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A synopsis of Chaenotheca in North America: including a new species from southern Ontario, C. selvae, supported by morphometric analyses

Journal: Botany

Manuscript ID cjb-2018-0042.R1

Manuscript Type: Article

Date Submitted by the Author: 10-Apr-2018

Complete List of Authors: McMullin, Richard; Canadian Museum of Nature, Research and Collections Maloles, Jose ; Credit Valley Conservation Selva , StevenDraft ; University of Maine, Biology & Environmental Studies Newmaster, Steven; University of Guelph

Keyword: Calicioid, Caliciales, Coniocybaceae, Great Lakes region, Old-growth forest

Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? :

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A synopsis of Chaenotheca in North America: including a new species from southern Ontario, C. selvae, supported by morphometric analyses

Richard Troy McMullin1, Jose R. Maloles2, Steven B. Selva3, and Steven G. Newmaster2

1Canadian Museum of Nature, Research and Collections, Ottawa, ON, K1P 6P4, 2Biodiversity Institute of Ontario, University of Guelph, 50 Stone Road East, Guelph, ON, N1G 2W1 3University of Maine at Fort Kent, 23 University Drive, Fort Kent, Maine, 04743 *Corresponding author: tmcmullin@musnature.ca

Abstract

We describe and illustrate a new species of calicioid lichen from southern Ontario, Chaenotheca

selvae. We place it in Chaenotheca because of its lichenized thallus with a Stichococcus photobiont, a

brown mazaedium, and ascospores that are brown, globose, and singlecelled (3.03.6 m in diam.). It

differs from other species of Chaenotheca by its photobiont, nonornamented ascospores, straight stalks, and producing orangebrown pruina on theDraft mazaedium, excipulum, and upper portion of the stalk when mature. Using a detrended correspondence analysis we identified 25 taxa, including the sp. novum,

analyzing 22 morphological characters. Our results provide evidence of chemical and morphometric

variation among species of Chaenotheca. We also provide a key for the identification of the 25

Chaenotheca species in North America.

Keywords: Calicioid, caliciales, Coniocybaceae, Great Lakes region, oldgrowth forest.

Introduction

Chaenotheca Th.Fr. is a charismatic calicioid genus that is widespread in North America (Tibell

1999; Selva 2014). Species typically occur in mature woodlands and are usually corticolous or lignicolous

(Tibell 1992; Selva 2003; McMullin et al. 2008). Chaenotheca differs from other calicioid genera by

being lichenized, having round singlecelled ascospores, and by producing ascospores in a brown

maezedium (Tibell 1980, 1999; Selva and Tibell 1999). It is also unusual among calicioid genera, and

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other lichen genera, because four photobionts are associated with different species (Dictyochloropsis,

Stichococcus, Trebouxia, and Trentepohlia) (Tibell 1975, 1999; Selva 2014).

The first comprehensive contemporary identification key for Chaenotheca in North America was produced for the boreal region by Tibell (1975). It included 14 species: C. brachypoda (Ach.) Tibell (as

Coniocybe sulphurea (Retz.) Nyl.), C. brunneola (Ach.) Müll. Arg., C. chlorella (Ach.) Müll. Arg. (as C. carthusiae (Harm.) Lettau), C. chrysocephala (Ach.) Th.Fr., C. ferruginea (Turner ex Sm.) Mig., C. furfuracea (L.) Tibell (as Coniocybe furfuracea (L.) Ach.), C. gracillima (Vain.) Tibell (as Coniocybe gracillima Vain.), C. hispidula (Ach.) Zahlbr., C. laevigata Nádv., C. phaeocephala (Turner) Th.Fr., C. stemonea (Ach.) Müll. Arg., C. subroscida (Eitner) Zahlbr., C. trichialis (Ach.) Th.Fr., C. xyloxena Nádv.

