Morphometric Analysis and Synopsis of Pseudognaphalium (Gnaphalieae, Asteraceae) in North America

Abstract

Pseudognaphalium Kirp. (Asteraceae, Gnaphalieae) consist of about 60 species mainly distributed in South, Central, and North America. As a first contribution toward a comprehensive taxonomic review of Pseudognaphalium, we perform here the first morphometric analysis of North American species, using UPGMA method for the construction of the dendrogram. Based upon these results we present a synopsis including a key to identify species and their associated synonymy. Thirty-seven species are recognized, two taxa are newly synonymized, Pseudognaphalium microcephalum under the name P. canescens and Pseudognaphalium semilanatum under the name P. semiamplexicaule, and two other names are confirmed as synonyms as previously proposed, Pseudognaphalium micradenium as a synonym of P. helleri and Pseudognaphalium crenatum as a synonym of P. viscosum. Lectotypes are newly designated for Gnaphalium beneolens, G. berlandieri (= Pseudognaphalium stramineum), Gnaphalium decurrens (= Pseudognaphalium macounii), G. leucocephalum, G. oxyphyllum, G. oxyphyllum var. semilanatum (= P. semiamplexicaule), G. semiamplexicaule, G. sulphurescens (= P. stramineum), G. thermale, and second-step lectotypifications are proposed for G. helleri and G. wrightii (= P. canescens). In addition, the first illustrations of Pseudognaphalium helleri and P. semiamplexicaule, and a colour figure of P. canescens and P. beneolens emphasizing the results of the morphometric analysis are provided.

Key words
Cluster analysis/UPGMA dendrogram; compositae; morphology; phenetics; taxonomy

INTRODUCTION

Pseudognaphalium Kirp. (Asteraceae, Gnaphalieae) was segregated from Gnaphalium L. by Kirpichnikov & Kuprijanova (1950). Estimates of the number of species ranges between about 90 (Anderberg 1991, Bayer et al. 2007) and 150 (Dillon 2005), and they are mainly distributed in South, Central, and North America, but some species also occur in Asia and Africa. However, as treated in recent revisions of the South American species (Freire et al. 2014a, b, 2018), the number of taxa accepted in Pseudognaphalium has been reduced to ca. 60. The genus is mainly characterized by its herbaceous habitat and the presence of disciform, heterogamous capitula in clusters arranged in corymbs or panicles, the possession of monochromous phyllaries with a divided stereome, truncate style branches with apical sweeping hairs, achenes either glabrous or with short oblong myxogenic duplex hairs, and pappus bristles free at the base. Studies of detailed morphological characters (Hilliard & Burtt 1981) and phylogenetic analysis inferred from morphology (Anderberg 1991) and molecular data (Bayer et al. 2007, Ward et al. 2009, Smissen et al. 2011, Galbany-Casals et al. 2010, 2014) have been interpreted as providing support for recognizing Pseudognaphalium as a distinct genus. However, the most recent molecular phylogenies (e.g. Nie et al. 2016, Acosta Maindo & Galbany-Casals 2018) recover Pseudognaphalium as polyphyletic in two independent clades: a clade containing mainly North American species (Canada, United States of America, and Mexico), and the other clade including mainly South American species. The generitype, Pseudognaphalium oxyphyllum (DC.) Kirp., was included in the North American clade with the other North American species. Both clades were nested within the southern African HAP clade (Helichrysum, Anaphalis, Pseudognaphalium, etc.) showing a southern African origin of Pseudognaphalium (Nie et al. 2016).

Taxonomic treatments and catalogues of North American species of Pseudognaphalium were carried out by Espinosa-García (1985, 2005, sub Gnaphalium), and Villaseñor (2016) for Mexican species, and by Nesom (2006) for North American North Mexican species. These treatments include several taxonomically critical groups of closely related species, as well as polymorphic species with complex intraspecific variation. As an example, the species Gnaphalium beneolens Davison, G. microcephalum Nutt., G. thermale E.E.Nelson, and G. wrightii A.Gray, were accepted by Ferris (1960), Munz & Keck (1959), and Munz (1968, 1974). On the other hand, Cronquist (1950) recognized Gnaphalium thermale as a variety of G. microcephalum while Douglas (1986) considered it as a subspecies. Stebbins & Keil (1992) and Stebbins (1993) merged all these species into a single polymorphic species, Gnaphalium canescens, recognizing G. beneolens, G. microcephalum, and G. thermale as subspecies, and placing G. wrightii into synonymy. Meanwhile, Nesom (2004) considered these four taxa as species under Pseudognaphalium, accepting Gnaphalium wrightii as a synonym of P. canescens. This taxonomic instability affects the identification of specimens of these and other species, and this prompted an investigation based on multivariate analysis to shed light on the delimiting morphologically close, infraspecific taxa or species classification. Cluster analysis and other multivariate techniques have been useful for solving taxonomic problems and circumscribing taxa from morphometric variability data. Several authors have used this type of study to identify entities based on morphology in diverse groups of plants (Owen et al. 2006, Lopez Laphitz et al. 2011, Grossi et al. 2011, Robbiati et al. 2014, Fernández et al. 2017).

The purpose of this study is to examine the current circumscription of the North American Pseudognaphalium taxa to define which species should be recognized based on the variation of morphological characters, including diagnostic characters, using cluster analysis for delimitation of species.

MATERIALS AND METHODS

Sample collection

A total of 297 specimens (including type material), representing 41 of the 43 North American species of Pseudognaphalium recognized in previous studies (Nesom 2006, Villaseñor 2016, Villarreal-Quintanilla et al. 2020), were chosen for the morphometric analysis (Appendix 1). Only two species, Pseudognaphalium altamiranum and Gnaphalium oaxacanum could not be included in the analysis since no material was available from these species. Herbarium material was examined from G, GH, M, MEXU, MO, NY, S, and UC (acronyms following Thiers 2020). Type images at high resolution from BM, C, G-DC, GH, L, LINN, MEXU, MICH, NDG, OS, P, RM, TEX, UC, US, and WIS were also examined (http://plants.jstor.org/).

Morphological data and cluster analysis

We analyzed a total of 36 characters: 13 vegetative and 23 reproductive. A list of the characters and their states can be seen in the Table I. All characters used to separate the North American species of Pseudognaphalium by former authors were included in our analysis. The measurements were performed on mature specimens. All specimens were studied by direct observation and by using a WILD Heerbrugg M5-26799 stereoscope; measurements were taken using a calibrated ocular micrometer. The leaf-blade observations were limited to the mid-section of flowering branches. Three florets per capitulum at same stage of anthesis from 2 or 3 capitula in each specimen were dissected. When the availability of specimens made it possible, replicates of three measurements for each character were obtained and the average was used in the ensuing statistical analyses.

Table I
Morphological characters and character states evaluated in the study.

Vegetative characters:
1. plant height (cm); 2. stem (1 = 0, 2 or 3 = 1, 4 or more = 2); 3. winged stem (absent = 0, present = 1); 4. filiform stems (absent = 0, present = 1); 5. stem pubescence (woolly = 0, lanuginose = 1, glandular = 2); 6. leaf length (cm); 7. leaf width (cm); 8. leaf apex (acute = 0, acuminate = 1, obtuse to rounded = 2); 9. leaf base (attenuate = 0, rounded = 1, decurrent = 2, clasping = 3); 10. leaf margins (flat = 0, undulate = 1); 11. leaf adaxial surface (lanuginose = 0, wolly = 1, glandular = 2, arachnoid = 3); 12. leaf abaxial surface (lanuginose = 0, wolly = 1, glandular = 2); 13. basal rosette of leaves at flowering (whitered = 0, fresh/persistent = 1).
Reproductive characters:
14. capitula arrangement (lax corymbs = 0, dense corymbs = 1, paniculiform corymbs = 2); 15. involucre height (mm); 16. involucre diameter (mm); 17. number of series of phyllaries (3 or 4 = 0, 5 or 6 = 1, 7 or more = 2); 18. phyllary number (#); 19. outer phyllary length (mm); 20. outer phyllary width (mm); 21. outer phyllary apex (acute = 0, obtuse = 1); 22. inner phyllary length (mm); 23. inner phyllary width (mm); 24. inner phyllary apex (acute or acuminate = 0, obtuse to rounded = 1); 25. inner phyllary color (white or creamy = 0, brownish = 1, yellowish or stramineous = 2, pinkish or purple = 3); 26. inner phyllary transparency (shiny = 0, opaque = 1); 27. marginal florets number (#); 28. marginal floret corolla length (mm); 29. disc florets number (#); 30. disc corolla length (mm); 31. disc corolla lobes (yellowish = 0, dark = 1); 32. disc corolla lobes pubescence (smooth = 0, papillose = 1); 33. achenes length (mm); 34. achenes diameter (mm); 35. achenes epidermis (smooth = 0, papillose = 1); 36. pappus length (mm).

We carried out a cluster analysis on all the OTUs. The similarity between two OTUs was calculated on the basis of Gower’s general similarity coefficient (Gower 1971) and the UPGMA method was used for the construction of the dendrogram (Sneath & Sokal 1973). The multivariate analyses with Gower’s (1971) coefficient are suitable for the analysis of mixed characters, e.g., qualitative (binary and ordinal) and quantitative characters to generate a distance/dissimilarity matrix (Mapaya & Cron 2016). Likewise, UPGMA is an accurate and spread method to deduce similarity/dissimilarity among OTUs (Radford 1986, Ward 1993). The qualitative characters were coded accordingly as binary (presence/absence) or a code was assigned for each character state (see Table I for the list of characters and coding). Cluster analysis was performed on all the OTUs using all 36 characters, to obtain information about general relationships and similarities between them. The analyses were carried out using the software’s PAST (Hammer & Harper 2006).

Concepts of species delimitation

The treatment here reported is based on the conservative and widely accepted morphological species concept. According to this concept, continuous variation of characters is allowed within the species, while discontinuous variation in more than one character define distinct species (Davies & Heywood 1967).

RESULTS AND DISCUSSION

The cluster analysis (UPGMA) (Figure 1a) showed that most of the specimens of each species grouped together in separated clusters. A few specimens were intermingled in four main groups (Figure 1b), each with little metric distance suggesting high similarity between the species of each group.

Figure 1a
UPGMA cluster analysis of the whole set of 297 specimens of the North American species of Pseudognaphalium , using a matrix calculated from 36 characters (Gower’s coefficient of similarity). I: P. canescens and P. microcephalum; II: P. semilanatum and P. semiamplexicaule; III: P. helleri and P. micradenium; IV: P. crenatum and P. viscosum.

Figure 1b
Groups I-IV of the UPGMA cluster analysis of the North American species of Pseudognaphalium (continued from Fig. 1a). I: P. canescens and P. microcephalum; II: P. semilanatum and P. semiamplexicaule; III: P. helleri and P. micradenium; IV: P. crenatum and P. viscosum.

The first group (I) includes Pseudognaphalium canescens (Figure 2a, b) and P. microcephalum. These species are morphologically similar with weakly discolorous leaves, which are oblanceolate, not decurrent and eglandular, and corymbiform capitulescences. Both are confined to United States of America and Mexico and are sympatric in part of their distributions, i.e. they are found together in Chihuahua (Mexico) and California (USA). Stebbins & Keil (1992), suggested that Pseudognaphalium beneolens (= G. beneolens), P. microcephalum (= G. microcephalum), and P. thermale (= G. thermale) could be treated as subspecies within a single polymorphic species, P. canescens (= G. canescens). Nesom (2004) considered these four taxa as separate species mainly by leaf base, shape of leaves, nature of tomentum, apex of phyllaries, capitulescence, number of florets, and size of capitula. Our results recovered P. beneolens and P. thermale in separate clusters. Pseudognaphalium beneolens (Figure 2c, d) is characterized by its linear to linear-oblanceolate leaves and paniculate capitulescences, and P. thermale characterized by its concolorous oblanceolate leaves, decurrent leaf base, and corymbiform capitulescences. However, the specimens of P. microcephalum and P. canescens appear intermingled. Consequently, based on this study, we consider Pseudognaphalium microcephalum as a synonym of P. canescens.

