02147nas a2200265 4500008004100000022001400041245013300055210006900188260001600257300001100273490000800284520132700292653002601619653002001645653002101665653000801686653001401694653001501708100001801723700002201741700002601763700002301789700002101812856004801833 2026 eng d a1055-790300aPhylogenetic reconstruction of African weakly electric fish (Campylomormyrus spp.) is suggestive of ancient introgression events0 aPhylogenetic reconstruction of African weakly electric fish Camp cJan-01-2026 a1084520 v2143 a

The African weakly electric fish genus Campylomormyrus belongs to the species-rich family Mormyridae and comprises 15 described species. Previous species trees that focus on the genus, have included at most ten species. The resulting phylogenies have been used to explain the evolution of the species-specific electrical organ discharge (EOD) and interpreted in the context of sympatric speciation caused by disruptive selection. However, available molecular phylogenies that are focused on the Campylomormyrus genus rely only on a limited number of genetic markers. Here, we used ddRAD sequencing to genotype 65 individuals from twelve Campylomormyrus species and Gnathonemus petersii at more than 8,000 SNPs. We reconstruct a robust species tree that is suggestive of ancient introgression events between different species, which might affect both genetic and trait variability in the genus. Different levels of genetic and EOD diversity in two significant Campylomormyrus clades reveals that i) certain species might be in early stages of their speciation process, ii) that hybridization/introgression occurs in mormyrids and iii) that EOD-related genes could be potential speciation genes. Our genomic approach demonstrates the significance of these findings in understanding the evolution of Campylomormyrus.

10aAncient introgression10aCampylomormyrus10addRAD sequencing10aEOD10aphylogeny10aspeciation1 aCanitz, Julia1 aKirschbaum, Frank1 aVendrami, David, L.J.1 aMamonekene, Victor1 aTiedemann, Ralph uhttps://doi.org/10.1016/j.ympev.2025.10845202679nas a2200241 4500008004100000022002500041245011900066210006900185260001600254300000900270520193000279653001602209653002402225653001602249653001202265653002002277653001602297100001802313700001602331700002202347700002102369856004702390 2024 eng d a0302-766X, 1432-087800aOrganization of the stalk system on electrocytes in mormyrid weakly electric fish Campylomormyrus compressirostris0 aOrganization of the stalk system on electrocytes in mormyrid wea cDec-20-2024 a1-173 a

The adult electric organ in weakly electric mormyrid fish consists of action-potential-generating electrocytes, structurally and functionally modified skeletal muscle cells. The electrocytes have a disc-shaped portion and, on one of its sides, numerous thin processes, termed stalklets. These unite to stalks leading to a single main stalk that carries the innervation site. Here, we describe the 3-dimensional layout of the stalklet/stalk system in adult Campylomormyrus compressirostris by differential interference contrast microscopy and confocal fluorescence microscopy. Using antibodies against Na+/K+-ATPase α-subunit and plasma membrane Ca2+-ATPase, we show that these ion pumps are differentially distributed over the stalklet/stalk system, with plasma membrane Ca2+-ATPase being enriched on the stalklet membrane. Stalklets are distributed and organized in a quite uniform pattern on the posterior face of the electrocyte disc and fuse to terminal stalks. The latter then unite in a mostly dichotomic mode to stalks of increasing thickness, with the main stalk measuring about 100 µm in diameter. We further analyse the structural organization of stalklets and stalks, with a characteristic cytoskeletal system of bundled actin filaments in the centre and nuclei in subsurface position. These results suggest that the stalklet/stalk system is adapted in its structural layout to generate an action potential highly synchronized over the entire disc-portion of the electrocyte, accounting for the short electric organ discharge in this species. Our results suggest that actin-related proteins overexpressed in electrocytes, as shown previously by transcriptome analysis, may be involved in the organization of the unique F-actin system in stalklets and stalks.

