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Morphological traits of the two Arisaema species. Length of (A) pseudostem, (B) peduncle, (C) petiole of terminal leaflet, (D) rachis between terminal leaflet and adjacent leaflet, (E) terminal leaflet. (F) Width of terminal leaflet. (G) Number of leaflets on upper leaf. (H) Length of spathe. (I) Length and (J) width of spathe blade. (K) Width of the spadix appendage at apex. Each trait was selected as per Fukai (2007), Murata and Kakishima (2008), and Hayakawa et al. (2010).
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Premise:
The genus Arisaema (Araceae) has rapidly diversified in Japan, and multiple species often coexist in the field. Although Japanese Arisaema species hybridize from artificial crossing, hybrid individuals are rare in mixed populations; suggesting the presence of effective pre-pollination barriers. We examined the following reproductive barri...
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Context 1
... verify the occurrence of putative hybrids with an intermediate morphology (Murata, 1962;Hayakawa et al., 2010;Matsuyama et al., 2013;Murata et al., 2018), we measured 11 morphological traits ( Fig. 2) with a folding scale and digital caliper for flowering individuals of A. sikokianum (80 males and 24 females) and A. tosaense (66 males and 19 females) from 5 May 2017 to 6 June 2017. Uneven numbers of male and female individuals were examined due to the extremely malebiased sex ratio in Arisaema populations ( Richardson and Clay, ...
Context 2
... estimate the frequency of inter specific hybridization, we compared 11 morphological traits (Fig. 2) by nonmetric multidimensional scaling (NMDS). We calculated morphological dissimilarity between pairs of Arisaema plants using the Bray-Curtis distance matrix. The number of dimensions in NMDS was determined based on Kruskal's stress, an indicator for the goodness of fit ( Quinn and Keuogh, 2002). The analysis was terminated when the ...
Context 3
... results of NMDS for the 11 mor phological traits (Fig. 2) were well rep resented in the two dimensions (final stress value = 0.11). The NMDS plot showed that the mixed population was clearly divided into A. sikokianum and A. tosaense (Fig. 4). Convex hulls of NMDS scores, representing intraspe cific variation in the morphological traits, did not overlap between the two Arisaema species ...
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Reproductive isolation plays an important role in maintaining the species integrity of sympatric close relatives. For sympatric Arisaema species, interspecific gene flow is expected to be effectively prevented by pre‐pollination barriers, particularly strong pollinator isolation mediated by fungus gnats. However, due to the lack of quantitative stu...
Citations
... Suetsugu et al. (2021) suggested that sexual deception could be more widespread, especially because some Arisaema species predominantly attract male fungus gnats, which may indicate the emission of pheromones that mimic female gnats. Nevertheless, there are notable exceptions, such as Arisaema sikokianum, in which the sterile appendix exhibits remarkable fungal mimicry, closely resembling both the appearance and scent of mushrooms (Kakishima et al., 2019;Matsumoto et al., 2019). ...
... Deceptive plants often experience extremely low pollinator visitation, making direct observation of pollinator behavior challenging (Widmer et al., 2000). Therefore, we adopted an approach to assess the pollinator status, drawing from previous research on Arisaema pollination ecology (Kakishima et al., 2019;Matsumoto et al., 2019Matsumoto et al., , 2021Suetsugu et al., 2021Suetsugu et al., , 2022Vogel & Martens, 2000). Our methodology involved examining the pollinator assemblages of A. thunbergii by collecting floral visitors trapped inside 62 spathes, comprising 39 male and 23 female ones. ...
... Additionally, the potential for pollen transfer among insect individuals within the trap complicates interpretations (Kubo et al., 2023). Therefore, to establish a clear definition of pollinators, we applied the following criteria to insect species, established from prior research: (i) they exhibited entrapment in both sealed male and female spathes, and (ii) they possessed a size conducive to passing through the exit hole of the male spathe, thereby facilitating pollen transport (Kakishima et al., 2019;Matsumoto et al., 2019Matsumoto et al., , 2021Suetsugu et al., 2021Suetsugu et al., , 2022Vogel & Martens, 2000). ...