Figure 2
Pseudognaphaliumcanescens.a. Habit; b. Detail of the inflorescence. c, d. Pseudognaphalium beneolens. c. Habit; d. Detail of the habit. (Photographs: a, b. John Avise, Asteraceae of Orange Co., California; C. Neal Kramer, CalPhotos-University of California, Berkeley; d. Michael Mitchell, montereywildflowers.com).

The second group (II) includes Pseudognaphalium semiamplexicaule (Figure 4a-g) from Mexico and Mesoamerica and P. semilanatum, endemic to Mexico. These two species have discolorous lanceolate leaves, clasping leaf bases, white shiny phyllaries and corymbiform to paniculate capitulescences. Villaseñor (2016) accepted both names as valid species. Espinosa-García (2005) considered Pseudognaphalium semiamplexicaule as a valid species and P. semilanatum as a dubious species. According to Pruski (2018) the identity of Pseudognaphalium semilanatum would be based on its adaxially stipitate-glandular leaves (vs. arachnoid-tomentose in P. semiamplexicaule). The present analysis shows specimens with glandular and eglandular leaves of Pseudognaphalium semilanatum and P. semiamplexicaule intermingled. Consequently, based on this study, we consider Pseudognaphalium semilanatum as a synonym of P. semiamplexicaule.

Figure 4
Pseudognaphalium semiamplexicaule.a, b. Flowering branches; c. Capitulum; d. Phyllaries; e. Pistillate floret; f. Bisexual floret; g. Achene [a-g. Rzedowski 36561, MEXU].

The third group (III) includes Pseudognaphalium hellerii (Figure 3 a-h) and P. micradenium from East of United States of America, characterized by narrow oblong-lanceolate leaves. Pseudognaphalium hellerii (= Gnaphalium helleri) was mainly described based on its densely glandular-pubescent stems and obtuse phyllaries. According to Weatherby (1923), the identity of Pseudognaphalium micradenium (= Gnaphalium obtusifolium var. micradenium) was based on its glandular-puberulent stems (vs. glandular-villous in P. helleri), linear or linear-lanceolate leaves (vs. leaves oblong-lanceolate in P. helleri), and acute phyllaries (vs. obtuse phyllaries in P. helleri). Mahler (1975), considered Pseudognaphalium micradenium (= Gnaphalium obtusifolium var. micradenium) as a variety of P. hellerii (= G. helleri; i.e. G. helleri var. micradenium). Cronquist (1980) distinguished Pseudognaphalium obtusifolium (= Gnaphalium obtusifolium) from P. helleri (= G. helleri, including G. obtusifolium var. micradenium as its synonym) by its wooly pubescence (vs. principally glandular in P. helleri). Nesom (2006) considered Pseudognaphalium obtusifolium, P. helleri, and P. micradenium as separate species, using leaf shape, pubescence, and the number of florets as a way to distinguish them. The present cluster analysis shows the specimens of Pseudognaphalium obtusifolium in a separate cluster, but those of P. helleri and P. micradenium appear intermingled. Consequently, based on this study, we consider Pseudognaphalium micradenium as a synonym of P. helleri as was previously suggested by Cronquist (1980).

Figure 3
Pseudognaphalium helleri.a. Plant; b. Capitulum; c, Inner phyllary; d-e, Foliar and stem trichomes; f. Pistillate floret; g. Bisexual floret; h. Bristle pappus [a-c, f-h. Fernald & Bayard Long 7696, PH; d. Bush 1000, MO; e. Demaree 16387, GH].

The fourth group (IV) includes Pseudognaphalium crenatum from Mexico and P. viscosum, from United States of America (Texas), Mexico, and Mesoamerica, characterized by its stipitate-glandular stems, linear-lanceolate leaves, white shiny phyllaries, and numerous pistillate florets. Villaseñor (2016) accepted Pseudognaphalium crenatum as a valid species, but McVaugh (1984) and Pruski (2018) placed P. crenatum into synonymy of P. viscosum. The present cluster analysis shows the type specimen of P. crenatum grouped together with the specimens of P. viscosum and confirms previously proposed synonymy.

Therefore, in agreement with our results Pseudognaphalium consists of 37 North American species (Figures 1a, b), and the corresponding synonyms for the three newly circumscribed species are indicated in the taxonomic treatment provided below.

Taxonomic Treatment of Pseudognaphalium in North America

Pseudognaphalium Kirp. (1950: 33). Type: Gnaphalium oxyphyllum DC. ≡ Pseudognaphalium oxyphyllum (DC.) Kirp.

Hypelichrysum Kirp. (1950: 33). Type: Gnaphalium heterotrichum Phil. Hypelichrysum heterotrichum (Phil.) Kirp. ≡ Pseudognaphalium heterotrichum (Phil.) Anderb.

Gnaphalium L. sect. Calolepis Kirp. (1960: 309). Type: Gnaphalium luteoalbum L. Pseudognaphalium luteo-album (L.) Hilliard & Burtt. Pseudognaphalium subgen. Laphangium Hilliard & Burtt (1981): 205 ≡ Laphangium (Hilliard & Burtt 1981) Tzvelev (1993[1994]: 105) ‘ Gnaphalium Kirp. subgen. Laphangium (Hilliard & Burtt 1981), (Sell 2006): 555. Type: Gnaphalium luteo-album L. ‘ Pseudognaphalium luteo-album (L.) Hilliard & Burtt 1981.

Annual, biennial or perennial herbs, woolly or glandular woolly. Leaves alternate, entire, sessile, often stem-clasping and decurrent. Capitula small, heterogamous, disciform, sessile or short pedunculate, in small clusters arranged in corymbs or panicles; involucre often campanulate; phyllaries in 3–many-series, papery, phyllary lamina monochromous opaque or hyaline and shiny, white, yellowish, brownish, straw, pinkish or purple coloured, stereome divided; receptacle smooth or honeycombed. Florets many, pistillate florets usually outnumbering bisexual; pistillate florets corollas filiform or narrowly tubular; bisexual florets corollas tubular, scarcely broaded above, 5-lobed, all corollas yellowish or whitish, lobes glandular and often yellowish or whitish; anthers with a small obtuse apical anther appendage; tails slightly longer or shorter than the filament collar; style branches truncate and penicillate. Achenes glabrous smooth or with imbricate microscopic papillae, rarely setuliferous with short duplex myxogenic hairs; carpopodium symmetrical, continuous; pappus monomorphic, of free capillary, barbellate bristles, apical cells sometimes inflated, bases cohering by patent cilia.

Worldwide distribution, mostly American, some African and Asian species. About 60 species of which 37 grow in North America.

Key to Species of Pseudognaphalium in North America

1. Leaves ≤ 1.5 cm long ……………..…. P. brachyphyllum

1’. Leaves ≥ 2 cm long …………...................................……..2

2(1’). Stem leaves strongly decurrent on stem (wings extending down beyond middle of internode)………………........................................................…3

2’. Stem leaves not to moderately decurrent on stem (wings never extending down beyond middle of internode) ……...........................…………….…..4

3(2). Leaves linear-oblong, adaxial surface arachnoid …………...........................………….. P. bourgovii

3’. Leaves lanceolate, adaxial surface stipitate-glandular ……….................................….…….….P. nataliae

4(2’). Leaves pseudopetiolate (long-attenuate at the base)………............................…P. ehrenbergianum

4’. Leaves sessile (shortly or not attenuate at base)……..…......................................................................….………5

5(4’). Adaxial surface of leaf blades glandulose, arachnoid or lanuginose, abaxial surface white-lanose …………………………......................................................6

5’. Adaxial and abaxial surface of leaf blades glandulose, lanuginose, glandular-lanuginose or lanose …………………….............................…………………..23

6(5). Adaxial surface of leaf blades conspicuously stipitate-glandular ….......................................……………..7

6’. Adaxial surface of leaf blades not stipitate-glandular .………..….……………........................................…..16

7(6). Bases of leaf blades not clasping ………………8

7’. Bases of leaf blades clasping ……..............……….13

8(7). Leaves narrowly linear to linear-lanceolate, margins strongly revolute ….................................………9

8’. Leaves lanceolate, oblong-oblanceolate or elliptic, margins slightly revolute ….....................….11

9(8). Phyllaries acute …………………P. austrotexanum

9’. Phyllaries obtuse to rounded ....……………………..10

10(9’). Phyllaries shiny; pistillate florets 200–400; achenes papillose ..............................…..… P. viscosum

10’. Phyllaries opaque; pistillate florets ≤ ca. 125; achenes smooth ........................... P. leucocephalum

11(8’). Stems lanose; leaves lanceolate to elliptic ……………….…...............................................…….… P. elegans

11’. Stems glandular; leaves lanceolate to oblong-oblanceolate ………….............................................…………12

12(11’). Stem leaves moderately decurrent...............................................……... P. macounii

12’. Stem leaves not decurrent ……………… P. helleri

13(7’). Phyllaries brownish …………………P. monticola

13’. Phyllaries white ……………………..........................……14

14(13’). Involucre narrowly campanulate, 2.5–3 mm diam; capitula ≤ 40-flowered …. ………………….……...............................................................………..… P. pringlei

14’. Involucre obconic, 3-6 mm diam; capitula ≥ 70-flowered ..……………............................................………..15

15(14’). Stems lanose; involucre 4–5(–5.5) mm high ……………….…........................................……. P. biolettii

15’. Stems glandular-lanuginose; involucre (5–)5.5–6 mm high …...........................… P. brachypterum

16(6’). Leaves narrowly linear 1–2.5 mm wide, 30–50 times as long as wide …..….… P. greenmanii

16’. Leaves linear, linear-oblong, linear-oblanceolate or lanceolate > 2.5 (rarely 2) mm wide, up 23 times as long as wide …………………….……17

17(16’). Phyllaries purple .….…………P. purpurascens

17 ‘. Phyllaries white, yellowish, purple, pinkish, brownish or straw coloured ….............................…….18

18(17’). Bases of leaf blades not clasping, not decurrent …...............................................................……….19

18’. Bases of leaf blades clasping or moderately decurrent ………......................................................…………20

19(18). Leaves lanceolate or linear-lanceolate, acute-acuminate …......................……...P. attenuatum

19’. Leaves linear-oblanceolate, acute or subobtuse …….……………....................... P. obtusifolium

20 (18’). Phyllaries white, occasionally pinkish…………………...................................................……… 21

20’. Phyllaries brownish …………….........…………....……22

21(20). Leaves lanceolate, apex acuminate …………..……….............................................… P. semiamplexicaule

21’. Leaves lanceolate-oblong to linear-oblong, apex acute ......................................................... P. roseum

22(20’). Capitula arranged in corymbs; leaf blade margins flat ….................................…..….. P. arizonicum

22’. Capitula arranged in panicles; leaf blade margins undulate …......................……….P. inornatum

23(5’). Annual herbs .………..............………………………….24

23’. Perennial herbs ..…………......................……………….25

24(23). Achenes setuliferous; leaves linear-oblong; stems 15–70 cm high ………… P. luteoalbum

24’. Achenes glabrous; leaves oblong-obovate; stems 5–30 cm high ….….........................… P. saxicola

25(23’). Capitula arranged in elongate to broadly columnar panicles …......................P. ramosissimum

25’. Capitula arranged in corymbs or paniculiform corymbs ………...............................................……..…………..26

26(25’). Phyllaries brownish …............……….…………..27

26’. Phyllaries white, whitish, pinkish, yellowish or straw coloured ……….....................................………….28

27(26). Leaves apices acute; basal rosette of leaves usually persisting at flowering …………..….……………………………………………..........…………… P. liebmannii

27’. Leaves apices attenuate; basal rosette of leaves withered at flowering …………… …………..…………………………………………….……..…….......…… P. conoideum

28(26’). Stem leaves obovate ……………………..………29

28’. Stem leaves predominantly lanceolate or linear …….……...........................................................………….31

29(28). Leaves oblong-obovate, 5–10(–20) mm wide, margins undulate .......................... P. nubicola

29’. Leaves linear-obovate, 4–10 mm wide, margins flat…………….................................................….….30

30(29’). Leaves concolorous, glandular-pubescent under wool ……............…....… P. thermale

30’. Leaves weakly discolorous, eglandular or with few short glandular trichomes hidden under wool …………………...................…...….P. canescens

31(28’). Stem leaves predominantly lanceolate ………………….…………......................................................……….32

31’. Stem leaves predominantly linear………….………34

32(31). Phyllaries opaque, white …….……………………33

32’. Phyllaries shiny, whitish, yellow-greenish or purple ………….......................................….…P. oxyphyllum

33(32). Leaves lanceolate, glandular-stipitate on both surfaces; pistillate florets 7-15 (24) times as many as bisexual florets …………...... P. californicum

33’. Leaves oblong-lanceolate, glandular-lanuginose on abaxial surface; pistillate florets 1–1.5 times as many as bisexual florets ………………….….……......................................................… P. chartaceum

34(31’). Phyllaries yellowish …………… P. stramineum

34’. Phyllaries white or whitish …………………..…………35

35 (34’). Leaves glandular on both surfaces, margins undulate …….......................… P. hintoniorum

35’. Leaves lanuginose on both surfaces or with short glandular trichomes hidden under the woolly trichomes on adaxial surface, margins flat ……………………….......................................................…….36

36(35’). Leaf apices acute, capitula 45–ca.100-flowered .………...................................…..….. P. beneolens

36’. Leaf apices attenuate; capitula ca. 140–270-flowered .………................……..… P. jaliscense

1. Pseudognaphalium arizonicum (A.Gray) Anderb. (Anderberg 1991: 147). Gnaphalium arizonicum A. Gray (1883): 1884: 3. Type: UNITED STATES OF AMERICA. Arizona, Huachuca Mts., “near Fort Huachuca along exsiccated beds of streams” [Cochise Co.], J. G. Lemmon 2757 (holotype: GH 00008290!; isotype: K 000978290! by Lemmon in 1882).

