[Mycology • 2024] Tuber itzcuinzapotl (Ascomycota: Pezizomycetes: Tuberaceae) • the First Edible Truffle reported from Mexico with Traditional Biocultural Importance

Tuber itzcuinzapotl de la Fuente & Rosales-Rosales,  
 

in de la Fuente, Rosales-Rosales, Martínez-González, Martínez-Reyes, Elizondo-Salas et Perez-Moreno, 2024. 

DOI:  10.11646/phytotaxa.635.3.2 

 Researchgate.net/publication/377700444

  facebook.com/TecNMZongolica

Abstract

Tuber itzcuinzapotl is described as a new species to science. This species is characterized by its pale brown ascomata, finely granular peridium, pale brown to gray gleba, and 22–52 × 15–40 µm alveolate ascospores. The new species is putatively associated with Pinus patula in conifer mixed forests in the state of Veracruz, located in eastern Mexico. Phylogenetic analysis based on the nrITS region places the new species in the Maculatum clade, closely related, but morphologically distinctive, to T. miquihuanense and T. mexiusanum. The new species consumed by the Nahua people, traditionally named “Itzcuinzapotl”, constitutes the first edible truffle with biocultural importance in Mexico. Macro- and micromorphological characterization, results of phylogenetic analysis, and photographs are presented. Ethnomycological aspects related to the species are also briefly discussed.

 

Key words: biocultural erosion, Ethnomycology, fungi, hypogeous fungi, Mexican truffles, mycorrhizal fungi, traditional knowledge

Tuber itzcuinzapotl de la Fuente & Rosales-Rosales, sp. nov. (Holotype).
A) Fresh ascomata. B) Dry ascomata. C) Peridium hyphal arrangement. D) Hyphae fromsterile veins. E) 3-spored ascus containing 2 mature ascospores showing alveolate ornamentation.

Bar: A–B= 10 mm; C–E=20 μm.

Tuber itzcuinzapotl de la Fuente & Rosales-Rosales, sp. nov.

Diagnosis:—The new species differs from other species within the Maculatum clade by its pale brown ascomata,finely granular peridium, composed of subglobose hyphae, forming pyramid-like structures, clavate terminal cells of11−30 × 6−10 μm, pale brown to gray gleba, and 22−52 × 15−40 μm alveolate ascospores. 

Etymology:—Refers to the name used by the Nahua people to designate this edibles species itzcuinzapotl (itzcuin = dog, and zapotl = zapote, a native sweet fruit). 

Javier Isaac de la Fuente, Wendy Rosales-Rosales, César Ramiro Martínez-González, Magdalena Martínez-Reyes, Andrea Carolina Elizondo-Salas and Jesus Perez-Moreno. 2024. Tuber itzcuinzapotl sp. nov. (Tuberaceae, Pezizomycetes), the First Edible Truffle reported from Mexico with Traditional Biocultural Importance.  Phytotaxa. 635(3); 206-216. DOI:  10.11646/phytotaxa.635.3.2

  Researchgate.net/publication/377700444_Tuber_itzcuinzapotl_the_first_edible_truffle_reported_from_Mexico

  facebook.com/100089681312095/posts/339977982334910

  facebook.com/TecNMZongolica/posts/679302027702523

[Fungi • 2023] Mallocybe pakistanica & M. pinicola (Agaricales: Inocybaceae) • New Species of Mallocybe from Pakistan, based on Morphological and Molecular Evidence

A Mallocybe pakistanica & B M. pinicola Saba & Khalid,

in Saba, Khalid et Sarwar, 2023. 