10aCa2+-ATPase10aConfocal microscopy10aelectrocyte10aF-actin10aPlasma membrane10aSodium pump1 aBaumann, Otto1 aCheng, Feng1 aKirschbaum, Frank1 aTiedemann, Ralph uhttps://doi.org/10.1007/s00441-024-03938-y00506nas a2200145 4500008004100000245007600041210006900117490000800186100002900194700001700223700002200240700002300262700002300285856005200308 2019 eng d00aSilencing the spark: CRISPR/Cas9 genome editing in weakly electric fish0 aSilencing the spark CRISPRCas9 genome editing in weakly electric0 v1521 aConstantinou, Savvas, J.1 aNguyen, Linh1 aKirschbaum, Frank1 aSalazar, Vielka, L1 aGallant, Jason, R. uhttps://mormyrids.efishgenomics.com/en/node/65402556nas a2200277 4500008004100000022002500041245011800066210006900184260001600253300001000269490000800279520169400287653002801981653001302009653002002022653002602042653002602068653001302094100001702107700002102124700002302145700001902168700002102187700002202208856004802230 2016 eng d a0378-1909, 1573-513300aReproduction and development in some species of the weakly electric genus Campylomormyrus (Mormyridae, Teleostei)0 aReproduction and development in some species of the weakly elect cDec-07-2016 a49-680 v1003 a

Reproduction in captivity of four species of the mormyrid genus Campylomormyrus was investigated. Cyclical reproduction was provoked by changing water conductivity (C) alone: decreasing C led to gonadal recrudescence, an increase induced gonad regression. Data on the reproduction and development of three species are presented. All three species are indeterminate fractional spawners. Spawning intervals ranged from 6 to 66 days in C. rhynchophorus, 10–75 days in C. tshokwe, and 18 days in C. compressirostris (calculated values). Fecundities (eggs per fractional spawning) ranged from 70 to 1570 eggs in C. rhynchophorus, 100–1192 in C. tshokwe, and 38–246 in C. compressirostris. Spawnings/ovipositions occurred during the second half of the night; no parental care was observed; no special spawning substrates were necessary. C. compressirostris successfully spawned in breeding groups, C. rhynchophorus as pair. Agonistic behavior in the C. tshokwe pair forced us to divide the breeding tank; therefore, only ovipositions occurred. However, injection of an artificial GnRH hormone allowed us to obtain ripe eggs and sperm and to perform successful artificial reproduction. All three species produce yolky, slightly sticky eggs. Egg diameter ranges from 2.3–3.0 mm. Hatching occurred on day 3, feeding started on day 11. Transition from larval to juvenile stage occurred at around 20 mm total length (TL). At this size C. rhynchophorus developed a higher body than the two other species and differences between the species in the melanin pigmentation of the unpaired fins occurred. Between 32 and 35 mm TL the upper and lower jaws started to elongate.

10aArtificial reproduction10aBehavior10aCampylomormyrus10aCyclical reproduction10aEnvironmental factors10aOntogeny1 aNguyen, Linh1 aPaul, Christiane1 aMamonekene, Victor1 aBartsch, Peter1 aTiedemann, Ralph1 aKirschbaum, Frank uhttp://dx.doi.org/10.1007/s10641-016-0554-103020nas a2200217 4500008004100000245015300041210006900194260001700263520226100280653001702541653001602558653001802574653001502592653001902607100001902626700002202645700002102667700002602688700001602714856007202730 2024 eng d00aA new type of spermiogenesis in teleost fish: formation of the aflagellate sperm in Campylomormyrus compressirostris (Osteoglossomorpha: Mormyridae)0 anew type of spermiogenesis in teleost fish formation of the afla c29 June 20243 a

Osteoglossomorpha, the bony tongue fishes, show great variation in morphology, behavioural strategies, reproductive biology and gamete ultrastructure. The order Osteoglossiformes is the only vertebrate taxon, in which four types of sperm (monoflagellate, biflagellate and aflagellate aquasperm and the complex introsperm) have been described. It is also the only vertebrate lineage in which aflagellate spermatozoa exist. The aim of this study was to analyse the structure of the testis and the process of spermiogenesis in the mormyrid Campylomormyrus compressirostris during the breeding season using light and electron microscopy (transmission and scanning). Males of this species have a single testis of the anastomosing tubular type. The tubules of the anterior part of the testis contain cysts with developing germ cells, and this region is much wider than the posterior part, which consists of efferent ducts filled with sperm cells. The cysts are filled with single or mitotic spermatogonia, primary and secondary spermatocytes and early spermatids. At the stage of spermatids with fine granular chromatin, the cysts rupture and successive stages of spermatid differentiation take place in the testicular lumen; we therefore characterise this process as ‘extracystic spermiogenesis’. Sperm development in C. compressirostris is extremely simple and involves chromatin condensation in the central region of the nucleus, a slight decrease in nuclear volume, the appearance of numerous vesicles in the cytoplasm that form a tubular-vesicular system at the base of the nucleus. Both centrioles and mitochondria are translocated to the peripheral region of the midpiece, which forms the opposite pole to the nucleus. There are many differences between the types of spermiogenesis described so far in teleosts and that found in C. compressirostris, including the loss of flagellum formation. This unique type of spermiogenesis is restricted to species of the families Mormyridae and Gymnarchidae, all of which possess aflagellate spermatozoa. Our data demonstrate that the spermatid differentiation and existence of the aflagellate spermatozoon are a unique phenomena not only among teleosts but also in the whole vertebrate lineage.