Distribution: Mexico and United States of America.

2. Pseudognaphalium attenuatum (DC.) Anderb. (Anderberg 1991: 147).

2a. Pseudognaphalium attenuatum (DC.) Anderb. var. attenuatum. Gnaphalium attenuatum DC. (Candolle 1838: 228). Type: MEXICO. Tamaulipas, Tampico de Tamaulipas, 1827, J. L. Berlandier 70 (holotype: G-DC G-00312671!; isotypes: G 00301179!, P 00704492!).

Distribution: Mexico and Mesoamerica [Belice, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama (Pruski 2018)].

2b. Pseudognaphalium attenuatum (DC.) Anderb. var. sylvicola (McVaugh) Hinojosa-Espinosa & Villaseñor (2014): 490. Gnaphalium attenuatum DC. var. sylvicola McVaugh (1972: 465). Type: MEXICO. Jalisco: Sierra de Cuale SW of Talpa de Allende, 1800-2250 m, 19-21 Nov 1952, R. McVaugh 14283 (holotype: MICH 1107414!; isotypes: BM 000574758!, BRIT 23669!, DUKE 10000803!, MEXU 00149481!; probable isotype: NY 00169487! by R. McVaugh and J. Sooby, Jr. 14283).

Distribution: Mexico.

3. Pseudognaphalium austrotexanum Nesom (2001a): 507. Type: UNITED STATES OF AMERICA. Texas. Jim Wells Co. 10.1 mi S of Alice, railroad right of way Santa Gertrudis Division of King Ranch, 24 Nov 1954, M. C. Johnston 542108 (holotype: TEX 00008978!; isotype: BRIT 23963!).

Distribution: Mexico and United States of America.

4. Pseudognaphalium beneolens (Davidson) Anderb. (Anderberg 1991: 147). Gnaphalium beneolens Davidson (1918: 17). Gnaphalium canescens DC. subsp. beneolens (Davidson) Stebbins & Keil (1992): 437. Pseudognaphalium canescens (DC.) Anderb. subsp. beneolens (Davidson) Kartesz (1999): no. 25. Type: UNITED STATES OF AMERICA. [California] Los Angeles Co., Crescenta, Sep 1917, F. E. Burlew 3275 (lectotype, designated here: GH 00008291!; isolectotypes: RSA 0001209!, RSA 0001210!). Figure 2c, d.

Observation: Nesom (2004): 782 indicated the type of Gnaphalium beneolens Davidson as ‘Type: U.S.A. California. [Los Angeles Co.]: Crescenta, 1 Sep 1917, E.E. Burlew 3275 (Isotype: GH!)ʼ. This citation of an ‘isotypeʼ (after 1 January 2001) cannot be corrected to a designation of a lectotype (Art. 9.10 of the ICN, Shenzhen Code; Turland et al. 2018), because they did not use the phrase ‘designated hereʼ or an equivalent (ICN, Art. 7.11, Shenzhen Code; Turland et al. 2018) in their associated statement. Consequently, Nesom’s typification for the name G. beneolens was not effective (ICN, Art. 9.23, Shenzhen Code; Turland et al. 2018). The specimen GH 00008291 which presents the most complete plant is designated here as the lectotype of G. beneolens.

Distribution: Mexico and United States of America.

5. Pseudognaphalium biolettii Anderb. ex G.L.Nesom (Nesom 2002): 149, replacement name for Gnaphalium bicolor Bioletti (1893: 16), hom. illeg., non G. bicolor (Lindl.) Sch.Bip. 1845 [= Xerochrysum bicolor (Lindl.) R.J.Bayer], nec Franch. 1896 [=Anaphalis bicolor (Franch.) Diels]. Pseudognaphalium biolettii Anderb. (Anderberg 1991: 147), nom. inval. Type: UNITED STATES OF AMERICA. California, San Diego, 25 Mar 1891, G. W. Dunn s.n. (holotype: UC 31964!).

Distribution: Mexico and United States of America.

6. Pseudognaphalium bourgovii (A.Gray) Anderb. (Anderberg 1991: 147). Gnaphalium bourgovii A. Gray (1883): 1884: 3. Type: MEXICO. Veracruz: Valley of Cordova, 7 Feb 1866, E. Bourgeau 1852 (holotype: GH 00008317!; isotypes: K 000500363!, L 0001972!, MSC 0091946!, P 00704482!, P 00704483!, P 00704484!).

Distribution: Mexico.

7. Pseudognaphalium brachyphyllum (Greenm.) Anderb. (Anderberg 1991: 147). Gnaphalium brachyphyllum Greenm. (Greenman 1907: 267). Type: GUATEMALA. Quetzaltenango: Cerro Quemada, 9000 ft, 8 Feb 1906, W. A. Kellerman 5301 (holotype: F, not seen; isotypes: OS 0000179!, US 00129533!).

Distribution: Mexico and Mesoamerica [Guatemala].

8. Pseudognaphalium brachypterum(DC.) Anderb. (Anderberg 1991: 147). Gnaphalium brachypterum DC. (Candolle 1838: 226). Type: MEXICO. Circà Victoriam, Nov. 1830, J. L. Berlandier 2189 (holotype: G-DC G-00469610!; isotypes: GH 00008318!, P 00704485!, PH 00012282!).

Distribution: Mexico and Mesoamerica [El Salvador, Guatemala, Honduras, Nicaragua (Pruski 2018)].

9. Pseudognaphalium californicum (DC.) Anderb. (Anderberg 1991: 147). Gnaphalium californicum DC. (Candolle 1838: 224). Type: UNITED STATED OF AMERICA. California: year 1833, D. Douglas 57 ‘... v. s. comm. ab hon. soc. hortic. Londin.’ (holotype: G-DC G-00469618!; isotype: BM 000839374!, probable isotypes: [D. Douglas s.n.] GH 00008292!, K 000978292!).

Distribution: Mexico and United States of America.

10. Pseudognaphalium canescens (DC.) Anderb. (Anderberg 1991: 147). Gnaphalium canescens DC. (Candolle 1838: 228). Type: MEXICO. Guanajuato, León, year 1829, J. Mendez s.n. (holotype: G-DC G-00312669!; isotype: P 00704486!). Figure 2a, b.

Gnaphalium microcephalum Nutt. (Nuttall 1841): 404. 1841. Gnaphalium canescens DC. subsp. microcephalum (Nutt.) Stebbins & Keil (1992): 437. Pseudognaphalium canescens (DC.) Anderb. subsp. microcephalum (Nutt.) Kartesz (1999): no. 26. Pseudognaphalium microcephalum (Nutt.) Anderb. (Anderberg 1991: 147). Type: UNITED STATES OF AMERICA. California, San Diego, T. Nuttall s.n. (holotype: BM 001010942!), syn. nov. Gnaphalium wrightii

A. Gray (1882): 214. Type: UNITED STATES OF AMERICA. Texas, [El Paso or Hudspeth Co.] between El Paso and Guadalupe Mts., Oct 1849, C. Wright 394 (lectotype [first step], designated by Nesom (2004): 783: GH; lectotype [second step], designated here: GH 00008314!; isolectotypes: GH 00008313!, US 00129567!).

Gnaphalium albatum Osterh. (Osterhout 1906): 141. Type: UNITED STATES OF AMERICA. Colorado. Larimer Co., canyon of Thompson River, [between the foothills and Estes Park], 16 Aug 1905, G. E. Osterhout 3158 (holotype: RM 0001102!; isotype: NY 00169466!).

Gnaphalium sonorae I.M.Johnst. (Johnston 1923): 99. Type: MEXICO. Sonora, Hermosillo, 1888, M. A. Crawford s.n. (holotype: GH 00008349!; isotype: US 00129558!-fragment and photograph of the holotype).

Gnaphalium texanum I.M.Johnst. (Johnston 1924): 86. Type: UNITED STATES OF AMERICA. Texas, Brewster Co., Mouth of Tarlinga Creek, Sep 1883, V. Havard 26 (holotype: GH 00008310!; probable isotype: US 00129562! [s.n.]).

Gnaphalium viridulum I.M.Johnst. (Johnston 1924): 86. Type: UNITED STATES OF AMERICA. New Mexico, Grant Co., Bear Mts., near Silver City, 2400 m, 19 Sep 1903, O. B. Metcalfe 742 (holotype: GH 00008312!; isotype: US 00129565!).

Gnaphalium albidum I.M.Johnst. (Johnston 1924): 84. Type: UNITED STATES OF AMERICA. California, San Diego Co., granite, in chaparral, 1850 ft [550 m], 11 Jul 1916, M. F. Spencer 69 (holotype: GH 00008288!; isotype: US 00129528!).

Observations:

1. According to the protologue, Gnaphalium canescens was based on the specimen ‘in Mexico ad Leonem ultrà Guanaxuato legit cl. Mendezʼ. We found at G-DC, where the original herbarium of Candolle is deposited, the sheet G 00312669 ‘Mexiq. (Léon à l’ouest de Guanajuato), 1829, M. Mendezʼ [sheet 1], which is in accordance with the protologue and is considered as the holotype of Gnaphalium canescens.

2. According to the protologue, Gnaphalium microcephalum was based on the specimen ‘St. Diego, Upper Californiaʼ. We found at BM, where the original herbarium of Nuttall is mainly deposited, the sheet Nuttall s.n. BM 001010942, which is in accordance with the protologue and matches the locality. Since Nuttall mentioned in the protologue ‘(I have seen but a single specimen)ʼ this specimen is almost surely the holotype. There are other two collections on the same sheet BM 001010942, i.e. Suksdorf s.n. and Macoun s.n.

3. Nesom (2004): 782 indicated the type of Gnaphalium wrightii A.Gray as ‘Type. U.S.A. Texas [El Paso or Hudspeth Co.] valley between El Paso and the Guadalupe Mts., Oct [1849], C. Wright 394 (Lectotype, designated here: GH!; Isolectotypes: GH!, US!)ʼ. This constitutes a valid [first-step] lectotypification. Since we located two sheets of this collection at GH, ‘second-step’ lectotypification is required (ICN, Art. 9.17. Shenzhen Code; Turland et al. 2018). The specimen GH 00008314 which presents the most complete plant is here designated as the lectotype of G. wrightii. Furthermore, this is the specimen that was labelled as the ‘lectotype’ by Kittredge in 2009, and shows that Nesom did not annotate either of the two sheets in GH.