DOI: 10.3897/mycokeys.99.86844

 Scale bars: 10 mm 

Abstract

Within the family Inocybaceae, many species of Mallocybe have been reported, but there are only a few reports of this genus from Pakistan. In this study, six collections of Mallocybe were studied by morphological and phylogenetic methods. Phylogenetic analyses, based on sequence data from two different loci (ITS and LSU) using Maximum Likelihood and Maximum Parsimony methods, have been performed to infer species relationships within Mallocybe. Results indicated that these six collections encompass two new species of Mallocybe i.e. M. pakistanica and M. pinicola, from Pakistan. Their detailed morphological descriptions and illustrations are also provided. In addition, comparison with morphologically closely-related taxa is also discussed. Previously, only two species of this genus have been recorded from Pakistan and, with this addition, the total number of reported taxa of Mallocybe has been raised to four from Pakistan. A key to the described taxa of Mallocybe from Pakistan is also provided.

Key words: Asia, molecular systematics, phylogeny, Pinaceae

Holotypes A Mallocybe pakistanica (MSM#0061) B Mallocybe pinicola (MSM#0060).

 Scale bars: 10 mm (A, B).

 Mallocybe pakistanica Saba & Khalid, sp. nov.

 

Diagnosis: Most similar to Mallocybe myriadophylla described from north-western Europe, but differs by the absence of a crowded lamellae, different pileal colouration and somewhat larger basidiospores. Phylogenetically separated from other species of Mallocybe due to unique ITS and LSU sequences.

Etymology: Referring to the country where it was discovered.

 Mallocybe pinicola Saba & Khalid, sp. nov.

 

Diagnosis: Most similar to M. siciliana and M. subtomentosa, but differs by the combination of pileal colour, absence of umbo, size of basidiospores, pyriform to broadly clavate, catenate cheilocystidia and an ecological association with Pines. Phylogenetically separated from other species of Mallocybe due to unique ITS and LSU sequences.

Etymology: Referring to its exclusive association with Pinus.

 Malka Saba, Abdul Nasir Khalid and Samina Sarwar. 2023. New Species of Mallocybe (Agaricales, Inocybaceae) from Pakistan, based on Morphological and Molecular Evidence. MycoKeys. 99: 171-186.  DOI: 10.3897/mycokeys.99.86844

[Fungi • 2023] Microstrobilinia castrans (Ascomycota: Leotiomycetes) • A New Genus and Species of the Sclerotiniaceae parasitizing Pollen Cones of Picea spp.

Microstrobilinia castrans Beenken & Andr. Gross,

in Beenken, Stroheker, Dubach, ... et Gross, 2023.

 DOI: 10.1007/s11557-023-01865-w

Abstract

The fungal pathogens of spruce are well known in Europe and elsewhere. Therefore, it was surprising to discover a new fungal species and genus in Central Europe that attacks the pollen cones of three spruce species. The new ascomycete forms apothecia on stromatized pollen cones of Norway spruce (Picea abies) and Serbian spruce (Picea omorika) in mountain areas and on West Himalayan spruce (Picea smithiana) planted in urban lowland regions of Switzerland, Germany, and Italy. It was also detected in France, based on metabarcode sequences deposited in the GlobalFungi database. Its sudden appearance and the different origins of the host trees in Europe and Asia leave the origin of the fungus unclear. The new fungus might be a neomycete for Europe. A phylogenetic analysis using SSU, LSU, ITS, RPB2, and TEF1 sequences classified the fungus as a member of Sclerotiniaceae (Helotiales, Leotiomycetes). However, it differs morphologically from the other genera of this family in having an ascus without apical apparatus containing four mainly citriform spores with 16 nuclei each. Furthermore, it is the only known cup fungus that parasitizes pollen cones of conifers by stromatizing their tissue and infecting pollen grains. The fungus does not seem to cause major damage to the spruce populations, as only a few pollen cones per tree are affected. All this leads us to describe the newly discovered fungus as the new species and new genus Microstrobilinia castrans, the fungus that castrates pollen cones of spruce.

