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The Cycads & Cycads Moths of KwaZulu-Natal
Michael Robert Cooper & Douglas Goode (2004)

Size:
Pages: xxii + 98
Pictures: Colour plates from watercolour paintings, maps.
Publisher: Peroniceras Press, 7 Ridge Road, 3610 New Germany.
Price: R250 excluding postage (US $38) from the publisher.
See also: http://www.peroniceras.com/cycads_kzn.htm
 Book review by: Prof. Piet Vorster, Department of Botany & Zoology, University of Stellenbosch, Private Bag X1, 7602 Matieland (As published in ENCEPHALARTOS)
I have always welcomed new books in these pages, perhaps because there were no books when I first became interested in cycads. This latest offering is co-authored by long-time Society member Douglas Goode well known for his previous books Cycads of Africa (1989); The cycad collection volume 1. Natal Province (1996); and Cycads of Africa, vol. 1 (2002, reviewed in ENCEPHALARTOS 69: 35—36, March 2002). However, this time he is neither first author nor, apparently, illustrator, that position being taken by Cooper, "Emeritus Professor of Geology at the University of KwaZulu-Natal and author of numerous scientific publications, particularly in the fields of palaeontology and evolutionary biology". He is not a member of this Society.

KwaZulu-Natal is not a discrete phytogeographical entity, in fact it is phytogeographically very diverse; but its inhabitants have always had a strong sense of provincial identity which is reflected in a number of natural history books covering the region. This is another one.

This book deals with both cycads and with cycad-associated moths of the family Geometridae. Yet this is no comprehensive treatise on cycad-associated insects or even moths. The all-important weevils, several of which are implicated in pollination, are only mentioned in passing, and on page 15 there is a drawing of the larva of Bunaea alcinoe which is said to be restricted to east Africa and thus has no bearing on the subject of this book. Apart from a brief mention in the introduction, the moths are treated separately on pages 76—93, some 9 species in all.
This section is reviewed further below.

The book commences with a preface, reminding the reader about the antiquity and present-day peril of wild cycads. There is a short explanation of this book, which explains how to interpret the symbols in the treatments of species. There is a strongly worded section on the evolution and classification of African cycads (about which more below) followed by description of new genera and species. These take into account the whole of the former Encephalartos, not only the KwaZulu-Natal species. Collectors will be delighted by the five new species to be collected. Of the xvii + 75 pages devoted to the cycads, the first 27 comprise an introduction, dealing with roots, stem, leaves, cones and seeds, pollination, reproduction, differences between sexes, seed dispersal, variation, hybrids, and age (longivity?), in general. There is a quite extensive treatment of the main vegetation types an climatic conditions in which cycads occur, but that has little bearing because cycads tend to grow in local mini-habitats quite atypical of the broad vegetation types in which they may be.

Some 19 species of cycads are treated. For each species there is a full-page watercolour painting of a whole plant with leaf detail. The cones are not illustrated on the plates, but those of some species are scattered through the introduction (pages xv—25), unfortunately without reference in either the main text or index.. The treatment of species is somewhat unorthox, appearing under the headings size, habitat, cone, habit, general identification, and conservation (in that order). The page is headed, not by the scientific name but by a vernacular name, which would be unfamiliar to most readers. For each species there is also a distribution map (covering KwaZulu-Natal only) plus a diagram showing the time of pollen- and seed release. Lastly, for each species vernacular names are provided in English, Afrikaans, German, French, Portuguese, Spanish, and Italian; but not in Zulu. Encephalartos aplanatus is claimed to occur in KwaZulu-Natal and included. Xantostrobus lebomboensis (Encephalartos lebomboensis) is included even though it is not known from KwaZulu-Natal. Encephalartos caffer, which occurs in the far south of the area covered, is not included.

The section on cycads is followed by one on the moths. The format is much the same as for the cycads, and the text for each species is under the headings eggs, larvae, pupa, larval foodplant, habitat, habits, identification, and conservation, in this order for reasons understood by the authors only. Again vernacular names in 7 languages are supplied. For every species there is a colour plate, which in some cases depict all the stages but in other cases only the adults.

