Journ. Hallori Bot. Lab. No. 69: 65-78 (Jan . /991)

A PHENETIC AND CLADISTIC STUDY OF THE CAMPYLOPODIOIDEAE

JAN-PETER FRAHMl

SUMMARY. Based on a phenetic evaluation of 7 gametophytic and 10 sporophytic characters, the Campylopodioideae have been redefined. The Campylopodioideae now consist of Atractylocarpus, Bryohumbertia, Campylopus, Dicranodonlium, Pilopogon and Sphaerothecium. The genera Microcampylopus and Campylopodium are placed in the Dicranelloideae, and Campylopodiella is placed to the Paraleucobryoideae. The sinuose seta has probably evolved several times independently. A key is given for the genera of Campylopodioideae. A cladistic analysis, based on the same character states as for phenetic analysis, reflects phenetic analysis.

INTRODUCTION The were first divided by Fleischer (190 I) into the Dicranelleae and Dicraneae. The former was defined by lacking alar cells and included Microcampylopus, Campylopodium and Dicranella. The latter was defined by with differentiated alar cells, including Dicranum, Dicranodonlium, Braunfelsia, Leucoloma and Campylopus. Brotherus (190 I), in his first edition of "Die Natuerlichen Pflanzenfamilien", took over Fleischer's concept, but treated Microcampylopus as a subgenus of Campylopus. In the second edition Brotherus (1924) split the Dicranaceae into a number of subfamilies: Trematodontioideae, Rhabdoweisioideae, Campylopo­ dioideae, Anisothecioideae, Dicranoideae and Paraleucobryoideae. Of these subfamilies, the Trematodontioideae have been treated as a separate family (Buck 1979). The Rhabodoweisioideae seem to be distinct in several gametophytic and sporophytic characters and well defined. However, the definition and delimitation of Dicranoideae, Paraleucobryoideae, Campylopodioideae and Anisothecioideae seem not to be sufficient. The Campylopodioideae were characterized by five characters ("meist dioezisch; Blatter von der Rippe gegen die Rander allmahlich diinner; Blattfliige1zellen meist differenziert; Perichaetialblaetter differenziert; Spaltoffnungen meist fehlend"), three of which were "mostly" expressed, the fourth character (leaves from the costa to the margin gradually narrower) is not applicable to most genera and the remaining character (differentiated perichaetialleaves) causes closely related genera such as Dicranella and Anisothecium to be placed in different subfamilies. Nevertheless, most authors followed the concept proposed by Brotherus (1924). Magill (1980), however, followed Fleischer's concept in leaving out the Campylopodioideae; placing Aongstroemia, Dicranella and Microdus in the Dicranelloideae; and all other

