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~ PHYTOLOGIA 


An international journal to expedite botanical and phytoecological publication 


Vol. 67 September 1989 No. 3 


CONTENTS 


, TELFORD, |.R.H., Notes on Sicyos (Cucurbitaceae) in the Hawaiian 


| S | ands SSSSSSSHSSSSSSHSSSSSSSSSHSSSHSSSSSSHSSSSSSSSSSGHSSSSSSHSSSSSSSSSSSOSSHSSSSOOSOSEOSS 209 


BRIDGES, E.L. & S.L. ORZELL, Lindera subcoriacea (Lauraceae) new to 


Al abama SOCSSSSSSSSSSSSSSSSSSSSSSSSSHSSSHSSSSSSHSSSSSSSSHSSSHSSSSSSSHSSSSSSSOSSSSSSSSSSSSOSOS ?1 4 


» BRIDGES, E.L., Carex walteriana (Cyperaceae) and Euphorbia inundata 
(Euphorbiaceae) new to Mississippi Pritiritiitiititiiiit tii) ?1 7 


. BRUCHER, H.E., Refutation: of recent creations of microspecies and 
hybrid taxa in Argentinian Solanum (sect. PetOta) secccersecssseeeseees 220 


/ HILSENBECK, R.A., Generic affinities and. typification of eleven species 
excluded from Siphonoglossa Oerst. (Acanthacede) ereesssecsseseeeee 227 


OCHOA, C.M., Change of name in a tuber bearing SO/aNUM eeecceseeveee 235 
OUDEJANS, R.C.H.M., Corrections of new combinations in Euphorbia 236 


BARKLEY, T.M., New taxa and nomenclatural combinations in Senecio in 
Mexico and the United States SSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSOSSSOSESOSE 237 


r RIEFNER, R.E., JR., Punctelia punctilla (Hale) Krog, new to North 


A me r i GY a SSOOSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSHSSSHSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSOOSSSOSSES 2 54 


Contents continued on inside cover. 


Published by Michael J. Warnock 
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Contents (cont.) 


/ COFFIN, D.P. & W.K. LAUENROTH, Small scale dis 
successional dynamics in a shortgrass plant commun 
of disturbance CHALraCtEListiCs cecccccccccccccccccscccccce 


WARNOCK, M. J a Book Reviews «ie esa saad ieeese a poceee 


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OCT 23 989 
NEW YORK) asa 
BOTANICAL GARDEN ~ 


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Phytologia (September 1989) 67(3):209-213. 


NOTES ON SICYOS (CUCURBITACEAE) IN THE HAWAIIAN ISLANDS 


Ian R.H. Telford 
Australian National Botanic Gardens, P.O. Box 1777, Canberra, A.C.T. 
2600, AUSTRALIA 


ABSTRACT 


Nomenclature and taxonomy of Sicyos L. in Hawai’i are clarified as 
a precursor to the Manual of Flowering Plants of Hawai’. Sarz H. St. 
John is reduced to synonymy under Sicyos. New combinations Sicyos 
herbstii (H. St. John) Telford and Sicyos alba (H. St. John) Telford 
are made and Sicyos hillebrandw var. anunu H. St. John given new 
status as S. anunu (H. St. John) Telford. Stcyos pachycarpus Hook. & 
Arn. is lectotypified and shown to be the correct name for the species 
known as S. microcarpus H. Mann. 


KEY WORDS: Hawai’i, Sicyos herbstiz, Sicyos alba, Sicyos anunu, 
Sicyos pachycarpus, lectotypification. 


INTRODUCTION 


In Hillebrand’s Flora of the Hawaiian Islands (1888, p 134), Sicyos L. was 
treated as the only indigenous genus of Cucurbitaceae with 8 endemic species. 
Since then, 5 genera have been segregated from Sicyos in Hawai'i and an 
additional 48 taxa described in Szcyos or its segregate genera. 

The delimitation of the segregate genera on fruit ornamentation was ques- 
tioned by Jeffrey (1978, p 361). His conclusion that they are congeneric is 
followed and developed in this paper. 

As a precursor to the Manual of the Flowering Plants of Hawai: (Wagner 
et al., in press), this paper attempts to resolve some problems of taxonomy 
and nomenclature in Hawaiian Cucurbitaceae. 


Status of the segregate genera. 


As discussed by Jeffrey (1978, p 361), four of the genera segregated from 
Sicyos, namely Cladocarpa(H. St. John) H. St. John (1978b, p 491), Sicyocarya 
(A. Gray) H. St. John (1978a, p 407) and Skottsbergiliana H. St. John (1974, p 
457), are better treated at sectional rank at most. All are consistent with the 
vegetative and floral syndromes of Sicyos, differing only in the fruit morphology 
which characterizes the sections. 

The following new combination transfers a recently described species from 
Cladocarpa into Sicyos. 


209 


210 PeYVT OL OG IA volume 67(3):209-213 September 1989 


Sicyos herbstii (H. St. John) Telford, comb. nov. BASIONYM: Cladocarpa 
herbstu H. St. John, Phytologia 63:185. 1987. TYPE: Hawaiian Is., 
Kaua'i, Barking Sands, D. Herbst 5691 (HOLOTYPE: BISH). 


Cladocarpa julesii H. St. John, Phytologia 63:185. 1987. TYPE: Kauai, J. 
Remy 542 (HOLOTYPE: P). 


Cladocarpa pauciramosa H. St. John, Phytologia 63:186. 1987. TYPE: 
Kaua’i, Polihale, C. Christensen 326 (HOLOTYPE: BISH). 


The single species of the fifth segregate genus, Sarz alba H. St. John (1978b, 
p 491) was segregated from Szcyos cucumerinus A. Gray because of its white 
fleshy fruit, despite mature fruit of 5S. cucumerinus being “still unknown.” 
St. John relegated Gray’s S. cucumerinus var. B to synonymy under Sarz. 
Recently collected fruit specimens of S. cucumerinus have been examined at 
BISH by the present author and the texture of the pericarp proved to be 
subfleshy. Certainly the character of fleshy fruit alone is untenable for generic 
segregation. This taxonomic view of Sarz in relation to Sicyos conforms with 
the treatment of Sechium P. Br. by Jeffrey (1978, p 360). In Sechium also, the 
pericarp may be fleshy, fibrous or woody. 

A new combination is provided below for Sarz alba in Sicyos. 


Sicyos alba (H. St. John) Telford, comb. nov. BASIONYM: Sarz alba H. 
St. John, Bot. Jahrb. Syst. 99:493. 1978. TYPE: Hawaii, Kulani, W. 
Gagne 698 (BISH). 


Vegetatively and in floral morphology, Sicyos cucumerinus and Sarz alba 
are so similar that not only generic but also specific separation must be ques- 
tioned. Field studies and additional collections of the two species are required 
before resolution of the species delimitation problem is attempted. 


The identity of Sicyos pachycarpus. 


The first species of Sicyos collected and named from the Hawaiian Islands 
was S. pachycarpus Hooker & Arn. (1832, p 83), collected by Collie on Dia- 
mond Head, O’ahu in 1826-27. A. Gray (1854, p 650, t. 80), in his study of 
the U.S. Exploring Expedition’s collections of 1840, misapplied the name S. 
pachycarpus to an as yet undescribed species from the Wai’anae Mountains, 
O’ahu. The specimen of S. pachycarpus sensu Gray lies far outside the circum- 
scription of Hooker & Arnott’s protologue. Gray must not have seen Collie’s 
type collection housed at Kew. 

Cogniaux (1881, p 896) followed Gray’s misapplication and compounded 
it by citing more specimens. Hillebrand (1888, p 137) recognized a problem 
existed: “It is somewhat doubtful, however, if Gray’s S. pachycarpus is the 


Telford: —. Notes on Sicyos in Hawaiian Islands 211 


same as that of Hooker & Arnott, for the latter authors assign a height of only 1 
1/2-2” to their fruit, which points to S. mzcrocarpa.” Unfortunately, Hillebrand 
did not resolve the problem but instead also followed Gray’s misapplication. 

Collie’s type gathering of S. pachycarpus raises another problem. The 
herbarium sheet at Kew originally consisted of two elements - one of these 
has been annotated by St. John as S. mzcrocarpus H. Mann, the other as the 
lectotype (unpublished) of S. pachycarpus Hook. & Arn. The ‘S. microcarpus’ 
element has been remounted on a separate sheet. 

Hooker & Arnott’s protologue agrees better with St. John’s ‘S. microcarpus’ 
element than with that proposed by him for lectotypification - ‘female flowers 
are numerous in each capitulum...the capitulum itself is on a peduncle, about 
three-fourths of an inch long...fruit is ovate, about a line and a half long, 
suddenly attenuated into a beak which is almost half the length of the broad 
portion’. 

The other element, i.e. St. John’s ‘S. pachycarpus’ element, differs in fewer 
female flowers per head, the peduncles longer and more slender, the fruit not 
so abruptly contracted before the attenuation. This element belongs in S. 
waimanaloensis H. St. John (1987, p 192) and is readily distinguished from 
S. pachycarpus by the slower expansion of the leaf laminas as well as the 
characters above. The two species are sympatric in eastern O’ahu. 

The ‘S. microcarpus’ element is here proposed as lectotype of S. pachycar- 
pus. 


Sicyos pachycarpus Hook. & Arn., Bot. Beechey Voy. 3, Sandwich Is.: 83. 
1832. LECTOTYPE (here designated): Diamond Hill, O’ahu, Colle 
(K). 


S. microcarpus H. Mann, Proc. Amer. Acad. Arts 7:167. 1867. 
The identity of Sicyos hillebrandii. 


The name S. hillebrandit H. St. John (1934, p 7), based on S. laciniatus 
Hillebr. (1888, p 138) non L. (1753, p 1013), with the type from Kula, Maui, 
has been applied to few collections from Maui and to those from several pop- 
ulations on Hawai’i. The Hawaiian material differs from the type in several 
character states - staminate flowers 5-8 mm diameter; staminate inflorescences 
on peduncles 12-30 cm long; pistillate flower heads on peduncles 4-12 cm long; 
fruit 15-20 mm long, as well as geographic separation, but has hitherto been 
recognized only at varietal rank. It is here raised to the status of species. 


Sicyos anunu (H. St. John) Telford, stat. nov. BASIONYM: S. hillebrandit 
H. St. John var. anunu H. St. John, Phytologia 63:187. 1987. TYPE: 
Hawai'i Is., Kapapala, C.N. Forbes 400.H (HOLOTYPE: BISH). 


212 PHY TODHG.GTA volume 67(3):209-213 September 1989 


S. hillebrandit H. St. John var. douglas H. St. John, Phytologia 63:187. 
1987. TYPE: Hawai’i Is., N, slope Mauna Kea, O. Degener & Greenwall 
21,864 (HOLOTYPE: BISH). 


ACKNOWLEDGMENTS 


I wish to thank Dr. S. Sohmer and staff of Herbarium Pacificum, Bishop 
Museum, Honolulu for assistance and hospitality during my visit to Hawai'i 
in 1986. I am grateful to John Obata of O’ahu and Robert Hobdy of Maui 
for guiding me to Hawaiian Sicyos localities. Mr. Charles Jeffrey of the Royal 
Botanic Gardens, Kew provided encouragement and kindly arranged for the 
photography of Leningrad Szcyos specimens. 


BIBLIOGRAPHY 


Cogniaux, A. 1881. Cucurbitaces, zn A. DC., Monogr. Phan. 3:325-951, 
953-951. 


Gray, A. 1854. U.S. Exploring Ezped., Botany, Phanerogamia 1. 


Hillebrand, W.F. 1888. Flora of the Hawaiian Islands. Hafner Publishing 
Co., New York & London. 


Hooker, W.J. & G.A.W. Arnott. 1832. The Botany of Captain Beechey’s 
Voyage, 3 Sandwich Islands. 


Jeffrey, C. 1978. Further Notes on Cucurbitaceae:IV, Some New-World taxa. 
Kew Bull. 33(2):347-380. 


Linnaeus, C. 1753. Species Plantarum, Facsimile ed. 1957. Ray Society, 
London. 


Mann, H. 1867. Enumeration of American Plants. Proc. Amer. Acad. Arts 
7:143-235. 


St. John, H. 1934. Panicum, Psychotria, and Sicyos, Hawaiian plant studies 
2. Bernice P. Bishop Mus. Occ. Pap. 10(12):7. 


. 1974. Skottsbergiliana new genus (Cucurbitaceae) of Hawaii Island, 
Hawaiian plant studies 41. Pacific Sci. 28(4):457-462. 


Telford: Notes on Sicyos in Hawaiian Islands 213 


1978a. New combinations in Hawaiian Cucurbitaceae, Hawaiian 


plant studies 66. Phytologia 38(5):407-408. 


1978b. The Cucurbitaceae of Hawaii 1. generic revision. Bot. 
Jahrb. Syst. 99(4):490-497. 


1987a. Diagnosis of new species of Cladocarpa (Cucurbitaceae), 
Hawaiian plant studies 142. Phytologia 63(3):185-186. 


. 1987b. Diagnosis of new species of Szcyos (Cucurbitaceae), Hawai- 
ian plant studies 143. Phytologia 63(3):187-192. 


Wagner, W.L., D.R. Herbst & S.H. Sohmer. (in press). Manual of the Flow- 
ering Plants of Hawai’. University of Hawai’i Press & Bishop Museum 
Press, Honolulu. 


Phytologia (September 1989) 67(3):214-216. 


LINDERA SUBCORIACEA (LAURACEAE) NEW TO ALABAMA 


Edwin L. Bridges & Steve L. Orzell 
The University of Texas Herbarium, Austin, Texas 78713 USA 


ABSTRACT 


Lindera subcoriacea, a candidate species for listing as federally en- 
dangered or threatened by the U.S. Fish and Wildlife Service, is re- 
ported new to Alabama. Its habitat in Alabama is discussed in relation 
to habitats for this species in adjacent southern Mississippi. 


KEY WORDS: Rare plants, Lindera subcoriacea, Lauraceae, Al- 
abama. 


In 1983, Lindera subcoriacea B.E. Wofford was described as a new species 
restricted to evergreen shrub bogs in southern Mississippi and adjacent south- 
eastern Louisiana (Wofford 1983). More recently, several populations have 
been discovered in the sandhills region of North Carolina by Julie Moore of 
the N.C. Dept. of Natural Resources, and in South Carolina and Georgia 
by Robert B. McCartney of Aiken, South Carolina. Additional surveys by 
the Mississippi Natural Heritage Program have resulted in a total of at least 
17 localities in that state. After concerted search efforts in North Carolina, 
South Carolina, Mississippi, Georgia and Louisiana, Lindera subcorzacea is 
now known from a total of 34 sites. It has been predicted to occur in Alabama 
but apparently has never before been found in the state (Gordon et al. 1986). 
While surveying potential seepage bogs identified by the authors through using 
soil survey aerial photographs to predict natural community types, we added 
Alabama to the range of this rare, distinctive shrub. The collection data are 
as follows: 

Lindera subcoriacea B.E. Wofford (Lauraceae). UNITED STATES. Al- 
abama: Mobile Co.: hillside seepage shrub-herb bog on S side of Beverly- 
Jefferies Rd (Co Rd 96), 7.1 mi W of int. US 45 in Citronelle, 1.3 mi W of 
Ramey Rd and 1.6 mi E of Escatawpa River bridge; NWQ, SWQ, NEQ, NEQ, 
Sec. 2, TIN, R4W, Citronelle West 7.5’ Quad., 31° 04’ 49” N, 88° 21’ 04” W, 
elev. 160-180 ft., 15 May 1989, Orzell & Bridges 9960 (FSU,MISSA,MO,NCU, 
SMU,TEX,VDB). 

Lindera subcoriacea is occasional in partial shade of evergreen shrub-tree 
thickets within an extensive series of mid-slope hillside seepage bogs. As- 
sociated species include Persea palustris, Magnolia virginiana, Gaylussacia 


214 ; 


Bridges & Orzell: Lindera subcoriacea new to Alabama 215 


mosiert, Ilex coriacea, Myrica heterophylla, Myrica inodora, Smilaz laurifolia, 
Nyssa sylvatica var. biflora, Rhus verniz, Pyrus arbutifolia and Osmunda cin- 
namomea. Although many typical open seepage bog plants (Sarracenia spp., 
Rhynchospora spp., Xyris spp., Eriocaulaceae, Orchidaceae) occur within the 
seepage bogs at the site, they are not found in the immediate vicinity of Lindera 
subcoriacea. These habitats and associates are very similar to those reported 
from adjacent southern Mississippi (Wofford 1983; Gordon et al. 1986), and 
include the constant rangewide associates Magnolia virginiana, Myrica hetero- 
phylla and Rhus verniz (Gordon et al. 1986). 

This record brings the number of Lindera species in Alabama to three. Lzn- 
dera benzoin (L.) Blume is common in north Alabama, but infrequent south- 
ward, reaching southwest to Sumter, Marengo and Conecuh counties (Clark 
1971), all at least 100 km from the Mobile County site for L. subcoriacea. Sim- 
ilarly, in Mississippi the ranges of these two taxa are separated by at least 120 
km (Wofford 1983). In Alabama, Lindera melissifolia (Walt.) Blume, is known 
only from a single 19th century Buckley collection from Wilcox County (Clark 
1971; Wofford 1983). Lzndera subcorzacea differs from these by its faint aroma 
and elliptic to oblanceolate, subcoriaceous leaves with obtuse to rounded tips 
(Wofford 1983). 

The flora and natural community ecology of the Gulf Coastal Plain of 
the southeastern United States remains inadequately studied and understood. 
This region undoubtedly still harbors many significant plant records. We have 
noted that many species of the Gulf coastal states which were previously known 
from very few collections, and presumed to be naturally very rare, are actually 
locally common in naturally restricted and isolated specific habitats. System- 
atic field work needs to be focused on this region while the opportunity remains 
to uncover previously unknown species populations and natural areas. Thirty 
meters from our Lindera site the landscape is being cleared and many sig- 
nificant plant records and natural areas are vanishing each year before being 
documented. 


ACKNOWLEDGMENTS 


We wish to thank Guy Nesom and Carol Todzia for their helpful review 
comments. Publication costs were provided by the Plant Resources Center of 
the University of Texas at Austin. 


He PHD OLO GLA volume 67(3):214-216 September 1989 


LITERATURE CITED 


Clark, R.C. 1971. The woody plants of Alabama. Ann. Missouri Bot. Gard. 
58:99-242. 


Gordon, K.L., R.L. Jones & J.B. Wiseman, Jr. 1986. Status report for 
Lindera subcoriacea B. E. Wofford. Unpublished report to the U.S. Fish 
and Wildlife Service, Jackson, Mississippi. 


Wofford, B.E. 1983. A new Lindera (Lauraceae) from North America. J. 
Arnold Arb. 64:325-331. 


Phytologia (September 1989) 67(3):217-219. 


CAREX WALTERIANA (CYPERACEAE) AND EUPHORBIA INUNDATA 
(EUPHORBIACEAE) NEW TO MISSISSIPPI 


Edwin L. Bridges 
The University of Texas Herbarium, Austin, Texas 78713 USA 


ABSTRACT 


Carez waliertana and Euphorbia 1nundata are reported for the first 
time in the published literature on the flora of Mississippi. Both are 
found in wetland savannas on the outer coastal terraces of Jackson 
County. These sites are the westernmost known for both species. Their 
habitats and range disjunctions are related to their rangewide distribu- 
tion and ecology. 


KEY WORDS: Mississippi, Carez, Euphorbia, floristics. 


