatisfactorily documented and understood diversity among Andean Pinguicula underline the need to continue with botanical explorations and taxonomic studies and intensify urgently needed conservation efforts. The actual diversity of the Andean flora, in particular that of Ecuador and Northern Peru (AHZ), is still not conclusively determined. The threats to natural ecosystems in general, and those to small scale mountainous habitats in particular, are very worrying. Pinguicula, which inhabits very specific ecological niches due to its peculiar habitat requirements, is particularly susceptible to the destructive effects of e.g. mining activities and infrastructure projects (Cross et al. 2020), which penetrate further and further into the remaining remote pristine refugia. To make things worse, climate change increasingly affects local precipitation regimes and temperature profiles, with potentially dramatic consequences for moist habitats such as the paramo wetlands of the high Andes and their unique flora and fauna.
FC-2020-0473). The expedition to Cerro Plateado in 2016 was supported by Secretaría de Educación Superior, Ciencia, Tecnología e Innovación de la República del Ecuador (SENESCYT, Arca de Noé Initiative; S. R. Ron and O.Torres-Carvajal, Principal Investigators) and in 2021 by the International Palm Society (IPS) Endowment Fund and by Claes Persson (University of Gothenburg), the expedition also received partial funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation program (grant agreement No. 865787,GLOBAL project). The Pontificia Universidad Católica del Ecuador and Herbario QCA provided logistic and technical facilities after the trips. The reviews provided by Daniel Montesinos Tubée and an anonymous reviewer are gratefully acknowledged. The Open Access Fund of the Leibniz Association covered the publication costs for this article.