Species richness and distribution of understorey bryophytes in

Journal of Bryology (2006) 28: 182–189

Species richness and distribution of understorey bryophytes in different forest types in Colombian Amazonia JUAN C. BENAVIDES1, ALVARO J. DUQUE M.2, JOOST F. DUIVENVOORDEN3 and ANTOINE M. CLEEF3 1

Universidad de Puerto Rico, 2Universidad Nacional de Colombia, Sede Medellıń, Colombia and 3Institute for Biodiversity and Ecosystem Dynamics, Universiteit van Amsterdam, The Netherlands

SUMMARY The first bryophyte survey results from Colombian Amazonia are presented. Bryophyte species, differentiated into mosses and liverworts, and further into four life-form classes, were sampled in 0.1-ha plots. These plots were distributed over four landscape units in the middle Caqueta´ area: floodplains, swamps, terra firme forests and white-sand areas. The total numbers of bryophyte species in the units were 50, 45, 45 and 32, respectively. The plots in floodplains and swamps were richer in moss species than the terra firme and white-sand plots, suggesting that coexistence of many moss species is favoured by high humidity. Moss species with fan lifeforms preferred floodplains. On the other hand, liverwort species richness was highest in white-sand plots, which suggests that light incidence controls liverwort species-richness, perhaps more than humidity. All plots from the floodplain of the Caqueta´ River differed remarkably in species composition (of both mosses and liverworts) from the other landscape units. This may be due to the unique properties of this varzea system where, during yearly flooding events, soil, dead logs and stems are covered with a fresh layer of nutrient-rich fine silt, enhancing the surface for colonization and improving the conditions for productive bryophyte growth compared with elsewhere in the middle Caqueta´ area. KEYWORDS: Amazon forests, bryophytes, growth forms, landscape units, species richness.

INTRODUCTION Bryophytes are important components of tropical forest ecosystems because they favour nutrient trapping (Nadkarni, 1986; Matzek & Vitousek, 2003) and regulate hydrological cycles (Veneklaas & van Ek, 1990). There is an ongoing debate about how the availability and longevity of substrata might affect local species richness of bryophytes. Contrasting views range from a positive relationship between species richness and time since disturbance, motivated by species coexistence in the absence of strong competition (Rydin, 1997; Kimmerer & Driscoll, 2000; Pharo & Beattie, 2002), to a negative relationship, motivated by favourability of intermediate-level disturbances for effective colonization and coexistence of species (Lu¨cking & Lu¨cking, 2002). Bryophyte species usually have wide geographical distributions, due mainly to the small size of the propagules allowing them to bridge large dispersal distances, and to their old age (Schofield, 1992). Air humidity, temperature, daily variation of UV radiation # British Bryological Society 2006 DOI: 10.1179/174328206X120040

and substratum structure are often considered as the main ecological factors determining the local patterns in species composition of tropical bryophytes (Richards, 1984; Florschu¨tz-de Waard & Bekker, 1987; Wolf, 1993). High sensitivity to microclimatic conditions (Richards, 1984) and dependence on nutrient acquisition from the substratum (Bates, 2000) seemingly bring about different floristic distributions at local and regional scales as compared with vascular plants (During, 1992; Kessler, 2000). Here, we present the first survey results of understorey bryophytes in upper Amazonia. Bryophyte species, differentiated into mosses and liverworts, and further into four life-form classes, were sampled in mature forests along a broad physiographic gradient related to humidity (seasonal flooding or stagnant water), and forest structure (forest height and related understorey light conditions) in four landscape units recognized in the middle part of the basin of the Caqueta´ River (Fig. 1). Bryophyte sampling was restricted to understorey habitats in order to avoid secondary gradients related to increasing forest height Received 27 June 2005. Revision accepted 30 March 2006

UNDERSTOREY BRYOPHYTES IN COLOMBIAN AMAZONIA

183

occurred mostly in well-drained terra firme and floodplain areas.

Fieldwork and species identification

Figure 1. Fieldwork area showing the location of the Caqueta´ and Mesay areas in Colombian Amazonia where the 0.1-ha plots were surveyed.

(van Dunne´, 2001). We hypothesized that between-landscape differences in the species richness and distribution of bryophyte species would be caused by landscape differences in humidity and forest height.

