PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN 



perfectae formed from sculptural elements and absence of sculpture 

 outside the contact areas towards the amb and over the distal surface. 



Occurrence. Rare in the Cantabrian Mountains, Argovejo, La 

 Vid. hemiesferica (H) to micrornatus - newportensis (MN) 



Biozones. Pfidoli to Lochkovian. In England (Shropshire & Her- 

 efordshire) where this species occurs in the Lower Pfidoli and spores 

 of this type have been found also in sporangia (Fanning eiai, 1991: 

 3.2 b). Similar spores are found higher in the sequence in the Lower 

 Lochkovian but the higher spores have a variably greater cover of 

 apiculate to spinose sculpture; in extreme forms the spore is com- 

 pletely covered with cones/spines (Richardson, unpublished). 



Remarks. In the case of Form 2 the curvatural bands are defined 

 by sculpture, sometimes barely discernible, sometimes forming 

 broad bands of variable grana and coni paralleling the trilete mark 

 and merging radially into curvatural sculpture. This species, 

 Apiculiretusispora arcidecits sp. nov. occurs in the Lower Pfidoli of 

 England and has been found in sporangia (Fanning et al, 1991). 

 Stratigraphically higher spores are variable but include specimens 

 with more pronounced sculpture. There is considerable variation; 

 variants in the Lower Devonian have dense distal sculpture, whereas 

 the proximal surface is completely covered with cones, or short 

 spines, with only three small inter-radial areas remaining smooth 

 (interradial papillae). A third species, Breconispoiitesl cf. conmadus 

 shows that zonate spores may also have a distinct proximal pattern of 

 sculpture similar to that of A. arcideciis sp. nov. 



Genus BRECONISPORITES Richardson et al. 1982 



Breconisporites? spp. (cf, Retusotriletes coronadus 



Rodriguez, 1 983, pars) PI. 2, figs 5-7; PI. 4, figs 2-A 



This form has equatorial wedge-shaped curvaturae extending 

 equatorially (into a zona?). On the proximal surface there is a band 

 of spines and cones in each of the inter-radial areas. These sculptured 

 bands parallel the curvaturae and join with the sculpture which 

 parallels the lips. The 'curvatural' sculpture is composed of fimbriae, 

 spines and cones. Sculpture size c. 1-6 |im. The proximal central 

 area, within the arcuate sculpture, may be depressed. There is a +1- 

 circular thickening over the distal polar area. 



Remarks. Specimens of Breconisporhes from South Wales (lo- 

 cus typicus) (Richardson et al, 1 982) have a bizonate cingulum and 

 some species have a proximal area of coni; the Cantabrian spores 



141 



have some similarities but are not distinctly bizonate. Rare laevigate 

 or proximally rugulate zonate spores that are more clearly bizonate 

 occur near the top (sample LV 13) of the sampled part of the La Vid 

 Section (PI. 4, fig. 5). In specimens with prominent proximal spines 

 (PI. 4, fig. 3) the distribution of the spines, in arcuate, 'curvaturate' 

 zones, linked to areas of spines bordering the lips, is similar to the 

 pattern of sculpture seen in Apiculiretusispora arcidecus sp. nov. 

 where the sculpture is much finer. The curvatural thickening is 

 extended to form a zona and there are three arcuate bands of spines 

 and cones on the proximal face. The latter spores, and the heavily 

 sculptured spores of type 2, have been found only in the Devonian so 

 far, and form part of a complex morphon, which is being further, 

 investigated. 



Infraturma MURORNATI Potonie & Kremp 1954 

 Genus EMPHANISPORITES McGregor 1961 



Type species. 



fig, 7). 



Emphanisporites rotatus McGregor 1961 (PI. 3, 



Emphanisporites splendens Richardson & loannides 1 979 

 PI. 2, fig. 8, PI. 3, fig. 9 



1973 Emphanisporites? pseudoerraticus Richardson & 

 loannides: 275-76, pi. 3, figs 12-15; pi. 4, figs 1-4, 7. 



1978c Emphanisporites? pseudoerraticus Rodriguez: 417, pi. 2, 

 fig. 17. 



1979 Emphanisporites splendens Richardson & loannides: 111. 



1983 ^Emphanisporites pseudoerraticus Rodriguez: 40, 41, pi. 

 3, fig. 20. 



Dimensions. 26-63|jm ( 1 7 specimens); 50-78|jm (24 .specimens) 

 (Richardson & loannides 1973). 



Remarks. The Cantabrian and North African specimens are highly 

 variable, but the specimens in this study include much smaller forms 

 than any found in North Africa. Also the 'annulus" is sometimes 

 formed of three inter-radial tangential thickenings which together 

 form a triangle resembling a proximal kyrtome. However, in some of 

 the Cantabrian and the previously described North African speci- 

 mens, there is no annulus but only irregular inter-radial thickenings. 

 In other specimens (PI. 3, fig. 8) with an annulus, radial ribs, and 

 polar, or near polar, thickenings that may be proximal, there is no 

 clear trilete mark. These are referred to herein as Emphanisporites 

 cf. splendens. 



PLATE 4 



Fig. 1 Aneurospora richurdsonii (Rodriguez) comb. nov. Tilt45°, Arg97/13/IP; la, BM 2118, oblique proximal compression: lb, BM 2119, showing 



distal coni, x 2000; Ic, BM130290, specimen in oblique compression showing rigid equatorial crassitude, Arg/92/13. 

 Fig. 2 Breconisporites sp. A. Sample La Vid 92/13/1; 2a, BM 132895, proximal view: 2b, BM 132896, distal proximal sculpture, x 2000. 

 Fig. 3 Breconisporites sp. C. LV92/13/1; 3a, BM 132899, proximal view; 3b, BM 132900, spinose proximal sculpture forming zones parallel with the 



equator and paralleling the trilete sutures, x 2000. 

 Fig. 4 Breconisporites sp. B. BM 130860, proximal view showing rugulate sculpture, x 500, LV92/13. 

 Fig. 5 Zonate spore. Proximal view, specimen with laevigate contact face, BM 130859, LV92/I3. 

 Figs 6, 10 Chelinospora cf. hemiesferica (Cramer & Diez) comb. nov. 6, BM 130308, Arg92/ 10; 6a, BM 130307, oblique compression; 6b, detail of radial 



folds around the hilum, x 2000; 10, BM 132469, oblique view with spaced uncluttered distal muri. LP92/19. 

 Fig. 7 Chelinospora cantabrica sp. nov. BM 135304, partial tetrad, upper specimen distal view showing large broad lacunae, lower specimen giving a 



partial proximal view of the equator surrounding hilate area and subequatorial sculpture, Ger92/2B/2. 

 Fig. 8 Chelinospora sanpetrensis (Rodriguez) comb. nov. 8a, BM 136891, distal view showing low muromate sculpture and narrow, occasionally 



branching lumena, Ger92/2B/3; 8b, BM 135042, proximal/oblique view showing granulate - micro rugulate curvatural sculpture, thin broken hilum, and 



subequatorial radial lacunae, Ger92/2A: 8c, BM 1 34063. spore tetrad showing sub-equatorial radial lacunae, sample Arg92/5A. 

 Fig. 9 Chelinospora hemiesferica (Cramer & Diez) comb. nov. BM 1 32687, oblique view, LV92/1 0. 



All figs x 1000, unless stated otherwise. 