 

Keywords: GlobalFungi database, Helotiales, Multinucleate ascospores, Neomycete, New genus, New species, Phenology, Pollen parasite

Microstrobilinia castrans:
a–e On Picea smithiana: a P. smithiana tree at type location, two P. omorika trees in the background; b fresh infected pollen cone showing first symptoms like deformation and brown discoloration; c infected pollen cone with apothecia in the first year after infection; d, e several years old pollen cones overgrown with mosses and lichens with fresh apothecia and remnants of last year’s apothecia.
 f–i On Picea abies: f P. abies at a typical habitat in the Swiss Alps; g infested pollen cones on twigs; h pollen cone with small immature apothecia; i pollen cone with mature apothecia. j Pollen cones of P. omorika with dry, mature apothecia

Microstrobilinia castrans, ontogeny of apothecia:
a Black primordium in center of a cone scale in early summer; b immature apothecium in summer; c mature apothecia in autumn with gray velvet outer surface; d dehydrated mature apothecia with black smooth outer surface in next spring; e, f hydrated mature apothecia in next spring; g apothecium in side-view showing the stipe; h dissected pollen cone scales with apothecium.

 a–d from P. abies; e, f, h from P. smithiana; g from P. omorika.

Scale bars: a, b = 0.25 mm; c, d, g = 0.5 mm; e, f, h = 1 mm

Microstrobilinia Beenken & Andr. Gross, gen. nov.

 

Etymology: The genus name refers to the substrate: “microstrobilus” means “small cone” in Greek and is the botanical term for the male pollen cone of conifers.

Diagnosis: Member of Sclerotiniaceae Whetzel (1945) emend. Holst-Jensen et al. (1997) (Helotiales, Leotiomycetes), different from all other members of the Sclerotiniaceae in 4-spored, cylindrical asci with a short stipe and an iodine negative apex without apical apparatus, mature ascospores containing 16 nuclei, parasitizing pollen cones of Picea spp.

Microstrobilinia castrans Beenken & Andr. Gross, spec. nov.

Etymology: The species name refers to the fact that the fungus prevents the male reproduction of its host: “castrans” is Latin for castrating/emasculating.

Short diagnosis: Apothecia brown, short stipitate, desiccation-tolerant, 1–5 mm in diameter, growing on stromatized pollen cones of Picea spp.; ectal excipulum of brown textura globulosa-angularis; asci cylindric, av. 108 × 14.6 µm when alive, iodine negative without visible apical apparatus, 4-spored, ascospores citriform, and almond-shaped, av. 25 × 13 µm when alive, containing 16 nuclei when mature.

 

 

Ludwig Beenken, Sophie Stroheker, Vivanne Dubach, Markus Schlegel, Valentin Queloz and Andrin Gross. 2023. Microstrobilinia castrans, A New Genus and Species of the Sclerotiniaceae parasitizing Pollen Cones of Picea spp. Mycological Progress. 22, 14. DOI: 10.1007/s11557-023-01865-w

[Fungi • 2021] Helvella jocatoi (Ascomycota: Pezizales) • A New Species from H. lacunosa Complex with Cultural Importance in central Mexico Abies religiosa Forests

Helvella jocatoi F. Landeros, R. Garibay-Orijel & L. Guz.-Dáv. 

in Landeros, Ferrusca-Rico, Guzmán-Dávalos, ... et Garibay-Orijel, 2021.

 DOI: 10.11646/phytotaxa.498.1.1

Abstract

Helvella lacunosa is a species complex, with Helvella lacunosa s.s. not currently distributed in America. The objective of this study was to resolve the taxonomy of specimens from central Mexico identified as Helvella lacunosa s.l. associated with Abies religiosa forests. The nuclear ITS and LSU regions were PCR-amplified and sequenced from dry herbaria specimens. Phylogenetic analyses were based on Parsimony, Maximum Likelihood and Bayesian Inference approaches. Sequences of Helvella from A. religiosa forests grouped into a well-supported lineage within the North American clade together with Helvella dryophila (associated with Quercus in western USA) and Helvella vespertina (associated with conifer forests in western USA). Therefore, we describe and illustrate Helvella jocatoi as a new species based on phylogeny and morphological traits. In central Mexico, this taxon is an edible mushroom known as “gachupín”, has high cultural importance and is sold in large quantities. The description of this new species restricted to A. religiosa forests has implications for its conservation since its habitat is endangered.