METHODS Study area The study area covers about 2000 km2 and is situated along the stretches of the middle Caqueta´ and Mesay Rivers in Colombian Amazonia, roughly between 72u37’ and 71u 18’W, and 0u55’S and 0u9’N, at an altitude of 200–300 m a.s.l. (Fig. 1). The mean annual precipitation in the study area is approximately 3060 mm (1979–1990), and the mean annual air temperature is 25.7uC (1980–1989; Duivenvoorden & Lips, 1993). The principal landscape units are well-drained floodplains, swamps (including permanently inundated backswamps and basins in floodplains or fluvial terraces), areas covered with white-sand soils (found on high terraces of the Caqueta´ River and in less-dissected parts of the Tertiary sedimentary plain), and terra firme (which is never flooded by river water and includes low and high fluvial terraces and a Tertiary sedimentary plain) (Duivenvoorden & Lips, 1993, 1995; Lips & Duivenvoorden, 1996; Botero, 1999). Duque et al. (2001) (see also Duivenvoorden & Lips, 1993) reported that forests from swamps and white-sand areas were mainly characterized by a relatively high density of low trees with small crowns while less-dense forests with taller trees

We conducted a survey of 40 0.1-ha plots that were located in the four landscape units mentioned above. In order to establish the plots, starting locations along the Caqueta´, Mesay and Cunãre rivers, and the direction of the tracks along which the forests were entered, were planned on the basis of the interpretation of aerial photographs and satellite images (Duivenvoorden et al., 2001). Along each track, the geomorphology was examined to identify more or less homogeneous terrain units. In these mature forest units, rectangular plots were delimited by compass, tape and stakes in areas that did not show signs of human intervention, and avoiding recent gaps of large trees. All plots were mapped by GPS and were located at a minimum distance of 500 m from each other. In each plot, bryophyte colonies were sampled at ten or more randomly located points. At each point, all bryophyte species which occurred on the forest floor (including mineral soil and litter), dead logs or tree trunks (including palms), shrubs, lianas, or herbs, were sampled within a circle with a horizontal radius of approximately 2 m and vertically up to 1.5 m height. When species showed low local abundances at a sampling point, bulk samples were taken to allow for the separation of individuals from different species afterwards. The species sampling for the entire plot may not have been exhaustive. Epiphylls were not included. Four different life-forms were recognized: fans (including pendants), rough mats, smooth mats and turfs (Ma¨gdefrau, 1982; Bates, 1998). Species were identified using Gradstein & da Costa (2003) and Churchill (1994), and with the help of specialists. Collections were deposited at the HUA herbarium. Nomenclature and taxonomic arrangement was according to Uribe & Gradstein (1998) for liverworts and Crosby, Magill & Bauer (1992) for mosses. The elevational range of the species was defined according to recent checklists (Uribe & Gradstein, 1998; Churchill, Griffin & Munõz, 2000; Churchill, 2003; Gradstein & da Costa, 2003). Species reported exclusively from neotropical lowlands below 800 m were labelled ‘lowland’.

Data analysis Presence–absence data of bryophyte species in the 40 0.1-ha plots were analysed. Species richness was analysed by species accumulation curves for mosses and liverworts, randomizing plot orders ten times (Colwell & Coddington, 1994). Between-landscape species richness of mosses and liverworts was analysed by one-way analysis of variance (ANOVA). Post-hoc analyses were made with Tukey’s honestly significant difference test (Sokal & Rohlf, 1995).

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Table 2. Life-form frequency of species of mosses (x 2 540.1, p,0.001) and liverworts (x 2 56.3, p50.39) arranged according to four landscape units in Colombian Amazonia. Life form Mosses Fan Mat Rough mat Turf Liverworts Fan Mat Rough mat

Floodplain

Swamp

Terra firme

White sand

16 14 12 22

5 18 5 29

0 27 5 60

1 4 1 10

10 39 7

9 49 3

27 92 20

14 38 9

RESULTS

Figure 2. Species-accumulation curves for bryophytes, mosses and liverworts in the four landscape units.