Societal Impact Statement
The research explores the complex interaction between flowering plants and their pollinators, specifically focusing on the genus Arisaema . Unlike most plants, Arisaema has a distinctive trait in which killing pollinators can be beneficial. Traditionally, this interaction has been viewed as highly antagonistic because it appears to favor the plants at the expense of the pollinators. However, new evidence reveals that a pollinator uses the lethal floral trap of Arisaema thunbergii as a nursery. Remarkably, some individuals probably even escape from the trap after laying eggs. This finding challenges the prevailing notion that deceptive pollination is the sole outcome in Arisaema , a genus known for its intricate lethal pollination mechanisms.
Summary
While many flowering plants engage in mutualistic relationships with their pollinators, flower‐insect interactions are not always mutually beneficial. Some plants exploit their pollinators by deceptively promising rewards without delivering them. Among such plants, the genus Arisaema (Araceae) is notable for its antagonistic behavior towards pollinators, as female plants invariably entrap them with lethal effects.
Here we tested the hypothesis that the succulent appendix of some Arisaema species, such as Arisaema thunbergii , may serve as a food source for pollinator larvae, leading to the possibility of nursery pollination. To investigate this, we retrieved adult insect corpses trapped within the spathes of A. thunbergii . Subsequently, we incubated the spathes after collecting the corpses to determine if conspecific insects emerged from them.
We present novel evidence that one of the main pollinating fungus gnats, Leia ishitanii , uses the lethal floral trap of A. thunbergii inflorescences as a nursery. Astonishingly, some L. ishitanii individuals may even succeed in escaping from the female spathe after depositing their eggs.
These findings challenge the conventional understanding that deceptive pollination is the norm within the genus Arisaema . The interaction between A. thunbergii and L. ishitanii , in which a significant portion of female adults die inside but still reproduction seems successful, suggests an intermediate stage between brood‐site deception and nursery pollination mutualism.
... In many regions, Arisaema species often overlap in their distribution (Matsumoto et al., 2018;Murata, 1995;Serizawa, 1988Serizawa, , 1997. It is generally accepted that interspecific gene flow among sympatric Arisaema species is effectively prevented by pre-pollination isolation (Matsumoto et al., 2019(Matsumoto et al., , 2021Murata et al., 2018). Divergence of habitat and flowering time reduced interspecific gene flow to some extent among sympatric Arisaema species (Matsumoto et al., 2019(Matsumoto et al., , 2021Murata & Ohno, 1989). ...
... It is generally accepted that interspecific gene flow among sympatric Arisaema species is effectively prevented by pre-pollination isolation (Matsumoto et al., 2019(Matsumoto et al., , 2021Murata et al., 2018). Divergence of habitat and flowering time reduced interspecific gene flow to some extent among sympatric Arisaema species (Matsumoto et al., 2019(Matsumoto et al., , 2021Murata & Ohno, 1989). In contrast to relatively weak ecogeographic isolation and phenological isolation, selective fungus gnat visitation resulted in strong pollinator isolation among some Arisaema species (Kakishima et al., 2019(Kakishima et al., , 2020Matsumoto et al., 2019Matsumoto et al., , 2021Suetsugu et al., 2021). ...
... Divergence of habitat and flowering time reduced interspecific gene flow to some extent among sympatric Arisaema species (Matsumoto et al., 2019(Matsumoto et al., , 2021Murata & Ohno, 1989). In contrast to relatively weak ecogeographic isolation and phenological isolation, selective fungus gnat visitation resulted in strong pollinator isolation among some Arisaema species (Kakishima et al., 2019(Kakishima et al., , 2020Matsumoto et al., 2019Matsumoto et al., , 2021Suetsugu et al., 2021). The strong pollinator isolation among certain Arisaema species might serve as a compensatory mechanism for weakened early-acting barriers, thereby ensuring the prevention of interspecific hybridization (Matsumoto et al., 2021). ...
Reproductive isolation plays an important role in maintaining the species integrity of sympatric close relatives. For sympatric Arisaema species, interspecific gene flow is expected to be effectively prevented by pre‐pollination barriers, particularly strong pollinator isolation mediated by fungus gnats. However, due to the lack of quantitative studies combining multiple pre‐ and post‐pollination barriers, it is not known whether pre‐pollination isolation is complete, and whether post‐pollination barriers also contribute to reproductive isolation among some Arisaema species. In this study, we quantified the individual strengths and absolute contributions of four pre‐ and post‐pollination barriers (phenological isolation, pollinator isolation, hybrid fruit formation, and hybrid seed formation) among three sympatric Arisaema species ( A. bockii , A. lobatum , and A. erubescens ). Although phenological isolation and pollinator isolation reduced the frequencies of interspecific pollen transfer among these species, the partial overlap of flowering times and pollinator assemblages resulted in incomplete pre‐pollination isolation. Post‐pollination barriers also contributed to reproductive isolation at the hybrid fruit and seed formation stages. We propose that, although pre‐pollination barriers are expected to contribute more to total isolation than post‐pollination barriers in Arisaema , pre‐pollination barriers may not completely prevent interspecific pollen transfer among some Arisaema species. Post‐pollination barriers, which are generally ignored, may also have contributed significantly to reproductive isolation in Arisaema .
