(1985) 31 BLUMEA 57-64 (Polypodiaceae) Synaptospory in the fern genus Pyrrosia Gerda+A. van Uffelen Rijksherbaiium, Leiden, The Netherlands Summary The by correlation Pyrrosia but may sporoderm sculpture between (1977) Kramer investigated is be present for the fern and life genus form of the Pyrrosia. This sporophyte correlation postulated as is not found in other fern groups. in Introduction An derm sis epiphytic order hand, tioned groups terrestrial wide a of spores 1975) correlated with may be groups live whole or a to means by Kramer, enhance higher), hypothesis holds for covers species the formerly as attributed also to his the surfaces a genus in On the other immediate no units like a number ( Dryopteris: Schneller, 1974, and an example This not men- phenomenon to Kramer is suggests these differences exist between larger single as a Firstly, genus. whole and (genus groups examine the relation between in hypothe- together synaptospermy in seed plants, and is Although to stick have that sporo- this to distance away. Leeuwen, 1929*). to sculpture ecology (Hovenkamp, the object to the Kramer mentions the genus exception van analogy sporophyte situation in to larger dispersal sculptured states sporoderm whether Kramer's whether it secondly, the groups within genus. become available chosen some Gastony, Docters interesting data about spores and genus is of their sporoderm it may be who According stable environment and more in (1977), spores, unable that may be cross-fertilization. and and substrate of the sculpture smooth sporangia (Cyatheaceae: dispersal ecology See trees by Kramer, Lecanopteris: level and * in less or Kramer by dispersal ecology. therefore they may have kept together by supposed ly more other to dispersal, called synaptospory As have spread easily to need of in is put forward interesting hypothesis sculpture proposed Tryon (1985) of of Pyrrosia Mirbel study. genera The genus Saxiglossum Drymoglossum rule. and Walker (Polypodiaceae) in prep.; Van Uffelen & (1985). in his Hennipman, Pyrrosia Ching have in also contains the and publication recent- press), Drymoglossum and states it this species to Presl. be an 58 BLUMEA - VOL. 31, No. 1, 1985 G.A. Synaptospory Uffelen: van MATERIAL AND The spores of all studied rosia by based absent thin roughly laesura and the in verrucae cristae echinulae In P. tightly presence the to by of while the types, one of spore. In all, Van type according thin with colliculate (figs. perispore by a by a the in with perispore and large characterized by nummulariifolia -type, and warts over solitary the P. rupestris- type, characterized smaller very the P. 1,2), 2) containing spherical hatch bodies, together 5) and characterized , from sculpture, perispore surface exospore the P. (fig. 4), 4) to princeps- type, rather a spherical (figs. 5,6), verrucae perispore the P. christii- type, characterized species some and/or bisculptate such the P. (fig. 3), 3) 1) characterized subfurfuracea -type, the follows: as that adheres perispore bodies the spore types arrange pronounced, to Pyr- have ornamentation. perispore on revision of Drymoglossum spore another yet in a five spore types within the genus Pyrrosia, main- Hennipman distinguish One may different two shows Saxiglossum in have been (Polypodiaceae) in connection with species formerly placed perispores showing sculptured species formerly placed Uffelen and five Mirbel Pyrrosia genus 59 METHODS Hennipman (I.e.) The by Hovenkamp (I.e.). elaborately ly in the species Van Uffelen and Pyrrosia in solitary large by verrucae a or (figs. 7, 8). princeps- type that an extent spores they will the exospore as be ornamented may cohere easily as spores with but (fig. 2) elaborately an not to ornament- ed perispore. RESULTS known are be to terrestrial impossible Of the 20 were rocks or for ence Fig. do 1-8. not (like 5: P. the but place plants) of — were growth; in the genus Species labels of the terrestrial Pyrrosia 123 specimens studied or epilithic. Forty-three whether this epiphytic (supposed or the on a - types P. in 52 terrestrial; 