Between-landscape differences in species richness ratios, which showed a strongly skewed distribution even after arcsin transformation, were examined with a Kruskal– Wallis test. A contingency table analysis with a x2 test was done to determine whether the number of species of different life-forms was associated with the landscape units. Patterns in species composition of understorey bryophyte, mosses, and liverworts were explored by detrended correspondence analysis (DCA; Hill & Gauch, 1980; ter Brak, 1987) utilizing presence–absence data. A one-way ANOVA followed by the above-mentioned Tukey’s test (Sokal & Rohlf, 1995) of the plot values along the first ordination axis was performed to examine how plots segregated between the landscape units. For each ANOVA homoscedasticity of variances and normality of residuals were checked. Analyses were done in SPSS for Windows (Release 11.0.1, SPSS Inc., 2001), apart from the DCA, which was done in in CANOCO version 4 (ter Braak & Smilauer, 1998).

In total, 37 species of mosses (24 genera, 14 families) and 47 of liverworts (22 genera, nine families) were recorded (Appendix 1). The Lejeuneaceae was the most-speciose family with 25 (30%) species, followed by the Lepidoziaceae with eight (9%) and the Sematophyllaceae with seven (8%). The most-speciose genera were Lejeunea and Ceratolejeunea with five species each. Sixty-two species (74%) were found in more than one plot, 47 (56%) in more than one landscape unit and 12 species (14%) in all landscape units. A total of 42 (50%) species was exclusively recorded from neotropical lowland rain forests. The average bryophyte species richness in plots hardly differed between landscape units (Table 1, Fig. 2). However, plots in floodplains and swamps were richer in moss species than terra firme and white-sand plots. Liverwort species richness was highest in white-sand-plots. The total number of bryophyte species was 50 in the floodplain, 45 in swamps and terra firme forests, and 32 in the white-sand areas. Life-forms of moss species were significantly associated with particular landscape units (Table 2). Thus, among mosses more species of fans were found in floodplains than in other landscape units. Life-forms of liverwort species were not related to the landscape units. The DCA ordinations diagrams revealed a consistent separation of five floodplain plots from the other plots along the principal DCA axes (Fig. 3). These five plots were exclusively situated in the floodplain of the Caqueta´ River. ANOVA tests of plot scores along the first DCA axes confirmed the distinct position of the floodplain plots (Table 3).

Table 1. Bryophyte species richness (mean number¡SD) recorded in the understorey of n 0.1-ha plots in forests of Colombian Amazonia. Homogeneous groups identified by Tukey’s test are indicated by the same superscript letter. Group

Floodplain (n58)

Swamp (n55)

Terra firme (n519)

White sand (n58)

ANOVA results (F3,36, p)

All bryophytes Mosses Liverworts

15.4¡4.1 8¡2.4a 7¡2.3 a

15.2¡3.0 7.1¡0.99 a 7.6¡2.1 a

12.2¡3.6 4.8¡1.7 7.3¡2.7

14.8¡2.5 3.2¡1.8 12.2¡2.6

2.3, p50.097 10.7, p,0.001 5.7, p50.003

b a

b b

185


Figure 3. Ordination diagrams of DCA based on species composition of bryophytes, mosses, and liverworts in 40 0.1-ha plots. The five plots separated towards the left side of the diagrams were exclusively from the floodplain of the Caqueta´ River.

DISCUSSION Species richness Despite the variation in forest structure between the landscape units in the study area (Duque et al., 2001),

significant between-landscape differences in the understorey bryophyte species richness were not detected. However, floodplain and swamp plots tended to show higher species richness than terra firme and white-sand plots for mosses, and white-sand plots were distinctly richer in species than the other landscape units for liverworts. It suggests that factors affecting bryophyte species richness differ between mosses and liverworts. The divergent strategies of these bryophyte groups might cope differently with disturbances, temperature, water, nutrients and light availability (Ma¨gdefrau, 1982; Richards, 1984; Cornelissen & ter Steege, 1989; Bates, 1998; Kessler, 2000; Proctor, 2000). The relatively high and constant humidity in floodplains and swamps might offer a benign environment for many moss species. The annual flooding in the floodplain during which the river water level easily reaches more than 1.5 m above the forest floor, however, does not seem to affect the species richness in mosses. Small-scale experiments under controlled humidity levels, together with detailed monitoring on substrate availability as function of daily and weekly disturbances in the forest understorey, might shed light on the role of colonization processes and interspecific competition in explaining species coexistence in mosses. The abundance of fans (and pendants) in the floodplains further suggests that mosses and these open life forms of mosses in particular are limited by water availability (Bates, 1998). The high number of liverwort species in white-sand forests might result from the high incidence of light at the forest floor, resulting from the low canopy and low forest stratification (Duque et al., 2001). This condition also allows the establishment of typical drought-tolerant epiphytes in the forest understorey, such as Drepanolejeunea palmifolia and species of Ceratolejeunea. Liverworts have been considered as indicators of humidity in mountain gradients as they are less tolerant to drought than mosses (Frahm & Gradstein, 1991; Wolf, 1993). However, in relatively exposed parts of the tropical rain forest canopy, the number of liverwort species was found to be higher than that of mosses (Cornelissen & ter Steege, 1989; Gradstein, Montfoort & Cornelissen, 1990). While spores of epiphytic liverworts on average have a shorter life span than those of mosses, they seem to be able to colonize exposed epiphytic

