PHYTOLOGIA 


A journal to expedite publication in plant systematics, evolution, 
phytogeography and vegetation ecology 


www.phytologia.org 
Vol. 91, No. 2, pp 189-258 August, 2009 


Hesperocyparis abramsiana 
var. butanoensis 


PHYTOLOGIA 


(ISSN 00319430) 

Phytologia, a journal for rapid publication in plant systematics, 
phytogeography, and vegetation ecology, is published three times a 
year. Manuscripts on plant genetics, plant breeding, plant physiology, 
and fungal biology and taxonomy will be considered if they are 
explicitly related to topics in plant systematics. 


Managing Editor Executive Editor 
micro-Molecular, Phytochemical General systematics 
and Multivariate Systematics and evolution 
Robert P. Adams Billie L. Turner 
Baylor University University of Texas at Austin 
Baylor-Gruver Lab Austin, TX 78705 
Gruver, TX 79040 billie@uts.cc.utexas.edu 


Robert _ Adams@baylor.edu 
Associate Editors 


Nomenclature General Systematics 
Guy Nesom A. Michael Powell 
2925 Hartwood Drive Dept. of Biology 
Fort Worth, TX 76109 Sul Ross State University 
www.guynesom.com Alpine, TX, 79832 
ampowell@sulross.edu 
Macro-Molecular and Ethnobotany 
Phylogenetic Systematics Martin Terry 
Andrea E. Schwarzbach Dept. of Biology 
Univ. of Texas at Brownsville Sul Ross State University 
Dept. of Biol. Sciences Alpine, TX, 79832 
Brownsville, TX, 78520 mterry@sulross.edu 


andrea.schwarzbach@utb.edu 


Secretary-Treasurer - Subscriptions 
Robert P. Adams 
Phytologia, Box 727, Gruver, TX 79040 
Robert_ Adams@baylor.edu 
Copyright 2009 Phytologia. Texensis Publishing, Gruver, TX 


Phytologia (August 2009) 91(2) 189 


Phytologia 
Contents 
W. H. Blackwell. Chromista revisited: A dilemma of overlapping 
putative kingdoms, and the attempted application of the botanical code 
Bis MINS REYES oc sch «vce cee cea eS MO cols Sie Se aie, rag eibla oR ee Sted 19] 
R. P. Adams and J. A. Bartel. Geographic variation in the leaf essential 
oils of Hesperocyparis (Cupressus) abramsiana, H. goveniana and H. 


mecrocarpa> Systematic implications../../0. Jad ..t eee ek eee 226 


R. P. Adams and J. A. Bartel. Geographic variation in Hesperocyparis 
(Cupressus) arizonica and H. glabra: RAPDs analysis................. 244 


G. L. Nesom. Solidago dispersa (Asteraceae: Astereae) replaces 
solidago ludoviciana as the correct naMe......22....66.25 a0 ad ceree eens 251 


B. L. Turner. Recension of the Mexican species of Sa/via (Lamiaceae), 
ee EA CON OCONIA. 2650.4) ete, Bee ee Ee ee 256 


D. B. Ward. Keys to the flora of Florida: 22, Dicerandra (Labiatae) 


R. P. Adams and J. A. Bartel. Infraspecific variation in Hesperocyparis 


‘\ 


goveniana and H. pygmaea: ISSRs and terpenoid data.................. 20a 


R. P. Adams and J. A. Bartel. Infraspecific variation in Hesperocyparis 


woramsiana: (SSRs and terpenoid data: &....:.2.d2c..ceeeilehe sh sdccke den cen 2a 


G. L. Nesom and J. O. Sawyer. Frangula betulifolia and F. obovata 


Geiiamimiiaceae )iare distinct SPECIES! 4H 2a -:c caves Saad dated Bandas ages 300 — 


Cover Photo: Hesperocyparis abramsiana var. butanoensis. Photos by 
Jim Bartel, see pp. 287-299. Se ee 
LIBRARY 


AUL it) 70NG 


NEW YOhuts 
BOTANICAL GARDEN 


190 Phytologia (August 2009) 91(2) 


B. L. Turner. Taxonomy of Asclepias hirtella and A. longifolia 


(Apocynacede) sc. <.coaadia dee he coe aoe Sede ee Se aes soci a 308 ~ 
B. L. Turner. Three new species of Koanophyllon (Asteraceae: a 
Eupatoricac) from Mexico....0.....5.0..010scvde- veo teaeewses a 312 

G. L. Nesom. Notes on non-native Asteraceae in Texas................ 325," 


G. L. Nesom. Thymophylla tenuiloba and. T. wrightii (Asteraceae: ze 
TAPS Re) sky esas es pcebhind pias ep an eat eRe Leas Joe ase eo ee 333 


B. L. Turner. Biological status of the varietal taxa of Thymophylla 4 
pentachaeta (Asteraceae: Tageteae)....:...2..-0- +. ecstess +0 +ses-s eee 340 


J. R. Reeder and K. Mauz. Panicum coloratum new for Arizona, and ¢ 
Echinochloa holciformis new for the United States...................0 347 


R. P. Adams. Variation in Juniperus durangensis and related junipers - 
(Cupressaceae): Analysis of nrDNA and petN SNPs.................... 3534 


Phytologia (August 2009) 91(2) 19] 


CHROMISTA REVISITED: A DILEMMA OF OVERLAPPING 
PUTATIVE KINGDOMS, AND THE ATTEMPTED 
APPLICATION OF THE BOTANICAL CODE OF 

NOMENCLATURE 


Will H. Blackwell 
Biological Sciences, The University of Alabama, Tuscaloosa, AL 
35487, USA 


ABSTRACT 


It was many centuries before it was realized that all organisms 
were not either plants or animals, and many more years before it was 
understood that the catch-all kingdom “Protista”—proposed to include 
predominantly unicellular, “non-plants” and “non-animals”—was 
heterogeneous and phylogenetically inadequate, encompassing both 
related and unrelated organisms. The probable unity of a particular 
group of protists—viewed as chromophytous algae or pseudofungi (and 
related protozoal forms), and often exhibiting a characteristic 
heterodynamic flagellar pattern—was gradually understood; these 
became separated from Protista, and recognized by various kingdom, 
subkingdom, phylum, or subphylum names. At kingdom level, two 
names and groupings—Chromista and Stramenopiles—have competed, 
among others, for these typically heterokont protists, in a_ partial 
overlay of descriptive information. Stramenopiles have the “tighter” 
circumscription, by virtue of definition based on the occurrence of 
unique, composite, tubular, flagellar hairs. Acceptance of 
Stramenopiles (as a more clearly monophyletic group) was beginning to 
hold sway over the more diverse Chromista (with its less obviously 
related major sub-groupings). However, recent evidence from plastid 
evolution has suggested that the larger, yet still generally monophyletic, 
assemblage of (mostly) heterokont protists—the Chromista—remains 
viable as a putative kingdom, much in the original sense of Cavalier- 
Smith (1981, 1986). Although the matter is still equivocal, the present 
paper notes a return to usage of the kingdom name Chromista— 
representing an assemblage including not only  stramenopilous 
organisms, but also plastid-related groups, i.e., haptophytes and 
cryptomonads. The naming of chromistans has fallen by tradition 
mainly under the botanical code of nomenclature, which, as the other 


192 Phytologia (August 2009) 91(2) 


major eukaryotic code (the zoological code), contains little regulation at 
kingdom level. Difficulty in properly establishing kingdoms, such as 
Chromista, or Straminipila, might be alleviated if a unified code of 
(biological) nomenclature were developed, with guidelines for 
determining/composing kingdom names. As a further point of present 
code deficiency, supra-kingdom ranks (to which yet larger groupings 
such as “Chromalveolata” might be assigned) are not recognized in 
existing, formal codes (botanical, zoological, or bacteriological)—a 
situation that could also be changed through code unification. It is 
important to examine current, proposed, ad hoc naming schemes in 
context of present nomenclatural codes (one of the points of this paper). 
It would be gratifying if systematists that produce future encompassing 
taxonomies (major eukaryotic schemata), and those involved with the 
development of a future code (or codes) of nomenclature, could work in 
consort toward the goal of improved, stabile systems of classification— 
systems not only modern and biologically accurate, but nomenclaturally 
appurtenant as well. Phytologia 91(2): 191-225 (August, 2009). 


KEY WORDS: Chromalveolata, Chromista, codes of nomenclature, 
code unification, complex plastids, Domain, endosymbiosis, 
heterodynamic flagella,  Heterokonta, holophyletic, Kingdom, 
monophyletic, Ochrista, Pseudofungi, Stramenopila, Straminipila, 
supergroup, Supra-kingdom, tubular mastigonemes, — tubular 
mitochondrial cristae. 


Botanical nomenclatural regulation is lax at ranks above 
Family, especially for names not automatically typified—consider 
Articles 16 and 17, /nternational Code of Botanical Nomenclature 
(ICBN), McNeill et al. (2006). Nomenclature at these “higher” levels 
can be confusing. At the rank of Class, three different terminations for 
names are suggested (Recommendation 16A.3, ICBN), depending on 
the “kind” of “plant” in question. This can result in rather closely 
related plant groups—e.g., liverworts, compared with their probable 
relatives, the charophytes (cf. Niklas, 1997; Blackwell, 2003)—having 
quite different sounding Class names. At Division/Phylum level, 
although name-terminations are reduced to two (i1.e., for “plants” vs. 
fungi’), the dual usage in botany now—1.e., Division or Phylum 
(Recommendation 16A.1)—is in itself confusing. At Kingdom level, 


Phytologia (August 2009) 91(2) 193 


there are essentially no Code instructions on how to name these major 
groups. There are, for example, no guidelines for kingdom name 
terminations. By convention, such names often end in “a”—e.g., 
Animalia, Archaebacteria, Biliphyta, Monera—but this is not uniformly 
followed, e.g., Fungi, Plantae, Viridiplantae. Jeffrey (1971) proposed 
that the suffix ‘‘-biota,” implying neither plant, nor animal, nor fungus 
(nor bacterium, for that matter), be used as a standard, “neutral” ending 
for kingdom names; however, this proposal was not widely adopted. 


There is, in fact, a paucity of nomenclatural regulation by the 
botanical code and (particularly) by the zoological code of higher 
categories such as Phylum (Division), Class, and even Order. In the 
case of the botanical code, such “weak regulation” has been attributed 
to the belief that these “upper groups” are too unstable or uncertain in 
delimitation (cf. Gledhill, 1989) for application of rigorous 
nomenclature—such as the principle of priority, or the type method 
(with the exception of automatically typified names, cf. Article 16.2, 
ICBN, 2006; but even this does not clearly apply to Kingdom). The 
dearth of rules and recommendations at the level of Kingdom, however, 
is perhaps more of an historical artifact. Until the mid-nineteenth 
century, it was believed that there were just plant and animal 
kingdoms—hence, no need for detailed regulation of names of 
kingdoms (because the matter was non-controversial). This changed, 
though, when authors such as Hogg (1860) and Haeckel (1866) 
recognized, additionally, a “protoctist” (or “protist’”) kingdom for 
(mostly) unicellular organisms, not clearly plant or animal. While some 
later authors (e.g., Copeland, 1956; Whittaker, 1959) continued to 
emphasize (and increase the recognition of) kingdoms, others (e.g., 
Christensen, 1958) came to believe that Kingdom was a relatively 
meaningless, perhaps artificial, grouping category, and recommended 
removal of Regnum (Kingdom) from the botanical code. Christensen’s 
proposal (1958) did not gain favor, however, and the ICBN (2006) still 
recognizes “Kingdom” (Article 3.1)—even though specific rules for 
“regulating” this category remain absent. 


By mid-twentieth century, the number of kingdoms generally 
accepted had risen to five (cf. Whittaker, 1969). More recently, six to 
nine kingdoms have often been recognized (cf. Edwards, 1976; 
Cavalier-Smith, 1981, 1987, 1993, 2004; Corliss, 1994; Blackwell and 


194 Phytologia (August 2009) 91(2) 


Powell, 1995, 1999; Blackwell, 2004). With proliferation of kingdoms 
came the additional complication of the supra-kingdom category, 
“Domain” (Woese et al., 1990)—a rank not sanctioned by codes of 
nomenclature. In any case, more kingdoms or kingdom-level groups are 
presently recognized than there are codes of nomenclature (There are 
three major organism-based codes, discussed below). Several different 
kingdoms, or parts of kingdoms, are under the umbrella of the botanical 
code alone (cf. Blackwell and Powell, 1999; Blackwell, 2008)—e.g., 
Fungi, Myxomycetes (these being Protozoa, cf. Corliss, 1991), 
Cyanobacteria, and most Stramenopiles (cf. Blackwell and Powell, 
1999)—in addition to plants, obviously the intended objects of this 
code. It may be controversial which code should control the 
nomenclature of a given group of organisms, e.g., Cyanobacteria—in 
this case, the botanical or the bacteriological code (cf. Blackwell, 
2008). 


So, questions remain: Were the kingdoms that we now 
recognize properly established, based on both biology and 
nomenclature? If not, how should they be established? Is it possible to 
determine if a given kingdom name is technically accurate and properly 
applied? Is, for example, the kingdom name-termination appropriate?— 
On what basis is this decided? And, under which code of nomenclature 
should each putative kingdom be “governed?” Cavalier-Smith (1978) 
and Corliss (1983) initially raised questions concerning possible 
nomenclatural consequences of creating multiple kingdoms of 
organisms, a situation readdressed by Blackwell and Powell (1999), and 
that pertains here. These sorts of questions do not necessarily have 
ready answers, nor will I seek to deal with all such questions here (and 
certainly not for all kingdoms). What I wish to address is a special 
confusion concerning two proposed kingdoms—Chromista and 
Stramenopila—that have similar, yet clearly non-identical, 
circumscriptions; i.e., they are descriptively over-lapping. I give, 
subsequently, particular consideration to determination of the usage and 
best application of these particular names and groupings, and to the 
various complications that are attendant. 


Phytologia (August 2009) 91(2) 195 


CHROMISTANS, STRAMENOPILES, AND 
THE BOTANICAL CODE 


The case in point here, Chromista vs. Stramenopiles, is which 
kingdom and name to recognize? In addressing this question, it should 
be decided under which code the nomenclature of these groups should 
fall. Organisms recognized variously as chromistans and stramenopiles 
are neither plants nor animals (cf. Patterson, 1989; Cavalier-Smith, 
1987; Keeling, 2004). Historically, however, the naming of the majority 
of the membership of either alleged kingdom has been in accordance 
with the botanical code (cf. Blackwell and Powell, 1999); thus, this 
question is decided (for now), by precedent, in favor of the ICBN. 
Although this decision is perhaps more clear-cut with Stramenopiles, it 
is nonetheless true that, since the Chromista are also largely 
characterized by chromophytous algal (e.g., chrysophytes, 
xanthophytes, phaeophytes, diatoms) and by pseudofungal lineages 
(e.g., Oomycetes), cf. Cavalier-Smith (1989), their nomenclature 
generally forfeits to the botanical code as well (further discussed 
below). To a lesser extent, some members of “both” putative kingdom 
groups have been named under the zoological code (e.g., certain 
amoeboid or colonial chrysophytes). 


We are saddled at present with the situation of having two 
separate eukaryotic codes, botanical and zoological, plus the 
bacteriological or prokaryotic code—hence, three major organismal 
codes (and, in addition, a code for viruses, and a specialty code for 
cultivated plants). In hindsight, it is apparent that none of these codes is 
a good fit (in biological context) for the naming of organisms 
considered herein (chromistans/stramenopiles). Furthermore, since the 
three main codes were each conceived (more or less independently) to 
facilitate nomenclature of members of the plant, animal and bacterial 
kingdoms respectively, the nomenclature in each code (with some 
exception in the bacteriological code) effectively starts be/ow the level 
of kingdom. The naming procedure for the category of kingdom is 
substantially neglected in the botanical code, and even more so in the 
zoological code. If, however, there were a unified code (cf. Cavalier- 
Smith, 1978; Patterson, 1986; Corliss, 1990; Blackwell and Powell, 
1999; Blackwell, 2008), the question of which code should cover which 
kingdom (e.g., Chromista) need not be asked, since nomenclature of all 


196 Phytologia (August 2009) 91(2) 


organisms (regardless of their “biology”) would be under one code, and 
rules would doubtless be in place for kingdom names. 


But since there is as yet no accepted unified code, the 
nomenclatural default in the case of all eukaryotes (regardless of 
relationships) is, at present, to either the botanical code or the 
zoological code. Whereas some organisms considered to be 
Chromistans (and Stramenopiles), such as the more protozoal 
representatives (e.g., the primitive Bicoecids), have often fallen under 
the zoological code, the majority are (as indicated) still under the 
governance of the botanical code—since most _ chromistal/ 
stramenopilous organisms, accurately or not, have typically been 
referred to as either “algae” or “fungi” (“groupings” traditionally 
covered by the botanical code). In the case of “fungi,” however, it 
should be noted that none of the “chromistal fungi” are actual Fungi, 
but rather are “Pseudofungi”—e.g., Oomycetes, Hyphochytriomycetes, 
and the more protozoan-like Labyrinthulids—these pseudofungi being 
relatively unrelated to true Fungi (Cavalier-Smith, 1986, 1989; 
Alexopoulos et al., 1996; Blackwell and Powell, 2000) and more 
closely related to types of (alveolate) Protozoa, such as ciliates, 
dinoflagellates, and apicomplexans (cf. Cavalier-Smith et al., 1995; 
Van de Peer and De Wachter, 1997; Keeling, 2004). Nonetheless, 
pseudofungi continue to be “covered” (as do true fungi) by the 
botanical code—evidenced, for example, by the reference to 
Oomycetes in item number 7 of the Preamble (ICBN, 2006). “Algae,” 
no longer considered a cohesive phylogenetic construct (cf. Van den 
Hoek et al., 1995, p. 9), are still also generally treated, operationally, as 
“plants” by the ICBN. It remains equivocal whether the nomenclature 
of certain groups of organisms (in the Chromista) such as the 
Pedinellids (cf. Patterson, 1989) and Silicoflagellates (cf. Tappan, 
1980)—difficult to pigeonhole as “algae” vs. “protozoa”—should be 
considered, presently, under the botanical or the zoological code. 


In opting at the present time primarily for the botanical code, 
based on overall membership of assemblages considered here, it might 
be assumed that we would thereby know what rules to follow in 
establishing the appropriate kingdom (Chromista or Stramenopila, or 
other competing names/groupings subsequently discussed). But since 
there essentially aren’t any “kingdom rules” per se—being as which 


Phytologia (August 2009) 91(2) 197 


name came first is largely irrelevant (priority is not binding at kingdom 
level, as extrapolated by comparing Principle IV with Article 11, 
ICBN)—and since none of these names, arguably, is really a typified 
name (see later discussion, however, concerning Chromista)—such 
decisions boil down more to a matter of informed preference than code 
“legality.” This preference is informed, mainly, by asking biologically 
based questions, such as: Which grouping is the most monophyletic 
(pertinent, if one is striving for phylogenetic nomenclature, cf. de 
Queiroz and Gauthier, 1992; de Queiroz, 1997, 2006; Cantino, 2000)? 
Even though a PhyloCode has not been formally endorsed by any 
official, international nomenclatural congress (other than, perhaps, that 
of those promoting the PhyloCode, cf. de Queiroz, 2006), it would 
nonetheless seem logical that at kingdom level we would wish to 
recognize a group that is phylogenetically inclusive (holophyletic)— 
with the caveat that all included sub-groups are not going to be equally 
related. If such can be determined, then, armed with this “phylogenetic 
knowledge,” we would perhaps next seek to assess which name is 
descriptively most appropriate, given the “special biology” of the group 
that it is desired to recognize. After answering biological questions, one 
would presumably want to determine how properly to compose this 
name, including the proper name-ending. Again, in these matters, there 
is no effective counsel from the ICBN at kingdom level. This seems 
ironic, if not a full-blown “Catch-22,” in that one is bound into the 
botanical code for guidance for naming, but there is virtually no 
guidance (in the case of kingdoms). Cavalier-Smith and Chao (1996) 
alluded to inconsistencies (confusion, or lack of instruction as the case 
may be) in the botanical code concerning the establishment of names at 
higher ranks. As suggested (e.g., Blackwell, 2008, and above), such 
situations could be addressed more forthrightly if there were a unified 
code of nomenclature, with clear rules for naming higher categories— 
including kingdoms. 


DISTINCTION OF PROPOSED KINGDOMS: 
CHROMISTA VS. STRAMINIPILA 


Whereas Chromista and Stramenopiles, both, are now rather 
well-known and often accepted names/groupings, usage of the name 
Chromista is slightly longer standing—and Chromista is the more 
inclusive grouping. The kingdom Chromista was formally proposed by 


198 Phytologia (August 2009) 91(2) 


Cavalier-Smith (1981), who subsequently (1986, 1989) provided more 
thorough expositions. The original Latin diagnosis by Cavalier-Smith 
(1981) emphasized: tubular mitochondrial cristae, chloroplast 
endoplasmic reticulum (complex plastids, with extra envelope 
membranes), and the presence of tubular mastigonemes (tubular 
hairs”) on at least one flagellum (cilium). The expansive content of 
kingdom Chromista (Cavalier-Smith, 1986, 1989) encompassed three 
presumably related phyla: I. Cryptophyta or “Cryptista” (the 
cryptomonads or cryptophyceans); II. Heterokonta: including, A. the 
“Ochrista” or chromophytous algae, such as_ chrysophytes, 
synurophytes, pedinellids, | dictyochophytes _(silicoflagellates), 
xanthophytes (tribophytes), | eustigmatophytes, — raphidophytes, 
phaeophytes, and bacillariophytes; B. “Pseudofungi,” 1.e., the 
Oomycetes, Hyphochytriomycetes, and the somewhat more protozoan- 
like labyrinthulids and the related thraustochytrids; and C. certain 
“protozoa,” such as the bicoecids (or bicosoecids) which Cavalier- 
Smith (1986) first recognized as a group, lacking plastids, under the 
Ochrista; and, finally, III. Haptophyta or “Haptomonada” (the 
prymnesiophytes, which include the stratigraphically significant 
coccolithophorids). As discussed below, the grouping which came to be 
known as “Stramenopiles” (Patterson, 1989) is generally equivalent to 
phylum II (Heterokonta) of Cavalier-Smith’s “Chromista.” 


The (usually) two flagella of heterokont chromistan motile 
cells are heterodynamic, with quite different actions or “beats” (cf. 
Sleigh, 1989). Tubular mastigonemes (typically tripartite, flagellar 
hairs)—sometimes known as “‘retronemes” (a more specific, functional 
term), because these generate a reversal of flagellar thrust (Cavalier- 
Smith, 1986, 1989; Round, 1989)—are found on the more anterior of 
the two subapical or lateral flagella (or on the only flagellum in some 
cases). These distinct, composite (three-tubulate) mastigonemes were 
determined to be often associated with a distinct organization of the 
flagellum-to-basal-body ultrastructure, viz. the “transitional helix” (cf. 
Patterson, 1989; Preisig, 1989). Such heterokont Chromista are, as 
indicated, known as “Stramenopiles’—in reference to the tubular 
mastigonemes (cf. Patterson, 1989). Photosynthetic representatives of 
Chromista typically have chlorophylls “a” and “c,” but not “b” 
(Cavalier-Smith, 1986; Jeffrey, 1989). More than one form of 
chlorophyll “c” may be present (Jeffrey, 1989). Distinctive carotenoids 


Phytologia (August 2009) 91(2) 199 


frequently occur (Bjornland and Liaaen-Jensen, 1989), imparting often 
(not exclusively) a golden-brown pigmentation to the plastids. 


Additional groups of stramenopilous (heterokont) algae, that 
is, groups added to the list of Ochrista (or chromophytous Chromista) 
since Cavalier-Smith’s treatments (1986, 1989), — include 
phaeothamniophytes, bolidophytes, and pelagophytes (see e.g., 
Blackwell and Powell, 2000, p. 71). Horn et al. (2007) proposed 
Synchromophyceae as a new class for an amoeboid “heterokontophyte” 
with a peculiar plastid complex. Relationships of certain other groups— 
such as the opalinids and the proteromonads—to Chromista have been 
postulated, but are equivocal (discussed in Blackwell and Powell, 
2000). Regardless, there is no question that the Chromista (sensu 
Cavalier-Smith, 1986) are a diverse assemblage, including forms 
ranging from diminutive golden algae, diatoms, and large brown algae, 
to water molds, slime-nets (labyrinthulids), and related “protozoa” (see 
website: http://www.ucmp.berkeley.edu/chromista/chromista.html). A 
preliminary cladistic analysis of Chromista was presented by Williams 
(1991), supporting relationships among heterotrophic (including 
pseudofungal) and autotrophic (chromophytous algal) members. 


The Kingdom (Regnum) name “Chromista” (Cavalier-Smith, 
1981) apparently stems in part from the Division (apparent Class) 
name, “Chromophycées” (Chadefaud, 1950)—cf. Christensen (1989). 
Other related names, though, are more directly equivalent to 
Chadefaud’s name, such as “Chromophyta” (Christensen 1962, 1989). 
Christensen (1989) formally proposed Division Chromophyta, 
including a Latin diagnosis (emphasizing the absence of chlorophyll 5). 
Cavalier-Smith (1986) had earlier, however, validated Chromophyta as 
a Subkingdom name (Latin diagnosis high-lighting the tubular 
mastigonemes and tubular mitochondrial cristae). Subkingdom 
Chromophyta (name meaning “colored plant”) represents a difficult 
concept, in that—being “above” phylum Heterokonta in Cavalier- 
Smith’s classification—it includes both chromophytous algae (which 
typically have colorful plastids) and pseudofungi (which lack plastids, 
and therefore often lack pigment or special color as well). According to 
Cavalier-Smith (1986), Subkingdom Chromophyta is typified by genus 
Chromophyton. There is indeed a “chrysophyte” genus name 
Chromophyton Woronin (Bot. Zeit. 38: 625, 1880), cf. Index Nominum 


200 Phytologia (August 2009) 91(2) 


Genericorum. However, it is not clear in the ICBN (2006) that 
automatic typification applies to Subkingdom and Kingdom level (see, 
for example, Article 16.2); nor is it clear, even if it did, that 
“Chromophyta” would be the correct name-form based on 
Chromophyton (see Christensen, 1989, but compare his view with 
Articles 10.7 and 16.4). In any case, the nomenclatural propriety of 
Chromophyta (be it considered a divisional or a subkingdom name) 
does not directly affect the legitimacy of Cavalier-Smith’s kingdom 
name Chromista—especially if Chromista is viewed as primarily a 
descriptive name (Article 16.1), viz. “colored protists.” 


Seemingly more pertinent to the question of whether 
Chromista should be the kingdom name recognized is that, prior to 
Cavalier-Smith’s (1981) Chromista, Jeffrey (1971) had proposed a 
similar (if somewhat more polyphyletic) Kingdom, the ““Chromobiota.” 
Jeffrey (1982), however, later modified this to a more monophyletic, 
Subkingdom grouping, the “Chromobionta”—ainserting an “n” into the 
name—a grouping more or less equivalent to phylum Heterokonta of 
kingdom Chromista (cf. Cavalier-Smith, 1986). But, Jeffrey provided 
no Latin diagnosis for either name, Chromobiota or Chromobionta, 
leaving them (technically) nomenclaturally invalid (cf. Article 36.1, 
36.2). Regardless, since priority is only a recommendation above the 
rank of Family (Recommendation 16B, even this not clearly applying to 
Kingdom), and since names (in specific reference here to subkingdom 
names) have no necessary priority outside of their original ranks 
(Article 11.2), there is no obligation (for one reason, or another) to 
employ Jeffrey’s (or Christensen’s, see above) name(s) at Kingdom 
level. Hence, the Kingdom name Chromista may be recognized, and 
attributed to Cavalier Smith (1981), with no requirement to reference 
other, perhaps similarly intended, names. Again, whether one can argue 
(spuriously, I believe) that Kingdom “Chromista” is an automatically 
typified name, based on the stated typification of Subkingdom 
Chromophyta by Cavalier-Smith (1986)—see paragraph above—is a 
matter of debate. However, this point is relatively moot to name 
selection, given the lax position of the ICBN on priority at higher levels 
(especially kingdom). Specific rules for naming Kingdoms (and 
Subkingdoms) would be a helpful addition to the botanical code—or 
better still, to a future, unified code of biological nomenclature. 


Phytologia (August 2009) 91(2) 201 


In consideration of the fact that Chromista, as outlined by 
Cavalier-Smith (1986), constitutes a diverse assemblage, somewhat 
vaguely defined—the name seeming to emphasize the “algal” or 
plastid-bearing representatives more than the “fungal” members— 
Patterson (1989) suggested that “core chromophytes” (primarily the 
heterokont assemblage of chromophytous algae), along with related 
pseudofungal and “protozoan” representatives, be recognized 
(informally, at the time) by a more uniformly appropriate name, 
“Stramenopiles.” Patterson (1989) coined this name (meaning, literally, 
“straw hairs”) emphasizing the distinctive, lineage-defining, tubular 
flagellar hairs (i.e., the composite, tubulate mastigonemes) possessed 
by members of this group. A more precise group is thus suggested by 
the name Stramenopiles than is the case with the more inclusive 
Chromista, although the overlap of these two large groupings is very 
substantial. As has been indicated, Stramenopiles correspond to the 
phylum Heterokonta (Cavalier-Smith, 1986) of kingdom Chromista. 
Haptophytes and Cryptomonads (both groups included in_ the 
Chromista, cf. Cavalier-Smith, 1986) are excluded from Stramenopiles 
(sensu Patterson, 1989, and later publications, e.g., Blackwell and 
Powell, 2000). “Algal” representatives of Stramenopiles—the ochristal 
heterokont groups (goldens, browns, xanthophytes, diatoms, 
pelagophytes, eustigmatophytes, etc.) listed previously—have 
informally been referred to as “stramenochromes” (Leipe et al., 
1994)—acknowledging the tubular mastigonemes as well as the often 
colorful plastids. There appears to be no comparable (“strameno---”) 
designation for pseudofungal or “protozoan” members of this 
heterokont grouping. 


Ultrastructural studies on Chromista (particularly 
Stramenopiles)—such as of the flagellar apparatus and transition zone, 
as well as the flagellar hairs—proved useful in_ establishing 
relationships of member groups, among (and between) Ochrista 
(chromophytous algae) and Pseudofungi (e.g., Hibberd, 1979; 
Moestrup, 1982; Beakes, 1989; Cavalier-Smith, 1989; Patterson, 1989; 
Preisig, 1989; O’Kelly, 1989; Owen et al., 1990a,b; Andersen, 1987, 
1991). Molecular confirmation of the “unity” of Stramenopiles (or 
organisms that would come to called such) was established, among 
others, by Gunderson et al. (1987), Ariztia et al. (1991), Bhattacharya et 
al. (1992), Leipe et al. (1994), Wee et al. (1996), and Honda et al. 


202 Phytologia (August 2009) 91(2) 


(1999). Based on morphological and molecular information, the 
Stramenopiles came to be viewed as a kingdom or kingdom-like 
category (i.e., a “crown” group) by Leipe et al. (1994), Blackwell and 
Powell (1995, 1999), Alexopoulos et al. (1996), Van de Peer and De 
Wachter (1997), and Sogin and Silberman (1998). The name 
“Stramenopile” (originating, as indicated, with Patterson, 1989) found 
its way into textbooks of phycology (e.g., Lee, 1999), and 
Stramenopiles were recognized in selected biological diversity texts, 
e.g., Barnes (1998). In their introductory college biology textbook, 
Campbell et al. (1999) put forward this group as a “candidate 
kingdom,” employing a formalization of the name, “Stramenopila.” 
Alexopoulos et al. (1996) had earlier made use of “kingdom 
Stramenopila” in correctly asserting that organisms morphologically, 
nutritionally and ecologically thought of as “fungi” actually encompass 
more that one kingdom—Fungi, Stramenopila, and various Protist 
groups (or Fungi, Chromista, and Protozoa, cf. Beakes, 1998). 
Blackwell and Powell (2000) presented a detailed consideration of (and 
support for) the phylogenetic integrity of the overall stramenopilous 
assemblage. Ideas on the filiation of the numerous member groups of 
Stramenopiles are found in Sogin and Patterson (1995, Tree of Life 
Web Project) and Blackwell and Powell (2000). Some authors have 
continued to use the name Stramenopiles (Reyes-Prieto et al., 2007), 
while others (e.g., Baldauf et al., 2000) recognized the stramenopile 
grouping, but employed other names—in this latter case the generally 
equivalent category, Heterokonta, of Cavalier-Smith (1986). 


In spite of the recognition mentioned above, it was apparently 
not until the book, Straminipilous Fungi, published by Dick (2001), that 
Stramenopiles were formally proposed (Latin diagnosis presented) as a 
Kingdom—viz., kingdom “Straminipila.” Dick’s circumscription 
appears primarily to include pseudofungal organisms (By whatever 
names employed, it is these that are enumerated)—although he spoke 
(pursuant to the diagnosis) of “coevolutionarily linked endosymbiont 
characters,” including plastid and chlorophyll features, in seeming 
reference to “algal” representatives. Dick does note in introductory 
discussion that “biflagellate fungi” and chromophyte algae, as well as 
labyrinthulids for example, are unified by the “straminipilous 
flagellum” —i.e., the anterior “tinsel” flagellum of previous discussion, 
bearing composite, tubular mastigonemes. It is plausible that Dick 


Phytologia (August 2009) 91(2) 203 


intended to include chromophytous algae (and bicoecids) by his 
statement in the diagnosis concerning “organisms that originally 
possessed, or evolved to possess” such features as heterokont flagella 
and straminipilous scales. In any case, it seems a little strange that Dick 
refers to straminipilous “fungi,” since, as he himself notes, these are not 
true fungi. As for nomenclatural detail, indication by Dick of the 
holotype of kingdom Straminipila, as phylum Heterokonta Cavalier- 
Smith [1986], is unnecessary since “Straminipila” is a descriptive, not a 
typified, name (cf. Article 16.1, ICBN)—and the type method does not 
otherwise apply above the rank of Family (compare Articles 7.1 and 
16); requirement for citation of type for validation purposes is, in fact, 
primarily at genus level or below (Article 37). Dick’s spelling of the 
name of this kingdom is unique, viz. “Straminipila.” He not only 
altered the spelling to “Straminipila” (from, presumably, 
“Stramenopila”), he listed his name (alone) as author of the kingdom 
(regnum). This assignment of authorship by Dick (to himself) is 
technically correct, although, as has been indicated, Patterson (1989) 
originated the informal name (and the concept of) “Stramenopiles,” and 
others (as mentioned), prior to Dick, used the name Stramenopila. 
Dick’s alteration of the connecting vowel in the name (from “o” to “i’’) 
is appropriate (cf. Stearn, 1983, p. 269). However, there was nothing 
incorrect about the spelling of the second syllable of the name 
“Stramenopila” (based on Latin, stramen), as given in the kingdoms 
listed by Alexopoulos et al. (1996) and Campbell et al. (1999)— 
although, these were, of course, not intended as formal kingdom 
proposals (no Latin diagnoses provided). If stramen, a noun, is (in the 
name Straminipila) employed adjectivally (cf. Stearn, 1983, p. 267), 
i.e., deriving from stramineus (cf. Simpson, 1968), then Dick’s spelling 
(Stramin-i-pila) would be acceptable. Dick (2001), however, indicated 
the etymological derivation to be from stramen [the noun]—this being 
equivalent to Patterson’s (1989) original usage. But, even if one accepts 
Dick’s kingdom, name and spelling, Straminipila, it would not seem 
inappropriate to cite authorship as Patterson ex Dick (cf. Article 46), 
since Patterson (technicalities aside) generated the name basis and 
originated the construct of what would become this “kingdom.” And, if 
kingdom Straminipila is recognized (regardless of spelling), it should 
be rendered convincingly more inclusive (i.e., formally emended, cf. 
Recommendation 47A)—in the sense of Stramenopiles as 
circumscribed, for example, by Patterson (1989), Leipe et al. (1994) 


204 Phytologia (August 2009) 91(2) 


and Blackwell and Powell (2000)—so that “chromophytous algal” and 
“protozoan” member groups are definitively included (listed, and 
accorded equal importance to the “pseudofungal” representatives 
emphasized by Dick, 2001). This suggested inclusiveness is especially 
pertinent given recent evidence of (not plastids but) plastid-associated 
genes in Oomycetes (cf. Tyler et al., 2006; Bailey, 2008; Sanchez- 
Puerta and Delwiche, 2008), indicating further relationship of algal and 
pseudofungal representatives of stramenopiles. However, formal 
emendation becomes truly important only if Straminipila is selected as 
the kingdom to best represent heterokont chromistans—rather than 
simply recognizing this group as, for example, phylum Heterokonta of 
kingdom Chromista  (Cavalier-Smith, 1986). If accepted, 
“Stramenipili”—the first half of the name based on the Latin noun, 
stramen (straw), and the second half based on the Latin noun, pilus, pili 
(hair, hairs), cf. Simpson (1968)—might be a preferable spelling (to 
Dick’s “Straminipila’”), and more comparable to Patterson’s original, 
informal “Stramenopiles.” Such orthographic changes are permitted (if 
justifiable) by the ICBN without invalidation of the standing name, 
authorship or date of publication (cf. Articles 32.7 and 60.1), 1.e., the 
validating author would still be Dick (2001). And, recall (first 
paragraph of text following Key Words), there is no rule (cf. ICBN) 
that kingdom names must end in “-a.” The ending, “pili” (of 
Stramenipili) is not only permissible, it would unambiguously satisfy 
the requirement that the name be treated as a noun in the plural (Article 
16.1). However, such points concerning spelling (as those concerning 
emendation) fade in significance if Straminipila (Dick) is not favored as 
a kingdom over Chromista. 