Norquist (1984) discovered two species that were apparently unreported 
for Mississippi while conducting field work for a comparative study of the soils 
and vegetation of savannas in the state. Although Ms. Norquist suggested 
in 1988 that the authors could report her records, we waited until we had 
independently, without knowing the previous specific locations, found these 
species in the state. The collection data are as follows: 

Carer walteriana Bailey (Cyperaceae). UNITED STATES. Mississippi: 
Jackson Co.: low pond cypress drainage swale on S side of US 90 at crossing 
of Franklin Creek, 0.1 mi W of Alabama state line, and 1.4 mi NE of int. 
Franklin Creek Rd exit from I-10; NWQ, NWQ, NEQ, Sec. 5, T7S, R4W, 
Kreole 7.5’ Quad., 30° 28’ 20” N, 88° 24’ 08” W, elev. 10 ft., 16 May 1989, 
Orzell & Bridges 10093 (MISSA,MO,NCU,TEX). 

The canopy dominant of this Carer walteriana site is Tarodtum ascen- 
dens, with Acer rubrum, Magnolia virginiana and Nyssa sylvatica var. bi- 
flora. Carez walteriana dominates the herb layer, with subdominants Ludwigia 
sphaerocarpa and Smilaz walteri. Other herbaceous associates include Erio- 
caulon decangulare, Ludwigia microcarpa, Osmunda regalis, Panicum hemito- 
mon, Rhynchospora corniculata, Woodwardia areolata and W. virginica. 

Carez walteriana is a common species of wet pond cypress or swamp black 
gum ponds, depressions and swales, and the edges of evergreen shrub-tree 
bogs and swamps. It ranges on the Atlantic and East Gulf Coastal Plains 
from the panhandle of Florida north to Massachusetts. The nearest records 


217 


218 PHYTOLOGIA volume 67(3):217-219 September 1989 


to Mississippi are in Geneva County, Alabama [Kral 38642; McDaniel 8918 
(VDB)} and in Okaloosa and Santa Rosa counties, Florida (Wilhelm 1984). 
Carer walteriana appears to be quite rare west of the Apalachicola River, 
although it is fairly frequent west to southwestern Georgia and in Florida to 
the Apalachicola National Forest. The author has collected C. walteriana in 
habitats similar to the Mississippi site in Liberty and Wakulla counties, Florida 
but has not seen it in apparently suitable habitats farther west in the Florida 
panhandle and southern Alabama. The Mississippi site is at least 120 km west 
of the nearest isolated station and 330 km west of the more continuous range. 

Euphorbia tnundata Torr. er Chapm. (Euphorbiaceae). UNITED STATES. 
Mississippi: Jackson Co.: wetland longleaf pine savanna on W side of Martin 
Bluff Rd, 0.1-0.2 mi S of I-10 bridge, ca 3 mi N of Gautier and 1.5 mi E of 
Gautier-Vancleve Rd int. with I-10; NEQ, NEQ, SEQ, Sec. 13 (or SE part 
Sec. 37), T7S, R7W, Gautier North 7.5’ Quad., 30° 26’ 07” N, 88° 37’ 55” W, 
elev. 25 ft., 16 May 1989, Orzell & Bridges 10101 (MISSA,TEX); acid low 
pine woods, ca 3 mi E of Ocean Springs, coastal flatwoods, 29 Apr 1967, S.B. 
Jones 11879 (VDB). 

Euphorbia inundata is found in open, generally treeless low swales within 
wetland longleaf pine savannas. Associated species include Aletris lutea, Calo- 
pogon pallidus, Centella asiatica, Chaptalia tomentosa, Ctenium aromaticum, 
Dichromena latifolia, Drosera capillaris, D. tracyi, Eriocaulon compressum, E. 
decangulare, Helianthus heterophyllus, Hypericum myrtifolum, Lophiola au- 
rea, Lycopodium alopecuroides, Rhezia alifanus, R. lutea, Rhynchospora bald- 
winti, R. oligantha, R. plumosa, Sarracenia alata, S. psittacina, Scleria bald- 
win and Xyris baldwinzana. 

Euphorbia inundata is nearly restricted to Florida, where it occurs in the 
Jacksonville area, along the west coast of the peninsula and in the panhandle 
(Michael Huft, pers. comm.). It is perhaps most frequent in the Apalachicola 
lowlands from Wakulla to Bay counties, Florida. The nearest records to Mis- 
sissippi are from Baldwin County, Alabama. The only other specimens of E. 
inundata outside of Florida are from Charlton and Ware counties, Georgia 
(Michael Huft, pers. comm.). It occurred with 10% frequency in sample plots 
on the Rains (Typic Paleaquults) soil series in Mississippi (Norquist 1984). 
Kral (1983) contrasts it with E. telephoides Chapm., a local Apalachicola 
endemic, and notes that E. inundata is usually found in moist to wet pine 
flatwoods savannas and borders of Hypericum dominated ponds. The author 
has noted E. inundata to be fairly frequent in wetland longleaf pine savannas 
in Liberty and Franklin counties, Florida; however, it is considered rare in the 


Pensacola region (Wilhelm 1984). 


Bridges: New Carez and Euphorbia in Mississippi 219 


ACKNOWLEDGMENTS 


Special thanks to Michael Huft of the Field Museum of Natural History 
for sharing his unpublished information on Euphorbia inundata. Thanks to 
reviewers Guy Nesom and Carol Todzia for helpful comments. Publication 
costs were provided by the Plant Resources Center of the University of Texas 
at Austin. 


LITERATURE CITED 


Kral, R. 1983. A report on some rare, threatened, or endangered forest- 
related vascular plants of the South. USDA, Forest Service, Technical 
Publ. R8-TP2. 1305 pp. 


Norquist, S.C. 1984. A comparative study of the soils and vegetation of savan- 
nas in Mississippi. M.S. thesis, Mississippi State University, Mississippi 
State. 110 pp. 


Wilhelm, G.S. 1984. Vascular flora of the Pensacola Region. Ph.D. diss., 
Southern [llinois University. 213 pp + bibliography. 


Phytologia (September 1989) 67(3):220-226. 


REFUTATION OF RECENT CREATIONS OF 
MICROSPECIES AND HYBRID TAXA IN 
ARGENTINIAN SOLANUM (SECT. PETOTA) 


H. Enrique Brucher 
Cas. correo 131, Centro Reg. Investigaciones Cientificas, 5500 Mendoza, 


ARGENTINA 


ABSTRACT 


Arguments are presented against taxonomic splitting within Solanum 
from Argentina. 


KEY WORDS: Solanum, Solanaceae, evolution, hybrids, Argentina. 


In recent years, we have been confronted with the splitting of well estab- 
lished taxonomic units of Argentinian wild potatoes and the proliferation of 
so-called “spec. nov.” of hybridogen origin. This deplorable tendency has 
created further problems for the understanding of the speciation of Solanum, 
a genus which traditionally suffers “per se” evolutionary and nomenclature 
problems. 

When working with wild potatoes, one should always bear in mind the 
biological fact that the majority of diploid tuber bearing species are necessarily 
outbreeders. Consequently, their sexually produced offspring are not identical, 
but show (more or less) morphological and physiological segregation, which in 
extreme cases may simulate speciation. Their self incompatibility is based 
on a rather simple S-allele scheme (oppositional factors of multiple genes, at 
the same chromosome locus) which inhibits self fertilization. This important 
fact should never be overlooked by wild potatoes collectors and should be 
borne in mind by taxonomists in their zeal to immortalize their names in 
the description of new species. Unfortunately, this problem arose recently 
in Argentina. Members of several series of tuber bearing Solanum, such as 
Acaulia, Cuneoalata, Megistacroloba and Tuberosa were involved in an undue 
“name creation.” We hope that those procedures may be discontinued. With 
regard to this, we may call attention to some recent publications (Okada & 
Hawkes 1978; Okada & Clausen 1982; Okada & Clausen 1985). 

In sect. Petota, there is a “two way system of propagation:” sexual repro- 
duction and vegetative clonal proliferation. To a certain degree, these have 
opposing effects on the frequency of genomic variability. Whereas sexual re- 
production encourages a limited genetic segregation and diversity of given wild 


220 


Bricher: Refutation of microspecies in Argentinian Solanum 221 


potato species, asexual reproduction has a more conservative effect. With the 
former process, self sown seedlings may cause a continuous slight segregation 
inside the offspring generation, apparently leading to “microspecies,” while 
the latter process of asexual-clonal reproduction allows survival of a chain of 
milieu adapted hybrid plants and many nothomorphic forms. This was already 
explained by Briicher in 1953 (the first case for Argentina). 

Even genetically infertile Petota hybrids, or sterile introgression biotypes 
can maintain, by asexual reproduction, their identity for a long time, perhaps 
for ages. 

Of course, between segregations and hybrids of this sort, neither can claim 
higher taxonomic status, nor do they deserve-in the light of a modern biolog- 
ical species concept (Love 1964)-any botanical name. Hawkes f.e. was quite 
right when he once expressed: “. . If these hybrids are to be named at 
all, they should be considered merely as nothomorphic forms of one species” 
(1963 p. 155). After having rejected his earlier, now outdated species concept, 
Hawkes wrote recently (1989 p. 58): “. . .in general the wildpotato species 
should be regarded as larger units that contain a wide range of genetic diver- 
sity. . .Such concept made it inadvisable, if not impossible, to divide potato 
species into the conventional infraspecific categories of varieties and forms.” 
Similar thoughts have been repeatedly expressed by other solanologists (Cor- 
rell 1962; Danert 1962; Dodds 1965; Ochoa 1984). 

Therefore, we wonder why there exists such a wide gap between good 
theories and objectionable practice related to Argentinian wild potatoes. 


PART I 
a) Hybrid: Solanum reche: Hawkes & Hjerting 


This “species” has a rather curious history, which began 60 years ago when 
the botanist Dr. Castellanos collected, in a farmer’s settlement (Guanchin 
Viejo) situated in the semiarid Dept. Chilecito (Prov. La Rioja), an apparent 
wild potato (now included in the herbarium of the Museum B. Rivadavia, BA, 
as number 28/345 BA). The locality, at 29° S, 67° 38’ W, is easily accessible 
by a side road, 10 km from the town of Chilecito. The plant was claimed 
by Hawkes & Hjerting (1963) to be a new species. It received the binomial 
Solanum rechet, so called in attention to a friend of the latter author, who was 
at that time a merchant in Tucuman. 

The short species diagnosis (1963 p. 146) claims a close phylogenetic affinity 
to Solanum maglia, a true species of long standing, which grows on the Pacific 
coast of the Chilean Republic. 

The authors considerably underestimated the geographic distance (700 km) 
between the “loci classic?” of the two epithets. Possibly, the similarity of 
names, Chile-Chilecito (which means little Chile) interfered subconsciously in 


222 PERT OL OG TA volume 67(3):220-226 September 1989 


the minds of the foreigners who created this new species and committed the 
incredible gaffe of declaring Solanum rechei as the ancestral form of Solanum 
maglia, a species which lives on the Pacific coast, in Chile, separated from the 
former by the high Cordillera. 

The term S. reche: has been created under the erroneous assumption of 
representing another endemic species of the Sierra Famatina (l.c. Bricher 1959; 
1965), a very extended mountain chain of difficult access, with a maximum 
elevation of 6200 m. 

Furthermore, the statement in the description that indicates that S. rechez 
occurs on several distant places of dry valleys in the province of La Rioja 
is not true. The truth, as we found out during several collecting trips to 
La Rioja, is that the so called S. reche: has a narrowly restricted manmade 
habitat in orchards of irrigated agricultural farms, called Guanchin Viejo, Las 
Tablas, Trapiche and a picnic place on the local automobile road from Chilecito 
to Mina de Oro. Our checking disproved completely, the pretention of the 
foreigners to have discovered a new species of Solanum. The results of several 
days inspection of the original locality and a comprehensive plant collection 
of more than 100 samples (available at our private herbarium in Mendoza), 
indicate that these “papas malezas” (= weed potatoes), as the local people 
quite correctly defined them, are the remainders of an earlier hybridization 
between Solanum microdontum (sensu lat.) and Solanum kurtzianum, two 
well known wild potatoes. 

Although this has been already published in two short notices in German in 
1969 and 1974, the authors of Solanum rechez did not eliminate their erroneous 
epithet, while they partially admitted their error and recognized the poor 
taxonomic value of this name in their book on Argentine wild potatoes (Hawkes 
& Hjerting 1969). But the last publication on the subject (Okada & Hawkes 
1979) restores the erroneous name with the statement “Solanum reche1 Hawkes 
& Hjerting es una papa silvestre. . .” 

To maintain a “holotype” of S. rechez is misleading because Nr. 28/345 at 
BA is a casually segregated individual from a huge hybrid swarm of uncounted 
different morphotypes. We discovered this when we collected at random, hun- 
dreds of plants in the orchards and at irrigation ditches of Guanchin Viejo. 
None of the plants matched the others. Neither should they be named “F, 
plants” (Hawkes) because there is no proof of how and when such hybrid F, 
populations arose. Keeping in mind that these orchards have been kept under 
irrigation for hundreds of years, and are plowed and harvested each year, the 
tubers of these hybrids swarms of F,, origin have been mixed and diffused by 
repeated cleaning and weeding of the irrigation ditches. 

The following table is a testimony to the heterogeneity of such “papas 
malezas.” This is only an abbreviated extract from our findings which shall 
be presented “zn ezrtenso” on another occasion. 

Publication of Part II follows in short. 


Bricher: Refutation of microspecies in Argentinian Solanum | 223 


Table 1 
Nr. of Plant Height Length X Width of Quantity of Stem Flowers! 
Sollection in cm Terminal Leaflet lat? int? Wings 
in mm Ifits Ifits 
1400 32 60 X 45 2 4 yes # 
1401 36 75 X 30 3 4 no & 
1402 40 40 X 25 4 6 no & 
1403 45 45 X 30 3 10 no & 
1404 40 30 X 22 4 al no & 
1405 30 40 X 20 4 4 no & 
1406 40 50 X 20 4 8 yes & 
1407 oe 45 X 22 1 6 no % 
1408 28 60 X 30 2 3 no # 
1409 22 40 X 18 5 8 no # 
1410 60 44 X 20 3 6 no # 
1411 18 40 X 20 2 2 no # 
1412 30 45 X 35 3 2 yes # 
1413 38 60 X 40 Z ?4 yes # 
1414 100 130 X 45 2 0 yes # 
1415 50 70 X 28 3 3 yes # 
1416 38 90 X 55 1 0 no # 
1417 25 80 X 60 1 0 no # 
1418 20 60 X 40 1 0 no # 
1419 16 70 X 55 2 0 no # 
1420 80 65 X 40 3 fe yes # 
1421 120 85 X 60 3 5 yes & 
1422. 90 70 X 45 3 5 no # 
1423 108 65 X 30 4 7 no # 
1424 110 65 X 25 4 6 yes # 
1425 95 70 X 30 3 4 yes % 
1426 90 60 X 35 3 4 yes # 
1427 98 70 X 40 4 8 yes % 
1428 80 80 X 40 3 5 yes % 
1429 100 110 X 450 2 2 yes & 
1430 120 95 X 335 3 4 yes & 
1431 100 90 X 35 4 8 yes & 
1432 110 65 X 25 Q 5 yes % 


2240 PeRYOT OD O'G TA volume 67(3):220-226 September 1989 


Table 1 (cont.) 


Nr. of Plant Height Length X Width of Quantity of Stem Flowers! 
Collection in cm Terminal Leaflet lat? int? Wings 
in mm Ifits fits 
1433 90 50 X 20 4 3 no # 
1434 100 70 X 30 3 8 yes % 
1435 110 60 X 30 t FS yes % 
1436 120 55 X 25 3 (4 yes # 
1437 120 70 X 35 4 12 yes 3 
1438 60 60 X 35 3 5 no # 
1439 100 85 X 45 4 6 no # 
1440 90 70 X 30 4 3 yes % 
1441 120 65 X 25 4 7 yes & 
1442 22 70 X 30 3 3 yes # 
1443 28 45 X 25 2 0 yes # 
1444 25 40 X 18 2 0 yes # 
1445 70 90 X 35 2 0 yes % 
1446 80 100 X 40 Z 7 no # 
1447 90 90 X 40 3 2 yes & 
1448 20 80 X 40 2 3 yes # 
1449 120 90 X 60 3 6 no # 
1450 100 100 X 60 2 3 no # 
1451 130 75 X 33 3 7 yes # 
1452 140 115 X 45 4 6 yes # 
1453 80 70 X 35 3 3 yes # 
1454 90 65 X 30 ze 3 no % 
1455 28 55 X 25 3 4 yes # 
1456 120 95 X 45 4 6 yes # 
1457 100 100 X 55 3 5 yes & 
1458 110 90 X 45 3 9 yes # 
1459 130 90 X 35 3 4 yes & 
1460 110 90 X 50 2 6 no # 
1461 80 75 X 35 3 0 no % 
1462 70 75 X 45 3 3 no # 
1463 90 85 X 40 3 if no # 
1464 100 110 X 55 3 6 no # 
1465 120 80 X 35 4 5 yes # 
1466 25 55 X 40 if 0 no # 
1467 12 50 X 30 1 0 no # 
1468 15 55 X 25 2 0 yes # 


Bricher: -  Refutation of microspecies in Argentinian Solanum 225 


Table 1 (cont.) 


Nr. of Plant Height Length X Width of Quantity of | Stem Flowers! 

Collection in cm Terminal Leaflet lat? “int? Wings 
in mm Ifits lflts 

1469 20 75 X 50 ear no # 

1470 lis 50 X 30 1 0 yes # 

1471 4p, ati hn tQcknds eect no 2 

1472 20 40 X 30 3 0 yes # 

1473 30 60 X 35 2 1 yes # 

1474 20 60 X 45 2 <0 yes # 

1475 iP} 55 X 35 1 0 yes # 
“Holotype” 

345 = 70 X 30 3 1 yes & 


1 & stands for present; % stands for aborted; # stands for absent. 
? lateral. , 
3 interject. 


LITERATURE CITED 


Bricher, H. 1953. Uber das nattrliche Vorkommen von Hybriden zwischen 
Solanum simplictfolium and S. subtilius im Aconquija. Gebirge. Zt. f. 
Vererbl. 85:12-19. 


1959. Kritische Betrachtungen zur Nomenklatur argentinischer 
Wildkartoffeln. V. Die Seria Acaulza. Zuchter 29:149-156. 


. 1965. Solanum kurtzianum, eine nematodenresistente Wildkartof- 
fel der argentinischen Halbwiiste. Rev. Botanica Kurtziana, Cordoba. 
21:212-215. 


—_____. 1974. Uber Art-Begriff und Art-bildung bei Solanum (sect. Tuber- 
arium). Beitr. Biol. Pflanzen 50:393-429. 


Correll, D. 1962. The Potato and its Wild Relatives. Texas Research Foun- 
dation, Renner, TX. 


Danert, S. 1962. Morphologie und Anatomie von Solanum tuberosum. in 


Schick & Klinkowski (eds.) Die Kartoffel VEB Vig. Berlin. — 


226 PAY T ODO LA volume 67(3):220-226 September 1989 


Dodds, K. 1965. The history and relationship of cultivated potatoes, in: 
Essay on Crop Plant Evolutzon. Cambridge. 


Hawkes, J.G. 1963. A revision of the tuber bearing solanums- Rec. Scott. Pl. 
Breed. St. 76-181. 


Hawkes, J.G. & J.P. Hjerting. 1969. The Potatoes of Argentina, Brazil, 
Paraguay and Uruguay. Clarendon Press, Oxford. 


. 1989. The Potatoes of Bolivia. Clarendon Press, Oxford. 


Love, A. 1964. The biological species concept and its evolutionary structure. 
Taxon 13:33-45. 


Ochoa, C. 1984. Karyotaxonomic studies on wild Bolivian tuber-bearing 
Solanum, sect. Petota. Phytologia 55:17-40. 


Okada, K.A. & J.G. Hawkes. 1978. Solanum X rechei, especie silvestre de 
papas de origen hibrido de la Sierra de Famatina (prov. de La Rioja, 
Argentina). Kurtziana 11:55-74. 


Okada, K.A. & A.M. Clausen. 1982. Natural hybridization between Solanum 
acaule Bitt. and S. megtstacrolobum Bitt. in the province of Jujuy, Ar- 
gentina. Euphytica 31:817-835. 