Tibell (1980) then added C. cinerea (Pers.) Tibell from northern Ontario and Chaenotheca gracilenta

(Ach.) Mattsson & Middelb. (as Cybebe gracilentaDraft (Ach.) Tibell) from the Lake Superior region of the

United States (Tibell 1984). Harris (1995) was the next to contribute to this list when he described C. floridana R.C.Harris from Florida. Rikkenin (1998) then discovered C. olivaceorufa Vain. in the Pacific

Northwest. Selva and Tibell (1999) increased the number to 20 when they reported C. hygrophila Tibell and C. servitii Nádv. from the northeast and C. sphaerocephala Nádv. From British Columbia. Tibell and

Koffman (2002) then described C. nitidula Tibell from Maine and New Brunswick. Selva (2013) also described Chaenotheca erkahomattiorum Selva from the northeast. Most recently, a widespread species in

North America was described by Allen and McMullin (2015), C. balsamconensis J.L.Allen & McMullin.

Chaenotheca cf. autralis Tibell was also reported from California by Williams and Sillett (2007), but due to the uncertainty of the identification we are not including it on our North American list.

The large number of new species and new North American records of Chaenotheca in recent

decades have been reported in numerous publications (e.g., Harris 1995; Rikkenin 1998; Selva and Tibell

1999; Tibell and Koffman 2002; Selva 2013; Allen and McMullin 2015). Selva (2014) developed a key to

many of these species (19 of the 25), those occurring in the Acadian Forest region of northeastern North

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America, and McCune (2017) developed a key to the 21 species in the Pacific Northwest. Here, we

provide a comprehensive key to all species in North America and describe a new species of Chaenotheca

from southern Ontario.

Methods

We examined specimens using standard microscopy techniques following Brodo et al. (2001).

We captured images of the ascomata with a Leica DVM6 digital microscope (Fig. 1A). Using a FEI

Inspect S50 scanning electron microscope (SEM) equipped with a tungsten filament we captured images

of the ascospores (Fig. 1C). We prepared specimens for the SEM using a stereomicroscope to remove

ascomata and place them onto an aluminumDraft stub using doublesided carbon tape. We placed the stub in a

Cressington Auto/SE sputter coater equipped with a gold target and deposited a thin gold film (~ 20 nm)

onto the stub before imaging.

Morphometric Analyses

We recorded 22 morphological variables (Table 1) from 25 species of Chaenotheca using

specimens at three herbaria (CANL, OAC, and UMFK) (Table 1). We used 49 specimens to generate

measurements for 19 species (means were calculated from 10 measurements). For species that we did not

have specimens we used previously published measurements (i.e., C. floridana, C. hispidula, C.

hygrophila, C. olivaceorufa, C. phaeocephala, and C. subroscida) (Tibell 1980, 1999; Rikkinen 1998).

We used a matrix of 25 species and 22 morphological characters in a multivariate analysis. Using

canonical ordination we detected groups of specimens and estimated the contribution of each variable to

the ordination. We used a principal component analysis (PCA) (ter Braak and Smilauer 1998) to identify

the length of the ordination axis and the need for either a linear or unimodal ordination technique. Using

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unimodal, indirect ordination detrended correspondence analysis (DCA) we explored variation in species scores in this study. We used a cluster analysis to classify the specimens, as it is better in representing distances among similar specimens, whereas DCA is better in representing distances among groups of specimens (Sneath and Sokal 1973). Our cluster analysis was performed with NTSYS (Rohlf 2000).

Using a distance matrix we generated an arithmetic average (UPGMA) clustering algorithm and standardized data based on average taxonomic distance subjected to the unweighted pairgroup method.

We used Pearson correlation analysis to assess the correlation among the 22 morphological characters and the respective DCA axes (SPSS 2014).

Results and Discussion

Draft

Morphometric Analyses

The ordination analyses utilized DCA in the separation of 25 species of Chaenotheca that were analyzed and provided a measure of the important morphological variables in the classification. Principal components analysis (PCA) provided a character gradient that was unimodal (2.89 Standard Deviations