I Fachbereich 6, Botanik, Universitiit Duisburg, Post fa ch 101503, D-4100 Duisburg I, B.R.D. 66 Journ. Hattori Bot. Lab. No. 69 I 9 9 I genera in the Dicranoideae. Walther (1983) in the most recent systematic treatment differentiated between Dicranoideae, Dicranelloideae and Campylopodioideae, a classification resembling that of Brotherus (1924) with the exception that Walther joined the Anisothecioideae and Dicranelloideae and placed Campylopodiella to the Paraleucobryaceae, Campylopodium in the Dicranelloideae and Microcampylopus to the Campylopodioi­ deae. Thus the present systematic arrangement of the genera of Dicranaceae is rather confusing, partly contradictory and not satisfying. A short discussion of the interrelationships of these genera has been published by Frahm (1986). To clarify the relationships between the genera included in these subfamilies, detailed studies of the generic concepts were lacking. Recent worldwide monographs or revisions of several genera provided a better overlook and a good basis for defining higher taxonomic categories. Monographs of Microcampylopus and Campylopodium (Giese & Frahm 1986, 1986a), and Pilopogon (Frahm 1983b) resulted in substantial reductions of the number of species. Revisions of Sphaerothecium (Frahm 1986a), Campylopodiella (Frahm 1984a), A tractylocarpus (Padberg & Frahm 1985), Campylopus subg. Campylopidulum (Frahm 1986b), subg. Thysanomitrion (Frahm 1984) and the Paraleucobryoideae (Muller & Frahm 1987) provided reinterpretations of generic concepts. Treatments of various regional floras of Campylopus provided a better ovcrlook of the species and their character states. In addition, the genus Bryohumbertia has been newly defined (Frahm 1982a). Altogether, character states used fo r the circumscription of these genera can now be critically evaluated, the range of expressions of these character states can be estimated, and this knowledge can be used as a good basis for defining higher taxonomic categories. For example, the past delimitation of the genera has been partially incorrect as shown by the revisions of Campylopodium and Microcampylopus, where several species described as Microcampylopus proved to be Campylopodium and vice versa, or had to be placed into to Dicranella or Campylopus. Furthermore the number of species has been much reduced, as in Campylopus from 720 to 180, in Pilopogon from 14 to 8, in Campylopodium from 12 to 2. On the other hand, genera regarded as monotypic (such as Bryohumbertia) proved to include several species. The following study comprises an evaluation of all character states and a character correlation found in the genera Campylopus, Pilopogon, Bryohumbertia, Microcampylopus, Campylopodium, Dicranodontium and A tracty!ocarpus, all of which were originally placed in the subfamily Campylopodroideae, as well as the genera of Paraleucobryoideae (incl. Brotheroideae, for revision see Muller & Frahm 1987). Dicranella (incl. Anisothecium) is considered here, too, because they are related in some aspects, although the 180 species of this genus have not yet been revised. The genus Bryotestua described by Theriot and Portier de la Varde and placed in the Anisothecioideae is, however, omitted here, since it was based on sterile plants, and cannot be compared in all repects with the other genera. Some other genera, such as Dicranum and Orthodicranum, are used as outgroups in the c1adistic analysis. J.-P. FRAHM: A phenetic and cladistic study of Campylopodioideae 67

The genus Earlleya described by Robinson ( 1966) is not taken into consideration because it was merely based on sterile material and the only species was placed into synonymy of Dicranella cerviculata (Crum & Anderson 1981). The monotypic genus Maireola described by Theriot ( 1927) and placed in the Dicranaceae has recently been recognized as belonging to the genus Ditrichum (Frahm & Seppelt 1987).

2. EVALUATION OF CHARACTERS 2a. GAMETOPHYTE The most conspicuous character of the gametophyte in all genera is the wide costa, filling half of the leaf width or more, although there is a broad range of leaf length between 2.5 and 15 mm and leaf shape. This feature is found in elsewhere only in the . Alar Cells For this character onl y the potential presence of alar cells has been evaluated. There are genera without any alar cells (Microcampy/opus, Campylopodium, Dicranella) or with very indistinctly differentiated alar cells (A rracly /ocarpus). In the other genera (Campy/opus, Dicranodontium) alar cells may be developed but sometimes only in special habitats or in some species. The absence of alar cells in some species may reflect the occurrence of these species in habitats where alar cells are not developed (such as habitats with high humidity). Field observation have shown that alar cells in the same species are not developed in habitats with high humidity such as rainforests (and uptake of atmospheric water through the whole ) and are well developed in exposed habitats with wet substrates (and water conduction along the stem as well as water uptake by the leaves through the alar cells). So these genera can have potentially well developed alar cells, which may not be expressed in some taxa.

Rhizoids Crundwell (1979), in his survey of the use of rhizoids for , made the interesting observation that Campylopus as well as related genera placed by Brotherus (1924) in the same subfamily (Microcampylopus, Pilopogon, A /ractylocarpus and Dicranodontium) have rhizoids born from the abaxial side of the costa and not from the stem as in other genera. In Atractylocarpus coslaricensis (now placed in Campylopodiella, cr. Muller & Frahm 1987, Frahm & Isoviita 1988) and Dicranodontium, rhizoids are borne on both sides of the costa . In contrast, Microdus, Dicranella and Campylopodium have stem borne rhizoids, and therefore Crundwell proposed to place these genera into the Anisothecioideae. The indications of leafborne rhizoids in Microcampy!opus by Crundwell ( 1979) probably goes back to the examination of species such as "M icrocampy!opus perpusillus" which in fact belongs in Campylopus (Giese & Frahm 1985). rn the 2 species of Microcampylopus that are retained in this genus after revision, no rhizoids were observed. Thus leafborne rhizoids become a character diagnostic of the Campylopodioideae and Paraleucobryoideae. 68 Journ. Hattori Bot. Lab. No. 69 199 1