—______. 1985. Natural triploid hybrids between Solanum acaule Bitt. and 
S. infundibuliforme Phil. in the province of Jujuy, Argentina. Euphytica 
34:219-231. 


Phytologia (September 1989) 67(3):227-234. 


GENERIC AFFINITIES AND TYPIFICATION OF ELEVEN SPECIES EXCLUDED 
FROM SIPHONOGLOSSA OERST. (ACANTHACEAE) 


Richard A. Hilsenbeck 
Department of Biology, Sul Ross State University, Alpine, Texas 79832 USA 


ABSTRACT 


The genus Siphonoglossa Oerst., as recently delimited, includes only 
ten taxa belonging to the typical section. This delimitation of Siphono- 
glossa sensu stricto is warranted because several misconceptions con- 
cerning the vegetative and pollen morphology of the genus have been 
disclosed. Of the taxa excluded from the genus, most may properly be- 
long to the closely related genus Justicia L., while others do not belong 
to the same tribe as Siphonoglossa and Justicia. 


KEY WORDS: Siphonoglossa, Justicia, Acanthaceae, pollen, typ- 
ification, systematics. 


The genus Siphonoglossa, previously comprising 25 species, has long been 
subject to erroneous concepts as to its proper delimitation. As such, it became 
an artificial assemblage of species having little affinity to each other or to 
the original concept of the genus as based on the generitype (Oersted 1854). 
Several of these species do not even belong together at the tribal (or subtribal) 
level. This was shown by Hilsenbeck (1983), who narrowed the concept of 
Siphonoglossa to ten taxa in seven species belonging to the strictly New World 
type section (see Henrickson & Hilsenbeck 1979). It is therefore necessary to 
exclude the remaining 15 taxa from the genus, and this study concerns the 
treatment of 11 of these. The purpose of this paper, then, is to present the 
generic and tribal affinities of the taxa erroneously classified in Stphonoglossa 
and to discuss their typification since some of the types were either destroyed 
or not designated. Additionally, it is felt that considerable nomenclatural and 
taxonomic confusion surrounding the genus will be significantly reduced by 
the following account. 

Six of the species here excluded from Szphonoglossa are African in distri- 
bution and, though belonging to the same tribe and subtribe as Szphonoglossa 
(Justicieae Lindau, Justiciinae Bremekamp), clearly do not belong to Siphono- 
glossa as recently delimited. The six species have an equally 5 parted calyx 
and conspicuously appendaged anther sacs, among other characters, which 


225 


Zee Pe TOL OOCTA volume 67(3):227-234 September 1989 


strongly ally them with Justicia and the taxa referred to Siphonoglossa sec- 
tion Pentaloba Hilsenbeck (in Henrickson & Hilsenbeck 1979). Because the four 
taxa of Siphonoglossa section Pentaloba |type, S. pilosella (Nees) Torrey] pos- 
sess morphological, chemical and cytological characteristics of Justicia rather 
than Siphonoglossa, they are to be formally transferred to Justicza ( Hilsenbeck 
1983; 1989a) and will not be dealt with further here. 

All of the above taxa were placed in Szphonoglossa under what I have 
termed the artificial “Torreyan concept” of the genus as discussed elsewhere 
(Hilsenbeck 1983; 1989a). In short, Torrey (1859) widened considerably the 
limits of Siphonoglossa (type, S. ramosa Oerst.) by transferring into the genus 
an American species known for years to European botanists as Adhatoda 
dipteracantha Nees (= Monechma pilosella Nees), calling it S. pilosella. To 
accommodate this species, Torrey had to enlarge Oersted’s original concept of 
Stphonoglossa to include characters that traditionally define the genus Justicia 
(see Leonard 1958; Long 1970). The inclusion in Siphonoglossa of S. pilosella, 
and later the African species S. tubulosa (E. Meyer) Bentham & Hooker [= 
Adhatoda tubulosa (E. Meyer) Nees] by Bentham (in Bentham & Hooker 1886), 
set a precedent for other taxonomists, particularly Moore, to classify elements 
in the genus based primarily on direct comparisons with S. pilosella (not S. 
ramosa) as representative of the genus. For example, in his description of S. 
rubra, Moore (1906), employing the Torreyan concept of the genus, states, 
“This plant. . . found in Tropical Africa, is quite unlikely any of its American 
and South African congeners. The flower has been compared carefully with 
that of S. pilosella Torr., and found to agree with it in all essentials of generic 
nature.” 

Excluded also here are five New World species placed in Szphonoglossa by 
Lindau. All of these species were so classified based on Lindau’s misconception 
of the pollen morphology of the group as discussed elsewhere (Hilsenbeck 1983; 
1989b). These five species properly belong in the tribe Odontonemeae Lindau, 
not in Justicieae, or following Bremekamp’s (1965) system, they would be 
placed in Justicieae, subtribe Odontoneminae. For example, on transferring 
Carlowrightia pringlet Robins. & Greenm. to Siphonoglossa, Lindau (1897) 
states that the species, “ist eine typische Siphonoglossa wie aus der Form 
der Corolla und dem Pollen (Spangenpollen) hervorgeht.” Spangenpollen is 
pollen that is basically prolate and tricolporate and is generally characteristic 
of Odontoneminae. The taxa of Justiciinae, including Siphonoglossa sensu 
stricto (Hilsenbeck 1983), are characterized by pollen that Lindau (1894; 1895) 
termed Knotchenpollen. This latter type of pollen is most often bilateral and 
2 porate. 

Therefore, all 11 species here excluded from Siphonoglossa were placed in 
the genus either upon: 1) the very broad and artificial “Torreyan concept” 
and are, for the most part, thus referable to Justicia or, 2) a misconception 
of pollen morphology and as such belong to a different tribe (or subtribe) of 


Hilsenbeck: Generic affinities of exclusions from Siphonoglossa 229 


Acanthaceae. I will not make the generic transfer of any of these species at this 
time, because I feel that they should be accorded more detailed study prior to 
their formal inclusion within other genera. It is clear, however, that these 11 
species do not belong to Siphonoglossa as based on the type, S. ramosa. 


EXCLUDED SPECIES 


Siphonoglossa gentianifolia Lindau, Bull. Herb. Boissier 2:370. 1905. TYPE: 
PARAGUAY: Gran Chaco prope Santa Elisa ad marginem silvarum, w/o 
date, Hassler 2841 (HOLOTYPE: B, destroyed; Phototypes: GH!,NY!). 


In the description of this taxon Lindau states, “pollinis granula, subglobosa, 
typica” in direct reference to this species having Spangenpollen which Lindau 
mistakenly believed typical for the genus. Judging from the description and 
type photographs, this species is very closely related to another of Lindau’s 
“siphonoglossas,” as he states, “Verwandt mit S. sulcata (Nees) Lindau, aber 
durch die grosseren und viel breiteren Blatter, die kurzeren Bluten und die 
breiteren Brakteen sofort zu unterscheiden.” The proper generic disposition 
of this taxon, of which the only material I have seen are the photos, will 
be discussed under S. sulcata. I have been unable to locate other specimens 
of Hassler 2841 on which to base a lectotype, and in the absence of other 
authentic material I cannot properly designate a neotype. 


Stphonoglossa glabrescens Lindau, Bull. Herb. Boissier 2:546. 1894. TYPE: 
MEXICO, Oaxaca: distr. Tlacolula, prope Zoquitlan, Jun 1888, Seler 
76 (HOLOTYPE: B, destroyed; Phototypes: GH!,MICH!,NY!). 


That this species is not a Siphonoglossa is plainly evident. The pollen 
was described by Lindau as ”pollinis granula typica” again in allusion to its 
prolate, tricolporate nature (i-e., Spangenpollen). From the description and 
the photo of the type, it is clear that this taxon is in reality Anzsacanthus 
quadrifidus (Vahl) Nees. Among other features, the species has a red corolla, 
the morphology of which is that of Anisacanthus, and conspicuously exfoliating 
bark also characteristic of that genus. 


Stphonoglossa (?) linifolia (Lindau) C.B. Clarke, in W.T. Thiselton-Dyer, 
ed., Flora Capensis vol. 5, sect. 1:75. 1912. Aulojusticza linifolia Lin- 
dau, Bot. Jahrb. Syst. 24:325. 1898. LECTOTYPE (here chosen): 
SOUTH AFRICA: Kalahari Region, Transvaal, mountain sides of Sad- 
dleback Range, near Barberton, 22 Feb 1890, E. E. Galpin 825 (BOL!; 
Isolectotypes: NBG!,US!). 


230 PUY re owo GLA volume 67(3):227-234 September 1989 


The holotype, Galpin 825 at Berlin, was destroyed. Therefore, Galpin 825 
at BOL, an isotype, is designated as lectotype. The only vague resemblance 
that this South African species bears to Siphonoglossa is a very long corolla 
tube. The calyx is equally 5 parted, the anthers are conspicuously appendaged, 
and in habit and characters of the fruit, this taxon is not congeneric with 
Siphonoglossa. In transferring this species to Szphonoglossa, Clarke placed a 
question mark between the generic and specific epithets and earlier states, 
“The question is greatly complicated by the arrival of a third South African 
species (Aulojusticia lunifolia) which has the corolla of Beleropone (he must 
have meant Beloperone), not of Siphonoglossa.” Even as widely as Lindau 
stretched the generic boundaries of Siphonoglossa, he did not place this species 
in it, instead erecting a new genus, Aulojusticia, to accommodate it. ] am in 
favor of leaving this distinctive species in Lindau’s monotypic genus until future 
study can perhaps better determine its generic affinities. 


Siphonoglossa macleodiae S. Moore, in Macleod, Chiefs & Cities Centr. Afr. 
304. 1912. TYPE: NIGERIA: N Nigeria, River Benue, Sep 1910, P. A. 
Talbot s.n. (HOLOTYPE: BM!; Isotype: MO!). 


This species clearly belongs to Justzcza subgenus Hujusticia, near sections 
Adhatodaand Tyloglossa of Lindau (1895), because of its flowers borne solitary 
and sessile in the leaf axils, equally 5 parted calyx, and spurred anther sacs. 
An illustration of the pollen is affixed to the holotype showing that this species 
has Knotchenpollen. 


Stphonoglossa migeodii S. Moore, J. Bot. 67:271. 1929. TYPE: TANZANIA: 
Tanganyika Terr., w/o date, F.W.H. Migeod 187 (HOLOTYPE: BM!). 


From all appearances, the relationships of this species clearly lie with the 
other African elements which have been mistakenly included in Stphonoglossa. 
It may be that the only extant material of this species is that of the type 
collection and two paratypes (Migeod 478, BM) as I have not seen any other 
specimens of this taxon, even from the South African herbaria from which I 
borrowed material. As with S. macleodiae, the proper classification of this 
little known species should await further study directed primarily at these 
Old World taxa. It appears, however, that this species has affinities with 
Aulojusticia linifolia through its corolla, inflorescence and fruit morphology, 
as well as with Justicia in its 5 parted calyx and conspicuously spurred lower 
anther sac. 


Stphonoglossa nummularia S. Moore, J. Bot. 18:40. 1880. TYPE: SOUTH 
AFRICA: “British Kaffraria,” 1860, T. Cooper 370 (HOLOTYPE: K!). 


As noted above, Moore accepted, followed, and even expanded upon Tor- 
rey’s artificial concept of the genus. Indeed, S. nummularia with its 5 parted 


Hilsenbeck: Generic affinities of exclusions from Siphonoglossa 231 


calyx and spurred lower anther sacs fits well within Justicia, not Siphonoglossa. 
Its most closely allied taxa appear to be the other South African “siphonoglos- 
sas” and section Pentaloba of Siphonoglossa which is currently being trans- 
ferred to Justicia (Hilsenbeck 1989a). 


Siphonoglossa peruviana Lindau, Bot. Jahrb. Syst. 42:173. 1908. TYPE: 
PERU. Amazonas: Prov. Chachapoyas, dstliche Talwand des Marafon 
uber Balsas, w/o date, A. Weberbauer 4269 (HOLOTYPE: B, destroyed; 
Phototypes: GH!,NY!). 


As with the other taxa placed in Siphonoglossa by Lindau, this species 
has Spangenpollen and thus more properly belongs in Odontonemeae. The 
type material unfortunately has been destroyed and J cannot locate any other 
material of Weberbauer 4269 with which to lectotypify this species. On initial 
inspection, it appears that S. peruviana may belong in the genus Yeatesza, 
having a very similar overall inflorescence and corolla morphology to this genus 
(Hilsenbeck 1989c). From the original and rather detailed description of the 
fruit and seeds, and the type photos, however, this species undoubtedly belongs 
in Tetramerium and Daniel (1986) has recently and correctly made the formal 
transfer. 


Stphonoglossa pringle: (Robins. & Greenm.) Lindau, Bull. Herb. Boissier 
5:622. 1897. BASIONYM: Carlowrightia(?) pringle: Robins. & Greenm., 
Proc. Amer. Acad. Arts 32:40. 1896. TYPE: MEXICO. Oaxaca: dry 
slopes Tomellin Canon, 30 Nov 1895, C.G. Pringle 6261 (HOLOTYPE: 
GH!; Isotype: CAS!). 


This species has Spangenpollen and as such should be in the Odontone- 
meae, well removed from Siphonoglossa. In his monographic treatment of 
Carlowrightia, Daniel (1980) included this species under Carlowrightia where 
it correctly belongs. 


Siphonoglossa rubra S. Moore, J. Bot. 44:88. 1906. TYPE: UGANDA: 
Entebbe, w/o date, Bagshawe 750 (BM). 


Although Moore states that the type of this species is at BM, I did not 
receive any material of it in aloan from BM that contained the holotypes of two 
other African species placed by Moore in Siphonoglossa. I have not seen, nor 
do I know the location of any material of Bagshawe 750 and have, therefore, 
been unable to properly lectotypify the species. I have, however, examined 
several other specimens of this seemingly polytypic, red flowered species. As 
stated above, Moore carefully compared the flowers of S. rubraand S. pilosella 
and found them to agree “in all essentials of generic nature.” It is indeed true 
that these two taxa have a similar “flower” morphology, including androecium 
structure, an equally 5 parted calyx and other features in common. However, 


232 PHYTOL OG hA volume 67(3):227-234 September 1989 


in characters of the inflorescence, corolla, fruit and seed, S. rubra more closely 
resembles the widespread tropical African and Asian Rhinacanthus nasutus 
(L.) Lindau (= R. communis Nees). Though clearly not a Siphonoglossa, it 
perhaps does belong in RAznacanthus, but as with the other African taxa herein 
discussed, its proper generic disposition should await further investigation. 


Stphonoglossa sulcata (Nees) Lindau, Bot. Jahrb. Syst. 48:19. 1894. BA- 
SIONYM: Jacobinia sulcata Nees in DC., Prodr. 11:333. 1847. Dr- 
anthera sulcata (Nees) Griseb., Goett. Abh. 19:224. TYPE: ARGENTI- 
NA: Rio Parana, w/o date, Tweedie s.n. (HOLOTYPE: K!). 


This species is clearly not related to Siphonoglossa but instead has a very 
close affinity to the genus Yeatesza Small, tribe Odontonemeae, of the south- 
ern United States and adjacent northeastern México (Hilsenbeck 1983; 1986c). 
This species and S. gentzanzfolia possess Spangenpollen and also have inflo- 
rescences, corollas, androecia, fruits and seeds characteristic of Yeatesza and 
the Old World genus Ecbolium and are clearly most closely related to (besides 
each other) these two genera. The proper generic classification of these species 
is under active consideration. 


Stphonoglossa tubulosa (Nees) Bentham & Hooker, Gen. Pl. 2:1110. 1886. 
This combination should properly be S. tubulosa (E. Meyer) Bentham 
& Hooker, but is cited here as listed by Bentham & Hooker (1886). 
BASIONYM: Justicia tubulosa E. Meyer in Drege, Zwei Pflanzengeogr. 
Documente 150,196. 1837. Rhinacanthus tubulosus (E. Meyer) Presl, 
Bot. Bemerk. 95. 1843. Adhatoda tubulosa (E. Meyer) Nees in DC., 
Prodr. 11:392. 1847. LECTOTYPE (here chosen): SOUTH AFRICA: 
Pondoland between St. Johns River and Umtsikaba River, 1837, S.F. 
Drege s.n. (K!; Isolectotypes: K!,MO!). 


Justicia suffruticosa E. Meyer in Drege, Zwei Pflanzengeogr. Documente 153, 


196. 1837. | 
Justicia prostrata Schlechtend. er Nees in DC., Prodr. 11:390. 1847. 


Gendarussa leptantha Nees, Linnaea 15:372. 1841. Adhatoda leptantha (Nees) 
Nees in DC., Prodr. 11:392. 1847. Justicia leptantha (Nees) Lindau in 
Engl. & Prantl, Naturl. Pflanzenfam. 4, 3b:349. 1895. 


Of the three sheets of this collection at K, two have been annotated by 
Nees von Esenbeck, and of these two, the most complete specimen has been 
chosen as lectotype. I think that this species most properly belongs in Justicia 
or in Adhatoda, if one accepts the latter genus. It is notable that Nees (1847) 
placed this species in the same genus with Adhatoda hyssoptfolia (L.) Nees (= 
Justicia hyssopifolia L.) and Adhatoda dipteracantha Nees [= Siphonoglossa 
pilosella (Nees) Torrey]. Justicia hyssoptfolia is one of the two proposed lec- 
totypes of the genus Justzcza (Stearn 1971). This points clearly to the close 


Hilsenbeck: Generic affinities of exclusions from Siphonoglossa 233 


affinity of S. pilosella, not only with Justicia (and Adhatoda), but with the 
African taxa improperly placed in Siphonoglossa. An instructive commen- 
tary concerning the generic status of this species was that of Clarke (1912). 
He states, “S. tubulosa was removed from Justicza to the American genus 
Stphonoglossa by Bentham (Benth. et Hooker, f. Gen. Pl. ii. 1110). S. Moore 
added S. nummularia which is beyond question congeneric with S. tubulosa. 
Baillon (Hist. de Plantes, X. 441) records S. tubulosa under Siphonoglossa but 
does not appear to have examined or considered it. Lindau (in Engl. & Prantl., 
Pflanzenfam. IV. 3B, 338) says that these two species can scarcely be referred 
to Siphonoglossa and (l.c. p. 349) records S. tubulosa (under a different name) 
as a true Justicia.” I agree with Clarke that S. tubulosa and S. nummularia 
are closely related, if not congeneric. I also agree with Lindau that these taxa 
should be excluded from Siphonoglossa (but for a different reason) and placed 
in Justicia, the former as J. tubulosa E. Meyer. 


ACKNOWLEDGMENTS 


I express my thanks to Marshall C. Johnston, A.M. Powell, B.L. Turner 
and Beryl B. Simpson for their advice and assistance in the preparation of this 
manuscript and to the curators of the various herbaria mentioned in the text. 


LITERATURE CITED 


Bentham, G. 1886. Acanthaceae. In: G. Bentham & J. D. Hooker, Gen. PI. 
2:1060-1122. 


Bremekamp, C.E.B. 1965. Delimitation and subdivision of the Acanthaceae. 
Bull. Bot. Surv. India 7:21-30. 

Clarke, C.B. 1912. Acanthaceae. Jn: W.T. Thiselton-Dyer, Flora Capensis 
5, sect. 1:1-92. 


Daniel, T.F. 1980. A systematic study of the genus Carlowrightia (Acan- 
thaceae). Ph.D. dissertation, Univ. of Michigan, Ann Arbor. 


1986. Systematics of Tetramerium (Acanthaceae). Syst. Bot. 
Monogr. Vol. 12:1-134. 


Henrickson, J. & R.A. Hilsenbeck. 1979. New taxa and combinations in 
Siphonoglossa (Acanthaceae). Brittonia 31:373-378. 