10aelephantfish10apermatozoon10aSertoli cells10aspermatids10aTubular testis1 aDymek, Anna, M1 aKirschbaum, Frank1 aTiedemann, Ralph1 aSiemiński, Krzysztof1 aPecio, Anna uhttps://www.sciencedirect.com/science/article/pii/S094420062400045X02463nas a2200157 4500008004100000022002500041245016500066210006900231260001600300520186800316100001602184700002202200700002202222700002102244856004002265 2023 eng d a0962-1083, 1365-294X00aDiet in phenotypically divergent sympatric species of African weakly electric fish (genus: Campylomormyrus)—A hybrid capture/HTS metabarcoding approach0 aDiet in phenotypically divergent sympatric species of African we cDec-21-20233 a

Ecological speciation within the mormyrid genus Campylomormyrus resulted in sympatric species exhibiting divergence in their feeding apparatus and electric organ discharge (EOD). This study documents the overall diet of the genus Campylomormyrus and examines the hypothesis that the Campylomormyrus radiation is caused by adaptation to different food sources. We performed diet assessment of five sympatric Campylomormyrus species (C. alces, C. compressirostris, C. curvirostris, C. tshokwe, C. numenius) and their sister taxon Gnathonemus petersii with markedly different snout morphologies and EODs using hybrid capture/HTS DNA metabarcoding of their stomach contents. Our approach allowed for high taxonomic resolution of prey items, including benthic invertebrates, allochthonous invertebrates and vegetation. Comparisons of the diet compositions using quantitative measures and diet overlap indices revealed that all species are able to exploit multiple food niches in their habitats, that is fauna at the bottom, the water surface and the water column. A major part of the diet is larvae of aquatic insects, such as dipterans, coleopterans and trichopterans, known to occur in holes and interstitial spaces of the substrate. The results indicate that different snout morphologies and the associated divergence in the EOD could translate into different prey spectra. This suggests that the diversification in EOD and/or morphology of the feeding apparatus could be under functional adaptation.

1 aAmen, Rahma1 aHavenstein, Katja1 aKirschbaum, Frank1 aTiedemann, Ralph uhttp://dx.doi.org/10.1111/mec.1724801433nas a2200145 4500008004100000022002500041245006500066210006500131260001600196520097300212100002301185700002201208700001701230856004001247 2023 eng d a0021-8782, 1469-758000aOntogeny reveals the origin of Gemminger bones in Mormyridae0 aOntogeny reveals the origin of Gemminger bones in Mormyridae cJul-25-20233 a

Mormyridae are well known and intensively studied for their weak electric organ discharges, which facilitate communication and orientation. The Gemminger bones of Mormyridae are located next to the electrical organ in the caudal peduncle; however, they have not attracted much interest until recently. Therefore, we investigated the diversity of Gemminger bones in mormyrids and studied their ontogenetic development in Mormyrus rume proboscirostris. Gemminger bones are paired, thin, elongated ossifications lying on the dorsal and ventral sides of the caudal peduncle, and usually reach anterior well below the dorsal and anal fin bases. Ontogeny revealed that they are not intermuscular ossifications, as suspected based on the anatomical position of this structure and the systematic position of the mormyrids. Instead, they are membrane ossifications that originate from the fin stays of the dorsal and anal fins.