Transverse Section of Costa Usually the costa shows an abrupt transition to the lamina in the way that the costa is prominent at the abaxial surface of the leaf. Genera such as Campylopodiella, Brothera, Paraleucobryum and part of the species of Campylopus subsp. Campylopus sect. Homalocarpus (those with lax ventral hyalocysts) show a gradual transition between lamina and costa. There are surprisingly gametophytic intergradations between part of the species of Campylopus and the Paraleucobryoideae in the structure of the costa (Frahm 1982a). However, the sporophytic characters of both groups are so different that a polyphyletic origin of Campylopus (in part from the Paraleucobryoideae) can be excluded and these characters common in both groups must be regarded as analogous. Most of the Dicranaceae show more or less the same type of transverse section of costa, with (often multilayered) bands of ventral and dorsal stereids, the outermost layer usually developed as an epidermal layer with larger lumens. Only the Paraleucobryoideae and again part of the genus Campylopus show hyalocysts in transverse section. Between Paraleucobryum and Camyplopus there is a gradual transition from dorsal hyalocysts to a dorsal band of stereids. This transition can be explained by the presence or absence of three cell divisions, but apparently does not reflect phylogenetic relationships because of the different sporophytes in the two groups. The transverse leaf section anatomy of Paraleucobryum is also linked with Orthodicranum and Dicranum (M iiller & Frahm 1987). Tn this case the sporophytes of Orthodicranum and Paraleucobryum are quite similar, suggesting a phylogenetic relationship. The amount of variation in the structures of the transverse section of the costae is different. Some genera (Microcampy lopus, Campylopodium, Dicranella, Atractylo­ carpus, Dicranodontium) show little variation. In the three species of Sphaerothecium, two have ventral stereids and one has ventral hyalocysts. This is the case in a presumably old genus, with a scattered but worldwide distribution of species. In Pilopogon, the most primitive (and only African species) has ventral substereids whereas in the neotropical species the most specialized species such as P. peruvianus (of deserts) and P. longirostratus (of rain forests and the only species with incrassate and pitted basal laminal cells) ha ve multilayered, small ventral stereids. The greatest variation of the structures in the costa is found in Campylopus, which is probably the reason for its enormous diversification enabling this genus to occupy a broad variation of habitats. Incrassate, Pitted Lamina Cells Firm basal laminal cells are found only in the genera Dicranodontium, Bryohumbertia and Sphaerothecium. In Pilogon, there is only one species with incrassate laminal cells, and in Campy /opus they are present only in species of subg. Campylopus, and in all except one species of subg. Thysanomitrion. All other taxa have thinwalled hyaline basal laminal cells, which are characteristic of species from humid habitats. J.-P. FRAHM: A phcnctic and cladistic study 01 Campylopodioidcac 69

Perichaetial Leaves Long, sheathing perichaetialleaves are highly diagnostic for the genus Pilopogon. This character is also found in part of the species of Dicranella these often placed in Anisothecium by some authors. The variation in perichaetial leaf form in such closely related groups as Dicranella and Anisothecium seems to indicate that his character is not of much use for an analysis of subfamilies. Hyaline Leaftips Hyaline excurrent costae as special xeromorphic adaptions occur only in some species of Campylopus and Pilopogon but not in any other genera.