There is a bibliography, not at the end but on page 27. At the end of the book there are no less than 8 separate indices, to scientific, English, Afrikaans, German, French, Spanish, Portuguese, and Italian names of both cycads and moths.

There is no glossary to explain how to distinguish pinnacanth and spine.

There is a list of subscribers on page vi, including local as well as overseas supporters. Apparently the prospectus was circulated rather selectively, certainly not to our members in general, and I for one have been completely unaware of the compilation of the book.

Goode is a professional artist and synonymous with paintings of cycads. Remarkably most if not all the paintings in this book are not by Goode (whose contribution is not clear), but by Cooper. They are really very good, but they do not include the all-imporant cones. They are therefore more decorative than informative. There are four different plates for Xanthostrobus natalensis (Encephalartos natalensis), each headed by a different fanciful vernacular name and each with its own set of vernacular names in 7 languages. I presume this is to show variation, but the variation in the leaflets could have been shown in a single plate.

The duty of a reviewer is to provide prospective buyers and users of a book with guidelines as to whether they can expect to find the work useful or otherwise. In this case we are confronted with, firstly, a splitting of the genus Encephalartos into 11 genera, and secondly five new species.

The authors claim (page ix) that the current classification of Encephalartos is not phylogenetic, i.e. does not reflect evolutionary relationships. That is quite true, not only for the cycads but for most of our plants. We can group species together on their similarities, but we are aware of the pitfalls and we don't really know how the evolutionary processes took place. Only two modern classifications, if they can be called that, exists: firstly that by Dyer in Bothalia 8: 405—515 (1965) which has been followed by subsequent writers of "popular" books without understanding the issue, and secondly that by Vorster in Walters & Osborne's Cycad classification concepts pages 69—83 (2004) in which available information but especially spatial information was incorporated. We hope that molecular data will cast more light on this scheme in the not too distant future, but we are not yet there. To rectify this deficiency, Cooper & Goode split Encephalartos up into 11 different genera which they suggest are natural groupings and sufficiently distinct. These are enumerated and discussed below.

To justify this act, they claim (page xi) that the Linnean system of classification is flawed because it only shows basal branching, while Darwin's method (which they claim to apply, whatever that may mean) is phylogenetic and shows all the fine branching. They also claim that Linnaeus worked on similarities and Darwin on differences. This is plain nonsense. Taxonomy is about similarities which is why we place all the African cycads (apart from Stangeria) together in the genus Encephalartos, rather than place every species in a separate genus. Furthermore, the Linnaean system of classification has been universaly obsolete for a very long time. We don't use any Linnaean classification system, but we do use the Linnean system of nomenclature. Darwin has never proposed any classification system. Cooper and Goode dogmatically state (page xi) that the "Toothed cone scales" of the genus Encephalartos in the sense of Cooper & Goode is of evolutionary significance, but provide no evidence.

The first question to ask, is whether the authors were successful and persuasive. They grouped species together within new groups, but completely neglected to motivate their groupings. They provided no clue to the perceived evolutionary relationships within or between the genera, and therefore failed in their proclaimed intention. The second question is whether the subdivision of Encephalartos is justified. It is often said that delimitation of genera and species is an arbitrary matter, subject to the personal opinion of the taxonomist involved. It is certainly not so simple, but there is some truth in this perception, within limits. The real question is to what extent the new genera are defined, and whether they demonstrably differ significantly enough to warrant separate taxonomic status. For each of the genera which they recognise, a description is provided but no indication of how it differs from others, and neither is there a key. This makes it quite impossible to understand how they distinguish these genera from each other. For instance, Encephalartos longifolius, E. arenarius, and E. latifrons are grouped together in the genus Rugostrobus, whereas E. princeps, E. lehmannii, E. trispinosus, and E. horridus are placed in the genus Acanthozamia; but it is totally unclear on which grounds they distinguish these groups as separate genera from Encephalartos, not to say from each other. Earlier on they make an issue of shared and derived characteristics, and in the first two genera treated (Encephalartos, Dracostrobus) several characteristics are thus indicated without any reason why, for instance, “woody [sic] to lightly woolly bracts” should be primitive and “dense golden brown tomentum” be derived.