... On the one hand, pre-zygotic barriers that limit interspecific gene flow between congeners could explain why multiple morphotypes can coexist. Such pre-zygotic barriers could include occurrence in specific habitats (Schnitzler et al., 2011;Anacker & Strauss, 2014;Mantel & Sweigart, 2019), divergent flowering phenology (Anderson, Alexandersson & Johnson, 2010;Newman, Anderson & Johnson, 2012;Matsumoto et al., 2019;Ramírez-Aguirre et al., 2019;Osborne et al., 2020), use of different pollinators or divergent use of the same pollinator (Peakall et al., 2010;Pedron et al., 2012;Ramírez-Aguirre et al., 2019) and divergence in mechanical features leading to character displacement (Queiroz et al., 2015;Zheng et al., 2017;Newman & Anderson, 2020). Studies of the reproductive biology of S. longicauda have shown that TELW, TFLP and TFLP are visited by nocturnal moths that differ in the length of their proboscis (Harder & Johnson, 2005;Jersáková & Johnson, 2007;Ellis & Johnson, 2010;Johnson et al., 2011Johnson et al., , 2019Duffy & Johnson, 2014). ...
Species-level taxonomy is traditionally based on herbarium collections that typically include few, or even single, representatives per site. This can lead to underestimation of diversity when there are sympatric populations of superficially similar plants belonging to different lineages. Satyrium longicauda (Orchidaceae) represents a taxonomic challenge for the delimitation of species boundaries due to the high degree of morphological variation detected within and among populations. Currently, just two varieties are accepted based mainly on length differences of the lateral sepal and nectar spur. However, there is extensive morphological variation within South African populations and evidence for several pollination ecotypes, indicating that this taxon represents an actively diverging species complex. Here, we evaluate intraspecific morphological variation through uni- and multivariate morphometrics and analyse internal transcribed spacer sequences for individuals sampled from 36 sites, including 14 sites where divergent morphotypes occur sympatrically. Morphometric analyses of 1802 individuals revealed the presence of eight morphotypes based on vegetative and floral characters. Up to six morphologically and genetically distinct morphotypes can coexist in sympatry. Morphological and genetic distances among populations were significantly correlated. Phylogenetic analyses of 120 accessions indicated that neither of the two varieties nor S. longicauda as a species is monophyletic, and provided evidence for the monophyly of some of the morphotypes including the newly described S. cernuiflorum. The presence of distinct morphological and genetic sympatric variants, which in several cases scale up to distinct evolutionary lineages, is consistent with the existence of different taxa according to morphological and biological species concepts. Our results therefore confirm that taxonomy based mainly on herbarium collections can grossly under-estimate actual diversity of disparate lineages, although further work is required to finalize taxonomic decisions. These findings have implications for efforts to estimate species diversity in groups that are in the process of diversifying and for conservation practice.
... Plants that mimic mushrooms, including the exceptional fungal mimetic Arisaema species (A. sikokianum), are visited by various fungivorous insects such as Mycodrosophila species (Drosophilidae) and staphylinids (Coleoptera; Kakishima et al., 2019;Matsumoto et al., 2019;Matsumoto et al., 2021). In contrast, many other Arisaema species (especially Pistillata taxa) are exclusively pollinated by a limited number of fungus gnat species (one or two) (Suetsugu et al., 2021). ...
... Moreover, Pistillata is considered to have radiated recently based on the limited genetic distances even among morphologically distinct species(Ohi-Toma et al., 2016). Therefore, there is particular interest in understanding how the diverse group has established reproductive isolation(Kakishima et al., 2019;Matsumoto et al., 2019;Matsumoto et al., 2021;Suetsugu et al., 2021).Gene flow among the Japanese Arisaema species is likely to be restricted by premating barriers. Indeed, even though artificial cross experiments have determined the cross-compatibility among species belonging to the Pistillata species(Murata et al., 2018), these Arisaema species often co-occur in apparent reproductive isolation. ...