2b. SPOROPHYTIC CHARACTERS Seta The seta is twisted in all of the genera treated here. In addition it is cygneous in Campylopus, Microcampy/opus, Campylopodium. Dicranodonfium, Sphaerofhecium, Bryohumbertia and straight in all other genera. The function of the sinuose setae may be to protect the young capsule during its development between the perichaetial leaves (Frey & Frahm 1987). Since closely related genera such as Campylopus and Pilopogon or Dicranodontium and Atractylocarpus have sinuose or straight setae, this character may not be as important as it might seem. Sinuose setae are sometimes difficult to recognize, especially in old sporophytes, since they are curved only in a certain stage of development, when the seta is curved down to place the young sporophyte between the perichaetial leaves. When the seta has curved upwards again, the seta is more or less straight again but still has a sigmoid curve in the middle (Frey & Frahm 1987 figs. 1- 7). This is apparently the reason that uncurved setae have been indicated for genera such as Dicranodontium or Campylopodiella (Brotherus 1924). Annulus A well developed annulus is found only in Para/eucobryum, Brothera and Campylopodiella and in part of the species of Dicranum, Orfhodicranum and Dicranella. It has been taken into account only if it is dehiscent. In some other genera such as Campylopus an annulus is visible but without function. Calyptra The shape of the calyptra is cucullate in all genera. In the first stages of their development, cucullate calyptras are symmetric and cover the young capsules all around. This can then lead to misinterpretations as in Brothera, for which sometimes a mitrate calyptra is indicated (cr. Crum & Anderson 1981). The base of the calyptra may be ciliate or not ciliate. Longly ciliate calyptras are diagnostic for Pilopogon and Campylopus subg. Thysanomitrion. There are, however, several examples in Campylopus in which the same species has been found with and without ciliate calyptras. In other genera, such as Campy/opodiella and Dicranodontium, species with or without ciliate calyptras are found. So the diagnostic value of this 70 Journ. Hallori Bot. Lab. No. 69 199 I character is not high. All other genera, however, seem always to have calyptras with entire bases. Peristome Teeth The peristome teeth are usually very uniform and of the so called dicranoid type. They are usually split into two prongs. Only Brothera and most of the species of Pifopogon have entire teeth. The inner surface of the teeth is usually papillose; only in Bryohumbertia is it smooth. The tips of the teeth are papillose and elongate, except in most species of Pilopogon, Campy lopus subg. Thysanomitrion, and Brothera, where they are fi liform. In Pilopogon the only African species has the typical dicranoid peristome but the 7 Andean species have long filiform teeth. In Campylopus subg. Thysanomitrion the only subantarctic species has a dicranoid peristome, but all tropical species have longly filiform teeth. The divided teeth are split half their length or more. In the common dicranoid type the peristome is split halfway down. Dicranodontium has peristome teeth split the whole length. In Campylopus only the filiform teeth of subg. Thysanomitrion are split more than the half. Lid The lid is obliquely rostrate in all genera and usua lly shorter than the urn. It is, however, as long or longer than the urn in Bryohumbertia, Brothera and Pilopogon longirostratus. Shape of Capsule The capsule shape is normally shortly cylindric. It is longer in Paraleucobryum, Orthodicranum, Dicranum, Campylopodiella and Pilopogon but globose in Sphaer­ othecium. The globose capsule shape is also found in Campylopus subg. Campylopidulum. The capsules can be symmetric or asymmetric. This character varies within genera, especially in Campy lopus or Dicranella. Curved, asymmetric (often strumose) capsules are characteristic for Bryohumbertia and Dicranum s. str., whereas in a ll other genera symmetric capsules are found. Stomata Stomata in the capsules are found in Paraleucobryum, Dicranella and Campylopodium. The val ue of this character may not be as high as expected since genera closely related in other characters differ by the presence or absence of stomata, as in Campylopodium and Microcampylopus. Spore Size Spore size of about 1311m is very constant in all infrageneric categories of Campylopus, Bryohumbertia, Dicranodontium and Pilopogon, indicating a close phylogenetic relationship, and also in Brothera. In Campylopodiella, one species (c. himalayana) has larger spores. Atractylocarpus has spore sizes of 14-1 8I1m. Spores sizes of more than 20l1m are found only in Paraleucobryum and Sphaerothecium. J.-P. FRAHM: A phenetic and c1adistic study of Campylopodioideac 71

Chromosome Numbers At present, chromosome numbers of genera of Campylopodioideae and Paraleuconryoideae cannot be used for systematic evaluation, since they are known only from a small percentage of species (cf. Fritsch 1982) and not even from all genera. Even the available information is doubtful considering misidentifications. At least, all chromosome count records (except for "Campy /opodium khasianum" with n = 35) are in the range between IO and 15 and thus represent the most common chromose numbers in the acrocarpous mosses.