Hilsenbeck, R.A. 1983. Systematic studies of Szphonoglossa sensu lato. Ph.D. 
dissertation, Univ. of Texas, Austin. 


234 PHYTOLOGIA volume 67(3):227-234 September 1989 


. 1989a. Systematics of Justicia section Pentaloba (Acanthaceae). 
Pl. Syst. Evol., (in press). 


. 1989b. Pollen morphology and systematics of Siphonoglossa sensu 
lato (Acanthaceae). Amer. J. Bot., (in press). 


. 1989c. Taxonomy of Yeatesza (Acanthaceae). Syst. Bot. 14:427- 


438. 


Leonard, E.C. 1958. The Acanthaceae of Colombia. Contr. U.S. Natl. Herb. 
31:1-781. 


Lindau, G. 1894. Beitrage zur Systematik der Acanthaceen. Bot. Jahrb. 
Syst. 18:36-64, pls. 1, 2. 


. 1895. Acanthaceae. Jn: A. Engler & K. Prantl, Nat. Pflanzenfam. 
IV (3b):274-354. 


. 1897. Acanthaceae Americanae et Asiaticae. Bull. Herb. Boissier 
5: 643-681. 


Long, R.W. 1970. The genera of Acanthaceae in the southeastern United 
States. J. Arnold Arbor. 51:257-309. 


Moore, S. 1906. Uganda Gamopetalae from Dr. Bagshawe. J. Bot. 44:83-90. 


Nees von Esenbeck, C.G. 1847. Acanthaceae. Jn: A. DeCandolle, Prodr. 
11:46-519. 

Oersted, A.S. 1854. Mexicos og Centralamerikas Acanthaceer. Vidensk. Med- 
del. Dansk Naturhist. Foren. Kjgbenhavn. 6:113-181. 


Stearn, W.T. 1971. Taxonomic and nomenclatural notes on Jamaican Gamo- 
petalous plants. J. Arnold Arbor. 52:614-647. 


Torrey, J. 1859. Botany of the Boundary. Jn: Report of the United States 
and Mexican Boundary Survey, under the direction of W.H. Emory. 


Phytologia (September 1989) 67(3):235. 


CHANGE OF NAME IN A TUBER BEARING SOLANUM 
C.M. Ochoa 

International Potato Center, P.O. Box 5969, Lima, PERU 
ABSTRACT 


Solanum chillonanum Ochoa is proposed as an avowed substitute 
for Solanum tenellum Ochoa, which is a later homonym of S. tenellum 
Bitter. 


KEY WORDS: Solanaceae, Solanum, nomenclature. 


Since the specific epithet tenellum that I recently published was already 
in use in Solanum by Bitter, I wish to replace my later homonym with the 
following combination. 


Solanum chillonanum Ochoa, nom. nov. = Solanum tenellum Ochoa, Phy- 
tologia 63:455. 1987. (non S. tenellum Bitter, Repert. Spec. Nov. Regni 
Veg. 11:219. 1912). 


235 


Phytologia (September 1989) 67(3):236. 


CORRECTIONS OF NEW COMBINATIONS IN EUPHORBIA 


Rob C.H.M. Oudejans 
Department of Experimental Zoology, University of Utrecht, 8 Padualaan, 
3508 TB Utrecht, THE NETHERLANDS 


ABSTRACT 


In order to technically validate two nomenclatural combinations 
made in a previous paper (Oudejans 1989), two corrections are made. 


KEY WORDS: Euphorbiaceae, Euphorbia, nomenclature. 


Due to a typist’s error in the editor’s office, a new combination and a new 
name were incorrectly listed in a previous paper (Oudejans 1989). In order 
to validate these names, they are repeated here. The editor apologizes to Dr. 
Oudejans for these errors and would urge all authors, whenever possible, to 
send manuscripts on computer diskettes. 


Euphorbia rockii Forbes var. grandifolia (Hillebrand) Oudejans, comb. 
nov. BASIONYM: E. clusiifolia var. grandifolia W. Hillebrand, Fl. 
Hawaiian Isl. 395 (as ‘clusiaefolia). 1888. 


Euphorbia smallii Oudejans, nom. nov., pro Chamaesyce pinetorum J.K. 
Small, Bull. N.Y. Bot. Gard. 3:429-430. 1906. TYPE: UNITED STATES. 
[Florida:] between Cutler and Camp Longview Small & Carter 836. The 
name pinetorum is not available in the genus Euphorbia. 


LITERATURE CITED 


Oudejans, R.C.H.M. 1989. New names and new combinations in the genus 
Euphorbia L. (Euphorbiaceae). Phytologia 67(1):43-49. 


236 


Phytologia (September 1989) 67(3):237-253. 


NEW TAXA AND NOMENCLATURAL COMBINATIONS IN 
SENECIO IN MEXICO AND THE UNITED STATES 


T.M. Barkley 
Herbarium, Division of Biology, Kansas State University, Manhattan, Kansas 
66506, U.S.A. 


ABSTRACT 


The following new species and new variety are described: S. billi- 
eturneri T.M. Barkley, S. lasiocaulon T.M. Barkley, S. porphyres- 
thes T.M. Barkley, S. pseudopicridis T.M. Barkley, S. scalaris var. 
carmenensis C.C. Freeman (all from México); and S. spellenbergii 
T.M. Barkley (from New Mexico, U.S.A.). The following new combi- 
nations are proposed: S. flaccidus var. durangensis (Greenm.) T.M. 
Barkley, S. hintonii (H. Robins. & Brettell) J. Pruski & T.M. Barkley, 
S. multidentatus var. huachucanus T.M. Barkley and S. scalaris 
var. parrasianus (Greenm.) C.C. Freeman. 


KEY WORDS: Senecio, Asteraceae, México, United States, sys- 
tematics. 


Studies in Senecto in North America have led to the recognition of four 
new species from México and one from the United States, plus the need for 
four nomenclatural adjustments. They are presented in this paper. 

Craig C. Freeman has kindly allowed me to present here a new variety and 
a new combination under Senecio scalaris in advance of the publication of his 
revision of the Aureoid species of Senecio in México. Thus, the names will be 
available for the forthcoming work on the Asteraceae of México by B.L. Turner 
& Guy Nesom. Author citation for the new variety and the new combination 
may be simply “C.C. Freeman” (instead of “Freeman in Barkley”), fide art. 
46.2 of the International Code of Botanical Nomenclature (1988). Likewise, 
the new combination proposed below by J. Pruski & T.M. Barkley may be 
cited simply by those two authors. 

The new species described here are known to me only from the collec- 
tions that are cited. The habitat and distributional information is that of the 
specimen labels from the various collections. 


Senecio flaccidus var. durangensis (Greenm.) T.M. Barkley, comb. et 
stat. nov. BASIONYM: S. durangensis Greenm., Publ. Field Columbian 
Mus., Bot. Ser. 2:275. 1907. 


237 


238 PHY POLO EGrA volume 67(3):237-253 September 1989 


S. ctenophyllus Greenm., Proc. Amer. Acad. Arts 43:20. 1907. (not S. cteno- 
phyllus Phil.). 


Senecio hintonii (H. Robins. & Brettell) J. Pruski & T.M. Barkley, comb. 
nov. BASIONYM: Roldana hintoni H. Robins. & Brettell, Phytologia 
27:420. 1974. 


John F. Pruski of The New York Botanical Garden noted the need for this - 
combination on some herbarium annotation labels several years ago. I agree 
with the utility of the transfer to Senecio and it is my pleasure to join John 
Pruski in presenting this combination. 


Senecio multidentatus var. huachucanus (A. Gray) T.M. Barkley, comb. 
et stat. nov. BASIONYM: S. huachucanus A. Gray, Proc. Amer. Acad. 
Arts 19:54. 1883. 


Senecio scalaris var. parrasianus (Greenm.) C.C. Freeman, comb. et stat. 
nov. BASIONYM: S. parrasianus Greenm., Ann. Missouri Bot. Gard. 
4:20. 1917. 


Senecio billieturneri T.M. Barkley, sp. nov. Figure 1. TYPE: MEXICO. 
Durango: Lecheria, about 6 miles W of El Salto, along read to Mazatlan, 
along banks and in shallow water of stream, elev ca 8500 ft, perennial 
herb with ray and disk florets both yellow, (no date), Howard Scott 
Gentry 10610 (HOLOTYPE: MICH; Isotype: MEXU). 


Senecio multidentato Sch.-Bip. similis, sed capitulis magnis (1-) 
4-10, disco 12-20 mm diametro et involucri bracteis 9-12 mm longis, 
necnon foliis caulinis medianis ac superioribus caulem circumdan- 
tibus semiamplexicaulibus, ulteris ecologia semiaquatica diversus. 


Coarse, soft stemmed herb to 6+ dm tall. Herbage mostly glabrate at flow- 
ering time, but with scattered short hairs on the upper stem, the pedicels and 
base of the heads; variously dense pubescent on the involucral bracts. Stems 
arising singly or 2-3 loosely clustered from an elongate rhizome with abundant 
fleshy fibrous roots; the roots unbranched or with a few thin, lateral branches. 
Basal leaves and those of the lower 1/2 of the stem of about equal size and 
not disposed in a clearly defined basal cluster, upper leaves few and somewhat 
reduced; the well developed leaves linear-lanceolate to narrowly lanceolate, 
the blade 8-15 cm long and 1-2.5 cm wide, tapering to a winged petiole about 
as long as the blade or shorter, the middle and upper cauline leaves becom- 
ing sessile and with the bases encircling and weakly sheathing the stem, but 
not at all auriculate clasping, margins subentire or obscurely wavy, with a 
few, minute, callose denticles. Inflorescence a corymbiform cyme of (1-)4-10 
heads; the ultimate branches of the inflorescence (pedicels) 3-7(-10) cm long, 
with 3-5 subulate bracts 5-7 mm long, equidistantly placed along the length 


Barkley: New Senecio in Mexico and United States 239 


of the pedicel. Heads cylindrical to campanulate at maturity, the disk 12-20 
mm across; principal involucral bracts ca 21, triangular-lanceolate, 9-11+ mm 
long, the central rib prominent, greenish and + permanently short pubescent, 
the margins scarious-stramineous and glabrous, the tip weakly attenuate and 
darkened, but without a distinctive blackish spot; calyculate bracts mostly 
5-8, greenish, subulate, (3-)5-9 mm long; receptacle flat or low hemispheric, 
sometimes infested with insect larvae, naked or with low, erose ridges among 
the achenes; ray florets ca 13, pistillate and apparently fertile, the ligule bright 
yellow, 10-15+ mm long in dried specimens; disk florets numerous, often more 
than 50, bisexual and apparently fertile, corolla yellow, 7-8 mm long, the throat 
separating the tube from the limb ca 2/5 the distance up from the base, the 
limb narrowly tulip shaped and flared upward to 5 small triangular lobes, ca 1 
mm long or less. Achenes 5-6 mm long (immature), angled, glabrous; pappus 
a single series of grayish white, minutely barbellate, capillary bristles, 3-5+ 
mm long but of uneven lengths. 

Paratypes: (All from MEXICO. Durango: along or near Mexican highway 
40, west of the city of Durango): Alrededores del Mil Diez, 2 kms al N de El 
Salto, Mpio. de El Salto, alt 2200 m, Junio 27, 1982, R. Hernandez M. 7414 y 
P. Tenorio (KSC); wet meadow in Pinus lutea and Pinus durangensis forest, 
10 miles west of El Salto on Route 40, elev 9000 feet, July 16, 1964, Miles 
& Wilma Johnson 1859 & 1861 (both WIS); 10 miles W of El Salto along 
Mexican Rte 40, pastured pine woodland, 27 June 1974, Marvin L. Roberts & 
David Keil 10819 (F,OS); 4.5 km al SW de El Salto, brecha El Salto-Pueblo 
Nuevo, Mpio. de El Salto, alt 2100 m, 3 de Julio de 1982, P. Tenorio L. 808 
y C. Romero de T. (KSC); Las Adjuntas, Mpio. de El Salto, alt 2000 m, 5 de 
Julio de 1982, P. Tenorio L. 829 y C. Romero de T. (KSC). 

Senecio bilheturnerz is referable to Group llc, Triangulares, in the scheme 
of Barkley (1985). It is similar to S. multidentatus Sch.-Bip. and the closely 
related S. huachucanus A. Gray (which is transferred in this paper to varietal 
status within S. multidentatus). It differs from them in having (1-)4-10 notably 
large heads, the middle and upper cauline leaf bases encircling and weakly 
sheathing the stem, and in a semi-aquatic habitat, or at least the ability to 
grow as a facultative aquatic. The species is further noteworthy in its rather 
long achenes in comparison to the length of the corolla and the pappus of the 
disk florets. 

Two of the paratypes noted above, Miles & Wilma Johnson 1859 & 1861, 
are distinctive in being monocephalous and in having few and reduced cauline 
leaves. Otherwise, they cannot be excluded from S. bilhieturneri as it is con- 
ceived here. 

It is a pleasure to name this species for Dr. B.L. Turner of the University 
of Texas, who has made notable contributions to the knowledge of the botany 
of North America, and who has been an inspiration for two generations of 
botanists. 


240 PHY. TODO GLA volume 67(3):237-253 September 1989 


Senecio lasiocaulon T.M. Barkley, sp. nov. Figure 2. TYPE: MEXICO. 
Durango: Fourteen miles west of Cd. Durango (Durango to El Salto 
highway), edge of limestone out-cropping dropping into arroyo south 
of highway, grasslands, with very sparse growth of oak scrub, eleva- 
tion 2000-2200 m, frequent, flowers bright yellow, June 1950, James H. 
Maysilles 7082 (HOLOTYPE: NY; Isotypes: MEXU,MICH). 


Senecio picridis Schauer simulans, sed foliis semper lanato- 
tomentosis, superioribus qual mediana minoribus, hic ultra 15 cm 
longis, necnon capitulescentia laxa diffusa, capitulorum numero 
(-15) diversus. 


Subshrub 2-5 dm tall. Herbage white, closely felted lanate-tomentose 
throughout, but upper side of leaves somewhat grayish and unevenly glabres- 
cent in age. Stems strict, branching only in upper 1/4, distinctly ligneous be- 
low but herbaceous upward, arising singly or loosely clustered from a ligneous, 
subrhizomatous caudex, with numerous branching, fibrous roots. Leaves about 
evenly distributed along the stem, the lowermost withering early, those of 
the upper 1/4 of the stem reduced in size; well developed mid-cauline leaves 
oblanceolate to linear-oblanceolate, tapering to a weakly distinct, winged peti- 
ole, 7-12(-15) cm long and (0.5-)1-1.5 cm wide overall, distinctly auriculate 
clasping at the base. Inflorescence a weakly compacted to loose corymbiform 
to subpaniculiform cyme of ca 7-15 heads, the ultimate branches of the inflores- 
cence (pedicels) 2-3+ cm long. Heads subcylindrical to narrowly campanulate, 
the disk 5-7 mm across; principal involucral bracts ca 13(-21), linear lanceolate 
to lanceolate, (4-)5-6 mm long, densely and permanently close tomentose along 
the median ridge, margins glabrous and stramineous, the apex minutely erose 
fimbriate and faintly to prominently darkened; calyculate bracts ca 5 or fewer, 
sometimes absent, linear subulate, less than 2 mm long, sometimes obscurely 
anthocyanic; receptacle ca 4 mm across, flat or low hemispheric, naked except 
for low, erose ridges among the achenes; ray florets ca 8(-137), pistillate and 
apparently fertile, corolla yellow, the tube ca 4 mm long, the ligule 5+ mm 
long in dried specimens; disk florets (15-)20-25, bisexual and apparently fertile, 
corolla yellow, ca 5-6 mm long overall, the lower 1/4 narrowly cylindrical, the 
upper 3/4 conically expanded and terminating in 5 short, triangular lobes less 
than 1 mm long. Achenes (immature) ca 2.5 mm long, angled, minutely hirtel- 
lous on the angles; pappus of abundant, minutely barbellate, white hyaline, 
capillary bristles in a single series, 5-7 mm long, but of uneven lengths. 

Paratypes: (All from MEXICO. Durango: along or near Mexican highway 
40 west of the city of Durango). Among boulders at base of cliffs in Mimbres 
Canyon, 40 miles W of Durango, July 24, 1958, D.S. Correll & I.M. Johnston 
20139 (NY); weedy meadow and roadcut in pine forest belt, 7 km NE of El 
Salto, alt ca 2800 m, July 21, 1969, B. & C. Marcks 1238 (DAV,WIS); 34 miles 
west of Cd. Durango, June 23, 1950, J.H. Maysilles 7072 (MICH); about 5 


Barkley: New Senecio in México and United States 241 


or 


¥ 


PLANTS OF $Q¥QRA, MEXICO 
TC: aB00 


Benecie -/' 


HERBARIUM OF KANSAS STATE UNIVERSITY 
MANIMATTIAN, KANSAS 


Sears bithe fureery TM. Bacisiey 
procotTyPe 


Annotated by Ju darhh, 9 ¥9 


Locelity’ Lecherta, sbout 6 miles ¥ of El Salto, 
mtuge along roed to Mssatles. 


Habitat. Along banks & io shallow water of strei. 
Blev. oa. 8500 fret. 
Poremnia) herd with ray & disk flowre ‘oth yellow. 


CB cat tes Hae are seott Garde we El, 


Figure 1. Senecio bilkieturneri - holotype. 


242 


PHY TOLO ESTA volume 67(3):237-253 September 1989 


HERBARIUM OF FANSAS SIF IL titvins iy 
MANHATTAN FANSAS 


Sanecio lnsiecauleon Tt. harwiny 
mocoTyPEe 


Annotated by J us Raheny, wad 


PLANTS OF DURANGO. MENIVO 
Heisenum of the University ot Michigan 


“ te 


Fourteen milee west of © Wurangy , Durange (o El Seite 
bighway), edge of imestuae vuterupping dropping into arr yy 
south of highway, grawlands. with very sparse growth of ak 
scrub. elesation 2) - IN m 


EVGsas aks Eo ae rk: 


Jone MW Meyeciiea, ws Pee 


Figure 2. Senecio las:ocaulon - holotype. 


Barkley: New Senecio in México and United States 243 


miles north of railroad at Coyotes (45 airline miles west of Cd. Durango), June 
28, 1950, J.H. Maysilles 7123 (MICH,MEXU). 

Senecio lasiocaulon is referable to Group 10c, Fruticosi, s. str. in the scheme 
of Barkley (1985). It is similar to S. pzcrzdis Schauer, a species chiefly of the 
eastern Trans-Mexican Volcanic Belt and the Sierra Madre Oriental, but it dif- 
fers in having permanently lanate-tomentose herbage, upper leaves distinctly 
smaller than the middle cauline leaves, well developed leaves that are regularly 
more than 8 cm long, and a loose, open inflorescence of fewer than 15 heads. 

The specific epithet “lasiocaulon,” is derived from the Greek roots “lasi-,” 
shaggy-wooly, and ”caulos,” stem or stalk, as an allusion to the conspicuous 
pubescence of the herbage. 


Senecio porphyresthes T.M. Barkley, sp. nov. Figure 3. TYPE: MEXICO. 
Tamaulipas: On mountain top 7 km SW of Miquihuana in forest of large 
pines, forest floor of low vegetation, elev 3430 m, (23° 40’ N, 99° 45’ 
W), Aug 5, 1941, L.R. Stanford, L. Rutherford & R.D. Northcraft 679 
(HOLOTYPE: NY; Isotypes: GH,MO). 


Senecio gerberaefolio Sch.-Bip. similis, sed ab eo notulis sequen- 
tibus diversus: capitulum solitarium et minus, disco 12-20 mm 
diametro et involucri bracteis 12-14 mm longis; folia subduplo mi- 
nora, 4-10(-12) X 1-1.5 cm; caules tenues nec robusti. 