The genus Encephalartos in the sense of these authors contain only E. villosus, E. aplanatus, E. cerinus, E. ngoyanus, E. caffer, and E. umbeluziensis.

Their genus Dracostrobus contain D. brevifoliolatus, D. cycadifolius, D. friderici-guilielmi, D. ghellinckii, their new D. mkomaasiana, D. humilis, D. lanatus, D. laevifolius, and their new D. dedekindii.

E. ferox is placed in a genus called Pyrrostrobus. It does seem as if E. ferox does not have close relatives, yet they undermine their argument by stating that it is close to E. arenarius, for which there is no evidence at all.

The genus Xanthostrobus contains X. natalensis as well as X. altensteinii, X. aemulans, X. msinganus, X. woodii, X. lebomboensis, X. senticosus, plus three new species viz. X. currachii, X. pietretiefii, and X. mgeniensis . Inexplicably E. transvenosus is excluded, yet it is morphologically so similar to E. natalensis.

The genus Inezamia contains I. paucidentatus, I. heenanii and I. relictus, but also I. transvenosus (Encephalartos transvenosus) which should be grouped with E. natalensis.

The genus Glaucostrobus contains G. eugene-maraisii as well as G. middelburgensis, G. dolomiticus, G. dyerianus, G. hirsutus, G. nubimontanus, and G. cupidus. None of the evidence provided distinguishes it from their Acanthozamia (q.v.). They make the extraordinary but unmotivated statement that “its affinities seem to lie with Dracostrobus”.

Dyerstrobus inopinus is the sole species in the genus. Again it is suggested, without any evidence, that it is most closely related to Dracostrobus.

The genus Viridestrobus contains V. manikensis, V. pterogonus, V. concinnus, V. chimanimaniensis, and V. munchii.

All the other tropical African species with well-emergent trunks are grouped together in the genus Tanzamia, while Congostrobus includes C. poggei, C. delucanus, C. schaijesii, C. marunguensis, C. schmitzii, and (surprisingly) C. barteri.

Should the reader use these generic names? At least some of them do reflect natural groupings according to our latest information, but in my opinion the differences are far too small to justify the recognition of separate genera. Acceptance of these genera are likely to make it very difficult to decide in which genus an unknown plant should be placed. Moreover, these names are not valid according to the International Code of Botanical Nomenclature. The authors didn’t do their homework.

At the CYCAD 2005 conference in Mexico I will discuss the evidence supporting this grouping of species, and the merits of the generic proliferation.

The five new species, which are described, are probably of more interest to our readers, especially to collectors yearning for names for all their plants.

Firstly, Dracostrobus dedekindii is the name given to a single plant from near Tugela Ferry, hitherto assigned to Encephalartos laevifolius. Plants from the Port Edward area, some years ago removed by the then Natal Parks Board amid much publicity and now practically extinct, are also included here. I have previously examined material of both, and concluded that both should be referred to E. laevifolius. It is very regrettable that no plant from either locality is in any public collection, because phytogeographically these outlying populations are interesting, and molecular studies of these could probably tell us a lot about the past history of this species.

Also in Dracostrobus, D. mkomaasiana is based on what was hitherto known as the coastal form of Encephalartos ghellinckii. If one looks only at the two extreme forms, the foliage is recognizably different, though the cones are practically identical in spite of what the authors claim. However, the matter is not so simple because of geographically and morphologically intermediate populations. Their respective maps of the distribution of Dracostrobus ghellinckii and D. mkomaasiana show a very small break between the distributions of the two, but the map on page 5 of Scott-Shaw’s Rare and threatened plants of KwaZulu-Natal (1999) shows the distribution to be continuous. I consider all the plants to be part of a continuous variation and distribution, and there is insuficient evidence for the recognition of taxonomically distinct entities at any rank.