Keywords: Abies religiosa, Ecology, Helvella lacunosa complex, Phylogeny, edible mushrooms, Fungi

Helvella jocatoi F. Landeros, R. Garibay-Orijel & L. Guz.-Dáv. 
 A) Holotype CB08326 (MEXU 25760); B) Castro-Castro 7 (IBUG); C) Landeros 3581 (IBUG); D) Hymenium; E) Ascospores; F) Ectal excipulum cells.

 A), B), C) Bars: 1 cm. D), E), F) Bars: 10 µm.

Helvella jocatoi F. Landeros, R. Garibay-Orijel & L. Guz.-Dáv. sp. nov. 

Diagnoses: Ascoma 40–160 mm high, pileus 20–60 mm, high 20–60 mm broad, with three lobes or irregularly lobed, dark gray to black. Pileus sterile surface smooth to slightly venous, white or slightly grayish. Stipe up to 120 mm high and up to 25 mm broad, cylindrical, lacunose, first white, then grayish even blackish. Ascospores 16.8–20.8 (–22) × 10.8–13 (–14) µm, hyaline. Solitary or in clusters, terrestrial or on mosses in Abies religiosa forests on the Trans Mexican Volcanic Belt. 

 Etymology: jocato (Spanish), is an acronym for the first two letters of the given name and two last names of José Castillo Tovar deceased at 2012 who was a prominent Mexican mycologist and teacher of the first author; i (Latin), pertaining to, indicates the genitive case in masculine form.

Fidel Landeros, Felipe M. Ferrusca-Rico, Laura Guzmán-Dávalos, Edgardo Ulises Esquivel-Naranjo, Noemí Matías-Ferrer, Cristina Burrola-Aguilar, Gala Artemisa Viurcos-Martínez and Roberto Garibay-Orijel. 2021. Helvella jocatoi sp. nov. (Pezizales, Ascomycota), A New Species from H. lacunosa Complex with Cultural Importance in central Mexico Abies religiosa Forests. Phytotaxa. 498(1); 1–11. DOI: 10.11646/phytotaxa.498.1.1

[Herpetology • 2020] Lycodon cathaya • A New Species of the Genus Lycodon (Serpentes, Colubridae) from Guangxi, China

Lycodon cathaya 

Wang, Qi, Lyu, Zeng & Wang, 2020

Huaping Wolf Snake | 花坪白环蛇 ||  DOI: 10.3897/zookeys.954.53432 

Abstract

A new species of colubrid snake, Lycodon cathaya sp. nov., is described based on two adult male specimens collected from Huaping Nature Reserve, Guangxi, southern China. In a phylogenetic analyses, the new species is shown to be a sister taxon to the clade composed of L. futsingensis and L. namdongensis with low statistical support, and can be distinguished from all known congeners by the significant genetic divergence in the mitochondrial cytochrome b gene fragment (p-distance ≥ 7.9%), and morphologically by the following combination of characters: (1) dorsal scales in 17–17–15 rows, smooth throughout; (2) supralabials eight, third to fifth in contact with eye, infralabials nine; (3) ventral scales 199–200 (plus two preventral scales), subcaudals 78; (4) loreal single, elongated, in contact with eye or not, not in contact with internasals; (5) a single preocular not in contact with frontal, supraocular in contact with prefrontal, two postoculars; (6) maxillary teeth 10 (4+2+2+2); (7) two anterior temporals, three posterior temporals; (8) precloacal plate entire; (9) ground color from head to tail brownish black, with 31–35 dusty rose bands on body trunk, 13–16 on tail; (10) bands in 1–2 vertebral scales broad in minimum width; (11) bands separate ground color into brownish black ellipse patches arranged in a row along the top of body and tail; (12) elliptical patches in 3–6 scales of the vertebral row in maximum width; (13) ventral surface of body with wide brownish black strip, margined with a pair of continuous narrow greyish white ventrolateral lines. With the description of the new species, 64 congeners are currently known in the genus Lycodon, with 16 species occurring in China.

