Copiapoa - Living on the Edge
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Cactaceae Systematics Initiatives

11: 23 - 29

Coping with Copiapoa

David Hunt

A taster for the 'Copiapoathon‘ to be run at the ICSG meeting in June

On the face of it there are significant differences of taxonomic opinion between various recent authors with field experience of this difficult genus. An analysis of these differences highlights several problematic groups where collaborative study of the problems could be helpful.

 

For the two published editions of the CITES Cactaceae Checklist, I have followed the advice of Nigel Taylor, who published a commentary on the genus some twenty years ago (Taylor 1981), illustrated with excellent half-tone photographs of plants in cultivation by Graham Charles, who also contributed notes on cultivation. Taylor (1989) subsequently authored the treatment of Copiapoa in volume 3 of the European Garden Flora and studied a good range of Copiapoa taxa (c. 70%) in Chile in June 1990. Graham Charles‘s own account of the genus, now beautifully illustrated in colour, was published three years ago (Charles 1998), after several visits to South America.

 

Meanwhile, Adriana Hoffmann published her illustrated guide to the cacti of Chile, attractively illustrated with drawings by Andrés Julliän (Hoffmann 1989). Adriana was helped by Roger Ferryman and Fred Kattermann, both of whom have made numerous field-trips to Chile over many years.

 

Next in line will be Fred Kattermann himself, whose revision and amplification of the other major Chilean genus Eriosyce was the first volume of Succulent Plant Research (Kattermann 1994), and whose treatment of Copiapoa will be the centrepiece of his new book, The Cacti of Chile, which he has asked me to edit and publish as a future volume in the SPR series.

 

Nigel Taylor‘s 1981 commentary appeared shortly after Ritter‘s treatment of the genus for the third volume of his Kakteen in Südamerika (Ritter 1980). Taylor credited Ritter with providing useful descriptive and distributional information, and with grouping the species into informal sections ,,which, by and large, make sense“, but thought many of the 36 new species Ritter had described (from 1959 onwards) were ill-defined. Taylor listed them all, but accepted only a quarter of them, along with various earlier-described species, in a total of 17. He also provided a map indicating their distribution and that of 37 other proposed species and varieties.

 

By the time of the second edition of the CITES Cactaceae Checklist (Hunt 1999), various species previously accepted, or not accepted, could be included as subspecies, and Taylor was inclined to accept a few more of the proposed species, making a total of 20 species plus 4 heterotypic subspecies.

 

In her account, Adriana Hoffmann also accepted 20 species; Graham Charles reduced the tally to 13 species hut admitted 17 heterotypic varieties; and in the manuscript for his book Fred Kattermann has proposed accepting 25 species, two of them (C. hypogaea and C. laui) each with one heterotypic subspecies.

 

Thus, disregarding the more liberal treatments of Ritter and Backeberg (c. 45 species), we have widely varying species-totals but some sort of consensus that there are 20—30 recognizable taxa (species + subspecies or varieties).

 

I have attempted to express some of the differences between the three most recent treatments mentioned in the form of a series of questions, and submitted them to the three protagonists, Graham Charles (GC), Fred Kattermann (FK) and Nigel Taylor (NT). Their responses (in some cases somewhat abbreviated) are given below.

D.H.

1. C. atacamensis. Is this really distinct from C. calderana, either as a species or as a subspecies?

GC: I chose to keep the two apart because of the large geographic separation. They are difficult to tell apart by description so subspecific status for one or synonymy would be botanically sensible.

 

FK: lt is not distinct from C. calderana and even less so from marginata, cinerascens, coquimbana, esmeraldana, fiedleriana, grandiflora, marginata, megarhiza and taltalensis. All these could be treated as one species, but you cannot just have C. atacamensis merged into C. calderana without merging them all. C. megarhiza and C. esmeraldana could be separated as a subspecies by their flower shape and C. cinerascens and C. grandiflora by their small stigma-lobes. In the US most of the group flower in spring, but C. cinerascens, C. grandiflora and C. megarhiza flower in midsummer (flowering time of cultivated C. esmeraldana not known). [Fred‘s statistical analyses of floral characters suggest N—S clinal variation in the group as a whole. There seems little variation in the seeds. D.H.]