Subscapose herb 10-15 cm tall. Herbage closely lanate-tomentose, but 
becoming loosely tomentose with long, arachnoid hairs toward the base or un- 
evenly glabrate; upper side of the leaves glabrescent, lower side closely and 
permanently short felted lanate. Stem simple, arising singly in a cluster of 
basal leaves from the end of a rhizome; uppermost 1/4 of the stem with 2-5 
linear bracts 5-15 mm long. Rhizome creeping, simple, nearly 1 cm in diam- 
eter, covered by the exfoliating bases of old leaves and producing numerous 
fibrous branching roots. Leaves all basal or nearly so, narrowly oblanceolate 
to subspatulate, the blade tapering to the winged petiole, 4-10(-12) cm long 
overall and 1-1.5 cm wide, somewhat coriaceous, the margin denticulate with 
callose denticles, weakly revolute to flat. Head single, broadly campanulate to 
obconic, the disk 12-20 mm across; principal involucral bracts ca 16-20, all of 
equal length but disposed in an inner and an outer series, 12-14 mm long and 
to 2+ mm wide, linear-lanceolate, felted lanate-tomentose on the outer side, 
the margins and tip prominently and permanently magenta or pinkish-purple; 
calyculate bracts few, reduced and without the distinctively colored margins 
and apex; ray florets 12 (or more?) pistillate and apparently fertile, the corolla 
tube ca 6 mm long, the throat with a prominent thickened collar; the ligule 
sharply defined at the throat, bright yellow, 10+ mm long and to 5 mm wide; 
disk florets bisexual and apparently fertile, the corolla ca 8 mm long, tapering 
upward, the throat indistinct, the limb gradually flared and terminating in 5 


244 PHYATOLOCGIA volume 67(3):237-253 September 1989 


small, triangular lobes 0.7-0.9 mm long. Achenes 2 mm long (immature), with 
appressed, flat, hyaline hairs (which presumably persist); pappus of both ray 
and disk florets of abundant, white, minutely barbellate capillary hairs that 
are nearly as long as the disk corollas. 

Senecio porphyresthes is referable to Group lle, Lugentes, in the scheme 
of Barkley (1985). It is similar to S. gerberaefohus Sch.-Bip., a species of 
the higher peaks in the eastern Trans-Mexican Volcanic Belt, but differs in 
having single and somewhat smalier heads, leaves only about half the size, 
and subscapose stems that are notably thinner. This new species is known 
to me by only three specimens from a single collection, but the aspect and 
combination of characters suggest that it is distinct. The purple-magenta 
color of the margins and tips of the involucral bracts is most prominent in the 
isotype in GH; the holotype in NY has the color somewhat faded. 

Senecio gerberaefolius has a chromosome number of n = 30 (vouchered on 
specimens of John H. Beaman 1948, México. Ixtaccihuatl, GH,;MEXU,UC, 
WIS), a number associated with cacalioid affinities and a distinctive suite of 
microcharacters (cf Barkley 1985). To date, cacalioid characters have not been 
specifically identified for S. gerberaefolius, but it will be interesting to learn if 
S. porphyresthes has n = 30 and/or cacalioid microcharacters. The disposition 
of the principal involucral bracts in an identifiable inner and outer series, as 
they occur in S. porphyresthes is often present with cacalioids. 

The specific epithet, “porphyresthes,” is derived from the Greek “porphyr,” 
purple, and “esthes,” garment, as an allusion to the conspicuous coloring of 
the involucre. 


Senecio pseudopicridis T.M. Barkley, sp. nov. Figure 4. TYPE: MEXICO. 
San Luis Potosi: Cerro Grande, 8 km al NW de Guadalcazar, ladera 
granitica con vegetacion de encinar, alt 2000 m, 25-IX-1955, Rzedowskz 
208 (HOLOTYPE: GH; Isotype: MICH). 


Senecio picridis Schauer simulans, sed semper herbaceus, et 
foliis primariis oblanceolatis, praeter denticulas callosas minutas 
integris, lamina in petiolum distinctum decurrenti diversus. 


Subligneous herb 2-4+ dm tall. Herbage unevenly arachnoid-tomentose, es- 
pecially upward and among the heads in the inflorescence, irregularly glabrate 
in age, the leaves persistently grayish lanate tomentose on the underside. Stem 
thin, stiff, branching upward from near the base, arising singly or 2-4 from a 
weakly ligneous taproot. Leaves prominently developed along the lower 2/3 
of the stem; upper leaves few and somewhat reduced. Principal mid-cauline 
leaves oblanceolate to narrowly oblanceolate, 6-8(-10) cm long overall and 1- 
1.5+ cm wide, tapering to a narrow, weakly winged petiole, margins subentire, 
with minute callose denticles, or sometimes with 1-3 prominent, rounded teeth 


Barkley: New Senecio in Mexico and United States 245 


HERBARIUM OF KANSAS STATE UNIVERSITY 
MANHATTAN, KANSAS 


Senecio perphyvesthey TM Bamiey 


Hocot yPL 
Annotated by yw. Banit~ 19 72 
SLATE. Us velarin te & 
PLANTS OF MEXICO 
Senecic 3>. (fide [.u. Join ton; 
On mountain too 7 kito. 3.4. oF s-lycbiuens bt 


forest of large 21.18; forest floor of lo. 
vege‘ at .on. 


Col. by LR Banterd 34 Bisse 5g 1968 
EL Beteertca 23 42 97 45 
BD Ronthaah me 679 


Figure 3. Senecio porphyresthes - holotype. 


246 PHYTOL GGT A volume 67(3):237-253 September 1989 


HERBARIUM OF KANSAS SIATE UNIVERSITY 
maT A 


Serecre preude picrides 7-A Baerley 
HoloryPe 


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Figure 4. Senecio pseudopicridis - holotype. 


Barkley: New Senecio in México and United States 247 


on the lower (proximal) 1/2 of the blade, the base expanded and auriculate- 
clasping, but sometimes inconspicuously so. Inflorescence a loose, corymbiform 
cyme of (5-)6-12+ heads terminating each main branch, the ultimate branches 
of the inflorescence (pedicels) 2-5 cm long, with 1-3 minute, linear bractlets 
equidistantly placed along the length of the pedicel. Heads cylindrical or 
slightly campanulate at maturity, the disk 5-8 mm across; principal involucral 
bracts ca (13-)21, linear-lanceolate to weakly subulate, 6-7 mm long at matu- 
rity, glabrescent, but with a few, scattered, minute, thick hairs, and sometimes 
with some persisting arachnoid-villous hairs, apparently anthocyanic toward 
the tip when young, the apex with a minute, darkened tip and a small tuft of 
arachnoid hairs; calyculate bracts 4-8, linear, spreading, ca 2 mm long, the tip 
darkened and with a tuft of hairs; receptacle flat or low hemispheric, naked 
or nearly so; ray florets ca 8 or 13, pistillate and apparently fertile, the ligule 
bright yellow, 6-8+ mm long in dried specimens; disk florets numerous, + 
25-35, bisexual and apparently fertile, the corolla 7-8 mm long, the lower 2/5 
tubular, the upper 3/5 conical and terminating in 5 small triangular lobes, ca 
1 mm long or less. Achenes ca 3 mm long (immature), angled, evenly short 
pubescent with minute, grayish hairs; pappus a single series of white hyaline, 
minutely barbellate, capillary bristles, 6-8 mm long, but of uneven lengths. 


Paratype: MEXICO. Zacatecas: 16 (air) miles E of Concepcién de Oro, on 
upper north side of Sierra de Astillero, ca 3 miles NE of Guadalupe Garceron, 
1/2 mile N of summit, in open pinyon woodland, frequent perennial, flowers 
true yellow, elev 8200 ft, Sept 22, 1973 (near 24° 38’ N, 101° 08’ W), James 
Henrickson 18845 (LL). 


Senecio pseudopicridis is referable to Group llc, Triangulares in the scheme 
of Barkley (1985). It is similar to S. picridis Schauer, a species chiefly of the 
eastern Trans-Mexican Volcanic Belt and the Sierra Madre Oriental. How- 
ever, S. pseudopicridis is fundamentally herbaceous, its principal leaves are 
oblanceolate, the blade tapering to a well marked petiole, subentire or nearly 
so, except for the minute, callose denticles. Clearly, S. pseudopzcridis ap- 
proaches some specimens of S. picridis, but the combination of characters and 
the gross aspect define it as a distinct species. Further information may alter 
the concepts of these two entities. 


The holotype and the paratype cited above both have leaf bases that are 
auriculate clasping, but the isotype (in MICH) has leaf bases that are scarcely 
expanded. However, the isotype is otherwise identical with the holotype and 
is evidently of the same collection, from the same population. It raises the in- 
teresting question of the stability of the character of auriculate clasping leaves. 
Herbarium studies indicate that other species that normally possess that trait, 
occasionally do have individuals with narrow based leaves. Experience with 
Senecio suggests that virtually no characters of taxonomic utility occur with- 
out occasional exceptions. 


248 PHY T,.0.L 0.4..A volume 67(3):237-253 September 1989 


Senecio spellenbergii T.M. Barkley, sp. nov. Figure 5. TYPE: U.S.A. New 
Mexico: Harding Co., Hwy NM 120, 16 miles NE of Roy, ca 11 miles SE 
of Yates, S side of Carrizo Creek, on white caliche in prairie, milepost 
91, only 2 seen in late flower, the rest past (and this was late spring); 
phyllaries of older heads becoming glabrate, dark maroon; heads rayless, 
dull yellowish; 1-6 rosettes in a patch; leaves convex above, concave or 
grooved beneath, May 29, 1983, Richard Spellenberg 7073 (HOLOTYPE: 
NY; Isotypes: KSC,NMC,TEX). 


Senecio werneriaefolium A. Gray simulans, sed tomento ap- 
presse coacto, foliis angustis arcte revolutis, capitulis eradiatis fere 
semper solitariis, necnon achaeniis hirtellis diversus. 


Dwarf, scapose herb, 3-5+ cm tall. Herbage closely whitish tomentose, the 
leaves with vestiture tightly felted into a velamen-like covering that irregularly 
exfoliates in age, upper herbage subglabrescent. Stems arising singly or some- 
times 2 from a tufted rosette of basal leaves; the rosettes single or 2-6 closely 
clustered, from a simple or branching, suberect or weakly creeping, coarse rhi- 
zome, 3-5 mm in diameter; principal roots fleshy fibrous, branch roots thin 
and thread like. Principal leaves all basal, coriaceous, linear, 10-15 mm long, 
strongly revolute and only 1-2 mm wide; cauline leaves reduced to 2-4(-5) 
linear-subulate bracteoles, 2-3 mm long. Head single (rarely 2) at the end of 
the stem, eradiate, the disk ca 10 mm across; principal involucral bracts ca 13, 
linear-lanceolate, acute, 6-8 mm long, purplish to deep maroon-purple along 
the wide midrib and toward the tip, margins wide and scarious-stramineous; 
calyculate bracts 1-4, linear-subulate, less than 4 mm long; receptacle low 
hemispheric, naked; ray florets absent; disk florets ca 20(+) in number, bisex- 
ual and apparently fertile; corolla yellow, ca 6 mm long, the throat ca 2/5 the 
distance upward from the base, the limb narrowly conical, terminating in 5 
triangular lobes 0.5-0.7 mm long. Achenes cylindrical to weakly fusiform, ca 3 
mm long, inconspicuously angled, hirtellous; pappus of abundant, white hya- 
line, minutely barbellate, capillary bristles 5+ mm long but of uneven lengths. 

Paratypes: (From same locality as type collection, but with following data): 
On white, nearly barren, very calcareous knolls in shortgrass prairie, elev 5500 
ft, July 2, 1981, R. Spellenberg, R. Soreng & T. Fisher 6053 (KSC,NMC). 

Senecio spellenbergii is assignable to the “Tomentosi” group of the Au- 
reoid assemblage, and it would key to Senecio werneriaefolius A. Gray in my 
treatment of Senecio in North American Flora and subsequent relevant papers 
(Barkley 1978; 1980; 1988). It differs from S. werneriaefolius in possessing a 
closely felted tomentum, basal leaves that are short, very narrow and tightly 
revolute, heads that are consistently eradiate and nearly always single, and ach- 
enes that are hirtellous. Additionally, S. spellenbergzz has a distinctive gross 
aspect, habitat and distribution. It was erroneously assigned to S. canus by 
me in the past, and the type and paratype collections are referred to S. canus 


Barkley: New Senecio in México and United States 249 


in a floristic paper by Spellenberg et al. (1986). (N.B. in that publication, the 
type collection is erroneously cited as no. 7203 instead of no. 7073). 

Senecio spellenbergit occurs on caliche soil, in a grassland habitat in the 
High Plains, some 150 km (90 mi) south of the nearest location of S. werne- 
riaefolius which is in the subalpine areas of the San Isabel National Forest 
west of Trinidad, Colorado, and there the plants are distinctively “typical” for 
the species. Senecio spellenbergii is superficially similar to “Phase no. 4” of S. 
werneriaefolius as described elsewhere by me (Barkley 1980). “Phase no. 4” 
includes several populations on clay soil in southern Utah, where the plants 
tend to have close, white tomentum, narrow leaves with revolute margins and 
a depauperate, sometimes monocephalous aspect. Herbarium studies suggest 
that these populations are but edaphic extremes in a continuum of variation, 
and clearly within the geographic and morphological ranges of a highly variable 
species. A notable example of this extreme form is the following collection: 
Utah: Garfield Co. Dixie National Forest, Paunsaugunt Plateau, low ridge 
system north of the East Fork of the Sevier River, 0.4 mi north of Utah Hwy 
12, near Coyote Hollow, on open exposed clay slopes with Pinus aristata, Sec 
2, T36, R4W, elev 7650 ft, 25 May 1968, J.L. & C.G. Reveal 1017(KSC). The 
specimens are small, about 5 cm tall, radiate and with glabrous achenes. 

Another relevant collection is: New Mexico: McKinley Co., Zuni Indian 
Reservation, SE of Zuni in Galestina Canyon, SW side, about 1/2 way up the 
canyon, R16W, T9N, elev 6500 ft, on N slopes of small box feeder canyon, 3 
June 1988, R. Spellenberg, D. Ward, J. Enote & S. Davis 9501 (KSC; the col- 
lection label notes that duplicates are also at ASU,NMC,NY,RSA,UC,Z.LR.). 
The plants of this collection are similar to S. spellenbergii in aspect, and they 
have both eradiate heads and hirtellous achenes. However, the mature plants 
are 7-10 cm tall, which is rather larger than those of S. spellenbergzt and the 
heads are frequently two per inflorescence. Furthermore, the field data sug- 
gest the collection site is not a distinctive caliche type soil, but rather, the 
plants are growing “on decayed red Wingate Sandstone overlayed by a gray 
Zuni Sandstone, with Pinus edulis, Juniperus monosperma, Cercocarpus, a 
few Pseudotsuga in upper cooler spots.” It is, therefore, easier to regard this 
collection as a southern extreme of S. werneriaefolius, an admittedly complex 
and polymorphic entity, and to keep S. spellenbergii as a restricted and easily 
circumscribed entity. 

It is a pleasure to name this species for Dr. Richard Spellenberg of New 
Mexico State University, who collected the type materials, and who has made 
important contributions to the knowledge of the flora of southwestern U.S and 
northern México. 


Senecio scalaris var. carmenensis C.C. Freeman, var. nov. Figure 6. TYPE 
MEXICO. Coahuila: Mpio. Villa Acuna, Sierra de Carmen, Canyon de 
Sentenela (= Canon del Centinel) on Hacienda Piedra Blanca, moist 


250 PHYTOLOGIA volume 67(3):237-253 September 1989 


stream side, July 6, 1936, F.L. Wynd & C.H. Mueller 546(HOLOTYPE: 
NY; Isotypes: GH,MICH,MO,TEX). 


Senecio scalaris Greene var. scalaris similie, sed foliis tomento- 
sis vel floccosis. 


Stout herbaceous perennial 2-5 dm tall. Lower surface of basal leaves and 
occasionally lower stem sparingly tomentose to floccose-tomentose, lower stem 
and basal leaves often faintly to deeply anthocyanic. Stems 1-3 from an erect 
or creeping and branched caudex. Basal leaves mostly pinnatifid, sometimes 
sublyrate, overall leaf dimensions (2.5-)3.8-11.1 cm long, 0.5-1.8(-2.3) cm wide. 
Inflorescence a loose corymbiform cyme of (4-)7-20(-35) heads; principal in- 
volucral bracts glabrous to sparingly tomentose at the base; achenes glabrous 
to hirtellous on the angles. 

Paratypes: (All from MEXICO. Coahuila): Higher western ridge on Sierra 
el Jardin, E of Rancho El Caballo, Chiang et al. 9835 (LL); Sierra Jardin, Flyr 
1188 (MO); Canon Humido on N side of Pico Centinel, Sierra el Jardin, 8 km 
E of Rancho El Jardin, Johnston et al. 11798 (LL); Sierra Maderas el Carmen 
at Campo 3, Wendt & Adamcewicz 502 (LL); Mpio. Ocampo, Sierra Madera 
el Carmen on upper slope and ridge of peak in upper portion of Oso Canyon 
between Campo 0 and Campo 5, Riskind & Patterson 1785 (LL); Madera el 
Carmen at end of rd above Campo 5 in saddle near viewpoint of W face of 
Sierra, Fryzell 2695 (LL,MO). 

Senecio scalaris var. carmenensis is referable to Group 11d, Aureoidei, 
in the scheme of Barkley (1985). It is restricted to the Sierra el Carmen, 
Coahuila, (sometimes written as “Sierra del Carmen”) where it occurs at 1500- 
2500 m in pine-oak woodlands, on sandy or gravelly loam, often derived from 
rhyolite. Three other aureoids are known in the same region: 5S. millelobatus, 
S. coahuilhensts and S. obovatus. Senecio scalaris var. carmenensis is similar 
to S. millelobatus but may be distinguished by its pinnatifid basal leaves that 
are pubescent beneath and its winged midrib. By comparison, S. millelobatus 
has pinnate or twice-pinnate basal leaves that are mostly glabrous and that 
have an essentially unwinged leaf midrib. The systematic position of this new 
variety is treated in detail by Freeman (1985). 


ACKNOWLEDGMENTS 


It is a pleasure to acknowledge the generous assistance of Dr. Patricia 
Davila and the herbarium staff of the Herbario Nacional de México (MEXU), 
Instituto de Biologia, Universidad Nacional Auténoma de México, in helping 
me to form an understanding of Senecio in the incredibly complicated flora 
of México. José Luis Villasenor, a diligent and capable young student of the 


Barkley: New Senecio in México and United States 251 


} 4 
¢ a HERBARIUM Of KANSAS STATE UNIVERSITY 
E IBANHATIAN, KANSAS 


# 
A Senece spellanbergii TH Berniny 
HovceoryPpe 


Annotated by J. aa Rake, wes 


cCanvs Hook 


Onis 2 im tate tlive , 
thiS was @ lete spring); 
hesds Ceconiry ,glatiate, care - 
Cayless, Gull perlusess; 
patch; leaves hoe ot 
grooved beneatn, Collen te 
as Szettenters, =<. er, 


voi, rc, mitt wott 


Asse 


EW YOr¢ 
BOTANICAL 
Garven 


ee 


= ster 


wat 


Figure 5. Senecio spellenbergu - holotype. 


252 PHY T'OLOGTA volume 67(3):237-253 September 1989 


UNIVERSITY OF 


PEXAS , 


HERBARIUM od 
f 


ISOTYPE of: 


Senet > alard as Var. ite ie tite 
C. C. Freeman 


Pet. Cruty C. Freeman lyts 


PLANTS OF MEXICO 
Gxtede do Coahuila 3 
Municiplo do «= Willa Acuns 
io acalhsnro Fontan, 
Ad § FRG 
Bierra del Garwsns Canyon de 
Jentenela on Baciends Pietra 


Blanes; Moist streem sitc. 
ye ~ Ne 6466 Dete July i, 1.36 
Vo \ Colected by F. Lyte Wyed oad CH bbealier 


Figure 6. Senecio scalaris var. carmenensts - isotype. 