Keywords: Colubrinae, Guangxi, Lycodon cathaya sp. nov., morphology, phylogeny, taxonomy

Figure 3. General aspects in life and close-ups of body scales of
A Lycodon cathaya sp. nov. (SYS r001542, holotype) from Huaping Nature Reserve, Guangxi, China
B L. futsingensis (SYS r002123) from Gaoping Nature Reserve, Shaoguan City, Guangdong, China,
and C L. ruhstrati (SYS r001631) from Huaping Nature Reserve, Guangxi, China.

Figure 4. General aspect of Lycodon cathaya sp. nov. (SYS r001542, holotype) in life when observed. 

     

Figure 5. Comparative characters of head scalation and color patterns (in preservative) of
A Lycodon cathaya sp. nov. (SYS r001542, holotype) B Lycodon cathaya sp. nov. (SYS r001630, paratype)
C L. futsingensis (SYS r002123), and D L. ruhstrati (SYS r001631). Scale bars: 10 mm.

Lycodon cathaya sp. nov.

Diagnosis: Lycodon cathaya sp. nov. can be differentiated from its congeners by the combination of the following morphological characters: (1) dorsal scales in 17–17–15 rows, smooth throughout; (2) supralabials eight, third to fifth in contact with eye, infralabials 9; (3) ventral scales 199–200 (plus two preventral scales), subcaudals 78; (4) loreal single, elongated, in contact with eye or not, not in contact with internasals; (5) a single preocular not in contact with frontal, supraocular in contact with prefrontal, two postoculars; (6) maxillary teeth 10 (4+2+2+2); (7) two anterior temporals, three posterior temporals; (8) precloacal plate entire; (9) ground color from head to tail brownish black, with 31–35 dusty rose bands on body trunk, 13–16 on tail; (10) bands in 1–2 vertebral scales broad in minimum width; (11) bands separate ground color into brownish black ellipse patches, similar arrangement in a row along the top of body and tail; (12) elliptical patches in 3–6 scales of the vertebral row in maximum width; (13) ventral surface of body with a wide brownish black strip, margined with a pair of continuous narrow greyish white ventrolateral lines.

Etymology: The specific name cathaya is a noun referring to the monotypic botanic genus Cathaya Chun & Kuang, 1958. The single species Cathaya argyrophylla Chun & Kuang, 1958 is an endangered relict plant, and was firstly discovered from Huaping Nature Reserve by the investigation team of Sun Yat-sen University. In memory of the predecessors and their contributions on the taxonomy of Chinese flora and fauna, we denominate this new snake species from Huaping Nature Reserve as Lycodon cathaya sp. nov. Its common name is suggested as “Huaping Wolf Snake” in English and “Hua Ping Bai Huan She (花坪白环蛇)” in Chinese.

Jian Wang, Shuo Qi, Zhi-Tong Lyu, Zhao-Chi Zeng and Ying-Yong Wang. 2020. A New Species of the Genus Lycodon (Serpentes, Colubridae) from Guangxi, China. ZooKeys. 954: 85-108. DOI: 10.3897/zookeys.954.53432

    

[Entomology • 2020] Beyond Wallace: A New Lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals A Journey Tracking Its Host-plants, Podocarpus (Pinopsida: Podocarpaceae)