It might be assumed that the name Straminipila (or 
Stramenopila, or Stramenipili—depending on interpretations of 
etymology and orthography) should be selected for the kingdom in 
question, because of the relatively cohesive phylogenetic 
circumscription of this group (cf. Patterson, 1989; Leipe et al., 1994; 
Blackwell and Powell, 1999, 2001; Blackwell, 2004). Stramenopiles 
are restricted to organisms that are actually “heterokont,” implying the 
presence of composite (usually three-parted), tubular mastigonemes on 
the more forward of two flagella (or the only flagellum in some cases). 
Such unique flagellar appendages are considered lineage-defining (cf. 
Leipe et al., 1994; Blackwell and Powell, 2000). Recent evidence has 


Phytologia (August 2009) 91(2) 205 


indicated that tubular mastigonemes of stramenopiles (Yamagishi et al., 
2007, studying Ochromonas) are not only — structurally but 
compositionally different from those of the simple mastigonemes of 
green algae, such as Chlamydomonas. In the interest of avoiding 
semantic confusion, proteins composing tubular mastigonemes (the 
mastigonemes, of course, externally attached to flagella) appear to be 
unrelated to tubulin proteins of actual microtubules (of which flagella, 
and certain other cytoskeletal elements, are composed). But, regardless 
of the seeming distinctiveness of Stramenopiles, there are 
complications. In the more broadly cast kingdom, Chromista, additional 
groups are included and must be considered—viz., the cryptomonads 
and the haptophytes—even if these have been placed in different phyla, 
or in some cases subkingdoms, from heterokonts (Cavalier-Smith, 
1986, 1989). Pursuant to Cavalier-Smith’s initial expositions, certain 
authors have apparently found haptophytes and cryptomonads to be 
relatively unrelated to the heterokont assemblage (i.e., to 
Stramenopiles)—see, for example, Daugbjerg and Andersen (1997) 
concerning haptophytes, and Van de Peer and De Wachter (1997) 
regarding cryptomonads. This viewpoint (including the consideration 
that “Chromista” was possibly too broad of a construct) would seem to 
support recognition of a separate kingdom Straminipila (as by Dick, 
2001) for truly heterokont organisms. Other authors (e.g., Bhattacharya 
and Medlin, 1995; Cavalier-Smith, 2002), however, have appeared to 
indicate a degree of relationship between heterokonts (stramenopiles), 
cryptophytes and haptophytes—and if this is so, a kingdom 
Straminipila would perhaps be too limiting, and a broader construct 
(Chromista) would be favored. So, how does one decide whether major 
chromistal groups are substantially related? 


The monophyly of the pseudofungal groups of chromistans 
has not (in recent times) been substantially in question (cf. Blackwell 
and Powell, 2000). Now, in consideration of “algal” representatives, 
information has come to light to suggest that there was a common, 
eukaryote/eukaryote (1.e., “secondary”) endosymbiosis—involving a 
red algal endosymbiont—connecting (through common plastid 
ancestry) the cryptomonad, haptophyte and heterokont ‘“algae”—cf. 
Cavalier-Smith (1992, 2002), Delaney et al. (1995), Delwiche (1999), 
Palmer (2003), Bhattacharya et al. (2004), Keeling (2004), Li et al. 
(2006), and Reyes-Prieto et al. (2007). Possibly, more than one such 


206 Phytologia (August 2009) 91(2) 


major secondary (or even a tertiary) endosymbiotic event was involved 
(Sanchez-Puerta and Delwiche, 2008). But in any event, the general 
consensus of references cited above (among others) suggests that the 
diverse “algal” (1.e., plastid-containing) representatives of the 
Chromista (sensu  Cavalier-Smith, 1986) are also, broadly, 
monophyletic (that is, with regard to origin of their plastids, Le., 
involving the same, an identical, or a very similar, secondary 
endosymbiosis). The kingdom Chromista, as conceived by Cavalier- 
Smith (1981, 1986, 1989), thus represents not only a larger grouping of 
organisms (than Straminipila), but possibly one that can still be viewed 
(by some measures at least) as monophyletic as well (Cavalier-Smith, 
2002)—even if not as obviously (clearly definably) monophyletic as the 
Stramenopiles. In other words, based on recent knowledge of plastid 
evolution (e.g., Keeling, 2004; Reyes-Prieto et al., 2007), it is not 
unreasonable to consider the Chromista as the more holophyletic—if 
plainly the circumscriptively looser and phylogenetically more 
diverse—of the two assemblages (Chromists and Stramenopiles). 
Therefore, if putative holophylesis (at least in the sense of containing a 
greater number of paraphyletic groups, cf. Bhattacharya et al., 1992; 
Schuh, 2000) is the guideline for kingdom selection, the nod would 
seem to go, for now, to Chromista (over Straminipila); however, the 
matter cannot be considered finally settled. 


IF NOT STRAMINIPILA, IS CHROMISTA 
THE BEST REMAINING OPTION? 


If deciding not to use the name Straminipila (or a related 
spelling) for this assemblage—because it is not the most encompassing 
group—then is one left with Cavalier-Smith’s (1981) name Chromista, 
with its attendant broad circumscription? Perhaps so, but there are 
additional problems. The name “Chromista” is without universal 
applicability of meaning, even within the heterokont assemblage. 
“Chrome” (Greek/Latin: Chroma, Chromus) implies the presence of 
color or pigment; principally, it came to connote the “brown” line of 
algae (Round, 1989), as distinct from “green” or “red” algae. However, 
as noted by Cavalier-Smith (1986) and Round (1989), not all members 
of Chromista are pigmented. Pseudofungi, such as Oomycetes and 
hyphochytrids, and pseudofungal/protozoan representatives such as 
labryrinthulids (as well as the “more protozoan” bicoecids), are without 


Phytologia (August 2009) 91(2) 207 


actual plastids (even though plastid genes may be present in 
Oomycetes, cf. Bailey, 2008). Also, the name “chrome” is vague in 
meaning (simply, “color’)—not precise given the various hues 
encountered in representative chromophytous algal groups (brown, 
golden, golden-brown, reddish-brown, yellow-green, almost grass 
green, and even other hues). And, any suggestion of “Protista” in the 
name Chromista (viz. “chrome-ist” abridging “chrome-protist”) is 
superannuated, since the hodgepodge “protist” or “protoctist” kingdom 
(Haeckel, 1866; Whittaker, 1969; Margulis, 1981; Corliss, 1984) is no 
longer phylogenetically tenable (Cavalier-Smith, 1987, 1993; Corliss, 
1994; Blackwell and Powell 1995, 2001). However, a name such as 
Chromista is not to be rejected because it is not compellingly 
descriptively appropriate (Article 51, ICBN, 2006). For that matter, the 
meaning of “Chromista” is not entirely inappropriate, being applicable 
generally to the chromophytous algal representatives (although there 
are colorless chrysophytes, e.g., as investigated by Belcher and Swale, 
1972). Furthermore, usage of the name Chromista (in the sense of 
Cavalier-Smith, 1986, 1989) has been steadfastly inclusive of rather 
diverse groupings that continue to seem suitable for inclusion (on 
plastid evidence, for example, cf. Keeling, 2004). Before finally 
accepting this kingdom name, however, it should be asked if other 
legitimate, descriptively appropriate names are available for use? 


A kingdom name that preceded Cavalier-Smith’s (1981) 
Chromista was Ochrobionta (Edwards, 1976). Edward’s “Ochrobionta” 
is loosely equivalent to the “Ochrista” (recognized later by Cavalier- 
Smith, 1986), and to “Ochrophyta” (Cavalier-Smith, 1997)—viz., 
“Ochrophytes” (Graham and Wilcox, 2000). In other words, 
Ochrobionta (Edwards) is composed mainly of what would come to be 
viewed as the chromophytous algal component of kingdom Chromista 
(including though, in Edward’s view, cryptophytes in addition to 
ochristal chromophytes). The kingdom name Ochrobionta, however, 
would not now be considered acceptable for several reasons. For one 
thing, “Ochrobionta” was not validly published (no Latin diagnosis). 
Secondly, organisms belonging to the pseudofungal group of 
heterokont chromistans were not covered by Edward’s construct 
(Ochrobionta). As a third point, dinoflagellates (““Pyrrhophyta”) were 
included in Ochrobionta by Edwards—not a desirable placement (as 
presently understood), since dinoflagellates, regardless of ultimate 


208 Phytologia (August 2009) 91(2) 


potential (multiple) plastid connections (cf. Keeling, 2004), are 
probably not as immediately related to chromophytous algae of the 
Chromista as they are to (other) Alveolate Protozoa (cf. Cavalier-Smith 
et al., 1995; Hausmann and Hiilsmann, 1996; Blackwell and Powell, 
2001; Yoon et al., 2005). Finally, Edwards (1976) spoke of “a 
preponderance of carotenoids over chlorophylls” in members of his 
Ochrobionta, seeming to downplay the role of chlorophyll which is still 
the primary photosynthetic pigment in these organisms—and, some 
chromophytes are indeed decidedly greenish in coloration, especially 
certain members of the Xanthophyceae (a fact which Edwards, 1976, 
acknowledged). A proposed kingdom mentioned previously, 
Chromobiota Jeffrey (1971), though similar to Edward’s Ochrobionta, 
did include some pseudofungi (i.e., as presently known). However, 
Jeffrey’s Chromobiota is otherwise beset with the same circumscriptive 
and nomenclatural problems as Edward’s “kingdom’—e.g., inclusive 
of dinoflagellates, lacking Latin diagnosis. Edward’s (1976) apparently 
partially patterned his kingdom (Ochrobionta) after Jeffrey’s (1971) 
Chromobiota. 


As for other “kingdom” name possibilities, a perhaps more 
serious candidate, Heterokonta, would appear to be available, and some 
recent authors (e.g., Baldauf et al., 2000) have employed this name. The 
name “Heterokonta” is descriptively applicable to the distinct, 
heterodynamic flagella—one forwardly directed pleuronematic 
(“tinsel” or “hairy”) flagellum (bearing tubular, reverse-thrusting 
mastigonemes), and one, sometimes trailing, smooth, whiplash 
flagellum (with a more typical flagellar motion)—of heterokont 
chromistan groups that are biflagellate (the majority), cf. Moestrup 
(1982), Cavalier-Smith (1986), Van den Hoek et al. (1995). However, 
there are problems. “Heterokonta” was formally established—Latin 
diagnosis focusing on tubular mastigonemes of the anterior flagellum of 
heterokonts: “algal,” “pseudofungal,” etc.—as a Phylum (Division) 
name by Cavalier-Smith (1986), not a Kingdom name. And, as pointed 
out, a name does not have priority outside its own rank (Article 11.2, 
ICBN)—even if we allow that priority carries any force at these upper 
ranks (kingdom, subkingdom, phylum, subphylum, etc.)—although a 
given descriptive name may in fact be used at different ranks (Article 
16.1). Earlier, Cavalier-Smith (1978) had informally (no Latin 
diagnosis) suggested ‘“Heterokonta” as a kingdom name—but for a 


Phytologia (August 2009) 91(2) 209 


heterogeneous assemblage, including not only chromophytous algae 
and Oomycetes, but also chytrids (which are true fungi), Myxomycetes 
(i.e., slime molds, which are Protozoa), and Foraminifera (also 
Protozoa); yet, this unwieldy grouping did not include cryptophytes 
(which are usually considered to be chromistans). Subsequent to better 
understanding, Cavalier-Smith (1986) abandoned Heterokonta as an 
overly diverse kingdom concept, in favor of the more circumspect 
divisional usage of the name (i.e., for a grouping generally equivalent 
to what would subsequently be termed Stramenopiles, cf. Patterson, 
1989; Leipe et al., 1994; Blackwell and Powell, 2000). Later, 
perplexingly, Cavalier-Smith (cf. 1995, 1997) “raised” Heterokonta to 
“infrakingdom” (= subkingdom?, cf. Article 4.2). In further 
complication, Cavalier-Smith’s (1986) phylum Heterokonta, though 
well-defined phylogenetically, is readily confused with the pre-existing 
(much older) name Heterokontae (cf. Luther, 1899; Pascher, 1925; 
Fritsch, 1935). Heterokontae, in the sense of these latter authors, is 
generally equivalent to the algal Class, Xanthophyceae (Tribophyceae, 
cf. Ott, 1982); the name (““Heterokontae’’) thus applies primarily to only 
a limited subset of Heterokonta (sensu Cavalier-Smith, 1986). 


Another (somewhat older) version of the phylum name 
“Heterokonta” (Cavalier-Smith, 1986) is “Heterokontophyta” (Van den 
Hoek, 1978)—likewise used as a phylum name (or seemingly so) by 
several authors (e.g., Moestrup, 1982, 1992; Van den Hoek et al., 1995; 
Horn et al. 2007; Sanchez-Puerta and Delwiche, 2008). 
‘“Heterokontophyta” has been applied more to algal than pseudofungal 
representatives of heterokonts. Cavalier-Smith and Chao (1996), 
however, pointed out that the name Heterokontophyta was questionably 
validly published, and favored use of Ochrista instead—It should be 
noted, though, that Ochrista (Cavalier-Smith, 1986) was published as a 
subphylum (subdivisional) name, not a phylum (division) name. 
Regardless, Lee (1999) used Heterokontophyta, de facto, in general 
correspondence to subphylum Ochrista of Cavalier Smith (1986). No 
matter the exact previous rank, name permutation, or usage employed, a 
pragmatic problem with a potential kingdom Heterokonta (or 
Heterokontophyta) is that not all chromistans are morphologically 
“heterokont,” as the term is precisely defined—implying not just 
flagella of (often) unequal length, but two structurally and functionally 
different flagella on the same cell (cf. Van den Hoek et al., 1995, 


210 Phytologia (August 2009) 91(2) 


glossary, re: “heterokont zoids’). “Heterokonta” was formally founded 
on this flagellar distinction (including the presence of tubular 
mastigonemes on the forward flagellum, cf. Cavalier-Smith, 1986). 
‘“Heterokonta” is, as a consequence, not an inclusive enough category 
for the entire chromistan assemblage. For example, Haptophytes 
usually have two, similar, “apical,” whiplash flagella, plus a 
“haptonema” (a central, superficially “flagellum-like,”’ sometimes 
coiled, appendage—cf. Sleigh, 1989; Van den Hoek et al., 1995). 
Haptophytes are not Heterokonts, yet they still appear to qualify as 
Chromistans (based on knowledge of plastid evolution, cf. 
Bhattacharya et al., 2004; Keeling, 2004). “Heterokonta” (sensu 
Cavalier-Smith, 1986) thus constitutes, even in broadest usage, too 
narrow of a kingdom concept to encompass all chromistans, as 
historically and presently recognized. 


In final analysis, ““Chromista” (as conceptualized by Cavalier- 
Smith, 1981, 1986, 1989) remains the most applicable name for the 
over-all group of heterokont and potentially related organisms 
discussed herein as a kingdom (Reference the listings in the first and 
third paragraphs of the preceding section: “Distinction of Proposed 
Kingdoms...”). The main consideration that might alter future 
acceptance of kingdom Chromista is not the appropriateness of the 
name, or the potential “challenge” of other competing names, but rather 
the question of the degree of relationship of the somewhat disparate, 
major member (chromistal) groups, that continue to be included 
(discussed below). 


CONCLUDING POINTS, CURRENT VIEWS, AND CONCERNS 
(Not only Kingdoms and their delimitation, but “Supergroups”) 


Though not representing an overwhelming consensus, the 
balance of currently available information indicates that the kingdom 
name Chromista Cavalier-Smith (1981; see also 1986, 1989) is the best 
option for proper application to, and implicit circumscription of, the 
presumed reasonably holophyletic assemblage of chromophytous algal, 
pseudofungal, and related primitive protozoal organisms discussed 
herein (see again, “Distinction of Proposed Kingdoms...” section, 
listings in first and third paragraphs). The Stramenopiles (as delineated, 
for example, in Patterson, 1989; Leipe et al., 1994; and Blackwell and 


Phytologia (August 2009) 91(2) 214 


Powell, 2000) are probably best viewed, presently, as constituting a 
major phylum of kingdom Chromista. It is possible (if one so wished) 
to use the kingdom name Straminipila Dick (2001)—by whatever 
spelling (discussed previously)—as a phylum (division) name, rather 
than Heterokonta Cavalier-Smith (1986), since priority generally does 
not apply above family rank (Article 11.1, ICBN, 2006), and since 
descriptive names (such as Straminipila) may be used, unchanged, at 
different ranks (Article 16.1). The suggested endings, “-phyta” or “- 
mycota,” for divisional names in the botanical code are, in the case of 
Straminipila (or Heterokonta, for that matter), not only inappropriate, 
but constitute merely a recommendation (16A.1)—firm rules for 
properly establishing a name such as this are lacking. Again, in these 
instances, one could wish for a unified code with well-reasoned, 
unambiguous rules for names of “higher” ranks. In any case, though, 
the possibility still exists that Straminipila Dick (2001), if emended to 
be more clearly defined and formally inclusive—e.g., as concerns 
chromophytous heterokont (i.e., certain “chromophytous algal’) 
groups—could eventually be accepted as a kingdom-level category, 
perhaps even replacing the more heterogeneous Chromista. However, 
this replacement would come to bear only if seemingly authenticated 
relationships of  non-stramenopilous chromistan groups to 
stramenopiles are not sustained (see discussions in Harper et al., 2005 
and Sanchez-Puerta and Delwiche, 2008). But, should future evidence 
indicate that cryptomonads and haptophytes (prymnesiomonads) are no 
longer tenable as members in an assemblage containing stramenopiles 
(1.e., within the Chromista), these groups would possibly revert to 
temporary systematic placement in the “catch-all” kingdom Protozoa 
(cf. Blackwell and Powell, 2001). 


Some recent authors have indeed adopted (or re-adopted, as 
the case may be) usage of “Chromista” in the sense of a kingdom name 
(e.g., Bhattacharya et al., 2004; see also the website: 
http://www.ucmp.berkeley.edu/chromista/chromista.html). Given the 
“super-groups” of organisms now recognized (Cavalier-Smith, 1999; 
Palmer, 2003; Bhattacharya et al., 2004; Keeling, 2004; Parfrey et al., 
2006), Chromista (as compared with the more restrictive grouping 
Stramenopiles) is the kingdom which appears more broadly suited 
(further discussed below) for membership within the supra-kingdom 
grouping, Chromalveolata (cf. Cavalier-Smith, 1999; Adl et al., 2005; 


212 Phytologia (August 2009) 91(2) 


Reyes-Prieto et al., 2007)—an assemblage encompassing not only 
chromists, but also the related alveolate protozoa, 1.e., dinoflagellates, 
ciliates and apicomplexans (cf. Bhattacharya and Medlin, 1995; 
Hausmann and Hiilsmann, 1996; Blackwell and Powell, 2001; 
Cavalier-Smith and Chao, 1996; Cavalier-Smith, 2002). 


Relationships between certain Chromists and Alveolates (still 
somewhat equivocal, cf. Sanchez-Puerta and Delwiche, 2008) were 
established, among others, by Cavalier-Smith et al. (1995), Van de Peer 
and De Wachter (1997), and Sogin and Silberman (1998)—a 
connection (based partly on plastid genetics, cf. Cavalier-Smith, 2002; 
Keeling, 2004) that has, so far, generally held up under scrutiny (Ad et 
al., 2005; Parfrey et al., 2006). However, such a phylogenetic 
relationship is perhaps one that is chimaeric (cf. Corliss, 1994; 
Cavalier-Smith, 2002; Parfrey et al., 2006), not necessarily taking the 
composite organism (“holobiont,” cf. Mindell, 1992) into account. 
Recent evidence (Harper et al., 2005; Sanchez-Puerta and Delwiche, 
2008), including evidence from genes additional to those involved with 
plastids, supports a closer relationship of alveolate protozoa with 
heterokont members of the Chromista (i.e., with stramenopiles) than 
with either of the other putative chromist groups: haptophytes 
(prymnesiomonads) or cryptomonads (cryptophytes)—see also Adl et 
al. (2005). Among chromistan organisms, thus, there may be only a 
distant overall (host-cell?) relationship between stramenopiles (true 
heterokonts) and either cryptomonads or haptophytes. On the other 
hand, there is some evidence of relationship between these chromistan 
groups: for example, the tubular flagellar hairs of cryptophyceans are 
similar (although bipartite, rather than tripartite) in morphology to those 
of true heterokonts (Moestrup, 1982; Cavalier-Smith, 1989); also, there 
is possibly a sibling relationship between cryptomonads and 
haptophytes (Sanchez-Puerta and Delwiche, 2008); and, some 
relationship of large-subunit (28S) cytoplasmic ribosomal RNA was 
indicated between chromophytous algae [stramenopiles] and 
haptophytes (Perasso et al., 1989). In any case, considering evidence 
pro and con, the recognition of Heterokonta, Cryptophyta, and 
Haptophyta as quite distinct phyla—yet these encompassed within the 
Chromista (Cavalier-Smith, 1986, 1989)—can be viewed as an 
assessment (by Cavalier-Smith) that was probably on target. As pointed 
out by Harper et al. (2005) and Sanchez-Puerta and Delwiche (2008), 


Phytologia (August 2009) 91(2) 213 


more data is needed before final establishment of membership of the 
Chromalveolata (and final re-establishment of the Chromista, in my 
view). Both support and doubt have been expressed concerning the 
“chromalveolate hypothesis” (see Palmer, 2003; Adl et al., 2005; 
Harper et al., 2005; Li et al., 2006; Sanchez-Puerta and Delwiche, 
2008)—1.e., concerning whether this very large grouping can truly be 
viewed as (even generally) monophyletic. Nonetheless, inclusion of 
Chromists and Alveolates in a common super-group is for the time- 
being reasonable (AdI et al., 2005; Yoon et al., 2005; Li et al., 2006), 
and comparable to inclusion of Fungi and Animalia in the Opisthokonts 
(Unikonts). Both “supergroups,” Chromalveolata and Opisthokonta, are 
arguably tenable based on selected morphological and molecular 
grounds (cf. Keeling, 2004; Adl et al., 2005; Parfrey et al., 2006; 
Reyes-Prieto et al., 2007). 


Given the above, the final question on the propriety of 
kingdom name selection (e.g., Chromista) concerns how well the 
kingdom fits (including considerations of phylogeny) with other 
kingdoms (or kingdom components) in the context of larger, super- 
group assemblages now recognized (e.g., Keeling, 2004; Adl et al., 
2005; Parfrey et al., 2006)—Amoebozoa, Opisthokonta, Rhizaria, 
Archaeplastida (Plantae), Chromalveolata, and Excavata. In the case of 
the super-grouping pertinent to the present paper—Chromalveolata 
(grouping Chromista with Alveolata)—the fit would appear to be 
relatively good, given improved, if still controversial, knowledge 
(especially plastid information, cf. Cavalier-Smith, 2002; Bhattacharya 
et al., 2004; Keeling, 2004; Reyes-Prieto, 2007) of phylogenetic 
relationships among major subgroups of these organisms. If it is 
eventually determined (see discussion in Ad et al., 2005; Harper et al., 
2005; Sanchez-Puerta and Delwiche, 2008), however, that haptophytes 
and cryptomonads are sufficiently phylogenetically distant from 
alveolates (and stramenopiles) to be marginalized or even excluded 
from the “Chromalveolata,” then a new supra-kingdom name may need 
to be formulated (based on a more restricted grouping). This new name 
could possibly derive from a combination of the names Straminipila 
(representing a more precise circumscription than Chromista) and 
Alveolata, since truly heterokont organisms (stramenopiles) and certain 
alveolates appear substantially related (Van de Peer and De Wachter, 
1997; Baldauf et al., 2000; Keeling, 2004; Harper et al., 2005; Reyes- 


214 Phytologia (August 2009) 91 (2) 


Prieto et al., 2007)—although the extent of their monophyly is not 
completely resolved (Sanchez-Puerta and Delwiche, 2008). 


As evident from discussion, some recent authors (e.g., 
Keeling, 2004; Adl et al., 2005; Parfrey et al., 2006) have utilized 
quasi-formal “Supergroups” (six in total, see above) instead of 
Kingdoms. Ad et al. (2005) considered these largest groups “similar to 
traditional ‘kingdoms’”—however, they are not (as may be judged from 
Table | in Adl et al., 2005; and Figs. 1, 2 and 3 in Parfrey et al., 2006). 
Rather, such super-groups are inclusive of kingdoms, among other (not 
necessarily coequal) groupings; e.g., the super-group Opisthokonta 
includes Fungi, Metazoa (animals), Choanomonads, and 
Mesomycetozoa. Use of six supergroups in the sense of kingdoms (Adl 
et al., 2005), and a contemporaneous recognition of six (actual) 
kingdoms (Cavalier-Smith, 2004), is potentially confusing (different 
names are typically used for supergroups vs. kingdoms). Parfrey et al. 
(2006) made a limited attempt to sort the matter out. Possibly, future 
codes of nomenclature should serve not only to establish rules for 
naming kingdoms, but should take supra-kingdom assemblages into 
account as well. Kingdom and Supra-kingdom categories should be 
clearly distinguished. In point of fact, however, no supra-kingdom 
ranking (super-group category) is presently covered (or “allowed,” 
depending on one’s point of view) by any official code of 
nomenclature. A related question (scarcely raised to date) 1s, whether 
such “super-groups” should be called “Domains”?—as_ previously 
applied by Woese et al. (1990) to the most major prokaryotic 
groupings, Archaea and Bacteria—a question (re: the six largest 
eukaryotic clusters) for nomenclaturists of eukaryotes to decide. There 
are other options (in addition to “Domain’”) for the appellation of super- 
groups (e.g., “Empire,” “Super-kingdom,” “Supra-kingdom”). Almost 
any option would seem preferable to the current designation of each of 
these largest assemblages by the informal, non-rank-identifiable term, 
“supergroup” (Parfrey et al., 2006). Regardless of which rank-category- 
name is ultimately selected, nomenclatural consistency would be 
desirable for such supervening taxonomic categories. 


With continued emphasis on kingdom and supra-kingdom 
categories, it does not seem that we (as taxonomists) are necessarily 
proceeding just in the direction of “rankless” classifications (cf., 


Phytologia (August 2009) 91(2) pA es 


Hibbett and Donaghue, 1998), toward a systematics of (only?) clades, 
not formal, named, taxonomic ranks. Whether stated or de facto, largely 
formal, hierarchical taxonomic systems still generally hold sway. Ad] et 
al. (2005) indicated that the eukaryote cluster-group system they 
employed is “nameless” and (although they asserted the inherence of a 
“ranked systematics”) is a system “without formal rank designations” — 
i.e., a system “not [formally] constrained...” However, the vast 
majority of the taxonomic groups, including the six main clusters, in 
Adl et al. (2005) bear what appear to be formal names—three of six 
main cluster names stemming from works of Cavalier-Smith. 
Significantly, traditional formal names (for what might be known to 
some as phyla, classes, orders, families, etc.) continue to be used under 
the main cluster names (Adl et al., 2005)—de facto signifying 
particular ranks, even though ranks are not explicitly stated. The 
reference by Adl et al. is useful and comprehensive. However, the 
(rank-unidentified) name mix can be confusing, requiring that one 
search the context of particular names (helpfully, sources are provided). 
In some cases, perplexingly, names of different ranks are apparently 
considered to be at a comparable taxonomic level, e.g., Schizocladia, 
Synurales and Xanthophyceae (see Ad et al., 2005, p. 429). 


The systematic descriptive enumeration of Ad et al. (2005) is 
unquestionably a valuable compilation; yet, it is not (I believe) the 
optimal, ultimate systematic approach to the higher level classification 
of eukaryotes. Though divided orderly into six clusters (of uncertain, 
though one would assume the same, “formal” rank), it is otherwise 
much of an “information-board presentation,” with items pasted from 
zoological, botanical, mycological, and protistological taxonomy. It 
would seem that, rather than attempting to piece together an ad hoc 
phylogenetic system such as Adl et al. (2005)—some potential 
nomenclatural pitfalls in ad hoc systems discussed in Blackwell 
(2002)—it would be prudent in the long run to take the trouble to 
render consistent the formal, upper-level nomenclature of eukaryotes, 
modifying not only the naming scheme employed but codes of 
nomenclature as well (an effort involving two at least partially different 
groups of workers—evolutionary systematists and nomenclaturists). In 
other words, the most desirable approach would be that of bringing 
both taxonomic/evolutionary schemata and nomenclatural codes into 
accord. If present codes of nomenclature are ill-tuned to the task, it 


216 Phytologia (August 2009) 91(2) 


logically follows that efforts to rebirth a formal BioCode might be 
appropriate, given the perceived need by some (cf. Cavalier-Smith, 
1978; Corliss, 1983, 1990; Patterson, 1986; Blackwell and Powell, 
1999; Blackwell, 2008) for code reform. The BioCode draft (Greuter et 
al., 1996—minor revision done in 1997) did not meet with success, and 
was not adopted. It is pointless to debate its merits here; however, the 
draft BioCode did at least purport to add the rank “domain” (above 
kingdom) to existing ranks (of the botanical code). Perhaps future 
BioCode efforts (cf. Hawksworth, 2007), no matter what form the 
document might take, could give detailed consideration to kingdom, 
subkingdom, and supra-kingdom nomenclature—addressing what ranks 
should be recognized, and how names appropriate to these ranks should 
be formed. By so doing, perhaps a mechanism could be provided 
through which the most comprehensive clades, e.g., “crown clades” (cf. 
de Queiroz and Gauthier, 1992), could be formally recognized as 
“crown taxa” (cf. Van de Peer and De Wachter, 1997). If a future 
version of the BioCode does not prove to be the answer to supra- 
familial nomenclatural problems, development of a unified biological 
code of nomenclature (even if it be, initially, a minimal or “skeleton” 
code) should nonetheless be pursued, hopefully to the eventual outcome 
of acceptance by all factions involved. If, in the process, it is desired to 
accommodate particular elements of the unofficial PhyloCode (cf. 
Cantino, 2000; de Queiroz, 2006) in a new, formal, unified code of 
biological nomenclature—thereby enhancing the ‘phylogenetic 
capability” of nomenclature, while maintaining sound nomenclatural 
practice and eliminating undesirable competition between these two 
possible future codes—this would seem a reasonable and appropriate 
way to proceed. 


ACKNOWLEDGEMENTS 


I thank Dr. Juan M. Lopez-Bautista (University of Alabama) 
and Dr. Heather A. Owen (University of Wisconsin — Milwaukee) for 
their thoughtful reviews of this manuscript—extremely valuable in 
manuscript revision. I also thank Dr. Martha J. Powell (University of 
Alabama) for critical reading of the manuscript, and for help with 
formatting the paper for publication. 


Phytologia (August 2009) 91(2) 257 


LITERATURE CITED 


Adl, S. M., A. G. B. Simpson, M. A. Farmer, R. A. Andersen, O. R. 
Anderson, J. R. Barta, S. S. Bowser, G. Brugerolle, R. A. Fensome, 
S. Fredericq, T. Y. James, S. Karpov, P. Kugrens, J. Krug, C. E. 
ane; L. A. Lewis, J. Lodge, D. H. Lynti, D: G. Mann, R.M. 
McCourt, L. Mendoza, @. Moestrup, S. E. Mozley-Standridge, T. A. 
Nerad, C. A. Shearer, A. V. Smirnov, F. W. Spiegel and M. F. J. R. 
Taylor. 2005. The new higher level classification of eukaryotes with 
emphasis on the taxonomy of protists. J. Eukaryot. Microbiol. 52: 
339-451. 

Alexopoulos, C. J., C. W. Mims and M. Blackwell. 1996. Introductory 
mycology, 4" edition. John Wiley & Sons, Inc., New York, 
Chichester, Brisbane, Toronto and Singapore. 

Andersen, R. A. 1987. Synurophyceae Classis Nov., a new class of 
algae. Amer. J. Bot. 74: 337-353. 

Andersen, R. A. 1991. The cytoskeleton of chromophyte algae. 
Protoplasma 164: 143-159. 

Ariztia, E. V., R. A. Andersen and M. L. Sogin. 1991. A new 
phylogeny for chromophyte algae using 16S-like sequences from 
Mallomonas papillosa (Synurophyceae) and Tribonema aequale 
(Xanthophyceae). J. Phycol. 27: 428-436. 

Bailey, C. 2008. Oomycetes among algae. Annual meeting of the 
Phycological Society of America, Abstracts, p. 20. 

Baldauf, S. L., A. J. Roger, I. Wenk-Siefert and W. F. Doolittle. 2000. 
A kingdom-level phylogeny of eukaryotes based on combined 
protein data. Science 290: 972-977. 

Barnes, R. S. K. (Editor). 1998. The diversity of living organisms. 
Blackwell Science, Oxford and London. 

Beakes, G. W. 1989. Oomycete fungi: Their phylogeny and relationship 
to chromophyte algae. in The Chromophyte Algae: Problems and 
Perspectives. J. C. Green, B. S. C. Leadbeater and W. L. Diver, eds., 
Clarendon Press, Oxford, UK. 

Beakes, G. W. 1998. Relationships between lower fungi and protozoa. 
in Evolutionary Relationships among Protozoa. G. H. Coombs, K. 
Vickerman, M. A.Sleigh and A Warren, eds., Kluwer Academic, 
Dordrecht, Boston and London. 


218 Phytologia (August 2009) 91(2) 


Belcher, J. H. and E. M. F. Swale. 1972. The morphology and fine 
structure of the colourless colonial flagellate Anthophysa vegetans 
(O. F. Miller) Stein. Brit. Phycol. J. 7: 335-346. 

Bhattacharya, D. and L. Medlin. 1995. The phylogeny of plastids: A 
review based on comparisons of small-subunit ribosomal RNA 
coding regions. J. Phycol. 31: 489-498. 

Bhattacharya, D., L. Medlin, P. O. Wainright, E. V. Ariztia, C. Bibeau, 
S. K. Stickel and M. L. Sogin. 1992. Algae containing chlorophylls 
a + c are paraphyletic: Molecular evolutionary analysis of the 
Chromophyta. Evolution 46: 1801-1817. 

Bhattacharya, D., H. S. Yoon and J. D. Hackett. 2004. Photosynthetic 
eukaryotes unite: Endosymbiosis connects the dots. BioEssays 26: 
50-60. 

Bjornland, T. and S. Liaaen-Jensen. 1989. Distribution patterns of 
carotenoids in relation to chromophyte phylogeny and systematics. 
in The Chromophyte Algae: Problems and Perspectives. J. C. Green, 
B. S. C. Leadbeater and W. L. Diver, eds., Clarendon Press, Oxford, 
UK. 

Blackwell, W. H. 2002. One-hundred-year code déja vu? Taxon 51: 
151-154. 

Blackwell, W. H. 2003: Two theories of origin of the land-plant 
sporophyte: Which is left standing? Bot. Rev. 69: 125-148. 

Blackwell, W. H. 2004. Is it kingdoms or domains? Confusion & 
solutions. Amer. Biol. Teacher 66: 268-276. 

Blackwell, W. H. 2008. “Anabaena,” “Anabaina,” and codes of 
nomenclature: A review of the feasibility of name correction, and a 
possible direction for the future. Phytologia 90: 324-354. 

Blackwell, W. H. and M. J. Powell. 1995. Where have all the algae 
gone, or, how many kingdoms are there? Amer. Biol. Teacher 57: 
160-167. 

Blackwell, W. H. and M. J. Powell. 1999. Reconciling kingdoms with 
codes of nomenclature: Is it necessary? Syst. Biol. 48: 406-412. 

Blackwell, W. H. and M. J. Powell. 2000. A review of group filiation of 
stramenopiles, additional approaches to the question. Evol. Theory 
& Rev. 12(3): 49-88. 

Blackwell, W. H. and M. J. Powell. 2001. The Protozoa, a kingdom by 
default? Amer. Biol. Teacher 63: 483-490. 


Phytologia (August 2009) 91(2) Pp 


Campbell, N. A., J. B. Reece and L. G. Mitchell. 1999. Biology, oo 
edition. Benjamin/Cummings (Addison Wesley Longman), Menlo 
Park, California. 

Cantino, P.D. 2000. Phylogenetic nomenclature: Addressing some 
concerns. Taxon 49: 85-93. 

Cavalier-Smith, T. 1978. The evolutionary origin and phylogeny of 
microtubules, mitotic spindles and eukaryote flagella. BioSystems 
10: 93-114. 

Cavalier-Smith, T. 1981. Eukaryote kingdoms: Seven or nine? 
BioSystems 14: 461-481. 

Cavalier-Smith, T. 1986. The kingdom Chromista: Origin and 
systematics. Prog. Phycol. Res. 4: 309-347. 

Cavalier-Smith, T. 1987. The origin of eukaryote and archaebacterial 
cells. Ann. New York Acad. Sci. 503: 17-54. 

Cavalier-Smith, T. 1989. The kingdom Chromista. in The 
Chromophyte Algae: Problems and Perspectives. J. C. Green, B. S. 
C. Leadbeater and W. L. Diver, eds., Clarendon Press, Oxford, UK. 

Cavalier-Smith, T. 1992. The number of symbiotic origins of 
organelles. BioSystems 28: 91-106. 

Cavalier-Smith, T. 1993. Kingdom Protozoa and its 18 phyla. 
Microbiol. Rev. 57: 953-994. 

Cavalier-Smith, T. 1995. Zooflagellate phylogeny and classification. 
Cytology (St. Petersburg) 37: 1010-1029. 

Cavalier-Smith, T. 1997. Sagenista and Bigyra, two phyla of 
heterotrophic heterokont chromists. Arch. Protistenkd. 148: 253- 
267. 

Cavalier-Smith, T. 1999. Principles of protein and lipid targeting in 
secondary symbiogenesis: Euglenoid, dinoflagellate, and sporozoan 
plastid origins and the eukaryote family tree. J. Eukaryot. Microbiol. 
46: 347-366. 

Cavalier-Smith, T. 2002. The phagotrophic origin of eukaryotes and 
phylogenetic classification of Protozoa. Int. J. Syst. Evol. Microbiol. 
52: 297-354. 

Cavalier-Smith, T. 2004. Only six kingdoms of life. Proc. Roy. Soc. 
Lond., B, 271: 1251-1262. 

Cavalier-Smith, T. and E. E. Chao. 1996. 18S rRNA sequence of 
Heterosigma carterae (Raphidophyceae), and the phylogeny of 
heterokont algae (Ochrophyta). Phycologia 35: 500-510. 


220 Phytologia (August 2009) 91(2) 


Cavalier-Smith, T, E. E. Chao and. M. T. E. P. Allsopp. 1995: 
Ribosomal RNA evidence for chloroplast loss within Heterokonta: 
Pedinellid relationships and a revised classification of ochristan 
algae. Arch. Protistenkd. 145: 209-220. 

