Behind the veil – exploring the diversity in Phallus indusiatus s.l. (Phallomycetidae, Basidiomycota)

Abstract Studies have demonstrated that many cosmopolitan species actually consist of divergent clades that present high levels of morphological stasis throughout their evolutionary histories. Phallus indusiatus s.l. has been described as a circum-tropical species. However, this distribution may actually reflect the lack of taxonomic resolution due to the small number of diagnostic morphological characters, which leads to the identification of new records as populations of P. indusiatus. Here, we examine the diversity of P. indusiatus-like species in Brazilian Amazonia. We show a clear congruence between detailed morphological data and ITS, nuc-LSU and atp6 based phylogenetic analyses and three new species are described within the Brazilian indusiate clade. These results highlight the importance of more detailed investigation, with the inclusion of molecular information, in Neotropical fungi.


Introduction
Te worldwide distribution o ungal species hypotheses has been questioned by modern molecular analyses.Studies have demonstrated that many cosmopolitan species actually consist o divergent clades that present high levels o morphological stasis throughout their evolutionary histories (Mueller et al. 2001, Bick ord et al. 2007, Geml et al. 2008, Davis et al. 2014).Phallus indusiatus Vent.-also known as the "veiled lady" mushroom -has been described as a circum-tropical species, with records or South and Central America (Dennis 1960, Saénz and Nassar 1982, Leite et al. 2007, Cheype 2010), Mexico (Guzmán et al. 1990), A rica (Dissing and Lange 1962, Dring 1964, Demoulin and Dring 1975, Dring and Rose 1977, Desjardin and Perry 2015), Asia (Dennis 1953, Liu 1984, Hosaka 2010) and Australia (Smith 2005).For some groups o ungi, spore dispersal mechanisms may support the idea o transoceanic dispersal connecting geographically isolated populations (Halling et al. 2008, Hosaka et al. 2008).However, the current distribution o P. indusiatus may actually re ect the lack o taxonomic resolution due to the small number o diagnostic morphological characters, which leads to the identi cation o new records as populations o P. indusiatus.Te insect-dependent mechanism o spore dispersal may also have played an important role in determining the current distribution o P. indusiatus.
As in phalloid ungi in general, ew morphological characters are available to delimit species in Phallus.In addition, most o the widely used diagnostic characters -such as colour and sizes -show high plasticity, another actor that may lead to misidenti cations and mask the real diversity within the genus (Kreisel 1996, Calonge 2005).As a consequence o these taxonomic uncertainties, a great number o synonyms are reported or several species o this clade.Phallus indusiatus is an emblematic example, where at least nineteen synonyms and several distinct orms have been described (Lloyd 1909, Liu 1984, Guzmán et al. 1990, Kreisel 1996, Calonge 2005, Das et al. 2007, Cheype 2010).
Due to lack o resolution when using morphological characters to identi y Phallus species, we believe that several specimens that have been identi ed as P. indusiatus might actually consist o independently evolving entities.In act, some new species with minimal, yet noticeable morphological di erences rom P. indusiatus, have been proposed.For instance, P. serrata H.L. Li, L. Ye, P.E.Mortimer, J.C. Xu & K.D. Hyde, described or China, di ers by the meshes o the indusium with serrate edges (Li et al. 2014); P. echinovolvatus (M.Zang, D.R. Zheng and Z.X.Hu) Kreisel has a whitish volva with mycelioid projections on the sur ace (Zang et al. 1988); and P. avidus Kreisel & Hausknecht, described or the Seychelles, has yellowish pigments on the receptacle and indusium (Kreisel and Hausknecht 2009).Some o these species were described with the support o molecular analyses, which rein orces the importance o this kind o analysis to resolve these taxonomic uncertainties.At least three species resembling P. indusiatus were described or Brazil: Phallus moelleri Lloyd, Dictyophora callichroa Möller and Dictyophora phalloidea Desv.In the original descriptions, they present some inherent characteristics that distinguish them rom P. indusiatus, such as the above-ground development o the volva in D. phalloidea and the orange receptacle and pinkish receptacle apex in D. callichroa (Möller 1895).Lloyd (1909) described P. moelleri based on a Brazilian species and synonymised D. callichroa with it.All three species are now considered synonyms o P. indusiatus by some authors and Index Fungorum (Lloyd 1909, Fischer 1928, Saénz and Nassar 1982, Calonge 2005, Kreisel and Hausknecht 2009).
Phallus indusiatus was described by Étienne Pierre Ventenat in 1798, based on a specimen rom Suriname.In 1809, Desvaux created a new genus, Dictyophora Desv., mainly characterised by the presence o an indusium, a skirt-like structure that expands rom the receptacle towards the ground.Ventenat's species was trans erred to Dictyophora and named D. indusiata (Vent.)Desv.Kreisel (1996) considered that the importance o an indusium or the taxonomy o the genus was overestimated, hence he downgraded Dictyophora to a section o Phallus.More recently, with the introduction o molecular data to the systematics and taxonomy o ungi, studies have shown that the indusium is a recurrent character, which independently emerged several times during the evolution o the group (Hosaka et al. 2006, Cabral et al. 2012, Marincowitz et al. 2015, rierveiler-Pereira et al. 2017).oday, P. indusiatus Vent. is the valid name or Ventenat's species.Phallus indusiatus is widespread in Brazil, with records rom our o the six Brazilian biomes (Magnago et al. 2013), but in ormation concerning its diversity and distribution is still incomplete.
In this study, we examined the diversity o P. indusiatus-like species in Brazilian Amazonia.We show a clear congruence between detailed morphological data and DNA-based phylogenetic analyses and three new species are described within the Brazilian indusiate clade.Tese results highlight the importance o more detailed investigation, with the inclusion o molecular in ormation, in Neotropical ungi.

