George Lindsay. The Genus Ferocactus.

[Лиана]

George Lindsay
The Genus Ferocactus

[admin]

из заслуживающих доверие специалистов я могу назвать лишь Лау

А как же - George Lindsay - http://www.cactuslove.ru/lit.php?id=11
Или это тоже афтар?

[petrovna]

Не читал. Суть не в имени, для кого-то и вон тот украинский тип, дай бог памяти... в защиту шыдевра которого чуть вторая оранжевая не случилась... тоже великий специалист )) Вспомнил - Буренков ! Читает потом очередной космонафт такую книженцию и с полной уверенностью доказывает, что ариокарпус чисто Бразильский эндемик Мне вот тут недавно в личку письмо написали(он себя узнает )) ) , так вот интересуется народ - где купить семян "правильных ариакактусов" , а я стесняюсь, незнаю что и ответить... такой род мне не известен... отстал...

[admin]

Не читал. Суть не в имени, для кого-то и вон тот украинский тип, дай бог памяти... в защиту шыдевра которого чуть вторая оранжевая не случилась... тоже великий специалист )) Вспомнил - Буренков !

Так господин Буренков вроде сам лично написал, что не претендует на монографию и что его книга это популярная литература. Так что шпынять его по каждому поводу и без поводу не стоит.

А вот многорафию George Lindsay о Ферокактусах - я лично читал и даже имею в наличии. Все никак не найду кто сможет ее перевести, а то бы уже давно выложил на сайте.
На мой взгляд, во всяком случае из всего того что мне попадалось о роде Ферокактус - это лучшая книга.

[vlani]

Разве Пилбим претендует на монографию? По моему обычный популяризатор, довольно добросовестный. Правда по его книжке всерьез и определить то растения не получится. Про классификацию он по моему вообще ничего не говорит - просто типа рассказывает что как называют да где растет. Из этого его утвержения что если двигаться вдоль ареала с севера на юг грацилисы плавно переходят в колоратусы очевидно что вид этот один и подвида никакого нет - но он это просто не обсуждает

Коллекции феров в этой стране наверняка есть, вот только похоже гденить в Техасе никто это за достижение не считаен и не кичится - подумаешь кактусы в огороде.

PS сеянцы понравились

[petrovna]

по каждому поводу и без поводу не стоит

Давайте каждый будет сам решать есть ли повод

это лучшая книга

Может быть выложить "Предисловие" , при условии, что оно написано автором и 1-2 странички о любом из видов ? Будет приблизительно понятно стоит ли заморачиваться.

гденить в Техасе никто это за достижение не считаен

Что то мне подсказывает, что хорошо выращенные из семечка именно в Техасе грацилисы - все же будут достижением. Этим сеянцам второй десяток, немногий род который у меня не очень получается растить быстро. Перувианус вроде тоже типа подвой, а вырастить до цветущего состояния - довольно не частое явление.

что вид этот один и подвида никакого нет - но он это просто не обсуждает

Все верно, но среднестатистическому космонавту , при его уровне подготовки, этого никогда не понять читая такую книгу, а более или менее подготовленный любитель едва ли будет воспринимать ее серьезно, разве только картинки посмотреть...

[admin]

Может быть выложить "Предисловие" , при условии, что оно написано автором и 1-2 странички о любом из видов ? Будет приблизительно понятно стоит ли заморачиваться.

Вот несколько описаний видов из этой книги:
(приношу извинения за ошибк, так как после FainRidera текст не проверял)

===============================================

1. FEROCACTUS FLAVOVIRENS (Scheidweiler) Britton and Rose, Cactaceae 3: 138. 1922.
Echinocactus flavovirens Scheidweiler, Allg. Gartenz. 9: 50. 1841.
PLANT cespitose, forming large mounds sometimes over 2 meters in diameter and nearly 1 meter tall; individual stems to 3 or 4 dm tall and 2 dm wide, light green, globular to elliptical. RIBS 13, acute, 3.5 cm deep, vertical, not tuberculate. AREOLES oval, 1.5 cm long and 8 mm wide, bearing slight gray tomentum which ultimately disappears, distantly spaced on ribs, from 2 to 4 cm apart. SPINES tan or gray, acicular, terete; central spines 4 to 6, radiating, the lower central directed downward, longer than the others, to 8 cm long; radial spines 12 to 20, radiating, the lower ones like the centrals but the upper ones sometimes lighter colored, bristle-like and occasionally twisted. FLOWERS yellow, produced on the youngest areole on the rib and thus in the center of the stem; scales of the ovary linear-lanceolate, 15 mm long and 3 mm wide, very ciliate with an attenuate-aris-tate tip, the cilia on lower portion dichotomously branched, those near apex simple, the tips of cilia and of scale hyaline, sharp; scales intergrading into the outer perianth segments, these 18 mm long, margin entire at base with upper portion like the scales; inner perianth segments linear-lanceolate, 20 mm long and 2 to 3 mm wide, margin at base entire but the acute tip serrate to ragged; filaments 4 to 10 mm long, very fine, anthers minute; style 15 mm long and 1.2 mm thick, yellow, striated longitudinally and divided into 13 unequal stigma lobes 1 to 4 mm long. FRUIT elliptical, 28 mm long and 18 mm wide, completely covered with long brown ciliate aristate scales, these arranged linearly giving the appearance of rows; wall of ovary fleshy, red, 2 mm thick, the locule filled with soft, red, acid pulp containing scattered black seeds; perianth persisting, also covered with scales, the withered perianth about 2 cm long. SEED small, black, 1 mm long, with fine polygonal sculpturing.
Neotype: Hills 9 km from Tehuacan, Puebla, Mexico, on road to Zapotitlan de Salinas, Lindsay 2596 (DS). The original description was of plants from "Mexico, Tehuacan, 6,000 ft. elevation."
Distribution: Vicinity of Tehuacan and Zapotitlan de Salinas, Puebla. Mr. Schwarz states this species extends from the Tehuacan region toward the Mistecas Altas of Oaxaca. Map number one. Representative specimens: MEXICO: PUEBLA: 5.6 miles from Tehuacan on road to Zapotitlan de Salinas, 6,000 ft. elevation, Lindsay 2596 (DS,SD); Zapotitlan de Salinas, Lindsay 2059 (DS.SD). Living material of Lindsay 2596 is at the Desert Botanical Garden of Arizona.
Scheidweiler described Echinocactus flavovirens in 1841, from living plants which had been sent from Tehuacan, Mexico, some years before. The collection, probably by Karwinsky, had been thought to represent E. robustus. Dr. Rose reintroduced Echinocactus flavovirens into cultivation in 1906, and in 1922 included it in Ferocactus. Two other names have been associated with this species, Echinocactus polyocentrus Lemaire (Salm-Dyck, Cact. Hort. Dyck. 1844. 22. 1845), which appeared in a list but was never published with description, and E. orthacanthus Link and Otto, the type of which came from Montevideo and the description of which is not of Ferocactus flavovirens.
Bravo (1937, p. 429) states the flowers are yellow. Britton and Rose did not know the flowers and fruit. I have not seen fresh flowers material, and have drawn the description from treated withered perianths on the fruit. The uniquely scaled, fleshy fruit, which is filled with red pulp resembling that of a Lemaireocereus, the scale-like spine-tipped perianth segments of the flower, and the very small seeds are unlike those of any of the other Ferocacti; in fact they do not fit well in any known genus. It is possible that a new genus should be established for this species, but I hesitate to do this without a better understanding of the flower.
Ferocactus flavovirens grows in the same area as F. robustus, and has the same associate species, but usually is found on higher limestone hillsides. Ferocactus robustus is much more common in the valley flats near Tehuacan, but in the mountains at Zapotitlan F. flavovirens is the most abundant, and F. robustus is scattered.