Chadefaud, M. 1950. Les cellules nageuses des algues dans 
l’embranchement des  Chromophycées. Comptes  Rendus 
Hebdomadaire, Acad. Sci. Paris 231: 788-790. 

Christensen, T. 1958. Regnum and subregnum? Taxon 7: 270. 

Christensen, T. 1962. Alger. in Botanik, Bd. 2, Systematisk Botanik, 
Nr. 2. T. W. Bocher, M. Lange and T. Sorensen, eds., Munksgaard, 
Copenhagen. 

Christensen, T. 1989. The Chromophyta, past and present. in The 
Chromophyte Algae: Problems and Perspectives. J. C. Green, B. S. 
C. Leadbeater and W. L. Diver, eds., Clarendon Press, Oxford, UK. 

Copeland, H. F. 1956. The classification of lower organisms. Pacific 
Books, Palo Alto, California. 

Corliss, J. O. 1983. Consequences of creating new kingdoms of 
organisms. BioScience 33: 314-318. 

Corliss, J. O. 1984. The kingdom Protista and its 45 phyla. BioSystems 
17: 87-126. 

Corliss, J. O. 1990. Toward a nomenclatural protist perspective. in 
Handbook of Protoctista. L. Margulis, J. O. Corliss, M. Melkonian 
and D. J. Chapman, eds., Jones and Bartlett, Boston. 

Corliss, J. O. 1991. Introduction to the Protozoa. Microscopic Anatomy 
of Invertebrates 1: 1-12. 

Corliss, J. O. 1994. The place of the protists in the microbial world. 
U.S. Federation for Culture Collections, Newsletter 24(3): 1-6. 

Daugbjerg, N. and R. A. Andersen. 1997. Phylogenetic analyses of the 
rbcL sequences from haptophytes and heterokont algae suggest their 
chloroplasts are unrelated. Mol. Biol. Evol. 14: 1242-1251. 

Delaney, T. P., L. K. Hardison and R. A. Cattolico. 1995. Evolution of 
plastid genomes: Inferences from discordant molecular phylogenies. 
in Chrysophyte Algae: Ecology, Phylogeny and Development. C. D. 
Sandgren, J. P. Smol and J. Kristiansen, eds., Cambridge University 
Press, UK. 

Delwiche, C. F. 1999. Tracing the thread of plastid diversity through 
the tapestry of life. Amer. Nat. 154 (Suppl.): S164-S177. 


Phytologia (August 2009) 91(2) 221 


de Queiroz, K. 1997. The Linnaean hierarchy and the evolutionization 
of taxonomy, with emphasis on the problem of nomenclature. Aliso 
15(2): 125-144. 

de Queiroz, K. 2006. The PhyloCode and the distinction between 
taxonomy and nomenclature. Syst. Biol. 55:160-162. 

de Queiroz, K. and J. Gauthier. 1992. Phylogenetic taxonomy. Ann. 
Rev. Ecol. Syst. 23: 449-480. 

Dick, M. W. 2001. Straminipilous Fungi: Systematics of the 
Peronosporomycetes including accounts of the marine 
straminipilous protists, the plasmodiophorids and similar organisms. 
Kluwer Academic, Dordrecht, Boston and London. 

Edwards, P. 1976. A classification of plants into higher taxa based on 
cytological and biochemical criteria. Taxon 25: 529-542. 

Fritsch, F. E. 1935. The structure and reproduction of the algae, Vol. 1. 
Macmillan, New York and Cambridge University Press, UK. 

Gledhill, D. 1989. The names of plants, 2"! edition. Cambridge 
University Press, Cambridge, UK. 

Graham, L. E. and L. W. Wilcox. 2000. Algae. Prentice Hall, Upper 
Saddle River, NJ. 

Greuter, W., D. L. Hawksworth, J. McNeill, M. A. Mayo, A. Minelli, P. 
H. A. Sneath, B. J. Tindall, R. P. Trehane and P. K. Tubbs. 1996. 
Draft BioCode: Prospective international rules for the scientific 
names of organisms. Bull Zool. Nomencl. 53: 148-166. (Minor 
revision of this “draft” was done in 1997) 

Gunderson, J. H., H. Elwood, A. Ingold, K. Kindle and M. L. Sogin. 
1987. Phylogenetic relationships between  chlorophytes, 
chrysophytes, and oomycetes. Proc. Natl. Acad. Sci. USA 84: 5823- 
5827. 

Haeckel, E. 1866. Generelle Morphologie der Organismen. Berlin, G. 
Reimer. 

Harper, J. T., E. Waanders and P. J. Keeling. 2005. On the monophyly 
of chromalveolates using a six-protein phylogeny of eukaryotes. Int. 
J. Syst. Evol. Microbiol. 55: 487-496. 

Hausmann, K. and N. Hiilsmann. 1996. Protozoology, 2™ edition. 
Georg Thieme, Stuttgart and New York. 

Hawksworth, D. L. 2007. Index Fungorum to Species Fungorum and 
the BioCode. Mycological Society of America Annual Meeting 
(2007), Published Abstracts: p. 71. 


222 Phytologia (August 2009) 91(2) 


Hibberd, D. J. 1979. The structure and phylogenetic significance of the 
flagellar transition region in the chlorophyll c-containing algae. 
BioSystems 11: 243-261. 

Hibbett, D. S. and M. J. Donoghue. 1998. Integrating phylogenetic 
analysis and classification in fungi. Mycologia 90: 347-356. 

Hogg, J. 1860. On the distinctions of a plant and an animal, and on a 
fourth kingdom of nature. Edinburgh New Philosoph. J., n.s., 12: 
216-225. 

Honda, D., T. Yokochi, T. Nakahara, S. Raghukumar, A. Nakagiri, K. 
Schaumann and T. Higashihara. 1999. Molecular phylogeny of 
labyrinthulids and thraustochytrids based on the sequencing of 18S 
ribosomal RNA gene. J. Eukaryot. Microbiol. 46: 637-647. 

Horn, S., K. Ehlers, G. Fritzsch, M. C. Gil-Rodriguez, C. Wilhem and 
R. »: Schnetter... 2007. ...Synechroma:., grande. speexi may. 
(Synchromophyeae class. nov., Heterokontophyta): An amoeboid 
marine alga with unique plastid complexes. Protist 158: 277-293. 

Jeffrey, C. 1971. Thallophytes and kingdoms—a critique. Kew Bull. 
252129 We209: 

Jeffrey, C. 1982. Kingdoms, codes and classification. Kew Bull. 37: 
403-416. 

Jeffrey, S. W. 1989. Chlorophyll c pigments and their distribution in the 
chromophtye algae. in The Chromophyte Algae: Problems and 
Perspectives. J. C. Green, B. S. C. Leadbeater and W. L. Diver, eds., 
Clarendon Press, Oxford, UK. 

Keeling, P. J. 2004. Diversity and evolutionary history of plastids and 
their hosts. Amer. J. Bot. 91: 1481-1493. 

Lee, R. E. 1999. Phycology, 3™ edition. Cambridge University Press, 
Cambridge, UK. 

Leipe, Ds D., P.O. Wainright,. J...H. Gunderson, Ds) PettesDiae 
Patterson, F. Valois, S. Himmerich and M. L. Sogin. 1994. The 
stramenopiles from a molecular perspective: 16S-like rRNA 
sequences from  Labyrinthuloides  minuta and _ Cafeteria 
roenbergensis. Phycologia 33: 369-377. 

Li, S., T. Nosenko, J. D. Hackett and D. Bhattacharya. 2006. 
Phytogenomic analysis identifies red algal genes of endosymbiotic 
origin in the chromalveolates. Mol. Biol. Evol. 23: 663-674. 

Luther, A. 1899. Uber Chlorosaccus, eine neue Gattung der 
Susswasseralgen, nebst einigen Bemerkungen zur Systematik 


Phytologia (August 2009) 91(2) 223 


verwandter Algen. Bih. Kgl. Svensk. Vetensk. Handl. 24, Afd. 3, 
No. 13: 1-22. 

Margulis, L. 1981. How many kingdoms? Current views of biological 
classification. Amer. Biol. Teacher 43: 482-489. 

McNeill, J., F. R. Barrie, H. M. Burdet, V. Demoulin, D. L. 
Hawksworth, K. Marhold, D. H. Nicolson, J. Prado, P. C. Silva, J. 
E. Skog, J. H. Wiersema and N. J. Turland (Editors). 2006. ICBN: 
International Code of Botanical Nomenclature (Vienna Code). 
Adopted by the Seventeenth International Botanical Congress, 
Vienna, Austria (July, 2005). Gantner, Verlag; Koeltz Scientific 
Books, Koenigstein, Germany. 

Mindell, D. P. 1992. Phylogenetic consequences of symbioses: Eukarya 
and Eubacteria are not monophyletic taxa. BioSystems 27: 53-62. 
Moestrup, @. 1982. Flagellar structure in algae: A review, with new 
observations particularly on the Chrysophyceae, Phaeophyceae 
(Fucophyceae), Euglenophyceae, and Reckertia. Phycologia 21: 

427-528. 

Moestrup, @. 1992. Taxonomy and phylogeny of the Heterokontophyta. 
in Phylogenetic Changes in Peroxisomes of Algae. Phylogeny of 
Plant Peroxisomes. H. Stabenau, ed., University of Oldenburg Press, 
Oldenburg. 

Niklas, K. J. 1997. The evolutionary biology of plants. University of 
Chicago Press, Chicago and London. 

O’Kelly, C. J. 1989. The evolutionary origin of the brown algae: 
Information from studies of motile cell ultrastructure. in The 
Chromophyte algae: Problems and Perspectives. J. C. Green, B. S. 
C. Leadbeater and W. L. Diver, eds., Clarendon Press, Oxford, UK. 

Ott, D. W. 1982. Tribophyceae (Xanthophyceae): Introduction and 
bibliography. in Selected Papers in Phycology II. J. R. Rosowski 
and B. C. Parker, eds., Phycological Society of America, Lawrence, 
Kansas. 

Owen, H. A., K. R. Mattox and K. D. Stewart. 1990a. Fine structure of 
the flagella apparatus of Dinobryon cylindricum (Chrysophyceae). J. 
Phycol. 26: 131-141. 

Owen, H. A., K. D. Stewart and K. R. Mattox. 1990b. Fine structure of 
the flagellar apparatus of Uroglena americana (Chrysophyceae). J. 
Phycol. 26: 142-149. 

Palmer, J. D. 2003. The symbiotic birth and spread of plastids: How 
many times and whodunit? J. Phycol. 39: 4-11. 


224 Phytologia (August 2009) 91(2) 


Parfrey, L. W., E. Barbero, E. Lasser, M. Dunthorn, D. Bhattacharya, 
D. J. Patterson and L. A. Katz. 2006. Evaluating support for the 
current classification of eukaryotic diversity. PLoS Genetics: 2: 
2062-2073. 

Pascher, A. 1925. Heterokontae. Die Susswasser-flora Deutschlands, 
Oesterreichs und Der Schweiz 11: 1-118. 

Patterson, D. J. 1986. The actinophryid heliozoa (Sarcodina, 
Actinopoda) as chromophytes. in Chrysophytes: Aspects and 
Problems. J. Kristiansen and R. A. Andersen, eds., Cambridge 
University Press, Cambridge, New York, and Melbourne. 

Patterson, D. J. 1989. Stramenopiles: Chromophytes from a protistan 
perspective. in The Chromophyte Algae: Problems and 
Perspectives... J. C.. Green, B.S..C. Leadbeater and Ws LE; Diver, 
eds., Clarendon Press, Oxford, UK. 

Perasso, R., A. Baroin, L. Hu Qu, J. P. Bachellerie and A. Adoutte. 
1989. Origin of the algae. Nature 339: 142-144. 

Preisig, H. R. 1989. The flagellar base ultrastructure and phylogeny of 
chromophytes. im The Chromophyte Algae: Problems and 
Perspectives. J. C. Green, B. S. C. Leadbeater and W. L. Diver, eds., 
Clarendon Press, Oxford, UK. 

Reyes-Prieto, A., A. P. M. Weber and D. Bhattacharya. 2007. The 
origin and establishment of the plastid in algae and plants. Ann. 
Rev. Genet. 41: 147-168. 

Round, F. E. 1989. The chromophyte algae—problems and 
perspectives. A summarizing view. in The Chromophyte Algae: 
Problems and Perspectives. J. C. Green, B. S. C. Leadbeater, and 
W. L. Diver, eds., Clarendon Press, Oxford, UK. 

Sanchez-Puerta, M. V. and C. F. Delwiche. 2008. A hypothesis for 
plastid evolution in chromalveolates. J. Phycol. 44: 1097-1107. 

Schuh, R. T. 2000. Biological systematics: Principles and applications. 
Comstock Publishing Associates/Cornell University Press, Ithaca 
and London. 

Simpson, D. P. 1968. Cassell’s Latin dictionary, 5" edition. MacMillan, 
New York. 

Sleigh, M. A. 1989. Protozoa and other protists. Arnold/Routledge, 
London and New York. 

Sogin, M. L. and D. J. Patterson. 1995. Stramenopiles. Version 01 
January 1995 (under construction). http://tolweb.org/Stramenopiles 
/2380/1995.01.01 in The Tree of Life Web Project, ttp://tolweb.org/ 


Phytologia (August 2009) 91(2) 225 


Sogin, M. L. and J. D. Silberman. 1998. Evolution of the protists and 
protistan parasites from the perspective of molecular systematics. 
Int. J. Parasitol. 28: 11-20. 

Tappan, H. 1980. The paleobiology of plant protists. W. H. Freeman, 
San Francisco. 

Tyler, B. M., S. Tripathy, X. Zhang, P. Dehal, R. H. Y. Jiang., A. Aerts 
and F. D. Arredondo. 2006. Phytophthora genome sequences 
uncover evolutionary origins and mechanisms of pathogenesis. 
Science 313: 1261-1266. 

Van de Peer, Y. and R. De Wachter. 1997. Evolutionary relationships 
among the eukaryotic crown taxa taking into account site-to-site rate 
variation in 18S rRNA. J. Mol. Evol. 45: 619-630. 

Van den Hoek, C. 1978. Algen: Einfiihrung in die Phykologie. George 
Thieme, Stuttgart, Germany. 

Van den Hoek, C., D. G. Mann and H. M. Jahns. 1995. Algae: An 
introduction to phycology. Cambridge University Press, Cambridge, 
UK. 

Wee, J. L., J. M. Hinchey, K. X. Nguyen, P. Kores and D. L. Hurley. 
1996. Investigating the comparative biology of the heterokonts with 
nucleic acids. J. Eukaryot. Microbiol. 43: 106-112. 

Whittaker, R. H. 1959. On the broad classification of organisms. Quart. 
Rev. Biol. 34(3): 210-226. 

Whittaker, R. H. 1969. New concepts of kingdoms of organisms. 
Science 163: 150-160. 

Williams, D. M. 1991. Phylogenetic relationships among the 
Chromista: A review and preliminary analysis. Cladistics 7: 141- 
156. 

Woese, C. R., O. Kandler and M. L. Wheelis. 1990. Towards a natural 
system of organisms: Proposal for the domains Archaea, Bacteria, 
and Eucarya. Proc. Natl. Acad. Sci. USA 87: 4576-4579. 

Yamagishi, T., T. Motomura, C. Nagasato, A. Kato and H. Kawai. 
2007. A tubular mastigoneme-related protein, OCM1, isolated from 
the flagellum of a chromophyte alga, Ochromonas danica. J. 
Phycol. 43: 519-527. 

Yoon, H. S., J. D. Hackett, F. M. Van Dolah, T. Nosenko, K. L. Lidie 
and D. Bhattacharya. 2005. Tertiary endosymbiosis driven genome 
evolution in dinoflagellate algae. Mol. Biol. Evol. 22: 1299-1308. 


226 Phytologia (April 2009) 91(1) 


GEOGRAPHIC VARIATION IN THE LEAF ESSENTIAL OILS 
OF HESPEROCYPARIS (CUPRESSUS) ABRAMSIANA, H. 
GOVENIANA AND H. MACROCARPA: 
SYSTEMATIC IMPICATIONS 


Robert P. Adams 
Biology Department, Baylor University, Box 727, Gruver, TX, 79040 
Robert_Adams@baylor.edu 


Jim A. Bartel 
U.S. Fish and Wildlife Service, Carlsbad Fish and Wildlife Office 
6010 Hidden Valley Road, Suite 101 
Carlsbad, CA 92011-4213 


ABSTRACT 


The compositions of the volatile leaf essential oils of 
Hesperocyparis abramsiana and its putative subspecies (Cupressus 
abramsiana subsp. locatellii, opleri, neolomondensis, and butanoensis) 
are presented along with H. goveniana, H. pygmaea, and H. 
macrocarpa. Most of the putative subspecies of H. abramsiana oils 
contained large amounts of umbellulone (16-21.8%), while the putative 
C. a. subsp. neolomondensis (type 2 oil) and H. pygmaea contained the 
unusual terpene karahanaenone (18.4, 2.2%). With the possible 
exception of C. abramsiana subsp. butanoensis, none of the subspecies 
proposed by Silba (2003) was supported. Phytologia 91(2): 226-243 
(August, 2009). 


KEY WORDS: Hesperocyparis abramsiana; Cupressus abramsiana 
subspp. butanoensis, locatellii, neolomondensis, opleri; Hesperocyparis 
goveniana; Cupressus goveniana subsp. gibsonensis; Hesperocyparis 
macrocarpa;, Cupressus macrocarpa subsp. lobosensis; Callitropsis, 
Cupressaceae; essential oil composition; taxonomy. 


Silba (2003) recently described four new subspecies of 
Cupressus abramsiana Wolf: C. a. subsp. locatellii Silba, Eagle Rock, 
CA; C. a. subsp. opleri Silba, Bracken Brae, Santa Cruz, CA; C. a. 
subsp. neolomondensis Silba, Majors Creek, CA; and C. a. subsp. 


Phytologia (April 2009) 91(1) 227 


butanoensis Silba, Butano Ridge, CA. In addition, Silba (2003) split C. 
goveniana Gordon and C. macrocarpa Hartw. into subspp. gibsonensis 
and /obosensis, respectively. Because these proposed new subspecies 
are morphologically rather indistinct, we collected samples of fresh 
foliage from five separate trees from all of the type localities from 
which we then extracted and analyzed the leaf volatile oils to gather 
additional genetic information. 


Recent DNA sequencing of Cupressus sensu lato (Little et 
al., 2004, Little, 2006) demonstrated that the Western Hemisphere 
species form a well-supported clade quite separated from the Eastern 
Hemisphere cypresses. As a result, Little (2006) not only confined the 
genus Cupressus to the Eastern Hemisphere, he also used Callitropsis 
nootkatensis and its generic epithet for the Western Hemisphere 
cypresses and Xanthocyparis vietnamensis. Debreczy et al. (2009) 
later argued, on morphological grounds, that Ca. nootkatensis is a 
monotypic genus. Sequencing by Adams et al. (2009) of two 
additional nuclear genes and petN-psbM further supported the 
recognition of Ca. nootkatensis as a monotypic genus. Because 
Callitropsis, therefore, should not be applied to the Western 
Hemisphere cypresses, Bartel and Price in Adams et al. (2009) 
described a new genus, Hesperocyparis, for the Western Hemisphere 
cypresses (exclusive of X. vietnamensis and Ca. nootkatensis). 
However, when referring to Silba’s subspecies, Cupressus is used 
throughout this paper to avoid creating any new name combinations. 


The volatile leaf oil of Cupressus macrocarpa has been 
examined by several authors: Briggs and Sutherland (1942); Zavarin et 
al. (1971); Briggs and Kingsford (1974); Malizia et al. (2000); 
Floreani et al. (1982); Cool (2005); El-Ghorab et al. (2007); 
Manimaran et al. (2007). However, only Zavarin et al. (1971) and 
Cool (2005) examined oils from trees native to California. Zavarin et 
al. (1971) confined their analysis to the monoterpenes, and concluded 
that Cupressus macrocarpa was distinct in its leaf oil. Cool (2005) 
focused on the sesquiterpenes of C. macrocarpa and identified several 
new sesquiterpenes. 


228 Phytologia (April 2009) 91(1) 


The volatile leaf oils of Cupressus goveniana appear to have 
only been analyzed by Zavarin et al. (1971) and that report was 
confined to the monoterpenes. 


The monoterpenes of the volatile leaf oils of Cupressus 
abramsiana were reported by Zavarin et al. (1971). Jolad et al. (1984) 
reported the isolation of cupresol from C. abramsiana. 


Cool et al. (1994) reported the occurrence of karahanaenone 
in trace or small amounts in Cupressus abramsiana, C. forbesii, C. 
goveniana, and C. stephensonii. However, they found individuals of 
C. pygmaea and C. sargentii whose oil contained over 20% 
concentrations of karahanaenone. 


No analyses have been made of the volatile leaf oils of the 
new subspecies of Cupressus proposed by Silba (2003). Thus, we 
present below analyses of the leaf essential oils of Hesperocyparis 
abramsiana (C. B. Wolf) Bartel, H. goveniana (Gordon) Bartel, and H. 
macrocarpa (Hartw. ex Gordon) Bartel and compare these oils with 
the Silba’s putative subspecies. 


MATERIAL AND METHODS 


Plant material - Specimens used in this study: H. abramsiana, 
Bonny Doon, Santa Cruz Co., CA, Bartel 1598a-e; C. abramsiana 
subsp. butanoensis, Pescadero Creek County Park, Butano Ridge, San 
Mateo Co., CA, Bartel 1605a-e.; C. abramsiana subsp. locatellii, Eagle 
Rock, Santa Cruz Co., CA, Bartel 1599a-e; C. abramsiana subsp. 
neolomondensis, Wilder Ranch State Park, Santa Cruz Co., CA, Bartel 
1604a-e; C. a. subsp. op/leri, Bracken Brae, Santa Cruz Co., CA, Bartel 
1600a-e; H. goveniana, SFB Morse Botanical Reserve, Monterey Co., 
CA, Bartel 1596a-e; C. goveniana subsp. gibsonensis, Point Lobos 
Ranch, Monterey Co., CA, Bartel 1595a-e; H. pygmaea, Albion Ridge, 
Mendocino Co., CA, Bartel 160la-e; Little River Airport, Bartel 
1602a-e; Casper Little Lake Rd., CA, Bartel 1603a-e; C. macrocarpa 
subsp. /obosensis, Point Lobos State Reserve, Allan Memorial Grove, 
Monterey Co., CA, Bartel 1593a-e, Point Lobos State Reserve, East 
Grove, Bartel 1594a-e; H. macrocarpa, Crocker Grove, Monterey Co., 


Phytologia (April 2009) 91(1) 229 


CA, Bartel 1597a-e. Voucher specimens currently are held in Bartel’s 
personal herbarium in Carlsbad, California. 


Isolation of Oils - Fresh leaves (200 g) were steam distilled for 
2 h using a circulatory Clevenger-type apparatus (Adams, 1991). The 
oil samples were concentrated (ether trap removed) with nitrogen and 
the samples stored at -20°C until analyzed. The extracted leaves were 
oven dried (100°C, 48 h) for determination of oil yields. 


Chemical Analyses - Oils from 5-10 trees of each of the taxa 
were analyzed and both average and individual values are reported. 
The oils were analyzed on a HP5971 MSD mass spectrometer, scan 
time 1/ sec., directly coupled to a HP 5890 gas chromatograph, using a 
J & W DB-S, 0.26 mm x 30 m, 0.25 micron coating thickness, fused 
silica capillary column (see 5 for operating details). Identifications 
were made by library searches of our volatile oil library (Adams, 
2006), using the HP Chemstation library search routines, coupled with 
retention time data of authentic reference compounds. Quantitation 
was by FID on an HP 5890 gas chromatograph using a J & W DB-5, 
0.26 mm x 30 m, 0.25 micron coating thickness, fused silica capillary 
column using the HP Chemstation software. 


Data Analysis - Terpenoids (as per cent total oil) were coded 
and compared among the species by use of the Gower metric (1971). 
Principal coordinate analysis was performed by factoring the 
associational matrix using the formulation of Gower (1966) and 
Veldman (1967). 


RESULTS AND DISCUSSION 


The leaf oils of H. abramsiana are dominated (table 1) by 
umbellone (16-21.4%), terpinen-4-ol (11.9 - 16.8%), and nezukol (6.1 - 
12.1%) with moderate amounts of sabinene (7.5 - 9.6%), and f- 
phellandrene (7.3 - 9.4%). However, the neolomondensis population 
sample contained 3 individuals (neol, table 1) with high amounts of 
karahanaenone and a-terpinyl acetate as found in H. pygmaea. In fact, 
the oils of the neolomondensis - neol plants share two unique 
compounds with H. pygmaea (pyg, table 1): (Z)-nuciferol and B-(Z)- 
curcumen-12-ol as well as similar quantities of sabinene, camphor, 


230 Phytologia (April 2009) 91(1) 


karahanaenone, terpinen-4-ol, 3-thujanol acetate, 4-terpinyl acetate, a- 
terpinyl acetate, and nezukol. 

The leaf oils of H. goveniana were dominated by sabinene 
(15.2 - 26.3%), terpinen-4-ol (9.5 - 15.7%) and nezukol (11.1-26.3%) 
with moderate amounts of y-terpinene (3.1-7.5%). Hesperocyparis 
pygmaea has also been treated as a subspecies of H. goveniana, but for 
this discussion it is treated as a species. The oil of H. pygmaea was not 
typical of H. goveniana in having a very high amount of karahanaenone 
(14.6%, table 1), camphor (8.7%), a-terpineol (3.2%) and a-terpinyl 
acetate (4.2%). 


Table 1 shows that both H. macrocarpa oils are high in 
sabinene (27.0, 23.3%), a-pinene (22.2, 19.8%), terpinen-4-ol (11.7, 
14.7%) with moderate amounts of y-terpinene (5.6, 5.1%), 
isophyllocladene (4.4, 4.9%), myrcene (3.6, 3.2%), B-pinene (2.6, 
2.0%) and phyllocladene (2.3, 2.0%). Of the 71 compounds identified, 
these subspecies seemed differ in only nezukol (0, 2.2%), citronellal 
(0.6, 0.3%) and piperitone (0, 0.3%). Clearly, the oils are nearly 
identical in both composition and component amounts (table 1). 


To examine the overall similarities of the oils, a Principal 
Coordinates Ordination (PCO) was performed on the mean oils of the 
eleven taxa. Figure 1 shows the ordination based on 23 terpenoids 
(each greater than 1.0% of the oil). Hesperocyparis pygmaea is quite 
separated from H. goveniana in this PCO (Fig. 1). As mentioned 
above, three individuals of neolomondensis had oils that were high in 
karahanaenone and a-terpinyl acetate as found in H. pygmaea. The 
mean values of compounds are designated as AN] in table | and figure 
1. The mean values of the other two individuals (low in 
karahanaenone) are designated as AN2 in table | and figure 1. The oil 
of neolomondensis, type 1 (AN1) is most similar to H. pygmaea (Fig. 
1), whereas neolomondensis, type 2 oil (AN2) is most similar to other 
H. abramsiana populations. Though not entirely unique, the Butano 
Ridge population grows on a sandstone outcrop surrounded by a dense 
canopy redwood forest. The oil of butanoensis (AB, Fig. 1) appears to 
be a little different from other H. abramsiana oils. 


Hesperocyparis abramsiana appeared to show some infra- 
specific variation (Fig. 1, Table 1). A PCO analysis of individuals of C. 


Phytologia (April 2009) 91(1) 231 


abramsiana from all five putative subspecies was made and is shown in 
Figure 2. The three individuals from neolomondensis, high in 
karahanaenone, group with H. pygmaea, whereas the other two plants 
of neolomondensis are imbedded with other abramsiana plants (Fig. 2). 
The individuals of butanoensis form a group somewhat distinct from 
other abramsiana individuals. The individuals of H. abramsiana, and 
putative subspecies Jocatellii and opleri are interspersed (Fig. 2). The 
PCO offers no support for the recognition of /ocatellii or opleri 


3(14%) PCO 


23 terpenes 
> 1% conc. 


(> 
iF ) H. macrocarpa 
H. abramsiana 


AB 


1 «— H. pygmaea 


a) 


ie 


H. goveniana 


Figure 1. PCO of Hesperocyparis taxa using 23 terpenes that occurred 
in 1.0% or greater concentration. 
AA =H. abramsiana, Bonny Doon 
AB =C.a. subsp. butanoensis, Butano Ridge 
C. a. subsp. /ocatellii, Eagle Rock 
ANI =C.a. subsp. neolomondensis, high karahanaenone 
C. a. subsp. neolomondensis, low karahanaenone 
C. a. subsp. op/eri, Bracken Brae 


232 Phytologia (April 2009) 91(1) 


PCO H. macrocarpa 


2/ terpenes 
>1% conc. 


H. abramsiana _fA. goveniana 
Me 

subsp. LE) 

“butanoensis" hy 


1. pygmaea 


3(10%) 


= abram., Bonny Doon’ = abram. subsp "locatellii" 
%= abram. subsp. "opleri” @= abram. subsp. “neolomondensis" 


Figure 2. PCO of individuals of H. abramsiana based on 27 terpenes. 


The initial analysis of H. macrocarpa subsp. macrocarpa and 
C. m. subsp. lobosensis average leaf oils (Fig. 1, Table 1) indicated that 
the oils were very similar with scarcely any differences (Table 1). PCO 
analysis of the individuals of H. macrocarpa confirm the overall trend. 
The individuals are interspersed (Fig. 3) implying that these two 
subspecies are behaving as one large population. We found no support 
in the leaf oil data to support the recognition of Silba's C. abramsiana 
subsp. /obosensis. 


Phytologia (April 2009) 91(1) 233 


2(15%) PCO 
28 terpenes 
H. pygmaea_ 79.5% conc. 


H. macrocarpa 


H. abramsiana 


% = macrocarpa 
@ =subsp. "lobosensis" 


Figure 3. PCO of H. macrocarpa and Silba's putative C. m. subsp. 
lobosensis individuals along with mean values of H. abramsiana, H. 
goveniana, and H. pygmaea. 


To examine differences among H. goveniana, putative C. g. 
subsp. gibsonensis, and H. pygmaea, a PCO analysis was made and is 
shown in figure 4. A slight separation exists between H. pygmaea and 
H. goveniana (Fig. 4), while there seems to be no difference between 
H. goveniana and Silba's putative C. g. subsp. gibsonensis, as these 
individuals are intermixed (Fig. 4). The three high karahanaenone 
individuals of neolomondensis were also included in the analysis and 
these seem close, but not conspecific with H. pygmaea (Fig. 4). 
Possibly these plants are the result of relictual or current hybridization 


234 Phytologia (April 2009) 91(1) 


2(11%) PCO 
33 terpenes 
>0.5% conc. 


%# = goveniana 
O = gibsonensis 


abamsiana subsp. 
neolomondensis 
high karahanaenone 


@ = Albion Ridge 
O = Little River 
© = Casper Little Lake 


Figure 4. PCO of individuals of H. goveniana, putative C. g. subsp. 
gibsonensis and H. pygmaea plus three plants of putative C. subsp. 
neolomondensis, high karahanaenone type. 


between H. abramsiana and H. pygmaea. Additional research using 
DNA markers is in progress to aid in resolving this situation. It should 
be noted that while each of the subspecies described by Silba (2003) are 
geographically isolated from one another, the individuals from the 
putative subspecies generally did not cluster geographically, but rather 
were randomly interspersed within each species as one would expect 
with an interbreeding population. 


In summary, the leaf oils of putative C. a. subsp. butanoensis, 
Butano Ridge, showed some differentiation from H. abramsiana, 
Bonny Doon. The three individuals of putative C. a. subsp. 


Phytologia (April 2009) 91(1) 235 


neolomondensis with high karahanaenone (and other components), 
seem intermediate to H. pygmaea. Additional research is needed to 
resolve this problem. 


None of the new subspecies proposed by Silba (2003), (C. 
abramsiana subsp. locatellii, C. a. subsp. opleri, C. goveniana subsp. 
gibsonensis, C. macrocarpa subsp. lobosensis) is supported by 
differentiation of their volatile leaf oils, except possibly C. a. subsp. 
butanoensis. 


ACKNOWLEDGMENTS 


This research supported in part with funds from U. S. Fish and 
Wildlife Service, Grant 814307J011. The findings and conclusions in 
this article are those of the authors and do not necessarily represent the 
views of the U.S. Fish and Wildlife Service. Thanks to Tonya Yanke 
for lab assistance, and thanks to Chuck Bancroft (State Parks Ranger at 
Point Lobos State Reserve), Tim Hyland (State Parks Resources 
Ecologist at Henry Cowell Redwoods State Park), and Mark Schneider 
(Parks Ranger at Pescadero Creek County Park) for providing access to 
the cypress groves. 


LITERATURE CITED 


Adams, R. P. 1991. Cedar wood oil - analysis and properties. In 
Modern methods of plant analysis: Oils and waxes. H. F. Linskins 
and J. F. Jackson, eds., pp. 159 - 173, Springler-Verlag, Berlin, 
Germany. 

Adams, R. P. 2006. Identification of essential oils components by gas 
chromatography/ mass spectrometry, 4th Ed. Allured Publ. Corp., 
Carol Stream, IL. 

Adams, R. P. 2008. Junipers of the world: The genus Juniperus. 2nd 
Edition. Trafford Publ., Vancouver, B.C., Canada. 

Adams, R. P., J. A. Bartel, and R. A. Price. 2009. A new genus, 
Hesperocyparis, for the cypresses of the western hemisphere. 
Phytologia 91(1): 160-185. 

Bartel, J. A., R. P. Adams, S. A. James, L. E. Mumba and R. N. 
Pandey. 2003. Variation among Cupressus species from the 


236 Phytologia (April 2009) 91(1) 


western hemisphere based on random amplified polymorphic 
DNAs. Biochem. Syst. Ecol. 31: 693-702. 

Briggs, L. H. and M. D. Sutherland. 1942. The essential oil of 
Cupressus macrocarpa. J. Org. Chem. 7: 397-407. (grown in New 
Zealand) 

Briggs, L. H. and M. Kingsford. 1974. Further terpene constituents of 
the essential oil of Cupressus macrocarpa. New Zealand J. Sci. 
17: 13-14. (cultivated in New Zealand) 

Cool, L. G. 2005. Sesquiterpenes from Cupressus macrocarpa foliage. 
Phytochem. 66: 249-260. 

Cool, L. G., K. Jiang and E. Zavarin. 1994. Karahanaenone in coastal 
California Cupressus species. Biochem. Syst. Ecol. 22: 857 - 

El-Ghorab, A. H., K. F. El-Massry and H. A. Shaaban. 2007. Effect of 
drying on the chemical composition of the Egyptian Cupressus 
macrocarpa (Hartw. ex Gordon) essential oils and their biological 
characteristics. J. Essent. Oil-Bearing Plants 10: 399- 
411.(cultivated in Egypt) 

Floreani, S. A., J. A. Retamar and E. G. Gros. 1982. Terpenoids of the 
essential oils from species of Cupressus. Anales Assoc. Quim. 
Argentina 70: 663-667. (cultivated in Argentina) 

Gower, J. C. 1971. A general coefficient of similarity and some of its 
properties. Biometrics 27: 857-874. 

Gower, J. C. 1966. Some distance properties of latent root and vector 
methods used in multivariate analysis. Biometrika 53: 326-338. 
Jolad, S. D., J. J. Hoffman, K. H. Schram and J. R. Cole. 1984. A new 
diterpene from Cupressus goveniana var. abramsiana: 5, B- 

hydroxy-6-oxasugiol (cupresol). J. Natl. Prods. 47: 983-987. 

Little, D. P., A. E. Schwarzbach, R. P. Adams and C-F. Hsieh. 2004. 
The circumscription and phylogenetic relationships of Callitropsis 
and the newly described genus Xanthocyparis (Cupressaceae). 
Amer. J. Bot. 91: 1872-1881. 

Little, D. P. 2006. Evolution and circumscription of the true cypresses 
(Cupressaceae, Cupressus). Syst. Bot. 31: 461-480. 

Malizia, R. A., D. A. Cardell, J. S. Molli, S. Gonzalez, P. E. Guerra, 
and R. J. Grau. 2000. Volatile constituents of the leaf oils from the 
Cupressaceae family: part I. Cupressus macrocarpa Hartw., C. 
arizonica Greene and C. torulosa Don species growing in 
Argentina. J. Essent. Oil Res. 12: 59-63. (cultivated in Argentina) 


Phytologia (April 2009) 91(1) 237 


Manimaran, S., S. Themozhil, M. J. Nanjan and B. Suresh. 2007. 
Chemical composition and antimicrobial activity of cone volatile 
oil of Cupressus macrocarpa Hartwig from Nilgiris, India. Natl. 
Prod. Sci. 13: 279-282. (cultivated in India) 

Silba, J. 2003. Field observations of Cupressus in central and coastal 
California, July 2002 to January 2003. J. Intl. Conifer Pres. Soc. 
10: 1-49. 

Veldman, D. J. 1967. Fortran programming for the behavioral sciences. 
Holt, Rinehart and Winston Publ., NY. 

Zavarin, E., L. Lawrence and M. C. Thomas. 1971. Chemosystematics 
of Cupressus. 1V. Compositional variations of leaf monoterpenoids 
in Cupressus macrocarpa, C. pygmaea, C. goveniana, C. 
abramsiana and C. sargentii. Phytochem. 10: 379-393. 


gia (April 2009) 91(1) 


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244 Phytologia (April 2009) 91(1) 


GEOGRAPHIC VARIATION IN HESPEROCYPARIS 
(CUPRESSUS) ARIZONICA AND H. GLABRA: 
RAPDS ANALYSIS 


Robert P. Adams 
Biology Department, Baylor University, Box 727, Gruver, TX, 79040 
Robert _Adams@baylor.edu 


Jim A. Bartel 
U.S. Fish and Wildlife Service, Carlsbad Fish and Wildlife Office 
6010 Hidden Valley Road, Suite 101 
Carlsbad, CA 92011-4213 


ABSTRACT 


RAPDs were analyzed from _ five Hesperocyparis 
(=Cupressus) arizonica and three H. glabra populations. This analysis 
supports the continued recognition of these taxa at the specific level. 
Phytologia 91(2): 244-250 (August, 2009). 