Xanthostrobus curachii is based on plants from the Ubombo Mountains. Furthermore, the Pongola valley plants hitherto placed in Encephalartos lebomboensis are described as yet another new species, Xanthostrobus pietretiefii. I cannot distinguish these two from Encephalartos lebomboensis or from each other by means of either the diagnoses or the illustrations, and consider the supposed distinctions to be spurious.

Lastly, the fifth species, Xanthostrobus mgeniensis, is a rather interesting thing. Plants have been traded under the name Encephalartos umgeniensis, and this is apparently what we have here. The problem is that the origin of the plants is unknown, all being in cultivation. Furthermore, all are males, and one can but wonder whether they aren’t all suckers of the same plant. The cones are indistinguishable from those of Encephalartos natalensis, but the leaflets have an arresting appearance: they are broad, quite blunt at both ends, heavily toothed, and the surfaces are wavy rather than flat. They are not unlike those of the so-called Highflats population of Encephalartos natalensis depicted in Goode’s Cycads of Africa volume 1: 128 (2001) and which is not mentioned in this book. What is to me really interesting, is that the leaflets of this X. mgeniensis are so very similar to at least the subadult leaflets of the so-called “false Encephalartos woodii”, likewise known only in cultivation and only from male plants. I am curious as to why the latter is not mentioned at all. While very interesting, I am not convinced that it is prudent to describe a new species from material known only from cultivation. I also feel that much more work should be done before it can definitely be recognised as a separate species. One only has to look at the bewildering range of leaflets shapes and leaflet orientations in the widespread Encephalartos natalensis depicted in Grobbelaar’s Broodbome pages 250 to 257 and including material from Highflats and of X. mgeniensis (as “Umgeniensis”), but especially his figure 6.29.2, to realise that it is irrational to single out this X. mgeniensis as specifically distinct. There is a map showing nothing, because the natural origin of the plants is unknown, though they are suspected of coming from the Mgeni River.

Regrettably these specific names are also invalid, and as detailed below, some of the epithets are gramatically incorrect.

We therefore have a set of fine decorative illustrations which in the case of many species don’t show the reproductive structures. We also have new generic and specific names which are not only technically invalid or incorrect from a nomenclatural point of view, but are also based on characteristics which make it impossible to reliably key out the species. Lastly the presentation is of a poor standard:

The table of contents is not representative of the text. For instance, it refers to the evolution and classification of African cycads, but does not indicate where new species are described. It refers to, amongst others, Genus Dracostrobus gen nov., but the generic description is not on the indicated page 32.

In the bibliography many titles have no bearing on the KwaZulu-Natal species, especially as there are no literature references in the text. Some very obscure titles are included, while extremely important ones such as Chamberlain (The Living Cycads, 1919), Norstog & Nicholls (The Biology of the Cycads, 1997), and Whitelock (The Cycads, 2002) are ignored. Hutchinson (1945) does not exist, and the reference to Vorster (2003) on page xi probably refers to Walters & Osborne (2004, not 2003). Phiney should read Pinhey. Read & Stolt should be Read & Solt. The reference to Stapf & Burtt Davy should read A manual of the flowering plants and ferns of the Transvaal with Swaziland, South Africa. Claasen should be Claassen. Excelsia should read Excelsa. Grobbelaar is consistently misspelt, also in the main text.

 

The date of publication is given as 2004. Because of the principle of priority, one of the six pillars of the International Code of Botanical Nomenclature, it is customary to indicate the date of publication to the day in works in which new taxa are published. The new genera lack a Latin description and are therefore invalid. This means that the new new species in these genera are also invalid, and the new combinations are invalid because they lack references to the basionyms. In the case of species it is not stated in which herbarium the type is lodged.