Table 3. Summary results of DCA based on species composition of bryophytes, mosses and liverworts in 40 0.1-ha plots in Colombian Amazonia. The right column shows the ANOVA results of the plot scores along the first DCA axis against landscape unit (floodplain, swamp, terra firme or white sand). Eigenvalues

Length of gradient

Cumulative variance (%)

ANOVA results(F3,36, p)

Axis 1 Axis 2

0.61 0.25

4.2 2.4

13.1 18.3

21.6, p, 0.001

Axis 1 Axis2

0.68 0.35

3.5 2.8

15.0 23.3

18.2, p,0.001

Axis 1 Axis 2

0.67 0.34

4.9 3.2

14.4 21.7

7.6, p50.0005

Bryophytes

Mosses

Liverworts

186


(canopy) substrata because of special morphological features such as water sacs and endosporic protonemata enabling them to avoid hydric stress (Gradstein & Poćs, 1989).

Distribution The bryophyte composition (both mosses and liverworts) in the five Caqueta´ floodplain plots is remarkable, and is probably related to a unique combination of two properties by which this varzea system differs from the other landscape systems: the seasonal flooding (which generally might last continuously up to 4 months per year, depending on the position relative to the water level of the Caqueta´ River; Duivenvoorden & Lips, 1993, 1995) and the yearly sedimentation of nutrient-rich fine silt (Duivenvoorden & Lips, 1993, 1995). Due to these two phenomena, more surface area of fresh and fertile mineral substrates is exposed for colonization each year and conditions for productive growth are better than elsewhere. Several species (Lepidopilum scabrisetum, Squamidium livens, Sematophyllum subsimplex, Syrrhopodon prolifer, Bazzania hookeri and Lophocolea bidentata) from the plots of the Caqueta´ floodplain showed a distribution which was not restricted to the lowlands (Uribe & Gradstein, 1998; Churchill et al., 2000). Possibly, the Caqueta´ River has served as corridor for bryophyte range expansion from the Andes into the Amazonian lowlands (Gascon et al., 2000). More quantitative bryophyte surveys, both in the Andes and Amazon basin, are needed to test this idea. ACKNOWLEDGEMENTS The authors thank Herbarium HUA and Fundacioń Puerto Rastrojo for logistic support, J. Uribe and T. Poćs for help with the identification of liverwort species, and S.R. Gradstein for correcting the manuscript. Comments from two anonymous reviewers greatly improved the manuscript. This study was partially financed by the European Commission (ERB IC18 CT960038), TropenbosColombia, COLCIENCIAS-LASPAU program, and the Netherlands Foundation for the Advancement of Tropical Research – WOTRO (WB85-335). TAXONOMIC ADDITIONS AND CHANGES: Nil. REFERENCES Bates JW. 1998. Is ‘life form’ a useful concept in bryophyte ecology? Oikos 82: 223–237. Bates JW. 2000. Mineral nutrition, substratum ecology, and pollution. In: Shaw AJ, Goffinet B, eds. Bryophyte biology. Cambridge: Cambridge University Press, 403–448. Botero PJ. 1999. Paisajes fisiogra´ficos de Orinoquia y Amazonia (ORAM) Colombia. Instituto Geografico Agustin Codazzi (IGAC), Bogota´ D.C. Churchill SP. 1994. Mosses of Amazonian Ecuador. AAU Reports 35: 1–211.