Chrysorthenches muraseae Sohn & Kobayashi, 

in Sohn, Kobayashi & Yoshiyasu, 2020

 DOI: 10.1093/zoolinnean/zlaa009

 twitter.com/ZoolJLinnSoc 

Abstract

A northward trans-Wallacean radiation is demonstrated for Chrysorthenches, a member of the Orthenches group. Here we review Chrysorthenches and allied genera resulting in a generic transfer of Diathryptica callibrya to Chrysorthenches and two new congeners: C. muraseae Sohn & Kobayashi sp. nov. from Japan and C. smaragdina Sohn sp. nov. from Thailand. We review morphological characters of Chrysorthenches and allied genera, and find polyphyly of Diathryptica and the association of the Orthenches-group with Glyphipterigidae. These findings were supported in a maximum likelihood phylogeny of DNA barcodes from ten yponomeutoids. We analysed 30 morphological characters for 12 species of Chrysorthenches, plus one outgroup, via a cladistic approach. The resulting cladogram redefined two pre-existing Chrysorthenches species-groups and identified one novel lineage: the C. callibrya species-group. We review the host associations between Chrysorthenches and Podocarpaceae, based on mapping the working phylogenies. Our review suggests that ancestral Chrysorthenches colonized Podocarpus and later shifted to other podocarp genera. Biogeographical patterns of Chrysorthenches show that they evolved long after the Podocarpaceae radiation. Disjunctive trans-Wallacean distribution of the C. callibrya species-group is possibly related to the tracking of their host-plants and the complicated geological history of the island-arc system connecting Australia and East Asia.

Keywords: Gondwana, Lepidoptera, phylogenetics, plant/insect interaction, taxonomy, Wallace’s Line

Superfamily Yponomeutoidea Stephens, 1829 

Famly Glyphipterigidae Stainton, 1854 

Genus Chrysorthenches Dugale, 1996 

Chrysorthenches Dugdale, 1996: 34. 

Type species: Orthenches porphyritis Meyrick, 1886, by original designation.

Chrysorthenches callibrya (Turner, 1923), comb. nov.

Distribution: Australia (New South Wales, Queensland). 

Host-plants: Possibly Podocarpus lawrencei Hook.f., Podocarpaceae (adult association).

 

resting adults of Chrysorthenches muraseae.
F–H, resting posture of adult (F, H, lateral view; G, dorsal view). I, close-up of adult head, lateral view.

Chrysorthenches muraseae Sohn & Kobayashi, sp. nov.

Etymology: The species epithet is dedicated to Ms Masumi Murase, who provided valuable information, collected specimens of this species and donated them to us. 

Distribution: Japan (Honshu, Shikoku). 

Host-plants: Podocarpus macrophyllus (Thunb.) Sweet., Podocarpaceae.

Chrysorthenches smaragdina Sohn, sp. nov.

Distribution: Thailand. 

Etymology: The epithet is derived from the Greek σμαράγδι, ‘smarágdi’, emerald, referring to the broad green patch on the forewing of this new species. 

Jae-Cheon Sohn, Shigeki Kobayashi and Yutaka Yoshiyasu. 2020. Beyond Wallace: A New Lineage of Chrysorthenches (Lepidoptera: Yponomeutoidea: Glyphipterigidae) reveals A Journey Tracking Its Host-plants, Podocarpus (Pinopsida: Podocarpaceae). Zoological Journal of the Linnean Society. 190(2); 709–736. DOI: 10.1093/zoolinnean/zlaa009

 twitter.com/ZoolJLinnSoc/status/1322221319505793024

[PaleoBotany • 2020] Krassiloviaceae fam. nov. • Reconstructing Krassilovia mongolica supports Recognition of A New and Unusual Group of Mesozoic Conifers

Krassiloviaceae Herrera, Shi, Mays, Ichinnorov, Takahashi, et al., 2020.

Krassilovia mongolica  
Herrera, Shi, Leslie, Knopf, Ichinnorov, et al.,, 2015

a branch of Krassilovia mongolica bearing terminal seed cones and alternately arranged leafy shoots of Podozamites harrisii. Mature and maturing cones are depicted distally showing the ultimate disarticulation of the bract-scale complexes and the dispersal of the winged seeds.

in Herrera, Shi, Mays, Ichinnorov, Takahashi, et al., 2020. 