Throughout the book species are glibly referred to as e.g., woodii. That is incorrect: the name of a species consists of a combination of the correct genus name with the oldest available epithet, and should always be used thus. It is also wrong to refer to a name as a sp. nov. [new species] in any place other than where it is originally described. It is incorrect to cite authors of names in recombinations as, for example (Bertoloni) [page 72]. The name of the author who first published the epithet should be between brackets, followed by the author who published the recombination, thus (Bertoloni f.) Cooper & Goode.

On page 8 it is stated that species of “toktokkies” [Tenebrionidae] are amongst the pollinators. That is news to me. It is also claimed that the weevil Metacucujus goodii is a pollinator of Xanthostrobus natalensis (i.e. Encephalartos natalensis) but I am not aware of any published proof of this assumption. On page 10 it is claimed that the sclerotesta of the seed is poisonous – I presume they mean sarcotesta. Stroboli on page xii should read strobili. Pinnules on page xx and elsewhere should be pinnae – there are no pinnules in African cycads.

It is stated (page 10) that Cycas thouarsii is 140 million years old: I would like to know on what evidence this is based, as molecular research on three continents showed that that all living cycad species which were examined are less than one million years old.

The International Code of Botanical Nomenclature allows names of genera and species and genera to be taken from any source whatsoever, or even to be made up arbitrarily. As such the names used in this book are not wrong, but rather quaint. Several of the new generic names in this book ends in –strobus, which has no significance – strobus means something which rotates. Do they perhaps mean –strobilus, pertaining to cones? As such Xanthostrobus would mean “yellow cone”, but Congostrobus would mean “cone from the Congo” which is a bit silly. Dyerstrobus and Viridestrobus are gramatically wrong. Some of the specific epithets are also strange: pietretiefii (page xviii) means “of [the man] Piet Retief”. Now why would they be so politicaly incorrect to name a plant after a Voortrekker leader eliminated by Dingaan 170 years ago and who, moreover and contrary to Recommendation 20A.1.h of the Code (which alo applies to specific epithets), had no connection with botany? Dracostrobus mkomaasiana should be D. mkomaasianus.

Vernacular names are those used by people who utilise plants. In practice useful or damaging plants have vernacular names, but those not used for any purpose don’t have vernacular names. There has been a deplorable move by some botanists to invent vernacular names for plants, apparently in an attempt to gain public popularity. This cannot be condemned too strongly: firstly, such invented vernacular names make it impossible to distinguish the genuine vernacular names, and secondly such invented vernacular names are unlikely to be adopted by real enthusiasts – I have never met a cycad enthusiast who couldn’t memorise and use names such as Encephalartos friderici-guilielmi or Lepidozamia peroffskyana. In this book the authors go overboard: not only are vernacular names invented for every species, but these are in English, Afrikaans, German, French, Portuguese, Spanish, and Italian – all spurious -, but there are no vernacular name in any Bantu language. These queer names head the treatment of individual species, and the scientific names are in the smallest type on each page. No synonyms, i.e. equivalents in Encephalartos, are given.

I am not against the inclusion of the cycad moths, however unwelcome these pests may be in the garden. These include one attempted new species, again invalid. The very brief text neatly summarises the information and the plates are a delight. I remind the reader of the excellent, if brief, chapter African cycads and cycad moths by Staude on pages 307—311 of Goode’s Cycads of Africa, volume 1. Again every species has a string of spurious vernacular names in every conceivable language.

In conclusion, this book is an irresponsible, incompetent, and negligent attempt to upset current nomenclature and taxonomy, and adds unnecessarily to the synonymy. It looks like a deliberate attempt to commit damage, and I cannot accept either the attempt at classification or the nomenclature. This is an example of where people, however impressive their qualifications outside the field of plant taxonomy may be, try to emulate the deceptively simple-looking results of plant taxonomy without understanding the philosophy behind it.

The International Code of Botanical Nomenclature (Articles 32.8 and 32.9) makes provision for a list of suppressed works, publications which are to be ignored. This book comes close to being a candidate for inclusion in that list.

Piet Vorster

Department of Botany & Zoology, University of Stellenbosch, Private Bag X1, 7602 Matieland.

 

  
 
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