2. FEROCACTUS ROBUSTUS (Otto) Britton and Rose, Cacteae 3: 135.1922.
Echinocactus robustus Otto, in Otto and Dietrich, Allg. Gartenz. 1:364.1833.
Echinocactus robustus var. prolifer Pfeiffer, Enum. Cact. 61. 1837.
Echinocactus robustus var. monstrosus Pfeiffer, loc. cit.
Echinofossulocactus robustus Lawrence, in London, Gard. Mas 17- 318 1841.
STEMS very cespitose, forming huge clusters of hundreds of branches, the clumps to 5 meters in diameter and over 1 meter tall; individual stems globuse to clavate, 8 to 16 cm wide, deep green. RIBS 8, prominent, acute, thickened under areoles, tuberculate. AREOLES widely separated on the ribs, round, 8 mm wide, with a superior floriferous portion, this becoming inconspicuous in age. SPINES purple, reddish, or horn-colored; central spines 4 or sometimes 6, erect, radiating, straight, annulate, square or angled in cross section, sometimes flattened laterally, the lower one longer, to 6 cm long; radial spines 10 to 14, radiating, the lower ones much like the centrals and the upper ones acicular or bristle-like and lighter in color. FLOWERS 3 to 4 cm long and about as wide, yellowish; ovary covered with broad, rounded imbricated scales, these intergrading into the outer perianth segments; outer perianth segments yellow with red midstripe, mucronate; inner perianth segments yellow, oblong, acute; stigma lobes 10, red. FRUIT yellow, fleshy, 2 to 3 cm long and 2 cm wide, with broad, widely separated, fleshy scales. SEED black, 1.5 mm long and 1 mm wide, finely sculptured with polygonal ridges.
Neotype: Zapotitlan de Salinas, near Tehuacan, Puebla, Mexico, Lindsay 2058, Sept. 4, 1951, (DS).
Distribution: Arid area in the vicinity of Tehuacan, Puebla. I have observed this species from Tepeaca to Zapotitlan de Salinas, Puebla. Map number one.
Representative specimens: MEXICO: PUEBLA: Zapotitlan de Salinas, Lindsay 2058 (DS). Living specimens are to be found at Huntington Botanical Garden, Desert Botanical Garden, and the University of California Botanical Garden at Berkeley.
Karwinsky apparently discovered this species in the vicinity of Tehuacan in 1828. He sent fruit and probably plants to Cells in France that year. The name first appeared without description in a list of cactus plants authored by Friedrich Otto in the Allgemeine Gartenzeitung in 1833, and this reference has generally been cited as the valid publication. Britton and Rose (1922 3: 135) cite Link and Otto as the authors, but the article was written by Otto, although both Link and Otto were editors of the periodical in which it appeared.
In 1837 Ludwig Pfeiffer treated Echinocactus robustus, crediting the species to "H. Berol.", and describing two varieties,pro///er and monstrosus. The first seems to fit the typical species, while var. monstrosus was an irregularly ribbed form which appeared among seedlings. This treatment appeared in Enumeratio diagnostica Cactearum, by Ludwig Pfeiffer, and also in Beschreibung und Synonymik der Cacteen, by Louis Pfeiffer, the same year. One of these works is apparently a translation of the other, and the treatment of E. robustus is the same in both. I do not know which was published first.
Labouret and Schumann listed Echinocactus galeottii as a possible synonym of E. robustus, but both questioned it. Echinocactus galeottii is obviously one of the large Echinocacti, probably E. grusonii.
The remarkable polycephalic habit of Ferocactus robustus is quite unlike that of any other species in the genus. The compact clumps grow to enormous size, sometimes fifteen feet across. These clusters are probably not from the same original root system, but are a complex which has developed from seedling plants which have grown up around the parent. The blossoming period is long. I have collected flowers and mature fruit in March, when they were scarce, and in September, when both were more abundant. The stems are firmly attached to the clusters and clonal reproduction is not important as it is in some clumping cacti with easily detached joints.
Ferocactus flavovirens, another cespitose species, grows in the same area and has been confused with F. robustus. The plants are quite unlike and belong to different subgenera. Ferocactus flavovirens has larger and fewer stems and very different fruit. In the flats near Tehuacan, F. robustus is the more common species, but at Zapotitlan de Salinas F. flavovirens is the most abundant.
Ferocactus robustus grows in calcareus soil, with Lemaireocereus stel-latus, L. hollianus, L. weberi, Pachycereus marginatus, Cephalocereus tetetzo, C. macrocephalus, C. hoppendstedtii, Ferocactus recurvus, F. flavovirens, Echinocactus grandis, etc.

[admin]

А вот описание из этой книги так всеми любимого в этой ветке GRACILIS.

================================================== ========

18. FEROCACTUS GRACILIS Gates, Cact. and Succ. Journ. 4: 323. 1933.
Key to the varieties
a. Often over 1 meter tall, the widest central spine usually less than 5 mm wide......................F. gracilis var. gracilis.
b. Less than 1 meter tall, the widest central spine more than 6 mm wide . F. gracilis var. coloratus.
18a. FEROCACTUS GRACILIS Gates var. GRACILIS
Ferocactus gracilis Gates, Cact. and Succ. Journ. 4: 323. 1933.
STEM simple, globose to cylindric, to 3 m tall and 3 dm wide. RIBS about 24, 2 cm high, slightly tuberculate and expanded under the areoles, deep green. AREOLES oval to elliptic, to 2 cm long and 6 mm wide, with light gray tomentum in youth. SPINES in two series; central spines 7-13, annulate, red with yellow tips, the 4 principal central spines cruciform, the upper and lower of which are flatttened and the lateral 2 subulate, the lower is the largest, to 6 cm long and 5 mm. wide, rather convolute and curved or hooked at the tip; in addition there are usually 2-3 secondary subulate central spines above the principal 4, and also 3 central-like radial spines at the lower side of the areole; radial spines white, flattened setaceous, twisted, the lower 3 acicular to subulate, and annulate. FLOWERS red, funnelform, about 4 cm long and 3.5 cm wide; scales on ovary widely spaced, not overlapping, to 4 mm wide and 3 mm long, green with reddish tip; outer perianth segments to 2 cm long and 6 mm wide, obtuse, red with narrow yellow margin; inner perianth segments linear lanceolate, 25 mm long and 5-7 mm wide, the midstripe lavender red and yellow serrulate margins; filaments 5-15 mm long, fine, appressed against style; style 22 mm long and 2 mm wide, yellow below and red above, the upper 1 cm divided into about 9 stigma lobes which are red on the outside and have an inner yellow stigmatic surface. FRUIT yellow, oblong, about 2.5 cm long, or 4.5 cm long including the persistent withered perianth; scales becoming sclerous; the fruit not opening by basal pore. SEED black, shiny, 1.75-2 mm long and 1.10-1.35 mm wide, rounded and not angled, with small round white hilum and a pit between the hilum and end of the seed.
Holotype: Hills west of Mission San Fernando, Lower California, Mexico, Gates 22, July 25, 1932, (DS).
Distribution: Central Lower California, from the southern end of the Sierra San Pedro Martir to below Punta Prieta, including the area between the 29th and 30th parallels. Map number six.
Representative specimens: MEXICO. LOWER CALIFORNIA. Hills west of Mission San Fernando, Gates 22, (DS, holotype); 20 mi. inland from Rosario, Lindsay 1846 (DS,SD).
Ferocactus gracilis is a particularly attractive cactus which was discovered by Mr. Howard E. Gates in 1928. Mr. Gates published the species in 1933. It occurs in northern central Lower California, from Rosario southward to where it intergrades with and is replaced by Ferocactus peninsulae var. viscainensis in the northern Viscaino desert. The section in which it grows is particularly arid, and the plant seems to have broad edaphic tolerance, from silt flats near the beach to alluvial gravel slopes and benches and rocky slopes. Associated plants include many with unusual forms, among them Idria columnaris, Fouquieria splendens, Pachycormus discolor, Pachycereus pringlei, Lophocereus schottii, Bursera microphylla, Agave nelsonii, etc.
Mr. Gates named the species gracilis because of its tall, slender form. It is usually less than one foot in diameter, but specimens nine feet tall have been found found. The plants do not ordinarily branch, but if injured may produce a cluster of heads. The red flowers get their color from the strong red midstripe of the perianth segments, which have a narrow yellow margin, and from the mass of bright red stamens. This color pattern is characteristic of most of the Lower California barrel cacti which occur to the south. The flowers appear in the summer in June, July, and August.
Ferocactus gracilis var. gracilis appears to be a stable form in the northern portion of its range, but becomes variable farther south. There it apparently intergrades with Ferocactus gracilis var. coloratus, which may actually represent a hybrid population between F. gracilis and F. peninsulae or F. peninsulae var. viscainensis.


18b. FEROCACTUS GRACILIS Gates var. COLORATUS (Gates) Lindsay, Cact. and Succ. Journ. 27: 169.1955.
Ferocactus coloratus Gates, Cact. and Succ. Journ. 4: 344. 1933.
STEMS simple, globose to subcolumnar, to 1 m tall and 3 dm wide. SPINES like those of the previous variety, except that they are usually somewhat shorter, and the flattened upper and lower of the four principal centrals are always 6 mm, and often over 1 cm wide. FLOWERS similar to var. gracilis, but apparently have less red pigment in the corolla segments, which are usually also more obtuse; the flower is not as broadly expanded and scales on the ovary are imbricate. SEEDS are more angular, irregular, and slightly larger than those of var. gracilis, and lack the pit adjacent to the hilum.
Holotype: Southwest of Aguaje San Andres, Lower California, Mexico, Gates 160 (DS, holotype of F. coloratus Gates).
Distribution: Known only from the area between Punta Prieta and Miller's Landing, Lower California, Mexico. Map number six.
Representative specimens: MEXICO: LOWER CALIFORNIA: Southwest of Aguaje San Andres, Gates 160 (DS, holotype); Lindsay 2013 (DS,SD); 20 mil S. Punta Prieta, Harbison s.n. April 19, 1940 (SD); San Andres, Lindsay DBG-83 (DES).
This variety was also discovered by Mr. Howard E. Gates, who considered it a distinct species and named it Ferocactus coloratus in 1933. It is very near the typical variety of F. gracilis, but has differences, already summarized in the above description, which are probably sufficient to segregate it as a varietal form. The population of barrel cacti in which var. coloratus occurs is variable, and the specimens which have been collected to represent coloratus are usually the extreme examples. It is possible that the population represents a hybrid swarm containing genetic material from F. gracilis var. gracilis, F. peninsulae var. peninsulae, which occurs in the adjacent Sierra San Borjas, and F. peninsulae var. viscainensis, which occurs in the Vizcaino Desert to the south.