Morphological data
Specimens o Phallus sp. with white indusium were collected during the rainy seasons o 2013 to 2015 in various areas o the Amazon Rain orest domain (Figure 1).We included in the analyses our additional specimens attributed to P. indusiatus borrowed rom the Herbarium o the Instituto de Botânica (São Paulo) and the Universidade Federal de Rio Grande do Norte-Fungos, which were collected in various areas o the Atlantic Rain orest domain.Other Phallus species were included in the molecular analysis to increase taxon coverage, both rom GenBank and newly sequenced specimens rom the Paleartic-Oriental region (Suppl.material 1: able S1).Species were morphologically described based on resh and dried material.Macroscopic characters were described based on eld notes and photographs, while microscopic details were obtained by mounting slides with ragments rom di erent layers and structures o dried basidiome in 5% potassium hydroxide (KOH) and/or stained with Congo red dye.We ollowed the speci c literature or species identi cation (Lloyd 1909, Kreisel 1996, Calonge 2005, Kreisel and Hausknecht 2009) and colours were described ollowing Küppers (1979).

DNA extraction, amplification and sequencing
DNA extraction ollowed Hosaka (2009).Te nuclear ribosomal I S and nuc-LSU regions, as well as mitochondrial atp6 region, were ampli ed using previously described primers and protocols (Vilgalys and Hester 1990, White et al. 1990, Kretzer and Bruns 1999).DNA ragments were visualised in 1% agarose gel stained with GelRed™ (Biotium) under UV light.Te ragments were puri ed using Ilustra ExoProStar (GE Healthcare) and then sequenced using the Big Dye erminator Cycle Sequencing Kit (Applied Biosystems) with the same primer pairs.A ter sequencing, some I S electropherograms presented double peaks; in order to resolve these, the I S PCR ragments were cloned ollowing Marincowitz et al. (2015).All ribotypes were included in the phylogenetic analyses.

Molecular phylogenetic analyses
We submitted each sequence to a BLAS search to identi y the closest relatives and to check or possible contamination.Te closest sequences resulting rom the BLAS search and sequences with genus names o Phallus or Dictyophora were retrieved rom GenBank and added to the dataset.All sequences were aligned and manually edited with Geneious R6.1 (Biomatters Ltd.).wo analyses were run, one or the I S dataset (I S) and the other with I S, nuc-LSU and atp6 concatenated matrix (CONC).Te I S nal aligned matrix contained 618 positions, while the concatenated matrix contained 1896 positions (571 or I S, 794 or nuc-LSU and 529 or atp6).Tese two matrices were analysed separately.Based on a previous phylogeny ( rierveiler-Pereira et al. 2014), species o the genus Mutinus were chosen as outgroups.Maximum Parsimony (MP) analyses were per ormed with PAUP* (Swo ord 2003) using heuristic searches with the BR branch-swapping algorithm; the initial tree was obtained by stepwise addition o random additional sequences repeated 100 times and 1000 replicates as bootstrap (bs) settings.For Bayesian analysis (BA), the substitution model o evolution was chosen with MrModel est (Nylander 2004).Te analyses were run in MrBayes 3.2.6, as ollows: two parallel runs were executed with our incrementally-heated simultaneous MCMC simulations over 5 million generations, with trees sampled every 1000 generations.Te consensus trees were reconstructed with the remaining trees a ter the burn-in stage, which was de ned based on the average standard deviation o split requency values.Te con dence values were estimated with posterior probabilities (pp).rees were visualised and edited in Fig ree version 1.4.2.All data are available in reeBASE under ID 21524.