KEY WORDS: Hesperocyparis (=Cupressus) arizonica, H. glabra, 
RAPDs, geographic variation, taxonomy. 


Hesperocyparis (= Cupressus) arizonica (Greene) Bartel and 
H. glabra (Sudw.) Bartel are two closely related taxa that have a 
variable taxonomic history. Table 1 summarizes the taxonomic 
treatments. Wolf (1948) recognized both taxa at the specific level 
(Table 1), while Little (1970) reduced C. glabra to a variety (C. 
arizonica var. glabra). Though Bartel (1993) and Eckenwalder (1993) 
included C. glabra within C. arizonica, Farjon (1998) followed Little 
(1970) in recognizing C. glabra as a variety of C. arizonica (Table 1). 
All these classifications were based strictly on morphology. 


Askew and Schoenike (1982) concluded that bark texture (fibrous 
and not peeling =H. arizonica versus smooth and peeling in thin plates 
or strips =H. glabra) correctly identified the taxa 89% of the time, 
while resin gland occurrence worked 85% of the time. However, Little 


Phytologia (April 2009) 91(1) 245 


(2006) separated these taxa using only resin glands (on < 5% of leaves 
=H. arizonica versus on >5% of leaves =H. glabra) in his key.. 


Table 1. Taxonomic treatments of H. arizonica and H. glabra. 


Treatment arizonica glabra 
Wolf (1948) C. arizonica C. glabra. 
Little (1970) C. arizonica C. arizonica var. 
var. arizonica glabra (Sudw.) Little 
Bartel (1993) C. arizonica (= C. arizonica) 
Eckenwalder (1993) C. arizonica (= C. arizonica) 
Farjon (1998) C. arizonica C. arizonica var. 
var. arizonica glabra (Sudw.) 
Bartel et al. (2003) = C. arizonica C. glabra Sudw. 
D. P. Little (2006) — Callitropsis arizonica Callitropsis. glabra 
(Greene) D. P. Little (Sudw.) D. P. Little 
Adams et al.(2009) |= Hesperocyparis arizonica Hesperocyparis glabra 
(Greene) Bartel (Sudw.) Bartel 


A recent analyses using RAPDs fingerprinting (Bartel et al., 
2003) found H. glabra to be distinct from H. arizonica (Fig. 1). 


Minimum Spanning Network 
83 RAPDs bands C. arizonica 


Dry Beaver Ck. 
Yavapa Co., AZ 


No Mazatzal Mtns. 
2 Gila Co., AZ 
co) 


Tonto Basin 
Gila Co., AZ 


Santa Catalina Mtns. 
Pima Co., AZ 


Dragoon Mtns., 
Cochise Co., AZ 


Pedregosa Mins. 
Cochise Co., AZ 


arizonica 


Greenlee Co., AZ 


Chisos Mtns., TX 


.99 


Figure 1. Minimum spanning network (from Bartel et al., 2003). 


246 Phytologia (April 2009) 91(1) 


Recent DNA sequencing of Cupressus sensu lato (Little et al., 
2004, Little, 2006) demonstrated that the Western Hemisphere species 
form a well-supported clade quite separated from the Eastern 
Hemisphere cypresses. As a result, Little (2006) not only confined the 
genus Cupressus to the Eastern Hemisphere, he used Callitropsis 
nootkatensis and its generic epithet for the Western Hemisphere 
cypresses and Xanthocyparis vietnamensis. 


Little (2006) found very limited nucleotide differences among 
any of the Western Hemisphere Hesperocyparis species. However, he 
did find differences that supported the recognition of H. arizonica and 
glabra (Table 2) and he maintained these two taxa as distinct species. 


Table 2. Summary of DNA sequencing support for the recognition of 
H. arizonica and H. glabra (from Little, 2006). 


Chloroplast genes Nuclear genes 
matK+rbcL+trnl nrDNA(ITS) NEEDLY 
60% support 56% support no differences 


Debreczy et al. (2009) argued on morphological grounds that 
Ca. nootkatensis is a monotypic genus. Sequencing by Adams et al. 
(2009) of two additional nuclear genes and petN-psbM further 
supported the recognition of Ca. nootakensis as a monotypic genus. 
Because Callitropsis should not be applied to the Western Hemisphere 
cypresses, Bartel and Price in Adams et al. (2009) described a new 
genus, Hesperocyparis, for the Western Hemisphere cypresses 
(exclusive of Xanthocyparis  vietnamensis and _ Callitropsis 
nootkatensis). In Adams et al. (2009), Bartel made the new 
combinations of Hesperocyparis arizonica (Greene) Bartel and H. 
glabra (Sudw.) Bartel. The present paper will analyze geographical 
variation within and between H. arizonica and H. glabra. 


MATERIALS AND METHODS 
Collection information for specimens utilized: 


Hesperocyparis arizonica: Adams 9268-9269, Boot Spring, 
Chisos Mtns., Brewster Co., TX, USA; Lab # 9378-9379, 


Phytologia (April 2009) 91(1) 247 


Bartel,1580A,B, upper Bear Canyon, 11.8 min of Houghton Rd along 
Catalina Hwy, N 32 21' 47.9", W 110 42' 50.3", 1695m, Santa Catalina 
Mtns., Pima Co., AZ; Lab # 9380-9381, Bartel, 1581A,B, Stronghold 
Canyon East, 7.3 mi from US 191, along Ironwood Rd., N 31 56 26.9", 
W 109 57' 27.8"1457m, Dragoon Mtns., Cochise Co., AZ; Lab # 9382- 
9383, Bartel,1582A, B, Rucker Canyon, 6.1 mi from Leslie Canyon Rd 
along Rucker Canyon Rd., N 31 45' 18.3", W 109 22' 39.5", 1676m, 
Pedregosa Mtns., Cochise Co., AZ, 9384-9385, Bartel, 1583A,B, 
Metcalf, w of Chase Creek, 9.6 mi from lower Eagle Creek Rd, along 
US191, N 33 08' 01.5", W 109 22' 38.7", 1683m, Greenlee Co., AZ, 


Hesperocyparis glabra, 9386-9387, Bartel,1584A,B, upper 
Slate Creek, 7.1 mi sw if SR 188, along SR87, N 33 57' 28.5", W 111 
24' 21.8", 1009m, Mazatzal Mtns., Gila Co., AZ, 9388-9389, Bartel, 
1585A,B, se of Tonto Natural Bridge St. Park, jct along SR87, nw of 
East Verde River, N 34 18' 58.6", W 111 23' 12.6", 1483m, Gila Co., 
AZ, 9390-9391, Bartel, 1586A,B, upper Dry Beaver Creek, 0.1 mi. e of 
SR 179 along Wild Horse Mesa Rd., N 34 46' 07.7", W 111 45' 46.4", 
1225m, Yavapai Co., AZ. 


Adams' specimens are deposited at BAYL herbarium, Waco, 
Texas. Bartel's specimens currently are held in his personal herbarium, 
Carlsbad, California. 


One gram (fresh weight) of foliage was placed in 20 g of 
activated silica gel and transported to the lab, thence stored at -20 C 
until the DNA was extracted. DNA was extracted from the leaves by 
use of the Qiagen DNeasy Mini-plant extractors. Ten-mer primers 
were purchased from the University of British Columbia (5'-3'): 131, 
GAA ACA GCG T; 153, GAG TCA CGA G; 204, TTC GGG CCG T; 
Meat GCG TGA C; 218, Cie AGC ‘CCA -G; 239, CIG AAG 
CGG A; 244, CAG CGA ACC G; 250, CGA CAG TCC C; 327, ATA 
GGG" CGT C; 338 CTC TGG CGG Tf; 346, TAG GCG AAC 'G; 347; 
TTG CTT GGC G; 389 CGC CCG CAG T; 413, GAG GCG GCG A; 


PCR was performed in a volume of 15 ml containing 50 mM 
Tris-HCl (pH 9), 2.0 mM MgCl2, 0.01% gelatin and 0.1% Triton X- 
100, 0.2 mM of each dNTPs, 0.36 mM primers, 0.3 ng genomic DNA, 
15 ng BSA and 0.6 unit of Taq DNA polymerase (Promega). A control 


248 Phytologia (April 2009) 91(1) 


PCR tube containing all components, but no genomic DNA, was run 
with each primer to check for contamination. DNA amplification was 
performed in an MJ Programmable Thermal Cycler (MJ Research, 
Inc.). The thermal cycle was: 94 C (1.5 min) for initial strand 
separation, then 40 cycles of 38 C (2 min), 72 C (2 min), 91 C (1 min). 
Two additional steps were used: 38 C (2 min) and 72 C (5 min) for final 
extension. Bands that occurred once or did not show fidelity within the 
two replicated samples of each taxon were eliminated. It should be 
noted that these bands contain very useful information for the study of 
genetic variance and individual variation, but are merely "noise" in the 
present taxonomic study. Bands were scored in 4 classes: very bright 
(=6); medium bright (=5), faint (=4) and absent (=0). See Adams and 
Demeke (1993) for details on electrophoresis and RAPD band scoring. 


Similarity measures were computed using absolute character state 
differences (Manhattan metric), divided by the maximum observed 
value for that character over all taxa (= Gower metric, Gower 1971; 
Adams 1975). Principal coordinate analysis (PCO) of the similarity 
matrix follows Gower (1966). 


RESULTS AND DISCUSSION 


Contoured clustering based on 83 RAPD bands (Figure 2) 
shows that the populations cluster by geographical proximity. The 
most similar H. arizonica populations are Santa Catalina Mtns. - 
Dragoon Mtns. (0.969), followed by Dragoon Mtns. - Pedregosa Mtns. 
(0.943), then Pedregosa Mtns. - Greenlee Co. (0.932). The Chisos 
Mtns., TX population is clearly quite differentiated (linkage of 0.926 to 
the Dragoon Mtns. population). 


The most similar H. glabra populations are Mazatzal Mtns. - 
Tonto Basin (0.959), then Tonto Basin - Dry Beaver Creek (0.937). 
The H. arizonica - H. glabra populations are finally linked by Greenlee 
Co. - Mazatzal Mtns. (0.916). 


The linkage of populations of both taxa by geographically near 
neighbors suggests that differentiation is due to restricted gene 
exchange perhaps leading to genetic drift. Due to the likely short 
distances of cone/ seed dispersal, it seems probable that pollen dispersal 


Phytologia (April 2009) 91(1) 249 


over long distances may be the principal agent of gene flow among 
these populations. 


Contoured Clustering 
Flagstaff 
a" 83 RAPDs 
Sedona, 


Dry —» 
Beaver 


eee @ glabra 


% arizonica 


|° 
= 
le 
o 
j= 
> 
® 
= 


Figure 2. Contoured clustering of populations of H. arizonica and H. 
glabra based on 83 RAPDs bands. 


250 Phytologia (April 2009) 91(1) 


ACKNOWLEDGEMENTS 
This research supported in part with funds from U. S. Fish and 
Wildlife Service, Grant 814307J011. The findings and conclusions in 
this article are those of the authors and do not necessarily represent the 
views of the U.S. Fish and Wildlife Service. 


LITERATURE CITED 

Adams, R. P., 1975. Statistical character weighting and similarity 
stability. Brittonia 27: 305- 316. 

Adams, R. P., J. A. Bartel and R. A. Price. 2009. A new genus, 
Hesperocyparis, for the cypresses of the western hemisphere. 
Phytologia 91(1): 160-185. 

Askew, G. R. and R. E. Schoenike. 1982. Identification of characteristic 
traits of two varieties of Arizona cypress. Silvae Genetica 31: 158— 
160. 

Bartel, J. A. 1993. Cypress family-Cupressaceae. J. Ariz. Nev. Acad. 
Sei) 27x195-200; 

Bartel, J. A., R. P. Adams, S. A. James, L. E. Mumba and R. N. 
Pandey. 2003. Variation among Cupressus species from the 
western hemisphere based on Random Amplified Polymorphic 
DNAs (RAPDs). Biochem. Syst. Ecol. 31: 693-702. 

Eckenwalder, J. E., 1993. Cupressus In: Morin, N.R. (Ed.), Flora of 
North America. Pteridophytes and Gymnosperms, vol. 2. Oxford 
University Press, New York, pp. 405-408. 

Farjon, A., 1998. World checklist and bibliography of conifers. Royal 
Bot. Gard., Kew, London. 

Gower, J. C., 1966. Some distance properties of latent root and vector 
methods used in multivariate analysis. Biometrika 53: 326-338. 
Gower, J. C., 1971. A general coefficient of similarity and some of its 

properties. Biometrics 27: 857-874. 

Little, E. L., 1970. Names of New World cypresses (Cupressus) 
Phytologia 20: 429-445. 

Little, D. P. 2006. Evolution and circumscription of the true Cypresses 
(Cupressaceae: Cupressus) Syst. Bot. 31: 461-480. 

Little, D. P., A. E. Schwarzbach, R. P. Adams and C-F. Hsieh. 2004. 
The circumscription and phylogenetic relationships of Callitropsis 
and the newly described genus Xanthocyparis (Cupressaceae). 

Am. J. Bot. 91: 1872-1881. 
Wolf, C. B., 1948. The New World cypresses. Aliso 1: 1-250. 


Phytologia (August 2009) 91(2) 251 


SOLIDAGO DISPERSA (ASTERACEAE: ASTEREAE) 
REPLACES SOLIDAGO LUDOVICIANA 
AS THE CORRECT NAME 


Guy L. Nesom 
2925 Hartwood Drive 
Fort Worth, TX 76109, USA 
www.guynesom.com 


ABSTRACT 


Solidago dispersa Small is the correct name for the species 
previously identified as S. /udoviciana (A. Gray) Small. The species is 
restricted to east Texas, western Louisiana, southern Arkansas, and the 
southeastern corner of Oklahoma. A distribution map is provided for S. 
dispersa and taxonomic with nomenclatural summaries for S. dispersa 
and its synonyms, S. /udoviciana and S. strigosa Small. Phytologia 
91(2): 251-255 (August, 2009). 


KEY WORDS: Solidago ludoviciana, S. dispersa, Asteraceae. 


The correct name for the species generally identified as 
Solidago ludoviciana (A. Gray) Small is S. dispersa Small, as 
interpreted here. The distribution of S. dispersa is restricted to east 
Texas, western Louisiana, southern Arkansas, and the southeastern 
corner of Oklahoma (Fig. 1). In Oklahoma it is known from only two 
collections. McCurtain Co.: disturbed roadside with windrows of 
bulldozed trees, 2 mi S and 0.4 mi W of Tom along Hwy 87, 18 Sep 
1976, Taylor & Taylor 23641 (BRIT, OKL); sloughy bank of Cedar 
Creek, 10 mi N of Broken Bow, 16 Oct 1937, Hopkins & Cross 2496 
(OKL). 


In resolving the nomenclature, observations on_ the 
morphology of Solidago dispersa are critical. Compared to S. arguta 
var. boottii, with which it is sometimes confused, the slender, scale- 
leaved, stoloniform rhizomes of S. dispersa are diagnostic. The two are 
sympatric in an area of northcentral Louisiana and adjacent Arkansas. 
The map in Fig. | is unambiguous and the coherence of the geographic 


252 Phytologia (August 2009) 91(2) 


distribution supports the recognition of S. dispersa. The leaves of S. 
dispersa are thicker with denser reticulation (smaller interstices) of 
tertiary venation, and distal cauline leaves are erect-ascending and 
basally attenuate (vs. spreading and usually with an abruptly delimited 
petiole-like base in var. boottii). Plants of S. dispersa in Texas (except 
for some counties bordering Louisiana) are completely glabrous below 
the capitulescence. In Louisiana and Arkansas scattered plants are 
similarly glabrous but more commonly the lamina of basal and 
proximal cauline leaves are abaxially strigose-hirsute to hirsute; 
sometimes both faces have hairy lamina. The proximal and mid-region 
stems of such hairy-leaved plants commonly are sparsely to moderately 
hirsute. Plants of S. arguta var. boottii very rarely have leaves with 
lamina hairy abaxially (never adaxially) and none has been observed 
with hairy stems. Morton (1973) also recognized that S. dispersa 1s 
variable in vestiture. 


As noted by Semple (2006, p. 132), "G.H. Morton annotated 
the type of [Solidago dispersal] as possibly being S. arguta introgressed 
with S. ul/mifolia. The application of the name remains uncertain." The 
type collection of S. dispersa, however, was made in western Louisiana 
(Sabine Parish, on the Texas border), outside of the range of S. arguta 
but within that of S. /udoviciana and in an area where the latter is 
common. 


The roots and lower stem of the type of Solidago dispersa 
were not collected and (fide Small’s protologue, p. 476) "The 
inflorescence is paniculate and very loosely disposed [hence, 
apparently, the epithet], while the branches of the panicle and the 
elongated peduncles are filiform or nearly so and quite weak." Small 
also noted that "It is peculiar in being glabrous or nearly so above and 
with more or less pubescence on the lower leaves." The vestiture is 
similar to that of many plants from Louisiana previously identified as S. 
ludoviciana. The inflorescence is uncharacteristic, but plants of a few 
other collections otherwise typical in morphology for S. /udoviciana 
have elongated peduncles and heads in a ‘loosely disposed’ 
inflorescence, appearing more diffusely paniculate than — short- 
pedunculate and strongly secund on spreading branches (e.g., Texas: 
Wood Co., Lundell & Lundell 11741 (SMU); Bowie Co., Correll & 
Correll 24789 (LL, SMU). These variants are very similar to the type 


Phytologia (August 2009) 91(2) 253 


of S. dispersa and the type is thus within both the geographic and 
morphological bounds of the species. There is no apparent evidence 
that Solidago dispersa hybridizes with S. ulmifolia, S. arguta, or any 
other species. Solidago dispersa was observed by Cronquist (1980, as 
S. ludoviciana) to be closely similar to S. arguta and S. tarda. Fernald 
(1936) used the name S. /udoviciana to identify plants now known as S. 
tarda on the Atlantic coastal plain. Taylor and Taylor (1984) identified 
S. dispersa in Texas and Louisiana as S. salicina Ell., which in their 
concept also included S. patula. 


Solidago dispersa Small, Bull. Torrey Bot. Club 25: 475. 1898. 
TYPE: U.S.A. Louisiana. No other data, but probably Sabine 
Parish, 1836-1838, Leavenworth s.n. (holotype: NY, internet 
image). Melines Conkling Leavenworth, an army physician, was 
stationed in Sabine Parish, Louisiana, in an active phase of his 
botanical work (1836-1838), during which he corresponded with 
John Torrey (ca. 25 letters) and sent him many specimens 
(McVaugh 1947). The type of S. dispersa probably was among 
these collections. Torrey and Gray (1842) cited collections by 
Leavenworth (and by Josiah Hale, see comments in typification of 
S. strigosa and S. ludoviciana) from Louisiana under "var. B" and 
"var. &" of S. boottii (see comments under S. strigosa). 


Solidago ludoviciana (A. Gray) Small, Fl. S.E. US. 1198, 1339. 1903. 
Solidago boottii var. ludoviciana A. Gray, Proc. Amer. Acad. Arts 
1721952 1882) LECTOTYPE GK. Small ex M:L. Fernald 1936, p. 
210): U.S.A. Louisiana. No other data, J. Hale s.n. (NY 00259711, 
internet image!). No indication of a type was given in the 
protologue but Gray (1884, p. 154) noted "W. Louisiana." Most of 
Hale's Louisiana collections, which began in 1838, were made 
from the vicinity of Alexandria (Rapides Parish) and sent to Torrey 
and Gray (Ewan 2005). "Hale's plants were not numbered, and so 
after Charles Short and others had divided the original specimens 
and exchanged a portion, the origin (‘Louisiana’) was often all that 
accompanied the specimen" (Ewan 2005, p. 2282). Comments 
following Solidago strigosa, a name closely associated with S. 
ludoviciana, explain the lectotypification. 

Solidago strigosa Small, Fl. S.E. US. 1198, 1339. 1903. Solidago 
arguta var. strigosa (Small) Steyerm., Rhodora 62:131. 1960. 


254 


Phytologia (August 2009) 91(2) 


TYPE: U.S.A. Louisiana. No other data, J. Hale s.n. (holotype: 
NY 00259966, internet image; isotype: GH 12437). Three Hale 
specimens of Solidago aff. arguta are housed at GH and NY: GH 
12437; NY ...711 (identified on the original label as "S. boottii ¢?") 
and NY ...966 (originally identified as "S. boottii between B and 
é"), each with an original "Torr. & Gray, Flora N. Amer." label. 
Gray studied the collections at both herbaria; he observed that NY 
...7/11 was different from the GH collection, noting at the bottom of 
NY ...711 "My specimen of this is hirsute" and on the GH sheet 
"The specimen in Hb. Torr. of var. €? is glabrous." Gray (1884, p. 
154) included both specimens in his concept of S. /udoviciana, 
describing the species as having "lower leaves and lower part of 
the stems sometimes roughish-hirsute or hispidulous with many- 
jointed hairs, or glabrous" [emphasis added]. 


Small apparently decided that NY ...711 was representative of Solidago 


ludoviciana and annotated it as such. Seeing that NY ...966, was 
different from ...711, he described ...966 as S. strigosa. Small's 
morphological and geographical descriptions of the two taxa 
(1903, 1933) support this interpretation. Fernald (1936) provided 
an account similar to the one here of the three Hale specimens; he 
also noted that "Small having selected the glabrous plant of Hale to 
stand as the type of S. /udoviciana, that point is satisfactorily 
settled, the hirsute plant of Hale being S. strigosa Small." 
Fernald's affirmation of Small's choice is taken here as 
formalization of the lectotypification of S. /udoviciana. Morton 
(1973) also concluded that the types of S. strigosa and S. 
ludoviciana are conspecific, despite the difference in vestiture. 


ACKNOWLEDGEMENTS 
Collections were studied on site at BRIT/SMU, MO, NLU, 


TEX/LL, and VDB. Loans of specimens from MISS and OKL were 
valuable in interpreting patterns of variation and distribution. 


LITERATURE CITED 
Cronquist, A. 1980. Flora of the Southeastern United States. Vol. 1. 
Asteraceae. Univ. of North Carolina Press, Chapel Hill. 
Ewan, J.A. 2005. Notes on Louisiana botany and botanists, 1718— 
1975. Sida 21: 2275-2296. 


Phytologia (August 2009) 91(2) 255 


Fernald, M.L. 1936. VI. Studies in Solidago. Rhodora 38: 201—229. 

Gray, A. 1884. Synoptical flora of North America. Vol. I, pt. IL. 
Ivison, Blakeman, Taylor, and Co, New York. 

McVaugh, R. 1947. The travels and botanical collecting of Dr. 
Melines Conkling Leavenworth. Field & Lab. 15: 57—70. 

Morton, G.H. 1973. The taxonomy of the Solidago arguta-boottii 
complex. Ph.D. thesis, Univ. of Tennessee, Knoxville. 

Semple, J.C. 2006. Solidago. Pp. 107—166, in Flora North America 
Editorial Committee (eds.). Flora of North America. Vol. 20. 
Asteraceae, Part 2. Astereae and Senecioneae. Oxford University 
Press, New York. 

Taylor, C.E.S. and R.J. Taylor. 1984. Solidago (Asteraceae) in 
Oklahoma and Texas. Sida 10: 223-251 

Torrey, J. and A. Gray. 1842. A Flora of North America. Vol. II, Part 
2. Wiley & Putnam, New York. 


Figure. 1. Distribution of Solidago dispersa. Sabine Parish, Louisiana, 
where the type was collected, is outlined in bold. 


256 Phytologia (August 2009) 91(2) 


RECENSION OF THE MEXICAN SPECIES OF SALVIA 
(LAMIACEAE), SECTION SCORODONIA 


B. L. Turner 
Plant Resources Center 
The University of Texas at Austin 
Austin, Texas 78712 
billie@uts.cc.utexas.edu 


ABSTRACT 


A recension of the Mexican species of Sa/via belonging to the 
sect. Scorodonia is rendered. Fourteen species are recognized, two of 
these described as new: S. pericona, a white-flowered taxon, from 
Oaxaca, and S. tenorioi, a yellow-flowed taxon from Puebla. A key to 
the various taxa is provided as well as photographs of the holotypes. 
Maps showing distribution of the taxa are provided. Phytologia 91(2): 
256-269 (August, 2009). 


KEY WORDS: Salvia, Lamiaceae, sect. Scorodonia, Mexico 


43. Sect. Scorodonia 
Sect. Atratae Epling 


Perennial herbs, shrubs or subshrubs, mostly 1-5 m high. Leaves ovate 
to deltoid, rarely cordate; petioles mostly short with a well-defined 
abscission line near the base, rarely not; blades often bullate, variously 
pubescent, the margins crenate. Spikes various, usually terminal, 
interrupted or not, rarely spicate-paniculate. Floral bracts lanceolate to 
broadly ovate, early deciduous. Calyx with upper lip 5-7 veined. 
Corollas mostly purple to blue, sometimes white, rarely yellow. 
Stamens arising from near corolla throat, the anthers not exserted. 
Style branches pilose, the upper lobe 2-3 times as long as lower. 
Nutlets glabrous. 

Type species: S. melissodora Lag. 


Epling (1939) notes that sect. Scorodonia is “A group of 
closely related forms difficult to distinguish save on the summation of 


Phytologia (August 2009) 91(2) 257 


minutiae in habit, yet the range both in structure and in distribution is 
such as to support the [treatment rendered].” So introduced, he 
recognized 11 species as occurring in Mexico. Ramamoorthy (1984) 
added to the assemblage S. boegeri; this, along with the two species 
described below bring to 14 the number of Mexican taxa currently 
recognized in the section. 


Epling (1941) subsequently combined sect. Urica with sect. 
Scorodonia, but I have opted to retain the two, mainly because of the 
lack of a clearly defined absission line at the base of the petiole in 
species of the sect. Urica. 


Key to Mexican species 


eeuas yellow? Hid: ..2.222:45:.452. 00 RAPA a S. tenorioi 
1. Corollas not yellow...(2) 


2. Corollas variously blue to purple...(4) 
2. Corollas white or creamy white.. .(3) 


seecholes 2-3.cm: long; Sierra Pericon, Oax......:2. 2.2.0.1... S. pericona 
aeons 0:5-1.0:cm-long; Pue; Oax:...c2n esi nean S. ramosa 
4(2). Upper stems pubescent with branched hairs .........S. melissodora 


4. Upper stems variously pubescent but w/o branched hairs (5) 


eee orolla tubes: 3=Samim lone... 2.26226... sdsc0sxseoreacens S. ramosa 
a Corolla tubes 5-13 mm long...(6) 


6. Corolla tubes 7-13 mm long...(13) 
6. Corolla tubes 5-7 mm long, or leaves 1.0-1.5 times as long as 
wide...(7) 


7. Calyx densely glandular-pubescent....................5 S. melissodora 
7. Calyx not glandular-pubescent.. .(8) 


oe -Wareer leaves 12-15 cm long; Oax..... 2.2.2.0. ey S. occidua 
8. Larger leaves 1-5 cm long...(9) 


258 Phytologia (August 2009) 91(2) 


9. Petioles 5-10 mm long; blades 3-5 cm long...(11) 
9. Petioles 1-5 mm long; blades 1-2 cm long; Pue...(10) 


LO: Calyx elandular-pubeseemt, 2 .:A0ij20e) «oss veka tees se S. paupercula 
10, Calyx densely, white-woolly:ajiiscot,.wih he .aooled «sacs S. boegei 
11(9). Leaves 1-2 times as long as wide; Nue, Tam to Hid.....S. keerlii 
lt... Leaves 2-3 times asilongyas wide; Mex.to Gue::....22- (12) 
12. Calyx densely ivory-white pubescent..................... S. breviflora 
12) Calyx not as described im the above... ......:ce0d50-52.52 S. melissodora 
13(6). Corolla tubes white, 10-13 mm long.................. S. semiatrata 
[340 -Coroella:tubes'purple; 7-9 mmilone..:2)0).4tageo eee (14) 
4: Foliage: eglandular-pubescent..... 0.05.2 06004).5.00 eds. (17) 
14. Foliage slandular-pubescemts, 4.42. 0200. 0c 2 ee es (15) 
15. Calyx 5-6 mm long; corolla tube 7-9 mm long............ S. rupicola 
15. Calyx 6-7 mm long; corolla tube 6-7 mm long..................66 (16) 
1G, Blamisioi Nayarit. 2... so. : baat caves teats jaueade sdeee age S. tepecensis 
LG, Jelantsi@t Simaloals :.).5.es0 cu. sae seoe mee Basta tide eee S. aequidistans 
17@4). Calyx minutely glandular; Pure. «12 acneieeceesese sceees S. pannosa 

li Calyx white-villous; Nue, Tam, Dur, San, Hid......... S. keerlii 


SALVIA AEQUIDISTANS Fernald, Proc. Amer. Acad. Arts 35: 512. 
1900. Map 1 


S Sin, between Rosario and Colomas (type material) and San 
Ignacio, ca 400 m; Jul. 

As indicated by its author, the species ““Scarcely differs from 
S. tepicensis save in the more lax habit, longer petioles, less dense 
pubescence and thinner leaves.” 


Phytologia (August 2009) 91(2) 259 


SALVIA BOEGEI Ramamoorthy, J. Arnold Arb. 65:138. 1984. 
Map 2 


Known only from Puente de Dios Molcaxac, Pue, 1800 m; 
Sep. 

According to its creator, the species “can be recognized 
immediately by its articulated petiole, almost capitate verticels [of 
flowers], and woolly white calyx.” The relatively small leaves with 
short petioles also distinguish the species, the latter characters 
suggesting a close relationship with S. paupercula, which is readily 
distinguished from S. boegei by its glandular vestiture. 


SALVIA BREVIFLORA Moc. & Sesse ex Benth., Lab. Gen. Sp. 274. 
1833. Map 2 

Salvia albicans Fernald 

Salvia nelsonii Fernald 


Mic, Mex, Mor, Pue and Gue, calcareous hillsides with 
juniper, 1000-1500 m; Aug-Oct. 


This is an attractive blue-flowered shrub 1-3 m high having 
white-woolly compacted spikes 
My synonymy follows that of Epling (1939). 


SALVIA KEERLII Benth., Lab. Gen. Sp. 263. 1833. Map 3 


Nue, Tam, San, Gua, Que?, Mic, Mex and Hid, pine-oak 
forests, 2400-3000 m, Aug-Oct. 


Salvia keerlii in northeastern Mexico is a relatively common 
blue-flowered shrub 1-2 m high; I have not seen specimens from the 
state of Durango, as reported by Epling (1939). 


SALVIA MELISSODORA Lag., Gen. et Sp. 2. 1816. Map 4 
Salvia dugesii Fernald 

Salvia scorodoniifolia Poir. 

Salvia scordoniifolia var. crenaea Fernald 


260 Phytologia (August 2009) 91(2) 


A widespread highly variable species distributed throughout 
most of central Mexico in relatively dry bushy habitats, 500-2500 m; 
Aug-Mar. 

As well noted by Epling, this taxon has two pubescent forms: 
plants with glandular hairs, and those with branched hairs. The name S. 
dugesii has been applied to the latter. 


SALVIA OCCIDUA Epling, Repert. Spec. Nov. Regni Veg. Beih. 
110: 173. 1939. Map 2 


Oax, coastal areas near San Miguel del Puerto, known to me 
only from Liebmann collections cited by Epling. 

A poorly collected taxon, readily distinguished from closely 
related taxa by its large foliage. 


SALVIA PANNOSA Fernald, Proc. Amer. Acad. Arts 40: 54. 1905. 
Map 3 


Southern Pue and closely adjacent Oax, xeric shrub lands with 
Juniperus, 1200-3000 m; Jul-Oct. 

Said to be a locally abundant purple-flowered subshrub 1-2 m 
high, relatively distinctive because of its bicolored leaves, the blades of 
which are rather lanceolate and truncate to subcordate at the base. 


SALVIA PAUPERCULA Epling, Repert. Sp. Nov. Regni Veg., Beih. 
110: 173. Map 4 


This is a poorly known taxon, reportedly from near Fort de 
Guadalupe and Rancho Posada, Pue, the type (US) collected by F. 
Arsene in 1909. 

Except for its vestiture this taxon appears closely related to S. 
boegeri, so noted under the latter. 


SALVIA PERICONA B.L. Turner, sp. nov. Fig. 1 Map 5 
Salvia ramosa Brandegee similes sed differt foliis multo majoribus 


valde bicoloribus ac plerumque cordatis, petioles longioibus (2-3 cm 
longis vs 0.5-1.0 cm), et corollis majoribus albis (vs. azureis). 


Phytologia (August 2009) 91(2) 261 


Perennial herbs or subshrubs to at least 0.5 m high. Stems purplish, 
pubescent with peculiar erect scattered 2-3 celled trichomes 0.2-0.3 
mm high, beneath these an understory of minute glandular hairs. 
Leaves (the larger) 5-8 cm long, 2-5 cm wide; petioles 2-3 cm long, 
having a distinct disarticulation scar at the base; blades deltoid to 
cordate, bicolored, the lower surface densely white-tomentose, the 
upper dark green and rugose, beset with small hairs throughout, the 
margins irregularly serrulate. Spikes terminal, 5-15 cm long, the 
flowers arranged 4-10 at interrupted nodes. Floral bracts (uppermost) 
broadly ovate, 6-8 mm long, 4-5 mm wide, soon deciduous. Flowering 
calyces 8-9 mm long, pubescent like the stems, the upper lip ca 3 mm 
long; pedicels 2-4 mm long. Corollas white, smooth within; tube 8-9 
mm long; upper lip ca 4 mm long; lower lip 4-5 mm long, reflexed. 
Anthers not excurrent; filaments ca 3 mm long; anthers ca 1.5 mm long. 
Style pilose, the upper branch recurved, 2-3 times as long as the lower. 


TYPE: MEXICO. OAXACA: Mpio. Tamazulapan, Cerro Pericon, 5 
km al N de San Pedro Nopala, “Suelo obscuro derivido de roca ignea.” 
2000 m, 6 Jul 1986, Abisai Garcia M. 2342 (with D. Frame, P. Tenorio 
and A. Salinas) (holotype: TEX). 


ADDITIONAL SPECIMEN EXAMINED: Same locality as Type, 
“Bosque de Encino con elementos de Matorral xerofilo, 2350 m, 13 
Nov 1985, Ramamoorthy 4778 (TEX). 


The label of Ramamoorthy 4778 describes the plant as a 
white-flowered herb 0.5 m high; the species name is taken from the 
Sierra to which it is possibly confined. Ramamoorthy apparently also 
recognized the taxon as new when collected, to judge from his 
annotation on the sheet concerned. 


SALVIA RAMOSA Brandegee, Zoe 5: 255. 1908. Map 4 
Salvia lantanifolia Mart. & Gal. ? 
Salvia variana Epling 


Southern Pue and closely adjacent Pue in dry shrublands with 
Juniperus, 1800-3000 m; Sep-Nov. 

As pointed out in detail by Epling (1939), S. ramosa is a 
highly variable taxon, especially in pubescence, possessing calyces 


262 Phytologia (August 2009) 91(2) 


with only branched hairs, or else pubescent throughout with 
multiseptate glandular trichomes. He also noted that S. ramosa was 
“Scarcely to be distinguished from S. mellissodora save in the smaller 
leaves, smaller flowers and finer pubescence.” I am unable to 
distinguish Epling’s S. variana, the latter reportedly having somewhat 
larger corollas and longer petioles than typical S. ramosa. Finally, it 
should be noted that the earlier name S. /antanifolia Mart. & Gal. might 
be better tagged upon the present, since its distribution and general 
habit, as judged by phototypes at TEX appear very similar to S. ramosa. 
Indeed, Brandegee assigned the name S. lantanifolia to the type of S. 
variana. 


SALVIA RUPICOLA Fernald, Proc. Amer. Acad. Arts 45: 420. 1910. 
Map 1 


Hid, vicinity of Ixmiquilpan, rocky hillsides; ca 2500 m; Jul- 
Aug. 

Epling (1939) thought this taxon perhaps but a variety of his 
concept of S. scordoniifolia, but subsequently retained the species 
(Epling 1944). 


SALVIA SEMIATRATA Zucc. in Abhandl. Akad. Muench. 1. 298. 
1832. Map 5 


Oax, where relatively common in pine-oak forests, 200-2500 
m; Jun-Oct. 

A very distinctive blue-flowered shrub 1-2 m high having 
mostly cordate, markedly rugose, leaves and relatively large flowers. 

Epling (1939) treated this species as the sole member of his 
sect. Atratae. | think it better positioned within the sect. Scorodonia. 


SALVIA TENORIOI Ramamoorthy ex B.L. Turner, sp. nov. Fig. 2 
Map 5 


Salvia ramosa Brandegee similes sed differt corollas flavis (vs azureis 
vel purpuratis), floribus in paniculis spicatis dispositis, et vestimento 
calycini trichomatum glandulosorum (vs trichomatum ramosorum). 


Phytologia (August 2009) 91(2) 263 


Shrubs 2-3 m high. Stems mostly pubescent with white, recurved 
hairs ca 0.3 mm high. Leaves (newly produced among the upper 
nodes) ovate to deltoid, markedly rugose, their margins crenulate; 
petioles 1-2 mm long. Flowers arranged in paniculate interrupted 
spikes, the latter mostly 4-6 cm long, the terminal panicle ca 30 cm 
high, 25 cm wide. Floral bracts (uppermost) lanceolate, 2-4 mm long, 
glandular-pubescent, soon deciduous. Flowering calyces 6-7 mm long, 
glandular-pubescent with viscid hairs ca 0.5 mm high; upper lip ca 1.5 
mm long, 5-nerved; lower lip ca 1 mm long. Corollas yellow; tube 
more or less straight, 7-8 mm long, papillose and/or rugose within; 
upper lip ca 3 mm long; lower lip flabellate, reflexed, 3-4 mm long. 
Anthers not excurrent, attached near the throat of the tube; filaments 3- 
4 mm long, markedly flattened and recurved or twisting at maturity. 
Style sparingly pilose, more so below, the upper branch 2-3 times as 
long as the lower. Nutlets ovoid, glabrous, ca 2.5 mm long, 1.5 mm 
wide. 