[admin]

Welcome to The Genus Ferocactus
This book has George Lindsay's 1955 doctoral thesis, "The Taxonomy and Ecology of the Genus Ferocactus," at its core. To round it out, there are updates and other contributions, all with Dr. Lindsay' s blessing. These include a biography, some interesting color photos, botanical illustrations, tips on propagation, an illustrated glossary, a table summarizing Ferocactus distributions, a listing of Ferocactus synonyms, and a chapter on current taxonomy research involving analyses of DNA to rearrange the taxa tree. The book can serve as a pocketable field guide, be useful to a gardener, and bring one up-to-date on recent laboratory techniques, which when backed with field work can be used to trace what was thought to be untraceable a decade ago.
Cactophiles are attracted to these New World plants for their forms, their hardiness, their floral varieties, their ecologies, and their biological adaptations. Ferocacti are one of the 200 plus cactus genera with about three-dozen species and varieties. George Lindsay's enthusiasm began at an early age, and he elected to leave ranching behind and pursue a life of botanical research and exploration. This is described in the biography by Larry Mittich.
It is generally agreed that Ferocactus have evolved from Echinocactus and that Ferocactus transformations into species and varieties have occurred according to some order. Tracing this order via DNA techniques was selected as a doctoral thesis topic by Hugo Cota, with Professor Robert Wallace at Iowa State University as his advisor. The genus Ferocactus was a fortuitous selection for this pioneering work for several reasons: the genus is large enough to find a pattern, but not so large as to be an impenetrable thicket; Lindsay's field work is a reliable guide to morphology and biogeography of the species. Hugo describes these reasons in his own way in his chapter. The techniques developed by the Iowa group will continue to improve and serve others who wish to pursue studies within other plant taxa. At the time of original publication of this book, the Ferocactus studies were incomplete and these published results should be considered only as indicative of future findings.
A few updating revisions in italic form have been included within Lindsay's text. They include the addition of two added species; the rearrangement of some species into varieties; and some details on Ferocactus acanthodes varieties, provided by Frank Thrombley, who has field collected and studied Ferocactus for many years. Since the positions of species and varieties are likely to change following Cota's work, we expect to be forgiven for an occasional disarrangement in our arrangements. These are exciting times!

The editors have supported the text with a few appendices. George Lindsay's thesis, while easy reading, uses botanical terms which may be unfamiliar to some readers. Hence we have provided an illustrated glossary. Madelyn Lee, Manager of the Grigsby Cactus Gardens and a longtime Ferocactus enthusiast, provided a list of past and present synonyms. The editors have included a list of Ferocactus localities.
It was important to us that this book reflect the beauty of Ferocactus and the work of all involved. We have included the remarkable color plates produced by Dr. Dallas Hanna and team several decades ago on ordinary office equipment; the illustrations of Lucretia Breazeale Hamilton, made for George Lindsay's doctoral adviser and friend, Ira Wiggins; the amusing rear cover caricature of George Lindsay and Reid Moran by Gerhard Marx; and the poetic front cover and end papers by Sandra Reed, our Art Director.
Charley Glass wrote us that he has been waiting for the Lindsay book for 35 years. A book is an experience. We predict that this experience will serve for a long time.


Acknowledgments
This book represents the efforts and contributions of many people. The core of the book, Dr. Lindsay's informative PhD thesis, has never before been effectively published, being available only as illustrationless copies, extracted from microfiche. The California Academy of Sciences typed most of the text on a word processor. The balance of text was scanned by OCR (optical character recognition) software and subsequently smoothed into English. The approximately 120 accompanying photos and figures were scanned into the text apart from the captions, which were separately blended in. In areas where there was new information, it was gently inserted as editorial notes or as appendices. The sources of this new information include Frank Thrombley, a frequent Baja California explorer, who wrote on Ferocactus acanthodes varieties; Madelyn Lee, Manager of Grigsby Cactus Gardens, who supplied a list of synonyms; and authorities such as Curt Backeburg's Cactus Lexicon; Nigel Taylor's articles on Ferocactus in Bradleya Nos. 2/1984 & 5/1987; Lyman Benson's The Cacti of the United States and Canada; and Ira Wiggins' Flora of Baja California.
George Lindsay and the editors were interested in how molecular studies being performed by Hugo Cota at Iowa State University would clarify the phylogenetic relationships of Ferocacti. Despite a heavy field, laboratory, and academic schedule, Hugo made time to write a chapter describing his current work. This work will bring about a rearrangement of the genus and will serve as a model for work on other genera.
Aesthetics were not neglected. The "Illustrators Gallery" appendix shows the botanical illustration skill of Lucretia Breazeale Hamilton. These were reprinted from "The Cacti of the United States and Canada" by Lyman Benson with permission of the publishers, Stanford University Press, (1982). Our Art Director, Sandra Reed, not wishing to duplicate the botanical illustration style of Lucretia Hamilton, created the fanciful and colorful front cover. The rear cover by Gerhard Marx makes us smile. The Hanna Color Plates were made decades ago at George Lindsay's instigation by an ingenious office copying process devised by an ingenious man and his helpers (more about Hanna in the book).
The lucid illustrations of cactus forms, flowers, and spines within the Illustrated Glossary foldout were done by Thor Methven Bock. They appeared in Cactaceae by W. Taylor Marshall and T. M. Bock (1941) and are reprinted with the permission of Charles Glass, publisher of the former Abbey Garden Press.
Galleys of the book were reviewed and edited by George Lindsay and all of the contributors. The editors welcome corrections and suggestions for future printings. The book was assembled by the editors, using a blend of old crafts and the latest in digital processing methods. This book now resides in digital form. It will be printed in small batches as needed, a process called "printing-on-demand." What we have learned during the publication process will be made available to others.
Eventually, the editors expect that publishing-on-demand will become a smooth process. At present there are many speed bumps. We were able to glide over them through the help and ingenuity of our premiere suppliers. They include: Copy Girls for endpaper printing, who made my overlays match up; McKibben Silk Screen Printing who easily handled what others couldn't; and Universal Reprographics for text printing, inputting our files through their system. Special thanks go to Jerry Kiley, Manager of Golden Rule Bindery, for exceptional help and ingenuity. Golden Rule assembles this book from components as needed.
The Editors
Reese Brown Mark Raptis Stan Yalof Fred Fox



The Taxonomy and Ecology of the Genus Ferocactus
Dr. George Lindsay Introduction
The genus Ferocactus belongs to the subtribe Echinocactinae of the tribe Cereeae, in the Cactaceae. Ferocacti are commonly called "barrel cacti" or "visnagas" and inhabit certain arid and semi-arid regions of Mexico and southwestern United States. They are bulky succulent plants, globular or cylindric in shape, with a covering of spines borne on parallel ridges or ribs. True leaves are lacking, and photosynthesis takes place in the superficial layers of the stem.
Ferocactus was one of the more natural segregate genera proposed by Britton and Rose in 1922, when they subdivided the large polyphyletic genus Echinocactus. Political unrest in Mexico, as well as the inaccessibility of some of the major cactus territories, prevented adequate field work by Britton and Rose. A number of the 31 species which they assigned to the new genus were imperfectly known, and several have been found to be synonyms or have been transferred to other genera. Recent plant exploration has added new species. A monographic study of the genus Ferocactus therefore seemed desirable.
Cacti present unusual problems for the taxonomist, and effectual work with them requires somewhat unorthodox research procedures. Ordinarily taxonomic research involves a review of the literature, the examination of a mass of herbarium material, and, if possible, field investigations of critical taxa or areas. The literature concerning the Cactaceae is fairly abundant but often of inadequate quality. Many species were described from a single, sterile, seedling plant, the geographical origin of which was unknown. Descriptions were often meager and before 1900 the preservation of type material was the exception rather than the rule. Even now cacti are poorly represented in herbaria because their succulence and spininess make field preparation and preservation slow. Those specimens which are available are often unsatisfactory because they consist of only flowers or fruit or fragments of vegetative material. Complete specimens may fail to show die characters of the living plant, these characters having been lost or altered with the drying of succulent tissue. Several species of Ferocactus apparently were not represented by a single specimen in any herbarium when this study was undertaken
I used the following investigational procedure. The available literature was reviewed and photocopies of pertinent information were made. The major cactus herbaria and botanical gardens of the United States were visited and critical and type material was studied and photographed. Specimens, photographs, and information were obtained from European herbaria through correspondence. Herbaria visited included those of the New York Botanical Garden, Missouri Botanical Garden, Desert Botanical Garden of Arizona, Biological Institute of the University of Mexico, University of Arizona, University of California, San Diego Natural History Museum, California Academy of Sciences, United States National Herbarium, Allan Hancock Foundation, Rancho Santa Ana Botanic Garden, Pomona College, and Dudley Herbarium of Stanford University. In addition, living plants were studied in the collections of the University of Michigan, New York Botanic Garden, Henry E. Huntington Botanical Garden, Desert Botanical Garden, the Botanic Garden of the University of California, Howard E. Gates Lower California Botanic Garden, and at the Biological Institute of the University of Mexico. I am indebted to the officials of all of those institutions for the assistance they gave me. Special thanks are due Charles Baehni, Director of the Botanical Gardens and Herbarium in Geneva; Sir Edward Salisbury, Director of the Royal Botanical Gardens at Kew; Dr. A. W. Ewell, Deputy Keeper of the Department of Botany of the British Museum; Prof. Guillaumin, Director of the Laboratoire de Culture of the Museum National d'Histoire Naturelle; and Dr. Reed C. Rollins, Director of the Gray Herbarium of Harvard University. All of these gentlemen were most cooperative in searching for critical specimens and supplying photographs, loans, and important information.
The major phase of my research was that conducted in the field. I traveled about 20,000 miles by automobile and truck and 5,000 miles by boat in order to visit the general areas where Ferocacti grow. There the species were studied, photographed, and collected as living and herbarium specimens. The living plants were grown and observed at Stanford and the Desert Botanical Garden of Arizona. The field studies were essential for any understanding of the plants which are poorly represented in herbaria, and for information about the variation within and between taxa. The collections of herbarium material have been adopted in the Dudley Herbarium, with duplicates going to the San Diego Natural History Museum.
Ecological data do not constitute a major portion of this paper, but some of the more obvious ecological factors are discussed. A comprehensive ecological study would have involved methods of field investigation incompatible with the schedule of collecting which was maintained in order to observe all of the species in their native habitats.
I want to thank the many friends who have helped investigate the Ferocacti. My major professor, Dr. Ira L. Wiggins, has been most generous with counsel and suggestions, has evidenced a deep personal interest in the problem, and has been my companion on field trips. Dr. Richard W, Holm has also advised and helped me in many ways. Mr. J. W. Sefton, Jr, sponsored a major expedition, the Sefton Foundation - Stanford University Expedition to the Gulf of California, which allowed me to collect on the inaccessible islands along the west coast of Lower California and the Gulf of California. Dr. Helia Bravo Hollis, noted authority on Mexican cacti, has given me much information about the species in central and southern Mexico, and has arranged trips in that area in order for me to study them in the field. Mr. Howard E. Gates, who discovered and named several Ferocacti in the Lower California region, has been a personal friend for many years and always most generous in supplying information. Dr. Faustino Miranda, of die University of Mexico, accompanied Dr. Wiggins and me on a survey trip through the arid regions of the states of Puebla and Oaxaca. Dr. Miranda's knowledge of the thorn forest and desert flora increased tremendously the value of die trip for us. Mr. Fritz Schwarz, of San Luis Potosi, has been very generous in supplying information about collecting localities of the Mexican mainland species, and has personally shown me where many grow. Dr. Reid Moran has been my companion on several botanical exploration trips and has been very helpful with ideas and information. Mr. Wm. Taylor Marshall, Director of the Desert Botanical Garden of Arizona, has raised and kept records of plants for me, and has also supplied data from his own knowledge of the family. Dr. Lyman Benson of Pomona College not only gave me the use of his splendid cactus herbarium, but also supplied helpful suggestions and ideas concerning various taxonomic problems within the Cactaceae, on which he has actively worked for many years. Mr. Thomas MacDougall, veteran plantsman of southern Mexico, collected with us in Tehuantepec, showed us where unusual species were found, and has made special efforts to procure rare plants for me. Prof. William C. Steere, of Stanford University, has given me friendly encouragement and advice and has been my companion on several collecting trips. Mr. Charles F. Harbison of the San Diego Natural History Museum has given me photographs and specimens of Ferocacti from isolated areas in Lower California which I have not visited. Some of the specimens and information which he obtained cost him considerable effort, and proved to be of great value. This is but a partial list of those to whom I am indebted, and to whom I wish to express sincere appreciation.