SD) violating the assumption of a linear model (ter Braak and Smilauer 1998). Consequently, a DCA was used to classify the Chaenotheca specimens into distinct groups representing 25 species. Examination of

the eigenvalues indicated that the first axis (0.363) and second axis (0.607) were of importance in

assessing variation among all the specimens. The first axis (DCA 1) is strongly correlated with 10

characters: asci arrangement catenulate, asci arrangement single, mean stalk height >1 mm, mean spore

diameter over 5.5 m, phycobiont Chlorococcales, spores smooth, stalk pruinose, stalk with yellow pruina, lignicolous only, and thallus usually episubstratal (Table 1). The second axis (DCA 2) is strongly

correlated with 8 characters: phycobiont Stichococcus, phycobiont trebouxioid, stalk pruinose, Stalk with

a KOH+ red precipitate, stalk with yellow pruina, substrate lignum only, substrate Trichaptum abietinum,

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and thallus usually episubstratal (Table 1). The DCA ordination of the first two canonical functions

identifies a distinct cluster of 2 specimens that represent the new species, C. selvae, in the proximity of

the respective allied species, C. brachypoda (Fig. 3). This study provides evidence of morphometric

variation among species of Chaenotheca, which allows further development of hypotheses concerning

phenetic species concepts.

Taxonomy

Chaenotheca selvae Maloles & McMullin, sp. nov.

Figs. 1-2

Mycobank #825505 Draft

Type: CANADA. ONTARIO: Wellington County, Guelph, University of Guelph, The Arboretum, Wild

Goose Woods, c. 100 m SE of the Wild Goose Woods Trail, 43.5368o, 80.2064o, 18 May 2011, R.T.

McMullin 7752. (holotype: CANL; isotype: OAC).

Description

Thallus immersed. Photobiont Stichococcus. Stalk smooth, dark redbrown to black, 0.31.2 mm

× 0.050.15 mm (n=15) (Fig. 1A). Mazaedium brown (Fig. 1A). Excipulum and mazaedium with orange

brown pruina that often extends down the top third of the stalk on mature apothecia (Fig. 1B). Pruina is

violetred with the addition of KOH. Paraphyses sparsely branched. Asci cylindrical to narrowly clavate,

1823 m × 3.84.5 m (n=15). Ascospores globose with a depression in the centre (appearing round

under a compound microscope), pale brown, 3.03.6 m in diameter (n=15), and lack ornamentation (Fig.

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1B & C). On decorticated deciduous stumps in undisturbed oldgrowth treedwetlands with standing water most of the year (Fig. 2A & B).

Etymology

The epithet “selvae” recognizes Dr. Steven Selva and his important contributions to the study of calicioid lichens and fungi in North America over the last 30 years.

Diagnosis

Draft

Our morphometric analysis shows that Chaenotheca selvae is a distinct species. Chaenotheca

brachypoda is most similar in the analysis, but that species has yellow pruina on the capitulum instead of

orangebrown. Chaenotheca gracillima is the only other species in the genus with somewhat similar

coloured pruina (redbrown vs. orangebrown in C. selvae), but its stalks are >1 mm tall (vs. <1 mm) and

flexuous (vs. straight). Species in the genus Sclerophora may also appear similar, but the photobiont of

that genus is Trentepohlia, instead of Stichococcus and it typically has paler stalks. Chaenotheca selvae

differs from its other congeners by its photobiont, nonflexuous stalks, and producing orangebrown pruina on the capitulum and upper portion of the stalk when mature.

Ecology and Distribution

Chaenotheca selvae is currently only known to colonize the hard lignum of deciduous tree stumps

in mature or oldgrowth treed wetlands with standing water for at least part of the year. Its current

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distribution is limited to two wetlands (The Nature Reserve and Wild Goose Woods), which have little or

no known history of timber harvesting and are approximately 1 km apart in The Arboretum at the

University of Guelph in southwestern Ontario. Both forests contain a variety of tree species, but, in The

Nature Reserve, the area that we located C. selvae is dominated by Thuja occidentalis, while it in the

Wild Goose Woods the area around the collection is dominated by Acer. It was initially located in Wild

Goose Woods. We then completed targeted searches in similar habitats in the region around Guelph (Fig.

1F) and only rediscovered it again in The Nature Reserve. This region has been heavily impacted by

agriculture, settlement, and historical timber harvesting. Few wetlands left in the region are associated

with oldgrowth forests, which may be the cause for its paucity. However, the region around Guelph has

been poorly studied by lichenologists (McMullin et al. 2014) and calicioids in particular are often

overlooked and understudied, which might be why this species has gone undetected. Guelph is within the

transition between two forest zones, CarolinianDraft and the Great LakesSt. Lawrence (Rowe 1972). In either

of these zones, further exploration for C. selvae should target treed wetlands with standing water much of

the year that contain mature or oldgrowth forests.