TYPE: MEXICO. PUEBLA: Mpio. Teontepec, “14 km al NW de 
Teontepec, brecha a Nopala...Matorral calcicola mixta...Suelo negro 
sobre roca caliza.” 16 Nov 1985, P. Tenorio L. & G. Dieringer 10648 
(holotype TEX). 


According to label data the flowers are yellow, and the plants 
are said to be abundant shrubs 2-3 m high. The species is named for its 
collector, Pedro Tenorio, this suggested on the type itself by 
Ramamoorthy soon after its collection. According to Dr. Fernando 
Chiang, Pedro is a diligent collector and photographer of the Mexican 
flora who formerly worked at MEXU. He assembled over 20,000 
numbers from throughout Mexico, and is well known for his collections 
from the area of Caltepec, Puebla where he was born and raised. 


Yellow-flowered Salvias are quite rare in Mexico, as noted by 
Ramamoorthy (!984). In his description of the yellow-flowered S. 
tuxtlensis he stated, “Of the estimated 275 species in Mexico only three 
have yellow flowers.” Actually, including S. tuxtlensis and the present 
novelty, five yellow-flowered species are known, Ramamoorthy having 
been unaware of the lovely S. madrensis of Sinaloa. Epling (1939) 
placed the yellow-flowered species known to him (S. aspera Kunth, S. 
madrensis, S. subhastata, and S. hidalgensis Mir.) in four Sections. 


264 Phytologia (August 2009) 91(2) 


Ramamoorthy did not assign his novelty to a Section but allowed as to 
how it might belong to yet another monotypic Section. By implication, 
in my Latin diagnosis I have tentatively assigned S. tenorioi to the sect. 
Scordonia, the plant concerned having the general habit and vegetative 
features of that assemblage. 


SALVIA TEPICENSIS Fernald, Proc. Amer. Acad. Arts 45: 420. 
1910. Map 1 
Salvia scordoniifolia var. subsessilis Benth. 


Nay and Col, mixed mesophytic forests along the Pacific 
coast, 300-900 m, Jun-Aug. 

This species is doubtfully placed in the sect. Scordonia, 
although maintained by Epling (1939); at least it appears to lack the 
distinctive disarticulation scar at the base of petioles found in most of 
the other taxa of the complex. Regardless, Epling positioned S. 
tepicencis in sect. Scorodonia, along with S. aequidistans, the two 
scarcely distinguishable 


ACKNOWLEDGEMENTS 


I am grateful to Guy Nesom for the Latin diagnosis, and for 
helpful comments following his review of the article. 


LITERATURE CITED 

Epling, C. 1939. A revision of Salvia; Subgenus Calosphace. Repert. 
Sp. Nov. Regni Veg. Beth. 110: 1-380. 

Epling, C. 1941. Supplementary notes on American Labiatae-11. Bull. 
Torrey Bot. Club 65: 552-568. 

Ramamoorthy, T.P. 1984. A new species of Salvia (Lamiaceae) from 
the Sierra de Los Tuxtlas, Mexico. Pl. Syst. Evol. 146: 141- 
143. 


Phytologia (August 2009) 91(2) 


SALVIA 
© aequidistans 
@ tepicensis 
* rupicola 


Map 1 


SALVIA 
e breviflora 


* occidua 
© tenorioil 


Map 2 


265 


266 Phytologia (August 2009) 91 (2) 


SALVIA 
@ keerlii 


® pannosa 


SALVIA 
@ Melissodora 


© ramosa 
%*% paupercula 


Map 4 


Phytologia (August 2009) 91(2) 267 


SALVIA 
@ semiatrata 


© pericona 
*& tenoriol 


thc Wagtee M41 Ge 
Map 5 <}: ae Bs 


268 Phytologia (August 2009) 91(2) 


HERGQARIUM 


tina 


Fig. 1. Holotype of Salvia pericona. 


Phytologia (August 2009) 91(2) 269 


UNIVERSITY OF 


amy, ERAS 


HERBARIUM 


‘Plantes de Miuex pyrma 


Herbaris Nacional (MEO) 


a! MW de Teontepec, Dreche @ Nopale 


coet. 


Fig. 2. Holotype of Salvia tenorioi. 


270 Phytologia (August 2009) 91(2) 


KEYS TO THE FLORA OF FLORIDA: 22, 
DICERANDRA (LABIATAE) 


Daniel B. Ward 
Department of Botany, University of Florida 
Gainesville, Florida 32611, U.S.A. 


ABSTRACT 


Dicerandra (Labiatae) is represented in Florida by three 
species: one, D. densiflora, is monotypic; one, D. linearifolia, consists 
of two varieties; and one, D. frutescens, is believed best interpreted to 
be formed of 7 varieties. Dicerandra frutescens var. christmanii, var. 
cornutissima, var. immaculata, var. modesta, var. savannarum, and 
var. thinicola are recognized as new combinations. Dicerandra 
densiflora and D. frutescens are endemic to Florida. Dicerandra 
Jrutescens (typical) and certain of its varieties are rated as endangered. 
An amplified key is given to the Florida taxa. Phytologia 91(2): 270- 
276 (August, 2009). 


KEY WORDS: Dicerandra, Labiatae, Florida flora. 


The task of the present number is to formalize the change in 
rank of certain taxa within the woody mint, Dicerandra (Labiatae), as 
needed for uniform presentation within an amplified key to the Florida 
species. 


Soon after arriving in Florida in 1958, the present author 
discovered in the herbarium (FLAS) specimens that seemed not to 
belong where they had been filed. Many had been collected by James 
B. McFarlin (1901-1969), from near Sebring, Highlands Co., and 
others from elsewhere in the peninsula. Trips to collection sites 
quickly found additional populations, and a small folder was assembled 
of notes and information that might eventually become meaningful. 
But in 1961 a request came from Lloyd H. Shinners (SMU), for the 


Phytologia (August 2009) 91(2) 271 


loan of the genus Dicerandra. With the advice of Erdman West, a 
selection of materials was made and dispatched to Texas. 


The result was Shinners' publication (Sida 1: 89-91, 1962) ofa 
synopsis of the genus, with the unnamed materials now named 
Dicerandra frutescens. Their distribution was from throughout the 
peninsula and showed no variances sufficient to bring attention. 


But within the year Olga Lakela, then in Tampa (USF), found 
plants near the eastern shore of the peninsula which lacked dots on the 
lip of the corolla; she named it Dicerandra immaculata (Sida 1: 184- 
185, 1963). Lakela was followed by Robin Huck who ranged widely, 
throughout Florida, Georgia, and the Carolinas; she found further 
variation in north-central Florida which she named D. cornutissima 
(Phytologia 47: 313-316, 1981). A student of the peninsular scrub 
flora, Steven Christman, then encountered a different form in 
Highlands County, and a team of 6 investigators pooled their resources 
to name it D. christmanii (Huck et al., Syst. Bot. 14: 197-213, 1989). 
A bryologist, Harvey Miller, with close contacts among vascular 
systematists, was inspired to find and describe another species, D. 
thinicola (Phytologia 75: 185-189, 1993). A degree of order and 
understanding was brought to this burgeoning profusion of new entities 
by Huck & Chambers (Edinb. J. Bot. 54: 217-229, 1997) who found 
different ploidy levels and distinct though often overlapping ranges for 
all described taxa. Loose ends were tidied up by Huck (Novon 11: 
417-420, 2001), in describing 2 infraspecific entities, D. frutescens ssp. 
modesta, and D. immaculata var. savannarum. A comprehensive 
summary of the entire nine-species genus, utilizing total genomic DNA 
and permitting the drawing of phylogenetic conclusions, was then 
assembled by L. O. Oliveira, R. B. Huck, M. A. Gitzendanner, W. S. 
Judd, D. E. Soltis and P. S. Soltis (Amer. J. Bot. 94: 1017-1027, 2007). 


This body of recent literature is far too extensive for 
summation here. Clearly, past geologic changes have served to isolate 
small populations of the woody mint that Shinners called D. frutescens, 
permitting alteration in gene frequency and small changes in 
morphology that astute observers have now been able to detect. 


272 Phytologia (August 2009) 91(2) 


Perhaps in time there will come an understanding of sea-level and 
climate changes that have influenced not only these taxa but other 
members of the complex Florida flora. 


But from the standpoint of a conventional taxonomist, either 
lacking knowledge of source or denying himself the unearned luxury of 
determining a species based almost entirely on where it was found, the 
described morphological differences are insufficient. Other species in 
related genera are not so finely dissected, and it is inappropriate that 
these taxa bear names well beyond the usual meaning of their 
morphological basis. In essence, Lloyd Shinners was right; the woody 
mints of this group are a single species, Dicerandra frutescens. 


Dicerandra frutescens L. H. Shinners var. christmanii (R. B. 
Huck & W. S. Judd) D. B. Ward, comb. et stat. nov. 
Basionym: Dicerandra christmanii R. B. Huck & W. S. Judd, 
in R. B. Huck, W. S. Judd, W. H. Whitten, J. D. Skean, R. P. 
Wunderlin, & K. R. Delaney, Syst. Bot. 14: 198. 1989. 
TYPE: United States, Florida, Highlands Co., Sebring, 10 Sept 
1987, Hansen & DeLaney 4825 (holotype, FLAS; isotypes, A, 
BHO, DUKE, F, FLAS, FTG; GA, K, MO, MSC, NEU, NY; 
SMU, TEX, UC, US, USF). 


Dicerandra frutescens L. H. Shinners var. cornutissima (R. 
B. Huck) D. B. Ward, comb. et stat. nov. Basionym: 
Dicerandra cornutissima R. B. Huck, Phytologia 47: 313. 
1981. TYPE: United States, Florida, Marion Co., Fla. 484 & 
I-75, 19 Sept 1980, Huck 2436 (holotype, NCU). 


Dicerandra frutescens L. H. Shinners var. immaculata (O. 
Lakela) D. B. Ward, comb. et stat. nov. Basionym: 
Dicerandra immaculata O. Lakela, Sida 1: 184. 1963. TYPE: 
United States, Florida, Indian River Co., U.S. 1, 30 Sept 1962, 
Lakela 25440 (holotype, USF). 


Dicerandra frutescens L. H. Shinners var. modesta (R. B. 
Huck) D. B. Ward, comb. et stat. nov. Basionym: Dicerandra 


Phytologia (August 2009) 91(2) 213 


frutescens L. H. Shinners ssp. modesta R. B. Huck, Novon 11: 
417. 2001. TYPE: United States, Florida, Polk Co., Dundee, 9 
Sept 1999, Huck 5555 (holotype, FLAS; isotypes, MO, USF). 


Dicerandra frutescens L. H. Shinners var. savannarum (R. B. 
Huck) D. B. Ward, comb. nov. Basionym: Dicerandra 
immaculata O. Lakela var. savannarum R. B. Huck, Novon 
11: 419. 2001. TYPE: United States, Florida, St. Lucie Co., 
Savannas State Preserve, 26 Oct 1996, Huck 5492 (holotype, 
FLAS; isotype, MO). 


Dicerandra frutescens L. Shinners var. thinicola (H. A. 
Miller) D. B. Ward, comb. et stat. nov. Basionym: 
Dicerandra thinicola H. A. Miller, Phytologia 75: 185. 1993. 
TYPE: United States, Florida, Brevard Co., Brandt Road, 5 
Nov 1987, Reifler s.n. (holotype, MU; isotypes, USF, FTU). 


The present paper, intended to address a subset of plant names 
which bear names at a level above that justified by their morphological 
differences, was initially limited to concern of the approach to take 
with Dicerandra, where a cottage industry has arisen to identify and 
name further variants of this heretofore unimportant woody mint. 
Every one of the many authors is (or has been -- Olga Lakela, 1890- 
1980; Lloyd Shinners, 1918-1971) a close associate or friend of the 
present author. One trusts that friendship will not be weakened by the 
present author's nomenclatural diminution of their discoveries. 


DICERANDRA Benth. Scrub Balms ' 


1. Annual (or persisting a second year and developing a thickened 
collar at soil line, but not woody), unbranched or with few 
spreading to ascending branches well above base. 

2. Peduncles absent or very short, the flowers numerous and crowded 
in axils of leaves or leafy bracts; anther horns abrupt, obtuse or 
barely acute. Pungently aromatic annual herb, to 0.4 m. Dry 
woods, pinelands, roadsides. North peninsula (Lafayette, 


274 Phytologia (August 2009) 91(2) 


Suwannee, s. to Levy, Volusia counties); infrequent. Fall. 
Endemic. Dicerandra densiflora Benth in DC. 


2. Peduncles (except at upper nodes) evident, usually well developed, 
few-flowered, the inflorescences rather loose; anther horns 
subulate, acuminate. Annual herb, to 0.4 m. Dry pineland, dry 
open hammocks. Fall. Dicerandra linearifolia (Ell.) Benth. 


a. Corolla pale pink to white; anthers yellow; leaves narrow. 
West and central panhandle (e. to Leon, Wakulla counties); 
infrequent. var. linearifolia 


a. Corolla dark purple; anthers reddish-brown; leaves wider. 
Central and east panhandle (Jackson to Suwannee), disjunct 
eastward (to Alachua, Duval counties); infrequent. 

var. robustior Huck 


1. Perennial, woody below, with numerous erect to ascending branches 
from near base. Low shrub, to 0.6 m. Sand pine scrub. Summer- 
fall. Endemic. ENDANGERED (Federal, State listings). 

SCRUB BALM. Dicerandra frutescens Shinners 


a. Corolla without spots; corolla tube smoothly curved. 


b. Habit upright; leaves narrowly oblanceolate, 1.5-4.0 mm. 
wide; corolla light lavender. East coast of peninsula (Indian 
River... St... Lucie. counties); very wate. Endemic. 
ENDANGERED (Federal, State listings). [Dicerandra 
immaculata Lakela] Occasional white-flowered plants have 
been named forma nivea Lakela. 

var. immaculata (Lakela) D. B. Ward 


b. Habit lax; leaves rhombic, 1.2-12.0 mm. wide; corolla vivid 
pink. East coast of peninsula (St. Lucie County); very rare 
(two populations, with 200 individual plants). Endemic. 
[Dicerandra immaculata var. savannarum Huck] 

var. savannarum (Huck) D. B. Ward 


Phytologia (August 2009) 91(2) ZIS 


a. Corolla with spots or blotches, at least on upper petals; 
corolla tube abruptly bent. 
c. Corolla reddish-purple; anther spurs >1 mm. long. 


d. Leaves very narrow (+1.0 mm. wide); style glabrous or with 


few hairs. North-central peninsula (Marion, Sumter 
counties); rare. Endemic. ENDANGERED (Federal, State 
listings). [Dicerandra cornutissima Huck] 

var. cornutissima (Huck) D. B. Ward 


d. Leaves broader (+1.3 mm. wide); style hispid. Eastern 


coastal peninsula (Brevard County: Titusville); very rare 
(three known populations). Endemic. ENDANGERED 
(State listing). [Dicerandra thinicola H. A. Miller] 

var. thinicola (H. A. Miller) D. B. Ward 


c. Corolla white or cream; anther spurs <1 mm. long; style with 


e. 


numerous stiff hairs; leaves narrowly oblong (1.3-2.5 mm. 
wide). 


Anthers bright yellow, with few or no glands on connective; 
crushed leaves with wintergreen odor. Central peninsula 
(Highlands County: Sebring); very rare (five known 
populations). Endemic. ENDANGERED (Federal, State 
listings). [Dicerandra christmanii Huck & Judd] 

var. christmanii (Huck & Judd) D. B. Ward 


Anthers white or lavender, with abundant glands on 
connective; crushed leaves with minty fragrance. 


276 Phytologia (August 2009) 91(2) 


f. Inflorescence with 1-2 flowers per cyme; corolla white or 
rarely pink. Central peninsula (Polk, Highlands County: 
Lake Placid); very rare (nine extant populations). Endemic. 
ENDANGERED (Federal, State listings). var. frutescens 


f. Inflorescence with 2-3 flowers per cyme; corolla pinkish 
white, turning pink with age. Central peninsula (Polk 
County: Davenport); very rare (two known populations). 
Endemic. [Dicerandra frutescens ssp. modesta Huck] 

var. modesta (Huck) D. B. Ward 


This paper is a continuation of a series begun in 1977. The "amplified key" format 
employed here is designed to present in compact form the basic morphological 
framework of a conventional dichotomous key, as well as data on habitat, range, and 
frequency. Amplified keys are being prepared for all genera of the Florida vascular flora; 
the present series is restricted to genera where a new combination is required or a special 


situation merits extended discussion. 


Phytologia (August 2009) 91(2) ZiT 


INFRASPECIFIC VARIATION IN HESPEROCYPARIS 
GOVENINA AND H. PYGMAEA: ISSRS AND TERPENOID 
DATA 


Robert P. Adams 
Biology Department, Baylor University, Box 727 
Gruver, TX 79040, USA 
Robert_ Adams@baylor.edu 


Jim A. Bartel 
U.S. Fish and Wildlife Service, Carlsbad Fish and Wildlife Office 
6010 Hidden Valley Road, Suite 101 
Carlsbad, CA 92011-4213 


ABSTRACT 


Hesperocyparis (Cupressus) goveniana and __ putative 
Cupressus goveniana subsp. gibsonensis plus H. pygmaea were 
analyzed by Inter-Simple Sequence Repeats (ISSRs). The ISSRs 
analyses support the continued recognition H. pygmaea and H. 
goveniana, but the recognition of Cupressus goveniana subsp. 
gibsonensis Silba was not supported. Phytologia 91(2):277-286 
(August, 2009). 


KEY WORDS: Hesperocyparis (= Cupressus) goveniana, Cupressus 
goveniana subsp. gibsonensis, H. pygmaea, ISSR, Inter-Simple 
Sequence Repeats, terpenes, DNA fingerprinting, systematics. 


The coastal California species of Hesperocyparis (= 
Cupressus) are often found in very small populations and several are 
endangered. The taxonomy of the western hemisphere cypresses has 
recently changed as DNA sequencing of the classical Cupressus 
species (Little et al., 2004; Little, 2006) demonstrated that the western 
hemisphere cypresses are a separate genus from the eastern hemisphere 
cypresses. The eastern hemisphere cypresses maintained the name 
Cupressus and Callitropsis was applied to the western hemisphere 
cypresses plus Callitropsis nootkatensis (D. Don) Oersted ex D. P. 
Little. However, Debreczy, et al. (2009) argued, on morphological 


278 Phytologia (August 2009) 91(2) 


grounds, that Callitropsis nootkatensis is a monotypic genus. 
Sequencing of two additional nuclear genes and petN-psbM (Adams et 
al., 2009) supported the recognition of Callitropsis nootkatensis as a 
monotypic genus. Thus, Callitropsis could not be applied to the 
western hemisphere cypresses, so Adams et al. (2009) erected a new 
genus, Hesperocyparis, for these cypresses. 


Silba (2003) proposed the recognition of several new 
subspecies of Cupressus abramsiana Wolf: C. a. subsp. locatellii 
Silba, Eagle Rock, CA; C. a. subsp. op/eri Silba, Bracken Brae Forest, 
Santa Cruz, CA; C. a. subsp. neolomondensis Silba, Majors Creek, CA; 
and C. a. subsp. butanoensis Silba, Butano Ridge, CA. Cupressus a. 
subsp. butanoensis was recognized (Adams et al., 2009) as 
Hesperocyparis abramsiana var. butanoensis (Silba) Bartel & R. P. 
Adams. Silba (2003) also proposed C. goveniana Gordon subsp. 
gibsonensis Silba and C. macrocarpa Hartw. subsp. lobosensis Silba. 
However, the proposed new subspecies are morphologically rather 
indistinct. If these new subspecies are accepted, then these taxa need 
to be considered under the endangered species act. 


Adams and Bartel (2009) examined the volatile leaf oils of H. 
goveniana (Gordon) Bartel, C. g. subsp. gibsonensis Silba and H. 
pygmaea (Lemmon) Bartel (Fig. 1). The leaf oils of these taxa appear 
to separate them into 2 groups (Fig. 1) composed of H. pygmaea and 
goveniana - gibsonensis. There was some variation among the three 
populations of H. pygmaea (Fig. 1) and a hint of differences between 
goveniana and gibsonensis. These differences appear to be in the 
nature of geographic variation. Adams and Bartel (2009) concluded 
that, based on the leaf terpenoids, there was insufficient evidence to 
support the recognition of Silba's Cupressus goveniana subsp. 
gibsonensis. 


To gather additional genetic information about the validity of 
these subspecies, analyses using Inter-Simple Sequence Repeats 
(ISSRs) were conducted. It should be noted that the leaf samples 
utilized in the present study were taken from the same trees analyzed by 
Adams and Bartel (2009). 


Phytologia (August 2009) 91(2) 279 


PCO 


33 terpenes 
>0.5% conc. 


% = goveniana 


abamsiana subsp. O = gibsonensis 


neolomondensis 
high karahanaenone 


— 


Lt 


ti 


f 


@ = Albion Ridge 
O = Little River 
© = Casper Little Lake 


Figure 1. PCO of Hesperocyparis goveniana, Cupressus g. subsp. 
gibsonensis, H. pygmaea plus three plants of C. abramsiana subsp. 
neolomondensis (from Adams and Bartel, 2009). 


MATERIALS AND METHODS 


Plant material - Specimens used in this study: H. abramsiana 
(C. B. Wolf) Bartel, Bonny Doon, Santa Cruz Co., CA, Bartel 1598a-e; 
H. abramsiana var. butanoensis, Pescadero Creek County Park, Butano 
Ridge Fire Rd., San Mateo Co., CA, Bartel 1605a-e.; C. abramsiana 
subsp. /ocatellii Silba, Eagle Rock, Santa Cruz Co., CA, Bartel 1599a- 
e; C. abramsiana subsp. neolomondensis Silba, Wilder Ranch State 
Park, Santa Cruz Co., CA, Bartel 1604a-e; C. a. subsp. opleri Silba, 
Bracken Brae, Santa Cruz Co., CA, Bartel 1600a-e; H. goveniana, SFB 


280 Phytologia (August 2009) 91(2) 


Botanical Reserve, Monterey Co., CA, Bartel 1596a-e; C. goveniana 
subsp. gibsonensis Silba, Point Lobos Ranch, Monterey Co., CA, Bartel 
1595a-e; H. pygmaea, Albion Ridge, Mendocino Co., CA, Bartel 
1601la-e; Little River Airport, Bartel 1602a-e; Casper Little Lake Rd., 
CA, Bartel 1603a-e; C. macrocarpa subsp. lobosensis Silba, Point 
Lobos State Reserve, Allan Memorial Grove, Monterey Co., CA, Bartel 
1593a-e, East Grove, Bartel 1594a-e; H. macrocarpa (Hartw.) Bartel, 
Crocker Grove, 100 mn of 17 Mile Drive and Madre Lane intersection, 
Monterey Co., CA, Bartel 1597a-e. Bartel specimens are help in his 
personal herbarium in Carlsbad, CA. 


One gram (fresh weight) of the foliage was placed in 20 g of 
activated silica gel and transported to the lab, thence stored at -20° C 
until the DNA was extracted. DNA was extracted using the Qiagen 
DNeasy mini kit (Qiagen Inc., Valencia, CA). ISSR primers were 
purchased from the University of British Colombia (5'-3 seq., annealing 
temperature used'): 807: AGA GAG AGA GAG AGA GT (50°C), 808: 
AGA GAG AGA GAG AGA GC (50°C), 811: GAG AGA GAG AGA 
GAG AC (50°C), 812: GAG AGA GAG AGA GAG AA (50°C), 836: 
AGA GAG AGA GAG AGA GYA (54°C), 840: GAG AGA GAG 
AGA GAG AYT (54°C), 841: GAG AGA GAG AGA GAG AYC 
(54°C), 847: CAC ACA CAC ACA CAC ARC (58°C), 861: AGC AGC 
AGC AGC AGC AGC, (58°C), 881: GG TGG GGT GGG GTG (50°C), 
886° VDV CIC TCT CIC TCT CT (S0°C)887: DVD TCM ie tea 
CTGC.TE (54°C), 895: AGA GIT GGT AGC TCT. TGA: TEGGGe): 
900: ACT TCC CCA CAG GTT AAC ACA (50°C). 


PCR stock solutions (Taq, primer, and buffer) were made in 
bulk so that all the PCR reaction tubes for a primer were prepared using 
the same bulk stock. This is a critical factor for minimizing variation in 
band intensities from sample to sample (see Adams, Flournoy and 
Pandey, 1998, for protocols to minimize PCR band variation). PCR 
was performed in a volume of 15 pl containing: 7.5 pl Epi-Centre 2X 
buffer E (containing 0.4 mM of each dNTP, final conc. = 0.2 mM), 0.75 
ul primer (0.6 uM final conc.), 0.75 wl Epi-Centre Fail-Safe Tag (0.75 
unit/rxn.), 6 wl genomic DNA (0.3 ng/rxn.). A negative control PCR 
tube containing all components, but no genomic DNA, was run with 
each primer to check for contamination. DNA amplification was 
performed in an MJ Programmable Thermal Cycler (MJ Research, 


Phytologia (August 2009) 91(2) 281 


Inc.). Samples were run in duplicate to insure reproducibility (Adams, 
Flournoy and Pandey, 1998). A temperature profile was obtained for 
each well of the thermocycler to be sure that no variation existed 
among wells in the heating/ cooling block. The thermal cycle used 
was: 94° C (1.5 min) for initial strand separation, then 39 cycles of 91° 
C (1 min), 50°C (or 54°C or 58°C, see above) (2 min), 72° C (2 min). 
Two additional steps were used: 91° C (1 min), 50°C (or 54°C or 58°C) 
(2 min) and 72° C (5 min) for final extension. The temperature inside a 
PCR tube containing 15 ul buffer was monitored with a temperature 
probe, quantitated and printed for each step for each of the 40 cycles for 
every PCR run (Adams, Flournoy and Pandey, 1998) to insure that each 
cycle met temperature specifications and that each PCR run was exactly 
the same. Amplification products were analyzed by electrophoresis on 
1.5% agarose gels, 70V, 55 min, and detected by staining with ethidium 
bromide. The gels were visualized over UV light and scanned to digital 
images. The digital images were size normalized in reference to 
pGem® DNA size markers before band scoring. Bands were scored as 
present (4 = faint, 5 = bright, 6 = v. bright) and absent (0). Bands that 
were inconsistent in replicate analyses were not scored. 


Associational measures were computed using absolute 
character state differences (Manhattan metric), divided by the 
maximum observed value for that character over all taxa (= Gower 
metric, Gower, 1971; Adams, 1975). Principal coordinate analysis 
(PCO) was performed by factoring the associational matrix using the 
formulation of Gower (1966) and Veldman (1967). It should be noted 
that problems of homology of RAPD DNA bands on agarose gels can 
be significant (Rieseberg, 1996), but these errors can be accounted for 
using multivariate statistical methods (PCO) (see Adams and 
Rieseberg, 1998). A minimum spanning diagram was constructed by 
selecting the nearest neighbor for each taxon from the pair-wise 
similarity matrix, then connecting those nearest neighbors as nodes in a 
network (Adams, et al. 2003). 


RESULTS AND DISCUSSION 
The use of 14 ISSR primers resulted in 98 scoreable bands 


among the taxa. PCO of the association matrix removed five 
significant eigenroots accounting for: 15.2, 14.0, 9.1, 7.1, and 6.2% of 


282 Phytologia (August 2009) 91(2) 


the variance among the OTUs. It is noteworthy that there was 
considerable variation among individuals and this is reflected in the 
relatively small amount of variance that was extracted by the first 3 
eigenroots. 


Ordination of the taxa shows that H. goveniana and H. 
pygmaea are very well resolved from H. abramsiana and H. 
macrocarpa (Fig. 2). 


There appears to be considerable variation among the H. 
pygmaea individuals (Fig. 2). In addition, there is also variation among 
the goveniana - gibsonensis individuals (Fig. 2). 


3(9%) PCO 
98 ISSRs 


H. abramsiana 


H. macrocarpa 
Face 


Fay 


H. pygmaea 


2(14%) 


H. goveniana 
and C. g. subsp. 
gibsonensis 


Figure 2. PCO of H. goveniana, Cupressus goveniana subsp. 
gibsonensis, and H. pygmaea with exemplars of H. abramsiana and H. 
macrocarpa. 


Phytologia (August 2009) 91(2) 283 


In order to better visualize the variation among populations of 
H. pygmaea, a new PCO was performed which contained only the H. 
pygmaea individuals plus two exemplar H. goveniana samples. The 
PCO using 78 ISSRs resulted in four eigenroots accounting for 16.6, 
12.8, 10.0, and 8.9% of the variance among individuals. Clearly there 
are considerable differences among H. pygmaea individuals (Fig. 3). 
There is a trend for the Casper Little Lake population (plus 2 
individuals from the Albion Ridge population) to cluster together (Fig. 
3). The balance of the Albion Ridge and Little River population plants 
are interspersed (Fig. 3). There was also a trend in the terpenoids to 
subdivide these populations (Fig. 1), but the pattern is slightly different 


PCO 
78 ISSRs 


H. pygmaea 
@ = Albion Ridge 
H. goveniana 0 = Little River 
© = Casper Little Lake 


3(10%) 


Figure 3. PCO based on 78 ISSRs for H. pygmaea. Two individuals of 
H. goveniana were included in the PCO. 


284 Phytologia (August 2009) 91(2) 


than in the ISSR data (Fig. 3). It seems likely that these populational 
differences are normal geographic variation and should not to be 
recognized as separate taxa. 


The variation between H. goveniana and C. g. subsp. 
gibsonensis was further examined using 75 ISSRs for their ten samples 
plus two individuals of H. pygmaea. PCO resulted in four eigenroots 
that accounted for 23.1, 19.5, 11.7, and 10.2% of the variance among 
these OTUs. Ordination reveals (Fig. 4) that H. goveniana and putative 
C. g. subsp. gibsonensis form a large group, with some separation 
between the taxa. However, as these putative subspecies are each in 


3(12%) PCO 
75 ISSRs 


% = goveniana H. pygmaea 


O= gibsonensis 


1(23%) 


Figure 4. PCO of H. goveniana and C. g. subsp. gibsonensis 
individuals, plus two exemplars of H. pygmaea. 


Phytologia (August 2009) 91(2) 285 


distinct (and disjunct) populations, geographic differentiation could 
well explain this clustering. 


In summary, taking both the terpenoid and ISSR data into 
consideration, there appears to be sufficient genetic differentiation to 
support the recognition of H. pygmaea, but there is insufficient 
differentiation to support the recognition of Silba's Cupressus 
goveniana subsp. gibsonensis. 


ACKNOWLEDGEMENTS 
This research supported in part with funds from U. S. Fish and 
Wildlife Service, Grant 814307J011. The findings and conclusions in 
this article are those of the authors and do not necessarily represent the 
views of the U.S. Fish and Wildlife Service. Thanks to Tonya Yanke 
for lab assistance. 


LITERATURE CITED 


Adams, R. P. 1975. Statistical character weighting and similarity 
stability. Brittonia 27: 305-316. 

Adams, R. P. 2008. Junipers of the world: The genus Juniperus. 2nd 
Edition. Trafford Publ., Vancouver, B.C., Canada. 

Adams, R. P., J. A. Bartel and R. A. Price. 2009. A new genus, 
Hesperocyparis, for the cypresses of the western hemisphere. 
Phytologia 91(1): 160-185. 

Adams, R. P. and J. A. Bartel. 2009. Geographic in the leaf essential 
oils of Hesperocyparis (Cupressus) abramsiana, H. goveniana and 
H. macrocarpa: Systematic implications. Phytologia 91(2): 226- 
243. 

Adams, R. P., T. Demeke and H. A. Abulfatih 1993. RAPD DNA 
fingerprints and terpenoids: clues to past migrations of Juniperus 
in Arabia and east Africa. Theoret. Appl. Genetics 87: 22-26. 

Adams, R. P., L. E. Flournoy and R. N. Pandey. 1998. Obtaining 
reproducible patterns from random polymorphic DNA 
amplification (RAPDs). in Conservation of Plant Genes III: 
conservation and Utilization of African Plants. R. P. and J. E. 
Adams, eds., Missouri Botanical Garden, St. Louis. 


286 Phytologia (August 2009) 91(2) 


Adams, R. P.. and L. H. Rieseberg. 1998. The effects of non-homology 
in RAPD bands on similarity and multivariate statistical ordination 
in Brassica and Helianthus. Theoret. App. Gen. 97: 323-326. 

Bartel, J. A., R. P. Adams, S. A. James, L. E. Mumba and R. N. 
Pandey. 2003. Variation among Cupressus species from the 
western hemisphere based on random amplified polymorphic 
DNAs. Biochem. Syst. Ecol. 31: 693-702. 

Debreczy, Z., K. Musial, R. A. Price and I. Racz. 2009. Relationships 
and nomenclatural status of the Nootka cypress (Callitropsis 
nootkatensis, Cupressaceae) Phytologia 91(1): 140-159. 

Gower, J. C. 1971. A general coefficient of similarity and some of its 
properties. Biometrics 27: 857-874. 

Gower, J. C. 1966. Some distance properties of latent root and vector 
methods used in multivariate analysis. Biometrika 53: 326-338. 
Little, D. P., A. E. Schwarzbach, R. P. Adams and C-F. Hsieh. 2004. 
The circumscription and phylogenetic relationships of Callitropsis 
and the newly described genus Xanthocyparis (Cupressaceae). 

Amer. J. Bot. 91: 1872-1881. 

Little, D. P. 2006. Evolution and circumscription of the true cypresses 
(Cupressaceae, Cupressus). Syst. Bot. 31: 461-480. 

Rieseberg, L. H. 1996. Homology among RAPD fragments in 
interspecific comparisons. Mol. Ecol. 5: 99-105. 

Silba, J. 2003. Field observations of Cupressus in central and coastal 
California, July 2002 to January 2003. J. Intl. Conifer Pres. Soc. 
10: 1-49. 

Veldman, D. J. 1967. Fortran programming for the behavioral sciences. 
Holt, Rinehart and Winston Publ., NY. 


Phytologia (August 2009) 91(2) 287 


INFRASPECIFIC VARIATION IN HESPEROCYPARIS 
ABRAMSIANA: ISSRS AND TERPENOID DATA 


Robert P. Adams 
Biology Department, Baylor University, Box 727 
Gruver, TX 79040, USA 
Robert_ Adams@baylor.edu 


Jim A. Bartel 
U.S. Fish and Wildlife Service, Carlsbad Fish and Wildlife Office 
6010 Hidden Valley Road, Suite 101 
Carlsbad, CA 92011-4213 


ABSTRACT 


Five Hesperocyparis (Cupressus) abramsiana groves were 
analyzed by Inter-Simple Sequence Repeats (ISSRs). ISSRs analyses 
revealed geographical differentiation among the groves (= Silba's 
subspecies) with the Butano Ridge grove being the most distinct in 
both ISSRs and terpenoids. Combined ISSR and terpenoid data 
support the recognition of Hesperocyparis. abramsiana_ var. 
abramsiana and Hesperocyparis abramsiana var. butanoensis Bartel 
& R. P. Adams, comb. nov. The recognition of Silba's subsp. 
locatellii, neolomondensis, and opleri were not support by these data. 
Phytologia 91(2): 287-299 (August, 2009). 


KEY WORDS: Hesperocyparis (= Cupressus) abramsiana_ vat. 
butanoensis, C. subsp. locatellii, C. subsp. neolomondensis, C. subsp. 
opleri, ISSR, Inter-Simple Sequence Repeats, terpenes, DNA 
fingerprinting, systematics. 


Hesperocyparis (= Cupressus, see Adams et al., 2009) 
abramsiana (C. B. Wolf) Bartel, widely known as Santa Cruz cypress, 
was listed (as Cupressus abramsiana) as an endangered species under 
the Endangered Species Act of 1973 (ESA) by the U.S. Fish and 
Wildlife Service (USFWS) in 1987. According to the ESA recovery 
plan developed for the species (USFWS 1998), H. abramsiana is 
restricted to five groves or populations (Fig. 1) that include a total of 


288 Phytologia (August 2009) 91(2) 


5,100+ individuals that collectively occupy about 142 ha (356 acres) 
within a 24-km (15-mile) range in the Santa Cruz Mountains in Santa 
Cruz and San Mateo counties, California, USA. Using an ESRI shape 
file of the grove boundaries provided by the USFWS, McGraw (2007) 
clarified that the areal extent of the cypress groves depicted in the 
recovery plan actually include 41.28 ha (102.0 acres). McGraw (2007) 
estimated that the areal extent of the five groves totals only 25.87 ha 
(63.9 acres). 


H. abramsiana 
distribution 


Butano Ridge 
a 


Eagle Rock ° acken Brae 


Boulder Creek 


Bonny Doon S 


Q 


i 1) 9 
Majors Creek 
a sestatabaienetbebaiaaitamheaianadionmesnaeianaieestaadaaaaiameadadididl 
dha Santa Cruz 


Figure 1. Distribution of H. abramsiana groves endemic to 
Santa Cruz and San Mateo counties, California, USA. Geographic 
limits of individual cypress stands within each grove, which are 
depicted in hatched polygons atop halftone circles, were derived from 
Fig. 2 from McGraw (2007). 


Phytologia (August 2009) 91(2) 289 


Using the ESA recovery plan and after visiting four of the five 
groves or populations, Silba (2003) subdivided Cupressus abramsiana 
into five subspecies with his descriptions of four new subspecies; C. a. 
subsp. /ocatellii Silba (restricted to the Eagle Rock grove in Santa Cruz 
County), C. a. subsp. op/eri Silba (restricted to the Bracken Brae grove 
in Santa Cruz County), C. a. subsp. neolomondensis Silba (restricted to 
the Majors Creek grove in Santa Cruz County), and C. a. subsp. 
butanoensis Silba (restricted to the Butano Ridge grove in San Mateo 
County). According to Silba (2003), the nominate subspecies is 
restricted to the type locality of H. abramsiana (i.e., the Bonny Doon 
Grove in Santa Cruz County), which Wolf (1948) detailed as “on the 
southwest slope of Ben Lomond, a mountain 7/10 mi. east of the 
Bonnie Doon School, elevation 1600 feet.” Given the lack of a 
taxonomic key or a set of clear morphological characters to separate 
the subspecies, and the overall poor quality of the self-published article 
(e.g., orthographic errors, illegibility), Silba’s (2003) new subspecies 
seem to be morphologically indistinct and separated largely by 
collection locality. 