[admin]

Key To The Species of Ferocactus
Editors' Note: Changes have occurred in the Ferocactus taxa since the publication of George Lindsay's 1955 thesis, such as the discoveries of the species F. lindsayi and F. reppenhagenii. There also have been field and laboratory studies which have attempted to group Ferocacti according to their evolutionary development from the appearances of their plant parts and seeds, most notably the work of Nigel Taylor [Bradleya 2/1984 & 5/1987]. Taylor has made several changes in taxonomy, including the changing of some species to varieties. Whether these changes will endure will probably be determined not from more studies of appearances, but from molecular genetic studies, such as those being performed by Hugo Cota, described elsewhere in this book. This book has as one of its goals the review of progress in Ferocactus taxonomy. Its major goal is to present Lindsay's work with updating additions, making the original work complete in its descriptions. Lindsay occasionally differs from others in naming and grouping. We have placed editorial notes to highlight synonyms. These differences are minor and probably unimportant as Cota's work will probably result in substantial rearrangements.
We have expanded the original key. We also have changed its form, which originally used eight levels of indents, to a simpler numerical form modeled after that of Nigel Taylor. The key includes Lindsay's descriptions, added to by descriptions drawn from Nigel Taylor (NT) and Backeberg's Cactus Lexicon (Bb).
The editors have taken liberties with this key by deviating from being dichotomous (having only two possibilities) in a few items. This deviation from usual practice was done in order to include species identified since Lindsay and some of the useful insights of Nigel Taylor while keeping to Lindsay's chapter arrangement. For example, Key item 24 has four species possibilities. Two of these, Ferocactus herrerae and Ferocactus townsendi-anus, appear as Lindsay species and chapter headings. Taylor considers them as varieties, hence are not included in his key. The genus Ferocactus is small, and we believe that such deviations from orthodoxy won't confuse the reader.
Following the Key are two additions: Ferocactus Distributions, a listing of Ferocactus species, varieties, and their distributions; Ferocactus Synonyms, a list of historical and current names.
I. Plants strongly cespitose, forming broad low mounds of many (more than 10) heads.............................2
1. Plants simple, or if cespitose normally with less than 10 stems, not forming low mounds..........................3
2. Fruit red, densely covered with linear apiculate, ciliate scales; stem not crowded; ribs about 13; (Tehuacan to Oaxaca) ....................................... 1. F. flavovirens
2. Fruit yellow, with widely spaced lunate scales; stems crowded; ribs about 8; (Tehuacan to Zapotitlan, Puebla).........2. F. robustus
3. Spines 1-10/areole, straight and needlelike, none strongly flattened above or recurved to hooked at apex.................4
3. Spines 10/areole, or at least one strongly flattened and/or recurved to hooked at apex, or stem ribs 13-16 and spiraled...........15
4. Seeds smooth or with reticulate markings, but not pitted.......5
4. Seeds pitted..............................8
5. Spines 1-6, more or less equal (+/-), to 2.5 cm. long, stems 15-50 cm diameter Spines usually 7, central longer, 3-10 cm; stems to about 25 cm diameter .............................7
6. Stem distinctly glaucous, blue green; fruit nearly white; seeds very smooth; (Hidalgo Mexico).................. 3. F. glaucescens
6. Stem dark green, largest specimens over 30 cm tall and broad, seldom cespitose; seeds with a reticulate pattern of raised testa-cell margins; spines 1-3, sometimes 5 in juvenile specimens; (from mtns. of Sinaloa in west Mexico)............... 5. F. schwarzii
7. Areoles well separated on the ribs, 2-4 cm apart; stems often clustered; stigmas 10-14; stems under 30 cm tall, often cespitose; (eastern Sierra Madre, SL Potosi and Hidalgo, Mexico).
...........................................4. F. echidne
7. Areoles about 1 cm apart or +/- confluent on the ribs; stem solitary; stigmas (about 7); (Michoacan, south-west Mexico).
.................................... 26. F. reppenhagenii
Editors' Note: F. reppenhagenii was not in the original text.
8. Fruit red to purple, indehiscent, and/or very juicy and deliquescent; ribs acute; stem not exceeding 1.2 m high.................9
8. Fruit yellow (red in 21b ) or dihiscing by a basil pore; ribs obtuse or acute; stem 30 cm-4 m high......................12
9. Flowers yellow; seed about 1 mm...................10
9. Flowers red to purplish pink; seed about 1.8-2 mm..........11
10. Stem ribs, 13-18; spines 6-7/areole; seed elongate, curved;
(Michoacan Mexico)........................ 27. F. lindsayi
Editors' Note: F. lindsayi was not in the original text.
10. Stem ribs 20-40; spines golden yellow, usually 7/areole, typically 9,1 central and 8 radials; seed +/- isodiametric; (Durango to Jalisco and Puebla, north-central Mexico)................. 6. F. histrix
11. Stem globular to cylindric, 30-120 cm high; flower 6-7 cm long;
(Puebla-Vera Cruz border)............. 28. F. haematacanthus
Editors' Note: F. haematacanthus was not in the original text.
11. Stem disc shaped, depressed globose, more than twice as wide as tall, deep seated in ground, to 10 cm tall; (central and southern Mexico). ...................................... 8. F. macrodiscus
12. Central spine, 1 per areole, conspicuous; seeds 2-3 mm...... 13
12. Central spines 4 or not distinct from radials; seeds 1.5-2 mm. . . 14
13. Flower to 4.5 cm long; stem to 1 m high (SW Chihuahua, SE Sonora, and N Sinaloa).........................l.F. pottsii
(listed as F. alamosanus, a synonym, in Lindsay)
13. Flower 6-7.5 cm long; stem to 2.5 m high; (S Arizona to N
Sinaloa, BC Sur)............................ 14. F. emoryi
(F. covillei synonym in Lindsay)
13. Central spine 1, always straight, 9-15 or even 25 cm long; r. sp. 8-12, upper ones stouter; flower yellow, 6 cm long; (San Ignacio to
Commondu, central BC).................. 15. F. rectispinus
(F. emoryi variety per NT)
14. Areoles with 13-20 hairlike whitish marginal bristles or setaceous spines, usually red, some +/- flattened or angled ; central spines all similar; stems often in clumps; (Central N Mexico). .......................................... 11. F. pilosus
14. Spines 22-25, similar in shape and size; stem short cylindric; ribs 24-31, wavy; flower yellow, reddish outer, 5 cm long(Angel de la
Guardia Island, BC)................... 24. F. johnstonianus
(Species/Bb ,F. acanthodes variety/NT).
14. Stem ribs 25-35; spines clear yellow, rarely reddish brown, terete, all of one type; stem solitary; (BC Islands).......25. F. diguetii
15. Scales on receptacle tube and fruit with long narrowly attenuated apices; stem biscuit shaped (or depressed globular); ribs about 21; (central Mexico).......................... 10. F. latispinus
15. Stem globular to short cylindric to 25 cm high; ribs spiralled, 13-15, 3 cm high; radial spines subulate, 8, 2 of these +/-compressed, banded, to 2.5 cm long; central spine 1, very stout, to 7 cm long; flower red, bordered white or pink; (Puebla and Oaxaca,
Mexico)................................... 9. F. recurvus
(F. latispinus var. per NT, sp. per Bb)
15. Scales not as for the preceding....................16
16. Fruit pinkish red and indehiscent, and/or very juicy, deliquescent, releasing the seeds in fluid.......................17
16. Fruit yellow or dehisciing by a basal pore when ripe........19
17. Spines straight or curved, but not hooked at apex; flower 3-4 cm short tubed..................................18
17. Spines, at least one per areole hooked at apex; flower 6-10 cm with a long tube; (El Paso, Cameron City TX to SL Potosi Mexico).
................................... 29. F. hamatacanthus
18. Flowers purplish pink to red, tepals with paler margins; seed to 2 mm. (Cumbre (10,000 ft., El Renosco)........8. F. macrodiscus
18. Flowers yellow; seeds c. 1mm...................6. F. histrix
19. Radial spines, 7 per areole, terete, only slightly thinner than the solitary terete or radially compressed central; (S AZ, Sonora).
.......................................... 14. F. emoryi
19. Radial spines 9 or at least some thinner than the one or more centrals. .....................................20
20. Perianth segments remaining +/- erect at anthesis; flowers to 2.5 cm diameter; stems often clustering, to 3 m high; principal spines 6-12, none curved or hooked at apex, often with marginal hairlike radials; central spines all similar; (northern to central Mexico).
.......................................... 11. F. pilosus
20. Perianth-segments spreading; flowers 3.5-6 cm diameter., stems rarely clustering; spines not as above or centrals and radials intergrading... 21
21. Flowers reddish to wine purple; principal central spine hooked or strongly curved at tip; (W coast BC).............. 21. F.fordii
21. Flowers green, yellow, orange, or red...............22
22. Spines clearly differentiated into stout dark colored centrals plus upper and lower radials, and finer whitish laterally directed radials, or the latter absent and seeds with tabular testa-cells............23
22. Spines in each areole intergrading in size and color; seeds with
concave testa-cells.......................... 25
23. Central spines red, 4, the principal one distinctly flattened instead of diamond-shaped, only the principal one twisted; flowers red; fruits to 7.5 cm long; (from Rosario to Millers Landing BC). ......................................... 18. F. gracilis
23. Central spines 4, twisted and tortuous (except rarely F. gracilis var. coloratus); radial spines about 16, radiating, terete; flower 6 cm diameter and long, red; (Smith Island group, Bahia Los Angeles,
BC)...................................... 19. F. gatesii
(Sp./Bb; var. F. gracilis/7Vr)
23. Largest central spine more flattened than the others or otherwise dissimilar, often strongly recurved or hooked; flowers red to yellow; fruits to 6 cm long.......................... 24
24. Ribs 20-30; stem wide and heavy; seeds with finely verrucose, +/- isodiametric testa-cells, the raised anticlinal walls at the cell margin not prominent; (from Sonora, Chihuahua, N Sinaloa, W Durango andAZ) ......................... 12. F. wislizenii
24. Central spines dissimilar in character, one or two more developed than others; radial spines 8, 2 of these white, the others flecked red; central spine 1, hooked initially, becoming straight; ribs, 12-20; stem thinner than F. wislizenii; flower funnelform, reddish, bordered yellow; (S Sonora, Sinaloa, and Durango)
........................................ 13. F. herrerae
(species per Bb, F. wislizenii variety per NT)
24. Central spines 4, cruciform, hooked; radial spines variable, usually about 11; seeds with tabular to concave, coarsely verrucose testa-cells, the verrucae few and separate, the raised cell margins prominent or the cells oblong; (central BC to Cape). ..................................... 16. F. penninsulae
24. Central spines 4, lateral two not distinctly flattened; radial spines bristle-like; flowers about 6 cm long, middle pink, greenish-yellow
border; (BC gulf and Isla San Jose)......17. F. townsendianus
(species per Bb, F. penninsulae variety per NT)
25. Flowers greenish-yellow; spines not hooked or strongly curved at tip; spines to 5 cm long; seed about 1.5 mm; stem not conical, as broad as tall; radial spines not bristle-like; (San Diego USA and nearby BC).............................. 22. F. viridescens
25. Flowers yellow, orange or reddish, or green but with other colors in the same population; spines to 5-17 cm long; seeds about 2-3 mm. ................................... 26
26. Central spines golden yellow or rarely bright red, usually
+/-diamond-shaped in cross section and twisted flattened and fairly uniform, central and radial spines about equal in number (10); flowers orange to red with 4-5 mm wide inner perianth-segments; (Isla Cedros and W Baja)................20. F. chrysacanthus
26. Central spines fewer than radials and 4 of the centrals are much larger than the others, to 7-17 cm long; inner perianth-segments 7-11 mm wide, flowers yellow; (SW USA, NW Sonora, NE BC). ...................23. F. acanthodes