1 aKoch, Ann‐Katrin1 aKirschbaum, Frank1 aMoritz, Timo uhttp://dx.doi.org/10.1111/joa.1393504135nas a2200193 4500008004100000022001400041245016700055210006900222260001600291490000700307520344900314100001603763700002203779700003003801700001803831700002203849700002103871856004903892 2023 eng d a1471-216400aA new genome assembly of an African weakly electric fish (Campylomormyrus compressirostris, Mormyridae) indicates rapid gene family evolution in Osteoglossomorpha0 anew genome assembly of an African weakly electric fish Campylomo cMar-20-20230 v243 a

Abstract Background Teleost fishes comprise more than half of the vertebrate species. Within teleosts, most phylogenies consider the split between Osteoglossomorpha and Euteleosteomorpha/Otomorpha as basal, preceded only by the derivation of the most primitive group of teleosts, the Elopomorpha. While Osteoglossomorpha are generally species poor, the taxon contains the African weakly electric fish (Mormyroidei), which have radiated into numerous species. Within the mormyrids, the genus Campylomormyrus is mostly endemic to the Congo Basin. Campylomormyrus serves as a model to understand mechanisms of adaptive radiation and ecological speciation, especially with regard to its highly diverse species-specific electric organ discharges (EOD). Currently, there are few well-annotated genomes available for electric fish in general and mormyrids in particular. Our study aims at producing a high-quality genome assembly and to use this to examine genome evolution in relation to other teleosts. This will facilitate further understanding of the evolution of the osteoglossomorpha fish in general and of electric fish in particular. Results A high-quality weakly electric fish (C. compressirostris) genome was produced from a single individual with a genome size of 862 Mb, consisting of 1,497 contigs with an N50 of 1,399 kb and a GC-content of 43.69%. Gene predictions identified 34,492 protein-coding genes, which is a higher number than in the two other available Osteoglossomorpha genomes of Paramormyrops kingsleyae and Scleropages formosus. A Computational Analysis of gene Family Evolution (CAFE5) comparing 33 teleost fish genomes suggests an overall faster gene family turnover rate in Osteoglossomorpha than in Otomorpha and Euteleosteomorpha. Moreover, the ratios of expanded/contracted gene family numbers in Osteoglossomorpha are significantly higher than in the other two taxa, except for species that had undergone an additional genome duplication (Cyprinus carpio and Oncorhynchus mykiss). As potassium channel proteins are hypothesized to play a key role in EOD diversity among species, we put a special focus on them, and manually curated 16 Kv1 genes. We identified a tandem duplication in the KCNA7a gene in the genome of C. compressirostris. Conclusions We present the fourth genome of an electric fish and the third well-annotated genome for Osteoglossomorpha, enabling us to compare gene family evolution among major teleost lineages. Osteoglossomorpha appear to exhibit rapid gene family evolution, with more gene family expansions than contractions. The curated Kv1 gene family showed seven gene clusters, which is more than in other analyzed fish genomes outside Osteoglossomorpha. The KCNA7a, encoding for a potassium channel central for EOD production and modulation, is tandemly duplicated which may related to the diverse EOD observed among Campylomormyrus species.

1 aCheng, Feng1 aDennis, Alice, B.1 aOsuoha, Josephine, Ijeoma1 aCanitz, Julia1 aKirschbaum, Frank1 aTiedemann, Ralph uhttp://dx.doi.org/10.1186/s12864-023-09196-603202nas a2200301 4500008004100000022001400041245015100055210006900206260000900275300001300284490000700297520224500304653001202549653001802561653001902579653001602598653002002614653003002634653003102664653001402695653003602709653002402745653001802769100001802787700002202805700002102827856005202848 2020 eng d a1932-620300aTranscriptome-wide single nucleotide polymorphisms related to electric organ discharge differentiation among African weakly electric fish species.0 aTranscriptomewide single nucleotide polymorphisms related to ele c2020 ae02408120 v153 a