3 These last substrate lateral 3: specimens, ground. certain Spore 80 and 3 is Pyrrosia; view. — 2: P. were guineae (Christ) two to specimens not scale were the means specimen be usual in Pyrrosia) growing dead wood, growing on tree indicate that in distinct. very 2 is represents 10 schimperiana (Kuhn) 1227, L), approx. (Brass 27921, L), - 8: P. two lateral Kew spores. piloselloides (L.) Price - some µm. — Alston 1: P. (Maas well as the species 685-69 princeps (Mett.) as on prefer- the view. hatch 6337), general - 4: P. christii over view. niphoboloides (Baker) Morton 6270A, L), Geesteranus lateral view, showing 7: P. on trunks Besides, in 9 of the 31 species of subfurfuracea (Hooker) Ching (Tsai 53252, A), Price spores. epiphytic, found were nummulariifolia (Swartz) Ching (cult. 5831, K), Pyrrosia. number of them large (Giesenh.) Ching (Nooteboom - generally, indicate the most remaining epilithic view. a in species DISCUSSION to assess. (Damask 7, L), lateral the epiphytes indicate that rather of the labels was of and substrate Spores AND - the laesura. 6: Price (Phengklai 1130, L), general view. P. novo- (Humbert 60 BLUMEA Table 1. Substrates Substrates on on which the which - 31, VOL. species (in ]prep.) (in prep, ) No. 1, 1985 in . Pyrrosia in Pyrrosia Species ABCDEFGH A B C D E F G H P. abbreviata P. 4 4 + + africana 11 + + P. P. albicans 4 4 + + P. P. _ — — _ - - _ — _ — as recognized by Hovenkamp Hovenkamp have been found. found. + + - _ — + + - + + 5 5 epiphytic or epilithic 11 epiphytic, epiphytic, sometimes 2 2 epiphytic, epiphytic, occ. occ. epilithic terrestrial or epilithic epilithic P. P. angustata angustata 4 + + — — — - — - 1 1 also epiphytic, epiphytic, epilithic or terrestrial terrestrial P. P. angustissima angustissima 3 P. P. assimilis 2 2 P. P. 4 asterosora P. P. boothii 2 P. P. 5 5 ceylanica P. P. christii 3 P. P. confluens 5 5 P. P. costata costata P. P. distichocarpa P. P. drakeana 1 4 2 2 + + _ - _ - _ - - _ - _ - _ - _ + - + + + + — - - - - — - - — + - - — - + + _ + + _ - — - - — - - - - - - - — — — — + + - _ - + + — 2 2 epiphytic or 1 epilithic, epilithic, occ. 2 2 epiphytic 1 1 epilithic epilithic or epilithic epiphytic epiphytic epiphytic + 2 2 epiphytic + + 2 2 mainly mainly epiphytic + + 3 epiphytic, epiphytic, - + + — 1 1 2 2 1 1 epiphytic or sometimes or or epiphytic, epiphytic, epilithic epilithic epilithic epilithic occ. occ. or epilithic epilithic epilithic epilithic terrestrial, sometimes epiphytic P. P. eleagnifolia 5 + _ — — _ + + + + 2 2 epiphytic or or epilithic, epilithic, sometimes terrestrial terrestrial P. P. fallax 5 + P. P. flocculosa 2 2 + + - - — — - - - — - - + + 1 1 mostly epiphytic epiphytic 2 2 mostly epiphytic, epiphytic, sometimes sometimes epilithic epilithic P. P. foveolata 5 5 + P. P. gardneri 3 + + P. hastata hastata 2 _ — - - - - - + + - _ — + _ — - 4 4 epiphytic, + + 4 4 epiphytic 2 2 mostly epilithic, mostly - _ — occ. occ. or or epilithic epilithic sometimes epiphytic 5 + P. P. kinabaluensis kinabaluensis P. 4 + P. P. laevis 3 P. P. lanceolata 5 5 heterophylla - + - - - - - — - - - _ — - + + - _ — 1 1 epiphytic 1 1 epiphytic, epiphytic, + 3 + 13 or epilithic sometimes sometimes epilithic epilithic probably mostly epiphytic mostly epiphytic, epiphytic, sometimes epilithic P. P. linearifolia 2 + P. lingua 3 3 + - _ — - - _ — _ - + + — 2 2 epiphytic or epilithic 3 epiphytic or epilithic, sometimes terrestrial P. P. longifolia 5 5 + — — - — - — - 2 2 epiphytic, rarely epiphytic, rarely epilithic or terrestrial P. mannii 2 + — — - — - — - 1 mostly epiphytic, epiphytic, epilithic sometimes G.A. Synaptospory Uffelen: Pyrrosia in 61 continued, 1 continued) 1 (Table van Species ABCDEFGH C D E F G A B H P. 5 5 — P. novo-guineae novo-guineae 4 + P. 4 niphoboloides niphoboloides nummulariifolia nummulariifolia - P. pannosa 2 P. penangiana 2 P. petiolosa 3 P. P. piloselloides 5 P. platyphylla 1 P. polydactyla _ - — - - + - - - - - + — - - - — + - + - - - - + - - - — - + — - - - - epiphytic 11 epiphytic 2 2 epiphytic 2 few + 11 + + 3 - ____ - 2 + or epilithic epilithic data, mainly epilithic epilithic mostly epiphytic, mostly epiphytic, epilithic epilithic, rarely rarely epiphytic epiphytic 11 epiphytic, ! 