[admin]

A Review of Ferocactus Britton and Rose
J. Hugo Cota
Department of Botany, Iowa State University, Ames, IA 50036
and Centro de Education Ambiental e Investigation Sierra de Huautla
(CEAMISH),
Universidad Autonoma del Estado de Morelos,
Av. Universidad # 1001, Cuernavaca, Morelos, Mexico.
Prelude
The following text, my contribution to this book, combines the field observations made by many people with the revolutionary molecular approach under way in taxonomy. This involves the analysis of DNA, such as sequencing and restriction site data. Field taxonomy often deals with the interspecific comparisons of morphological and physiological traits, both subject to convergence. The study of DNA provides new evidence which supplements conventional taxonomic studies, and lessens the amount of homoplasy (where two or more organisms independently acquire the same character state due to convergent evolution). DNA sequencing is the source of many new characters (the order and position of bases in the strings of DNA). Many of these characters are stable and highly conserved during evolution; some are evolving rapidly. From a comparison of similarities and differences between species, genealogical trees (cladograms) are built, with a distant relative (outgroup) serving to anchor (root) the tree. There are several methods to build and analyze trees. The method using "parsimony" (principle of maximum parsimony) constructs cladograms with a minimum number of evolutionary steps (changes). The construction of these trees from the sequencing and restriction site data was done objectively, by the software within the computer. My work involves the study of Ferocactus chloroplast DNA. Differences between cladograms, such as branching patterns, occur for several reasons, e.g., different number of species sampled, unequal rates of evolution between the regions of the genome being investigated, hybridization, gene duplication, and lineage sorting. As with all statistical methods, confidence increases with an increase of sampling. At the end of this section I have included a glossary of terms which may be unfamiliar to the reader.


Introduction
More than four decades have elapsed since Lindsay (1955a) wrote his doctoral dissertation, a study of the taxonomy and ecology of Ferocactus Britton and Rose. Lindsay's studies represent a valuable source of field documented material in which he includes both morphological and ecological data, and his own concepts about the classification of Ferocactus. His extensive field work at that time was challenging especially in remote regions of the Baja California Peninsula and surrounding islands as well as in mainland Mexico in the 1950's. The extent of his field work proves that there was no obstacle which could stop George from accomplishing his goals and conducting his field studies. His passion for cactus taxonomy and adventurous spirit have inspired other people to follow his example. George has been one of my mentors and his support and advice during my graduate studies and career in cactus taxonomy have been meaningful. Many fruitful discussions with him have given me a broader perspective of the genus and encouragement to seek the answers of the taxonomic relationships and evolutionary history of Ferocactus. I feel honored to contribute with this chapter of the book, in particular because of my personal interest in Ferocactus and because Lindsay's pioneer field studies of the genus represent an excellent example of his love and devotion for plants and his understanding of their taxonomic relationships.
Subsequent to the conclusion of Lindsay's dissertation, several authors have conducted separate studies dealing with diverse biological aspects of the genus. Such studies have considered a single species or major taxonomic groups within the genus and have referred to nomenclatural, biological, ecological, or physiological matters (e.g., Blom and Clark, 1980; Bravo-Hollis, 1966; Cota and Wallace, 1996; Nobel, 1977; Ruffner and Clark, 1986; Taylor, 1979a, 1979b, 1980, 1984, 1987; Taylor and Clark, 1983; Unger, 1992). Lindsay contributed descriptions of new taxa, nomenclatural changes, and ecology of Ferocactus (e.g., 1942, 1955b, 1955c, 1955d, 1964, 1965, 1968).
In the following sections, I briefly describe the main topics that have been investigated in Ferocactus from 1955 to the present. My objectives in this chapter are to summarize the knowledge of the genus in different areas of study and to discuss the importance of understanding its phyloge-netic relationships at the interspecific, generic and tribal levels. It is my intent to review Lindsay's taxonomic treatment and associate it with the results from modern techniques (molecular studies) used in phylogenetic analyses of Ferocactus. I hope that the up-to-date view of Ferocactus presented here is useful to anyone interested in cacti.
Because of the diverse array of topics investigated since 1955, several studies have been omitted not because they are irrelevant, but because their goals are considered distant from the main issues addressed in this chapter. In additon, to mention every paper dealing with the genus would imply the discussion of several areas of research that are not my specialty. Therefore, and with my apologies to those authors who have contributed to the knowledge of die genus in the topics excluded herein, I restrict myself mainly to those studies that in one way or another are more closely related to the understanding of the biosystematics of Ferocactus. At the end of this chapter there is a list of additional bibliographic references dealing with various aspects of the genus.
Ferocactus is distributed in arid and semiarid regions of central and northern Mexico and southwestern United States (Fig. 4), and it has been hypothesized to occupy a critical phylogenetic position in the classifications oftheCacteae(Fig.5)(Buxbaum, 1951,1958;Barthlott, 1988; Barthlott and Hunt, 1993). Within the Cacteae, Ferocactus is the third largest genus in number of species after Mammillaria Haw. and Coryphantha (Engelm.) Lem. (Gibson and Nobel, 1986; Barthlott and Hunt, 1993).
The understanding of the phylogeny of Ferocactus and its phylogenetic position within the Cacteae (the largest tribe within subfamily Cactoideae in North America), along with its patterns of speciation, is significant to understand the relationships at the interspecific and tribal level (in particular between the tribes Pachycereeae and Cacteae). The development of a robust phylogenetic hypothesis for the genus will provide a basis for the subsequent determination of its evolutionary patterns with other genera of the Cacteae as well to potentially clarify the relationships between the tribes Pachycereeae and Cacteae. Likewise, a methodical study of Ferocactus will help to explain the interspecific phylogenetic relationships and most likely provide a scenario of the evolutionary patterns of radiation.