Barkley: New Senecio in México and United States 253 


Mexican Asteraceae, has shared with me his field and herbarium knowledge 
of the group. Dr. Rupert Barneby of The New York Botanical Garden kindly 
set the diagnoses of the new species into Latin. 

This paper is contribution no. 90-56-J of the Kansas Agricultural Experi- 
ment Station, Manhattan. 


LITERATURE CITED 


Barkley, T.M. 1978. Senecio, N. Am. Flora II(10):50-139. 


. 1980. Taxonomic notes on Senecio tomentosus and its allies (Aster- 
aceae). Brittonia 32:291-308. 


. 1985. Infrageneric groups in Senecio, s.l., and Cacalia, s.l., (Aster- 
aceae: Senecioneae) in Mexico and Central America. Brittonia 37:211- 


218: 


. 1988. Variation among the aureoid senecios of North America: a 
geohistorical interpretation. Bot. Rev. 54:82-106. 


Freeman, C.C. 1985. A revision of the aureoid species of Senecio (Aster- 
aceae: Senecioneae) in Mexico, with a cytogeographic and phylogenetic 
interpretation of the aureoid complex. Ph.D. dissertation. Kansas State 
University. Manhattan. 


Greuter, W. & J. McNeill, eds. 1988. International Code of Botanical Nomen- 


clature. Regnum Veg. 118). Koeltz Scientific Books, D-6240 Konigstein, 
Federal Republic of Germany. 


Spellenberg, R., R. Worthington, P. Knight & R. Fletcher. 1986. Additions 
to the flora of New Mexico. Sida 11:455-470. 


Phytologia (September 1989) 67(3):254-257. 


PUNCTELIA PUNCTILLA (HALE) KROG, NEW TO NORTH AMERICA 


Richard E. Riefner, Jr. 
Assistant Research Biologist, Museum of Systematic Biology, 
University of California, Irvine, California 92717 USA 


ABSTRACT 


Punctelia punctilla (Hale) Krog, previously known only from South 
Africa, is reported from southern California. Habitat data and the 
disjunction are discussed. 


KEY WORDS: Punctelia punctilla, California, North America. 


Few Punctelia species are found outside temperate or subtropical regions, 
most occurring in Africa and the Americas (Krog 1982). One species, P. 
rudecta (Ach.) Krog, is known to have a remarkable range that spans several 
continents in both hemispheres. Thus, it was not surprising to find a closely 
related species, P. punctilla (Hale) Krog (Figure 1), in coastal southern Cali- 
fornia. Punctelia punctilla has previously been known only from South Africa 
(Krog & Swinscow 1977). Hale & Cole (1988) list P. borrerx (Sm.) Krog, P. 
stictica (Duby) Krog and P. subrudecta (Nyl.) Krog from California. Of these 
species, P. punctilla may resemble P. stictica at first glance because of the dis- 
tinct, white rimmed pseudocyphellae and the darkening lobe tips. However, P. 
punctilla is clearly related to P. rudecta due to the presence of isidia, lecanoric 
acid and a pale lower surface. Punctelia rudecta (which is absent in the western 
states; M.E. Hale, pers. comm.) is a larger, corticolous species with smaller 
pseudocyphellae and abundant isidia which are coralloid or squamiform with a 
glossy cortex. The isidia of P. punctzlla differ in being low, papilliform or spar- 
ingly branched with a dull surface. Punctelia punctilla is strictly saxicolous. 
Additional range extensions may be discovered if collections of P. rudecta from 
rock are carefully examined to detect misidentifications of P. punctilla. 

In California, P. punctilla occurs rarely in the Pt. Mugu-Camarillo area 
of Ventura County. The climate at this site is Mediterranean, and the pre- 
dominant vascular plant community is coastal sage scrub. As with most of 
the southern California coastline, agriculture and urbanization have left lit- 
tle natural habitat intact. Punctelia punctilla is apparently restricted to a 
rocky hill region about five kilometers from the ocean at the base of outcrops, 
mostly on east facing slopes thickly vegetated with Opuntia littoralis, Core- 
opsis gigantea, Salvia leucophylla and Artemisia californica. Characteristic 


254 


Riefner: Punctela punctilla in North America 255 


lichens of these outcrops include Xanthoparmelia mezicana (Gyelnik) Hale, 
Flavopunctelia flaventior (Stirton) Hale, Physcza callosa Nyl., Niebla ceru- 
choides Rund. & Bowler in ed., Leprocaulon microscopicum (Vill.) Gams ez 
D. Hawksw., Dimelaena radiata (Tuck.) Hale & Culb., Acarospora schleicheri 
(Ach.) Massal., Thelomma mammosum (Hepp in Hartung) Massal., Lecanora 
gangaleozdes Nyl., Caloplaca bolacina (Tuck.) Herre, Lecanora muralis (Schre- 
ber) Rabenh. and Buellia halonra (Ach.) Tuck. It is interesting to note that 
the author has collected P. borreri from similar saxicolous communities along 
California’s southern and central coast. Punctelia borreri is also known from 
South Africa (Krog & Swinscow 1977). 

How can one account for such an enigmatic South African-coastal Cal- 
ifornia disjunctive range? This outlying population may be the result of 
chance long distance dispersal, or it could be considered a relic of an ear- 
lier and broader range due to vicariance events, e.g., the allopatric distri- 
bution of closely related taxa due to events in geologic history. Long dis- 
tance dispersal may be the more reasonable explanation when a sorediate 
species is involved, since isidia are relatively heavy and only transportable 
over short distances (Ott 1987). Although it is not easy to identify a specific 
vector/dispersal/pathway, chance long distance dispersal cannot be dismissed 
as a real possibility. Plants might have been carried along with ballast to 
nearby Pt. Mugu Naval Station. Also, Croizat (1952) discusses a variant of 
tracks of the African Gate that conceivably connects southern Africa to points 
beyond in the New World. On the other hand, a hypothesis supporting vicari- 
ance is strengthened when this distribution is viewed as a possible example of 
a classic pattern (a New World/southern Africa disjunction) for plants which 
evolved before the continents drifted apart (Hale 1987; Culberson 1972) What- 
ever the explanation, this new record represents an emerging distribution for 
P. punctilla that may add insight to lichen biogeographic study. 


ACKNOWLEDGMENTS 


I am indebted to Mason E. Hale for identifying the Punctelia and allowing 
me to publish the record. I also thank Peter A. Bowler and Charis C. Bratt for 
giving useful criticism of this manuscript, and David M. Williams for photog- 
taphy. The University of California, Irvine’s Museum of Systematic Biology 
provided Jaboratory space and other support for this research. 


256 PHY TOLO GIA volume 67(3):254-257 September 1989 


Figure 1. Punctelia punctilla (Hale) Krog from California, Ventura Co.: 
At base of conglomerate rock on E. Potrero Rd. near Lewis Rd. Riefner 87- 
69 (COLO,IRVC,SBM,US), TLC: atranorin and lecanoric acid. At base of 
outcrop off Lewis Rd. and SE of Camarillo State Hospital, Riefner 89-20 
(IRVC,WIS). TLC: atranorin and lecanoric acid. Solvent systems G, A, C 
of White & James (1985) used for TLC analyses. 


Riefner: Punctelia punctilla in North America 250 


LITERATURE CITED 


Croizat, L. 1952. Manual of Phytogeography, or an Account of Plant-dispersal 
Throughout the World. Uitgeverij Dr. W. Junk, The Hague. 


Culberson, W.L. 1972. Disjunctive distributions in the lichen-forming fungi. 
Ann. Missouri Bot. Gard. 59:165-173. 


Hale, M.E. 1975. A revision of the lichen genus Hypotrachyna (Parmeliaceae) 
in tropical America. Smithsonian Contr. Bot. 


Hale, M.E. & M. Cole. 1988. Lichens of California. University of California 
Press, Berkeley, California. 


Krog, H. 1982. Punctelia, a new lichen genus in the Parmeliaceae. Nord. J. 
Bot. 24:287-292. 


Krog, H. & T.D.V. Swinscow. 1977. The Parmelia borreri group in East 
Africa. Norw. J. Bot. 24:167-177. 


Ott, S. 1987. Reproductive Strategies in Lichens. Jn: Progress and Problems 
in Lichenology in the Evghties. Bibl. Lichenol. 25:81-93. J. Cramer in 
der Gebr. Borntraeger Verlagsbuchhandlung, Berlin-Stuttgart. 


White, F.J. & P.W. James. 1985. A New Guide to Microchemical Techniques 
for the identzfication of Lichen Substances. British Lichen Society Bul- 
letin No. 57. British Museum of Natural History, London, England. 


Phytologia (September 1989) 67(3):258-286. 


SMALL SCALE DISTURBANCES AND SUCCESSIONAL DYNAMICS 
IN A SHORTGRASS PLANT COMMUNITY: 
INTERACTIONS OF DISTURBANCE CHARACTERISTICS 


D.P. Coffin & W.K. Lauenroth 
Department of Range Science and Natural Resource Ecology Laboratory 
Colorado State University, Fort Collins, Colorado 80523 USA 


ABSTRACT 


The effects of interactions among disturbance characteristics on the 
successional dynamics of small scale disturbances in shortgrass plant 
communities of north central Colorado, USA, were evaluated by com- 
paring naturally occurring and artificially produced disturbances of dif- 
ferent type, seasonality, size, and location by soil texture. 

The species composition on the two naturally occurring disturbances 
(western harvester ant mounds and small animal burrows) were similar 
one year after plant recovery began, and were different from the compo- 
sition on the artificially produced plots. The high density and cover of 
perennials on ant mounds and animal burrows indicated that perennial 
organs were not killed by the clipping activity of harvester ants or by 
the pile of soil from burrowing animals. The majority of the cover on 
the artificially produced plots for two sites differing in soil texture, and 
most dates and sizes, was attributed to annuals. The importance of 
seasonality was indicated by the low cover of annuals and perennials on 
artificial plots produced in September. 

Annuals and perennials responded differently to disturbance size. 
The density of annuals was not affected by the size of the disturbance 
while the significantly smaller cover values on the smallest compared to 
the largest plots indicated the effects of competition on the growth of 
annuals by plants surrounding the plots. Vegetative ingrowth from the 
edge of the plots resulted in significantly greater cover and density of 
perennial grasses and sedges on the smallest plots for most dates and 
both sites than plots of the two larger sizes. The cover and density of 
the most important perennial forb on the plots, Sphaeralcea coccinea, 
were highest in the center of the largest plots, or the farthest distance 
from potential competitive interactions with plants in the surrounding 
undisturbed community. 

Although long term monitoring is necessary to evaluate the time 
required for each disturbed area to be dominated by a shortgrass plant 
community, the short term results suggest that animal burrows will have 


258 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 259 


the most rapid recovery time of the disturbance types studied and the 
largest artificially produced plots will have the slowest recovery time. 


KEY WORDS: Disturbance, grassland, Bouteloua gracilis, animal 


burrows, ant mounds, succession. 
INTRODUCTION 


The disturbance regime of a plant community is composed of a number of 
disturbance types, each with its associated characteristics (Pickett & White 
1985). The rate and pattern of reestablishment of plants following a distur- 
bance is dependent on the life history characteristics of plants available to 
enter the disturbed site and the characteristics of the site (Sousa 1984). Most 
studies in grasslands have focused on the independent effects of different dis- 
turbance characteristics, including size (Davis & Cantlon 1969), seasonality 
(Perozzi & Bazzaz 1978) and type, such as ant mounds (King 1977), small 
burrowing animals (Platt 1975; Hobbs & Mooney 1985), drought (Albertson 
& Weaver 1944) and comparisons of several types (Collins & Barber 1985; 
Belsky 1987; Milchunas et al. unpublished). 

Interactions among disturbance characteristics are also important (Collins 
& Uno 1983; Collins 1987). However, the nonuniformity of natural distur- 
bances in characteristics such as size and seasonality causes difficulties in eval- 
uating the separate effects of interacting disturbance characteristics. Several 
researchers have used manipulated plots to control for the heterogeneous char- 
acteristics commonly found associated with natural disturbances (Rapp & Ra- 
binowitz 1985; Belsky 1986), but these simulated disturbances are likely not 
representative of all conditions found on natural disturbances. A comparison 
of natural disturbances with artificial disturbances of comparable, yet uniform 
characteristics, is necessary to evaluate the effects of different disturbance 
characteristics on successional dynamics. 

In shortgrass steppe plant communities of the central and southern Great 
Plains of North America, most successional studies have focused on the in- 
dependent effects of large scale disturbances, such as abandoned agricultural 
fields (Savage & Runyon 1937; Judd & Jackson 1939; Costello 1944; Judd 1974; 
Reichhardt 1982). Small patch producing disturbances are also important, but 
they have been largely ignored (Coffin & Lauenroth 1988). In shortgrass com- 
munities dominated by the perennial grass, blue grama [Bouteloua gracilis 
(H.B.K.) Lag. ez Griffiths], disturbances that kill at least one individual B. 
gracilis have the largest potential effect on community structure (Coffin & 
Lauenroth 1988). 

Two examples of small scale disturbances of sufficient intensity and spa- 
tial scale to always kill at least one Bouteloua gracilis plant are western har- 
vester ant mounds [Pogonomyrez occidentalis (Cresson)| and burrows from 
small mammals, such as skunks (Mephitis mephitis and Spilogale putorius) 


260 PHYTOL O GPA volume 67(3):258-286 September 1989 


and badgers (Tazzdea tarus). The effects of these two disturbance types on 
shortgrass communities are different. Western harvester ants remove the vege- 
tation from around their mounds by clipping the plants below the soil surface 
while small mammals produce a mound of soil at the surface that covers and 
kills the vegetation. 

Disturbance characteristics, such as size and seasonality, are correlated 
with different disturbance types and very likely affect succession. The texture 
of the soil on the disturbed site is also important to plant recovery in short- 
grass communities because of the effects of soil texture on plant community 
composition (Anderson 1983). 

Our objective was to evaluate the effects of interactions among distur- 
bance characteristics on the successional dynamics of small scale disturbances 
in shortgrass steppe plant communities by comparing naturally occurring and 
artificially produced disturbances of different type, seasonality, size and loca- 
tion by soil texture. 


METHODS 


Data collection was conducted at the Central Plains Experimental Range 
(CPER) in north central Colorado, approximately 60 km northeast of Fort 
Collins (40° 49’ N latitude, 107° 47’ W longitude). Mean annual precipitation 
over the past 45 years was 311 mm (sd = 79 mm) and mean monthly temper- 
atures ranged from -5° C in January to 22° C in July. Moderate grazing by 
cattle occurs throughout the area. 

Two disturbance locations based on soil texture and with similar climate 
were chosen: a site with a coarse textured soil (sandy loam) and a site with a 
fine textured soil (clay to clay loam). Disturbance size was studied by produc- 
ing artificial plots comparable in size and shape to the range of sizes observed 
for ant mounds and animal burrows (0.2-1.8 m?). Seasonality was evaluated 
by four recolonization dates: September 1 (1984), March 1, May 1 and July 1 
(1985). Recolonization date refers either to the date when the artificial distur- 
bances were produced or to the date when ant or animal activity ceased and 
plant recovery began on the mound or burrow. 


Western harvester ant mounds 


Previous studies in Wyoming (Kirkham & Fisser 1972) and observations 
at the CPER indicated that harvester ant mounds rarely occur on sites with 
fine textured soils; therefore, the location chosen to study plant recolonization 
on ant mounds was a site on a sandy loam soil. The surface area of each full 
size mound in a 1.5 ha area was estimated by measuring the longest diameter 
of each mound and the diameter perpendicular to the first diameter. Eight 
mounds were randomly assigned to each of the four recolonization dates (Coffin 
& Lauenroth unpublished). 


| ; 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 261 


During the time that western harvester ants inhabit a nest site, the ants in- 
hibit plant growth on the mound by clipping the plants below the soil surface. 
Therefore, it was necessary to remove the ants in order to study the recolo- 
nization of plants on the mounds. Environmental conditions in the nest were 
altered by placing a plywood box over each of the eight mounds, two weeks 
prior to the September, May and July recolonization dates. This resulted in 
the ants relocating their nests. The plywood boxes were ineffective for the 
March recolonization date since harvester ants overwinter below the soil sur- 
face and do not become fully active until late March or mid April (Lavigne 
1969). A biocide (Diazinon) was used to kill the ants of the eight mounds for 
the March date. 


Small animal burrows 


Observations at the CPER indicated that small animal burrows occur pri- 
marily on sites with coarse textured soils with a relatively low frequency of 
occurrence (0.5 burrows ha’ y~' from Coffin & Lauenroth 1988). Therefore, 
a large area (5-10 ha) on a sandy loam soil was examined during the time of 
the study (September 1, 1984-July 1, 1985) for the occurrence of newly formed 
burrows. Seven burrows were found for one recolonization date, July (1985). 
Newly formed burrows were not found for the other three dates. Observations 
at the CPER also indicated that small burrowing animals do not inhibit plant 
growth on the mound of soil produced by the burrow; therefore plant recov- 
ery began immediately after the burrow was produced. The surface area of 
the mound of soil was estimated by measuring the longest diameter and the 
diameter perpendicular to the first diameter. 


Artificial plots 


Artificial plots were produced on two locations differing in soil texture: a 
coarse and a fine textured soil. Circular plots of three sizes (50, 100 and 150 
cm diameter) were produced by removing all above and below ground plant 
material to a depth of 10 cm. The soil was sieved using a 1 cm mesh screen 
to remove all perennial organs, then replaced. Eight plots were produced for 
each site, date and size. A randomized block design was used where plots 
were located within eight 16x20 m? blocks of homogeneous soil and vegetation 
separated by 10 m borders of undisturbed vegetation within a 0.75 ha area at 
each site. For each recolonization date, one plot of each of the three sizes was 
randomly assigned to one of the 20-16 m? cells within each block. The result 
was that the blocks and plots were separated by undisturbed vegetation to 
allow each plot to be near a source of propagules from the undisturbed plant 
community. 


262 PH YT OLMG TA volume 67(3):258-286 September 1989 


Data collection and analysis 


The number of plants were counted and the percentage canopy cover was 
estimated by species in early June and late July (1985, 1986) on each of the ant 
mounds, animal burrows and artificial plots. The sampling dates were chosen 
to correspond to the maximum development of cool and warm season species. 
Density and cover values were obtained within concentric circles of 25, 50 and 
100 cm in diameter to determine the relative location of each plant within 
each of the disturbed areas. Total density on each disturbance was found by 
summing the densities of each circle. The short term effects of the disturbance 
characteristics on plant recovery were analyzed using data collected one vear 
after plant recovery began on the disturbed areas. Therefore, the data col- 
lected for the two sampling dates in 1985 were used for the September (1984) 
disturbances, while the data collected in 1986 were used for the disturbances 
of the three dates in 1985. 

Because plant recovery on western harvester ant mounds and animal bur- 
rows was only monitored on coarse textured soils and the average surface area 
of ant mounds (0.92 m*) and burrows from small animals (0.98 m*) were most 
similar in size to the 100 cm diameter artificial plots (0.78 m*), the density 
and canopy cover of plants on ant mounds and animal burrows were compared 
with the 100 cm diameter artificial plots on the site with coarse textured soil. 
The effects of the location of the disturbance by soil type, recolonization date 
and size were evaluated using the data from the artificial plots. 

Analysis of variance was used to evaluate the effects of disturbance type, 
location by soil texture, date and size on the density and cover of four groups: 
(1) all plants, (2) perennial grasses and sedges, (3) perennial forbs, shrubs 
and succulents, and (4) annual grasses and forbs. Tukey’s Q values were used 
to compute least significant ranges (LSR) and to evaluate the significantly 
different means at the P<0.05 level (Sokal & Rohlf 1981). 