1 epiphytic sometimes or 2 - - - - + + 11 epilithic, 2 + — — - — - + + 7 epilithic P. 1 1 + — + + 6 6 mostly epilithic epilithic terrestrial or epilithic epilithic P. porosa - also epiphytic or sometimes epiphytic, epiphytic, terrestrial terrestrial princeps + — + also also P. rasamalae 4 + — — - — - 11 — terrestrial, or epiphytic epiphytic, occ. occ. epilithic epilithic or on earth banks P. rhodesiana 2 + P. rupestris 5 5 + P. samarensis 4 + P. 1 1 + P. serpens 5 5 + P. sheareri P. 2 + + P. P. sphaerosticha 4 P. P. splendens splendens 1 1 P. P. stigmosa 1 1 schimperiana + + - - P. P. stolzii stolzii 2 + P. P. 2 + A: subfurfuracea Spore type (Van wood. D: studied. which well - half of the image than terrestrial of tris-type between or specimens Pyrrosia elevation of one as + 2 2 epiphytic + 5 epiphytic, also also 1 1 epiphytic, sometimes 2 epilithic, + 3 epiphytic, + 1 few data, - - - - - - + - - - - - - - - - + + - - - - - - - - + - - - - + - - 1 epilithic 4 epiphytic epiphytic 2 epilithic, epilithic, specimen as was the epiphytic - B: were place of some contradicts pronounced sporoderm sculpture and appears a epilithic epilithic epiphytic epilithic epilithic epiphytic occ. epiphytic C: On dead Epiphytic. G: Number of specimens of those on dead wood) (table 1). Nevertheless, growth Kramer's occ. logs Hovenkamp (in prep.). found species and on trees mostly mostly epiphytic or or studied, epiphytic (incl. genus still in press). Unknown. described by epilithic specimens of which in F: epilithic less often less often + - or also also - Hennipman, sporoderm sculpture spores, epilithic, - - preference therefore epiphytic 1 - - E: Terrestrial. an 6 + — - - Uffelen & Epilithic. H: Substrate more as + + — - is known to agree were epiphytes, with the Pyrrosia, especially hypothesis as about the terrestrial mode of life. facts. as over the The in P. rupescorrelation 62 BLUMEA the As in elevation of Pyrrosia, the Kramer's among lariifoliaAs the valid no conclusions attributed his to proposed Cohesion of vation the of sporoderm have taken sculpture. I place Lugardon the I genus (1981) separated from each other perispores from more the two states an closely together position. that in most during development by to the assump- on novo-guineae (fig. 6), with spores that pronounced a and in cohering formation perispore This agrees with the ideas of cristae ferns exception correlated with the ele- in P. three or high, certainly species formerly whole is highly cohering origin of perisporal specimens in some adjacent some species spores of Bolbitis, probably are kind of barrier, studied of the different spore types according Spore type* 4 Epiphytic On dead wood Epilithic Terrestrial % preventing 25 14 40 6 3 19 8 23 1 6 1 3 . - - - - 44 - 12 34 nr. 2 - D - E Total % nr. % nr. % nr. % 12 11 50 21 64 52 42 1 5 2 2 20 16 3 2 - 7 arranged substrate. C % nr. 1 7 to B A nr. Unknown cohere is than so nummu- merging together. Table 2. Numbers of Variable a of synaptospory is based to with the spores in this about the as does specimens only not Pyrrosia found spores fitted perispores that seems of set niphoboloides (fig. 7), species found never epiphytes ar- found are substrate is unknown is very this However, hypothesis P. rasamalae and P. Hennipman (1977) although The clearly more with other types of spores. species which spores tend observed the cases must — to sporoderm sculpture. the but that pronounced sporoderm sculpture (P. and it hypothesis spores. degree P. samarensis, studied of the different spore types, of which the be drawn. can Drymoglossum, rule. tion that than in percentage of specimens confirm Kramer's not specimens spores with very rupestris -type) 1985 differs very much between spore types substrate. These data show to 1, No. may still hold for these smaller groups. Table 2 gives hypothesis species having and P. 31, VOL. sporoderm sculpture numbers and percentages of