DOI:  10.1371/journal.pone.0226779

 Illustration: Pollyanna von Knorring. 

Abstract

Previously unrecognized anatomical features of the cone scales of the enigmatic Early Cretaceous conifer Krassilovia mongolica include the presence of transversely oriented paracytic stomata, which is unusual for all other extinct and extant conifers. Identical stomata are present on co-occurring broad, linear, multiveined leaves assigned to Podozamites harrisii, providing evidence that K. mongolica and P. harrisii are the seed cones and leaves of the same extinct plant. Phylogenetic analyses of the relationships of the reconstructed Krassilovia plant place it in an informal clade that we name the Krassilovia Clade, which also includes Swedenborgia cryptomerioides–Podozamites schenkii, and Cycadocarpidium erdmanni–Podozamites schenkii. All three of these plants have linear leaves that are relatively broad compared to most living conifers, and that are also multiveined with transversely oriented paracytic stomata. We propose that these may be general features of the Krassilovia Clade. Paracytic stomata, and other features of this new group, recall features of extant and fossil Gnetales, raising questions about the phylogenetic homogeneity of the conifer clade similar to those raised by phylogenetic analyses of molecular data.

 Fig 1. Seed cones, cone axis, bract-scale complexes, and winged seeds of Krassilovia mongolica and associated leaves of Podozamites harrisii.
(A–C) Articulated seed cones showing tightly imbricate interlocking bract-scale complexes (A: PP55848; B: PP59064; C: PP59065). (D) Isolated cone axis; note conspicuous spirally arranged abscission scars (PP59066). (E) Incomplete leafy shoot showing a cluster of three attached leaves (one represented only by the leaf base); the fourth leaf (left) was attached to the axis when discovered (PP56218). (F) Three detached strap-shaped leaves; note variation in leaf size and shape, and conspicuous parallel venation (PP56226; PP56223; PP56222). (G) Detail of A showing tightly imbricate interlocking bract-scale complexes. (H) Detail of leaf apex showing converging veins (left; PP56228); leaf base showing the absence of a clearly differentiated petiole (right; PP56230). (I) Three isolated bract-scale complexes showing abaxial (top) and adaxial (bottom) surfaces; note three prominent, spiny, distal lobes and two prominent, spiny, proximal lobes (PP59067; PP59068; PP59069). (J) Two isolated seeds showing narrow wings and variation from more or less symmetrical (top; PP59070), to strongly asymmetrical (bottom; PP59071).

Scale bars: E, F = 1 cm; A–C, G = 5 mm; D, H, I = 2 mm; J = 1 mm.

Fig 6. Reconstruction of Krassilovia mongolica.
(A) Complete mature seed cone showing the strongly imbricate spiny bract-scale complexes. (B) Isolated bract-scale complex in adaxial view showing five seed scars (top left), isolated bract-scale complex in adaxial view with five seeds (top middle), isolated bract-scale complex in abaxial view showing the inconspicuous leafy bract (top right); isolated bract-scale complexes in lateral view showing seed scars and leafy bract (bottom). (C) Isolated asymmetrical (top) more or less symmetrical (bottom) winged seeds. (D) Isolated seed cone axes showing prominent abscission scars

. Drawings not to scale. Illustration: Pollyanna von Knorring. 

Fig 7. Reconstruction of a branch of Krassilovia mongolica bearing terminal seed cones and alternately arranged leafy shoots of Podozamites harrisii. Mature and maturing cones are depicted distally showing the ultimate disarticulation of the bract-scale complexes and the dispersal of the winged seeds.

Illustration: Pollyanna von Knorring.

Systematics and nomenclature

Class: Coniferopsida

Order: Voltziales

Family: Krassiloviaceae Herrera et al. fam. nov. 