 

NT: I agree with Graham that C. atacamensis is difficult to tell apart by description so subspecific status might be preferable. I cannot agree with Fred‘s view and wonder what he uses that long list of names for.

2. C. bridgesii. Is this sufficiently distinct from C. marginata, or better treated as a subspecies?

GC: These two are very similar and grow in a similar habitat not far apart. I remain uncertain that C. bridgesii is correctly applied by Ritter to the plant north of Chañaral.

 

FK: C. bridgesii can be included in C. marginata. lt is a very localized population, only a few km further N than the last population of C. calderana (var. longispina). The anticlinal wall of the seeds is more bowed than straight. Is this a good character? I must also point out that Pfeiffer‘s picture has nothing to do with the plants Ritter calls C. bridgesii but is a form of C. echinoides.

 

NT: Reference to my comments on the application of the name C. bridgesii (Taylor 1981: 50) and Fred‘s remarks raise doubts about the correct application of the name. lt may be best abandoned, since Pfeiffer‘s illustration is more relevant than Ritter‘s neotype, which could easily be overturned. Taxonomically, I agree that differences with C. marginata are more conspicuous than significant, but I recall that one of the plants (?,C. bridgesii‘) has mucilaginous stem tissues, while the other (C. marginata) does not. If we agree that these two entities are close enough, then the solution might be to redescribe the ,C. bridgesii‘ as a subspecies of C. marginata typified by a modern collection corresponding with horticultural usage of the name.

3. C. conglomerata. Should this name be rejected as ambiguous? lt is referred to C. solaris in CCC2, but Kattermann takes it up for C. varispinata.

GC: I followed the view that C. conglomerata is uncertain so should be discarded. You know that my view is that Ritter‘s C. varispinata is a variety of C. humilis and that there are two species of Copiapoa at Botija which are undescribed. The one on the beach is a miniature variety of C. cinerea, the other ,sp. Botija‘ is very different from any other species in a number of important features notably the unique method of branching. I climbed to the top of the coastal mountain and examined a large population of this unique plant. (I could show you pictures at the June meeting if you like.)

 

FK: The description of C. conglomerata fits C. varispinata. I suspect Philippi recorded a suspect latitude. His map does not show Chaguar de Jote but his report suggests that it is Q. Botija at about 400 ft alt. He does report seeing an Echinocactus in the coastal sand but nothing else. I believe that the name applies and could be used.

 

NT: I would abandon the name C. conglomerata and under no circumstances re­apply it to something as clearly identifiable as C. varispinata (which I have seen near Q. Botija and is rather unusual more like a Parodia than a Copiapoa!). C. conglomerata‘s protologue data are confused and most important, there is no type or illustration to guide us. No good will be served by trying to apply it now. I agree that there are two undescribed taxa in Quebrada Botija. Graham is right that one is a miniature relative of C. cinerea, near the mouth of the Quebrada. The second is higher up and not common except in one part to the north side, where the colonies of multi-stemmed individuals have to my eyes a close relationship with C. marginata, though nonetheless specifically distinct. I studied both and the other three species found therein in June 1990.

4. C. fiedleriana. Is this distinct from C. coquimbana, or better treated as a subspecies?

GC: C. fiedleriana is certainly distinct from C. coquimbana, intermediate with C. echinata perhaps. lt has a very limited distribution and may be a hybrid?? lt has tuberous roots like C. echinata although this not a consistent character in some Copiapoa populations so may not be a way of separating it from C. coquimbana.

 

FK: C. fiedleriana first the overall C. marginata complex. Lumping or splitting? I studied many populations trying to reach clear and precise differences. There are none. In Eriosyce I set myself the goal to find at least three solid characters to separate species (a few have only two and a half) but this does not work in Copiapoa.

 

NT: I am unsure what C. fiedleriana is.

5. C. laui. Is this distinct from C. hypogaea, or better treated as a subspecies?

GC: I followed Adriana Hoffmann‘s view that it is a variety of C. hypogaea; this was supported by Roger Ferryman finding an intermediate form at Pan de Azucar. Other than size, the two are very alike.