ranged according - 41 - 1 - 1 6 7 41 - - 5 - 9 41 - 2 - - 6 - 2 6 3 2 8 24 43 35 Total number of * specimens A: P. 16 princeps-type. 35 - nummulariifolia -type . — B: 17 P. E: P. 22 subfurfuracea- type. rupestris -type. - 33 C: P. 123 christii-type. - D: P. G.A. van As the spores that I observed of perispore in twos it is formation, and threes have spores leave the tetrad that in Pyrrosia observed even from with Pyrrosia have lain side must 63 side by the tetrad, same that is way subsequent gametophytes. actually prevents the recombination of unless it is assumed that the supposed Cohesion of spores from the material: within genetic gametophytic selfing (Klekowski, in homozygotic the first although it would In both place. be not ferns combination in as from different tetrads cases, exact result of different copies of the one to same in sporophytes are are the other; inter- selfing plant between that com- produced polyploids (Klekowski, the place, 1973). Re- follow if spores usually only can tetrad tetrad there little recombination would take very synaptospory been rarely identical gametophytes genetically intergametophytic impossible, especially a or between would result in plementary gametophytes spores in 1969) sporophytes, have enhance cross-fertiliza- to pairs of genetically identical spores, each pair complementary would result beginning hanging together pronounced perispore sculpture a haphazard the at mixed up after exospore formation. It follows tion between the two in that many of the spores thus and get figure cohere in the to cohering probable arisen spores Synaptospory Uffelen: involved. CONCLUSION Synaptospory be an accidental amount position as observed in remaining of variation in of this (= Pyrrosia) such should be are more as an serve found in the spores and to exception to his a clear purpose, but regard to Kramer cover not the As the refers already I do seems to perispores. indicates Pyrrosia as hypothesis, It could still rejected. consistent with not intricately sculptured perispore sculpture pothesis groups that of together genus with respect moglossum as does Pyrrosia unique to Dry- suggest the hy- the situation in many other sporoderm sculpture and ecology. ACKNOWLEDGEMENTS I want to aptospory thank Prof. ecology thank by Prof. Prof. Dr. Dr. Dr. C. Kalkman of the E. K. U. Hennipman Kramer for for and for reading drawing my critical manuscript the attention reading of the and Mr. P. to the publication manuscript. Hovenkamp I on syn- also want to for data about the species. REFERENCES DOCTERS VAN Deutsch. GASTONY, LEEUWEN, Bot. Ges. G. J. Series 2, HOVENKAMP, KLEKOWSKI ations. 1973. considerations. 1977. A E. no. Leiden P. Jr, W. iiber Ameisen-Epiphyten aus Java. Ber. in the Cyatheaceae. I. The perine and sporangial capa- Amer. J. Bot. 61: 672-680. monograph 1969. Bot. J. Linn. Genetic load Mitteilung of the fern genus Bolbitis (Lomariopsidaceae). Pyrrosia Mirbel Leiden Bot. Univ. Press. 331 pp. (in prep.) E.J. 1929. Kurze 90-99. Spore morphology 1974. city: general HENNIPMAN, 47: A monograph Soc. 62: in of the genus Reproductive biology of the (Polypodiaceae). Pteridophyta. II. Theoretical 347-359. Osmunda regalis populations. Amer. J. Bot. 60: 146-154. consider- BLUMEA 64 KRAMER, K. U. LUGARDON, phytes. Gard. Bull. 1981. B. Pollen SCHNELLER, Synaptospory: 1977. pteridophytes. J.J. et Les Spores 3. 31, hypothesis. a 1,1985 No. possible A function of spore sculpture in 30: 79-83. globules des filicinees, homologues des corps d'Ubisch des sperma- 23: 93-124. Untersuchungen 1975. filix-mas-Gruppe. Sing. VOL. - an einheimischen Farnen, Okologische Untersuchungen. Teil. Ber. insbesondere Schweiz. Bot. Dryopteris der Ges. 85: 110— 159. TRYON, 3 A. F. 1985. Spores of myrmecophytic ferns. Proc. Roy. Edinb. Soc. 86B: 105-110 + pi. UFFELEN, SEM G. A. VAN study. WALKER, T. G. Pollen 1985. & et E. Spore viewpoint. Proc. Roy. HENNIPMAN. The spores in Pyrrosia Mirbel (Polypodiaceae), a Spores (in press). filaments Soc. Edinb. in 86B: the ant-fern 111-114 + 1 Lecanopteris pi. mirabilis: an alternative