The composition of the undisturbed plant communities (controls) was de- 
termined using fifty 0.25 m? quadrats randomly positioned on transects in the 
vicinity of the disturbances at each of the three sites (artificial plots: coarse 
and fine textured soils [Coffin & Lauenroth 1989]; ant mound site |Coffin & 
Lauenroth unpublished}]). The density and cover values for the plant commu- 
nities on the site with coarse textured soil and the ant mound site were used 
as the control for the effects of disturbance type and seasonality. Because the 
animal burrows were located in an area that included the ant mound site, 
the control for ant mounds also represented the control for animal burrows. 
The number of individuals were counted and canopy cover by species was es- 
timated for each quadrat. The data were collected at the same time as for 
the disturbances (early June and late July, 1985 and 1986). The values for 
the four sampling dates were used in an analysis of variance to evaluate the 
effect of soil type on the density and cover of the four groups of species for the 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 263 


undisturbed plant communities. 
RESULTS 


The undisturbed plant communities of the ant mound site and the sites 
with coarse and fine textured soils for the artificial plots were typical of short- 
grass communities (Sims et al. 1978) by having greater than 60% of their total 
density and 80% of their total cover attributed to perennial grasses and sedges 
(Table 1). The plant communities were dominated by the perennial grass, 
Bouteloua gracilis which contributed greater than 70% to the total cover. 
Other important perennial grasses and sedges were Carer heliophila Mack. 
(sunsedge) and Buchloe dactylordes (Nutt.) Engelm. (buffalograss). [Nomen- 
clature follows McGregor (1986)]. 

The plant communities at the site with coarse textured soil and the ant 
mound site were similar and significantly different from the plant community 
at the site with fine textured soil for all groups except perennial grasses and 
sedges. Density and cover of annuals [primarily Vulpza octoflora = Festuca 
octoflora (Walt.) Rydb. (sixweeksgrass)| were significantly higher on the site 
with fine textured soil, while the density and cover of perennial forbs, shrubs 
and succulents were significantly higher on the other two sites. The succulent, 
Opuntia polyacantha Haw. (plains pricklypear) and the perennial forb, Sphaer- 
alcea coccinea (Pursh) Rydb. (scarlet globemallow) were important contribu- 
tors to density and cover on all three sites, while the shrubs Chrysothamnus 
nauseosus (Pall.) Britt. (rabbitbrush), Gutzerrezia sarothrae (Pursh) Britt & 
Rusby (broom snakeweed) and Atrzpler canescens (Pursh) Nutt. (four wing 
saltbush) occurred primarily on the site with coarse textured soil and the ant 
mound site. 


Species composition and disturbance type 


The density and cover of the four groups of species were different for 
the three disturbance types, although the species found on the disturbances 
were similar. The first year after western harvester ant mounds were va- 
cated, most of the density (>70%) and canopy cover (>80%) of plants on the 
mounds were perennials (Figures 1;2), including forbs (Sphaeralcea coccinea, 
Oenothera caespitosa Nutt. (evening primrose) and Picrodeniopsis oppositifo- 
ha (Nutt.) Rydb. (plains bahia), the succulent (Opuntia polyacantha), the 
subshrub [Artemisia frigida Willd. (fringed sagewort)], the grasses [Sporobolus 
cryptandrus (Torr.) A. Gray (sand dropseed) and Sitanion hystriz Nutt. (squir- 
reltail)| and the sedge Carez heliophila. Important annual species were Vulpza 
octoflora, Plantago patagonica Jacq. (Patagonian plantain) and Lepidzum den- 
siflorum Schrad. (prairie pepperweed). For most dates, the density and cover 
of perennial forbs, shrubs and succulents on ant mounds were greater than 


264 PHY DO LOG A volume 67(3):258-286 September 1989 


Table 1. Average density and canopy cover of four groupings of species for 


three undisturbed plant communities. 


Fine Coarse 
Textured | Textured | Mound 
Soil! Soil! 


Density (nom) | 
Perennial grasses 81 72 
and sedges 
Perennial forbs, shrubs 16 19 
and succulents 
Annual grasses and forbs 14 10 


ol Pe | 


Canopy Cover (%) | 
Perennial grasses 38 38 41 
and sedges 
4* 7 8 
Annual grasses and forbs ? ig 1 1 
er a ef 
* indicates significance at P<0.05 among sites for each species group 


' from Coffin & Lauenroth (1989) 
* from Coffin & Lauenroth (unpublished) 


Perennial forbs, shrubs 
and succulents 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 265 


perennial grasses and sedges while the reverse was true for the surrounding 
undisturbed vegetation. 

Most density (71%) and cover (53%) of plants on the mound of soil from 
small animal burrows were from perennial grasses and sedges (Figures 1;2). 
Large contributions to density (23%) and cover (44%) were also made by 
perennial forbs, shrubs and succulents. The same species of perennial plants 
were found on animal burrows as for ant mounds with the addition of a large 
contribution to cover by Bouteloua gracilis (20%). Relatively few annuals 
were found on the burrows and were primarily the forb species, Chenopodium 
album L. (lamb’s quarters). Although the density of perennial grasses was 
significantly higher on the burrows than in the surrounding vegetation, the 
cover of perennial grasses was significantly lower (Figures 1;2). 

In contrast to ant mounds and animal burrows, the densities of annuals, 
perennial grasses and perennial forbs, shrubs and succulents were compara- 
ble within and among dates on the 100 cm diameter artificial disturbances 
on the site with coarse textured soil, while the total densities of plants were 
comparable among dates (Figure 1). Greater than 43% of the total cover on 
the plots was due to annuals and more than 33% was due to perennial forbs, 
shrubs and succulents (Figure 2). Important species on artificial plots were the 
perennials: Sphaeralcea coccinea, Oenothera caespitosa, Picrodeniopsis opposi- 
tifola, Carez heliophila, Sporobolus cryptandrus, Sitanion hystriz and Buchloe 
dactyloides, and the annuals: Vulpia octoflora, Lepidium densiflorum, Plan- 
tago patagonica and Chenopodium album. Significantly lower density and cover 
of perennial grasses occurred on the artificial plots than for the surrounding 
undisturbed vegetation and for most dates, the cover of annuals was signifi- 
cantly higher on the plots than for the controls. 


Disturbance type and recolonization date 


The highest total density (228 plants/m*) and canopy cover (30%) of the 
three disturbance types were found on animal burrows (Figure 3). For most 
dates, ant mounds had intermediate density (66-227 plants/m*) and cover 
values (14-25%) while artificial plots had the lowest density (29-70 plants/m’) 
and cover (8-20%). Significant effects of recolonization date were the result of 
low cover values on artificial plots produced in September. 

The density (162 plants/m*) and cover (16%) of perennial grasses and 
sedges on animal burrows were significantly higher than on ant mounds or 
artificial plots (Figure 4). Most of the perennial grasses and sedges on the three 
disturbance types were from individuals of the rhizomatous species, Carez 
heliophila. For most dates, the lowest density and cover values were found 
on the artificial plots. The low cover values for the September artificial plots 
(0.2%) were the significant date effects for perennial grasses and sedges. 

Ant mounds and animal burrows had higher density (30-116 plants/m”) 
and cover (9-17%) of perennial forbs, shrubs and succulents for all recoloniza- 


266 Bay? OL Cra volume 67(3):258-286 September 1989 


DENSITY (no./m’ ) DENSITY (no./m’) 
5) kagome 8 6B Ble 8) 3S) ee lee Sees 


ie) 
° 
= 
2) 
v 3 
m 5. 
$2) 
7 ee 
v 
o 53 
40 
o “3 
= 
uv 
2 
=] = 
Bowie UA ou 
+ 
5 
oO SmMame 
> spmoemé 
5 fF Ze? 
a 3$55 6 
pie eo ees 
+ o.e a> 
< 2m 
uv ozs 
m maw 
@e@-f 
c a ee) 
3 Cher 
e 
oO 
3 
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= > 
eo £3 
9 334 
ale 
3 . 
c 
v 
Pad z 
sf 
Dok © 
n 
+ 
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P *] o 
ro) = 
m a 
= 
< 
Uv 
m 


us 


Figure 1. Density of plants of four groups of species for three disturbance 
types and the control for four recolonization dates (a) September, 1984 (b) 
March, 1985 (c) May, 1985 (d) July, 1985. Use LSR4 for significance of an- 
nuals between disturbance types and the control and within dates, LSR,- for 
perennial forbs, shrubs and succulents, LSRg for perennial grasses and sedges, 
and LSR7 for total density. 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 267 


L8A 


BURROW 


ARTIFICIAL 
PLOTS 


MOUNDS 
SPECIES GROUP AND DISTURBANCE TYPE 


CONTROL 


LSA 


AND SUCCULENTS (F) 


E2529 PERENNIAL GRASSES (G) 


Co SOTorax (1) 


KS PERENNIAL FORBS,SHAUBS 


GEEB ANNUALS (A) 


MOUNDS 


(c) 


ARTIFICIAL 
PLOTS 
SPECIES GROUP AND DISTURBANCE TYPE 


CONTROL 


10 
i) 
60 


o 
o 


40 
20 
10 

0 


° oc ° °o 
© v ” N 


(%) HBAOD (%) HBA0D 


Figure 2. Cover of plants for four groups of species for three disturbance 
types and the control for four recolonization dates (a) September, 1984 (b) 
March, 1985 (c) May, 1985 (d) July, 1985. For explanation on use of signifi- 
cance ranges (LSR) sce Figure 1. 


268 PHY TOLOGIA volume 67(3):258-286 


(a) 


240 


September 1989 


200 
a 
= 160 
°o 
Cc 
— 
= 420 
o : 
Z “ 
wi Res 
Q 80 bs] 
~ 
BS 
RS 2 
ES 
BS 
40 ~~ 
bes 
4 
se 
2 
oO aA 
50 
(b) 
% EZ4A ARTIFICIAL PLOTS 
ro CJ ANT MOUNDS 
40 Re G8 3=6ANIMAL BURROWS 
rS<) 
3 
% 
e. st 
x] 
x 30 ote 
prs ‘ 
> are 
ve) set 
oO 20 sf 


204 


> 


a 
POOP OOOO 


10 


CONTROL SEPT MARCH MAY JULY 
DISTURBANCE TYPE AND DATE 


Figure 3. Density and cover of plants for three disturbance types and the 
control for four recolonization dates. Use LSRz for significance of type within 


dates and LSR, for significance of date within types. 


LSR 


) 
I 


| 
} 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 269 


240 (a) 
200 
~~ 
2 160 
° 
= 
— 
> 
2 120 
mh} 
ry 
80 
8 
40 bs 
“ 2 4 
<] 
re) 
50 
(b) EZ4a ARTIFICIAL PLOTS 
CJ ANT MOUNDS 
8) Ss ANIMAL BURROWS 
BP 
a 
fea 
wi 
> 
(eo) 
Oo 


CONTROL SEPT MARCH MAY JULY LSR 


DISTURBANCE TYPE AND DATE 


Figure 4. Density and cover of perennial grasses and sedges for three dis- 
turbance types and the control for four recolonization dates. For explanation 
on use of significance ranges (LSR) see Figure 3. 


270 fFoayrroryoGia volume 67(3):258-286 September 1989 


tion dates than the artificial plots (Figure 5). Sphaeralcea coccinea was the 
most important species in this group for the three disturbance types. Sig- 
nificant date effects were the result of high densities on the March plots and 
low cover values on the September plots. The density of annual plants was 
comparable on the three disturbance types, except for the high densities of an- 
nuals found on ant mounds vacated in September (Figure 6a). The percentage 
cover of annuals was significantly higher on artificial plots than on mounds or 
burrows for all dates except September (Figure 6b). Most of the cover on the 
artificial plots was attributed to two species, Lepidium densiflorum and Vulpia 
octoflora. 


Disturbance location, date and size 


Similar species were found on the artificial plots at the sites with fine and 
coarse textured soils, although the density and cover values of the four groups 
of species were significantly different. The total density of plants on artificial 
plots at the site with fine textured soil (33-211 plants/m?”) was higher for all 
sizes and all dates than the density of plants on plots at the site with coarse 
textured soil (34-81/m”) (Figure 7). Densities on the disturbed plots were 
comparable for most dates and sizes within each site. The percentage canopy 
cover on artificial plots at the site with fine textured soil was significantly 
greater than plots at the site with coarse textured soil for all dates except 
September (Figure 7c). 

Similar density and cover values of perennial grasses were found for both 
sites, although the values on the plots were less than on the controls (Figure 8). 
The 50 cm diameter plots had significantly higher density values, due to Carer 
helophila on the site with fine textured soil and Buchloe dactyloides on the 
site with coarse textured soil, than the 100 cm or 150 cm diameter plots for 
both sites and most dates. 

The density of perennial forbs, shrubs and succulents on the artificial plots 
for both sites were comparable to. or significantly greater than the controls 
(Figure 9a,b). The March plots on the site with fine textured soil had signifi- 
cantly greater density values for all three plot sizes than the plots for the other 
dates primarily because of differences in the densities of Sphaeralcea coccinea 
(Figure 9b). In most cases, the 50 cm diameter plots had the smallest density 
values of the three sizes within a date and site. Although the percentage cover 
of perennial forbs, shrubs and succulents was significantly different for the 
two controls, significant site differences were not found for the cover of plants 
on the plots (Figure 9c). The plots produced in September had significantly 
smaller cover values for all three sizes than plots for the other three dates. 

The density and cover of annuals was higher for the three plot sizes on the 
site with fine textured soil and the control vegetation than for plots on the 
site with coarse textured soil (Figure 10a,b). Significantly greater density and 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 271 


240 (a) 


120 


100 


DENSITY (no./m2) 


50 


(b) 


[ZA ARTIFICIAL PLOTS 
CJ ANT MOUNDS 
EB ANIMAL BURROWS 


“COVER (%) 


CONTROL SEPT MARCH MAY JULY LSR 


DISTURBANCE TYPE AND DATE 


Figure 5. Density and cover of perennial forbs, shrubs and succulents for 
three disturbance types and the control for four recolonization dates. For 
explanation on use of significance ranges (LSR) see Figure 3. 


272 PHY? COLOGTA volume 67(3):258-286 September 1989 


240 (a) 
N 
E 
S 
° 
= 
— 
> 
= 
0 80 
z 
TT] 
Q 60 
aa 2 4 
20 
fe) 
50 
(b) 
ZZA ARTIFICIAL PLOTS 
CJ ANT MOUNDS 
S28) =6ANIMAL BURROWS 
So 
Ce 
fea 
Ww 
> 
re) 
oO 


CONTROL SEPT MARCH MAY JULY LSR 


DISTURBANCE TYPE AND DATE 


Figure 6. Density and cover of annuals for three disturbance types and the | 


control for four recolonization dates. For explanation on use of significance 
ranges (LSR) see Figure 3. 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 273 


220 (a) 
a. 180 
£ 
S 
2 140 
a 100 
g 
w 8660 234 
a 

20 

(6) 

CI 50cM 
= CSI 100CM 
a 180 mmm 150CM 
E CZ) CONTROL 
2 140 
‘a 100 
g 
wi «=s«a6O 
Q 

20 

SEPT MARCH MAY JULY LSR 
DATE AND SIZE 
= % (c) C1 COARSE SOIL 
4 (<1 FINE SOIL 
B35 + | |S 
z K 
= 4 
& 25 ‘I 
re) 
oO 15 : 
3 S$ 
i 


CONTROL SEPT MARCH MAY ~— JULY LSR 
DATE AND SOIL TYPE 


Figure 7. Density of plants on artificial plots of three sizes for four dates, 
and the control for two sites (a) coarse textured soil (b) fine textured soil. (c) 
Cover of plants on artificial plots of three sizes for four dates. Use LSR» for 
significance of site within size and date, LSR3 for size within site and date, 
and LSR, for date within site and size. 


274 PRY TOLO GYA volume 67(3):258-286 September 1989 


3Zis QNV 3LVd 


3ZIiS ONV 3LVd 


COVER (%) DENSITY (no./m*) 


ge 

or 
z 
9 


= - -_ 
o > a 
°o °o o 


(e) 
ozz 


id38 ‘OHLNOD 


AVA HOUVAN 


ane 


ust 


1d38 OWLNOD 
(9) 


HOUVAN 


AVN 


G3 
wooot Cy 
ey 


ate 


WO 09 


os 
z 
m 
e 
2 
> 


° 
° 
> 
2 
a 
m 
o 
2 
r 


WD Oot 


ust 


Figure 8. Density of perennial grasses and sedges on artificial plots of three 
sizes for four dates, and the control for two sites (a) coarse textured soil (b) 
fine textured soil. Cover of perennial grasses on artificial plots of three sizes 
for four dates, and the control for two sites (c) coarse textured soil (d) fine 
textured soil. For explanation on use of significance ranges (LSR) see Figure 
fe 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 275 


el ik 
a 180 
Pas 
o 140 
i= 
~— 
> 
- 100 
g 
= 60 
a 2304 
20 
220 CI 50cm 
(b) SSS 100CM 
a 180 MH 150CM 
“ts CZ CONTROL 
S 
2 140 
— 
> 
FE 100 
7) 
ai 
H 60 
20 
SEPT MARCH MAY JULY LSR 
DATE AND SIZE 
45 
(c) CO 50cCM 
BSN 100CM 
35 Me 150CM 
a COARSE SOIL 
x (2777 FINE SOIL 
— 
fea 
Ww 
> 
oO 
oO 


CONTROL SEPT MARCH MAY JULY LSR 
DATE AND SIZE 


Figure 9. Density of perennial forbs, shrubs and succulents on artificial 
plots of three sizes for four dates, and the control for two sites (a) coarse 
textured soil (b) fine textured soil. (c) Cover of perennial forbs, shrubs and 
succulents on artificial plots of three sizes for four dates, and the control for 
two sites by soil texture. For explanation on use of significance ranges (LSR) 
see Figure 7. 


276 PHY PO 0 GiA volume 67(3):258-286 September 1989 


cover values of annuals (primarily Lepidium densiflorum and Chenopodium 
album) were found on the 150 cm diameter plots than on the 50 cm diameter 
plots for the site with fine textured soil. The September plots had significantly 
lower cover values of annuals than plots for the other three dates (Figure 10c). 

An analysis based on the density and cover of plants located in the 50 cm 
diameter center circle of each artificial plot was conducted to further evaluate 
the effects of disturbance size on plant recovery. Other factors besides perime- 
ter and surface area, such as microenvironmental gradients and the ability of 
plants to disperse over the entire plot, may change as plot size changes, and 
would be indicated by a difference in the density and cover of plants in the 
center of the different sizes of plots. 

The total density of plants located in the center circle of each plot was 
higher on the 50 cm diameter plots than the 100 cm or 150 cm diameter plots 
for both sites and most dates (Figure 1la,b). Although total density was higher 
on the 50 cm diameter plots, percentage canopy cover was significantly higher 
on the 100 cm (18%) and 150 cm (19%) than on the 50 cm diameter plots 
(11%). Significantly higher density and cover values were found for perennial 
grasses and sedges (primarily Carez heliophila and Buchloe dactyloides) for 
the 50 cm diameter plots than for the 100 cm or 150 cm diameter plots on 
both sites (Figure 12). Size was not a significant factor for the density or 
cover of perennial forbs, shrubs and succulents except for the high density and 
cover values on the March 150 cm diameter plots, due to Sphaeralcea coccinea 
(Figure 13). Although the densities of annuals were not affected by disturbance 
size, the cover values were significantly higher on the 100 cm (10%) and 150 
cm diameter plots (11%) than on the 50 cm diameter plots for all dates (3%). 


DISCUSSION 


The species composition on the two naturally occurring disturbances (west- 
ern harvester ant mounds and small animal burrows) were similar one year 
after plant recovery began, and were different from the composition on the 
artificially produced disturbances. This distinction between natural and hu- 
man caused disturbances was also found for grasslands and savannas of the 
Serengeti National Park (Belsky 1987). The high density and cover of peren- 
nials on ant mounds and animal burrows indicate that perennial organs, such | 
as rhizomes of Carez heliophila and tap roots of Sphaeralcea coccinea, were | 
not killed by the clipping activity of harvester ants or the pile of soil from | 
burrowing animals. Similar growth of perennial grasses and forbs on badger 
mounds was observed in the first growing season in a tallgrass prairie (Platt 
1975). 