 

FK: I do not have a strong argument against it being a variety except that it is easily identified just by its small size and it is the only Copiapoa where the side branches form their own roots.

 

NT: I remain convinced that C. laui is close to C. hypogaea, but whether you lump them (as subspp.) or not is a matter of choice. However, it is important to be aware that at Pan de Azucar there are two different plants mixed up together. One (the smaller of the two) is C. laui, or a form of it, but the other is a dwarf geophytic form of C. humilis. They have very different flowers corresponding to those typical of these unrelated complexes.

6. C. longistaminea. Is this a good species or a subspecies of C. calderana or C. cinerea?

GC: I am now inclined to think that C. longistaminea is probably a good species rather than a variety of C. cinerea. lt grows together with C. cinerea columna­alba without hybrids which supports this view. There is a also a case to argue that columna-alba is synonymous with the type, being just the southern form of C. cinerea. C. longistaminea also has tuberous roots suggesting an affinity with C. calderana rather than C. cinerea.

 

FK: C. longistaminea does not belong to the C. marginata group. The seed suggests a close relationship to C. cinerea and the flowers are cinerea-like. lt could be merged into C. cinerea without recognition of a subspecies.

 

NT: My views on C. longistaminea should be obvious since I combined it with C. calderana. There is no question of it being anything to do with C. cinerea, for the same reasons as Graham observes. However, it looks just like a fat C. calderana and has mucilaginous stems like that taxon. I do not subscribe to Fred‘s very broad species preference/relationships.

7. C. malletiana. Should this problematic early name be rejected? Its original application was arguably to a form of the later-named C. cinerea, so unless treated as separate species it would displace that well-known name.

 

GC: I followed Nigel in rejection of C. malletiana but I understand that it could have been the first description of the prominent and easily found C. dealbata. I was probably wrong to treat this as a variety of C. cinerea since I am told it has rather different seeds which I did not know at the time. C. malletiana could be resurrected for this species without threat to the popular name C. cinerea.

 

FK: C. malletiana has nothing to do with the C. cinerea group. The seed suggests a stronger relationship to the C. marginata group and of course to C. echinoides. They are very different from those of C. cinerea. Thomas Bridges travelled by ship from Coquimbo to Caldera, then back to Coquimbo via Huasco. Carrizal Bajo is a natural harbour between Caldera and Huasco. Most ofhis collection numbers are from the Copiapo valley. he did not record anything from Chanaral or further north of Chanaral where the distribution of C. cinerea starts. If you do not like the name C. malletiana, C. dealbata is OK with me.

 

NT: As I suggested in 1981, the application of this name is not so uncertain as some might think and knowledge of Bridges‘s itinerary and Salm-Dyck‘s protologue argue for C. dealbata. If C. dealbata (syn. C. carrizalensis) is treated as a good species I would advocate the use of C. malletiana as the oldest name, but if it is not distinct from C. cinerea then I would hesitate to displace this better-known but younger name. Perhaps the best solution would be to conserve C. cinerea over C. malletiana or reject the latter as a source of potential nomenclatural instability.

8.  C. montana, C. olivana and C. rupestris. Are these distinct from C.taltalensis?

GC: I have no more information about this since I wrote my book. The most difficult thing for me was differentiating forms of C. hypogaea from forms of C. humilis since the two are very similar. I cannot include C. rupestris with these because of its ability to grow into huge mounds (C. desertorum). The other names you list are all similar, like humilis or hypogaea. The Quebrada Oliva would appear to contain plants like humilis and hypogaea and both have been collected and distributed as C. olivana, I think a name best left as a synonym of C. montana.

 

FK: C. montana, C. olivana and C. rupestris are close to C. atacamensis and C. taltalensis. Graham Charles illustrates two different plants. His fig. 79 is what grows north of Taltal and fits Ritter‘s description, but his fig. 80 is not the same and could be C. humilis.