... Indeed, even though artificial cross experiments have determined the cross-compatibility among species belonging to the Pistillata species(Murata et al., 2018), these Arisaema species often co-occur in apparent reproductive isolation. Pollinator specialization contributes to the reproductive isolation of these species(Kakishima et al., 2019;Kakishima et al., 2020;Matsumoto et al., 2019;Matsumoto et al., 2021;Suetsugu et al., 2021). For example,Suetsugu et al. (2021) reported that the mycetophilid fungus gnats trapped by inflorescences of the sympatric, co-blooming, and closely related species Arisaema angustatum and Arisaema peninsulae differed significantly. ...
Female individuals in the genus Arisaema are the only plants known to imprison their pollinators permanently. Their lethal pollination system has long attracted the interest of evolutionary biologists and naturalists. Nonetheless, the mechanisms of how Arisaema plants lure their victims to the brink of death are only just being unveiled. Here, I briefly review the unique reproductive biology of these plants, focusing on the possibility of sexual deception to lure pollinating victims, and its role in the evolution of new species.
... Moreover, there are many examples of spontaneous and artificial hybrids among Pistillata species; most of these are highly fertile (Murata et al., 2018), except for the crosses between parent plants with different chromosome numbers (Kobayashi et al., 2005), indicating that post-zygotic isolation among Pistillata species is generally low (Murata et al., 2018). Nevertheless, several recent studies have revealed that many sympatric species of Pistillata are reproductively isolated by species-specific pollinators, which are exploited by the deadly trap mechanism of the plants Matsumoto et al., 2019Matsumoto et al., , 2021Suetsugu et al., 2021). Suetsugu et al. (2021) suggested that the appendix of the Arisaema inflorescence probably emits olfactory cues for species-specific pollinators. ...
... Okuyama and S. Kakishima, personal observations). Likewise, in Pistillata, pollinators have been reported for at least 12 species (summarized in Figure 3; Barriault et al., 2010;Kakishima et al., 2019Matsumoto et al., 2019Matsumoto et al., , 2021Suetsugu et al., 2021) but their deception mechanisms remain unknown, except for the only well-examined mushroom mimicry system in Arisaema sikokianum (Kakishima et al., 2019). Suetsugu et al. (2021) suggested pollination by sexual mimicry in two species of Pistillata, Arisaema peninsulae and Arisaema angustatum, and it is possible that sexual mimicry is more widespread in the lineage. ...
The adaptive radiation of flowering plants as manifested by the floral diversity has long been considered associated with the diversity of plant–pollinator interactions, because changes in plant–pollinator interactions are hypothesized as one of the major mechanisms driving plant ecological speciation. To understand the relative contributions of various mechanisms for plant radiation, including pollinator changes, it is useful to study a plant group for which comparative study of the species life history across the whole lineage is feasible. To this end, we will focus on the plant lineages that have presumably radiated in the Japanese archipelago, namely, the genera Asimitellaria, Asarum, and Arisaema. By comparing these three genera, we will comment on the possible modes of adaptive radiation and diversification among the endemic flora of Japan. In this paper, we focus on the three plant lineages that have presumably radiated in the Japanese archipelago, namely, the genera Asimitellaria, Asarum, and Arisaema, in which detailed studies on phylogeny, reproductive isolation, and character evolution are underway. By comparing these three genera, we discuss the ecological mechanisms underlying the contrasting diversification modes among them.
... In such cases, multivariate morphometrics can be an effective tool for delimiting taxa (Marhold 2011, Kaplan andMarhold 2012). Several recent studies have successfully discriminated some Arisaema species using multivariate approaches (Kobayashi et al. 2003, Murata and Kakishima 2008, Matsumoto et al. 2019). In the Serizawa (1980a); 12 Serizawa (1980b); 13 Serizawa (1981a); 14 Serizawa (1981b). ...