Additional Specimens Examined

CANADA. ONTARIO: Wellington Co., Guelph, University of Guelph, The Arboretum, The Nature

Reserve, lignicolous on a deciduous tree stump, 43.53708o, 80.20867o, August 2010, J.R. Maloles 79

(OAC).

Key to the 25 North American species of Chaenotheca

1. Apothecia with yellow, yellowgreen, red, reddishbrown, brownishorange, or brown pruina on excipulum, stalk, or both excipulum and stalk ………………...... ………………. 2 1. Apothecia with white pruina on excipulum, stalk, or both excipulum and stalk, or epruinose ………………………………………………...... ……………..…………………….. 13

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2. Thallus with Trentepohlia as photobiont; pruina yellow or brownish; most spores spherical…...…… 3 2. Thallus with Stichococcus or trebouxioid photobiont; pruina and spore shape variable………………. 4

3. Mature spores 3.0–4.5 m in diameter, with smooth surfaces that may have a few irregular cracks ……………………...... ………………………...……… C. olivaceorufa 3. Mature spores 4.5–7.5 m in diameter, with irregular reticulate cracks surrounding polygonal areas ……………………...……………………………...... ……………………. C. hispidula

4. Photobiont with spherical cells measuring 7.0–14 m in diameter, i.e., trebouxioid ...... 5 4. Photobiont with short, cylindrical cells, which sometimes form short filaments, or with small, irregularlyshaped cells in colonies (Stichococcus) ……...... ………... 8

5. Spores spherical, 6.0–7.0 m in diameter …………………………………...... …………. 6 5. Spores, at least in part, ellipsoidal to cylindrical ...... 7

6. Thallus minutely granular, white, diffuse and/or with flat greenishbrown verrucae at the base of apothecia ……………………………………...... …...... ……………….... C. subroscida 6. Thallus squamulose to verrucose, greyish to greenishbrown …...... …… C. phaeocephala

7. Thallus bright yellow, usually welldeveloped,Draft granular to verrucose; spores spherical to short ellipsoidal, typically <8.0 m long …………...... ………………… C. chrysocephala 7. Thallus immersed or squamulose to verrucose, greyishgreen; spores nonseptate and ellipsoidal, or cylindrical, septate, and typically >8.0 m long (and as long as 34 m)...... … C. laevigata

8. Apothecia with red, reddishbrown, or orangebrown pruina ……………...... …………….. 9 8. Apothecia with yellow or yellowgreen pruina ……………………………...... ………….. 10

9. Pruina red to reddishbrown; apothecia flexuous, typically >1.0 mm tall …...... ….… C. gracillima 9. Pruina orangebrown; apothecia with straight stalks, typically <1.0 mm tall …...... …… C. selvae

10. Thallus episubstratic, farinose, bright yellowgreen …………….…...... ………. C. furfuracea 10. Thallus immersed or episubstratic, but not both farinose and bright yellowgreen …...... 11

11. Apothecia typically >1.5 mm tall, with thin, flexuous stalks …………...... ……..… C. servitii 11. Apothecia typically <1.5 mm tall, with straight stalks ………………...... ………….…….. 12

12. Spores spherical or squarrish, 3.0–4.0 m in diameter; thallus immersed or thin, green and poorly developed ……………………………………………………...... …..….… C. brachypoda 12. Spores shortellipsoidal, 4.0–10 × 3.0–4.5 m; thallus welldeveloped, green to greygreen, granular to verrucose ……………………………………...... ……….……….. C. chlorella

13. Photobiont with short, cylindrical cells, which sometimes form short filaments, or with small, irregularlyshaped cells in colonies (Stichococcus) ..………...... ……….. 14 13. Photobiont with spherical cells, either Dictyochloropsis (cells 15–20 m in diameter) or trebouxioid (cells 7.0–14 m in diameter) ………………………………...... ………..… 19