4. The types of new species described by Osterhout were deposited in his private herbarium and, upon his death, all his sheets were bequeathed to the Rocky Mountain Herbarium in Laramie, Wyoming. Consequently, the only collection kept at RM (RM 0001102) is considered as the holotype of Gnaphalium albatum.

Distribution: Mexico and United States of America.

11. Pseudognaphalium chartaceum (Greenm.) Anderb. (Anderberg 1991: 147). Gnaphalium chartaceum Greenm. (Greenman 1904: 95). Type: MEXICO. Jalisco: Hillsides and banks near Guadalajara, 22 Nov 1888, C. G. Pringle 1827 (lectotype, designated by Pruski (2018: 225): GH 00008319!, isolectotypes: BR 0000005317776!, K 000500362!, M 0029851!, MEXU 01215400!, MICH 1107416!, NDG 59113!, UC 88344!, US 00129536!; syntype: Puebla, near Tehuacan, 22 Dec 1895, C. G. Pringle 7033, GH 00008320!).

Distribution: Mexico and Mesoamerica [Guatemala (Pruski 2018)].

12. Pseudognaphalium conoideum Anderb. (Anderberg 1991: 147), nom. nov. Gnaphalium conoideum Kunth (1818, folio edition): 67; (1820, quarto edition): 85 p., hom. illeg. non G. conoideum Lam. 1788. Type: MEXICO. ‘Crescit prope urbem Mexici, alt. 1170 hex. Floret Majo’, F. W. H. A. Humboldt & A. J. A. Bonpland s.n. [4157] (holotype: P 00322315!; isotypes: P 00704488!, P 00704489-fragment!).

Distribution: Mexico.

13. Pseudognaphalium ehrenbergianum (Sch.Bip. ex Klatt) Hinojosa-Espinosa & Villaseñor (2014): 490. Gnaphalium ehrenbergianum Sch.Bip. ex Klatt (1878): 127. Type: MEXICO. [Hidalgo] Zamaltepan [=Zacualtipán], Ehrenberg 24 (P- fragment 00704465! ex herb. Schultz Bipontinus 1845).

Distribution: Mexico.

Observation. Since the material, ex herb. (Schultz Bipontinus 1856) is not annotated by Klatt one could assume that he saw material in B, now destroyed, in order to provide some parts of the description.

14. Pseudognaphalium elegans (Kunth) Kartesz (1999): no. 28. Gnaphalium elegans Kunth (1818, folio edition): 63; (1820, quarto edition): 81, p. 30 Type: ECUADOR. Pichincha, 2450 m, Jun 1802, F. W. H. A. Humboldt & A. J. A. Bonpland 3001 (lectotype, designated by Freire et al. (2018): 330: P-HBK 00322306!; isolectotypes: P 00704547!, P 00704548!).

Gnaphalium poeppigianum DC. (Candolle 1838: 227). Type: PERU. Dpto. Huánuco, Cuchero, Sep 1829, E. F. Poeppig n. 34 diar. 1368 (lectotype, designated by Freire et al. (2018: 330): G-DC G-00469598!; isolectotypes: F 881500!, F 970435!-fragment, B† photo F0BN015138!, GH 00008361! fragment, GH 00008362!, HAL 112159!, NY 00169513!, P 00704530!, P 00704531!).

Distribution: Mexico, Mesoamerica [Costa Rica, Guatemala, Honduras, Nicaragua, Panama (Pruski 2018)] and South America [Colombia, Ecuador, Peru, Venezuela (Freire et al. 2018), Guyanas (Boggan et al. 1997)].

15. Pseudognaphalium greenmanii (S.F.Blake) Anderb. (Anderberg 1991: 147). Gnaphalium greenmanii S.F.Blake (1931): 329, nom. nov. for Gnaphalium linearifolium Greenm. (Greenman 1897): 308, hom. illeg. non G. linearifolium (Wedd.) Franch. 1892. [= Quasiantennaria linearifolia (Wedd.) R.J.Bayer & M.O.Dillon]. Type: MEXICO. Jalisco, rocky hills near Guadalajara, 7 Oct 1889, C. G. Pringle 2342 (lectotype, designated by McVaugh (1984): 455: GH 00008329!; isolectotypes: CAS 0002858!, CM 2400!, GOET 001569!, MEXU 01215406!, NDG 59124!, UC 88345!, US 00129543!, UVMVT 027543!, UVMVT 027544!).

Distribution: Mexico and Mesoamerica [Guatemala, Honduras (Pruski 2018)].

16. Pseudognaphalium helleri (Britton) Anderb. (Anderberg 1991: 147). Gnaphalium helleri Britton (1893: 280). G. polycephalum Michx. var. helleri (Britton) Fernald (1908: 94). G. obtusifolium L. var. helleri (Britton) S.F.Blake (1918: 72). Type: UNITED STATES OF AMERICA. Virginia: Southeast Virginia, Norfolk Co., 23 Sep 1892, A. A. Heller s.n. (lectotype [first step], designated by Mahler (1975): 30: NY; lectotype [second step], designated here: NY 00169470!; probable isolectotypes: E 00385958!, GH 00008294!, MO 2168385!, NY 00169472!, PH 00012602!, PH 00012603!, PH 00012604!, US 00129539! [all these probable isolectotypes bear the number 741]. Figure 3a-h.

Gnaphalium obtusifolium L. var. micradenium Weath. (Weatherby 1923): 22. G. helleri Britton var. micradenium (Weath.) Mahler (1975): 32. Pseudognaphalium helleri (Britton) Anderb. subsp. micradenium (Weath.) Kartesz (1999): n 29. Pseudognaphalium micradenium (Weath.), Nesom (2001b): 618. Type: UNITED STATES OF AMERICA. Massachusetts: Barnstable Co., 7 Oct 1917, M. L. Fernald 15870 (holotype: GH 00008300!; isotype: NEBC 00231216!).

Observation: Mahler (1975): 30 indicated the type of Gnaphalium helleri Britton as ‘Lectotype selected. Northwest Norfolk County, Virginia, A. A. Heller s.n., 23 Sep 1892, annotated by Britton (NY). Syntypes. Northwest Norfolk County, Virginia, A. A. Heller s.n., 23 Sep 1892, annotated by Britton (NY, MO) …ʼ. By referring to the collection at NY as ‘lectotypeʼ, this constitutes a valid [first-step] lectotypification. Since Mahler (1975) mentioned two sheets of this collection at NY, ‘second-step’ lectotypification is required (ICN, Art. 9.17. Shenzhen Code; Turland et al. 2018). The specimen NY00169470 which has two complete plants with inflorescences and a label of 1974 in which Mahler annotated ‘lectotypeʼ is here designated as the lectotype of G. helleri.

Distribution: Eastern United States of America.

17. Pseudognaphalium hintoniorum (G.L.Nesom) Hinojosa-Espinosa & Villaseñor (2014): 490. Gnaphalium hintoniorum Nesom (1990): 414. Type: MEXICO. Nuevo León: Mpio. Galeana, Cerro Potosí, pine forest, 3350 m, 15 Sep 1969, G. B. Hinton et al. 17263 (holotype: TEX 00373729!; isotypes: ANSM 094528!, CIIDIR 017091!, GBH 017263!, MICH 1107417!).

Gnaphalium flavocephalum Nesom (1990): 413. Type: MEXICO. Coahuila, Mpio. Arteaga. Sierra del Coahuilon, open limestone hillsides, 3200 m, 28 Jul 1985, G. B. Hinton et al. 18902 (holotype: TEX 00373728!; isotypes: ANSM 064097, CIIDIR 022434, GBH 018902).

Distribution: Mexico.

18. Pseudognaphalium inornatum (DC.) Anderb. (Anderberg 1991: 147). Gnaphalium inornatum DC. (Candolle 1838: 225). Type: MEXICO. Morelos: ‘Cordillère de Guchilaque, 8bre 1827ʼ, J. L. Berlandier 1146 (lectotype, designated by McVaugh (1984): 456: G-DC G-00312682!; isolectotypes: G 00301178!, P 00704472!, P 00704473!; syntypes: J. L. Berlandier 309 G-DC G-00312681!, J. L. Berlandier 740 G-DC G-00312660!, J. L. Berlandier 1195 G-DC G-00312680!; isosyntypes: J. L. Berlandier 309 HAL 0112155!, P 00704469!, P 00704470!, J. L. Berlandier 740 HAL 0112154!, J. L. Berlandier 1195 P 00704472!).

Distribution: Mexico.

19. Pseudognaphalium jaliscense (Greenm.) Anderb. (Anderberg 1991: 147). Gnaphalium jaliscense Greenm. (Greenman 1904: 96). Type: MEXICO. Jalisco. Gravelly soil near Guadalajara, 9 Aug 1902, C. G. Pringle 8628 (lectotype, designated by Nesom (2001c): 1187: GH not seen; isolectotypes: C 10007405!, CAS 0002857!, CM 2399!, E 00433304!, ENCB 003704!, JE 00000746!, K 000500360!, LL 00373730!, M 0029852!, MEXU 01215402!, MEXU 01215403!, MICH 1107418!, MIN 1001224!, MSC 0091947!, NY 00169490!, P 00704477!, P 00704478!, PH 00012605!, RSA 0001212!, UC 88403!, US 01101249!, UVMVT 027540!, S 07-11985!).

Distribution: Mexico and United States of America.

20. Pseudognaphalium leucocephalum (A.Gray) Anderb. (Anderberg 1991: 147). Gnaphalium leucocephalum A. Gray (1853): 99. Type: MEXICO. Sonora: Bed of mountain torrents near Santa Cruz, 23 Sept. 1851, C. Wright 1285 (lectotype, designated here: GH 00254927!; isolectotypes: GH 00008297!, K 000978299!, NY 00169474!, NY 00169475!; syntype: Texas, Pecos County, Western Texas to El Paso, New Mexico [80 miles beyond the Pecos], 21 Aug 1849, C. Wright 393, GH 00008296!, GH 00008299!, GH 00282538!, K not seen).

Observation. In the protologue of Gnaphalium leucocephalum, Gray (1853) mentioned ‘G. polycephalum Gray, Pl. Wright p. 124. no. 393, non Michx.– Bed of mountain torrents near Santa Cruz, Sonora; Sept. (1285)’. Stafleu & Cowan (1976) stated that the herbarium types of Asa Gray are at GH. We located at GH two specimens of C. Wright 1285 (GH 00254927 and GH 00008297) and three other specimens of C. Wright 393 (GH 00008296, GH 00008299 and GH 00282538). Only two of these specimens, Wright 1285-GH 00254927 and Wright 393-GH 00008296, were there when Asa Gray described the species, the remaining are duplicates transferred to GH from the Boston Natural History Society at a later date. We selected the specimen C. Wright 1285 GH 00008297 as lectotype of Gnaphalium leucocephalum, because it bears the name of the new species written on it in Gray’s own hand with the annotation ‘n. sp’ and it is better preserved. There are other two collections on the same sheet GH 00254927, i.e. Thurber 924 GH 00254926 and Xantus 66 GH 00008298.

Distribution: Mexico and United States of America.

21. Pseudognaphalium liebmannii (Sch.Bip. ex Klatt) Anderb. (Anderberg 1991: 147). Gnaphalium liebmannii Sch.Bip. ex Klatt (1887): 89. Type: MEXICO. [Veracruz], Pico de Orizaba, 10.000 ped., F. M. Liebmann 310. (holotype: C 10007406!, isotype: GH 00008328-fragment! together with a pencil sketch (by Klatt) of the holotype and

a full Latin description as appearing in Leopoldina).

Gnaphalium vulcanicum I.M.Johnst. (Johnston 1923): 100. Type: MEXICO. [Veracruz], Citlaltepetl (=Pico de Orizaba), near timber line, Sep 1907, C. A. Purpus 2782 (holotype: GH 00008352!; isotype: US 00129566!).

Distribution: Mexico and Mesoamerica [Guatemala (Pruski 2018)].