Adams and Bartel (2009) examined the volatile leaf oils of H. 
abramsiana and Silba's subsp. butanoensis, locatellii, neolomondensis, 
and opleri (Fig. 2). The leaf oils of these taxa appear to separate H. 
abramsiana into 2 groups (Fig. 2) composed of butanoensis and the 
remaining four groves. However, note that 3 trees of butanoensis had 
oils similar to H. pygmaea (Fig. 2, see also Table 1, Adams and Bartel, 
2009). Adams and Bartel (2009) concluded that additional research 
was needed into the nature of infraspecific variation in H. abramsiana. 


To gather additional genetic information about the validity of 
these subspecies, analyses using Inter-Simple Sequence Repeats 
(ISSRs) were conducted. The leaf samples utilized in the present study 
were taken from the same trees analyzed by Adams and Bartel (2009). 


MATERIALS AND METHODS 
Plant material - Specimens used in this study: UH. 


abramsiana, Bonny Doon Grove, Santa Cruz Co., CA, Bartel 1598a-e; 
Butano Ridge Grove, (butanoensis), Pescadero Creek County Park, 


290 Phytologia (August 2009) 91(2) 


2(18% 
PCO ne H. macrocarpa 


2/ terpenes 
>1% conc. 


H. abramsiana las goveniana 
subsp. 
"butanoensis" 


1. pygmaea 


3(10%) 


= abram., Bonny Doon = abram. subsp “locatellii" 
%= abram. subsp. "opleri" @= abram. subsp. “neolomondensis" 


Figure 2. PCO of H. abramsiana and Silba's subspecies based on 27 
terpenes. From Adams and Bartel (2009). 


San Mateo Co., CA, Bartel 1605a-e.; Eagle Rock Grove, (/ocatellii), 
Santa Cruz Co. CA, Bartel 1599a-e; Majors Creek Grove, 
(neolomondensis), Wilder Ranch State Park, Santa Cruz Co., CA, 
Bartel 1604a-e,; Bracken Brae Grove, (opleri), Santa Cruz Co., CA, 
Bartel 1600a-e; H. goveniana, SFB Botanical Reserve, Monterey Co., 
CA, Bartel 1596a-e; H. pygmaea, Casper Little Lake Rd., CA, Bartel 
1603a-e. Bartel specimens are held in his personal herbarium in 
Carlsbad, CA. 


Phytologia (August 2009) 91(2) 29] 


One gram (fresh weight) of the foliage was placed in 20 g of 
activated silica gel and transported to the lab, thence stored at -20° C 
until the DNA was extracted. DNA was extracted using the Qiagen 
DNeasy mini kit (Qiagen Inc., Valencia, CA). ISSR primers were 
purchased from the University of British Colombia (5'-3 seq., annealing 
temperature used): 807: AGA GAG AGA GAG AGA GT (50°C), 808: 
AGA GAG AGA GAG AGA GC (50°C), 811: GAG AGA GAG AGA 
GAG AC (50°C), 812: GAG AGA GAG AGA GAG AA (50°C), 836: 
AGA GAG AGA GAG AGA GYA (54°C), 840: GAG AGA GAG 
AGA GAG AYT (54°C), 841: GAG AGA GAG AGA GAG AYC 
(54°C), 847: CAC ACA CAC ACA CAC ARC (58°C), 881: GG TGG 
Sia GGG, GIG. (50°C); $86: VDV CIC TCT CTC’ TCT Cr Ge'C), 
tan ov TCT CT@ TCT'CIC TC 64°C), 895; AGA GTT GGT AGC 
er TGA TC (50°C). 


PCR conditions and numerical methods - see Adams and 
Bartel, 2009. 


RESULTS AND DISCUSSION 


The 12 ISSR primers resulted in 89 scoreable bands among the 
taxa. A minimum spanning network was constructed using the matrix 
of associations and is shown in figure 3. Individuals of each of the five 
groves of H. abramsiana cluster together. Hesperocyparis goveniana 
and H. pygmaea are well resolved (Fig. 3). However, the Butano Ridge 
grove actually clustered after 1. pygmaea clusters (Fig. 3). 


PCO of the association matrix removed six significant 
eigenroots accounting for: 13.74, 11.84, 10.51, 7.89, 7.70 and 6.67% of 
the variance among individuals. Ordination shows a similar pattern 
(Fig. 4) with H. goveniana and H. pygmaea being distinct from H. 
abramsiana. Most of the H. abramsiana groves are not resolved. 


Removing H. goveniana and H. pygmaea from the data set and 
running a PCO focuses on the H. abramsiana groves. Ordination 
shows that all five H. abramsiana groves are resolved (Fig. 5). A 
minimum spanning network (dashed lines, Fig. 5) shows that Butano 
Ridge population is the most divergent grove (0.730). While three 
groves consist of asingle largely distinct stand of H. abramsiana, the 


292 Phytologia (August 2009) 91(2) 


Minimum Spanning Network 
89 ISSR bands 


Eagle Rock 


Majors Creek 


© 

c ® 
s&s 2 
ym © 
Ec 
fs 
Q Gb 
GO ® 
i 
+ 


Bonny Doon 


Butano Ridge 


Figure 3. Minimum spanning network based on 89 ISSR bands. Gov = 
H. goveniana, Pyg = H. pygmaea. 


Phytologia (August 2009) 91(2) 293 


2(12%) PCO 
89 ISSRs 


abramsiana 
groves 


abramsiana 
Majors Creek 
< 


pygmaea 


goveniana 


abramsiana 
Butano Ridge 


Figure 4. PCO based on 89 ISSR bands. 


Eagle Rock and Majors Creek groves are made up of three and two 
stands respectively (USFWS 1998, McGraw 2007). Though Silba 
(2003) used the groves or populations as identified in the ESA recovery 
plan to delimit his five subspecies, he also included within C. a. subsp. 
neolomondensis two collections (B-256 and B-257) from “a small grove 
of less than a dozen trees” within the Bonny Doon Ecological Reserve. 
This collection site apparently is between the Bonny Doon and Majors 
Creek groves, yet west of Laguna Creek and across Martin Road from 
the largest cypress stand and type locality. Much of the ecological 
reserve, including most of the cypress grove, burned in June 2008 in the 
210-ha (520-acre) Martin Fire. 


The five groves range from about 2.2 to 25.7 km (1.4 to 16.1 
miles) apart, though the populations generally fall within three 
watershed basins. Going south to north, the Bonny Doon and 


294 Phytologia (August 2009) 91(2) 


PCO 89 ISSRe 
H. abramsiana 
groves 


Majors Creek Bracken Brae 


Eagle Rock 


Bonny Doon 


Figure 5. PCO based on 89 ISSR bands analyzing five H. abramsiana 
groves. 


Majors Creek groves lie between Mill Creek (a tributary to San Vicente 
Creek) and Majors Creek with Laguna Creek bisecting both groves. 
The Eagle Rock and Bracken Brae groves largely fall within the 
watershed of Boulder Creek and its tributary Jamison Creek. The four 
southernmost groves drain generally south toward Santa Cruz or 
immediately to the northwest of Monterey Bay. However, the Butano 
Ridge grove, well isolated from the other groves, is located within the 
Butano Creek watershed, which drains west and empties into the 
estuary in Pescadero Creek State Beach. With this hydrology in mind, 
the four southernmost groves are very similar (0.747 - 0.754), while the 
divergence of the Butano Ridge grove (0.730) correlates with its 
relative isolation from the other populations. 


Phytologia (August 2009) 91(2) 295 


In summary, both the terpenoids (Fig. 2) and ISSRs (Figs. 3-6) 
data show that the Butano Ridge grove is differentiated from the four 
southernmost H. abramsiana groves. Little variation was found among 
the four groves in the terpenoids (Fig. 2). The three individuals (Fig. 2) 
that had leaf oils similar to H. pygmaea were not similar to H. pygmaea 


H. abramsiana 
Minimum Spanning 
Network 

Butano Ridge 


Eagle Rock 2 acken Brae 


eg” - 


153 


Boulder Creek 


* Ben Lomond 


. 754 \ 


Bonny Doon - Felton 
»: 


) 
QO 
Majors Creek 


8 km (5 miles) 


Figure 6. Minimum spanning network (dotted lines) superimposed on a 
geographic map of H. abramsiana groves. 


296 Phytologia (August 2009) 91(2) 


in their ISSRs (Figs. 4-6). None of the subspecies proposed by Silba 
(2003) are supported by terpenoid or DNA data, except the Butano 
Ridge grove. This taxon is recognized as a variety: 


Hesperocyparis abramsiana (C. B. Wolf) Bartel var. butanoensis 
(Silba) Bartel & R. P. Adams, comb. nov. 

Basionym: Cupressus abramsiana C. B. Wolf subsp. butanoensis 
Silba, J. Intl. Conifer Pres. Soc. 10: 34. 2003. 

Type: On sandstone slope, Butano Ridge, Santa Cruz 
mountains, north of Big Basin. (The cypress area includes the common 
comer of sections 11, 12, 13, 14 of township 8 South, Range 4 West, 
Mount Diablo Base and Meridian, Santa Cruz Quadrangle.), San Mateo 
County, 1 Sep 1951, C. McMillan 1620 (with P. McMillan, R. 
Bacigalupi, L. Heckard and H. Dutton) (holotype - NY). 


McMillan (1952) reiterated the above locality the holotype 
with his description that a "trek of approximately one-quarter mile 
down the slope leads directly into the cypress area, which probably 
includes the common corner of sections 11,12,13, and 14, of Township 
8 South, Range 4 West, Mount Diablo Base and Meridian, Santa Cruz 
quadrangle." However, the grove does not occur near this common 
corner of sections, but rather the cypresses are largely centered at N 37° 
14.512', W 122° 15.127' on land managed by Pescadero Creek County 
Park. 


McMillan (1952), in his article reporting the discovery of a 
third H. abramsiana grove, the Butano Ridge grove, noted that a 
“striking difference among the [then three known] populations is to be 
found in the size of the female cone.” After randomly sampling 100 
seed cones from each grove, McMillan (1952) reported that the average 
length of the Butano Ridge grove was 28 mm, Eagle Rock grove was 
24 mm, and Bonny Doon grove was 21 mm. With the exception of the 
Bracken Brae grove (n=11), we measured 45 to 70 cones per grove to 
validate and update McMillan’s data (Table 1). Not only were seed 
cones from Butano Ridge consistently longer than the other four 
groves, but the cones also were consistently wider with only four cones 
from the other groves falling within the measured range of the Butano 
Ridge grove cones. 


Phytologia (August 2009) 91(2) Pte 


Table 1. Grove-by-grove comparison of H. abramsiana seed cones. 
peeks aN 


- Butano | Eagle _ Bracken Bonny Majors 


Ridge Rock | Brae Doon Creek 
meancone | 27.0 | 221 | 167 | 220 | 199 
_ length (mm) | | | 
"mean cone 25.5. |.200 16.1 1:8 wh ness 
_ width (mm) | | 
“meannumbrr = 4.7, | «42 | (3.6 ZG ah i 
ofscale pairs | | | 
_ per cone | | 


Like cone size, the number of scales per cone pairs varies from 
cone to cone. McMillan (1952) reported that the Butano Ridge grove 
averaged 5.2 scale pairs per cone with 5 and 6 pairs “common.” While 
48% of our sampled cones had 5 and 6 scale pairs, the remaining cones 
had 4 pairs per cone. In contrast to Butano Ridge, McMillan (1952) 
reported that the Bonny Doon and Eagle Rock groves “were 
predominantly of 8-scaled [4-paired] cones, although the average 
number for the hundred cones was found to be 8.5 [4.3 pairs] and 8.7 
[4.4 pairs] respectively.” However, because we found that the Bonny 
Doon (31%) and Major Creek (45%) groves often have 5 scale pairs, 
only cone width and length (especially the former measurement) appear 
to reliably differentiate the Butano Ridge grove from the four other 
groves. 

McMillan (1952) noted that seed color (dark brown to dull 
black), presence of glaucous seeds, foliage texture (fine versus coarse), 
and foliage color (dark green versus yellow green) failed to 
differentiate any of the then three known groves. With the inclusion of 
the Bracken Brae and Majors Creek groves, our observations confirm 
that these morphological characters do not consistently differentiate any 
of the five groves from one another. While the average number of 
cotyledons per seedling clearly separated the Butano Ridge and Eagle 


298 Phytologia (August 2009) 91(2) 


Rock groves from the Bonny Doon grove (McMillan 1953), additional 
work is needed to determine whether this character has any taxonomic 
merit for H. abramsiana groves. 


ACKNOWLEDGEMENTS 


This research supported in part with funds from U. S. Fish and 
Wildlife Service, Grant 814307J011. The findings and conclusions in 
this article are those of the authors and do not necessarily represent the 
views of the U.S. Fish and Wildlife Service. Thanks to Tonya Yanke 
for lab assistance and thanks to Tim Hyland (State Parks Resources 
Ecologist at Henry Cowell Redwoods State Park) and Mark Schneider 
(Parks Ranger at Pescadero Creek County Park) for providing access to 
the cypress groves. 


LITERATURE CITED 


Adams, R. P. 2008. Junipers of the world: The genus Juniperus. 2nd 
Edition. Trafford Publ., Vancouver, B.C., Canada. 

Adams, R. P. and J. A. Bartel. 2009. Geographic in the leaf essential 
oils of Hesperocyparis (Cupressus) abramsiana, C. goveniana and 
C. macrocarpa: Systematic implications. Phytologia 91(2): 226- 
243. 

Adams, R. P., J. A. Bartel and R. A. Price. 2009. A new genus, 
Hesperocyparis, for the cypresses of the western hemisphere. 
Phytologia 91(1): 160-185. 

Adams, R. P. and J. A. Bartel. 2009. Infraspecific variation in 
Hesperocyparis goveniana and H. pygmaea: ISSRs and terpenoid 
data. Phytologia 91(2): 277-286. 

Bartel, J. A., R. P. Adams, S. A. James, L..E. Mumba and R. N. 
Pandey. 2003. Variation among Cupressus species from the 
western hemisphere based on random amplified polymorphic 
DNAs. Biochem. Syst. Ecol. 31: 693-702. 

Gower, J. C. 1971. A general coefficient of similarity and some of its 
properties. Biometrics 27: 857-874. 

Gower, J. C. 1966. Some distance properties of latent root and vector 
methods used in multivariate analysis. Biometrika 53: 326-338. 


Phytologia (August 2009) 91(2) 299 


Little, D. P., A. E. Schwarzbach, R. P. Adams and C-F. Hsieh. 2004. 
The circumscription and phylogenetic relationships of Callitropsis 
and the newly described genus Xanthocyparis (Cupressaceae). 
Amer. J. Bot. 91: 1872-1881. 

Little, D. P. 2006. Evolution and circumscription of the true cypresses 
(Cupressaceae, Cupressus). Syst. Bot. 31: 461-480. 

McGraw, J. 2007. Distribution, abundance, size structure and 
conservation status of three populations of the endangered Santa 
Cruz cypress (Callitropsis abramsiana). Unpublished reported 
prepared for U.S. Fish and Wildlife Service, Ventura, California. 
29 pp. + appendices. 

McMillan, C. 1952. The third locality for Cupressus abramsiana Wolf. 
Madrono 11: 189-194. 

McMillan, C. 1953. Variation in seedlings of Cupressus abramsiana 
Wolf. Madrono 12: 28-30. 

Rieseberg, L. H. 1996. Homology among RAPD fragments in 
interspecific comparisons. Mol. Ecol. 5: 99-105. 

Silba, J. 2003. Field observations of Cupressus in central and coastal 
California, July 2002 to January 2003. J. Intl. Conifer Pres. Soc. 
10: 1-49. 

U.S. Fish and Wildlife Service. 1987. Endangered and threatened 
wildlife and plants; Determination of endangered status for 
Cupressus abramsiana (Santa Cruz cypress). Federal Register 52: 
675-679. 

U.S. Fish and Wildlife Service. 1998. Recovery plan for the Santa 
Cruz cypress (Cupressus abramsiana). U.S. Fish and Wildlife 
Service, Portland, Oregon. 51 pp. + appendices. 

Veldman, D. J. 1967. Fortran programming for the behavioral sciences. 
Holt, Rinehart and Winston Publ., NY. 

Wolf, C. B. 1948. The new world cypresses - Part I: Taxonomic and 
distributional studies of the new world cypresses. Aliso 1: 1-250. 


300 Phytologia (August 2009) 91(2) 


FRANGULA BETULIFOLIA AND F. OBOVATA 
(RHAMNACEAE) 
ARE DISTINCT SPECIES 


Guy L. Nesom 
2925 Hartwood Drive, Fort Worth, TX 76109, USA 
http://guynesom.com 


John O. Sawyer 
Department of Biological Sciences, Humboldt State University, Arcata, 
CA 95521, USA 


ABSTRACT 

Frangula betulifolia var. obovata occurs in northern Arizona, 
Nevada, Utah, and Colorado and is geographically disjunct from var. 
betulifolia, which occurs in southern Arizona, New Mexico, Texas, and 
northern Mexico. The two taxa are consistently different in leaf shape 
and texture, and with their genetic isolation, each is appropriately 
treated at specific rank. A new combination to this effect is made here: 
Frangula obovata (Kearney & Peebles) Nesom & Sawyer, comb. et 
stat. nov. Phytologia 91(2): 300-307 (August, 2009). 


KEY WORDS: Frangula_ betulifolia, F. obovata, F. xblumeri 
Rhamnaceae, taxonomy. 


Frangula (Rhamnus) betulifolia (Greene) Grubov has been 
treated without formal variants by Johnston (1971), Johnston and 
Johnston (1969, 1978), Cronquist et al. (1997), and Welsh et al. (2003). 
In contrast, Rhamnus betulifolia var. obovata Kearney & Peebles was 
described by botanists working in Arizona, the only state where both of 
the putative varieties occur, and a recent treatment for Arizona 
maintains them as separate (Hill 2008). Although Cronquist et al. 
(1997) identified the plants as R. betulifolia, the corresponding 
illustration in his treatment depicts var. obovata. 


According to Kearney and Peebles (1960, p. 532), “The 
typical plant [of Frangula betulifolia], with elliptic or oblong leaves, is 
limited in Arizona to the south-central and southern counties. In the 


Phytologia (August 2009) 91(2) 301 


northern part of the state is found var. obovata Kearney & Peebles, the 
type of which was collected on Navajo Mountain, Coconino County. 
This variety is apparently common in and near the Grand Canyon and 
Havasu Canyon, and extends into southern Utah and Nevada, thus 
being well separated geographically from the main area of R. 
betulifolia. The variety is characterized by more or less obovate leaves 
with thicker, more prominent veins.” 


The current study corroborates the observations of Kearney 
and Peebles. The two taxa are consistently and discontinuously 
different in leaf morphology. Var. betulifolia occurs from northern 
Mexico into southeastern Arizona, southern New Mexico, and trans- 
Pecos Texas; var. obovata is geographically disjunct and occurs in 
northern Arizona and adjacent Nevada, Utah, and Colorado (Figs. | and 
2). In view of their morphological and geographic distinction, 
recognition of each at specific rank is appropriate. 


1. Leaf blades elliptic to oblong, elliptic-ovate, or narrowly ovate, 
1.6—2.6(—2.9) times longer than wide, relatively thin or slightly 
thickened, paler beneath, lateral veins (8—)9—13 pairs 
EP Ree. Se oe «suse te «Pe eR ecko eaea te Frangula betulifolia 

1. Leaf blades obovate to oblong-obovate or oblong, 1.2—1.8(—2.5) 
times longer than wide, distinctly thickened and nearly coriaceous, 
evenly colored on both surfaces, lateral veins (S—)6—8(—9) pairs. 
PET ee hed NSCRL ER OEEN S sncuedeetes Me sansa nese Frangula obovata 


Frangula betulifolia (Greene) Grubov, Trudy Bot. Inst. Akad. Nauk 
S.S.S.R., Ser. 1, Fl. Sist. Vyssh. Rast. 8: 268. 1949. Rhamnus 
betulifolia Greene, Pittonia 3: 16. 1896. Type: USA. New 
Mexico. [Catron Co.:] along streams, Mogollon Mountains, 20 
Jul 1881, H.H. Rusby 63 (holotype: US-digital image!; 
isotypes: MO!, NY-digital image!). 


Shrubs or small trees 1-4 m, unarmed, stems brown to gray- 
brown, glabrous or pubescent. Leaves deciduous, alternate, petioles 
(2-)5—-16 mm, blades elliptic to oblong, elliptic-ovate or narrowly 
ovate, (4—)4.5—10 cm x (2—)2.5—5.5 cm, 1.6—2.6(—2.9) times longer than 
wide, thin or thickened but not coriaceous, green above, yellowish and 
paler beneath, hirtellous to hirsutulous on both surfaces, glabrescent, 


302 Phytologia (August 2009) 91(2) 


lateral veins (8—)9—13 pairs, margins serrate to subcrenate, not revolute, 
apices acute to obtuse, sometimes slightly acuminate, bases obtuse to 
truncate or rounded. Flowers bisexual, 5-merous, 2—20(—38) in 
pedunculate axillary fascicles, peduncles (flower) (0—)1-10 mm, 
pedicels (flower and fruit) 3-7 mm. Stigmas 3-lobed. Drupes 
globose, 5—10 mm, black, stones (2—)3(-4). 

Flowering Apr-Jun. Cliff bases, ledges, moist canyons, 
ridges, roadsides, rocky slopes, stream banks, Gambel’s oak, oak-pine, 
pine-walnut-maple, white fir; 900-2750 m.  Ariz., N.Mex., Tex.; 
Mexico (Chihuahua, Coahuila, Durango, Nuevo Leon, Sonora, 
Tamaulipas). 


Powell (1997) observed that “The leaves of the specimens 
from the Guadalupe Mountains are smaller and thinner in texture than 
those of the Davis Mountains population.” Such a difference has not 
been confirmed here among numerous specimens examined from both 
areas. 


Frangula obovata (Kearney & Peebles) Nesom & Sawyer, comb. et 
stat. nov. Rhamnus betulifolia var. obovata Kearney & 
Peebles, J. Wash. Acad. Sci. 29: 486. 1939. Frangula 
betulifolia subsp. obovata (Kearney & Peebles) Kartesz & 
Gandhi, Phytologia 76: 448. 1994. Type: USA. Arizona. 
Coconino Co.: Near Rainbow Lodge, N end of Navajo Mt., 
1920 m, 11 Jun 1938, R.H. Peebles 13930 with E.G. Smith 
(holotype: US-digital image!; isotypes: MO!). 


Shrubs 1—2.5 m, unarmed, stems red to brown or gray-brown, 
glabrous or pubescent. Leaves deciduous, alternate, petioles S—14 mm, 
blades obovate to oblong-obovate or oblong, (4—)5—9 cm x 3.2—6 cm, 
1.2—1.8(—2.5) times longer than wide, distinctly thickened and nearly 
coriaceous, green and minutely puberulous to hirtellous on both 
surfaces, glabrescent, lateral veins (S—)6—8(—9) pairs, margins minutely 
serrate to nearly entire, not revolute, apices obtuse to truncate or 
rounded, bases truncate to subcordate. Flowers bisexual, 5-merous, 2— 
12 in pedunculate axillary fascicles, peduncles (flower) 3—8(—20) mm, 
pedicels: (flower and fruit) 3-10 mm. Stigmas 3-lobed. Drupes 
globose, 5—8 mm, black, stones 3. 


Phytologia (August 2009) 91(2) 303 


Flowering Apr-Jun. Canyon bottoms, cliff faces, stream and 
creek banks, hanging gardens, talus, seepage below cliffs; 1350-2350 
m. Ariz., Colo., Nev., Utah. 


Frangula obovata has been collected infrequently in Nevada, 
but the identity of the plants there is unequivocal. Nevada. Clark Co.: 
Sheep Range, Grapevine Spring area, Rhamnus-white fir, 6300 ft, 19 
Sep 1978, Ackerman 31463 (TEX); Charleston (Spring) Mountains, 
Kyle Canyon, gravelly side of ravine with Pinus ponderosa var. 
scopulorum and Cercocarpus ledifolius, 2425 m, 11 Aug 1937, Clokey 
7579 (LL, TEX). Wolf (1938) cited the same Clokey collection as well 
as one other collection of “Rhamnus betulifolia” from Kyle Canyon (24 
Jun 1926, Jaeger s.n., CAS). 


Other plants from the Charleston Mountains are nearly 
identical to many of Frangula californica var. ursina over its wider 
range, with coriaceous, abaxially whitened leaves with a dense, close 
tomentum of stellate hairs: Clark Co.: gravelly wash a mile N of 
Wilson’s Ranch, Larrea belt, 1200 m, 13 Jul 1939, Clokey 8415 (MO-2 
sheets, TEX); Excelsior Canyon, 1200 m, 7 Sep 1941, Clokey 8762 
(MO, TEX); La Madre Mts, Willow Spring, 3 May 1988, Liston & 
Meury 740-2 (TEX). 


Harrington (1954) noted that Rhamnus betulifolia “has been 
reported close to southwestern Colorado [in Utah] and may be growing 
in that part of the state.” Most listings of this species in Colorado 
perhaps have been based on Harrington’s inclusion, but recent accounts 
of the Colorado flora (e.g., Weber & Wittman 1992; Hartman & Nelson 
2001; Snow 2007) have not included it. A collection from La Plata 
County in southwestern Colorado is identified in the forthcoming Four 
Corners Flora (Spence 2008) as Frangula betulifolia, but it is here 
recognized as F. obovata: La Plata Co.: Fort Lewis College on Ft. 
Lewis hill, 28 Jun 1976, V. Murray s.n. (SJNM 2118). 


Wolf (1938, p. 78-79) noted that a collection from Cochise 
Co. in southeastern Arizona named as Rhamnus blumeri Greene 
appears to be a hybrid between Frangula californica var. ursina and F. 
betulifolia. Both the holotype (US) and an isotype (DS) “have two 
pieces of material on them: the one is a vegetative branch which is 


304 Phytologia (August 2009) 91(2) 


obviously R. betulifolia, the other the material upon which Greene 
based his species. The latter resembles R. californica ursina, but is 
larger in leaf size and has less pubescence. In 1928, I collected around 
Paradise and obtained material of R. betulifolia (C.B. Wolf 2595). The 
other collections were made from large bushes resembling R. 
californica ursina in habit but only lightly pubescent on the under 
surfaces of the leaves. These suggest intermediates between R. 
californica ursina and R. betulifolia, but are slightly different from the 
type of R. blumeri: Collections: C.B. Wolf 2592 (RSA), 2593 (RSA).” 
Study of an isotype of R. blumeri, specimens of Wolf 2592 (MO), Wolf 
2593 (MO), Wolf 2595 (MO-2 sheets), and Wolf & Everett 11384 
(TEX), essentially corroborate Wolf’s observations. From numerous 
other collections of both species from the Chiricahua Mountains, 
however, we conclude that if hybridization has taken place between F. 
betulifolia and F. californica in Cochise Co., it apparently has not been 
a common occurrence and there is no evidence at hand of introgression. 


Frangula xblumeri (Greene) Kartesz & Gandhi, Phytologia 76: 
448.1994. Rhamnus blumeri Greene, Leafl. Bot. Observ. 
Crit. 2:266. 1912... Type: USA. ‘Arizona Cochisem@ea:, 
Chiricahua Mountains, Paradise, small tree near creek, 5300 
ft, 28 Aug 1906, J.C. Blumer 1290 (holotype: US, digital 
image!; isotypes: DS, MO!). 


Johnston (1971) noted that Frangula betulifolia is “extremely 
similar to, and probably conspecific with [Frangula caroliniana (Walt.) 
A. Gray]” of eastern North America. The two taxa are closely similar, 
but they are allopatric and have never been formally merged into a 
single species. 


ACKNOWLEDGEMENTS 


We are grateful to the staffs at MO and TEX for help and 
hospitality during visits there, to Ken Heil for information on SJNM 
collections, and to Dr. Billie Turner for his comments on the 
manuscript. 


Phytologia (August 2009) 91(2) 305 


LITERATURE CITED 


Albee, B.J., L.M. Shultz and S. Goodrich. 1988. The Atlas of the 
Vascular Plants of Utah. Utah Museum of Natural History. 
Digital version <http://www.gis.usu.edu/Geography- 
Department/utgeog/utvatlas/>. Accessed January 2009. 

Cronquist, A., N.H. Holmgren and P.K. Holmgren. 1997. Vascular 
Plants of the Intermountain West, U.S.A. Vol. 3, Part A: Subclass 
Rosidae (except Fabales). New York Botanical Garden Press. 

Harrington, H.D. 1954. Manual of the Plants of Colorado. Sage 
Books, Denver, Colorado. 

Hartman, R.L. and B.E. Nelson. 2001. A Checklist of the Vascular 
Plants of Colorado. Rocky Mountain Herbarium, Univ. of 
Wyoming, Laramie. <http://www.rmh.uwyo.edu/colorado/ 
colcklist.word.doc> Accessed January 2009. 

Hill, M.-E. 2008. Frangula. In Christie, K., M. Currie, L.S. Davis, 
M.-E. Hill, S. Neal and T. Ayers. Rhamnaceae. Vascular Plants of 
Arizona Project. Deaver Herbarium, Northern Arizona University, 
Flagstaff. Accessed Jan. 2009. 
<http://www4.nau.edu/deaver/VPA%20RHAMNACEAE. pdf> 

Johnston, M.C. 1971. Manual of the Vascular Plants of Texas. Texas 
Research Foundation, Renner, Texas. 

Johnston, M.C. and L.A. Johnston. 1969. Rhamnaceae, in Flora of 
Texas 2 (pt. IT):357-388. 

Johnston, M.C. and L.A. Johnston. 1978. Rhamnus. Flora Neotropica 
20:1-96. 

Kearney, T.H. and R.H. Peebles. 1960. Arizona Flora (ed. 2 with 
supplement by J.T. Howell, E. McClintock and collaborators. 
University California Press, Berkeley. 

Powell, A.M. 1997. Trees and Shrubs of the Trans-Pecos and 
Adjacent Areas. Univ. of Texas Press, Austin. 

Snow, N. 2007. Checklist of Vascular Plants of the Southern Rocky 
Mountain Region (Version 2). 
<http://www.conps.org/pdf/Plant%20Lists/SRMRChecklist_2007_ 
Version2_Final.pdf> Accessed Jan. 2009. 

Spence, J. In press (2008). Rhamnaceae. In Heil, K.D., S. O'Kane, 
and L. Reeves (eds.). Flora of the Four Corners Region: Vascular 
Plants of the San Juan River Drainage—Arizona, Colorado, New 
Mexico, Utah. Missouri Botanical Garden Press, St. Louis. 


306 Phytologia (August 2009) 91(2) 


UVSC Virtual Herbarium. 2008. Utah Valley State College, Orem, 
Utah. <http://herbarium.uvsc.edu/Virtual/> Accessed Jan. 2009. 

Weber, W.A. and R.C. Wittman. 1992. Catalog of the Colorado Flora: 
A Biodiversity Baseline. University of Colorado Museum, 
University Press of Colorado, Boulder. 

Welsh, S.L., N.D. Atwood, S. Goodrich and L.C. Higgins. 2003. A 
Utah Flora (ed. 3). M.L. Bean Life Science Museum, Brigham 
Young University Press, Provo, Utah. 

Wolf, C.B. 1938. The North American species of Rhamnus. Botanical 
Series No. 1. Rancho Santa Ana Botanic Garden, Claremont, 
California. 


@ Frangula obovata 


@ Frangula betulifolia 


Figure 1. Geographic distribution of Frangula betulifolia and F. 
obovata in the U.S.A. Map points are from specimens at LL, MO, 
SJNM, and TEX, augmented by records from Hill (2008), UVSC 
Virtual Herbarium (2008), and Albee et al. (1988). 


Phytologia (August 2009) 91(2) 307 


Sets 


@ Frangula betulifolia | 


Figure 2. Geographic distribution of Frangula betulifolia in Mexico. 


308 Phytologia (August 2009) 91(2) 


TAXONOMY OF ASCLEPIAS HIRTELLA AND 
A. LONGIFOLIA (APOCYNACEAE) 


Billie L. Turner 
Plant Resources Center 
The University of Texas 

Austin, Texas 78713 
billie@uts.cc.utexas.edu 


ABSTRACT 


The taxonomy of Asclepias hirtella and A. longifolia 1s briefly 
reviewed. It is concluded that they are best treated as two very distinct 
varieties: A. longifolia var. hirtella (Pennell) B,L, Turner, stat. nov. 
and A. longifolia (Raf.) var. longifolia. The two taxa intergrade to a 
limited extent in southwestern-most Louisiana and closely adjacent 
Texas. Typical members of var. /ongifolia in Louisiana are confined to 
the five easternmost counties of that state; all other collections from 
elsewhere in Louisiana and eastern Texas are essentially typical 
elements of var. hirtella. A distribution map of the complex in North 
America is provided. Phytologia 90(2): 308-311 (August, 2009). 


KEY WORDS: Apocynaceae, Asclepias hirtella, A. longifolia, Texas, 
Louisiana 


The present contribution was occasioned by the appearance of 
a paper by White (2008) entitled, “Asclepias hirtella (Apocynaceae) 
newly documented for the flora of Texas.” In this, he noted that the 
taxon had not been previously reported for the state of Texas, 
documenting the occurrence by a collection from Lamar County. He 
further commented that Turner et al. (2003) had mapped a Lamar Co. 
collection as A. longifolia, which he took to be A. hirtella. White fails 
to report that the latter had been treated as a subspecies of A. longifolia 
by Farmer and Bell (1985). Indeed, the latter workers annotated all of 
the Texas material on file at TEX as A. /. subsp. hirtella (Pennell) 
Farmer & Bell, annotations of which I concur, except for the rank 
rendered, as discussed below. 


Phytologia (August 2009) 91(2) 309 


It is likely that White’s misconception of the names concerned 
was occasioned by his reliance upon Woodson’s (1945) seminal 
treatment of Asclepias. In this, Woodson failed to map (or cite) A. 
hirtella as occurring in Texas, not having seen sheets from the area, 
most of these assembled after his treatment; however, he did state, 
“Asclepias hirtella and A. longifolia are so closely related that they 
might better be treated as subspecies.” Thus, the treatment of Farmer 
and Bell who, after the study and annotation of numerous specimens 
from 14 or more herbaria (including LL-TEX), made formal the 
infraspecific names concerned. 


BASIC TAXONOMY 


Asclepias longifolia dates back to 1803, typified by plants 
from the eastern seaboard, “in sylvis Georgiae occidentalibus.” 
Asclepis hirtella is typified by material from Jasper County, Missouri, 
this collected by Pennell himself and first published in 1919 (as 
Acerates hirtella). Since then it has been accounted for by a bevy of 
workers and is readily distinguished from A. hirtella by a number of 
characters, as well documented by Pennell in his original description. 
Indeed, A. hirtella is easily distinguished by pubescence alone, 
possessing a spreading (hirtellous) pubescence along the pedicels, A. 
longifolia having an upcurved-appressed pubescence, such vestiture 
characteristic throughout its range. 


My examination of specimens at LL-TEX suggested that the 
nomenclatural treatment of Farmer and Bell was sound, but I was 
curious to know the extent of possible intergradation of the two taxa in 
Louisiana, hence my loan of material from LSU. I was surprised to 
find that typical elements of 4. longifolia were to be found only in 
easternmost Louisiana (Parishes: Livingston, St. Helena, St. Tammany, 
Tangipohoa and Washington), the remainder referable to A. hirtella, at 
least by the pedicel vestiture mentioned in the above. Among the 
Louisiana specimens referred to A. hirtella, a few in southwestern 
Louisiana (Natchitoches Parish: Lynch 3884, LSU; Featherman s.n., 
LSU) and closely adjacent Texas (Jasper Co.: Orzell & Bridges 5678, 
TEX; Newton Co.: Correll 36544, LL) were found to have an 
intermediate pedicel vestiture; these are mapped in Fig.1. It is likely 


310 Phytologia (August 2009) 91(2) 


that such plants are a result of ancestral genetic contamination, and not 
extant hybridization, since I did not find the two taxa coexisting. 


Because of the apparent intergradation, however limited, of A. 
hirtella and A. longifolia | prefer to treat the two taxa as infraspecific 
categories, as proposed by Farmer and Bell, but would instead count 
these as varieties, as argued by Turner and Nesom (2000) and yet 
others. 


Asclepias longifolia var. hirtella (Pennell) B.L. Turner, stat. nov. 


Based upon Acerates hirtella Pennell, Bull. Torrey Bot. Club 46: 184. 
1919. 


ACKNOWLEDGEMENTS 
I am grateful to my colleagues Barney Lipscomb and Guy 
Nesom for reviewing the manuscript, and to LSU for the loan of 
specimens. 


LITERATURE CITED 


Farmer, J. and C.R. Bell. 1985. A new combination in Asc/epias. 
Phytologia 57: 380. 1985. 


Turner, B. L. and G.L Nesom. 2000. Use of variety and subspecies and 
new varietal combinations for Styrax platanifolius (Styracaceae). 
Sida 19: 257-262. 


White, M. 2008. Asclepias hirtella (Apocynaceae) newly documented 
for the flora of Texas. J. Bot. Res. Inst. Texas 2: 1495-1496. 


Woodson, R.E., Jr. 1954. The North American species of Asclepias. 
Ann. Missouri Bot. Gard. 41: 1-211. 


31] 


gia (August 2009) 91(2) 


Phytolo 


ee ‘ee 
eee Feee ol. 


; var. 


Fig. 1. Distribution of Asclepias longifolia: var. hirtella (dots) 


longifolia (open circles) 


; possible intergrades (small circles). 