Results
A total o 19 recently collected specimens o Phallus spp. with white indusium were studied, 15 o which were collected in Brazilian Amazonia, while our other specimens were collected rom the Brazilian Atlantic Rain orest (SP and UFRN-Fungos herbaria) (Figure 1).Additionally, we obtained sequences rom 21 Phallus specimens rom Japan, Russia, Vietnam and Tailand.Te collection localities, herbarium vouchers and GenBank accession numbers can be ound in the Suppl.material 1: able S1, as well as in species descriptions.

Phylogenetic analyses
We obtained 95 sequences, amongst which 54 were I S, 19 were nuc-LSU and 22 were atp6 (Suppl.material 1: able S1).Te I S nal aligned matrix contained 618 positions, while the concatenated matrix contained 1896 positions (571 or I S, 795 or nuc-LSU and 530 or atp6).Maximum Parsimony and Bayesian analyses with both matrices (I S and CONC) resulted in trees with the same intraspeci c relationships, but with di erent topologies (Figures 2, 3; MP trees in Suppl.material 2: Figures S1,  S2).For Maximum Parsimony analysis, o the 618 positions rom the I S matrix, 382 were in ormative and resulted in a most parsimonious tree with 2006 steps (CI = 0.458, RI = 0.859, RC = 0.394), while o the 1896 positions rom the CONC matrix, 502 were in ormative and resulted in a most parsimonious tree with 1097 steps (CI = 0.547, RI = 0.709, RC = 0.388).In all o the phylogenetic trees obtained in this study, the Brazilian specimens o Phallus grouped together (I S: pp = 1, bs = 94%; CONC: pp = 1, bs = 100%).Tis clade can be divided into six groups, which correspond to the our morphospecies identi ed and described here (coloured clades on Figures 2,  3), Phallus cinnabarinus (W.S. Lee) Kreisel ound in Amazonia (Cabral et al. 2015) and one specimen rom southern Brazil (P.indusiatus ICN 176960), or which we do not have morphological in ormation.Sequences under the name P. indusiatus (and D. indusiata) retrieved rom GenBank, all rom Asia (China and Japan), as well as those collected by us in this study, orm a paraphyletic clade with intercontinental disjunct distributions.Based on morphological similarities and the geographical proximity to the type locality (Suriname) o the Amazonian specimens collected and supported by the molecular data, one Brazilian clade (blue on Figures 2, 3 Diagnosis.Tis species is characterised by the campanulate receptacle slightly constricted at the base, pale yellow, reticulated, with a prominent apical pore, epigeous development o basidiome, volva varying rom white to dark brown and spores up to 4.6 × 2.5 µm.Notes.Phallus avidus Kreisel & Hauskn.could be comparable with P. denigricans by the conical receptacle and the indusium size; however, P. avidus has smaller spores (up to 3.6 × 1.8 µm), the sur ace o the volva is light grey with an orange ush and the indusium is cream to yellow (Kreisel and Hausknecht 2009).Phallus impudicus var.obliteratus (Malençon) Kreisel has a reticulate white receptacle and a rudimentary white indusium; Phallus denigricans also has a poorly-developed indusium, but it is very di erent rom P. impudicus var.obliteratus, where the indusium is hidden under the receptacle (Calonge 2005, Kreisel andHausknecht 2009).Phallus callichrous (Möller) Lloyd is a species described rom Brazil, with white indusium and di ers rom P. denigricans by having an orange to pink receptacle and reddish-violet rhizomorphs.Recently, another indusiate species was described or Brazil, Phallus aureolatus, but it di ers rom P. denigricans mainly by the strongly developed pore and the merulioid sur ace o the receptacle ( rierveiler-Pereira et al. 2017), in addition to its di erent phylogenetic placement (Figures 2, 3).Phallus echinovolvatus (M.Zang, D.R. Zheng & Z.X.Hu) Kreisel is another white-indusiate species, characterised mainly by the volva covered with echinulate hyphae projections; in P. denigricans, hyphae projections on the volva sur ace can also be ound in some specimens, but they are smaller than in P. echinovolvatus (Zang et al. 1988).In P. indusiatus, the receptacle is campanulate, the immature basidiome is hypogeous, so that the volva is buried under the ground when the basidiome is ully developed, the indusium is completely developed reaching the ground and the volva and rhizomorphs have pinkish pigments (Ventenat 1798).On the other hand, in P. denigricans the campanulate receptacle is constricted at the base, the basidiome has a completely epigeous development, the indusium is poorly-developed reaching only 2/3 o the basidiome and the rhizomorphs and volva are white to brownish.