22. Pseudognaphalium luteoalbum (L.) Hilliard & Burtt (1981): 206. Gnaphalium luteoalbum L. (Linnaeus 1753): 851. Gnaphalium conglobatus Lam. (Lamarck 1779, 1788): 64. Helichrysum luteoalbum (L.) Rchb. (Reichenbach 1829): 1460. Filaginella luteoalba (L.) Opiz (1852): 44. Gnaphalium luteoalbum L. var. incanum A.Rich. ex Endl. (Endlicher 1833: 50), nom. illeg. Dasyanthus conglobatus (Lam.) Bubani (1900: 199), nom. illeg. Laphangium luteoalbum (L.) Tzvelev (1994): 105. Type: EUROPA. ‘Habitat in Helvetia, G. Narbonensi, Hispania, Lusitania’ A. van Royen s.n. (lectotype, designated as holotype by Hilliard & Burtt (1981): 206: L, sheet 900. 286-294).

Gnaphalium trifidum Thunb. (Thunberg 1800): 150. Type: SOUTH AFRICA. Cape of Good Hope, C. P. Thunberg s.n. (UPS 19278!).

Gnaphalium multiceps Wall. ex DC. (Candolle 1838: 222). Gnaphalium luteoalbum L. var. multiceps (Wall. ex DC.) Hook.f. (Hooker 1881: 288). Type: NEPAL. Mar 1821, N. Wallich 2949-59A, (lectotype, designated by Freire et al. (2018): 339: K 001118249!; isolectotype: BM 000521869!, probable isolectotype E 00531206!).

Gnaphalium luteofuscumWebb (1849): 143. Syntypes: ‘Hab. In petrosis supra medium Montis Verede ins. S. Vincentii (Vogel, n. 38. 55. 56. Junio, 1841, sp. florida et fructifera.)’ (Vogel 55, 56, K).

Gnaphalium luteoalbum L. var. pallidum Hook.f. (Hooker 1881): 288. Type: INDIA. ‘G. pallidum, Ham. in Wall. Cat. 2953.’ (lectotype, designated by Freire et al. (2018): 339: K 001118256!; isolectotypes: K 001118255!, K 001118257!).

Gnaphalium luteoalbum L. var. compactum Kirk (1899): 298. Type: [New Zealand]

‘Lake Lyndon, Enys and Kirk’.

Observation. In the protologue of Gnaphalium luteoalbum var. compactum, Kirk (1899) mentioned ‘Lake Lyndon, Enys and Kirk’. We located one specimen at K […of Lake Lyndon, 2800 p, New Zealand, T. Kirk 1093 com. 9/1884] K 000975815, which totally corresponds to the protologue in its detail, including agreement for the locality. However, since we were able to see only one sheet, we prefer not to lectotypify the name at the moment.

Distribution: Eurasian species with cosmopolitan distribution, being adventitious in Mexico and United States of America (Pruski 2018).

23. Pseudognaphalium macounii (Greene) Kartesz (1999): no. 30. Gnaphalium macounii Greene (1902): 278. Type: CANADA. British Columbia, Chilliwack Valley, B.C., 2500 ft, 29 Jul 1901, J. M. Macoun 26487 [26847 in lit.] (holotype: NDG 59133!).

Gnaphalium ivesii A.Nelson & J.F.Macbr. (Nelson & Macbride 1916): 46 as nom. nov. for Gnaphalium decurrens Ives (1819: 380), hom illeg., non G. decurrens L. 1759. Type: Plate s.n. sub. ‘Gnaphalium decurrens’ (Ives 1819) (lectotype, designated here).

Observations. The protologue of Gnaphalium decurrens Ives does not include any reference to the collector: ‘This plant was observed by me in company with Mr. C. Whitlow, in July, 1817, by the margin of a brook, a few rods north of Mr. E. Whitney‘s gun manufactory, near New Haven. It it is also found on the margin of the Housatonick about thirty miles from Long Island soun where it was observed by Dr. Alfred Monson, the last summer’. The name of this species is accompanied by an illustration, which is here selected as the lectotype.

Distribution: Canada and United States of America.

24. Pseudognaphalium monticola (McVaugh) Villarreal, Estrada & Encina (2020): 5. Gnaphalium vulcanicum I.M.Johnst. var. monticola McVaugh (1972): 466. Gnaphalium liebmannii Sch.Bip. ex Klatt var. monticola (McVaugh), Nash (1974): 74. Pseudognaphalium liebmannii (Sch.Bip. ex Klatt) Anderb. var. monticola (McVaugh) Hinojosa-Espinosa & Villaseñor (2014): 490. Type: MEXICO. Jalisco. Sierra de Manantlán, 25-30 km SE of Autlan, pine forest near summits between El Chante and Cuzalapa, 19°35’N, 104°8-15´W, 2750 m, 20-21 Mar 1965, R. McVaugh 23126 (holotype: MICH 1107419!; isotypes: CAS 0002859!, ENCB 003706!, NY 00169500!).

Distribution: Mexico and Mesoamerica [Guatemala (Pruski 2018)].

25. Pseudognaphalium nataliae (F.J.Espinosa) Villarreal, Estrada & Encina (2020): 6. Gnaphalium oxyphyllum (DC.) var. nataliae F.J.Espinosa (1983: 17). Pseudognaphalium oxyphyllum (DC.) Anderb. var. nataliae (F.J.Espinosa) Hinojosa-Espinosa & Villaseñor (2014): 491. Type: MEXICO. Mexico, Municipio de Amecameca, San Pedro Nexcapa, 3000 m, 21 Sep 1979, F. J. Espinosa 766 (holotype: MEXU 00343482!; isotypes: ANSM 030402!, ENCB 003705!).

Distribution: Mexico.

26. Pseudognaphalium nubicola (I.M.Johnst.) Anderb. (Anderberg 1991: 147). Gnaphalium nubicola I.M.Johnst. (Johnston 1923): 98. Type: MEXICO. Rocks above timber-line, Mt. Ixtacuhuatl, Nov 1905, C. A. Purpus 1524 ‘typeʼ (holotype: GH 00008332!; isotypes: GH 00008333!, UC 134562!, US 00129545!).

Distribution: Mexico.

27. Pseudognaphalium obtusifolium (L.) Hilliard & Burtt (1981): 205. Gnaphalium obtusifolium L. (Linnaeus 1753): 851. Type: UNITED STATES OF AMERICA. [Habitat in Virginia, Pennsylvania] Kalm, Herb. Linn. No. 989.64 (lectotype, designated by Reveal (1998: 362): LINN).

Gnaphalium obtusifolium L. var. praecox Fernald (1936: 231). TYPE. United States of America. Maine: York Co., 2 miles east of Walterboro, 17 July 1927, K. M. Wiegand & W. E. Manning 3301 (holotype: GH 00008301!).

Distribution: United States of America and adjacent Canada.

28. Pseudognaphalium oxyphyllum (DC.) Kirp. (Kirpichnikov 1960): 33. Gnaphalium oxyphyllum DC. (Candolle 1838: 225). Type: MEXICO. Villalpando au Sud est de Guanajuato, J. Mendez s.n. (lectotype, designated here: G-DC G00469505! sheet 4, isolectotypes: G-DC G00469505! sheet 1-left hand plants, sheet 2-left hand plants, sheet 3).

Observation: The specimen G-DC G00469505 consists of four sheets. However, on sheet 1 and 2 the three plants on right of sheet correspond to Gnaphalium oxyphyllum var. semilanatum. Because the sheet 4-G-DC G-00469505 has a label on it, identifying it as the type and matches the locality provided in the protologue, it is here formally designated as the lectotype of Gnaphalium oxyphyllum DC.

Distribution: Mexico and Mesoamerica [Guatemala (Espinosa-García 2005)].

29. Pseudognaphalium pringlei (A.Gray) Anderb. (Anderberg 1991: 147). Gnaphalium pringlei A. Gray (1886): 387. Type: MEXICO. Chihuahua: cool shades slopes, rocky hills near Chihuahua, 25 Oct 1885, C. G. Pringle 305 (holotype: GH 00008341!; isotypes: AC 00319433!, BR 0000005317783!, CM 2402!, COLO 00368399!, CORD 00004577!, E 00433303!, FI 005289!, GOET 001573!, JE 00000749!, K 000500355!, MEXU 01215407!, MO 191020!, NY 00169497!, NY 00169498!, P 00704570!, P 00704571!, P 00704572!, PH 00012614!, RSA 0001213!, TEX 00373731!, UC 88343!, US 00129552!, US 00930967!, UVMVT 027542!).

Distribution: Mexico and United States of America.

30. Pseudognaphalium purpurascens (DC.) Anderb. (Anderberg 1991: 147). Gnaphalium purpurascens DC. (Candolle 1838: 225). Type: MEXICO. 1833, M. Mairet s.n. (holotype: G-DC G-0469517!; probable isotype: P 00704573!, without collector).

Distribution: Mexico.

31. Pseudognaphalium ramosissimum (Nutt.) Anderb. (Anderberg 1991: 147). Gnaphalium ramosissimum Nutt. (Nuttall 1848): 20, non G. ramosissimum Sch.Bip. 1856, hom. illeg. [=Achyrocline ramosissima Britton ex Rusby]. Type: UNITED STATES OF AMERICA. California. Monterrey (“Monterey”), W. Gambel s.n., -Types: GH 00008306!, GH 00008307!, K 001096583! “Nuttall from Gambel”, written on sheet.

Observation. Stafleu & Cowan (1981) stated that the types of Nuttall are mainly deposited in BM and PH with additional specimens or duplicates in GH, K, MO, NY, OXF. Since we not able to see Nuttall specimen at PH and BM, we prefer not to lectotypify the name at the moment.

Distribution: United States of America.

32. Pseudognaphalium roseum (Kunth) Anderb. (Anderberg 1991: 147). Gnaphalium roseum (1818, folio edition): 63; (1820, quarto edition): 81, p. 33. Type: MEXICO. ‘Crescit in covalli Guanaxuatensi, alt. 1700 hex. (Regno Novae Hispaniae) Floret Septembriʼ Guanajuato, F. W. H. A. Humboldt & A. J. A. Bonpland s.n. (holotype: P 00322305!).

Distribution: Mexico, Mesoamerica [Belice, Guatemala, Honduras, Nicaragua (Pruski 2018)], and United States of America.

33. Pseudognaphalium saxicola (Fassett) H.E.Ballard & Feller (2004: 777). Gnaphalium saxicola Fassett (1931: 75). Gnaphalium obtusifolium L.var. saxicola (Fassett) Cronquist (1946: 121). Pseudognaphalium obtusifolium (L.) Hilliard & Burtt 1981 subsp. saxicola (Fassett) Kartesz (1999): no. 32. Type: UNITES STATES OF AMERICA. Wisconsin: Adams Co., sandstone ledges, cold water canyon, Dells of the Wisconsin River, 22 Sep 1929, N. C. Fassett, F. M. Uhler & W. T. McLaughlin 9590 (holotype: WIS v0011634!; isotypes: F 0050259F!, GH 00008308!, NY 00169480!).

Distribution: United States of America.

34. Pseudognaphalium semiamplexicaule (DC.) Anderb. (Anderberg 1991: 148). Gnaphalium semiamplexicaule DC. (Candolle 1838: 228). Type: MEXICO. ‘Vittoria a Tula, Nov 1830ʼ, J. L. Berlandier 2188 (lectotype, designated here: G-DC G-00469541!; isolectotypes: G 00223947!, G-DC G-00469542!, GH 00008348!, K 000500347!, P 00704575!, PH 00012624!). Figure 4a-g.

Gnaphalium oxyphyllum DC. var. semilanatum DC. (Candolle 1838: 225). Gnaphalium semilanatum (DC.) McVaugh (1972): 465. Pseudognaphalium semilanatum (DC.) Anderb. (Anderberg 1991: 148). Type: MEXICO. Villalpando, au Sud est de Guanajuato, J. Méndez s.n. (lectotype, designated here: G-DC G-00469505! sheet 1-right hand plants; isolectotypes: G-DC G-00469505! sheet 2-right hand plants, probable isolectotype: G-DC G-00469537!), syn. nov.