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JUAN C. BENAVIDES, Departamento de Biologıá, Recinto Universitario de Mayaguëz, Universidad de Puerto Rico ALVARO J. DUQUE M., Departamento de Ciencias Forestales, Universidad Nacional de Colombia, Sede Medellıń. E-mail: [email protected] JOOST F. DUIVENVOORDEN, Institute for Biodiversity and Ecosystem Dynamics (IBED), Centre for Geo-Ecological Research (ICG), Universiteit van Amsterdam. ANTOINE M. CLEEF, Institute for Biodiversity and Ecosystem Dynamics (IBED), Centre for Geo-Ecological Research (ICG), Universiteit van Amsterdam.

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APPENDIX 1. Bryophyte species found in 40 0.1-ha plots in the middle Caqueta´ and Chiribiquete regions of Colombian Amazo nia. TF5terra firme, FP5floodplains, SW5swamps, WS5white-sand forest. The distribution is based on Uribe & Gradstein (1998), Gradstein & da Costa (2003), Churchill et al. (2000) and Churchill (2003). Life form assignment follows Bates (1998).

Mosses Bartramiaceae Philonotis sphaerocarpa (Hedw.) Brid. Calymperaceae Syrrhopodon incompletus Schwa¨gr. incompletus Syrrhopodon prolifer Schwa¨gr. Syrrhopodon rigidus Hook. & Grev. Syrrhopodon simmondsii Steere Daltoniaceae Lepidopilum affine Mu¨ll.Hal. Lepidopilum scabrisetum (Schwa¨gr.) Steere Lepidopilum surinamense Mu¨ll.Hal. Dicranaceae Leucobryum martianum (Hornsch.) Hampe ex Mu¨ll.Hal. Octoblepharum albidum Hedw. Octoblepharum erectifolium Mitt. ex R.S. Williams Octoblepharum pulvinatum (Dozy & Molk.) Mitt. Octoblepharum stramineum Mitt. Fissidentaceae Fissidens elegans Brid. Fissidens pellucidus Hornsch. Hookeriaceae Brymela parkeriana (Hook. & Grev.) W.R.Buck ˚ ngstro¨m Callicostella pallida (Hornsch.) A Callicostella rivularis (Mitt.) A.Jaeger Thamniopsis cruegeriana (Mu¨ll.Hal.) W.R.Buck Hypnaceae Ectropothecium leptochaeton (Schwa¨gr.) W.R.Buck Vesicularia vesicularis (Schwa¨gr.) Broth. Meteoriaceae Squamidium livens (Schwa¨gr.) Broth. Neckeraceae Neckeropsis disticha (Hedw.) Kindb. Neckeropsis undulata (Hedw.) Reichardt Phyllodrepaniaceae Mniomalia viridis (Mitt.) Mu¨ll.Hal. Phyllodrepanium falcifolium (Schwa¨gr.) Crosby Pottiaceae Hyophila involuta (Hook.) A.Jaeger Sematophyllaceae Colobodontium vulpinum (Mont.) S.P.Churchill & W.R.Buck Hypnella pallescens (Hook.) A.Jaeger Sematophyllum adnatum (Michx.) E.Britton Sematophyllum subsimplex (Hedw.) Mitt. Taxithelium planum (Brid.) Mitt. Trichosteleum fluviale (Mitt.) A.Jaeger Trichosteleum papillosum (Hornsch.) A.Jaeger Stereophyllaceae Pilosium chlorophyllum (Hornsch.) Mu¨ll.Hal. Stereophyllum radiculosulum (Mu¨ll.Hal.) A.Jaeger Thuidiaceae Cyrto-hypnum involvens (Hedw.) W.R.Buck & H.A.Crum Liverworts Aneuraceae Riccardia amazonica (Spruce) Schiffn. ex Gradst. & Hekking Calypogeiaceae Calypogeia miquelii Mont. Calypogeia tenax (Spruce) Steph.