° 


ai2 Phytologia (August 2009) 91(2) 


THREE NEW SPECIES OF KOANOPHYLLON (ASTERACEAE: 
EUPATORIEAE) FROM MEXICO 


Billie L. Turner 
Plant Resources Center 
The University of Texas 

Austin, Texas 78713 
billie@uts.cc.utexas.edu 


ABSTRACT 


Three new species of Koanophyllon are added to the Mexican 
Flora: K. coixtlahuacum B.L. Turner, from Oaxaca; K. concordianum 
B.L. Turner, from Sinaloa; and K revealii, from Guerrero and Oaxaca. 
In addition, a new varietal combination is proposed: K. solidaginoides: 
var. filicaulis (Sch.-Bip. ex A. Gray) B.L. Turner, a widespread taxon 
in eastern Mexico. Distribution maps are provided, along with 
photoholotypes. The several taxa are keyed along with yet other 
Mexican taxa in the format of Tumer’s 1997 treatment of 
Koanophylion in which 21 species were recognized; the current account 
brings this total to 24. Phytologia 91(2): 312-324 (August, 2009). 


KEY WORDS: Asteraceae, Eupatorieae, Koanophyllon, Mexico. 


Routine identification of Mexican Asteraceae has revealed the 
following novelties: 


KOANOPHYLLON COIXTLAHUACUM B.L. Turner, sp. nov. 
Fig. 1, Map 1 


Koanophyllon richardsonii B.L. Turner similis sed differt laminis 
foliorum multo majoribus, capitulescentiis magis congestis, et 
receptaculis glabris (vs pubescentibus). 


Shrubs 1-2 m high. Mid-stems minutely puberulent to glabrate. 
Leaves opposite throughout; blades broadly deltoid, 6-10 cm long, 4-7 
cm wide, 3-nervate from the base, glandular-punctate beneath, sparsely 
hispidulous above, the margins irregularly serrate; petioles 2.0-3.5 cm 


Phytologia (August 2009) 91(2) 313 


long. Capitulescence a terminal congested corymbose panicle of 
numerous heads, 3-6 cm high, 4-6 cm across, the ultimate peduncles 2- 
5 mm long. Heads ca 7 mm high. Involucres ca 4 mm long, 
composed of ca 11 slender, nearly glabrous, subequal bracts, their 
apices gradually attenuate. Receptacles ca 1 mm across, glabrous or 
nearly so. Florets 13-16 per head; corollas white, glabrous, ca 5S mm 
long, the lobes ca 0.5 mm long. Achenes 2.3-3.0 mm long, markedly 
hispidulous, especially along the ribs; pappus of ca 40 tawny-white 
bristles 4-5 mm long. 


TYPE: MEXICO. OAXACA: Mpio. Coixtlahuaca, "Concepcion 
Buena Vista. Km 94.7 de la carretera Tehuacan-Oaxaca (cuota) y de 
este puntoaproximademente 2 horas a pie montana arriba hasta base de 
paredes verticales en la cima de cerro." 1680 m, (18 06 58.5 N, 97 19 
47.1 W), 27 Oct 1996. Jose L.Panero & Ismael Calzada 6760 
(Holotype: TEX). 


The present novelty, in habit and leaf shape, resembles a 
species of Fleishmannia, but it clearly belongs to Koanophyllon, where 
it finds no clear relatives. Panero identified the type as K. gracilicaule, 
which it superficially resembles. 


The species is named for the Municipio Coixtlahuaca, from 
whence the type. 


In my account of Koanophyllon for Mexico (Turner 1997), I 
treated all of the latter within a broadly circumscribed Eupatorium. | 
now follow the treatment of King and Robinson (1987). Below find a 
modified key to the Mexican species of Koanophyllon, including the 
three novelties described herein. 


KOANOPHYLLUM CONCORDIANUM BLL. Turner, sp. nov., Fig. 
2., Map 1 


Koanophyllon reyrobinsonii B.L. Turner similes sed differt foliis 
ovalibis (vs ovatis vel deltoideis ad medium latissimis et flosculis per 
capitulum paucioribus (4-5 vs 7 vel plures). 


314 Phytologia (August 2009) 91(2) 


Perennial suffruticose herb or sprawling subshrub to 1 m (?) high. 
Stems densely pubescent with mostly upswept hairs. Leaves opposite, 
5-7 cm long, 3-5 cm wide; petioles 3-6 mm long; blades oval, widest 
near the middle, 3-nervate from the base, nearly glabrous and 
atomiferous-glandular below, glabrous above, the margins pubescent, 
crenulate. Capitulescence a terminal corymbose panicle ca 15 cm 
high, 6-10 cm across, the ultimate peduncles 1-6 mm long, pubescent 
like the stems. Involucres 3-4 mm high, composed of 5-6 subequal, 
glandular-atomiferous bracts. Receptacles ca 0.5 mm _ across, 
pubescent. Corollas white, atomiferous-glandular, ca 2.5 mm long; 
tube ca 1.5 mm long; lobes 5, obtuse, ca 0.2 mm long. Achenes black, 
5-ribbed, ca 1.5 mm long, appressed-pubescent; pappus of ca 30 
persistent bristles ca 5 mm long. 


TYPE: MEXICO. SINALOA: Mpio. de Concordia, "El Capomito, 
Ejido Los Ciruelos, Comunidad La Guasima," (23 18 10 N, 105 56 12 
W), tropical deciduous forest, 341 m, 7 Jan 2006, A.L. Reina G. et al. 
2006-122 (Holotype: TEX). 


ADDITIONAL SPECIMEN EXAMINED: MEXICO. SINALOA: 
Mpio. Concordia, “La Cuesta Blanca, la Bajada de Campo Redondo, 
Comunid La Guasima,” ca 13.2 km NE Concordia, 393 m, 25 Nov 
2008, Reina G. 2008-661 (TEX). 


This novelty is markedly distinct, having the pubescent 
receptacles of those taxa centering about K. Jongifolium and K. 
reyrobinsonii, but possessing the capitulescence and heads of K. 
palmeri, to which it is perhaps more closely related. 


The species is named for the Municipio de Concordia, from 
whence the type. 


KOANOPHYLLON REVEALII B.L. Turner, sp. nov., Fig. 3, Map 2 


Koanophyllon gracilicaule (Sch.-Bip. ex B.L. Rob.) King & H. Rob. 
similes sed differt capitulescentiis minoribus (6-12 mm altis 6-12 mm 
latis vs ca 20 cm altis 20 cm latis) pedunculis ultimis brevioibus (3-6 
mm longis vs 8-15 mm) et setis pappi numerous (ca 40 vs 20). 


Phytologia (August 2009) 91(2) 315 


Shrub or small tree 1-4 m high. Mid-stems purplish-brown, minutely 
hispidulous. Leaves opposite throughout; blades ovate to ovate- 
deltoid, 4-6 cm long, 2-4 cm wide, 3-nervate from the base, glandular- 
punctate, the margins crenulate; petioles 2-4 cm long. Capitulescence 
a terminal corymbose panicle, 6-12 cm high, 6-12 cm across, the 
ultimate peduncles hispidulous, mostly 3-6 mm long. Heads 
numerous, 6-7 mm high. Receptacles ca 1 mm across, glabrous or 
nearly so. Involucres 3-4 seriate, imbricate, 1-4 mm long, densely 
brown-hispidulous throughout, linear-lanceolate, their apices abruptly 
acute. Florets ca 15 per head. Corollas white, glabrous, ca 4 mm 
long, the lobes deltoid, ca 0.5 mm long. Achenes ca 2 mm long, 
subglabrous to sparsely hispidulous; pappus of ca 40 tawny persistent 
bristles 3-4 mm long. 


TYPE: MEXICO. GUERRERO: Mpio. Atoyac de Alvarez, "along 
the Millpillas-Atoyac road via Puerto del Gallo, about 48.5 miles 
northeast of Atoyac and 6.8 miles southwest of Puerto del Gallo, in 
mixed deciduous forest with scattered tree ferns on steep slopes," ca 
6800 ft, 19 Oct 1975, J.-L. Reveal, K.M. Peterson, R.M. Harley & C.R. 
Broome 4346 (Holotype: TEX). 


ADDITIONAL SPECIMENS EXAMINED: MEXICO: 
GUERRERO. Mpio. Atoyac de Alvarez, "below Puerto El Gallo 
along road to Atoyac." 2255 m, 10 Oct 1986, Breedlove 65118 (TEX); 
38.5 km NE El Paraiso, rumbo a filo de Caballo, 7 Sep 1983, Villasenor 
555 (TEX). 

OAXACA: Mpio. Santiago Juxtlahuaca, ca 5 km del 
poblado El Manzanal, 17 13 0.20 N, 38 04 33.7 W, 22 Aug 1996, 
Calzada 21163 (TEX). 


In my treatment of the Koanophyllon complex for Mexico 
(Turner 1997) I included the above collections within K. gracilicaule 
(to which it is clearly related). The latter is typified by material from 
Tlacolula, Oaxaca (GH!), first collected by Ehrenberg in 1839. 
Koanophyllon revealii differs from the latter in having much smaller 
capitulescences, with shorter ultimate peduncles, somewhat smaller 
heads, and pappus with more numerous bristles, as noted in the above 
diagnosis. Distribution of the two taxa is shown in Fig.4. 


316 Phytologia (August 2009) 91(2) 


The species is named for James L Reveal, Systematist 
extraodinare, still kicking up nomenclatural novelties and academic 
miscellany at the age of eighty. Bravo! May he dance on. 


KOANOPHYLLON SOLIDAGINOIDES (H.B.K.) King & H. Rob., 
Phytologia 22: 151. 1972. Map 3 
Eupatorium solidaginoides H.B.K. 


Weak-stemmed, arching or clambering, shrubs 1-3 m high; 
stems striate, densely puberulent; leaves 6-10 cm long, 2-5 cm wide; 
petioles mostly 1.5-4.0 cm long; blades deltoid to decidedly cordate, 
3(5)-nervate from at or near the base, densely minutely glandular- 
punctate beneath, glabrous except along the major veins, the margins 
crenulate to dentate; heads white, numerous in both terminal and 
axillary, loose or congested, corymbose racemes, the ultimate 
peduncles 2-10 mm long; florets 10-15 per head; achenes ca 2 mm 
long, the pappus of 40-50 bristles 2.5-3.0 mm long. 


A widespread, highly variable, species but readily 
distinguished by its weak clambering stems and cordate leaves (rarely 


deltoid). 


Two varieties are recognized in the complex for Mexico, as 
follows: 


Ultimate peduncles 3-7 mm long; heads 5-7 mm high; eastern 


MICKISON os susdetieae ds ahah. etter cea ARS var. filicaulis 
Ultimate peduncles 1-3 mm long; heads 4-5 mm high; western 
REISS ot rare hd Soe Lak a revit chase aaNet ga eta var. solidaginoides 


var. filicaulis (A. Gray) B.L. Turner, comb. & stat. nov. Map 3. 
Koanophyllon solidaginoides var. filicaulis (Sch.-Bip. ex A. Gray)B.L. 
Turner, comb. & stat. nov. Based upon Eupatorium filicaule Sch.-Bip 
ex A.Gray, Proc. Amer. Acad. Arts 21: 384. 1886. 


San, Ver, Oax, Cps and Guatemala southwards, in barrancas of 
montane cloud forests 20-2600 m; Nov-Feb. 


Phytologia (August 2009) 91(2) 317 


In Mexico, the two varieties are quite distinct; in Central 
America, however, they appear to intergrade, especially in northern 
Guatemala (numerous specimens in and about Tikal, LL-TEX), hence 
my treatment of these at the varietal level. 


var. solidaginoides Map 3. 


The type of this taxon is from Ecuador. In Mexico, it is 
known only from Chiapas, the latter populations easily recognized from 
the typical var. by its much shorter ultimate peduncles and smaller 
heads. Eupatorium  solidaginoides’ var. armourii B.L. Rob. 
(photoholotype FM!) from Palenque, Chiapas appears to be a form of 
this taxon having markedly deltoid leaves and somewhat larger heads. 
Additional field studies might show the name concerned worthy of 
recognition. 


Key to Mexican species of Koanophyllon 


1. Leaves 3-parted or trifoliolate on mid-stems (a few leaves simple 


POM SIE MPPEMSLEINS i. 2 d <ee-is .wxesdeewseeabeoeee esses K. tripartitum 
Pee eee eM SII IS cia Hoa «dsodwigu aasueomembaniiens Savane delnaagtace. Somaeeaes (2) 

Pemkcaves pinnately veined wecc.evceiniev sees oeideca sess nase K. pittieri 

2. Leaves with 3-5 principal veins from, or near, the base......... (3) 


3. Heads arranged in ball-like clusters, the involucres with only | or 2 
ih DRE GG A I RR RR 2 Cats ak ale i wet Re K. monanthum 
3. Heads not as above, the involucres with 4 or more florets 


4. Pappus about 0.5 mm long or less; heads in congested, spike- 
like, capitulescences, the ultimate peduncles mostly 0-1 mm 
LCE GCS Any Ale nae aR aA A ee ew ge K. ravenii 

4. Pappus 2-7 mm long; heads in mostly open or loosely arranged 
capitulescences, the ultimate peduncles mostly 2-10 mm 


318 Phytologia (August 2009) 91(2) 


\o 


] 


—" 


] 


— 


13, 


. Leaves with petioles 1-4 mm long, the blades appearing to clasp; 


Webcal K. pseudoperfoliatum 
. Leaves with petioles 3-40 mm long, the blades not appearing to 
CLASP i. . ssc+cesncdesaccorks: sexed spicsdcbusigeeinne aha RS- serene ce) eee (6) 
6:.. Petioles mostly 12-40 mm long... ...2.. ..22.sncuses-00tonesepeeeee (7) 
6.. Petioles, mostly 2-15(20) mmilong. .... «<5 ..0....00000eeeeee (14) 
Heads 8-10 mm high; achenes densely pubescent with soft 
dppressed hairs: Baja\Cale. ooo too. ce eeenmensensers K. peninsularis 
Heads 4-7 mm high; achenes glandular or sparsely hispid; not in 
Praja (Cd oc nk ace a ar detain sae Ont nieces inieae aaa (8) 
S." Heads 42S ain Wie 22 022.6. o aes de oee oo go ee (12) 
S. Meads'G=6 imi lien. 55% 00 io..y deecs cece seus + 2 = eee (9) 
. Receptacles glabrous or nearly so; Gue, Oax, Cps................ (11) 
| Receptacles pubescent: Tait to Hid)... 2... sets <as0nce-eeeeeee (10) 
10. Leaves cordate; achenes hispid; Nue............ K. hintoniorum 
10. Leaves ovate, obtuse or truncate at base; achenes to some 
deeree plandular-pubesctent.......5..5.... 42008 K. richardsonii 


.Involucral bracts densely pubescent, abruptly 


PE 8c = coeds Sesto bole ea ee ee K. revealii 


.Involucral bracts sparsely pubescent, if at all, gradually 


attenuates cent wee athe. Cowen ae eee K. coaxtlahuacum 


12.Capitulescence of axillary or terminal spike-like branches, the 
heads arranged in ae. or loose interrupted corymbs 
psscoee ..K. solidaginoides 
12: G@apitulescence nee as ee foauael aa fee in pyramidal 
COPYEND OSE PAN CN ES inp Suey ital seis istians hms eos eee (13) 


.Blades of leaf pubescent above and below, (rarely glabrate); 


corollas 2.5-3.3 mm long; involucral bracts sharply acute; 
CSS: ST ec ora ak censor eee oS K. sinaloense 
Blades of leaf glabrous or nearly so; corollas ca 3.5 mm long; 
involucral bracts broadly acute; San, Ver, Gue, Oax 

eiscas PON easativae te Staias AAMRe eee eno eee nate get K. gracilicaule 


Phytologia (August 2009) 91(2) 319 


Pete SecOIACIe SIADTOUS cast eedes. Uses sceavereueresseaaceves geaveee (17) 
14. Receptacle decidedly pubescent, rarely not; Sin, Nue, Tam, San, 
RIPEN eer ree ear aty eons ts chia ncevee tons cwurangdemseessas'os (15) 
15. Leaves oval, widest near the middle; florets 4-5 per head; 
MMi at nao srccke Saks tetesene seeared Lee Pesos cee K. concordianum 
15. Leaves ovate to triangular, widest near the base; florets 6-15 
BEM Sas ae me aint stile wie tt fas He eaNe | eee db wan 2 Wa webe tages (16) 


16 . Blades of leaf uniformly pubescent beneath; petioles mostly 7-15 
man long: corollas ca 2:mm long..............0...+... K. longifolium 
16. Blades of leaf sparsely pubescent along the major veins; petioles 
mostly 3-6(8) mm long; corollas 2.5-3.0 mm long 
Ee co cas ecw ete Setsomicns ome Aeaao ee MES eM RETe URE K. reyrobinsonii 
17. Suffruticose herbs or arching, weak, shrubs, 0.5-3.0 m high; 
older stems not white; foliage to some degree pubescent.....(19) 
17. Tree-like shrubs, small trees or woody vines, 2-10 m high; older 
Semis white: foliage PIADTOUS /.5...550i0.50.0d<50 cers. va oteb tues (18) 


18. Woody vines; capitulescence pyramidal, axillary; leaf blades 
emi. 15-20 em long: Gues.. .o.i05 << s2ccs Seow K. guerreroanum 

18. Shrubs or trees; capitulescence corymbose or pyramidal, terminal; 
leaf blades ovate-elliptic, 6-15 mm long; widespread 

Re 258 ahd Ju ick Seats Coa be ngs ewes Save eenemboneveamene K. albicaule 

19. Leaf blades densely pubescent above and beneath; San 
MEAs eh vo Hew seve Moen meme ds dh eae sean astedy K. rzedowskii 


19. Leaf blades not densely pubescent except along the veins 


Benieain: Widespread: <..002 5 a..tdet sonst ansmntes cates eneatneenes (20) 

20. Leaf blades variously lanceolate, ovate, or deltoid, but not clearly 
ee ete: SUN COTGALG: fn Bsc,y cece tape ashes Ge roe sc tases dens sdageeunend (22) 
20, eat blades neatly cordate or subcordate....... 2c o.5...dennecsscnes (21) 


21. Involucral bracts rigid with narrowly acute (acicular) apices, 
fot at allsearmus; NGS...) 5. 2..ccb. Stet Soh aceee K. galeanum 

21. Involucral bracts not rigid, scarious marginally, the apices 
obtuse or rounded; widespread but not in Nue 

K. solidaginoides 


ee ee ey 


320 Phytologia (August 2009) 91(2) 


22. Leaves often alternate above, the blades epunctate beneath; e Chi, 
Bear. Za .cushiceace ext dese: aeeteee ewe see K. solidaginifolium 

22. Leaves opposite throughout, the blades glandular-punctate beneath; 
Sonyw ‘Chi; Dur, Jal, Col; Mie wijscijesccncsmssctincss ewsont see eee (23) 


23 Jneadsa 9 mim hirel@pss..V Ais este sosiciewnn-aayeaerdeae K. coulteri 
23. Heads 4-6 mm ise w Chi, Sin, Dur, ae Jal, Col, Mic, Gue 
esata lee meat eniawalecece Meas va Ete Oa deactuets exreemaae soe: ..K. palmeri 
ACKNOWLEDGEMENTS 


As always, I am grateful to my long-time colleague, Guy 
Nesom, for the Latin diagnoses and helpful editorial comments. The 
Field Museum provided selected types, for which I am grateful. 


LITERATURE CITED 


King, R.M. and H. Robinson. 1987. The genera of the Eupatorieae 
(Asteraceae). Monographs in Syst. Bot., Missouri Bot. Gard. 
22; 1-581. 

Turner, B.L. 1997. Koanophyllon, in The Comps of Mexico. Phytologia 
Memoirs 11:1 10-111. 


KOANOPHYLLON 
© coixtlahuacum 
@ concordianum 


Map 1 


Map 1. Distributions of K. coixthlahuacum and K. concordianum. 


Phytologia (August 2009) 91(2) 321 


KOANOPHYLLON 
@ revealii 
© gracilicaule 


Map 2 


KOANOPHYLLON 
solidaginoides 
© var. filicaulis 
®@ var. solidaginoides 


Map 3. Distributions of K. solidaginoides var. filicaulis and var. 
solidaginoides. 


322 


UNIVERSITY OF 
amy, FEXAS 


HERBARIUM 


Phytologia (August 2009) 91(2) 


wan Hetero (DEMS 
Koanoptiy thon coitiahuacun BL. Turner 


bes, BY. Twas WO 


PLANTAS DE MEXICO 


Won minyten them: atin hoe bane 

Ba Towa 
Koamaphytton sracthicawte Mi Bip. ox B 
1 Rob. Mi Kaa de bh Bot 


OARNAG A: Coieeriabosanein 6 


Fig. 1. Holotype of Koanophyllon coixtlahuacum. 


Phytologia (August 2009) 91(2) 323 


HERBAR 


Ken epta ion cen cotdie noes 


ee 


AT 008 4 


PLA OF StKLoR, KERLOS 


Fig. 2. Holotype of Koanophyllon concordianum. 


324 Phytologia (August 2009) 91(2) 


UNIVERSITY OF 


1 


HERBARIUM 


7 


Kemme phy ten weve ali, Ra Tonue 


Fig. 3. Holotype of Koanophyllon revealii. 


Phytologia (August 2009) 91(2) 325 


NOTES ON NON-NATIVE ASTERACEAE IN TEXAS 


Guy Nesom 
2925 Hartwood Drive 
Fort Worth, Texas 76109 
www.guynesom.com 


ABSTRACT 


Documentation and/or commentary is provided for 16 of the 
73 species of non-native Asteraceae reported to occur outside of 
cultivation in Texas. Arctium minus, Logfia (Filago) arvensis, Madia 
elegans, Matricaria courrantiana, Sanvitalia procumbens, and Sonchus 
arvensis are removed from the current account of plants naturalized in 
Texas. Carduus tenuiflorus 1s more appropriately and consistently 
identified as C. pycnocephalus. Phytologia 91(2): 325-332 (August, 
2009). 


KEY WORDS: Texas, flora, adventive species 


73 species of non-native Asteraceae have been recorded as 
occuring outside of cultivation in Texas (Nesom 2009). About 37 of 
these were reported after publication of the Manual of the Vascular 
Plants of Texas (Correll & Johnston 1970). The present report provides 
taxonomic clarification and voucher information for a number of the 
latter, where ambiguity existed or documentation was lacking. 


1. Arctium minus Bernh. 

Hultén and Fries (1986, Vol. 2, Map 1853) mapped the 
species for Texas, at the southern extremity of its adventive range in 
North America, placing a dot near the center of the state. Its occurrence 
in Texas may be expected, as it is documented to occur in New Mexico, 
Oklahoma, Arkansas, and Louisiana (USDA, NRCS 2009), but no 
voucher or other documentation has been located for its occurrence in 
Texas (Keil 2006a) and the species is removed from the current account 
of Texas plants. 


326 Phytologia (August 2009) 91(2) 


2. Carduus acanthoides L. 

Listed by Jones et al. (1997); mapped by Turner et al. (2003) 
for Collin and Schleicher counties. Vouchers are at TEX. 

Collin Co.: Farm Rd 98ljust E of jct with Hwy 78, 
bottomland forest, 13 May 1992, Saunders 3418 (TEX). Schleicher 
Co.: Westernmost portion of Eldorado, weed along Hwy 190, 8 Jun 
2001, Turner 21-770 (TEX). 


3. Carduus pycnocephalus L. 

Identification and taxonomy of Carduus pycnocephalus must 
be considered in conjunction with that of the very similar C. tenuiflorus 
Curtis. Cory (1940) collected plants of C. pycnocephalus in Sutton Co. 
in 1939, in pastures and fields growing with plants of Silybum 
marianum (L.) Gaertn. and Carduus nutans L., all three of which 
apparently arrived there in a shipment of hay from California. In a later 
collection from Sutton Co. (Cory 53662, 5 May 1947, SMU), he had 
changed the identification to C. tenuiflorus. Carduus tenuiflorus, as it 
has been subsequently identified in Texas, now has spread over a large 
region of central Texas (Turner et al. 2003). 

Dunn (1976) identified the Texas plants as Carduus 
pycnocephalus, and Keil (2006b) included Texas among the states in 
the known range of C. pycnocephalus (“Aggressive weed of waste 
ground, rangelands, pastures, roadsides, fields; Ala., Ark., Calif., Idaho, 
Miss., N.Y., Oreg., Pa., S.C., Tex.”). He including only California and 
Oregon in the North American range of C. tenuiflorus, noting that it has 
been reported from New Jersey, Texas, and Washington but that he had 
not seen specimens from those states. 

Comments on the subtle and quantitative distinctions between 
Carduus pycnocephalus and C. tenuiflorus were provided by Keil 
(2006b). Keil also noted that “Published chromosome counts (2n = 18, 
31, 32, 54, 60, 64, 80) for C. pycnocephalus from a variety of Old 
World localities indicate that this is a complex species in need of 
further investigation. ... The two species sometimes grow in mixed 
populations and at times appear to intergrade. Hybridization has been 
reported in Europe ... and is suspected to occur in California.” The two 
also have been treated as conspecific, the younger name becoming C. 
pycnocephalus L. var. tenuiflorus (Curtis) Fiori. 

It seems clear that only a single evolutionary entity is 
represented among the Texas plants, whether from the initial 


Phytologia (August 2009) 91(2) 327 


introduction in Sutton Co. or from multiple introductions. The 
morphology and variability in Texas are similar to that characteristic of 
California plants of Carduus pycnocephalus as well as all or almost all 
California collections at BRIT and TEX identified as C. tenuiflorus. 
The Texas plants are more appropriately and consistently identified as 
Carduus pycnocephalus rather than C. tenuiflorus. 


4. Carthamus tinctorius L. 

Mapped by Turner et al. (2003) for Brazos and Tarrant 
counties, evidently based on the following collections. 

Brazos Co.: College Station, 8 Apr 2006, Reed & Reed 3086 
(TAMU). Ellis Co.: Waxahatchie, in yard near a house along FM 55, 
not in vicinity of a bird feeder, ca. 7 plants in area 15 feet long, 11 Jun 
2002, Hill et al. s.n. (BRIT). Tarrant Co.: Fort Worth, weed in yard, 
probably from bird seed, 19 Jun 1996, Sylvester 2414 (BRIT). 

Carthamus lanatus L. is known in Texas from Gillespie, 
McCulloch, Kimble, Schleicher, and Coryell counties (collections at 
BRIT, TEX), and Llano, Menard, and Tom Green counties (as 
additionally mapped by Turner et al. 2003), where it has been found in 
pastures and gravel pits and along roadsides. 


5. Centaurea benedicta (L.) L. (= Cnicus benedictus L.) 

Listed by Jones et al. (1997) and included for Texas by Keil 
and Ochsmann (2006). The Texas record is based on the following 
(fide Steve Hatch at TAES): Callahan Co.: “grown in a pot from a 
wheat field,” Mar 1993, J. Landers s.n. (TAES). The plant apparently 
was grown to identify a weed submitted for identification. 


6. Cotula australis (Sieber ex Spreng.) Hook. f. 

Recorded by Johnston (1990) based on plants from the Texas 
A&M campus, as documented here by collections at TEX; the report by 
Hatch et al. (1990) probably based on the same or similar collections. 

Brazos Co.: College Station, campus of A&M University, 
lawn weed with Soliva, 27 Apr 1970, Correll & Correll 38493 (LL, 
TEX); Texas A&M University campus, an Australian weed widely 
adventive in W hemisphere, the first population known in Texas, 23 
Apr 1970, Johnston s.n. (TEX). 


328 Phytologia (August 2009) 91(2) 


7. Leontodon hispidus L. 

Mapped by Turner et al. (2003) for Panola Co. An apparently 
corresponding voucher at TEX is from adjacent Harrison Co.: 
Marshall, SE corner of yard at Grover Hall, East Texas Baptist College, 
16 Jun 1983, Fleetwood 6 (TEX). 


8. Leontodon taraxacoides (Vill.) Mérat 

Vuilleumier (1973) included “Texas” in the statement of range 
for the species; mapped by Turner et al. (2003) from El Paso Co. (as 
Leontodon “taraxacum’’). A voucher at TEX is from El Paso Co.: El 
Paso, 7400 block of N Mesa, in watered lawn, 12 Aug 1993, 
Worthington 22279 (TEX). 


9. Logfia (Filago) arvensis (L.) Holub 

This species has been included by the PLANTS Database 
(USDA, NRCS 2009) in the Texas flora based on Johnston (1990). 
Johnston, however, noted only that his description of Stvlocline 
micropoides Gray in the Texas Manual (Correll & Johnston 1970) was 
“wholly inappropriate, [and] instead applies to Logfia arvensis.” The 
real Stylocline micropoides, which is a native species, is vouchered 
from El Paso. El Paso Co.: NW part of El Paso at small andesite hills 
0.3 mi. N of jct Belvedere w/ Westwind, 4200 ft., mostly a creosote 
bush community with several substrate types and arroyos, 29 Apr 1983, 
Worthington 10130 (TEX, UTEP). The real Logfia arvensis has not 
been observed in Texas (Worthington 1989, 1997), and the species is 
removed from the current account of Texas plants. 


10. Madia elegans D. Don ex Lindl. 

Turner et al. (2003) mapped the species for Grimes County, 
evidently based on early collections from Plantersville, a town founded 
in the 1830’s and whose population has fluctuated around 200 since 
about 1900. The Atchison, Topeka and Santa Fe Railway runs through 
the town and might have been a source of the seeds, but the label data 
do not indicate that the plants were naturalized or waifed in 
Plantersville, and it seems most likely that they were garden 
collections. Madia elegans has not been reported from anywhere else 
outside its native range in California, Nevada, Oregon, and 
Washington, and the species is removed from the current account of 
Texas plants. 


Phytologia (August 2009) 91(2) 329 


Grimes Co.: Plantersville, Fall 1927, Bletsch 42-77 (TEX); 
Plantersville, Fall 1927, Bletsch s.n. (TEX). 


11. Matricaria courrantiana DC. 

Kearney and Peebles (1960, p. 937) included “Texas” in the 
description of geographic range for the species, and Arizona and Texas 
remain the only two states recorded for the occurrence of this species in 
the USA (fide PLANTS). Although the observation by Kearney and 
Peebles surely was based on real evidence, until a voucher for the 
Texas record can be located, the species is removed from the current 
account of Texas plants. 


12. Sanvitalia angustifolia A. Gray 

Recorded by Johnston (1990), with the following comment: 
“Found once in the campgrounds of the Chisos Mountains basin, 
perhaps waifed through automobile dispersal from its native home in 
Mexico.” Based on a TEX collection. 

Brewster Co.: Big Bend National Park, Chisos Mts. Basin 
Campground, at the first campsite on the rt. after the paypost, only two 
plants seen, 13 Oct 1978, Henrickson & Bontemps 12444 (TEX). 


13. Sanvitalia procumbens Lam. 

Noted by Strother (1979) to occur “in trans-Pecos Texas in 
Chisos Basin (J. Henrickson pers. comm.),” with further comment 
(Strother 2006, p. 71): “Sanvitalia procumbens has been seen only 
sporadically in the flora; it probably is not a resident.” Strother (pers. 
comm.) comments that for the FNANM treatment, he “saw no 
supporting specimen(s) for Sanvitalia procumbens from California or 
Texas,” and it is highly probable that Henrickson’s communication was 
intended to refer to S. angustifolia. Sanvitalia procumbens 1s removed 
from the current account of Texas plants. 


14. Scorzonera laciniata L. 

Recorded by Johnston (1990) to occur in Randall Co., without 
other documentation. Collections of the species have been made from 
three Panhandle counties. 

Hutchinson Co.: LAMR, Harbor Bay area, directly W of Fritch 
on Harbor Bay road, roadside and steep red-sandy slopes with dolomite 
boulders from caprock, 3190 ft, 22 Apr 2002, Nesom & O’Kennon 


330 Phytologia (August 2009) 91(2) 


LAMR 16 (BRIT); LAMR, lawn and roadside of NPS Ranger Station, 
just W of Sanford-Yake road on short spur, near jct of High Plains 
Road, 25 Apr 2002, Nesom & O’Kennon LAMR110 (BRIT); W side of 
Borger along Hwy 136, grassy areas bordering Wal-Mart parking lot, 
28 May 2002, Nesom & O’Kennon LAMR243 (BRIT). Lubbock Co.: 
Canyon Rd, lower slopes and along the bottom of a “lone profile” 
canyon, 9 Apr 1990, Rose s.n. (LL). Randall Co.: Camp Don 
Herrington Boy Scout Camp on Palo Duro Creek, mixed shrub- 
grassland community, 24 Apr 1985, Higgins 1527] (TEX); Canyon, E 
side of town on vacant lot, 29 May 1988, Worthington 16584 (TEX). 


15. Solivia anthemifolia (Juss.) Sweet 

Turner et al. (2003) mapped Soliva anthemifolia for Bastrop 
Co. and Leon Co., separately from “Soliva mutisii,’ which is shown 
with a considerably broader range. Soliva mutisii Kunth, however, 1s a 
synonym for S. anthemifolia (Watson 2006), and in the Flora of Texas 
Database (http://www.biosci.utexas.edu/prce/Tex.html), identifications 
of TEX/LL vouchers for these records have been corrected to S. 
stolonifera (Brot.) Sweet and S. anthemifolia (Juss.) Sweet, 
respectively. Nueces Co. is added to the two other counties for the 
distribution of S. stolonifera. Soliva sessilis Ruiz & Pavoén (= S. 
pterosperma [Juss.] Less.) also is commonly and widely naturalized in 
east Texas. 


16. Sonchus arvensis L. 

Hatch et al. (1990) noted that a previous claim that this species 
occurs in Texas has not been confirmed. Jones et al. (1997) also listed 
it, but no voucher has been located and Texas was not included in the 
range of the species in the FNANM treatment (Hyatt 2006). The 
species is removed from the current account of Texas plants. 


ACKNOWLEDGEMENTS 


I am grateful for helpful comments from Monique Dubrule 
Reed, Steve Hatch, David Keil, John Strother, and Richard 
Worthington and to staff at SMU/BRIT and TEX/LL for their 
hospitality during studies there. 


Phytologia (August 2009) 91(2) 33] 


LITERATURE CITED 


Correll, D.S. and M.C. Johnston. 1970. Manual of the Vascular Plants 
of Texas. Texas Research Foundation, Renner, Texas. 


Cory, V.L. 1940. Six thistles recently introduced into Texas. Madrofio 
5: 200-202. 


Dunn, P.H. 1976. Distribution of Carduus nutans, Carduus 
acanthoides, Carduus pycnocephalus, and Carduus crispus in the 
USA. Weed Sci. 24: 518-24. 


Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1990. Checklist of the 
Vascular Plants of Texas. MP-1665, Texas Agric. Expt. Station, 
Texas A&M Univ., College Station. 


Hultén, E. and M. Fries. 1986. Atlas of North European vascular 
plants north of the Tropic of Cancer. Vols. 1, 2, and 3. Koeltz, 
K6nigstein, Federal Republic of Germany. 


Hyatt, P.E. 2006. Sonchus (Asteraceae: Cichorieae). Flora of North 
America North of Mexico 19: 273-276. 


Johnston, M.C. 1990. The vascular plants of Texas: A list, up-dating 
the “Manual of the Vascular Plants of Texas” (ed. 2). Published by 
the author. 


Jones, S.D., J.K. Wipff, and P.M. Montgomery. 1997. Vascular plants 
of Texas: A comprehensive checklist including synonymy, 
bibliography, and index. Univ. of Texas Press, Austin. 


Kearney, T.H. and R.H. Peebles. 1960. The Flora of Arizona, 2nd ed. 
with supplement by J.T. Howell, E. McClintock, and collaborators. 
Univ. of California Press, Berkeley. 


Keil, D.J. 2006a. Arctium (Asteraceae: Cynareae). Flora of North 
America North of Mexico 19: 168-171. 


332 Phytologia (August 2009) 91(2) 


Keil, D.J. 2006b. Carduus (Asteraceae: Cynareae). Flora of North 
America North of Mexico 19: 91-94. 


Keil, D.J. and J. Ochsmann. 2006. Centaurea (Asteraceae: Cynareae). 
Flora of North America North of Mexico 19: 181-194. 


Nesom, G.L. 2009. Literature Reviews of the Texas Flora: Non-native 
species recently discovered to be naturalized in the state; Non- 
native species in the state, complete list; Native species new to 
science for the state. <http://guynesom.com/Texasplantsweb.htm> 


Strother, J.L. 1979. Extradition of Sanvitalia tenuis to Zinnia 
(Compositae-Heliantheae). Madrofio 26: 173-179. 


Strother, J.L. 2006. Sanvitalia. (Asteraceae: Heliantheae). Flora of 
North America North of Mexico 21: 70-71 


Turner, B.L., H. Nichols, G. Denny and O. Doron. 2003. Atlas of the 
Vascular Plants of Texas. Vol. I—Dicots; Vol. Il—Monocots. 
Sida, Bot. Misc. 24, | and 2. 


USDA, NRCS. 2009. The PLANTS Database. National Plant Data 
Center, Baton Rouge, La. 
<http://plants.usda.gov> 


Vuilleumier, B.S. 1973. The genera of Lactuceae (Compositae) in the 
southeastern United States. J. Arnold Arb. 54: 42-93. 


Watson, L.E. Soliva (Asteraceae: Anthemideae). Flora of North 
America North of Mexico 19: 545-546 


Worthington, R.D. 1989. An annotated checklist of the native and 
naturalized flora of El Paso County, Texas. El Paso Southwest 
Bot. Miscell. No. 1. [581.974 T355WO 1989] 


Worthington, R.D. 1997. Nomenclatural changes, additions, and 
corrections to an annotated checklist of the native and naturalized 
flora of El Paso County, Texas. El Paso Southwest Bot. Miscell. 2: 
1-27. 


Phytologia (August 2009) 91(2) 355 


THYMOPHYLLA TENUILOBA AND T. WRIGHTII 
(ASTERACEAE: TAGETEAE) 


Guy L. Nesom 
2925 Hartwood Drive 
Fort Worth, TX 76109 
www.guynesom.com 


ABSTRACT 


Thymophylla tenuiloba has generally been treated to include 
four varieties. Among these, 7. tenuiloba var. wrightii (A. Gray) 
Strother is distinct in morphology and sympatric and non-intergrading 
with 7. tenuiloba var. tenuiloba. It is appropriately treated at specific 
rank as 7. wrightii (A. Gray) Small. Thymophylla tenuiloba var. 
treculii and var. texana differ from var. tenuiloba in minor features of 
pappus morphology and populational variation occurs in the same 
features; each 1s geographically distinct, however, and these three taxa 
are maintained at varietal rank. Phytologia 91(2): 333-339 (August, 
2009). 