Observations:

1. According to the protologue, Gnaphalium semiamplexicaule was based on the specimen ‘in Mexico inter Vittoria et Tula legit cl. Berlandier pl. exs. n. 2188ʼ. We found two sheets at G-DC, G-DC G-00469541[sheet 2] and G-DC G-00469542 [sheet 1]. We propose the specimen G-DC G-00469541 which bears the number ‘2’ written in pencil in the top right hand corner, as the lectotype of Gnaphalium semiamplexicaule, since this is the only one that bears the annotations provided by Candolle.

2. Candolle (1838: 225) mentioned ‘β. semilanatum, foliis subtùs albo-lanatis, ̶ Cum var. α mixtum in pl. Mendezianis. (v.s.)ʼ. We located at G-DC, where the original herbarium of Candolle is deposited, the collection ‘ Villalpando, au Sud est de Guanajuato, J. Mendez s.n. ʼ G-DC G-00469505 of four sheets (see observation of Pseudognaphalium oxyphyllum). On sheet 1 and 2 the three plants on right have discolorous lanceolate leaves, with abaxial surface white lanate. We propose the sheet 1- G-DC G-00469505 as the lectotype of Gnaphalium oxyphyllum var. semilanatum.

Distribution: Mexico and Mesoamerica [Belice, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama (Pruski 2018)].

35. Pseudognaphalium stramineum (Kunth) Anderb. (Anderberg 1991: 148). Gnaphalium stramineum Kunth (1818, folio edition): 64; (1820, quarto edition): 82. Type: MEXICO. ‘Crescit locis temperatis inter Moran et Omitlan Mexicanorum, alt. 1300 hex.ʼ, F. W. H. A. Humboldt & A. J. A. Bonpland 4108 (holotype: P 0322314!; isotype: P 00704565!).

Gnaphalium berlandieri DC., Prodr.6: 223. 1838. Type: MEXICO: 30 Jun 1827, J. L. Berlandier 471 (lectotype, designated here: G-DC G-00469612!; isolectotypes: G-DC G-00469600! 24 Jun 1827, HAL 0112152! 29 Jun 1827, LP 001900! 24 Jun 1827, MICH 1107415!, 30 Jun 1827?, P 00704481! 24 Jun 1827, P 00704479!, P 00704480! Jun 1827, P 00704481!, 24 Jun 1827).

Gnaphalium gossypinum Nutt. (Nuttall 1841): 403. Type: UNITED STATES OF AMERICA. Oregon [Río Columbia], 1834-1835, T. Nuttall s.n. (not seen).

Gnaphalium chilense Spreng. var. confertifolium Greene (1897): 400. Type: UNITED STATES OF AMERICA. California. San Francisco, 27 May 1893, E. L. Greene (holotype, NDG 59239!).

Gnaphalium sulphurescens Rydb. (Rydberg 1900: 415). Type: UNITED STATES OF AMERICA. Yellowstone Park, Lower Geyser Basin, 7500 ft, 4 Aug 1897, P. A. Rydberg & E. A. Bessey 5135 (lectotype, designated here: NY 00169483!; isolectotypes: E 00433320!, GH 00008309!, K 000978305!, NDG 59192!, NEB-v-0000112!; NEB-v-0000111!, PH 00012625!, RM0001109!, US 00129561!, US 00130793!).

Gnaphalium lagopodioides Rydb. (Rydberg 1900: 416). Type: UNITED STATES OF AMERICA. Yellowstone Park, Lower Geyser Basin, 7500 ft, 4 Aug 1897, P. A. Rydberg & E. A. Bessey 5134 (holotype: NY00169473!; isotypes: GH 00008295!, NDG 59122!, NEB-v-0000110!, US 00129542!).

Gnaphalium proximumGreene (1902): 279. Type: UNITED STATES OF AMERICA. Wyoming. In moist ground about warm springs, Mammoth Hot Springs, Yellowstone Park, 21 Jul 1899, A. Nelson & E. Nelson 6036 (holotype: NDG 59162; isotypes: NY 00169479!, RM 0001107!, RM 0001108!, US 00129553!).

Observations:

1. Rydberg (1900: 415) mentioned in the protologue of Gnaphalium sulphurescens: ‘Yellowstone Park: Lower Geyser Basin, August 4, 1897, Rydberg & Bessey, 5135; Hot Springs, 1884, Tweedy, 172; Mud Springs, 1871, Hayden Survey’. We propose as the lectotype of G. sulphurescens the collection Rydberg & Bessey 5135 kept at NY which is a widely distributed collection (duplicates seen at E, GH, K, NDG, NEB, PH, RM, and US), and the specimen kept at NY, NY 00169483, because, according to the preface to Rydberg’s paper, ‘Rydberg & Bessey, 1897’ type materials are found in this herbarium.

2. Candolle (1838: 223) mentioned for Gnaphalium berlandieri: ‘circa urbem Mexico media aestate legit cl. Berlandier pl. exs. n. 471! … (v.s.)’. We found two sheets at G-DC of the collection Berlandier 471 G-DC G-00469600 [sheet 1], 24 Jun 1827 and G-DC G-00469612 [sheet 2], 30 Jun 1827. We selected as the lectotype of G. berlandieri, the specimen G-DC G-00469612 with the pencil number ‘2’ in the top right hand corner, which is the only one annotated by de Candolle and has a annotation on the original Berlandier label indicating there were ‘14 ex’ [= duplicates].

Distribution: Canada (Pruski 2018), Mexico, United States of America and Mesoamerica [Guatemala, Honduras (Pruski 2018)].

36. Pseudognaphalium thermale (Nelson 1900, 2004): 781. Gnaphalium thermale Nelson (1900): 121. Gnaphalium microcephalum Nutt.var. thermale (Nelson 1900) Cronquist (1950: 47). Gnaphalium microcephalum Nutt. subsp. thermale (Nelson 1900) G.W. Douglas (1986: 2726). Gnaphalium canescens DC. subsp. thermale, (Nelson 1900) Stebbins & Keil (1992): 437. Pseudognaphalium canescens (DC.) Anderb. subsp. thermale (Nelson 1990) Kartesz (1999): no. 27. Pseudognaphalium microcephalum (Nutt.) Anderb. var. thermale (Nelson 1900) Dorn (2001: 375). Type: UNITED STATES OF AMERICA. Yellowstone National Park, Norris Geyser Basin, 25 Jul 1899, A. Nelson & E. E. Nelson 6139 (lectotype, designated here: RM 0001110!; isolectotypes: G!, GH 00008311!, K 000978306!, NDG 59193!, NEB-v-0000113!, NY 00169484!, P 00704566!, RM 0001111!, US 00129563!).

Gnaphalium johnstonii. Jones (1938): 159, 176. Type: UNITED STATES OF AMERICA. Washington. [Thurston Co.], upper valley of the Nesqually, open, barren ground, 15 Sep 1896, O. D. Allen 223 (isotype: WTU-V-001136!).

Observation: Nesom (2004): 781 indicated the type of Gnaphalium thermale E.E.Nelson as ‘Type. U.S.A. Wyoming [Park Co.]: Yellowstone Park, geyser... A. Nelson & E. Nelson 6139 (Holotype; RM; Isotypes: GH!, US!)ʼ. This citation of a ‘holotypeʼ (after 1 January 2001) cannot be corrected to a designation of a lectotype (Art. 9.10 of the ICN, Shenzhen Code; Turland et al. 2018), because they did not use the phrase ‘designated hereʼ or an equivalent (ICN, Art. 7.11, Shenzhen Code; Turland et al. 2018) in their associated statement. Consequently, Nesom’s typification for the name G. beneolens was not effective (ICN, Art. 9.23, Shenzhen Code; Turland et al. 2018). The specimen RM 0001110 which presents the most complete plant is designated here as the lectotype of G. thermale.

Distribution: United States of America.

37. Pseudognaphalium viscosum (Kunth) Anderb. (Anderberg 1991): 148. Gnaphalium viscosum Kunth (1818, folio edition): 64; (1820, quarto edition): 82. Type: MEXICO. [Distrito Federal] ‘Crescit in radicibus montis Chapoltepec juxta urbem Mexiciʼ, F. W. H. A. Humboldt & A. J. A. Bonpland 4153 (holotype: P-HBK, not seen; isotype, P-00704569!).).

Gnaphalium hirtum Kunth (1818, folio edition): 64; (1820, quarto edition): 82. Type: MEXICO. ‘Crescit in radicibus montis Chapoltepec juxta urbem Mexici, alt. 1170 hex. Floret Apriliʼ F. W. H. A. Humboldt & A. J. A. Bonpland s.n. (holotype: P 00322309!; isotype: P 00704568!).

Gnaphalium gracile Kunth (1818, folio edition): 65; (1820, quarto edition): 83. Type: MEXICO. ‘Crescit in Regno Mexicano in declivitate montis Serenae, Guanaxuatum inter et fodinam Belgradensem alt. 1100 hex. Floret Septembriʼ, F. W. H. A. Humboldt & A. J. A. Bonpland s.n. (holotype: P 00322310!).

Gnaphalium tenue Kunth (1818, folio edition): 65; (1820, quarto edition): 83. Type: MEXICO. ‘Crescit in Nova Hispaniaʼ, F. W. H. A. Humboldt & A. J. A. Bonpland s.n. (holotype: P 00322311!; isotype: P 00714567!).

Gnaphalium leptophyllum DC. (Candolle 1838: 226). Type: MEXICO. 1831, L. Alaman s.n. (holotype: G-DC G-00469611!).

Gnaphalium crenatum Greenm. (Greenman 1904): 96. Pseudognaphalium crenatum (Greenm.) Anderb. (Anderberg 1991): 147. Type: MEXICO. Jalisco, near Guadalajara, 5000 ft, 15 May 1901, C. G. Pringle 9524 (holotype: GH 00008322!; isotypes: CM 2398!, ENCB 003703!, K 000500361!, MEXU 01215401!, NY 00169489!, TEX 00000466!, UV MVT027538!).

Pruski (2018: 230) indicated the type of Gnaphalium viscosum Kunth as ‘Holotipo: Mexico, Distrito Federal, Humboldt y Bonpland 4153ʼ, erroneously as ‘P-Bonpʼ, pers. comm.

Distribution: Mexico, United States of America, and Mesoamerica [Guatemala, Honduras (Pruski 2018)].

Excluded species

Gnaphalium chilense Spreng. (1826: 480). Gnaphalium sprengelii Hook. & Arn. (1841[1833]: 150), nom. superfl. TYPE. ‘Chili. Chamisso. Peru?’ (P 00704592!).

Gnaphalium sprengelii Hook. et Arn. (1841[1833]: 150) is a nomen supefluum because this name was used to replace Sprengel’s Gnaphalium chilense (1826: 480), adopted by Lessing (1831): 525 for Californian plants. Rémy (1849): 228, mentioned Gnaphalium chilense Hook. & Arn. (1841[1833]: 31), non G. chilense Spreng. (1826), as a synonym of G. falcatum Lam. [= Gamochaeta falcata (Lam.) Cabrera]. One sheet of Gnaphalium chilense Spreng., ‘Chili. Chamisso. Peru?’, that bears the label ‘Sprgl. herb. no.: 827, Syt. III, 480… 168’, was located at P, P 00704592, where most of the original material used by Sprengel for his new taxa, in Systema Vegetabilium, is deposited. According to the digital image, the two plans on the sheet correspond to Gamochaeta falcata by having capitula arranged in spikes, pappus bristles basally connate, and linear obovate to obovate leaves.

Names of Dubious Identity

Pseudognaphalium altamiranum (Greenm.) Anderb. (1991: 147). Gnaphalium altamiranum Greenm. (Greenman 1904): 95. Type: MEXICO. Morelos, Mountains above Cuernavaca, 2640 m, 2 Feb 1899, C. G. Pringle 8041 (holotype: GH 00008316!; isotypes: CM 2397!, E 00433305!, G 00301190!, G 00301191!, GOET 001566!, K 000500364!, LL 00373727!, M 0029850!, MEXU 01215398!, MEXU 01215399!, MIN 1001223!, MSC 0091945!, NY 00169486!, P 00704490!, P 00704491!, PH 00012284!, RSA 0001208!, S07-11955!, UC 88249!, US 00129529!, US 01101248!, UVMVT 027539!).