... Based on previous studies using multivariate morphometrics (Kobayashi et al. 2003, Murata and Kakishima 2008, Matsumoto et al. 2019, I selected 13 morphological traits as follows: a) length of spathe blade (cm); b) width of spathe blade (cm); c) length of spathe tube (cm); d) auricle width of spathe tube (mm); e) diameter of spadix appendix (mm); f ) length of peduncle (cm); g) length of pseudostem (cm); h) length of petiole (cm); i) petiolule length of terminal leaflet (cm); j) length of rachis (cm); k) length of terminal leaflet (cm); l) width of terminal leaflet (cm); m) number of leaflets of first pedate leaf (correspondent to Fig. 3a-m). From 17 May to 6 July 2018, I measured these traits for A. angustatum (9 ♀ plants, 32 ♂ plants), A. maekawae (18 ♀ plants, 38 ♂ plants) and A. pseudoangustatum var. ...
The Arisaema serratum group sensu stricto (Araceae) is a Japanese species assemblage
characterized by a well-developed leaf rachis and a long pseudostem. It has been
regarded as a complex including many species with few decisive traits. They were previously considered as a single species due to their wide intraspecific variation, but
recent monographs have recognized about 13 species. In this group, A. maekawae and
A. pseudoangustatum var. pseudoangustatum were recently divided from A. angustatum
from the type localities. An earlier study found sympatric populations of the three
species in the Chugoku region, far from the type localities, and distinguished them
by qualitative observations, but there are no quantitative data to delimit the species
boundaries objectively. Since high phenotypic plasticity hinders species delimitation
in the A. serratum group s.s. using just a few quantitative traits, I investigated the morphological discontinuity among them using principal component analysis with eleven
quantitative characters. In an ordination space including the three species, the morphological variation was roughly divided into A. maekawae and the others, and largely
overlapped between A. angustatum and A. pseudoangustatum var. pseudoangustatum.
However, the morphological variation clearly distinguished the two latter species when
A. maekawae was excluded from the PCA. To conclude, the three Arisaema species
appeared to be morphologically distinct. The strong reproductive isolation reported
previously also ensures the species distinction according to the biological species concept. The discriminating traits between A. angustatum and the other two species partly
differed from those in the type localities, suggesting that examination of intraspecific
variation is needed between geographically isolated populations. This study showed
that a combination of quantitative traits effectively supports tentative species identifications using qualitative traits and reinforces the usefulness of multivariate morphometrics for future systematic studies.
... First, the quantification of pollinator visits is easy in Arisaema because all visitors can be trapped in the pitcherlike spathe (Fig. 1) by artificial manipulation (Barriault et al., 2009(Barriault et al., , 2010. Second, pollinator isolation caused by selective visitation of fungus gnats (Vogel and Martens, 2000) and contrasting fly pollinators (Kakishima et al., 2019;Matsumoto et al., 2019) has been reported in sympatric Arisaema species. Third, Arisaema section Pistillata rapidly diversified in Japan (Ohi-Toma et al., 2016) and several species often coexist in the field (Murata, 1995;Serizawa, 1997). ...
... (2) phenological isolation caused by interspecific differences in flowering phenology (Matsumoto et al., 2019); and (3) pollinator isolation caused by selective pollination by fungus gnats (Vogel and Martens, 2000). We quantified their strength and contribution to the total reproductive isolation using the unified calculation method described by Sobel and Chen (2014). ...
... To investigate interspecific differences in floral visitor assemblage, we collected all floral visitors in the spathe using a hand-made aspirator simultaneously with the investigation of flowering phenology, and identified the arthropods captured by inflorescences of the five Arisaema species. The exit holes of male spathes were closed with Teflon tape, before they opened, to prevent the escape of attracted insects (Matsumoto et al., 2019). In 2018 we collected 429 arthropods from A. maekawae, 292 from A. angustatum, 383 from A. peninsulae, 630 from A. ovale and 837 from A. pseudoangustatum (Supplementary Data Table S1). ...
Background and Aims
Interspecific difference in pollinators (pollinator isolation) is important for reproductive isolation in flowering plants. Species-specific pollination by fungus gnats has been discovered in several plant taxa, suggesting that they can contribute to reproductive isolation. Nevertheless, their contribution has not been studied in detail, partly because they are too small for field observations during flower visitation. To quantify their flower visitation, we used the genus Arisaema (Araceae) because the pitcher-like spathe of Arisaema can trap all floral visitors.