 

NT: I‘m not sure what the other protagonists here think, but for my money C. montana, C. olivana and C. grandiflora are all closely related and might be the same thing in more or less geophytic forms (C. hypogaea is perhaps also allied). C. rupestris for me is a member of the C. marginata complex, like the undescribed plant referred to above, from Botija.

9. C. taltalensis.  Is this distinct from C. humilis?

GC: C. taltalensis is a C. humilis form as far as I can tell from description and pictures. I have not seen this plant in nature.

 

FK: C. taltalensis, which has ribs, is distinct from C. humilis, which has its ribs completely dissolved into tubercles. If you want to merge C. taltalensis with something it has to be C. marginata.

 

NT: C. taltalensis, was, I thought, a clear member of the C. humilis complex, as defined by Ritter, but have I missed something? What does Fred know by this name?

10.  C. tenuissima and C. tocopillana. Are these distinct from C. humilis?

GC: My view remains what I published, that they are both C. humilis forms although C. tenuissima looks more like C. hypogaea which I would have considered as the species if not for the geographic separation.

 

FK: Both could be treated as subspecies of C. humilis. Carrying this a little further, C. hypogaea (incl. C. mollicula) could be another.

 

NT: C. tenuissima and C. tocopillana are undoubtedly related to C. humilis. lt is merely a matter of taste or convenience as to whether you lump them or keep them as species, but either way the former still awaits a valid name. I fundamentally disagree with Fred‘s assertion that C. humilis and C. hypogaea are related. As noted above I believe they grow together and don‘t intergrade/interbreed and their flowers are unmistakably different, which is in itself unusual in a genus with so little floral diversity. Rather, I suspect that each species is allied with a different larger-growing relative, i.e. C. hypogaea with C. montana l C. cinerascens and C. humilis with C. tocopillana l C. varispinata etc.

Proposals for updating the Checklist

  1. C. atacamensis. Recognize this as a subspecies of C. calderana.
     

  2. C. bridgesii. Redescribe the false, C. bridgesii‘ as a subspecies of C. marginata with a modern type corresponding to horticultural usage of the name.
     

  3. C. conglomerata. Reject this name.

  4. C. fiedleriana. Further study/discussion needed, and of the synonymy listed in CCC2 and C. megarhiza.

  5. C. laui. Reduce to subspecies of C. hypogaea.

  6. C. calderana ssp. longistaminea. No change.
     

  7. C. malletiana. No change at present, but reject the name if the taxon is subsumed in C. cinerea. (The preferred name would be C. cinerea ssp. dealbata (Ritter) Slaba 1997/Kaktusy 33 (speciäl): 3.)

  8. C. montana (syn. C. olivana) and C. rupestris (syn. C. rubriflora). No change at present. (The question was prompted by Fred‘s proposal to include all these, plus C. hornilloensis and C. desertorum in C. taltalensis).
     

  9. C. taltalensis (form of C. humilis). No change at present.
     

  10. C. tenuissima and C. tocopillana. Validate the former and reduce both to subspecies of C. humilis.

References

CHARLES, G. (1998) Copiapoa. The Cactus File Handbook 4. Pp. 80. Southampton: Cirio Publishing Services.
HOFFMANN J., A.E. (1989) Cactäceas en la flora silvestre de Chile. Santiago: Fundación Claudio Gay. [Pp. 94—131]
HUNT, D. (compiler) (1999) CITES Cactaceae Checklist, ed. 2. Pp. 315. Richmond, Surrey: Royal Botanic Gardens Kew.
KATTERMANN, F. (1994) Eriosyce. The genus revised and amplified. Succ. Pl. Res. 1.
RITTER, F. (1980) Kakteen in Südamerika 3. Spangenberg. [Copiapoa: pp. 1044—1107]
TAYLOR, N.P. (1981) A Commentary on Copiapoa. Cact. Succ. J. Gr. Brit 43(2/3): 49—60.
 --- ,, ---  (1989) Copiapoa. In Walters, S.M. et al. (eds), The European Garden Flora 3:252—255.Cambridge University Press.
 

© 1998-2006  David Hunt and individual contributors

All material, except where otherwise credited, is Copyright
 © 2001-2006 Paul Klaassen

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