Gnaphalium nubicola (I.M.Johnst.) Anderb. var. panniforme (S.F.Blake) F.J.Espinosa (1983: 20). Gnaphalium panniforme, Blake (1943): 268, replacement name for Gnaphalium pannosum A.Gray, hom. illeg., non Gnaphalium pannosum (DC.) Sch.Bip. 1845. Syntypes: MEXICO. ‘In the mountains near San Luis de Potosi, Schaffner 227, Parry & Palmer 420ʼ (both collections on sheet GH 00008339!; isosyntypes Parry & Palmer 420: NY 00169496 and US 00811084).

Gnaphalium oaxacanum Greenm. (Greenman 1904): 96. Type: MEXICO. Oaxaca. 1750 m, Jul-Aug 1900, C. Conzatti & V. González 1012 (holotype: GH 00008334!, isotypes: MEXU 00525673!, TEX 00000467!).

Since we were able to see only the images of type specimens of Pseudognaphalium altamiranum, Gnaphalium oaxacanum and G. nubicola var. panniforme, and we have seen no other material that can be assigned to these species, the identity of them is not clear to us.

ACKNOWLEDGMENTS

We thank the anonymous reviewers for their valuable suggestions on an earlier and the revised version of this paper. Special thanks are due to Nicholas Hind (K) who kindly helped us with nomenclature aspects. Appreciation is expressed to the directors and curators of the herbaria G, GH, M, MEXU, MO, NY, S, and UC for the loan of specimens that made this study possible. We thank María Alejandra Migoya (LPAG) for the illustrations of Pseudognaphalium helleri and P. semiamplexicaule. Also thanks to ‘Adumbratio Ilustración Científicaʼ for designing the figure 1 (a and b). Finally, we thank John Avise for making avaible to us the photographs of Pseudognaphalium canescens and Neal Kramer and Michael Mitchell the photographs of P. beneolens. Financial support (PIP 112-201501-00843) was provided by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina, and Universidad Nacional de La Plata, Programa de Incentivos, Secretaría de Políticas Universitarias, Ministerio de Educación, Argentina.