Methods
We evaluated floral visitor assemblage in an altitudinal gradient including five Arisaema species. We also examined interspecific differences in altitudinal distribution (geographic isolation) and flowering phenology (phenological isolation). To exclude the effect of interspecific differences in altitudinal distribution on floral visitor assemblage, we established 10 experimental plots including the five Arisaema species on high- and low-altitude areas and collected floral visitors. We also collected floral visitors in three additional sites. Finally, we estimated the strength and contribution of these three reproductive barriers using the unified formula for reproductive isolation.
Key Results
Each Arisaema species selectively attracted different fungus gnats in the altitudinal gradient, experimental plots, and additional sites. Altitudinal distribution and flowering phenology differed among the five Arisaema species, whereas the strength of geographic and phenological isolations were distinctly weaker than those in pollinator isolation. Nevertheless, the absolute contribution of pollinator isolation to total reproductive isolation was weaker than geographic and phenological isolations, because pollinator isolation functions after the two early-acting barriers in plant life history.
Conclusions
Our results suggest that selective pollination by fungus gnats potentially contributes to reproductive isolation. Since geographic and phenological isolations can be disrupted by habitat disturbance and interannual climate change, the strong and stable pollinator isolation might compensate for the weakened early-acting barriers as an alternative reproductive isolation among the five Arisaema species.
... Antennae. 19. Antenna, structure (dorsal and ventral views): (0) normal, moniliform to filiform ( Figures 9D, 64B and 68F,G); (1) weakly modified in association with social insects (i.e., flagellomeres weakly flattened and contiguous, concealing each stem pedicels; Figure 9C); (2) strongly modified in association with social insects (i.e., flagellomeres strongly flattened and adjacent ones contiguous, concealing pedicels) ( Figure 26G); (3) weakly clavate or clubbed apically ( Figure 23B); (4) extremely slender, verticillate. ...
Tachyporinae are one of the most phylogenetically problematic subfamilies in the mega-diverse rove beetle family Staphylinidae. Despite its high diversity and abundance in forest micro-environments, with over 1600 species worldwide, several previous studies had refuted the monophyly of this subfamily and its largest tribe, Tachyporini. Based on the results of morphology-based phylogenetic analyses and direct examination of specimens encompassing two extinct and all forty extant genera, a new suprageneric classification of Tachyporinae is proposed here, with the removal of the tribe Mycetoporini into a newly recognized subfamily Mycetoporinae stat. nov. Four tribes with two subtribes are arranged within Tachyporinae sensu nov.: Tachyporini sensu nov. (Tachyporina stat. nov., sensu nov. and Euconosomatina stat. rev., sensu nov.), Vatesini sensu nov., Deropini, and Tachinusini stat. rev., sensu nov. (= Megarthropsini syn. nov.). Urolitus syn. nov. is placed as a junior synonym of Sepedophilus, and Palporus stat. nov. is raised to a distinct genus from a subgenus of Tachyporus sensu. nov., and †Mesotachyporus syn. nov. is treated as synonym of the latter. Mycetoporine Bobitobus stat. rev. is resurrected from synonymy with Lordithon sensu nov., and considered as a valid genus. My revised classification provides a novel framework for taxonomic inventories and ecological studies of these groups.
... It is noteworthy that the F 1 hybrids of artificial crosses of Japanese Arisaema species have high germination rates and high pollen fertility (Murata et al. 2018). Despite such weak postpollination barriers (Murata et al. 2018), multiple Arisaema species often coexist in the field and maintain reproductive integrity (Matsumoto et al. 2019). Therefore, gene flow among Japanese Arisaema species is probably limited by effective prepollination barriers such as interspecific differences in pollinator assemblages (Matsumoto et al. 2019, Kakishima et al., 2020. ...
... Despite such weak postpollination barriers (Murata et al. 2018), multiple Arisaema species often coexist in the field and maintain reproductive integrity (Matsumoto et al. 2019). Therefore, gene flow among Japanese Arisaema species is probably limited by effective prepollination barriers such as interspecific differences in pollinator assemblages (Matsumoto et al. 2019, Kakishima et al., 2020. ...
The genus Arisaema contains approximately 180 species of deciduous or evergreen perennial herbs characterized by spathaceous inflorescence and one to several leaves that emerge from underground stems (Murata et al. 2018). Most Arisaema species are distributed in subtropical to cool temperate regions of Asia, although several are endemic to North America and tropical East Africa (Murata et al. 2018). The genus is particularly diverse in the Japanese Archipelago (Murata et al. 2018), where 53 species have been recorded.