It is not rare to nd Phallus specimens with a blackish volva; recently, a new species was described, P. uscoechinovolvatus (Song et al. 2018), but it is quite di erent rom P. denigricans mainly by the strongly echinulated volva.Phallus merulinus (Berk.)Cooke and P. atrovolvatus Kreisel & Calonge are very similar, di ering by the volva colour -that is black in P. atrovolvatus and white in P. merulinus -and the habitat (Calonge 2005).In our I S phylogenetic analyses (Figure 3), specimens identi ed as P. atrovolvatus and P. merulinus grouped together in the same clade, indicating a possible identity between these two species.However, no type material was analysed here, which prevents a reliable determination o the species boundaries between P. atrovolvatus and P. merulinus.Similarly, in P. denigricans, we ound specimens with white and pale white to brownish volva all grouping in the same clade in phylogenetic trees (Figures 2, 3).Tis suggests that the volva colour might change due to the soil properties or with the maturity o the basidiome.Tere ore, this speci c characteristic -pale or darker volva -should be care ully analysed be ore it can be used as a diagnostic character in Phallus species.
In Diagnosis.Tis species is characterised by its large basidiome (up to 200 mm), the indusium reaching 2/3 o the basidiome, the purplish volva and rhizomorphs and the thimble-like and strongly reticulated receptacle.
Habitat and distribution.on soil, in a ragment o upland secondary orest.It was ound in the municipalities o Manaus (State o Amazonas, Brazil) and Sinop (State o Mato Grosso, Brazil).
Etymology.with re erence to the volva becoming purple.
Notes.Tis species is the most distinctive amongst our collections, mainly due to its large basidiome, the purplish volva and rhizomorphs and the strongly reticulated receptacle.Phallus rubrovolvatus (M.Zang, D.G. Ji & X.X.Liu) Kreisel is one o the largest white-indusiate species (up to 330 mm); it di ers rom P. purpurascens by the deep red volva, the ragile indusium, by larger reticulations on the receptacle and smaller spores (3.7-4 × 2-2.5 µm) (Liu 1984, Calonge 2005).Additionally, in the phylogenetic analysis (Figures 2, 3), P. rubrovolvatus does not group with P. purpurascens, which con rms their separate identities.Phallus callichrous has an orange to pink receptacle, reddish-violet rhizomorphs and orange receptacle with pink margin (Möller 1895, Kreisel and Hausknecht 2009), which di er rom the white receptacle, purplish volva and rhizomorphs o P. purpurascens; un ortunately, there is little in ormation available or this Brazilian species (Calonge 2005).Phallus multicolor (Berk.& Broome) Cooke is similar to P. purpurascens in the purplish volva and rhizomorphs, but it di ers by the cream to orange indusium and the light yellow pseudostipe (Lloyd 1909, Calonge 2005, Kreisel and Hausknecht 2009).Phallus indusiatus di ers rom P. purpurascens by the smaller basidiome, the hypogeous development o the immature basidiome and smaller spores (up to 4.1 × 2.2 µm), the well-developed indusium reaching the ground and the campanulate receptacle with wider reticulations (Ventenat 1798).Te phylogenetic analyses show specimens o P. purpurascens grouping in a clade with high support values (I S tree: pp = 1, bs = 100%; CONC tree: pp = 1, bs = 98%), con rming its distinct identity.
Phallus purpurascens was ound in a ragment o secondary orest, in an extremely threatened area o the Amazonian orest domain in the State o Mato Grosso, Brazil.Tis state was the second most de orested in Brazil in 2018 (INPE 2018), meaning that species in this area may be su ering the consequences o habitat ragmentation, which is one o the main causes o decline in ungal species (Courtecuisse 2008) 6 Diagnosis.Tis species is characterised by its immature basidiome and volva with a squamous sur ace, white receptacle with shallow reticulations and a wide pore.
Habitat and distribution.ound growing on sandy soil, in a ragment o ombrophilous orest in the Atlantic Rain orest domain.
Etymology.with re erence to the volva covered with small scales.