3. PHENETIC ANALYSIS For an evaluation of the phenetic relationships of the genera, a cluster analysis was performed. Within Campylopus, the subgenera Thysanomilrion and Campylopus have been used separately, since Campylopus is much more variable than any other genus taken into account, and the concept of Campylopus would be too broad. The cluster analysis was performed on a desktop computer using a program described by Frahm & Rettig (1984). F ig. I shows I. a clear distinction between three main blocks comprising genera which fit into Dicranelloideae, Paraleucobryoideae and Campylopodioideae. 2. The genera Microcampy /opus and Campylopodium are placed into the Dicranelloideae. 3. Brothera and Campylopodiella and next related and separated from

Campylopoaiella

l'1i crocampylopus

Dicranella

Or thod lcranum

Bry ohumbert 1a

01 cranod ont l um

Atractylo carpu s

.---+----- Spha erot hecl um

T hysano m i t rl on

P I 1op og on

Campyl opu s

,------Paraleucobryum

~ CampylopoCllella

Brothera

FIG. I. Phenetic analysis of the genera ofCampylopodioideac and Paraleucobryoideae. 72 Journ. Hattori Bot. Lab. No. 69 199 I

Paraleucobryum, but since these three genera are separated from all other on more or less the same level as Dicranelloideae and Campylopodioideae, there is no evidence to split both genera into a separate subfamily, Brotheroideae, as proposed by Takaki (I968), separate from the Paraleucobryoideae. 4. Within the Campylopodioideae, Dicranodonlium and Atractylocarpus as well as Bryohumhertia and Campylopus subg. Thysanomitrion are related. This would suggest that Thysanomitrion should be distinguished at the genus rank. However, since sterile specimens of Campylopus subg. Thysanomilrion cannot be distinguished from certain species of Campylopus subg. Campylopus, this separation is not followed here mainly for reasons of practicability. 5. Genera with cygneous setae occur in all three clusters recognized here as subfamilies: Microcampylopus and Campylopodium within the Dicranelloideae, Camylopodiella within the Paraleucobryoideae and all genera except for Pilopogon and A traclylocarpus within the Campylopodioideae. This character also occurs in Cynodontium gracilescens (Web. & Mohr) B.S.G. which clearly shows that it has been developed several times independently. Proposed Classification Derived from the phenetic analysis, the following modified classification is proposed: Paraleucobryoideae Brotherus, Nat. Pft . 2, 10: 191 , 1924 emend. M iiller & Frahm, Nova Hedwigia 45: 283- 314, 1987. Description, key to the genera and description of genera see M iiller and Frahm 1987.

Campylopodioideae Brotherus, Nat. Pft. 2, 10: 180, 1924, emend. Frahm K EY TO THE GENERA Seta erect when wet 2 Perichaetial leaves long, sheathing the seta ...... Pilopogon Brid., see Frahm 1983 2* Perichaetialleaves indistinct ...... Atractylocarpus Mitt., see Padberg & Frahm 1985 I * Seta cygenous when wet 3 Seta 3-4 mm long, inserted in the perichaetial leaves 4 Spores 1311m in diameter ...... Campylopus subg. Campylopidulum Vital, see Frahm 1986b 4* Spores 20- 2211m in diameter ...... Sphaerofhecium Hampe, see Frahm 1968a 3* Seta 5- 15 mm long, exserted 5 Seta 15 mm long, lid as long as the capsule ...... Bryohumbertia P. Varde & Ther. emend. Frahm, see Frahm 1982 5* Seta 6-12 mm long, lid shorter than the capsule 6 Upper laminal cells more than 6 times longer than broad ...... Dicranodontium Brid. 6* Upper laminal cells quadrate to 4 times longer than wide 7 Basal laminal cells incrassate 8 Capsules symmetric, scabrous at base ...... Campy/opus subg. Thysanomifrion, see Frahm 1984 J.- P. FRAHM: A phcnetic and cladistic study of Campylopodioideae 73

8* Capsules curved, smooth at base ...... Campylopus subg. Campy/opus sect. Campy/opus 7* Basal laminal cells thinwalled ...... Campy/opus subg. Campy/opus sect. Homalocarpus