African weakly electric fish of the mormyrid genus Campylomormyrus generate pulse-type electric organ discharges (EODs) for orientation and communication. Their pulse durations are species-specific and elongated EODs are a derived trait. So far, differential gene expression among tissue-specific transcriptomes across species with different pulses and point mutations in single ion channel genes indicate a relation of pulse duration and electrocyte geometry/excitability. However, a comprehensive assessment of expressed Single Nucleotide Polymorphisms (SNPs) throughout the entire transcriptome of African weakly electric fish, with the potential to identify further genes influencing EOD duration, is still lacking. This is of particular value, as discharge duration is likely based on multiple cellular mechanisms and various genes. Here we provide the first transcriptome-wide SNP analysis of African weakly electric fish species (genus Campylomormyrus) differing by EOD duration to identify candidate genes and cellular mechanisms potentially involved in the determination of an elongated discharge of C. tshokwe. Non-synonymous substitutions specific to C. tshokwe were found in 27 candidate genes with inferred positive selection among Campylomormyrus species. These candidate genes had mainly functions linked to transcriptional regulation, cell proliferation and cell differentiation. Further, by comparing gene annotations between C. compressirostris (ancestral short EOD) and C. tshokwe (derived elongated EOD), we identified 27 GO terms and 2 KEGG pathway categories for which C. tshokwe significantly more frequently exhibited a species-specific expressed substitution than C. compressirostris. The results indicate that transcriptional regulation as well cell proliferation and differentiation take part in the determination of elongated pulse durations in C. tshokwe. Those cellular processes are pivotal for tissue morphogenesis and might determine the shape of electric organs supporting the observed correlation between electrocyte geometry/tissue structure and discharge duration. The inferred expressed SNPs and their functional implications are a valuable resource for future investigations on EOD durations.

10aAnimals10aelectric fish10aelectric organ10aElectricity10aGene Expression10aGene Expression Profiling10aGene Expression Regulation10aPhenotype10aPolymorphism, Single Nucleotide10aSpecies Specificity10atranscriptome1 aCanitz, Julia1 aKirschbaum, Frank1 aTiedemann, Ralph uhttps://mormyrids.efishgenomics.com/en/node/62601616nas a2200169 4500008004100000022001400041245015000055210006900205260001600274520099000290100002601280700002501306700002001331700002101351700002201372856005201394 2022 eng d a1432-135100aIntergenus F1-hybrids of African weakly electric fish (Mormyridae: Gnathonemus petersii ♂ × Campylomormyrus compressirostris ♀) are fertile.0 aIntergenus F1hybrids of African weakly electric fish Mormyridae  c2022 Feb 043 a

Hybridisation is an important element of adaptive radiation in fish but data are limited in weakly electric mormyrid fish in this respect. Recently, it has been shown that intragenus hybrids (Campylomormyrus) are fertile and are able to produce F2-fish. In this paper, we demonstrate that even intergenus hybrids (Gnathonemus petersii ♂ × Campylomormyrus compressirostris ♀) are fertile. Three artificial reproduction (AR) trials, with an average fertilisation rate of ca. 23%, yielded different numbers of survivals (maximally about 50%) of the F1-hybrids. The complete ontogenetic development of these hybrids is described concerning their morphology and electric organ discharge (EOD). Two EOD types emerged at the juvenile stage, which did not change up to adulthood. Type I consisted of four phases and Type II was triphasic. The minimum body length at sexual maturity was between 10 and 11 cm. Malformations, growth and mortality rates are also described.

1 aKorniienko, Yevheniia1 aNzimora, Kingsley, C1 aVater, Marianne1 aTiedemann, Ralph1 aKirschbaum, Frank uhttps://mormyrids.efishgenomics.com/en/node/60802043nas a2200205 4500008004100000022001400041245014400055210006900199260001600268300001400284490000800298520134200306100002601648700001701674700002701691700002001718700002101738700002201759856005601781 2020 eng d a0340-759400aIntragenus F1-hybrids of African weakly electric fish (Mormyridae: Campylomormyrus tamandua ♂ × C. compressirostris ♀) are fertile0 aIntragenus F1hybrids of African weakly electric fish Mormyridae  cJan-07-2020 a571 - 5850 v2063 a

Hybridization is widespread in fish and constitutes an important mechanism in fish speciation. There is, however, little knowledge about hybridization in mormyrids. F1-interspecies hybrids between Campylomormyrus tamandua ♂ × C. compressirostris ♀ were investigated concerning: (1) fertility; (2) survival of F2-fish and (3) new gene combinations in the F2-generation concerning the structure of the electric organ and features of the electric organ discharge. These F1-hybrids achieved sexual maturity at about 12–13.5 cm total length. A breeding group comprising six males and 13 females spawned 28 times naturally proving these F1-fish to be fertile. On average 228 eggs were spawned, the average fertilization rate was 47.8%. Eggs started to hatch 70–72 h after fertilization, average hatching rate was 95.6%. Average mortality rate during embryonic development amounted to 2.3%. Average malformation rate during the free embryonic stage was 27.7%. Exog- enous feeding started on day 11. In total, we raised 353 normally developed larvae all of which died consecutively, the oldest specimen reaching an age of 5 months. During survival, the activities of the larval and adult electric organs were recorded and the structure of the adult electric organ was investigated histologically.