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14. Thallus thin, farinose; capitulum spherical, with poorlydeveloped excipulum …...... ……. 15 14. Thallus immersed or welldeveloped, but never farinose; capitulum obconical to lenticular, with well developed excipulum ……………………………………...... ……………….. 16

15. Apothecia typically <1.5 mm tall and with straight stalks; bottom of capitulum pale brown, which contrasts with the darker brown mazaedium, the consequence of having spores caught in what is left of the excipulum; thallus glaucousgreen ………...... …. C. stemonea 15. Apothecia typically >1.5 mm tall and with thin, flexuous stalks; bottom of capitulum the same color as the mazaedium; thallus greyishgreen ……………...... ……… C. gracilenta

16. Apothecia epruinose; tall and with flexuous stalks; conidiophores typically present…….….. C. nitidula 16. Apothecia whitepruinose; stalk not flexuous; conidiophores absent …………...... ………... 17

17. Stalk of apothecia pale at base …………………………………………...... ……….. C. cinerea 17. Stalk of apothecia not pale at base ……………………………………………...... ………... 18

18. Thallus immersed; excipulum, upper part of stalk (and often into the lower part) with thick white pruina; base of excipulum and uppermost part of stalk with anticlinallyarranged hyphae extending outward from surface; lignicolous ……………...... ………...Draft C. xyloxena 18. Thallus rarely immersed, typically welldeveloped, greyishgreen; excipulum and, rarely, the upper part of stalk, whitepruinose, though not thickly so, rarely epruinose; base of excipulum and uppermost part of stalk with periclinallyarranged hyphae, K+ red precipitate absent; corticolous, lignicolous, or growing on the small polyporous fungus Trichaptum abietinum.…………...…...….. C. trichialis

19. Growing on the small polyporous shelf fungus Trichaptum abietinum, on dead gymnosperms; thallus usually immersed; stalk mostly epruinose, K± red precipitate; mature spores smooth...……………………………………………………...... ……….. C. balsamconensis 19. Growing on bark or wood of angiosperms and gymnosperms; thallus immersed or not; spores smooth or ornamented ………………………………………………………..……...... ………… 20

20. Thallus typically welldeveloped, white to greenishwhite, and with scattered orangy patches or granules that are K+ red; excipulum welldeveloped, often with gold flecks in mazaedium; spores spherical and mostly >6.0 m in diameter ………...... ….. C. ferruginea 20. Thallus immersed or welldeveloped, but without orangy patches or granules that are K+ red; excipulum poorly or welldeveloped, without gold flecks in mazaedium; spores, if spherical, <6.0 m in diameter …...... 21

21. Thallus typically immersed, rarely episubstratic, grey to greenishgrey and granular; photobiont Dictyochloropsis, which is conspicuously wrapped in multiple layers of hyphae; stalk black, shiny ……………………………………...... ……….. C. brunneola

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21. Thallus typically welldeveloped; photobiont trebouxioid, which may or may not be wrapped in multiple layers of hyphae; stalk dull or appearing thinly white to greyish pruinose….…………….……. 22

22. Excipulum poorlydeveloped; asci typically attached endtoend, in chains, and of irregular shapes (search under dim light) .………………………...... ………….. C. sphaerocephala 22. Excipulum typically welldeveloped; asci cylindrical or of irregular shapes, produced singly .……………...………………………………………………………...... ……….…….. 23

23. Thallus minutely granular, greenishgrey …………………………...... ……….. C. hygrophila 23. Thallus leprose to granularleprose, whitish …..…………………...... …………………. 24

24. Spores of two distinct shapes—spherical (4.5–6.0 m in diameter) and ellipsoidal to shortclindrical, simple to septate (most >5.0 m long); thallus leprose, PD+ deep yellow to orange (barbatic and obtusatic acids) ……………………...... ….….. C. erkahomattiorum 24. Spores spherical to slightly angular, 3.0–4.5 m in diameter; thallus granularleprose, PD+ redorange (virensic acid) …..…………………..…...... ………………. C. floridana

Acknowledgements We gratefully acknowledge ChrisDraft Earley for habitat images, Arboretum staff for providing permission for us to conduct our study, and two anonymous reviewers for helpful

comments on the manuscript.