Phallus indusiatus
Habitat and Distribution.ound on sandy soil, in dense old-growth orest.It has a questionable circum-tropical distribution, with records or South and Central America, Mexico, A rica, Asia and Australia, but we believe that the distribution is restricted to South America.
Other specimens examined.BRAZIL.Pará: Belterra, Floresta Nacional do apajós, Jamaraqua Community Notes.According to Ventenat's original description, P. indusiatus is characterised by the hypogeous volva, the campanulate and reticulated receptacle and by the indusium reaching the ground.Te indusium is white, but it can become reddish as it matures.Ventenat does not give in ormation on the colour o the volva and rhizomorphs, but some authors state that the volva can be light pinkish and rhizomorphs can be pinkish to violet (Calonge 2005, Kreisel andHausknecht 2009).Our collection presents the same characteristics o the original description and those in the key or indusiate species presented by Kreisel and Hausknecht (2009a); in addition, some o the specimens are rom the State o Pará, which is geographically close (about 970 km in a straight line) and with the same orest domain as the type locality (Suriname).Tereore, we believe that the specimens that are nested in the same clade in the phylogenetic trees (Figures 2, 3), all collected in the Brazilian Amazonian and Atlantic rain orests, correspond to P. indusiatus sensu stricto.Since Ventenat's original description does not designate a type specimen and, consequentially, it is not possible to nd the original material in herbarium or comparison, we designated here a neotype or Phallus indusiatus, in accordance with the provisions o the International Code o Nomenclature or algae, ungi and plants (ICN) (Article 9.8) ( urland et al. 2017).