1 aKorniienko, Yevheniia1 aNguyen, Linh1 aBaumgartner, Stephanie1 aVater, Marianne1 aTiedemann, Ralph1 aKirschbaum, Frank uhttp://link.springer.com/10.1007/s00359-020-01425-702385nas a2200157 4500008004100000022001400041245011300055210006900168260001600237520163800253100002601891700002101917700002001938700002201958856024701980 2020 eng d a0021-996700aOntogeny of the electric organ discharge and of the papillae of the electrocytes in the weakly electric fish0 aOntogeny of the electric organ discharge and of the papillae of  cDec-08-20203 a

The electric organ of the mormyrid weakly electric fish, Campylomormyrus rhynchophorus (Boulenger, 1898), undergoes changes in both the electric organ discharge (EOD) and the light and electron microscopic morphology as the fish mature from the juvenile to the adult form. Of particular interest was the appearance of papillae, surface specializations of the uninnervated anterior face of the electrocyte, which have been hypothesized to increase the duration of the EOD. In a 24.5 mm long juvenile the adult electric organ (EO) was not yet functional, and the electrocytes lacked papillae. A 40 mm long juvenile, which produced a short biphasic EOD of 1.3 ms duration, shows small papillae (average area 136 μm2). In contrast, the EOD of a 79 mm long juvenile was triphasic. The large increase in duration of the EOD to 23.2 ms was accompanied by a small change in size of the papillae (average area 159 μm2). Similarly, a 150 mm long adult produced a triphasic EOD of comparable duration to the younger stage (24.7 ms) but featured a prominent increase in size of the papillae (average area 402 μm2). Thus, there was no linear correlation between EOD duration and papillary size. The most prominent ultrastructural change was at the level of the myofilaments, which regularly extended into the papillae, only in the oldest specimen – probably serving a supporting function. Physiological mechanisms, like gene expression levels, as demonstrated in some Campylomormyrus species, might be more important concerning the duration of the EOD.

1 aKorniienko, Yevheniia1 aTiedemann, Ralph1 aVater, Marianne1 aKirschbaum, Frank uhttps://onlinelibrary.wiley.com/doi/abs/10.1002/cne.25003https://onlinelibrary.wiley.com/doi/pdf/10.1002/cne.25003https://api.wiley.com/onlinelibrary/tdm/v1/articles/10.1002%2Fcne.25003https://onlinelibrary.wiley.com/doi/pdf/10.1002/cne.2500301976nas a2200169 4500008004100000022001400041245012300055210006900178260001600247520138800263100001601651700001901667700002101686700002201707700002101729856005601750 2020 eng d a0269-765300aMorphological differentiation in African weakly electric fish (genus Campylomormyrus) relates to substrate preferences0 aMorphological differentiation in African weakly electric fish ge cApr-04-20213 a

Under an ecological speciation scenario, the radiation of African weakly electric fish (genus Campylomormyrus) is caused by an adaptation to different food sources, associated with diversification of the electric organ discharge (EOD). This study experimentally investigates a phenotype–environment correlation to further support this scenario. Our behavioural experiments showed that three sympatric Campylomormyrus species with significantly divergent snout morphology differentially react to variation in substrate structure. While the short snout species (C. tamandua) exhibits preference to sandy substrate, the long snout species (C. rhynchophorus) significantly prefers a stone substrate for feeding. A third species with intermediate snout size (C. compressirostris) does not exhibit any substrate preference. This preference is matched with the observation that long-snouted specimens probe deeper into the stone substrate, presumably enabling them to reach prey more distant to the substrate surface. These findings suggest that the diverse feeding apparatus in the genus Campylomormyrus may have evolved in adaptation to specific microhabitats, i.e., substrate structures where these fish forage. Whether the parallel divergence in EOD is functionally related to this adaptation or solely serves as a prezygotic isolation mechanism remains to be elucidated.

1 aAmen, Rahma1 aNagel, Rebecca1 aHedt, Maximilian1 aKirschbaum, Frank1 aTiedemann, Ralph uhttp://link.springer.com/10.1007/s10682-020-10043-3