REFERENCES

• ACOSTA MAINDO A & GALBANY-CASALS M. 2018. Pseudognaphalium aldunateoides de nuevo bajo Gnaphalium (Compositae: Gnaphalieae). Collect Bot 37: 1-27.
• ANDERBERG AA. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot 104: 1-195.
• BALLARD HE, FELLER JR DS & NESOM GL. 2004. Cliff Cudweed at specific rank in Pseudognaphalium (Asteraceae: Gnaphalieae). Sida 21: 777-780.
• BAYER RJ, BREITWIESER I, WARD J & PUTTOCK C. 2007. Tribe Gnaphalieae. In: Kadereit JW & Jeffrey C (Eds), Kubitzki K (series editor), The families and genera of vascular plants, Flowering plants - Eudicots: Asterales, Berlin, Springer-Verlag. 8: 246-283.
• BIOLETTI FT. 1893. Two New Californian Plants. Erythea 1: 16-17.
• BLAKE SF. 1918. Notes on the Clayton Herbarium. Rhodora 20: 21-73.
• BLAKE SF. 1931. Nine new American Asteraceae. J Wash Acad Sci 21: 325-336.
• BLAKE SF. 1943. Ten new American Asteraceae. J Wash Acad Sci 32: 265-272.
• BOGGAN J, FUNK V, KELLOFF C, HOFF M, CREMERS G & FEUILLET C. 1997. Checklist of the Plants of the Guianas (Guyana, Surinam, Fench Guiana), 2nd ed. Biological Diversity of the Guianas Program, Smithsonian Institution, Washington, D.C, 238 p.
• BRITTON NL. 1893. New or Noteworthy North American Phanerogams-VII. Bull Torrey Bot Club 20: 277-282.
• BUBANI P. 1900. Flora Pyrenaea per Ordines Naturales gradatim digesta, Ulricus Hoeplius, Milan, vol 2, 718 p.
• CANDOLLE AP DE. 1838. Prodromus systematis naturalis regni vegetabilis, Sumptibus Sociorum Treuttel et Würtz, Paris, vol 6, 687 p.
• CRONQUIST AJ. 1946. Notes of the Compositae of Northeastern United States. III. Inuleae and Senecioneae. Rhodora 48: 116-125.
• CRONQUIST AJ. 1950. Notes of the Compositae of the Northwestern United States. Leafl W Bot 6: 41-56.
• CRONQUIST AJ. 1980. Vascular flora of the southeastern United States. Asteraceae, University of North Carolina Press, Chapel Hill, vol 1, 261 p.
• DAVIDSON A. 1918. Gnaphalium beneolens, N. Sp. Bull S Calif Acad Sci 17: 16.
• DAVIES PH & HEYWOOD VH. 1967. Principles of angiosperm taxonomy. Oliver & Boyd, Edimburgh and London, 556 p.
• DILLON MO. 2005. Asteraceae. In: Sklenář P, Luteyn JL, Ulloa C, Jørgensen PM & Dillon MO (Eds). Flora Genérica de los Páramos. Guía Ilustrada de las Plantas Vasculares. Mem New York Bot Gard 92: 96-180.
• DORN RD. 2001. Vascular Plants of Wyoming, ed 3. Mountain West Publishing, Cheyenne, Wyoming, 412 p.
• DOUGLAS GW. 1986. Nomenclatural changes in the Asteraceae of British Columbia. II: Astereae, Eupatoreae, Inuleae, Lactuceae, and Heliantheae. Canad J Bot 64: 2726-2727.
• ENDLICHER SFL. 1833. Prodromus Florae Norfolkicae. F Beck: Wien, 100 p.
• ESPINOSA-GARCÍA FJ. 1983. Notas taxonómicas y observaciones sobre algunas especies mexicanas de Gnaphalium (Compositae). Bol Soc Bot México 45: 15-21.
• ESPINOSA-GARCÍA FJ. 1985. Gnaphalium. In: Rzedowski J & Calderón de Rzedowski G (Eds), Flora Fanerogámica del Valle de México, Escuela Nacional de Ciencias Biológicas del Instituto Politécnico Nacional e Instituto de Ecología, México, DF 2: 507-523.
• ESPINOSA-GARCÍA FJ. 2005. Gnaphalium. In: Calderón de Rzedowski G & Rzedowski J y colaboradores, Flora Fanerogámica del Valle de México. 1a reimp, Instituto de Ecología, AC y Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. Páztcuaro (Michoacán), México, 2 ed, p. 840-856.
• FASSETT NC. 1931. Notes from the Herbarium of the University of Wisconsin-VI. Rhodora 33: 72-75.
• FERNALD ML. 1908. Notes of some plants of Northeastern America. Rhodora 10: 84-95.
• FERNALD ML. 1936. Contributions from the Gray Herabrium of Harvard University-No. CXIII. VIII. Varieties of Gnaphalium obtusifolium. Rhodora 38(450): 231-233.
• FERNÁNDEZ M, EZCURRA C & CALVIÑO CI. 2017. Species limits and morphometric and environmental variation within the South Andean and Patagonian Mulinum spinosum species-group (Apiaceae-Azorelloideae). Syst Biodivers 15(5): 489-505.
• FERRIS RS. 1960. Compositae. In: Abrams L & Ferris RS (Eds), Illustrated Flora of the Pacific States, Stanford Univ Press, Stanford 4: 98-613.
• FRANCHET AR. 1892. Observations sur le groupe des Leontopodium. Bull Soc Bot France 39: 126-136.
• FREIRE SE, BAYÓN ND, BAEZA CM, GIULIANO DA & MONTI C. 2014a.Revision of the genus Pseudognaphalium (Asteraceae, Gnaphalieae) in Chile. Gayana Bot 71: 68-107.
• FREIRE SE, BAYÓN ND & MONTI C. 2014b. Pseudognaphalium. In: Anton AM & Zuloaga FO (Dirs), Zuloaga FO, Belgrano MJ & Anton AM (Eds) & Freire SE (Coord), Flora Argentina, Asteraceae: Anthemideae-Gnaphalieae, Estudio Sigma SRL, Buenos Aires, Argentina 7(1): 495-509.
• FREIRE SE, MONTI C, BAYÓN ND & MIGOYA MA. 2018. Taxonomic Studies in Pseudognaphalium Kirp. (Asteraceae, Gnaphalieae) from Peru. Syst Bot 43: 325-343.
• GALBANY-CASALS M, ANDRÉS-SÁNCHEZ S, GARCÍA-JACAS N, SUSANNA A, RICO E & MARTÍNEZ-ORTEGA MM. 2010. How many of Cassini anagrams should there be? Molecular systematics and phylogenetic relationships in the Filago group (Asteraceae, Gnaphalieae), with special focus on the genus Filago. Taxon 59: 671-1689.
• GALBANY-CASALS M, UNWIN M, GARCÍA-JACAS N, SMISSEN RD, SUSANNA A & BAYER RJ. 2014. Phylogenetic relationships in Helichrysum (Compositae: Gnaphalieae) and related genera: Incongruence between nuclear and plastid phylogenies, biogeographic and morphological patterns, and implications for generic delimitation. Taxon 63: 608-624.
• GOWER JC. 1971. A general coefficient of similarity and some of its properties. Biometrics 27: 857-871.
• GRAY A. 1853. Plantae Wrightianae Texano-Neo Mexicanae. Smithsonian Contr Knowl 5: 3-119.
• GRAY A. 1882. Contribution to North American Botany. II. Novitiae Arizonicae, etc.: Characters of the new plants of certain recent collections, mainly in Arizona and adjacent Districts, etc. Proc Amer Acad Arts 17: 199-230.
• GRAY A. 1883. Contribution to North American Botany. I. Characters of new Compositae, with revision of certain genera, and critical notes. Proc Amer Acad Arts 19: 3-73.
• GRAY A. 1886. Contribution to American Botany. 2. Sertum Chihuahuense. Proc Amer Acad Arts 21: 378-409.
• GREENE EL. 1897. Flora franciscana: an Attempt to Classify and Describe the Vascular Plants of Middle California, Part 4. Cubery & Co., San Francisco, p. 353-480.
• GREENE EL. 1902. Some New Northwestern Compositae. Ottawa Naturalist 15: 278-282.
• GREENMAN JM. 1897. Descriptions of new or little-known plants from Mexico. Proc Amer Acad Arts 32: 295-311.
• GREENMAN JM. 1904. New and Otherwise Noteworthy Angiosperms from Mexico and Central America. Proc Amer Acad Arts 39: 69-120.
• GREENMAN JM. 1907. New or noteworthy Spermatophytes from Mexico, Central America and the West Indies. Publ Field Columb Mus Bot Ser 2: 247-287.
• GROSSI MA, GUTIÉRREZ DG, BERRUETA PC & MARTÍNEZ JJ. 2011. Acanthostyles (Asteraceae, Eupatorieae): a revision with a multivariate analysis. Aust Syst Bot 24: 87-103.
• HAMMER Ø & HARPER D. 2006. Paleontological Data Analysis. Blackwell Publishing, Oxford, 351 p.
• HILLIARD OM & BURTT BL. 1981. Some generic concepts in Compositae-Gnaphaliinae. Bot J Linn Soc 82: 181-232.
• HINOJOSA-ESPINOSA O & VILLASEÑOR JL. 2014. New combinations in Pseudognaphalium (Gnaphalieae-Asteraceae) of Mexico. Bot Sci 92: 489-491.
• HOOKER JD. 1881. Compositae. The Flora of British India, vol 3. Lovell Reever and Col, London, UK, 712 p.
• IVES E. 1819. Description of a New Species of Gnaphalium by Professor E. Ives. Amer J Sci (New York) 2(1): 380-381.
• JOHNSTON IM. 1923. Diagnoses and notes relating to the spermatophytes chiefly of North America. Contr Gray Herb 68: 80-104.
• JOHNSTON IM. 1924. Taxonomic records concerning American Spermatophytes. 2. New or otherwise noteworthy plants. Contr Gray Herb 70: 69-86.
• JONES GN. 1938. Flowering plants and ferns of Mount Rainier. Univ Washington Publ Biol 7: 1-192.
• KARTESZ JT. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland, 1 ed. In: Kartesz JT & Meacham CA, Synthesis of the North American Flora: Nomenclatural Innovations, Version 1.0. Chapel Hill, NC, 688 p.
• KIRK T. 1899. The Student’s Flora of New Zealand and the Outlying Islands. J Mackay, Government Printer, Wellington, NZ, 408 p.
• KIRPICHNIKOV ME. 1960. Botanicheskie Materialy Gerbariia Botanicheskogo Instituta imeni VL Komarova Akademii Nauk SSSR, vol 20. Leningrad. Leningrad [St. Petersburg].
• KIRPICHNIKOV ME & KUPRIJANOVA LA. 1950. Morphological-geographical and palynological contributions to the understanding of the genera of the subtribe Gnaphaliinae [In Russian]. Acta Institutum Botanicum VF Komarov Academiae Scientiarum URSS, Ser 1 9: 7-37.
• KLATT FW. 1878. Die Gnaphalien Amerikas. Linnaea 42: 111-144.
• KLATT FW. 1887. Beiträge zur Kenntniss der Compositen. Leopoldina 23: 88-91.
• KUNTH CS. 1818. In: Humboldt A, Bompland A & Kunth CS (Eds). Voyage de Humboldt et Bonpland. Sixième Partie. Botanique. Nova genera et species plantarum. Ed. Folio, Maze, Paris, vol 4, 247 p.
• KUNTH CS. 1820. In: Humboldt A, Bompland A & Kunth CS (Eds). Voyage de Humboldt et Bonpland. Sixième Partie. Botanique. Nova genera et species plantarum. Ed. Quarto, Maze, Paris, vol 4, 312 p.
• LAMARCK JBPA. 1779. Flore françoise ou description succincte de toutes les plantes qui croissent naturellement en France, l’Imprimerie Royale, Paris, vol 2, 684 p.
• LAMARCK JBPA. 1788. Gnaphalium. Encyclopédie Méthodique. Botanique, vol 2(2). Panckoucke, Paris & Plomteux, Liége, p. 739-763.
• LESSING CF. 1831. Synanthereae. De plantis in expeditione speculatoria Romanzoffiana disserere pregunt observatis Ad. de Chamisso et D. de Schlechtendal. Linnaea 6: 83-528.
• LINNAEUS C. 1753. Species plantarum. L. Salvius, Stockholm, 1200 p.
• LOPEZ LAPHITZ RM, MA Y & SEMPLE JC. 2011. A Multivariate Study of Solidago subsect. Junceae and a New Species in South America (Asteraceae: Astereae). Novon 21: 219-225.
• MAHLER WF. 1975. Typification and distribution of the varieties of Gnaphalium helleri Britton (Compositae-Inuleae). Sida 6: 30-32.
• MAPAYA RJ & CRON GV. 2016. A phenetic study of the Emilia coccinea complex (Asteraceae, Senecioneae) in Africa. Plant Syst Evol 302: 703-720.
• MCVAUGH R. 1972. Compositarum Mexicanarum Pugillus. Contr Univ Michigan Herb 9: 459-484.
• MCVAUGH R. 1984. Compositae. In: Anderson WR (Ed), Flora Novo-Galiciana: A Descriptive Account of the Vascular Plants of Western Mexico, The University of Michigan Press, Ann Arbor, vol 12, 1157 p.
• MUNZ PA. 1968. Supplement to a California Flora. Univ. California Press, Berkeley, 224 p.
• MUNZ PA. 1974. A Flora of Southern California. Univ. California Press, Berkeley, 1086 p.
• MUNZ PA & KECK DD. 1959. A California Flora. Univ. California Press, Berkeley, 1681 p.
• NASH DL. 1974. Studies in American plants, VI. Fieldiana, Bot 36: 73-75.
• NELSON A & MACBRIDE JF. 1916. Western plant studies. III. Bot. Gaz. 61: 30-47.
• NELSON EE. 1900. Some new species of Wyoming plants. Bot Gaz 30: 117-122.
• NESOM GL. 1990. Two new species of Gnaphalium (Astercaeae: Inuleae) from the high peaks of northeastern Mexico. Phytologia 68: 413-416.
• NESOM GL. 2001a. Pseudognaphalium austrotexanum (Asteraceae: Gnaphalieae) a new species from southeastern Texas and adjacent Mexico. Sida 19: 507-511.
• NESOM GL. 2001b. Notes on variation in Pseudognaphalium obtusifolium (Aseraceae: Gnaphalieae). Sida 19: 615-619.
• NESOM GL. 2001c. New records in Pseudognaphalium (Gnaphalieae: Asteraceae) for the United States. Sida 19: 1185-1190.
• NESOM GL. 2002. Lectotypification of Pseudognaphalium biolettii (Gnaphalieae: Asteraceae). Sida 20(1): 149-150.
• NESOM GL. 2004. Pseudognaphalium canescens (Asteraceae: Gnaphalieae) and putative relatives in western North America. Sida 21: 781-790.
• NESOM GL. 2006. Pseudognaphalium (Gnaphalieae). In: Flora of North America Editorial Committee (Ed), Flora of North America North of Mexico, New York and Oxford: Oxford University Press 19: 415-425.
• NIE ZL, FUNK VA, MENG Y, DENG T, SUN H & WEN J. 2016. Recent assembly of the global herbaceous flora: Evidence from the paper daisies (Asteraceae: Gnaphalieae). New Phytol 209: 1798-1806.
• NUTTALL T. 1841. Descriptions of new species and genera of plants in natural order of the Compositae, collected in a tour across the continent to the Pacific, a residence in Oregon, and a visit to the Sandwich Islans and Upper California, during the years 1834 and 1835. Trans Amer Philos Soc 7: 283-454.
• NUTTALL T. 1848. Descriptions on plants collected by Mr. William Gambel in the Rocky Mountains and Upper California. Proc Acad Nat Sci Filadelfia 4: 7-26.
• OPIZ FM. 1852. Seznam rostlin květeny české. [List of flora of Bohemia]. Fr. Řivnáč, Praze [Praha], 216 p.
• OSTERHOUT GE. 1906. Colorado Notes. Descriptions of new species. Muhlenbergia 1: 139-143.
• OWEN E, SEMPLE JC & BAUM BR. 2006. A multivariate morphometric analysis of the Symphyotrichum boreale - S. nahanniense - S. welshii complex (Asteraceae: Astereae). Canad J Bot 84: 1282-1297.
• PRUSKI JF. 2018. Pseudognaphalium. In: Davidse G, Sousa Sánchez M, Knapp S & Chiang Cabrera F (Eds), Fl. Mesoamer., Asteraceae, Missouri Botanical Garden, St. Louis 5(2): 222-231.
• RADFORD AE. 1986. Fundamentals of plant systematics. Harper and Row, New York, XIII+ 498 p.
• REICHENBACH HGL. 1829. Handbuch der Gewächskunde. In: Mossler JC & Reichenbach HGL, Hammerich, Altona, 2 ed, vol 2, 6318 p.
• RÉMY EJ. 1849. Compuestas. In: C. GAY (Ed), Historia física y política de Chile según documentos adquiridos en esta República durante doce años de residencia en ella. Botánica, Privately Published, Paris, vol 4, p. 5-317.
• REVEAL JL. 1998. Typification of Linnaean specific and varietal names in the Compositae (Asteraceae). In: Jarvis CE & Turland NJ (Eds). Taxon 47: 347-370.
• ROBBIATI FO, ANTON AM & FORTUNATO RH. 2014. A New Synonym in Senna, Series Aphyllae (Leguminosae, Caesalpinioideae). Syst Bot 39 (4): 1120-1126.
• RYDBERG PA. 1900. Catalogue of the Flora of Montana and Yellowstone National Park. Mem New York Bot Gard 1: 1-492.
• SCHULTZ BIPONTINUS CH. 1845. Verzeichniss der von Claussen in der Brasilianischen Provinz Minas Geraes gesammelten Compositen, welche von Hohenacker herausgegeben worden. Botanische Zeitung (Berlin) 3: 171-172.
• SCHULTZ BIPONTINUS CH. 1856. Lechler’s neueste Sammlungen aus Peru und Chile. Bonplandia 4: 50-54.
• SELL PD. 2006. Campanulaceae-Asteraceae. Flora of Great Britain and Ireland, vol 4. Cambridge University Press, Cambridge, United Kingdom, 652 p.
• SMISSEN RD, GALBANY-CASALS M & BREITWIESER I. 2011. Ancient allopolyploidy in the everlasting daisies (Asteraceae: Gnaphalieae): complex relationships among extant clades. Taxon 60: 649-662.
• SNEATH PHA & SOKAL RR. 1973. Numerical taxonomy. The principles and practice of numerical classification. Freeman WH and company press, San Francisco, XV+, 573 p.
• SPRENGEL K PJ. 1826. Systema Vegetabilium, ed. 16 [Caroli Linnaei ... Systema vegetabilium editio decima sexta], vol 3. Classis 16-23. Sumtibus Librariae Dieterichianae, Gottingae [Göttingen], 936 p.
• STAFLEU FA & COWAN RS. 1976. Taxonomic literature, ed 2, vol 1: A-G. Bohn, Scheltema and Holkema, Utrecht, 1136 p.
• STAFLEU FA & COWAN RS. 1981. Taxonomic literature. Lh-O. Bohn, Scheltema and Holkema, Utrecht, 2 ed, vol 3, 980 p.
• STEBBINS GL. 1993. Pseudognaphalium. In: Hickman JC (Ed), The Jepson manual higher plants of California. University California Press, Berkeley, p. 270-271.
• STEBBINS GL & KEIL DJ. 1992. New combinations in Gnaphalium (Asteraceae: Inuleae). Novon 2: 347.
• THIERS B. 2020 (continuously updated). Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/: 10.07.2020
  » http://sweetgum.nybg.org/ih/: 10.07.2020
• THUNBERG CP. 1800. Prodromus Plantarum Capensium, quas, in Promontorio Bonæ Spei Africes, annis 1772-1775 collegit Carol Pet Thunberg, vol 2. Edman, Uppsala, Sweden, p. 85-191.
• TURLAND NJ. 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code). Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten.
• TZVELEV NN. 1993. Notes on some Caucasian Asteraceae and Araceae. Byull Moskovsk Obshch Isp Prir Otd Biol 98: 99-108.
• VILLARREAL-QUINTANILLA JA, ESTRADA-CASTILLÓN AE & ENCINA-DOMÍNGUEZ JA. 2020. Dos cambios de rango taxonómico en Pseudognaphalium (Gnaphalieae, Asteraceae) de México. Acta Bot Mex 127: 1-16. DOI 10.21829/abm127.2020.1582.
• VILLASEÑOR JL. 2016. Checklist of the native vascular plants of Mexico. Revista Mex Biodivers 87: 559-902.
• WARD JM. 1993. Systematics of New Zealand Inuleae (Compositae-Asteraceae)—2 A numerical phenetic study of Raoulia in relation to allied genera. New Zeal J Bot 31: 29-42.
• WARD JM, BAYER RJ, BREITWIESER I, SMISSEN R, GALBANY-CASALS M & UNWIN M. 2009. Gnaphalieae. In: Funk VA, Susanna A, Stuessy TF & Bayer RJ (Eds), Systematics, evolution, and biogeography of Compositae. International Association for Plant Taxonomy (IAPT), Vienna, Austria, p. 539-588.
• WEATHERBY CA. 1923. Some critical plants of Atlantic North America. Rhodora 25: 17-23.
• WEBB PB. 1849. Spicilegia Gorgonea; or a catalogue of all the plants as yet discovered in the Cape de Verd Islands, from the collections of J.D. Hooker, Esq. M.D.R.N., Dr. T. Vogel, and other travellers. In: Hooker WJ (Ed), Niger Flora or, an enumeration of the plants of western tropical African, collected by the late Dr. Theodore Vogel, botanist to the voyage of the expedition sent by Her Britannic Majesty to the River Niger in 1841, under the command of Capt. H.D. Trotter, R.N. etc. Hippolyte Bailliere, London, p. 89-197.

Publication Dates

• Publication in this collection
  25 Nov 2022
• Date of issue
  2022

History

• Received
  06 Feb 2020
• Accepted
  09 Sept 2020