Taxonomic Background
The taxonomic history of Ferocactus is addressed by Lindsay earlier in this book, and no further discussion is needed. At present, the taxonomy of the genus remains in a somewhat confused state. Different taxonomic philosophies, disagreement as to species boundaries, and under-repre-sentation of some taxa in herbarium collections have contributed to the complexity of its taxonomic circumscriptions. Another factor involved is the relatively high level of morphological homoplasy, which obscures the degree of relatedness due to convergence. Additionally, cactus taxonomy in general is complex because morphological variability"is associated with gradients in habitats and geographic ranges (Cody et al., 1983). In Ferocactus, the use of primarily vegetative morphological characters (which are likely to display different degrees of plasticity in response to local environmental conditions) has also contributed to its taxonomic difficulty. As a result, natural subpopulations subject to varied selective pressures may diverge both morphologically and genetically, occasionally providing discontinuities that are useful to distinguish taxonomic complexes in the process of speciation. As a consequence of natural variation and taxonomic delineations, the number of species circumscribed in the genus has been a matter of disagreement and is reflected in several taxonomic treatments (Table 1).
Prior to Lindsay, Britton and Rose (1922) proposed a taxonomy in which they included 30 species [Lindsay (1955a: 40) indicates 31]. Lindsay's classification (1955a) considered a total of 25 species and ten varieties. More recently, in a taxonomic treatment of Ferocactus sensu stricto Taylor and Clark (1983) included 29 species; Taylor (1984) recognized 23 species and 20 infraspecific taxa, while Bravo and Sanchez-Mejorada (1991) accepted 29 species and 25 varieties. Finally, Unger (1992) provided a review of the genus in which descriptions and distributional data are discussed. Although his taxonomic delimitations were based on previous treatments, he presented a classification scheme which included four sections (Table 1) and a list of naturally occurring hybrids. In the broad sense of the genus, Taylor (1980) transferred some species of Stenocactus (Schum.) Berger to Ferocactus, based on morphological affinities such as similarities in rib pattern and presence of glandular areoles, and created the subgenera Ferocactus (in which are placed all of Lindsay's species) and Stenocactus.
Despite the substantial amount of taxonomic work conducted during the past seven decades, the controversy in species boundaries within the genus is evident: no classifications are similar yet the number of species are similar (e.g., 30, 29, 28, 25, 23). Neither Britton and Rose (1922) nor Lindsay (1955a) used infrageneric categories, whereas Bravo and Sanchez-Mejorada (1991), Taylor (1984), Taylor and Clark (1983), and Unger (1992) classified the genus using other taxonomic hierarchies, such as sections, groups and subgenera (Table 1). A common feature of these classifications is that the delimitation of species has been based primarily on gross morphological features of the stem, flower and fruit.
Of these taxonomic treatments, those by Taylor and Clark (1983) and Taylor (1984) are of special importance because they have correlated gross morphology with micromorphological seed coat characters. These studies have led to the establishment of sectional boundaries and the consideration of new ideas about the evolution of Ferocactus. In spite of Taylor's (1984) comprehensive treatment combining all the data available at that time, a robust phylogeny of the genus has not been presented to date, and key issues regarding its origin and radiation remain unresolved.
The extensive use of varietal rank in the different classifications is further indication of the taxonomic confusion of this genus. To some extent, the use of varietal rank may be helpful in distinguishing infraspecific groups in Ferocactus, and might accurately reflect the relatively recent origin of some species groups, especially for some Baja Californian species. Indeed, Lindsay (1955a, 1965) and Taylor (1984) indicated that those species occurring in mainland Mexico are taxonomically well-defined, while several taxa from northwestern Mexico and the Baja California peninsula remain problematic, probably due to incipient processes of speciation.
In the following sections, I review new data from cytological and molecular sources that reveal preliminary information about the relationships at the interspecific level as well as the inferred relationships of Ferocactus with the columnar cacti of Tribe Pachycereeae. Both sources of information are used to discuss the issues of phylogeny and hybridization. To avoid confusion and to facilitate die taxonomic circumscriptions of Ferocactus, the specific epithets and authorities used in this chapter are those included in Lindsay's classification for two reasons. First, to honor him for contributing to the understanding of the infra- and interspecific categories of the genus and second, because the other taxonomies (Taylor and Clark, 1983;Taylor, 1984; Bravo and Sanchez-Mejorada, 1991; Unger, 1992), also recognize the majority of species proposed by Lindsay. This is not meant to discredit the valuable taxonomic contribution of these authors. Also, it is worth mentioning the outstanding contributions of Taylor (1979a, 1979b, 1984, 1987) in the nomenclature of the genus clarifying the application of several specific epithets and authorities that should be discontinued. I must repeat, however, that while I am aware of these nomenclatural changes, here I follow those names used by Lindsay (1955a) for the reasons indicated above.
To date, the most modern taxonomic treatments of Ferocactus are those of Taylor (1984) and Taylor and Clark (1983), in which phylogenetic ideas about the evolution and radiation of the genus based on vegetative and reproductive characters, geography, and macro- and micromorphology of the seed coat have been proposed. This information is essential to understanding the modern concept of classification of the genus, and these recent evolutionary hypotheses will be used to discuss phylogenetic aspects of Ferocactus, especially in the section which includes molecular studies.


Reproductive Biology in Ferocactus
Reproductive strategies in plants are numerous and they are reflected in an array of floral morphology and breeding systems, which have evolved in response to genetic, environmental, and selective forces. In sexually reproducing plants, as is the case in Ferocactus, breeding systems determine, in part, patterns and levels of genetic variation. Although little is known regarding the pollination biology and breeding systems of the genus, in the following paragraphs I summarize our current knowledge of the topic. Floral Biology and Pollination
The flowers of Ferocactus are perfect and bowl-shaped (Fig. 6); they open during the day, and have a slight fragence (Bravo and Sanchez-Mejorada, 1991). The color spectrum varies from yellow, orange, red, pink, lavender, to purple (Fig. 6); the stamens are numerous (Fig. 6A), usually with yellow anthers that mature before the style becomes receptive, thus they are protandrous (Richards, 1986). The plants bloom from early spring to late summer, and the flowers remain open for several days (Bravo and Sanchez-Mejorada, 1991).
In Ferocactus there is an apparent coadaptation of pollinating agents and its flowers. The floral attributes such as the funnelform to bell-shaped, diurnal, bright yellow, medium-to large-sized flowers, with nectar and little or no fragance at all, are characteristic of bee pollinated flowers (Rowley, 1980a). The flowers of Ferocactus are pollinated by several species of bees. Bee pollinated flowers have been documented in F. wislizeni (Engelm.) Britton and Rose (Grant and Grant, 1979a) which is usually pollinated by medium sized bees of the family Megachilidae (Megachile sidalcea Cock-erell and Lithurge echinocacti Cockerell) and Apiidae (Diadasia australis Cresson) (Grant and Grant, 1979b). Cockerell (1900) also indicated that Auglochlora neglectula Cockerell visits the flowers of F. wislizeni. Unidentified bees have been observed crawling around the stamens and stigmas of the flowers of F. histrix (DC) Lindsay and F. robustus (Otto) Britton and Rose in areas of Central Mexico (pers. obs.). Although the flowers of Ferocactus are also visited by other species of insects, the pollination syndrome has evolved to promote bees as the principal vector. In additon to bees, beetles are common visitors of Ferocactus flowers and feed on pollen and may accidentally pollinate the flower (Grant and Grant, 1979b) acting as secondary pollinators.
Mechanisms such as herkogamy (the spatial isolation of stamens and stigma lobes) and protandry in the flowers of Ferocactus, coupled with the active behavior of bees, are important factors that discourage self-fertilization and promote gene exchange. In addition, there are several stigma lobes that expand when they are receptive and provide a landing platform for pollinators (Fig. 6), facilitating cross-pollination. Therefore, open floral morphology and full exposure of anthers and stigmatic surface probably allows pollination by more than one bee species. Similar floral attributes have been reported for Echinocereus Engelm. (Cota, 1993) and the cactus flora of the southwestern United States (Grant and Grant, 1979b).
Other floral characteristics in Ferocactus are nectar and pollen rewards. Nectar production seems to be relatively low and despite the lack of experimental tests, measurable amounts of this fluid are difficult to detect (at least to the human eye!). Similarly, low nectar production has been reported in the flowers of Echinocactus grusonii Hildm. and other genera of the Cactoideae (Scogin, 1985). Low nectar amount has also been observed in Echinocereus engelmannii (Parry) Lem. (Cota, 1993) species which shares similar floral attributes to Ferocactus. It is likely that low nectar production in the flowers of Ferocactus is compensated for by the correspondingly abundant pollen (as evidenced by the numerous stamens) which probably is the major floral reward. The same pattern (i.e., little nectar and abundant pollen) has been reported in Opuntia lindheimeri Engelm. (Grant and Hurd, 1979). Moreover, pollen amount is also associated with pollinator efficiency: high amounts of pollen produced per flower correlate with higher pollinator efficiency (Richards, 1986). In the wild, the anthers of the numerous stamens in the flower of Ferocactus produce massive amounts of pollen which satisfy demands by pollinators and assure its transfer to another flower. Correspondingly, seed production usually occurs in large numbers, which seems to be sufficient for the perpetuation of the species in nature.