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Tables

TABLE 1. DCA analysis of 22 quantitativeDraft variables (taxonomic characters) for 49 specimens (classification of 25 Chaenotheca species). Pearson correlations indicate the characters significant to the classification (** = p value < 0.01, * = p value < 0.05). DCA axis X DCA axis Y Characters P Corr. Sig. (2 P Corr. Sig. (2 tailed) tailed) Asci arrangement catenulate 0.667(**) 0.001 0.019 0.932 Asci arrangement single 0.516(*) 0.014 0.027 0.904 Excipulum developed 0.401 0.072 0.162 0.484 Mean spore diameter over 5.5 m 0.683(**) 0 0.085 0.699 Mean stalk height >1mm 0.41(*) 0.042 0.257 0.216 Phycobiont Chlorococcales 0.568(**) 0.003 0.16 0.446 Phycobiont Dictyochloropsis 0.373 0.066 0.085 0.685 Phycobiont Stichococcus 0.383 0.059 0.558(**) 0.004 Phycobiont trebouxioid 0.21 0.314 0.557(**) 0.004 Phycobiont Trentepohlia 0.047 0.824 0.208 0.319 Pruina on entire stalk & capitulum 0.278 0.178 0.035 0.867 Spores ornamented 0.273 0.197 0.059 0.783 Spores smooth 0.838(**) 0 0.126 0.559 Stalk pruinose 0.552(**) 0.004 0.531(**) 0.006 Stalk with a KOH+ red precipitate 0.264 0.203 0.407(*) 0.044 Stalk with white pruina 0.106 0.613 0.179 0.391 Stalk with yellow pruina 0.641(**) 0.001 0.434(*) 0.030 Substrate lignum only 0.61(**) 0.003 0.549(*) 0.010 Substrate Trichaptum abeitinum 0.264 0.203 0.407(*) 0.044 Thallus KOH+ 0.195 0.361 0.285 0.177

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Thallus PD+ 0.117 0.585 0.53(**) 0.008 Thallus usually episubstratal 0.552(**) 0.004 0.248 0.232

Figures Draft Figure 1. (A-D) Chaenotheca selvae. (A) Apothecia, Maloles 79 (OAC), scale = 0.5 mm. (B) Ascospores under a compound microscope at 1000x, McMullin 7725 (CANL), scale = 13 µm. C) Ascospores under a scanning electron microscope, McMullin 7725 (CANL), scale = 7 µm. (D) Distribution, blue = present, red = absent in target searched localities.

Figure 2. (A-B) Chaenotheca selvae habitat. Images are from the same location, but facing opposite directions. Yellow arrows indicate the location of C. selvae at this site in the Arboretum at the University of Guelph, McMullin 7752 (CANL).

Figure 3. Scatter plot of the first two axes from a detrended correspondence analysis (DCA) for 22 quantitative variables (taxonomic characters) of 49 specimens (classification of 25 Chaenotheca species). The new species, Chaenotheca selvae, is bolded and highlighted.

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Draft

Figure 1. (A-D) Chaenotheca selvae. (A) Apothecia, Maloles 79 (OAC), scale = 0.5 mm. (B) Ascospores under a compound microscope at 1000x, McMullin 7725 (CANL), scale = 13 µm. C) Ascospores under a scanning electron microscope, McMullin 7725 (CANL), scale = 7 µm. (D) Distribution, blue = present, red = absent in target searched localities.

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1.5 C. balsamconensis C. hygrophila 1 C. ferruginea C. C. C. floridana sphaerocephala 0.5 erkahomattioru C. stemonea m C. brunneola C. cinerea C. laevigata C. olivaceorufa 0 C. subroscida C. trichialis C. chrysocephala C. gracillima DCA 2 DCA C. phaeocephala C. servitii C. nitidula C. gracilenta C. xyloxena -0.5 C. chlorella C. furfuracea

C. hispidula -1 C. brachypoda C. selvae

-1.5 -1.5 -1 -0.5Draft 0 0.5 1 1.5 2 DCA 1

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