KEY WORDS: Thymophylla tenuiloba, Thymophylla wrightii, 
Asteraceae, Tageteae, Texas 


In Johnston’s taxonomic overview of Texas Dyssodia (1956), 
he noted that D. tenuiloba (DC.) B.L. Rob., D. wrightii (A. Gray) B.L. 
Rob., D. texana Cory, and D. treculii (A. Gray) B.L. Rob. “are more 
closely related to each other than to other species. In details of 
involucre they are nearly identical; they differ in pappus-form, and to 
some extent in habit.” Strother (1969) emphasized the similarities 
among these four taxa by combining them as varieties of a single 
species, D. tenuiloba. And so they have been treated since that time 
(Strother 1970, 2006), except for Turner (1996) and Turner et al. 
(2003), who combined var. treculii and var. texana with var. tenuiloba, 
simplifying the species to var. tenuiloba and var. wrightii. Robinson’s 
treatment (1913) of D. wrightii and D. treculii at specific rank reflected 
his general transfer of names from Hymenatherum Cass. to Dyssodia 
Cav. rather than a refinement of species concepts. Strother’s transfer 


334 Phytologia (August 2009) 91(2) 


(1986) of all these taxa to Thymophylla Lag. has been supported by 
molecular evidence (Loockerman et al. 2003). 


In contrast to the generally accepted taxonomy, there is good 
evidence to treat var. wrightii at specific rank. As documented by 
Strother (1969) and as confirmed here, var. tenuiloba and var. wrightii 
are sympatric (Fig. 1), and I find no evidence of hybridization where 
the two occur together. The two taxa also differ slightly in pappus 
morphology (see key below) and they are consistently different in leaf 
morphology. Var. wrightii has entire, mostly linear leaves while var. 
tenuiloba has pinnatisect leaves, and there is no indication that toothing 
or lobing appearing rarely on proximal leaves of var. wrightii results 
from gene flow from var. tenuiloba. 


The two have been collected at the same site: Karnes Co.: 12 
mi S of [Wilson] county line on Texas Hwy 80, 15 Apr 1965, Strother 
137 (TEX)—Thymophylla_ wrightii and Strother 138 (TEX)—T. 
tenuiloba. Refugio Co.: 18 mi S of Woodsboro, 10 Apr 1965, Strother 
128 (TEX)—Thymophylla_ wrightii and Strother 129 (TEX)—T. 
tenuiloba. At both localities, Strother identified the plants as different 
species when he made the collections; he later treated the two taxa at 
varietal rank. The two also have been collected at very close though 
not identical localities within their area of sympatry (e.g., Bee Co., San 
Patricio Co.; Fig. 1). 


Chromosome number reports for Thymophylla wrightii all 
have been diploid, 2 = 16 (Strother 1989). Plants of 7. tenuiloba var. 
tenuiloba may be diploid, triploid, or tetraploid (2n = 16, 24, or 32), 
and populations may include a single ploidy level or mixtures of two or 
three ploidy levels. One population of var. tenuiloba from Webb Co. 
was observed to include diploids, triploids, and pentaploids. In the 
region of sympatry with T. wrightii, populations of T. tenuiloba are 
diploid or triploid or a mixture of the two levels (Strother 1989, Fig. 2). 
Circumstantial evidence indicates that triploids and perhaps other 
polyploids produce seeds apomictically. Knowledge of this aspect of 
biology strengthens the observation that 7. wrightii and T. tenuiloba are 
genetically isolated where they occur in sympatry. 


Phytologia (August 2009) 91(2) 335 


Status of Thymophylla tenuiloba var. texana and var. treculii. 
In contrast to Thymophylla wrightii, T. tenuiloba var. treculii 


and 7. tenuiloba var. texana differ from typical 7. tenuiloba only in 
minor features of pappus morphology, and intergradation and 
populational variation occurs in the same features. Var. treculii and 
var. fexana are geographically distinct (Figs. 1 and 2), however, and are 
appropriately treated at infraspecific rank within 7. tenuiloba. Limited 
sampling indicates var. fexana to be diploid, var. treculii to be 
tetraploid and pentaploid (Strother 1989), 


Almost all collections of var. treculii in Texas have been made 
very close to the Rio Grande, where it occurs in close sympatry with 
var. tenuiloba. In its broader range in Coahuila and Nuevo Leon, var. 
treculii occurs alone, thus the area of sympatry along the Rio Grande is 
where the ranges of var. treculii and var. tenuiloba meet (Strother 1969, 
Fig. 19). Mixed populations and intergrades appear to be common in 
the area of sympatry, although most plants display one or the other of 
the pappus expressions. Northern outliers of var. treculii in Crockett, 
Sutton, and Uvalde counties may be relatively recent adventives 
dispersed along roadways, as hypothesized by Strother (1989) for var. 
tenuiloba. 


Var. fexana is rare in Texas and restricted to a few west- 
central counties, far disjunct from Mexican populations in referable to 
this entity (Strother 1969, Fig. 20). It apparently does not intergrade 
with other expressions of Thymophylla tenuiloba in Texas, but in 
Coahuila, plants technically referable to var. fexana do apparently 
intergrade with var. treculii. 


Taxonomic overview. 
Morphological criteria for recognizing these taxa, as in the key 
below, are similar to those of Strother (2006). 


1. Plants erect to ascending; leaves relatively lax, entire, oblong-linear 
to filiform, rarely those on the proximal 1/6—1/2 of stem with 2-4 pairs 
of linear teeth or lobes; pappus of 10—12 unequal pales 2-3 mm long, 
each terminating in a single bristle-like awn or (less commonly) pales 
of outer series bifid and terminating in a single bristle-like awn; Texas. 


336 Phytologia (August 2009) 91(2) 


Thymophylla wrightii 


1. Plants usually diffusely spreading to decumbent, sometimes erect; 
leaves rigid, pinnatisect into 7-11 subulate, filiform divisions; pappus 
vatahle Woh tee... enter ce RE Thymophylla tenuiloba 


2a. Pappus of 10 pales, each 3—S-awned; Texas and Mexico 
(Famanlipasyineis.ct Meas. Sabo Gehl aekek T. tenuiloba var. tenuiloba 


2b. Pappus of 10 pales in two series of 5 each, those of the inner series 
2.5—-3 mm long and each 1-awned from the middle of the often bifid 
apex, those of the outer series 0.8—1 mm long and awnless; Texas and 
Mexico (Coahuila, Nuevo Leon, Tamaulipas).T. tenuiloba var. treculii 


2c. Pappus of 10 pales of subequal length in two series of 5 each, all 
awnless, or the inner 5 slightly longer and occasionally | or 2 of them 
1-awned; Texas and Mexico (Coahuila)......... T. tenuiloba var. texana 


1. Thymophylla wrightii (A. Gray) Small, Fl. S.E. U.S., 1295, 1341. 
1903. Hymenatherum wrightii A. Gray, Mem. Amer. Acad. Arts, 
n.s, 4(1): 89. 1849. Dyssodia wrightii (A. Gray) B.L. Rob., Proc. 
Amer. Acad. Arts 49: 508. 1913. Dyssodia tenuiloba var. wrightii 
(A. Gray) Strother, Univ. Calif. Publ. Bot. 48: 76. 1969. 
Thymophylla tenuiloba var. wrightii (A. Gray) Strother, Sida 11: 
378. 1986. TyPE: USA. Texas. In dry post oak woods between 
the Rio Colorado and the Rio Guadalupe, C. Wright s.n. (holotype: 
GH). 


2. Thymophylla tenuiloba (DC.) Small, Fl. S.E. U.S., 1295, 1341. 
1903. Hymenatherum tenuilobum DC., Prodr. 5: 642. 1836. 
Dyssodia tenuiloba (DC.) B.L. Rob., Proc. Amer. Acad. Arts 49: 
508. 1913. Type: USA. Texas. “In Mexico circa Bejar,” [between 
Laredo and San Antonio], Berlandier 2063 (holotype: G-DC; 
isotype: GH). 


2a. Thymophylla tenuiloba var. tenuiloba 


Phytologia (August 2009) 91(2) 5oT 


2b. Thymophylla tenuiloba var. texana (Cory) Strother, Sida 11: 378. 
1986. Dyssodia texana Cory, Rhodora 49: 162. 1947. Dyssodia 
tenuiloba var. texana (Cory) Strother, Univ. Calif. Publ. Bot. 48: 
76. 1969. TyPE: USA. Texas. Taylor Co.: Abilene, Camp 
Barkeley, grassland in stony clay soil, 26 Apr 1943, W.L. Tolstead 
7030 (holotype: GH; isotype: SMU!). 


2c. Thymophylla tenuiloba var. treculii (A. Gray) Strother, Sida 11: 
378. 1986. Hymenatherum treculii A. Gray, Proc. Amer. Acad. 
Arts 19: 42. 1883. Dyssodia treculii (A. Gray) B.L. Rob., Proc. 
Amer. Acad. Arts 49: 508. 1913. Dyssodia tenuiloba var. treculii 
(A. Gray) Strother, Univ. Calif. Publ. Bot. 48: 75. 1969. TyPE: 
USA. Texas. “SE Texas [near Eagle Pass], A. Trecul s.n. (holotype: 
GH). 


ACKNOWLEDGEMENTS 


This study is based on study of collections from SMU/BRIT 
and TEX/LL, the latter originally loaned to BRIT for study by a staff 
member who found other employment before she could complete the 
study. I’m grateful to John Strother for comments, though we don’t 
agree on the taxonomy. 


LITERATURE CITED 


Johnston, M.C. 1956. The Texas species of Dyssodia (Compositae). 
Field & Lab. 24: 60-69. 

Loockerman, D., B.L. Turner and R.K. Jansen. 2003. Phylogenetic 
relationships within the Tageteae (Asteraceae) based on nuclear 
ribsomal ITS and chloroplast ndhF gene sequences. Syst. Bot. 28: 
191-2007. 

Robinson, B.L. 1913: Diagnoses and transfers among the 
spermatophytes. Proc. Amer. Acad. Arts 49: 502-517. 

Strother, J.L. 1969. Systematics of Dyssodia. Univ. Calif. Publ. Bot. 
48: 1-88. 

Strother, J.L. 1970. Dyssodia. Pp. 1680-1683. In D.S. Correll and 
M.C. Johnston, Manual of the Vascular Plants of Texas. Texas 
Research Foundation, Renner. 

Strother, J.L. 1986. Renovation of Dyssodia. Sida 11: 371-378. 


338 Phytologia (August 2009) 91(2) 


Strother, J.L. 1989. Chromosome numbers in Thymophylla 
(Compositae: Tageteae). Sida 13: 351-358. 

Strother, J.L. 2006. Thymophylla. Pp. 221—222, Vol. 21. In: Flora of 
North America Editorial Committee (eds.). Flora of North 
America North of Mexico. Oxford University Press, New York 
and Oxford. 

Turner, B.L. 1996. The Comps of Mexico. Vol. 6. Tageteae and 
Anthemideae. Phytologia Mem. 10:1—93. 

Turner, B.L., H. Nichols, G. Denny and O. Doron. 2003. Atlas of the 
Vascular Plants of Texas. Vol. 1—Dicots. Sida, Bot. Miscellany, 
Vol. 24. 


© >-~ 


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© Thymophylila wrightii 


@ Thymophylla tenuiloba 
var. tenuiloba 


Figure 1. Distribution of Thymophylla wrightii and T. tenuiloba var. 
tenuiloba in Texas. Collections were first mapped on a large-scale 
highway map, then transferred to this format (also for Fig. 2). Var. 
tenuiloba ranges into Mexico; it also occurs as an adventive in the 
U.S.A. in Alabama, California, and Louisiana. 


Phytologia (August 2009) 91(2) 339 


@ Thymophylla tenuiloba 
var. treculii 


® Thymophylla tenuiloba 
var. texana 


Figure 2. Distribution of Thymophylla tenuiloba var. treculii and var. 
texana in Texas. 


340 Phytologia (August 2009) 91(2) 


BIOLOGICAL STATUS OF THE VARIETAL TAXA OF 
THYMOPHYLLA PENTACHAETA 
(ASTERACEAE: TAGETEAE) 


B. L. Turner 
Plant Resources Center 
The University of Texas at Austin 
Austin, Texas 78712 
billie@uts.cc.utexas.edu 


ABSTRACT 


The biological status of Thymophylla |Dyssodia| pentachaeta 
is evaluated; the species was treated by Strother (1969, 1986) as having 
four infraspecific taxa: var. belenidium, var. hartwegii, var. 
pentachaeta and var. puberula. After examining most of the 
specimens that his taxonomy was based upon, it is concluded that the 
all of the taxa are worthy of specific rank, except for var. belenidium, 
which is treated as a synonym of 7. pentachaeta. Since all of these taxa 
were previously treated as species, no new names are required. 
Reasons for the dispositions are given, along with maps showing their 
distributions, these based upon numerous specimens assembled since 
the seminal study of Strother. Phytologia 91(2): 340-346 (August, 
2009). 


KEY WORDS: Asteraceae, Tageteae, Dyssodia, Thymophylla, T. 
pentachaeta, Argentina, Mexico, U.S.A. 


Strother (2006) provided a_ systematic treatment of 
Thymophylla pentachaeta (DC.) Small for the Flora of North America, 
this largely based upon his doctoral study of the genus Dyssodia (s.1.). 
In this, he recognized a subsp. hartwegii, this having but a single var. 
hartwegii (A. Gray) Strother; and a subsp. pentachaeta, this having 
three varieties: var. belenidium (DC.) Strother, var. pentachaeta (DC.) 
Small and var. puberula (Rydb.) Strother. 


In my treatment of the Comps of Mexico (Turner 1996) I 
inappropriately recognized Thymophylla pentachaeta as possessing 


Phytologia (August 2009) 91(2) 34] 


only two varieties: var. pentachaeta and var. hartwegii. My detailed 
reexamination of the group has led to the present treatment in which 7. 
puberula is resurrected, leading to the recognition of three species in 
the complex. The biological status of each of these is discussed below. 


var. belenidium 


The type of this taxon is from Argentina, based upon 
specimens obtained in the province of Mendoza by Arnott prior to 
1838, the year of its publication. Strother accepted the taxon as a valid 
variety, and assumed it to be confined to Argentina and_ the 
southwestern U.S.A. and adjacent northern Mexico. In his key to taxa, 
he distinguished the variety from var. pentachaeta by its shorter 
peduncles (2-5 cm vs. longer); outer phyllaries nearly free to the base 
(vs. not so), these bearing 3-6 pairs of marginal glands (vs. “fewer 
glands”). If one applies such key leads to specimens from Argentina 
(on file at LL-TEX) it will be found that both var. belenidium and var. 
pentachaeta occur in that country, but such is not noted by Strother. 
Presumably, he believed Argentina to lack specimens referable to var. 
pentachaeta. Further, I found so much variation in the characters called 
to the fore by Strother, that I was unable to map a coherent var. 
belenidium in either Argentina or North America. In short, I take the 
two taxa to be synonymous. Fig. | shows the distribution of the two 
taxa as interpreted by Strother. I would map these as but a continuous, 
highly variable, var. pentachaeta, both in Argentina and North America 
(Figs. 1, 2 and 3). 


var. pentachaeta = Thymophylla pentachaeta Figs. 3, 5 


The type of this taxon is from the state of Nuevo Leon, 
Mexico, first collected by Berlandier in the vicinity of Monterrey prior 
to its publication by De Candolle in 1836. As indicated in the above 
account, I consider 7. belenidium to be synonymous with var. 
pentachaeta, the characters separating these are highly variable and 
when mapped as a syndrome do not stand up to meaningful morpho- 
geographical interpretations. 


342 Phytologia (August 2009) 91(2) 


var. puberula = Thymophylla puberula Fig. 4 


The type of this taxon is from the state of San Luis Potosi, 
Mexico, first collected by Schaffer in 1877 in the Valley of San Luis 
Potosi. As indicated by Strother (his Fig. 18), this taxon is sympatric 
with 7. pentachaeta over many a mile of Mexico (Fig. 4). In spite of 
the numerous populations sampled, very few intermediates between the 
two taxa have been detected in the field or in the herbarium, either by 
Strother (at least by annotations on specimens) or myself, this in spite 
of the fact they often grow in close proximity. Indeed, numerous 
specimens assembled since Strother’s study has shown the two taxa to 
be easily recognized, intermediates being conspicuously absent, 
suggesting specific status for both. 


var. hartwegii = Thymophylla hartwegii Fig. 6 


This species is easily recognized by the characters called to the 
fore by Strother, hence its treatment as a monotypic subspecies by the 
latter author. It is known to grow with or near both 7. puberula and T. 
pentachaeta without the propensity to form recognizable hybrids with 
either. For example, in Cochise Co., Arizona 7. hartwegii is said by 
Barr (63-130, TEX)) to occur “as [a] distinct population but adjacent to 
Dyssodia pentachaeta.” In short, it appears to be a good biological 
species. 


A complete synonymy for all of the above taxa is given by 
Strother (1969). 


ACKNOWLEDGEMENTS 


I am grateful to my colleagues, Guy Nesom and Jana Kos, for 
reading the paper and offering helpful suggestions. 


LITERATURE CITED 


Strother, J.L. 1969. Systematics of Dyssodia Cavanilles (Compositae: 
Tageteae). Univ. Calif. Publ. Bot. 48: 1-88. 


Phytologia (August 2009) 91(2) 343 

Strother, J.L. 1986. Renovation of Dyssodia (Compositae: Tageteae). 
Sida 11: 371- 378. 

Strother, J.L. 2006. Thymophylla, in Fl. N. Amer. 21: 239-243. 


Turner, B.L. 1996. Dyssodia, in Phytologia Memoirs 10: 7-18. 


Fig. 1. Bicentric distribution of Thymophylla pentachaeta. 


344 Phytologia (August 2009) 91(2) 


Fig. 2. Distribution of Thymophylla pentachaeta in Argentina, by 
Provinces (data from http://www.tropicos.org). 


Phytologia (August 2009) 91(2) 


THYMOPHYLLA 
pentachaeta 


{includes T. belenidium} 


Fig. 3. Distribution of Thymophylla pentachaeta in Mexico, as 
envisioned by Turner (present account). 


THYMOPHYLLA 
puberula 


[= T. pentachaeta 
var. puberuta]} | 


Fig 4. Distribution of Thymophylla puberula in North America as 
envisioned by Turner (present account). 


345 


346 Phytologia (August 2009) 91(2) 


THYMOPHYLLA 
pentachaeta 


Fig. 5. Distribution of Thymophylla pentachaeta in the USA as 
envisioned by Turner (present account). 


THYMOPHYLLA 
hartwegii 


Fig. 6. Distribution of Thymophylla hartwegii. 


Phytologia (August 2009) 91(2) 347 


PANICUM COLORATUM NEW FOR ARIZONA, 
AND ECHINOCHLOA HOLCIFORMIS NEW FOR THE 
UNITED STATES 


John R. Reeder and Kathryn Mauz 
Herbarium (ARIZ), Herring Hall, P.O. Box 210036, 
University of Arizona, Tucson, AZ 85721 


ABSTRACT 


Two new records for Arizona, one of which is new to the 
United States, are reported. The introduced Panicum coloratum L. has 
previously been found in neighboring states, and is now known from 
one location in southern Arizona. Echinochloa holciformis (H.B.K.) 
Chase is known from several Mexican states but has not formerly been 
documented north of the international border. Phytologia 91(2): 347- 
352 (August, 2009). 


KEY WORDS: Panicum coloratum, Kleingrass, Echinochloa 
holciformis, Poaceae, Arizona. 


Author’s note: John Raymond Reeder (1914-2009) had been 
planning a note about these two new records when, in January 
2008, he and his wife, Charlotte, were seriously injured in a car 
accident. Sadly, John never fully regained his former vigor. I 
am happy to have collaborated with him in the field and 
herbarium, and to have been included in discussions of these 
two records. I have prepared this brief communication, at long 
last, as yet another of his contributions to the grass flora of 
North America.—KM 


Introduced from Africa, the perennial Panicum coloratum L. 
was first documented in Texas from nursery collections in the 1940s 
and 1950s, and has more recently been collected in uncultivated 
situations in Texas (Flora of Texas Database, www.biosci.utexas.edu/ 
prc/Tex.html, Feb 2009) and in New Mexico (New Mexico 
Biodiversity Collections Consortium, nmbiodiversity.org, Feb 2009). 
The plant, commonly called Kleingrass, was grown in the Soil 


348 Phytologia (August 2009) 91(2) 


Conservation Service Nursery in Tucson, Arizona, in the 1940s, and 
has been collected in Sonora, México, just south of the border with 
Arizona (University of Arizona Herbarium Database, 
ag.arizona.edu/herbarium/ search, Feb 2009). The collection reported 
here is believed to be the first record of the grass outside of cultivation 
in the state of Arizona. Although favoring the tropics and subtropics, 
and often found in wet ground (Freckmann and Lelong 2003), 
occurrences in the southwest region tend to be in ruderal settings but 
range in elevation up to 1600 m (5250 ft) above sea level. 


SPECIMEN: USA. ARIZONA. Cochise County: Parker Canyon Lake 
parking area, surrounded by vegetation of oaks, junipers, etc. Several 
clumps, this specimen from one ca. 80 cm diam.; the plants to | m tall. 
Elev. 1600 m. 22 Aug 2002, J.R. Reeder 9846 & C.G. Reeder (ARIZ, 
US) (Fig..1). 


The New World native Echinochloa holciformis (H.B.K.) 
Chase habitually grows in saturated substrates, often in shallow water 
and standing a meter or more tall. It is known from several Mexican 
states including Aguascalientes, Durango, Guanajuato, Jalisco, México, 
Michoacan, Nayarit, and Puebla, as well as south to Guatemala 
(McVaugh 1983; Rzedowski and Rzedowski 2001). The nearest 
occurrence to Arizona documented by herbarium specimens comes 
from neighboring Sonora, México (A.L. Reina-G. 98-1371, 19 Sep 
1998, ARIZ 349502!), about 325 km (203 mi) south of the international 
border. Since E. holciformis was first found in Arizona in 2002, several 
more collections have been made of this species from three localities, 
all in southern Santa Cruz County; these are, to-date, the only known 
records for the United States. 


Echinochloa holciformis is distinguished from congeners, 
particularly E. polystachya (H.B.K.) Hitche. and E. oplismenoides (E. 
Fourn.) Hitche. that are both known from the southern continental 
United States (Michael 2003), by the combination of glabrous culms, a 
conspicuously hairy ligule, an empty sterile (lower) floret, and awn of 
the sterile lemma ranging 3.5-5.0 cm long (see McVaugh 1983). Gould 
(1975: 533) reported a specimen from Texas, cited as “Williams in 
June, 1950 (US)” of which he wrote, “appears to be E. holciformis 
(H.B.K.) Chase. This is similar to E. polystachya in being a perennial 


Phytologia (August 2009) 91(2) 349 


with well-developed, hairy ligules and large spikelets, and with stamens 
in the lower floret. ... The specimen from Jefferson County, Texas, has 
puberulent culm nodes and awns to 20 mm long.” Although we failed 
to locate this specimen, it seems apparent from Gould’s description that 
the plant represented was, in fact, consistent with E. polystachya, and 
was not E. holciformis. The annual species E. oplismenoides was, like 
the records of EF. holciformis reported here, first documented in the 
United States by a collection from southern Arizona (Fishbein 1995); 
although broadly sympatric, it is readily distinguished from FE. 
holciformis by a glabrous ligule and shorter awns. 


Echinochloa holciformis is regarded as a perennial in its 
southern range (McVaugh 1983; Rzedowski and Rzedowski 2001), 
however the plants in the southern Arizona populations are clearly 
annual, with individuals dying completely to the substrate in the fall 
and populations varying greatly in size from year to year. The plants 
typically grow in shallow water, and often develop stout, fibrous roots 
from the lower nodes, but we have not observed structures that could be 
interpreted as creeping stems or rhizomes among the populations 
reported here. 


SPECIMENS: USA. ARIZONA. Santa Cruz County: Canelo Hills, ca. 1 
km N of Canelo Pass summit, rocky slope in oak-juniper area. Growing 
thickly on the margin of a cattle tank full of water. Elev. 1650 m. 10 
Oct 2002, J.R. Reeder 9894 & C.G. Reeder (ARIZ). Canelo Hills, ca. | 
km S of trail head to Arizona Trail along FS-799. Around a cattle tank a 
short distance W of the road, base of plants in the water. Elev. 1550 m. 
7 Sep 2004, J.R. Reeder 10002 & C.G. Reeder (ARIZ); 7 Sep 2004, 
J.R. Reeder 10003 & C.G. Reeder (ARIZ); 22 Sep 2004, J.R. Reeder 
10006 & C.G. Reeder (ARIZ, MO, US); 20 Sep 2005, J.R. Reeder 
10020 & C.G. Reeder (ARIZ, CAS, MO, NMC, TEX, US). San Rafael 
Valley, just S of entrance to Little Outfit Ranch. Grassland with a few 
scattered oaks and junipers. On margin of cattle tank a short distance 
from the road. Elev. 1550 m. 20 Sep 2005, J.R. Reeder 10021 & C.G. 
Reeder (ARIZ, US); 15 Sep 2006, J.R. Reeder 10028 & K. Mauz 
(ARIZ, MO) (Fig. 2). 


350 Phytologia (August 2009) 91(2) 


ACKNOWLEDGEMENTS 


I thank Charlotte Reeder and the staff of the University of 
Arizona Herbarium (ARIZ) for assistance in locating materials used in 
the preparation of this note, and for the specimen images. The 
comments by reviewers Charlotte Reeder and Philip Jenkins are also 
appreciated. 


LITERATURE CITED 


Fishbein, M. 1995. Noteworthy collections: Echinochloa oplismenoides 
(Fourn.) Hitche. Madrofio 42: 83. 


Freckmann, R.W. and M.G. Lelong. 2003. Panicum L., pp.450-488. In: 
Flora of North America vol. 25 — Magnoliophyta: Commelinidae 
(in part): Poaceae, part 2, M.E. Barkworth, K.M. Capels, S. Long 
and M.B. Piep, eds. Oxford University Press, New York and 
Oxford. 


Gould, F.W. 1975. The grasses of Texas. Texas A&M University Press, 
College Station. 


McVaugh, R. 1983. Gramineae. Flora Novo-Galiciana vol. 14, W.R. 
Anderson, ed. University of Michigan Press, Ann Arbor. 


Michael, P.W. 2003. Echinochloa P. Beauv., pp.390-402. Jn: Flora of 
North America vol. 25 — Magnoliophyta: Commelinidae (in part): 
Poaceae, part 2, M.E. Barkworth, K.M. Capels, S. Long and M.B. 
Piep, eds. Oxford University Press, New York and Oxford. 


Rzedowski, G.C. de and J. Rzedowski. 2001. Flora fanerogamica del 
Valle de México, second edition. Comision Nacional para el 
Conocimiento y Uso de la Biodiversidad, Patzcuaro. 


Phytologia (August 2009) 91(2) 35] 


Fig. 1. Panicum coloratum L. Near Parker Canyon Lake, Cochise 
County, Arizona, 22 Aug 2002 (ARIZ). 


352 Phytologia (August 2009) 91(2) 


389910 


Fig. 2. Echinochloa holciformis (H.B.K.) Chase. San Rafael Valley, 
Santa Cruz County, Arizona, 15 Sep 2006 (ARIZ). 


Phytologia (August 2009) 91(2) 353 


VARIATION IN JUNIPERUS DURANGENSIS AND RELATED 
JUNIPERS (CUPRESSACEAE): 
ANALYSIS OF nrDNA AND petN SNPs 


Robert P. Adams 
Biology Department, Baylor University, Waco, TX 76798, USA 
Robert_Adams@baylor.edu 


ABSTRACT 


Recent discovery of a low shrub from Topia, in the state of 
Durango, Mexico that appears similar to both J. durangensis and J. 
jaliscana, prompted the analyses of nrDNA and petN-psbM (cpDNA) 
SNPs. The plants from Topia differed from J. durangensis by 2 indels 
but were shown to be closely related as shown in a minimum spanning 
network. Phytologia 91(2): 353-358 (August, 2009). 


KEY WORDS: Juniperus durangensis, J. monticola, J. martinezii, J. 
flaccida, nrDNA, petN-psbM, SNPs, Cupressaceae, geographic variation. 


Juniperus durangensis Mart. is a tree or large shrub to 5 m that 
generally branches near the base (Adams, 2008). It is often found on 
rhyolite, a nutrient poor rocky volcanic substrate, in the mountains of 
western Mexico from Sonora and Chihuahua southward to 
Aguascalientes. Juniperus durangensis is in the serrate leaf margined 
junipers and appears most closely related to J. martinezii Perez de la 
Rosa and then to J. flaccida Schlecht, J. jaliscana Mart. and J. 
monticola Mart. (Fig. 1). 


nrDNA + trnC-trnD Figure 1. Clade from the serrate leaf 
jaliscana margined junipers, based on nrDNA + 
monticola trnC - tmD (cpDNA) data from Adams 
(2008) showing the putative 
) relationship of J. durangensis to 
durangensis | closely related junipers. 


martinezii 


flaccida 


354 Phytologia (August 2009) 91(2) 


Recently, a low growing shrub was discovered near Topia, 
Durango that seems to be related to J. durangensis, although it has 
some characteristics of J. jaliscana. To further investigate the Topia 
juniper, sequencing of nrDNA and the petN-spacer-psbM cp DNA 
region were performed to obtain SNPs to reexamine the relationship of 
the Topia juniper to J. durangensis and other closely related junipers. 


MATERIALS AND METHODS 


Specimens collected (GenBank #: nrDNA; petN-psbM): J. 
durangensis, Adams 6832-6834, (FJ948469, FJ948473) 52 km w of El 
Salto, on Mex 40, Durango, MX; Adams 11420-11421, (FJ948469, 
FJ948473)Topia, Durango, MX;, J. flaccida, Adams 6892-6893, 
(FJ948470, FJ948476), on Mex. 60, 19 km E. of San Roberto Junction, 
Nuevo Leon, Mexico; J. jaliscana, Adams 6846-6848, (FJ948466, 
FJ948475), 19 km E of Mex. 200 on the road to Cuale, Jalisco, Mexico; 
J. martinezii, Adams 5950, 5951, 8709, (FJ948471, FJ948474)10 km s of 
Mex 85 on road to La Quebrada Ranch, Jalisco, MX; J. monticola f. 
monticola, Adams 6874-6878, (FJ948467, FJ746736) 1 km n of Mex 105, 
9 km nw of Pachuca, El Chico National Park, Hidalgo, Mexico. Voucher 
specimens are deposited at BAYLU. 


One gram (fresh weight) of the foliage was placed in 20 g of 
activated silica gel and transported to the lab, thence stored at -20° C 
until the DNA was extracted. DNA was extracted using the Qiagen 
DNeasy mini kit (Qiagen Inc., Valencia CA). 


SNPs obtained from DNA sequencing 
ITS (nrDNA) and trnC-trnD amplifications were performed in 

50 pl reactions using 10 ng of genomic DNA, 3 units Qiagen Taq 
polymerase, 5 ul 10x buffer (final concentration: 50 mM KCl, 10 mM 
Tris-HCl (pH 9), 0.01% gelatin and 0.1% Triton X-100), 1.75 mM 
MgCl, 20 ul Q solution (2X final), 400 uM each dNTP, 1.8 uM each 
primer and 4%(by vol.) DMSO. 
Gene __ Primers 2x buffer annealing program size bp 
nrITS ITS-42F/ITSB+57R K 50°C (94-50x30) 1270-1272 

etN__petNSF/psbMI11IR_ EE 50°C —_—((94-50x30) _ 839-845 

Primers (5'-3'): 

ITS: ITSA = GGA AGG AGA AGT CGT AAC AAG G; 


Phytologia (August 2009) 91(2) 355 


IYSB= CTT TTG'CTC CGC TTA TTG ATA TG. 
ITSA and ITSB primers from Blattner (1999). 
additional ITS primers (based on Juniperus sequences): 
ITSA-42F = GAT TGA ATG ATC CGG TGA AGT 
ITSB+57R = ATT TTC ATG CTG GGC TCT 
petN - psbM: 
petNSF = AAC GAA GCG AAA ATC AAT CA 
psbM111R = AAA GAG AGG GAT TCG TAT GGA 
petN and psbM primers were based on conserved sequences from 
Juniperus species. 


The following PCR conditions were used: MJ Research 
Programmable Thermal Cycler, 30 cycles, 94°C (1 min.), 50°C or 57°C 
(2 min.), 72°C (2 min.), with a final step of 72°C (5 min.). The PCR 
reaction was subjected to purification by agarose gel electrophoresis 
(1.5% agarose, 70 v, 55 min.). In each case, the band was excised and 
purified using a Qiagen QIAquick gel extraction kit. The gel purified 
DNA band with the appropriate primer was sent to McLab Inc. for 
sequencing. Sequences for both strands were edited and a consensus 
sequence was produced using Chromas, version 2.31 (Technelysium 
Pty Lid.). Alignments were made using MAFFT 


(http://align.bmr.kyushu-u.ac.jp/mafft/). 


SNPs analyses 

Aligned data sets (nrDNA and trnC-trnD) were analyzed by 
CLEANDNA (Fortran, R. P. Adams) to remove invariant data. 
Mutational differences were computed by comparing all SNPs, divided 
by the number of comparisons over all taxa (= Gower metric, Gower, 
1971; Adams, 1975). Principal coordinate analysis was performed by 
factoring the associational matrix using the formulation of Gower 
(1966) and Veldman (1967). A minimum spanning network was 
constructed by selecting the nearest neighbor for each taxon from the 
pair-wise similarity matrix, then connecting those nearest neighbors as 
nodes in the network (Adams et al., 2003). 


RESULTS AND DISCUSSION 


Analyses of the nrDNA sequences revealed 26 mutational 
events that included a 2-bp indel (CA) that was present in the three J. 


356 Phytologia (August 2009) 91(2) 


martinezii individuals and absent on all other taxa. In addition, one of the 
J. flaccida individuals (6893) contained an insertion (A) that was absent 
in all other samples. Thirteen of the mutational events were single events 
and 13 were multiple occurring with fidelity within populations. A 
minimum spanning network was constructed based on 13 SNPs 
(including one indel) and is shown in figure 2 (left). The Topia shrubs 
had no SNPs different from J. durangensis. Overall, these taxa appear to 


Minimum Spanning Networks 


13 SNPs, ITS sequences 12 SNPs, petN-psbM 
sequences 


J. martinezii 


J. flaccida 


J. flaccida me 
J. jaliscana 


1 


J, monticola J. jaliscana 


Figure 2. Minimum spanning network based on SNPs from nrDNA (left) 
and petN-spacer-psbM (right). The number of SNPs are next to the links. 


be very closely related, with only J. martinezii having appreciable SNPs 
differences. 


Phytologia (August 2009) 91(2) 357 


Analyses of a petN-spacer-psbM (from cpDNA) revealed 14 
mutational events, with 6 of these being indels. Two events occurred in 
single individuals. Twelve SNPs (including 5 indels) were used to 
construct a minimum spanning network (Fig. 2, right). The Topia plants 
had 2 indels (an A at 401 and a deletion at 666) not found in J. 
durangensis (or other taxa). Juniperus martinezii is separated by 4 SNPs 
(Fig. 2, right) and J. flaccida-J. jaliscana-J. monticola are separated from 


J. durangensis by 6 or more SNPs. 


Combining the nrDNA and petN-s-psbM data resulted in the 


minimum spanning network shown in figure 3. Notice that species are 


Minimum Spanning Network 
Combined 13 ITS SNPs + 12 petN SNPs 


/ 
J. martinezii ( 2) 


“ 


Topia ( 
popn. 


J. monticola e ») 
oe 


J. flaccida 


J. jalisciana 


Figure 3. Minimum spanning network based on combined SNPs from 
nrDNA and petN-s-psbM sequencing. The dotted line is the second 
shortest link for J. monticola (9 SNPs). 


358 Phytologia (August 2009) 91(2) 


separated by from 4 to 11 SNPs. The Topia plants are quite near typical 
J. durangensis in these two nucleotide sequences. However, it is clear 
that conclusions based on a single sequence might be misleading (cf. Fig. 
2, left vs. right). Additional collections and analyses of the leaf essential 
oils of the Topia plants, as well as sequencing additional genes, should 
shed light on the scope of differentiation of this population and its 
affinities to other junipers. 


ACKNOWLEDGEMENTS 


Thanks to Andrea Schwarzbach, Socorro Gonzalez and Billie 
Turner for manuscript reviews. Thanks to Tonya Yanke for lab 
assistance and to Socorro Gonzalez for samples of the Topia juniper. 
This research was supported in part with funds from Baylor University. 


LITERATURE CITED 


Adams, R. P. 1975. Statistical character weighting and similarity 
stability. Brittonia 27: 305-316. 

Adams, R. P. 2008. Junipers of the World: The genus Juniperus. 2nd 
Ed., Trafford Publ., Vancouver, B. C. 

Adams, R. P., A. E. Schwarzbach and R. N. Pandey. 2003. The 
Concordance of Terpenoid, ISSR and RAPD markers, and ITS 
sequence data sets among genotypes: An example from Juniperus. 
Biochem. Syst. Ecol. 31: 375-387. 

Adams, R. P., E. von Rudloff, L. Hogge and T. A. Zanoni. 1980. The 
volatile terpenoids of Juniperus monticola f. monticola, f. 
compacta and f. orizabensis. J. Nat. Prods. 43: 417 - 419. 

Blattner, F. R. 1999. Direct amplification of the entire ITS region from 
poorly preserved plant material using recombinant PCR. 
BioTechniques 27: 1180-1186. 

Gower, J. C. 1966. Some distance properties of latent root and vector 
methods used in multivariate analysis. Biometrika 53: 326-338. 

Gower, J. C. 1971. A general coefficient of similarity and some of its 
properties. Biometrics 27: 857-874. 

Veldman D. J., 1967. Fortran programming for the behavioral sciences. 
Holt, Rinehart and Winston Publ., NY. 


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