Ants and Extrafloral Nectaries
Extrafloral nectaries are glands that secrete nectar from different parts of the plant. These glands are characteristic in Ferocactus and they occur apically in the areole, in particular in actively growing areoles where new floral structures are in the process of development. The number of extrafloral nectaries per areole varies from one to five in F. gracilis Gates (Blom and Clark, 1980). These glandular structures produce an ant-attracting nectar of varied sugar composition (glucose, sucrose, and fructose) of which only glucose has been detected in higher concentrations early in fruit production in F. acanthodes (Lem.) Britton and Rose (Ruffner and Clark, 1986).
It is generally accepted that extrafloral nectaries play a role in a mutu-alistic relationship between plants and protective ants (Formicidae) (Ben-tley, 1977a, 1977b; Pickett and Clark, 1979). In this protectionist scenario, ants are attracted by nectar produced in these glands, and while foraging for nectar they protect the plants from potential herbivores. Ruffner and Clark (1986) indicated that ant visitation and activity is correlated with nectar production and phenology of the plant. For instance, in F. acanthodes var. lecontei (Engelm.) Lindsay, the composition and amount of extrafloral nectar changes according to plant phenology, and ant visitation is more common during the blooming and fruiting periods. Also, ants depending on liquid food in desert environments may respond to fluctuations in the water content of their food (Ruffner and Clark, 1986).
Though several species of ants visit the plant, some species are more frequent visitors than others. Among the species of ants visiting extrafloral nectaries in Ferocactus, the honey ants (Crematogaster depilis Wheeler) have been reported to be the most frequent in F. gracilis (Blom and Clark, 1980) and F. acanthodes var. lecontei (Ruffner and Clark, 1986). In addition, del Castillo (1982, 1988) documented the presence of Irydomyr-mex and Dorymyrmex in F. histrix in mainland Mexico. Other species associated with the extrafloral nectaries of F. gracilis are Iridomyrmex pruinosum, Camponotus ocreatus, and Pheidole grallipes (Blom and Clark, 1980). Unidentified ants have been observed visiting flowers and extrafloral nectaries in wild plants of F. histrix, F. latispinus (Haw.) Britton and Rose, F. recurvus (Miller) Y. Ito, and F. robustus (pers. obs.).


Seed Morphology
Both seeds and pollen grains are important sources of morphological characters useful for elucidating evolutionary trends and phylogenetic lineages. The importance of the architectural design of these structures is evident when they are applied to the classification of taxa in which characters based on gross morphology are not the best source of data to assess the degree of relationship.
In the Cactaceae, several studies of seed macro- and micromorphology have been successfully applied at different taxonomic levels (e.g., Barthlott and Voit, 1979; Friedrich and Glaetzle, 1983; and Glaetzle and Prestle, 1986). The shape, variation in the testa cells and wall sculpture of seeds provide information of taxonomic importance. In Ferocactus, studies on seed coat attributes are limited; the most relevant is that of Taylor and Clark (1983), in which the importance of seed characters was demonstrated in the classification of the genus. In their study, the authors correlated characteristics of the fruit and the hilum-micropilar rim (HMR) which allowed them to define the two sections of Ferocactus, namely section Ferocactus (seeds with massive HMR, fruit dry, dehiscent by a basal pore, and glossy seed coat) and section Bisnaga (Orcutt) N. P. Taylor (seeds with narrow HMR, juicy fruit and shiny seed coat). To date, the most complete description of the distribution of seed coat attributes in 31 species of the genus is found in Taylor and Clark (1983), which also provide hypotheses about the evolutionary trends of Ferocactus based on micromorphology of seeds, plant habit, and stem morphology.
In terms of the current knowledge of macro- and micromorphoplogical seed characters, the following are some of the defining seed attributes of Ferocactus (variations to this pattern should be expected due in part to convergent evolution, malformation, and genetic factors among others). According to Taylor and Clark (1983), the seeds of Ferocactus subgenus Ferocactus are in general mussel-shaped, and range in size from 1 to 3 mm; the seed coat may be smooth to rugose (with testa-cells tabular to tabular-concave). Some species (F. robustus and F. wislizeni) have a reticulate pattern with verrucose periclinal walls. In the case of F. rectispinus (Engelm.) Britton and Rose and F. acanthodes var. lecontei the seeds have tabular testa cells (Fig. 7A, C, D). The seed coats of the species analyzed exhibited cracks (Fig. 7B) probably due to the exposure of seed to the beam of the Scanning Electron Microscope; the same situation was reported by Taylor and Clark (1983).



Pollen Morphology
Palynology, the study of pollen grains and spores, is another source of comparative data for taxonomic purposes practically at all levels of hierarchy. Reliable palynological data is related to the pollen grain itself, e.g., aperture number, shape and position of apertures, and exine sculptural patterns. In the Cactaceae, it has been shown that the distribution of some morphological features of the pollen grain are useful in defining major lineages. For instance, the three subfamilies (Pereskioideae, Opuntioideae and Cactoideae) can be distinguished based on pollen types and exine structural design (Kurtz, 1948; Tsukada, 1964). Moreover, pollen grain morphology is relatively uniform at the tribal level (Leuenberger, 1976).
At the specific level, pollen analyses are limited in Ferocactus. According to Kurtz (1948) and Tsukada (1964) the pollen grains of F. acanthodes, F. covillei, and F. wislizeni are tricolpate with a punctibaculate ektexine (finely pitted exine), and equatorial diameter ranging from 48-68 (85) urn. Whether these morphological patterns in pollen are common throughout the genus remains unclear, until further studies are conducted in a broader range of species.