Discussion
Molecular and morphological analyses, as well as geographical distributions, support the description o three new species within the Phallus indusiatus-like specimens rom Brazil, with partially overlapping distributions.Our results suggest that a great number o species might be hidden within the circum-tropical P. indusiatus species concept, since the sequence data obtained rom GenBank are clearly polyphyletic with di erent relationships with other Phallus species (Figures 2, 3).In a similar way, several studies have unveiled cryptic ungal diversity hidden within species complexes, especially a ter the integration o phenotypic, single-DNA and next-generation sequencing (NGS) data (Geml et al. 2006, Jargeat et al. 2010, Kasuya et al. 2012, Kõljalg et al. 2013, Sousa et al. 2017, Accioly et al. 2019).For instance, Geml et al. (2008) revealed that at least eight phylogenetic species are ound in the worldwide distribution o Amanita muscaria (L.) Lam, with strong intercontinental genetic disjunctions and intracontinental phylogeographic structure.Sousa et al. (2017) revealed our species within the pepper pot Myriostoma (Phallomycetidae, Basidiomycota), which has always been considered a monotypic worldwide genus.Long-distance dispersal and cosmopolitanism seems not to be the rule in ungal geographical distribution and, or this reason, there are ew species with truly worldwide distributions (Salgado-Salazar et al. 2013).Peay et al. (2016) afrm that climate, environment and dispersal play important roles in shaping ungal communities, where endemism is the most common result in continental and global-scale studies, instead o cosmopolitanism.Tis becomes clear when analysing the P. indusiatus s.l.distribution.As in all gasteroid ungi -basidiomycetes that produce spores inside the ruiting body -this species has a passive mechanism o spore dispersal (statismospory) (Wilson et al. 2011).Phalloid ungi have developed a peculiar spore dispersal mechanism that depends mainly on insects as vectors or dispersal and this actor, together with environmental conditions, should limit P. indusiatus s.l.geographical distributions, generating the species mosaic observed here.
Regarding the Brazilian indusiate clade, we suggest that species within this group are, in act, divergent entities that maintained the general ancestral phenotype (P.indusiatus s.l.) throughout their evolutionary history, due to high levels o morphological stasis.Tis would explain the high requency o taxonomic uncertainties, which generates a great number o synonyms o P. indusiatus.Te maintenance o a conserved morphology due to low rates o phenotypic variation has been widely discussed in evolution (Davis et al. 2014).wo main mechanisms have been proposed to explain the small levels o morphological change through time: genetic and developmental constraints may restrict the appearance o phenotypic variation; or there is strong stabilising selection or a phenotype (Lee and Frost 2002, Geml et al. 2008, Davis et al. 2014).In our hypothesis, because the di erent species in P. indusiatus occupy similar niches and, there ore, they are in similar environmental conditions, they are likely to experience similar selective pressures.A similar pattern was ound by Mueller et al. (2001) in two disjunct and paraphyletic populations o Suillus spraguei (Berk.& M.A. Curtis) Kuntze, that presented no noticeable morphological di erences, probably as a result o stabilising selection.For the uture, this could be tested or other Phallus indusiatus-like species rom other continents and alternative methodologies should be applied, such as ancestral state reconstruction.
When studying phalloid species, it is noticeable that macro-characters are more variable than micro-characters.For instance, spores are o ten cylindrical to bacilloid and smooth throughout the order (except or Gastrosporiaceae), probably as an adaptation or dispersal, since they are dispersed through the gut and do not adhere on the bodies o insects ( uno 1998, Oliveira andMorato 2000).Te presence o rounded crystals in globose cells amongst hyphae o the volva and rhizomorphs was reported or Phallales species (Io sidou and Agerer 2002), but it is not a commonly used character in species descriptions.Probably these crystals consist o calcium oxalate, as ound in other Phallomycetidae species, such as Gastrosporium simplex Mattir.and Geastrum Pers.(Io sidou andAgerer 2002, Zamora et al. 2013), but urther studies about unction and composition in Phallus are needed.Tese crystals are present in most o the species described here, although on di erent parts: only on the volva o P. purpurascens, only on rhizomorphs o P. squamulosus and on both volva and rhizomorphs in P. denigricans.Further studies are needed in order to evaluate the taxonomic value o the presence o crystals in phalloid ungi.For instance, the presence, shape and the arrangement o oxalate crystals were ound to be important characters to delimit species in Geastrum (Zamora et al. 2013).
On the other hand, macro-characters, such as the shape, sur ace and colour o the main structures (receptacle, pseudostipe, indusium, volva and rhizomorphs), are important characters or in rageneric classi cation (Kreisel 1996).In this study, the phylogenetic clades o P. indusiatus-like species were di erentiated, based on these eatures ( able 1), con rming their importance as diagnostic characters.Given that these diagnostic characters are lost once phalloid specimens are dehydrated, it is extremely important that newly described species and new records should be well illustrated with coloured photographs o resh material.In addition, we believe that molecular data are indispensable or delimiting and describing species in Phallus.

Figure 1 .
Figure 1.Currently known distributions o the Phallus species described in this study.Highlighted areas are the Brazilian Biomes (IBGE 2012): Amazonian Rain orest, Cerrado, Caatinga, Atlantic Rain orest, Pantanal and Pampa.

Figure
Figure 2. Phylogenetic tree obtained by Bayesian analysis with I S. Brazilian clades corresponding to the new species and P. indusiatus are indicated (the holotype o each species is in bold).Posterior probabilities and bootstrap values are on the nodes (pp/bs), values o pp < 0.95 and bs < 90 are not shown.Te black dots indicate specimens under Phallus indusiatus deposited in Genbank and downloaded or this study.
Figure 2. Phylogenetic tree obtained by Bayesian analysis with I S. Brazilian clades corresponding to the new species and P. indusiatus are indicated (the holotype o each species is in bold).Posterior probabilities and bootstrap values are on the nodes (pp/bs), values o pp < 0.95 and bs < 90 are not shown.Te black dots indicate specimens under Phallus indusiatus deposited in Genbank and downloaded or this study.

Figure 3 .
Figure 3. Phylogenetic tree obtained by Bayesian analysis with concatenated data (I S, nuc-LSU and atp6).Brazilian clades corresponding to the new species and P. indusiatus are indicated (the holotype o each species is in bold).Posterior probabilities and bootstrap values are on the nodes (pp/bs), values o pp < 0.95 and bs < 90 are not shown (except or P. denigricans clade).

Table 1 .
Morphological di erences between the new Phallus species described here and Phallus indusiatus.Inovação e Comunicação (303851/2015-5) or scholarships.We thank Dr. Doriane Picanço Rodrigues or coordination o the Laboratory o Applied Evolution at the Federal University o Amazonas, where part o the molecular data was obtained.We are grate ul to Ricardo Braga Neto and Dirce Leimi Komura or collecting support.