Type: Krassilovia Herrera, Shi, Leslie, Knopf, Ichinnorov, Takahashi, Crane et Herendeen. Int. J. Plant Sci. 176:793, 2015.

  Krassilovia mongolica Herrera, Shi, Leslie, Knopf, Ichinnorov, Takahashi, Crane et Herendeen. (Figs 1–4 and 6–7).

Familial diagnosis: Leaves distichously arranged on slender deciduous shoots, borne helically on small persistent leaf cushions, but flattened into a single plane by twisting of their bases. Leaves narrowly oblong to strap-shaped, with multiple conspicuous veins. Seed cone with helically arranged, imbricated, and tightly interlocked bract-scale complexes on a slender central axis. Each bract-scale complex consisting of an inconspicuous bract partially fused to the stalk of an ovuliferous scale. Ovuliferous scale with five conspicuous spine-tipped lobes; three distal (always pointing away from the cone base), the other two proximal (always pointing toward the cone base). Bract scale complexes bearing three to five winged seeds. Leaves and bract-scale complexes with thin, delicate cuticles. Outlines of epidermal cells frequently arranged in two pairs, sometimes forming quartets. Stomatal complexes of both organs transversely oriented, paracytic and monocyclic. Outlines of the guard cells not sunken, flanked by the outlines of two lateral subsidiary cells.

Note: The family includes the seed cone genus Krassilovia and the leaf species Podozamites harrisii Shi, Herrera, Herendeen, Leslie, Ichinnorov, Takahashi et Crane.

Conclusions: 

Krassilovia and the Krassilovia Clade suggest the need to reevaluate current models of conifer evolution and reassess the significance of unusual morphological traits in living and fossil conifers. Current concepts of “conifers” as an evolutionary meaningful group may have been unduly influenced by their simple leaves. Furthermore, their other potential unifying feature, the compound ovulate shoot, is not diagnostic and occurs in other groups of living and fossil plants. In our cladistic analyses, the Krassilovia Clade appears to be close to the conifer crown group but it likely evolved from a paraphyletic and diverse assemblage of ancient conifers or conifer-like plants. Morphological differences among the seed cones of Krassilovia Clade, from elongated and lax in the Cycadocarpidium-Podozamites and Swedenborgia-Podozamites plants, to compact and tightly interlocked in the Early Cretaceous Krassilovia-Podozamites plant, highlight the diversity within the group. However, their conifer-like features, combined with their potential similarities to Gnetales, suggest new lines of investigation to further examine the close gnetalean-conifer relationship inferred from DNA data.

The analysis presented here provides only an initial assessment of the potential relationship of Gnetales, given the few alternative phylogenetic positions that were possible for the Dechellyia-Masculostrobus plant with such limited taxonomic sampling of other potentially relevant seed plants. Nevertheless, it is interesting that the Dechellyia-Masculostrobus plant is resolved close to the Krassilovia Clade even when most extant and fossil placeholders for extant families of conifers are excluded (S5 Appendix). Also, recognition of the Krassilovia Clade, which combines conifer-like cones with leaves that have transversely oriented paracytic stomata, highlights similarities to both conifers and Gnetales, as also do features of the Dechellyia-Masculostrobus plant. Ultimately, conifer monophyly may or may not be supported, but a more definitive understanding will require incorporating more fossil material into morphology-based phylogenetic analyses, not only putative conifers, but also other Gnetales, Bennettitales and Erdtmanithecales, as well as corystosperms and similar plants.

Fabiany Herrera, Gongle Shi, Chris Mays, Niiden Ichinnorov, Masamichi Takahashi, Joseph J. Bevitt, Patrick S. Herendeen and Peter R. Crane. 2020. Reconstructing Krassilovia mongolica supports Recognition of A New and Unusual Group of Mesozoic Conifers. PLoS ONE. 15(1): e0226779. DOI:  10.1371/journal.pone.0226779