4. CLADISTIC ANALYSIS A c1adistic analysis was performed with the same data matrix as used for the phenetic analysis. For the calculation, the IBM-PC version of the program "Treesearch" written by R . W . Bruggeman (University of Utrecht) was used. The program Treesearch uses a combination of parsimony as well as compatibility analysis, whereas similar c1adistic programs use either parsimony (PAUP, PHYLIP, H ENNTG 86) or compatibility (CLINCH). As outgroups, the genera Dicranum and Orthodicranum have been added. In contrast to the phenetic analysis, the c1adistic analysis requires the evaluation of primitive (plesiomorphous) and derived (apomorphous) characters. An indication of modern taxa with apomorphous characters can be taken from a phytogeographical analysis. For example, Microcampy lopus and Campylopodium and also Dicranodontium and Atractylocarpus seem to be sister groups. As an analysis of their distribution shows and seem to have a common evolutionary history. Dicranodontium is a mainly holarctic genus confined to forests, which invades only part of the tropical mountains, whereas A tractylocarpus is an alpine genus and present in a ll mountain systems of the world of Tertiary origin, thus probably younger than Dicranodontium. Microcampylopus is present in South America, Africa and SE-Asia with each one species whereas Campylopodium is confined to SE-Asia and therefore younger. Many genera have worldwide ranges, either through the tropics or the holarctic, and can therefore be regarded as old. Examples for such old genera in the tropics are Sphaerothecium (3 species each in South America, Africa and SE-Asia known from only extremely small ranges, cr. Frahm 1986a), Bryohumbertia (each one species in South America, Africa and SE-Asia), and Campylopodiella (disjunct in the Himalayas, New Guinea, and the Andes). Examples for widespread holarctic genera which go into the tropical mountains with few species are Dicranum, Orthodicranum, Paraleucobryum, Dicranella, and Dicranodontium. Only a few taxa indicate a younger origin. A tractylocarpus is found with about one species each in any of the major mountain systems of the world (Padberg & Frahm 1985), which are of Tertiary age so that at least the species (but not the genus) are of younger origin. In Pi/opogon, 7 of the 8 species occur in the Andes and probably have evolved there. For Campylopus it could be shown (Frahm 1988) that the genus was probably present in the southern part of the Gondwanaland continent and has spread from there to the tropics. This is especially true for the subg. Thysanomitrion, of which the probable ancestor occurs in the subantarctic, one species in the Neotropics, but 5 in Africa and 7 in SE-Asia (Frahm 1984). If the distribution maps of both Pi/opogon and Campylopus subg. Thysanomitrion are plotted together (fig. 2), it shows a balanced distribution (7 species of Thysanomitrion ---J..

1,,1 I

iJ 0- :; " i I ::t: ~ ) Ol e: 0- II ::l . I:Il ~ n'0 r '"0- Z ~ 0"- I -.0 ~-~~i~· ~~~~~ L 4I +- -1 r- I __ I.I i -I I

i - r" '

Thysanom 1tr1 on Bryohumbert1a P11opogon

D1cranodont1um Atractylocarpus Campylopus ,.------Campylopodle ll a

Brothera Paraleucobryum Mlcrocampylopus Campylopod1um Dlcranella

FIG. 3. C1adogram of the genera of Campylopodioideae and Paraleucobryoideae. plesiomorphic but straight capsules as apomorphic. Peterson also regarded polysety and asexual reproduction as apomorphic. If these character states would be included in the calculation, Campylopus would get a higher rate of apomorphous characters. The tree derived from the computer program (fig. 3) reflects the same ordination as given by the cluster analysis, grouping together the genera of Paraleucobryoideae, Campylopodioideae and those genera of the Dicranelloideae taken here into account. It supports the placement of Microcamyplopus and Campylopodium into the Dicranelloideae. The Paraleucobryoideae included Paraleucobryum, Brothera and Campylopodiella in both, the phenetic and cladistic analysis. Orthodicranum has been included in the analysis which seemed to link Dicranum and Paraleucobryum (Muller & Frahm 1987). In this tree Orthodicranum shows more affinities than Dicranum, whereas Brothera and Paraleucobryum can be derived from Orthodicranum on the same level of divergence but with other character states. Campylopodiella is derived from Brothera.

ACKNOWLEDGEMENTS Part of this stydy was done during a stay as visiting professor at the University of Alberta. I wish to thank Dr. D. H. Vitt for providing working facilities, support and discussions on the cladistic analysis and correcting the English text.

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