Cytological Studies
In general, the classification of the Cactaceae has used traditional methods and has been based on shared morphological characters. In the last 40 years, several tools to address questions in plant systematics and reconstruct phylogeny at different taxonomic levels have blossomed. Cytotax-onomy and modern molecular techniques based on DNA analyses are clear examples of the revolution that plant systematics has experienced to produce genetic based phylogenies.
The study of chromosomes (structures that contain the genetic material) is a good source of comparative data in plant classification. Cytological data of various types (chromosome number, behavior at meiosis, size and shape of chromosomes, and DNA amount) are widely used in plant taxonomy. In the Cactaceae, cytological studies have provided insight into polyploid complexes, hybrid lines, chromosome morphology, and chromosomal rearrangements. Additionally, studies of mitotic and meiotic chromosomes have shown that species of Ferocactus have a base number (lowest haploid humber) of x = 11 (Fig. 8) (which is also the base number for the family; Pinkava et al., 1977). Thus far, all chromosome counts for Ferocactus reported in the literature (17 species) are consistent with the basic number for the family (Table 2). The number of taxa analyzed to date represent 68% of the total of species recognized in Lindsay's taxonomy.
Overall, the morphology of mitotic chromosomes among the species of Ferocactus is homogeneous: primarily metacentric with a few pairs of submetacentric chromosomes, and size ranging from 3 to 7 urn (Figs. 8B-C).
The chromosomes of Ferocactus happen to be the largest among those taxa surveyed in the Cactaceae (Cota et al, submitted); however, the significace of large chromosomes is unclear. Some taxa, Ferocactus acanthodes var. lecontei, F. chrysacanthus (Orcutt) Britton and Rose (Fig. 8B), and F. wislizeni, exhibit satellites or secondary constrictions (small extensions at the end of the arms Of the chromosomes) (Cota et al., submitted). Similar patterns in chromosome morphology have been observed in unrelated genera of the Cactaceae such as Echinocereus Engelm. (Cota, 1991; Cota and Wallace, 1995), Mammillaria prolifera (Miller) Britton and Rose (Johnson, 1980), and Nyctocereus (Berger) Britton and Rose (Palomino et al., 1988).
The uniformity in shape and number of chromosomes and lack of satellites does not necessarily reflect evidence of close phylogenetic relationship. Chromosomal rearrangements at the molecular level may account for the distinctiveness of some species. Alternatively, the presence of satellites in some northern species, such as F. acanthodes var. lecontei, F. chrysacanthus, and F. wislizeni indicates that satellites might be used as taxonomic markers (Cota et al., submitted), and that further analyses of chromosome morphology might be advantageous to understand interspecific relationships in Ferocactus.
As mentioned previously, some of the taxonomic confusion that characterizes Ferocactus has been attributed to hybridization and introgression events. Lindsay (1955a) and more recently Taylor (1984) have both indicated that Baja Californian species are taxonomically problematic due to probable introgressive processes, which brings new genetic attributes to actively evolving species. This hypothesis is based upon observations of individuals with intermediate morphologies in areas where two species are sympatric. Indeed, cases of intermediacy have been suggested in two related species groups: F. gracilis and the F. peninsulae (Engelm.) Britton and Rose group (Taylor, 1984). Polymorphism in spine shape and color in addition to intermediacy of stem and flower characters are common in individuals of the same population. At present, it is unclear to what degree this variation is genetically and/or environmentally induced. Analyses of meiotic configurations in different individuals of F. gracilis (Fig. 8 A) and F. peninsulae failed to provide evidence for hybridity (Cota et al., submitted) at least for the populations examined. Contrary to those hypothesized introgressive events that have contributed to the taxonomic confusion in some Baja Californian species, chromosomal analyses of these taxa at different stages of meiosis show normal pairing and formation of bivalents at diakinesis, which may suggest the lack of hybridization in sympatric species (Cota et al., submitted) or simply a lack of genetic barriers. In addition, previous meiotic counts of some Baja Californian and Mexican mainland species (Beard, 1937; Pinkava et al., 1973, 1977, 1985) have not reported abnormalities during meiosis. To date, in spite of cases of morphological intermediacy, there is no cytological evidence such as univalents, trivalents, chromosome bridges, fragments, lagging chromosomes, or any other abnormalities during cell division to support the possibility of hybridization. Also, hybrids may not show meiotic abnormalities because there is little chromosomal differentiation between species. It should be mentioned, however, that even though there is no evidence of hybridization based on chromosome behavior and morphology, this event can not be ruled out until further analyses are conducted on the remaining species, especially in those problematic populations indicated by Taylor (1984).
In addition to hybridization, polyploidy has been indicated as an important factor in the evolution of certain cacti, e.g., some species of Opuntia (Tournef.) Miller (Pinkava et al., 1985) and Echinocereus (Cota and Phil-brick, 1994; Cota and Wallace, 1996). Both processes are yet to be documented in Ferocactus and is likely that they have not contributed substantially to speciation events in this genus (Cota et al., submitted). The frequency of polyploidy in the subfamily Cactoideae in which Ferocactus is circumscribed is low: approximately 12.5% (Pinkava etal., 1985). Unlike some other genera of the subfamily in which polyploidy is relatively common, e.g., Echinocereus (Cota and Philbrick, 1994; Pinkava etal., 1992; Weedin and Powell, 1978), to date all the species of Ferocactus which have been cytologically investigated are diploid (Table 2). Moreover, F. robus-tus, one of the putatively least derived species (Taylor and Clark, 1983), has not experienced changes in chromosome number. The same apparently stable diploid number has been observed in putative sister genera of Ferocactus such as Echinocactus grusonii and Stenocactus (Echinofossulocac-tus) crispatus (DC) Berger (Katagiri, 1952); Echinocactus polycephalus (Engelm.) Bigel. (Pinkava et al., 1977); E. horizonthalonius Lem. and E. texensis Hoppfer (Weedin and Powell, 1978); and 5. pentacanthus (Lem.) Berger (Pinkava and Parfitt, 1982). The homogeneity of chromosome numbers in Ferocactus and allied taxa does not provide useful information for understanding interspecific and intergeneric relationships by correlating il with patterns of chromosomal evolution. Cota et al. (submitted) suggest that the relatively recent origin of the genus may account for the lack of polyploid individuals in Ferocactus and its suspected sister taxa (Echinocactus Link, and Otto and Stenocactus), and that changes in chromosome number have not played a major role in the evolution of Ferocactus. They have hypothesized that chromosome evolution in the genus is likely taking place at the molecular level, and consequently, chromosome rearrangements i i-main cryptic. Indeed, the non-coding region (intron) loss of the chloroplast jicne rpoCl has been reported in representative members of the Cactoideae, including F. flavovirens (Wallace and Cota, 1996). Further cytological investigations with a wider taxonomic sampling within Ferocactus may be useful in determining the extent of the diploid condition and the occurrence "I chromosomal rearrangements. In addition, studies of crossability to examine the genetic basis of taxonomic characters and isozyme analyses to estimate the genetic distance between taxa would certainly be effective detecting cases of natural hybridization and/or introgression in populations where individuals with intermediate morphology are common.
On the other hand, and in spite of the lack of evidence to document natural hybrids based on the study of meiotic figures, artificial hybridization is an approach from which cactophiles obtain bizarre specimens. Intergen-eric hybrids involving Ferocactus are precious novelties, and documented cases are those crosses between F. acanthodes and Leuchtenbergiaprincipis Hook, to produce the hybrid X Ferobergia (Glass, 1966), and the cross between Stenocactus {Echinossofulocactus sp.) X Ferocactus sp. that produced XFerossofulocactus (Rowley, 1980b). Disregarding fertility issues in these intergeneric hybrids, the ability to recover viable offspring from intergeneric crosses probably indicates a relatively low degree of genetic divergence; therefore, the apparent lack of reproductive isolation allows chromosomes to pair at meiosis to create artificial hybrids. If artificial and even naturally occurring intergeneric hybrids are created within certain groups of cacti, then it is quite likely that interspecific hybrids of Ferocactus do occur in the wild as suggested by Lindsay (1955a) and Taylor (1984). Nonetheless, sources of evidence (other than intermediacy) are still missing.


Current Molecular Phylogenetic Knowledge
Several molecular biological techniques involving macromolecules (in particular chloroplast and nuclear DNA), are being applied to the study of phylogenetic relationships and evolution of certain plant groups. Because the advantages of molecular tools in plant systematics and evolution are discussed at length in several papers (e.g., Avise, 1994; Palmer, 1987; Palmer et al., 1988; Olmstead and Palmer, 1994; Whitkus et al., 1994) I will only mention two of their strengths: 1) they provide virtually unlimited number of characters to analyse and compare in relatively short periods of time, and 2) they are less subject to convergent evolution, so that homoplasy is minimized and phylogenetic reconstruction is less problematic. Previous papers (Wallace, 1986, 1995a, 1995b; Wallace and Cota, 1996; Cota and Wallace, 1996) have explained the importance and application of macro-molecular techniques to the study of cactus evolution in more detail.
Pioneering studies using molecular techniques in the systematics of the Cactaceae at different taxonomic levels were initiated in the early 1990s at Iowa State University under the direction of Robert S. Wallace, and are providing new insights into the systematics, evolution, and phylogeny of the cactus family. In this facility, various molecular methods [in particular gene sequencing and restriction site variation studies of chloroplast DNA (cpDNA)] are being applied to both North and South American groups of cacti in order to generate a phylogeny of this family. Among the numerous promising projects in progress in this lab are:
1. Within the subfamily Cactoideae, a major study using different regions of the chloroplast genome to elucidate the interspecific relationships in Ferocactus and its relationships with North American columnar cacti of Tribe Pachycereeae. The preliminary results of this project are discussed in the following sections.
2. The assessment of the phylogenetic relationships and major lineages between North and South American columnar cacti.
3. In the subfamily Opuntioideae, a study of the interspecific phylogenetic relationships in Opuntia and related taxa (Pereskiopsis Britton and Rose, Pterocactus MacDouglas and Mir., Quiabentia Britton and Rose, and Tacinga Britton and Rose), using several regions of the chloroplast genome represents a major effort which is providing new insight to clarify the taxonomy of this complex group.
4. The phylogenetic study of the Tribe Rhipsalidae (including Hatiora Britton and Rose, Lepismium Pfeiffer, Rhipsalis Gartn., Rhipsalidopsis Britton and Rose, and Schlumbergera Lem.) to determine the putative terrestrial lineage from which this epiphytic tribe evolved and the pattern for divergence within the tribe.
5. Analyses of the major lineages within the tribe Hylocereeae to clarify their phylogenetic relationships and intergeneric boundaries.
6. An intergeneric study in members of the South American tribes Tricho-cereeae, Cereeae, and Browningieae to investigate their phylogenetic relationships has been initiated recently. Additionaly, several other projects that will complement those outlined here are being planned for the near future.
Relevant descriptions and preliminary results of the main projects in progress in Wallace's lab in diverse lineages of the Cactaceae, and a review of the major techiques in the molecular systematics of this family using cpDNA are provided in Wallace (1995a, 1995b), Wallace and Cota (1996), and Cota and Wallace (1996). The reader is advised to consult these references for a more comprehensive understanding of terminology and methods used in the following section. Molecular Studies in Ferocactus
The putati vely basal phylogenetic position in which Ferocactus has been placed in the Tribe Cacteae of the Subfamily Cactoideae (Fig. 5) (Buxbaum, 1958; Barthlott, 1988; Barthlott and Hunt, 1993) suggests the possibility that several lineages evolved from it. However, the pattern of intergeneric relationships within the members of the Cacteae tribe and possibly taxa of other tribes remains unclear despite a few shared morphological characters that suggest some degree of relationship. The phylogenetic reconstruction of Ferocactus confronts a series of issues, some of which are related to the........


Ну и т. д.

[admin]

(Figs. 8C).

Этот смайлик не я вставил. Видимо текст совпадает с кодом этого смайлика:blink:

[vlani]

Диссертация 1955 года с небольшими поздними добавлениями..
Можно попытаться перевести, по крайней мере выборочно. Написано нормальным языком - переведется более-менее легко

Еще бы вот эту статейку:

Chloroplast DNA Evidence for Divergence in Ferocactus and its Relationships to North American Columnar Cacti
J. Hugo Cota and Robert S. Wallace
Systematic Botany, Vol. 22, No. 3 (Jul. - Sep., 1997), pp. 529-542

[admin]

Диссертация 1955 года с небольшими поздними добавлениями..

Так это может я чего не то отсканировал.
Так как там полно всего и эта диссертация могла быть приведена как одна из глав книги.

Если готов взятся за перевод, то могу отсканировать и выслать. Или можно попробовать PDF файл сделать и где-нибудь выложить для скачивания и соответственно для перевода.

[vlani]

Читая по диагонали -похоже в основе книги как раз та диссертация, но есть еще какой то материал для не-ботаников, и какие то выжимки из новых работ, плюс новые иллюстрации.

Для перевода хорошо бы интерпретированный текст. Тут вроде есть еще люди кто умеет это делать - так что эту часть работы можно делегировать. Одно скаирование - это уже много работы для одного

Можно потихоньку выкладывать перевод - хоть в форуме. Все сразу не выйдет - уж очень много страниц