Landscape unit

Life form

Distribution

FP

Turf

–

Turf Turf Turf Turf

– – – Lowland

FP FP FP

Fan Fan Fan

Lowland – Lowland

FP, SW, TF, WS FP, SW, TF TF FP, SW, TF, WS SW, TF

Turf Turf Turf Turf Turf

Lowland Lowland – – Lowland

FP SW, TF

Turf Turf

– Lowland

SW FP, SW TF FP

Rough Rough Rough Rough

FP FP

Rough mat Rough mat

– –

FP

Pendant

–

FP, SW FP, SW

Fan Fan

Lowland –

SW SW

Turf Fan

Lowland Lowland

SW

Turf

–

FP WS FP, FP, FP, SW FP,

Mat Mat Mat Mat Mat Rough mat Rough mat

Lowland Lowland Lowland – Lowland – –

FP, SW, TF SW

Mat Mat

Lowland Lowland

FP, WS

Weft

Lowland

FP, SW, TF, WS

Mat

–

SW, TF FP, TF, WS

Mat Mat

Lowland Lowland

FP, FP, FP, FP,

SW, TF, WS SW SW, TF TF, WS

SW SW, TF, WS SW, TF SW, TF, WS

mat mat mat mat

Lowland Lowland Lowland –

189


Mnioloma cellulosa (Spreng.) R.M.Schust. Mnioloma parallelogramma (Spruce) R.M.Schust. Cephaloziaceae Odontoschisma falcifolium Steph. Geocalycaceae Lophocolea bidentata (L.) Dumort. Lophocolea martiana Nees Lejeuneaceae Aphanolejeunea camilli (Lehm.) R.M.Schust. Aphanolejeunea clavatopapillata (Steph.) Reiner-Drehwald Archilejeunea crispistipula (Spruce) Steph. Archilejeunea ludoviciana (De Not. ex Lehm.) Geissler & Gradst. Ceratolejeunea confusa R.M.Schust. Ceratolejeunea cornuta (Lindenb.) Schiffn. Ceratolejeunea cubensis (Mont.) Schiffn. Ceratolejeunea fallax (Lehm. & Lindenb.) Bonner Ceratolejeunea sp. Cheilolejeunea rigidula (Nees & Mont.) R.M.Schust. Cheilolejeunea trifaria (Reinw. et al.) Mizut. Drepanolejeunea palmifolia (Nees) Steph. Drepanolejeunea subdissitifolia Herz. Lejeunea boryana Mont. Lejeunea controversa Gottsche Lejeunea flava (Sw.) Nees Lejeunea intricata J.B.Jack & Steph. Lejeunea phyllobola Nees & Mont. Lopholejeunea nigricans (Lindenb.) Schiffn. Lopholejeunea eulopha (Tayl.) Schiffn. Pictolejeunea picta (Gottsche ex Steph.) Grolle Pictolejeunea sprucei Grolle Stictolejeunea balfourii (Mitt.) E.W.Jones Stictolejeunea squamata (Willd. ex Web.) Schiffn. Xylolejeunea crenata (Mont.) X.-L.He & Grolle Lepidoziaceae Arachniopsis diacantha M.Howe Bazzania aurescens Spruce Bazzania hookeri (Lindenb.) Trevis. Bazzania pallidevirens (Steph.) Fulf. Bazzania phyllobola Spruce Micropterygium leiophyllum Spruce Micropterygium trachyphyllum Reimers Zoopsidella integrifolia (Spruce) R.M.Schust. Metzgeriaceae Metzgeria aurantiaca Steph. Plagiochilaceae Plagiochila cf. distinctifolia Lindenb. Plagiochila montagnei Nees Plagiochila subplana Lindenb. Plagiochila cf. vincentina Lindenb. Radulaceae Radula sp.

Landscape unit

Life form

Distribution

WS TF, WS

Mat Mat

– Lowland

FP, TF, WS

Rough mat

–

FP TF

Mat Mat

– –

TF TF FP, TF SW, TF, WS FP, SW, TF, FP, SW, TF, FP, SW, TF WS SW SW, TF, WS WS TF, WS TF FP, SW SW, TF SW SW FP, SW, TF, FP FP, WS TF FP, SW, TF FP FP, TF FP, SW, TF,

Mat Mat Fan Fan Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat Mat

Lowland Lowland Lowland Lowland Lowland Lowland Lowland Lowland

SW, TF, WS TF, WS SW, TF, WS FP, TF, WS TF FP, SW, TF, WS TF SW, TF, WS

Mat Fan Fan Fan Fan Rough mat Rough mat Mat

– Lowland – – – Lowland Lowland –

FP, SW, WS

Mat

Lowland

FP, SW, TF, WS FP, SW FP FP, SW, WS

Fan Fan Fan Fan

Lowland – – –

FP

Rough mat

WS WS

WS

WS

– – Lowland – Lowland Lowland – Lowland – – Lowland Lowland Lowland Lowland – –