The recolonization of western harvester ant mounds was primarily by peren- 
nials, and the density of annuals on the mounds was significantly greater than | 
on animal burrows and most artificial plots. The activities of western har- | 
vester ants may affect the recovery of plants on abandoned mounds in several 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 277 


oO 
a2 (a) 
eb 180 
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CONTROLS SEPT MARCH MAY JULY LSR 
DATE 


Figure 10. (a) Density of annuals on artificial plots of three sizes for four 
dates, and the control for two sites by soil texture. (b) and (c) Cover of annuals 
on artificial plots of three sizes for four dates, and the control for two sites by 
soil texture. For explanation on use of significance ranges (LSR) see Figure 7. 


278 PHY TOL O.GLA volume 67(3):258-286 September 1989 


130 


(a) 
110 
¥ 90 
ee 
= 
a 
= 50 
ra) 
oa 234 
10 
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DATE AND SIZE 


Figure 11. Density of plants in the center 50 cm diameter circle of artificial | 
plots of three sizes for four dates and two sites (a) coarse textured soil (b) fine | 
textured soil. For explanation on use of significance ranges (LSR) see Figure | 


a | 


{cc 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 279 


as0 (a) 
110 
= 
N 
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Figure 12. Density and cover of perennial grasses and sedges in the center 
50 cm diameter circle of artificial plots of three sizes for four dates. Use LSR; 
for significance of size within date and LSR, for date within size. 


280 PAP ODO'GTA volume 67(3):258-286 September 1989 


130 (a) 

<& 110 

E 

S90 

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SEPT MARCH MAY JULY LSR 


DATE AND SIZE 


Figure 13. Density of perennial forbs, shrubs and succulents in the center 
50 cm diameter circle of artificial plots of three sizes for four dates and two 
sites (a) coarse textured soil (b) fine textured soil. For explanation on use of 
significance ranges (LSK) see Figure 7. (c) Cover of perennial forbs, shrubs| 
and succulents in the center 50 cm diameter circle of artificial plots of three) 
sizes for four dates. For explanation on use of significance ranges (LSR) see 


Figure 12. | 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 281 


ways due to the interactions of ants with their environment during the 30-60 
years that a colony may inhabit a nest site (Wiens 1976; Coffin & Lauen- 
roth unpublished). The foraging and clipping activities of the ants affect the 
microenvironmental conditions on the nest as well as the availability of repro- 
ductive and vegetative propagules. 

Nutrients and seeds are concentrated in a nest as a result of the foraging 
and storage of seeds and other organic materials in underground chambers by 
harvester ants, and soil water is significantly higher (P<0.05) in a nest than 
at points away from the nest due to the clipping of all plants from the mound 
(Rogers 1974; Rogers & Lavigne 1974). The significantly greater (P<0.05) 
standing crop of plants surrounding a mound compared to points located away 
from the mound (Rogers & Lavigne 1974) may also result in an increased avail- 
ability of seeds and vegetative propagules to an abandoned mound compared 
to disturbances that are recolonized immediately after being produced (artifi- 
cial plots and animal burrows). The number of germinable seeds found near 
the soil surface of mounds in September, 1984 (6756 seeds/m* from Coffin & 
Lauenroth unpublished) was significantly greater (P<0.05) than the number 
of seeds in the soil at the site with coarse textured soil for the same date (2748 
seeds/m? from Coffin & Lauenroth 1989). The length of time these conditions 
persist on an abandoned mound is unknown. However, the largest effects on 
plant recovery most likely occur in the short term. 

The 6% cover by the dominant plant species, Bouteloua gracilis, on animal 
burrows was primarily due to the vegetative growth of B. gracilis plants par- 
tially covered by soil. Bouteloua gracilis was not found in significant amounts 
on ant mounds or artificial plots. These results suggest that the time required 
for B. gracilis to dominate the plant community on animal burrows will be less 
than for ant mounds or artificial plots, and less than the estimated recovery 
time of greater than 50 years for B. gracilis to dominate abandoned agricul- 
tural fields (Reichhardt 1982). The recovery time for abandoned fields was 
based on the observation that it took 33 years for B. gracilis to occur on the 
fields and another decade to reach 2% in frequency of occurrence (Reichhardt 
1982). 

In contrast to ant mounds and animal burrows, the majority of the cover 
on the artificially produced disturbances for the sites with coarse and fine 
textured soils, and most dates and sizes, was attributed to annuals. This is 
similar to plant communities found on old roads and abandoned agricultural 
fields in the shortgrass steppe region within five years after the beginning 
of plant recovery (Shantz 1917; Costello 1944; Judd 1974; Reichhardt 1982). 
Perennials colonized the artificial plots in the first year as indicated by the 
comparable densities of annuals, perennial grasses and sedges, and perennial 
forbs, shrubs and succulents. 

The recolonization date and size of the disturbance had important effects 
on the density and cover of plants on artificial plots. The effects of recoloniza- 


282 Pry Io b OGrA volume 67(3):258-286 September 1989 


tion date on short term successional dynamics are due to interactions between 
the availability of propagules to the site and the microenvironmental condi- 
tions on the site relative to the requirements for germination, establishment 
and growth of the propagules (Sousa 1984). The low cover of annuals and 
perennials on artificial plots produced in September may be a combination 
of: (1) the seasonal dynamics in the relatively few germinable seeds stored in 
the soil at the two sites (964 seeds/m? averaged over two years from Coffin 
& Lauenroth 1989), (2) the low and variable patterns of precipitation found 
in the shortgrass region (Sala & Lauenroth 1982), and (3) the timing of the 
precipitation events relative to factors such as temperature. Because of the 
variability in precipitation within and among years, conducting the study in 
another year would most likely indicate the timing of the disturbance to have 
important effects on plant recovery, however, the specific effects observed in 
this study may not occur. 

Although annuals and perennials responded similarly to the effects of re- 
colonization date, they responded differently to disturbance size. The size of 
the disturbance may be important to the availability of propagules to the dis- 
turbed site and the resources available to the propagules. As disturbance size 
increases, propagules must disperse over larger distances to colonize the entire 
disturbed area, while competition from plants around the edge increases as 
the size of the disturbance decreases (Sousa 1984). 

The source of propagules for annuals included seeds stored in the soil and 
the dispersal of seeds onto the plot. The density of annuals was not affected 
by disturbance size while the effects of competition by plants surrounding the 
plots on the growth of annuals was indicated by the significantly smaller cover 
values on the smallest compared to the largest plots. 

Perennials may recover either from seedling establishment or vegetative 
growth. Relatively few perennial grass or perennial forb, shrub and succulent 
seeds were found stored in the soil compared to annual seeds (Coffin & Lauen- 
roth 1989). Most of the perennials on the plots were the result of vegetative 
growth by Carez heliophila, Buchloe dactylotdes and Sphaeralcea coccinea. The 
recovery of C. heliophila by rhizomes and B. dactyloides by stolons occurred 
as ingrowth from the edge of the plots. The greater perimeter to area ratio 
of small compared to large plots (Miller 1982; Sousa 1984) resulted in signifi- 
cantly greater cover and density values of perennial grasses and sedges on the 
smallest plots for most dates and both sites. 

The cover of the perennial forb, Sphaeralcea coccinea, responded similarly 
to disturbance size as annuals by having the highest values in the center circle 
of the largest plots, or the farthest distance from potential competitive inter- 
actions with plants in the surrounding undisturbed community. Although the 
density of annuals was not affected by disturbance size, the density of S. coc- 
cinea responded similarly to the cover values. The recovery of this important 
forage species (Hyder et al. 1975) was primarily from deep tap roots either 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 283 


from plants at the edge of the plot or by regrowth of plants on the plot. 

The interaction between disturbance size, recolonization date and location 
by soil texture was also important to the cover and density of Sphaeralcea 
coccinea. The period of growth of this species (April and early May from 
Dickinson & Dodd 1976) corresponds to the significantly greater density and 
cover values on the largest plots cleared in March on the site with fine textured 
soil for perennial forbs, shrubs and succulents than for plots of the other dates. 
Similar results on the importance of the timing of the availability and growth 
of propagules relative to the seasonality of the disturbance were found for the 
recovery of an annual grassland following small disturbances (Hobbs & Mooney 
1985). 

Although the long term monitoring of the disturbed sites for the three dis- 
turbance types and their associated characteristics is necessary to evaluate the 
time required for each disturbed area to be dominated by a shortgrass plant 
community, the results after one year of recovery suggest that animal bur- 
rows will have the most rapid recovery time of the disturbance types studied 
because of the presence of Bouteloua gracilis and other perennials on the bur- 
rows. Artificial plots, and in particular the largest plots, will have the slowest 
recovery time since relatively few perennials were found on the plots compared 
to ant mounds or animal burrows. 


ACKNOWLEDGMENTS 


A number of people assisted with the field work, including R.D. Wuertz, 
C.A. Lee, L. Yimin, D.G. Milchunas and D. Hazlett. G. Richardson provided 
assistance with the statistical analyses. We thank two reviewers for comment- 
ing on the manuscript. This study was supported by the National Science 
Foundation, Shortgrass Steppe Long Term Ecological Research Project (BSR- 
8114822) and the Colorado State University Experiment Station Project, Long 
Term Grassland Ecosystem Research. 


LITERATURE CITED 


Albertson, F.A. & J.E. Weaver. 1944. Nature and degree of recovery of grass- 
land from the Great Drought of 1933 to 1940. Ecol. Monogr. 14:394-479. 


Anderson, M.D. 1983. Soil and vegetation pattern on shortgrass catenas. 
M.S. thesis. Colorado State University, Fort Collins, Colorado, USA. 


284 PHYTOLO ERA volume 67(3):258-286 September 1989 


Belsky, A.J. 1986. Revegetation of artificial disturbances in grasslands of 
the Serengeti National Park, Tanzania. I. Colonization of grazed and 
ungrazed plots. J. Ecol. 74:419-437. 


Belsky, A.J. 1987. Revegetation of natural and human-caused disturbances 
in the Serengeti National Park, Tanzania. Vegetatio 70:51-60. 


Coffin, D.P. & W.K. Lauenroth. 1988. The effects of disturbance size and 
frequency on a shortgrass plant community. Ecology 69:1609-1617. 


. 1989. Spatial and temporal variation in the seed bank of a semiarid 
grassland. Amer. J. Bot. 76:53-58. 


. (submitted). Vegetation associated with nests of western harvester 
ants (Pogonomyrez occidentalis Cresson) in a semiarid grassland. Amer. 


Midl. Naturalist. 


Collins, S.L. 1987. Interaction of disturbances in tallgrass prairie: A field 
experiment. Ecology 68:1243-1250. 


Collins, S.L. & G.E. Uno. 1983. The effects of early spring burning on 
vegetation in buffalo wallows. Bull. Torrey Bot. Club 110:474-481. 


Collins, S.L. & S.C. Barber. 1985. Effects of disturbance on diversity in 
mixed-grass prairie. Vegetatio 64:87-94. 


Costello, D.F. 1944. Natural revegetation of abandoned plowed land in the 
mixed prairie association of northwestern Colorado. Ecology 25:312-326. 


Davis, R.M. & J.E. Cantlon. 1969. Effects of size area open to colonization 
on species composition in early old-field succession. Bull. Torrey Bot. 


Club 96:660-673. 


Dickinson, C.E. & J.L. Dodd. 1976. Phenological pattern in the shortgrass 
prairie. Amer. Midl. Naturalist 96:367-378. 


Hobbs, R.J. & H.A. Mooney. 1985. Community and population dynamics of 
serpentine grassland annuals in relation to gopher disturbance. Oecolo- 
gia (Berlin) 67:342-351. 


Hyder, D.N., R.E. Bement, E.E. Remenga & D.F. Hervey. 1975. Ecological 
responses of native plants and guidelines for management of shortgrass 
range. United States Department of Agriculture-Agricultural Research 
Service Technical Bulletin Number 1503. 


Judd, I.B. 1974. Plant succession of old fields in the Dust Bowl. Southw. 
Naturalist 19:227-239. 


Judd, I.B. & M.L. Jackson. 1939. Natural succession of vegetation on aban- 
doned farmland in the Rosebud soil area of western Nebraska. J. Amer. 
Soc. Agron. 39:541-547. 


Coffin & Lauenroth: Disturbance and succession in shortgrass prairie 285 


King, T.J. 1977. The plant ecology of ant-hills in calcareous grasslands. I. 
Patterns of species in relation to ant-hills in southern England. J. Ecol. 


65:235-256. 


Kirkham, D.R. & H.G. Fisser. 1972. Rangeland relations and harvester ants 
in northcentral Wyoming. J. Range Managem. 25:55-60. 


McGregor, R.L. 1986. Flora of the Great Plains. University Press of Kansas, 


Lawrence, Kansas. 


Milchunas, D.G., W.K. Lauenroth, P.L. Chapman & M.K. Kazempour. (sub- 
mitted). Community structure relationships along a multi-disturbance- 
type gradient. Vegetatio. 


Miller, T.E. 1982. Community diversity and interactions between the size 
and frequency of disturbance. Amer. Naturalist 120:533-536. 


Perozzi, R.E. & F.A. Bazzaz. 1978. The response of an early successional 
community to shortened growing season. Oikos 31:89-93. 


Pickett, S.T.A. & P.S. White. 1985. Natural disturbance: The patch dynamics 
perspective. Academic Press, New York. 


Platt, W.J. 1975. The colonization and formation of equilibrium plant species 
associations on badger disturbances in a tall-grass prairie. Ecol. Monogr. 


45:285-305. 


Rapp, J.K. & D. Rabinowitz. 1985. Colonization and establishment of Mis- 
souri prairie plants on artificial soil disturbances. I. Dynamics of forb 
and graminoid seedlings and shoots. Amer. J. Bot. 72:1618-1628. 


Reichhardt, K.L. 1982. Succession of abandoned fields on the shortgrass 
prairie, northeastern Colorado. Southw. Naturalist 27:299-304. 


Rogers, L.E. 1974. Foraging activity of the western harvester ant in the 
shortgrass plains ecosystem. Environmental Entomol. 3:420-424. 


Rogers, L.E. & R.J. Lavigne. 1974. Environmental effects of western har- 
vester ants on the shortgrass plains ecosystem. Environmental Entomol. 


3:994-997. 


Sala, O.E. & W.K. Lauenroth. 1982. Small rainfall events: An ecological 
tole in semiarid regions. Oecologia (Berlin) 53:301-304. 


Shantz, H.L. 1917. Plant succession on abandoned roads in eastern Colorado. 
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Sims, P.L., J.S. Singh & W.K. Lauenroth. 1978. The structure and function 
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characteristics. J. Ecol. 66:251-285. 


286 PHY TOLOGIA volume 67(3):258-286 September 1989 


Sokal, R.R. & F.J. Rohlf. 1981. Biometry. Second Edition. Freeman, New 
York. 

Sousa, W.P. 1984. The role of disturbance in natural communities. Annual 
Rev. Ecol. Syst. 15:353-391. 

Weins, J.A. 1976. Population responses to patchy environments. Annual 
Rev. Ecol. Syst. 7:81-120. 


Phytologia (October 1989) 67(4):287-288. 


BOOK REVIEWS 


Michael J. Warnock 
Department of Life Sciences, Sam Houston State University, Huntsville, 


Texas 77341 USA 


Atlas Florae Europaeae Distribution of Vascular Plants in Europe, Vol. 
I, Part 1, Pteridophyta (Psilotaceae to Azollaceae); Part 2, Gymnosper- 
mae (Pinaceae to Ephedraceae). Jaakko Jalas & Juha Suominen, eds. 
Cambridge University Press, 32 East 57th Street, New York, NY, 10022, 
USA. 1989, 121 pp. (Part 1), 43 pp. (Part 2), + 4 pp. appendix, $59.50, 
cloth. ISBN 0-521-34270-8. 


Atlas Florae Europaeae Distribution of Vascular Plants in Europe, Vol. 
II, Part 3, Salicaceae to Balanophoraceae; Part 4, Polygonaceae; Part 
5, Chenopodiaceae to Basellaceae. Jaakko Jalas & Juha Suominen, eds. 
Cambridge University Press, 32 East 57th Street, New York, NY, 10022, 
USA. 1989, 122 pp. (Part 3), 69 pp. (Part 4), 124 pp. (Part 5), + 4 pp. 
appendix, $69.50, cloth. ISBN 0-521-34271-6. 


Atlas Florae Europaeae Distribution of Vascular Plants in Europe, Vol. 
III, Part 6, Caryophyllaceae; Part 7, Caryophyllaceae (Silenoideae). 
Jaakko Jalas & Juha Suominen, eds. Cambridge University Press, 32 
East 57th Street, New York, NY, 10022, USA. 1989, 168 pp. (Part 6), 
237 pp. (Part 7), + 4 pp. appendix, $89.50, cloth. ISBN unknown. 


These three volumes, which parallel volumes of the Florae Europaeae series, 
compile records of geographic distributions of the plants of Europe. The range 
of each taxon is shown on a map of Europe (or in some cases a map of part 
of Europe). Distribution information is coded onto a 50 km grid system and 
a symbol placed on the map corresponding to the grid location if a represen- 
tative of the taxon has been reported from within the unit. Due to landform 
irregularities (particularly coastlines and islands), some deviations from the 
strict pattern of the grid system are necessary and these deviations are cited 
in the introductory matter for each volume. Taxonomic differences from the 
Florae Europaeae are also noted in each volume. The methods employed in 
producing these maps have some inherent shortcomings, as were pointed out 
by Holub (Norrlinia 2:107-115 {1984]) and summarized in the prefaces of vol- 
umes J and II. Many of these shortcomings are surmountable and should allow 
any future editions to be much improved. The maps are easy to read and plant 


287 


288 Pony PCO OCG aa volume 67(3):287-288 September 1989 


distributions may be observed quickly. The appendix in each volume is a set 
of 2 maps showing the location of the grid units used in the mapping system. 
A single page landform map in each volume might be helpful to workers not 
familiar with European landforms. Also, a single set of page numbers within 
each volume would make the series easier to use and simplify citations. The 
most significant difficulty I experienced with these three volumes is that the 
first 58 pages of my copy of volume III are the first 58 pages of volume II. | 
trust that this is an error in binding which did not occur in many copies. 


McGraw-Hill Concise Encyclopedia of Science and Technology, 2nd 
ed. Sybil P. Parker, ed. McGraw-Hill Publishing Co., New York. 1989, 
Ixxvi, 2222 pp., illus., price unknown, cloth. ISBN 0-07-045512-0. 


“Concise encyclopedia” strikes me as a non sequitur, much like “jumbo 
shrimp” and “army intelligence.” It is probably impossible to be both ency- 
clopedic and concise in the same document. This book suffers from the attempt 
to achieve that goal. The terms that are treated in the work are given lengthy 
definitions and descriptions of their use, and many are accompanied by il- 
lustrations. However, in order to treat terms at length and cover the entire 
spectrum of science and technology, only a relatively limited set of terms may 
be considered. Therefore, many useful terms are left out. The method for 
selection of terms is not clear. It would seem that defining the more general 
terms and omitting the more narrow would be appropriate for this work but, 
this does not seemd to be the case. For instance, lichens are included, symbio- 
sis is not. On the other hand, superfluous information is included for some of 
the definitions. For example, the entry for each element in the periodic table 
is accompanied by a copy of the periodic table, occupying considerable text 
space. Another copy of the table is found in an appendix. Significant space 
would be saved by simply referencing the table in the appendix for each ele- 
ment. McGraw-Hill’s Dictionary of Scientific and Technical Terms (reviewed 
in Phytologia 67([2]:208. [1989]) would be a better purchase. 


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